Reptiles of

b
Central America
-%A?
4
Cover photos:
Front cover, leR: Polychrua gui3umsus (Rio San Juan, Nicaragua).
Right top: Bdhdechis aurifer (Guatamala).Photo: M,Burger
Right bottom: Diploglossrru mncrtmpis. Photo: R. W,Van Devender
Photo p. 4: Cloud forest ((CerroK i l W , Jmotega,Nicaragua).
Back cover: Rain forest 15aslaya National Park, AtlWco N o h , Nicaragua).
Lampropeltis trhngulum (Patuca National Park,Blancho, Honduras).
MAW
GZL
656
Al
K6L\ )IC
200 3
KiShler, Gunther
Reptiles of Central America
Ofbnbach:Herpeton Verlag, 2003
ISBN 3-936180-02-4

Rditorial &am% rd Gutman,WBat fZartford; J. R.1McCmniefi P e k StdRrd, London;

HarZaa Walley, bkalb; Lg12.3David w
itson,M i d .

AU dghC resewed.
No part of thiB publicertion may be m p r o d u d or trr&tM in any form or by any m e w ,
electronic or mecPlaxrical, inel*. mw,
phet;a~~py, or m y iof~~mation atorage and
reMwal system, wikhout p~rd~rgien in writing &am the publiahe~
No liability is asrsmed with res@ to the w e .of infornwtiun con- herein,

Q 2003 Herpetan, Verlag Elke KZlhler,

Rohtr. 22, D-63075Offenbach,Germany
Photo8 and drawhgefe:Gunther K&hler,if not otherwise indicated
Layout: Eke Ktihler, Wenbach
Reptiles of Central America
Gunther Kohler
Gunther Kiihler

Reptiles of Central America

644 Color photographs

103 Drawings and diagrams
210 Maps

H ~RP~TVN
Verlag Elke Kiihler
Contents

Foreword ........................................................... 6 -WormLizards (Amphisbaenia) .. 166

Intdudion ............-....... ............................... 9
Comments on the Urns o f t h i s Book ..... 10 Snaker (Herpentee) .....................-*....168
Claseiileation of the Reptilee of Blind Snakes (Scolecophidia) ............ 169
Central America ............................................ 12 Anomalepididae .......................................1.69
The Environment .................e..,........nmnmnm16 ..... Leptotyphlopidae ..+....... .*...............
*-. ...... 171

Geological Hietory and Zoageo~aphy Typhlopidae ..,............,............*....... ..u..... 172

of the Reptile Fauna ......,&........................... 24 Macrostmata .......................".... ..............
174
Boidae ...............................~................ . . 174
hmemidae .......................................... 179
.
.
Tropidophiidae ...................... ..............179
Turtles (Teskdhssi) ........,......... ..... 33 .
Colubridae ......*..........*.. ., ..*......*......181
Chelydridae ........................ 34 ............ 281
Elapidae ..*................*......v.~+.....~.*.+
Dermatemydidae -................., , , .. .
........... 35 Viperidae ......-.....,.,..... -... .................. 291
m.u.

- ............ 36
Emydidae .* ....................*....-...**...
Kinostemidae .................................................. 41 Table of &lation Charactera in
Testudinidae ..................................................... 46 S n h .........................................................306
Cheloniidae, Dermochelyidae .....*.."......... 47 Snakebite - Prevention and
First Aid ..........-....... ...... 312
Liszardr (Sauria) ..-... ...........................-..
64 Mimicry in Snakes ................................. 316
An(yidae ....................................
~~............... 56 Acknowledgments .......... .#. ...-.................
8gO
Eublepharidae ................................................ 67 Literature Cited ............*.......................... 322
Gekkonidae ............................................ ......... 68 Glossary ...........*.........*........... ......*...... 368
..ws

*..
Gyeanophthalmidae .........*..........., .......... 80 ..
Index ................. .......................l...,..t......~...
361
.....
Helodematidae .........n....n.r .,.* .................. 89
....
Iguanidae ..a.....w............a"* ........ .......... 91
. ...........
Seincidae ..................-.................., ...,. 147
. . .164
Teiidae ...,....~...........................V..+............
,..
Xantuaiidae ....*..U.................,........ .............162
.
Xeno~auridae............-.............. .......~.......‘.......
164
Foreword

Middle America constitutes one of the adopted language, English. Although the
biodiversity hotspats of the world- region book is entitled The Reptiles of Central
characterized by a high level of species America,it covers the same area as its pre-
richness and endemurn. It is understand- decessors. The presentation of thi~sinfor-
able, thus, that thh area, extending from mation in English should increase its
the Isthmus of Tehuantepec in Mexico to appeal to a broader readership.
the southern extent of the Panamanian
isthmus, should have attracted considera- The two previous bocks dealt with 3 mco-
ble interest in this era of biodiversity dilians, 25 turtles (including six marine
decline, The reali~ation among natural s p i e s ) , 215 b a r d s , three amphiebaeni-
scientists that the earth's biota is under ans, and 268 snakes. The present work
attack by force5 most humans would increaser the species number by fihen
describe as involving simply the business lizards and eight snakes to a total of 537
of living has generated an unprecedented speciea. This increase of 23 species has
interest in the cataloguing of the extent of occurred, however, within the last two
that biota before it dieappears. years! With the continuation of the herpe-
tofaunal surveys currently underway
The herpetofauna of Middle America, too, throughout Middle America, the number of
is being studied by a multinational group constituent species will continue to rise. As
of specialists, including herpetologists it stands now, however, the current total of
&om the length and breadth of Mexico and 537 epecies is only 55 fewer than the 692
Central America, the United Stabs, and, species listed on the Center for North
in the case of the present work, Germany. American Herpetology website for all of
T h i ~study has resulted in the publication the United Stat= and Canada! Clearly,
witbin the last decade of a number of Middle America has a very rich herpeto-
detailed faunal work by Julian C. Lee, fauna.
James R. McCranie and me, and Jay M,
Savage and mrnmary works by Jonathan The present book cuntinues the publishing
A. Campbell, Celina DueAas and co- tradition begun by Dr. Kfihler in 1993 with
worker~,Roberto Ibaiiez and ca-workers, a book on b ~ i l i s k of
s publishing his major
and the author of the current work, works under the banner of Herpeton
Guether KBktler on varloug portions of Vexlag EIke Ktihler, the publishing com-
Middle America. pany operated by his spouse. In the last
decade, Dr. Ktihler has produced, some-
These works have formed the basis for the times alone, sometimes with coauthors, an
two predecessors of the c m n t work, as impressive series of colorful boob of both
well as this work itself. In the year 2000, scholarly and popular interest on subjects
Dr. KWer published the first volume of ae broad-ranging as reptile egg incubation,
the Reptilien und Amphibien Mittel- systematics of spiny lizards (Sceloporus)
amerikas, which dealt with the craco- and spiny-tded iguanas (Ctenosaum),and
diliana, turtles, and lizarde of this region. the systematics and distribution of the
He foll~wedthis book in 2001 with a members of the Nicaraguan herpetofauna,
second volume dealing with the snakes of T b Herpeton Verlag wdbsite, in fact, cur-
h~liddleAmerica. These two books, written rently lists ten books under Dr. Ktihler's
in Dr. KZlh1er"s native language, are now authorship, two of which are German and
combined, updated, and tramdated inta his Spanish wrsiona of a treatment of the bio-
logy af Iguana igwrna, tihe Green X@tuaaa. the twelve bh,100 popular articles,and
%vwd of them books am dimtad toward 110 ~ E i ~ ~ ~ tpaper8 i f i c on reptiles and
h&rpehulturheSr,whereas othem target amphiktians he h a p d u d to data
p d k h n a l md matera herwI;o@k
0 x 1 a more prsonal naEe, I am pleased
Gnntber mhlw is the Head of the call Cunthwr KWer both cull- and
hp&ment af HerpebIogy at the friend. Remnt1~we bBth h d a h o e to
Rmearcb 1~1~CituteSendsenberg in Piaptidpa@ in the Memmsriean W e w
F F am Mdn,~ Germany, d Wotk~fiopfor the IUCN-enpportsd Global
Direahor bf the C@memakiomM e e t Utfla Amphibian Worhhop, which OM:^ at
Iguana, TXdmm. HPi~ continuing a ka- the La Selva Bialogical5,tatiw in eastern
dition ehat ~rIEtmdsback t;o the days d Cmta Ria. W e them, in addition t~par-
O s h r Btittger and, morvz m n t l s Robert tidpathg in very impartant work of
Mertens, mhler ig wnthuing and deb- t h conservation
~ &atus of the
expanding cra tha inbest M w k m had in almwt 700 q a ~ i e gof amphibians rnctapy-
kbe herpebfama of Middle Aznwkea, @ape= ing M&CQ and Central b e d c s , ave bad a
dally that of El Balvador, Currently, &an@ b O BM ~ W p while sam-
Kiihler ia carrying: on sw&al lnajor pro- bling Wt in arearcb d amphibians and
je& dealing With the bez-petafaauna d E f . reptllw to photagraph an the station
Salvador (in ~01labtlraiit-m with Eli grounds. During the five day$ we apnt at
ICTmenbaum md i?dIiImVBSI~~), Hmdufa~ itshe station, the m e we had k e n in
(in mllabmtian with James R. McGPaafa h &Id together, I b a m e f m p l c e ~ dwith
auzd ad,md N i m m , and the biology Dr. EE3Jhler'~ emthusiwrn, dedication,
and mmm&on af the Utila Spiny-tailed b a d t h of bwldge a h t t b Middle
Iguana, Cenosaum baksni. Siaee 1995, American hergetafama, and even Ma m-
when he was a w M his Ph.D. and be- deathl. EEe holds h t h %.D. and DVM
m k d the Re~ear&httitute 8-m- degmt3i
berg, be bas nipidly made a narne fir him-
self @ a msy'orplayer in the studies on the It was &Q dur!iq this time at La Laelva
herpettofauna d bipldiverse Middle that we to trace tbts outh~biof a
America. T b aeflesw~n,t i~ mideneed by ppsjsCt that PVill l d t;O mothar book, tbiB
one an the am@biw and reptiles af the
Honduran Bay Islaoda and Caym
Cwhinos, with our mutual eolleaguie
Jamea It. Wraefe. I Ibok farward to thb
pmjact with mticip~ttion.
The book f~ which this fe the fmword ia a
maaterfd elpkhesis of information fram
noti only the ttffio b k e that preceded it, but
a h o from the broad 812'ay d ElcienM~1 liter-
Zmctnu, &a$ appwmd bath before and &
tbe publication of its predecessors, produe
ing a Literature Cited seotion*ofmare than
1000 entries. T ~ spsrtiedi
B U s and k e y for
identifhdon have 41 beren revised and
updated and will prOve useful to mpne
who hm a dmbt about what herp is in
hand The text is accompanied by profes-
sional quality color phato&rarph%of about
400 of the 537' specie& and expertly dime
 Foreword

Rg. 2. Bothriechis bicdor (SanM e 1 Pie de la Cueata, San Guatemala).

line drawing8 of diagnostic features. En This foreword wm written while I w w on a

addition, there are many attractive habitat research Ieave at the University of Miami,
photos and generdized distribution maps which leave was supported by NIGMS
that will broaden the reader's understand- (National Institute of General Medical
ing of the places these creatures live. The Science) Grant No. 1R2SGM0083-02to Dr.
6 and 112 by 9 inch eize also i~ convenient Michael S. Gaines, chairperson of the
for portage into the field. Thua, finally, Department of Biology.
after more than three deeades, we have a
fitting modern treatment of the herpeto-
fauna of Middle America, which has been Larry David Wi1.son
wanting since the cIassic work by Jamea L.
Peters and his oo-workers, covering only the Miami, Florida
squaraate reptiles, appeared in 1970 and 12 January 2003
quickly grew out of date. Middle America is
an important reservoir of berpetodiveraity,
much of which evolved in aitu, and it de-
serves the kind of attention Gunther
Whler has provided. Thie book will prwide
the basis for more extensive treatments of
this region's herpetofauna that are needed
to demonstrate the vast biogeographic
significance of thia region and the need to
protect it for paptuity.
Introduction

The present book, which is baged on

KC)HLF:R(2000a, 2001a), attempts to pm-
vide a summary of the widely scattered
information on Central American reptiles.
The region that ie covered by t h k h k
includes the seven countriels that masti-
tute political Central America (Belize,
Guatemala, El Salvador, Honduraa,
?
isthmus d
Teh
9
wntftpec
ns ~ ~ ~ I O J
Nicaragua, Costa Rica, and Panama), ae
well as the aoutheaatern part of Mexico
(the area east of the Isthmus of PldRc Owan
Tehuantepec, including the Yucatdn
Peninsula; Fig. 3). This part of Mexico
belongs ta the zoogeographically impar-
tant Central American landbridge (SAVAGE fig*3. The soo~eogra~hic region of Central
1982). Although the Swan lslands belong America (shaded).
politically to Honduras, they are not inclu-
ded because they are biogeographically
part of the West Indiee, instead of Central
America. (erg, ERNST& BARBOUR1989, IVER~ON
1992), lizards (e.g, UZZELL 1965a, 1966b,
The history of herpetological research ie ECHTERNAC~ 1971, OITEDAL1974, HARRIS
Central America datea back to the early & KLUGE1984, HARRIS19441,snakes (e.g.
nineteenth centq Early mileetones in DUELLW 1958, Dams 1967, MYERS
the history of Central American hewtolo- 1974, CAMPBELLL, LAMAR 1989, CWBELL
gy were the 'Herpetologia Mexicans" of' & BRODIE1992, R ~ Z E1996, SAVAGE &
Arend F. A. W ~ G M A N N(18341, the SLOWINSKI 19963.Additionally, a number of
"Biologia Centfali-Americani" of Albert C. regional checklists and wmpilatury works
L. G. G ~ H E (1885-1902)
R and the (MEYER& W a s o ~1973, H. SMITH& S m
"Mission Scientifigue au Mexique et dms 1973, 1876, 1977, 1979, SAVAGE& VU
l 0 h 4 r i q u e Centrde" of D~M&RIL,1986, CAMPBELL& VMWINI 1989, KC)HLER
Bocom~& MOCQUARD (1870-1909). Early 1 9 9 9 2001b,
~~ P E ~ SZ m s 1999), well
mile~tonesof Central American snake as thorough monographs of selectad wun-
literature include BOULENGER'S Cata- tries, have been published (TAYLOR 1961,
logpee on the turtlea, liza& and snakes 1954, E 1996, CAMPBELL 1998a,
in the British Museum (BOULENGEB 1885, STAFF~RD & MEYER2000, SAVAGE2002).
1889, 1893, 1894, 1896a). In 1970, Finally, in 1999, a masterful taxonomic
important w o r b such as the two volume reference for the snakes of the world
"Catalogue of Neotropical Squarnatan of (Colubriclae not treated in the first volu-
Jaznes A. Fkmms and m-workers appeared me) was puhliahed t M c D w m et al.
In the interval of about SO yews since 1999). I generally follow the taxonomic
these two publications, a large body of arrmgement proposed in that work.
papers dealing with the variow mpects of
Central American reptiles has been Central America is the link between North
pubhhed. Thia body includes worh on and South America and is af great zoogeo-
certain genera or larger groups of turtles graphic importance because its fauna con-
9
b i n s species fram both continents, in addi-
tion to a high number of endemic species.
As pmaently understood, the reptile fauna
of Central America consists of 533 species
representing 163 genera and 29 families
(Table 1). This enormous diversity is due
to the exceptional vegetational variation
found in the region, which makes it a
mosaic of rain forests, pine forests, cloud
forests, dry forests, cactus forests, eavan-
nw,swamps, and mangroves. This divemi-
t 5 and the fact that some areas have so far
not been ecientifically explored, renders
Central America a great challenge for her-
petological research. Much research
remains to be done on the diversity,
systematics, and ecology of Central
American reptiles. The recent discovery of
new taxa has even necessitated the
description of new genera ICAMPBU &
SMITH1998, KOHLER e t a]. 2001b).
However, both scientific investigation and
efforts to pmseme biudivermty are in a
race against time as the pace of habitat
destruction increases in Cenkal America.
Table 1,Taxanomic composition of the reptile fauna of Central America.
Comments on the Usage of
this Book
The intent of this work is to provide an
identification guide for both scientists and
travelers ta the numerous reptile species
found in Central America, Many Central
American s e e s of reptiles are are distinc-
tive in coloration, as well as body size and
ahape, that an animal in question often
can readily be identified by photographs or
drawings alone. If the identification remains
questionable by this method, identification
keys should be consultd. Additionally, the
distribution maps can be included in the
identification process, because many
species have very restricted distributiom.
The identification keys included in the
present book consider all families, genera,
and species that are known to occur in
Central America, even though this is not
explicitly stated in the respective key hea-
dings. Subspecies are mentioned i x ~the
species accounts, but are not considered in
the identiflation keys or in the distribution
maps, The keya are dichdomouq meaning
that they provide paire of contrasting
1
alternatives. .Such a key works using step
by step elimination of alternatives until
only a sifigle alternative remains. It ie ed by the citation af the original descrip
important to remember when wing the tion and bype locality; the maximum size
idmtBcation key that the juvenilee of (carapace length, total length rrL] or
maay lizard and snake species have dif- snout-vent length [SVL]),and the known
ferent coloration and pattern than the geographic distribution. For wme ~petriea,
adults. Any key baaed on dhtinguiehing additional notes on ecalogy and reproduc-
coloration will not be able tb definitively tion are included, Detailed anatomical
identi@ the juveniles. The technical terns dehimiptioas are beyond the scope of the
and rnorphologicd character d e s i p a t i mbaak. The recommendations for further
used in the keys e m be ht;rating to non- reading a t the end of each generic account
speei&ts, but I am optimietic that anyone m e the readar as a guide to the addition-
willing to inveet gome effort in learning theal refeferencecr.The apeciea accounh am eha-
terms will be able to uee the keys with con- ded with gray for easy recognition. The
fidence and succeae. The glossary included index at the end of the book will help in
in the book will be helpful in this regard, as
finding the pages w h the keye, speciciee
win the illuak.ertians accmpanyiag the accounts, photographs (bold page nrun-
~ Y B . bra), md distribution maps (italic page
numbere) of the respective species are loca-
Following the introduction to each major ted. The keys and maps included in t b i ~
m u p of Mtrd American reptilee, the f ~ l - book are based mostly on the information
lowing information i6s pmvided in the spe- pmvided in the publications listed at the
cies ac~ountathe current scimtifi: name end of each generic account supplemented
with author and year of description, follow- by my own unpublished data.

I
Fig. 4. Bothriechis schlegelii (Bartola, Rfo San Juan,Nicaragua).
Classification of the Reptiles of Central America

Classification of the Reptiles of Central America

The taxonomic level~x

Clas~ Family
Order Genus (inparentheses is the number of species
Suborder .thrrt occur i
n Central America)
C M m t i o n of the Reptilea af Central America
Claeaiflaadon af the Reptiles of Central h e f i a t

r-

Agkistrodart (
Atropordes ( 2 )
Bothrrerhrs ( 9 )
Rothrnps 12)
C e r r u p h r d ~ o(~2~)
Crutalus (1)
van Lncllests ( 2 1

Boa (1)
Comllw (2)
Epicrutea (1)
 Claseification of the Reptiles of Central America

The taxonomic levels:

Suborder
Ifiaorder
Family
Subfamily
Genus (in parentheses is the number of species
that occur in Central America)
The Environment

The Environment

The Enviropment
A chain of mulr;hly 100 volcanoes stretches
along the Pacific coast of Central America
h m eastern Chiapas, Mexico, to central
Panama. Of these, about 25 retain signs of
activity. The volcanic peabs are highest id
the north, where two in Guatemala maoh
elevations of more than 4000 m. The
various mountain complexes that form the
physiogeography of Central America can
be divided into two major ma~sifaThe
northern block includes the Mesa Central
of Chiapas, the highlands of central and
mu&m Guatemala, and those af Honduras,
El Salvador, and northern Nicaragua.
These are separated from the muthem
highlands by the Nicaraguan Depression. (Honduras).
The southern highlands include the
Cordilleras de Guanacaste, Central, and
Talamanca. The two large Nicaraguan Sandy beachea
lakes (Iagpa de Managua and Nicaragua) Sandy beachels (Fig. 6;) extend intermit-
are the largest inland bodies of water in tantly along much of beth the Pacific and
Central .fmericat with surface arras of the Caribbean coast, especially in bays
1036 k m and 8167 km2, respectively. The that ars protected from the breakers by
diverse landscape of Central America s u p offshore mfs. Commonly, a narrow atrip of
pods very different regional climatic coconut palms ~ c c malong the beach.
situations (DUELLW 1966, STUART 1966). Dunes aupport the growth of scrubby xero-
On the Pacific versant of nuclear Central phytic vegetation, such as low shrubs, aga-
America the dry season extend8 from ves, and matted cacti (Rhipsalie sp.).
January through April. A subhwnid corri- Among others, the helidthermic race run-
dor, made up predominantag of dry forest n m (Cnemidopharus), apiny-tailed igu-
or the remains thereof, stretches along the anas (Ctertosaura), and spiny lizards
Pacific versant of Central America from (Scelaporus) inhabit the sandy beaches.
the Isthmus of Tehuantepee through These exposed sandy habitats are also
northwestern Costa Rica. Generally, the important nesting sites for sea turtles and
Caribbean versant experiences an equa- spiny-tailed iguanas (Ctenosaural
ble, tropical climate with relatively mesic
conditions throughout the year. Mangroves
Exceptions are the northern part of the Mangroves (Fig. 6) a m found along the
Yucatan Penineula, ~IIwell as some interi- comtline at many localitiee in Central
or valleys, wbich experience a more arid America. The red mangrove (Rhbpkora
climate. The wettest regions of Central mangle) is more aalt water tolerant, and
America are found in southeastern usually occurs immediately along the
Nicaragua and northeastern Costa Rica, edges of lagoom and bays. Behind the
which receive an excess of 6000 rnm preci- Btand~of red mangrove, the much taller
pitation per year. black mangrave (Avicennia germinand
 pmlonged drought periods. Dry fornets
consist of a t&ick ground cove* of succu-
lents and drought-reeistant shrub^. Tree
heights generally do not ex& 20 m, and
many of the trees are armed with t h o m ~ ,
A curiou~laspect of thk formation ia the
presenw of tree cacti, which often reach
b e i t s equal to that of trees (Fig.71.

Savama
The eavanna that occurs in Central
America includes wet savanna (e,g., on the
Honduran island of UtilaJand pine rsavan-
na ("&banas de Pinos Caribe"), Pine

Fig. 6. Mangsove with epiphytes cm Isla de

Utila, Honduras.

and white mangrove (Luguzaculariarace-

rnosa) ofeen form dense swampy forests. A
tangle of aerial and adventitious mote
characterizes thi~ecosystem. Mangrove
m m p s provide habitat for the American
crocodile (Crocodylus acutus), the Utila Fig.7. I3ry forest at Tehuantepec, Mexico
anole UVomps utikml),and the U t h iguana (406m elevation).
(Ctenosaura b a k e d .

Dry forest
Dry forests are typical of the Pacific ver-
sant of Central America sad a few interior
valleys of Honduras and Guatemala. The
climate is characterized by law precipita-
tion (500 to 1200 mm) that falls primarily
during the rainy seamn from June
through November. During the dry season,
when precipitation is negligible, a majori-
ty of the trees shed their leavea, permit-
ting sunlight and wind to desiccate the
ground layer, Only the large rivers contain
water throughout the year, where- smal-
ler streams are intermittent or dry during
the dry aeason. Plante and h a l a of this Fig. 8. Pine savanna at Alamikamba,
extreme habitat are well adapted to the Nicaragua (100 rn elevation).
 savanna is found on the Caribbean ver-
sant of Honduras (&om the Laguna &ban0
southeaetward) and Nicaragua (from the
Rio Grande de Matagalpa northward).
W habitat irs duminakd-by open stan&
of Caribban nine (Pinus caribaea) that
abruptly cham& aver ta broadleaf foreat,
II
Rain forest
Evergreen tropical rain forest ia among
the most diverse e c o s y s m on the planet,
and in many regions its scientific explora-
tion ie still in the beginning stages. In the
northern parte of Central America, rain
forest is found mainly in the Caribbean
lowlands. fn Costa &ea and Pmambl, it Rg.10.Buthwm tra+ in the rain fm~tat Bm-
ale0 covem large meas of the Pacific rer- toh, Rfo San Juan,Nicarapa (30 m elmatid,
aant, During the day, temperatuna in the
tropical rain forest reach 90-35°C. Eha+q
the night, tempraturea drop to about
25°C. Relative humidity is very high 190- 100%). W a l l tiends ta wlsiderable
throughout the year, and heavy rains are
most common in the early afternoon,
Several lemL of vegetation can be dieting-
uished in tropical rain forest, from the
dimly lit forest floor, the scrub level, the
@ trees of different ~izes,up to tb8 t~11mt
trees of the forest, which r&ch high above
the c a n o ~The
~ . diver~ityof tree meeies is
extra&6afy and can ixeeed 56100 per

I
Fig. 9. Rain forest at Bartola, Rio San Juan, Fig. 111. Cloud forest at Voldn Mombacho,
Nicaragua (30 m elevation). Nicaragua (1100 m elevation).
18
Fig. 12.View of the cloud forest on Cerro %lamb&,Nicaragua

hectare. Many reptilm occur exclusively in than that of the lowlands, but there is a
primary lowland and premontane rain high numher of endemic species, i.e., spe-
forests and disappear with deforeetation. cies that only occur in one particular area,
such as a murunit or a mountain chain.
Cloud forenst
A world of ita own, cloud forest is the type
of natural vegetation that can be found
above 1200 m elevation in many of the
mountain ranges of Central America. In
thia type of habitat, temperatures are rela-
tively low (5-16 "C,depending on altitude)
and humidity is oomtant1y high. The trees
in cloud forest are densely cavered with
epiphytes, such as bromeliads, orchids,
ferns, and mosges. Tree ferns, which can
reach several meters in height, are aleo
found frequently. Small atream channel
the water on its way to the lowlmds. At
very high altitudes along exposed ridges, a
special eub-type af cloud forest grows, the
so-called elfin formt. Subject ta low tern-
peratuw, high precipitation, and high wind
speeds, trees in the elfin forest usually
attain leas than 1-3m in height. The fauna mg. 1.3.A cloud fomst trail. at Montevede,
of cloud forests tends to be less diveme Costa Rim I5600 m elevation).
The Envi-ent

Elevation Map
 The Environment

lVIap of Vegetational Zones

Climate in Central America
climate in Central America is high-
ly location depend~nt,with the yearly
cycle largely characterized by variations
in precipitation levele. OR the
Caribbean side, the heravimt rains fall
during Deceder and Jmua~y,while
the Pacific side receives virtuaUy no rain
dmbg the northern winter (the dry sea-
son on the Pacific ~ i d elasta from
January until April), In general, the
Caribbean side of Cantral America is
more humid and reeeivw more precipi-
tation than the Pacific mde.
The fallowing climate diagrams UUB- line
trate the yearly climate in seleet~d
are-. The mean monthPy precipitation
is indicated by bars, The mean daily
temperature can ber read from the upper
curve and the mean nightly brnpmfure
-from the lower curve (data fn,m M i j L m
1996, except far Utira, which was collec-
ted per~onally).
&an Salvad~r,El Salvador ' agua, Nicaragua

iI

ID

II

0
A M J J A S O N D
Geological Hl8talay

Geological History and Zoogeography of

the Reptile Fauna of Central America

SAVAGE (1966, 1982, 2002) provided became cooler and dryer during the late
masterful analyses and discussions on the Cenozoic era, they became restricted to
composition, origins, and h i r y of the tropical Mesoamerica. Examples of the
Central American herpetofauna and "Middle American Element" include the
developed a t h e m explaining the present Kinostemidae, Helodermatidae, Untusi-
dietribution pattern of the amphibians idae, and Xenoaauridae. These familie8
and reptiles. Ewlier biogeogmpher~have are all known aa fossils from early
regarded Central American aa a faunal Tertiary deposita in North America near
tramition zone between the American a n - 40" latitude north (QAVAGE 1966).
tinente (e,g. SIMPSON 1960, K. SCHMIDT
1954, DARLINGTON 1967, STUART1864). A m n d major historical unit is the 'Old
However, SAVAGE(1966, 1982, 2002) Northern Element" (SAVAGE 1966, 19821,
demonstrated that the Centrd h e t i c a n which is compasd of genera with a pri-
herpetofauna forms a distinct zoogeo- marily extratropical distribution in
jpphic entity, both in regard to its history Eurasia and/or North America. During the
and pre~entcomposition. The following esrly Tertiary period, this group'^ anma-
discussion is baaed largely on the latter tors experienced a more or less continuous
three publications, supplemented with and circumpolar diatribution on the
information *om works on the geology of northern landmasses. The gradual clima-
the region by MALFAIT & DINKELMAN tic changes during the Cenozoic era forced
(1972) and MAWHALL et al. (1979). this group's ancestors southward and
caused their fragmentation into geograph-
The Central American herpetofauna is ic isolates. h a result, this stock contains
composed of genera with an origin in the a unique Mesoamerican component of
region (autochthonoue genera), as well as endemic taxa that have evolved with the
those that dispersed from North and autachthonous "Middle American Element"
South America into Central America. from the Eocene epoch onward.
SAVAGE(1966, 1982) concluded that three
major and one minor historical source A third major historical unit is the "South
unite had contributed to the Central American Element" (SAVAGE 1966, 19821,
American herpetofauna. which is composed of genera with primari-
ly South American distributions and rela-
The most important historical unit is the tionships. Their ancestors evolved in
"Middle American Element" (SAVAQE 1966, South America and migrated various
1982) that is made up of genera that are d i s h c e s into Central America after the
primarily tropical Mesoamerican in distri- clomre of the Panamanian portal during
bution (many Central American endemics) the Pliocene. Consequently, this unit is a
and have their closest relatives either in rather recent contributor ta the current
Central or South America. The ancestors Central American herpetofaunal diversity.
of this historical unit u s d y evolved in
the region itself or in South America, and Finally,a fourth lew important historical
reached far into North America during tihe unit, the *Young Northern Elementn
early Tertiary period when a tropical (SAME 1966,1982) can be differentiated.
rnesic climate with w m temperatures This unit k represented by a few genera
prevailed. When the climate generally (includingSceloporus and Umsaurus)with
a primariIy extratropieal cfistributien IS&m lSS4 10821, The latbr gtrmp
d a t e d with semiarid ta add m&;i0118
in reached Middle America in t h ~Eecsne
aauthweatezp North Amedca. through migration wuthward along the
developing RoclQl Mountdm arid became
A hrief summargr d the gaola&i.cdhidmy iaoleted from their congeners in the
of Central America fadlihtm the under* eastern and weatern BPBWof what i~ now
atanding of the cumeat distribution pat- the Unibd state^ by the grdud develop-
tesns of reptiles in the region. The land ment of a mare arid climate atroes north*
connection that existed between the ern Mexico, In the Oligocene epoch, condi-
American continent8 during the lata tione remained similar te thorn of the
h-ue and early Tertiary pride (70 E m n e in Nudeat Centrad Ameria-a, al-
to 60 million yeam ap3 bmb up in the though aome uplift d the northern moun-
Ewme epoch (64 to 36 d l i o a yeare ago) taie mersrsifa may have ocaunwd.
due to inund~ttionof tba I&hmi&tlLlnk
(P lower C e n M Amdeal. Thie so-& The Miooene epoch (28 to 6 million years
Panamanian portal separated Nuclear agar was the time of great& change
Central America from South America during the Tertiary period in Cmtral
during a period of time &hath a for h e r i m , bioth in regard ta climats and
about 46 million yearn It wm not Mre vegetation, as well aa changes to the land
the early Pliocene b o r n 5 milli~nyeam relie£ In the Miocene, with the disappear-
ago) &at the dosing Bf the Paa;amanign ance of the Panamanian p o w a contin-
portal m ~ e c t e dthe American conti- uous laad c o m ~ o between
n the eonti-
nents. The uplift of the Central American nents was re.wtablished for the fir& time
mountain mawif@began in the Plimm since the early Eaceae, makin(g faunal
and apntiaus~rtoday. TIIB &or phyaiopeo. mbwmge possible,
gmphio &an$& caused ei@c:$nt c l b a -
tie &Rs. During the Plioc~ne,the climats The 8auth heriminfluence is present-
baGseme genmdly woler agd mare arid. ly re~ltrictedmo~tlyto lower Central
Pleietmene gladah add
Additiaaally, t h ~ America (i,e., &here* of what i~ today
inbrglsciab w m e3t.tmmly important Co&a and Panamal, Only a few ape-
mmt. cieus d'South American arigiu haw mached
northern Central America (e,g., Gymnoph-
In the early Drtkary period (a,g. during thlnetrs ttpecwsus), where&& mwt South
the Paleocene e p h , 86 t~ 54 millicrn g e m h e r i o a n elements dispmed only ink
ago),the North aad Soueh American mnti- Panma (e.g., the genera Marutaasaurm
nents were connechd by a land bridge and Zlwchyboal or Costa, Wca CB.~., tPle
with low relief that supported a general- genera Anadia, Dactyloa, and Nau-
ized tmpieal reptile fauna thet extended sbicurus). Many species of Central or
northward to at least 40e htitude. It North American origin, have migrated fm
Iw wsumd that dl f&~a of reptdlsa into South America (e.g, the genera
that cumen* inhatbit the ETeot-ropicswere CmtaZw, &1cctna, Noraps).
alneadg present at that b e . During the
Eaeme, flaodhg of the d m parts of The Pliocene (5 to 1.8 million yeam ago)
th~3intercontinental land bridge Efmm weu the mwt imporhant period in the wow
what iia now m t r a I Nicaragua to north- luti~nof the Central American mptrile
arrn Colambia3 eliminated the comectim fauna that led to the modem apecia cam-
of N d and South America, Soon, two pwition. kt the begjnrdng of this epoch,
p u p s were pmsent in the northern m mprmenZativm of both historfcal units
gion: the Widdle American Element" md (Middle American Element and Cenkal
s aabunit of the Old Northern Element, Ameriaan Complex) were widely &Mb-
the so-called "Central American ComplexA uted o ~ e rmu& of the C e n k d American
Za~ggographyoZ the Reptfle Fauna
region, with speeiation e a t e r s in the
uplifting motantain mass* Ths mom-
taiae formed a climatic divide that led to
the differentiation of s p e c i ~communities
~daptedto the more arid mnditions on the
, well as those asmciated
Pacific v e ~ a n tas
with the more mesic habitats on the
Caribbean ver~mt.
T ~ B modern
B~ epecies dispersed along
muhs that were controlled by climatic
and physiopo%paphic factors (Fig, 14).
Momaina raatrided the migratiod of low-
land specie@and provided d i ~ p e r ~mute8
al
for highland ~species,Two tgpes of barrier8
Fig, 14.Principal diapmal routas of amphibi-
am aad mptiles in Central America during
the pwt two million years. ARsr SAVAGE
619681.
are e f f d v e far lowland apecies: highland
massif8 and dimatidly unauihxb1.e area$.
An example of the latter type of b d e r is
the xeric n o r t h ~ mpart of the Yucatan
Peninsula thet prevented mesic-adapted
species fmm moving from the Peten
f o m k inb the region. An area of very
complex lowland climatic filters occurs
dong the Pacific ve-t of Costa Rica and
wehltern Panama.
From the late Pliaeene onward, the o w a l l
picture of relief and climate in Central
America mrnbIed, for the mwt p- the
present physiogeography, with the eprcep-
tion of the marine. embayment of
Nicaragua and fie &tillflooded northern
part of the Yucath Penin~ula,both of
which became more and more dry due to
decraasiag sea levels,
During the glacial periods of the
Pleigtocene (1.8 miIlion years ago to the
present), the sumrnitrs of the highest
Central Amedcan mauntaina were cw-
ered with ghciere. Based on recent studies
on fmeil pollen profiles ~ C ~ ~ ~ N 1999,
VAUX
1446), it is aesmed that the average
temperature in Central America dropped
6 to 8% below the preeent level during
glacial peuiodg, resulking in environments
in the lmlanda mmt closely resembling
&age ~f midelevation forests f6W to 1600
m)at presmt (SAVAGE2502). M n g these
periods the dietributiom of the highland
species were oompmssed downward (by
abmt 800 ml,where they co-occurred with
lowland species (SAVAGE 2002). During
interglacial periods, the highland species
moved back up the cnrdillem as tern-
perat- returned to levels eqvquivdllent to
those at present (WART 1961, CAMPBELL
1984, SAVAGE 20021. It ie aaeumed that the
cooliag and warming cycle8 in the latter
part of the Cenmic, in wociation with
the continued uplift of the cordilleras,
provided a major impetu~for epeciation
(SAVAGE a002).
During the turbulent geological W r y of
Central America, eeveral effects have eon-

Mbuted to the evolution of the region's
reptile fauna (SAVAGE
19661:
A characteristic of penhaulas is that spe-
cies can invade, but are hindered from
further migration by the surromding sea.
Nuclear Central America formed a pen-
insula during moat of the Tertiary perid
and accumulated successive invaders from
the north. The Panamanian portal served
as an effective barrier, prmnting these
invadera from migrating southward. From
the Eocene thmugh the Pliocene epochs,
this peninsula sewed as a center of spe-
cies differentiation.
W n e portal effects
The formation of marine portals can frag-
ment the former continuous distribution
of a species, &us stimulating independent
differentiation of the isolated populations.
With the inundation of the Isthmian Link,
species that were formerly continuously
distributed in South and North America
became isolated on the two sides of the
Panamanian portal, with differentiation
Zoogeography of the Reptile ]Fauna
Old Northern Element
GeneralizedTrapical Fauna
Middle American Element
- &uth Amerisan Element
Central American Complex
Fig. 16.Wlogieal history of
Central America aince the
Paleocene. A f k SAVAGE(1988).
proceeding in Nuclear Central America
independently from that in South
America. Examples for such sister groups
are the genera Ames'va / Cnemidophorus,
Caiman I Crocodylus, and Dactyloa /
Norops .
I d a r eBects
Due to ieolation, islands stimulate specia-
tion. During the Tertiary period, p h i c u -
larly in the Miocene epoch (24-6 million
years ago), the numerous islands associa-
ted with the Isthmian Link / Panamanian
portal zone of instability supported insu-
lar evolution.
Land bridge effects
The fornation of land bridges makes pos-
sible the invasion of species previously
hindered h m immigration by the exist-
ence of marine p o M . ARer the re-estab-
lishment of the Isthmian Link in the
Pliocene, invasion from the south by spe-
cies previously excluded from Central
America began and continues to the pres-
ent.
 Crocodilians (Crocodylia)
The order Crmdylia is represented in
Central America by three apedm in two
families. Typical of dl species of the w&rL
these are large, robu8< aemiaquatic rep:
tiles, They inhabit lakee, lagoom, Bwampe,
and large rivers, Theee atrictly c a r d -
voroua reptiles fdon invertebratea md
vertebrates and exhibit necmphagy and
e m caanibalism (LEE1998).
Fig, 16,Adult C r d y E w acstus lRio ?%moles,
Costa R i a ) . Photo: A. FlPischea&&ger
Fig. 17. Adult Morelet's mocDdile (Crwor6ytw
momktii). Photo: R. Cedeiio 'I!
28
In Central America, the American m c o -
dile QCmcod,lrtsacutus) inhabits predom-
inantly brackish coastal environments
(lagooia, marshes, m d m a n m e swfunpe)
with individuals penetrating further
inland alang major rivers. Morelet's
crocodile ~Cmodylusrnoreltii) and the
spectacled or brown cdmm (Caiman
crocodilu8) live primarily in freshwater
habitats, such ast lakes, ponds, mad rivem.
Fig. 18.Spckcbd caiman Eabum c d i E u 8 1 at
CafIo Negrm, Costa Rim IPRata: H.M m
Fig,ls.Speekded caiman (Caiman c7.oOordi/w31at
Cafio Negro, C a t . Em. Ww H.Maaear
Fig. 20. Juvenile Crocadylus a ~ u t u s(Rfo Fig, 21,Juvenile Spectacled caiman (Bartola,
Patuca, Honduras). Nicaragua).

AH crocodilians are oviparous and lay
hard-shelled eggs (usually eeasonally, and
a single clutch per year). The spectacled
caiman and Morelet's crocodile build
mod-type nesb made of dead vegetation,
sticks, and mud for their eggs, whemm the
American crocodile excavates a ahallow
hole in the ground. In Chiapas, Mexico, the
egg-laying season of the American cmc-
odile ia during the months of March, April,
and May with individual nests also
obeemed early as February (ALVAREX
nn, TQRO1974). The female digs a hole,
approximately 770 cm in diameter m d 20
to 60 cm deep. Preferably, nests are made
on sandy e m b m b e a t s above the water-
line or high tide mark. C r d y l u s acuEu8
produces 15 to 105 (usually 20 to 60) eggs
per clutch that hatch after an incubation
period of86 to 111 days (K0ar;E~1997).
ssason from early August through early
November; GRENARD1991) produces clut-
ches of 10 to 23 egga that hatch after an
iacubakion period of 88 to 104 days. In
Cmodylus moreletii (egg-laying season
fkom April through h e ) , the clutch size
varies from 20 to 45 eggs and incubation
takes 78 to 98 daye (ALVAREZDEL TORO
1974, KOHLER 1997).
As far ahc we know,all crocodilians exhibit
parental care (GREER 1971, KU~HLAN &
SIMON 1981). During the incubation peri-
od, the female remains in the immediate
vicinity of the nest and attach potential
egg predators (MODHA1967, Cow 1871).
The female responds to vocalizations
made by juveniles from within the nest by
opening it and tramporting the hatchlings
to water (POOLEY 1917).

The mound-type nests of Caiman crocDdilus In crocodilians, the incubation tempera-

can reach a height of one meter and a ture determines the sex of the hatchlings.
diameter of more than two meters (STATONIn most species, high and low tem-
L DEON 1977). After oviposition, the fe- peratures produce male hatchlings, where-
male closea the nest cavity and crawle over as intermediate temperaturea produce
it several times to condense the material. females (DEEMING & FERQUSON 1988,
However, air pockets prevail in the nest K O ~ 1997).
R
throughout the incubation period. Due to
fermentation and deoomposition of the In Central America, all three species are
plant material, constant humid tempera- endangered due to overhunting and
tures of 32-34 "C are produced within the destruction of habitat, National laws to
mound. Caiman crocodilus (egg-laying protect crocodilians exist in all comtrieg.
 Fig. 22. Female Spectacled caiman with neonate^ (Bartola,Nicaragua).

Furthermore, Cmcodylus acutus md C. ~ l l i ~ ~ t ~ ~ i d ~ ~

more'etii are listed in Appendix and [>ll,)lll,l (Ir,
l ' l ' l ~(l(ll//(h Y N,\[:I IS I ~ ~ HSI V
. ~~CII
Caiman crocdilue in Appendix 2 of .the N:lt u , , , ~ ,,i 10. 20r;; tjl,(. I ~ ~ unkrlow~l : ~ I ~ ~ ~
Convention on International Trade in hinxr mum trltal lrnfilh 2 111 S u u l h e r ~ lMcsr t7rl
Endangerad Species (CITES). IIOII~ r1111~t01' t h t bL'o:%s~+ or ('<,I~Lv:II
~IIIIC~~IC;~
111to Sntrth A n l r r ~ c , ~ . Iovrl t o 200 In c,lt.v,~-
\(>;I

Crocodylidae
l ~ ~ , s ('! IVII<I{ 1807%Arin. XICIS,
( ' r ( ~ c ( ~ i i y (zc~ittis
Ilist. Nnt., Pal-is, 1 0 : 55; type 1rrc;rlity: S i ~ n t u

LID t.rl f i nil. I+'t~im

soulhrrlj Floridil. LISA.

rlnd Venrzu(bl;t), sr:i I(xvc~lto 50(l 11) t*lr>v;lt

ion

ion.
Fig. 23 Juvenile American cromdile
(Crocodylusacutus) at N o Patuca, Honduras. C
30
 Fig.24.Crocodylus acutus submerged in a
small stream (Rio Patuca, Olancho, Honduras).

L
FIg. 26, C d y i u momletii
~ GWn,Guatemala).

Craeodylus acutus
 I

I
I
~ l ur s q -
-
- -.
- -

Fig. 2% Charachrhtiea important in the identification of cmmdyliaos. Drawinga by M. Ves*

Key to crocodilians 'T

1 a A transverse ridgejust anterior to eyea on
top of head (Fig. 28); no teeth of lower jaw
visible when mouth closed .......................
/
...........................,...
Caiman crocadilus
b No tranlvens ridge jlld ehrh
to eyea
on top of head (Fig.26); 4 tooth of lower
a. Cmmdfl~&
acutus

jaw lies labally in an open groove and is

vieible when mouth is c l o d ...................
2
2 a Primary aubcaudal whorls not interupted
by incomplete ewndary whorls (Fig. a7a);
snout relatively lender, about twice aa
long a wide ...............
C~rlaodyluuactstw
b Primary subcaudal whorls interupted by
inooglplete secondary whorls (Fig, 27b);
snout relatively broad, length about 1.8
timeswidtb ...........
C d y L u n nsoreZeti1
b. Crocodylus moreletil
Further Reading
WERMUTH& MERTERNS 1961, NELL 1971,
B-s 1Q73,ALVAREZ ~ ~197* Ross 8t
f l TORO
Ross 1974; BUFFETAUT 1979, ROSE 1987,
DENSM~BE & OWEN 1989, ROB & GARNE~ Fig. 27. Ventral surface of tail, Incomplete
1989, ORENARD lQ91, F%REZ-HXG~~REDAet al. secondary aubcaudal whorl8 in Crocodylug
1991,EWT et al, 1999. moreletii ahadd.
Turtles (Testudines)

nuchal
marglnel plates

axllQr plate Inguinal plate

cm moveable Fig. 88. Charaaterietiosr
important in the identifi-
cation of tartlea
a. lateral view
b. Dorsal view;
c. Ventral view;
M=marginal plate;
P=pleural plate;
V=vertebrd plate,

aMomln% piatate
/ \
rammat ptats
\anal plate

Twrtles are represented by 25 specie8 in 14

genera and 7 families in Central America,
the kinosGrnida and emydids being the
mast speciose with 8 species each.

Key to genera oftur&le~

1 a Forelimbs m&ed a8 omlike flipper&; % ~ ~ + P l ~ ~ ~ .
two or fewer claws on each foreleg; manic Mar WERnauTH &
species Sea Turtle8 (see key on page 63) Mmmw IlWlI.
b helimbs not modified as owlike flip
pers; more than two claws on each foreleg;
species living on land or in fresh or
brackiah water habitate .......,.......... .,.... 2
2 a PIaetFon with 11 or fewer plates .............8
b Plastron with 12 plates .......................... 7
n reduced and not cruciform ..4
3 a P l ~ ~ t r onot
cmcifom (Fk. b, Chelydns
v v a
b Plastron very reduced ad
.
2%-d) ............................................. 6
c. S&umtypug d. Cl@&r,,
4 a Pbtran with two mumable hinges (Rg.
28c and 90k chin barbels pwent
.....................,....***....,. ...........Hhx~Gmom
.......
b Plaahxi ried, without moveable binges
(Fig.296t);cFun barbeh absent ....................
..............................l3wmatem {in part)
B a Chia barbels absent ..................
Ck&d~a
b Chin barbela preset (Fig.28c) 6 ......,.......,,
6 a Plastron rigid, without mweable m a
mg.29d);bridge mme&ed to mrapaee by
ligament5 ................................
Chdiw
b PSa%tron with moveable hinge; bridge
connected to carapaee by an o~seousmhn%
..................................,.......S t a u ~ r s s
I a Plastron with moveable hinge; plastron
and mapace in contact, bridge &went ......
....................
,.......,.............Terrapen@
b Plastmn rigid, without mweable hinge;
Wdge present ..........................................B
Fig,31. Chelydra serpandinu (Ria San Juan,
8 a Hind I i m b ~elephantine (calm>; cara- Nicaragua)
pace dome-eh~ped;without webbing bet-
ween ..........................
Ckmhekme
b Hind Embe not colrtmnar; carapace rela-
tively flat, only slightly convex; wehbhg
hetween toea more or lesa developed 0 .......
9 a Limb unieolor dark gray; #lout slightly
upturned,wmmhat tubular ......................
...............
,..,..,., Demtwrw.9 (in part)
b Lknh with some kind of pattern (mwtly
pale stripes);snout not rn above 10 ...........
10a Pattern of carapace m i s t s of yellow can reach 47 cm carapace length with a
rings with dark center, bind feet with weight of 20 kg and should be handled
extedva webbing ................. l h c h p
with great care,It is able to deliver quick
b Pattern d carapace without yellow rings, strikes with its powerfid jaws. Snapping
or, if yellow riqgs are men&then hind
f ~with t littl8or no WB bing
.................,..
g ........,............
turtles inhabit pander, lakea, and slow-
.........*..........Rhl3~1~lemws moving rivere with muddy bottams.
Mostly nocturnal, t h e e animals usually
rest in shallow water with only the
nostrils and eyes protruding above the
~nrfaceduring the day. At night, this turt-
le walks slowly on the bottom of it^ watery
habitat searching for food. Snapping turt-
leg feed on dmoet anything they can ewal-
low, partidatly insects, shrimp, crabs,
fish, and &om,but also crocdle and turt-
le hatchliqp, snakes, bird@,and small
mammals, as well as plant matter such as
algae and other water plants and Fruits.

Fig. 30. All apeues of Kinoaternon have a plas- The female dig a nea~tfor her eggs (in
tmn with hing~that enable a wmplete occlu- April t b u g h June1 on a sandy embank-
sion of the shell (K.leuco&omum~. ment above the waterline, preferably in a
spot with some protection by low vegeta-
tioa. Twenty to 50 (exceptionally up
to BS egp in very large femalw) am laid
per clutch. The egp measure 23 to 33 mm
(weight 7 to 16 g) and the j u v d e n ( m a -
pace length 24 to 31 mm) hatch after an single pnue with one species, Dermat-
incubation period of 55 to 126 daye (Emm smys m w i t is found in lagoom, rivers, and
& BARB~VR 1989Z lakes from southemtern M e x h through
Belize. The species also talerates brackish
water. Them noeturaal turtles are highly
adapted to their aquatic environm~nt,and
are capable of performing prolonged dives.
On land, hawever, locomotion is difficult,
aa is liRing the head, Adult Central
American river turtles feed almost
exclmively on plant rnslterlal (algae and
other water pletnta and W t a thd fall into
the water). The egg-laying period laab
from April through Septemba with the
female8 buryinaC their clutche~of 6 to 20
eggs (67-70 x 30-34 mm) on sandy
M a r Reading embankments near the shoreline,
ERNST& GIBBONS
1988

'ig32.Central American river turtle

Demtemys muwti) in Tabasco, Maxica,
Photo: J, Haft
 Emydidae eng&.w zwnu"
nodweltern Erma

himlemmys puleIremima (GR~Y

'Ihe species of hnznocletrtmy$ are distribu-
ted across the tropics and subtropics of
Mexico, Central America, and northern
South America. Some species le.g.,R. fune-
Shield Rept. Brit. Mua. 1: 26; tm locality:
Mexico. Catapaee length to 200 mm. Sonora,
Mexico, to Coeta Rica,sea level to 1180 m ele-
vation, The subspecies Rhinoelemmays p, put-
ohemha (Guerrero and Oaxaca, M&cQ), R.

I
red me highly aquatic, whereas the
majority of Neotropical wood turtles have
a semiaquatic or even terrestrial lifeartyle.
In the Bioephere Reaeme, BosawBs,
Nicaragua, I encountered several 8.annu-
p. i n c h (BWOURT M68)(Oaxaca, Mexico, to
narthern Nicaragua), and R, p, manni DUNN
195h (southern Niwragua ta Costa Eta)
occur in Central Ameriea.
Rhinoclemnaya rubida (Cope 18701, Roc.
Amer. Fhilos, Scc, 11: 148; type l d t y :

J lata within the rain forest, Borne 100

metera from the nearest body of water. On
the other hand, I have recorded R,funerea
Jvchith, O~axaca,Marim. Carapace length to
230 mm.Pacific wmt of Mexico from Jab00
to w e e m Chiapas, sea level to 750 m. Only
the wminate aubspedahl murs in Central
in the same general area, but exclusively
in rivers. There, it can frequently be seen
basking on partially submerged logs.
Neotropical wood turtles are highly herbi-
vorous and only occasionally inge~tin-
sects. They o h n forage on land at night,
feeding predominantly on broadleaf
plants, grasses, and h i t s . Moat Central
American species produce small clutches
with one or two eggs; only R,melanosterm
and R. pulchrrima lay larger clutches (up
to 6 eggg per clutch) ERNS ST & B~RBOUR
1989,KOHLER1997).

Fig. 33. Rhinoclemmys rubida,

Photo: R. Wicker

Fig. 34. Juvenile Rhiwlemmys fururea (Rio

San Juan,NicaraguaE.
Fig. 36.Rhinoclemmys pulchem'ma a n I
Patuca, Olancho, H ~ n d d ) ~

rig, ap. +En&raaoiemmysareolata.

Fhota: 8.D. Bartlett

--

Fig. 37. R h i n o c b y s oreolota. (Cdakmd

Biosphere Reserve, Campeehe, Mexisol.
Phota H.Bahena B.

-
Fig. 38,Rhinaclernmys annulata is mom
terrestrial than other species of the genus.
&g to Rhfmlenanzy8
X a Hind h t with exten~ivewebbing a ...........
b Hind feet with Uttle or no webbing ......... 4
B a W pale gray without tFanscrerae dark
bar; past&or surface of thigh with verti-
cal orange bar8 ....................
..
..........
................
R h i n o c h meknmternas
~
b Iris with k ~ ~ ~ v e dark r n e bar; pattern of
f&b not aa above ..........................., .8 .....
8 a Qne or two red supratemporal stripes
pre8ent on each 8ide reaching snout;cam-
pa= with or without ocenakd pttesn
..................RAilsocEemnys pslchereS:ma
b NQ red supratemporal attipea; c a r a m
without ocellated attern ......................
...........................&&-ye mm
.. * nrmpme camiinrs yuwtana
~ i g40.
4 a ~ ~ ~ of head
a *t.h
l large C C d b d B ~ Q Bm@em%
P ~ empeche,
@low marking (&en horne~h~~-sIraped;Mexico)+ Photo: H,Bahena 3.
Kg.33%bridge darkly pigmented ............
..........................,.
Rhb0C-r~ tub@
b Dmal cruxface of head not as above;
bridge pigmented or not ......,....................
eS
5 a Dorsal eurface d head mtfonnly dark
-
bmwq supratempd stripe, if present,
indistinct; car& ace with broad keel
.......................
b One or
dlna0ha98tzwdkta
pala su~rabmp~rd s t s i ~ fiur
~ of the
-
nn
nrrapem
present on each, aide d head posterior to
orbit; k-1 an c a a aa, if prwnt, n-w mmentlS feca@zed, with only camiina
........................
~ R k - 8 ama~ata o c h g in southern ~ e x i c o(ERNST &
MCBREEN 1491). On the Yucath
MhBr m a w Paninaula, this specie8 inhabits wet
EEKST1978,1S&Oa-c, 19Bla-e, PEEEZ-N~GAREDA B B V m B and P&W land, well as
& S m 188'7. thorp forest and evergreen rain fore~t,
with a preference for open habitats (LEE
1996).Although box turtles d b i t a ter-
restrial lEe style, individuals can be ob-
served sitting in hallow water. Jweniles
appear to have a predominantly carni-
while adults are predominant-
vorous dietst,
ly herbiv~mus.The carnivorous diet of box
turtle8 includes snails, w o r n , inwcta, spi-
ders, salamanders, and frogs.
Copulatian is preceded by an exbnsive
courtship, which can be divided into three
phases CEvm8 1953):(1)a circling,biting,
and shoving phase; (2) a preliminary
mounting phase;; and (8) a copulatory
phase. The females produce clutchera of 2
Fig. 99, R i L i d n n a y a Pylchrrima [near to 4 eggs that hatch after 60 to 70 dap,
PoFtedpe, Gumcaste, Cmta Rim). depending on ambient temperature
Phota H. Bringsoe (KCIHLER 1997L
 n m p m OmOIIM L L W
Naturaa. ed. 10: 198:m e locaj
Car&& l a @ to 2a-nrm. Wiady diatribu-
ted over the central and eastern U.S.A. with
popdatioa in Mexico (Tamatdipae,
iaolatRdtRd
San Luua PotosfpV e m , and the Yucat8n
Peninsula), sea level ta 100 m elmatian. The
subspecies E c m h m (GRSY1849b) % (-I
to V ~W o o 3~and T, c, y w -
? ~ G ~ N Q E R 1885) OIuaUn Phlin~uk)
ocmvintbeareacuvssedbyUlisbook

1
Fig. 4-2, Juvenf le %a&eayg acrip;ta
bUmihmba, Niearagual,

lac slider turtle Cr).acbmys twiptaS h

the widest geogmphic distribution of axq
nom-B turtle ME?% Due tQ ik pow-
larity aa a pet, it repreeenh the rnmt w d I
h o r n tnrtle s m e e.At the same time,
them is stil considerable dieagreem~nt
8mmg &enti& authorities reganking its
tamnonay an both the generic and e u b p
ci&o h l .I follcnv GGtL R. E m (in
GIBBONSIQQO:pp. G7-87), who places thihl
species inthe genue k J i e m y s and recog-
&ea the Flubape~ie.~Z a. venu&a
(Carl- vemant of wuthern Mexim
and Y w t h Fkniwula).,I Ia -pi Chdfic
yremant of tha Isthmus of Tkhuantqw ta
werstenz Guatemala), and 'P, a, oraada
(Honduras to P m w ) in Central
America. The pwit.ion of JORN Me~ G L E R
bin ~ I B B 1 ~0;pp. 82-1Oti1,lKha=
Q99 & b ar
I the e@et+ to the genus Pseuhnys, with
Fig. 41, !&mpm
caro2ifaa y W a m the followingOantral American ~uhpede8
t C w e , Wcoh Photo: J, C. Lee l? a v~nusta(Caribbsm ve~azlntfrom
I Testud.: i6;type localitg: unknown. Chapace
h i g b to BO cm. Widely distributed acroae
mnkd md e W m U.SA and southward
zhraugh Mexico and Central America t o
2oIombia and Veneauela, sea lemd to 1000 m
elevation, See taxt abwe Por subspecies.
 repwbd to eetivata sequeatemd in mud
during t;he dzy maam and resume activfty
with the m e t 09 the rainy seamn
Wv- DEL Tom 1983).

&BS is C. angwfatus, the chopontil Dr

narrdw-bridged mu& turtle. Narrow-
bridged mu& turtles we charractwizadby
a small p 1 m h that is mllnected to the
cBrapwe by 3-h. Th588 +iwtIes are
Emown for their gumeiow attitude, and
sbuld be hmdld &kh @mest are in mder
ta amid severebites. Cladius aw8&&& Me &*
BEmy lgEIO,
h rtn aquatic s@ea that principally in-
habit9' mqaa with eeaa~ndlydoDded
gramlancb and f a r m in th&e grm-
lax& during the QoadingQ ~ r n e - V n m u
& Zuc 19%). It is mmt active m u n d
dam. f2aasdius angttetatus ia ~triotly
carnk~rnusand prep on hvi~rte-brata
arrd mall v e r k h t i s . X)lxrina; the rainy
season, some inMdualrs *b b0IlBider-
able distances and can then acawidly
be ixmmtered on land during the day-
time (Lm1996l.Abr the f l d s , C. angu8-
&&a mpradlaw. The clutch tze varies
from 1 tn 8 egg& which hat& after an
incubation perid of 98 b 150 days
&pending on mviromentd conditions
CEmST C BABBOUR1989, ICd,= 199.71.
me Narrow-bridged musk turtle ia

(Tabam,
Fig. 44,CaaIldiua cdng~~s$atus
Mexico), Photo:J. H a Quietma Rao, Mexico), Photo: H.Bahena B.
Mud turtles (genusfi7a-osternola; a b u t 20
species) are mall to medium-sized, pre-
dominantly aquatic cheIoniana that are
widely distributed aver North, Central,
and South America. They are character-
ized by one or two maveable hinges in the
plmtroe that enable complete cloeure of
the shell,The Central h e r i e a n specie8 d
Mud turtles live aquaticallyLmostly in
ponds, h e m , olnd marshes from sea level
to slightly mom than 1000 rn elevation. Fig. 47, K i w t o m n letlco&ornum (ma Ban
Frequently, individuals can be encam- Juan,Nicaragua).
tared on land when they wander lirarn one
body of water to another, On 'the morning
of 220 April 1990, during a stay at
'Ibrtugue&, Costa &a,I observed a spec-
imen of Eimstemora leucostomurn in a
clearina within the rain forest. When a
mud &le is lifted, it wudly r e h a b into
it8 shell md clo~esthe ahell completeIy
with the aid of the p l a e h n binges (Fig.
30).After a whilet some individuab might
etick out their heads and attempt to tribe
whmver holding it. This ia often accem-
panied by the excretion of a malodorme
fluid fromtheir cloacd bladdexe.
In regions with a pronounced mnual &-
matic cycle, mud trxrtEw spend the dry ~ a -Fig,48. Mnostemn creaeeri ECalaksnul,
son estivating while buried in mud te&., Campeche, Mexico). Photo: R.Cedefko V,

Kinosternon
-- . creugeri in the northern
Yucat4n Peninsula; LEE 1996).
Kinoakernids feead mostly on aquatic inver-
tebratm, such as snail#, warms, and in-
seds. Some s m e a (e.g., K leucostom~m)
also consume a large quantity of plant
material (seeds, fi-uits*flowers,and leaves);
however, whenever available, they also
prey on amdl vertebrabs &ha, hgs,
lizmh, aad an&ed including carrion.
During &e reproductive season, the fe-
male produces aeveral clutchw, whioh,
7 dlependmgj an the species, contains one ta
KiMsgerwn @%a$&WWW3 TBSiQ, Eke,
Pap. Mu*1Fool+,UaiT*
Id& *iZw mib
m h I&* ?i%mtatl," NSeca iCyqw!B
length tr, Ul m. Ntn%hern ~CECIOD; of
Y m b h Eenhdfil e4lrr level ba 100 m deva- Fig. 49. Kims&rnan worpkide8 (Gualaea,
kion,
Chiriquf, Panama). Photo: M. Lundherg
Fig.60.Kin&mon mrpioidea (near Porte- Fig. 51. Kindstenon leucostomurn (Tartuguero,
golpe, Guanawts, Costa Rim). Cwta Rica).
Photo: H. I3ringsoe

Key to Ximaternon
1 a Bridge very n m w , its width lees than
21% of carapace length ..............................
....................gi:nastemon angustipom
b Bridge less narrow, its width more than
21% of carapace length ............................
B
% a Length of p l a r scuta about half the
length of anterior moveabIe lobe of pEas-
tron (Fig.52a,b) ..................................
8
] fixed I&

b Length of gular scute distinctly less than posteriM

]?
half the length of anterior moveable lobe
of plastron (Fig,52c,d) ..........................
4
S a Anterior moveable lobe of plastron as long
as or shorter than b e d lobe (Fig. 52af b, K. creased
............................
Kdnosternom ocutltm
a, Knacdum

b Anterior moveable l o b of plastmn con-

spicuoualy longer than faed lobe (Fig.
52b) ....................
Xinosternon c m e d
4 a Anterior moveable lobe of plastmn con-
epicuou~ly longer than fixed lobe (Fig.
62c); axillary scute usually not in contact
with inguinal scute; carapace never with
more than one keel; a pale postorbital
stripe usually present (Fig. 47) ...................
.....................Efmaternon Ceumetomm
b Anbrior moveable lobe of plastron aa long
as Exed lobe (Fig. 52d); axillary scute
usually in contact with inguinal @cute; EBucOSlwnum d. K scarpi~ides
carapace usually with three keels; no pale Gs

...
postorbital stripe .............,.....,..........a.
......................
$i'nostemm empioides
Fig 62. Plastron of Kimternon species.
m a r y and inguinal ecutes man&.
Further Reading
I\~ER~OW I976 (Kinor~temnin Belize); IVERSON Adopted after LEE(1896).
1980a, b, I983b.
Biaummm
The genw Stattrotypas inincludes two sjm-
cies, the Pacific caast giant musk turtle
(S. aalvinii) and the Mexican giant musk
turtle (El.triparcatus), bath of which oecur
in Central America. These are aggressive
turtles that need to be handled with great
caution. With their strong, sharp jaws,
these animals can cause serious bite
wounds. The strongly develaped d o r d
kwh help to stabiliae the carapace, there-
by making it more difficult for predators
(e,g., C d y l u s naoretetii) to break the
etrong ehell. The speoies of this turtle
genus are strictly aquatic inhabitants of Fig. 63,Staurotypus triporcatva (Tabasco,
lakes and elow moving rivers with soft Mexico). &ta: J. H&
bottoms and rich aquatic vegetation.
Giant m u k turtle^ are not great swim-
mers;these animals prefer to walk mund
on the bottom of their aquatic habitat. Stcwm&pw d v i n i i GUY 1864, Roc. Zoot
~h~~ are c ~ V O m u sfeeding
, on crabs, Sot. London 1864: 127; type locality:
Huamuchil, QuaeEnala. Carapace length to
worms, snails, aquatic insects, amphib- ZliO -, central
ians, and fishes. Additionally, smaller oaam brldcto0,~1 mador,
turtles (eepeciallp of the genus Kino- ma leva1 to 700 m elevation.
stemn) are pmyd upon @ant Staumtypus t i p o m t w (WEGUN 1828),
mu& turtle and their shells broken with ~8 ~n 21: w;
the aid of thdr strong jaws (CAMPBEU Alvamdo, Vera-, tgXe
locllity:
Mexico. mapaw length
1998aI.During the repmdmtive seami, 3'79 On ~ ~ b vemant b e fmm ~
the female buries several clutches, each Veraornz, Mexico, through the base of
w i t h three to ten eggs, on the riverbank.
Tlse hdchlingrs open the eggshells &ex
Yueafi
.
W ~ Q M .. to wmfern MORdUr88, Bea
9lBVBtion.
120 to 207 d a p of incubation. At that
time, they have a she11 length of 25 to 31
mm B.8alttinii) or 30 to 36 mm (S.tripor-
catu8) (EWST & BARBOUR 1484, KOmm
lW71.

Fig. 64, Staumtypue soluinii I v e n t d view;

near Iztapa, Escuintla, Guatemala),
Key hStalamtypus
1 a DoxwIaterd hl~ extend the length of
cara ce from antedor t o po%teri~rmar-
ging(F&.66a); antaxior moveable lobe of
lastMn eh~rtertban posterior mweable
L .................statr-ptse *m**
b Dossolataral keels do not extend the
length of carapaoe, but end on 1st rtnd 4th
aleurals (Fig. 66b;k);antejtjor moveable lobe
of plastron hnger than poetmior moveable
lobe ......................
Staurotypur salvinil

Fig, 87, Adult Geochedone arEronariu (DariBn,

Panama). f ie .H.Birkahn

Tesrtudinidae
-

-
The red-footed tortoise (Ge~cheionecarbo-
a. ~aumtypustripmatus b. steum&pus ss~vinii naria), which occurs in Panama, is the
only South American tortoise that reaches
AM.55. Carapace with keels I~haded). m far north as southern Central Aglerica
(UGLER 1963). It is otherwise distributed
moatly in tropical South America. The spe-
cies was introduced by humans on several
Caribbean islands, ~ u c hae Isla de Maiz
Grande, Nicararrua (PRKTCHARD & TREBBAU
1984). 1n ~an&ia, the red-footed tortoise
-1 is f m d mostly in fore~tedareas and does
not occur in the humid savannas (LEGLER
1968). These diurnal animals dig burrows
in the ground into which they retreat at
night. Red-footed hrtoise~feed mainly on
grasses, flowere, fallen fruit, and carrian.
In Panama, this tortoise is hunted and
eaten by the local people, as i8 the case in
other parts of itsr range to the south, The
egg-laying season is from June through
September. During this period, the fe-
males bury their clutches with 2 to 15 eggrs
in the ground. Females probably produce
several clutchea per season. The eggs are
aomewhat elongate (40-59 mm x 34-48
Fig.56. Sbaumtypue sdwinii (near Izhpa, mrn) and have a rigid shell. After an incu-
Eacuintla, Guatemala). bation period of 116 to 185 days, the ju-
veniles hatch; they have a shell length of
39 to 46 mm (ERNST& B m o m 1989,
KOHLER1997). Reproductive maturity is
-
reached at a shell length of 200 to 260 mm ,
(LEGLER19631, I

H e r Reeding
LEGLER1B63; EWT & LEUTERITZ
1999.

Fig,59. Hawksbill ma turtle <Erefmochelya

imbricatal at a coral reef. Photo: J.Pichler

7
(Chelolniidae, Damochelyidae)
Sea W e s were already to be found around
the world during the Cretaceous period,
with fossil finds dating back at least 200

i
million years. Thus,the ancestors of our
modern-day sea turbles lived side-by-side
with the dinosaurs, but were able to
outlive the gigantic Plesiosaurus and
Ichthyosaurus. The modern speciea came
into existence around 10 - 60 million
yeam ago and eompriee, along with marine
h a n a s and sea ~nakes.the onlv marine
kptiles. All species of 'sea tugle~have
forelimbs that have evolved into paddle-
ahaped fins, with only one ar two- claws.
Further adaptations to the aquatic life
style are salt glands, by means of which
superfluom salt, which has been ingested
through feeding in sea water, is expelled.
Fig, 68. Juvenile Geoche~ curbomria These animals can reach adult weight of
(Cbrdoba, Colombia). r'hotu: M, Lundberg bebeen 35 and 600 kg.
The females crawl up onto certain beaches
at dusk, in order to dig a nest and deposit
their egg8. After filling in the nest, the
female returns to the sea during the same
night. The young sea turtles hatch after an
incubation period of approximately two
months and abandon the nest together far
the sea during the night. After attaining
sexual maturity a number of years later,
the adult femalea return to the original
beach for nesting. However, this generali- % ,$*
zation of a sea turtle's life cycle only applies
in a general way, The eight known apecies
of sea turtles vary considerably with
> * b..A - - - ~
Fig. 60. Green turtle8 (Chelonia mydare)
regard t o their geographical location, copulating. Photo: J. Pichler
mating cycles, and behavior. Moreover, the
various populations of a particular species
exhibit a number of differences due to geo-
graphical distribution. The following in-
formation about the nesting aeasona of the
individual species relates in each case to
Central America. Six species of sea turtles
from two families and five genera nest an
the beaches of Central America. There
u e d to be mass landings of sea turtles (so-
called "arribadaa" or "arribazones"1 in
many Central American countrie~(Costa

I
Rica, Honduras, Nicaragua, and Panama
to name a few), in which thousands of
females of the species Lepidochelys olfva-
cea and others came onto land all at once
in order to lay their eggs on one beach. Fig. 61. Lepidocklys oliuaceu neating at Playa
This phenomenon only takes place in Bavaria, Guanacaste, Costa Rica.
Costa Rica presently (CORNELIUS 1995).
Among the most important factors that
have led ta the rapid decrease in sea turtle
populations are their massive commercial
use (the colledian of eggs and killing of
adult turtles), the plundering of nests by
feral animals (dogs and pigs), shrimp
fi~hing,and sea pollution (ca~eaof death
due to ingestion of tar and plastic refuse).
Turtle products are offered in numerous
tropical and subtropical countrise as sou-
venirs for tourists. It is completely ir-
responsible and illegal to sell and abuse
sea turtles or products made h r n these
animals as souvenirs, in light of their
endangered status. Fig. 62. b a turtle hatchlings on their
dangernus walk to the sea. Phot~:J. PlchIer
Fig. 63. Green turtle (Chelonia mydas) with Fig.64. Loggerhead sea turtle (Caretfa
shark suckers. Photo: J. Pichler caretta). Photo: B.T).Bartlett

T h e loggerhead sea turtle (Caretta are located along the coast of the Mexican
caretta) is found worldwide, primarily in state of Michoacdn, as well as in El
subtropicalregiona, and less in tropical cli- Salvador CJiquilisco), Guatemala, and
mates. It i~ found primarily on the Costa Rica (Tortuguero). Single females
Caribbean side of Central America. The come on land in many areas along the
moet important nesting grounds of this Central American Pacific and Caribbean
species are located in Florida and South coasts to nest. During one nesting w w n ,
Carolina in the U.S.,while in Central which can last from August until January
Arnerica few specimens come onto land to (Pacific rside) or April untiI October
lay their eggs, mostly in the northeastern (Caribbean side), a female can produce 2 -
portion of the Yucatan Peninsula. In 5 clutchee (rarely up to 8) at intervals of 12
Carettrs carettu, egg laying has been docu- - 14 days, each with 38 - 120 (rarely up to
mented from April to July. After a single 195) eggs, h r n which the young hatch
short mating phase, a female produces 2 - after 50 - 75 days (MAGNUSON et al. 1990,
6 clutches, each with 40 - 190 eggs at MARQuEZ 1490). The animals reach sexual
internal6 of approximately two weeks. maturity between eight and t h i i n years
From these, the young will hatch after 65- of w. A female does not come on land
68 days of incubation (MAGNUSON et al. every year for nesting, rather every 2 - d
1490,MARQUEZ 1990). years. Green turtles subsist mainly on
vegetarim fare (various sea algae and sea
While it was earlier proposed that two ape- grass), but also feed upon sponges, mol-
cies of the genus Chlonia (green turtles) luscs, jellyfish, and fish,
populated the coaatal regiona of Central
America (12.m y d a on the Caribbean side The hawkebilY sea turtle (Erectmochelys
md C. agassizdi on the Pacific side), it Es imbricata) is found around the world in all
currently blieved that only a single s p - tropical seas and on both the Pacific, as
cies is represented in the genus (ERNST & well as the Caribbean side of Central
BARBOUR1989,KARL 8t BOWEN1999).The America, Important Central American
shell of C. mydas can measure up to 140 nesting grounds for this species are found
crn in length. Important nesting grounds on the coast of the Yucatan Peninsula, as
 30th species of the genus ZepidmheIyd
(ridley sea tawtle) are found on the
coasts of Central America, with the
Kemp's ridley sea turtle (I;, kempii) ody
faund on the Y u w t h Peninsula in thle
region, and the olive ridley sea turtle &
olirrcscea.) along the entire Pacific coast of
Central America. The latter is the most
common ~peciesof sea turtle an the Pacific
coast of Central America, with its mmt
important nesting &tee in Nicaragua
(Masachapa, Pachamil, Boqdta) add
Costa Rica (Nancite, Oationall (CORNELIUS
1896). Playa Nancite is EI beach in the
Ganta Roaa National Park in Costa Rica
that is naturally protected by the s h q
rocks and thorny bushes that surround it.
There, one can experience thrt spectacular
Fig, 65,Kemp'~ridley sea DudIe @pkkhelys "arribadas"from July until Swternbe1; in
which up to 8000L,oliuacea females crawl
up onto tha beach to lay h i r eggs within
a span of a few day@. Important nesting
gmmds for L.k~mpiiare located along the
well as near Tort~guera(Conta Rita), Gulf Coast, with the beaches near Rancho
while nesthg specimens have o d y been Nuevo (Tamaulipm, Mexico) of by far the
sporadically reported elsewhere. The egg- greatat importaaoeoeh the 1940"gp the
laying seaeon is from April until amateur fiImmaker Andm58 Hemm d m -
November, but it e e a according to re- mermtd a masa landing of appro-tely
gion: Meicn and Belize - April to Auguat; 40,000 &. kempii near Rancho Nuevo
-
Guatemala May taNovemhr; Hand- (MAGNUSUNet d.1890).Tbis type of mas8
- June to September; Nicaragua - May to landing is known only in the case af tha
O & k , Costa Rim - May to November specie# of the ganw LepidocWys, It is
(WUEZ 1996).During m e seaaon, each msumed that the simdt8neous hatching
f a a l e wmes on land o w or twice to busy of rnariy young increases the chances that
dutches cmaisting of 93 - 223 eggp each. the necessary number of turtlee survive, in
A femaIe only nests every 2 - 3 para. The epite of numemu predators. LepidoeheZys
incubation period lasts 47 - 75 days and kempii also diffess fmm the other sea turt-
the young have a @hell-lengthof 38 - 46 les in that it comes onto the beach during
mm. (MARQUEZ 1990). The hawksbill sea the day to lay its egp, while the other a p e
turtle is mainly carnivomus, and its food ciee nest at night. Alao remarkable is the
ranges from crabs, sponges, sea anemones, fact that the females of both these specie@
~ ~ and fish ta sea a& nearly every yew producing 2 - 7
jellyfish, m o l ~ u s cBQuid,
algae. Hawever, there are apparently clutche~,each with 60 - 140 eggs in one
great geographical ljifferena~with regard seasan. After an incubation p d a d of 46 -
to its range of faoh. According to the in- 66 days, the young hatch with a ahell
vestigati!ona by CARR Br STANCYK (19451, le@ of 36 - 45 nun, W Q U E Z lQ9O).
the hawksbill sea turtles in the Tartu- Ridley ma t d e s feed on crabs and o U l e ~
guem region (Costa Biw) feed primarily invertebrates (sponges, sea anemones, jel-
upon certain sponges. lyfish,and squid).
Fig. 66. Leatherback sea turtle (Lkrmochelys
mriacea) nesting at Playa Naranjo, Coeta Rica.
Photo: M,fianz~n
The largest turtle living today is the
leatherback sea turtle (Derrnmhelys
cori~cea),with the largest specimen ever
measured having a shell length of 256.5
cm.and a mas8 of 916 kg.(MAEQUEZ 1990).
Most specimens,however, have shells with
a length of 180 - 170 cm. This speciea can
be found all over the world, as they are
able to populate cooler seas than other sea
turtle apedes. Leatherback sea turtles
usually stay in the open ocean and are
obserrd only occasionally near the coast.
They can submerge to great depths.
Important beaches for neating in Central
America are located in Costa Rica (Playa
Naranjo, Playa Tarnarindo, between
Tortuguero and Lim6n) and Panama.
While the nesting seasan on the
Caribbean side lasts from April 20 October,
the ne~tingseason on the Pacific side lasts
from October to March,The females come
on land to nest every !&3 y e m so that
they can b u y 4 4 clutches of 61-126 eggs
each in a ~hg1.esearron. The incubation
period ie 60-78 days and the young have a
shell length of 61-68 rnm (MARQW 1990).
Leatherback sea turtles are mainly cami-
varous and feed on jellgtfish and pelagic
crustaceans, and occasionally also on sea
algae.
Fig. 67. Cheloraia rnydarr. Photo: J. Pichler
Cheloniidae
Camtta mmtta ~LISYNAES~~
-
l758X Sy~tema.
Nstuw, ed. 10: 197; t locali* u h d a a r I
T
hericanas.* Campam ength to 106 m, 1
Worldmida:in Central Anmica &v in the
~aribbsgn~
CbZolaie & (LtMNxtms 17Mt,
Naiame, ed. 10: 19% type lodity:
Peksi; imlm Adscmdoaibl.'' Crrrapace
leggth to 140 om.Worldmide.
E&mnehelp imbrhta ELINNA$ua 17061,
Sp- Naturae, ed. lk SM1; type locality:
"Md Amesi~ano,h t i c o . " Catapace length
to 114 cm.Worldwide.
kpidocIseEy~ knapii ( ( 3 m l a ) , Bull,
Mu& Cpmp. Zool. & 138;type l d t y : of
Mexico. C ~ Olength B to '76 om. Gulf of
BkkSim
Lepidochelye oliuscea @N~CHOL'PZ 18231,
zcwl. Atlaa 1: a; typa wts". hlallih Bay,
Philippiins Carapace len&l~to 76 an,World-
wide; m Centsal Am&a only in the Padc.
Dammhelyidae
Dermsckelya eoriaeea ILmmus 17661,
L
8y1bma Naturae, ed. l5k 350; type locality:
4Lmatier Qmbni o m m agro Lauwntiana."
Carapace length to %6,6 cm.WmIdwide.
 . imbricata.
rlg. b ~~retmochelys Photo: d. Pichler
Key ta eea twtlear
X a Shell mered with a continuous layer of
undivided skin (Fig, 70a) .........................
............................ De-&€ye EOWJ~
b fie11 cwered wikh distinct horny SCU~RS
.
(Fig.7Ob-f) ...............,.-..........+.q,..,...........
3
a. Chelonle
B a Four pleural (= costal) acukea resent on
each side ideg. ?Oe,d); n u c d kmhl)
ecute not in contact ~ i t hfirst pIefural
Fig w.Lateral view of herd (prekntal scalea
brmn). ~~halsothe &~k.]& in
mm .....................................................a EretmmhEya
b E'iue or more plwral(= watal) s c u k pres-
eat on each aide (Fig. 7(Je,fl;nuchal
(cervical) s a t e in contact with first pleu-
A SCUM ............................... ...........
,..,4 Carap- d sea
8 a One air of p r e f s o n a beak not hawk-
~ i k e!fig. BW; w t e s of rarapsos non-
imbricate .....................C b h i a e
b 'hKO pair^ of prehntals (Fig. 69b); beak
hawk-like;seutee of carapace imbricate
.......................
EmtnmheZyu i d r i c a t r s
4 a Bridge u a d l y with t h i & m a l
acytea, each withwt pores; three or more
Mamandibular scale3 behind beak on
each aide of lower jaw ....Conetca oafst€a
b Bridge usually with few infrarnenal
pe*q ea& ~ t ahpore gt msbriw "0 Demhelm
~ ffaa b. C@m' em"
W r :siagle large infrarnrndibular8 4 %
behind beak on each side of lower jaw ....B
8 a Usually 6 pleural scutes (Fig. 70e): ground
color $say ..............X4danJae&u kempii
b U~ually6 or more pleural wtes (fig. 700;
gmund color olive hplaEacirelye div-a

Further Reading
CARR3867, H~RTHlQ8Oq b, muTcNARD 1980,
4.1880,M&QUEE1990,WILSON
b h 0 ~ 8 0 P Iet
i Zuo 1091, CORNELIUS1995, ZUG L Ems
1948. c. Chef& mydes d. EmtmocheIys Irnbrhta
 Lizards (Sauria)

There are 230 species of lizards in 10 fami-

lies and 60 genera that occur in Central
America.

