A Key To The Tadpoles of The Continental

Herpetologists' League
A Key to the Tadpoles of the