Key ta families of lizards

1 a Third and fourth toe about same length
.................................
Helodermatidae
a- view
suboculan
b ~ourthtoe distinctly longer than third tee
............................................................
....!a
B a Two pairs o f sealea between mstral plate
and firat unpaired enout plate (Fig, 7Sa)
...............................................
Anguidae
b Fewer than two pair^ of ~calesbetween
rostral plate and &st un abed snout
plate (Fig. 73b) or snout ec ea small and 3
irregular ...................................... 3
9 a Scales on dorsum and venter of body simi-
lar in shape and aize ...............................
4
b Scales on dorsum and venter of body dif-
ferent in shape and eiae, ventral ecalea
mually larger than dorsal scales ............5 b, view

4 a No eyelids present ................................... Kg. 72. Head scalation characters important

............,..
Gymnophthalmidw (in pad) in the identification of lizarda.
b Eyelib present ..................... Bainaidae
6 a Venter covered with large plates (Figs, TI
b-d) ...............
............. 0
b Venter covered with mall scales (Fig.
?la) ................................ .........,". 8

n
a. Dactyk~18tRmns deppllj
b. &pld~w~@Iis
a. Celestus b. Mabuycl

Fig. 73. Scalation of dorsal surface of head.

Fig. 74.
Scalation of
dorsal ~Urpace
of head Imtral
brown;fronto-
llgSal orange),

Fig. Tl. Ventral aods in lizards. b. Anedie

54
8 a No eyelids premnt ...............Xmmtwiida#
b Eyelids p m n t ....,..............................1
1 a htamasals present, rostraI separate Brom
frontooaasal (Fig. 74a) ..............,....lbiidae
b Wfthout internmala, roetral in contact
with $$antenasal (Fig, 74b).......................,
...............QymnophthWdae(inpart>.
8 rt h a l d a c e of head covered w i t h sub
equal granular 8caIes ..............................9
b Dorsal surface of head covered with non-
granular s c a b of M a r e n t aims ........-10
9 a No eyelib present ...............Wdsonlldae
b Eyelids present ..............Eublepharidae
10a Doramn of body with conspimow beaded
d e s ................................
ILersogauFidae
b Domum of body without conepicuowa
beadd 1pcales ..........................
Iguaddm
Fig.77. Abmniu smidhi (El Triunfo, Chiapas,
Mexico). Photo: k Ramirez

An&& are cosmopolitan, with six genera

occurring in Central America. Most alli-
gater Eizds have square scales that are
arranged in ring8 md strengthened by
bony dermal plates (oateodems). While
representatives of the genera Mesapis and
&whonotus are found more commonly7
the other species are known from only a
few specimens. This could be due to the
fad that the "rarenspecies Iive in habitats 3 a Longitudinal
(i-e., treetops) where, for the mmt part,
they evade our observation.
Fig, 76. Lateral view of toe (oaly sheath
shaded).
Eig, 76.LateraI
head inM e s p i a
~preocularbro~11
and euboculers
orangel.
Key to genera of hguidae
1 a Body without di~tinctlongitudinal fold 2 ..
b Body with a &tinct longitudinal fold ....8
I a Clawa almost completely covered by scales
(Fig, 76a) .........................., Dip10gkl88~8 .
b Claws not. cavered by males (Fig, 76b)
..
...................... .........................
Cehaturr
fold between ear opening
and shoulder reduced or absent; head
more or lees flattened, often helmet-like
...................................................
Abrvniu
b Longitudinal fold between ear opening
and shoulder conspicuously developed;
head not flattened, never helmet-like 4 ....
4 a Suboculare differentiated from preoculars
and postoculars (Fig. 761, not in a single
continuou series; h e pairs of large and
one pair of small chin shield8 ...........
........................ ,.,. ,
................Mentapin
b SYboculars, preoculars and patoculars in
a aingle mntmuous series of similar 8ca-
les; four or five pairs of large chin shields -
6 a 10 Iongitudind rows of ventral scales; 12
nuebds; five paim of large chin shields;
usually two pstmtrals ........
Coloptylchon
b 12 or more longitudinal rows of ventral
scales; 10 nu&&; four pairs of large chin
shie1dB;0-1 postm~trals........Gerrhonobue
 - -- - - -

The tree-dwelling alligator lizards of the

genus Abronia are distributed in Mexico
-
P
and northern Central America, While
most speciea live in highlands between
1500 and 3000 m above sea level in pine,
oak, or cloud forest, the species of the
I
,
bogerti group (A. bogerti, A, chkari, R
ramirezi) are inhabitants of more tropical
pre-montane habitats between 360 and
1400 rn above sea level (H. S m & SMITH
1981, CAMPBELL1994),Abronia are diur- Fig. 78. Abranta rcarniwi ( C e m La Vela,
nal and stay mostly up to 40 m above Chiapas, Mexico). Photo:A. Ramirez V.
ground level in trees that are overgrown
with epiphytes, where they can climb

II
among the bmmeliada, ferns, and orchids
(CAMFBELL& FROST1993). The long pre-
hensile toil senree the alligator lizard as a
fifZh leg and, thus, coslsCitutes an impor-
tant climbing aid. Occasionally, however,
the animals will descend to the mound,
whsre they have been found bene&h piec-
es of tree bark and tree trunks. and alao I
sitting openly on the leaf- or needle-cover-
d ground (BOGERT& PORTER 1967, I
C A ~ ~ B E&LLFROST 1993,F R A N &
1999, MCCRANIEpers. comm. 1997).These
lizards have a calm nature. CAMPBELL
~N

FROST(1993) reported a gravid female A.

gt
I
Ii
-
r9,
nia 5riata IAlta 8,
Guatemala). Photo: J.k Campbell
fimbriata that they observed in the quiet
inlet of a smdl mountain stream. All epe-
ciea of the genw Abronia, far which repro-
ductive data exist, are live-bearing, with
the number of young per litter varying
between one and twelve (WERLER1951,
MARTIN 1955, H. SMITH & ALVAREZDEL
To~o1962, ~ V A R E ZDEL TORO 1973,
CAMPBELL& FROST 1993).The coloration of
the juveniles varies from species to spe-
cies. but consists mostly of dark lateral I
dorsal stripes on a pale background
~CAMPBEU & ~ O S T19931. Due to their I
restricted geographical ranges and the
riek of habitat destruction, most species of I
I
Abronia are conaidered amongst the most
endangered species of lizards anywhere.
One can assume that some specie8 have
even Fig. 80.Ab row.- campbslli (Jalapa,
& IFkOm 1993).
be discmered (CAMPBELI, Guatemala). Photo: J. A. CmpbeU
 Abmnia enzwtoi C m m & 199Sl
Bull. Amerr Mus. Nat. Hiet. 5116: 22; type
locality: Finca Roeario Vllsta Hermoaa, 1219
m elevation, on S slope of Volch de Agua,
Eeeuintla, Guatemala. SVt to 135 mm.
h t h e r n slopes of Volcrh de Agua in south-
ern oentral Guatemala, 1220-2290m elevati-
w in cloud foreat.
Abmnia autdta (COPE18691,Pme.Acad. Nat.
ki.Philadelphia 80: 306; type locality: Forest
of Verapaz, Guatemala. SVL to 126 mm.
Central Biahlanda of Guatemala, 2000-2660
Im e1~vati&in pine-oak foreat. A female ve
birth to 12 iwenilas between 28 anp28
J&uay I~)~~(CAF&PBELL i% FROST 1993).
II
Abmnla Bagerti TIHEN 1964, h e r . Mus. Fig*81,Ahronia lythmhilc Cnm §an
Nwitates 1687: 3; type l d i t y : N of Niltapc, Criet6bd de las Cmw, Ghiapaa, Mexical.
between Cerro Atravesad0 and Sierra Madre,
Oazaca, Mexico. SVL to 64 mm. Emtern
Qaxaca, Mexico, 760-1570m elevation in rain
hrest and lower cloud forest. I

bI
Abronh cnmpbslli BRODE& SAVAGE 1983,
H etolagica 49: 421; type locality: C e m
h%n ds lu Minu nw Pmtoda, 14*4W,
8Do67'W, 1800-1900 m elevation, Jalapa,
Guatemala. SVL to 127 mm. Eastern
I~uatemala.1800-1900 m elevation in h e -
oak foreat.
Abronia fhnhr-iata (COPE1886),Prw. h e r ,
Pbil. Soc. 18:971;type locality: Pine forast of
Alta Verapaa, Guatemala. EVL ta 130 mm.
Central-eastern Guatemala, 1400-2100 m
elmtion in cloud forest, A female gave birth
ta four juvsniles (33 mm SVL) on 15 June ;. .,
(CAMPBELL & FROBT19931, 1

~ b mti~ ~ sAs4
b ACEVEDO & Fig. 82. Abmnia mntecristoi EQuebrada Grande,
MENDEMON1998,Herpetologica Bid: 222;type C Q P i30~dura)-
~ Photo: J. R. McCranie
locality: along road to Patalcal, 2835 m eleva-
tion, 15"51'lS'TJ, 91°31'02W, 5.9 km (by
road) NIN interadon of Guatemala Road 9N
(near San Mateo Irtetdnl, S i m de Lm
Cuchumaknea, Huehuetenango, Guatemala.
9VL to 101 mm.Northwestern Guatemala,
2836 m elevation in cloud forest. F': j
,.,,,::;,i\ihi\Lj$
Abmnia rabphantasnaa CAMPBELL & FR08T
1893, B&. Amer. Mus. Nat. Hid. SIB: 1s;
type locality: vicinity of La Unib Barrios,
II
1600 m elevation, on W alopa of Cerro Verde,
Baja Verripa, Guatemala SVL to 110 mm.
Ceritral-eastern Guatemala, 1600-2360 m
elevation in pine and cloud forest.
Abmnia EeumEepb CAMPBELL & FROST 1993,
Bull. Amer. Mug. Nat. Hi&. $16;36; type lo-
cality: Santa Rosa, near Cornitth, Chiapae,
Mexim. SVL to 106 nun. Only known fmm =
the localit%amta in the vicinity of Fig. 83.Abmnb m a t h i (Aguas&orginas, Quetz-
Comith, in -tern Chiam,Medco, 18011- dun- Guatemala). Photo: L.Melendez
2300 m elevation in cloud forae'
57
 Fig.84,Abmnia aurita [El :1&8,
Guatemalal. mob:J. A. Campbell

t a w I-tmp;UwJ 1Q&8 thFitlp.

IWL N& HM. UW* ZWM I& &;
Lad$ Man*). - m a a,
d m a. datqsxf-yaaan,
El Wvdarr. SVL iio BQ slaz, N a W
park BBOIIMB~ at the SumStwe oP
~ w a d b p ) I o a 4 mm ~d~ ~a*
$Pebrada G d r e in mtem Bmhrug
W ?&m Om tdwct4m h d d Parest.
b P a (- m tat&?o
a%w3,Anal,Inet.& L , W 10:8s.P tgp
Lac&@ Bmfa Rpaa, QamitBa3 ~dqbm
lkkdmi m !3? am.
M&m, 1~00-$900m *abon
ht.5 m
tntm
a
Ftmwk
Key to Abm~da
1 a 10 longitudinal ventral scale row8 .......
...................................Abmnh mid
b or more longitudinalventral scale rows
.....................,.
. . . . . . . . . . . . . . . . . . . . .m .,..*..,... B
B a Cona imus cmpra-auriculm spin- pre-
mnt k g . 81) 8 ................................ ...
b Withcut supra-sruridar spiaea (Fig.88)
................................... .,........................18
S a Buprana~alsin contact with each otheq
thereby byparating =briar and posterior
paira of internuah; &ontonasal reale
a b n t .................... Abmm& fpmbrtatat ..
otlier* therefore anbrimr a d postmior
paiFs of internasals in contact; frontanam1 Fig. 86. Abmnio 64gerti ( C m o Beta, O ~ X ~ E B ,
scale preeent or absent 4 ............................ Mexico), Photo: T.Bile
4 a 12 long'ihdinal vmfial scale rows 6 ..........
b 14-16 longitudinal ventral scale rowrr .... T
5 a Frontonasalabmnt; h a l body d e s with- b A bright ring around eye present, this
out keeIs;poshrionn& Malabial cornpic- color stdckgly different from that of rest
l z ~ w &elongate .........Abrunia hmEspl;e of head ......................... ..,....................
9
b Fmptoeaeal present; dorsal body soales 8 a 14-15 longitudinal dorsal scale rows;dor-
keeled; posteriormost infralabial not can- sal color variable ...Abronia lythmhlZa
spicuously e l w g a t ~ 0 ................................. b 12 longitudinal dorsal scale mws;doreurn
8 a Posterior s u h l a r in contact with lower brownish .......Alimnia galophsmtmma
antexior Cempord; prsapriculm granular 0 a 32-36 transverse ventral scale rows;
and in several rows; three primary temp- ground wlor gray to brown; circumorbitrtl
orals; no bright yellow ring around eye region pale brown to cream color ...............
present ................Abmmiss ochpkmfd ...................................
Abmnia campbeig
b Posterior eubacular not in contaot with b 35-38 transveree ventral scale rows;
lower anterior temporal; preauriculara ground color green; ckcumarbital region
usually subimbricate to imbricate md in yellow to orange .................................. IQ
two m e ; usualty 4-5 prImary temporah
a bright yellow ring araund eye presm$ 10a Dorsal body scales only sparsely pigmen-
thb color strikingly Merent from that of d with black ............
i Abmnia anzuetoi
mt of head .....................Atrmnia s d k l b Dorsal body scales heavily pigmented with
T a Cirmglmbital region not colored different-
black, anterior portion of scaled almost
ly frommt ofhead 8 .................................. entirely dark ......................................
11
Ila 9-11 supra-auricular spines ..................
...............................
Abrunio metMona
b 4-7 supra-auricular spines ...................
18

1 Abronia 1Ba Dorsal body scales green in life with black

pigment ........................
Abmaia aurita
b Dorsal body scales yellowish green to tur-
quoise in life with black pigment
.............................
A h n h vcsecombsii
13a Supranasale very large and in contact
with each other, thereby separating anter-
ior from posterior pairs of internasals;
posterior and doreal edges of dmsd scales
distinctly pale colored Abmndu o m h i
b Supranasals, if reaent, small md not in
contact with eacE other, therefore anterior
 and posterior pairs of intermeale in mtact;
coloraticm o f d o i d s c a l e e v d e ..........14
14a Parietal in contact with supraoculars 16 .,.
b Parietal scale not in mntact with mpm-
o d ..,............17
m ......,..........,..............,.
16a Dorsal body patkern c & t ~ of distinct
altermaking white Or yellow trmaveree
bars; 12 l o n g i t u ~
doreal scale rows; 28-
32 tranmma dorsal soale rows; 33-36
tranwezse ventral scde rows ...............
.......................................
Abm?mB f h s t i
b Dorsal body pattern different from that
indicated above; 14-18 longitudinal dorsal
male rcws; 38-41transverse dorsal scale Fig 88.Abmnia lytkmhila (near San
rows; 96-65 i5anmeme ventral a l e row

- 16rr Supra-auricular B&BS slightly prbjacting

but not s he-like; prefrontal not in
mntack nit! auperciliarh; 4 i-dary
C r i s t 6 d de lae Cam, Cbiapas, Mexioo). Note
supra-auricular spines (arrewl.

and 4 tertiary te~dparals.6 r o w of nu-

scales ...........................
~kmnia mat&cri
b Sum-auricularscale%not diffbrentiatd;
psefrontd in cantact with m p ~ a r i 0 63;
secondary and 3 tertiary temporah; 8
b
rows of nuchd ~ a b e .....................
...............,.................,.... .,.
Abrrrntdc bogeN
14a 6 occipitals; 8 primary temporals in contact
W i t h posbcdme ...A&ronialtl~mtecriaai
b 1-8&pitale; 2 primary
with lao~t~eulw
~~in ~ontast
....Abromia s a w &
Fig: 88.Abmnia ~.aabvladoremis(Zacate Blanm,
sacunde-ry temporals primary temporal8 Intibud, Hondu~as), Phah J.R. McCranie
\ / ~ostDeulara .loreal

Fig. 89. Cele~tuemdk (Quintma b,

Mdw). Photo: B.Eahena B.

Farther RRladins
TIREN1956,CAMPFELL & FROST 1993,K~HLER
Fig, 86.W t i o n on l a M and dmsal aurf~108of 19%k, C A M ~ E LetLel. 29B8,-C et al,
head in&m& m&&. 1998,MCCW & WILSON1999
 13 species of Mexican and Central
American anguids (11 of them in Central
America) share the common trait of a
cylindrical body without a longitudinal
fold. These epecies are placed into two
genera (Celestus and Diploglosswms) by
aome authom (PETERS & DONOSO-BARROS
1970, SAVAGE& LIPS 1994, MCCEEANDE &
Wmo~14461,while others (CWBEZ;L &
CAMARIUO1994)combine.all these species
in the genus DiplogEossus. UntiI more is
known about the evolutionary relation-
ships of these species, I believe that it is
beat to retain the two genera concept. Of
the ten described species of the genus
Celestua, eight are found in Central
America, whereas two (C. legnotus and C,
enneagrarnmus) are found $&her north-
I west in Mexico. While most Celestus spe-
cies are highlands inhabitants that live
more than 1000 m above sea level in the
pine-oak forest (i.e.,C, bivittatus, C.cyano-
chloris, C, montunu& C. orobius, and C.
scansorius), C. cyanochloris, C. hylaius,
and C. rozellae are found in lowland rain
forest. Some species are obvious tree-dwel-
lers (i.e.,C. atitlansis, C. mzellae, and C.
scansorius), which spend their time at a
considerable height above ground.
However, individual8 of these same spe-
cies have been observed basking on tree
trunks only 50-250 cm above the ground
(UE 1996, MCCRANIE& W ~ 1996).
N
Other species (e.g.C. bivittatus) have been
found predominantly under fallen logs
(Wnson et al. 1986).Celestu~rozellae is a Fig.90. Celestua hylaim (Cornadre, Costa Rica).
livebearer, producing clutches of 3-6 juve- Photo: R. W.Van Devender
! niles during the months from May to July
I (ALVAREZDEL TORO 1983). A female
Celestus bivittatus from Honduras gave
b i h to tee juveniles (SVL28.2-30.7 mm)
with bright red-orange tails on the 1~~ of
March 1980 (WIUON et al. 1986).

Fig. 81. Cdestus scanmrius (near La Fortuna,

Y m ,Honduras). Photo: J. R McCranie

Fig. 92, Celeatus bivittatua (Honduras).
Photo: J. R. McCranie
Pedro Bnla, Honduras, SVL to 98 m.
Marthwerrtern Honduras and adjacent
&atemala, 915-1370m elevation,
?eleatus ombiug WAGE %r. ZrPb: 1994, RePv.
Bbl. Tmp. 41: 823; type lacalitg: area near
- Kohnsia, h l m a and Fortuna on the
Carretera Intararnericana, XM10-2000 m el*
vation# Canton Wms-Zkledirn, Cordillera de
I'alamanoa, San Jorrk, Costa Rica. SVL to 88
mm. Cordillera de Talamanea, C o ~ t aRicsl,
1600-2000m elevation.
7elesbus rozellm S m 1942b, Froc, U.a
Natl. Mu. @&372, type locality: Palsnqu~,
Chinpas, Mexico, SVL to 102 mm. Swthern
Mexico through northern Guatemala ta
Behe and the Sierra de Eepirita Santo near
Lbe Guatemalan-Honduran borderI sea levwl
k~1350 m.
Eelesturr 8cansoriu.s MCCRAME& WILSON
1996, Rsv. Bid. Tro 44 260; type locali*
1.5 airke Yim d La Fort-
Slo19W, 1560 m elevation, Cordillera
16O26.N,
Yombre de Dios, Yoro, Honduras. WL to 111
m. Northwedrn Hondurae, 1560-1590go
atevation.
1 a Eluboculars and postoculars in a single
continuow aerie8 Cebstus eyanoclilde
b Subocular and postocular series juxta-
posed .......................................................
3
2 P 14-18lamella* under 4th tae ..................S
b 20-27lamellae under 4th toe .................. 4
3a 17-18 lamellae under 4th toe; SVL to 110
mrn .....................CeIestus atiIZoneneis
b 14-16, rarely 17,lamellae under 4th toe;
SVL to 90 mm ..........Celestus biuihlrrdue
4 a Three almoet subequal prefrontal plates,
median prefiontal separated &om supra-
oculars; Bank pattern consists of occelated
blotches ...................
Celeetue mondanrss
b One single large prefrontal plate, in
contact with supraoculars; flank pattern
.
.
not as a hove ............................. ....8
5 a Fewer than 70 transverne r m of dorsal
scales ...........................
CaLshce oroMtrrs
b 73-81transverse rows of dorsal scales .. 8
8 a No pale dorsolateral stripes; pale vertical
bars on neck present ...Cekstrrs maellae
b Pale domdateral stripes preeent; pattern
on neck not as above ...............................
7
7a 84-92 transveree row8 of ventral ~cales;
76-81 tramverse rows of dorsal scales
..................................
Celeshs hylaiw
b 76-78 transvame rowe of ventral scales; 74
transvmae rowe of ventral e c a h ............
.............................Celaahs seansoriue
Fwther Reading
Vws, & WIWN 1988,CAMPBELL& CWILLO
1994,SAME & LIPS1994, M c C w & W ~ O N
1996.
I
 The genus Ccdoptydm m~ists of a single
~ ~ eC. rs b d, i f i r @%mms 1877). The
holatype of C, rbmbifer was eoIlectd in
the praviaoe d GhMqui in Pawam. Since
the ~poies'original desmiption mom hhm
120 Y W 8 w,0nly tW0 a d d i t i d SP&-
me= d C, r b d i f i r have been diecover-
d,b& mf them in the rainfomts of
mthe&em Cwta R i a (G?ooD ISM}. The
n am btgr of thh 4pedw 53 *ually
unknwvn. C~lopbckonrbrnbifkr h a au
elongate 15ender b d p and a aompicuou%11y
1 hangtail Crsakhg &a and half h a s the
SVL). A P ~ ~ B Pof: aW ~
93,aIPwaa lwhm hll* er
S P ~ * a phch ip Goon (19481.~ Y BLateral
I : vim d
men efthis sped- WW pubwed by WD had (from 11vw@m et Q. 1870-1909).
[1988). The domd b d y pattern oonsiats of
eight pala mbbiangalrtr m ~ ~ b e c onrs a
I dark ba&gmmd. The tail is a h banded,
1 wh~raatlthe 1- &W O ~ Y indistinct C

mottling. DBbEOsBw
~pe:dm
'I% $~lfthe genm DiplogIrrgsw are
staut-bodied lizards, the epwies El.
mendrepis b e i the mixst ~ ~ 1 add
0 ~ 1
also the large& angolid bard of Central
America, maehhg a maximum SVL of 215
men. The- lizardsl me mostly &round
dwprller~,but can dm be observed c W -
ing on wi%habundant vine growth, A
female of 8, BiloBiatu~w a ~discwend
Bene~ththe trwlk af a tre w h it
guarded a clutch of sirr e&gs (T~yz0.RlE€i).
 1
Key to DigEogloseue
1 a A pair of prefiontals and one median
frontonad (Fig. 96a); dorsal scales dis-
tinctly keeled;SVL to 215 mm ...................
.....................,..Diptaglaerrus moiwtmpte
b A aingle large prefnontd (Fig. 96b); doreal
d s s not or anly faintly k e e l 4 SVL ta
93 ..........,.........
~ . , . . . . q . . . . - 4 . . ~ . ~ + r r r r r r r r r2

First wpraIabial Beparates nasal from

M B I ~ ; 01312 p~~tIlM81 ..............................
.................DtpEogEoaeun montieilvestdn
I b Nmal i.m n M with maYal; two pm8a-
B& btlobahde
s.,rtm~-vrtr...Diplo~loa~a~~
Fig. 95. Dipluglossus nomtmgia (Ri9 San
Juan, Nicaragua).

DipEogEossas b-w .~~'$I~AUQHNE~

1874), Ann. Mag. Nat. Riat, (4) 14: 257; type
lacali* Costa Rica. SVL to 96 mfn, C a ~ t a
R i a and wetitern Panama, =a level to 1380
m elevation in rain farest.
Diplcgicmsua monatropia ( K U ~ ~ L 15201,
.
&ib&pmu EooZoologie und vergle~chenden
h l a n r i e : 128; type l&tyl unknm.SVL
to 236 mm. Southern Nicaragua to weetern
Eeuad6r, eea level to 1000 rn elevation in rain
fore&
Diphg$@mue montidvcst~is M ~ C W 1973, &, 0.mommpls b. GI. bfkrbahrs
Amer. Mw. Novit. 8689: 3; type locality: SE
elope of Cerm Pine, 1440 m elevation, ~ g96.. Bcalation of dmal surfam of head
Serranla de Pirre, DariBn, M a . SVL to (frcmtod pmfronm -1).
100 mm.E m r n Panama,near b d 6 r with
Colombia, 14.40m elavation m daud f o m b
R ~ Q ~ ~ ~
TAWR 1966,M ~ R 1973. S
 color, this species exhibits irregular dark
brown cross bars on the dorsal surfaces of
the body and tail.The venter is dirty white
Four species of Grrhonotus are currently to pale yellow, The short legs are wed only
recognized, only one of which, G. lioceph- for slow movement, whereas for more
alus, occurs in C e n t d America GOOD rapid movement, when, for example, it is
1994). The other speciw are distributed in trying to escape, it will dart away with
Mexica and the. southern USA. undulating body movements, with the lega
Gesrhonotus liwephalus is a very elongate adpressed to the body. Gerrhonatue lioce-
lizard with a very long and breakable taiI. phalus can be found both on the ground
Because of its extremely slender appear- and on low vegetation (bushes and small
ance, this lizard is called 'lagarto culebra" treea). When climbing, it utiIizes its pre-
(= "make-lizard*) in Chiapas (ALVAREZ DEL hensile tail as a fifkh limb to hold onto the
T o ~ o1960). On an olive-brown ground vegetation. It feeds mostly on insects,
which are slowly approached and then
grasped with its strong jawa in a fast for-
ward motion of the body. The females pro-
duce clutches with 10 to 16 eggs that are
about 13x8 mm in size (A~VAREZ DEL TORO
1960).

F'urther Reading
~vATCZSDEL TOM 1983,&OD 1994.
 The genus Maaspis ki composed of ~ i x
species, two of which o c m in Central
America: Mesaspis rnonticola inhabite the
mountain regione of Costa Rica and
western Pmama, M.moreletii is found in

-
the highlands of northern Central
America (GOOD 1988). Both species are
ground dwellem that prefer open pine
forest, p & m o and clearing8 in the cloud
forest at elevations up to 3260 m (e.g. on
the Cemo de la Muerte, Coata Rica). Like Fig. 98.MesagpZs manticola (Cam de la

- most Central American anguids, these Muerb, Costa Rita).

agile lizards prey mostly on insects and
other arthmpods, Both species are vivi.
pamm. Once a year, after a gestation peri-
Photo: M.Franzen

od of about twelve months, the female M.

monticob will give birth to a litter of 2 to
10 young. The newborn M. manticola
measure 60-57 mrn 'IZ with a SVL of 23-26
nun (FITCH1970,1973a, VIAL& STEW^
1986, WICKNICK1993).

Meaaspia rnareletii IBQCQURT16711, B31.

Nouv. h h . Mm. Hist.Nat Paris 7: 102; type
10caliB: k M~
~2
~ w a ~ d u m mii
s probably northern
, Fig,99.Meampid nwmletii
" (El PiW, H ~ ~
ndw)+

N i m 1500-50M+ m elevation.

Key ta Meeaepis
1 a 14-16 longitudinal rows of darsd wales;
prefrontal wale usually present .................
.................................Menampia m n t i c o k
b 18-20 longitudinal rowa of dorsal scales;
prefrontaI scale usualiy absent ..................
..................................MBBQIP~Smmhtii
Further Reading
TATLOR1966, FITCH1970, 1973a, V w &
Qmm1088, Goon 1988, 1989, W I ~ C K
1993,VE~ELY
& KC,HLER 2001.
Eublepharidae
Currently, the geckos with moveable
eyelids are separated aa, a digtinct family
(Eublepharidae) fmm the other geckos
(family Gekkanidael (KLUGE1887, GRB-
l k l ~19883.
~ The family Eublepharidae t
widely distributed in the Old World but in
Central America represented only by the
genus CoEeonyx. Banded geckos are char-
acterized by moveable eyelids and a strik-
ing banded domal pattern. Their subdigi-
td lamellae are neither broadened nor
adhesive.

-
Two species of the genus Cokeonyx occur in
-
.
Kg.101. CoIeonyr tnttratua (SaeSalvador, El
Salvador).

Central America. Coleonyx slegans and C

rnitratue are closely related. Both are tsr-
restrial nocturnal lizarde that hide
beneath mcks and fallen tree trunks and
in hollow tree trunks durlng the daytime.
Their diets consi~tmostly of insects and
qidem. The movements of these attrac-
tively patterned geckos are very graceful,
almost cat-like. When alarmed, they
expand their t h a t and asgume a stiff
posture with the body held high, well off 1
the ground. Both species are oviparous
and produce clutches consisting of one or
two eggs each. At a temperature of 28-
1
30°C. the earn hatch aRer 62-68 dam (C.
elegahs) a i i 46-65 days (C. mitritus), Fig. 102. Coleonyx efqans (near Chetumal,
respectively ( ~ B E R L A H 1996, ~ H L E R Quintana Roo, Mexico). Photo: H.Bahena B.
1997).

Cdeonyx skgans GRAY 1846, Ann. Mag.Nat.

Fig. 100.Coleonyx mitmtus (Isla de Utila,
Honduras).
Ket. (11 16: 163; type l d t y : Beliza. SVL to
97 mm. On the Fa& veeaat from southern
N@t, Mdeci, to w e a m El Salvador, aad
on the Caribhn veraant fhm Yamma,
Mexioo, through the Yucatan PBninda b
northern Guatemala and Belize, sea level to
1000 m elevation.
C&ORYX mitdw (Pm%~s 1863), Monatsber.
l&g. Ahd. Wim. Berlin 188s: 4% type lo-
cality: Panama.SVL to 91 mm. Guatemala b
C n s h Rica on the Pacific vereant and
norWesn H o n d m on the Caribbeian ver-
aant, rea h e 1 to 700 m elevatim,
 G s c h of the family bkkardd~eorxrur
tbroughaut much of the tropical and ~ u b -
tm@cal regione of both tbe Old and New
Worlb. In C m b d America, thia family is
represented nille genera with 24 ape-
cia,Most & specie5 are chara&rl~d
by broadened adhe~livelarnellae bemath
fimaad -8 (d10Wh~ *W B J h d 8 to
Fig.103. Cde~nyxe@sw / M ~ M ) clikb on smooth,~ertieafgnrface8)~ m well
fiQta:R* Ihtlett as by the absaam pf miwoable eyaW
(their eye8 are m e n d by a traxwparent
spctade inatead). &me gecko apeciee are
crepurscular or nocturnal (e.g., Hemi-
dactylus and P h y l E d a r ~ ~ y i ~whereas
s$,
&By ta C o k m x othens: such as wciea of the gmwer.

J
~ o n a l t d and
s rSphmrodac@Jus arrt aativs
a 4 9 eheath ahmitt(Fig. 1W;qfet sublabid
mmpialy b~tri- a
ang;ular; aa d f ~ e of
~ dd l mrch d d n g the day. All Central America
W d "s"...... 6 cdmysekggns ge&o s&ea are ariparwa, us& pro-
Ib Claw8 not: wmed by a s d y ghmt;h (Fig. ducing one m two ~ E Pp~ clua.
IO4b); $rei;ahlabid s q d h ; no field of
emdl d e bhhd ................
......,,,.....,..,,...,.,.,~~~ internueJsn~rldrr~tass Kay tv genera ofG d & o a f h
I* Digits lGri& ddensd auk& tal lamellae, at
least partially Wgta, 106a-
b Digite without widened su tal lmdhq
B f .....................
lender thmughaut their ength (Fig@.
10&h-j31or with a sinfie & m d y enl-ed
""a
brmiml subdigital acde (Figg10Bgl 7 .........
Wlltffll
vlsw
Ba TriIatim of eubdlgital lamellae mkiW to
two ~ymmeM[rallg e d w g e d p ~l a ~h
(I& low ..........................Pk@WactyZac8
a' sIm' mb.Btus
b. CdfBonYX b Dilaeon of subdigital lamellae extend8
EPa. 104.The@( d yabmkh &sideti]. Dm* throughout most of digit (Figs. IOla-ctl .....B
If9 Claw in contat with or only sWtlsrbepnd
widend eubdigital lamsllae IRg, lob3
......................
-.-.,..*.............T & ~ @ Z U
b Claw mu& b e y 4 dbtd of vPi&ned
g-
suBdigital Iamellae (Fig&1 6b-e) .............4
4a AU subdigla1 l m ~ l l a eundi~derd(Fig.
105~) *...,....................
,..I...I .-..Ikriatdliger
b At: Ilea& the di~rtrrleobdigiEal lamella8 divi-
ded m arranged in paira (FW. 106b,dp) 12 .,
Ba Iihdian aubc81udals nut enlarged, not
aligned in a median aeries ..............
...........................................**ryh
b Median auhcrtudals enlarged and
aligned in a media wrim ...,.,.....,.,......* . 8
Ba DomJ soales hebrogenaua with large
tubmdw ..............................
RenoW@Jm
b Dm& scales hamagenelruar, no large
tubercle ........
,........*......... ...<..*.....#Aym
Ta A eingle extmmly enlarged spherical ter- I
d a l subdigital scale present (Fig. P06g)
.......................,..,.,.,. S&mmhe&1&r
b Terminal subdigital ecde not larger than
other subdigitd lamellaa .............. 8 ,,..........
Ila Two supralabtak to a pint below anterio~
margin of eye (Ffg, 1MaR head of adult
malea eolored more ar legs Bimilar to body
.........................................
LepWbkphad*
b ! k m eupralabials to a poiat below anted-
pr margin of eye (Fig. 1Of3b);head of adult
malea orange colored, contrasting with
grayid3 body ................,.............
Qo-8

Fig. 107. ArisdeIEigw gmrgeeneis ( M a Cozurnel,

Mexico). Photo: J. C. Lee

The genus Ariartelliger, with six epecies,

has a distribution centered mostly in the
Antilles. Only rr single species occurs in
Central America. AristelEiger georgeensis
is mostly found in the vicinity of h u m
settlement&,At night, these large g e h s
cw be observed on home walls and the
t r u n k s of palm trees, where they hunt for
f. Q. h i e c t e , They prey upon mything they can
Phv,mw'Us SPhe*rodactrlw subdue, even feeding on smaller l i e d s
( D mL SAXE1950).Like many geckus,A.
geosgeensia is also capable of producing a
vocalization that ia described as a screech
or a low chirp (DUEUMAN1965, SCHWARZ
Fig. 105. & HENDEWON 1991).The species is ovipa-
Under.. mus (LEE19961.
sides of
h. 1. 1. toerr in
L e ~ ~ ~ d s
Gonetodes Colmyx gedxos. Mbedlker georgeensis (BOCOURT 1313),
Waa. W. Mex., Kept. 1875:41;type Iocality:
St. George Idand, near Edbe City, Be&,
BVL to 115 mm. Embm coast of YucattIn
PBnineula including o f i a r e islands, also on
IsLae Pmvidencia and Sau A n a s , aea level
ta 60 m elevation.

a. Lepldoblepharis b. Gonetodes Further hading

K. S c m 1941, BAUER& RUSSELL1993a, b,
Fig.106. Lateral view of head (labiala orange). LEE1998.
The s p i e s of the genus Gehym are wide-
ly distributed in Madaga~car,Southeast
Asia, Australia, and Oceania. Like many
gecko species, Gehyra rnutiiata has been
intduoed to nUMeMU8 places ig.the tropics
worldwide, including Central America,
where the species is known fPom Chiapas,
Mexico IALVAREZ ~m TORO1983). These
crepuscul~and nocturnal lizards prey
moatly on ihs& and spiders, but also eab
reweet fruit. Geiayra rnutibta is oviparous. fig. 1i)ur.Adult male of Gonotodes albogulat-ia
and produces clutches with I or 2 eggs.
Gehym mutilato (W~Q- 18861, Acta
A d . Caee. b o p . C a d . Nat, Cur. 17: 238;
"Mdlaa[Luzon, PhjJigpineal.SVL ;to 64mm.
Southeasthia, - 0 and India; dispemd
in the Tropics worldwide.
Further Reading
ALVAREZDEL TORO1988,JOHNSON 1990.
Fig. X08. Juvenile of &hym nutiksta
('Ihpachula,Mexi@). Photo: T.BiUe
(Bartola, Rlo San Juan, Nicaragual.
The speciea of the genus f3onato&s are
diurnal, attractive colared geckos that
hare their main distribution in tropical
South America. Of the more or less 20 spe-
cies, only one (G albogula'aris) occurs in
Central America. Claims that G.annularis
reaches southern Central America (VILLA
s t al. 1988,A m 1994) am erroneous and
are probably based on PETERS gt DO NO SO^
BARROS(1970),&matodes annularis is
restricted to the northern Amazon Basin
(AVILA-PIREG 1995, HOWM~ED pew. comm.
1999). Unlike most other gecko species,
the representatives of the genus
Ganat&a have round pupils and do not
have braadened adhesive lamellae
beneath fingere and toes. Jn Gomtodes,
there is a &mag sexual didichromatism,
with the male exhibiting a strikingly
colorful pattern, whereas the more or less
grayish-brown females are less con-
~picuous.Gonextodes aEbogulsr6s can be
found in many different habitah, including
human settlements. These graceful gecko~
are highly thermophiiic and, partidarly
in the morning hours, can be observed
basking in the em on tree trunks, walls,
and rocks. The females lay single eggs
Fig. 110. Hernidectylylllsgmadi [Florida, USA).
Photo: K Krysko

several times per year, Thejuveniles hatch

after an incubation period of two to three
months ( K O ~ 1997).
R

r f o m t d ~ralbogulwis (Duwlm~% B ~ R O L .
18381, Erp. a n . 8: 415; type locality:
bktbique. SVL to 40 mm. C e n t 4 America
and northern hutb America, sea level to
1000 m elevation. The sub!apcies G.a. fumua
~~

e
ELummu1865)oomus in brim-

Further Heading
TAYLOR
1956;F r r c ~1973a, b.

Five s&ee of Old World house geclros

(genus Henaidactylus) haye been intro-
duced to numerous localities in Central Pig. 111.Hemi$actylus mubouia,
America (K~HLER %)Old, FRANKLIN 200Q,
MCCRANIE& WILSON2000, GF~ENBAUM
2002). Thew geckm are usually aseociabd
with human habitation.At night, they can
be observed in houses, where they prey
upon arthropods. They are capable of
vocdization and their call sounds like
"gec-gec-gec."The clutches coneist of two elwatioa,
egp each, with the young hatching after
an incubation period of about two months. 1898,Erp. Gh. §M;tspe lo& :Java and
Photo: R. D.Baxtlett
HemMoctytue b m k i i GMY 1845, Cat.
LizardsBrit. MuB.: 168;type- l d t g : Borneo
flnd AwWa, SVL to 67.6 mm. Zntmduned ta
Panama and H o n d m , mea lwel to 1000 m
H e r n i d w ~ Ifrenabua
~ D&m % BEBEOM
LYmor, 8VL to 80 mm. Intmdu f
b wed
ocalitiw in Central America, Bea level to
1250 m el~lfttitm.
Flemidoebyiw gwnotii Dtn&m 8t BUBBON
.
11830,E QBn. 8: 888, type locality: W t i ,
~ r e n c hesia. x ~SVL ta 66 mm, Introduced
to Coda Rim, sea level to 1160 m elevation.
Hemi&&ltls &ia (Mo-u DE Jo-B
1818), Bull. 80c. Philam. M e 1818: B8;tpPe
l d t s l ; Antilles. SVL to 76 mm. htmduced
bo Panama and Hondurae, ma level to 1000 m
eleYabion.
Hemidactylus m i o w (Laramus 17581,
l@gstemaNakuas, d,10: 20% type loealty:
,Weatem. SVL tn 6Q nun. Intduced ka
Panama and M d w CYucgt8n Fedxwda) I
 1 a Tail with demtirmlate fringe; twa or t h ~ ~
p a h of enlarged chin shields, paateriox
Ma) not in contact with infPalabW
(Fig.1lk) ..........
HmtPlacSyClcrsgnmwMt
b T d witbout dentidate fringe, although
widely spaced ventrolaterd spines can be
psesent; two paifa of enlarged chin shields,
both pairs in contact with infralabials
(Fig. 114a-b,d-e) ............................. .......
2
a a marg, mbdigital 1-llae of 4~ a. Hm?Edw&l'us meba~ia b, Hemidacfyu8 Frerlel~5
not reach b e of toe (Fig, 113a) ..........
........................
IIemidrPc@tbeanubouia RE.113.~ n d e m i d ofdth e toe 0amdlae shaded).
b Enlarged subdigital lamellae of h
reach base of toe (Fig' 113b) ..,................ 8 K-3
3 a No tubemlea on upper surface of bind
l h b and above ear opening
......................
.,,Bemi&ctjriw p.eloatus
...............
b Mltinct tubernlea on upper surface of hind
limb and above ear opening ...........,.......4
'
I'
4 a Each tubercle on dor~uglamunded by
10-12 mall scalaa; male^ with 21-32 pre-
anofemoral p e s H m i d i m t y l ~b d S
b Each fuhrele an don- surrounded by
4 16-20 ~ m d ~ulea; l
.....
mdm with 3-10 pb
anofemoral pores ....................................
..................., HemldQE&4w turctew
Further Reading Fig. 114. Chin region
McCoy 1970,IVLQRCELLINI 1971,SALVADOR
1981, in HemiaIactylw.
POWEU8G MAXEY 1990,AUTH 1994,LEE 1996,
POWELLet al. 1998. Anterior chin slhielde
orange,po~terioranea
brown,

Fig, 112.Hemidactylus f r e t a a t ~(David, Fig.115.Hemidactylus b m k i i [Cbrdoba,

Chiriquf, Fanama). Photo: M. Lundberg Colombia). Photo: M. Lundberg
Fig, 116,~emidactyE~8
trarcicm Fig. 117.k p i d o b h p h n k xanthoetigma
Photo: k NUllert (Nwagandi, Panama),

Lepi$obtephare'a
The species of the genus tspidoblepharis
are small ground dwellere that live in the
leaf litter on the rain forest floor, mostly in
close vicinity ta mtten tree trunka, large
mats or decaying plant material that pro-
vide suflicient humidity. The females pm-
duce clutchee considing d single eggs,
which they lay in organic debris under
roots or loose bark several times per year.
Both Central American speciee of Fig 118.Leptdoblepharis saractmmartae
Lepidoblepharirs appear to have an exten- (Panama). Photo: M.A. Guerra
sive reproductive season,
Further Reading
BARBOUR1923,Durn 1 9 4 0 a , T ~ m ~ RWAs
1956,
FUENUAYORet al. 2002.

Key to Lepidublepkarde
1 a Dorsal wales distinctly imbricate ..,.,
..........Lepidoblepltarise a m t a e w
b Dorsal scalea juxtaposed ....................
...........Lepidublepkrb x r u r t h o 8 W
 Leaf-hd geckos are distinguished from
dl other Central American geckonid p-
era by a pair of enlarged leaf-like terminal
lamallae on the finere and tms, whereas
the more proximal subdigital lamelbe are
not widened. Leaf-toed g&os are mepus-
cular and nocturnal inhabitants of old
trew with loose bark and can deo be
found on house d s . During the daytime
in ita natural environmeat, these geckos
like to hide in tree crevices and beneath
Fig. 119.Lepiclordactylu8l w b r i s 031Vakle de loose bark, while in human habitsrtim
- M n ,Panama). they will make uee of any dark refuge that
provides gome Bind of protection. The call
of thig speciea can he d e d b e d as a aeries
of kias-like s o n d e (ALVAREZDEL TORO
1983). Duringthe c o m e afthe year1%VET-
al c1utchea cansiating of twa eggs each are
glued to the surface in crevkw or other
Lepidddactylw Iugubd~is an animal that prahted sites. In cases where several
has apread with the advanw of ddization females use the eame egg laying dte, mars8
and been intmdused worldwide, The fret aocumulations of egge and ewhells can
pubIished record of this epwies being be found ( ~ H L E Rupubl. obwm. on Utila,
established in Centrd America a p p e d Hondurae).
in the early 1960s (H.SMITHL GIGAMT
1961). Theee authors collected a subadult
female in Paaama. Meanwhile, L.lagubris
is known fram additional localities in
Central America, including places in Coata
Rica, as we11 as the Corn Islagcis off the
Caribbean mast of Nicaragua ( V Uet al.
1988, K0m.x~199%). The specim repro-
duces parthenogenetically, with the fe-
males producing clutches oomisting of one
or two eggs that batah after 60 to 100
days, depending on incubation tempera-
ture I ~ 1997).B

Em Zugubris (Du~&IL& BBgKp

BN,8: 304; type lwalit~' M
[= Tahiti, accordisg to F%mm L Do~oso-
BAE~ROB19701 8VL to 42 mm. Southea~tAsian
and Indoau~kalianregion; &per& world-
wide, sea level to 700 rn elevation.

Further Reading
H.S m L% G w 1961,HENDERSON
et al. 1976,
mu 2002,
74
Fig. 120,Phyllodcactyl~urinsularis (Half Moon Fig. 124. Phyllodac~ustsahrculosua (Isla
Cay, Bebe). Photo: J. C. Lee Ornetape, Nicaragua).

1 a Dorsum with 14 or fewer longitudinal rows

of enlarged tubercle-like scales ...........
................Phyllodactylue tubem~Eo9us
b Dorsum with 16 or more longitudinal rows
of enlarged tubercle-like scales ....,..........B
$2 a 48-60 ventral scales from throat to vent;
39-47 tubercle-&.e scales in a parmerte-
bral raw from occiput to base of tail .......
.........................
P l c y E ~ t y L uplrrre-trs
s
b 60-66 ventral scales h m throat to vent;
46-52 tubercle-like scales in a paraverte-
bral row from occiput to base of tail ............
.......................
PhyZld&cbyLu# itnrvlarz~

Further Reading
DEON1960,1964.

Fig, 1'22,Egga d egg shells of Phyll&bgrEw

pdmeus (Isla de Utila Hondnrm).
 Sphmdactylus
With some 160 recognized epecies, t h e
least geckos (Sphaerodactylus),with their
center of diversity and distribution in the
Caribbean (about 70 ~pecies),form the
largest of all geckonid genera. Nine spe-
cies of least geckos occur in Central
America, where they are restricted to low-
land regions (sea level to about 1000 m
elevation),These are very small lizarda (in
fact the smallest amniote is a Sphaem-
dactylus; HEDGES & WOW 2001).Most
Central American Sphaerodactylus have a '
-
maximum SVL of 25 5 30 rnm, except for
*_
the ielmd endemics S. msaurm and 8. Rg.126.Sphaerodactylus m~uurae(Ids de
pacificus, which reach 40 mm. Least Roath*?30ndurw).
geckoe are typical leaf litter inhabitants;
they dart through the leaves, skillfully

i-
hunting small arthropode, euch as spring-
tails, cricket larvae and little spiders. The
best way to spot them emall g e c b ia
when s h d i n g slowly through the leaf
litter carefully watching the ground. These
geckaa especially l i i ta be in the proximity
of the roots of large trees. The population
density can be very high at certain sites,
reaching two or three individuals per
square meter. On the Islas de la B a a ,
Honduras, I found 5.rnillepunctatus and
S. rosaurae syntopically, whereas the
former epecie~ocsurs together with X
dunni at Pica Bonito on the Caribbean
clutches
coast of Honduras, of single
consisting Least geckos
egg5 that
produce . S p h a e d c t y l u s mil2epumtatue
lQ126,

hatch &T 60 to 90 days. (Matamoms at Rlo Cuyamel, Hondurnal.

Fig. 127. Sphaemductylve dunni (National

Park Pico Bonito, Atlmtida, Honduras).
 --

'Fig, 129. S p h a d c t y l u s hornolepis-

(Tortuguero,Cmta &caZ
 Fig. 132.SphaemdacEyiw a r m WanddIe,
Man-, Jamaica)* Photo: 3.C. Lee

Key to SphaerwlacQlus 4 e One rupratlaeal scale (Fig. 131al ...........

5
1 a D o m m with a median band of amall b Two supranasal scale8 (Fig. 131b)...........7
granular scales beween large: imbricate a Mediansubeaudds and
dorsal scales (Fig.134a) ........................... in a median series (fig. 133);dorswn with-
...................Splzaerodactylua mtaunae out dark mas-ban& ..................................
b All dorsal acsilels about the same aize, ...........,B p h a e W Q Z u s gmptolaemtae
without
scalea (Fig, a 134b)................................,..
of small 2 b Median subeaudals not enlarged, not
2 a Dorsvm of adults with distinct large dark
- -
aliwed in a median aeries:. iuveniles with
dark cross-bands on dorsum ....................0
blotcheh or mottbng; S n 47 mi B a norsaland rrcdes keeled;
endemic on Isla de Coca, Costa 'Riea .......... stlbcaudals arranged altewakely; 48-67
....................Sphtcanodactylsrr pacifiue dorsale between lerela of axilla and groin
b Dorsal pattern not ae d e b e d above;
SVL to 33 mm .........................................8
................Sphae&c@ltue hm@b
S a Enlarged spine-like superciliary situated
posterior b level of rnideye (Fig. 131a);
snout conspicuously convex in lateral view
........................
Sphaer~dacSylwadunni
b Enlarged spine-like superciliary situated
anterior to level of rnideye (Fig. 131b);
snout flat, in lateral view running in a
atrru'ghf line from eye to tip of snout ......4

Fig. 131. Scalatiun of the enout region (supra-

nasals shaded); note also the different position
of the apine-like superciliary (arrow). Ahb. 134. Dorsal scales in Sphaemdmtylus.
 b Doreal mdee weekly keeled,
rcaleer g"""
uaually smooth; ru candala
arrangrrd in eequeneee of one mall d e ,
one large BE&, and a palr of mall wales;
62-81dmala betwean Iwele of milla and
groin ......... S p h m & l u s lirawkrbtcs
7 a Dorsal scales smooth ................................
....................... S p h w M Q E w glamw
b ..
Dorsal sides keeled ..................,,,....., 8
8 a 38-50 dorsals between Iw& of axilla m d
groin; domum with longitudinal rows of
ooellqted blotches .......................A,...... ,.,.,.
..........................Sphrsemdm@Zrar t w m s
b 47-74 domala between lavela of rixilla and
gmia; doma1 patterm variable$but without
longitudinal rowe of oeellated blokhes Fig, 138 Tkr&ru!tytul~mpicauda t W a ,
........ S p h m d a o t y l w m i t k p a c ~ # ~ t u .Rfo $an Juan, Nicaragua)

~ e r ~ g
HARars & KLU~E1984 ($phm&tyEus of
Central America), Ktl- 30010.
Thecadmtylue
- -
,nocturnal,
~ c a d a e t y l u s,,picauda is
arboreal inhabitant of the tropical rain
forest. At night, these large geckos can be
observed on the tnxnks of large treea some
3 to 6 rn above the ground, while during
the daytime they retreat into bark crevices
and hollow branches. Other typical day-
time hiding places include piles of leaf
litter among the buttress mots of large
trees. Turnip-tailed geckos are also corn-
rnonly encountered on house walls. They
feed on invertebrates (mostly crickets,
moths, grasshoppers, and spiderd and
Fig. 135. ThcadactyEus rapicauda (Quintma small lizards. Like mest geckos, T, rapi-
Roo, Mexico). Photo: H.Bahena B. cauda is capable of vocalization and its
call sounds like "chack-chack-chack"
(BEEBE1944). During the reproductive
season, female T rapicauda lay several
clutches each comisting of a single egg.

.TlwcdxdyIus rc~ptcrsrrda( H o r n 17821,

wet. vlbhgwl8:
1s.""hT
vdL
'l$t&m
cab Wq. ByL
~ ~ l me*^
401
v a ~ +a
tmoi~alSouth Amedm also on the h e r

Further Readfng
TAYLOR1986, AV~LA-PIRES
1995, RUSSEL &
P R~UER
2002.
l?m~ac~ (1483)suggested separating the
w-called "microteiMsminto a a p m t s
family (Gprmophthdmidae) fram the eo-
b Frontopatietah pmmnt, frontal,therefare,
&eparat.e&om p d t a l s (Fi& 140bl;auter
ear opening pregeat; exhmitiea normal g
I
&led "mamteii&" (Teiidae). Thih, view
has been accepted by the rn@orityof her- 3 a the
PDBteFior edgea of the parietal ecale~and
interparietal form a straight line (Fig.
petelogists workhg in Central America 141a); upper surface of tongue cavered
(ILWEZ et al. 1996,SAVAGE 2002, WESON wit6 plicsls .......................
P&cksgbsr*s
pem. comm., McCRANIE pers. comm.1, b Middle parietal smIe clearIy protrudes
whereas others p r e M eot tg recognize pas& the posterior edges of the arietal
the Gyrrmophthhidae as a W n c t faglily s d w (Fig.141b); Upper s d c e o t o d p
covered with papiIlae ..........................4
F
(HARRIS1986, MYERS& DQNNELLY 1996).
The lizards of the family Gymnoph- 4 a Dorsal scalation heterogeneoua (large
thaIzgidae, including about 30 genera with tubercular a d m sprinkled fn with the
h e r ground acalation] ,..........................6
approximately 169 species, are distnluted
in temperate, subtropical, and tropical
areas of s~uthern Mexico, Central B a Prefrontds and frontal frqrnmted iato
America, and South America. Limb reduc- &mallirregular scales (Fig. 14%); a clearly
enlarged scale beheen fir& f h p r and
tion hae occurred several times within faiat ................................Nerretiaww
Gynmophthalmidae, and in m y of t b e b A.eTrontals and h t a l form large shields
it is amompanied by body elongation. AD (Fig, 192al;no clearIy edargsd sade be
gymm~ph'thdmidsare avipmus, u s d y $ween first finger and wrist .....................
producing two eggs per dutch. .....,..,.................................
E e h t ~
6 a Doseal scales amoeth ..................M i a 1
Xey to genera of 7 a A double longitudinal row of widened
Gymnophthalddae gular scaler; a vertebral pale atripe mu-
1 a Without ffontoparietale,frontal, therefore, ally present ...................
MRO&@&IW
in contact with parietds (Fig. 140a); n o b No double longitudinal row d widened
outer ear opening present; extmmitiea g u h d e r ; no vertebral pale stripe ......
greatly reduced ...........................
Bachia .............................................
..u.. hposonaa

Rical.
It
~
I
Fig, 137, Neu.$tkurusapo&rnw CPalmar, Costa Rg,138,L e p ~ m muthi
Photo: R. W. ,Van Dwend~r Panama). ,
a (Nuagandi,
 y "-.I I
frontal
supraffiulaw
parleiaka superctlladeg

Eig. 140.
mmnm of
head (fiontal
brown;
parietals
or*).

head (interpet-
r i d brown; Fia. 143.Anadia ocelEata (Rara Avie, Coeta
Photo: T.benders

In Central America, two species of the

Fig,142. g e n u Anediu, A oceliata and A, vittatu,
Dorsum of are found (OETEDAL19745.Both are very
had(pf~fron* alender diurnal tree dwellers with long
*brown; prehensile tails. The habitat of these
hntala lizards ia the rain forest where they prefer
orange).
treea with a plentiful supply of bmmeP-
iads. Large brorneliads aho serve these
b. Echinasaum b, N ~ U S ~ ~ C U M
animals as hiding places. Their diet can-
gists primarily of insects aed spiders.
 slender p m d dwellers,Two species dthe
predominantly South American genus
reach the ~outhernportion of Central
Gmerica (Darohl 1973, MCD~ARMED &%
DEW!&= 1977). Bacliiar blairi wm conaid-
e d by DEQN ('1975) to be synonymous
with B. palZMiceps, a decfsion for which
there was no evidence. MCDMMZDL
DEWEESE(1977) were able to show that
Fig. 144.Anadiu ' a I 1 ~ t u(ProYhcia Limb, Bachia &laird was a separate species,
Coeta Rim), Photo: R,W.Van Dcavender
which differ8 from B. pallidiceps in
coloration, build, and several scale
& t d a oceliata GRAY
fW6,Cat. L h d s Brit.
charadem. Rppreaentatives of this gmw

"tT
to 36 m r ~ ~ o~ta
.
1 ~ ~ ' ~ . o p i i d - ~ ,
and w ~ t e panama;
500-1310 m elevation in pnammbm rain
~ p
are characterized by the absence of an
external em openigg and by the tendency
to reduced limb &he,On the front limbs, B.
fmt. blat'ri aad B. pallidiceps have f w r tcms+
M a uittota B O ~ C ~ 1913,E R PmcProc h l . whereas the rear limbs have only + b e e .
90c, Loadon 1918: 1033; tJlpe W i t y : hfla They are highly secretive ground dwellers
L.iaa, 300 f i g ChocQ Colombh in the forest. %se I i ~ d as~al.
to
Calornbia.,
67 0-190 rn elevation in lowland
to muthem
rain ly found in looaa dirt a under beet3 or l a p
h& ( D m1940a).
Key to & d u A#%&* sat
1 a Pale dorsolateral strr:pee maximum of two 6ci. Philadelphia @% 115; type locality:
scdea wide; obvious ooelli on body sides; herto Armuellee, Chiriquf, Paaama. sVL to
27-32 d a s at midbody; males with a total 70 mm. butheastern Costa RiEa and adja.
of 8 1 1 femoral pores .....AnQdZatlcellota cent areas in Panama, sea level to 40 rn ele-
vation.
b Pale dorsolataral stripes at lsaet three
males wide;ocelli, if preaent, not, obviow; Bachia pallidiceps {COPE1862e),PBX.Atad.
30-37 scales at midbody;males with a total Nat;. Sci. Philadelphia 1Q 366,type l d t s f :
.
of 11-14femoral pores .. ..A n d l a vittafa Rlo aVanda mpim, Colombk. SVL to 79
mm. Eaetern Panama to northwestern
Further Reading C&n&& 888 1 4 49 1QQm dwatioa.
OFTIDU 1874.
Bq to &u!hicc Although usually found in the vicinity of
1 a Pale dom1atera.l atdpea prmnt; 22.24 st-reams, these lizards live samewhat mom
~ midbody; 46-48 m e rows of dor- terrestrially than Chs semiaquatic Newti-
s c a l at
sal scales; 39-33 cross r o w of ventral curus CUZZ~ELL1966a, b). According to
males .....................,,Baohia p d k t d c e p DUNN(1944), BREDER(19461, and U z z ~ u
b Without pde dorsdateral stripas; 2426 CU65a),active Echinomura can be obisenred
wdee at midbody; 43-48crorsn rwa of dor- during tha day ae well a ~ at + night; how-
sal d e e ; 29-31 m@row5 .s of ventral
acalee ...............................
Baehfa bWri ever, they appear to be predomhntry noc-
turnal tlTzau. 1966a). The animals me
a d v e among the leave- of the gmund
cover or hide under mcks and in rotting
plant material, $everd raporta etate that
Eehirtogaurck will become absolutely rigid
when handed I U m u 1466~LEMTOM ig;
ANDERSON 1966).The m h d s will sWen
their bodies m d press their limbs- againat
their sides. If they are turned on their
backs they will quickly turn over and
freeze again. Aa with most micmteiid~,
Echinosaum prduces clutches, each wikh
two soR-shelled eggs ( U Z ~ L1986a).
L

Fig. 146.Bachia palldiceps (southeastern

Costa Rica). Photo: R W.Van Devender

-
The lizards of the genus Echinosaura are
small terrestrial "microteiidsnthat are dis-
tributed from western Panama across
northwestern Colombia to central-western
Ecuador. In his revision of the genus,
U ~ L (1965a)
L recognized a single species
(E, horrih) divided into three subspecies.
Recently, FRlm et al, (2002) described a
new species of Echinosaura and elevated

I
the former subspecies to species.Thus, as
currently understood, the genus Echino-
saura contains four species, two of which
(E. palrneri and E, panamensis) occur in
southern CentraI America,
Fig. 146.Echinosaum panamensis (ElCog,
Panama). Photo: R.W.Van Devender
Fig, 147.Echinoscaura panrameraark (El Co*, Fig, 149. Gymnophthdnue apeciosua (near
Panama). Photo: R. W. Van Devender Guanagasapa, Escuintla, Guatemala),

Key to Ecfciraoeawrct vestigations indicate that such a dividon ier

1 PFrontal plate transversely divided (Rg. not justified (KClrntE~19Ef6c).The usually
1 4 W ............,Echiawaum pamameneie intensely red tail is particularly conspic-
b Frontal plate single (Fig. 148b).................. uous, and gives this small lizard the local
. .. .
...,........ ........, . EchSnoaaara plnped name "Cola Rqia" ("red tall"). These are
typical undergrowth dwellers that spend
Further Reading their days searching for prey (mostly inver-
D m 1944, UZZEL~ 19658, FJWITS
et d.2002. tebrates) under the wilted leaves. The
population deneiky ie apparently seasonal,
with an increase during the dry season
(TELFORD 1971).According to the investiga-
tions of TEWRD (1971) the female of G.
Gymnophthalmus speciosus is the ole epeeiosus in Panama produces clutches of
representative of the micmteiid~to extend 1-2,sometimes as many aa 3 or 4 eggs. The
as far north aa Mexico. Whereas this ape- egg-laying seaeon, during which the fe-
cies has, a t timea, been divided into a num- males produce up ta three clutches, lasts
ber of subspecies (MEIUTENEI 1962c,PETERS,from October to March, with hatchlings
& Do~oso-Bmos19701, more recent in- being ob~erved in the months from

Fig, 148, M a 1 head d a b in E c h i ~ u m c

mntd omgel.
 Like most of the rnicroteiids, the genus
Leposoma is also most heavily distributed
in South America. Only Leposoma rugiceps
and I,. gouthi mur in southern Central
America, The malee of L. mubhi generally
display an intsnaive orange coloration on
the underside of the body. Similar to spe-
cies of the genera Ptychoglossus and Gym-
nophtibalrnus, the Lepoaorna species are
aIso predominantly u n d e r g ~ w t hdwellem
that are found between the wilting leaves
on the rain f o r e ~ tfloor, The following
details on the reproduction of Lepssoma
rugimps are based on the inveatigations of
TELFORD(1971) in Panama. During the
February to May (TELFORD1971, ALVARFX egg-laying Beason, which lasts for several
DEL TORO1983).The young measure 17-19 months (May to November) during the
mm at hatching. rainy season, the females produce clutche~
of two eggs each. The young hatch with a
SVL of 16-19mm. They are sexually ma-
ture within six to eight months.

Fnrther Reading
COLEet al. 1990 (evolution and syatematicg of
the genus); TELPORD 1971 (ecolagy, =reprodue-
tion).

I
(Panama).
Fig. 161. Lepl,soma r u g b p ~
Photo: M, A Guerra
Gymnophthalmfdae

Colombia). Photo: M Lundberg

I Fig. 184. Lepomca southi (mutheastern Coata
Rica). Photo: R. W.Van Devender

Key to Leposonsa
1 a hnntomal divided ........Lepmmu soutAi
b hvrntonasd undivided &pmmm mgkpa

Further Reading
R m IN2, U z m ~ t BARRY
1971 (taaronorny),
TELFORD1971 (ecology).

Abb. 153. Leposoma rugiceps (Pearl bland&,

Panama). Fob: C. W.Myem

Fig. 165, D d head rcalatim in Leposoma

(frontonasaleorangel.
 E J h ~ : u r wa-mu8 U ~ I-, L B d .
Amer. MUB.Mat. Hist, 138: 298; type l d t s ;
16 km BW San Edro del General, 866 m elei
vation, isan JoeB, Coata Ha,SVZ to 41 mm.
sou^ portion of the Pubtrvenw Provinw,
Cats Rlca, new sea level to 900 m elwatim.

Fbrkher Reading
U- IS65b.

uruy one epscies of this predominantly

South American genus PrionadactyZus
Fig, 158.Newticurw apdnzua (Fdmar, Costa reaches the border m a between Panama
Rlca). Photo: R. W*Van Dwender and Colombia in the extreme southern
portion of Central b e r i m . Them agile
Liz-& are ground dwellem in the rain
fore& where they s d for prey (mostly
insects and widem) during the das To
mpmduw, the female P. vertebral& pm-
duw~ dutches o f two eejy each, which me
Water tegus of ths genm Neusticunsg have only mpeffidally buried (UZZEU1973).
their primary distribution in South
America, with only one species,
Neusticurus a p d e m w , reaching muthem Priotcoda~tylua
18791, b.
venkbralis ( ~ ' ~ w G w B ~
Mag. Nat. Hiat. (6) 4: 298, type
Central America in Costa R i a AB with all locality: Intac, Ecuador; SVL ta 68 mm.
species of this genus, N. apodemtts is Ezrtreme eastern Panama (horde*
semiaauatic. found alone f a t flowing Colombia) to northweetern Peru,700-2500 m ,
streamk in the rain forest ueually wa& 1 -
tegus are found on rocb OF roots by the
edge d the atman, and they flee into the
water at the slight& &turbmceC8Some F u e : 9 k &
active X apodemtls can also be found in
leaf litter on the forest floor (UZZELL
196Sb). Although representatives of the
genus Neus#icurm are active primarily
during the day, A? apodemtcs haa been
observed swimming in the water at night,
as well a&during the day ( U ~ L1Q6frb).
L
While swimming, N e d c i w w will fold in
their legs alonpide the body and hold only
the head above the water's surface while
making malee-like mwemente with the +
body and tail. Nothing is known abolrt
their reproduction, but most TikeIy N.
-
tzpm&tnaus, as with ather Naustidurus spr+
ciw, pmdwa taFo egm per clutch, which F
are deposited by the females mtting mg, 157.p r i o n h t y d u g oedebmlh ( ~ e m o
logs ( U m 1965b). Mdi, Darih, Panama). Photo: C.W.Myers
Fig. 168.Ptychoglossua festae (Nwagandi, Fig. 159.Ptychoglosau~pdicatug in Costa Rica,
Panama). Photo: R.W.Van Devender

The primary distribution of the genus

Ptychoglossus is in northwestern South
America, with only a few species reaching
Central America. Whereas l? ~ l i c a t u a
reaches at least EEI far as nartheA Costa
R i a , P. fa~tueand l? myersi are mstricted
in their distribution to eastern Panama,
Ptychoglossus festae i~ most easily distin-
guished externally by the presence of small
lateral scalea between the dorsal and ven-
tral scales. However, the only definitive dif-
ference between the other two srpecie~can
only be seen in the hemipenial morphology
(HARRIS1994). The speciee of the genus
Ptyych~lo$susare small diurnal under-
growth dwellers. Thus,they occupy an em-
logical niche similar to that of members of
the genera Lep~sornaand Qynanophtfaalmw.

l&ey to Ptycht3gbiT*tcs
1 .a A longitdid fold ~eparrttaed d ecdm
&om ventral scdes <Figr,lala) .......
............................P & ? fhtw
,,.,.
b Witbat a IongitnW fold betwem damd
m a b and vmkd s d a (F'ig. 161b3 .,..,., 1
B a Hemipenia with 23-28 rhq E M S ..........
..............*.........-*-pa-
b E d m with 9-1%ring f ~ l b................
............................Bych#krm#rqytPm2
Of the tm surviving @es of the genw
Bekaderm, only one (2% h~rriduuntS is
found in Central America,These me mbmt
lizard8 thet can reach a total bxlgth of over
70 cm, d t h mm~ivehead8 and short,
.powerful limbs. They have fat tail%w&&
are nat capable of mtattimy ar reganera-
tion. The bactr is c o t r d with h g a mEed
knobby scales The bmgne iil flw,but
Fiked at the tip. The H e I o d k m & p w b s
are the only 8tWiving v s m m l i d .
Elorgated, multi-laM poison gEm& are
located along both aidm of the lower jaw, A
aeparab opening from each labe of the
g l a d lea& to of ths grow&
~tb,w?dchmup2io8mm.inl~.
When the teeth elmp dam, the venom is
out ofthe g l h by the p m m
d the jaw mwlea and flows through the
grotwed teeth into the wound primarily by
capillary action. B e c a w Gila rnnnetem
are shy and live veq saeretitr&lyIt h q pma
no *a.t to humam. Biting idcidenter
o~cwonly when the animals ate prwoked
ar captufied, Thew oth~rwiaecalm lizards
can laumh them_%elveswith unexpected
apeed and biter m y ~uddenlp.A M a mom-
abr d l not readily deasle anything that
amem between its powerful jam, and it
will bite down fmiblv. Gila rnva~hrW k a
ahould be t k m .ye& ssriowly and can
l~adto Iocalised se well as syastemic m
tow of envenomation, whi& abwfute y
require medical &ention (EdEBs 1992).
P-

;
L
!

I
rn@
a FSEhwm
Abb. 161.Latsral bod$ edeai,
b- @'*odQw% @bet@
In Mexico and Dentrd America, three sub-
specie8 of Hehdwma homidm ape recog-
nizd. R. h. Worridtbna WISGW 1829;)
dmg the Pacific aide &M&o ftam &on&-
ern Sonora t a Chiapae, H: R. alvarezi
B o r n & MAirrlx DEL CAMW 1866 ia the
Rfo: aTijdva Bmin, CMapaa, and £
c h r L e s m GAMPBEU $ T J m
A,I,
lWB in
the Rfo Motagua Badn, GuakrnaIa. Gila
rnaastsrs ilre ~pwmtlwand no&mmd
ground-dwellera in dry f m t that bgcame
activejust aRef dwk. On cloudy days, they
can sometimes be abrervd during the day
I outside af their hiding place8 C b m w q
(El&end, Zacapa, Guatemala). (Mixtequilla, Oaxaca, Mexico).

which they have either excavated them-

selves or were pre-existing). They hunt
down their prey, which consists of neetling I
t
rodents and birds, aB well as bird and rei-
tile eggs, with comtant tongue flicking.
Gila monsters can survive for several
month without food by living off the fat
stored in their tails. During the rainy sea-
son, these animals are more active than
during the d r i ~ rmonths of the year.
Females will produce one clutch per year
(during the months from October to
December),consiahing of 4-15 elongated,
thick-shelld ems,which hat& &r four
to five months (ALVAWUDFL TORO1860,
C
Fig. 164. Hel&ma horridurn oluarezi (Tuxtla
GutiBrrez, Chiapas, Mexico).
F r r c ~1970).Gila monsters are long-lived
animals, reaching an age of over 20 years.

Further Reading
BOGERT& MARThY DEL CAMPO 1956 (mmpre-
henaive wurk on the biology of this genmj.
 b SuhdigM l a r ~ ~without l l ~ knobs (Fig,
1Ebk serCim1fin or h h t on bead B .......
a a T4e9ofhin.dibatwithfringeofflatecalea
Lrniing ~ e m t mqia
s mg, 166b);latard
bdy c d e s Btrmog~neoue ........
&rrr&lcs B ~ ~ N A U~q.t~;tas
B Larntn-
b Rwa of bind fbet withonk fkhm af flat pI-8 m b s
acdw formingwimte margin; 1 x 1bosy
@ale@hekm0gem.w .........~ p Ag. 1 8 8.Lateral
~ view of
u haad.
% a Tail plpith whmk ofedmged spinow maim
(Figs,MBa, 171) ...............................,.......
1
b W1Withopt ~piaai ...................................
8
2 a A large, e l o ~ g ~ suboeuEar
te d e pmmt
Fig. 16&j ...,..........................
Cte#mtama
b Orbit ktrderd b e k by tm arc d several
aubeqnal arrales ................Mom-m
8 a Large p d u l o w dewlap prwwt. m e or
two compiouow31y enlaged rounded scales
~ ~ ~ p ~ ~ m ; r n t o 5 a ~ . . . ~
b No 1- psnduloua dewlap; no conspie- Fig. 163, a. SeeIaticm of tail in Cteaoamnr
uolll~lyenlarged wales blow tgmpanw quiagrte~arinata;
mdl to m & w ~ s Wlizards ............-... B b. Lateral b a d seales in h a s a u r a similk
Ienlawd m W a d e orange),
9 a V e q large interparietal piate p w f t t W.
Ira4 .............,........,...,..,.....,.I .........I O
b InterparEetd plate, if praseat,emdl (FigFirr
0 .. S ............................11
. ' .

1l)a A doreolated la 'tudhal fold alongside

bod pmerit, ,m3ig. with Mma ML.llal
&; I ~rn*l~b lzxmwms pular Bld,
diffemltiated fFom adjacent guk a d % s
byvery Fine d m ..................
Umatcmm
b No dmelataral longitudinal hid alongside
body; M, trruamersla @ar Eald ...,......
............................................S ~ ~ p O ~
ilr !lM abut three the^ m,no v&abral
mw of enI& see ..............J9@duw?
b Tail 1- than twice SVL,a vertebral row of a Ses- (Inter-
parietal ml$shaded)
enlarged wales present ......... Enyalioidse
Fig. 170, Domd h a d malation (interparid1
0 ~ 1 ,

Armw paints to &r fold. Fig, 171.TaiI of Mwunaeaurua annularb.

 .-
Anoles (genera Analis, ctgrconotue, relationahips of the anolm is still too
limiked to accuataly define the natural
mu~ings*
The anoles dAnoIk sensu lato), with more Most ~pecieeare bush or tree-dwellers, al-
than 300 valid species, repreaent the most though same are only found amongst the
diverse bard genus, with more gpecieg leaves on the ground. Even some semi-
being described almost every year. They aquatic species tha* occur only in the vici-
are a conapieuotz~component of the fauna nity of water, such as N D M ~bark&, S M.
wherever the^ occur,and throughout their liomntus, N.poecilapus, and A? aquaticus,
range it is common for several species of are f d in Central America. mid Ied-
moles to occur bgether. Despite the fact dwellers in rain f m t are eame 8peciear of
that anolea have hem the focw af interest the tmpidonotus group (prfiicnlarly N.
by herpakilogists for more than a century, humilie and N. uniformis],whieh in some
the phylogenetic re la ti am hip^ and twon- areas reach a bigb population density
amy of tbis group of lizards xe& sub- ( H a m 1998~)~ A more robugt represent-
ject to eerioue coatrover~yin the m n t ative of the genus ia N,capido (SVL to 96
literature. Indeed, our understanding of mm). These l i z d are typical sit-and-
the systematics of thew Lizards ia still wait predators, which are Micult to pot
poor. In the present work, I follow the clas- when sitting oh a branch or tree trunk.
sification proposed by GWER 8t SAVAQESimilarly large (to about 100 mrn SVL), ihl
(1987,1992) and SAVAGE& GUYER(1988)in the bright green colored N.bigoreatus that
using the gsneric names Anoli% Ctsno- inhabit8 the- tmpical rain fore~t,usually
mfug h t y l w , and NQTOPB for the mole on trees at the forest edge, in clearings and
species of Central America. Thia decision at the shore of ponh and lakee. The true.
reflects ihe strong support: far the assertion giants, however, amongst the Central
that Norops repwsenb a rnonophyletic American d e s are found ia the genus
lineage.Nomps is the beta section of moles Dactyloa, such m D,insignis (SVLto 160
of ETEUCFU~E (10691, whose specie8 are mm). These are trdwellera that are dif-
chmacteri~edby aubtomic caudal verb- f i d t ta spot in the rain fornet, With luck,
brae with tran~lvereeprooea8es that are they can be observed on the t& of
directed antarolateral$; see figure QA in large treets where the animals usually sit
ETHERIM~E (1957). However, the appm- with their heade pointed down. According
p h t e l l e ~ sof GWER and %VAGE'B (1987, to S m et al. (19781,t h s s animals u~ual-
1992) other proposed genera (the alpha ly remain: on tree t m d w at heights bs-
p u p of ETHERIRUE19591 appfU' more Ween 40 and 200 em above the ground,
uncertain. The anole classification of although specimens sitting up higher
GUYER 44 SAVAGE(1987) was heavily would be Micult to observe (SAVAGE &
criticized by CANNATJZLLB 8z DE QUEIRO~ Tmm 1978).The species of the ~ U H C M E U -
(1989) and by E. WILLW~SC1489). mtw gmup bg., Norops dtae,h?exaul, N.
fuscoauratu&,N.kemptoni, N Eirnifim N.
For moat specie^ groups" of moles, aa pro- pandoensis, N polykpis N, I-odrigmii,
p ~ e by d variaua researchers in the litera- and N. Z ~ U ) are slenderly built anoles,
ture, evidence that these actually repre- which live mostly in buehes or low trees#
mnt monophy1etic groupa is meager. Some where they are som&imesM c u l t to spot.
of them probably are n a k d groups of spe- They are predominantly lowland species,
dm, while others repreaent artificial usually living in border areas and clear-
groupinp of species that are supeffilcallgr ings in the rain forest, where they tend to
Bimilar and me placed together bemuse of be quite plentiful. The species of the cms-
particular character cumblnations, At this sulus group bg., N, ampfisquamosus, N.
time, our knowledge of the phylogenetic anisolepis, lK crassulus, A? htquei, N:
he&ropholidotua N. laeviventris, A? mural- -,, a,,w,, - ,,, . , , , ,,,,
la, N; ntbribarba* N: smintha, and N. E n g l d ~ 1 club . 10: 68; type l&ty:
wermuthi) and the schiedei p u p (e.g., h? Coxen Hole, Ruatari, Bay Manda, Honduras.
cobanensis, N.hubrt~mithi,N.jahnmeyeri, SVL to 87 mm. Body green, dewlap pi&.
N. pijpbmis, a d N. ~ u l p u r m ~ ~ r ) I R Guamja
~ and Roatb, C a p Cochinoe,
NOndUTaat as well ae in and on Cuba,
mostly highland specie8 (above 1000 rn el@- 1-1 300 elmatian.
vation), m u l y wikh small -I) of distri- CtenoMtus Fisbtelb (Dvm B-N
bution in sad forf%t.It be- 1887). Etp, mn.1: 143; type locdity;
lieved that the apedea of the &e&i ~p MartiniqUe[in emr~,SVL to 70 -, ~d~
originated h r n the fragmentation of what brown, dewlrtp greeni~hyellow with orange
was the ~ n & u o&&$bution
~ mar@n. Puerto Rim and Virgin Islands;
of an anwBw species due of fntrodud in Costa Rica, mea level to 750 m
elevation; oomurenee on Isb C m e l doubt-
mountain mamifa during the %diary peri- (-
od (McC- et al. 1B93b).
Dmtylaa W O B E ~I~ANEZ N A ,& DE
Sousa 1992,Rev. Biol, Trop. 39: 268-262; type
All m l e species mp~duceby ems, l d t y : Mdrgenes de Quebr& ha&, 1100
and, during the breeding season, will pro- m, 8*44Yt 82"13W, Resema Forestal de
duce mveral elutchee, each condeting of a Fortuna, CMqui, Panama. SVL ta 108 mm.
8.6 ob ' ue bm bands
a w e egg,at intefvds of one to two w e e k
% ovan. and rm @ideof the
B&
(0' %
formed o f a d l a ~Ut@ '1 in-pned
blue-outlined yelAow patchee; wme female8
body are altematgly active. ib soon a ~ one
, b ~ a ebroad, dark b m mid-dmal stripe
egg ie laid fmm the right &duct, for extending h r n nape to bme of tail; dewlap
example, el new foUic1.e is prcrduced by hha M y cream with broad yellow scale row5
lee ovary passea through the lefi im@=ll, i n m p e r ~ dwith ~~~
rald green d e a . Cordillera Central in the
dud, it ie fertilized and 1he1led p,~,,, ckriquiand Boeae del n m ,
within a few days, More beiflg ldd. In the Panama, 1080-1400 m elevation.
meantime* *t OVug pmdu* a Doctyh ehlon. ( B m a r u w 1898), Roc.
new foUicle, which L w d y to ovulate. h zool. sot, h n h1898: 110; t ~ r p elocality:
long as th~a is enough food avdable and Parwba, Ewdm. SVL tp g0 mm. Body
the wmther conditions are ~ u i ~ l ethis p n , dewlap
, white, pale blue
cycle d l rnZ1tjnueend eggs a be laid at . ~ k d o r 1Eatern
~ Panama to nwthweabrn
Ecuador.
regulw intemals PITCH

-
1982).

-
hdyiw c h r u m ( W w s & I~UEUWN
the folfOwingkt, information fab d 3 le671, Bwviora, Mus, Comp. Zool,236:2; type
localiky Rfo T u h at Quebrada La PIah, 100
and dewlap coloration are for live spci- ,, DhBn, SVL to 79 in males,
mew; accept wh~rea t h m i s ~indi~ttsd* to 73 in femal~,B& en with or
d d a p colbration pertaim omly to ad& without darts seen obliqw E d * ; dewlap
m&, firops bi?w&~a$ (PETERS 186313)is Pale Or- with Pn l i n ~bathern
. Costa
excluded became there appeare to be no Rica to Colombia, 100-1100 rn elevation.
valid record for this sp&es in Central Drrctyhu fmnuta (COPEl8B9), 8Ci. Bull.
America. BARB~UR
this
(1823) emneously
-Onfor lasedon
&w*z byiF--ino
males. Body green with several dark green
mbiBentlsed N; uitt&wus (KOHLEBunpubl. a(- o u t l i n e d d t h p e ~that
)
ly & and form dark oblique bands.,
data).
dmh urdformely cream9olored (oocaaional-
lyni4 black shading). Costa Rioa to nmth-
ern Coloanbia, mlwel to 820 m elmation in
lowland a d premontane rain foreata
Da@lorr iwhb (COPE1811), Roc. Aad.
I Nat. Sd. PbiladelpEa Ba: 213; type 1 ; d t ; y :
Ban JOE&,Costa Rim. SVL to 160 mm in
, malee, to 135 mm in females, M y yellowish
Rg.172. Andie allis~nd(Isla de Roatln, Fig, 176.Dactylw chmorum (Cemo Sapo,
Honduras). Darih, Panama), Photo: C.W.Myers

Fig,173. Ctenomtua cristatellus. Fig. 177. DactyEtm frenata (Costa Rica).

Phoh: R. D. Bartlett Photo: A. Fliachendrliger

.)a.,
Hg. f 74. Dactyloa casildae (Fortuna, ,,,
Photo: EZ. Lips

Fig. 176. Dactylom chlari~(Rio Jaque, Ilaridn, Fig. 178.Dactydoa insignis (La8 Crucea, Punta-
Panama). Photo: C, W.Myem renas, Costa Rica), Photo: &-H.J@er
 gray to tan or greenish with a distinct large,
pal~ce~ltered dark-outlined blot& abwe
ehouldep; hw1ap o m g e red, with or without
~everalhorhental green or white bars and
dark spotting and with a white to greenish
h e margin, Costa Rica and Panama, ma
level to 1600 m elevation.
D a c t y h latifnom ( B m m m 18451, Nachr.
Ckorg-Augusta Ueiv. und K, Gee. Wiea
02lttidgen 8: 39; type laeality: western
Colombia (see MYER~ & BII- 1996).8VL to
124 mm in mdes, to 97 mm in fanah. Body
green with a distinct large, p a l e - m b d ,
dark-outlined b l o a above &odder; dewlap
ale grseniah gray with pale yellow aeales.
& a & mpanama to nartewrnbrn ~cuadm.
& c Q h ?nicrd:ue (COPE 16713, k.Acsd
7
Nat. Sci, Philadel hia a4: 214; type locality:
8an J d , Cwta h a . WL to 111 mm in
d e a , to 104 mm in females. M y yellowish
gray tn tan with a e v d pale brown-centered
dark bande that may cbaege ta p n on
fl& and a distinct ptarbital dark-bor-
dered pale &ripe; dewla salmon with white
d e ~Costa . Rica .nd 425-1500 r
elwation.
Norope altae (DVNN 1930131, Proc. N ~ T
England Zool, Club 12:22; type locality: Fin-
Acosta, 7000 R, Volcaa Wba3Costa Ricaca
SVL to 62 mm. Body grayish brown, dewlap
rust orange grading to yellm m h e masgin.
Highland8 of Cmta Rica, 1229-2000 m elm
tion.
-- -
Fig. 179.Dactyllra latifins Illear Nomami,
DariBn, Panama). Photo: C. W.Myers
96
Nomp8 aivamdeCtomi (NIETO MONTES DE
OCA 3LQ961,J. Herpeto1,1I#):20; type locality:
18,6 km N, 8,1 km W Ocoul~oautk,940 m,
1Bn66'N,9757W,Chiapm, M d m . SVL to 66
mm. Body grayish b m , dewlap darls red
with a suBMon of black pigment. lbgian of
El Omto, CBiapmqMexico,910-840 m eleva-
tion.
Norope arnplkqwmoaur McCM~ W ~ ~ S O N
& W m m m 1992, Carib. J. Sci. S& 209; type
locality: Fin- El Cuaum, 1860 m,15 31W. 88
12W, 8.6 lun WSW Buenoe Aims, Sierra de
Omoa, Cork& Hoduraa. SVL to 48 mm.
M y grayish brownv dewlap ye11ow-aranp.
Only known from the type localityt 1530-1720
m elevation.
Norops raniaolepb (SEXITI~, BURLIEY & FarPTS
1W81, J. Hespetal, 8: 147; type 1 4 10 mi
8E Ban Chrbtdbal Las Csaaa, L a p = .
Meuico. SVL to 47 nun. Body &sh hwn,
dewlap pinkish red, Meseta Central de
Chiam, Mexico, 1MH)-2600m elevation.
Norops aquatieus (TAYLOR1966), Univ.
Kansaa Sei. Bull, 88: 141; type locality:
Palmar, Puntareraas, COBta Rica. SVL to 71
mu Body grayish ellowiah green with b m d
transverse brown L a d s and a pale prsmi.h
yellow lateral longitudinal stripe; dewlap
Ted-orange with pUow ~hdpe%.Comb Rim
and Panama, 30-1170m elevation.
Noraps auPactw (DAUDIN18021, Nist. Nat.
Rept.: 88; type lm$ity: unknown. SVL to 52
graph brown, dewlap blue.
mrn.
Central anama and northern South
Amarkst, sea level to 2000 m elevation.
T Field Mw.
N o w b d e r i ( S C B A ~19391,
Nat. Hisrt. Publ. W .Ser. 24: 7; type localiw:
Gmmjal, upper Uspampa River, V ~ a m m ,
Mexico. SVL to 101mm.Body a d brown,
dewlap d a r ~orange with YBfTol~iahm a p .
margin. Caribbean versant of kthmus of
Tehumtepsc (Veram, Om8~8,%bascxl,and
Chiapaa, Mdm), 200-1100 ep elevation.
Nomps bhorum X ~ M ~1996b, R Senckee-
bergiana bid. 76: 21; type l d t y : trail to
Rock Harbour, about 3 km north of the tam
of Utila (16' B,34W, $15~69,94W),Iala de
Utila,hlae de la Bahfa, HondurasqSVL to 75
nun. Body grayhh b m , dewlap orange with
euffu~lonof black pigment centrally and with
white ~calea.Endemic on Isla de Utila, k1m
de la Bahia, Rondunte, sea level to 100 m ele-
vation.
Nomp~tbiporcattle CWmw 18845, Herp.
Mex. 473 1dB.8anta Rhss de Pamas
[=Pmuo113ta Venpaz, Guatemala. 8VL to
l(E2 mm. Body green (capable of rapy mlor
Fig, 180.Nornps a%varez&lfor?oi
(Chiapas, Fig. 184.Norops aumtus (Canjilones. Gualaca,
Mexico). Chiriqul, Panama). Photo: M. Lundberg

Fig. 181.Nomps amplisquarnosus (Cusuc Fig.185.Norops bankeri (Chiapas, Mexico).

National Park, Honduras).

Fig. 182. Norops a n i ~ o l e p (Ban

i Crist6bal de Fig. 186.Norops bicaorum (Isla de Utila,
las Casas, Chiapas, Mexico). Honduras).

Fig. 183.Nomps aquatic~s&as Cruces, Punta- Fig. 187.N o r o p ~biporcatug (Barbla,

renag, Costa Rca). Photo: K.-H. Jungfer Nicaragua).
 change ta b-1, dewlap blue w i t h dark7 W ~ o grayish
orange margin. Southern Mexico through
Central America to northern South America,
sea level to 2000 m elevation. The nominate
rubrpecies occurs in Central America,
KO- & RAWR 12001) demonetrated that
the holotype of Nurops bipmatw WEQW
1834) i~ conspecific with the ayntypea of N.
pteraii ( B o c o a ~
1873).In order to stabilize
hhe m n t and long established uage of the
names A? bip~rcatus and A? petersii, tht
International Commieaioa on Zoological
Nomenclature (ICZN) set aside the statm of
the original holotype of h? biporcatus and
dmignated MNHM 2426 (holotype of Anolk
copei B w o w 1873) m the neotgpe of N.
biporcatu (International Cornmiasion on
Zoological Nomenclature 2002).
Norops cap20 ePETER8 1863b1, Monata.Akad.
Wiss. Berlin 1868: 142; type I d i t y : Costa
Rica. SVL to 96 mm. Budy grayish brown
with green elements, dewlap y e l l o w i ~ h r + Chiapm, Mexico, and w e a m Guatemala.
Caribbean ver~ant of southern exlco
through Central America, sea level to 1800 rn
elmtion.
Norops carpenteri (ECHEL~E, ECIIELLE
FITCW1971), Herpetologica 87: 358; t q e lo-
& localiT
cality: Rio Reventazon, 7 km ESE h a l b a ,
Cartago, Costa Rica. SVL to 45 mm. Body
greenish, dewlap orange. Nicaragua, Costa
Rica, and Panama, eea level to 1109 m deva.
tion, Anolir procellarig MYERS 1571a 2
regarded to be a eponym of N,carpenten
I
according to lWmH et al. (1976).
N o q s cobanemis (STUART 19421, Occ. Pap.
Mu.bl.Michigan 4.04 6; type locality: 3
km S f i c a Samac (6 &line Irm of Cob&),
W60 m,Alta Verapas:, Guatemala. SVL to 60
mm, Body papiah bro~vn,dewlap purple.
Highlande on the Caribbean vemant of
Guatemala,31300-1410 m elevation.
N o w compressicauda ( S M Bt~ K E R ~ E R cality, 1650-1936 rn.
1966), He tologica 11: 193; type locality:
N e u La &a, 0-m. Msndm. ~ Y tLa M
btalhuleu,
,.,,: L.Melendez tion.
98
d ~ brawn, mewlap -om
rose to purple. Disjunct populations in
eastern Oruraca and western Chiapas,
Mexico, 6QO-1200 m elevation.
Nompe cancolor (COPE 1862b), Proc. Aced.
Nat. Sci. Philadelphia 14: 180; type localitg:
Nicafagua, SVL to 46 mm, hlae hovidenc~a
and ElanAn-, Colombia (off the Caribbean
coast of Nicaragua).
Nomp8 craarsdwl (COPE18643, Pmc.Acaa
Mat. Sci. Philadelphia 18: 173; type loealiQ
C o b , Alta Verapaz, Guatemala. SVL to 69
mm. Body grayish brown, dewlap red.
Highlands of northern Central America
(southern Mexico, Guatemala, H o n d m , a d
El Salvador), 1300.8000 m elevation.
Norogs crietifir (SMFTH 198&aP,Trane. Kama
Acad. Sci. 71: 195; type locality: nmr
Ipcacoyagua, Chiapae, Mexico. SVL ta 68mm.
Body grayish brown. Pacific vereant o
Norops cupreus (WOWELL 1&%1),Pror
Acad. Nat. Sci. Philadelphia 11: 481; typ
Nicaragua. SVL to 57 mm. Body
grayis b m , dewla h w n with orange
margin. Hondura~ta 8 o ~ t Rica,
a sea lewl b
1250 m elevation.
N a m p cuprinus (BB6FTH 19641, Herpetologica
O~W
PO: P ,
locality, Vicinity d Zanabpec,
31;C ~ &CO. SVILto 89 mm. ~ o d grayish
brown, dewlap red. Eaetern Oaxaca and
y
wedem Chiapas, Mexico, 200-600 m eleva
tian.
Noropa eusuco McCW, KLI- &W ~ O !
2000, Senclrenbergiana biol. 80: 214; type
locality: Parque Nadmal El Cueuco Centro
de Visitante%15"29.92'N, 88'12.88W,1568 n
elevation, Cort.4~~ Hondura~.SVL to 46 rim.
Body grayish brown, dewlap unioolw dirty
white. Known only from vicinity of type lo-
Nomps ddtfisianw l B o o o u ~1873X ~ %.
Sd.Ma.,Rept. 1093:84,type locality: Volch
AtiUh, 1200 m, Guatemala. SVL to 44 mm.
Body grayish brown, dewlap yellow. Pacific
vemant of eatern Chiapae, Mexico, an?
westem Gu~tsrnala,200-1200 m elevation.
Nom s exaul (ARI)SEIMENA& IBANEZ D. 19941,
carikl J,Sci, 2% type locality: *orillm del
Ria Chidquf, a uaas 100 m de la deaembwa-
dura dde la Quebrada Bijau (8"45'll"N,
82"11'0794), e n h 1050-1076 m eobre nfvel
del mar," Reeerva Fore~talde Fortuea,
Chiriauf. Panama,SVL to 48 mm,Body mw-
&i b k , dewlap oranga red vPikh-$lldw
hlcale~.Chiriqui, Panama, 1050-1076m eleva
Fig, 18% Nompa capita (Bartola, Rio San J m , Fig. 193. Noropa cmaaulurr (nearBanta h a
Nicaragua). E'ass QuichB, Guatemala).

Juan,Nicaragua). Matagalpa, Nicaragual,

Fig 191,Norape m & a m i a(PasadoEl Fig. 195. Nompa c u s w (Gem Cuauco, Co&s,
Quwal, Bqa Verapaz, Guatemala), Handwe),

Fig. 192.N o m p ~comprea~icauda(ElOwb Figb198.N m p s dolifmianus (Resews Natural

region, Chiape Mexico). L ~ Tarrahs,
B Suchitepbquez, Guatemala).
 17"12'68"N, Q2"57'47W, Municipality af
RaySn, at the border with MunIupetlity of
Pueblo Nuevo Solistahuacdn, Chiapas,
M&w& SVL ta 49 mm.Body m y k h brown,
dewlap rose, Highlands d northwestern
Cbiapaa. Mexicot 1525-2000 m elevation.
Nomps h~dmlli~l(&E.PERS l803b), Monaw.
Akad. Wh. Berlin 1868: 138; type locality:
Vemgua, Panama.SVL to 44 mm. W y gray-
ish brawn, dewlap redst.ange witb yellow
margin or dewlap uniform red orange (popu-
tations in eastern Panama), Northern O&a
Rica along both vemmta to central Panama
east of the Canal Zone, sea level to 1600 m
elevation.
Norops johnmeyela (WILSON & M c C ~ E
l$B2),Trrme. K m m s kcad Sci. BB: 189; type
: km WSW Bmnoa &a,
I d t ~ r 5.6 1580 rn,
El Cusuoo, Cortee, Honduras. SVL to 70 mm.
Body grayish brown, dewlap orange red with
Iargs centad blue blotch. Femalw with well
dwelo ad dewlap (yellow with latlfe mtml
blue dot&). Sierra ds Omoa Dptaa. Cop&
and Cort&~)l,Honduras, 1410-1996rn eieua-
tinn.

Norope karnptod I Dm l W a ) , Pro~.Acad.

NaZ. Sci. Philadelphia PB: 111;type locality.
Enea Urida, W q u f , Panama. SVL to 56
mm. Body grayish b m m dewlap rsd with
armge blotch &ntertorly, and with white
scales, Highlands of Cbirlquf, Fangma, 1008
1800 m eIevation.
Narope b u t a i MSJCRA~E, K~HLER &W~ON
2000, Senckenberpiana bid. 8Q: 218; type
l d t p 2.5 & h e km NNE La Fortma,
lti026'N, 87*"rW, 16YO-11690 m elmation,
Pi,Hmduras. SVL to 51 nun. BxIy grayieh
brawn, dewla pale ~ 0 1 1 with
0 ~ purple males.
K n m only the type loeal~ty.
Narop8 laeviventris ( W r E a U N 18341,
Heqetalogica Mdcana: 47; type locality:
Mexim. SVL to 5a 19m.Body grayish brown,
dewlap dirty white, Southern Mexico to
-tern Panama, 1160-2000 m elevation.
Baaed on the studies of B m m i1@99)and
KUmim.I2001el I coddwArr01isr i~terrnedius
l W b , Anolk nannordes Corn 1M4,
and &adis botlrgeaet' Bmom 1875 to k
of N. hviw&5s.
N o w knztctzm~(COPE18gld1,J.Acad, Nat,
Sci. Fhiladelphia 18: 213; type locality:
V~rggm,Pazlama.SVL 20 76 mm. Body gray-
i&h brows, dewlap red (offen with black
scaler). Central Veracruz, Mexico, along
C L U ~versentP ~ oF C e e M America to oerm-
rig. r a r .iwurrrprr I uawurubu.r ~WULUIU,
tfd Pmimq also on Pacific v w ~ a nin
t Costa
Rica a d Fsrnama, EM level tb 10SO m eleva-
Paataza, Emador). tion.
Rg. 19.8.Nomps h~aeterophob~dotw
(El Htal, Fig. nu& NOW ~ ~ u i u e n t(Salva
r i ~ Negra,
Hondurasl). Matagalpa, Nicaragua).

Fig. 1Sg. Nomps humit& (El Vdle de ARMd, Fig, 008. Nomps femurin- ((Bartola,Rig San
Panama). Juan, Nicaragua).

Fig. 200. Nomps johnneyeri (Cusuco National Fig. 204. Norops limifrons ( C e m Saslaya,
Park,CorGs, Honduras). Atlantic0 No&, Nicaragua).

Fi Vop~ :emptoni (north of Boqu Fig. 206, Norops l~onotus(near Puealo Wieo,
Photo: 1. A l e Jinokega, Nicaragua).
101
Nompa lidfrom (COPE1862b1, h. Acad. ~ o m p pachypue
s (COPEl87%),J. Acad. Nat.
Nat. Sci. Philadelphia 18:178; type locality: Sci. Philadelphia (2) 0: 122; type locality:
Vemgua, Panama, SVZ to 48 mm. 3ady gray- Slopes of Pico Blanco, Costa Rica. $VL to 50
ish brown, dewlap dirty white with a b a d mm. B d y grayish brown, dewlap red-orange.
orange.yellow spot or (populations in central Highlande of Costa Rica and Volch Chiriqui,
and e a e t m Panama) dewlap dmost mm- Panama, 1370-1580 m elevation.
pletely orange-yellow. Emtern Honduras to Nomp.9 pmndoewis SAVAGE 8z GUYER1998,
Panama, eetr level to 1350 m elevation. AnoEisRev. Biol. Trop, 48: 8Oti; kype locality: La
bkutiger T ~ 1956R and Anolis g d m a n i
Changuinolita, 1980 m elevation, Cant6n
BOULENQER1886 are both regarded to be Coto Brus, Puetarenaa Province, Costa Rica.
8ynonyms of N.l i m i n a a m d i n g to SAVAGE
SVL tcr 60 mm,Body grayish brown to olive
(2002).
green,dewlap pink with orange anterior mar-
Nomps lbmtus (COPE186ld1, J. Aad. Mat. &. Cordillera Watl~ancaon both ddes near
Sci. Philadelphia 1%:210; t e locality: the border between Costa Rlca and Panama,
Cucuya~de Veragua, Panama. & to 78 mm.
Body gray%& brown, dewlap yellow to yellow-
1350 to 1950 m elevation.
orange, Earitern Honduras to central Nomps panricinctdatusr (ALVAREZDEZ TOM &
Panama, sea level to 1200 m dwation, Baaed 8 ~ 19561,~ Herpetologiea
3 18: 5; type local-
on the studies of O B m (19@8),1consider ity: El Suspirq 1200 m,Chiapae, Medco. SVL
Nomps o&ophus COPE1876 to be a eponym to 50 mm. Body grayish brown, dewlap
of N.liorwtus. yellowiah orange with a red central blotch.
Centd-eastern Chiapae,MBxico, 500-1200 m
Noropa Eoueri$gei (SCHMIDT 1936a1, ROC. elevation.
B i d Soc. WdGngbn 49: 47; type Locab* Norops pentaprion (COPE1&62b),Roc. Acad.
Portill0 Grande, 4100 R, Yoro,Honduras. SVL
ta 118 mm. Body grayish brown, dewlap or- Nat, Sci. Philadelphia 14: 178; type locality:
ange with pink streaka. Mountainow regions Truando River, Colombia. SVL to 79 mm.
in the Dptos. de YOM and Atlbntida, Body mayish brown, dewlap red. Along the
Honduras, 650-1100 m elevation. Caribbean veraant of Central America h m
Tamdipas, Mexieo, to Panama, including
Norops mtaccropha6lus (WERNEB 39171, Mitt. Yucam Peniesda; also on Pacific versaat in
Zool. Mw.Hamburg Sd: 31;type locality: S m Coeta Riea and Panama, sea level to 900 m
JOB& Guatemala. SVL to 46 mm.Body gray- elevation (to 1780 m in Veracruz, Mexico,
ish brown, dewlap roae with a orange gellow according to V~rtret al. 19971. TWO subspe-
basal blotch. Pacific versant o f northern cies are mco&ad: Nomps p, pentaprion
Central America (Guatemala and El (Caribbeanv m a n t of Central America from
Salvador),48-1320 m elevation. eastern Hondnraa to Panama)and N m p s p.
hk?$(BOULENGER 1881) (Caribbean ver-
Noraps matudai I S m 19561, Herpetologica sant from Tamaulipas, Mexico, to Honduras;
I& 1; type locality: rsgioa of sow nu&^, including YucatBn Peninsula).
Chiapaa, Maim. SVL to 42 mm. EMy gray-
ieh brown, dewlap purple. Sierra Madre de Norops petersii (BOCOUBT 18731, Mea. Sci.
Chiapae, Chiapae, Mexico, and adjacent high- Mex,, Rapt,: 79; type locality: Alta Verapaz,
land~in Guatemala, 200-1000 m elevation. Guatemala. SVL to 102 mm, Bcdy greenieh,
dewlap urple with pale yellow margin and
Nomps mumEla KOHLEB, MCCW gt pale yelfow gorgetd sealea. an ~ u i Potosi,
s
W I L ~ ~1999,
N Amphibia-Reptilia a0: 286; Mexico, to Honduraa, 200-2130 m elevation.
type l d t y : along trail to C m de Enmedio,
1500 m, Parque Nacional La MuraUa, Noropa pijolensia MGCRANIE, w~~k3014&
Olancho, Honduras. 8VL to 56 nun. Body WILL~AM~ 1993b, J. Herpeh1. 97: 893; type
grayieh brown, dewlap red. Parque Naciond loc&ty: Ea-et s l o p of Pico Pijol, 2060 m,
La Murda, Hondurae, 1440-1740m elevation. 56VOT, 88°SW, Montaiia de Pijol north-
west of Tegucigalpa, Yom, Honduras, 1860-
Nomps oeelloscapularis E C ~ ) A ~ RMCCM
, 2060 m elevation. SVL to 80 mm. Body gray-
gt W ~ PPOOla,I Heptologica 6F 248; type ish brown,dewlap Mae with pink id-.Only
t d t y : near Quebrada Grande off a trail to known &om the type W t y . Note that the
Laguna del Cerro, 1tiD04,82'N,88"68.45W, species name hart been emended from the
1200 m elevation, CopBn, Honduraa. SVL to
47 mm.Body ayish brown, dewlap orange.
r o
Known anly m vicinity of type locality,
&"
original apellin p&kneeA to " p W n s i s * to
corre~pondto e rnmdhe gender of the
genm name N o m p (following H . S ~ .Bi: H
1150-1870 m elevation. LriflsEN 1974a).
r lg. hvv. rvuruprr ruvvr I W F ~ \avrw-u.ups ~f Fig.210,Norops petersii (Chiapas, Mexico).
C e m Bdfalo, Atlantida, Honduras).
Phato: J. R, Cranie

II
Fig. 207, Noropa macmphallus (near Fig. 211. Norops puo as (east- R 01 n c o
Ouanagasapa, Escuintla, Guate~mlm.). Pijol, Yore, Honduraf Photo: ,.McCranie

Fig. 208, Narops ocelloscupullaris (holotype). Fig. 212. Norops poecilopus t Nusagandi,
Comarca de San Blas, Panama).

Fig. 209. Norops pentaprion (Bartola, Ko San Fig.213. Norops pol.ylepis (Esquinas,
Juan, Nicaragua). Puntarenus, Costa Rica). Photo: V. Maass
Fig*215.Noropa quaggulw (Gem El Tom, Fig. 219. Noreaps 8 e t i c e u ~( h c h o Akgre,
Atlstntico Rcfrte,Nicaragua). Chiapaa, Meleimh

Fig. 218.Norope salprei (Florida, USA), Fig. 222. Noraps tropidogaater (Cbrdoba,
Colombia). Photo: M.Lundberg
 blow by a black gtripe; dewlap dark brown.
Iala del Coco, Costa W, sea level to 630 m
elevation.
N m p s tro idagader C H A L M ~ L L 18661,
h c . had. &at.Sd.Philadelphia 8: 224; type
locality: Colombia. SVL to 66 mm. Body pay-
iah bmwn, dewlap redsraslge with white
scalea. Emtern Panama and northern South South Americat aea l d to 900m devatia~,
America, .sea level to 7Ml m elevation,
N o q e voctfkra~CMmt8 1971b),Am= Mus.
Noraps- tropidokpi~( ~ o l m a ~1886), m Cat. Novitat. 2471: 16:type h d t y :4 km W cenro
Zizards BnL M a . 21L ed, 23: 53; type localits: Pan* 1828 m, W q * Panama, 8VL ka 64
Vol& Iraeu, Coata Rica. SVL to 59 mm. lgna B e g r w h brown, dewlap red w ith
Body gray%&brown, dewlap purplih red to white sale& Costa Rica and Pansma, 1400-
pink. Vol& frazu aed Monbverde, Coeta 1830rn elmation.
Rica, 1220-2500 m elevation,
N m p a i.uampwt(~iaMCCRAPJ~ & 3hr.m
Noro s tmpidomtus ( ~ R 1863b1, S Monats. 2001,Sen&-- biol., ZhnMwt a M.,
~kab: W3.s. Berlin 1- 156; t m (U2X 22% type Mts;a o h n . OfRfos ~ ~ ~
Huanuseo, VB~BVU~, vl"
Mexico. 8 ta 'O
Body grayish bwwn, dewlap dark orange
m"mm.
60 : Aner and Wmpfi, 16"0S'N, 86*0r"W, 110 m
elevation, Ohmihu,Iimdm.BVLto Umm.
with yellow to yellowieh orange m~tr
central dark ~ t r e r l .South- G~dz
nostbern Nicaragua, 90-1740 m elevation.
Body gmyh'h b ~d a d ~ t3mngered
,
pllaw margin. K e r n a y from ths type
l d t p d96-110 m elmation.
with

Nomps uni$nnh (COPE 18851, Proc. h e r . N o m p ~wsrmuthi I.EaEIL?Ei& OB~MEZ$R1W8,

Phil. Soc. SB: 392; type locality: Yueat-gn, Se&nLwgha biol. '7% 129; type 1-
Mexico. 8VL to r10 mm.Body graybh brown, road &om M a w p a to Jinoteg;a at lrtg 146
dewlap rose with purple epots. Southern (18"01.996Wv86 $6.84@#)), 1400 m elwati-
Mexico to weatem H o n d w , sea level to on, Dqwrtmwnto JIln~tega,Nicamma. SVL
1800 m elevation. to 64 mm, ayita%brawn, *lap red
Norope utileneis K C ) ~ RlW&, Sencken- with
~i~ pals to d&r brown ades.
1250-1600 elerva~on. North-
bergiana biol. 76: 24; type lacrtlity: 2 kni NNE
of the town of Utila, Isla de Utila, Isllas de la Nolwrp~d l i (mlf440a1, Proc. Acad, Hat,
Bahia, Honduras. SVL to 69 mm. Body gray- M..PhikWphi~Q2: 110; lo&@: El
ish brown, dewlap red with white scale^. Isla
de Utila, Hondurae.
Vok&n, Chk&&, f,k %
13 trr 100 nus
Body grayish bmwn, dewlap dark-orange,
brownish olive BF bl& (apeb1e of color
Nomp. villoi (RTcs & HWDER~ON197B), -1. ptrn-B soeta mGa,
Milwaukee Pub. Mus. Conkrib. BioE. Geol. 9: 1;
+-

~mul panBznaI
, ~ ~ B ~ Z MmU ]dWat;iW,
locality: Great Ialpnd, Nieu-. ~ ~ 1 0iBm . 1 d~ to ~
55 mm. MY fFaYish brown, dewlap be a ~~~ o f l ~w, d f a ~ a- bp SAVA~
czoozj. -
N0mp8 y~r~e?%Sb McCWIE, N E H O ~ &N
Kt3m.a~ ZOM, Amphibia-Reptilia 466;
m
typa locality: L6 atrlins km La Fb-,
1600 m elevation, l&"B%N, BTRIBfUr,
Cadillera N m h de Dim, Yon, &dm.
SVL to 47 mm. Body greyieh brown, dewlap
pnicolar o q . Northwestern Honduras
(Yore), 1180-1600 rn elevation.
Nom 8 &us K ~ W R 8t MCCABNE 3001,
~eo$nba@am Sol. Bl: 288: Qp1 4 t y :
Be National PIco Bonita, 90 m
-3
dm9tian, hkida, H&-.
m. Body $r&dl lmmm, dewlap d m l m
SVL kr 44
Idixty white. Caribbean eemaat of the
I ~ g 2 . fLngosts~ (northdope of C&
8~ornp. $v$~ in no*hem
Pando, 1450 re, Bema deI Tom, Panama).
Photo: C. w.Myers
196
 Fig;. 222. Norops vitdigerua (Cbrdo'ba,
Elena, Guanacaste, Costa Rita). Photo: Bille Colombia). Photo: M. Lundberg.

Fig.226, Norope tropidonottss (male; &&a Fig.229, Nampg dropidonntlta Bmale; Rancho
M e m , Ghiapas, Mexico). Alegre, Chiapas, W * o ) .

Fig. 226. Norops uniformis (Lacandona Forest, Fig. 230,Nomps wermuthi (Cerro El Toro,
Chiapas, Mexico). Atlbntiee Norte, Nicaragua).

Fig. 231.Nomps waodi (Monteverde,Coab

WaI, Photo: H.B-oe
 'tf
---4
 Ctetwnotus cristatellus Norops aluarezdeltoroi

Not-aps amplisquamosw Nompa anisolepis

g e2,*- -
;,.C$4
v:;.....
-3*;$g*
%'',
*
,,.'S*.=.
kq&
c:;., "'.
,-.$?><.',. *:......
..;:
:.:
,"...k.-+.
. r-."":
,,:, ,.:;
:{<2:,t>,-5
, 3

,
. .- . .
, ,
.v

b..

. . .* .. .
,
,.
. .
'$
Norops hiporcatus Norops capito

Norops carpenteri Nomps cobanerteis

Fig. 932 bb 273. Anolea with eartended dewlaps, Because in most epeciea fenalee have no or very
) males mare shown.
malh dewlqw (mexception in this ~eapgctia N o m ~ J b h n m q e donly

110
 Nomps liolbotus

Nomps pentaprbn Norops paecikqus

Norops polytepis (normal Nompo potylepis (rare variant) Nompe quaggulus

Norops rodriguezii
Norops sermnoi

I
Norops wernuthi Norops zeus
 Key to moles (generaAraoli'ig h t y .
Elaborating an identifiation key all Carrhal
American moles ia a dBcnlt task, partly
because m e s p 5 m are known &om a single
eneeimsn or a &w individuals. In theere
&i ;ariation of aauppwely diagncrstic character:
iaticr is insufficientlydocumented. $ o m apecim
are extremely similir in their external m@o-
logy m d coloration, but can be disthguiehed
readily by their very different hemlpenial
morpk;olo#, which, in &me case4 is inte$reted
as evidence for onlv diakmt relatiomhi~sof
thew cryptic ap+&e~le. N o m p cuprew &d N.. or and posterior
&
mocmphollua;K O ~ RKREUTZ1999). The fol-
lowing information (including EIVZI refere ta
of tympanum.
adulb ~nlv.Juveniles of m a n y mole aneues are
extmmelyly&f8cuItta identi$Also, there is m-
eiderable intraee~cidc (nartieularlv sexual)
vaxiatian in ,w& apeciae.BOme~ref~m, few ~ p e -
ciea appear more than once inthe fdwving key.
Statements regarding coloration (of body and
dmlap) refer to Bpecimens in life; dewlap color
mfera to addt males, if not othervim spec-ifid
I ( f d e e of moat anole species have nQ or only a
rudimentam dewlad. Bubdieital lamellse
1- caunta are &en for-phalangeeii t o iv of the
fd tm (not including distal phal-1, if not
otherwise specified.

1 a Mostly men in We (capable of rapid color

chana to alive brown): in adult male%head
mmp"imausly large &d elongataf dewlap
pink, 2-3 ~caleabetween nasal an rortrd;
BVL 66-87 mm; short-legged (le toe of
adpwaeed hindlimb reaches usually to ear
ope*, in some *individuals to point be-
tween ear opening and eye); dewla pink
.*. Anodte Jk@,i
Fig. 276&Ventral scales in a n o h :
b Combination of charactera dXerent from
that above .......................................... 4 a, 'rmbricats, strongly keeled, mumonate
b. imbricate, weakly keeled, natl-mumonate
E, smooth, non-imbricate
d. granular,not keeled
e. imbricate, smmth
f. slightly imbhcata, &moo&

Fig. 277. Nor* wermuthi.

a (leR).Mid-domum; note several rows of enlarged
Fig. 274. Nosops troptdonotus; now tube-hlce d d scalm
d l a q pocket Earrow). b (right).Htttarogeneous lateral body scales.
114
la A deep tube& d a r y pocket present b Ventral8 to distiElctly keeled; supra-
(FTg. 274) ....................................................
9 caudal scales of equal size (Fig, %Ob-d);
b No deep tube-like d m N dewlap dmk omge, 0 r ~ F ofdor-
et present,
at most, a shallow w A k y depres~fon 8 .... with pink streaks ...............................
.,. 10
8 a M e 8 mruior to em opening flat and h e - LOa 2-3 scales between eupraorbital semi-
ld etnd abaut twice as large a8 acales &dm; aupraoeularer smooth ....................
........................... ,,...,.,...... Norap k M
posterior to ear o
usually grt~dar;Q 2 7 5 ~ ) r'"......,..............
b Scalss sulterior to ear opening about same
are4 b Either 0-1 or 4-6 sfill8s between m ra-
d i t d ~snicimles;auprapCular~kw1d 11
eiw as ecalea posterior to ear opening (all lla 0-1 scalea between sugraorhital semi-
granular;Fig. 275b)................-.........,...,..6 circles; maIes with enla ed postanal
4 a Dewlap rose to ~nrg1ejt d tn sad@ 3
............................... O F o p s sag&
mdes conspicuously comprwsed .......... b 4-6 scales between eupraorbikal semi-
..........................
Noropo o o n y r ~ n i a a u d a r circle~;malea without enlarged postanal
ades .......... N o m p ~krw-el l h part)
b Dswlap orange red with y e h margin
(with or withoot a rrentral dark streak); tail 1Ba Head conspicuously broad and stout, snout
not compiauoualy wmpremsd ..................
8
r:
ni rtr blunt in lateral aspi& long-legged
6 a Adult males 60-60 q n 8VZ, adult femalm (4t trre c& adpremed hindlimb reachee at
45-65 mm SfrL; dewla orange red with a least fa anterior bwder d eye); usuany
p n o n margin and a & k c t an- dark with a pale band acmw chin (mwt diatin~t-
stre& ..................
nor up^ hrrpfd~nobue injuvenilee;Rg, 279);domal eedm smooth,
juxtaposed, mostly pentagonal or hem-
b Adult mgles 40-60mm SVL, adult fmale8 gonal; tip of tail enlarged, not tapering, no
42-61mm 9VL; dewlap orange red with a enlarged poahnal soales in males; SVL 80-
yellow margin, but without a m t t d dark 96 mm; dewlap uniform nieh p1Iow
streak .....................
N o ~ c l praampweml
~ ........................,..............f?amps capat0 I-
B a Dewlap rms with purple spots
....................................
Norope uni.polPnirr
................. b Combination of characters different %om
that above .......................................... 13
b Dewlap red orange with yellow margin or 18a Shmt-legged (qth toe of adpregred
dewlap u n i f a m red orange without y d a w hindlimbreaches unually to a r opening,in
margin (popdati.oaein eastern Panama) 'il some individuh to point betwsen ear
4 a Hemipenia r~latiively large with well- opening and eye); lar adult size (SVLto
developed elongate l o b and with a 100 d; 9-11 @upra!?Ibiah to level below
strongly Ealyculat& ~urfaoepn both the center of eye; males without enlarged
politanal malea; dorsal5 small, not mu&
truneus and the l o h ; maximum SVL 4-62?
mm in mdea and 80.0 anm in b d e r larger than Qank ecales ............
..,,,.,,.. .. 14
......................
N m p o Isudlie (im part) b Combination of charactera dtkent &om
that above ................................. I6 .......-
s ralativel mall with sthort end
2$E* and nid0.t. ItmnEly c . 1 9 0 ~ -
late ~urfacson &her the t r u n OF ~ the
l k Domal head 6calee faintly b l e d ; ventrda
only slighty keeled with mundd paterim
lobes; maximum SWL 87.0 nnm in males margins (Fig,276b), 62-78 ventrals ba-
and 41.0 nua in famalea .......,...,.......
tween levels of d a md groip; 120-14
...............................
Norups quww E w wales around midbody; throat region with
8 a Tail disnctly compressed laterally md dark,-B dewlap purple with pale
with a crest of enlarged e d e a le~peelally geIlow marlpin and pale ellaw gorgetal
vhible in males) ., 8 ............................ ,.&OrT)PII pefmti
....,............. ...........,...
b Tail not disbctly compressed laterally, b Dorm1 head malee strongly keeled;
although a 5ligbtlg carbate crest can be ventrals str~ng1ykeeled and mucranate
pm~ant ......,..,......, .,.,
................la (Pig, 276a), 40-54 between levelr of d l l a
and p i n ; 100-112 d m wound midbody;
9 a Ventrals smooth; tail with a m&of supra- no dark stmalra in throat region; dewlap
caudal acalea of unequal e h hequenas of blue with drtrk orange rnwgin ..............
wnaller reales that increw in b the ....................................
Nomps bdpmria#w
following lateral whorl; Fig. 280a; dewlap
greerdsh pllm with orange margin .......... 1Ba Zris metallic blue in life; large a d d t size
.......................Ctemnotws cI.dntateIb (SVL to 100 mm); body often with dark
green or brown ban& with numerous
small do&;2-3 scales betwsen supraorbital
 Sedbin:les; 1 scde ketween subaaular8
mdx
g&
abial~;males with enlarged
scales; dorsal umd color dd1
red m dark olive in?%, pale yellow to
tan with numlerou~emall punctatinne ia
rewmtive; dewlap Iargs, dark-o
F m m oliw ~ or blao~(cs able
c h q ) .........,...................
i$ amps d i
0 Combinahn d h e t e r &ff~ent ~ hzn
Fig. 278. Enlarged poet- Fig. 279. Nomp-
anal scalea in Nomps capita: ventral surface
that above .........................................
18
tropidog~ster. of head. 14% Adult size more than 100 mm ................
17
b Adult size less than 90 mnm ....,..,.,.,,,,..
M
Y~ntrab&thotly keeled; 1-2 d e s be-
tweea s u b a d a r ~and BU dabids; d s
rlwt m*rpd P r n M T d W ..............
.....................Nompa Ewes-l Ifn part)
Ventds smooith;suboEulam and suprala-
bide in contact; males with enlarged
p a a l d e ~ .......................
,,..,..,..,.
8bort-legged 44& tca of adprewed hhd-
limb reaehea usudg te ear ope*, C
BQme indSviduds ta point batwean w u openg
ing and eyek brown md meam / w h i b isl
mlwation, a h a with a greenish cast En life
......................
...S
". ...................................
I@
e a s e d (& toe of adpressad
reaches at least to oenter of ew:
Fig. 280. Lateral sealation of tail (scales of
w h ~ r l sshaded) 19a Inteqgietd plate not didnct, m afield
e)s; 4 to^ nitb fewer tban 60 BuBi~agital
lan3db Cf3-mham d digit to claw); &ult
eize t o 111 (males) or 104 m m (fa-
malea) .,......... ............Drrc&Eo;a mbmtas
b A wdl dewqlopd in26rpdet.d pla* pres-
ent, distinctly larger tJqg erdja~e~lt
scalm,
parietal eye pre~ant;4 toe with more
than 60 ~ubddtallamella%(from b e af
digit, tE)~law3;adult size t o 168 rnr~l(malea)
or 155 mm (femaleal
..'........*...m....... .............. .....................
D g o t y I o ~insig~ris
ma All of tapper margin of eye with edmged
s u g ~fm ~ h~g as stiff [Fig.
a. No~aprrM~n81a
zsza) *...................... Da~&&c% W ~ I M
b Enlarged aupwrseiliariew re&ieted to mtw-
281.Ventral ewrh.ce8 d h O . ior half of eye, f ~ ~ l l mbyds m d gtmulw
scales postjeriwb, not farming a stiff ridge
(Fig. 282b) ...........................,.., 81 ........
Sla Dimm with 4 5 oblique bmwn btm6 (not
fmctd of adjacent o~elliliatetape~~ed by
blue-outlined yellow a b b ; dewla dirty
aeam wit& broad y e l k am1e mat8 k gu-
larlp inteqsemd with a d e r m d d
&rm rroalba; mal in contact with rwtml
a. Dedyloe ktlfmns b. Dectyloa fmnste or lseparated by one a& ..........................
..................................D 1 ~ ~ r y 3caddm
oa
Fig. 282, Eye region (enlarged auperciliaries
orange). b Domum with a e v d dark green ocelli
tmmS a u h d with mIlaw1 that are clo~e-
 ly arranged and farm dwk oblique hB., b NBbony parietal pmtuhe~amea;wales
dewlap dull, creamsolorad (owaaionally alang d o 4 surface of tail Easinate (a-
with black hibdingl; nasal ~eparated&om largeti or n&) ..........,..............................
80
rostral by ane or two aEalee .........,............
.............,.,......... Dcfctyf0tl~100Jy1. 2Rn Venfmls keeled; a dark intembital baF
pmmnt; mderr with snlargd pa5tand
ma 6-9 ac&s between interparietal and eeales ........................ Nomps v o c t f k m ~
auprawbiital d & c k (Fig.283bl; body b Ventrals ~rneoth;no darls interorbital bar;
ydlowieh
r
f n, with bmad trammeme
brown lnzn e and a pale gmmish yellow
lateral 10x1 tudinal etripe; dewlap r d -
mdaa without enlarged p o e t d ecales Zf4
29a Proximal subdigital -lee Dftoee granular
or- witf oblique yeUaw stripes ...... [PTig. %&la); ~ d e as l q domal Burface of
........................... .
N m p a aqmtkue tad carinate, but not enlarged; SVL ta 59
msn ......................... N~ro~g utZkFwk
b 0-4 scales heheen interparietal and
supraorbital s d c i r c l s e (Fig. 2S9a); body b Prodma1 euWigitaI sctrlea of h e differen-
t;teFn diffment than above; dwlap color tiated as alight$ broadened lamellae CF'
t erent than above .......................... ..
E3 Iglb); rcdes dong don* surfam of t8
ma Bright w e n fa life ....................+.......
,....%4
b Mmtly ptay or bmwn {at most dull olive
wen) in life ......................................26
V ~ t d sdiatindly kwId; shart-legs4
(4 tos of adpnesaed hindlimb daea not
reach to aye); SIT to $0 mm ................
..................................h t y b ehloda
b Vmtrals smooth; long-legged (4th Coe of
adgrsead hiadlimb reaches at leaat; to
anterior border of eye); SVL +a73 mm .......
, .....,....................... Daetyfoa ehocorum
3% EFMb l n in ~ lip% very long-IegfgBd (4th toe
of adpreased hindlimb ra~ehesbeyond
antrtrior border of eye); distal halam of
f b g w 5 arid Me&ranrrpicuouerly Kng, about
the e m length a~ m m length~ of
penultimate and antetpendthate phalan-
gea; d d ground color olive b r m with a
greenish ca&; lateral body with 3-6 oblique
or vertical pale line* ...................
..
................... Nampa adlvamzdelbnrd
b Combination of charact~rsdifferent than
tAat &ope, particulw1~distal ghdamr of
flnpra and toes nab conspicuously lcmg,
much ah&@ than combid le
penultimate and antepenultimate p a h -
gee
Td
....................................................
E4l
%@aVentral5 nnlm or emaath and eubirnbri- b. N o w 8 WuaNcuS
cab; flGalr!s hornogmmou8; veny 1hort-
legged 14 t w of adpreseed hindlimb ma- Fig. 283. Dtrrsal view of head,
&ee not much f&r than efioulderh IB or
fewer hmhantal rows of lomala;mprtimu-
l ~ mm o d k liniag of throat black .........
t p m t ~ p hgroup) i ~ ....,.........................
87
k Combination of &@actera Werent h m
that above .......................,.......,.
......81
Bra A pair of emall bony parietal protuber-
ances pasterior and lateral to interparietal
plate; seaha along dorsal 5uFi$ce of tdi
carinake, but not enlarged ..........................
...................*...................
Nopops %rngostls
 carinate and distinctly e n l w SVL to 79
mmn ..........................................................
3Oa Adult males with a wall-developed serra-
b d middoreal meet (Fig. 285a); SVL ta 888
80
1
..............................

J
nun Nampe c ~ f i ~ r
b A double-row of @lightlyenlarged mid-dor-
sal sealea, but no -ted creet present
(Fig, 286bl; SVL to 79 mm ......................
......
............................. Norope pentapden
81a Maat dmaal head and body sales strongly a. &mps c ~ r
b l e d ; lateral body sales homogeneoue;
ventral seales strongly keeled and
mumnate; 7-12 distinctly enlarged dorsal
scale mws; males without enlarged paat-
anal smlee; dewlap red orange with yellow
mar& or dewlao uniform red a r a w
withkut gp:Uaw margin (populations h
eastern Panama); maximum SVL 46.2 mm
in malea and 60.0 rnm in females
......................Npmps ?bumilk (inpart)
I
b C o d b t i o e of characters Merent from
that above .....................
,.. ............82 b. Wonyla pntapibn
SSa U d y a conspis-uaus lyrffom marking in
&pjtal e o n , can pe.fSapeqfRd (pr* Abb. 285. Middorsal scales in Norops criebifer
sent In about 80% of mchuldualal, ventral8 and N , pntaprlon
~trunglykeel&and mumonate; relatively
long-legged (4 toe of adpressed hixrdlimb
r
r e a h at least to terior border of eye);
males without e WIW~~ m a scales;
dewlap red, red-omge or brown-red
l M a Dewlap orange red, with suffusion of black
sent centrally; malea average 66 mm
(lemurhue group) ................................ 99 (maximum 76 mm), females 64 mm
b Combination of character^ Merent &om SVL (,rridmum 67 mm); endemic on Iala
that above .........................................-87 de Utrla ..................... Namps bicaomrn
58a 10-20 dorsal scale raws irregularly a- b Dewlap pink red, with a d h i o n of black
Iarged; flank scalem usually beterageneom pigment mntrdy; malea average 65 mm
(enlarged scale8 scattered among ~maller SVL (maxim- 62 nun), females 67 mm
scales); SVL 80-60 mm (mdesl or 55-72 SVL (maximum63 mm); endemic on Isla
mm (females& dewlap red-orange with a de ................N ~ m p rsu d r u r e d
dark blotch .................N m p s v i ~ ' g e m d 57a Lateral body scales h&empn8ow,solitary
b Only the two medial dorsal scdea mws enlarged keeled or elevated d e s (often
slightly enlarge& flank scdea homoge- whitieh) scattered among smaller [meetly
neaus; dewlap without e dark blotch, al- granular) later& [Fig, Z'i'7bl; malee with
though o h n with blaek prgetal scales 33 enlarged patanal scales; suboculars and
aupralabials mostly in contact ( c m g ~ l u e
M a Dewlap red tu red-orange without suffu- gmupl ...................................................88
aim of black pigment, o b n with Mack gor-
tal males; mala, on the average, smaZ1e-r b Combination of characters different from
%an ferndm ...............~ a w gi m that above, p d d a r l y , lateral body scales
homogeneous ..........................................
. w~inrc8
49
b Dewlap red ta pU-red, with 5llaLsion of
black pigment centrallg; gmgetal scale8 88a Enlarged dorsal scales abruptly digsrmt
white; males, an tfie average, lagar than in she from much emaller lateral body
females, w both about the same size scales; dewlap orange-yellow .................
......................N-o ampC2equla~wn
5-8 h o r i m n ~ms of lorpala; hemipeais b Ed ed doreal scales ade into smaller
unilobate; Pacific veraant of northern l a b ~ b a d ~g c d m 2"
(fig. 7.h dewlap color
Central h n m i ~ a ...........
Nonops a e ~ m 0 2 different from that a t w e ...................... SB
b 8-10 horimntal row8 d lore&; hemipRnie BBa Dewlap dirty white, pale ;yellow or pale
bilobate; endemic on Idas de la Bahfa, my; ratio tail length J SVL 1.9 or lesa 80 ..
Honduras ..........................................
36
 b Dewlap color &@sent; ratio tail l a b / tively few gwgetel 8cales
.................................
..............
SVL gmatlerth 1.8 .......,...................
48 Norop% w m t M
4th Dewlap extencling ~KIabout lml of d a e 49a Ventpals perfectly smooth, flat and
......-. ..................... Nowpa &H Jaentrb disthotlv imbricate (Fa.276e): 2 medial
b Dewlap ~~ ..........................
or to level of d a e
wall onto &e& pest&-
41
domd d e towe &&d; canthus ro&r&
iia dietino$ supraaculara distinctly keeled
[ d f e d s f proup) ..,.,.......,..*...................M)
4la DwIap pale yaUm with purple gwgetd b Combination OF characters different h m
amles; 6-7 e d e s between second castbale that above ........................-.,,,....... BiB
..................................... Nonopa Ikmukf
BOa Dewlap of maim orimpred w i th a large
b Dewlap dirty wMke or pale gray with gor- central 'blue blotch; dewlap of femde~well
getd s d e a of 3ame color; 7-10 males FH4
Ween second wthah .....N m p C~U~YMO developed, ydow with a lar mtrd blue
4% Ventrda 8trwgly keeled; B or more rows of
~
Mot& SVL to 70 mm (m eg) or $5 mm
(Omlee) ..................
P
Nomp&hnnwyeri
dist,jnctly e d w g ~ dd m d s d e s ..........-88 b D m k p color differat; ftsmales without a
b VmtFal~smooth or oaly U t l y keel& 6-8 well developed dewlap; SYL to 60 mm 61 ..
. .
.................*... . ....
rows of dietipEtIy enlarpd &areala d s
............ 48 6la Parietal ridges usually we11 developed, a
&&in&parietal d e p d o n present; males
48a Ratio tail length / SVL 2,8 or m b r ; 4 usuallg with @lightly enlarged poetand
a d - batween recond canthala; B raws of scales ......................
.,& ....................
l2
distinctly enlarged dorsal a d e s ; about 29 b Parietal ridges absent to only sligbty
ventral8 in one Bead langth ............ developed, parietal deprea&ionabsent or
...........................
Nonops did- only slightly developed; males usually
b Ratio tail I@h / SVL s m & ~ thm 2.4 6- without enlarged postand ecales ...........I
9 scales between second cmithds; 10-23 6 h 6-8~ c d e br t w % ~ sscond n canthala; snout
Emody B-1&1 m a of diskinctly enlarged aeales midticarinate; 52-62 dorsal r&s
dbslsal acales; 15-37ventrals in one h a d
le* .........*...
............*............................ 44 between levels d &Ila and gnlin, 7-9
(rarely 63 horbontd r a m of laraab;
44a U d y fewer than 27 dared$ i n me head clawlap purple . ... N pum1cmkaSs
~ ~
length (17-28,mean Z4); no nuohd mt b 10-11 scalsa between second canthala;
with d a g e d 8 & ~ ...Namp O ~ # S U ~ ~ & @ snout scale0 ununiearinatq 48-64 dbrsal
b Usually more than 27 doreale in one head e d e s between levels of axilla and witr;8.
length 12149, mean 50); a n u W crest 7 ~ ~ n traws s lof lomals; dewlap KW
with enlarged wales u s d y m t l t 45 ....
with purple spot ..........
Nomps p#oknabs
4Ba 1-2 males between supraorbital semi- 85a Sealea of aupraorbital a e m b h l ~
circles; 22-26 lamelhe beneath 4. toe; SVL ly smaller than wake bettween d c i r c l e s ;
to 4T mm ..................Nompe w b l e p d s dawIap yeUowlsh orange with a red central
b 0-1 scales between supraorbital semi- blot& ................No- pdai&aW
chdes; 23-50 lme11ae beneath 4, toe; SVL b Ekdw of aupraorbitd remicirole~about
k~ 83 gam .........................Nonlps iraguei equal in size to, or lasger than s ~ l e be-
tween semicimlea; dewlap color different
s
43a Ventrda perfectly smooth ..-............. 41 a..
....................
.........................................
54
b Ventral8 faintly beled .......,........*.,,,..... a 64a Subocularm and supralabids uarwll in
47a M d i a l dm& scales Warm in he, with-
out inLre& m a l l scales
antset; mually 3 4 f&t pale oblique
...................
on flank (especially in males); ratio shank
LI
......................,
Namp hdemphakidmba length/ head length 1.04-1.18;dewlap pur-
b Small scales h larly interspersed pls .............................. N m p mbmensi~

......................
2
among the enlarg medial dorsal seatea
....... Nomps m m l h
b Usually one complete scale row wpmathg
~lubocularaand supralabial; no faint pde
obli ue lines on flank; ratip shank length !
4Ba Hemipertie wit$ one asdcata-side pMEe58; heel length variable; dsnlap color
dewlap af adult malea of moderate dm
with relahiely numemu gorgecal scales
variable .................,................................
(I&

.....................................
No-a d n t h u a 56s Nasal and postrostra1 males f u d ; 6 or
fewer medial rowa of dorsal e o a l ~sli t
b Hewiper& with two asulateeide pmeas-
M* dewlap of adult mdm small, with rela- enkarged (dlstinotly larger than $3
 Combleatian d &~~a&ersWerent &am Wia 2-4 m s of elightly enlarged xn- d d
that above ...........................................
89 d m ; males with enl- poatanal r c d e ~
...........................
..... .".....,..
78 ...................
8-10 T C ) d ~ d o d scdw digtiaotly en-
larged aad abnrpkly larger than much b W~thout rows d enlarged medial dmd
emaller leterala; mdea dhout enlermd scales; malea dthaut edm& paekml
postanal d e s ; dewlap np"* xed @call313 ....,......,....,..,..
79
--*..*.I3* .................
..................................... msP@PYtm-3-
-flu..MMU.

7th 2.3 males between supraorbital mmi-

4% rows of dored miles s1'itly edmgd, cirdm; dewlap o f n m b a very large (peer-
usually grad@ into ~rnalllateral scales; ior bedion at midkvanter).),Qran
males with eakwpd poetanal m a k or not;
dewlap color different TO
with ge~lowgo-1
.............................
gin
sales and a
.........................
N m p e fiFttrwnei8
2iZ!
leg@ ( 4 % ~ of adpm~edhindlbb b 1-2 *lee between supraorbital eemi-
~ X L.........................
bepad eye),
at least to oenbr of eye* uwalb
....ad..ad.....
7l
UPD~~%&dewlap of IlId@aS&
b d i m at lavel of chest), pink with white
gorgetal scales and orang@ margin
8horBhgg~rd (4th toe of adpressed
hindlknb does lrat reach to eye, uer- not
........,..............,......Na~opsptmthmrsa
bmymd ear op5ning) ................ Ttl 711a Dewla of mdm sp3all (pwrterior iaaertion
.*.,.*,a

ebnt Re1 or shoulder), rust orange pa-

Ventrala disanctly keeled and aliphtly ding to yellow on frae margin .....................
imbdmte .......................... .........
.....*. 42 ..........................................
M i e alkra
Ventads usually m o ~ t b and non-imbriata b Dewlap d male&larger f tc?riorituserticia
...........................................................
73 %butlevel or c b t ar Gmd), color dif-
An 4 a C e d isboulder gpot resent ......... ferent ................... '..*,., ................. 80
.....................-..Namp. o a & m l r p r r h r * 808 Dewlap pink to giakah-brown ..................
NPcmellabd shoulder apot ,,..,,.,.,..,..,
.................................
NOFISPII
ynlridmta
,,..,+),., ., ..........................N b c e p ~~~~~
b Dewlap color &iffrentfrom that above 81
2 elongate, overlapping suprdliariee:SVL Sla 2 wales between s u p t ~ o ~ b i t a 1 ~ ~ m r c l e ~
Co 57 rmn;dewlap orange with a d a r k dewlap of mdes orange with red mar*
h a 1 portion or dirty white witb 8n, mange g;orgetal scalee yellow; SW to 47 ram
basal portion .................
Nomp~ po&kp& .........................................
Noncps wart
1 eloeggte supedliarsf* SVL to 48 mm;
dewlap color vaiiable ,.............................
74
b 1 mle beWwn ~upraorbitdtalmiihlea;
dewlap of males red with orange qmt at
of ad*& hindlimb ma- rrntwtier mat*, prptal m i l e e wMW
4th
center of eye; dewlap maage with a darker
Co
SVL to 64 rmn .............
Noroprs hm@mi
barral portion .............
Nolrrpe &gwdi
4fi of adpte~sdhinumb pesche%
beyond eye; dewlap color different . 75 . ,-.. Meirl'teading
Dewlap dirty whits with a b a d oran* I
T m 199Pb, BAEBOUR1982, 19M, H.S m &

yellow Bv L
emtern an-)
or (po ulatione in e t r a l and
wlap d m s t complete-
ly mange-yeow; fernalee with W 9 d m
IEmmm 1954 STUART 1965, E m m ~
S~YAOE& TALBOT
1959,
M ~ & RWILSON 1871, E. BVILL~MS 1076,
1$78,G m P S~AV~QF: lS87,
ZM2, &WE & Crmm 1989, E. WILLWB 1989,
B& in one head IengthNompu J W h a e
1 M c C w et al. 1992, 1945a,b, ~RWEMA&
Dewlap mihndy dirty white, no b a d -1 1994, Kljrm~l W $ a . ~ lWDa,
, K ~ W Rb
orrurge-yellow epot, fa& with 49-69 M c C w 1988, KOH~~R & O B Elm ~, ~
d0rd3 illom head length Hoopie mIdu ... $kvm & GWR 1998, K ~ H ~ EetRal. 19B,
Body penlsh in fife (inpmsematim gel- KCE&LHI& MCCRANIE2001, KOHLERet al. MQk,
low* wth 11u-u~ mall dark p m h - MCCRAN~E & F?~HLEB 31001, M c C w et el.
tions,~esultingia a f h a spckled appear- 2000,0001, NICH~WON et &I,20Q1 Imorphology,
ance);venter usually with a Ucbnvw re- ayetanaticsl;SIEXTDN et nl, 1971,F f i c 1979a,
~ b,
ticulum; dewlap wdt%Pmly orange ............ Frrcsrr et aL lf37@,hTCEJ 3982 ( . 8 o o l *lo@);
~~
.........................,,........
Norrrps t m p e ~ t e r d F~A~CHENDMQER
bandry and b&g).
Q W m m m 1997 (captivehue-
BbtEg in lifh gray or brown (in pmsemtiilre,
vulabl. .
with no Elns speckled apmum);dewla
....*..*.......*..... .
............K
All four h w n speciea of basilisk are
found in Centrd America Earlier claims
that the di~tribution of the Ecuador
basilisk U3a8iEiscus galeritus) reached as
far ae Costa Rioa ( M1989) G proved ta be
erroneous, probably d u ~to misidentlfica-
tion of a juvenile B, brasilbcw (J.M.SAVAGE
per$. eomm. 1996). However, this species
haa been colleded in the Darien region of
Panama (R. pers, c a m . 2000).
Basilish, with their head and dorsal
ma,which give them a truly dragon-like
appemanm, ate among the mmt impres-
sive and bizarre Lizards of Central America.
Their ability to run on their hind legs even
across rapidly flowing wat~rhas earned
them the l d name " I g u m Jesucrista*.
T w o important adaptations that allow
these animals to walk on the w~ter'ssur-
face me broad Mnges of ekin on the toes,
which dietribute their body weight wer a
larger area and the relatively heavy tail,
which act8 as a counter-balanee to the
u p r a i d body.These attractive lizada are Fig. 286. Adult male of Elasiliscw uittatw (Iala
found in a number of habitats, wherever de UtiIa, Honduma)
water in the form of & e m , ponds, or
swamps is present, and are quite numer-
ous in many areas. On the Idae de la
Bahia, Honduras,BwiZiseue vitdatus is vir- Female baeilisrka produw thrw to seven
tually ubiquitous. B a d i s b f e d on imects, clutche~of eggs per seaBon, each with 418
epiders, and emall vertebrates, as wel as eggs, which hatch in 2-3 months ( K 0 ~ n . x ~
the occasional bit of vegetable matter, 1993b3.
Fig. 287. Juvenile Basiliscus basiliscus Fig. 290. Adult male ofBosiliacus basiliscus
(Manuel Antauio, Puntarenas, Coda R i d . (Manuel Antonio, Puntarenas, Coeta Rica).

Fig. 288, Adult male of BQsiliscus phrnifi.on6

(near Pueblo Wiso, Jino&ga, Nicaragua).

Fig.289. Adult male of Basiliscus galeritus

(Ecuador)
 Key to ~ M t s c u s r
1 a Vmkd acadmkeeted;1-2m ~ t &elthin
d
con&& d t h i&dabiale (Mg. 2gh); h a d
ereet of duIt males WIe, +ahgal$r in
cmtline ..........,..,., ......
hm'tkwnit-
b VenhL e d e s l mmoth; 3-4 mental shield8
aa each slide in omtact with MWbiaIs
282h); Bead m t of adult malera dif-
f ~ ~ B b Q V e . r , .............................
P
B P A series of enlarged, r a i d middoreal
scales,rngdatly separated &Im each other Fig. B!2,Arrangement of chin Wlds (oran@.
by 2-4 emdl malea (Fig.2Sl; head creet of
ddt males &glel f l W y t J d b ~ e dat
h e ; so d o r d 5n ..BattBinem # W a t t r s
b Middomum wit% etigktly enlarged s&s
she* ns *e of t.egplar spachf$
b e d m a t ofadult mdeg Merent; a b e
dm~d En pwmt ...............................
3
ff a h u n d mlar grew, on8 or two--I
nal m a of pale guts along ides of b&y; Fig. 298;-lidid&raal ecaleta in Basilkus galeIdtua
head ereat of adult male? oomprisabgM a
lobs; drin region af juv~nnead o r m
.B&Ib%@ plwwiffrme
.*.,.,......*.,...,...~~.
b Gsomd color b w n ; c0llr;inuouspale long$-
tvdind sttri along eidw of bodg; head
B"
mt uf adu t mdes singlee,offen ex&ndbg
ae a thin process pb9terirnly; &a ne&ionof'
jmteniIes with distiinct longitudinal dmk
ekeah ..................
B ~ B ~ Z ~ W
basb#mvrl

Further Reading Corydaphaw& I

MATUAANA19162. L m o 1988. K O m 1989e.
199Qb.
The helmeted iguanas are among the rela-
tives of the basilisks and their distribution
is restricted to Mexico and Central
America,All helmeted iguanas have a later-
ally flattened body and delicate proporti.
om. Even at eingle locality, the coloration
of the C. crietatus em v m from dark
brown to olive green and yell& with irreg-
ular blackish brown spots or crossbands.
The ventral lside ia pale brown in color with
a few dark spots. Males are easily diatin-
guished frolm the females by the hemipenid
bulge at the base of the tail, larger helmets
end aomewhat larger heads.
The helmeted iguana is a peaceful tree-
dwelIer that spends moat of its time in par-
tial shade, lying in wait for its predomi-
nantly insect prey. Analysis of stomach
Fig;.291.Shape of h e h e t in Bes2iaisca.a. eontents reveals that C, cristatus in the
wild rarely, if ever, eat during the day
124
Fig.294. Corytophunes percarinatus p
( ~ Imposible
1 National Park, Ahuachaph, El
Salvador)

L
Fig. 296. Corytophanes hernan&,sii (El Omte
region, Chiapas, Mexico).

(AND~EWS19791, Helmeted iguanas pro-

duce 4-11 eggs per clutch, each weighing 2-
4 g and m~aaadng10-16 x 18-30 nun
(REAM 1965, BOCK1987). Depending on
tempnature, the egg& will hatch after an
incubation period af 60-155 days ~KCIHLER
1999b1,Hatchling C. criatatus have a total
length of57-76m and a SVL of 25-30 mm
IREAM 1966,K~HLER et d.1494).
 l W m dmhk
~#.B~F*~PL~I"B$~, r
I
B ~ m M l b l I n W m W ~ 4
riaJlaWdhfbPerrtu
Fig.897. Corytaphaneslcristatus (Piw, Bonito,
Aflantida, Honduras Nationalpark).

Key to Corytophanee
1 a Crest of helmet continues without inter-
ruption into the dorsal crest ..................B
b CFest of helmet not continuous with dorsal
crest ,,,..,..,..Coiytophanes hemn$e&i
3 a Helmat remcted to bony skull p r m ~ a ; a. C. cristetus
scales on dorsal surface of head keeled or
..
at least clearly lined ..................... ,........
................Corytopiraneep e d n a t u s
b Helmet extends pmt bony skull proceBa; F
ig,298, $hap of
scales on domd surface of head ~mooth head (lateral vim1 in
..........................
Corybophanescdeaatus the spcierr of
Carytophsanes.
Further Reading c C.hemmndesii
LANG 1989,K~HLEB
et al. 1994,K W R 1998b.
L .
C#enosarsm
Depending upon e e s , spiny-tded igwm
,' as can reach a total length of anywhere
fhm 26 to over 100 cm.These stPiking ig-
uanas are found in the lowlands of Mexim
and Central America fiom the south-
eastern Baja Califonria and the southern
hnora in waertm Mexico, also more or
lem above the Tropic d Cancer in eastern
Mexico as fw as Fanama, and on certain
i~landsin the e&em Fadic, the western
Caribbean and the Sea of Cortez.
Uf the larger species (maximum total
lengkh over 100 cm), C. cacanthum and C.
pectdnata rea& only the north of Central
America in the area of the I s h u e of
Tehuantepec, Mexico, where= C.sirnilis ie
distributed a11 across the Central
American l a d bridge. Thew three ~pecies
art, predominantly gr9und dwellers, but
they will alm climb in t r s a The aaa21er
dmo~aus~peeiee,as a rule, ma trmdwel-
lecrs, men thmaltened, they will retreat
into knotholes, and block the entrance with
Ohsir o piny tails. The bauWhI1y colored
Yucatan Spiny-tailed Iguana (Ctenosaura
defenaor)i~ a gmuad dweller in the rocky
terrain of the northem YucatAn hnimsula.
Limeatone erevices eerve as hiding places.
All apiny~t;ailediguanas lay egg0, with field work p m d e d evidence that it is
clutch size5 ranging from 5-15 (mall spe-
ties) and 20-88 IC. dmilis), Suitable
neating s i g h are open, sun uposeii
areas where up to five fernales will 4
A
uw a oommon ay~temof holw to
1e.y their agga ( F r r c &
~ IFAc~E~TH-
J o m 19831. Three nest tunnels
examined by ~ E & HACKFORTH
I JONES
(1983)were 11-22 m long and reached 20-
36 cm under the surface of" the
ground. The e m require an incubation of
approximately three montha (KOHLER ~
1%93a), The hatchlings ofmosrt epiny-tailed
iguana species are predominantly green in
color with green m&.bgs. One exception
in this regard is the Utila Iguana IC.bake-
ri), whose HatcYJing~sare grey-brown with
virtually no marltinge, possibly due to the
habitat (harder mnea of mangrove swamp)
in which the hatahhg~of this species are
found. Spiny~taiIediguanm reach eexual
maturity at two to thme years of age.
Based on resewch of the geographic v9tia-
bility d the spiny-tailed iguana formerly
designated as Ctetsosatsm pabaris,
Buc- & Axmcu (1997)have deamibsd
the Honduran populations as a separate
species, C. melanesterna. Thus C. palearis
is re~trictedin its distribution to the
Motagua Basin of Guatemala. Similmly,
the populations of whrtt wm formerly
known BB C. guinguecarimta have been
siplit into tw6 species. Because of the hoIo-
type's identity, the name quinquecarinata
has to apply to the Nicaraguan and Costa
Rican populations and the Mtsican popu-
lations received a new name, 12,ommana
( E ~ ~ u &tN KOHLER 2001, K ~ H L E R&
HAssm 2001). Recently, the distributional
range of the three large cknmaur species
CC. acanthutui, C. peetinatca, and C. sirnilis)
in the area of the Isthnzu~of ' h h u a n t e p
and in the Central Depression of Chi-apas
wae studied (K~HLER & MENDOZA
maau13criptI.Contrary to earlier views that
amsidered the ctenoeaur species in the
Central Dspmsaion of Chiapas to be C.pee-
tinata (A~VAREZ DEL TURO 1960, 1983;
J O ~ S O1989,
N 1990, STAFFORD19901,our
actual13 C.acanthura.
~ ~ ; ~ ~ f
C & A O S Oma~fbuirz
~~ (BEAT 18021, Wn". C t a - r i i I(kLvidudia IC0mm-t & KUWMER
ZOQI,, London (9: 2x6 t lotali& rmg,oam. l a SaItmandra 80:1 W type l d t y : I km
8 ~ ~ b 8 1 6 m m . h n g % ~ v i b b s s n v ~ a n tS La Fan, 760 m elevatian, 1491f3W,8T040W,
from eenbd Tamaulfpas Ca rroutbm La Pse,Hondurm. 911L to 170 mm,Dlsjunet
Y~~acrua, M d m ; dm ia. the Central populatians &om erarotern Guatemala b
Depression of Chiapq M m b , asa 1-1 ta eastern El Sdvr\dst and eoutbe~tern
lOQO m elmatian. Handurae ta the CotnqmguaV a e y in mtlth-
central Honduras;ST, to 1010 m elevation.
Cteaosuum stl~dschmfdbiK O H ~ Rl-lia,
Bslmzandra 81: 6; type localits: 70 kno E Creno~ammekrnostam Em- L Amm&
Ihcamxap m road ta C h M vCamphe, 1897 189;type I d * P km 8
M&w, SVL te 170 m.Southern Cnrpsche, lae7,
Coplee O D TYm,Hondmae. 8VL to 310
enkral,
Mdm mm. V d e de Ma w a n and Cayw CocKnos,
Hand=&, sea level tu 260 m e1watiw.t-
m-um hkd S-ER 1W1,PPoG US,
Natd. Mw. Sk 46T; type localiw. Irrla ds Utih Ctetto~arsra oamcana K~IXGER 8t HASBUN
I e h de la Bahia,Wmdum- SVL to 300 mm. 2001, Senckenbergiana bialogica 81: BBO;
Endemic to Isla de UtiPa, Mas ds la Wa, type lmdity: 'ikhnantepec, Oaaaca, Mexim,
Hordgrm, eea level to 90 m elenrakio~ 8VL to 170 mm. Rsatricted to the Pacific mr-
sant of the Isthmus af Tehuarttepec, Ow-
Ctemxwum &@am (COFE1866),Proc. k d . Mdw;sea Imal to 250 m elevation.
Mat. Sci. PhiiadeIphria I& 126;lype hcaNQ~
Yacatbn, Ibkdw. SWL to 146 NoFtbm C t m u w r a oecdirhina RE Q ~ I R O I1587a, I
P on d the %cat& Peninsula,Made4 sea
wd to 100 m elwatien,
Copeia 1W7: 898; type looality: approle. 4.8
lun W Roa* m ath tn tamers Bay, Ma de
Pa
ha*, I b d. Bahl., HOnd-. m to
270 mm Iela de Reat& and its satellite
kllanda I&anta E1ena, Barbareta, Big Pigeon
Cay), I s b do la B a a , Honduras, 8ea level b
100 m elevation.
C&~asaumpale& E ! ~ % ~ E G E R 18941, Proc.
US.Natl. Mw,81: 381, type locality: Gualan,
Gnabmrda. SVL to 300 mm. V d e ds Fkfo
Motagua, Guatsmala, 150-900rn elevation.

#erpeblopici Mexima: 42;; type l o c a l i ~

Fig.900. Ctenasaura acanthura (Monte Bonito,
Chiapw, Mexico).
unknm. SVL kv 365 mm, Southern Sinaloa,
Mdco, along the P a d e versant to western
Chiapas, w s level to 2000 m elevatian.
C t e m w a quinqwcori~ata(WY
1M2),
Zool. bfbc. 1842: 69; type locality:
"Deme~fvb?"[based on morpholagical and
molecular genetic data, NAssm L KOHLIW
2001 dlmtad the C. uui'nawmrinat~hala-
type to Wiwagt.mn and ~ k t &can a poptila-
ti-], SVL to 200 mm. Imlated populations
in Niwqgua and northwwtsm CoM. Rica,
eea level to 260 m elevatim.
C t e ~ u m csimUiar (GRAP18311, Bynopah
In G R ~Cumers ,
om 9: 38; typa l d t y : un-
mnl. I a t h r u of
IEshumkprac (central Tabatmu an Carib&
verrmaat and central Uaxaca on h d i c ver-
mt),Maxieo, acraw all of Central h r i c a
.(excepttrighlan& and very humid rqgion5)to
Fan-, sea level to JS20 m elevation.

Fig. 301.CTtenoseura aE&d8chmi&i (mnthmn

Campde, M h ) ,
Fig,502.Cdenogaura bakeri (Isla ds Utila, Fig. 306.Ctenosaura oazmana (Oaxaca,
Honduraal. Photo: W.Steiger Mexico),

Fig. 303. Ctenosaura defernor (near Fig.au.4. r;temmura oedirhina uua ae

Telchaquillo, YucatBn, Mexiclo). bat&+ Honduras).

II
Fig. 304. Ctenonaum flavidomlis (near La Fig. 508. Cdenoscsum palearis (upper Motagua
Paz,Honduras). Valley, Guatemala). Phota: J. A. Campbell

Fig. 306.Juvenile of Ctenosaura melanosterna. Fig.309. Ctenorraura pectinata EOaxaca,

Mexico).
Key to Ctenosaura
1 a W l at the point of xnaximurn circumfer-
ence wider than high (width-height ratio
-
1.66 1.7%);ratio of tail length/SVL less
than 1.5;maximum SVL to 200 mm;distal
dark tail band0 not tnuch longer than
proximal ones ..................................... 2
b Tail at the point of maximum circumfer-
ence appraximately as wide as high
(width-heightratio 0.90-1.11); ratio of tail
length/SVL greater than 1.6; maximum
SVL greater than 250 mm;posterior dark
tail bands twice as long as anterior ones 6
rlg. a l l . brerwuraura quinquecarinata (near
2 a Only about 213 of tail covered with whorls Teustepe, Boaco, Nicaragua). Photo: F.Schmidt
of enlarged spinoua scales; parietal eye
clearly visible with naked eye; keels at the
base of the 3rd toe not fused together .....3
b Entire tail covered with whorle of en-
larged spinous scales; parietal eye barely
vieible with naked eye; keels at the base of
the ard toe f u d together .......................6
3a Adult males with middorsal crest spines
<2 mm high; ueua gaps prerent along
%
the crest row in nu a1 and anterior tho-
racic region; 2 or 4 poatmentds; doreal
coloration ulsually with some yellow ele-
ments ............
Ctemnaurrr ~ v i d o l u c l b
b Adult males with middorsal creat spines I
llgually 22 mrn high; nuchal and thbacic
creste continaoue; 2 postmentals (Fig.
314a): absence of yenow elemenb in dor-
I
r BE. u u . bcanvaoud ~ i m i f i(Isla
s de Utila,
eal coloration .......................................... 4 Honduras).

Fig.310. Dorsal view of anterior tail. Fig. 313. Mid-doreal scale&

 4 r TaiJ lsss spinnus (paramedian mud91
aroaltlr eplnwe an. pMximal 3.7 imsm
! v a h 451 mudd wh0~1eik ueuall body
I
I
pattern of a kin& pale micidc~r~band,
arhirrh may be fksgmentd into Iwge
J
I
blot ah^^; STFL to 179 mm in males, 124
mm i female8 ,,.Wmemra og.rocams
b ls;il mare spbus Ipqamdim aaudd
~cal@ spiaw on 419 Em~an I. -vrawm& b, cltenos~um~lrni#~
value 8.41 caudd whorls); ueuall no
brsd p& nxidet~~gal b d ddmk S& to pi^ 3 ~m
. a L t i a n (pmtmtnla o m ) .
235 a m in males, 145 mm. in fm&s
..............C-afsm quirop-WD
5 a 27-28 tail whorls; ratig af len&BVL
0.7%O088;m a of medid dorsals uniform
aad ig~&&qto pelvia Wdn; parf~tals
mfmd wilih d parietal ep; ad^
the &5t OS htempace8 betworen C a d
sphotl~without complete m a of mall flat
I scdae; SVL to 17O nun
...................
&-urn aLWWt3
.......................
b 23-24;tail whorls: tail I m m
@f
0.66-d.74; row d enlarged medial dads

I
z ar and reduaed; p~~
tal eye not di-abIei
no mrnpI* mn of e n l p lpilqr
~a firat&&t t wbr~;
mduoed,
W ~

sm Q
a,
J GYBWU~ m i ~ i s b. qujng-m
Fig. 331. D o d view of anterior tail Cmk-
:I 14am .............
,. C b s m r defineor
~ ealay m h onage)
6 a 3147 dorsal apirw, dmlap rrum ax lw
I 8b.Onglyd ~ ~ I ~ p e d .............................+..
7
b $6-106 dmQ epinw; nn dewk rather
j t s t a tFaawerae &M mat48 *B &oat 9 #,

*
7 a Two complete raws of m a l l intsmdsty
&H between whorls of enlarge$"pinous
s dietaIIy Estmthg aRar T ta the
tail whor16); mwkinga and colaration
fent
miwlpl:areu-brown
dorr)sb*t
of dorsal @pineseonfami ta that af aqa-
D.
mbakserf
I . Dn-rn ~ a c t r t n ~ CDmwurar &#/s

b ~ ) a eciP~Bte row of rmltll interca~my e, . a t i o n on M sdaea of shank.

SL~m
ecdrss betiwe= dl wbrla of enlarged
spinom aides (MB rows of hhrcalaig
s d e a m8en.k b e i m w mt tlr&mP 8
whorl~-hHome s p ~ h ~~
and

.-
mlmtian sf d a d spmm do nut wlnforrn
to that d divant d o r d &: e n a d r w .
wide gcey ta &lackm a bmda m'dmum,
sepwt9d by raws of amall pals spats 8 .....
% a Vmtral area p-tzy Ma& in eolor-
................,.........'CM-..................
a% 9.12 supn&ibi& .,..,,
pa-
b Venttsl m a brewn in cdoration; 840
mpralabials C # i w m m l ~ ~ ~ ~ h m k ~
9. a Enlarged heavily beled scdes on the dm-
~d8urfam of&& (FYp. 916~);matat -t
older individda blunt and rounded aff
%; [
@%g. 3175;9- ;h &@ aneg Fig,317, In older individuals of C, wdi&i~~o
betaem tihe whrrrla, only Dhe mout is blunt aad r o d & oft
 om mmpIeh ruw Qf mall iatttm- tail crest; in area ke$wsen?3Mand tail
..,..I...,.....,. ~ ~ o osd5rh~tra
u w who&, twa mmpleb m s of m a n inter-
b Scalcur on the dorsal mu%w o f ~ h m knot
W g e d aad only sli&W h e l d ; snout
...,. ...
dery wales ., ,.. C ~ O % Q -aimilk
b No dark oramkids on dmpmi d a d
not blmt and rounded 0% 4%fmm& mest btersupted inp l v i c r p m mtspkia
ps;intharwbtwmtha4 and6 thearea beheen the 3 and 5 tail
bail wEUlFla, at l w t tvPo complete m uf whorIFE, Q mrnpleb m s of mall btw-
dinbmdaq s d w ,,.,, ..., .......,,.,.
,...10 calEuy s d e c ."....G t t m 0 1 c r r s~~0
IOa sadel3 of tail WM vasy spiky (angle of
keeL greater SO degrees); dorsal
m&t w d y intmmpte8 in plPi6 mghq
in so%e iapwxgrimIrow8 Of 1ca1ea bdwwn
t h e 6 d 8 tdwhorlareduoedtame
...,
crompleta mw Qm - OE&-
b Scales af. tail whorls moderately
langls of kwh lmer *an 20 degl-sl wj#
k681~running d i m a l l y (Fig. 818ali
dorsd c e t in pelvic &on h b m p t e d or
nu$ dwaye at l& M complete mwcr of
wdw between tail whmls ...............,...I1
i l a h e r d dark i ~ a m h n on
6 dofaam, whith
w d l y have a pale center dang the doP-
actmedial Une; h a l crest cont=inunuswith

Fig,318, Dorsal view of anterim tail.

132
 -
Fig. 319, Erayalioidee hetciwlepis (Lita San Lorenzo, Esmeraldas, Ecuador). Photo: K.-H. Jungfer

$nyddaidesL t e d p i e (J~OCOUR'P1874, Ann.

Sci. Hat. Paris (6) 19: 1; tSrpe localitJP.Vem-
- pa, Panama. ENL to 198 mm. Panama to
From the primarily South American genus northwestern Eouador, -1 rn elrntio11
~nyatioid;.s,only-one species, E. &tern-
lepis, reaches as far as southem Central -her Read-
America. So fw, the existence of this spe- NICHOLSON
1988,LIPS 199%
cies has only been establish4 at a few
localities in the Panamanian provinces of
Bocas del Tom, Colbn, CoclB, Panad, and
San Blas INrc~otso~ 1998, LIPS 1999,
KOKLERunpubl. data). The species of this
genus are it-and-wait predators in the
rain fore~tthat primarily feed on iasecb
and spiders, At night, these lizards will
sleep at a height of 9.5-1.6 m an tree
trunks or lianas, sometimes even in the
rmettea of the aerial roota of palm trees.
Iguana -
GS
The green iguana (Iguana iguana) is ce&
tainly the bast known l i z a d in Central
and South America. The preferred habitat
of this magnificent giant lizard is l o w h d
forest, near streams, rivera, or lakes.
Whereas adult iguanas spend a Iot of their
I
time in the treetops, juveniles are often
found closer to the jyound in bushes.
However,I have also observed adult green
iguanas basking in the vegetation along
the shoreline in Tortuguero, Costa Rim.
Green ilfuanasl are irnpreasive svpimmem
and direrB, they have demomtrahd by 1%. 3x0. i W m (near Pueblo wise,
leaping into the water h m as much as a Jinotega?Nicarama)b
10 m height when I approached within a
few metars of them. In the wild, gmen ig-
uanas comume an almmt exclwively vege-
tarian diet, consisting m a d y of leaves.

-
At the beginning of the dry season, the
female iguana will excavate a nesting site,
which,depending on how hard the ground
is, will be from 39 ta 200 cm long (RAND
1968, RAND & DUGAN1983). Clutch size
varies from 20 to 60 (seldom to 80) eggs,
primarily depending upon the gize of the
female. The young will hatch following an
incubation period of approximately three
montha, which usually correspands with
the oneet of the rainy sea~on.Thue, the --
hatchling &Urn= are ensured a ready Fig. 321. Iguana iguana rkinohpho (Isla de
supply of tender leaves as nourishment. utda,~ ~ ~ d ~ ~ ~ ~

Further Reading
& RAND 1982, KCIHLER19986.
BURGFLARDT
The caaquehead iguanas,which are found
only in Central America and Mexico, are
remarkably well adapted for life in the
trees, which they leave only to lay eggs.
Predictably, caaquehead iguanas are
=cult for the human observer to spot
amongst the green leaves. Several timea a
year, Laemnetus femalesl will produce a
clutch of 4-9 eggs, which hatch &er ontv
two month&at-&.incubation temperature
of 28-30 O C . Freshly hatched Laernanctua
Zongipea on average, have a SVL of 60 mm
and a tail length of about 179 mm, Fig. 322. Luernanctua Eongipee. Photo: F,Ride'

Lcasnaawctue longips WIEOIIMNN1834,

Reptflia and Batamhie. Biologia Cmhli-
Americana: 48; t a looalit~Jalapa [a
.ttpJ* Megico* 8% ta 160 mm. Ot3nl
V,- Mexico, to Niamgw, new sea
lmd to 1200 m elevation irr m& M e t and
rain ffirest, In Cantral America, tbe sub~pe-
ti& hmrsmptts I, &barref Botmwam 1877
bthmus d %bu&nt&pc to Nimgua) and
&, L waldensi Qb- I W b (n&hw$~tesn
Bm-1 DEm.
J L w ~ r r c t mmhssl
~u~ COPE 1864, Proc.A d .
Hat. EM.Philadel Ma IS: 178; 1 .d W
h i U b a Valley, bexico. Sn to190 rm.
Tamaulipae, Me&* the Atlantic w t
3
aa far ae northwestern ondma~i,mar srta
level to 1500 m elevatim L dry h t . Ia
F,"&peclE$$myi
zCgz Pig. 313. Locrnanetus s~rmtua(famale).
a rnccoyi P E R E Z - H ~ G ~LAVWT
cwutl!m v-1 mxw,
Ker to Luem~tetue Aocording ta the ober~atiomofC ~ ~ ~ S ( E I X
1 a Pbstsrlw of head wlCh epiky sEa3.a; dia- &td.(1986),the animals live in b u m & . )
tinct dvmd crest pteunt; d e a on dmsd which they smaoate in tkts floor dthe rain
eurface d asnout Im-ge and regular fore~t.Dudng the day, thw h d s will
. l t l t l l ? , I . . . . . U . l . . l * l%Brmtm
*,t~~m9~
eit modonlem in front of the enhances ta
b PDBtahr dhmd without rtpiky .scdw;no their burrows attentively watching thar
dorsal crest present: aclrp~son dmal s
w
fa= of mmt mall rmd k e g u k
....,...
"..** ...*.........
lhwmwwmb&pes
..,.. smun&g area, If they feeI threataned,
they d l disappear with Egh* 5peed
into their b u r n o , blocking the entrance
Further Raading with their epiky taila. PFedatons, ~ucha~
McCoy 1968,LANG1989,KBHLER199913. aaakeeI are thue prevented from f&g
their m y into the bmows, Morma~auras
a n n d h mpducee by l&ng e m .

Two speciear make up the genus

M o ~ u n c sM, : annularb in primon-
tan% mas on the h a m $ideof Emador
md Peru and M.groi, whicb ims distribu&d
aer narth as muthwn Central dmeriea.
They are diurnal g~nmd-dweUe~~ of
&tackybuild that Eve in the shady areas of
the rain fw&. Both aped= haw a wezy
h e t e r n ~ ~ omilation
w on the domil and
lateral edaesr, of ths tmdc, with lmge
conical scaIles scatkmd mang the granu-
Iar scale^. The tail ie equipped Mth re@-
larlp arrmgedwhorls made rrp af enlarged
spi& acalebs, which becoma in~eashg1y
1mallef didally and am bar@ &tingukth-
able at the tail tip, &for26n@cturaas;groi IEE
predoaslinaatly reddishbrown in color. The
W y has dark bmwn cmabands, whit$
reach to the midpoint of t haid# a d then
break hto amaU irregular ~p0tfl.
W the twelva speGie~of horned lizard that
am distributed across Mexico and the
USA, only the giant horned lizard
(Phrynosama a s h ) reaches as far as the
north of Central hmiw (REEVE 1952).
Their presence in Guatemala has not been

I
verified by any find8 in thki cantury. In
Daxaoa and Chiapas, the giant horned
bard is found in qarm forest aud rocky
semi-desert ( B m & M~NTANUCC~ 1998,
H~EH;ER per$, oba, 19833.Phsynosom mia
feeds on hects and spidem, with the larg- Fig. 3260 p h r ~ m m f~~~
i
e ~portion
t of their diet, as with all horned
l i d species, corlglisting of ants (MONTA-
~ I This is one of the egg-laying
C 19119).
spiwieis of the genw, with clutch aize ranp
ing from 15 to 24 egm, Young measuriag
32-37mm SlVE will hat& aRer an inenba-
Mom~~uccr
1998).
hiychrw gutturnus, the only bush mole
~pecieain Central b e r i c a , lives in the
N~T

i
Phrylaosanaa ash COPE1864, Proc.Aead. crown wi?bn of the lowland rain foreat
Sci. Philadelphia 1%: 178; type locality: and,apparently,only W W B in low popula-
Colima, MerpIm. SVL to 124 mm. C Q ~ , tion dedtiea, B-uae they move very
Mexico, along the P a d c mde of Mexico to
Chiam, me= sea led 760 in W l e , bush anolm are difllcult to deteot in
~ e v aridm dry forest the le@ tangle of the rain forest tnea On
the 9 af Max ROBERTS (1997)obgerved a
copulating pair of R gutturnsus about 2
M e r Rs9dimg metem up a tree in La Selva, cost^ R i a . A
F€mvE195% MmrANuccI 1987,1-, & female alpable eggs was
M o m o c r 1988.
eramind on the 24& of July (ma~ars

Fig, 326, Adult male of FoEychnca gutturnus-
lRfo Ban Juitn, Ncarslgua),
I species reproduce by laying egp whereas
Fig, 327.Adult fernale of Pblyc&m guttumeus
(Rlo San Jurtn, Nicxra@mj+
Further EIBadhg
TAYLQR
1856.
m h e dry regions of Mexico and Central
America, negt to apiny-tailed iguanas,
spiny biaarde w e the dominant igu-
group.The gems contains over 70 species,
with increasingly fewer varieties occur-
ring towards the south of Central America
(only three species occur south of
Guatemala). Spiny lizards are & m a 1
small to mid-sked lizardg, meet of which
are covered with e&rsmeIy epiky scaltx,
They feed on various mthropade, as well
as smaller vertebrates. Among the
Scelaporu~species are ground-dwellera
and 5articolo~species,as well as prinaari-
ly arboreal epeciee. Mmt of the Sceloporua
the species of the ~ ~ F ~ O Bgroup,
L ~ I F as well
rn S. sevifer (tarquestus group), are livs-
bearer@.
The spetri~~ of the uariabilis group are
ground dwellers in the lowlande and, in
some areas, occur in high deneities.
Frequently thma animale can be seen on
1
.tree stumps or rocks. The males, aa with
an aped88 of the genus, we highly tenitmi-
a1 and attempt tx, intimidata th& rival5
1
1
with pueh up-lilre movernenta of the upper I
My. AU species of this group lay eggs (2-7
e g e per clutch depending on species).
The systematic@of the speuw of the fbr-
US group have yet ta be 8atlsfacbriIy
resolved, as the morphologioal variation,
as well as geographic diMbution of cer-
tain species, L as yet unknown. Thus,
STUART(1971) claimed that; mnmpruisollls
between populatiom of Scelapom mala-
chiticus produce ul..) only a hopelem jum-
:ble d random variatiom". In hi8 generic
revieion, S MC1939a) ~ considered mala-
chiticus and smrcq&dInua aa subspeciee of
formoaus. Subsequently, both tma have
been designated a@full qmAm (STUART
1971, VUiLA et al. 1988, K ~ H L E K19901.
Also, the exact distribution of the various
apmiea d the firmosus group remains to
be well dwwented. S m et al. (2001)
slated that 8. aea~thirausraeges into El
Salvador, wherea~ Kt)- L HEMS
(20021, partly basejd on ~ d v bk m Eric N
Smith Ipem, comm. to P. Heimes), refermd
a31 malachite Sceloperus hEl Salvador to
S.mlwhitiom, and I have used the latter
name far a11 El Salvadoran populations
herein.
With a v e & i d distribution of 600-3800 m
above sea level, X malachiticue axld S.
smarcsgdintss are typical highland dwel-
ler~[MAEION & SEXTON1971, K O m a Fig. 328.Adult male dSce60poma gctadainus
1940). They are famd on t r w , fence (near Gu~nagampa,Ewrlinth, Guatemald.
posts, and on the roofs of b o w s I W R ~
1962e, ROBINSON1983). Usually onllg a
-1e male will -bit a treeswhich it
will defend From rivals I~IERT?GNS 1962~).
Aaimala found at higher eleratione tend
more towards a ternstrial lifeatyls
IRosnvso~1983). On Gena de la Muerte
(Cordillera de Talamanca, Cwta IEica,
80OQ m above sea level], S,mekachiticm
will inhabit natural stone walls and r d
ewered with momes and ferns.At thils de-
~atim, S.malachiticus i~active only when
t b BU shines. The air temperature mea-
aured 160 cm above the ground barely
reache6 to 20 "C despite the e m . These
lizards, t h e r b e , have derpted to optimal Fig.329. Seelopoms ~ n ~ m i(male;
s ; near
uae of t3u micmclimata They flatten their a n Cristdbal de lae Casas, Chiapaa, Mertiw).
bodies and turn vary: dark while basking
on mcke protectad h r n the wind. The
f a a h of S. molaclait&us p d u o e one
brood of 3-10 young per pear (~LWON & found on individual stone blocks or on
SEXTON 1971,K~HLER l990,1993d), while pilea of r&. This species lam 4-6eggs,
at lower elevations, birth o c c u more or which hatch afbr 60-63days at an incu-
lesa rergularly throughout the year. In bation temperature of 29 "C. The young
populations ebwe 2000 rn elevation, measure from 17.0-18.6 fnm SVL.
however, a h
te e is a highly m a ~ o n d
r mpm-
ductive cycle, wikh birth occuning prim- As the @olerepramntative of the Corqrdatue
arily in late January and February group in Central America, Scaioporus ser-
(MARION& SEXTON 1971). rifer reachesl Gu~temalaand Belize. This
species can be e m in great numbere at
The dniferua group is repmsented in the Mayan mhs in YucatBn, althaugh the
Central America by three epedes, all of ankale are quite shy and will flee at the
WWam ground dwellem in hat, dry ellghtmt dhturbmce. These attractive
envimmentsr, 1x1the Valle de Comayagua, Ezardsr bask an treecl, as well ae on roch
~ ,squarraomn is quite common- and brick walls, and will dieappear into
H a n d m ~8.
ly seen (EIS~NBERG & KCSEBR 1996). Th-e the nearest crack or opening when threat-
spiny l h d e in this area are generally en&.
 Smlopatrsa acurathinus B o c o u ~1873,
~ .Am.
W.Nat.,Zool. (5) 17(61: 2 4 locality:San
-tin, near VoMn de ME, Guatemala.
5VL to B9 mm. REifIc versant of Chiapas,
Mexico, and Guatemala, 400-1800 m eleva-
tim.
Scebporus carinaha H. S m l986b, h.
Biol. Soc. Wr&tin@m 49: 88; type locsSit .
near ~uti~rm ,
~hiapaa, SX
~s.iea
to 55 mm. Grijdva Valley of Chiapa~,Mexico,
and acljaaant G u a W , 500-1000m elevation.
SceEopoms chry~mtictusCOE% 1866, PMC-.
A d Nat. Sci. Philadelphia 18: 126 type
l d t y : Yucatlh, Melth. SVL to 60 mm.
YucaW Peninsula fkom central Belize and
no& Guatemala northwards in dry foreat,
,eah e 1 to 2OD m elevation.
dcelopoma coeumeEcse JONEB1921, Occ. Pap.
Mw*Zaol., Univ. W g m 1- 1;type locali-
ty: Ida Cozumel, Quhtana Rm, Mexico.SVL
to M mm+ Coastal &re8 of the northern
Yucatan P a h u l a , including the ornore
islanb Cozumel, Isla Conto5 and Ida
- Mqjents in drJl foreat and open rocky mas1
aea level to MI m elevation.
S c e b p ~ m&ardtaylml H. SMITHl936a,
Ermpetologi~~ 1: 6; type I d t y : Ixtepc (Sari
Gednimo], Uagaca, Mexico, SVL to 107 mm.
Pacific w t of the Isthmw of Tahuanta c
( ~ a x s c a ,~ e x i c o )in dry fmest, sea B
lvB K
760 m elevatioa
Sceloporua inbernaeaIi8 H . Smafiff &
BUMWLHEM1555* Hwpetolagica 11: 118; type
oealiQc La Gloria, Oaxaca, Mexico. SVL to
100 mm. Lo8 !Cuxtlas region in southernVera-
m z , M d o o , southward through -me
eastern Oaxaca to northwestam Chiapae, ma
level to 2000 m elevation in oak foreat.
%6elopom Eunaei Bocom 1873, Ann. EL.
Nab, Zool. (6) 17 (10):1;type l d t y Plateau
of Guetada. SVL to 95 mm. Mo-a
and Baja Verapaz, Guahmda, 150-1000 ele-
Bash
vatim in dry forest.
S c e l o p w lundelli H. Shnrrrr 1939a, Field
Mu. Fubl. Zool. Ser. H:66;type locality:
Cohune RidgeI 28 miles SE Benque Vie'a,
Belize. SVL to 100 mm.Yucatin P e n i d a ,
eea level to 300 rn elevation in dry forest,Two
subspeciei: Scelo ma d, drs&Ili (baseof the
Yucath fedinsU a) r and lsceloporus I. gaigeae
H.S m US9 (northern past of the YucatAn
hninsuaal.
Scebpoms malcu:hitktts Corn 1864, Roc.
haaa Nat. Sci. Philadelphia Is: 178; type
locality: "Arriba"PthiFS name apparently wae
applied loosely ta the Meaeta Central"accord-
ing to SAVAGE 1974:7711, Coda Rim. ;SSrL to 98
mm. El Salvador and Honduras acrose
Nicaragua and Goeta Riea to Panama, 800-
3800 rn e l d o n in pine and cloud fomt.
Sceloporus melanorhinus Bocou~r 1876a,
Ann. Sci. Nat. Zool., Paris, (6) 8, mt. 12: %;
P e locality: Iethmue of Tehnantepec,
axam, Mexico. SVL to 105 mm. Southern
Nayarit, Mexico, along the Pacific aide of
Mexico to the Isthmus of % h u a n t e ~
well m in the Rio W d v a Baeia, Guatemala,
ma level to 2000 m elevation in dry forest. In
Central America, the subpadee Sdopoms
m. mela~~)rkinua (Mexico) and 5,m. $twrti
S m 1948 (Guatamalal accnr.
Seelopoms gewifkr COPE 1868, PMc,h a d .
Nat. Sci. Philadel hia 18: 124; type localitg:
&
~ o c a a~, e x i ~ o . to 100 mm. ~ e r ais*
lated populations from eouthern Tamauliparr,
l
Mexico, as far aa CSuabmala and Beliae, as
well asl in the north of theYucath Peninsula,
60-2300 m elwation in dxy forest. In Central
America, the euhpecies &el0 ms a. ~errifir
Ptmineula) and ~n&Pw a prety-
g w H.S m 1942b (southweatam Guate-
rrlala) M.Jrmr.
Swloporus sinif-ua COPE 1870, Pmc.Amer.
Phil. 1Soc. 11: 159; type locality: N c ver-
srrnt of Isthmus of Tehuantepz, SVL to 71
mm. From w&am Qwmtrn, Mexico, do
the Pacific C& to eaetem Chiapae, sea lev
to 2080 m elevation in dr+yf m t .
3
liYg, 330.SceIoporwl carinattee (RanchoAl-,
Wapas, Meadmi).
 Fig.335, Scebport~alundelli gaigem ( C o k
:P,Hsimes QuintmaRoo, Merricol, Photo: J. C. h e

Fig.333. Scelopgrue edwardtqlori (Oaxaca, Fig. 337, SceEeporu~melanorhinus (Wuautla,

Mexico). Moreloe, Mexico). Phato: P,Heirnes

E
Fig. Sw, 1 ~ c a m p mrn-iaar ttiawmaco, Rg.938. SmZeporu~serrifir (Ruinas de
Verauuz, Mexico), Fhota: f! Heimes Mayaph, YcetAn, Mexico),
s c e c o ~ k g d i n BWOURT
w 1878,Ann.
344. Nat., h l . (63 17 (10): 1; type l d w :
Zuabmda. in the vicinitv of 801014,
htonieap&, anrl Quefd-. 8VL to 86
mm, Highlmda of Guatemala and Chiarm,
2000-4000 m elmatian in pine a d dapd
Forest.
Smlaporw srniiflci HARTWEQ & O ~ 1937,
R
Doc. P a p a MUI. Zool. Univ. Michip 368: 1;
type locality: Quiengola Mountain, about 6
miles NW af the town of Tehuante ee,
0-a, Mexico. SVL to 71 mm. P a d 3 o si%of
the Isthmus of Tehuantepec (Oaxaca,
Mexico), sea level to 600 m elevation in dry
r m- -t- .
---
Fig. 339.Sceloparus siniferus Inear Mixte-
bloporra~sguamrrewr BQCQUET1874,M i ~ s . quillav O-cav Me*).
SU. MW.,~ e p t ,am;: type loc~lity:~ n v i m m
de Guatemala et de 1*AntiguawGVolcBn
Antigua+Guatemda). SVL to 69 nun. From
weatern Chiapas, Mexico, dong the Pacific
side of Central Amexica into northwestem
COB^ Rim. aea Level to 1000 m devatim h
dry forset. '
h
E
Sc8Zoparua twnhnemia Cow 1886,
Am. Phil. Sac. &
%.
: m e locality:
3W:
I I

I
Guatemala SVL ta 80 mm: CGtral
of Cbiapas, Mexico, to central Gu~&B,
1200-2600 rn elevation in phe and doud
fmt.
Sceb F U teupmk
~ GONTEIP~R1890, Reptilia
and l%mchia. Biologia CsnW-Ame-
76; type localits: ?Ltapa>B h w , Mexico. SVL
70 m.Fmm m~a, Vermru and Fig,540, Soeloporus s;maragdinw (btw. Soloma
eastern Owacaj and Smta Eulalia, Guatemala). Photo: T. Bille
Guatemala as far as Bebe, sea b d to 1000
m elevation in aavanaa and dry foreat.
Sceloportls wriahilirr WIEQMANN 1834,
Herpetal 'ca Magicana: 5% type lucality:
Mexico. % 8 to 74 mm. From mas, USA
ong the Caribbean dde of Mexico into

f orth~~~ltern Coeta Ria,sea lewd to 1700 m

levation (to lBU0 m elevation a c c o d h ta
OHNRON 1BSg) id saoanna and d f g hrest. In
Central America, only &a nominate subape-
uea occutrr (of which dZaporun H.Wf!l!~1937
is a agnonp).

Fig.341. Sdoporus srnithi (near Mixtequilla,

Oaxaca, Mexico).
Fig. 542. S c e b p o w equamosw (Sanka Lucia,
Intihca, Handuras).

Fig. 344, Swlaporurr uarirmbilis /Caflan de

Elurnidem, Chiam, Mexiool.
ICey tar S ~ e l o p r t l s
1 P P o s ~ ~ m o rpocket
d present (Fig. $471
Cm*b3is jpwtrp) ................................
O
b Withont postfernoral pocket .....................
S
1 a Malee uniform pale belws; no c a M beny
pakkteq endemic to tha north ~f the Yuca-
tAn Pe&ula .....
S P B L ~ P ~~~
b Mdes d t h obvious varrtral marWp, em-
sirrkbg of two latge pwpleceloned belly
patches, ueudy with black berders ........8
8 a 49 or fmw dorsal sales posterior of Fig. W ~ E ~piew
Wof head in8ceE~ponag
head to baae d tad; 7-9 rows denhged r n ~ h . i t ~
domal rcdw in the pelvic m l s s in
the inguinal region relatively large
...........................
~ I d p o * ~ c&?qensL
ff
b MI ar mom dmd wales hmpaet&or of
head ta h e of tail; 10 or rnm mws of
e n b g d d-al wales in the pe1P.i~
acalrs in inguinal mgian reI&velg amall 4
4 a M u n d cakr wary bark, n e d y blackt \Kith
very obviw5 yellow w wite l*tudind
6.ttipes (Fig. &Ma!, wfLich are 2-2.6 ncalea
wilde; mf out; obwwsl dark di& bm
w dor~umand body aiderr;ksels of the dor-
sd scales form dbhd ffns m b a ; hatch-
h g with
......... .....................
~ pink colored tail
...................... Buetoporw unrdchf
b Chrrund d m grey to brown, with Leas
distinct pale longitudinal abipeg (Fig. Fig.W, Bomal ~ m of v head int3ceEoporus
348'b),WU are 1,B-2 d m Mde; disthot chrymkictta
dmk di~gonalbafdl an domum and body
s i b p m n k $eeb ofthe doreal a& do
not form diskid fins combs; batchl*
with grey brown t d ...........................
,,.,,..........*........,..** SEek,frc,m aaridi2k b 12-11 femord pmeq 42-57 dorsd aePes
b m po~tefiorof head to bwe of tail
6 8 Males untfm pala balm7no clolwful beUy
patdm tsin3@m and c ~ ~ m ' i e t t c e
......................
SoeSo~c m ~m~ * b
gmupai ........................I...................-
6 B a A &tinct dark &band pmemt 10..-..&..,

b Male5 with mn~~icuow d o r e d beUy pat- b No dsrk neckband preaent (rrpd~tos

&es; if without ealarhl belly patchtrs gpwpl ......,..,.,.,,.,,,....- '..,..,,.,....A 1e ........
b%dop~maed~udrdtayEt)ri,only on the 10a Dark neckband on both @idespale grey or
Isthslw of Tehmbpec), then large, mxy
epiky-smled species ...........................,.
*. B
blue bordered [Co~lcirbrPs-~upl
........,....................&lo- %ed&r
b mkbmd p& bordered
awmgroap) ..........................................
11
7 a 9-12 f e m d pares; 38-44 d d stdm 2la In t b anout m a lbetw~enthe m a l s and
h m p a w o r of head to basa of tail between roatral and median bnkrnet~al)
.......................... S e e l a p m s rmrhrrhcsr npmy mall scale%(Fig.BEOa); -ally two
wales b t w e a n n m d snd rostra1
b 3-7 femoral por~ts;28-87 d o r d ealee from
posterior of hsad to h ~ ofe tail ............ ............,.,., .......
~oebpolpeph L ~ a 1 i s
......................... S e e h p w rquarnourcs ~3 IIX the snout region, ab w large d e r ,
SM)b1;;u d l y 1 wale betwwn nasal and
8 a 3-11 femu,ral porm; 81-46d ~ 8 dd m
from -&or of head to barn of t d
rostra1 ......................... ....v+.q. .....
...........,................ Bdmwu~ rjapifim 1% Usually, d y a aingls anthal. ...........,..
13
b U~uallytwo canthala .............................
16
 Bupraoeulars very large aad wmged in e.
maueuatly om ar more In contact with
medial head sdw; 51438 dwal slcale8
from posferiar dhead b barn uftaiI; low-
laad ~peEies...... ...,.,..
Scabpnm l u d
S a p r d a r smaller
~ a d maged in C o
Fig.347. Poet- m r rreldom h contact with medial head
g
fmd *t. sealea; 30-51 d d 8calm ~sterior
ARer (1999aI. af head s baae of hil:kdgklan8 n p d n

30-39 d m d errales Wrn paa-r

of head
to baee oftaiJ, kbpom m a W W w n

16a 3439 dorsal ~ B fircm B porrt&or of b a d

..
tb baare of tail ,.%e&porm acantk#~wr
- - - .
b 59-47 dad ~ d e &om e postmior 04 head
tobaeeof~ilk3tZe- tas-
Rg.348. Dorsal cdoration. 1Ba Maim without colared v%~ltrala ta
................,....
s c e b p e m admrdtay ri i
b Malw with c9nspimouil ealomd ventral
gppt .,.,...,..,....,,.....,........+ ..................
17
17a 17-24 f~eraralpores
.... ......... & ~ + F ~ c B m%lunarkbu~
....,..,................
,.,.
b $la fm~al go= &L~~POIOC% l ~ ~ d e t l t

a, 5.
-8r Readixyr
H. ShnfiH 1QB89atRmkim a f Mexiom and
Fig. 549. heral vim of head ic~nthds
mwqp). G ~ a t r bd e r i m Sceioporw species); firm &
D ~ l. N982, H. SMITHet d. 1393, K~ELEB
lW4.4, H.Sna~rmst al, 1998b ESoeZopr~vwiu-
&l& pup); H, B m 8k B v ~ I%@, m
STUART I.gBi, 235- 1980,119d,H. 8 m C
E D ACuScetop~usfo~rnogw
P ~ ~ ~ Z - H I G A B1992
QMGN 1W7, K b m ~ e t al. 1998;
% Hzwm m 2 (8- of hi@lt?g&
care md W i n g ) .

a. S. lntemasalis b. S. melachFtilcus
Fig, 360, Dorsal view of head in Scelopoma.
 \

rrnoeaurrra

The delicate, small harde of the

Urosaurus are distributed predominantly
in the US and Mexico. Of the nine
Urosaurus species, or& one reachea the
northern, portion of CentraI America,
namely ti bicarinatua (-MAN
WENS 1999). Phylogenetically, the
Urnmuma species are cloaeIy related to
@nus

1942,

the Spiny Lizards af the genus $celoporua

r . P. dJ

(ETRERIDGE& DE Q U E ~ Q Z1988).
Umaur26s biearinixtw ia an imaeepicuous
pde grey colored bush and tree dweller in Fig. $61.U r n u r n s bimrimtw Inear
dry areas, urnally found oa the outer, thin- Mixtequilla, O w c a , Mexiko).
ner twige. Thus,thorny bushes are a pre-
f e d habitat. The cheat of khe male is
adorned with twa bright blue erpot9, while
the throat irs yellowriah in mlor, The threat
behavior of thae &mitorial lizards can.
dsta of pushuplike movsmenk, in which
L the whole body ia pushed vertiaallg
upright and back dawn again, The diurnal
Umaurm f d a on mall invertebratea
(inaectrr and spidere, among otheref. For
egg laying, which, in Chiapas, occurs pti-
marily in June, the females mme down
&om the t r m ~in order to bury their
elutebee of m u n d ken em (apgrox. 10x7
mm in ~ b ( A)LY~Z DELI Tam 1960).

Further Reading
E/IETFLEMAN 1942, WIENS 1993.

Fig. 352. Umsaurus bicarinatus (near

Mixtequilla, Oaxaca, Mexilro).
Saincidae
From the large family of skhb, only five
genera with a total of nine spedea occur in
Central America. Most of the remaining
~peciesare found intbe OId World. Skidus
typically have an, elongatad head, body
and tail* with relatively short leg8 and
very smooth wales. Immediately under
the scala are small dannal bonee ( o a t e
denml.

Key to Skhk Genera

1 a Without supranads, therefore rostra1 in
complete contact with first unpaired wale ~ i *3,
on dorsal surface ef ~ n o u(Fig.
t 365a,b).* 2 h,4a, Honduras), PhoW E. KaMer
b Supranasale present, therefore rostra1
entirely or partially separated from first
unpaired scale on dorsal surface of anout
(Fig.355c,d) .........................................
8
B a One frontoparletal (Fig.356b) ................
..
............................ .....Spkenomorphue
b Two frontoparietals(Fig. 35Bal ................
.
.
.............................. . . .Bctnealla
ScinceNa
8 a Scales in temporal region same B ~ Z Bas
body Bcales (Fig.35Sb) ................M d u y a
b Scales in temporal region cumpicuously
enlarged (Fig.353a) ..............................
-4
4 a Scales along the dorsomedial line not
obviouslylarger than paramedian domals;
with t h pale longitudinal stripes, the
middle of which branches on the middle of d. Eurneees
the head (FSg.3663 ................... 1Fumeae~
b Scales along the dormmedial line obvious- Fig. 356. Dorsal head scalation in ski*
ly l a r p r t h a p v m e & m dorsals; mark. ( ~ ~ p r ~ n morawl
a l s h n t o ~ a r i e t d brown).
s
ings different than these above (Fig.
356b,c)............................. lldt?80tWiFWU8

a. Elrmeces b, Mabuya a. E, sumichrastl b. M. maneguee c. M. schwatizei

Fig. 353. Lateral view of head (temporals pale Fig.356. Doreal coloration in Eurneces and
brown). Mesoscinctbs.
 L

Based on a phylogenetic analysia of spe-

cise traditionally grouped under the gen-
eric name Ewzeeee, GRIFIWH at al. 12000)
provided evidence that this assemblage
was not a natural grouping. Conagquently,
they split the Eumecea sensu lab intc
several genera, Acoording h this new
classification, one Central Plmerican gpe-
cies (E. sumichrasti) remains in the genus
Eumces, whereree two species (M, mna-
guw and M. achwadsei) ape assigned to
tihe genw M e ~ o w i m s , Fig. 367. Juvenile Eumces ~urnichmstifrom
Honduras. Phob J. R. McCrania
A clutch of 11 eggs &om E. ipumkhmti
was discovered in a f?whly fdlen bread-
fruit tree in Belize (MILLER1997). The
eggs Q6x11 mm, a bit lesa than 1 g in
weight) lay in a hotbole, which would
have baen about 20 rn above the ground
before the tree had collapsed. They were
buried in a substrate mix of fallen bark
and other organic matsrial. A female E.
sumichrusti I92 mm SVL) stayed by the
clutch,possibly indicating that the apeties
practice8 brood care, aa is the case with
other specie8 of this g e m ( F r r c ~1964,
VITT$ COOPER1985, SOMU & FAWCETT
1989). h s h l y hatched E. sumichm~H
have a total length of 70 mm with a SVL
I
of 26 mm and a weight of 0.7 g (MI~.&EB
P W ) . While adult E. sunichrcsstl have a
pale brown ground color with black longi-
tudinal stripes, the young are aomewhat
more brightly colored. The latter have a
bright blue tail, and several orange apots
on the head, a ~ well
s as three orange t9 yel-
low longitudinal stripes on the body
(MILLER1997).

M,Nat. Sci. Wiladelphia lsBB: 821; type

& e d i t Orhaba, Verauw, M e x h . SVL to BF
mm. &n:entra~vma-, ~esim,ta
3 3 0 ~ale0 ~ nmthearpttampwtian
; Opyutatan
h u l a , sea lwd to 1OOO m elevation.

M h e Reading
~
et al. 2000.
TAYLOR1936,GR~FF~TH
While earlier authors (e.g, TAYLOR1956)
pmpoeed that, in Costa Rica alone, three
epecies (alltacea, braehypodo, unimargi-
nata) of the genus Mabuya occurred,
recent author6 conclude that only one spe-
ciea, M, uunimrginata, is distributed all
across Central America (SAVAGE& V u
1986, VILLAe t al,1988).However, the geo-
graphic variation of Central and South
American representatives of this genus
hae hardly been examined. Therefore, the
c m n t taxonomic division can only be Fig. 359.Mabuya uninmrgin~--.Bar --.-,
Rio
considered temporary, Thi~hisinsectivorous San Juan, Nicaragua).
kink occurs commonly in many locations
in the lowlands ef Central America, These
animals are generally found on the
ground, but they will climb tree trunks in
order to bask.Mabuya unimarginata is a
livebearer, with lit& size varying h m 2
to 7 young*

Y a b ~ y eunimargintrkr COPE 1Bf32I.2,Proc;

Acad. Nat. [aci. Fhiladelphia 1k 187; type
WiyPanamzl. SVL to 90 mm. M h to
B w ,a m lev61 to 1W m devatiod
Further Reading
TAYLOR1956,Webb 1968.
Fig.360. Mabuya urrimrgdnata Wdeh
Maderaa, Isla Ornetape, Nicaragua),

Fig. 358. Mabuya unimagincdta Inear

Guanagasapa, Escuintla, Guatemala).
 found under lmm bark on upright tree
-* trufl3rs. The Mmo~cimuas p d e ~are shy,
~~cretiv liaada
e that are rarely encounter-
L .- ed in nature,

Key to Meso~cincw
1 a With twa wide pale 1ongItudinal stripee,
- which hegin at the tip of snout and fuse
into a broad band in the anterior dorsal
area [Fig. 356~);21 scales at midbody;
front and rear limbs touch, i.e., averlap,
The genus Mesoscincus wae recently pro- when laid at the aide of the body ..............
posed for three species of kinks formerly .........................Mesomimcwn mliuxz&ei
assigned to the diverae genus Eumeces b Domum b m with dark b r m langitudi-
nal lines,whiEh kl some individuals break
(GRI~ITH et al. 2000). TWO of the apecies up into spots (Fig, 366b); 17-19scales at
included in Mesoscincus occur in Central midbody;front and rew limh~do not touch;
America, M,mnaanaguae and M,~schwartzei. when lsid at tbe dda of the M y ..............
These ski* are ground dwellers in dry ........................
H e m b u s managarae
and seasonal rain forest, where they can
be seen basking on roots and fallen logs. In F d e r Reading
July 1999, I discovered an adult specimen '~'AIWR 1935, 195.8, L~TUZet a], 1979, m m
of M managuae at the foot af the Volcb lggO* et 2000.
Co~iguina,Nicaragua, under a muety tree
t d . Sometimes these lizarda are also

Fig, 363. M e ~ i n c r r nonreguae

s (Rfo BandiIlal,
Coats Meal. P h m R.W.Van Dwender
Fig,364. Mesaaeimw sckwavteei 119 h N
Temoehn, YucaW, Meximl.
Photo: Section of Waxpetalogy, Natural History
Mumum of Lm Angeles County,
lFjg,366.Scimcedla genamingeri (Hidalgo,
Mexico).

According ta the tgw)nomic division by

GRE~R (1974), one Centd American
species, 8.gemmilagest h m the subfamily.
Lygosominae, L included in the genm
Scincella, where- the w e e aarsatua,
chew%, ihceF'tuS and mru8 are placed in
the geeue Sphemomorphus (MYERS &
DONNELLY1991). Seineelkt gernmitzgeri
has 60-68 dofsal acalea from the parietal
the base of the tail (at the level of the
posterior border of the thighs) and 26-28
longitudinal rowe of dmeal scales (H.
S M1949). ~

EkirrcelEfc p,~~nilrygerS(COPE lW), R=OG

Acad. Ngt, M.P hh
de1m 16: hiQw
!z!T& f k h x , ~C
wgo*Vm-
' ,-). WE& SVL a0 4
a d oaac% ld&a,
aoO-2UoU m elevation in pine hnervt,
5

I
 Tbe Central h e r i t a n herpetafauna in-
clude$ three spmenfly &~eb-~@hted,
small skhk s@es fFom the erubfLlm..
Lygaa0mfeer.e(mwtt68,& e e i , a d incar-
kt681that, until the b&ning ofthe 1970'~,
mre varioudy iPOluded in the genm
@gamma b,g. SWT I940), Sci~celEck
(0.g: H, 8mm & TAYLOR 1960) or
Lehlopismu (e.g, TAWR 1956, STUART
1963,Fm%mgG D O N O S Q - BN701. ~ In
his revhim of genera related to &,967, Sphemmrphw c R e m (Qen'o
LeioIopisnwa, Q ~ (1874) B placed the Swlapa,A t h t i m Nor% Niaragml.
@ee above-maned species in &0 pnue
Sphenomrphuo, s view which is widsly
held M a y aYIFfTCH 1983,V u et al. 1980).

C
Recently, M m m & ~MNNELLY IlQ91)
~ BaS fourth
M species from thia genw
in Pariaatla, LR mm8, that iifl h ~ onlym
fffina a few apchmA
1 memas Sphemmo*ua chrrici are
diurnal jpnmd dwellam in the under-
growth of the Pdlis h a t , 2.3. assa&io is
m o w an W i t a n t of suhhurnid areaa,
The ~hortl e s are a~siateainfirnard l o w
midtion by makelike mwementa of the
body, empBci6111y when incmwing speed, At
dl' banimal5
l?2~~8bE m~~
btS
WwIL
active and $7
in 'C, I
the 1- 388. S p l ~ , ~ m a r p has*w (near
mo-t while they hide the &mnagmapa, hlentiptla, Guatemata).
midby lsun hG Q O ~ W phiC8B. h t b d k -
nwn, the agile S,pLnomorphus will
undertalse a t h e r period of activity - - -
IFrnr:~1985). Diet eon&ta of eeaaIf &luw
pDdrr insect^ and apidere), which they
track primarily with their olfactory smw.
h g e prey animals a m pmfemx$ and
wren colenibtllism 48 b o r n among thew
akinkr, CFrtv~1883). T h b @pecksof the
gmm SpknoktasfjIs~~ repMdw o v i p
o t ~ d y ,with femalm producing mveral
clutcihai per sleasoa, each wi& m e to three
(& fimf 0gg$. w k e m S p h - h
in the constmtly moist Caribbean law-
lands areas reproduce tboughout the
y~ar,pqml8tion8 at hi@w dtitude~or
mesa with &tinct dry pedode am higW Fig. at%. S a m of5phenomosphlcs
mistma1 in tbir egg plloduct'i~n.rl(hlls) &&&ltBnB RQo, Maim). Flreto:H.Bahma B.
 F ~ C(19833
H rspofied that the mproduc-
live activities of S. cheniei in the region of
Tunidba, Costa Rica, etagnate in the
month &om November to April. With an
SVL af approx. 22 mm, the hatddhjp of&
chrriei me already 40 % of their adult
size. Within seven months, the young are
sernrally mature at an 8VL of 48 mm.The
averagrj duration of a gemration of S.
cherriei is approximately one year.
Key to Sphwnwwhrcs
1 a Withoat prefmntd scale, therefore,
Mntact o m m between hntonasal and
&ontal...............8pketsommph ram
b frefhntal scale pmmt*fron~nasdnot
.
in conbad with h n t d ..................... B
B n 30-36 sealeg at fpidbod~sfront and rear
extremitiee touch when laid at side of
body; tail brPwn in adult specjmene, blue
....
in jwenilw Sphenomotpkrra c h d e i

-
Countlese predatarrs (primarily snakes,
birds and mammals) prey upon these
~ m d &inks,
l AU Sphnonaorphug spedea
wilI discard their tails if they are seized.
Cam ZWli $rp
Acad. Nat. Sci, Philadelphia 16: 178; type
V o l h Izaloo, El Salvador. SVL to 60b Fewer than 66 domd ecdes between
Calima, Mexico, to El Salvador and
b 2431 Scalee at midbhdy; front and rear
exbemities do not touch when laid at ids
of body;tail red browa in adult .~pecimen$
i a W red in juvdlee .......................... 8
8 a 6T-77 dwml E& bstw~enparietal and
baae of tail tat lwel of p~eteriarborder of
thighg); 27-31scales at midbody ................
.........,...........,
Sple~u)ltcorphueamatroe
parietal and h e of tail (at level af poster-

' uthem Honduraa WCCRANE& KORLER

MQa),eea level to 1500 m elevation (to 2600
I m slevahm a c a d h g to J~HNSOM 1989).
ior border of thighs); 26 or fewer d e e at
midbody ........SpItefu1lno~phwincertus

Sphnomorpkaa ebrriai (COPE 18931, Pro@, Further Reading

a
Amm. Phil. Soc. 81: 340; type l d t j r : TAYLOR1937a, STUART1940, H. SMITH1946,
jPalmsr, C00ta Rim. 8VL b Bg mm. Tabasg 1951,GREER1974,W w w & MCCRANIE1994,
Mexico, to western h a m a , h n sea level t& MCC~ANIE% KOHLER1999a.
1800 rn elwatim in lowland aad premontam
rain foreat*
I
irscertus ( S ~ ~ A1340),
R T Oca
Univ. Michigan $81:10;
%
Tajumdui, Guatemala. S
40 67 mm. Guatemala and Honduras.

5 ' p b n ~ m r p h wrams MYERS L I~NMELI

1991,h e r . Mu.Nov. 8087: 2; type localit:
h e d w a h of Rio Gwbo, 780 m elevb-
tion, 8"47'N, 82"11W, Boca~l del To*,

1
C
Fig. 370 Sphmmwphtrs mrue Iholotme), A Sphenomorphus rams
w.
phoh:C. &$yiam ISphenomoiphus ohem'el
 Teiidae
The lizards of the f d l y Teiidae, including
ten genera with approximately 110 ~pecies,
w e widehlpr~adin temperate, &ubtrspical,
and tropical areas of the USA, Mexico,
Central A m e r i c ~ South
, America, and the
Antl.lIa, Most specie8 are active termtrial
lizards, and all are &diurnal, All teiidsl are
oviparous; parthenogenesis is preualent
among Aspidoecelia

Eey to genera ofTeiihe F

ig.373. Amiua utldulata (ElImpmfble
1 a One mw of greatly enlarged ~ e the
d e on National Pruk, Ahuachapctn, El Salvador).
outside of the upper eu?n (Fig, 37+); tor-
sal pattern with five OT fmer loegltudrnd
stripes; a fleshy eheath enclaaing baee of
tongue (Fig, 971a) ........................
h i m
b Tbre or more m w s of mderatelg en-
kg4 &w on the outaide of the upper
arm (Fi.3ST2bh dorad pettern ueually
with six or more pale longitudind stripes,
or wit& pale spoh no fleshy sheath en-
closing base aftanpe (Fig,371-bl ..........2
B a Qne intmpaietal and 4 parietals (Fig.
3%0 ..........................Cneddhpkarm
b b e iakerparjet;al and 2 paFietals (Fik,
386b,e)............................
.,..hpddo%ce&u

Fig.374. Ameiua q u o d l r i l i ~ a(near Nelly,

Cwta Rica).

Pig. 971. Floor of mu&. Note b g u e &heath

tamwl inddw. I

Fig, 372.Foreleg (eda~p;ed Wedl.

154

Fig,37'6. Ameiua udulata (SenJdian,
Suchltepli.quez, Guatemala).
Fig. 877. h i v a ameiva. Phsrto:R.D.Bartlett
I l a e t ~129-137daye. ThejuGenilen of A fbt-
' haJe
de San Bles,Panama).
In Central h e r i w , there are six ameiva
specie&of which A. fiediva and A utadula-
ta are the most widely distributed
(ECHTERNACIFT 1971).Ameivas are diurnal,
very agile lizards that are m m a n in
many placsa, primarily in open serxrndar~r
forests along the paths d in clearinge,
The ~unbightand heat lwing ameivm keep
their body temperature arwnd 37 *Cduring
their active phase (HBW 1963, l i h ~ ~
1969,F r r c ~1973a). Cooler and rainy days
are spent in burrows in the ground.
Ameivaa are tenacious hunters, who will
wander around their ter-ritam l o o k for
prey, scratching in the leavG, digging. in
the ground, and searching in the under-
growth and under bar$, k u m f n g any
creature they can overpower, p z h a d y
ineects md araehnide.The lizards are eon-
~tantlyflicking their h & u e s , checking
went8 in their s u n u l l m ~ Although
.
t h e e lizards are gmund dwellere, they
will also c b b tree tnmks and low vents-
tion ta baek or to reach prey. Important
predatom of the ameivaa lare the diurnal,
quick colubfide, including Comphh linea-
&a, DryadopMs mlamEomm, Drymobius
nargaritifints, and Mmticuphts mentova-
ti^.
The C~mbalAmerican ameiva specie8 pm-
duce several clutches of agge per year, with
the eize of the Female ~
clutch eize dependent upon sgeciea and
a m ~ v a1-9,A,
&diva 2-4, A, undubata 1-7; E ~ M H T
1083,~OHLER1997). At 28-30'C, the incu-
bation period for the e m of A. ameiua
iva. A auadn'lilaeata. and A. undulata
bluwlorea t&, jut l b the
juvenile whiptail Aspidosoelis deppii. Up
to three ameiva specie8 can occur in a
aingle area Iayrnpatric) (ag.,the species
festiua, leptophrya and quadditceata in
southemtern Coeta Rica), wibh each
making me of a slightly different ecologi-
cal niche to reduce the competition
I(ECHTERNACIPT 1971,1983,FTKX 1973a).
A m - ~ L u m m u u 1 iooj, uya~-iI
N a m e . ed. 1Q: 202: Pme localitv: Brazil.
BV]& to '197 mm ~ d e s 'and
i (fe
d e g ) , Cwta Rica Panama ab well as large
parts of tropical Sonth America, sea lwel to
TOO m elevation. In hnmm, the eubepecler
Ameiua a. ptaeeignia (BAIBD& GIRARD1852
occurs.
AmEva ehaitaarni ~ A E 1942a, T Pmc. Biol.
Soc. Washhghm 65: 143;type ha1it.y: along
Cahab6n-Lanquin trail about 2 km N Fin-
Caniher, about 38 km ENE Cobtin, klta
Verapaz, Guatemala. SVL ta 85 mm (males)
and 76 rnm (females). Eastern Chiapas
Mexico, ae well ae emtern and centra

1I
Guabmla, 800-2000 m elevation.
ARaeiua fesfiva (I~XT~~NSTEIN L VON MULTENS
18661, Nomenclator Reptilium
Amphibinnun Musei Zoologici Berolinemk:
18;type locality: Veragua, Panama. SVL et e
114 mm (males) and 129 mm (females)
Masca, Mexico, across all of Centra
America I. Ear as Cdombis, aea level to l a d
m elevation. In Central America, three sub
species r n h i v a f: fiativa (Panama anc
northern Colombia); Ameivra f: dwcsdsii

9I
BOCOURT 1673 (Iethmns of W ~ ~ l l t e p etoc
Uiwmgua); Amfua f: -dentalis Tml%
1966 (Costa Rial.
Amiva Eeptophrys COPE 1893, Proc. Amer.
Phil. Soc. 31: 341; type locality: Buenoe Air-
Cogta Rim. 9VL to 183 mm Cmdes) and l2
~ u Ifemalea).
n Southesatern Costa Rica
emtern Panmu, sea level to TOO m elevation.
Ameiua qum?rilinccata (HALLOWICU 18611,
b c . Acad. Nat. %. Philadelphia 12: 483;
type locality; Nicaragua. 6VL to 88 mm
(males)and 82 mm (femalw). Southeastern
Nicaragua ta western Panama,~a level to
1060 rn elevation.
Ameiua undulata (WEG~~ANN18341,
Herpetolngha Mexiwa: 27; locality:
&Xh. 8VL to 129 mm [ M ~ ~ G Ba) d 111 mm
(feglalehl).Naywit, Mexico, to Cmta Rica, se
l m l to 11600 m elevation. In Centra
America, six subspuies olrrur:Anseivar u. gai-
geai SMITH L LAWE 1946 (northern Yucath
Pmimula), Ameiva u. Saarmr 1940
[Ram of the Yucatdn Penineula and northern
Honduras), Amsiua u. m i d k BARBOUR &
Z~VEWDGE 1Q29a [Isla del M a h Qrande),
Ameiva u. p a w B A R ~ L O ~NOBLE1915
[Pacific aide of Central America), Anaeiva u.
palchm WWELL 1861 (Honduras to Coata
Rica along the Caribbean aide), Ameiua u.
t h m s i S m & UUFP: 1946 ( u ~ e r Rio I
Key to A m e k
1 a 10-12 longitudinal rows of ventrd =ales;
no obviou enlarged throrrt d e s (Fig. a. A. amelve
379a1, rather gradlkal increase in size
posteriorly; wales on the nadetrride of the
neckband (mesoptychial aedael net
obviously enlarged .........Amclva ame3w
b 8 longitudinal rows of ventral emlea; 2-5
throat scales enlarge scales an the
undereide of the neckbud (mesoptycbid b. A, %dw
rcalee) abviody enkrgd (Fig379bdI..B
2 a Anterior throat scale6 smaller than p a -
tgrior throat scalea (Fig. 3820) .................
3
b Anterior and posterior throat d e s of
equal aim (Rg, 379b,d) ......................,.....
4
3 a M e t a l a and firmtopapietds separated by
one or more scalea (Fig,38th); a d @ at C. A. lepfophve
mid-throat greatly enlarged .................
................................... Ameav~krp-hv~
b Perrietals in contact with ADntoparietds
(Fig, 380b); ~ c d e eat mid-throat only
slightly enla@ Amwiva qua&dlhata
4 a Scales at mid-throat w t l y enlarged and d. A. quedmInesia
net arranged in a longitudinal row (Fig.
378231; a pale dmeomedid Ionj&tudinal
stripe preaeet (may be absent in very
large specimens) .............
Anreiva fbtiva
b Scales at midbthroat greatly enlarged and
arranged in a lmgitud.iaa1 row or or&
slightly enlqpd and irregularly
(Fig 379d); without pale dorsome IJ y ..Lum*&
longitudinal stripe ..................................
6
6 a SVL to 86 mm (malw) and 75 mm (fe-
male>;paravertebral stripes very narrow;
do~.solawalspoh in males fueeddfifie Fig. 518, Ventral view of bd in h i v a .
g
d o d a t g r d stripas . . , h i mcbSfzami
b SVL to 129 mm (males) and 111 mm
(females); pavertebral stripe8 wide; dor-
solateral spots in malee, if pment, not
fused with pale doreolateral stripes ..........
.................................. A m e I unduwu

a. A. bpnqPhfp
1
b. A, qusr#Iine&
Fig. 380.Dorsal view of heed in Ameilro
1 (&oultoparletale orange, parietals brown).

Further R43ading
E ~ A C E 1971,
F P
 Whiptails are the most common lizards in
the dry areas of Mexico and Central
America. REEDIERet al. (2002) proposed a
new classification of the whiptails based
on a phylagenetic aaalysis. Because the
diverse assemblage of species that was
traditionally grouped under the generic
name Cnemidophorw was demonstrated
to be paraphyletic, REEDER et aL (2002)
~ p l i tthe p u p into several genera. The
lemniswtus group retained the genaric
name Cnemkiophorus, whereas the other
North American and Central American
speciee were transferred to the resurrec-
ted genus Aspidoscelis. The genus
Aspidoscelis contains at least 87 currently
recognized bisexual and unisexual h a
and is distributed throughout moat of L .-
North America (except Canada), south Fig. 382.Aspidoscelia deppii (male abwe, fe-
through Mexico into lower Central male below) at Playa T m n d o , Cogta Eta.
America. In Central America, the genus is
represented by six species (DUELLMA~M &
WEUMAN 1960, VILLAet al. 1988, LEE
1996).
in comtant motion. As heat-loving ani-
Whiptails are active huntem that will eat mals,the whiptails are out in the open pri-
everything that they can overpower, most- marily an hot, sunny days, whereas they
ly insects and other arthropods. To the retreat to their burrows when the sky is
human observer, thege lizards seem tre- cloudy, Most of the 189Aspidoscekis deppii
mmdously restless, as they are virtually examined by FITCH ( 197313)had body tern-
peratwee from 29-42 "C.Aspidoscelis are
me& active in the morning, Most of the
Aspidoscelis species produce 2-6 clutches
per year, each with 1-5 eggs, which they
bury supeficially in the sand. In the dry
season, reproductive activity comes almost
entirely to a atandstill, whereas the egg-
laying period begins with the onset of the
rainfall. The incubation period lasts &om
60-80 days. Aapidoscelia deppii becomes
sexually mature at the age of 5-6 months
(ECHTERNACHT 1983).Worthy of note i~ the
fact that several Aspidoscelis spsdss,
including the Central American A COZU-
mela and A. mdecki, reproduce partheno-
I genetically, In theere two species, only
Fig.381.Aspiduscelis motaguae. females, which produce fertile eggs with-
Photo: R. D.B d e t t out any previous copulation, a m known,
 Fig. 384. Aspidascelie aragusticeps,
PhotQ:R,Cedeaa V,

Fig.383.Aspidoscelis cnzumelae m s l i n i Rg.886.Aspidoscelis gutbata (mde right, fe-

(Banco Chincham Reef, Quintana Roo, male left) near MiKtequilla, Oaxaca, Mexico.
Mexico). Photo:H. Bahena B.
Kay tQ Aepzdoacelis n
1 a 4 supracrdars ..................... ..............B

"-I
b 3 suprauculars ........................................
8
4 a Dormm of juveailea with aix pale
Ioxtgieal5tripes on a dark brown back-
ground; in adulta lSVL 100 mm), d ~ 8 d I
etripee usua11.y corn lebly replaced by
epots .................b p
b Domum o f j u w h with six or seven pde
lmgitudinal strim an a black back-
ground; adult pattern uariabl$ but e r n e
trace of skiping usually retained
....................... Midmm~is
.......... 387-Aspidowlis c c ~ [Isla u ~
Cwtunel, Merdoo). Photo: R. Cedefia V.
8 a FmntopaFietalrs divided from parietab by
.....
one or more scaleg .................. 4 .#

b Fmnto+Ms in contact with parithala

..........................
.....,,.,,,,,,....,......,..,.,~~,.,r,,, 6
4 a Dasal attern with indimtinct para-
P
vertebra &ripear, dimnthuaus or abaent
.an the poeterior bady half .........................
. ......
........,,.,,.. ,... AapidoweZC nodaclkf
b Duraral pattern with foru distidct contin-
uous paravertebral %tripes ......................
.......................n. A e p W I t a CPmmWkr
6 a Fewer than 1160 s a l e s around midlsody,
not counting veatral d e s .....................
,.....,.*,,.,..,.,..,*........AepmdDecelss gu;mdb
b More than 166 scale&around midbody,not
countitlg ventral .........................
.................................Aepidamelb &ppH

a.
388. k a l head d a t i o a (~upsaOCdm8
orange;p a r i e l s ILIQWII)
in &pa;do8~&? and
Cmmirdaphomrs,
Fig,388.Adult male of Cmnaidophorus Fig. 389.Adult female of Cmmidophorus
lernniscatus (Isla de Utila, Honduras). lenniscatus (Islade Utila, Honduras).

4 nemidophorza lernnigcafurr (LINNAEUB

7681, Syctterna Naturae, ed. 10: 209; type
Iocality: 'Guineaw [probably in error for
WEDER et al. (2000) proposed a new Guhna]. BVL to 104 mm. Central America
classification fm the whiptail lizards and +om Guatemala ae far aa northern South
transferred the majority of species from herim east d t b Anderr, iectuding many
Cnemidophorus to A~pidoscelis.As cur- c&huxe ialaade, aa well rrrr variou~Caribbean
ialande (e.g+,%Mdd and Tobago), ma lwel to
rently recognized, C. lemniscatus is the ~tlleaslt loOD m elevation. OnZy the nomimte
only Central Amsican representative of subrspeeies in & p h l Amnrirrsr
the genus Cnemidophoru~.
M e r Reading
Cnemidophorus lemniscatus is a typical KOHLER19966,COLE& DESSAUER 1993,Wright
diurnal inhabitant of the open mastaI & Vitt 195S,REEDERet d.2002, MOJ~CAet al,
areas, but will deo advance inland aIong 2003
large rivers, hardly ever penetrating the
forest. These active lizards are often seen
in great numbers. They run a bit, stop ab-
ruptly and execute rapid turning motiom
with their forelegs, which has earned them
the name "Shaky-Paw"amongst the inha-
bitants of the Islm de la Bahia, Honduras
WILSON & HAHN 1973, IC13i-m~1996d,
1998fl. Although Cncrnidophorus bmnis-
catus feed primarily on insects and other
arthropods, juveniles have also been
obgel-ved eating cactus blossoms (Opuntia
waratiana) (MWARIB-URRUTLA et al. 1997).
C. lemni~catuaproduce several clutches
per year, each with 1-5 eggs, which hatch
after about two rnontha IKOHLER 1897).
Xantusiidae
The family Xantueiidae colmiets of three
genera (Xcuttusia, Lepidophyma, Crico-
8aura), with only Lepidophyma muring
in Central America M BE^ & CAMABILLO
2002). The distribution ranges of mast
night lizard8 are extremely fragmented,
and populations are o b n isolated from
one another by hundreds of kilometers.
Such disjunct distribution has been taken
a~an indication of the great geological age
of the group (BEZY1972).

Fig. 390.Lepidophymra 8mithii (El Refugio,

- Ahuachaphn, El Salvador).
Lepidophymci-
l n e I r species of the genus Lepidophyma
are distributed &om Nuevo Le6n in north-
eastern Mexico to as far south as Panama.
They are bizarm-looking mid-size lizards,
which are mcretive ground-dwellers that
are seldom observed outside their hiding
places (beneath decaying tree stumps an;
loge, in rock crevices and caves). They 1
have no eyelids, rather having, similar to
many gecko species, so-called "spectacles".
The sides of the body are decorated with
I
countless, irregularly arranged tubercular
scales, whereas the tail is equipped with
whorls of slightly enlarged scales.
During the day, these lizards hide under
rocks and fallen logs, which provide a Fie 391.Lepidophynaa flauinaaculatum (near
moist microclimate. These ani- Pueblo Wiso, Jinotega, Nicaragua).
mals are sit-and-wait predators whose
prey consists mostly of insects and spi-
ders. The remarkable thing about this
genus is the livebearing method of repro-
duction. There are also populations that L. chicoasense
consist entirely of females that reproduce
parthenogenetically. During the months
from April to July, the females of L flaui- , 1' L, reticulaturn
maculatum bear 2-5 (seldom as many as ( ' L. fievlmeculetum
8) young, which have a SVL of 35-38 mm
at birth (ALVAREZ DEL TORO1960, TELFORD
& CAMFJBEU1970).

Lepldophyme
&mty I
hptdop m a m u w B ~ 1 9 ~ h ~ a
Sd.
C0;ntrr.k 1; type w
nmr Chin@$, 145 m elervatian, Altr
Gwtelplala SVt ta %$6prm
Eil~ruvdtlrelmwsl~~kb
h de 11oacku$-kpm%b~~
and Bern
af H u & u & s ~ ~
l d k3ta VmpiWl, C z l a m .dm Mayr
Ym* dmka,1 0 0 m nr elm-.

Fig. 592. Dorsal head scalation in two species

of Lepidophymu,

Fig. 3H. Lspidqphyma flovimacudatum (El

Ocote region, Chiapas, Mexico),

Fig. 3W. Lateral head Elcalatlon in two epodea
dlRpidophpnza. we~betwee11
i Epd p0St;osbi-
tsal eup~dabidm d ptoeulaF ommqa.
4 a Dorsal scales (occiput to base of tad) 179
or fewer; more than 10 longitudinal rowe
of ventrala on posterior half of body
..............................
Iapidqhyma tipetzl
b Dorsal acalea usually 180 or more; if fewer
than 180, 10 or fewer longitudinal rows of
ventrals on posterior half of body
5 a 0-1 Scales between second oetorbital
supralabial and postoc111am (kg.
394a);
laterd tubercular males arranged in 33-
46 mws .................kpidophyma m y a e
b 2 or more d e s between a w n d post-
orbital supralabial and poetoeular scales
(Fig. 394b); lateral tubercular scales
arranged in 24-32 rows ........................
8....
8 a Throat region without abvious dark retic-
ular markings .................................
..Lepddophyma retleulatum
...................
b Throat region without obvious dark retic-
ular markings, although some diffuse
shading may be prewnt ........................
.............bptdophyma flevimaculattlm
Further Reading
Fig. 395. Xenosaurus gmndis (near Cordoba,
Veracruz, Mexico. Photo: R.E. Ballinger
The family of knob-scaled lizards has a
fragmented distribution in Mexico and the
northern portion of Central America, as
well as in China,which ahows it to be a
relictrial group, known fmm fossils since
the Upper Cretaceous.
In Central h e r i c a , only one knob-scaled
lizard species occurs, ~ekosaurus grandis.
The ability to autotomize and regenerate
the tail is not present in the species of the
genus Xenosaurus. The typical habitat of
Xenoaaurus grandis is open rocky land
with hiding places in the form of rock crev-
ices. Actually, all the grandi~observed
by BALLLNQER st al. (1995) in Veraemz,
Mexico, were in rock crevices; none were
found in the open. Only one knob-scaled
lizard lives in each crevice, Social rela-
tionships and interactions could not be
determined in field ~tudies,which leaves
the suspicion that these animals are very
solitary. Horizontal rock crevices are pre-
ferred over vertical ones (BALLIN~ER et a],
1996). The assumption in earlier Iiterature
that knob-scaled lizards are nocturnal
(KING & THOMPSON 1968)could not be sub-
stantiated by more recent field studies. At
nigh4 all of the observed X,grandis dept
Fig. 3346. Xenasaurus grandis.
Photo: R.D.B d e t t

with their eye8 closed and did not react to

light stimulation (BALLINGER et al. 1995).
The first activities k g . opening the eyes,
looking out of the rock crevice) were deter-
mined to take place about two houm a h r
sunrise. Xenosaurus grandis belongs to
the thigmothermic lizards, which make
use of the heat in the smounding sub-
strate, but not the heat of the sm'~ rays.
The body temperature of active knob-scaled Fig, 397. Xenosaurus grandis (El0oot.e region,
Chiapw, Mexim). Photo:A. Rarnbrez V.
lizards corresponds to the temperature of
the surrounding rock and lies between
20.0-26.g°C (average 22.7"C); (BALL~GER
et al. 1995). In general, these lizards are
not shy and seem h o s t lethargic. The
radiue of activity is reported to be less
than 1 m (ALVAREZDEL TORO1960). If &
knob-scaled lizard is grabbed, it will epray
a considerable amaunt of fluid from the
cloaca1 bladder as a defense mechani~m,
eimilar to many fro@ and bade. Knob-
scaled l i z d ~feed p r i m d y an many
types of insects, but will occasionally eat
small lizarda (RWSCH 1981, E~UUNGER et
al. 199B). Xenosaurus grandis reproduces
ovwivipmusly, with litters containing 3-7
young (FRrrr~1966). A t v m nEL TORO
(1960)reported that births amongX gran-
dis occur in Chiapaa, Mexico, h r n March M h e r Reading
to July, and that newborns have a h t a l F m s 1966, PRESCW1981, BWINGERet al.
length of 45 mm. 1995.
Amphisbaenia

Worm Lizards (Amphisbaenia)

Although the taxonomic position of the they can be found in the underground colo-
worm lizards has yet to be satisfactorily nies of lmf-cutter ante (GANS 1969).Worm
resolved, this group is generally regarded lizards are equally able to crawl back-
as an independent suborder related to the wards and forwarb in their tunnels.
lizards (Sauria) and snakes (Serpentes). Amphisbaenids are skilled hunters, prey-
Evidence exists that the worm lizards had ing upon insects, spiders, and rodents, as
already separated from the snakes and well aa worm snakes and other reptiles
lizards (order Squamata) during the (GANS 1969).
Cretaceous period, even prior to the Tiae of
modern lizard families (GANS1969,1978).
Earlier authors had considered the worn AnSpI&dnwna o Eh 176B,8-
Lizards to b a lizard family (VANZOLINI Matnrae, ed. 10 229; type locality: America.
1951).A number of anatomical characters &VLta 670 mm. Tropical South America east
of the Andes. Oixummo8 in Panama ques-
distinguish the worm lizards from both tionable.
the anakea and the lizards. These include
the reduction of the right lobe of the lung Amphiebaena fuliginosa L~NNAEUS 1758,
Systema Naturae, ed. 10: 229; type l d t y :
and the exhtence of a middle p r e - m d - America, SVL to 600 mm. Panama, tropical
lary tooth,mi well as a characteristic skull Swth America emt of t h Andes,
~ as well aa
and middle ear morphology (GANS1969). weatern Colombia and Ecuador.
AmphLbaena s urrelii (BQWLEN~ER 1916),
Worm lizards are the true burrowers ~ m oZ. ~ Ih,
. k n h n U I 659; ~ type l o d i -
:Andagoya, at junction of Rfos Condoto and
among the reptiles. Living in selfen-
structed, permanent tunnel systems, Ian Juan, Colombia. SVL ta SO0 mm.
Panama, northem Colombia, and Venesuela.
ampbisbaeniana have an elongated body
and, with the exception of one genus
(Bipes),no exhrnal limbs. Rather than use
already existing tunnels as worm snakes
wiI1, or etaying in loose soil or aand, they
we in the habit of tunneling through com-
pact earth. They seldom come to the eur-
face and then only at night. Frequently,

Fig. 398. Ampkiabaeno fulighwa (Peru).

Photo: E.Lehr
---
3 a pdedarg ~po*@ m m t h n
four preclwd ...............................
....................~ k b M f i w -
,

-
?'
b Without spatting Eour pmclaaoal pores
...................
t t , 4 . . d m p .punrlli
R~
,
'
V m m lMI,QWB TsgZa, b, 1967, Ham-
as- 1073, Gma @ NCA- 1977, G m s 1978

Fig,400. Amphisbaena albra.

Pboto: R.W.Van Devender

Fig*401,Amphisba;encsa h .
Phob R.W.Van %vender

Fig. 399. Ampbisbuena spurmlbi. Modified Flg. 402.Amphisbaenu ftlligihosa (Nueagandi,

fmm EOULENGEB
(1915). Comarca de San Has, Panama).
 Sexpentea

Snakes (Serpentes)
\
antedor temporalp . c e b ~ t o c u ~supraocular
mtmrlor temporals / prefrontal

Fig. 403. Scalation

characters on the
head of a colubrid prenasal
snake.
F mental
postnasal

- 8
infra,abra,s WJ'al'bl'ls '

1st dorsal row

Tm=%w
ventrak
Internasal lntralablals

chln shleldrs

9- m
13
-
groove
mental

In Central Amedca, the snakes are well 8 a Ventrda do not extend across the entire
represented by 276 different species, body width (one or more dona1 rows on
the ventral side); uaually all or most sub-

-
I
Key to Snake Famillea
1 a Tail conspicuou~lyflattened laterally (Fig.
.
686) ,, .,.EIapidae (inpart: we snakes)
b Tail more or lem mund in crass-section 2!
caudals undivided (exception: Larocemus);
often with cloaca1 spurs on either side of
the cloaca ..............................................
b Ventrals extend across the entire body
width (no derads on the ventral side);
7

uaually all or most subcaudals paired

2 a Ventral scale8 not distinctly enlarged,
scales a m u n d body all approximably of
equal aize ....................
.......................
8
b Ventral scales distinctly larger than dorsd
scales ....................... . . ..................6
3 a 14 scale rown at midbody ......................
.................................Leptotyphlopidae
b 18 or more scale rolwa at midbody ...........4
4 a 18-20 scale rows at m i d m one preorm-
lat; no subocular ...............
Typhlopidae
b 21-22 scale rower at midbody; preocular
present or not; subocular present or not
..................................Axlomalepididae
B a One tube-like loreal pit between nostril
.
.
and eye ..................... ............Viperidae
b Without tube-like loreaI pit between
nostril and eye ........................................
8 loreal present or not;coloration and mark-
(exception: Pseudoboa); no cIoacal spurs
...................
present ............................
. . B
7 a MI m most subcaudals paired ....................
..........................................Lorwemidae
b All or most eubcaudale undivided ...........B
B a Fewer than 36 d o r d rows at midbody
...
................... ...,.......... Tropidophiidae
b More than 36 dorsal rows at midbody ......
.
.
........................ ........................
Bofdae
It a Upper jaw with 2 immobile, enlarged
teeth, othezwise, upper jaw without teeth,
no loreaE; body pattern with complete
black rings ....................
,. ..........
..............*..
...........Elapidae (inpart: ooral mudma)
b Upper jaw with many teeth, none compic-
uously enlarged in h n t of upper jaw;
i n g variable .......................
Colubridae
 Seolecophidia

Blind Snakes (Scolecophidia)

Bey to Anomalepididae
1 a Dorsal surface of head with two large pre-
frontals behind the rostral, bro~dly in
contact with each other and followed
posteriorly by a large frontal (Fig.406a) ....
........................................
Anomdrpdr
b Scalation on dorsal surface of head diffe-
rent than that above .............
,,...,.,.., ...... 4
2 a Prebntds behind the rostra1 in contact
with each other (Rg.40%) tlehhthop,kte
Fig. 404. Scalation character^ on the head of a b PMtfrantals not in contact with each other,
blind enake. separated by conapicuowly large ro~tral
(Fig. 4 0 6 ~..........................
) LbQphhp8
Three M e s of comparatively primitive
snakes are grouped under the term blind
snakes (Scolecophidia). They are charac-
terized by shin5 ~moothscales of fairly
uniform size surrounding the entire body,

Fig. 406. D a d head s c a b

(pmfmntalsyellow;
parietal b m ) .
Fig.405. Anomalepis mexicanus (Maritza,
Guanrtcaste, Costa Rica). Photo: M. Franzen

Anomalepididae
The family AnornaIepididae i8 made up of
four genera CAnonurlepis, Helminthophi$ l'he genus Anomalepis includes four spe-
Liotyphlopq and Qphlophia; the last not cies EKOFRON 1988a1, one of which occurs
in Central America), with a total of 15 spe- in southern Central h e r i c a . Anornatepis
cies. Little is known about the habita of micamus is oviparous,as verified by the
these small underground-dwelling snakes. dissection of a femala, which contained
In Central America, the genera two eggs (KOFRON 1988a).
Anomalepis, Helrninthop his, and
Liotyphlops are each represenkd by a
single species.
Further Reading
TAYLOR
1938,DUNN 1941,~ B &B6- 1966,
K Q ~ 1988a,
N

Fig. 408. Llotyphlops albirostris (Isla Pachaca,

Pearl Islands, Panama), Phob: C. W.Myera

HeImin tho+his frontal i.s (PETERS18601,

Monata, M.Wh.Berlin 1860: 617; type
Z d t y : Cogta RiEa, '
I%to 160 mm. Central
Coda Rica and southwestern Paaama, 90-

i 1440 m elevation, ~
Fig. 407.Helminthophi~
Costa Rica).
$2lett
ff'~g:t:$g, ~ 9 ~ ~ " , i n &
~ ,-,mRC H 1997.

Lio#yphEope .-
I
- -
Eielminthphi@
L According ta the most recent revision of ,
Little is known about Helminthophis fmn-
telis, other than that it lives as a secretive
Liotyphlops by DXON & KOPRON(19841,
this mue represented in Central i
ground dweller. lt is distributed in
em Central America.
America solely by the species L. albiros-
tris. The other seven species of thia genus i
are di~tributedin South America. 1I
Liotyphdops albimatria (PETERS18571,
M a t s . Akad. Wm. Berlin 1881: 402;type
locality: Pa-. TL to 223 mm. Southern
Central America (mtheaetarn Cogta Rica,
Panama) and aorthwesCern South America
(Ecuador,Colombia, Venezuela), as well es on
C u r w o (Netherlands Antilles), near sea
lme1 to 1640 m elmation.

Further Reading
DIXON& KOFRON1984.
Leptotyphlopidae
The family Leptotyphloddae contains two
genera &ptdtyph~ope-md Rhimkptue)
with ra total of 80 species worldwide. TPlrO
spciw of Leptotypht~psw m in Central
AmRri68.

The taxonomy of the Central American

Lep-to@phEops is poorly undsratood.
I I O ~ (2001a,b)
R and SAVAGE(2002) treat- %if. 409. LeptotwkEo~agoud~ti.ii(Isla de
d kpto&phlops rtamlis and L. abr, re- Guan4a*Hondwm)*
epectively, aa valid speeiea. However;these
decidona appear to be d y weakly support-
ed and I prefer to not recognize them taxa
until a comprehensive review o f the
Central hesimm Lepfo&phlops has been
published. hpt~typhlopsmealis has been
described baed on a single specimen &om
Managua, Nicaragua, which differs &om
the widely distributed species L. go~datii
by a ~ h g l distiz@sh&g
e scale character,
which might a180 bs, due to individual I
L
anomall~r.Shce t.n m a l i ~was first describ-
ed over 60 years ago, m W h e r apecimem
that could b w~igeedto this t m have
been found. Leptot$pfrlope goudutii is the
most common kptotyphtops found in I
Central America. These not even pencil- Rg.410, L e ~ t W ~ M o o pifoudofii
a be famd
thick makes have a bright yellow spot on * nests.
both the; anterior and the posterior end. As
the very mall head is not well differentia-
ted from the bodx it ia difBmIt at fir&
glance Q d i ~ t i n ~the
s h heed from the tail
iip, These blina snakes live secretively in
rotting logs and in the mounds of termites
or ants. which make UD the bulk of their
food intake I K a m i498fl.Them little
snakes moat o%n eat their prey whole.
However, they aleo have been observed
5wk.ing out the mfi W y conhta of ineecte
and leaving behind empty exoskeletons
(
S C m198Sc). Leptotyphhps golldotd'i are
oviparou~and produce clutches of 8-12
egga (ALVAREZDEL TORO19831,

Calima-and ~eracrus;, Mexico, throu h
Central Amerira to Colombia and ~ene&
9ea level to 1800 m elevation in drg hmst and
wet forest.
Leptotyphlops maemkpis I P ~ M18671,
Momts. Akad. Wiss. Berlin 1W-R 802;
locality: Cmacss and Puerto Cab%
Venezuela.ZZ to 400 mm. Panama and north-
am South America.
gtly to Leptofyphlopa
1 a Without: ale longitudinal fines CPTg. 4llh
supaJarB tw large w p d m ~
...
and fmntel Leptdypirhpr nuwmkpts Key to Typhlopidaer
b Pale lon@rih&nalUna preeent Pig. 410);
eupraoeulam @mewhatlarger than pre-
frontal and b t a l ........................
......
..........................Leptofypffacspa m
Fig. 412. Lateral head scalation in nphlops
and Ramphotyphlops (rostral brown, prefron-
tals yellow).
The family Typhlopidae is repmentad by
two genera in Central America, one of
which ~Rampltotyphlops).haa been intro-
duced in several countries. The species ia
this family only have teeth on the upper
jaw and not on the lower jaw,u&e the
Leptotyphlopidae and Anoaalepididae.
Similar ta them species, however, they are
also small,worm-likemakes that live very
samtively underground. Their dorsal and
ventral ~ctllmme shrdsimilarly ~mgothand of
eqnd size,
.
1 a More than 346 dorsaI scalerr between
rostral aad spike; scale suture between
nosttil roatral
and on a diagonal (Fig.
4m,c3...................................
i
b Fmer than 245 dorsaI scales between
rostral and tail apw suture between
nostril and mstral almost vertical (FQ.
41Pa) ...........,..........,. ....
Rompb&phbps
Fig. 413..Dorsal head scalation in Qphlops
costarkensis (rostral brown, prehntala
yellow).
 The wus Fyphlops is found throughout
the tropic@and subtropics worldwide. In
Central America, there are four species of
this genus, one of which (T stadelmuni)
has only recently been resurrected
(MCCRANIE & WILSON 2001b). These blind
snakes live very secretively buried under-
ground,where they feed on termites, antar,
and other small insect^. The Central
American species of this genus are avi-
parous.

Fig.414. ,.-. t p b Q p h l o p brcarninlcsl.

Photo: R.D.Bartlett

-----
L'

Ramphotyphlope braminus is a species

native to Southewt Asia, but has been
introduced throughout the tropica world-
wide. In Central America,this speciea haa
been reported in Mexico,El Sahadm and
Guatemala, where it has recently become
indigenous I D n r o ~& I~ENDRICXS1979,
1996, DUERASet al. 2001). It is assumed
that t b small snake is t r m s ~ o d e dinto Fig. 416. r5phEops stadelmani (near Quebrada
foreign countries along with the soil on Grande, Go-, Honduras).
plants. As it is parthenogenetic, only a
single female ie required to establi~ha new
population (NUSSBAUM 1980). Rurnpho-
typhlops braminus produces clutches of 2-
3, possibly up to 6 eggB (NUSSBAUM 1980).

RarnapWpMapr br~laniruca( D A ~
Hist. Mat. b p t . WI: 279: type to
Vizagapatam, - India. TL - to - 178 mm.
Southeast Asia, Mca, Madagascar, Z n U
Owan, Hawair, South P a a e and AwEralia;
introduced in several locations in Central

Further Reading
DIXON L WDRI(SILS~ 1980,OTA
1979, NUSSBAUM Fig. 410. Qphlops rnicmtomus (Quirl~w~aRoo,
et al. 1991 Mexico). Photo: H.Bahena B.
 ; %P~LP@ I

............................ .................................
1 a 20 d m atm3db&
mhlope cogt&nsSa
b 16slcalwat midbody .............................
B
B r 1 5 6 0 ~than
~ 470 dorsal scales between
r a ~ t r a l dtail spike,; e ~ b o e u -eat
l~
........................."rgphtopm c & ~ 1 & g k , m
b F e r n than 480 d ~ males d between
&d rrrrd tiail epib; mahcdar absent ,,.3
S a 861441 doreal aoaIee betweea mdml aad
tail qike; body with bmm a p t s ..,....,.,...
.....................................
QphaQpf &rn&
b 841-389dorsral s d betmex~ ~ mtral mid
tail 8pike;kmdy safid mior pink I[ialifiirl and
pale b m n (inprar~mative] .........,........,,..
.............................!Q@llqpeskrc%8isrtaaf
. F ~ e r ~ a ~
V U 1978, D ~ D NdB HENDRICM1879,
MCCRANIE
& WILIK)N2001b

Macrostomata

i
According to McDumm et al. (19991,~lix
fmnilies of early snakes (e.g., Boidae,
- Bolyeriidae, Loxocemidae, Pythonidae,
Trapidophiidae, and Xenopeltidae) are
included in the Macrostomata.

Boidae
The species of the family Boidae are wide-
ly dietributed in the tropics and subtropics
of the Old and New wad, ~haracterbtic
of the b i d s are the head, which is clearly
diaerentiated h r n the neck, vertically slit
pupils, remnants of the pelvic girdle
(which in some species is recognizable by
externally visible cloaca1 spure) aa well as Fig.417.B w comtrictor (Bartola, Rio San
the presence of functional lungs on both Juan*Nicmaf~a).
the left and right side^. All boids are non-
venomous and kill by constriction, using
the muscular coils of their body to anffci-
cate prey. The jaws we equipped with ~urt,hw &ding
stmng teeth. All boids are live bearers. KLUGE1991,ZAHER
1994
 Fig. 418.Boa constrictor (Isla de Guanaja, Hondurae).

b y to Boidae of leaf litter among the buttress roots of

1 a All, awlerr on dorsal surface of head small large trees. Boas are not too particular
and irregular, no enlarged d e a on the about their vertebrate prey and will con-
I
snout; no labial pita ..........................
Boa
aume any lizard, bird, or rnammnd they
b Snout wifh several enlmged ahields;labi- can swallow. If they feel threabned, these
II al pits present ................................... Z
well-armed snakes will hiss with their
2 a More than 246 ventrals; more than 66 sub- mouths wide open, and they are capable of
caudaIs;usually 8, rarely 2 lmeals ............
.
. inflicting bloody and painful injuries with
.................... ..,......~..,....... Corrallue
their powerful jaws. Often pieces of the
b Fewer than 240 ventral&;fewer than 66 long teeth will break off and remain
subcaudals; usually one large loreal, sel-
dom 2 Ioreals ........................... lodged in the wound. Boa constrictor is a
EpicFatea
live bearer and produces litkers of 10-64
young ( G R E E 1983).
~ On September 2,
-
boa
1897,I di~covereda female Boa constrictor
slightly more than 180 cm in total length
in the hollow stump of a dead coconut
Boa constrictor is potentially the largest palm on UtiIa. The snake was coiled up
as well aa the best-known snake in and had obviously j u ~ tgiven birth, as
CentraI America. It is a large, heavy- there were 13 newborn snakes, with the
W e d snake that can be found both on the remains of umbilical cords, in the back of
ground as well rn in the branches of large the hole. The total length of the young
trees. Although it is predominantly cre- boas was between 49 and 52 cm,their
puscular and nocturnal, it is occ~ionally weight around 62 g.
encountered ouhide of its hiding place8
during the day. Typical daytime hiding
places include hollow trees, as well M p i l e ~
Boid

h a comtrietor L ~ A E I J 1758,
B Hptema
iatwaeI ed,10:215;type locality: *Indian[ia
%omllras annulatus (COPE18761, J,Acad. Nar
Sci. Philadelphia (2) 8: 129; type l d @ r
I
error]. TL to 4450 mm ( W A T R L N ~ - C O&L ~ Co~taR i a . TL to 133.6 mm. Didunab &om
LEENDERS 2003); however, most apecimene mstsrn Guatemala to aouthwatern Ecuador,
meaaure lesa than two meters). Tamaulim sea level to 400 m elevation in rain forest.
and Sonora, Mexico, to Peru, Bolivia,.&d
Argentina, as well aa in the Lesser Antdlee, Corullua mschenbsrgerii (COPE 18761, J.
level to 1000 m elevation in Bry forest, Acad, Nat. 5%. Philadelphia (2) 8: 129;
wet forest, and rain forest as well as man- locality: Panama. TL U, 810 mm. So%

r es. In Cenkal America, the sub~gecfes

oa c. imperator DAUDIN 1808 (Mexico,
Central America ~xc1ucling Toboga Island,
&ma, and northweetern South America)
md Boa c. mbogae (BARBOUR 1906) (%bag
kland, Panama) occur.
M e r Reading
S ~ L1936,LAZELL 1964,GREENE1983,W ~ O N
& MEYER1985
westem Cmta Rica to Panama, as well BB
icross northern South America east of the
Andes, sea level to 1000 m elevation in rain
Key ta Coraldue
1 a Nasal shielda eeparated from one another
by rostral; scales poetmior to n d shiel*
not enlarged; fewer than 90 subcaudals
....... ..
... ........... ...,..,, Carrrllur a n n u h
b Nasal shields in eontact with one mother;
scales posbrior to nasal ahielde usually
distinctly enlarged; more than 90 aub-
..
caudab . .....,.Camtlus ~ b n b e r g a r i i

g.419,Snout d a t i o n in CorabEus (nasal

~ieldrsorange).

There are seven diflerent species in the

genus Corallus, all of which are crepuscu-
lar and nocturnal tree dwellers in rain
forest mas. The two Central American
species are extremely variable in termls of orailus ennufaius
coloration and pattern; this is particularly
the case with Gorallus ru~chenbergeriii.
Corallas species will consume frogs,
lizards, small mammals, and birds, which
they kill by constriction. Corallus ate
usually very aggressive, and a bite &am
their particularly long front teeth can pm-
duce a deep wound.These snakes are live-
bearers, producing litters of 7 t
a 30 young
(Ross& ~ ~ A K Z E1990).
C
 (Cdrdoba, Colombia).
Photo: M.Lundberg

Fig. 421.Comllus m h e n b e ~ r i (Rio

i Sierpe,
Cwta Rica). Photo: R.W. Van Devender
kh,

Fig. 422. Comllu~annulatue (near'Ibrtuguem,

Limbn, Costa Rim). Photo: M.Burger

Fig;. 423. Corrrllue annulatus (Limbn Province,

Costa Rim). Photo: R.W.
Van Devender
 Only one specie8 of the genus Epicrates is
found in southern Central America, E, cen-
chria, whose common name, "Rainbow
Boa", comes from its iridescent coloratian.
T t is a slim, arboreaI, crepuscular, and noc-
turnal specie@found in the rain forest. It is
a livebearer (15-28 per litter; Ross &
MARZ'Ec 19901 that will eat fiogs, bards,

-
and small mammals.

F p h t e a m h * W A E V B 1,UW, UJ ~ ~ a a
Naturae, e& 10: 215; type localitg: Suhm.
TL to 1600 mm. Coata Rica m d Panama, ae
well a~ widely distributed thmugh tropical
South America; sea level to 500 m elevation
in rain foreat. The subspaciea E p i ~ ~ t c.
maurus GRAY 18498 o m s in Central
America.
es

Ihg.
-
426. lolonmus bicolor fmm Costa Rim
M L bioodor h m Mexim (Playa A d ,Osutacal
Further Readinn
KLUQE1989.

Fig. 424. Epicrates cench Guangca~b,h t a

*I. Phc R. W. Van Devender
178
 LittIe b known about the two erpeGies of
tbe g e m aturchyboa. InEcuador, together
Tfie mexican 'burrowing python with R. SEIpp, I fomd a epecimen of !l'.
(1;Qlt.Qcemm i3Wor) i8 the only species h bouhngen' that waa lying well eamorzfla-
the genus. If,lives in dry foreat and ~ivan- ged and m~tiodesson the ground duxing
aa habitat and can b sesn hoth drnringthe the day. The animal behaved quite d d y
day as well ae at nigbt outside of its hiding: and made no attempt to flee. Rather, it
place (parsanal observation). It digs seemed to &iffen into whatever shape it
undeqgmund pmeagea with ib F-aismd and w m manipulated.The species is live bear-
pointed snout m d hunt8 whiptails ing and pmduce~6-7 young per kittar
bbpidmelis) and small mammals. This 1Ito85 gt hlkRZEc 1990).
s n d is also well known .far sumwefully
haelring down and raiding iguana nests ~ l k h p b o abau€mgeri PERACCA 1910, Ann.
(Mom 19871,b m m s bicol~rreprodrz- d h ~Zaol, ~ . univ.N8poli 8 (121:1;type Z d t y :
cea ovipamu~lily,with the largeat d m m - j d o w n . TL to 480 enm. Chocd regim of
ted clutob ~lntainiqgfour eggs CROSS L ~,F%ma Paefic Colombia to western
, level to 750 m elevation in rain
B c d o rserr,
1990). Ifomst.

P
at. 6ci. Philadelphia 18: 77; t p loadits:
o m . TL to 1690 mm. Nayadt, M h ,
&ng the Raclfic coast to n~rthwseterrnC ~ t a
'w iaolatsd omreaces elso on the
~anibeanarid$ sea lwei to 600 m dwation
+
1
Jin dry foreat aad savanna.

Further Reading
NEWN & MEYEa 1967

The species of the family Tropidophiidae I

enakes that ~sMbubd in Fig. 426. lhchybaa boulengeri from Ecuador.
southern Mexico, Central America to
northwestern Pacific South America. rtnd
0x1 the West Indies. Contrary to an ehlier --
mangement in which the~emakes were v UngaEiophis mntJnentalis
panam8nsis
included as a subfamily (Tropidophiinae)
in the Boidae, the tropidophiida w e now
recognized as a &&tinctfamily (GREENE
1997, M c D w et a1 1999).
I
Key to Tropidophiidae I
1 a Dormla keeled; more than 27 doreal mws
at midbody ......................... Tnaehyboa I
b Domala emooth; fewer than 27 doreal mws 'I Trecfrm
at midbody .......................Ungrsliophf -
The dwarf boas of the genus Ungrrliophis
are represented by two species, which are
distributed from Mexico a m ~ Central
s
America and into northeastern Colombia.
Little is known about these secretive
snakes. In Selva Negra, Nicaragua, about
two hours after duak, I discovered an U
continentalis on a tree overgrown with
tilland~isias.The animal was flicking its
tongue as it hung about two meters above
the pound. Other observers have reported
Ungdiophis as a nocturnal tree-dweller
that to live in dense epiphytic
growth (CORN19741, These mare attractive
snakes that are completely harmleas and
make no attempt to bite when handled. A
specimen of U continentalis that was kept
at the Senckenberg Museum for several Fie;. 428. Urqgaliophis contimentalis (5elva
year8 ate new-born mice. UngaEiophis con- Negra1 Matagalpa, Nicarrrgua).
tinentalis bears 6-6 young per litter (Ross
& MARZEC 1990).
U lidpkia ~~IFXi~ntali-a 'dlMth~ 11882,
VX N a t u r f d Oe.. 1-
W , ICetdhdBu,
7: la;tpp
TZ to
Quahmala, 482
mm. 8h i a m &dw ta norkbm Niwmgva*
10#-2130 m eleuatioa in foreet and
wrjt
cloud k t .
U ~ a l b p l b f pe w m n s t ' e QCIEWIDT 193236,
Bmithaon, Wac,CoU. 88 (13:12;type locadits:
Cerro Brqja, Panama. TL to 470 mm.
Southern N w a g u a t~ wwtern Colombia,
ma level to 2 UY O B rn devatim h rain f ~ m t
and daud fmt.

Key to Ungaliophis
5 a Rostrd and prefrontal broadly in contact,
therefore, internasals separate from one
another; blotches on body sides oval; 25
dorsal rows at midbody, 15 doma1 rows one
headlength anterior of cloaca ...................
.....................
Ungatiopkis continelptalia
b Roatrd and prefiontal not in contact,
internasals in contact with one anather;
blotches on body eides triangular; 19-23
dorsal rows at midbod& 17 dorsal rowe one
headlength anterior of cloaca ........
.....................Ungalhphb p a n a m m l
Further Reading
Fig, 427. Ungdiophispananwnsie (Tortwguero, BOGERT196881, CORN 1974, VILLA& WLLSON
L i d n , Costa Rica). Photo: M.Burger 1990
Approximately 80% of the known enah
qwciars belong to the fam'iy Colubtidae.
Thh family is diMbutd worldwide and,
on almmt all continents, it is the dominant
family in bms af both the number of ape-
cies and their frequency. The one ezc~eption
is Au&dh, where the cobra-e, haits and
dapid~( I f d y Elapidae) ape mast em-
monly mpreeerlted. There is some contm-
verq regarding the intrafamiJia1 rstmctw
cr;F Colubridae, In Central Am~rica, 64 - - .
genera &am Ws family are known. The
m w eHeB (&dded among 270 enera F i e @.Dipms breuifack (muthem of C&R,
worldwi&) of the family Colubridae dis- Qlrinaa Roq bhico3. Phato: J, C. Lee
play tresrnmdou~marpho1~ddand ecolog-
ical variety, Thus, in Central h e r i m ,
there are ~lnderlybuilt climbing. species
f0rybeh ~pp.,teptophh app., I m W s
spp.1, ground-dwellers with stocky
(zu8d0ft ~ B d e e e p k lOr ~ Vefy
) S h
$hapa lLepCodrymus p~kdedrnue)and
apeciee that b e in atreams or awamp~
(Tretanorhinw ~sigroIuteu~, Hydrornor-
p h w t6ton;~oIool;Aferodia r?iumb~m). Some
epwies maeh W I Btwo ~ meters in total
lm&h CClelh clelia, Lspfaphia akwulla,
M@iaophis naen.tollariue),while others are
fully grown ak 1m than 30 mn (e.g. & o p b

Some Coluhrida have enlarged grooved FFg,430, OzybeIis raeme~ts(Idade Rotit&,

fangs in the baek of the upparjaw md pro- Hondw88)+
duce a secretion that i s mdertrtely venom-
ous (Conophi~lineatus, thybeti8 ~pp.1,
Although the coluKd m&a of Central
h r i c a do noti pose an$ mortal danger to
humane, their bite ean had to serious
locdmd swelling and considerable pain.

Fig.431.Nerodia rhombifem on branches

above the water. Photo: I? Kornacker
 Key to Colubridras
1 a Doma1 surface d head m e r e d with small,
irregular scales (Fig.492) .......
Nbtliqwh

L
b Domal s ~ ~ r f rofmhead c w d with large
~ymmetricalshields (Fig.4381 .................
S
a a k t r a l Bhirld pmtrudipg and pointed (Fig.
456) ......................
... .
.. ..., 8
b IEoetral shield normal (Fig*483); ...............
4
8 a Fewer than 160 WD.W & a d acute
divided (Fig. 43%); no lomd ........Plcimia
b More than 166 ventraler; doacal Fig 482. Nob;h,,.a n~gaaua;r-- the sma&
........
undivided; lored p r ~ m ~ t Phinrophta irregular d e a on top of head.
Photo: R.W.Van Devander
4 a Dowal mdea in wen number of rows at
midbody (campare EYg. 4361 ..................
6
b Daraal scale8 in uneven number of rows st
midbody ..............................................8
8 a 10-12doml raw& at midbody; fewer than
170 ventmls; cloaca1 @cuteusually divided
(Fig.&7b) ..................
b iela h r ~ g l m at
...
s mid-
......ChtrOnius
mom w
E
190 ventrals; cloacal mte undivided (Fig.
437a) ...................*..........,..... , . . . spibbicrr
I a Without longitudinal mental @move (Fig.

b I m d t u M m e a d -8
...
488).................................... .....Ddp~gg
present .+..--..7
- . *&phis
~ i g&-. mlawlomve; n ~ b
7 a ~crdy a l d e r ; hmd 1- and s m e t r f c d shields top of head-
distinct &om thin (Rg.4a4aE; with
h g e pmtrudk eyes; 196-263 ventrde;
........,...,....
1 M178 subcaufala I
-

w
--
b Combination of cltar~ctersdifferent from
that above ................................ ............ 8
a. hmanttxk8
8 a Number d dorsal rows conatant from d d -
body t~ tsloaca, no reduction ................... 9
b Number of doreal rows one headlength
anterior to c h c a at I-t two rows fewer
tfiag at midbody ..,....,,...,..,...,.. .....,.......87 h. Oxymqpus

d
Fig. 435. Ficlmia
9 a Cloacd @cuteundivided [Fig. 437& .......10 FIG 434, Head shape publiu ( h m D W a a
b Cloaca1mtadivided (Fig,M7b) ............
17 in two colubride, ek al. 1870-1909).
10s No anterior temporal .................a.W......... 11
b One or two anterior temporals ...............
l3
1161 U dorsal mwa at midbody Sibon Gin part)
b 1817 dorsal ~ Q W Bat midbody ......i h p h b
I B a k a d sealee,etronglykeeled ........... Ndm&
b Dorsal scales smooth or anly slight1
I keeIed ................................... ., d
13a Unimlor black (adults) or red with pale
neck band (juveniles) ...... CIeIia (inpart1
b Coloration &%rent Prom above ............. 14 Fig, 438. HoHl wmt dorsal scale rmfl; ha
54s Preocular present; more than 210 ventrala example &ma 21 dorsal scale row@ in a
.,.,...........,.,...,.......,..,...,..,,,,.,.Siplbdopkla RWinaea.
 r
b Usuall without preocular; fewer than 210
ventra s .................................................
16a 13-16 dorsal rows at midbody
11
.................
Fi 437.
~3cauials
can be paired
(as shorn) or
........,. ........s t h (inp h )
....................... m-d.
b 17-19 dorsal rows at midbody ................18
1Ba Sixth supralabial not conspicuously en-
larged, not in contact with poBtocular, an-
terior and pposterior temporal; fewer than
40 subcaudals ............................. Atractw
b S W aupralabial enlarged and In contact
with postocular, anterior and posterior Fig, 438. Stemrrhina
temporal; more than 40 subc~udals.......... degerahardtii;note that
...................................... TmpiCMipaas the prenasal is fuaed to
the adjacent internasal
1% h a l scales keeled (Figa.441c,d) Stomria (orange).
b Dorsal scales smooth (Figs.Mla,bf .......18
18a h a s d fwed with bordering internasal
(Fig,438) .................... ... .............. 18 Fig,439. Ninia sebae;
b Renasal not fused with bonking intema- the loreal (orange)
...........................
.......................ao reachea to the eye;
there isl na preocular.
1Ba 17 doraal rows at midbody; fewer than 65
subcaudala ......................... Stenowhima
b 15 dorsal mws at midbody; more than 90
subcaudds ........... 15ymphimur (in part)
aOa Body with complete black rings .............21
b Body without complete black rings,at mmt
dark bands that are not clolosed on the veru-
tral side .................................................
2s
2 l a 17 dorsal rows at midbody; fewer than 150
ventral8 ..................................
Plhemue
b 15 dorsal rowe at midbody; more than 160
ventrb .................................................
22
%%aOne posterior temporal; black rings at
regular intervals; in life, a bright orange-
red dored blotch between each of the black
rings (Fig. 440); in presemative, without
theee blotches, only with black and white
rings ...................................
Scolecophb
b Two posterior temporals; black rings not at
regular Intervale, either a pair of black
rings or rings ~ e somewhat
t h g u l a r l y or
partially dong the vertebral line; without

-
orange-red blotch dorsally between the
black rings in life ...........Erythmlampms
23e 15 dorsal rows at midbody .....................
24
...
-
- '-
.-
--'
I..

\
.: I-"*- - "' -1

b 17-21dorsal rows at midbody ................

2V
2Aa Preocular and loreal present, therefore, at
least t h m scalea between n ~ t r i and --
l eye c. sentidis
..................................10
...................
d. 'lhamnophis

b Either premulb~or loreal missing, there- Fig 441.Doma1 scales

fore, only two scales between nostril and
eye ....................................................
80
a. smooth, without apical scale pits
b. moaith, two apical sale pits
c. keeled, with two apical scale p i b
d. keeled, without apical scale pit0
26a Fewer than 160 ventrale; fewer than 80
subcaudda ............Trimgdopon (inpart)
b More than 160 veetral~;more than 90 eub-
caudals ................. SymphSrmre (in part)
lbsa No preocular; loreal reachea to eye ........87
b Preocular present; no loreal ...................88
27e Fewer than 160 ventrala; fewer than 611
subcaudala ............................ AhZpAiaoe Fig. 442. Hydromrphus concolor;inte-als
b More than 110 ventnls; more than 80 5ub- 223:
pmfiontdS &led,
caudals ...................... .... .....
Elzuliaphia
M a Fewer than 21 maxillary teeth, the poster-
iormost pair grooved; more than 12& vee-
trals or, if fewer than 128 ventrals, then
without pale spots on eupraoculam and
prefrontds .......................
., ,,. Tantilb
b More than 2 1 maxillary teeth, the poster-
iormoet pair not grooved; fewer than 116
ventral8 or, if 116-126ventrals (I:brevtssi-
ma),then with small pale spots on supra-
oculars and prefroetals ............Ta?hHE#ta Fig, 443.&yh&fuWu.8the ; loreal is
29a 19-2-1dorsal rows at midbody .................. the pdmtals (or-) contact the supralabbh.
..............................Rhadi- (in part)
b 17 dorsal rows at midbody ...................SO
SOa Anterior portion of body doradiy patterned
aimilar to coral snakes (paired black ban&
with pale spaces in htween on red ha&
posterior portion of body either
colored h e anterior or predominantly mi- Fig,444.Dorsal
color brown) ...............S~aphiodolstophds acalea in
Coloration Merent than above ............. 81 Diaphomlepk
Number of dorsal rows one headlength
t h BOGERT1964).
nosterior of head two rows fewer than at
&body ............................ ChraphpRh
Number of dorsal rows coestaet from head
.
.
to cloaca ....................................... 88
No preocular; loreal reachea to eye (Fig.
439) ....................
...............................
3S
h o c u l a r pre~ent;loreal m n t or not 84
Anterior temporal present; ratio of tail
length / TL r 205%.......................
EmaIiua Fig.445, BnuEius bifoueatue.
b No antedor temporal; ratio of tail length /
TL / < 10% ........................
Omoiadip~
34a Domal scale@in neck area with apical pits
(Fig. 441b); dareurn of head noticeably B6a Prehatalg not fueed into a singte shield;
paler in mlor than dorsurn of body; with usually more than 50 aubcaudals ...............
.
distinct canthw roatralis ,,Anreehidiacrn ........................
., Rhadinuea (inpart)
b Without apical pits (Fi . 441a); head colo- b hefrontala fused into a single shield or
distinct
1
ration variable; cmt us rostralie not
.............................................
,. 36
not; if not, then usually fewer than 50 sub
caudds ................
Mnretopon (inpart)
35a Usually obvious white line along the f i s t 97a Body with complete black rings ............. 38
domal rows ............................Umtheca b Body without complete black rings, at
b No obvious white line along h t dorsal most with dark bands that are not closed
....
row8 .................... .........................38 on the ventral side ..............................
38
88a Each shield on the dorsal surface of head
black with sharply contracting pale edging
(Figs.446,609); more than 100 subcaudals;
cloacal scute divided ......RhinoboEhryacm
b Head coloration different than above;
fewer than 80 subcaudals; cloacal acub
yndiyided ...................
...,. Lampmpeltds
3Pa Scales of the vertebral row with two keels
(Fig.444) ..........................
Dlaphomhpis
b Scales of the vertabral *ow smooth or with m,
.
.
only one keel ........................ ........,..... 446.Rhinobothryum bovalllii.
40 Drawing: M. Vesely
40a Prefrontale fnsed into a single shield (Rg.
442), or three prefrontal shields p m e n t
..........................................
H~morphlcs
b Two w o n t a l s shields preamt ..............41
41a No loreale, prefrontals in contact with
supralabiala (Fig,443) .......................... 4%
b h e or more loreals, prefrontals not in
contact with supralabiala ..................... 45
4%a Fewer than 85 subcaudals; 19 (rarely 17)
dorsal rows at midbody ...........M m b p t e
b More than 85 subcaudals; 15-17 dorsal
rows at midbody ................................... 43
% 17 dorsal rows at midbody ................... . Fig. 447. Dorsal ~caleein Xeaodon.
...................
.............., 0.wbelis (inpart)
b 15dorsal rows a t midbody .....................
44
44a Cloaca1 acute undivided; 19 dorsal rows 50a 17 dosad rows at midbody ...................
one headlength anterior of cloaca ............. ......................
Dendrophidion (inpart)
.................................Orybelie (inpart) b 19 domal rows at midbody Thanemphis ..
b Cloaca1 s a t e divided; 11 dorsal rows one
headlength anterior of cloaca ..................... Sfa Fewer than 66 suboaudale; dorsal scale
..................................Lephphin (in part) rows run obliquely towards the rniddor-
s u m (Fig. 441) ...........................
Xenodon
4 s Three loreals; an obvious lyre-shaped b More than 66 arubcaudala; dorsal scale
marking dorsally behind the head (Fig. rows run lengitudindy ...................., 52
638); 26-27 dorsal rows at midbody .............
............................. ..... IIPdm#phQdon 52a Dorsum with dark cross bands .............58
b One or two loseals; no obvious lyre-shaped b Doraum without dark emss bands ........Ii4
m&ng dorsally behind the head; number
of dorsal rows at midbody variable ........40 65a 17 doraal rows at midbody ..Mpanorgos
M a Cloacal scute undivided ....................... 47
b 19 dorsal rows at midbody Osyrhopus .....
b Cloaca1 acute divided ...............,........... 56 64a Subcaudala not paired ........... Peeudoboa
b Subcaudals paired ............................ 65
47a Dorsal scales keeled ..............................48
b Dorsal scales smooth .......................,..,...61 =Ba UnicO1Or black (adulta) Or red with pale
neck band (juveniles) .....CMda (inpart)
&a More than 28 domal rows at midbody ......... b Coloration different than above (Figs. 496,
.
.
................................. ............Pltuophis 498) ................................
Drymarchla
b 26 or fewer dorsal rows at midbody .......49
66a 25 or more dorsal rows at midbody .......87
4sa 19-25domal at midbOdY; more than b 23 or fewer dorsal rows at midbody .......61
180 ventrala ...............................fieusten
b 17-19 doFeal at midbay; if 1g dorsal 5% More than 230 ventrals .......................58 ..
rows, then fewer than 180ventral5 ........80 b Fewer than 220 ventrals ........................
59
Ma UswQy 8 su d a b i l s , two bnrdering the
grq
B
mj d m d b atche4 if pr%sen&bmm sl:
.....................
.
.i.
a.... ** ......&3mt&Qhfs
b U~udIy9 awpdabials, k e bmdering the
eye; d d lAatahes-h ..........E W

............................. ...............
43Oa Bodywith dark spots
tepk&?im(inpart)
,,..,...,.,,...,.
b Body with d d langitudinal ahipes .., .,..,
......................-.
Co&pkanee Il.npart)

ma Nostrils directad doraqE8:tedy ................

........,......., ,... ..,. .....*tcu)~rhSw
b N O S ~d i~nBs d laterally ....................tW
b 17 h d rows at midbody ..............,...Ba
&4a Body with n m w & a p d stripss,pmter- FIg. 449. Pattan of d a d bodp in €%nophk
ior m a l o n @ u w striped o~ complete- l h a b W)and t2Phmt~pLLt z m e m R j
ly micolor IFSga,476-4803; resr d l l a r y t~ig?itl. b*: M. Vwdy
teeth obvioudy %al&
.........................
hruhph%asfon finpm%l
............................
b Bady ullimlor -1 in lifefblm in p e e r -
vatfive) OF vdth largg dark a* @T a pale
spot on each dorsal s d e m that M y sur-
f"" ars wtimly spdded (Pigs. a 3 -
6053;a mgldllm-yteeth etppruxbately the
armme #be .....,.........................
IbymmhZtc~ bF a than 195' ventral^; fewer than 140
rubmudallr;'two po%Mars Tl .,. ...........
n a h e amall pmubocular helm the p m u -
b 15 or mom d w d m m one beadlength
....... ............................,,.....*.
laF ,. Col*r
mcrior of the eloerca ............................
a7
8Ba Body without dark 1 o * e a l ak
....................
,ttt~t,tt.t...t..tttttMw~p Is 781a Pupile "R"
d c s l l y em; domm with large
dark b l d e a or broad dark diagmal
b Body with dark bngitndinal &ips
....,.......................................
4ibbdam
ban& .,. ..............
&ptodeflptl I i partl
b Pupila rnued; daxstrm longitudhdp
strspd, unitdm or witb dark diagonal
b 0-1 anterior tempale ..............
7% banda
,.+..,...,....
.........,.......................,....................
R
T2?a Dorsal d e s with apical pib, at le& at
tmhior podon d the bodg; bozly fi
dark di@ ban& or, if l o & t u ~
fi B d g ~ 0 1 0 1 0 ~with 8 w e d dark l@&t~- pad then with Ewo pdm of Iaqghdi-
nd ed s,with the p ~ d d i % N n n i
&al eMpe on pale backgmund (Fig; 4443
..st i..-.r...r ..*............................
t2a110phtsr along s &*cent 7 and B d J
rowst whe ear the lateral stripes rua
%"
b Body dark with 88vwal pals h4gitudinal
etripes ** Q45t) ................
C*p&b
along the 6%%row Z&pAtr ......................
b Dar~lalscales withrrut apical pi* body
70a Mom tJmn 1W veahde; more l;han, 144 with dark langitudiad stxf~ea,-arranged
..............,..................,.... *..................
5 u h u d ~ &three poahdars
Lep&?odry~cuar
WeflentIy than a h e ox unimlor (Figs,
4f3M42) CmiaphhpCen E ...........
b part)

The genus MeZphicoe ie rgpreeented in
southern Mexico and northern Central
America by six species ( C ~ B E L&L FORD &theastern
tion in pineoak foreet.
1982, Cwm & BRODIE 1988).These are
small and highly secretive snakes. They
are found in rain, cloud, and pine forest at
elevations up to approximately 2200 m
above sea level. By chance, while digging a
hole about 30 cm deep, I came upon a spe-
cimen of Adelphicos quadra'virgatus at Pim
Bonito, Honduras. Otherwise, they are
usually found by turning over rotting lags
or decomposing plant material. LWDY et
al. (1966)reported fmding an A. quadrivir-
gatus in the ledlitter st the base of a large
tree. These nocturnal snakes feed predom-
inantly on earthworms (LANOYet al. 1966).
The qeeiee of this genue reproduce by lay-
ing eggs. A clutch of three A. quadrivir-
gatus eggs was found in a nest of tree ter-
mitee about one to one and a half meters
above the ground (PI~ESHIGAREDA
S M19891, ~ ?tvo weeke latex, a snake, 86
&
mm TL, hatched out of one of the em.In
Guatemala, the eggs are laid in the early
rainy aeason (May and June) (CAMPBELL
1998a1.
Fig. 450. Aaklphicos nSgriZatus IChiapas,
Mexico). Photo: P.Heimes
o w -
C i p h b davi CAMPBELL & FORD1982,
.Pap. M w Nat, Hist. Univ. b
tim, 5.6 km
Suatemda,
~~
-
a s 100:3;
l d t y : San Jmge MUM, 1844 m ele-
W 8an J&TL Pinula,
Guatemala. mm,
to 674 DpEo,
185&2186 m eleva-
logicm 4 4 41Q type locality: 3
c a s k m g v , 2100 m elevation,
batween Paxot and Caminibal,
mala. TL ta 521 ram. Southern
uatamal$, 2000-2100 m elsvati-
ifiupciatw Lmca & 8 m lBW,
Acad. Sci, 86): $6 typ lmlity:
north of Zanate Oaxaca,
~ 1 mm7 wmtern r & a adr re
apm la Oaxaea, Mexico, 1600-2000m
on in 0 ~ farest*
0 ~
tua E ~ T H1942a, Proe.
.8: 182; tppe locality: San
jkist4bal ds 1m Cams, Chiapae, Mexico. TL
&I 461 mm. Meaeta Central in Chiam,
,@&o, 2 2 0 0 4 W m elevation in cloud famt
lipld pinsoak foreat.
P Iphica quudrivirgahre JAM 1862, Arch,
ol.Auat. I%. 2: 18,t locality:M h . TL
T
t &I0 mm. Central eracrm to northern
~ k u r a son the Caribbean vereant; in
&asternChiapaa and w a t e f a Guatemala 01
'$he Pa& vmant, Sea level to 1500 m ete
Won in rain h m ~ and t pine-oak h e s t ,
MiChigana:G;*
daxmc, 7 Enn W Cobth, Alta
a, Guatemala TL t o 461 mm. Central
mah I&ntatiar de Cuilm, Sierra de
Pace Minas), 1200-2200 m elevation in cloud
$&restand pines& foreat.
Key to Melphicos
1 a Third infralabial absent or much reduced;
chin ehields greatly expanded toward lip
(Fig.461) ,.,, Adelphiam q u ~ v l r g a ~ e
b TMrd infralabial barely or not at all re-
duced;chin shields not greatly expanded 2 [
2 a More than 42 [males)or 32 (fernale-a1sub-
caudals; vertebral tripe d ~ encloses
p~amedianmale row;venter unpigment-
ad ...................
o
AdolphZoea &zt€m*e
b Femr than 42 (males)or 32 Ifmales1 aub-
I
on median d e TOW; venter unpigmenbd
or not ...................................... . . . S Fig.$62, AtEelphicm qumdriuirgatlca (Chfaw,
Mexiool. Photo: k Ramirez V.
3 rr Venbr unpigmentd or only darker along
the midline; doreal ground calm pale
brown or bright red-omme .....................4
-
b Anterior border@ afvenbil shields conspic- *:
u o d y dark; dored pound color gray a Am~Hfc-- -
h

4 a 113-128(males) ur 123135 (females) van- Contrary to tihe earlier view of the genus
M E ;ratio of anout length / eye diameter Arna~tridium ae monotypic (Wmon I&
1.4-2.8 ................
Addpkicaer nigrlldw MEYE1 1969, 'WILSON & R O B ~ W 1971, N
b 136-138ven- (%malea)(data for males WILSONl98&), the current view k that
not available); ratio of snout length I eye them are two different species, one of
diametm 2.9-5.2 A&Zlmfr&oe b b a m m which k distributed in northern Cental
B a Venter not conspisuowly darker along America (A. sapperil, the other in the
midline; ventrals 120 (only kttown make) or south CA. velifirum) (J,SAVAGE pen.mmm.
128-132 ( f d e a ) ; aubmudala 28 (&5) 2000). This small solubrid is rarely found,
OF 19.22 (fmdes);fkontal at leaet w wide
as long .........................
Adelpkhroe Plrs* thus little i~ known about i b habits. It is a
diurnal p m d - d w U w in the Iwflitter of
b Venter noticeably darker along the mid-
line; vsnkds 120-159 (malea) or 152-142 the rain forest, but it is also famd on oof-
(females);~ u b c 9 . d29-41 ~ trnaksl or 24- fee plantations ( W a s o ~& MEYER1969).
31 (&males);&mtd usually bwr thm Sparse observations on the nutrition cf A
wide ....................A d e E p l t h vempacie v d i f e ~ ~ lindicate
m that frog8 and lizards,
as well as invertebratels,such a~centipeh,
M h e r Reading play ct role in their diet (MARTIN1966,
H, SMITH
1942a, CAMPBELL & FORD 1982, BLANEY& BLANEY 1970, CAMPBELL1998a).
CAMPBELL.$ BRODIE
1988

Fig. 451. Ventral vim of head in Adadedphicm

q u a d d v i v ; noCe the WS@CUOU~)~~large
chin sbiel&.
 Colubtidae

Only a few of the approximately 70 species

in the almost excluaively Smuth American
genus Atractus occur in Central America,
where theee are extraordinarily rare.
Recently, M m (2009)described four new
species of the genus A C ~ U C ~mUPanama
fro ~
and reported upon a Colombian specimen
of the poorly known A. clarki. Two of the
Panamanian epecies are known from the
Fig. 464. Amcastridiuna vebiferum (La Selva, low uplands east of the Canal Zone and
C&a Rim). three are from the Barik. The species of
the genus Atractus are small to medium
size fossoria1 snakes that exhibit shiny
bodies and tiny eyes. The following
a,~~&Ht"-tafi m m WEW@l19U8&AH%, 9, Atractus key is based on MYEM (2003).
B w r . Akad. Wias. Z k 349; type locality:
Guatemala. TL to 726 mm. Nwvo Le6n and
Oaxaca, Mexico, to Honduras, 100-1000 m
elevation, :w.Comp, Zoal. 88: 8; type l d t y : Mine a
s n b Crue de Cana ['approx, T046'N
Amastridium wlifsrum COPE 1881a. Prac. r041W,500 ma according to MYERS 20031
h a d , Hat, Sci. Philadelphia 12: 370; type arihn, Panama. TL to 318 mm. Eaten
local it^^: Cwuyas de Varaguaa, N. Grenada mama and the Colombia Chocd
[= Cmyas, Pmamd, TL to 726 nun. Nica-
rarma to Panama, 200-800 rn elevation. irtractus darknensia MYERS2003, Amer. Mus
Novit. 8591: 16;type l d t y : north end of tha
$emanfa de Firre, epprox. 8°00X, 7To43'Ui
Rey to rlmaetridium 008 m, DariBn, m a . TL to 846 mm
1 a Fewer than 140 ventrala;no loreal ............. Known o d p from the type 1 d i Q ,
........................
Amuehzhzdi~m velifr!mrn Atmtw &ptr?88imlllta Mmm 2003, he^.
b More than 1411 ventrale; loreal premnt .... Mus. Novlt. 3981: 20; type locality: Cem
...........................
Arnaedridium s a p p d Azul [Cem Jefel, approx 9"14'N, 79"23W,
800-800 m,P d ,Panama.TL to 750 mm.
Xnown only from the type locality.
Further Reading
DUNN 1924, WILSON& METER1969, WILSON& &mtw hostilitractus MYERS 2003, h e r .
F~OBINSON 1971,WILSON1988c Mus. Nwit. 3SB1: 22; type l d i t y : "Murti
I.Iydmn,8"62'28'W, 77"64'19'W, about 100-
200 m, N o Mwtl, Darien, Panama. TL to 374
mm. h o a m d y from the type locality.
&ractuar imperfictus MYERS 2003,Amer. Mus
Novit. 5891: 25;type locality: Piedrar-Pacora
Ridge, Pan-, Pan-. Known only horn
t21e type l d t y .
 -
Key to Atnzctzav Chaphphls
1 a Venter pale; lower glcde rows with pale +
08nbem; eye length noticeably greater than
dietanm to Iip ................ m t r a e~rs&i genus C h ~ d m ~ ha hn w g the
ventsr with pig- ~peciesC.xanthochilus, was described in
mmt; no pals drrehea on lower a d e mw5; 1998 (CAhPEELL & SMITH 1998). This d u -
ey.e length lsae than or scarcely greabr brid is a diurnal gramd-dwller fmm b
&an d i s ~ m to lip .................................. I cloud farest that go far ia known fkom only
2 u Neck brown like body, with black trsna- t h m swimem.
veme liner nat greatly wider than on rest
of body; eye very am& contained about
three times in loreal plah ................
........................A h a t r s depms~hellus @~brrrw h i n a p h i @ ~ ~ n t h o c h e iCmP~~Lh
m a , HI M O ~ ~MM(2): 2 1 s
~~s &
b Neck bIaek with pale rings, or +Kith black Fli~ 6 irm ~%es~lncaMirtmda on to
bands wider thwe dm Vega L-a, 2300 m devatim, Baja
P D ~ ~ O I : ~ size modera% w p m , Gaat-. TL to 590 -, Wesbm
going1esa than 2.6 tima in l o d plate 8 on of Sierra de lars Nnae in central
8 a Snout &ed in profi1s; 1mdappmaohing
or touching internasal; labiala mostly
#? atemala, 1M0-2800 m elevation in cloud
bat.
black ..,.*,.,.,..,,,,Al.redtae ~ ~ ~ r m d z i a
b Snout bluntly pointed in prome; lored
well ee-pareted from internaaal; labials
mostly white ................*..... ............
. , . . 4
4 a Neck witti black ~arldles;rpstral plate
naffowly vkible in dwml view; eye length
lem than 80% oddiatance to lip ..................
..............................LktmtuaWetlemtie
b Neck black with paIe bm or ringlike
mar-; roetral plate well vbible in dm-
4 mew; eye length about equal to
distance to lip ...... Atl~~etus #mperjktrae

Further EeadLng
D m & BAILEY1939, MYERS2003 Fig. 465. Chapinophis xunthocheilrrs (near
Chilasco, 1856 m,Sierra de lag Minae, Baja
Verapaz, Guatemala). Photo: E.N.Smith

.
"
Fig. &%, 3uvede Chisoniu8 granokiaq~anis
(Bartola, Rio Ban Juan, NicmagueL
Fig. 467. Chimnius exolatus (Pavones de
Turrialba, Cartago, Cwta Rica).
Photm R.W. Van CLevender
u
Fig. 468. C h i m i u s carimtua (Prov.Punta-
sen=, Costa Rim). n o t o : R,W. Van Devender
Chfmniae
The specie%of the genus Ckimntus are
large, predominantly tredweUbg* harm-
lees colubrids. They are the only
Neotmpical makes with 10 or 12 domal
scaie rows at midbody. Only one s p e c i ~in
this genw has advanced ae fw north as
Honduras, whereas the other 12 specie&
are distributed in South America and in
southern Central Amarica, where three
species occur in Panama and Coeta Rica.
For the most p&, they live in primary rain
farest, but can &O be found in areas with
m d a r y growth. ?%en they are haadled,
they wiIl usually respond with gaping
mouths and defensive strikes.
Large eyes with round pupils are charac-
terietic of these diurnal colubrids. Their
primary food source ie fhgs Emoatly tree
fmga), followed by lieards and, lees fke-
quently, sal8manda-s ( T k n o ~1951, DEON
& Sam 1977, DUELLEXAN 1978, DIXONet d,
1995).At night they can be found coiled up
d e e p in shrubhl and trees. AU species in
thie g e m are oviparous and produce
clutches of 415 eggs (DDEON et al. 1993).
Peg Lo ChirO?mZ~#
1 a 'LO d a ~ rows d at midbody .....................
..
............,..... Chimnfrsd@*-is At peewnt, three species of the genus
b 12 doma1 rows at midbody ........,,.....,.,... CkIia are recognized in Central America
B a Dmd color olive brown with w h i t s or Qf these, 6: cklia ie di%Ferentiatedfrom h-
pilow spots on m a ~ tof the doreal s d e ~ &ex two species by the presence o f 19
............................Cktmniue oadawbun than 17 dorsal rows at midbady, a
b Dorsal mior unimlor olive brown or dive rather charader that is poasibly more variable
green {adult specimens) or with pale nar-
row cmse band5 tu&g diagonally than previously thought ~CWBEU
(juvenEle3 ................ C h i m i a e exoktu8 19988).

CJdia clelita is a very large snake with a

relatively calm ternpermtent. Juveniles
Fig. 459. Adult have oompletely different caloratiori (red
Chironius gran- w i t h a yellow neck band) than the adults,
disquamk which am unicalm ahioy black. The eye8
Photo: R.w.
Van Devender are sglall md dark and the head is not
dearly distinct from the neck. C. cEdk
fee& primarily on other makes, but also
an liaclrds (predominately badisks md
ameivas), small mammals, and p d b l y
Further Reading frogs ( S ~ m m 1999). Other patentid prey
DMONet aL 1993 items inelude large pitvipers, as verified by
the case of a 180 cm long B~thmpaaamr
that was swallowed by a 2 meter long C,
cldia (CAMPBELL 1998a). C. cielia ifi pre-
dominantly nocturnal; thue, tthis specieti, ie
seldom ~mauteiide of its hiding place
duxing the day (~TAFFORD 1BsQa).

I b No apical pita; ventral. 202-20'1 (males) or
Fig.460.Adult Clelia clelia (upper Raspado
River*Maya Mountains, Behe).
~ h o t oP
: .Stafford
Gklia clelk (DAUDW 18031, Hist Nat. Rept. 6:
330;typa locality: Surinam.TL to 2MH) mm.
Guatemala and Belize to northweatern
Ecuador as well as to Argmtirra, sea level to
1000 m elevation. In Central Ameriea, the
nominate aubpPciea occurs.
Clelia epuatoriana (AMARAL 1924), 5.
Waahbgton Acad, Sd. 14 201; type locality:
Guayaqyil, Ecuador, TL to 1575 mm. Costa
Rica through Panama and C o l d i a to north-
westem Ecuador.
Cklia acytalinu (Corn X867b1, h. Ad.
Nat. Sei. Philadelphia 10: 820; type locality:
Near Tabmw, Mexico. TL to 1600 mm.
Southern Mexico, Guatemala,and Belize; sea
h e 1 to 1000 m elevation.
Fig. 461. Juvenile C l e h scytalina (Tapanti,
Cartago, Cmta Riea). Photo: R.W. Van Dwender
Colubridae
Key to cte1i.a
1 a 19 dorsal rowe at midbody ....Clelia clelia
b 17 dorsal rows a t midbody .......................2
2 a Two apical pits; venbais 203-216 (malee)
wr 218-221 (females); subcaudals 78-90
(males) or 75-80(femalels) ...........................
.........................................
Ck12a acytalim
200-217 (females): subcaudals 76-80
Furthe* mading
~ ~ ~ S c m 1983a, ZAHER
GWER 1 9 6 6 , B 1970a,
1996%
S T M F 1499
~ ~
Coluber constrictor is a slender, fast, diur-
nal hunter that prefers savannas and open
dry forest; it feeds on insects (e~peciallyas
a juvenile), lizards, frogs, and rodents
(WRIGHT & WRIQHT1957). Thia snake
reproduces oviparoualy, producing clutches
of 2-16 egga, which hatch in about two
months (Frrc~ 1963,19703.
Coluber eonetrkbor LINNmua 1768, Bysterna
Naturae, ed. 10: 216; type localit *America
septentrionala" TL to 700 mm. fhributed
widely acmes North America and Mexim; in
Central Amedca south to Behe and north-
I
Fig. 462. Colukr c. constrictor (North
Carolina, USA). Photo:P. Kornacker
 Fig.463. Conipphanes aEvareti (ElLaureI,
Chiapas, Mexico). Photo: A. Rmirez V.

I
hlubermetnrctw

-
The species of the predominantly
Neotmpical pnua Coniophanes are @rnaZl Abb. 484.Caniopham~bipunctatus ( C a t e m ,
to medium size mlubd& that are highly Veracm, Mexico). Phota P.Heimee
secretive. While many speciea are particu-
larly nocturnal leg., C. bipunctatus, C.
guinqwvbttatus; see C ~ L l338a>, L at
least C. fimidena, C. i r n p d i s and C.picei-
&ti$ are also active during the day MYERS I
1969, personal ebearvation). They are
included amwg the rear-fanged (opistho-
glyphous) snake& but are usually not
aggressive. Caution is still recommended,
a~ their bite can lead to locdhd ~wdling
and pain (BROWN1933). Most species are
gmund-dwellertrs in forest areaa, while
some (e.g. C. bipumtatu8, C. quinqwvitta-
tw)am bslieved to be semi-aquatic / h ~
1996, CAMPBELL 1998a). The Coniapharses
epwies are k n m to feed on fmga, eala-
m d e r s , lizapdg, and aniilfee, aa well as . Coniopkrae.ghi&m [near
~ i g485.
reptile and bird e w . They will algo eon- Quanagasapa, Errminth, Guatemala).
Fig. 466. Juvenile Coniophanes imperialis (Isla Fig. 467. Adult Coniophalaes imperialis (Ida
de Utila, Hondurad. de Utila, Hondurwb

sume invertabrates, such as earthwarma

Caribbean maat except fir the Tucatkm
and ineect larvae (MMTON& S M1960, ~ Pedmula), C o b hame fl o b d e t m MIHTQN
LMDY et al. 1966, S6m 1985, PUTT &
RAINWATER19981, To escape predators,
& smm la60 A*d* Bwnw Aims md
TurrSdba, Puntaranma, Co&a Rica), and
Conwplrame j5 pu~b~t&darisC m 1880
theae colubrids will throw piece6 af their
tail if they are mized. This explains why
many specimens are found with incorn-
(along the W C
&&&#@38
cloast) occur.
f
hJWld&bf m c o l f i 1 8 8 9 ) , 4
plete t d s (Zuct et al. 1!3791. The species of Bmm Raptiha of the B d a r y : 25; type
this geneta are oviparow and lay 1-10eggs
per clutch, usually under rotting logs or
roots (H.$MLTH 1940, WERLER1949,
L ~ & PECW
Y 1953,PITCH 1970,ZUG
et a]. 1979, CAMPBELL 1998a).

Cmia k n e 8 dvami CAWPBELL 1889,Proe, 1- 885; type l d t y : O m Pirre,

Bid &OF. Washmfm 103 (U 10861 type la- PaeaEa e d of tha 600-lMm elma
d*: 11.3 km E E Tso ' a, 2078 m eleva- tion, in montane rain forest and dwd for&.
tion, Chiam Medm. 5-21mm. In tlre
a~ea of%phm, Cbis Maaria, 2010-2138
m s h t i o n in pineamst.
ConiopFsgnee bipunctatw ( O m 18581, 179; type 1caEt;v: MBrida,
Cat. s l l 4 3 h Brit. Mus. 1EIBB:SB; t;grpe locality: TZ to 376 mm. Northern half of the Yuea
u n k n m . TL to 760 mm. SouthernV e r m r ,
Mexico, to Panama, lea l e d to appmxb~tte-

II
ly 400 m elevation in rain fore& wet forest Conbopb~e8 keiuinis COPE 1870, P m .
and dry fcrreak Arne. Phil. fw. 11: 14% type locality:
Chihuitan, Oaxaea, W m . TL do 570 mm.
Coniophlsss /%B&IWB ( G T ~ w 1~86 R,Cat.
~81 Tanmutipa% and aUerrer0, Mexico, to Coeta
Sealawe Brit. Mu. 1- 86; type lwaliQ: &a, ma level to appmxhately 1000 m ele-
ME&&. TL ta 795 mm. Soud~fllVerac~ur vation ht dry forest.
and Omaca, Mexico, tbm h Central
Amalca ta Colombia pad Edz& sea level Coniopiramn qulngueufttutus (DuwiBm,
Z
to approximate 1809 m elevation in rsin
mid wet &reeta tlentral America, the sub
~ ~the
speEiee ConwpIbOne~f: f i s (along
EWN % JhnaW~186IL1, &p. OBn. 7: $76;
toe looalitv: unknown.. TL to 600 mm.
 Guatemala, 8ea level to a p p m d a t e l y 200 m
elevation in rain foreat.
Coniophenes &midti b u y 1937,Om.Pap,
Mus.h l .Univ. Michigan 1; type locali-
ty: Chichen I t d , Yueath, Mexico. TL to 400
mm. Northern Chiapas, Merdm, w well as the
YucaMn Peninsula,sea level to approximata-
ly 900 rn dwation in dry forest and wet
f0~5t.

Key t o Cm36phmnee
1 a 17 b a d rows at midbody .......................
B
b 19 or more dor~alrowa at midhady ....,.....
S
2 a Mom than 70 subcaudah; pale, dark-bor-
dered stripe running along side of head at Fig.469. Lateral head in Cont"ophamcs.
eye height h r n snout tip to posterior tern- Drawinp: M, Vesely
poral ecalq pale neck band, which may be
incomplete ventromedially, present ...........
......................Coniophanes meridenus
b Fewer than 60 subcaudals; pale, dark-bor-
dered atripe rumin don side of head at
mouth height h m Lhinb lower eyelid to
neck; without pale neck band .........
..
.......................... Cuniophanesjoantw
3 a More than 150 ventrals .., ..,.,. .,., 4
b Fewer than I60 ventrals .rlu.*i..,)rr+~tt~~;~B.... 0
4 a At outer end of each scale a large dark
spot, such that a symmetrical row of spots
runs along each ventral edge; subcaudals
63-70 .....Conlophanee qwinqusvittatros
b Venter
4. r
mimlar pals colw; aubcaud& 78

5 a Lateral dark longitudinal stripes at sider

4-15 scale8 wide, lower e& color ~ h a r p l ~
contracting fmm color of venter .................
........................ C Qf0phm&?8
~ pkshttfk
b Lateral dark longiturlinal stripes 0.6-1.6
scales wide, lawer edge color grad- into
B C ~ W
v a t r a l colmatian Canliophnlse~
8 a 21 dorsal rows at midbody ........................7
b 19 dorsal rowa at midbody ,.,,.,..,.........,. 8

Fig. 468, Ventral pattern in Coniophanee.

196
 Colubtidae1

7 a Venter llnicolor pale or with irregularIy

di~tributed6ne dark %pots(Fig.488a) .,,,,..
............Comiophanee fenatdm tin part)
b At outer end of each ventral wale a large
dark spot, guch that a symmetrical row of
spots runs along each ventral edge (Fig.
488b).............C o n h p k s blpumtahce
3 a Dorsum unicolor brown without lonpitu-
din4 stripes ........Coniopkanes alvamzl
b Dorsum with longitudinal stripe ............B
9 r Pale, dark-bordered stripes running along
aides of head &om snout tip above eye to
pogterior temporal acales ...........................
......................ConZophalces impsrdcrlle
b Head markings different than those above Fig. 470. Contopharse~jmnae (holotypel.
(Fig. 469a1, if a dark-bordered stripe is Photo: C,W.Myers
present, it is limited ta the temporal *&on
............Csnlophanen fieidene (inpart)
Further
I 3 m u 1939,McCoy 1969, Mmm 1969, S m
1985,CAWBELL1989,SOL~RZANO 2001.
- r--

h a
Fig. 47 1. uoniophnes piceiuiktzs r v o ~ c h
Maderw, Isla Ometepe, Nicarama),

1
L-
.Conjo@n.8plreiv&s
Conmphanes schmldli

Fig.472. Coniophuws schmMti (nearXpujil

Cmpeche, Mexico). Photo: J. C.Lee

Both e@es of the gmtu1; CampAi~~ t r e
m d m g h e d , vexy irritable re~rfaaged
snalres, with a very painful bite. They are
d i m a l and hunt mostly liliwd~
(Seeloporus and Cnernidophorw) and
other apgtiaa of andtea*whigh are inj-d
with powerful venom when bitten by the
long rear fangs of these predators.
Although the bite of a Conophis iEl not crit-
ically dangemug to humanrr, it can still
Fig, 478 Conophh vittatm (near Mixtequilla,
Oaxaca, Mexico].
cause painful localized swelling and bleed-
ing (MERTENS 1962c,WELLWN 1463,SCOTT
N LEE1996).Captive-
1983b,J O ~ S O1980,
held Conophis will eat frogs and toads
(M~EMAN 1944, as we11 as ~nakea
(Thammphis and Stareria). Rodents am
abo part of the diet of these snakeq cte
demonstrated by the fiading of a mauae in
the stomach of a dhsected C. Ihmtus
(WELLMAN1963). Both species ropmduee
ovipamusly, with a clutch size of 5 ta 10
egg8 ~ V A R E ZDEL TOR0 1960, LEE 1898,
persond observation).
Conophda liwrstwr (DUMI~RIL,BIBROH&
DLIM@#ElW), Erp. a n .7: type locali-
Mdm. 'I% to 1170 mm. Oaxaca anrl
l e d to appdmatelg 1000 m elmtion m
dry b e s t and mavatma. In Central hitrica,
the aubspeeiear CompBie L mlrwsm COPE
1867 & c a m F%mbula) and Conophh I.
Saarmr 1842b i m u t h m m Qaxaca,
M d e o , ta Coeta Riea) occur. Conophia pd-
COPE 1 s S g k ~ d t u b e a a
of C, l h a t u s according to WILSON gt Mima'
(1986).
C~mphhvit-Botus ~ E R 1860, % Monatsber,
Brig, Alrad. Wlas. B e F h 618;tgpe Is-
mliv unknown, TZ to &aQ mm. Padfie side of
Mawu fram saut;hern Durango aer far sodl3
a9 Chiapas, sea level to approximately la00
m elmatinn in drg forest and sawmum.
Key to Conophwr
1 a Body with 3 or 4 1cmgituW black stripes,
each at least two s c d raws ~ g d e ; usually
7 supralabials, the ar and 4 of these in
contact with eye ........Conophis vittataa
b Bady with 6 to 10 l ~ n g i t black u ~ atrip,
that are ueually loss than two scale m
wide, or entirely without longitudinal
s pea; u ually 8 supralabials, with the
4 and 6& of these in contact with eye ,...
....,....,......................
Conophin Ilneulrre
Further Reading
H.SMITH 1949, WGLL~~
1941a, 1942b,SAVAGE
1969,SCOTT
1983b
Fig. 474. Conophis lineatus (Guanacaste
National Park,Costa Rica). Photo: M. F'ranzen
included in the genus C'onophis ( W u
1963), but at the beginning of the 1470'8,
was placed in ite own genus (VU 19711,
This fast colubrid lives in dry areas,prefer-
ably close to a eource of water where it can
hunt h g e and lizards.

C-tLia wumannZ ( D m lWTs2

EspeialW7:?1& ~,l%Poerc&
A.saiCaIrnr%.oolbdBoldgee~ Fig. 476. Crisantophis neuemanni feedlng on
M R Q Q & . ~ ~ O @ O - ~ - a race runner (Ometepe, Nicaragua).
hW&I W a w b level to appm Photo: P.van Beest
h W y 1260 m ~lwatj&m
ia dry fbreat anr
&aana*

Further Reading
WELLMAN1463,VILLA1969b,1971,1988.
Theae medium-siaed, slender colubrids me
dbtributed from southern Mexico hwugh
Central America and far into tropical
South America They are diurnal forest-
dwellers that feed primarily on lizards and
frogs. Dendrophidion species reproduce
oviparou~ly;females of D. pernardnuturn
and D,uinifor in lower Central America
have an extended or possibly continuous
reproductive cyde and may produce more
than one clutch per year, Clutch shes
range h m 3-6 in D,uinitor to 4-6 in D.per-
carinaturn and 5-8 in D,nuchole (STAFFORD
in press).

Dedr0pkidi-m pueWncrtunr (Gam 1894),

PFDC. Am.3. Ma*M.lwhXl4- 48; 90%
type lac&@: Zrr Capdeltatie, Bruno
Carrsrs~ea*GO* RTca* TL ta llao -.
Cordillera Central in "Ma Rica and
%dillera de l'damma E'crskIR5m and jn
westem Pauams
 Dendrophidcon pmrznoturn (Cam 1893),
b, Amer. Philos, %c. 81: 548; type locality;
Bonrca and Buano~Airea, Caata Rica. TL to
1130 mm. Northern Honduras ta north-
w~tern Ecnador, 30-MO m elevation in rain
fowst.
Dendrophidion vinitw S m 1Q41d, PIYIC.
Biol. Soc. WashingLon 84: 74; type locality:
Piedras Negras, Cluhmala. TL to 980 mm.
Southern Veracruz, Mexico, to western
Colombia, 60-1300m elevation in rain foreat,

Key to DendmpAid$on
1 a Reduction of scales on doreal side of tail
fro eight to six rows occurs before the
26& aubcaudal scale ............................
2 Fig. 476, Dendmphidwn nwhale (Cuxta Bani,
b Reduction of scdea on the dorsal ~ i d eof upper Raspado River, Cayo District, Belize).
tai from eight to six rowa occurs after the
A
25 aubcaudal scale ............................
8
Photo: P,Stafford

2 a 179-195 ventrals; 119-139 sub caudal^ ....

.........Dendraphidton purccid&um
b 165-166 ventrals; 147-163 subcaudala
.............DslzdmphMion penearina#um
3 a 152-183 eubeaudals; cloaca1 a t e mually
divided; first dorsd scale row strongly
keeled; dark-bordered pale cross bands on
neck, if preeent, leee than one scale row
wide; primary color of tail abmptly paler
than that of body .............................
..
...................... DendrvphMlIon ~sclsa€.e
b 111-128 subcaudala; cloaeal scuta undivi-
ded; firat domal wale row mooth or only
slightly keeled; dark-bordered pale m a
bmds on neck more than one scale row
wide; primary color of tail not paler than
that of body .......DendropiZdZon aimitor-

I Further Reading
LIEB 1988,1991a-c,1996 Fig. 477. Detadrophidion paucimrinatum
(Gem Cacao, C o ~ t aRica). Photo: M. kanzen
 The poorly known genus Diaphorolepis
consista of two species, of which one, D,
wagneri, occurs in eastern Panama.
Nothing is known about the habits of this
species.

DiaphomJ~pbwagmri JAN l W b , Elenim

m w Ofidi: 98, type locality: Andes aC
~dor,TLto600~.Kngwntaoecurir
Ecuador and in ea&sim Panama,

Fig.478. Dedrophidion vinitor (north of El Further Reading

Cop6, CacM, Panama). Photo: C.W. Myera BwEm 1964

Fig. 475, DedmpMdion winitor (Volch d,

Gmmcaste, Costa Rim). Pbota: F! Stafford
Fig. 481.Diaphorokpis wagneri. Frnm BOGERT
{1964).

Fig, 480,Dendroplaidion pesetarimturn ( N o

Patuca,Olancho, Honduras).
 Approximately 30 species are included in
a- pms wtkulata (COPE18681, Proc. Acad.
t. Sd.- Phhdelpbia
-- 24h 136;type l&w
% ~ ~ C U B , 'Costa RimAtactually i
d

n Panamal.
@ to 712 m..Southemtern N i m g u a to
the genus Dipsas, eight of which occur in %eatemPanama ma level to appscutimaw
Central America. They are small to mid- L
l o m elevation rain *st.
size, harmless eolubrida that can be found biedcr (GONTWER18961, Biologia
at dusk a d at night crawling around in ntrali-Americaaa. Re- and B a h W
vegetation, particularly in the rain. They 8; type loCali* Chontales Mines,
feed (exclusively?) on snails and mlugs. TL to 482 mm. N a
to northeastem Cash Rica, sea
These small colubride have been observed byel h approximately 600 m elevation in
flicking their tongues and following the Foreet.
slime trail of snails over several meters ,@peas brwri&cies (COPE18881, Proc. A d
( S w 1989). Slugs are grabbed in the &$at, Soi. Philadelphia 18: 117; type l d t y
middle, pulIed out of the subatrate and Tkcattin, M d c o . 'ST, to 628 mm. Nwtbem
usually swallowed tail first. Ends are also @ i f of the Yucatkn Peninenla (southern
pulled out of the substrate first, and held Qhmpeche, Mexico, and central B e b north-
&el, 8ea level to approximat& 300 rn ele-
tightly in a coil. The snake will grab the i t i o n in dry fare&.
foot of the snail with the teeth of the lower
jaw, press the shell against the upper jaw &@was rnezillaria 19091, ZooI.
(WERNER
8 8 : 2 7 9 : t g Ioeal;itJr:Tkbmeo, Mexim.
and then uae the lower jaw to pull the st& b 405 mm. bme0, &dw1,6W l e ~ d
body- -parks out of the shell. Up to a thin3 of P O m alBvation in rain foreat.
the snake's lower jaw is shoved inta the tempm (Wmwsn 1910),
shell during this procedure. ARer the snail N ~ MW.B-~ m:241;~tgpe .
~
is pulled out,the snake will drop the shell d t y : &mad&, Ecuador. TL to 680 mm.
and swallow the soft body parts. All Dipgas b t r a l k w m a to Em.~ador.
species are oviparous and produce clutches 1
of 1-5 (usually 2-3) eggs (FITCR1986,
~ R CAMPBELL1998al. In
S C H U M A C1996,
captivity, the eggs of Dipsas articulata
require just three monthe to hatch.
(SCHUMAGHER 1996 [misidentified s s D.
1 tenuiasimal).

r D. terrnpec811s
r a. vatfegate \' ..
r D. B I ~ I C U I ~ ~ B
Fig. 489.Dipsas bicolor (Siquirrers, Costa
-- - - -_ - . Rieal. Photo: R.D.B d e t t
Fig. 483- Dipsos bravifacies (Quintma Rcu, Fig. 485, Dipsas brevifmies (Corrillo Puerto,
Mexico). Photo: J. C.Lee Quintana Roo, Mexico). Photo: A. Ramirez V.

Dipsas tenuissima TAYLOR1954, Unii..

E a s e s W. Bull. 80: 771; type locality: on
DornWcal b a d in swamp, a prox. 16,Ian
WSW &n i~idradal Genenl, eat.Rca. TL
to 666 mm. M f l c aide of muthern Costa
R h and of weabrn Pmama, sea level to
appmrcimately 600 rn elevation in rain f-t.
Dipma varkguba (Du~Pnn, Bmao~ &
D W M L18M),Erp, GBn,7: 473; type lwali-
Surinam. TL to 821 mm. Eastarn Panama,
ea well as all acmw nurth~nlSouth America,
eea l d to approxhnatelgr600 m elevation in
rain fbrest. In Cerikd America, the riubspe-
cies Dipsas a nichdgi (DUNN 1983b)occura.
Fig. 486. Dips# bemmralis (Ria Jaqu$ / rtia
Dipma viguieri (BWLTET 18BO1, Bull. Soe, Imamadi5, Dad&, P a n m ) ,
Philorn Paris (71 8: 136; type locality) Photo: C, a! Myers
bthmue of D d n , Panama. TL to 637 mr
Eastern Panama d weatarn Colombia.

irrr4- I
Fig. 484. Dipsus bicolor (Tort~guero~Costa Rg. 487. Dipsas articulatu (Cahuita, Coata
Rica). Photo: M.Burger Rica), Phoh: R.W.Van Devender
Jrey to Dipau
1 a Dorsum with large lateral dark bmm
spots that alternate and do not meet along
the vertebral h e .........
Dip#- aadegafa
b Domum with reguIar dark browa ban&
t h ~reach
t b m one side to the other P ......
2 a Two or three large unpaired chin ~hieI&
(Fig. 48Ba) ........................
aip8aa b h & r
b Two or t h e pairs of chin sMel& (Fig.
488b) .........................................................
3
3 a Fewer than 188 ventrale; fewer than 1Dl
submud& ............................. .......*v.. 4
b More than 188 ventralsf more than 101
submudala .....................................,.... 5
4 a 8 or more suprdabials .........,......,..........,,.
b
.................................Lldpeaa bmvijkks
Fewer than 8 eupralabials ........................
.................................DipiBa8 w l X a r 2 8
5 a Mental abiald in contact with firat p a i ~of L.
chin shieldg (rustpair ofinfralabialenot in Rg.489,Dipsas adkulata (Bartola,Rio Sltn
contact with each other) ................... Juan, Nicaragua).
...............................
Dipsaw tempnmlia
b First pair of infralabiak in contact with
each other, thereby separating mental
shield from h t pair afohie shields ........ 8 Further Reading
PETERS1960, K O ~ N
1982, K b m ~%I
6 a No p~%oculm;usually 2 pod.odma .........
VWLMETPER
2002.
..................................D i p m ~~
b One preocular;usually 8 posltDcvlars .......7
7 a More than 210 ventrals; doraal d a c e of
Lad unicolor dark brown ........................
............................... D$pecrs ~ t r i e s i m o
b Fewer than 210 ventrals; dorsal surface of

B
head with b e white spatting ..................
.................................. Dipeas v i , W

Fig. 488, Scalatian of cbin region in D i p m Fig. 490. DryarEophia mZanolomus Wol&
(chin ~lhield~orange), Mombacho, Granada,Nicaragua).
The name Mastigodryas has mpeatedIy
been used for the species of the genw
Dryadop his (PETERS& OREJAB-NIIRANBA
1970,SAVAGE& VILLA1986).A r m w (1964)
and H. Smm & LAR~ENt1974b) have
shorn substantial morphological differen-
ma between lGfa.&lgodryas danieEi [the type
epecies of the genus Mastigodryas) and the
species that are included in the genus
Dryadophis, and have augge~tedthat both
genera are valid. Thia interpretation has
been widely accepted (WESON & M E ~ RF ~ ~ ISelva Negra,
ig.492. D Y A P dorsalis
Matagalpa, Nicmagua).
1985, VILLA et al, 1988, LEE 1996,

While adult specimens of the three Central

American species of the genlls Dryadophis
are all one color or have longitudinal
atripes, the juveniles display pale moss
bands (see Fig. 491). All Dryadophi8 spe-
c i e ~are agile and very irritable, These
diurnal colubride live in savannas, dry
I
forest and rain farest,where they hunt pre-
dominantly lizards (in particular,
Basiliscu4 Nomps, Seeloporus, and Aspido-
scelis), but also frogs, snakes, and rodents
(SEXTON & HEATWOLE 1965, HIMMELSTEIN
1980, SEIIJ1984). All D~*yadopkisspecies
lay eggs (2-5 per dutch), with larger fe-
males producing larger clutches. Female D.
Fig. 493. Dryadophis rnelnnnlornus (Isla Utila,
Handurasl.

Fig, 491, Juvenile Dtyadophis nsehnolomw Fig. 494. Adult Dryadophis melanolornus
(weatern Hondurae). (Nelly, Costa Rica).
melami~musreach aexual maturity at Xey to.Drydophis
appmximately 580 mm SVL, at the age of 1 a No vertebral longitudinal atripe ................
at leaet 18 mosthe (CENSKY& MCGOY .....................D-pkte ~ ~ r n
1988). b A dark medial IongitudinaI atripe p m t
at lea& on the anterior end of the body
(sdditioeal longitudinal stEipm may tre
present posteriorly) ........*....
b....,....ee... 2
$3 a Only om dark medial longitudinal stripe is
formed; darb lateral etripea p a e a t or not
............................
Dydaphiu & d f s
b Three dark longitudinal stripes that run
close to one another anteriorly and join into
a single ~txipeposteriorly; in addition, a
dark l h r a l stripe present on each side ....
...................................
Drycrdophie p h i

Furth~rma-
MART 1841a, LYNW& E S m 1966, SMITH
&
3974b

Fig,496. Dryadaphis pleei ( C W b a ,

CEolombial. Photo: M.Lundberg
 - -
Drymarchon
letugc, snakes (Drymarcha) comprise a
widespread gmup of large colubrid makes,
distributed in eouth-eastern North
America, Central America and parts of
South America. Throughout most of the
2oth century, Drymanchon has been m-
sidered monotypic k g . , AMARAL1 9 2 9 ~
MCCRANIE 1980).Recently, two former sub-
species (e.g., couperi and rnekanurus) were
raised to the status of full snecies and new
s p i e s , D. catc&mmulatlltr; was &cribed Fig.496. Drynaarehon cords unicolor (Saeta
from venezuela (corns 1991,woSTER et Roaa National Park, Guartaeaste, Costa Rita).
Photo: M.Franwn
al. 2001). However, I prefer to recognize
melanurus aa a aubspeciee of D. c o r d
until the s y ~ r n s t i c sof the Indigo snakes
is better understood.
Xndigo snakes are quick, diurnal eolubrids
that feed on a variety of vertebratee kg.,
fish, frage, turtJes, lizards, rodents, and
amall birds). Drymarchon corais reprodu-
ces wiparouslg, pmduchg clutches of 4- 11
eggEl, which hatch in 70-80 days EKOHLER
1997).

Fig. 498. Drymarchon cotvair~unicoiar (San

Marcos F'alqjunoj, Quetzaltenango,Guatemala).
 Fig. 499. Dvmobtua~chlomtbus (Selva Nagra, Fig. 502, Dymabiur ma crritifirue (~wtula,
atagalpa, ~icmagm). o ~uan.~ i c ~ g u 3 ,
~ i ~a.n

- - -
_.
The genus Drymobiua includes only a few
species that are distributed in Mexico,
Central America, and northwestern South
America. They are slender, quick, and
ucludy rather irritable snakes that are
diurnal. They live as pound-dwellers in
savanna ;and forest, in both primary and
secondary vegetation. Whereas D.chlomt-
icus is mostly a highland apeeiea, the
othersr live predominantly in the lowlands,
Fig, 600. Drymobius rhomi . - (Ecuador). usually close to a body of water, Despite its
bright green coloration, D. chloroticus is
t debris
quite difficult to spot m o n g ~ the
and contour^ of the fore& floor. Although
their diet consigts primarily of frogs and

-7 I
toads (DUELLMAN 1963, HENDERSON
HOZ&RS 1877b). they will s b o eat Liz&.
reptile eggs and rodents, All Drymobiua 1
&

species are oviparous. Drymobitss margarit-

ifems produces 4 7 eggs per clutch, with
young hatching in 64-68 days ( W ~ L E R

I
19497 SOL~RZANO & CERDAS 1987). In
Guatemala, freshly hatched snakes of this
species can be seen from June until
dctober (CAMPBELL 1998a).

Rg.601.Drymobius melanotropis ( C e m El
Toro, Atldntico Node,Nicaragua).
Dtynaabtus chbroticwr (COPE IsgBaI, Pmc,
Amer. PW. Snc, e3: 278; type locality:
Guatemala; restricted by COPE (1887) to
C o w ,Alta Veraw, Guatemah. TL to 1200
man. San LUi8 PPtQaiand Oaxaca, Maim, to
central Nicaragua, 600-2600 m elevation in
pine-oak foreat, montane rain f m e t and
doud foreat.
Drymobius m r g a ~ t i f i m sCBCIIL~EL18371,
Essai Physion. Berpens 3: 184; type locality:
"New Orleaam[in error]. llto 1540 mm.
Southern Texas,USA,aad Sonora, Mexico, m
far as Colombia, aea level to 2000 m elevation
in savanaa, wet and rain fmwt. In Cen-1
America, th@? subspecie8 DtynubEus m,mar-
garitiferbce (Texas,USA,along the Caribbean
coast as far as South America), Drymabiue m.
rnaydis VILLA1968 (Great Corn Island,
Nicaragua) and Dvmobiw rn. midentalis
B o c o u ~ T1890 (Chiapaa, Mexico, dong the
Pacific coa~tto F1 Salvador) occur.
Drymobiw mhmtropie (COPE18781, Journ.
Amer. Phil.Soc. (2) 8: 134;type l d t y Costa
Rica. TI. to 760 mm. Eastern Honduras to
southeastern Costa Rica, sea level to 1400 m
elevation in rain forest.
Dryrnobius rhombifer ( GONTHEH1860), Proc.
Zool. Soc. London 1880: 236; type locality:
Esmeraldas, Ecuador. TL to 1270 mm.
Nicaragua, Costa Rica, and Panama, as well
as northern South America (Peru, Ecuador,
and Venezuela).
Pig. 503. Dryrnohius chloruticus (Selver Negra,
Matagalpa, Nicaragua).
Key t;o Drynsobdw
1 a Domum green in Me (blue in preservative),
without large dark btomes or yellow-
green spots ................................................
2
b Doram with large dark blatcher or yellow-
.,.
graen &pot$........., ...........................
.3
9 a 107-126 mbcaudals; kwla of the middle
three do& wale rowr not black ................
.....,.............,..... .D~ymobtua~hlomttol~~
b 91-94 subcauW, keelr of the middle three
dorsalscale rowsblack ................ ..,........
,
...................... D?ymobdrrs mekuurfropt
S P On each doreal scale, a amall yellow or yel-
low-greenspat, so that the domum appem
ewered with fine specklee; 103-138 eub-
caudala ........ Dqwmbiaca mwgda;l&ms
b Dorsal pattern condata of large dark
........................... ..
blotch-; 84-102 mbeaudals .,,. ............
Dq~nwbkrsrhonrbij%r
Further Reading
WILSON1970,1974,1975a-c
? ~ h m o b i u srhombifer
 The species of the genus Elaphe are
distributed predominantly in the Northern
Hemisphere. Many of them have very
attractive markings,as does the yellow-red
rat snake (Eluphe flavirafu), which o ( x m
in Central America. This speciea is semi-
arboreal and live8 in savanna, wet forest
and dry forest, hunting at ~ g h predom-
t
inantly for rats, but also birda and lizarda,
In captivity, a female produced a clutch of
five eggs, which was incubated at 28 "C
and hatched within 65-66 days [ENSPCEROTH Fig. 505. Elaphcr flava'rufa (Calakrnul,
1991). Cmnpeche, Mexico). Photo: H,Bahena B.

Eluphe fbavirufa (Cow: 18671, Proc. A

Nat. M.Philadelphia 18: 3 19; type locality
fi~[tt&, TL to 1660 mm. 'lbaulipm an
Chiapas, Mexico, to Nicaragua, sea level to
1600 m elevetion in seasonal rain and dry
1
forest. In Centrd America, the aubpeciee
Elaaphe fauvirwfa jhvirufa (Tamaulipas,

P
Mb8i00, to the bass of t
h~ U C E ~ ~M
& Iu l a ) ,
Elcsphe f parddim TPE:TEB~ 1868b) (Alta
Verapaz, OuatemaLa, and Bslize to Nicara-
gua),X k p h 8 matwhi S m 1941a (PacSc
veragnt of O a x m and Chiapa$ Mexico), and

XJ
E6 he p h m m D o m 1952
of e ucabinPminsula)OCCUK
~ Cthe north

Fig. 508. Blaplre flauirufa (Cancun, Quintana

Roo, Mexico). Woto: k Rmimz V,

I Fig.604. EEapk flrsuirufh (Isla de Utila,

Honduras).
The genus Enuliophis was recently estab-
lished for the species sclateri which was
previously included in the genua E n u l i u ~
(MCCRANE & VILLA19933. Not much is
known about this speciee other than that it
ia highly eecretive and livee in savanna,
dry forest and wet forest (TAYLOR1951,
Pemz-SANTOS & MORENO1988).
Enulhphh mlatera' (BOYLEN~R 18s4a1, Cat.
Snakes bit. Mm. 3: 851; type bcdity: Sou
America. TL to 504 mm. Nicaragn
Colombia, aea level
forest, and we$ far&
Further Reading
DUNN1938a,MCCRANE& VILLA1993
Fig. 601. Enuliophis scloteri (LaSelva,
Heredia, Costa Rica).
The species of the genus Enulius are mdl,
harmleas colubrids that are highly secre-
tive. They inhabit the rotten limbs of dead
trees and coconut palms, which termite8
have left qonge-like, riddled with hole8
that make ideal hiding places for the little
snakee. Termites and ante are their pri-
mary aaurce of food, although SCOTT
(1983~) mainitains that E. flavitorques lives
on a specialized diet of reptile eggs.
m
Enullus bifbveatu* MOCRANIE& KOHLER
1999b,Carib. J, W,88: 15; type locality: be-
twaen 8avanna.h Bight and East End
i~~29,19W, 85*60.3~ll+1eee than 10 m elavm
tion, Iala de Guanaja, Islaa de b B d a
Honduras. TL to 321 mm, Ida de Guam&
Hondutas, ma 1 4 to 60 m ebvation in we!
meadow and wet f m t .
Eadiun flavitorpea (COPE18691,Proc. h d
Nat. Bci. Phitadel hia 30: 307;type loeatity
NO ~ a g d d y bl~mbia.
, TL to 49s mm
30uthen-iJahsco, M d e o , aa far as Calombia
sea level bo r r p p z b ~ t d y1000 m inlevatior
in rain fbrw$ wet fomt and dry for&. Ir
Central America, the hiubepecies EnJiua 1
&vitotqms (Quatemeb ta Colombia) and
Edttcs f: su&hras# B ~ o 1883 ~ T(south-
eastern Oaxaea and southern Chiapae
Meaim) ocou.
Enulb matamnab Mc&&NE St IC~)HUX
IgWb, Carib. J. W,811: 17; type l d t ~ rnear:
Mud Hole Bay, 16"20,83'N, 06'92.OEW, Lsa
than 10 m elmation, rela de Roath, hlw de
la B&k, Honduras. TL tD 546 rsm. Iala de
Roatsln, Honduras*p e a level to 50 m elevatior
in w& meadow and wet Eoreat.
ICeg fo Enrcliw
1 a Doreal soales with two apical pita; pale
n~ckband intempted or bgmeneed Pig.
808b) ................,........
EtrrElw blfuuimtm
b ]Dorsal scaleswith one apical wale pi@pale
neck band compleh or entirely a k n t ,.B ..
4 r Pale neck band preeent (Fig.5091; ventrale
166-190in d e s , 184-216in females; sub-
eaudalr 100-117 in males
........................,....&wtium~ V $ $ O
...................
~ E
b Without paIe neck band IFig, 508a); veatrals
165 in a d knmn nrale, 176 in only known
female;sdcaudale 121inmde ...................
................................
~ n ~ l t mats-ia
us c-1
FigP009. &hdiwbffoueatu (Isla de Guanda,
Honduras).

a. E,m~tanensls b. € blfoveetus
Fig. 508. Dorsal head view.

Fig. 610. Enulius flavitorques (Costa Rica).

Photo: R.W.Van Devender

Fig.511. Enutius matanensis ( b l de

~b a a ,
Hondurasb
Zur? false coral SI.LL=B- of the .enue
Erythralamprus (altogether 7-8 ~pecies)
are cormpicuoua far their remarkable color-
ation, They are diurnal ground-dwellers
that feed on snakes, fiogg, lizards, and
small mammals (SAZIU & Am 19911.
Their clutches of 6-9 eggs are hidden under

Boos 19963.

Erythro~rnpr~s b b n a JAN 1863a, Arch. Fk. %$hmia~rus m i m u {Montaa L@

Galia*Jinote@%Nicaragua'-
".
Zwl. h a t . Fie, 3: 514:. t w e Iocalitr *Bahia.
Meesico, Po ayan, Cayenne, Brasile,
B
Montevideo, olombia". TL ki 1000 mm.
Caata Rica ta Colombia and Venezuela, sea
levd to approximately 1600 m elevation (in
ColomMa dm to Z6Oil rn elevation) in rain
foreit.
LlytmmZramprus mimus (COW 18891, Roc.
Acad. Nat, Sci, Philadelphia 80: 307; type
locality: "?Zghre 'ons of Ecuador or New
arenab.. TL to 1 8 0 mm. ~ondvrasb ~eru,
600-1500 m elevation [in Colombia also td
2000 m elevation) in rain forest. In Central
America. the subeaeciea Ervthralarnrrrw n.
imrrcar ~ G 1636a ~ (~oindurae
T tb Costa
Rib) and Erythrolamprus m,mkrurue DUNN
L BAILEYI939 (Panama to weatern Ecuadorq

Key to Erythrolampm8
1 a One black neck band, in some specimenrr Fig. 513. ErythroEampsus bizona (Valle Be
partly divided by one pde band;black body M n , Panama).
rings ~ingle,may b divided laterally by a
pale band and have a tendency to become
offset along vertebral line; infralabfals and
pcales in throat area without black pig-
ment; 42-51 aubcaudds ............................
.........................
Etythmlampw mSms
b Two black neck banda, wmpletely divided
by one pale neck band; paired black body
rings not offset along vertebral line;
infialabiale and scdea in throat region
with black pigment; 51-60 subcaudale
.................,,..,.....
E~throlumpncsb h n a

Further Reading
J. HARDY & Boos 1995
Fig, 614.Ficirnia pubdia (nearLa Ceiba,
Atlantida, Honduras).

Key to Rcimia
1 a Intervals between dark ape bss than
Rcirnia
twice as wide as the diameter of the s ts
..................................... r
EYclmlo pu 150
b htervalo between dark spota at least !mice
The genm Ficimicn contains six apscie~, rn wide aa the diameter of the spots ..........
two of which occur in Central America, and ................................ B i m t ranelmi
the remainder further north (L.HARDY
1975). The species of this genus are Further Reading
dist*gui&d by a that i~ H.SM~TH 1941,L. HARDY 19763 1979,
&TAYLOR
upturned and pointed. They are cornpIetely 1980,1490
harmlese, nocturnal ground-dwellers that
feed primarily on spider8 (LEE 1936).
Fkirnia publia, is oviparous, as demonstrat-
ed in the report by G m (1966)of a fe-
male that laid two eggs.

Fig. 615. Fm%imlapuhISa; w e individual

arc in Kg,E11.
The species of the genus Gophis are small,
harmless colubrids that are distributed
across Mexiw, CentralAmerica, and north-
western South America. In Central
America, the genus is well represented by
18 species. The diet sf these s~cretive GedpAS dowwi WVm 1981, Copeia 1@81:
snakes comists of earthworms, slugs, and 64% type ImaliQ: Lm G a s FhFieId Station
small arthropods (CAMPBELL& MURPHY and Weal Garden, 4 Irm 8 San Vita de
1977, CAMPBELLet al. 1983). At the Jab* ISM m elevation, Canton C o d m ,
Biosphere Preserve Bosawb, Nicaragua, I mtarenaa, Caeta Rim, TL ta 268 mm.
South8tp O f the Cordiuera Goes- in
found two Ceophis hoffmanni under a sauthwesternW t a Rim, 1100-1200m elm-
large rotting log in the rain forest. It ia
otherwise reported that most specimens of
thie genus are found under rocks, mots, or
tree branches, and rarely in the open
(CAMPEELL & MURPHY 1977, CAMPBELL et
d.1983).All Geophis species are believed
to be egg-layers, with a reported clutch size
of 3-6 eggs documented for G. Bmchyceph-
alus (SMA 1993).The eggs d G.brachy-
cephalus incubated at 26-28 "C took 109
days to hatch; the hatchlings had a total
len& of 136-143 nun (SASA1999).

erlevation, Panam& Panama. TL to at)l mm.

K d m only 1B m
. tJrpe l d t y . Fig. 616. Geophis brachycephalus (Monteverde,
ChpAis bwhyce hdw8 tmPE 1871X Pm. Costa Wca). Photo: R.W.Van Devender
&ad. Nat. ad. hadelPhia 99: 411; tw
localitr: new Sari Joe4 Ga* Bia. TL to 860
mm. h t a Ria to em- PEteama, 260-
UHH] rn elevation.
+hb wdkkre 194lg, 8mi-
Mi=. Coll, 99 (19): 1 ' ; type localiw
Chichamas,Chpas, M d c o . !l'L to 410 mna.
Southe&rn Chiapas, Menico, and mth-
we- Guatemala, 600-a000 m eIevakjon in
montane min foreet.
Geophk earinaam Smm 1B410, Qcc,
MUM.Zml. U n i Mkhigt~d
~ 463% & type 1 3 -
.
ty: Firm Ban Fr~~eieoa, about 1176 m m a -
tian, 2T km NE Neb&,El Qdsa$Guatemala.
TL to 278 rrrm. Veraeroz, Mdm, to weefern
Gu~Wala,200-2000 m eleveti011 in wet
f&, , p i n 4 W C and daud forest.
&phi8 chanapioni Bo- 1884, Cat
Fig*517. Geophia bmchppIralu8 (Tapanti,
Snakes Brit, Mue. 81: 921; ts;pe lacality
C4wtaRica). Photo: R.W.Van Devender

Fig. I. Geaph amiani (holotype).
Photo: J,R. McCranie Wh.Wien 134: 60; type lwdity: Sarepigui,
b p h b dunni %mmm1992b,Copeia 1-
8;type locality: Matagalpa, Nicaragua.TL to
587 mm K n m only from the type Iocality,
Oeophfa fidrrqguttatue MER'PENS1962b,2001.
Gnz. la 134; type locality: Hacienda
Montacristo, 2200 m elmtion, G O ~ ( V Z L
Meta bn, Saata Ana, El 8dvad.o~l% to 388 Geurphb seledoni TAWR 1954, Unk ~GWW
ma. korthweatcm ~1 ~ a l v d o r westem
EIondm~,1700-2200 m elevation.
&phis goctmuni B o u o m 1894, Cat.
Snakes Brit. MUB.8: 322; type l d t y : Irazd,
Cwta R h , TL to 401 mm. Central Coeta
Rica ta western Panama, 1300-2100 m eleva-
J~E
I. ade em, munioipatity of
C h i ~ p m ,Mexico. TL to 384 mm. Mess
Central in Chiapas, Mexico, 600-2000 m el&
vation in wet forest, pine-oak foreet and cloud
for&.
Geophis m I i s GQPE 18681, Proc. Acad, N&
Sci. Philadelphia SO: 160; type locality: mru
Guatemala City, Guatexnala. TL to 360 mm.
Pacific aide of Guatemala and Chiapas
Mexioo, 40-1830 m elevation ip rain f m t
wet for& and dry forest.
GeophC rhodogaster (COPEl8W, Pmc. kcad,
Nat. Elci, Philadelphia m. 1SD; type locality
OLthe elevated country in the neighborhood d
tFU3 city of Guatemalam.TL to 377 rmn,
Eastern Chiapas*Mexico, to northwestem El
Mvador, 1600-53500m elevation,
Qeophia rubhyeni WERNER1826, Sitz. A M
Brazil (in e m far Smpiqui, H e d a , Casts
Rim, accoding ta DOWNS1967). TL to 26€
mm. Northern and central Cogta Rica, 560
1600 m elevation.
-his tabamancae L m & SAVAGE lQB4,
k.Biol. SIX. W ~ r r h h g k iXOT: 410; type
locality:Erica Jaguar*1800m elevation, Zwra
Pmtethw h Tablm, Cant6n Coto B r q
Pmhnap, Cmta Riea. TL to 218 mm,
Cordillera de Talamanca, Costa Rica, nea~
the border of Panama, 1800 m elevation.
M.Bull. 88: 698; type locality:h c a Zeled6a
between Vdch Barba and Vol& P Q ~H, Nl
R, Casta Ria.TL to 397 mm, Vullrae Po&,
--
Goeta Riea, 1800-2000 m elevatiota.

tion.
Geophis hoflmanni (PEm~81 1869),
MonaQk mng, W .Wh. k l i n 1 m
278; type l d t y : Corta RiEa a d P u e h
Caballo, Goerta Riw bed aa Coeta RSca by
l e a t a t . designation in DOWNS(18873. TL to
800 mm. E&mm Hondurae to &-mntral
Panam& pwibly to northern CoIombia (we
M m 2009),SM)-5OO m e l m t i o ~
Geo his imnwuE&1cs 5)0m$198'7, Mitlc.
hbp. M ue. Zool.Upiv. Michigan 181: W type
lwaliQ: l?ha Lorena, ca. 1709 rn elevation,
& u & a l t e m l Oua-a. TL kr 305 mm.
I Chiapa~, Mexico, and southwestern
Guatemala, 1MIO-230 m elevation in wet
firrest and cloud for&.
QmpRb Eatict'nctw Skum L WILuiwS 1963,
Hezpekdagica 1% type locality: Colonia Fig. 619. h p h b nasaIis (ElRincbn,
I Guatemala). Photo:T,Bill6
216
 ts Qectphie
1 r 17 doraal scale rmsr ........................
...,.... 2
b 15dorsal s d e rows ........................,.,.,. T
$' 4 a Dorsal males prominently keeled, at least
in the posterior half of body ....................8
b Dorsal scales smooth or only &&ly
keeled only over the cloaca1 area ...,w,,,,B
3 @
, Dorsura pale with dark spots or ban&
.
..
................... .............h p k k dunni
b Doraum unicolos brown or black .............. 4
44 41-48 subcaudals; loreal same length or
shorter than pre- and postnasal together;
length of intarnasal shield approximately
3/4 that of prefiontal suture (Fig.520a);
venter banded ...........Geophts eorinoeue
b 23-37 subcaudals; loreal longer than pre-
and postnasal together; length of inter-
nwal shield lees than y2 that of prefrontal
suture (Fig. 520b); venter not banded
.,.....,
...........................,. tkophia naealia
6 a Dorsurn dark with y e h ~ wor pale bmwn
spots ................. Geophie filvogu#abus
b Domum unicolor .,.,. $
, .,
d4.,,+.,*U..w, 6
8 a Without s u p r a d a r shields, frontal ~ i e l d
reaches to eye .......Gsophts rhodogaster
b Supraocular shields present, h n t a l shield
doe8 not reach to eye .......................,......
.................,...., Geophiu immacultue
7 a Without supraocular shields, parietal
shield reaches to eye ...............................
8
b Supraocular shields present, parietal
shield do= wt reach to eye ....................
9

internasal suture

prefrontislsuture

Fig.520. Dorsal head view,

 8 a %%lwon domd side of tail and pcwrterior
third of b* m ~ h t a y%*ha; ~8ntrh
322-P353 eubcaudda 41-48; ro&d and
p m d s m e color an adjacent ecale~
......................................
amp?& aEMonsd
b Soalee on dored aide of t&l and p M o r
€hidplf bady smmth: rrentrah 182-14&
suhudale 2B-36; mstraI and p m w b
white. or yeuaw, in aharp eonb-t ta
darker duration d adjamnt a d m
.......*..........................
OIHlpw
Male; venterr predgminadly pale
in co oration .............b..L..-.....*........-a......I0
lSCW.p
b 6 aupralabialB;venter pale OF d d oolored
b Eye a
.rkmoderate, ratia e 5 lengOh I I a d
lcDpb PO.?: dormm d boar &nn black
ar with pale b d a on posterim- @; no
..... white band a m r rear sf head in adrr1La
(can be p m m t In indi'9idwl8 4160 mm
WL) CS8ophJrr Grec&wpkcslrrr (inpartl
lla Snwt pointed; mtrd lnar
~MD bMeenZX intern%%%
Y
........... pohted .........................
W h km t & d
b Snout rounded; rogtsal b m d y prodwe8
geroteriory between inkernads; mental
1~1mded................. Ghphls h&mamaa
4 4

1Oa D m 4 wale&mmoth or di@tIy keeledjust

m r a i d areta ......
C7mp& ho-f
b Dorsal BFalw markedly keeled ...................
...,...
G B O Ph~ r~ a ~ b h h tin c past)
~
l l a Irnbmmd~h d w i f i prefmntals; venter
&ormly pale In~ Q T
................................
...............................
a q @ h b#xanml-
b Internasals not h d with pdrantah;
venbr with dark pigmentation .............
U
1 %Dnr~Um
~ bl& with n m w pder bands
tbmghout length of bsdy; renter black
with a few pale spats .......,.............w.. 18 .....
b h s a I a d o r ventral ~10krra'b:pidSment
than above; if pale bands atre psent,
these are mtricted to po~teriotportion
of body ...............................................
,. 14
lSa Mom than If50 v a k d fewer ~ than 37 8th- 5%X.&uphie Wua Qlolotspel,
c a d .................*.aeoph2s J r t * w M c, w,Myers
b Fewm thpn 160 vepWa; mra than 88
) ~ubudale .................
i h p h h dbmkrn3
14a h~gd d s mark* keeled rrt l& on
the ~ O S ~ hdfofbody
~ O T ..................... IB
b DmJ scalm $mmt.h or slightly W e d
...........................
1 justoverc1oacal area
la h e a l longer t h n prb and p w m d
to@haq ~~ of intemasal ehkfld lm
than 1J2 that of refrontal em- hntaI
18

ehield me wrii longer than pui.t.1

ruhm length ....................................... IS
b Lored erne lm&h w 5hortsr than pm-
and poetnaaal -er; length of interm-
ml ~ W l d componda to pmfrwtal srutllre
leqih; h n t a l *la twim as long as
padoa ~uturalength ...........................17
I& lzp verp id* ratio o ~ 1. s I b a I
h q t h 40.8; b a d artd bady
black a h md k&w, with a l~onq~cuon%
o m ~~
whib bsnd aims1 raw of haad
...*...............
~.*..,.,...<,# tkipbb I)elZw
...........
 --
Hydmmorphue
-- - I
' ~ r l t : water colubrids of the gem^
Hydromorphus are mid-sized, harmless
snakes, of which one species (lk, concolorl
is distributed widely in Central America,
whereas the other (H,dunni), is known
only from one location in western Panama.
In western Honduras, I found a H.concoior
in a small stream during a nightly rain.
Otherwise, these colubrids prefer to
remain in the area of piles of detdtug in
streams and ponds (SAVAGE & D O ~ L Y
1988,persond observation). These animals
mly leave their aquatic habitat during Fig.523. Hydmmorphus concolor (western
nightly rainfalls, a&also supported by the I-I~nduras).
observations of VENCESet al. (1998).
Nothing is known about the diet of H. eon-
color, however, they most likely feed on fish
and frogs. Thie species is oviparous, aa
demonstrated by a captive-held femal H - W i l d 0 ; ~ C r w Pi?m%818lH
who laid a clutch of four eggs on the 2gna
&~~i,~~~~
&
of June 1876, om which two Young
hatched on the 11 of December 1976
gashm
m elevation in
to mnkdpanama, 100
m d wet forest
(SAVACE & DONNEL~..Y1988).Further repro- @drorrwf.ph c h d i DLTNN 1942 ie regadet
ductive data have been supplied by b be a o n P of H.~ 0 m d o raccordi;llg
SOL~RZANCI et al. (19893; a female laid *VAQE DONNEUY (lgM1s
seven em,which were incubated at 19-24 H dromorphudi dunni S ~ V I 1842, M hoc
"C (dwing the day) and 17-21 "C(at night) & h d . 8ci. a 4?r:.tp locality:vicinit:
aorth of Boquete, C h q u f , Farnama. TL tc
and fhm a young @g -, W e s m panu the area
after 94 days, with a total length of 166 meb w, qu ~fBl,Em~O
mm.

Key to Rydmmorpltw
X a One prefrontal shield; one internasal
shield;dorsal ~ c a l e sin 17 rows at midbody
. .. .
, .. . . .............. H ~ m r p h u r eoncotor
r
I
b Three prekontal &Melds;two internaaal
ehielda; dorsal ecales in 15 rows at mid-
body ...............,......

.
I
I1 Hydromorphars dunni

Further Reading
I Hydrwnorphus coneolor
NELSON1966, V u 1970, SAWE & DONNELLY
1988, VU 1990
A Hydromwphus dunni
-
' I"
 -

Imantodes Imntadtrs t?mclcoa Ctnm~ua176B1, B~firtema

' T a m , ed. 10: 216;type I d W - ArrraFica,
Z to 1108 man. Southern Tmmidipaa and
lne speees of the genvs Imantodes are &axaca, Mexico, to ~~, ma I d to
dietinguished by a very slender body, a approximately 1500 rn elevation in rain
large head clearly distinct from the neck, forset, wet farest and dry for&.~~
MBRS (13833 and W m & Mrmm 11!3&6),
tw
and large, protruding eyes. They are cre- the df*ian into eubspecise saggabd by H.
puscular and nocturnal ~ h r u bdwellers, S~am( I W b )a d m~4 & Ow=-
which are entirely harmless md rarely (19701ie insuBciently supparted.
attempt to bite. During the day, they hide I m n t o d e ~gcnnini@tratur(COPE1861e1,Roc
in bromeliads, hollow trees, under loose Aead. Nst, Sei Ph&de1p& 15: 29%
bark, under rotten lop, and in similar pla- locality: n6m kalco, 8an Salvador, ,
ceB (HENDERSON & NICKERSON 1876). At 8ahrad0~.TL to 889 nun, Nwthern Veracm
night, they can be seen, usually etretched and dlontrhesn Sanora, Maria, to Colombia,
out, gliding slowly through the vegetation, einmdrg level to approximately 1600 m elevation
fmst md wet forest. Aocordhg to
particularly along streams on the lookout 116- U9@3 aad W m & Mmsa (19851
for moles (Norops), which make up the the &vidcig into eube b? .."
bulk of their prey (M~ERS1982). They will ZWEIFPIL(It3693 and
d s o consume small fhgs and reptile eggs Z'dmMm~~1B70) iB iasaPBcjdy eup
h ha Central America, and I ps not O
(LANDP et d. 1966)-All species of this
genus are oviparous, producing clutches of
1.8 (usually 2-3) eggs IFITCH 1988,
CAMPBELL1948a).A female I. cenchchoa cap-
tured by Mmm (1982) laid a clutch of 3
eggs on the 3oth of May, which, aRer 69
days of incubation, contained embryos that
were nearly ready to hatch. It is otherwise
reported that the egg-laying period for t h i ~
species in the northern portion of Central
America is from A p d until June (STUART
1948).

w Lr
Ir
Fig. 625. Imantodes pharatnmu (holotypel.
Phota: C. W Myers

= !. tenuisslmus
-
'

I Fig. 524 Imantodes cenchoa (Bartola, &o Spin

Juan,Nicaragua). .
I. phentesma
.
 Fig.527. Itnuntodm gemmistratus (Monkverde,
Costa Rica). Photo: R.W.Van Devender

-am
EL-

Fig.526. Imantodea tenuissimue h e r t o Fig. 528. Irnantdes c e n c h (LaS~lva,

Momlos,Quintana Roo, Mexico), Heredia, Costa Rita)
Photo: R,Cedeiio V.

Fig.629. IrnontodBs inormhs (Tortuguero,

Lim6n, Costa Rica).
Key to Inran#&s
1 a Doreum with large, dark saddle-like spob
{Fig, li50a) ..........................................~....
b Dora- unieol~rpde bmwn or with irreg-
ularly dietPibuted dark e ts, in&-
dark i h @or pale *ddl*Ee apt. wig.
&Bob) ................................................
-4
4 a S d e of~ the d i d dmal m s distinckly
enlarged (8-4 times as wide aa the adjacent
0&; Fig,531a) ......XmMlt&s m m h
b Scab8 of the medial d o r d rows not or only
&lightlyenlarlqed (L5-2.0 M wide fu
the adjacent oealw, F'isS31b) ..................
8
8 a Dark dorsal spob flBt h*mpted at aides;
more than 240 rentrala .......,.................
.........................~~~8 kawtssfmus
b Dark do& spot0 offen hberrupted at
aides, especially posteriorly; usually f ~ t 3 r Fk.691.h d scales in Imantode~.
than 240 ventrals ..,....................
*...,.
4 a Fewer than 220 v e n t d g fewer than 140
dammi uaicolor e brown or
~~bcaudala;
with irregularly dlstrtbu
................,... ts8"
dark spots
..,......Intantodm h m h i a
b More than 220 v~ntrals;more than 140
subcaudal~; dorsum with pale saddle-
epat13 ......~ r n c c Q ~ ~ p h 4 M B d a ~ r n Y r

.-

b. I, inomatus
b*wheae
dkipm* a! -
l s ; E ~ q ~ i p k :dtP- M
N m m inthe qtum R @ UFmkr J
--ram8
w S d ZL
-W*
Fig.630, Dorsal pattern in Imaatodes.
Fig. 632.Lampropeltis triangulum (Rio Patuca, Olancho, Honduras).

Lnmpmpeltre tirrlrgtldum (LAc&P~DE 17891,

Hist. Nat. Qua& Ovip. Ssrp. 2: 8QS,type local-
ity: America. TL b 2000 mm, Dbtributad all
across the central 4emtarn USA ae weU aa
amahi Mexica, %tml America and north-
western South America. In Central America,
the eubspeeiea hmpnopaltia t. Dbrurmrrr (Bo-
mum 1886) (central and nofitam Guate-
mala, muthem Bsrk~l,t a ~ r a p e l td.i ~b h -
chamli STUAXF 1536b (YucatBn hninsula),
Lamprapeitis t. e&eae DUNN193735 (south-
emten costa fliu and we8tern Panama),
Lampmpelbis s: hodurensw WUIAMB 1978
(Caribbm side ofHondunw, Nicaragua mid
nartheaatera Cmh Rita), LampmpeItis h
mlcmpblis COPE 18614 (mtralPanama to
naFtbw%~tern South America), Lrwnpmpeltis
t. digo8ona B O C ~ ~ 1886
R T [PaciRe aide of
ehia as, Mexico, to eoutheastern Guate-
mala! h r n m w i a t, pdymnn corn 18806
~ e r a c m zGexico,
~ to w a t l m G m t a r n d I
and Lumpmplti t. stuarti W m 1378
(Pacific side of southeastern Guatemala ta
c e n t 4 Costa Nca) occur. Fig. 533. k t ti$ triawulurn (nearLa
- -- Ceiba, Atlantida, Honduras),
 Fig. 584. LampmpeEth ;hd&@urn hodurn?&* Fie 636.Ltpt&t'm finatcm (Rchucalca,
Photo: U,Huch Chlapas, Me&). F h t a A, Ramirea V.

I
I

1
Fig. 636. b m ~ ~ i t tdr ks ~ a l u mm i c m ~ m b . Fig.537. L,pto&ira fiigrofa8~iata(Liberia,
Phok~: ILuch Guanacaste, Costa Rica). Photo: R. D.Bartlett

Fig. 638.LBpfodeim septentrionalis (Tortuguem,

Limdn, Costa Rica).
M h e r Reading
B w 1973,K WILKAM~
1978,1988,1994
224
The species of the genus Z e p t d i r a
hare the common name cat-eyed
snakes, due to their vertical pupils.
They are nocturnal illubrids that
are harmless as a rule, and rerely
~-
attempt to bite. They are found in a broad
range of habitats, wen in secondary vege-
tation doee to human habitation.At night,
theee semi-arbreal snakes are on the look-
out for their preferred prey of frogs, small
snakes, and lizards,Lept&ira annulata is
in particuJar far cmsumhg
spawn that bas been deposited on leaves
(e~pecially that of red-eyd tree-frogs
lAgalychnis collidryosl and glasa frogs
[family Centrolenidael) (C-BELL 1998a).
All h ~ t o d e i ms p i e s reproduce oGP'-
ously, with clutch sims between 2 and 12
eggs (DUE- 1968, F ~ 1985).H The
incubation p e a , depending on tempera-
ture, is 56-86 daye (KOHLEB 1997,
CAMPBELL
1998a).
L 1935 bouthem half of the Yucatan Peninsula,
- Leptodeim ennuiata
=Leptadelre frenata
~i~ 534,k P ~ i r r7a
mc
a (Ida Cozumel,
Quintma Roo, Mexico).
Lee
Lapthira anlaukta [LINNAEUS 1758),
sptm~~ Natme, ed. 10: 216; type l d t ~
Amwon Baain. TL to 870 mm, TamauGw
z&I1zzf
, ;di.~~?$t8nB~$B~f
level to appkaMy 1000 elBVatim
in rain forest, wet forert and ctry forest. In
Central America, Che subspecies Leptod~ira
a- cmsairis ~~1368 w a d i p a s and
hmto CK~PU,Mexico) m d h ~ d i m
a. rhmbifsm G U m m 1872 (Guatemala to
panama~OMN,
Leptotkim F.emba (Corn 188Bb), h a . U.6.
Nafl. Mus. 9: 184; Jalapa I=Xalapal,
Veracmz, Mmieo. TL to 715 mm, Vera-,
Mexiw, to northern Guatemala, aa well as
the entlre Yucatsln Penineuls, sea lwsl to
1000 m elevation in Fain forest, wet f m t
and dry fom~t.In Central America, the eub-
epeciw Leptodeim fi m~lkiaiDUNN h STUART
as well aa don the eaet ooast of the Yucatsln
Peninauk) an! hptadaira fi yumtanensis
COPE 1887 (the northern portion of the
Yucath Fenineula) =nu.
Fig. 640. Leptodeira rubricata (ProvincePunta-
renas, Costa Rica). Photo: R. W. Van Devender
 Iw.
-
Leptodpim laigm)h?htaQONTBBR 1868, A n n
Nak. Hist. (4) 1: 425; tgpe hdi*
N r c a r m . Tt b 660 mm.Guerrw~~ Mexico.
don &e h ~ % cm t to C d a hca; ixi
onf fur as &o on the Cariibem tide,
level to appmxhmtely 180Q m, slevatiw in
d r y M and wet fwsst.
hptdeirra rubripatrr (COPE 18981, P r a ~ b e d
Fhilm. 8ac. 81: 347; type locality: Boea m a ,
Goeta Rim. 'llto 790 mm, Cwta Rica, in
mmgmves.
h p t d a ' m ~ptemiotwrlIs~KENNZC~TP 18591,
in EUm Reptilea of the 'Boundary 6 16;type
I
l d t y Matarnmas, Tamaulipm, Mdm. TL Fig. 642. Leptodeirrr alnnukta (Bartola,Ria
to1WS mm. Texa~, U& and Sindoa, San Juan,Nicaragua).
Mexico, ka Peru, -sea lwd to a p p d m a b l y
1600 m el~vationin rain brest, wet fmwt
end drg forest, In Central Amrtrica, the rub-
@BB Leptodeim a o m & ~BOCQURT 1884) Key to Lepfocteirts
(- & ~0rthw6~taI'n South h&Cd I a, 19 doreal row8 at midbody ........................ P
and & p W i r a a poEysticta 1895
(Veram and hemru, bQeEioo, to Co- b 21 or more domal rows at midbody ..........8
Rim) otcur IPD~mm with 10-21 dark ban& that eEtend
to the ventraIs; 64-7d aubcaudala
.....................Lepdadeh migrr,-i&.s
b Domum with 50-70 dark bands that do nok
extend to the vent& 72-107 subeaudah
.....Leptd%im~ b s r p t ~(in~part) l f .
8 a One dark poetorbital stripe pregent, in
broad eontact with k t body a p t : a single
dark pot in the center of each a r i d
(mg.541e) .................
~ c p d Rh rrrdQ
b Dark pborbital &ripe, if present, not in
b m d contact with firat bady spot; ma&-
ings QII the p&t& different than a b m

4 a VentraIcr usually more than 186; 72-107

snbcaudale; medial neck etrip, if prasent,
extends on1 3-8 scales behind the
pui- =dl d t~h h h
h a lI0 E o Z I ~ G
body spot [Fig, EiQlb);dark dorsal Bpot :
without pale border (Fig. W a ) ..................
.....Leptdinz mpi:entrtonabs (inpart1
b Venttab fewer than 186;6-4-89subcau*, ;
medial neek atrdge usually extende from
the parietal8 to h t body pot, with which
it is c d u e a t (Fig. 841a); dark dorsal spPt .
with pals b d e r [Fig,643b) ..................6
B a Dnrsm with 28-51 dark 8pm (urn@
with pal%borders;Fig.M3bI on pale b m
ground color or s p h blend into a z i p g
pattern; venter mam colored in lit%...........
............................. Leptde1v-a o n n n b
b D o r m with 67-67 dark, regolar cross
bands (without pale borders) on red
Fig.541. Dorsal head pattern in three spedm of gmund color; venter red in life ...................
Leptodeirn (afterDUELWIAN 1958). ...........................tspdwCeIm Flbbdcuta
226
 As the species name pulcherrimu~implies,
the only species of the genua Leptdvmus is
a particularly graceful, beautifully ~ f i p e d
snake, remarkable for the vivid green
coloration of the d a d surface of its head.
L. pulchrrimua is a fast moving colubrid,
at home in dry areas. It is a nocturnal
ground dweller,whose main diet consists of
lizards.

Fig.643. Doma1 body pattem in Zeptadeim

(aRarDmuam 1968).

Further Reading
D m w 1958

Fig, 544. Lepikdettm wptenWwlh (Cape

Cochinos, Honduras). Phato: M.Lmbarg

fig.546. Leptod fymus pulckerrimmt

Photo: R. W.Van Devender
 Colubridae

Iap#ophis
-

-
-
The genus Leptophis (parrot snakes) in-
cludes eight species that populate the
forests, mangroves, and savannas of h k i c o ,
Central, and South America. Leptaphia
depressirostris is a magnificent leaf green
snake, as are some of the subepeciea of L.
ahmtdla. Aa their large, round-pupilled

arboreal have an irritable tempera- Fig. 643. Leptophb nebulosus (Mmteverde,

ment, When handled, they will respond Costa Rica). Photo: M.Fxanzen
with a threatening gape, substantially
expanding the lower jaw, hence their Latin
name. They will often remain in this pose
for an extended period of time, but rarely
make use of their numerous pointed teeth.

Parrot snakes consume primarily fiwga, as

welJ as the occasional sdaznander, lizard,
snake or bird egg ((3- 1948, STUART
1948, SEXTON& HEATWOLE 1965). All
Leptophis species are oviparous and pro-
duce clutches of 2-11 egga (STUART 1948,
SEXTON & HEATWOW1966, CENSKY&
McCoy 1988,CAMWELL 1998a). DUEUNAN
(1978) discovered a combined clutch of
Leptodeira and Leptophia ahQ.etuZEaeggs in
a b a m b h t e d e about 12 meters off Fig. 548. h p t o p h i s nebutosus (Krahkra,
the ground. In northern Central America, Gracias a Dios, Hondurae). Photo: J. R. McCranie
the egg-laying period occurs h m March to
October (CWBELL 1998a1,

Fig. 546. Zeptophia ahraetulla (Selva Negra, Fig.549, bptophis ahaRtuEla (Rio Patuca,
Matagalpa, Nicaragua). Olanchu, Honduras).
228
-
kptophis ahaetulla (LWUS 1768);l
Bysterna Naturae, ed. 10: 226; type locality:
'!Asia, America". 'lZ to 2250 mm, Veracnu
and Oaxaca, Mexico, to northevertern
Eauador west of the Andm, aa well as

I
Argentina and Brad east of the Andes, sea
level to approximately 1300 m elevation in
rain fore&, wet foreet and dry forest. In
Central America, tha subspecieg Leptophls a;.
occi&ntalie (GIMTHEIR 1859) (Nicaragua to
Panama as well ae western Venemla tc
Ecuador) and teptophia a, mestuns (COPE
1869) (Vemruz, Mexico, to lfondmad m.
Pig. 10,LDptop his : (near
Pueblo Wiso, Jinotega, Nicaragua).

Eeptaphh depreseiroetris (COPE186lb1,Roc.

&ad. Nat. Sei. Philadelphia 12: 657; type
locality: Cmyas de Veraguw, New Grenada
[k Cocuyaa, Panamal, TL to 1400 mm.
Ecuador.
XIIcaragua to w e ~ h r n
Ljeptophis diplotmpia ( G ~ 18T21, R Ann
Mag. Nat. Hist. (41 0: 25; type locality
%huantepec, Mexico, TL to 1410 mm
&uthwe~tern Chihuahua and aoutherr
hnora to southeastern Oaxaca, Mexico, sea
b e 1 to app~loxhately1000 m elevation in
dsy h t and wet fbmt.
,&ptophia micartus D W w , BIBRON&
I
&JX&RJL 1864,Erp. Ct$n., 7: 636;type l d h y -
PaXico. TL to 1270 mm. Waulipaa am
%sac4 Mexico, to Coata R i a , sea lwel
sppmgimately 1700 m elevation in dry foresl
and wet fmwt, In Central M c a , aa well ar
B e nominate subepecier, the subspecier
Jkptophis m. hoeverai HENDEMON1978
fmdemic to the Turneffe IsIandeI Bebe)
&cur.
Lptophh modashrs (GUNTHEX18721, AM.
lufag. Nat. Hist. (4) 9: 26; type 1Wtr Rio
Wsoy I=W o y or N w l , below the tuwn of

I
Cubuloo, Baja Y e r a p , Guatemala I e m m
ewa according to M c C w & WILSON 19431.
TL to 1720 mm. Northeastern Chiapea and
central Guatemala to northwestern El
Salvador and southweetern Honduras, 1800-
2000 in cloud forest.
Leptaphis nebarlasuar OLIVE%1942, Occ. rap.
I
Mm.Zool. U n k Michigan 488: 12; type lo-
Witg: Cariblmca, Coata Nca. TL to 884 mm
H e e m t e r n Honduras to Costa Rica
&pto his rtveti DE~FAX 1910,Bull.Mus.Hist.
Nst Faria 7: 36s; type locality: Gualapuiza,
S O m elevation, Ecuador. TL to 400 mm
butheaatern Coata Riea to Ecuador west d
&heAndea, as well aa Peru east of the Andes,
a%elevations of 100Q-1800m.
Key f o Leptaphk
................................... .,. B
1 a Loreal present. .
b NQlmal ................................................
6
a a Keelle pregent or& on the paravertebral
sdentors ...............................................
3
b Keels an dl the doreal ~calerows, e m p t
forthefirst m ................,.....................
4
3 a Fewer than 160 ventmls; 9 aupralabialr
.............. ....
., Legbophis dqpmmi1~~hde
b More than 160 v e n h h ; 8 euprdabials .,..,.
.............................LephopM~d $ h f m p i ~
4 a Unicolar green; v e n W without lateral
keel;168-183veatrale ............................
.................Leptaphis mQdenhzs (in pwtl
b Mid-dorsum b m e colored, borderad by a
dark longitudinal stri * vent& dwap Fig. 561. Leptopftia ~ h a e ~ u l(Sslva
lu Negra,
with a weakly defined E&rd keel; 11S474 Matagalp, Nicaragua),
ventrab ................ hpbplblu W a n u s
B a Fewer khan 150 vent&; dl 111mal acalso
-.
keeled ....................... Leptophie riue#i
b Mom than 150 ventrals; @dmofthe first
domal caw withut lwek ...................
,,..,I
8 E A bIue-green or blue longitudinal d p e
d n g dong the haecond and thirul or
third and fourth damal mwe .......................
.............................tRPtttphi~B ~ ~ I ~ U B
b Unioolor green or, if a bllle-green or blue
IongitudinaI strip is present, it runs dong:
the third to fdth dorsal rows .................... 7
7 a Unicolor peen; 168-188 wntrds; in the
highladds (150Q-2000m elevation) .............
................ LepttFphlla mods~~trre tin part1
b Unimlor green or with a b l u e - p n OX blue
longitudinal 5trfpe along the third to fifth
h a 1 mwe; 160-183ventrds; in the low-
land and premontane areas (gea level ta Fig, 552. Leptvplrle mxkenus (Ida cEe Utila,
a h t 1300 m elevation) .............................. Honduras).
..........
...................,. hptophk &tuZla

Fig. 565, Leptophis diplotmpb ( C a m de

Sumidem, CBiapas, Mexico).
Photo: A. Ramirez V.
4 I

Liopkis - -
The genus Liophis includes around 36 spe-
cies, only two of which IL epimzphelus and
L. lineatus) reach Central America (Dnror~
1989).In the past, Liophis epinephlus was
placed in the genus Leimadophis (J.
PETERS & O R E J A S - M ~ ~1970,
D A SAVAGE 8t
VKLA 14861, whereas L. Uneatus was
included in the genus Lygophis (J.PETEM
L OREJAS-MIRANDA 1970). Both these spe-
cies are ground-dwellers that feed on Fig. 564. Liaphie Iineatue (Cdrdoba, Colombia),
lizards and frogs, and also on invertebrates Photo:M.Lundberg
ag hatchlinge (TAYLOR1951, LANCPNI &
KORNACKER 1989). They reproduce ovipar-
ously, producing clutches of 1-8 eggs.
-1

Fig, 555. Liaphis epinepblus (Coata Rica).

Phata: R.W.Van Ilwender

I
Key to Liophirr
I a lT dorsal rows at midbody; It'ewer than 156
ventral8 ................
t h p h L epinephehs
b 19 dawal rows at midbody; more than 155
ventrals .......................
Liophie lineaha

/ Pip"566. L h p h i ~epinephslu8 (C- Carmo,

Guanacaate, Cmta Rita), Photo:M,Franmcen
 The genus Manolepis is monotypic,contain-
ing the species M, putmmi. Interestingly,
thisl specie8 displays a pronounced sexual
dichromatisrn: while both genders have the
dark dorsal stripe typical of the species, in
the male it is uniformly dark, and in the
female, it bae a pale center (WERLER&
SMITH1952, JOHNSON 1978). Little is
known about the habits of this colubrid.
They live in dry area8 and ndpmduce by
laying eggs (HARTWEG& OLIVER1940).
Further Reading
WERLER& ~ M U I I T1962,
H JOHNSON 1978
Rg,667. Manolepis prrtmmi Monte Banito,
Chiapm, Mexico). P h ~ t a : b . ~ V -.
232
Masticophis, which is closely related to the
genus Coluber, include8 eight mid-eixe to
large, very agile colubrid species that are
distributed all across the southern half of
the United States, aa well as acrorJs
Mexico, and one species that is distributed
in Central America, Isolated populations of
this species also exist in Colombia and
Venezuela. Masticophis mentovarius is a
diurnal ground-dwelling colubrid that will
dart off at great speed if dieturbed. If
handled, they will g a p with wide-open
mouths and not he~itateto make warning
strikes and bite. Their diet consiste mainly
of lizards bbpidoscelis and Sceloporus),
but they will d s o hunt rodents curd blrda
(HENDERSON & HOEVERS1977b, h v m z
n m T o ~ o 1983, CAWELL 19913a).
Egglaying has so far been documented in
March, with clutch sizes between 7 and 30
eggs (WRLER 1961, M r m ~& M ~
1991,C ~ E L 1998a).
L
Mastimphis mentovarius (DUW~NL, BIBR~N k
I L Erp. GBn. 7: 187;type locali-
I W ~ ~ 18H1,
ty: Mexico.TL to 2600 mm. Sonora and San
Luis htosf, Mexico, to Coeta Rica;a h dia-
junct in Panama, Colombia and Venezuela,
sea level to approximately 2000 m elevation
in savanna, plne fore& and dry forest. In
Central America, the eubspeciea Maeticophht
rn. naentovasius (Jaliacu and San Luis PotwiI
Maxico, to C o ~ t aRical and Masticophis- m.
centralis (ROZE 1963) (disjunct in Panama,
Colombia and Venezuela) occur.
Illl Manolepis putnami
Masticophis rnentovarfvs
 - - --
- 1 frogs and freshwater
WRIGHT
(Wluoar &
shrimp as well
B
1967, NLQNJARREZ
Nerorlia rrhombifera
W f A S - G m f ~1991).
is predominantly nocturnal,but is occasian-
ally seen basking during the day on
branches jutting out of the water.They are
very irritable snake8 that will not hesitate
to bite when caught. This species is a Eve-
bearer, with the tropical populations bear-
ing fewer babies than their northern con-
sGcifics I ~ ~ L D R ~etGal.
E 1996).According
to these authors, the litter size of N.
rhoidifera in Vetamz, Mexico, is 8-35
(mean 17)-Females reach sexual maturity
Eg.658. Masticopki~men with a snout-vent length of 670 mm,males
Otoro, Handwas). at 475 mm (ALDRIDGEe t al. 1996).

N e d i a rhombifim (HALLOWELL 1862), Pro1

Further Reading &ad. Nat. Sci. Philadelphia B: 177; type lt
ORTENBURGER
1928, eality: the Arkaneaa river and its tributarie-
1977.1982 near the northern boundary of the Creek
Nation. TL to 1200 mm. Iowa, lllinoia m d
Indiana, USA, to Tabaaco and northern
Gbiapas, M d w ,sea lwei to 600 rn elevation.
T h e subspeciee Arerodia r. werlen' (CONANT
1953) occnra in Central America.
-
Nemdiq -
-
Only one species of the genus Nerod&I R
rhombifera, reaches as far a5 the northern
portion of Central America, whereas the
other nine apecies are distributed across
North America. As with aL1 species of this
genw, it is semiaquatic, living in rivere,
lakes, ponds, and flooded meadows.Its diet
consieta primarily af fish, but indudea

Fig. 653, A,. d i a rhombifera (Veracruz,

Photo: P.Heimes

Further Reading
Nemdia rhambifeta CONANT1969,M C A U I m 1935,AtDRflSixet el.
! - - -
-
Fig,680.Ndnia s e k [ElImposible Natiomal
- Park, Ahustchaplln, El Salvadorl.

The genus Ninia includes small, secretive,

harmlees snakes thd, in many places,
make up an important part of the (&if-lit-
ter herpetofauna. Very often, these snakes
are found under logs and in rotten roots on . .1-

coffee plantations, which is why they are

known as "coffeesnakes".All Ninia species
ara predominantly nocturnal m d eat
earthworms, leeches, elugs, and anails
1975). Ifthreaten-
(TAYLOR1961, GREENE
ed, the Ninia will flatten their
bodies, Bide their heads under
their bad^ liR their tails Kg, 561. NinbarbDe (near Duanagmaw,
in a epiral (GREENE 1975). ~ ~ - ~~ l ~ , ~ ~ b
Occasionally, they w i l l also lift the
frank of their bodies "cobra style"
in order to &play to potential
r predator IHENDWON & HOE-
1977a, compare Fig. 662 and 566).
All Ninia species are oviparous and
July and December (GAIGE1936, GREENE
1976, ALVAREZ DEL TORO1983). The ju-
veniles of N. sebae hatch after an incuba-
produce clutches of 1-4 eggs between tion period of 76-79day8 (GREENE1975).
Ninia strata (KsLLo- la@,Proct Amd.
Nat, Sci. Madelphia 8: 126;type locality:
within 200 miles dCaracris,Venezuela. TL ta
450 mm. Eastern Panama and northern
South America (Ecuador, Colombia and
Venezuela), 830-1360 m elevation in raie
foreat and thud fomgt.
Ninia cela& MCCRANIEBt WILBON1996, J.
Herpetdl. 88 (2): 225; type locality: Cinchom
(Isla Bonita), Atlantic side of Vela Pods,
1200 m elevation, Alajuela, Costa Eta. TL to
360 m. North-central Coeta Rita and r
we&m -, 830-1600 m eltyatjm in C

I
rain forest and cloud f a s t .
Ninia diabmato BAIRD& GIILARD 1863, Cat
N. Amez Fkpt.: 49; type locality: Orizaba
Mexico, TL to 421 mm. San Luie Potosf and
Omca, Mexico, to m t r d Hondurae, sea
level to 2200 rn elevation in rain forest and
cloud forest. According to WLSON gt IvDmm
(19861, the diviaion into suhpetiea a u w t e d
by B T ~& ~ WWERR
R (1954) ia ioeu5ciently
mpported, and I prefer not to rewgniae any
eubspciea.
Mnia espinali MCCW & WILSON 1996, J.
Herpetal. 28 (2): 228; locah El P d l l o
de OclDtepequs (14'2% 89'd%, 1810 m
elevation, btspeque, Honduras. TL ta 610
mm. Western Hodurat and northern El
Salvador, 1690-2270 m elevation in doud
fmst.
Ninlh mcubata ( ~ R 18613, B Monateber
Ktilzg. Akad. Wiss. Berlin lE81: 924; type lo-
cality: Goeta R i a . TL to 352 mm. Eastern
Honduras to e a s h n Paaama, sea level ta
1760 m elmation in rain forest and cloud
formt.
Ninia pavimentata @ W O ~ R T 18831, b.
Sci. Mex., Rept.: 649; type l o d t y : Alta Vera-
,Guatemala.TL to 380 mm. Fringe ofhi&-
E d s of Guatamala, 1800-1SW rn elevation.
Ninh se hota (COPE 18783,J. Acad. Net, Sci.
~hil&l$m (21 8: 146; type loc&W: higher
points on Pioo Blanco, 5000-7000 R, C@
Xea TL to 494 mm. Cwta Rlca and weatam
Panama, 1200-2770 m elevation in rain forest
and cloud forest.
Mnia sebQe ( D m ,BIBR~N& DUMPZUL
18841, Erp. W., 7: 616;type locality: MBxico.
TL ta 386 mm. V e r m z and Oartaca, Mexico,
b central Cwta Rica, rsea level to 2200 m
devation in wet f ' s t , rain forest, and cloud
fomat. becording to W m B MEYER(19851,
the division b,to subspecies ernggehd by
SCHMIDT & RAND (1957) is insufficiently
BU ported, and I prefer not to reognke any
suEsPeciea.
 Fig, 567. tia mael
MatagaJ,-, Niwag-, .

Fig. 566. Nznb psephota (Guanacaste, Costa Fig. 568. Ninia psephota (Monteverde, Costa
Rica). Photo: R.W. Van Devender Rica). Phato: R. W,Van Devender

Fig.568. Dhplag behavior in Ninia Awlsoni Fig. 569. Ninh diQdewta (new Quebrada
(Ecuador). h d e , Ca*, Honduc~e).P h o h J. R,Mdranie .
Kes to 1V"Enk
.......................
1 a 17 dwadmwloat midbody
..........................
b Isdmal
2 a VenCer with dmk, aftan r n w
...,. *Ntn2apa@wta
ab ddbody .....,. ,.

spots ,................,.,...,.,....,.,,.,a
b Vfabnftrkrgelg unpigmznted
..........
..............
aicpa~
B
S a
.............,.....
6
p- - -

8 a Fmer thaa @2 subcaodrrla

...*........~~.~..A.....v..~,,,..s.~,. WkkrmmvZata
...
.,...,..
b Horn than 6B e h t l d a b .................. ,.,,,,. 4
4 a Bmum reddi&-hmm 4 t
h narrwp, dmk
band8 ....,........Nrnk prrv-a
Dr(l$hi
bDonnim dark brown or Ha& bub .ppi&aut
dark mW.5 banda .........
NbL did#-*
Rg.570, Nni& otr~bcst(?drdobalColombiaZ
6 a Doreum red or reddish b m m in life (pde Photo: M. LundlsEtrg-
ta dark brown in p m t i v e e ) 2
b d t y e h to pale brown in Tel with
poW0~bIackbordtm m t . h b d w not
!I?=
&ex parietals (Fig.5'?2a) .....&Mu&boe
b Doraum black or dark if a pale neok band
is formed, it aleo cove= the pattenor half
of bparietal8 (Fig.672b) .......................
6
6 a Without pale neck band .,.Ndnia enpdnali
b W e neck b d dhtinet ...................., ...
7
7 a More tban 190 venhIs; 3s-70 aubcsludds
......................*..... &w&
i.ll.l,t.~Il...iCNt;tffC1
b Fewwthan ZSQ verrk& 33.45 t+ttbcaiadqile.
............. m a
v*LL*****+*I*I**IL1.vIl.*.al+*NW

Further Reading
I
BLIRGER & WERLER1954, K. S c m & RAND
1957, SAVAOE& LAI-IANAS 1991, MCCRANIE8t
Wmopr 1996, E. SMITH& C ~ P B E L L1996.
M c C m i i et al. 2001, Fig. 571. Ninb atsnta (Lac Moralw, CGdFdoba,
Colombia), Photo: M.Lundberg

int k w rpeGiea ofNhk,

Fig, 57% Dorsal head ~p~;#am
 The genus Nothopsis is monotypic contain-
ing the species N,rugosus.Both the shape
of the head and the presence of emall imp
ular scales an the doreal surface of the
head give this mlubrid a rather bid-like
appearance. Even t;oday, our knowledge of
the biology of A? rug08us ie very isketehy
and the epecies is scarceIy represeated in
museum collections. F,SCHMIDT asd I &-
wvered a specimen of N; mqo$us lying
motionless on the ground at 820 rn eleva-
tion at Cemo Saslaya (northeastern
Nicaragua) during a rain &owsr at night
K
(C m& E&mm20011,When we picked
1
it up, it made no attempt to bite. Ths few
other existing notea on the habitg of t b
The genus Omoadiphs (KOHLER et al.
2001b), containing the species 0.aurula,
was described very recently. So far, this
species is only known from a single speci-
men ~ollectedby the Director of Cueulca
National Park and given to me in 1996,It
was found on a coffee plantation near the
village of Buenos Aire~below Cusuco
National Park.It is a very delicate colubrid
that preeumahly diga in the ground and
leada a highly eecretive existence.
I
I

species indicate that it i~ a llpcbunalmund- I

dweller ( D m& DOWLING1967).

Further Reading
D m Bc DWLINQ1857, O ' S m 1986,K O ~ LR
2001.

H Fig. 574. O m d i p h w rclcnrla [holotype).Fnom

KOHLERet al. (2001) I

Fig. 678. Nothapsis m@sw (Gem Sarrlaya,

AtlgfLticoNo*, Nicaragua),
238
Vine snakes are characterized by their
very slender bodies and conspicuously
narrow and pointed heads, AU species also
lack a loreal. They are rear-fanged snakes,
with mild venom that does not pose a
serioue threat ta humans, but can lead to
localized swelling and itching (CRIMMINS
1937).Large specimens need to be handled
with paxticular caution. When a vine
snake is hmdled, i t will react with a
threatening gape, exposing the bluealack
interior of its mouth; it will also bite with-
out any warning. Aa a defensive reaction,
this colubrid will continuously hold its Fig.577. Oxybelis breuimstris (Cerro Saslaya,
outstretched tongue rigid with the two tips Atlhtico Norte, Nicaragua).
of the tongue pressed together (compare
Fig. 675. Oxybelis fulgddus (Gamboa Forest
Besorb, Panam&,Panama). Photo: M. Luxidberg
'I
1 parously, laying clutchee of 3-5 (0.aeneue),
I
Fig. 576. OxybeEk aenaua (Bartola, Rio San
Juan,Nicaragua).
Fig. 577). The precise adaptive advantage
conferred by this unusual behavior
remains a mystery to scientists. All
Oxyblis species are diurnal and spend the
night cailed up asleep in shrubs or trees up
to 3 metera above the ground. They inhabit
savanna, dry forest and wet forest, where
they spend most of their time in ahrubs,
high grass, and treea anywhere &om 50 to
300 cm high. They can also be seen on the
ground (FRANzEN 1996, persona1 observa-
tion), In vegetation, they am well-camou-
flaged and very difficult to spot. A large
psrt of *
diet made up of
(Norops),but other lizards are also eaten
along with frogs,rodents, birds and insects
(SWART1943, H. SMITH& GRANT1958,
HENDEWON 1982, CAMPBELL 1998a). OR
Utila, Honduras, I observed a green vine
snake (0, filgidus) as it caught and swal-
lowed a mouse. Vine snakes reproduce ovi-
8-10 (0.fulgidue) and 8 eggs (0.wilaomi),
respectively, during the egg-laying period
in the northern portion of Central America,
which fdls between March and the begin-
ning of September (SEXTON & HEATWO~
1966, G ~ L D E1983,
R CENSKY8t McCoy
1988, GROVES1995, CAMPBELL1998a). A
clutch of 0. filgidus hatched after an incu-
 Colubridae

Key 'ts: W b e l i s
1 a 16 dorsal rowa at midbody; cloacal ecub
undivided; mually 6 aupralabials ...............
..............................
Qayblie BmdrosMe
b 17 dorsal row8 at midbody; cloacal cute
divided;wually 8-10 supralabials ...........
L
2 a Dorsal surface of head, body,and tail green
in life (Mue in preservative) ........................
.................................
Oxybelia ftsigidulr
b Dorsal surface of head, M y and tail yel-
low, brown, or p a y ................................
8
8 a Dared surface of head, body, and tail uni.
form1 yellow or yellow-brown;uaudy 10
nrpr&bials .................. Orybelis w i h 1
b Doraal surface of head, body, and tail
?ig.678.~ ~ x y b e l wil80nk
is (Isla Ue HOaWl, brown or gray with amall dark f l d ,
londurars). Photo:J.R. McCranie wually 8-9 sugdabials Owbelie menew

Further Reading
BWERT c%t OLIVER 1946, m ~ f 1974,
l ~ ~uaa,
bation period of slightly more than three 1991, VILLA& M c C 1995,
~ GROVES 1996,
months (CONNEB 1909), as compared ta ZEN 1996
an incubation period of 92-94 daya for eggs
of O. wilaoni (GROVES 1945).

Oq+WL U,,IJS ( W A ~ E Il ~w ) , &,I Nm.

Sew. Brae.: 12; type l d t g r : forest adjacent
to the Solimoens River, near Ega llbf~l, *
Brazil. TL to 1455 m.Southern Tamaulipas,
Mexico, and southern Arizong USA, ae far as
southern BradJ sea level to approximately
1600 m elevation in savanna, wet fomt, and ,
dry forest. No subapeciea are Tet25gIIi~ed
(KEISER 1974).
Oxy belis brevirorctria (COPE 186lb), Proc.
Acad. Nat. Sci Philadeiphia la: 866; type
locality: "Veragum, New CrPeneda"llbamal.
TL to 1200 mm. Eadltern Honduras to
weatern Ecuador, sea level to approximably E EO~yk[isbrevJtm~II.I*
$00 rn elevation in rain fareat.
Cheybelia fulgidus (DAUD~N 18051, Hiat. Nat. I
. Oxybells aaneus

Rept. 8: 362: locality: neipborhd of -

Port-au-Prince anto Domingo m arrorl. TL
to 2000 mm. lethmus of Tehuantspec,Mexico, I
to Argentina, sea lave1 to approximately 1000 -.- W'
m elevation in savanna, dry foreat and wet
fomt.
O*.ybelia widsoni VU & MCCW lase,
Rev. Biol. Trap. 49 (1-3): 297; type locality:
Roclry Point, b t a Elena on the wtem end
of Iala de Roatdn, Idas de la Bahia,
Hondurm. TL to 1982 mm. Endemic on Isla
de Roath, Honduras, 8ea level to 200 m el* A ~ x y k l i s&SBCM/
vation in dry foreet and wet for&, Oxybells fulghclu~
4
- . -1
240
The Neotmpical genus OxyrFsopus consists
of about a dozen specied, which are distrib-
uted predominantly in South America.
Only one species, 0.petah, occurs in
Central America. In the past, there was
some discussion as ta whether Lfpetola'' or
"petoEariusPshould be used a~ the specie^
name for the Central American Oxyrhopta
species (BAILEY 1986, H. SMITHet al.
1986b). B m (1988)was able to show
that "pedola*was correct. This colubrid is a
strong, cmpuedar and nocturnal ground- Adult O W ~ ~ W"o:P U ~(Germ
dweller. diet is up primarily of Fig. 579. Atldntlw N o h , Nicaragua).
Smlaya,
lizarda, snakes, and rodents. Oxyrhpws
petob is o v i p m w and produces clutches
of 5-10 eggs, which hatch after a period of

I
approximately three months into young
that are about 20 cm long (FITcH 1970,
CAMPBELL1998a).

I
C?~~hopus pemh ( m m s 17583,Systems
Watmm, ed. 10:225; i$p Ih
l d t y : "Ah.ie=aA
emorl, TL t~ 2200 m.Varaatlz, Wee, ta
Balivia m d Nartb- Brmd, sea h e l to
approximately 800 rn elevation in rain forest.
In Centrd America, the mxb~pecie~
QzyrAopus p. wbae D m = , BIBRONst
DUMBRft 1854 oormr.

Further Reading
BAILEV
1970b,1986,H.SMITHet al. 1986b

Dhimphk guianensis
5xyrhopus petola Fig. 680. Juvenile Owrhopus petola (4 km
--- -
, u-7 m t h of Ayapal, Jinotega, Nicaragua).
Colubridae
- 2
Phimophie
The genw Phimophie includes four mid-
size rear-fanged snakes that are character-
ized by their protruding, painted ro8tral

-I
shield. Only one species, l? guianensis,
o w e in so;thern central ~ m k c aIt. ie a
secretive ground-dweller about which little
is known.There are reports that thig enake
is predominantly noctumd, though it can
also be seen during the day, and that it
m
(LANCIN' '
feeds on lizard8 and small mammals
KORNACKERlQB9)' to
these authors, .?? gudalkensis reproduces by
Fig 582. Pituophir Eineatimllia (LaCepultura,
800,, Chiapag, Mexlm). Photo: A, w e e V.
laying eggs.

-
Ph0phi8
Only one species of the genus Pituophis, P,
Eineaticollis, reaches to northern Central
America. The others are distributed in
North America.
Further Reading
MARTINS Bt OLNEIRA1998,ZAHER 1999

M e r Reading
I STULL1940, D m w 1960

Fig. 681.Phimophis guianerssis (Colombia).

Photo: H.W.Greene

Fig. 585. Pitwphig ZineatteOllia tJalknango,
2000 m,Chiapas, Mexico). Photo: Ramha
Belize). P1hercw ezapQidrs
Fig* m&"gya
Fig. 687. Plimem~seuryzrrnut~(LaSelva,
Hesedia, Cwta RicaJ.
9 :
Pli~cema8 -
The false coral snakes of the genus
Pliocercue were placed by SAVAGE &c
CROTHERC19891, together with the species
of the Rhadinarea kateristriga group (sensu
MYERS1974)in the genus Urothca. While
the partitioning of the lateristriga group
into a genw separate from Rkdinaea is
justified, it also seema meaningful to sepa-
rate the PZhercus species into their own
genus (which would presumably form a
sister genus to Umtbca).h i d e from mor-
phological differences,this decbion is justi-
fied by the evolutionary history of
Pliocereuii a8 a mimic of the true coral
snakss (genus Micrurus), which fa not the
with the Umthca s p i e s (MYERS &
CADLE1994, H. S m et al. 1995a). H.
SMITH& CHI- 11996) have presented
the most extenlsive m d recent revision of
the genus Piiocemu, although it has been
strongly criticized [WILSONgt MCCRANIE
1497).
The lung tail, which will break off easily if
it ia gripped tightly, ie characteristic of
these colubrids. The fact that most s p i -
men8 of Pliocercus that have been found
have incomplete tails suggests that the
breakable tail is a survival strategy to pro-
tect these species &om predators (SAVAEE
& CROTHER 1989).STUART(1948) found the
tail of a Plioeemua dapoidea in the stomach
of an Indigo Snake (Drrmamhon coraisl.
Although lthese colubrilds are generally
harmless, they will bite if they feel
threatened. 1n humans, the bite of a
Pliocercus will, at most, cause a paink1
localized reaction and swelling of the
mandera, frogs, and amphibian spawn
(STUART N LEE 1996).
1948, D ~ L L M A1963,
During the rainy season add dm during
the dry sewon, these colubrids will pm-
duce clutches of 4 8 e g g ( C m m 1998a,
STAFFORD 199913).
Fig. 584. Hio;dercus eupyzonus (CMoba, Fig. 585. Juvenile Pseudoboa neuwkdi (road
Colombia). Photo. M. Lundberg David-Ctualaca, ChiriquI, Panama).
Photo: M. Lundbe

Nat. Sci. Philadelphia W 283;type lotalitg

4
Near Jalap, Vera-, M d m . TL to 86
mm. Oruraca and central V e r a m , Mexico,
o e r ~ h Honduras
l and El dalvadcr, em 1
to 2000 m elevation in wet foreat, dou
foreet, aod dry fbrest. In Central A t n e the
aubspeciae P. e. aequdis ESALVIN l86lbl
(easternChiapw to E B ~Honduraa
M aa well
ae all amma the Yucatan W a ) , P, e,
undrewai S m 194% (the north of the
Yucatan Penimula) and P. e. dfastema
(Bocom~188;g) [Oaxaott, Mexico,along the
Pacific eide to wtera El S a l ~ o roccur.
l
Pkiooemun eurymnm COPE1862a, Proo. h a d .
Nat. Sci. Philadelphia 14: 72; type locality:
R&cin of 'l'hmnh,New Grenada, C a l d i a .
TL to MM mm, North-tern H d u r a s to
CdomMe and Ecuador, aea level t6 1300 m
elevation in rain f m t and cloud foreat. In
CenW America, the BU~BJ&W R e. dimid-
iattrs CSom 186.8 (northern Nicaragua t o
ewtesn Cwta RSca) a d l? e. bughadti
u~ & c-3mM.B W96 (Panama) oomu.

Key to Pliacereus
1 a Body with three colom (black-red-yellow
or black-red-white rings) ........................
...............................
Plbocemue ebpaidee Further Reading
b Body with only two colors (black-red or SAVAGE& CROTHER1989, S m L CHI~ZAR lW,
.
black-whiterings) , PEiOCBmw ~ ~ M CWWN B et al. 1998,WWON Bt MCCRANIE
lW?L
 rwmces ie a Neotropical genus with four

!
I I
Fig.588. Adult Pseudoboa neuwiedi 1Chnboa
Forest Resort,Panam&,Panama).
photo: M. Lundberg
recognized species, one of which is distri-
buted throughout Central America.
Pseustespwcilonotus lives primarily in old
growth rain forest, where i t can be found
both on the forest floor, as well as in
shrubs and trees. If it feels threatened, it
will inflate its upper M y , open its mouth
wide and bite without hesitation, which, in
view of the size d aome specimens, can
inspire considerable
cil~norus respect.
is diurnal and feedaPseustes
primarilypoe-
on
birds, tree-dwelw
bird eggs (SEXTON& HEATWOLE 1965,
ALVAREZDEL Tom 1983,CAMPBELL1998a).
The speciea of the genus Pseustes repro-
duce wiparously, with females laying 7-14
eggs (GOODE1988).

P#m&duboa -
The genus Pseudoboa,which is distributed
:I m W l t & B pt?eCr60- (lim~
Snakas BRk. Mw, 1W. 86; typ@l d t y :
Hondure~ and Medca; reetricted to
~d-
Lag.

b u ~ l h~ t shp e d e a t ; i b n by
predominantly in South America, includes BOW.,EMOER (1BHal. TL ~ 2 1 0 mm.
0 Sari L&
four species, only one of which, 19 m u - 1 Potoaf aad Mexcot rn far aas Brad,
wiedii, reaches to the southern extreme of ~ g ~ ~ e t ~ ~ ~ ~ t l
Central America. This rear-fanged snake is @Mm intg 8uw to
nocturnal and lives as a ground-dweller in Bt m-imx
(19Sa) seems prablamatie (prmo~
dzy forest and rain forest, generally in the 1986). Pbynonnz shropshimi gi;
vicinity of bodies of water. Aside from 1ahregarded to k a SgeOn- of
lizards. their diet also include8 small m m - I? pmiEotwtus accordingto hvmm (2002).

Further Reading
W m s & O w m 1958, ZAHER1999

Rg,589. Psewtes poecilonohcs (Bartola, Rfo

San Juan,Nicaragua).
Further Reading
AWARdL 192%
Fig. 543.R M i n m o rogemnaani (Gem
Saeiaya, At1Bstico Norte, Nicaragua).

Fig,184, R W i m d&c@er ~ C Chompip~

m
Cash Rica). Photo: R.W.Van Devender

596. R M i m e o sdadelnaani (new

CMgntla, Cardillera de 10s Cvchurnatanes,
Hwhuetenango, Guatemala]. Photo: T*Bile

kh d i m h h r y m m a (Corn 18701, PMC-
ar, Phil. 80c. 11: 164; type W t g : un-
owl.TL to 443 mm, 5.0ut8eaehrn Chiapas,
esico, and mutl~weetemGaatarnda as wen
as in the Sierra de Agalta in eaat-Eentrel
M u r w (me M O C w st WWON 2001a1,
00-3000m elevation in pine-oak fomt.
1960, J. Washington A d . Sci. 89.:
Rio G r a d e , Om-, M ~ c QTL . to 292 mm.
P a d c side of the Iathmu~of lbh~~~ltepec 1370-1600 m elmation in rain fwd to weabrn Panama,
B astern O-a,
'm in rain forest%
Mexico), ~ 2 0 0 m0elma-
R W w m pibltaorum (STUART 1964), Rure. 8&, Mm,Camp. Zool, 88: 10; type locall*
Bid. Soc. - W 6% 176; type 1rmEQc
Finca La Gloria,abut 960 m elevatim, abut
12 km northeast of Chiquimdilla, 8anta
ROBP, Guatemala. TL tt, 310 m.
m
Souhaatern Guatemala and wsstern
Salvadrrr, 670-9513m elevatian in wet forest.
R b d i m a osacbi (8- 19363, Pm.
Zalif h d &. ii& 19; tgp.LOFaliw aauthem
dope of Volch Zunil, Buohitqquex,
Guatemala. 9% to %8 mm, Southw~~tern
Guatemala, 6881800 m elevation in wet
forest.
R M ~ c r e t pulueriventris
a B~WNQER 18&,
Cat. Sna%ee, Brit. Mw.8: 636; type l d $ :
Aaahar de c- C*Rica T L b MltL
m r ~
Central Costa Biea
and
clwd fom&.
Rhod5naea r r r g ~ r ~ ~ m a m i & McCRAME
lma, Benckenbergima bid. rB. 244;
locality: 8 a I i p d C e m BuLga
3fi002.8W), 1460 m elevation, AtlaetIoo
Norte, Nicaragua. TL to 288 mm. E C n m only
from the tm localit;y,
R M i m wwnti Dmm i% BAILEY 1989,
"sUu e d - E & p mUp, ~ ~ ~ 1880 ft, near W
f oPep~Paaam~~.TLtoSldmm
&the EandZam in central Panama, 300-770
n elwation in rain foreat.
 - a
x-

Fig. 696. Rhadinaea kanalchzltchan (8.8mi SE Fig. 697.R h a d i m a pulverivenbris (Fortuna,

San CrisMbal de lam Casas, Chiapas, Mexico). Chiriqui, Panama). Photo: C. W.Myera
Photo: C. W, Myers

I B h , Rnc.
M: 21% typa ,
locality: near Ban J&* Costa Rea. TL ta 441467
mm. Central Costa Ria,1220-1460rn
tion in rain farest and cloud foreat.
Rhadintrea staddmani STUABT 1&
UP; *t y e l d WMw*
1W1, CXoc, Rip.
vatim. uebuete-.
:
Zool.
woe
Unh.

460 mfn. CJjerra der 1

s'; C u e h . m ~ t a n eend"
ad cuilm im westem o ~ a t o m ~ l a (
~antaaps
~ l s o ~ l it omi p o l a ~ rHOLM
~~ CRUZ D?
~
1 I
1994, H ~ ~ I 6&~ 1% ctype a hadits: Ow&
lan N and 1.6 km E C m San Ekaneieao iric,
theCwrdilEeraNomfnradeDios* lm melava-' Fig. 6B8.Rhudinaera decotvata IBarhla, Rfo
tiotl, Aubntida, Hondurm, TL 381 m! %,J~,, Nicaragua).
h a m only 6om the type lcdity [cloud
fometl.
R h a d i ~ l e a vemlulaticepa (COPE 18BOd).
R o c &ad. Hat. M.PWdelpbia I& 249;
type lmalitr Veragua, Panama TL ta 374
m. Weetern Central PBnama, 700-5DO m
dwatim fn rain Pored..

I
I Rhadheea Fig. 699. Rhaditaasa decorata (Nusagmdi,
I
Corn- ds San Blaq Panama).

1 o 19 or 21 dored rows at midbody .,,.,,,,,.,,,.9
b 17 domal rows at midkdy .......................8
2 a 21 dorsal m a at midbody .......................
b 19 dorsal row et midbody ...................
8 a Venter white or yellow; pde nneok band pre-
sent (may alea be fr ented) ...................
..........................
.;;%&-a
b Vmtw red-oran without pale neck band
gdmani
......................g&n- ~ m m a n i
4 a Very wide and eontsaeting dark longitudi-
nal etripea present, lateral atri e5 (on ard
dmal row and a+cent
ea (an the ends of the ven-
tmlaterd
trale @Ti'
and lB amd tow1 ..........................
.......................RIradfmmw mo-tieti
b L0ng;itndind strim narrower than above
and nat m oobviourhy contrarag aa above
6 a A mnspieuous, narrow, dark longitudina
106 h w e r (1st to 3rd) dorsal d e rows nni.
color black, eharply ~ontraatipgwith pale
rnediodoreal area; mnapicu~ll~
lar markings on top af b a d
paIe reticu-
...............
8 .............................
Rhadiaam aargsnti
,.,,4 b Dorsd and head markings diffemnt than
thome above ................. .. ...................
11
Ila Domum either with a dark d m o terd
k
e dong tlm border of both the 4 and
@
dmsd m a m predominantly unim10r
brown ........................................
. . . ...........
......
R W i u a m pulmrJvsntrds [inpart)
b Damd and head matbga different 1B .,..,.
ven- 1Ba Two pale spots, one of which is diF&ly
bhind the eye, the other an the neck theae
spot@may dm blend into a pale stripe that.
ruasho&wntd posCerim1y ..........................
...................... .....
RhruSbaw dmorato
-
b k o m the eye, a pale e M p e runs obliqaely
a m s a temporal region ........................
.,
.....................
l i h d i i ~ a e r nracdougolli
r

&ripe t p ? u z z g deag the borders of the 184 18a A wries of dark am,bam or a con&uous
and 2n dorsal m e ............................,........
stripe along the ventmmedid b e
........................Rhod-
.............
.........................Rhaddnaea mrgrwmte~ ea&@u~der
Without a longipdinal atripe dong the Ventral markings different .................... 14
bardem of the 1 aad Zn doreal m s , d- At leaat ma obviaus dark longitudinal
though othier longitudinal stripes may be
present .................................,................
8 tripe p w n t M aide of body .,.,..........
16
Rody without obvious &ark longitudhal
Mpe mmhg along striper or an$ with dark vertebral tripes
vents& ...........R~~
dorsal rcnxts; 169-191
utodeZmamS
................................................... m
~ t b w t ~ & a ~ W s t r i p o n h d t h , mostly D d c verbbral stripe short and restricted
7 or 8 dorsal m e ; 166-1'76ventrals 7 to neck area; a black &ripe pmmnk
on dorsal &ee of head .....................,.,
7 a 13-16 maxillary teeth sWpsd OX UTljcolc~r
.......................
R W n a e a kmpefeadw
.....
RftadSnaaaplslrrerd~~entdu (inpark)
b Dwgd aed head markings diflbrent than
b 16-18 maxillary Wtb; witbout lo^^- thase above ...*....................
... 18
~~
..a,....

ma1 stripe8 h a n a l c h t e k n l e a V&F rad tD read-orange (in life); doreurn

8 a A poatormlar pale spot or longitudinal with three dark l o n ~ t &ripe@ u ~ (verb-
stripe begimbq at the upper posterior m- b stripe plus laterd at ripe^ dong the
ner ofthe eye (Fig. 6Ma) .........................
B 4fff dorm1 row md djaeerit areas af the
b If a ppatocular pale wet or longitudtnd zrd rwrw, ............
J3hzd1naea tol-m
atripe iB pment, it b e g b at the lower b Venter white or meam colored; dorsal
pa&teriorcorner ofthe eye (Fig.$0(3b3 18 .... rnwIrin@dXomnt than those above ...,. 27
8 a Braad dark verteW .stripe intermittent l7a Without a large, pale a p t on each fro4tal
dong the middle of a pale line, mch that shield, dorsal snrfaee of head ~lnifowlp
the markin@ could b mistskm for a pair dark (Fig 807a); fewer than 159 v e n m
of paravertebral skipea; cons EGuow pale one postcedar ....................................
.I0
reticulum on dor~al.surfkc8&Read
.................
.....,......
~ ~ # ~ C E L W * U
, dark spot on each frontal shield
b Ausu1 % y clearly formed (E'ig. 607b);if with-
b Dark vertebral strips, if present, not inter- out pale h n t d spat, then more than 1M
mittent along the middle of a pale Sins; ventrale; uauaUy two postormlars ..........18
pale reticulum en dmsd amface of b a d l&aOne paetdor temporal; dark vertebral
preeent or not ........................ ............
,. 10 atripe covem the v e r t e M and part@aftee
Fig. 600. RIracEinaea serperaster (Monteverde, Fig. 603.Rhdinaea rogemmani (Cerro
Goeta Itica). Photo: U.W.Van Dwender Saslaya, AtllSntico Norte, Nicaragua).

Fig.601.R h a d i m ZacfLrynaatbs ( P hLa Pi- Fig, 604. RWilucea godnnani (northern

mch9, Sierra de &aha, O h c h o , Honduras). Hondur~t~). Pboto: J.R.McCranie
Phok J. R.McCrani~

Fig. 602. R h a c k ~inkelini

~ ~ (14 km north of Fig. e05.Rhadinaea pilonaorurn (Fin= La
Marcala, Honduras). Photo: J. R. McCranie Giralda, La Libertad, El Salvador).
Photo: E. Greenbaum
 b Two posterior temlpody dark vertebral
stripe rmtrieted ta medial third of the vm- The genus RFainsbotkryum indudea two
t&al row ..*...,.*, Rhud&uwa bnmkdlat &ee: R bovallii, which mculg in Central
1Ba Dored pattau donei~atsdby palm of la- America, and the South American spades
Wal and $81ikrolabrdb n g i t u a atbiw R Eentiginoarvrn. Rhlnahthrytsm bov&llii is
........................R;rsluthaatrsChymam a ssldam-ee~n, nocturnal rear-faaged
b Damal pattern mom complm and not make* noted for the rernwkatrIs conlmwt-
domia8t;ed b~r I% of lateral and ventro- ing coloraticna on i b head: each shield on
lateral longitu LEL e.ipes ........................
............................R W m W a mktF3Sni the dorsal surface d the head ha% a hlaclt
spot lximmd with white. It is a t r e d w d -
Ier in primary rain forerst that fee& pre-
dominantly on arhd lieards, frogs, and
d mammals. Rhimhthrytbm bavcaZIii
rapmduces by laying sg@;s as verifEd by
ths Q ~ E W V ~ ~ ~ Oof
- ZJ[OHPXEISTEXE caaov.
c m P B ~ wBt
. '6rAmam (1BSgE list this 9pe-
elm for the PeEn R@m af GuatemaIa, a
claim that w w not later suht~ntizlted
(CWW 19%~) which, therefore, must
be regadd aa queathnable.
/'- a. R. decorate

Fig. 606. Lateral head pattern in RhQdirusea.

Fig,608,Rhimhbhty~rnGovaZLii (Costa RicaX

Phota: R. D.Bartlett

a, R. montearfstl b. R,klnkeffnf
Fi 607, Doreal head pattern in RMinuea.
 -
Sdvhra
'fhe specie8 of the genus SaIvadorce all
have a canspicuowly large metral shield,
that extends further upward on the anout
than in other colubrid~.These are agile,
diurnal snakes that inhabit the dry are88
of the southwestern US and Mexico, Only
one species, % Eernniscata, ranges as frrr as
the narthern portion of Central b e r i e a . It
f i primarily on l i a & ~ B ~ G E R T
1999.)
and will lay c h h e s of 6-12 eggs, which
hatch after 3-4 monthe of incubation (OBST
-I II et al. 1984, D.SCHMIDT 1990).
Photo: ''W'
Fig. 809. Rhimboth~ytsrnbovallii.
Devender
fialvndam h n i s c o t a (COPE18961, Trans.
Amex. Philos. Soc. 18: 203; type locality:
weahern Mexico. to 1300 mm. Alon Fade
wmt of Mexieo fmm Guensm ta dhihiapas
also in Grijaba Valley of Chiapas and adja

1
cent Guatemala; sea level to 1000 m elwati
on on savaxum and in dry forest

Further Reading
BOGERT1939, W E ~ E &
R S M I1982
~

Fig. 610.Rhinobothryum bovoUii (Coata Rica).

Phoh: M. Burger

~hinobothryumbovi-N 1816
Medd. Nteborga M u . 2001.Afd. 9: 32;t p
locality: Giquirres, C a ~ t aRim. TL to 130C
mm. Southern Honduran to Ecuador and
Venezuela, sea level to 600 m elevation in
rain foreet,

Further Reading
HOHMEISTER2001
Fig. 611.SaEvadora lemnbcuta (near
kktequilla, Oaxaca, Mexico).
Scaphiodontophis- -
-
( h v m13mT ~ R gt 1958,& v m
O SMITH
DEL "hE0 1983, HEWERSOH 1984,
Amrding to ccuncmt howledge, the genus GAMPBW 191)8a). to the apciw of
Scaphisdonhpht is mm~tspie,contahiqg anndutuls d e o has a very l U?athea,
tbe genera PlZocercus
~ n g
tail
$:
that;
the 5pecies 5.anmda~s, with S, uenwtis-
d m viewad E ~ B8 8yIlbny~athemf EQAVAGE will M y h a k off ifthe mindis m i d .
L QLOWM~KI l W B ) , ScapAiodontophis have Most of t h spedrnem
~ ofthis epscieg fwnd
anttdatua Tivea on the leaf dmwn p m d
incomplete tails.
of the r $ h fsraElt and ie an agile, diurnal

--
hunter, that feeds primarily on lizard9
@3mllzml$%X, ~ E B ~ O1984). N Their
h i q d and rshovel-ahaped teeth are parti.
mlmly well suited to holding onta their
p * ~prey, tihe hnnwth, ale- aW
Sphenomarpk lkg cherriei, SeapJaiocEot&-
0pJlis allmtatl~~ m@uel &pmu~1y,
laying clutches of 1-10 OU.(B- 2.4) egg8 H E ~ I W D N
1984,H.SMITH att al. IlQ&dd,SAW
under rotting or in o k auikb1e L ~ L U ~ B KlsCrs I
hiding places d&g the raioy seaeon

Fig.618. Scaphiodontophis annubtus

(Veracruz, Mexioo). Photo: P,Heimer
Fig. 612.Scaphbdontophie annulatus (Finca
La Giralda, La Libertad, El Salvador).
Photo: E,Greenbaa
8~1lecaphie&wim$us is a mpimtm
gmund-ddw in the & are- along the
M c ~ o a eofCentFalAmdea.Tbemmge
t
red spot on its ba& is so briget that it
mema to have been painted on with fluor-
emant paint. The spwiw is diurnal and
feeds prkmdy on bae& rtnd araWds.
species repduees oviparowely
aE@amm 2WO).

-
T b genus &aticokda was e~W2shed&
D f ~ m& o FEIES(198371 for apedm
triaspb,which wasl formerly .bgiuded hi
the gpus Elapk. The young of this epe-
cie~are mxlspiduou~lyspotted, as oppased
to the IEnicolar yelIowish bmwn or olive
grem ddt spxhnens. SeadicoZL @hspEs
i~ ti nodxrmd, t
e r
r
m w i=awmi-arboreal
colubrid &st eat# p r i m e md anam-
nzaZ~a d the occasi~nd1E~ardOF bird
CLlMw~rno19&0,CAMPBELL 1W8a), Pemdeis
ma& ssxual maMty mu;ndthe age OF
bm, %dth a snoutcVmt lm& of at leaat
626 mm (C-EW. ltd9tW. This wipamtw
rspecis predumsc clutcheg of 3-9 egpd
which hat& afbr 75+100d w at an incu-
bation krnperatum of 2ti.29 T! (i%mm
im1.

Fig.816,Adult SatsChiia frhw L (El Im@le

National ~k ~hnact~a* i s~~~IM'I.

Pig. 817,Juvenile Senticolis trimpis (El
Imposible National Park, Ahuachaph, El
Salvador).
The genus Sibon Indudes about a dozw
epeeim that are distributed in Mexico,
Central America, and South herica.
SMITH(1982)pointed out that the gender of
the genus name Sibon is maemline, thus
the epederr ahodd be called annulatw,
dimidiatus, and nebulatw. Similar to the
epeciee of the genera Dipme and
!hpidodipmtz, the Sibon s p c i e ~apecialh
in snail8 as a food s o m e . The morpholog$-
lcal adaptations that enable these cdubdcb
to pull their slimy prey out of their she&
include partidslly long teeih in the lower
jaw and the con~picuouelyshort snout of
most of the Sibon species, All species of
this genue are nocturnal and l i e predomi-
nantly in trees and ah rub^. They reproduce
by laying 2-6 eggs per clu&h &OFRON
1983,1987).
Sikon a n n u l a h (GONTW lBm),AIla Mag.
Nat. Hbt. (4) 0: 30; type Iocality: Near
Garbgo, Corn Rica. TL ta 667 mm. Northarn
Nicaragua to emtern Panama, 8ea level to
about ZXHl m elevation in rain foreet, I
8 i h n .n.nthrom (COPE 1868). Pme.
N& Sol. ~ h i l a p h i aPO: UB;type
hadl
Central America. TL ki 664 mm.Pacific v
eant hEl Salvador to C a b Rica;
mala (Rfo Motagm Valle ) and Hondurae
1
(mo va~ay) on Earibban v~tgant,~
sea lwel to about 500 m elevation in
foreat. I.
Fig 618,Senticolitr t r i m p i g (Rancho A h p ,
CAiapas, Mexico).
 - - - - - -
Colubridae

Sibon a r m (COPE
Philadelphia (2) 8:
R i a . Tt to 690 mm*Southeaeternh t a Rica
to &rn Panama, ,wa Ievel to h t 600 m
elevation in rain foreat.
Siban carri ( S m IDtiXI, Copeia 1M1:M;
type l d t y : EscueIamcola Panamericana,
Central El or
n w TeguCipI ,H o n d ~TL . to 408 mm,
ta toaut&ern Hand-,
sea level to about 1OOO m elevation in dry
fore%t.
Sibon d i m W w [(GONTHER 18721,Ann. Mag.
Nat. Hist. (4) 9 31;tspe l d t ~ rM.: TL
to 680 mm. Vemcnm and Oaxaca, Mexico, to
southeastern Costa Rica, sea level kr about
1600 m elevatiw in rain fbwt and cloud
fmt.

NrttI. Mw.86: 457; type locality:

del !hro, Padama, TL to 624 mm.
vemant of eastern Hondurw Cash M a ,
western Panama, 4W-600 rn elevation in r
b 8 t .
9ibm d u h b u s (LINN~Eus1758
Naturae, ed 10: 222, typa locality:
TL to 890 mm. Nayatit andvsracruz,M e h ,
to Ecuador and Braail. ma level to about 1500
rn elevation in rain &d dry forast,The sub- I
species Siban n. wbdaturp (M&w to central
Panama) and Sibon R. ~ u c o n m eIBOIJLEN-
~
QER t898b)(eastern b~northwem-
Ecuador) occur in Centralh a r i c a .
S i h n mnnhia ( C a r 18671, Rac. A d . Nat.
Sci. Philadelphia 18: 318; type locality;
I
Yucahh, Mexico. TL to 430 mm. Norkhern
and Eastern portions of the YucaUn
PleRinda, sea level to abut 960 m elevation
in dry folreet. The aubspecie~Sibon a 8annio.
la (noithem portion of Yucatsln Peninsula)
and Sibon w tmei6ld HENDERSON, H O ~ & W
W n s o ~ 1977 {Belize and narthewtern
temala) occur in Central America. -
-

Fig. E Sibon a n d h k w (nt ?ueblo Wieo,

Jinotega, Nicaragua).
Key to SZboa
1 a 13 dorsal rows at midbody .......................2
b 16 dad mwa a t midbody ....................... 8
3 a Without anterior temporal, first p&ndar
supralabial in contact with parietal (Rg.
621d .....................................
Sibus nand
b b e anterior temporal,no supralabial in
contact with parietal .,S i b m t h m p s
8 a Dark dorsal ban& or apoQ reach at most
to outer edges of ventrals, although duk
B ~ mw. W be p w m t on venter; en-
wis, ,,
iar ed penultmake aupralabl in contact
..*............................................. 4
b Dark dorsal h d s or apob reach well onto
venter or even across venter; enlarged Fig 620. Sibon ~nebulatw(El Impoeible
p e n u ~ ~ aupralabial
te not in contact National Pa&, Ahuachap& El Salvador).
with eye . .
...,.......................... ............ 3
4 a 181-201 ventrals; 112-121 subcandale ......
..................................... S i h argua
b 161-175 ventral^; 82-108 rrubcaudals .........
...,...,..............,..........S i b o m Eom&fi.etsis
I a Firat infralabials in contact with e a 1
other behind themental S $ k n n&uktwr
b Firat infrdabiale sleparated from each
oteer by ode or two postmentalsbehind the
mental (Fig, 822) ....................,.,,.... ........
B
Ia 8-1Q eupdabiala, t h a e in contact with
eye; 145-162 ventrals; 57-88 suhaudals
.,...........,........................
Bibm samnioIe
b 7-8 su~dabiala,two in contact with eye;
161-200 ventrals; 106.133 eubcaudals ..... 7
'7 a Usually one postmental present (fig.
622a); 175-200 veatrals; d m d surfam of
head usualIy exte9pii~elyunicolor dark
brown; stout body; ratio of head b. b neMatus
lengthhead width >Id; ratio of head
"" 8iben *ww' Fig, 621. Lateral view of head in two sped= of
b Usually two poetmentala present (Fig. sibon. Drawingar: M Veeely
622b3.;161-192 ventral@;dorsal aurface of
head gray-brown with irregular dark
brown s p b ; delicate build; ratio of bead
lengthhead width e l . & ratio of head
w i d w e & width r2.7 S d h annulatas

FtahwRaading
etNal. 1977, K O ~ O N
PeTElw 1960,H E ~ E R B O
1985,1485, McCw 1@$6,PtgPaa~1990,SAVAOE
& MCDWD 1992, MCCWIE et d.2001

a. S.dlmEdlatus b. S. annulabs

Fig. 6m.Scalatiun of chin &on Ipostmen'talil

orange).
 a Colubridae

Fig. 623.Sibon annulatus (Cerro Saslaya, Fig. 626. Sibon anthracops ( E l Salvador).
Atlintico Narte, Nicaragua).

Fig. 624. Sibon sanniola (near Coba, Quintma Fig. 627. Sibon dinaidiatus (Selva Negra,
Roo, Mexico). Photo: J. C. Lee Matagalpa, Nicaragua).

Fig. 625. Sibon dimtdiatuar (near Quebrada Fig. 628. Sibon lzebulatus (Bartola, Ria San
Orande, Ceph, Hondurm). Juan, Nicaragua).
 The only species of the genus Siphlophis
that occurs in southern Central America is
S, cervinus. V u et al. (1488)emneously
listed S. longicaudatus from Panama;
however, its distribution is restricted to
Brazil IJ. PETER^ gt OREJAS-MIRANLFA 1970,
WEZ & SOLIS19931.Siphlophis ceruinus
is a tree-dweller in warm and wet lowland
rain forest and feeds on frogs, lizarda, and
small mammals (COSTA PRUDE^ et al.
1998).The ~peciesreproduce^ oviparously
/Pfimz-Smos & MORENO 1988).
Fig.629. Siphlophis ceruinus (Peru).
Siphlopiris cervinus (LAURENTI 17581,
Synopsin Raptillurn: 88; type l d t y :
America. TLto &15 mm. Panama to Boliviaaa
well on Trinidad.

Further Readin
H m 1964,COSTA b UDENTE et al.1498 through vegetation, thw its Spanish
name, %ladoraw (the flying one). It fix&
primeuily on mall mammaIs, birds, and
bird eggs (HENDEFSON & H O E V E197Tb,
~
LEE l!396). We saw a large S, pulktus at
the fbot of the Mombacho Volcano
CNi~~~agua), that was in the prows o f
swdowhg a Mexican Porcupine ISphig-
T'he genm Spilotm conaista of one widely. gems mexicanus) (K~KLER & SEPP 1999),
distributed Bpecim, S.pullatus. This large Spilotes pdlatus produce^ aviparody,

I diurnal dubrid can be faund bath on the producing clutches of 7-26 eggs, which
pound add in. the branches of large trees. hatch in about two months I H V
This snake is extremely agile at moving 1986, D.SCHMIDT 1990,CAMFBELL 1998a).

S Go&* p u l h t w (LmxAEU8 17681, ~ y s t e d

dturae, ed. 10: 3%; type lmlitg: 'Asia"
m r 1 . TL to 2600 mm. Tamaulipw end
Qaxaea, Mexi? tq &qn2ina, sea level b
1500 m elevgt~mm ram foreet, wet for&
and dry forest. Aaco* ta WilsPn %G MEwm
(19861, the diwleicm intO aubapdw euggeate8
by Aktm~,(1929a) ie insu8Btiently ~uppmb
4, and I prefer not to m m p i z e any eubspe-
*.a$ k&kC d d -
Fig,630. Spihtes pull- trying to swallow a Me;niean Forcupine ( V o l h kbmbdm, Nicaragua).

Stenomhtna fmminvilbi D&m, B~BRON %

lk&m 1854, Fap. GBn. 7: 868; hype locality:
Mexico. TL to 700 mm. Veramz and
~ 0 t hspecies of this genus are distributed G ; Y e m , Mdco, to Costa Rim, ma level k
widely across Mexico, Central Ameriea, 2209 m elevation in dry famtand wet f-t
and northern South America. Tt is charac- According to Smm (1963) and W m o ~8
teristic of this genus to have the prenasal M r m I~l985),the divieion Into subpecies L
inad5ciently supported, and I prefer not tc
scale fused with the internasal scale.These remgnber any subspecies.
are harmless, rear-fanged makes, which
will not attempt to bite. As nocturnal
ground-dwellers, they live predominantly
in dry areas. They feed d m w t exclusively
on spiders and scorpions ( D m m 1963,
SEXTON& HEATWOLE 1966, personal ob-
mrvation). Both species are oviparous and
produce clutdie~of 4-9 eggs during the dry
season (STUART1935a, CENSKY& MCCOY
1988).

Sfenomhim i%genhardtii (E~snrom18461,

Nachr, krg-Auguilt~Univ. und K Des.
m ~Gaminpen
~ . 8: a9; type locality: wwtem
Colombia (see MVEBs & BOmm 1996). TL to
~~
800 mm, Southern Veraerua, Mexico, t o I
Ecuador and Veeexuela, aea level to 1850 rn
elevation in dry forest and wet forest.
According ta STUART(1963) and W n s o ~k
Mmm (1985), the- divi&ioninto 8ubpdee ia u
huEciently su ported and I prefer not to Fig. 631.StenorrhSraa degenhadtid Peramus,
recnenim anv nuftnmcirc~. M~co), Photo: P.Ehimea
Colubridae -.,..
- - L
Shmda - L '

The genw Storeria includes twe s p d m

that are distributed widely amoee North
America, with the species, S.&krayi macb-
ing only to the northern poFtian of Central
America. These are ~ m gmuad-dweling
d
colubrids that feed aa invertebra- such
as earthworms, slugs, and s& hse&, At
the beginning ofthe rainy seiwon,S: dekayi
gives birth to between 3 and 24 live ywng

-
(WRIGHT &WRIGHT1957, CPLMPBEU 19983.

'storeria tlekayi [HOLBBOOK 1~142),North

Key to Stenomhinu
1 a MQWJthan 169 ventral~;dor~umlongitudi-
nally striped or unioolor ......................... COPE1885 OWWB.
......................
SfenorrMna fheminwillei

1
b Fewer than 160 rentrala; dorsum with fine
spotting or unicolar ................................. Further Reading
....................
Sfemmhfnadegenhadil TRkPIDO 1 9 4 4 , A ~ D ~ 1461,
~ s o ~C?EUTN%N1982

Fig. 633.Stenarrhina degenhardtii (CSrdoba,

Colombia). P h o h M. Lundberg I
Fig. 634. S t o r e r i ~dekayi (Hondura~,Dpto. de Fig.635. Symphimus leucostornus (Oaxaca,
IntibucA). Photo: J. R. McCranie Mexico). Photo: W.F. Duellman

Both species of this genus ate diurnal and

semi-arboreal, inhabiting dry areas. The
species mayue was earlier included in the
genus Ophedrys (H.SMITH& TAYLOR
19461, until R O ~ ~ M A
& NSGRAEFER (1974)
placed it in Symphimus. Symphimtls
mayae is known to feed on insects (mo~tly
crickets and grasshoppers) (ROSSMAN&
SCHAFFER 1974). The species is oviparous
and produces cluWnes of 2-4 eggs during
the wet season (STAFFORD perer. c o r n . ) . Fig. 336, Symphirnus mayae (Cdakmul Bio-
ephere Reserve,Campeche, Mexico),
Photo: H.Bahena B.
I

Key to Symphdmua
1 a Prenanal fused with adjacent internaaal;
.........................
..............
162-181ventrals ..............................
Syntphirnue kmastormss
b Prenarsal not fused with aqiacent kite-&
150-186 vwtrals .........Synephimm m q m

This species contains 58 species, 22 of
which occur in Central America. These are
without exception small, secretive, harm-
less oolubrids that will rarely attempt to
bite when they are handled, They are most
commonly found by turning over rotten
loge or rocks in the forest, as well as in
meadowe and on coffee plantations. They
feed on invertebrates, auch as beetle Iarvae
and centipede8 (VAN DEVENDER & COLE
1977,LEE 1996, C ~ l r d ~1998b).
m All spe-
cies are oviparous and produce clutches of
1-3 eggs (VANDmmm & COLE 1977,
F ~ 1985).
H RntiUa E ~ C L , BWIZ~ON
! k n t i l l a ~ o m 1926, Occ. Pap.
Boston Sot, Nat, W t . Q: 166: tme localitv:
Barn Colorado laland, Gatun h&,~ammk,
TL b a12r mm. Barro Colorado Isbind,
Central Panama, around 100 m elevation.
Kuawn only from the holoQp8.
,I
Nat. Hist. PubL b l , Her, !M: 3%type locality:
M&ida, YumUn, Mesico. TL ta 220 mm.
Norkhern Guatemala and the Yucatsn
Peninsula,sea lw0l to 60 m eIevation.
TbntiUa im e m C ~ C 1998b, L Su. Pap.
Hat, Hi&, &us. u&. ~ U1: B: I
T1&-
ty: AIdea San Miguelito, 480 m e evation,
1512'N, 88*43W,Sierra de Crud, Municipio
de Mordee, Imbal, Guatemala. T L to 726
mm. Ewterm Chinpas, Maim, to weatem
Honduras, a w level to 1600 m elevation.
Tontikla jrsni (GUNTHEE B96), Biologia
Centrali-Americma.R e p W and Batrachia:
148; type l d i t y : Ouaternala. TL to 242 mm.
Pacific versant of Guatemala near the
M a i m bwder at around 1000 m elevation,
~~ 1982 h regtuded to be
a aynonym of T.jani according to CAMPBELL
(IsS8b),
'Aantiilalempinr WmoN 8t M m A 1980,Mem.
San Die Sot. Nat. Hist. 11;25; type locality:
41 km & Tegucigdpa, Fnndaca Morplln,
Hondurae. TL to 25.4 mm. So~~th-centsaI
Honduras,1460 rn elmatian.

Mag. Nat. Hist, ('TI 14: 9W, p e w@

~ +.
!&mtiEla alticolu ( B O U L E N G19091,
h t a Rita, northern MedeUu? olombia. TZ
to 827 mm, Central Nicaragua to north-
western Colombia, 90-27QO rn elevation in
rain foreat and doud f m t .
TbrtfilIa rsmiJlta Corn 1876, J. h a d . Nat.
6ci. Philadelphia (2) 0: 143; type locality:
Middle Costa Ria. TL ta 4Wl mm. On the
Pacific veraant from Guatemala to the
Meaeta Cetrad Occidental ofCosta Rica; also
on the CarlBbean versant in Honduras; aea
level to 1430 rn elevation in dry forest and
rain forest. Ybntilla amiElaja wasi regarded
aa a synonpn of T melmmephrsb by WWN
L M m (1980)but r e s d ae a valid ~ p e -
u r n by SAVAGE I2002l.
Thntillrs bcsirdi &ART 1941%Doe. Fap. Mun
Zool. Utliv. U l G g m 458: 1; type localiw: 2
km SE Finca Chichh, about 1560 m eleva-
tion, 10 &line km S CoMn on Chameloo
ttail, Alta Verapm, Guatemala. TL to 465.
Alta Verapaz and Baja Verapaz, Guatemala,
1800-1550 m elevation. T. lempira
ItmtiDa breviwtldn 1962b, Zool.
Anz. 14g: 137;type locality: El Mtot1510 rn
elevation, Los Angeles, La Libertad, El
Sdvador. TL to 171 mm. South-central
Guatemala to n o f i e ~ ~ t aEl
r nSalvador, 600-
17M) m elevation.
Tantilla
Fig. 638, TbntilEQ:britareniatu &la de Gwqja,
Honduras). Photm J. It.McCrania

Thntilh mfteeps wag regarded as a synonym
of E melanmephaJa by WITSON & bbH&
(1980) but rmlureckd an a valid apeci~st
SAVAGE (2002).
2kntilla schistosa (I~~COURT 18831,Miss.
Mex., Rept.: 584; type l d t y : dta Vereps,
Cluatemala, and Mexico. TL to 293
Veracruz and Oaxaca, Megco, to
Mama, sea level to 1600 m elevaCion.
Tnatillu supracincta PETE^ -1
Momtaber. Wng. Akad. Wise.Berlin 1 - b
272; type locality: Guayaquil, Ecuador. TL tB
582 mm,Southeastern Nic~vagaato Emada~e; 8 a
sea level ta 860 rn elevation. aZzntiEla aranul&-
tu 30- 1892 is regarded to be a syn@
nym of T supdncta: armding to WIL~CHY'
1987~).
hntikla taeniata (BOCOURT 18831,Mb. Sci,
Ma., Rept.: 587; type I d t y : Guatamala. TL
to 416 mm, Antigua Basin and the m a -
eatern higblanda of Guatemala amwe @
Salvador and Hoadurae ae far south @i b
lorthem Nicaragua, sea level to 1650 ah
:lwatioll.
h t i l l a t a p e WWN 1988, J, Herptol, lb
jq; type locality: F i c a Ban Jerbimo, 7.5 Is@
R (by road) Cacaahatb I= Cacahoath ~ l r B a
Cacahuatdn], 760 m elevation, Volck
TacanP,Municipla de Unidn Juhez, Wpm@,
Mexico. TL to 317 mm. Southeastern
Zhiapae, Medca, 760-960 m elevation. 1- b
h n t i l l a tecta CAMPBELL8t SMITH 1997,
Be tologicti M. 333; type l d t s slope
R x gthe NE side of LsgunaYaxhA, 220 rn
olev~tion,17"03'48"N, 89"23'12"W, PetBn,
Guatemda. TL to 222 mm. Northemtern El
Pet& Guatemala, 220 rn elevation in dry
forest.
Tantilla trhwriuta SMITH EQ SMPPEI
1961,h c .
Biol. Sac. Waehiagton 84: 97; type E d & :
Coatlb, Oerxaca, Mexico. TL to 203. South-
central Oaxaca, Mexico.
TcmtiJZa tritmniatu SMITH BE Wn~vinas1966,
3outhwe&. Nat. 11: 483; type locality;Isla de
Guaaa~'a,Islm de la Bahla, Honduras, TL to
I 8
278 Ida de G u w a , Efondruas.
?bmbiEa vermipomb (HALLOWELL 18603,Proc.
h a d . Nat. Sci. Philadelphia 12: 484; type
locality Nicaragua. TL to 157 mm. Pacific
aide of El Salvador to nartbwmtern Costa
Rica, sea level to 620 rn elmtlm.
Thntillu vudcani CAMP= lM8, 8ci. Pap,
Nat. Rat. Mus.Univ, Kansm 7: 11;type 1c+
d t y Fin= El Carmen,618 m elevation, km
197.5 on CA-2,Quetzaltenango, Guatemala.
ll . +n ad6 mm PnFiPir side of Guatemala.
I
Key to TsntdMcc
1 a Dmum with ale darkbordered crow
bands (m.
...nutilk M ~ ~ Y I O ~ ~ I E S O
b D a d mlm~tioridifferent Chan that abme
.......................... B
2 a Dorsal surface d head aad fht one or two
doreale whib to er~a~n-colorsd with indis-
tiact dark shadM along the frontal-p
I
frtrnhal autuaetl; dwum uai~01ordmk olive
brown; ma postocular IbnMilb &&e,pa ..
Head d m dwsd coloration Wemnt
ikarl abow; u e d y two p a M a r s .,..,,--9
Dareurn and venter uniformly dwk brown
to bkek; wmpicrsous bhi&t yellow or whit-
ish head band pre~ent(cover8 parietala
grid the fimt 2-7 dmal mwsl
..*.*......,.......................
Ikslpti*
..,
m f a
...................
b Venter paler in color than d6r~itlg;pale
head band present or not ........................
4
4 a Dark vertebra1 stripe present ..................4
No dark vertebral strip present ...........B .&
., ............,....,..-...
I
b
6P Fwer than SO ~ub.cgudals
...........................
FloamtitIa v e m i f i m b
More than 80 subraudals
F'eww thm 165 ventrale; d e neck s t
8 ........................ 1
indistinct and P d ~ m i n a n t on
g the S C ~ B
bebind the parieals, pde lateral rtriper
with Imer dark hrder TanttUGF k m p h
More than 154 ventrals, or 8 fewer than
156 venbda, pale neck pot distinct and
prdt)*tl on the p a r i d s ; pale la-
teral atripe, i{present, wifiout lower d~
harder. .C
.L. . . . . . . . . . . . . . 7 ........................
...
T a Venixals 155-174 (mais*), 158-177 (L-
Ifemale& dmum of bead usually dark
brown or black with a &tinct small pde
I
aj& on each parietal IfanMla armlaosbrr
b Venttala 159-149 (males), 143-166 (fa-
malesh s u h u d d a W-83 (rnal~d,68-76
(faalw); d9mm af head uauall~pale
brown with lwge indhtinct pale spots on
parietals .....................
,Tanbflb*uj%s&Ul
.
I
a aorfium without pale longitudinal sfripea
...................................................................B
b Dotsum with pale longihdmd stripes
,..,....,..........,~,.,,.,,..,...~............ 1 8 .
B a Snout with pale ca1oreth-cpale wtoW
spat p w e n t .................
TmtiLh a U h l a
b Snout oolw s h i h ta reat of dorsal bead
d a c e , or d y slightly paler; without pale
pa@.Qcylar@pot .......................................
10
XQa Mare than 62 anbudals TanftEL m h
F%g641.Tantilla s~pmincttz(Gomadre, (Limdn, Rg.644.lhntitaa v e - f i n i s (Ouanac&9teI
Cmta Riea), PhoW. R. W. Van Deve~der Wta Rica). Photo: R.W. Van Devender
267
 b Fewer than 52 snbcaudaler ......................
11 ma& as far as taiI .............%nH& terEta
lla Mare than 155 i6entrala ..Tw~tbIkb a l d b Vertebral stripe reduced to a ssriea dpale
spots or ahnost eomphtely absent; pde
b Fewer khan 155 ventrds .........................
18 laterd stripes fade on posterior half of
lap Mom than 43 mbcaudals; XAQ-I54v a t r a h body ...........................
T&& rruhml
.........................*.............
M l k tayme lsa FaL IongitudinaI atripe only hdislitinct;
b Fewer than 48 aubcaudals; 11T-147 ven- fewer than 6B subcaudal8 ...........................
twls .....................,,...
mtzik ~ e i s k r ~ a ..,.,.,..,.,,, lbntilb cunSculloior C h i part1
1Sa FQW~P than 30 subcaudale ........... b Pale longitudinal stripe distiaet; mom
...................,. TmUZZa breaduauda than 68 subcaudale ...................
.,.*. .....20
b More than 30 submud.& ................... .,.. 14 Ma Qround color ~o dark that dark border of
pale lo@tudinal stripe nat distin
No pale v d b r a l s t i p e&""t
14.~1
..............Tantl&a can r&hr (inpart)
..
e...,.
able; Bmt dorsal row Iparavmtrals} ee of
dads pigmeahtion, at least an anterior
9""
b Pale vertebml atripe present ..............,..
16 half of body ................
T d t b t&eeFdato
IIa Edges d ventral malea darkly p e e n e d Pale longitudinaI stripe di~tiictlybordered
b
ar with heavy dark spotking ...................1& by a dark line; upper half of first dorsal
b Edges d ventral seala unpigmented or row {paraventrdsl darkly pigmented .,.,81
only wit&bolatad rrpots ...............,..... ..... 10 S1a
162-171 ventrdq pale vertebral stripe
sesrtrioted to vartebrd row; pale hteral
16a Mom Ohgn 159 ventrals; more than 65 sub d p e s (yellow in life1 very distinct over
caudab ........... ., ..... T m t t t k ~ t i a u f ~ t a
entire length of body; pale neck band wm-
b Fewer than 167 ventral@;fewer than 65 pltlste or intempted dorsally ......................
subcaudals ....................
B
..... ......... 17 ....................
....... Tgntilk incpenee
17a M e neck band reduced Co two neck spots b 143-161 ventrals; pale vertebral sMpa
................................
,.., bbilkEJsrrt covers verhbral row and adjatent third of
b Pd6 neck bagd c ~ m p l & @ ...........,......
,.....18 paravertebral rows; pale lateral stripes
less $istin& pde neck band complete .,.542
l8a Pale vertabral stripes continuous at least ma Pale verhbral stripe bsgins three 8eal~s
on anterior h d f o f b o d ~pde lateral ~ t r i p a behind the parietal5 and is not namwer at
the b e g i d g than posteriorly;pale l a w
&tripebegins three scales bddnd the 6.d!
supralabial an cove the adjacent two
WB t
of the Sr and 4r;% dmal rows ..........
..................
...,.., ~~hrltaanirrfa
b Pale vertebral ertripe be- e e scalm
behirid thr parietals and is narrower at the
beginning than posteriorly; pale lateral
&@p begin^ five or six scales behind the
6 supralabial andp~"vmg l y the aolja.
cent Wves of the 3 and 4 doreal m a
....*..........*..Tercdtlb tasntrrb
a.....,.....

a. T reticulate
M e r Reading
H. 8MlTH 1942a, & COLE1967, WILSON&
MEYER 1971, Wnsoa L V w 1878, VAN
mrnm & C w 1977, WWON & MENA 1980,
Wmom 1982a,b, 1986a-e,1!386,1987a-c,
1$8EJ&,
1SBOa-c, 1992, CAMPBELL& QMITH 19Q7,
CAMPBELL1998ki,H. SMITRet al, IQB8,W ~ O N
lW, WILSON & McCBANrrc 1999,D n a ~et al.
z m , S~VAOE:
2002.

Fig.648. Domal coloration in 'll.cntilla.

Fig. 847. Ykntii2a mbm (Tuxkla Guuma,
Chlapw, Mexiool. Phota: A. R a m h z V.
Fig.648. Tantilla breuicauda (ElSalvador).
Photo: T.Leendere
Abb, 64%.TrrnMIita fistani (Germ El Tom,
Atlbntico N o h , Nicaragua).
$.648. %ntidlib Iintoni (Las Cuevas,
Be me]. Phota:P.Word
Three very mall, secretive colubrid spe-
cies, which are very similar to the Rzntilla
species, are included in the genus
TQntillita. They usually stay on the
ground, under rotten logs or in soft plant
material, In nodeastern Nicmiragua Iin
the vicinity of Pueblo Wiao), I startled a
specimen of Tttntilkita lintoni in the wet
leaves on the forest floor during the day
(KC)HLER 1999d3. These small snakes are
probably nocturnal, feed on small inverte-
brates (termites?) and are oviparous (LEE
1946,CAMPBELL1998a).
 There are approximately 30 species of gar-
ter snakes distributed predominantly
across North America, with only four spa-
ciea occurring in Central America. The
nominate form T, surnichrasti eerebrosus,
W B h m& (€2- 1W61,J, &ad. Nat. described by H,SMITH (1942b)on the bmis
Sd. $hiladelpMa I31 C 1.44; type M t g r : of three specimens from "Esquintla,
M&, TL t;s 175 mm, YgcraW Guatemala",cannot be aasigned to any of
YU%, am lmd ts W209 91 ele~fatim
the known Central American species. Since
I l s d i t a i%ntmi@- W),FEae. Sm1, no further specimens have been found, this
w.Waah, 85: 61; type I d W : Plerdras is currently regarded as a nomen dubium
W ~ E l ~ b e a ~ . m t o 2 0 0 x u n *
v i m , l&3&0, to FiprthmXJi.cBt-m1 8eB (name of dubious merit) (ROSSMAN1991).
lwd Q 400 m nsalevstiahIn Cen,tralh*
Garter snakes are semiaquatic, harmlees
snakes, that will expel a vile-smelling
secretion from the cloaca as their first line
Key to Tantillita of defeme hefore they will bite. They live in
1 a 116-126 ventrals; 28-84 aubcaudals: pale ponds, lakes, swamps, streams, and rivers
neck spot present Trrntillita brew kaima and feed on tadpoles, frogs, fish,and earth-
b 103-115 ventrala; 32-66 subcaudala; with- woms (RQSSMAN et al, 1996).They are pre-
out pale neck spot ................................. 2
dominantly diurnal, but are wcaeionally
2 a 32-44 subcaudals; pale vertebral stripe also seen at night (CAMPBELL 1998a1, All
present; pde arem present on dorsal sw-
fa= of head ..................
Titntillita muEa species are livebearers, producing litters of
b 43-68 subcaudale; no pale vertebral etripe; 6-22 young (WRIGHT & WRIGHT lN7,
d o r d surfaue o f head unicolor brown R~SBMAN et al. 1996).
...........................,.,.. TantSlEita lintanb
~ h a r n m ~ ~I-MCOTT l r ! 11(Z(OJ)
Further Reading h Awl. Nat. Sci, Philadelphh 25k 333;
WILSON 1988b, d type locality: Rinconada, Coahuila, MeHim.
9% ;to 1146 nun.Utah and Colotado, USA, to
Guatemala, aea leva1 to 2700 rn elevation. Jn
Central America, only the subspeeim
Z!kamnqphds 6,d.iaris ISAH mmm.
Th;amwphis Bluw (WURT l W ) , Mia9.
Sci. M ~ L Rept.:
, 777; type locality: Alta
Vempae, Guaknda. TL to 6B2 i2CenW
Chiapas, h h h , 00 s o u t h w e s ~Hondutge
and aa-k El Balvadar, 14004981iO m eleve
kim*
!Fluamno his mwcktmus ( B m & @ ~ & a g ~
llMBl, &k. No& h e r . hR.:88: tgpe k
ealite:ECedRiPer;PLr TLto1090mm.
Kmem and California, USA to nortbern
BBca, asta Iwel b 1640 m el8m.f;ion.In
CanW A r i m k h subqdea ~ m n o p h b
m. b a f t i DWNN 1Wb CNiemgm and north-
ern Rita) snd Zkzrnrac hie m. pme-
mu&& I3ocmm 1892 (p,pni%rn in th.
YucatBn hninmla, Uuatemala, and
Abb. 660. Thrarnnophia cyrtopsis (New Mexico, Hondwul mu.
USA). Photo: k NGllert
Key fa ThmnophZs
1 a P e l o n ~d d stripe on the side covers
d
B and 4'Ef: doreal m9 at laaat in Wr-
ior kd area; without dark epate an the Two of the three aquatic gpeciee belonging
f
suprala i&; no pale ne& ~pote...............
...............,.......... to the genus Tretanorhinu~occur in
~ 1 1 0 p ~ p r a r i m t s sCentral America. On the Bay Ielmds
b fale lon ' udinal a 'pe on d if p r m t , (Honduras), there are specimens of Z
8
w e r e 2Ri and 3 or just 3 doreal row, taigmlute~cswith gray-brawn underbsllim
at least in anterior body area; d d x spota
on the supralsbials present;with pale neck as well as some that have brilliant orange
...... ..
rpota ................................. B coloration on their ventral shields. These
colubrids are rarely aggrerssire and live in
d
$4 a Pale laqgitudin stripe on dde, if recent,
L
coves only 3 c l o d mw, at a t In pondB, lakes, and swampa, a@well as
anterior body atea; aterior mpralabida muddy 8tream rund river@with alow flow-
disti~~ctly P"
darker co ored (similar to tem- ing water; only rarely w i l l khay leave their
porals) than m t e d o ~eupralabids ............. aquatic homes,At dusk, they kmme active
......................,mc~mpMsmol.ctanus and can be observed sating small fishes
b Pale 1 098 d m' a l a ri on side, if present, and tadpoles. They will not, however, eat
msw 2 am^ 3 $mal m,i t l e a t i n
anterior body ma; dl suprdabiale simi- the freshwater shrimp, which are often
larly colorsd .........................................
9 abundant in their aquatic habitat (Wmm
S a No dark dorsal ~ m spat k pres9lt, pale ver- & HAHN 1973). All Ttetamrhitaue speciea
tebral etripe h m c t or abmnt, at least reproduce ovipamu1y, pmducing dutche~

.............................. ~ a m w p h &law
h -
in posterior M y arw t o n p completely of 6-8 eggs (CAMPBELL
blaclr; 68-80 eubcaudrlls ..........................
1998a),

htanorhinrm seigdutew COPE18Ble, Froc.'

b One broad dark dord neck spot present; ,had. Nat, Su. Philadelphia 11: 298;
pale vmkbral stripe@distinct as far as tail Mi@: Ban Juan del N o h ,
ti^ tangus red with black tips; 76-101 8~rrmt actually 'hpinwalf, Panama," accord-
subcaudda ..........Tham~zophts -pat* 'bgto PETBRB& OREJAB-- (199O)I.TL
a 765 am.Southem Vmamm, Mexico,to
?mama, sea level to 750 m elmation.
Further Reading According b Wrrslo~6& MEYER (1886)the
SHREWgt GANS 1968, ROSI~MAN 1970, 1971, division into aub~peciespromulgated by
W ~ 1980,1982,
B R o s s w et d.1948 VU (1969a)II imuBcientlg supported, and
I prefer not to mcdgdze any subspedea.
W m r h i n u e moequardi I3ooom.s 1891, Le
NaeUraG5te (21 101: 13% type locality:
PanamP [= Panama C1 according to Drnrrv
Panama.
8
19591. TL to 730 rim. enttvll and eaalteF

Key to lh#wurRinsts
1 n D a r s ~ mwith three longitudinal stripes;
more thm 166 ventral8 ............................
.................... lktarnorltlnug m o c q u ~
b Domum with two longitudinal mwrc of dark
~pois;fewer than 156 ventral8 ................
...................Trefanorhinm n&plst%m
Fig. 666.Betanorhinus nigroluteua (Isla de Fig. 857. Tkimetapon simile (Monteverde,
Guanaja, Honduras). Costa Rica). Photo: M.& P.Fbgden

TZimtqm b a r k & Durn 19iWa, O ~ G Fap.

Hmtm goi;. Nat. B E t . 6: I56 trpe l d t
~ e d r o l a , g dcanalk*-mto2&
~blaa.kwb-

ma.
!Hat.
lkh d qPhilade
Sci. m p l T h COnC
4U:202; Roc
1% kyp W&
M.
faaa(S*Cm~TLtclmmm*Ea&
WeadPadarna
niiimbofron dm& Dmm UXk, W.Pap.
BPatrm h M& Bi& di: 551; hype lo*%&
I & ~ Y€bets. I Rim,
~ TL to 166 mm.
-M G o k Rim.
Fig. 666, Ttu3tanarhinusnigmluteus (Isla de
Guanajija, Honduras). m~~ mwH I W h , Proc. Aurrf.
NaL 8& FhSZttdeW '@: ll7: @pe hxdm
near W q 4 Pamma.TL to 2%
mm. CetlW- b t r t Riea md w&%m
FanaW.
!l%h&rnn vimwi D m lHFad f3oda

The species of the genus Trirnetopon are

mall, gracile colubrida, about which little
is known. Certain sllecies which had ear-
lier been included in'this genus (e,g.,hann-
.
*
Trimetopwr slevini
TrimetoponbanSouri
rlfmetopon $rn&
Trimtopon pliolepis
TR'metqoon simile
steini, pilonaorurn and posadasi; see J.
PETERB & OREJAS-MIRANDA1970) have Trimetopon viqmzi
been placed in Rhadinaea (MYERS197411.
The species of TErnetopon are known to
feed on salamanders and lizards (SAVAGE
2002). At least Trimetopon sdevini is
oviparous and produces clutches of 1-2
eggs (DUNN1940a).
Key ta Tn'mtopon
1 a 15 doreal rowa at midbody d! ..
......................
b 17 dorsd m e at midbody ........................4
..............
I
2 a kareparate prefrontds present
............................,,,
h'mstopon barbousi
b Prehntala fused into a single s s e l d ......8

II
3 a More than 136 ventrals; one postocular;
dorsum with indistinct dark vertebral
stripe; pale nuchrtl collar, if present, not
continvoug with throat color .....................
...................................TrimekOpOng m i t e
b Fewer than 130 ventrda; two postocdars;
dorsum uniform; pale nuchal mliar pre-
sent and continua6s with throat color ,I. , .
................................Trimopon simile
. Fig 659, Dimorphadora biscutatue (mar
Mixteauilla, Oaxaca, Mexico).
4 a Prehntab fwd into a eingle Bhield; a
pale nuchal collar present, eometimes bro-
ken by a vertebral dark tripe and contin-
uous with throat color ........................ ..
...........................
, .. ~ m d o p o m p l i o ~ p l s
b Two separate prefrontah present; no pale
nuchal mllar, although a pair of pale neck
spotscanbepresent .......................... ,..,.,6 . - -

5 a More than 40 subcaudale; a pair of ale 22immh*cton

neck spots; body with a vertebral Bark -
e w e and 2 or 3 lateral dark stripes ........
Lyre snmes me n~cturnar,semi-arboreal
...............................'Sri~h9p0n~ b ~ d n colub,.&
i that distributedall acmes the
b Fewer than 40 ~ubmudds;without pale &-y regions of the sout;hwestern U.S.,as
neck Wts;bDdy With a dark well as through Mexico and Central
stripe and a single Iahral dark stripes ...
..............................Trimtopon viqueei America (excluding the Yucatan Pen-
insula). They feed on lizards, small mam-
mals and bilds, with spiny-tailed iguanas
Further Readirrg (Ctenosaura)being their preferred prey in
DVNNl930a, 1940a, TAYLOR1961,1954
some areas. Swallowing a large spiny-
tailed iguana, however, can prove life
threatening for these snakes,as de~cribed
by R~M~REZ-BALTTISTA & URBE(1992).They
obserwed a 175 cm long lyre snake (weight
1183 g) in Mexico as it seized a large male
Ctenosaura pecbiwta, strangled it, and
wailo owed the lizard headfirst. After wrig-
gling around restlesely in the eame spot for
about 20 minutes, the snake wddenly
stretched, flipped onto its back and died.
Necropsy revealed that the gullet and sto-
mach of the snake had been dit open by
the sharp tail spines of the iguana.
7kimorplrodon biseutatus reproduces ovi-
paromly during the dry season; clutchea of
7-21 eggs will hatch after an incubation
Fig. 658. Wmrphodon bimutatu~(dorsal view ~riod of 79-106 days (FITCH 1970, KoHLER
of head). Drawing.: M. Vesely 1997).
 -
lWmrph&n biscrrWua (Dvns$RIL, BIBROW
& I~uM&& 18541, Erp. Wn.7: 1 1 4 type
bdty: Mexico. TL to 1750 mm.
S o a t h w e %USA
~ to Costa Ifica, eea level to
1500 m eletvatia AcCordin@t to Wum &
MIZYE.R(1985)the division into subepedw at
' least in Central Amaim ie ina&dently sup
p a w
qeeiw,
,mci I prefer not to mgnize any
Further Reading
H. ~ M W H 1941b, GEWACH 1971, Sc0l"C &
Mcl31~~bfm
1984
I
Abb. 660. !&imorphadon biscutatus (Apoyeque,
Managua, Nicaragua), Photo: A. 2.Quintma
Fig. 881. Tripanurgos c o m p ~ s ~ u s .
Photo: R.D.Bartlatt
e
, genus Trtpanurgu, .,
monotypic.
Pipanurgos compressus is a nocturnal
colubrid that feeds predominantly on frogs
and lizards. They live primarily in the
branches of treea (GREENE1997).This ape-
ciea is seldom seen, therefore hardly any-
thing is known abut their ecology Shortly
aRer nightfall, E.LEHR(pers. c o r n . 2000)
encountered an adult E compressus, crose-
ing a path in the lowland rain forest (230
m elevation) in central eastern Peru. The
animal seemed entirely harmless and
made no attempt to bite. RILEY& i
WINCH
(Ig06)reported on egga of T. compressus,
which they had discovered in a nest of leaf-
cutter ants on Trinidad, Because two
distinct size claeses were present m a n g
the eggs,it was suggegted that two OP more
females oviposited in the same nest.
Wpr;mugas m p r e e e u a ~DAUDM 18081, Hist.
Nak. RepL 0: 347;type lwaliw Surinam.
to 750 mm. Southeastern Costa Rica tn Pe
3
and f i l i h , aear Iwel to 1000 m elvation ('
South America t,1600 m almti~s].
Rmy & WINeH 1985, Z A H E ~lggg
-
Colubridae

Fig.662. Fmp iipsas sartorid (Campec , Mexico). Photo: J. C,Lett

~znpidodiplolzafmciatus (GOllrrm~Imal,
Cat. Snake?- Brit. Mus.: 181; type fix&*
Me9ico. TL t o 717 mm. Diejunet k F m
The ~peciesof the game Tropichdipirraa Gmnwre and Vcr$mus, Eidexim, aa far ae;
are clo~elyrelated to the Sibon epecies and C h i a m bkim, and in the north rd the
were earlier included in the latter genus by Yuetlt8n Eedmida, sea level bBM10 m dma- '
tion in rain fwd. L Central Am&, or&
Komo~1985. WALLACB (1996)showed that tlw ~ ~ ~ ~ m o c r m z e .
lhpidadipsae was distinguishable as a
Tivpiddipaae @a&eri B o w m a 1894~1,
separate genus from Sibon, Like the epe- eat, §nabs Brit. Mue. 2 typ M t p :
c i e ~of kbe genera Dipsas and Sibon, the clnE4C& aZ to 862 ~ P O!mea, J
YYupidadipsas apeciea feed primarily on MeTioo, &o toatem El Safwader d ~&h-
sn& and dugs ( D m m et al. 1986,LeE western Hrnduma, la3000 m elevation in
1996). All 2YopWipsas species are noc- pine+& foresFt md 3ad forest.?kro anibepe-
dw are mnqgnkd %pWCrs f fischd
tmd, mostly terrestrial. They reproduce (mnw dhtrbt40n as k h Bpa7iea ax@e t, fm
ovipmu~Iy,laying 3-6 egg8 per clutch rut. ~ i q a ,
hbrn.1.) an4 %&a,
t.mwmle the end ofthe dry season or at fithe F Mdden ST~ART 19420 ( A h Vsrapat,
beginning of the rsiny magon (KomoN fihateasalaln
1987,G ~ ~ E1998a). W I 7 h p i d a d W mr&iors:COPE
i 1w, Prw.A d ,
'kt,&If. Pktibidphi8 35: lOO; type l d *
M h d o ~%raw%%.)
~ M d w , 'I%t.4 &7 mm.
Wum Mexioa, ice N i w aea l e d
tie appmxhteIy P W m elentation in drg
fmst and wet fornet, k Cen~alAmria%
tidy th& TMmindb SUb~peCi$81Om3Um. I
 Fig.666. ,. ~ ~ i p s afasciatwrts
sr (Calakmul,
Camphe, Mexico), Phata: RCedcfioV

Key to TnopidodZpsas
1 a Doraal scales completely smooth ...............
........................Tmpidodipsaa fh9chtrr8
b Dorsal wales weakly keeled .....................
2
2 a Dorsum with 12-30black rings,that reach
as far as venter; epaces between the dark
ban& cream ccrlamd or orange red in life;
white in preeervatire (male tips may be
black) ................Tmpidwlip#a8 sartortl
b Borsum with 20-64 dark brown bands that
do not reach to venter; spaces between
dark bands pale brown to gray-brown........
.........................~ p ~ fiechert
~ p

Fig. 664.Wpidodipsm fischeri (GuatemJa).

Photo:L. Melendez
The genus name UrotIteca wm revived at
the end of the 1980'a for the spsciw of tihe
IZhlzdi~aeadai%&t~&& lpaup ~?@?wu MYERS
1974 (SAW & CROTHEI~ 1989). Char-
acteriatically, them colubrids have a con-
spimmdy long tail capable of auhtomy,
unique hemipene morphology, anand a white
line along the first dmaal rows that is
generally quite distinct.The tail will break
off easily if it is seized. In fact, most
specimens of tbi8 genue that have been 7
f w d b v e h-pleb ms, w ~ & sup- Fig. MB.Uratlseaarnentheri(Quebrada
po* the --ptim that the bredable 540 m elevation, Col6ns Hond-1,
Photo: J. hcZ McCranie
taiI ie a ~ m d v a~tmtegy
l to protect against
predators UVWEBS1974, SAVAGE & CROTHER
1489).Moat ofthe spdes ofthis genw are
thought to be diurnal, although Urotheca
ftdkuisep~has been Been out in the open at
ni&t (MYEM 19743. AJ1 Umtheca eusciea
are ground-dwellersthat feed on ad&a-
dew, frogs9 and bar&. They mproduce
wipmusly (MYERS39743.

5 1 n r t h jwv' mm 10r%& Pt06 BIPUB~.I

~he~oe.48:X$P.l*~slyna
h 489 mm, Gm€a 11 to nwthw-~
~*~~~fOabQUt6;00md~
rain fk&.

U W h myed & e M 119@*

Copedalm893;mhaIitlp:1te~rhdrs
~Bli6tl~,aptmr~t,$.$krvxSbyroadaf
rn ~ f w $ 8 ~otfm b- a h
C wI n w - (Rues 21, CantmJ
l=Em&dedan, 42wdin~t"ade 'lldmmna, SBin
~ O ~ ~ k ~ T L . & t ~ m m ~ P a & v e
%%1~ttJfchra&mde-hd
mxd d a r n ~~ma,1BOa-m m dm@-
tian in cloud fbYJe&,
I

am lCom 28?1P), 3. Am& N&

L?@ad%a (2)s: 146; Bp w*
~ o l n s r s u 8 ~ I l m 6 n , - ~ k r
MYWm 3974, aada IZ b "la6 mma
b W - h t a w c # ~ - ~ ~ - l
lwQ~~rnanel~rrninr9ix
Fig.,867. Umthsca dei5piam.a (north dopa of Cerro Panda, 1620 m elevation, Bocas del Taro,
Fanama), Phata: C. W,Myem

Key to Umtheca 4 o Without whits ocelli at aide of back of

1 a Dorsal aurface of head pale yellowish head; venter white in life; ventral8 132-
brown, in sharp contraat to the darker 137 (males) and 122-140Ifemalee) ............
body coloration, .................. ..............................U m t k a &cipknn
......................a
b Coloration of doma1 surface ofhead aimilar b White mllus formed on each side of back
to tbat of body ......................... ,. .............4
of head; venter reddieh color in life; ven-
trals 135-164 (males) and 147-176 (fe-
2 a Pale head coloration reaches 3-4 domal maled .........,...........
Umtkeea guenthd
scalas past the parietals; li shields inten-
eively dark .potted ... urnti800fiLIuiceps
b Pale head coloration reachea 1-2 dorsal Further Reading
ecalea pa& the arietals; lip shields, if at MYERS1974, SAVAGE .%i CROTHER 1989, SAVAGE
gt
all, only with ingmtinct pots ...................3 WAS 1989, MGCRANLE et 4.2001
8 a One narrow, dark neck hand present
...
..................... ..............Urvthetw w e m i
b Without dark neck band ..............................
.................................
Umthecapachyum
Fig, 668. Xenodon mbdocepkal~(upperBladen Ever, Ek Xux, Toledo Di~trict,Belize).
Photo: PaStafford

X e d o n mbrdoeephdus (WED 18241, Isis von

The species of the genua Xenodon are
strong, heavily-built rear-fanged anakea
that are dmilar in their external appear-
Bken 6: 668; type lccdity: Brazil. TL ta 790
mm. G u e m a and Veracruz, Mexico, tp
Ecuador, Bolivia and Brazil, sea level b 1300
rn elevationin dry fomt, wet fimt, and rain
forest, kocol.ding to W ~ % MWER
N (19861,
the division inb anhpdes is in~dBEiatly

1
ance to the dangerous fer-de-lance, suppmbd, and I prefer not to reoo@e any
Bothrops aaper, and are known in Spanish aubapeeia~.
as Ufalso terciopelo". If tfie falae fer-de-
lance feels threatened, it will flatten its
body dor~oventrallyand gape with an open Further Reading
Wnsolu & MEYER1986
mouth. &nodon rabdocephalua feeds pre-
do-tly on fmga and toads (LEE1996).
Toads in particular are known to innate
themselves when threatened in order to
make it more difficult for predators to
swallow them. Xenodon mbdocephalus is
able to puncture its prey with the enlarged
rear teeth of the upper jaw, so that they
deflah and me e a ~ i eto
r swallow. This spe-
c i a reproduces by laying eggs in clutches
of 9-10 during the rainy Heason (CAMPBEU
1498a). Freshly hatched young have been
seen during the months from June until
November in the northern portion of
Central America ( C ~ B E L1998a).
L
In addition ts f i e true oord makes (gem
Mierrtrm), which am dietrlbuted in
America, the family ElapIdae a l ~ includes
b Mi@mrwspesim, extreme caution L ad-
the cobras, mambas, and haib in Asia and
AAica, the Awhalim venommuis snake4
and the sea makes,At various inthe
past, the latter have heen split off h m tba
Elapidae fnta a sepalrate f-ly ("Hydro-
phiidaev), however, subjected to zncra
m n t inve~tigaUarus,t b division ha9 not
been retained CQrt~~imi199?), Char-
a d d t i c of all membre d the family
Elapidae is a pair of deep13 grooved faog;e,
located at tlw &ant of the lzpperjavp, which
are relatively immobile.
Fig. 669. M~~ diaetemra (VwPlateau, Bslize).
H view of the highly potent venom of the
vised in d d n g with them. Those rafe
instances in which a Mkruws bite iEl
eustgided tend ka be fa4al. The Temm is
predominantly newtoxic,acting ae a pm-
alHc agent. h Central America, there aner
36 rspeeies sf tha genw Micruraa, dl af
which have v e q o(1n~piouowand ~trik-
in@ycaloM coloration, Cord make@ W
pred~minanwon &BT &a&=, bnt:
a h eat eels, cae@ms, and lie&. !Fhe
species of the geana Mkrtlrus mproduce
ovl'pmusly, and produce clutchw d 1-13
egge, which hat& a h r 70-80 day& IIhom
1996). The b y ta Mimurua below is based
largely an mm (l$!mE.
Photo: P. Stafford
281
MKfym h i flaamm UB@kt,Field MUB, affemrn@i2irgaw tJm 1&1581,Bev. M8g. Zd*
Nak, H4k. kl.Bar. sk mi;t$pe *
l Rf?J (%3 1U &%4iwWw M d c ~ 1 T L t o785
mq 7 miles g$(Y98 &ma* A t l t w h bjrtr, m3n. CmW V m m , Mtaxieo, to Alta
Niwagua. TL ta 1165 nzm, JhsCemr Vere m,Guelmn& 101).1700melmtdtmio
Haaaw to @£m* ,pwtm£h eea leml to ~?&rmdW$mt8aestfacemPd~%
1.600malevatienmrarnSormk tho w w a mem~z?@ ~ a, lJmaI5paoZB
Sf+aaumlBif5c mmm.
Mtcnma a m d h lJAN 1872), h JAM 8t
So- IWtL- uphid., b.a: pl. A 43. M a r h' m@ @%EB~18631, Mpidab
2; $ype localZ$: I3hndm*IT, ta l e a raaL Atad. w m Trlin xm: mi Ipe I d *
EWam Panama to southw- Earadar, saatcr TaYrAh €9 I'tmtU *&!I de ail%€%
~IdClOOamel~v~inrain~t,I smdigg
n fa3 PETERS Q urn@--
c~atnrl Amwl* tbe fiabsped~W&rrsrusa, 19703, i2Wxmda, 'I'L to 710 mna, Belike sad
JiCkrdSmmn I916 -. ewbm Gw&mala, eeri level to @
.tiOninEaIn~t;andglne~t'
m sba-
&I
Mb~"scrudba@& BmE 16)83,h e r . Ildw.
Hi&. &SSP: 8; load& 2btigoh-a
Qwg~lmhl,mt ofPu& Angel, 6-,
-00, mto 736 %ide ofthe
~u~d-~1Vlanieo,~lwdto
m dmQmin b &re&
d s m &mami i?kmmT %G SMmi 1943, wst aireat.
Field Mw.Nat. Hk9t. Rrbl. M.em. W:mh

brawni (Bi~triWtmaame a~ 6pmim wmpt

for khe Antigua Basin Ijhtrtbmala) and
AKbimm h i w ~ l c k 9 I@%?&Antigua
BwiT&wm>-

1 m 1W Wka V a p
c2wt"amalal*it&le:TW't48
1wr (@&m0 -
d. 80pprf (WERNER 1008b)(the wuthern por-
tion of the f i c a t h Penin~ulaloccur.
C01ornbia"l.TL to lo00 mm. Ea~ternPanama
and northwestern South America, sea level to
860 m elmtion in rain foreat. h Central
America, the nominate auhpeGias wmug.
Mkrurus multi~chttssr(JAN 18681, Rw.
Mag.2001. (2) 10: 621; type l o c e k Central
America. TL to 1130 mm. Nicaragua to1
CentraI Panama, 5ea level ta 1200 m eleva-
tion in rain foreet. The sub~peeieeMicrurug
m. multifusrciatus (Central Panama) md,.
Micrurws m hertwigii (WERNER 1897)
(Nicaragua to northweatarn Panama) are
reco&ed.
Micmrwr nigrminctus ( G W 18541, Pmc.
h a d . Nat, Sd.Philadelphia & 228; type lo-
cality: Taboga Island, Bay of Panama.TL to
1150 mm. Southern Chiapas, Mexico, and'
northern Honduras to northwestern
Colombia (excluding the Yucath Penhula;
STAFFORD 2000),sea level to 1500 m elevation
in dry forest, wet forest, and rain forest. In
Centssl America eix subspecie8 are recog-.
Micrurus n. nigminctlcs (Pacific aide
of Nicaragua to northwestern Colarnbia),..
Micmrue n. Irabesparl R ~ Z E 1967 (Corn
Idanda, Nicaragua), Micrunrs n. coibemis
SCHMIDT 1936b (Coiba
Mlcrurua n. divarlcatm
(northern and central Honduras). Micruru
i, mos uitemi~sCmm 1933~icaribbe
aide of%icaragun to nmtbwastcm Fanma
and Micrum n. zuniknsis S ~ H 1932 M ~
&them Chiapaa, Me-, to El Salvad
OcCuT.
Miemrcse matranus (QOl"NTHER 18951, Bi
Centrali-Ilmericana R e p W and fSak
180; type locality: bla de b a t h , Ial
M a , Hondursa. TL to 681 mm.
haw,Hondwaa.
Micrurm etewarti BARBOUR & llnanRAt 1928
Bull. Antivenin In&. Amsr, 1: 100; type 1
cality: Nombre de Dios, Sierra de la Brrlj
4
Panama. TL t.a 888 mm. Central Psnam
500-1200m elevation in rain fore~t.
Micrurtds stuarti ROZE 1967, h e r . Mus,
Novit. 85431: 47; type locality: Finca La Pal
[L 1460 m elevation, 18 km N Coatepequel,
Ban Marcog, Guatenaala. TL to 74& mm,
Southweatern Guatemala, 800-1600 m eleva-
tion in wet fomt and cloud forest.
1
 Fig.670. Micncrus alleni (Costa Rical. Fig. 673. Micrurus browni ITuxtla Guti&rea,
Photo: R W.Van Devender Chiapas, Mexiool.

I Fig. 671. Micrurus clarki (Comarca de San

Blas, Panama). Photo: C. W. Myers
Fig. 674. Micrurus diaste
Chiapas, Mexico).

284
Fig. 672. Micrurus dicastema (near Quebrada
Grande, Coph, Honduras).
Fig. 675. M h r u s slegans (El Ocote region,
Chiapas, Mexico).
1
Fig, 676. M i c r u m hippocfepik (Liviqgkon, Fig. 679. Micrurtds n&mf~actu~
(Cramboa
Izabal, Guatemala), Rreat Resort, Panama), Phato: M.Lundbeq

Fig. 677. Micmrua nigminctus (Cem Fig.680. Mimurun rrral'antis (Isla de RoatAn,
Saahya, Atl$ntica No*, mcara@~). Hondura~). Photo: J. R.McCranie

Fig. 678.Mkrurus rnultifasciatue(CostaRich Fig. 681.Micrurwt stewarki (Panama).

Lirn6n Prov.). Photo: R. W.Van Devender Photo: M. A. Guerra
286
Kay to Miemmu laa Black;ee halic oap ~ompletelyor almost
1 a Black rings mangeql in triadil ...............a. 54 mp1ets4 ewsn the p u l w WuWg
b BI& rhga not amu@ in triads 4 ........... b the pmtmor tips ............
Mkmms CEarkt.
B M cephalic cap do- nd aver the
8 a b&mpalekdMandArskd* po~tariortip8 d ths parietat~ ................
I8
ody a &rag161black ring
................. .....................
. ,,., H 3 e m &gaCwraua l* b Usually F ew €?an10black body rinfp ........
...,..14
b pe%w-ee;n p+e h a band ( W M C ~ glay be more than B (~eldom8 or 9) bI&
mtempt9d ~nthe middle ofhead) and h t MyringB ......,.............*.*,. ..............1T
r e d ~ o n a ~ o r a t r i a d o f b l a a l r ~149 8 B i d k neok band w c h e ~8-13 dorsals
8 a &#id large137 md (Pig,684ak betypeen p d e
ptmiady ......
,.,~,.,.,,,..aaa.a.aaaaaaaa,,... I6
Izaadbanddik&drlngahiadaf b Blacls nerrlr band reaeba Eewer than 8 dor-
black ringa .............I h m m~w m J k r gals posteriorly .....................................
16
b Xead largely blaek (Rg.BWd; L h a l6a malee with mqm-mdtubenelm (Fig. 682k
p r r l e h a d b d andfiratdFiflgone pat females with fewer than 213 wtntsaEar
..
of blmk rings , , .,.,. ,..M&JTL~SOB elegana .........................
ML~UMB WprUaiaaarS
4 a Body wi& an& two w1m (bIa& and red BF ltr Malee without su a-mI tuberoles fe-
black and w b b ) ............................
......,.,,..6 mdes with more
.............
M t e m
Cm
PIP vemtrh
Bipprrcmpie tta pW1
.........
b h d y with h e ~ 1 0 Sbhck-md-ydow ~ 6 m
black-red-white) ...................... ........
...,.*. 10 1Ba Black neck baad reaehtts 2-9 dsrrals
6 P Wikh altepn-
Fin@ .. ..,..,.....,,.,..,.
longer and shorter black
..............................
b Ail blaok r;ing~of equal lengbh .................7
8 rteriorly; Fed body d e a with or without
ttle bkck pigment
.............
M b m w k~~
............................
(inpart3
b Black tlwk band reaches 4-5 doraaler
6 a Ventrala 178-188 (male~land 1BS-203 teriarly; tip of FBd bady r a ~ a 1wit& ~ ~
........
( f d ~ l ,,.*,. Mbmw mfunm Kelt .pot or 2 h m smlm d t h itmgdar&
&&Tibut& black pigment .*' ...............
b Ventral8 200-207 (arde~3)arrd 224-228
Zftrmalee) ..................
MSearrus s f a w d .................
Mhmms~ t m Ifn mp d
riqp; 8-18 black. 1% B h k nak b a d m a h the ~ &etala @&.
? a Mare than 40 black
tail ringg ...Mkmnrrr:bbttenra (ingart) 6a3)................
....,., ......... . . .
. . ' 18
b BIack neck band doer not reach tL
b &merally fbmr €ha8 blaok taul ria@; if
8, then fmer than 80 blftEk ring8 .. 8 parietal8 (Fig. d%, k) ..........................
10
S a KI.4 bwly rings st l a s t twice ae long m 1% Malm with supm=analtub& (Fig.W
..... ..................,..,...,..,.....~..... females with &wer than 40 aubcaudtlla ...
bla& r i q ~ ,
............MiF~rrrantgwEhtwr (in p&) .,..........MIfcrurorsR & F Y M ~ & U Ib prtrt)
b M& withoat s u p r a - 4 t u h r ~ l ~fe-,
b Red cn ~irhiitebad rhga lem than daub1e
.................... maim with more than QO aubcsmdala ....,....
*ledbhci~* "*I)
................
.&fLmm~ dfBgksmatin pad
@ a M y with black and white ringer; Hack 19a Red body S ~ B without S EOIU&IICUQW M&Gk
neck band mu& reaches to the b p ofthol ~
pmWds O?
t b two o~sals aaga. ;n. e darsal h p lees
i
.....................
............................. M k m l l l r ~m @ a h l t u e b tip8
tips, at most with very tmdl dots or
bramhh ahding .............................
Red body scab5 with mmpirn~wblack
W

b Body witb bla& and red rings; Ma& neclr ......................................................

24
band usually doas not m I the tige of tbe
etttlrs (Pig, 6$+&f); pale &opsal r5np at
P"
east two dam& long
.........................
&fbTW?T#
@
..............................
#W b m @ w 8
10a Head withoat pale brand& ...................
."..............Y i m s Wca&ema:(Inpart)
b H a d wkbh at least on12pale baad ...........11
lla Bl& eephalb cap mntinum as a mrrm
gtripe d q ths parietal suture /Fig. B84bl
.........,........*...-....... hi~mm*h i
,..m.

b H ~ a dmarkings ~ ~ those
thm i abwe :
*..................,.....................,....,...,.......1%
 V a k d r u d l $ ma% ;thw 10T W e s ~
ai3mme t$mn %Icf&i3wdmj *.'S1 ., .........
h Vamidt fmm &an [maled 4~ B
Wm2;<W& .....................
p,.*,,"B2
T

2la W~BB
with stfgra-anal t&sAm; Eedw

l&s *
b M a l vtft;bepC
~ a u m d $abPEfe~
15-ll$bf$i&$Qdgm& ,,.,..,...,
ha-
....................,,....., -.*.* M ~ M .hTD@&~
a. M,anma&
BMC I W C ~w at l e a 3
O T ~ ~ $Wad€O P bod3 ~ ~ rings ~tleaat
lo fimt r%rd uwalltllg Jwse
tbaa 19 1- melg. wnh &up*
4 t-88 ua.m~ln..a*uvrr
..,,,,,,,., Mi4mm-a nQmciWgq tin
a~n*-*.a+.

m)
irrruirrrvs

b Black rim& readhea 2-3 domds

paderfbrh d d ' U o l Ph@ xir@ mad-
Iylw &mi3 d c m M & l ~ h $d*
& mme thm B dmOrB$s 1% rnalbs
a.th~tltSU-X,FB&~ k U k d @ S ,..........~..
............. -~pocrsg%*parfS
me Nd d red
a*
9
@a nlth black tip, d, u bmwni
f*@~lm y &&ribtedI w$h
7

back id- ip,othm sat

*,,. .*. ,..*mv
"....... ....
aiwttma IPllph3r
**, .,..
b Pilt red b d y mmlee with mgdar black tip
br..m.*u.u.r.t.mrruqrrr *arrr*.q.t*tujry-..-rau-.urrr*s 24
Ma V@* mually Qewer&in 906 W~@B)
and &wertkm D 3 /&males)

L
5% .ec.,i,d,

b Wntrde rnban g08 We?arid mare

tawim3 Chale~l.,,.,,,.,., ,I,,+u.r..nv.nfM
ma XI-"t, ueway m e than lg hhck w g
rim & d
.
kh a p p - a a l imbepalw
nhw&drpr Ckmn4 -
1I
!gens h M. latjfescietus I. M. mrparfitus

B h k tip on red W e b &(bmttwbmtfrregn*

lru, large (me abut #4 of the %sd
wa188f; blaels n d band d e 6 6-8
C s a 71doma p W o s 1 y
.rt~*r+tu~rue.rt..a-n+r..+r*e**.r MSa-3
...........,.,,...,,.
-S
b%l&k on scalw mgJl
Cower about Y4 of the ssd gcgtak b l d
n$& baaid r&&es 46 tuaua;llp $1 d m &
pt%riwQ ,4.&7" a,,,.,., *.A. rn ................ (I, M,&&z k M. fl&nmim I. ns, *w#
 been mpcrrtod BQ fm have not been sup-
ported by any epedmem. If any speeimena
of Latkauda coiabrina have reached
Only one ofthe appmxha&ly 66 epgcies of Cents4 America in ths past, then them
sea d m o m s don&the Pacific ma& of ware likely isolated st-ray specimen^ hhat
Central America, the pelagic eei3 make have nat e&ablished self-~wtainingpopu-
U%bnais pZatu1~1).Their absence dong latiom.
the Caribbean Coast implia that this ape-
ides reached the Pacitic fewer than four The pelagic sea make rERelam's plattsrus)
million years agot after the Panama land has a striking appearance+with ib pUow-
bridge was formed and separated tbe two black colaration, It is- remarkably well
oceans. Another sea anah, the banded em adapted for a life at sea, with its Batt%ned
snake (Lathauda colubrinal, wae reported rudder-like tail, nostfila located on k b
in Niwagua by V w (19621 and, ae a upper Bide of the snout, and 4 t g h d g
result, ineluded in variaw ch~~ktigts for under the tongue.
the region W &mma 82 U R E J ~ M I ~ A M D A
1970, VLLA 13M, VILLA et d. 1988). PeEanais plrttunts ucnually is men floating
Hawever, the &at%nce of this snake is oan- an the euFfao8, whioh is why aome au-
s i d d &bly ~11likeIyWfa~rt199901, thors haye dwribed them as pelagic
particulm1g as the sigh- that have (Gmm 1W) . parkirmlarly like t o
They

Fig. 685. Pelamis platurus (Playa El Coco, Costa Ria). Photo: R.U.Bslrtlett
289
dwell. in ah waters with a lot of small
fish, &their p r M food. Tbey catch
their prey with quick sid~waysbites,
eometimea swimming rapidly back-
wards. This species baa highly effective
venom at ib dhpo~al,whicb it uses to
catch prey and to protect iteelf from pre-
dators. Bites from, these sea snakes
should Be taken very seriously, wen
though the few docuslented cwea have
h e n accompanied by relatively mild
sympt~toms,aucb w localized pain and
swelling (KR~PACH1975, S O L ~ ~ O
1995).

PeIagic sea m&ea will mah floating on Fig 686.fihmie platurn [ P l ~ E al

Cmp
Costa Elica). Photo: R.D.Bartlett;
the eurface of the water and d ~ bearo
their yomg in the watert Sea snake8
~ b atdhsquent ~ B T V(approximate-
B~B
ly &very 3-6 weelm), among other rea-
sons, to rid themselves of unwaxlted
extarnal pmaaites (algae, as well a8
barnacles and mussels).In order to h @ d
their old skin, the enakea will form
complex Imab with their bodies, and
then crawl out through them.The ahed
skin b usually left behind still tied iu
kmte IGREENE 1997). Apparently
PeZamis pleturus B a g few natural pm-
datm~,perhaps due to its warning color-
ation and Mehly toxic venom.Sea birds
have been oberved eakhing pela&c @a
snakes and thein dmppLng them again
&r a short time ( G m 1987).The
most deadly c b m s t m c e fax a &hrnit~
platurw is to be wmhed up mthe beach. Fig. 687, AEernis platrama ( P 1 ~ ; aLa Rw,
The animal would bave a very fiterd Nicaragua).
chance of baing abIe to retunz ta the
water under ita own power artd would
die &r a short t h e .
 awing down and farraard when the animal
is inthe procat~~ of &rib#.The venom of
Earrpt far the .s@eleies af the genus moat ripera disrupts both the Csrculatorg
Atmctasgk all venomau s&o that have SYBW, M well 88 timue and, ineome w e a
enl-, highly movable, t*m lldbw hg., south Amerioan ra#&lees&ea,
fangs bave been plamd in the fmily Crotulw duri&~us&r&feu81, also has
Viperidfie. About 160 of thee are &trowr i m t o d e Meeta. With the mep-
cham* by the p m m c e of pit organs tixln of the htlshma~~ters (genuer b h a 8 i d ,
and have been sombined in the ;subf&~ which reprodu~eovipaxon~ly,dl other
C r o W h ~ f 3(pibigem or laaceheads), tu N&pid pitvipers am l i v e M s .
which gZ1 Ceatml Amerioan vipem helong.
Pit organs me I.reat~aemitiPa,tube-like -ear-
depra~~iona fow1.d between the eye end Wla C W E L f r & LAMAR 1m, 1@2, ~~ 1992,
nosf& They aadd the s d c in~ tracking ~ Wrmim 1998,IS@& S r n ~ A oet d.k W
down warm-blooded prey, su& u mall
mmnmal~. AIl v i p r ~have a highly evolverd
venom delivery apparatus. Wh6n the
mouth ofthe emke i s dosed,$helong fangs
are protected in a fold csf skin and they

3
Fig.688. Bothrkchis nigrouiridis (Costa Rica). Photo: R.W.Van Devender
Key to Viperidae
1 a Tkailend ~ithrattle,,..,.,,~., ........CrofczI:~~~
b Tail end without rattle .............................. 2
2 g Underside of tail tip with 4 5 rows d nar.
row epiny scale% mg.690a) ....... LuaJiaais
b Underside of tail with 1-2raws of relative-
ly wide, flat malea (Fig.Wb,c) ...............8
8 a Snout covered with large symmetrical
shielda (Fig. 69la) .................
tlgkhtrodon
b Bnout covered with small irregular milea
(Fig,691b,e] .........................................4
4 a kI1 or most subcadtila paired ........,..-.
.............................................
Bo&rope / \mental
b All or most anhaudals single ...................5 #8k~8~
lint infmlsbial
5 a Eyelid with projecting, elongatedI pointed
ecalea ..................Bathrhhb (bipart) Fig. 689. Scalatien charahre an the head of a
viperid snake (Cemphicltotz g o d m i ) .
b Eyelid without projecting, elongated, Drawing by M.Veaely
pointed scales .........................................
8
8 e Urhedm green in life [blue in preama-
tiveh arbreal with prehensile tail ..,,.,,.,,,.,
.............................
Bothriwkia (in pa%)
b Ground color r e d - b m , gray, or bmwn;
terrestrial without prehellgile tail ...,...... 7
u Domal pattern mually made up of oppoaik
or alternatiy dark r ~ t e b r e Ispots
rrnd one pa e verte ral stripe .........
...................................,.......
Portkidftrm
b Doma1 pattern diffment &om that above
( o k n a dark zigzag pattern present) -

8 a 1-7 scalea between the ~iupraoculars;body

scales keeled,but mt conspicuously rough
19-29 dorsal rows at midbody ..................
...........................,............
Gewaphtdkrn
b 9-12males between the a u p m d m (Fig.
691b); scales very mugb; 23-31doma1 Fig.6WW3, halation of distal portion of tail
rows at mdbody ....................
hpoide. (ventral view) id some pitvipere.

Fig, 691,Sedation of
dorsal surface of head.
Fig. 692. Agkistrochn hilineatus (Taxieco, Santa Roea, Guatemala).

tion iP dry f5mst and savanaa In C M

h e r h a , the 11ubagw4es&Ai&rwlm h biiin.
(daap t b M e mu& of 0f to El
The four species of the genus Agkistrodon Balvadml* & M a b, halwa-kd~
are restricted in their distribution to North M e ymt from
and CentraI America. A g k i s t d o n bilinea- and & ~ M o h II.
tus is a mid-size powerful pitviper that pre- #-U&?dlsg GLmD 1m mcat8n hnins&
fers to live in dry areas, often near water.
At nightfall,it will become active and leave
its daytime hiding place. A g k i s t d o n bilin-
eatus feeds primarily on rodents, but will
also consume fish, from, lizarde, and
snakes I G m & CONANT1990). The spe-
cies is a livebearer and produces 6-20
yi3111Igper (~VAREZ DEL TORO1983,
GLOYD& C O N1999). ~

IGF-
Agkistm$rrn bi'&~~&# 18831,Ann.

Further Reading
Gmn & CONANT1990,P A R K Met
~N al. 2000
298
 Fig. 693.Afrspoi&a pic&i. Fig. 894. Atmpoidse, nunamifkr (near Quebrada
Photo: RW.Van k d e r OPanda, Cop&, kIondwas)

~trnpotdes Mps, h & n k r & a:rm!RB%w~,

~lrrap~~tosurii-8~
81% QpSaEaIfW
vogh.
M~.FLCBBOm.SrUYL&Pcrtdltfand
The three species that are included in the Me&@, $o mM i#
genus Atropoides are ~kkily-builtground- fBO0 m elem* drg! &f setst h t
dwelling pitvipers that are distfibuted in m Il&)ltrt, Fkm Bid.
Mexico and Central America (WERMAN 16% lotz~lizS:La
1992). Two species of the genus occur in Mag, tO ~ W ~ E Y
Central America. They are u ~ u d l yfound in vmztiam Famath IEifE1M)Q m
remote areas of extensive ~rimarvmwth.
As juveniles, their diet mkiats if insecteta. Key to AtnoPoidea
from, and lixards, whereas adults mually
hunt small mammals (A~vmmDEL TORO 1 a Mom than 138 ventral8 ,.................. .,..
1983'' D-g the rainy numnG
....*.....
........*..., .,..,..,...AtropoIde~ptcatloi
b Fewer than 138 ventral. ................... ,.,.,..
fir will give birth to 6-34 young with a tatal ,......,........,.......... Akopoi&e nrtmnrifir
s

length of 178220 mm (SOL~RZANO 1989,

LEE1996). Neonate Costa Riean A. nummi-
fer exhibit a marked sexual dichromatism
in head pattern (SOL~RZANO lQg0).
Eb'Lyz 1888, BOLORWNO 1989,
WERMAN 1492

* Atmpoldes picado/ i - kbb.695. Juvenile Atropaides numrnifer

(Unibn Barrios, Baja Verapax, Guatamda),
294
The genus Bothriechis includes a series of
small (generally reaching maximum total
lenpths betwoen 700 and 850 mm. rarely to
106mm),arboreal, preheneile-kiled ;it-
viper species, which occur predombntly
in Central America. Only one species, B.
schlegelii, also occurs north of the Isthmus
of Tehuantepec and south of Panama. This
species and the Co~lta Rican endemic
Bothnhchis supraciliaris are characterized
Fig, 696, Bothhchis rnarchi (Cusuco, Co~5,
Honduras).
Fig. 697. B o t h k h i e schlegelii (near
'hgucigalpita,CortBs, Honduras).
I
pig. 698, ~ ~ t ~ ~ a e s~i n (MOntaaaB
ua h de ~
la Unibn, Zacapa, Guatemala),
by spiky, enlarged, and protruding eyelid
acales. All other species of Bothriechis,
which are mostly green colored in life,
occur in isolated, wet, montane forests,
mually above 1000 m elevation, but two
species (B. bicolor m d 3. dhulassinus)
descend to about the 500 m contour.
During the day, these pitvipem rest, usual-
ly well-camouflaged, in the branches or on
tree t r u r b , making it prudent to check
carefully before leaning on a tree or reach-
ing for a branch when in Bothrkchis habi-
tat. Although relatively small, B, sehlegelii
posseases highly toxic venom, and bite0 by
this 'pecies can be life-threatenin~ to
humane (SEZFERT 19833, The B o t h h h k
species feed on firrgs, lizards, small mam-
mals, and birds (DUELLMAN 1963, A ~ v m z
DEL TORO 1983, CAMPBELL1998a).
Bothrkchik morehi specialiw in a diet of
frog8 of the genus Eleutherodactylus, and
in Cusuco National Park (Honduras) has
nearly vanished since the Eleuthem-
dactylus population8 dong the stmame
there have plummeted in numbers CJ.R,
MCCRANIE pers. comrn. 1999). All Bothri-
echis species bear live young. For R schle-
gelii, the litter size variee from 12 to 20
young (AMTONIO 1980, CAMPBELL1998a).
Bothriechis nigroviridis haa a dbtinctly
seasonal reproductive cycle, producing 6-8
295
young (1B8-177nun Wd le@ during
the r&ny eewon, with coloration identical
to that of the adulb (Bod-0 1Q97).
The young of 3. h k d i ~on, the athm
hand, m bmwn in color I H o ~ R
1896)
Fig. 699.Bothriechia aurifer (Uni6n Barrios, Fig. 702, Bothriechis aurifir.
Cem Verde, Bqja Verapaz, Guatamala). Photo: R. D. B d e t t

Rg.670. Bothriechis nigmuin'dis (Costa Rica). Fig. 703. Bothriechis nigmviridis (San Gerardo,
Photo: R. W.Van Devender Costa Rim). Photo: R. W.Van Devender

Fig. 701.Bothriechb binolor (La Concordia, Fig. 704.Bothtkhirr rowleyi. Photo: M. Burger
Chiapas, Mexico). Photo: A. Ramirez V.
 low; snout covered with relatively lage
scales; 1-5 scales between the aupraocu-
lars;iris yellowish green or bronze ............
.............................Bothdeckis aulifer
b Domum with heavy black mottling; venter
with black speckling;tail tip black; snout
covered with m a l l scales; 4-8 scales be-
tween the supraocularq iris blackish ........
........................Botlrriechiu migmviridia
6 a Venter yellow-green with yellow or white
paraventral stripes; tail tip yellow; gener-
aIly 23 dorsal rows at midbody ..............
..............................Both&vhin htemlls
b Venter yellow-green without pale paraven-
tral stripes; tail tip black: fewer than 23
dorsal raws at midbody ......................... B

IBa Iris p e n in life; 7-11, rarely 5, scales be*

tween the s u p r a d a r s ; generally 21 d o r d
rows at midbody; wales on top of head
small, each scale with a single keel, adja-
cent keel parallel .................................. 7
b Iris yellow in life; 3-7 scales between the
Fig. 705. BothrieEhie late&& (Monteverde, supraoculam; generally fewer than 21 dor-
Costa Eca). Photo: B.W.Van Devender sal rows at midbody; scales on head (ante-
rior to level of supraoculara) h e w in
shape, many larga and flat or with irregu-
lar and multiple keels at dBerent angler
...............................................................8
Key to Bothllie~hie 7 a 27-35 interrictals .....Bothrdecktr btmbr
1 a Eyelid (W?erci1iafie8) scale@ spinelike* b 23-26 iaterricbls ....................,........be......
enlarged, and protruding ..............,.........
2 ........................ Bathriechie thahainua
b Eyelid sealer (auperciliariee) not spinelike,
enlarged,andprotruding .....................,... S a Snout 'meBothdechie ...........
and grnooth m
.............................. ww
2 a Venbr d t h dark spotting; dorsal mark- b Snout scaler smaller and numelvur,
ings diffime with irregular spots, skin be- some keeled ...........Bothrieeiris mamhi
tween these markings with numerous
small dots or flacks of secondary pigmenta-
tion (except for the &color yellow
"Ompel"color variations);ventrds 138-186 Sm gi 1964, cAMPmLL $ LAMAR
(146-167in specimens from Costa R i a ) ..... 1489, CRoTHER et d.1992,WrrsB~ MCCw
............................l30thdechSs ucklagelii 1492.SOUSwO 1497.SOMRZANO et d.1998.
b At least the anterior two-thirds of venter
without dark spotting; dor~umwith clearly
bordered spots or crossbmds, skin between
these markings unicolor and w,ithout
secondary pigment (groundcolor untfom);
.......................
...
ventral4 141-160.................... ...............
Bothrlechi8 ecrpnaciliah
8 a Dorsal surface of head with dark spots or
black specklea; a black poatocular stripe M
generally present ................................4
b Dorm1 surface of head unimlor without
black epots or specklerr; Hack postocular
atripe generally not present, if a postadar
stripe preaent, it is bluish .................... 6
r Bothmps puncfatus
4 a Dorsurn without heavy black mottling; Bofhmps a s p r
venter unieolor yellow-green; tail tip yel-
 with newborn young f d primarily in
Babhmw- -- t$a P&T wa@ont * u m o 8t C ~ A S
ISBBE,B&~FO~pB u ~ is ~a large,
m aed
The " t e ~ i m~ "barb ~ ~ ~ mo d~' a * , arbmalpitviper *ache8 mly the
Bolhmps uspw is known in Cent~d km portion
America, is the most common and nimt
daxlmm mom I ~ inBthe m@m.
ft lives net only in fmwM area&,but aha
in sec6ndarJr -mdergmwth near buman
habitation, preferably clwe t o watm
l%we, thia n o c t m d pitviper will hunt
I
&=a, W e , mid small mammLg, This
spedes is highly pmliii~and can produce

1 a Dorsal pattern with dark khgular rg~@

fdmer than 70 eub.021uW 23-25 dored
m m at m-i .............
Batlbrrrpr Bsger
b m a 1 pattern with dwk bandm armwed
in p t r h more than 70 subcaudals; 16-29
d w d m a at midbdy .......................
.............................
& r & h p sprntotu#m

Fig. 706. Juvenile Bothmps asper (Corwvado

National Park, Cwta Rica). Photo: M. b n

707. Bothmpe -per (Baealar Lagoon, Fig. 708. Bothrope punctutw (Choca, Colombia).
Quintam Roo, Mexico). Photo: H. Bahena B. Photo: M.Lundberg
299
The genus Cerrophidion iecludee four spe-
cies, two of which me distributd in high-
land Central America (CAMPBELL & LAMAR
1992, LOPEZLUNA et al. 1999). They are
small, ground-dwelling pitvipera that
usually have a dorsal pattern of dark spots
that are often EUBd into a ~ g z a g&ripe,
Cerrophidion godmani feeds primarily on
insects, salamanders, frogs, lizards, and
emall mammals (CAMPBELL & S O L Q ~ O
1992). The species of the genus
Cerrophidion are livebearem; C. godrnani
produces litters of 2-12 young (CAMFBEU&
1992).
90~6~24 ~0 Fig. 710.Cermphidkn godmani bouth-
wentern Handura~).
Cemphidian @mni (GUNBER 186;3),Ana.
Mag, Nat. b t , (5) I& 864; type lacaUW
Duenw and other prh of tableland af Key to Cemphidion
C h a b d a . TL ta TE4 mm. Z]i@m& from
wutheaetern O m 8 , 119sa;ico, to wwtern 1 a Most of the lateral dark body spots ara 6-7
hmma, lgOa4m m ~1ewa.tioninrain farest scales high; dark postorbital band not pale-
and doud fmsst, as well arr in pin&@& LO&, ..
edged above and below .........................
.........................CemphMdon godmcurl
CermphMdot~tzoMorum O~AMFBELL19861,J. b Most of the lateral dark l d y spots are 2-3
Herpetor. 19: 48; type lac
%r 10.8 h ESE
8rra Criat6bal de ku Casts, 20 m elmation,
C b p u . Mnderr. TL Bo WO m. Meseta
scalea high; dark poetorbital band dietinct-
ly paleedged above and below ...............
GmW de C hi
am Mexi04 2000aOD rn ele- ....................
Cemphidfon tzotztlomm
vation in p i n d breat
Further Reading
CAMPBELL gt LAMAR 1992, CAMPBELL&
1992, LBPEZLUNAet al. 1999.
SOL,~RZANO
Only one rattlesnake species, Crocalm
durissus, occurs in Central America. It
lives as a ground-dweller in dry areaa and,
depending on the season,is diurnal, cre-
puscular, or nocturnal, Usually during the
cooler season, they can be seen during the
day outside of their hiding places (LEE
1996). Because farmem will usually kill
nearly every rattlesnake that they came
mom,this species has become extremely
rare in many areas. Crotalus durissus
feeds primarill' on mdents, but wfi Fig. ipU Qmtcdw duri.nu, tmbcan (Sian
consume lizards and ground nesting birdg Ka'an Biosphere R e ~ e ~Quintana
e, Roo,
(KLAUBER 1966, CAMPBELL 1998a). The spe- Mexico). Photo: B.Bahena B.
cies is a live bearer, producing 16-47young
per litter (ARMSTRONG & MURPHY1979,
CAMPBELL 1998a).

F'urther ]Reading
KLAUBER1956, CAMPBELL& LAMAR1989,
MCCRANIE
1893a Fig. 712.Cmtalu8 d. durissus (Depto. Jutiapa,
Guatemala).

Fig. 713.Cmtalus d. durissus-(Banta Rosa

National Park,Costa Rica). Photo: M. Franzen
 -
genus Lachesis was earlier seen as
monotypic, until RJFA(1994)and ZAMUDIO
& GREENE(1997)established the morpha-
logical and molecular genetic differences
and were able to show that Lachearts was
made up of at least three species, two of
which occur in Central America. The
b u a b a s t e r is the liegest of the New
World venomous snakes and the only pitvi-
pet there that lays eggs and actually will
brood them. They live as crepuscular and
nocturnal sit-and-wait predators in pri-
mary rain forest and feed predominantly
on mammals. While L. stenophrys can be
considered quite calm, L,melanocephla i~
very nervous arid aggressive, particularly Fig. 714.L a c h i s stelatrphrys (Costa Rim).
Photo: R. W.Van Devender
\
the males during mating season (RIPA
1994). Irritated specimens of L, melano-
cephala will sometimes even follow a per-
son for eeveral meters, GRing their heads short time in order to drink IRPA 1994).
cobra-like up to 50 cm above the ground. In h s h l y hatched bushmaster babies have a
the case of a venomous ~nakeover two total length of about half a meter (RPA
meters in length,thie behavior is certain to 1994).
impire a substantial degree of fear (RIPA
1994).The females will retire to a dark and
secure hiding place in order to lay their
eggs, and appear to like abandoned aputis
b m w s for egg laying (RIPA 1994). The
bushmaster will lay 9-19 eggs per dutch,

3
which are guarded by the female during
the 60-77 days of the incubation period.'Fhe
animal surrounds the eggs with the coils of
her body, not eating during the entire
period, and only leaving the clutch for a

Key to Luchesie
I a Dorsal surface of head black; 209-216 ven-
trale ..............
Lttckesis ncelaloocephda
b Dorsal surface of head pale brown; 198-209
ventraIs .................LQchesia ~ t e n o p h v

Further Reading
VIAL & JMENEZ-PORRAS 1967, MART~NEZ&
Barnos 1982, SOL~RZANO
(PE CERDASX986rRIPA
1994,Z m ~ LoGREENE1997
 I The genus Pbrthidium cxllmiats of a gwup
of small (to 900 mm total length), dender,

II
terrelstrial pitvipecs, some af which have a
snout tip that points cm~picuouelyupward
(these are commonly refemd to ae the
hognosed pitvipers). Recent revision^ of
the genus Portht'dium (CAMPBELL & Wwi
199%~WEW 1992, M c D ~ D et al.
1999)have restricted the generic name to
I the seven species (those listed below, plm
the rroork b w n rp. htwelo C ~ B E L L
&rocthe Mexican atate of 6olirna).Despite
Fig. 716. Lachasis m E a m p ~ CSan
a Vita, their modest body aim, bites from the Bpe-
C o d Rica), w.
Phatn; R, Van a%venhr ciw ofthi8 pnua can be life-threatening to
humans.

The Porthidium species are generally cre-

puermlar and nocturnal,but can be found
outside of their hiding places in ahady
during the day. Their diet ia made up
of h g ~ b, a d e , blind snakes, and small
marimale, with the young including inver-
tebrates, such as earthworms (PO- et
al. 1981,Lm 1996).All Ebrthidiuna species
bear live young; litter size: l?naeutum 8-18
young (CAMPBELL 1998a1, Z? ophryomgas
19 young (LAWSEN 19971,E! yucatanicum C
10 yoang UvfcCcw & CENW 1992).Sexual
dichromathm has been reported in Z?
FYg. 718. h h e s i s stmophrys lDari6n region, ophysmegas, with the ground celor being
Panama). noto: E* pay in males and tan to grayi~hbbrow in '
femalers laside from differeneers in color
pattern; see QOL~RZANOet al. 1988, LAW~EN
iog?).

Porthidium

r P.lensbetgii .
Fig. 717'. Lachsis sbanophrgrs I S i q u h , Coata
Rim>. Phoh R,D. Bartlett L .. . - -
 Ft
%.%p*
Ium unn
'
Mk. Tw$cr
~ ~ n 3 v P
lmali*: mmte 0
-
an
UVm3

Mexico.
%?to 610 mm. Bou&ktarn 0- to
--v M d a , ma 1 4 to 500 m
m1evathn in dry m a t .
firthWmz barmbergdi (SCIHI*E4(EEL 18413,Mag,
%ad. Rept. ( l 4 k pl, 1; type lode:Turbaoo,
Co1omb TL ta 900 mpa Fad& side d
e a s l ~ ~ a s w a ~ n o s t h e r n
Amrim em Pew1 ts 1370 m elmation in dry
~ L f r a l d w e ~ ~
Mhiddum z%4wt4jrun[Bwm1W), Aml,
W. Nat. F'az& (6) 10
Fmis&,anb&dRfo
TL to $300 m. Morthwmtem ehiapa$
Maxi~~*ton&- 3&* w a l e d te
900me~aionhrain~
firth- op&q~mega#CBOCOVRT 1WKi,
Ana 94i. N& Parfa (5) 10: 2Ot;tlpr kelif$
Fig. 718.Porthidium ophryonaegae (ElRosari~,
Zacapa, Guatemala).
@onw-
slaps d mountaSnri at
[act
&&a L-
-thd
TL to T 7 4 3 m m , ~ t h e e n t r a l Q y ~ 1 ~ t
k t a Rirta (ocollrrenee in Panama qtaestb*
~,~lwdtOlQOOtrt~a~in
f0rea-t; and sawma,

w & d % ~ C
~ w t a, R i s T L
ta $239nun.Vdh de:General in the .smt$mst

EbrthMiacrn yumtalsicam I S m 194lf),

2%mk&m 88: Bg
P hi?&@: C
Yuealh, Mdm. h &HI ma^ The d m
po~ofbY~toInRePinanzla,
w IW,
aaa level to
2BO m al~vetionin drg forest.

I
Rg.719.Porthidium dunni (Arriaga, Chiapas, Fig. 720.fbrthiditbrn lansbergii.
Mexico). Photo: A. Ramirez V. Photo: R. D.Bartlett
gey to RDrthMium
I a Snout tip pointe conspieuousIy upward
(Fig.721) ................................................
.1
b Snout ti does not point conspicu~taly
vprd Re 722, ................................ 3
2 s 123-145 ventrdr; outer rims af sapraocu-
lars form a afiarp edge .......,...........,......
.......,.....................Pw#AidIunt Raertma
b 159.161, as s d e mare than 145 ventral&
outer rims of suprawdam rounded OP
..........................hrthidimt kurab~rgi~
8 a 141-158,a8 a rule fewer than 158 v~etFals4
b 15&173ventraIs ....................
,..,,.,,,,0
4 a %I-23dmd FOW8 at d d b a d y I fewer than Fig.7%. fiflWivm m a a n d m
21 dorsal rows one headlength anterior o f b, ~ d ~ ) , Photu: H. Bahena B.
cloaca ..........................
Por#Ibidbn darnnt
b 83-27 d 0 ~ 8 dlDW8 at rpjdbody; dorsal
rows one headlerrgth anterior to cloaca
.............*........
maidkrnr yuretmdeum
......
I a More thag 40 dark do& qmt& 25-88 sub-
caudal&............PoFtbidiarm v o l c u n b ,
Further L i t e r a m
AMARAL 1929b, PORRAS
e t al. 1981, WILSON &
MCCRANIE1984, CAMPBELL& LAMAR 1989,
McCoy & CEN~KY1992, SOL~RZANO
1994

Fig. 724. Arthidiuna ylscabanicum (Dzibalchen,

Carnpeche, Mexico). Photo: R. Cededo V.

Fig, 732, hrthicdium aphryomegas, Fig. 7%. M d i r t m madurn (Gem &lapa,

ma*, M.VeaeIjr A~~ NicmagmI.
Scalation Characters in Snakes
Data from W m N 8t 1985,~IIEZ-SANTOS 8z MORENO1988,EIEE 1996, MZE1896, GAMPBU
1998,origlnd clescriptiuxm (in poorly known speGi~~), JAYWAGE (pen.c o r n 2000)and Khhler
unpubl. data. Sm &B ad blind sn&a rtre not inlcuded, kbbredatians: Dor~ale-MBInurnber
of dorsal m e at mid-tpody),DorshHC (number ddoxaal row5 one head length anterior ta d w
hied on posterior body).
car, k-post ( d a d ~ s d m

Ehidae
Boa constrictor amcloth undivided
Corallua annulatus smooth undivided
ComlEus ruschenbergerii BmOOth undivided
Epicnates cenchrh 8rnDOth undivided
h c e m u s bicolor 19moOth divided
Dachybaa boudengeri hled undivided
Wngaliophis cantinentelts gm00t.h undivided
Ungaliophis panamerasis smooth undivided

Colubridae
Acieiphicos daryi divided
Adelphicos i b a m ~ u n a divided
h l p h i c o s latif'cksciatua divided
Adelphicog nigrilatus smooth divided
b l p h i m s q wdriuirgatus ~rnootsl divided
Aoklphicog uencepacia smooth divided
Amaetridiurn mpperi emoath divided
Amastridium veliferum ~mooth divided
Atractus clarki undivided
Atractus darienensb undivided
Atraetus &pnessiocellus undivided
Atractus hastilitmctus emth undivided
&ractw imperfectus ~mooth
Chapinophis mnthocheilus l3nwth divided
Chimnius carinatus hded divided
Chironius exdetus keeled variable
Chimniu~ grandi8quamb bed variable
CleEia clelia ern6 undivided
Clelie equatoriana amooth undivided
Cielia scytarmlina undivided
Coluber comt&tar divided
Coniophanes aitramzi divided
Coniophaw bipunctatus divided
Coaiophanes fissidena divided
Coniophanw imperialis smooth divided
Coniophanesjmnae amOOth divided
Coniophenes meridunw emookh divided
 Coniaphanes piceivittis 78-115 smooth 19-21 divided
Coniophanes quinqueuittatus 63-70 smooth 17-21 divided
Coniophanes schmidtb 83-115 smooth 19 divided
Conophis lineatus 56-80 smooth 17 divided
Conophk vittatrwr 66-76 ~mooth 17 divided
Criwntophis nevermanni 71-89 smooth 17 divided
Dendrophidion nuchale 132-163 k d e d 16 variable
Dendrophidion paucicarinutum 119-139 keeled 16 variable
Dendrophidion percarinaturn 133-164 keeled 16 divided
Dendrophidim vinitor 111-128 keeled 16 variable
Diaphodepis wmgneri 133-138 keeled 17 undivided
Dipsas articulata 108-135 smooth 16 undivided
Dipsas bicolor 111-129 smooth 16 undivided
Dipsaa bsevifbciee 70-101 s m d 15 undivided
Dipma maxiElwb 84 ~ m t h 15 undivided
Dipsas ternpodis 125-132 am& 15 undivided
Dipsas tenutssima 128 lsmaoth 16 undivided
Dipsas uariegata 84-93 ~11~00th 15 undivided
Dipsas &uteri 102-127 smooth 15 undivided
Dryadophis domalis 109-137 ~rnooth 16 divided
Dryadophie meEanolomus 85-136 amooth 15 divided
Dryadophis pleei 83-105 smooth 16 divided
D r y m a d o n corais 66-88 mmth 13-16 undivided
D~yrnobiwbshloroticu~ 107-125 b l e d 16 divided
Drymobiua rnargaritiferus 103-138 EraeId 16 divided
Drymobius rnelanotropis 81-108 h l d 16 divided
Drymobius rhonabifer 84-102 M e d 16 divided
Elephe fiuirufa 96-122 k - p t 19-26 divided
Enuliophirr miuteri 88-109 smooth 16 divided
EnuEius bifoveatus 100-120 ~ m m t h 16 divided
Enulius flavitorques 86-121 smwth (15-)I7 divided
E n J i u s routanensis 103-121 smooth 17 divided
Erythrolarnprus bkona 47-60 ~mMIth 16 divided
Eryttbmlamprus nairnus 42.61 smooth 15 divided
E'lcirnia publia 26-44 smmth 17 divided
Ficimia rumimai 38 emoath 17 divided
Geophis bellus 33 k-poet 16 undivided
&aphis brachycephalus 30-51 keeled 16 undivided
Geophis cancellatuar 21-23 smooth 16 undivided
Geophis cerimsus 4148 k-w& 17 undivided
Geophis chlampioni 23-33 ~m00th 16 undivided
Geophia dambni 41 mooth 16 undivided
Geophia downsi 41-46 k-post 18 undivided
Geophis dunni 36 kwled 17 undivided
Gophis fulvag~lttatus 34-36 mmth 17 undivided
Geophis godmani 26-36 smooth 15 undivided
Geaphis hoffinanrsi 2437 mooth 15 undivided
307
 Geophis imrnaculatus smmth 17 undivided
Geophia laticimtus srnaoth 16 undivided
Geophis nasalis keeled 17 undivided
Geophig rhodogaster emooth 17 undivided
Geophis ruthveni k-rn 15 undivided
Geophis talamancae k-post 15 undivided
Geophis aeledoni smooth 16 undivided
Hydromorphus concolor SMOO~~ 16 divided
Hydromorphus dunni s m d 13 undivided
lmantodes cenchoa grnoath 17 divided
Imantades gemmistratus mooth 17 divided
Imantdee inornatus ~rnooth 17 variable
Imantoales pkantasma smooth ( 16-317 divided
Imantade~telnuissimus emooth 17 divided
Lampropeltis trdangudum smooth 17-1s undivided
L e p t d i n a annulata moatll 15-19 divided
L e p t d i m @nu fa ~lnooth 13-17 divided
Leptodeim nigrofasckta a m d 16-17 divided
Leptodeim rubricata rrmootb 17-19 divided
Leptodeim septentrionalis smooth 11-17 divided
LepEodrymus puleherrimus smooth 15 divided
Leptophiar ahaetulla keeled 11 divided
Leptophis depressimstris keeled 11 divided
Leptophis diplofropis keeled 11-13 divided
Leptophis mexicanua Beeled 11 divided
Leptophis modestus- hled 11 divided
Leptophis mbuIosus bled 11 divided
1 Leptop his riueti
Liaphis epinephelus
Liophis lineatus
bled
mooth
11
~ ~ g ~ ~ l t h 16
15-17
divided
divided
divided
Manolepis putnanai mmth 16 divided

I
Masticophis rnentovarius am& 13 divided
Nemdia rhambifem h b d 21-25 divided
Ninia atrata keeled 19 undivided
Ninia diademata keeled 19 undivided
Ninia celata keeled 19 undivided
Ninia espinali lreeld 19 undivided
Ninia rnaculata h l d 19 undivided
Ninia pavimentata keeled 19 undivided
Ninia psephota keeled 17 undivided
Ninia sebae kd0d 19 undivided
Nothops% rugosus keeled 22-26 undivided
Onoadiphns aurula smooth 17 divided
Oxybelig oeneus keeled 13 divided
Oxybedlis breuimtris keeled 13 undivided
Oxybelis filgidus keeled 13 divided
Ckybelis wilsoni beeld 13 divided
 Oxyrhopw petola smooth 19 undivided
Phimphia guirml~ensis smooth 19 undivided
Pituophis Eineaticdlis bled 27-31 undivided
Pkiocereu;~eEapoides smmth 17 divided
PEiocerctts eurymnus smwth 17 divided
Pseudoboa taeuwiedii moo& 19 undivided
PWU#~~?S JWC!C$O?&~%~@B keeled 19-25 undivided
Bhadinaaca ana~hcrreda emootb 17 divided
Rhadinaea calZ&aster smooth 17 divided
Rhadinaea &comta smooth 17 divided
Rhadinam gdnaani emooth 21 divided
R h a d i w Isornn~teini amooth 17 divided
Rhadinaea hempstedae em& 19 divided
Rhdinaea kurta2&utchan mooth 19 divided
Rhadinaea MnLlini smooth 17 divided
Rkadnuea Zachrymana smooth 17 divided
Ahadinaen, macdsugalli smooth 17 divided
llhadinaea neontecristi smooth 19 divided
Rhadimetz pihmorum smwth 17 divided
RhQoliwup d 4 i smwth 17 divided
Rlaadimu pulwriuentris smooth 17 divided
Rhci3-a ragemmani smooth 21 divided
Rhudinaea aargenti emooth 17 dividi3d
Ahdinma selpem8ter smooth 19 divided
RMiraaea ataddmani emaoth 19-21 divided
Rhdinarea t o l p m m m em& 17 divided
Riadinarea uermi~ubticeps am& 17 divided
Rlaialaobathvyum bouallii k-pat 19 divided
Sorlud~mEenanbixzta gmooth 17 divided
Sixphiadontaphis annukaturr smooth 17 divided
Scolecophis atrocinctus mooth 15 divided
SentkelL triaspis k-post 27-39 divided
Sibon annulat.w smooth 16 undivided
Sibon antlamcops emooth 13 llndivided
S i h n argw emoc~th 15 undivided
Sibon mrri smooth 13 undivided
Sibon ditimidiatw mmth 15 undivided
S i h n lo&f.eaig mmth 15 undivided
S i h nebuiatui4 smooth 16 undivided
S i h n 8annwJe rpmooth 15 undivided
SiphEophis wrvifim ~mooth19 undivided
Spilotes pullatus hled 14-18 undivided
Stelaorrhina dagenhardtii eaaooth
17 divided
Stmomhine bminviElei smmth 17 divided
Storeria dekayi Wed 15-17 divided
Symph fmus kumtomus smth 16 divided
Sympkimus maym ammth 16 divided
309
llltntilla aibiceps divided
rtzndilla alticola divided
llzntilla armillaka divided
nntilla bairdi mooth divided
TantZEla brevicauda smooth divided
RntilZa cuniculator divided
lbntilla impenw divided
%ntilla jani divided
Tbntilla lempira divided
TantilEa mwsta divided
%ntilEa reticubta divided
Tantilla rubm divided
Tantillu mficeps divided
antilde schistma divided
Tantillu supracincta ~rnwth divided
Tantiltca taenkata smooth divided
Tantiliar tayrae divided
?batilia kcta divided
Tantilla triseriata divided
Tantilla tn'taeniata smaoth divided
IRanfilla wrrnifirmis smooth divided
Rntilla vulcani smooth divided
Rntillita breuissima smooth divided
Tantillita canula smooth divided
Tantillita lintoni smooth divided
Thamnophis cyrtopsis keeled undivided
Thamnophis filvus keeled undivided
Tharnnophis marcianus hled undivided
Tttrsmnophisproximus keeled undivided
Tretanorhinwr nigroluteua w ed divided
Tretanorhinus mcquardi bled divided
Trirnatopon barhuri e m d divided
Dimetopon gracile divided
Tn'metoponpliolepis divided
Trirnetopon simile divided
Dimetopon elevlni smooth divided
%metopon viquezi smooth divided
7kirnorphodon biscutatus k-pat variable
W p n u r g o s compressus smooth undivided
'IFopi&dipsas fasciatus smDOth undivided
l?vpidodipsa fischri bled undivided
I).opidodipstw sartorii keeled undivided
Umtheca hcipiens smooth divided
Urothca filuiceps divided
Urotheca wenthen' divided
Umtheca myersi divided
 180-188 104-124 em& 17 - 17 - divided'
1!U-153 35-62 rrmooth 10 17 undivided

em* 15 divided
smaoth 15 divided
a n d 15 divided
Miemrus broulni ~mwEh 16 divided
Micrurw cehrki smooth 16 divided
Micrurus diustema emoath 15 divided
Mkrurus dissoleucus smooth 15 divided
Micrurus elegans mwth 16 divided
Micrusus hippucrepis smooth 15 divided
Micrurtls latifasciatu smooth 16 divided
Mdcmnte mipartitus emaokh 15 divided
Micrurus rnultifascietus smooth 16 divided
Micrurus niglracinctw smwth 16 divided
Micrurua ruatanus smooth 15 divided
Micrurue stewarti Emloath 15 divided
Micrurae stuarti smooth 15 divided
Viperidae
Aghistmdon bilirseatus keeled 23 undivided
Atropoidt!~nunamifkr keeled 23-31 undivided
Atmpides picadoi bled 23-29 undivided
Bothriechie aurifer bled 18-21 undivided
Bothriechis bicolor keeled 21 undivided
Bothriechia Eateralis
Bothrkchb marchi
Bothriechis nigroviridia
Bothriechis rowleyi
Bothriechis achlegelii
M1
4d
b l d
k d d
Wed
keeled
21-23
19-21
17-21
19
21-25
undivided
undivided
undivided
undivided
undivided
I
Bothriechis supracilhris bled 21-23 undivided
Bothriechis thalassinus keeled 21-23 undivided
Bothrops asper kded 23-26 undivided
Bothrops punctatus hled 25-29 undivided
Cemphidion godmani keeled 21 undivided
Cerrophidion tzotziEorum blad 19-23 Undividd
Crotalus duriseus k ~ l d 26-33 undivided
hchesis nelanocaphda k~l d 36-40 undivided
Laches& stemphry8 keeled 33-38 undivided
Porthidiurn dunni keeled 21-23 undivided
Porthidiurn lansbergii keeled 23-26 undivided
Porthidium naeuturn keeled 21-27 undivided
Parthidiurn ophryomegas W d 23-27 undivided
Portkidiun volcanicurn k d d 26-26 undivided
Parthidium ytdcatanteurn k-14 23-27 undivided
311
 -
dnakebite Prevention and ~ mAid
t -I

Snakes in the tropics pose a health riek

that should not be overlooked. Life-threat-
ening snakebites in Central America occur
most commonly among natives working k
the field, with the highest mortality report-
ed in incidents involving pitvipers (partic-
ularly Bothrops asper) (D. HARDY 1994).
Taking appropriate precautione can dimin-
ish the risk of makebite, A study of the
incidence of makebite amongst field biolo-
gists provides evidence that the risk of
being bitten by a pitviper, even in areas
with a high population of these snakes, is
slight. The use of proper equipment and
procedures within make habitat will en-
sure safety. Fig. 727. Bothrops aepr (Nicaragua).

Bafety Precautions

Suitable clothing: long, loose-fitting
panta, ~turdyfootwear, preferably rubber
boots, and a hat for protection against
tree-dwelling slnakes. A large pitviper,
however, can bite thmugh rubber boob.
One should only proceed through areas
where it ie clear that there are no
snakes; da not walk through dense bush
or high grass. Likewise,be certain there
are no snakes under or behind fallen
logs before stepping over them.
Only hold onto tree trunks or branches
once one has determined that there are
no sn&es present, Unless it movee, el
pitviper can be virtually undetectable in
the twilight of the rain farest, due to ita
cryptic coloration. Snakes reflect light
quite easily, and at night can be spotted
with a flashlight from a distance of
several meters,
When suddenly faced with a snake, it iB
important to remain as stiIl as possible,
as snakes have relatively poor eyesight
and will mostly strike in the direction of
a moving object, sudden movement must
absolutely be avoided. One ~houldthen
calmly withdraw.

Recommendations for Fir& Aid

The elaboration of a snakebite protocol can

improve the outcome of a venomous snake-
bite in the field, This protocoI should at
Fig. 726, Uiflicult to spat:a juvenile Bothrops least include (HARDY 1994):
asper (Tbrtuguem, Costa Rica),
Possible Resting Places of Pitvipers
 Fimt aid measures to be used and thbae specific tharapy far venomous snakebiite,
wMch will not be used. hymen o h n have the impreasion that
b u t e and meam of t m m e t i o n for once the antivenin hw been idminiabd,
watmtioa~name and lmtion of n e m t the patient ie out sf danger. However, the
medical f a c i l i t ~ ~ w e of antivenin carPiea its own ricrk.
Occaeionally, a patient may dmlop symp
Lht of medical c a d b t i c m resources toms of life-threatening anaphyla&ic
with ezpeIttiae in makebite treatment ah& facial swellingsresphtary d i s t r e ~ ~
in~1ucringkm*m aad telephone num- rapid drop in b l d preswe, and pallorF
bers bpth in m t r y and ovmeaa
Variaue snakebite kits (e.g,, "Sawyer
The psy&ologiad reaction of the snakebite ElrtractoF", *Aspivanin') am available m
r i c t h will greatly &6et the outame of the trade. They have been proven useless
entteil~mtim~ The victim should b ~ W U F under field eonditiona (D. MFBts per&
aged to remain calm and avoid @c. comm. 20'000). Barn extractam can lead fio
Under no s h d a pereon in3uriea and actually be dangemus: in
who has b n bitten be led% alone, situations where the perrom produces an
anticoagulant fleet on the bl~od,bmoF
The afB8cted extremity ehould he immDb2- rhahpe em o w r . There is a popular mh-
i d ( a m in o sling:,leg splinted, ars appro- conception that the venom oan be somshow
priate), in order ta e l m the 8pmad of bhe remmd by removing the m.mvuadingti&
venom. Hhgs and braceleb should be sue (e.g. by amputating a finger or making
immehtely removed to amid atting off an Weion h the skin). Most snrgid
the circul~tkiona ewellinJSoccurs. intervantiom for =*bit& actudly eatwe
fwthetr iqjurg artd either d~layar entirely
Whenavw pm~ible,the snake ~ h d ba d prevent moovery (NIEm20001.
identifled in order to da@rmine epecifit:
theraps Capture and killkg of the animal Under no ciraurnstaaces should p m s
should a d y be undehken if na threat is d ~ , s l z c h a s ~ d a n , ~
tO the vi& or helpm. Do mt handle applying suction, be perfomed an the
a seemingly d ~ a dmake. B n h are ex- wound site (am d e o =&bite kits). Such
perts at feigning death, They may even measmrr ofban cause damage to the largsr
twi& thenselva u @ d ~down and lie with vessele a d cauere the venom ta spread
thekr mouth open and the tongue l o k g more rapidly through the circulatory
out. Also, a fiesbly kiZled snake o h $ill 899tem. There is alao the dmwr of semd-
has some kind of bithg mfkes and might a q i d i o n , m d tissue damage tag lead
be able b deliver bits to hemorrhage if the venom b an antico-
agulant &ed. Ttra venom of most Central
The snakebite victim needs to be brought American pitvipers hae 330th pro-cosgulmt
to the n-st paint of m e d i d treatment and anti-cog_eflrlastproper tie^, which Eead
wi qui& as passible, At any time, respira- to the cmsump~oon ofthe blaad'~c l o t t h g h
th%b,l ~ d a d CO&OWI~QB& of tbrs (in a ~POC~PJJB h o r n a~ conlumption
the victim negddl to be manitorsd. A bfief 00agrOpatmy or diarseminated h h v h
outline of the drcum-ces of the 4- clotting). In humans, the fibrinalytic
dent, tha p m g m ~of the envmm8tiorn7 sptem irs activated to prevent clottingpbut
inf01"~ati;on about the vidhn, any evidence, once the blood clotting factbts are ussd up,
p a m o d observations, and measwear the b l d becomes inuoagulable, If hemor-
taken can k be1pfi.d for later treatment. rhagic ftrctorer la,g,, enzymw that perfbrake
DUB to paEdble3 daonpw.tione, k p e & c the b a d membrmea of blwd vesseb) m
measures should only ba undertaken by a pwent in the wenoan, this can lead ta
quaWed phyElitlian. Antivenin iis the only gpentanmuar major internal md &d
blmdiug. Tying off the a & e M adremi* is
not r m m d s d , as t b will diamppt the
blood flaw and l e d to tissue damage.
Extreme heat (c~utarhationlar 6oolbig of
the wound sit% are equdy dertrimmtal.
Applidm or iqjectim of home remadies,
a ~ k a t i o of n alcbhol or coffee, or the
ingestlon of food (which may cause v d t -
ing) we d a o oon.traindimM.The apphi-
tion af a pmwtm bandage is not advimble,
a~ a great deal of ~mllingwill mmlt &m
a Piper bite aed t h eiroulation
~ be a t
off (b,MZBS peps. comm. ?2000). Fig. 7 H . Cbmphk ~ i ~ un&e ;the snlarged
too* of this rewfarrged make.

Rear-fanged Snakes
Many colubrida produce a secretion in the
Duvernoy gland, located in the upper jaw,
that in some species is highly toxic. The
rear-fanged snakes have enlarged, grooved
teeth in the rear of the upper jaw (compare
Figs. 729 and 7301, that enable them to
inject this secretion into their prey or into
potential predators. While some Old World
rearfanged snakes are classified ae dead-
ly (e.g., the boomslang, Dispholidus t y p w ) ,
there have been no known fatalities frvrrn
envenomation by such snakes in Central
Fig. 730. Skull of Xenahdora m b ~ p h a ~ u s ;
America N -
( 1990). the enlarged tooth in the upper jaw is clearly
evident, Photo: R.W. Van Devender
Yet, the rear-fanged snakes of Central
America cannot be considered entirely
harmleas as certain epecies (e.g., Conio-
phanea spp., Conophis spp., Pliocercus
spp., Xenodon spp.) produce a highly
potent venom, which causes not only local-
ized ~lymptoms(pain and swelling) but
toxic systemic effect^ (headache and
pereistent bleeding) (MINTON& MEBS
1978, GREENE 1997).In order to avoid bitee
from rear-fanged snakes, these animals
should only be handled, if at all, with
leather glovea and protective, long-aleeved
clothing.

Further Reading
CAMPBELL & LARlAR 1989. MINTON 1990. D.
HARDY 199% n. H ~ D Y BLVA fig. 731.Xenodon rabdmephaltka (Costa Rica).
1998,MEBS 1992,2000 Photo: R.W. Van Devender
3 16
Mimicry

Mimicry in Snakes
Mimicry is defined as deceptive imitation
by an organism with regard to form, color-
aiion, or-behavior, ueu&ly to frighten off
potential predators, but, in some cases, to
attract prey Bateaim Hlirnicry (two similar
species, of which one, the model, is veno-
mous or inedible and the mimic i8 non-
venomous and edible) is differentiated
ficom MulIerian mimicry (both ~peciesare
venomous and are model and mimic of
each other). The concept of Mertensian
mimicry la mildly venomous model, a high-
ly venomous and a non-venomous mimic)
has been dismissed as ~uperfluousby some Fig. 732,A true aord snake (Micrurras hi
authors (GREENE& M C D ~ I 1981).D pocrepis) from Belize. Photo: P.&cud
In the latter half of the lgth oentury, re-
searchers had already noted the remark-
able similarity between venomous and
nea-venomous coral snakes and certain
colubrids (COPE 186Od, WALLACE 1867,
1870). The possibility that the brightly
colored true coral snakes (approximately
50 species of the genera Mkrurus and
Micruroides) and the many similarly col-
ored non-venomous or mildly venomous
colubrids of several genera (among others,
Erythdamprus, Lampropeltis, Oxyrhopus,
and Pliocercus) are involved in a complex
mimicry system has created much cantm-
VersY (DUNN 1964, BRATTsTEOM 1966, Fig.733. Pliocercus elapoides (a specimen from
J3mxr~& I\~ARIEN1956, MERTENS1956).No Belize) exhibits a very similar color pattern as
fewer than 115 species of colubtid (18% of several species of venomous coral snakea,
dl Amaricaa snakes) digplay elements of Photo: P.S W d
coral snake coloration and might be con-
sidered as mimics of Micrurus (SAVAGE &
SLOWINSKI 19921, The coral snake mimicry
hypothesis states that the conspicuous
colors of the venomous species act as a
warnina ta potential predators and these
fll
predat&~611 also avbid similarly colored
non-venomow and mildlv venomom snakea.
One of the primary a r g e n t s against this

Fig. 934. Eryihtvlamprtbs blzana has pairs of

black ringa, whereas in the Central American
Micrurus species, the black ringa are either
single or in triads.
hypotheaia, that the true coral snakm The idea that Mcrurus spedea are not just
s e m se a model for a W b l e number of mimicked, but themselves mimic certain
oolubrido, i~ that. a potential predahr leas venornoue rear-fanged snakes
would ad SURive ~ bite of sr M i c r m ~ r (&rythrulanapras, Plioc~mwslneedla tcr be
and,thus, h e f i t ttm hhe acquired e.qed- examined MEETENS 19661. Dgepite plenti-
ence. Gama 8k M c (1981)~augges- ful evidenoe that mimicry is the muse of
t d w v d possiMlii5es for chuglwnw the above-named ~ i r n i l d t l ~between
s
this 'deadly model*, a d report Ul& t b many unrelated ualubdd m d Mbrurtt#
bite of a Mferww need not be deadly, even spades, leshl obviaus explanations fbr We
fOr emdl vertebratm. In predatom that type of color similarity s h d d not be d i m -
hunt in graups, an imlividuai caa l e m h g d d . Tbe wmspondence may be the
avoid tb.e color;a$im of cord a d s by result af otber m m m t a l fhtctors that
observing the &fate of a compcifio that has have nothing t o do with mimimy?or it may,
been b i t h by a Micrurwr. The possibility in fact, be ooincidence. Evidence contrary
of the innate avoidance af snakes with to them alternative explanationsis pravid-
bright red and bl& ban&, whieh aw the ed by the fad that the suppasad madele
w d g colors of coral ernalras, should not and mimia inhabit different mimohabi-
be oubr1ookad. Tho@ individuda with a les- tat& to diffetrent environmental fan-
ser innate h d h t i o n to attad md makss tore yet they wmspxmd w e t l y ta the go-
have a higher pmbability of e d v i n g , aad, graphic vadability of models and mimics
therefme, a -selective advantap over m- with a b m d range, There are also o w e d
specifics that are not deterred by wflrr-l documented cam of supposed mudel and
make coloration. Contrary ta pr.evt.~ua mimfc pafrsl in wBi& unwaal color v&a-
assertione that they am ~trict1ynoctamd, tions mxw which are othemi~tzunknown
coral ~n.aakeaW h r u s and the bulk of in their respective genera, as for example
their aapgosed mimics) are also active apotted Micmrust and Plkce~eusas well as
during the day, so that pTedatum can eaai* black Miwtwzu and 8sythrdrxmtpr~ls.
137 epot their bright mlor~.
Finding8 to date suggest that certain epe
In recent literatw~,several impcatant cies s;F Abractus, JytlamLampncs,
arguments far mimicry mmg New World LQmpmpeEti~,Mim~mides,M k w u s and
sn&~ with blaak-rerl or black-flow-red PEiamrccsa and other genera am integrated
r n a k h g ~have bean publbhd. Expan- intn a ctimplerx ayataxn of mimicry whom
m~ntalwork relating to me phenommon dpamiea and relatiowhips hava yet to be
ha@ also been pedbrmed (GREENE& investigated CGREENE& 1McIhmm 19811,
M c D m 1981, SAVAGE L Smwmsm The phenomanon of mimicry is not Ilmlbd
1992, BROIIIE N Bmx,m & ammg makee in C e n h l America ta the
Bt J A ~ E1995,
EvZoom 1996,Wasorv & d.1996). An exact abwe mentimed venolaws and noa-veao-
geographic c o r r e ~ ~ n d ~etndl~st9e among mom coral snakm. Even the highly
several supposed model-mimic pairs dmgerm~pihipem CBothmp8 and related
[GREENE& ~~CDIARMII) 1981,Wmoa e t d. species) dao have imitatom, su& as 18-
19961, F%qr)rted mimics m a g wltlbrid tain Sibon s&w aa wen as Ceptodeir~
speciee with a brmd xmge display conspic- fenata, and especially the 'Falso
uow color variatios that correspond to TeraiopeloH tXemdsn rabdoeephaluid,
their ~ u p p o s dmodels. !Fhh is p m t i h k l p which at & ~gt I m appears qaite similm
notable in the various Pliwercw and tQ the pifvipm,
Zrythr&mprw populations, whoas calm
cmespmd markably to thoae of gympat-
ric M k r t k t ~e~p b .
Mimicry I

F'ig. 735. Scaphiodontophis annulabus (near Fig. 738. Rhinobothryurn bauulbii (Mo
Lim,6n, Costa Rica). Photo.A Hohmeister Siquims, Costa Rica). Photo: A Hohmeisbr

Fig. 736. Erythrolamprus bizonu (EUo Fig. 739.Erythrolamprus rnirnus (near Pueblo
Siquirres,Costa Rica). Photo: A Hohmeister Wiso, Jinotega, Nicaragua).

Fig. 737. Scolecophis atrocinctus (Nicaragua?). Fig. 740, Ezntilda suprecincta (Osa Peninsula,
Photo: R.W.Van Devender Costa Rica). Photo: B.W.Van Devender
 Mimicy

Fig. 741.Plimercus euryzonus (Selva Negra, Fig.744, Lampropeltis triungulum naicropholis.

Matagalpa, Nicaragua). Photo: U.Kuch

fig. 742. %pldodip~e sartorii Inear F'ig. 745. Oxyrko u9 petola (La Fortuna, PMV.
Escaruega,Campeehe, Medoo). Photo: J.C. Lee #i
Alejuela, Cwta ica),
Phm R.W. Van Dwenderr

Fig. 743. Rcinaia publia (La Ceiba, Hondum). Fig. 746.Dipam bmvifacies (Yuca*, Mexhol.
Phoh C. h e

319
Aclmowldgmente

Acknowledgments

Numerous people have assisted me in the herpetology a t Senckenberg Museum,

various phases of my work in Central Frankfurt, Germany,have tested the iden-
America over a more than 16-year period tification keys included in thia book. Their
and contributed to thia boak project. help is greatly appreciated.
I appreciate the cooperation I have re- I am grateful to n u m e m u s colleagues for
ceived from forestry and wildlife officials in photos of Central American reptile
Centre3 America. Collecting and exporta- species-these photographers are acknow-
tion permits were provided by R. Rivera ledged in the captions of their reapctive
and Cruzcaya, Autoridad Nacional de Am-
biente (ANAM), Panama City, Panama; J.
Guevara Seaueira and R. Tenorio Jimdnez,
Ministerio dB Recursos Naturales, Energfa
y Minae, Saa Joe& Costa Rim; M. Fonseca
Cuevae, S, Tijerino, M. G. Camacho, and C.
Peres-RomBn, Minisbrio del Arnbiente Y

I
10s ~ e c u r a & Naturalee (MARENA~,
Managua, Nicaragua; A. Barahona, A. P.
Martinez, T* Garcia, and C. Romero,
Corporacidn Handurefia de Desarollo
Forestal ICQHDEFOR), Tegucigalpa,
Honduras; L. R. Arevalo and A. Sanchez,
Ministerio de Agricultllra y Ganaderfa, El
Salvador; J.I. Gdlvez Duifianez, Conaejo
Nacional de Areas h t e g i d a s (CONAPI, - --

Guatemala City, Guatemala; F. Ramirez fig. 747. Expedition t o Pim Bonito, Honduras
Ruiz de Velaem and L. Lozano, Secretaria (March 1996). Leff to right: Webb Mufloz,
de Medio Ambiente, Recurnos Naturales y Emesto Rodriguez; Twan henders, Eke
Pesca, MBxico D.F., Mexico. f26Her, gun the^ Kiihler,

For logietic support for fieldwork I thank

Robert Dilger, Mauke Hoops, Siegfried
Kaetl, Mausi and Eddi Kiihl, Juctn Carlos
Martinez Skchez, Ivan Ortega, Roger
RomBn, and Francisco Serrano. For field
assistance, I thank Manuel E. Acevedo,
Martin Dblling, Jorge h r r a r i , Martin
Jansen, Elke Kijhler, JBrg Kreutz, James
Randy McCranie, Fernando Mendoza,
Kirsten Nicholean, Michael Obermeier,
Ardiel 2. Quintana, Antonio Ramirez
Vaaquez, Fabian Schmidt, Rabert Seipp,
Jorge Sob, Pablo Soto, and Jose Manuel -
Zolotoff:I am especially thanm to these Honduras (Owher 1998). Left to right: James
~ e o p l efor their valuable help. Many B ~ U - Randy McCranie, Elke Ktihler, Gunthsr
dents and associates of the department of Kijhler.
photos. Many of the excellent pen m d ink
drawings are the painstaking efhrts of
Milan Vesely. 1 I grateful to Mo&
Laudahn for technical help while process-
ing the collected specimens.
For the loan of ~peeimensor access fo the
collections under their care I thank Linda
Ford and Darrel R. Frost, American
Museum of Natural History IAMNHl, New
York City; Colin J. McCarthy, The Natural
History Museum (BMNH), Landan; Jenrs
Vindum, California Academy of Sciences
(GAS), San Francism; Alan h e t a r and
Harold Voria, Field Murseum of Natural
History (FMNH), Chicago; John E.
Simmons, Eli Greenbaurn, Linda TRleb,
and William E. Duellman, University of
Kansas Natural Hishry M u e m (KU),
Lawrence; David Kialrian and Kent
Beaman, Natural Hietory Museum of h s
Angeles County (LACM), LOB Angeles;
Douglae A. Rossman and Frank T,
Burbrink, Musetun of Natural Science,
Louisiana State University (LSUMZ),
Baton Rouge; Jose Rosado and James
Hanken, Museum of Comparative Zoology
(MCZ), Harvard, Aiain Dubois, Ivan heich
and h e m a r i e Ohler, Mu~eumNationid
d9Histoire Naturelle IMNHN), Paris;
Eunice E. Echwerria, Museo de Historia
Natural de El Salvador (MUHNES), San
Salvador; David B,Wake and Carla Cicem,
Museum of Vertebrate Zoology (MVZ),
Berkeley; fianz Tisdemann, Natur-
histarisches Mmem Wien (NMW), Wien;
Janalee Caldwell, Amy D. Eetep, and
Laurie J. Vitt, Sam Noble Oklahoma
Museum of Natural Hiatory (OMNH),
Norman; James R. Dixon and Katherine
Vaughan, Texas Cooperative Wildlife
Collection (TCWQ), College Station; F.
Wayne King and Kenneth Krysko, Florida
Mueeum of Natural History (UFI,
G a i n e d e ; Christopher A. Phillips, John
Petzing, and Steven D. Smka, Museum of
Natural History, University of Illind~lat
Urbana (UIMNH), Urban&; Ronald
Nmsbaum, Arnold Kluge, and Greg
Schneider, University of Michigan
Museum of Zoology (UMMZ), Michigan;
Ronald Crombie, 'Ronald Heyer and Rob&
Wlleon, Smitbonian h ~ t i h t i o nNational
,
Museum of Natural History (USNM),
Washingbn DC; Jonathan Campbell,
University of Texa0 at Arlington (UTA),
Arlington; Victor J. Hellebuyck (VH),San
Salvador; Wolfgang Btihme, Zoologisches
Forschwgsinstitut und Muaeum Alexan-
der Koenig [ZFMK), Bonn; and Rainer
Gfinther, Museum fiir Naturkunde der
Humbddt-Univeraitiit ru Berlin (ZMB),
Berlin; Frank Glaw and Dieter hchs,
Zwlogische Staatssammlung Miinchen
(ZSM), Munich.
I owe an immense debt of gratitude to AJ
Gutman, Jamea Randy McCranie, and
Lany David Wilson, who patiently sup-
ported this project and read the various
stages of the manuscript; the final manu-
rscript veraion was also read by Helmut
Diethert, Sflvia Mmchall, Udo Schweers,
Peter Stafford, and Harlan Walley; dl
individuals and pmvided valuable com-
ments and advice that significantly
improved the text. Manuel E. Acevedo,
Antonio Ramirez Vasquez, Jay M. Savage
and Hobart M. Smith have reviewed and
imprwed the distribution maps for the
species included in this book, I especially
wish to thank AJ Gutman who tramlated
the major part of the text fkom Geman to
English, James Randy McGranie, Charles
W. Myers, Jay M. Savage, and 'Carry David
Wilson generously shared their extensive
knowledge on Central American herpeto-
logy with me m d provided important
unpublished data,
Special thanks are due to my friend and
mentor,dlfred A, Schmidt, for his constant
support and encouragement.
Finally, I am especially grateful to my
beloved wife Elke KWer for her longtime
emotional support md inspiring encour-
agement, Elke often accompanied me in
t h e field and provided valuable a s s i b c e .
She &o merit6 special thanks for her very
careful editing of the manuscript, My in-
debtedness to her cannot be exaggerated,
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(1992k Tanlillo melcanoeephala
- Cat. Amer. h p h . b p t , 647: -
genus Ilmntilla. Herpetologica 2'I: 1160.
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WILSON,L.D. & D.E. HARN(1973): The hwpeta- -
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W%sa~,LAD,& J.R. McCaANn (1982): A new Additional ~pecimeesof the colubrid snake
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G1-m
Glossary
abdominal acute (plate)in turtles, a paired
scale located on the plastron betwaen the
pectoral scale and the femoral scale (Fig.
28c3
and e p w anal claw; in boids two claw-like
structure8 on either side of the cloaca,
which form the e x t d l y visible remnants
of rudimentary hind limbs
and male the hindmost pair of scales on the
plaatmn of turtle8 (Fig,2&); in lizards and
snakes, a aingle or divided scale carsring
the cloacal ope* (Fig.497)
apioal at the tip [apex)
arboreal living in trees
a d h a t i o n the promss of climatic drying
pods animals with jointed appendages
(insects+ spiders, crustaceans)
autoohthon evolved in a particular location
autotomy ability to drop body parts Ee.g. tail)
as a protective measure
adIlmy acute a small scale on either aide of a
turtle Bell between the bridge and the pec-
toral @cute(Fig.28c)
d a r y pocket tube-shaped pocket of skin in
the Bhoulder area (Fig.274)
b l locatd at the base
bmh foundation, a w i n g point
biotope habitat
bridge in turtles, the segment joining the
carapace and the plastron (see Fig. 23a)
canthd sealea on the canthua r o s t d s bet-
ween the eye aye the nagal opening (Figs.
72,345,349)
carapmce dorsal shell
carnivore meat-eater
caudal on the tail,pertaining to the tail
conepeui6~: of the same species
convex curved toward the outside
q t f o coloration coloration matching the
surrounding environment
distal further from the center of the body or
h m the point of reference
dorsal on the back, towards the back
dmolateral transition from the back to the
aide
endemic restricted in distribution to a parti-
cular location, limited distribution
femoral poree specifically arranged glandu-
lar opening8 on the undereide of the upper
thigh
femora4 male thigh scde; a paired ~ c a l one
the plastron between the abdominal scale
and the anal scale (Fig, 2&)
frontal (pl, frontds) frontal scale, scale(s1on
the top of the head between the eyes (Figs.
86,139,348,403,8913
frontomad (pl, frontonasah) scaled between
the rostral md prefkontals (Fie. 74,86,
96,139,346)
g~anuIarsmall, conical, non-overIapping (Fig.
276d1
guPar rhidd the fare& 5cde on the p k h ,
may be paired or unpaired (Fig. 28c)
habitat the phyaical and biotic characteristics
of an area in which plante or animals live
hemipnir (pl. Lmipenes) paired copuIatory
orgm in lizards and anakea
herbivore plant-eating
heterogeneuwa non-uniform (heterogeneous
ecalation see Fig.27%)
homogeneow similar, uniform
humeral wale a paired scale on the plastron
between the gular scale and the pectoral
scale (Fig. 284
imbricate overlapping (Fig, 276a,b,e)
houbation developmental phase of eggs,
brooding of eggs
infralabial in&dabid scale, scales on the
lower lip (Figs. 72a, 86,345,999,403,689)
Idramarghde scales on the bridge below
the marginal scales on a turtle shell
inguinal male a small scale present on both
sidee of a turtb shell between the bridge
and the femoral scale (Fig. 28c)
immtivore insect eater
intsrxalarlala 3ushed in ~cales",in iguanar,
the mall ~calesbetween the whorls of
enlarged spiky tail scales (Fig.315)
internasal intarnasal scde, scales between
the nasals (Figs. 843,403,442,691)
interparietal scale on the damal Burface of
the head between the parietals (Figs. 72,
88, 139,170,346)
interrictals scales counted horn am oorner of
the mouth across domum of head to the
corner of the mouth on the opposite side;
 this count includes the ultimate supralabial
an each aide
invertebratesanimals withaut a vertebrd
column
javenfb young animal
keeled with a raised ridge (Figs. 976a,b,
441c,df
labial pits in gome bids le.g,Comllu$
Epicrater) chara&ristjct tubular depressi-
on in the lip soalea; infrared sensing organa
lateral at or perkinhg to tha sida of the body
or a structure
loreal acale(s) between the canthala and
supralabiala inl i z - 4 (Figs, 7&, 86,346!;
scale(a1btween the nasal and pfeocular m
snakes (Figs. 403,689
lomal pit for pit viper% (subfamily crotalinae)
ch&r&christic,tubular deuressione between
the nmal ape&g and & ye (Fig. 689);
i&md sensing organ
lorilabial scales between the Ioreals and the
supralabiala (Fig.345)
marginal -tat e&ll males on the Lateral
edges of the carapace (Fig.28a,b)
metdian in the middle
mental chin scale, foremost, u s d y particu-
larly large male on the lowerjaw (Rg0,
ma, 345,399,403,404,689)
mental grmve extensible groove betwean the
pairs of enlarged &hin scale$ (Fig 403)
mrrnophyletic fbrraing a closed category der-
cended &om a common ancestor
monotypfo s@ a specks not divided into
subsp&ee
monotgpia genw a genus consisting of only a
aingle
- speeiea
.
marpholopic concerning the external form
muc:~onatebearing a psojecting spine on the
rear edge (Fig.276a)
&nose s d e , scale encirelig the noatril
(Rg.346,399)
nuchal d e neck sea14 the faremoat scale on
the carapace (Fig.28b)
d p i k a l scales at the back of the head behind
the parietals (Fig.86)
omnivore an animal that eats both plant and
animal food
oeteoderm supri5cial dermal banes under-
lying the epidem1 scales
o v d v i p m r m live bearing from eggs;
development of fertilized eggs in the womb,
with young hatching at or shortly afkr egg
laying
ocelli eye markin@, markings consisting of a
h ht ring with a dark center or a dark ring
J
t
h a lipht center
paravertebralof or pertaining to the area
immediately lateral ta the darral midline
parietal e e pineal eye, light sensitive w&an
below &e intapadetal e d e (Fig,392)
parietal parietal scale, s d e on the top oftbe
head posterior to level of eyes (next ta the
interparietal,if preeent) (Figs. 139,140,
141,380,403,691)
pectoral ecale thoracic d e ; a paired s d e
on the p b t r a n between the humera1 acale
and the abdominal scale (Fig.28c)
phylopenetio concerning evolutionary history
phydogmphy landscape featurea
plaatson ventral shell
-
p P e W swats rib scale, the large lateral scalm
of the plastron between the marginal scales
and the vertebral mates (Fig, 28a,b)
popul~~tioa a group of individuals of the same
~lpeciealiving in a particular area
poetanal scales posterior ta the cloacal alit; 2-
4 enlarged scalee are pmsent in t b area
in Borne lizard specie8 (Fig.278)
po6tfernoral poeket slitlike skin pocket at
the posterior insertion of the upper thigh
(Fig, 5471
postmental scales posterior to the mental
(Fi. 284,314,399)
poetnasal patnasal acale, scale between the
nasal opening and the 1oreaI or, in it8
abmace, the pnsocuIar (Figs, 403,6891
po6tocular Ipl. postocuh) postocular scale,
scale posterior to the aye (Figs, 86,403,
6891
po&orbital located postsriar to the eye
preavzioular (pl. pmauridam) scales
anterior to the external ear opening
pmftmrtal prefrontal d e ( s ) , lying anterior
to the fmntalr, (Figs. 96,139,346, S9%,403,
404,442,443,691)
pmaloacal p~res specie8 specific arrange-
ment of gland* ope* anterior to the
doacal slit (Fig. 3993
pmnad prenaeal scale, scale between the
rostra1 and the nasal opening (Figs. 403,
438,689)
p d a r scale(s)anterior to the eye (Figs.
86,346,403,439,6891
prorlmal situated near or toward the point of
reference
revision literaly ~t "new perspectivem(on a pro-
blem), comprehensive treatment of a taxo-
nomic p u p
roetrak rostral ecale, ugually particularly
enlesgdwl scale on the tip of the upper jaw
(A@. 72,73,74,86,139,345,346,399,41)3,-patria
404,689,891)
rudimentary diminiled, degenerated
s a k e (pl. B C U ~ S large
) flat scale
W a g g a t i a lifestyle partidy adapted to an
aquatic environment; most semiaquatic
species live at the edge of a body of water
a d spend part of the time in the water
d a r l d half dry
sefllal dichrurma;tisrn differential coloration
of males and ferndm in a species
subcaudal subcaudal scale, scales on the ven-
tral SW&CB of the tail; may be paired or
unpaired (Figs, 437.690)
eubdigitd lamellae widened rcalee on the
underside of the toes (Fige.M6d, 281)
subiaabrioateonly very slightly overlapping
(Fig.2788
mkuIsrr @ a l ebelow the eye (Figs.72a,
169b, 546)
aupediarg. scalels) above the eye at the
edge of the eye opening (Fige. '728, IS@, 282, ventrolateral h transition fmm the belly to
845)
srtpmand tubercle tubercular keels on the
dorsal scales abwe the lwal of the cloaca
(Fig. 682)
~upra-auricularscales acales above the
external ear opening; these may be enlarged
and spiky
uupreaurIcular a p h r enlarged spiky scales
above the external ear opening (Fig.871
aupnsEabial supralabial wale, upper lip scale
(Fig&.72% 86,345,399,403,689)
eupranarmal mlele) abwe the nasal (Fig &6)
mpmormlar supraocular scale, d e m the
d m a l aurface of the head above the eyes
(Figs. 139,548,405,691)
mpnaorbital aemfoirclee usually slightly
e d a g d scales on the dorsal surface of the
head arranged in a semicixcuktr pattern
bordering the inside of the e u p r a d m
(Fig.72b)
BVL anout-went length, distance from anout tip
to c l o a d opening:
co-mcuring in the same geographic
area
synonym two or more naaea far the same
taxon; in ~ystematiw,a ae~liorsynoap is
the sarlieet available name for a taxon; a
junior eponym ia any available name
other than the eenior sgnoeyrn and ia
invalid
sgptopic o&ng together in the w e loeation
tagonow study of the systematic claa~ifica-
tion of o r g d m s
temporal temporal scale, lateral head scales
in the temporal region behind the post-
oculars, between the supralabids and the
parietal (Figs.86,353,399,403)
tubercle a small, rounded scale or bump on
the skin scale
v e n M on the belly, towarda the belly; of Or
pertaining to the lower eurface of the body
or other structure
ventral ventral iadea, belly 4 e s
the lateral surface of the body or other
rtructure
vertebral d e central scale, scales running
along the midline of the plmtmn or body
(Figs. 28a,b)
vertebra&¶ &ds with a vertebral colomn
viadance event pmcesa in which a p r w i w -
ly uniform population is divided into two or
more aubpopulations; e,g. by the fomakion
of barrier&flooding or mountain upheaval
vivlpawua live-bearing
xemphytio suited to dry areas and climate
soogeomphy study of the diskibutien of
animalsl
Index
Bold number refer to figures Ipht&graphsand line drawings).
A - undulata gaigeai 156
Abrorrie 13,55, SBf
- amzuetoi 67,5@
- aurita 67,68,69
- bogerti 56,8T, 60
- campkili 56,67, IS
- ckizari 66
- fimbriata 56, I%, 67, E9
- frosti 67,W
- gaiopbnkama 67,&a
- kurolepis 57, I S
- iythmhilu 68,67,69, SO
- matudai 58,67,80
- rneledona 68,69
- montecriatoi 08,57,60,80
- ochoterenai 58,69
- ornslasi 68, I@
- ramirezi 56, Bg, 88,69
- salvedorensie 60, 60, BO
- srnithi 55,& 59
- uesconcelosii 58,5@
Aiklphicos 15,184,187f
- daryi 187,188
- ibumrum 187,186
- Iatifasciatus 187,188
- nigrihfus 187,187, I88
- quadrivirgatw 187,188,188
- weruepacis 187, 188
Agaiychnis callidryas 225
AgkBtrdon 14,893,292,298
- biiineatw as2, am,293
- biianeatus bilineatus 293
- bilineatus hwadgloydi 293
- btlineatw russeolus 293
Alligatnridae 12,
Anursiridium 18,184,188f
- aapperi 188,188, 189
- uelifemrn 188,189, 189
Amsrrlua E3,27, W, 164,164,
155f
- amiva 181,166,167,157
- ameiva pmestgnie 156
- chaitzami 186,167
- festiuu 154,1M, 155,156,157,
157
- festiua edurardsii 166
- festiua festtva 166
- festiva acci&ntaJis 156
- leptaphrys 156,166,157, 157
- gundrilineata lW, 156,156,
167,157
- undulata l M , 166,155,156,
167,167
- undulata brtwegi 166
- undulata rnbdis 156
- undulata pama 166
- undulata pulehm 156
- ~ndulatathomasi 158
- undulata undulata 156
Amphfabaeno 14,166f
- alba 8,166,167,187
- filiginosa 168,166,187, 167
- spurreEli 166,107,167
Amphisbaenia 14,166f
-la 13,25,54,80,81f
- oceilata 81,81,88,82
- vittata 81,82
Aaguidae 13,64,56E
h l i u 13,81,91,93f
- allisorti 04,88, 110,114
- aktenwtrarr 106
- biecutiger 102
- copei 98
-godmarti 102
- interrnediue 100
- nannodes 100
- ozyiophus 102
- procellaria 98
Anomalepididaa 14,168,169f
Anamalepin 14,169,169
- mexicanus 169,169, 170
AR'&lllger 13, BB, 69
- georgeemis 891,69
Atrachpis 291
Aspi&.weU~ 1&4,164,IbBf,
161,205,232
- anguatimps 159,159, 160
- angusticeps angwticeps 169
- angustimps petenensis 159
- cozumda 158,169,160
- cozumeh ccozuda 169,160
- cozumela mcmslini 169,159
- deppii 64,156,168,168,159,
160
-guttata 159,159,160
--mo6aguae 168,159,160,160
r d c k i 158,159,160
Aii.ao#uu 15,183,317
-clarki 189,190
- daldenensie 189,190
--
depressiocellus 189,190
hostiditmctua 189,190
- irnpelpctus 189,190
Adrnpoide~14,2911,292,294
- nummtfer S494,294
B
Buchio 13,80,81,82f
- blairi 82,83
-
pallidkeps 82,89,83
Baailfrcue 13,91,91,99,92,
m2zc 206
- bsilirrcus 122,188,123,
134,124
- galeritus ls8, 125, l M , 124
- plumifins 188,128, U4,
124
- uittatus 13S, 122,123, I N ,
124
Bipes 166
Bw 1 4 175f
-constrictor 176,175,176,
176
- constrictor imperator 176
-constrictor sebqgae 176
Boidae 14, 168,174f
Bolyeriidm 174
Bothdechie 14,88a, 292,
296f
- aurifir 296,397,298
- bkelor 8,295,236,297,298,
318
- la&n*lds 296, m,298
- marchi !Wi, 296,29&,298
- raigrouiridis a91,296,296,
B97,298
- n~twuiridismacdougalli 298
- rowleyl2QB,ae7,298
- schlegelii 11,286,296,290,
288,818
- supmciliarie 296,296,298
- thalaasinua mE, 296,296,
288
W h r o p r 14,=, 292,299,
317
- asper 192,280,W ,2W,
814,312,818
- puncbatus aP9,299
C
CaJmcur 12,2T, 28f
- cwodilus 28, Z8f, SO, SO, SZ
- yacare 30
CaMa 12,49f
- ceretb 48,49,51,83
CeEesfur 13,54,56,55,61f
- dtlawnsis %1,82
- bivitbtt.as61,B$62
- cycanoc.hlorie61,$2
- enneagmmmus 61,82
- hyykriw 61,61,B
- legnotus 61,82
- n s a b n u s 61,82
- arobiua 62
- ~ ~ e l l WI,
a e 6a
- acanmriua 11,62
Cenmic W
Cerrophtdlon 14,282,300
- godmani 8W,
- t~otzilorasna800,300
chcmhni, Ameiva
Chupinophb 15,184,180
- x a n t h h ~ h r180,190
Chelarria 12,48f
- agaasirdi 49
- 48,49,49,Bl,61,51,
B2,BB, 58
Ghalaniidae l2,47f
Chelydm 12,88,34f
- sewntiw 84,3Qf
- aerpenh'mocutimbricr 85
- serpentiw rossignonii 95
Chelydddrre 12,94
ChZmniue 16+18B,191f
- carnirrhds lB1,191,142
- carniatru ftlrvopdctur191
- exoletua lSl,191,192
-grandbzquarni~191,191,
1%2,192
CITES 30
Clatorddua 12,M,41
- a-utus dl, 41
CkEia 16,182,186,182f
- cWia MI,192,19B,kQ3
- equrston'ana 193
- sey@Lokina 188,193
climate 2 a
Chenaidaphorw 13,16,27,
la,lM,lfiB,161
- Iemniscatus 160,l&l,161
eOIeonys 15,87C09
- elegans 67,67,69,B8
- m i t r a m 67,67,68,68
CobpSyEkdn 13,68,63
- rhomblfir &5,&3
C o t u k 16,188,1SSf
- cons^)" 1m,193
- ~Owtrkfwmwcoe 194
Colubridae 168,181f
Colubrhae 15
Corolopiruwe 16,1%6,1W
316
- &a& 184,I!%, 197
- bip~cnetcabue184,194,195,
197
- *idens lad, 194,196,IBB,
197
--flssi&ns fiseidana196
ftsidene obgaletue 196
-
-flusidenspmct~an's196
impridla 194,lBB,1S6,1$7
- imperial& clauatur 195
- joanae lS6,196,1fl7
- meridanwr 195,196
-piceivi* 1B4,18S,lW, 146,
197
-- quinguevitf&ua 194,198,196
schmidti lS8,197
C o q h h 16,186,II,316
- l h t u 18b,181,188,1@8,
lBS,916
~
-
-
linaatug mn c&r 198
liwatrrar dunni 198
-puLchw 188
- uitdadue 198,198
-Camihe 14,175,176
~~ulatU.8178,176,117
- ruschenbergerEi IT&, 176,177
Ce@ophusea 13, W, 92,12Qf
- nristatua lZ4, I S , 186,128,
12rS
- h e m d z i i 1SB,126,128
-prcari~utuslaB, 196,126
Crieoewm 162
Wan#@pht$lb, 186,199
- neuermwvzi 1 W ,I-, lsB
C d y l i a B,25f
Cnmdylidas 12,30
Cmmdylrra 12,2Y
- aclahrs 17,S,28f,S9, SO,81,
98
- momktii 38,28f,
$1,88,4i3
htaEw 14,26,292,301
- durissus 301
- durisaus Wl, 301
~UF~IIMS
- durbsw irrifwwr 291
-Ctenolrotws
dtrrbsw tzabean 801,a01
13,81,91,9%f
- criakete1Jue 94, 915, 110,118,
11s
C t s m w u m 19,1d,@!4,92,
127% 274
- acanthus 127,f aS,128,
Isa, 132
- aljbdmhmidti 1Z8,lW, 181,
131
- bakeri 1'7,128,IS@,131
- Ifinsor128,1W,131
- fiavidorsalia128,12S,180,
130
- nzebnastema 157,1%8,la,
181
- oammna 137,128,lW,180,
131

- melandomus dehwnue 206
-pleei !206,206
D m m h o n 16,185,207
- cau&m[tcdatw 207
- cornis 207,243
- couperi 207
- cornis melanuma 807,207
- corais mbidus 207
- corak unicodor M ,907
D m I w 16,186,208f
-
-
chloroticw 808,208,SW,209
mar~arittfiru156,808,m8,
209
- margaridifems mcsrgaritifew
209
- margarififemsmaydis 2M)
- margafitifb~8O C C ~ & R & ~ ~
209
- nadanotmpb 808,208,209
- rhombifer808,209
E - dawnsi 215,218
&kdnowrut 13,80,81,83f
- horrida 83
- paImri 83,&4,84
-panurnelasis M,89,&4,&4
Elqhe J16,186,210
- flovirufu210,210
- flavirufaFvimfh:210
- flavirufarnahldai 210
- flavirufapadalina 210
- flavirufaphraemtrs 210
Elapidaa 14,168,281f
Eleuthemdactylltlr246,296
Emgdidae a,38
EawlfopMe 15,184,211
- ~clateriHi, 211
End3uu 16,184,211f
- bifovmtw 1&4,211,all, 212
- fravidorgues211,B U , 212
- flavitorpuesflavitarques211
- JZtcvifoquessurnlchmdi 211
- matanensis 211,B U , 212
EnyaNddea 13,92,138
- hstemlepis 138,133
Eoeene 27
Epkrabeo 14,175,178
- cer~~hldrs
176,178
E ~ e m A B l y 12
e Helimps
- inbriccsta 47,49f,Bl,Sa,08,
53
1FryOkmlamprue 18,183,213,
316,317
- b k u f~18,218,SIB,918
- rnirnus 818,213,818
- mimus impar %I3
- rnimus micrurus 213
Eublepharidaa 18,Bb;,87f
-- Index
- b m k i T1,78,72
- finatus 71,72,72
- ganotii 71,71,72,72
- mabasria 71,71,72,72
- tunicus 71,72,72,IS
H@rtm€arph~816,185,219
- clarki 219
- concotor 181,l&L,21@,219
If - dunni 219
-Gehym 19,68, 70
rnutilata 70,70
Bg,
Oekkonidae IS,%, 68f
I
I m n a 13,26,92,134
O w c h L n e 12,46f - igmlaa 184,134
-Wcarbanaria 88,46E,41 - iguana iguana I34
p h i r 15,182,216f - iguana rhinolopltra 91,154,
- bellw 215,srs,z1s
- bmehycephalus 811,215,218
la4
Iguanidae 13,55,9W
- ~ancrllatue216,218 Imrmt0d.e~15,181,la, 182,
- carinosue 216,217 a20f
- championi 216,218 - cenchors 22Q,220, 221,W2,
- ddmiani 216,316,218 222
-gemmistratus 220,aQ1, a28,
- dunni 216,217 222
- fulrogubtnt~216,217 - i?tOmtus881,221,m,222
-gdrnani 216,218 -phQntmma aBO,221,222
- hofianni 181,216,216,817, - inuissimus 881,221,222
B1%, 218
- irnmoeuhtus216,817,217 K
- Iaticiraetus216,218 Kinosternidae 12,Qlf
- nasalia fie,216,217 Kinasteman 12,41f,48
- rhalogasker 216,217 - acutum 43,4& 44,44
- rwthveni 226,218 - angwtipans 43,44
- blomanme 216,218 - maaeri 4P1,4Z,A3,44,44
- z e f i w i 210,218 - teucoakomurn 844% 42,44,
&~~homha W, 6 6 , e 44
- lioaephdue 86,66 - teuco8tomurn lruc~stomum
--Gonu&&# 13,09,69,70
albogularis 10,713
43
- leum~tanaumpostinguinde
dbogularisfuscua 71 43
- annudcaria 70 - mrpbides 43,43,44,M
Gymnophthalmidae 13,54,66, - mmpioi&~ arbaxillan?43
80f - scqiokfee albogulare 43
laynmqpktkalmvs 13,BO,81, - qioides cruenfatwn 43
84f
- apeclo8us 26,84, $4, &8,86
H
habitata 16f
HelmintkopkSs 6,I@, 169
- frOntaEi8170,170
R~l&mm 13,89f
- horridurn 89,BO
-
-
horridum alvareei 89, SO
hommdumclwmrksbogrrtdd 89,90
- horridurn horridurn B!3,80
HeLodematidae 13,5A,89f
RemkhcbEw 13,68,89,71

- sanctuemartcre TS,73
- xnnthopaigm 75,73
Lepidoolrelyu 12,Mf
- kempi 60, SO, 51,68,53
- divama 48,60,61,18,53
L&pWrrrtyIru~13,68,89,74
- ZU@%bnS 14,?4
Lepidcphynrcr 13,162f
- chicoaeers~163
- flavimcuEatum 18B, 162,
183,163,1&4,1M
- lipebzi 163,164
- rnayae 163,184,164
- mticulutum 163,164
- gmithii 162, la,163
- W l a e 168
Eepwotnra 13,80,81,85f
- rugbps 8LC,88, BB, 1
- mudhi BQ, 86, B8, €16
Lsptmhima 16,188,225t 228
- m u l a t u 226,226, a87
- annulata cuselldrie 226
- mnnulat~rhembifira 228
- frenata
317
a84,SS8,226,228,
- b n a t a nollebi 2%
- fimtta Y W & W ~ S ~ %226
-- nigmf08cciata M,W6
rubrkata 9215,226
- aeptentrionnlis a!&&,828, E37
- eepGantrienalis orrtata 288
- saptentrionalis poIystisda 22-28
Leptodrpw 15,186,227
- nubherrimus 181.327.227
M - nigmtnctus divarkutuer 283
Mttb:uya 13,64,147,147,149
- unimarginuta 64,147,149,
149
Rlacm~towta14,174f
Meno&pin lli, 185,232
-putnami 28% 232
M&pBte 15,186,252
- mentouariue 156,181,232,
22B
- ment6varius centratir,232
- mnbouariua mentouariulp
292
Mwtigdryas 206
- &nkli 208
Me+& 13,66,66
- montbla H , 6 6
- mreletii W, 66
M e s o w i n c ~147,160
~~
- mamugwe 64,141,148,180,
160
- whwartzei 147,148,185,1%1
Micrum*~ 316,317
Mbrrcrua 14,243,281c 316,
317
- alkni 282, aBQ,286, a88
- a ~ o m t i 282,286,
s
- a m a d k j ~ n 282
i
- boge& 282, S 8 , 2 8 8
- browni 2&a, 9W, 288,288
- bmwni bmwni 282
- bmwni iraprhLnu8 282
- clarki 282,264,226
- d k s t e m 281,282,%4,288,
%88,288
- diastem q l ~ a p 282e
- dimema alienus 282
- diadltenaa cxpkztua 282
- d i d e m a macdougalii 282
- dhterna sappnri 282
- d h s t e m browni 282
- dissoleucus 282,288, iB3
- dkmleucus dunni 282
- etegaw 282, SM,2 8 6 , m
- elegans vewpacl282
- kippocrepis 282,2238,288,2SbI
318
- k t i f w i a t w 282,288,288
- mipartaitus 282,2841,380
- muEtifasrciatue 289, W ,
2&8
- rnulfifasciatusherhuigii 289
- mtdkif~sciatusmdbij%s&trsa
283
- nigrocinctus 283, W ,888,286.
387, 2%
- nigminc6us babospul285
- nigrocinctus coibeneis 283
- nigrocinctua masquiknsis 285
- nigrocinctus nigraeinctur 283
- nigmcinctua eunilensis 285
- rwtantas 283,286
- atewarti 283, PW,284 288
- stwrti 283,288
Miocene 26,P
M o ~ p ~ 13,25,98,92,
~ a ~ r
138
- annularis @a,138
- groi 186,196

- altae 93,96,121
- advamz&Etomi 96,87,110,
117
- ~ ~ ~ Q U C ~ M 94,96,97,
O B U B
110,118
- animlepis B4,9B, 97, 110,119
- aquaticwr 93,1)6,%7,117,117
- alsmtw 96,9+7,120
- barkeri 98,96,67,110,115
- bicaorum 96, W ,110,118
- binotatus 94
- b i p ~ m t u B3,96,87,
s 110,
114,115
- capifo 93,B8, a,110,115,116
- carpent& Q8,BB, 110,121
- cobanensis 84,98, W, 110,11$
- compres~icauda98, gB,110,
115
- concolor 98
- cmrraulw 98, W, 111,119
- cristifer I,98, l l a , 118
- cupmus 98, H, 111,114,120
- clsprinus 98,220
- c~asuco98, SQ, 119
- doll*ianue 98,88, 1X1,120
- w u t 93,9B, 121
- firtutrensis 100,121
- fingosus 100,108,117
- fuscoaumtu8 98,fW,100,111,
l a1
- haguei 94,100,119
- heteropkolidutus 94,100,101,
111,119
- hobartemithi 94,100,120
- humilk 93,100,1Q1,111,114,
115,118
-johnmeyeri 94,100,101,111,
119
- benaptonlai 93,100,101,121
- krertn' 100,119
- laeviuentris 84,100,101, I l l
- kwrinua 100,101,111,118
- Eimifrone 93,104 101,114,
114,121
- Cionotus 93,101,101,1lB, 120
- lowridpi 102,108,1l.5,116
- naacpophllw 102,1WI,118,
114,lao
- rnatudai 102,120
- muralk 94,102,119
- oceltoecapularis 102, lMl 121
- pachypzcrr 102,121
- pandoensis SS,102,121
- parvicireulatus 102,119
- pentuption 102,208, l B , 116,
l l g , 118
- pe-ii 98,102,108,118,114,
116
- p ~ o b m i s94,102, LO#, 114,
119
-poecilopus 98,108, 104, 113,
- 120
polyhph 93,108,104,118,
116,117,121
-purpurguldria 94,104,1O.i,
119
--p y m u e 104,121
--quaggulus 104,105,112,116
nrtamnsk 104,108, 118
rodriguexii 99,104,10&,118,
121
--
rubribrarbaTis Q4,104,118
sagmi 104,105,118,116,11%
--
mAceus 104,105, llB, 120
-
semnoi 104,105,118, 118
sminth-us 94,104,108,119
- tawmendi 104,180
- tropirhgaster 106,106,216,
120
- topidalepis 106,107,120
- tropMonotw 106,107, 118,
114,116
- uniformh 93,106, lOT, 118,
-
116
ntilensb 17,106,1Q7,118,
114,116,117
- wampuensis 108,116
- villai 106,120
- vittfgerue 94,106,107,118
- vuclfemns 106,117
- warmudhi 94,1P6,107,118,
114,117,119
- woodi 106,107, 113,llB
- yoroensis 106,117,121
- zetcs 93,106,113,121
Noflropsaia 1&,182,238
- rugoaua 1831, BSS,238
Index
-platurus 8BB, 289, %90,290
Phimophle 16,182,242
-grc.ianensia U2,242
P h r y n o w shmphirei 246
PhFyno~onro13,91,137
- ww 181,137
PhylEodactyZus 13,68, W,
74f
- insularis 74,76,76
- palmeus 74,71,75
- tube~.cullosus74,76,76
Pihrophh 16,18BY242
- Idneaticollk 242,242, W
- liwaticollis gibboni 242
Pleitrtooene 26
Plfoeereus 16,l83,249f, 264,
315,516,317
- edapoas 248,244,916
- elapaides aequalie 244
- elapodes andrewais 244
- elapoides diostema 244
-eurytonus 66,M, a44,244,
318
- eurJ"zonusburghardti 244
- wryzonus dimidiatue 244
Pliocene 25
Polychms 19,Ba, 92,137f
- gwtturoeue 137,ISB
Porthidirun 14,292,503f
--
dunrai 8Q4,304,304,306
Reapre 303
- hnarbergii 8Q4,304,805
- msutum 303,304, WS, 305
- ophryomegaa 803,804,304,
305
- valmnieum 304,306
- yucabanicurn 304,806,305
Ph-lus la, 8 4 81,
87
- verkbrulta 87,87
P~s&boa 15,188,185,248
- neuwiodii 244, !U5,246
PssuaSen 15,186,245
-poecil~notua846,245, a46
RychogLarnus 13,80,81,88f
- festru, 88,88,gS,89
- myersi 88,88,89
-plicatus 88,88,8&, 89
qrthonidae 174
- dsmradra 247,848,260,968
- gortmni 247,26O,Z61
- hannsteini 247,252
- hempsdeudae 247,260
- kanalchutchan 247, !449,269
- kinkelki 247,261,242,252
- Zrachrymms 248, M l , 262
- macdougalli 248,260
- Iratedtrigu 245
- mantecristi 248,260,2&4
-pilonaorun 248, ?401,282
- psadaai 248.282
-pulveriventrk 248,849,260
- mgerrornand 247,248,260,
261
- sargenti 248,250
- serperQster 249,250,5461
- atadelmant' 846,247,249,
250
- toipanorum 249,260
- vermiculaliceps 249, a0
RMnobothqwa 15,186,262f
- bsuallii 188,21,252, a85,
263,819
RhbCkn~n~ u,
8 96f
- annulota 3%,87,38
- a m h t a 36,81,38
- frttaema SiJ, 36,38
- mlanoaterw 56, $8
-puZckndnac.~36,87,88,38
- pulchena'naa incba 36
-pulcherrim m n n i 36
- pulckrrirna pulchewima 36
- rrcbida Bll,a6,38
Ehinoleptus 171
- lundedli gaigme 140, 141
- EundelZi Iundelli 140
- m t a c h i ~ u 138,139,140,
s
141,146,145
- melanarhinlts 140, 141,145
- melanorhinus melanorhinus
140
- melanorhinus stuardi 140
- aerrifir 138,139,140,141,
144
- wrrs'fer preaygudl 140,
- awrifir serrifir 240,
-ainifirw 140,142,144,146
- a~na;rcsgdinw188,199,142,
142,146
- srnithi 142,142,144,146
- SQUat?t5SUS 139,142, 1.18,
144,146
- treniocftemb 199,142,146
- bapenais 142,143,184
- variablilia 142, lM, 144,146
- uariablilia otEopoms 144
- uariablilia vrariabtilis 144
Soineella 13,147,147,151
-gemrnfngeri 161,151
Scincidae 13,64,147f
Scolecopbidia 14,169f
S o o h p h l e 16,256
- a ~ ~ n c 189,183,855,
tw T
266, B18 TantilEa 15,184,246,264f
Scolecophidia 169f - albiceps 264,266
&nHooLb 16,188,166,266 - altkola 264,2%5,266
- t r b s p k m,255,f66,2B6 - armilkata 264,266,267
Serpentas 268f - bairdi 264,288
8 h m 16,182,183,266f,317 - brevimuih 264,268, 9gI)
- annulatus 266, S 7 , m. - cuesta 284
288, S B - cuniculatw 264,268
- anfhrucops 268,258,269 - escubitor 270
-aqua 257,268 - i m p m 264,817,268
- carri 287, m,258 -jani 264,268
- dimiolia$ua 21i6,257,288, - bmpim 264,266
258, % 3 1 - mhnorephlatcm 284,266
- langlfru!nie 267,268 - m s t a 265,266,287
- nebulatia W , 267,258,268, - reticulato 266, am, a68
- rubm 286,266,889
- llebulatus LucomeJaa 261 - rujicepa 266,266
- nebulatw nebukatus 267 - schiratoaa 181,266,268,208
- m n i o l a 261,268, NO - mpracinefa 266,887,260,388
- 8annioia mranniola 267 - taeniata 286,267, %g% 268
- mnnida &Eli 267 - t a y m 2@6,268
81Epkhphk 16,182,260 - tecta 26%, 268
- ceruima 880,260 - triaeriata 266,268
- blagimudatus 260 - fribaenbta 286,268,268
S p h a s r d w & b e 13,68,169, - vemiforaia 266,266, M7
T6f - vukani 266,268
- argue 11, Tg, 79 TantUlUu 15,184,269
- dunni 78,77,78 - bravi&sim 269
- glaucus W , T 7,79 -cant& 269
- --
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ISBN 3-936180-02-4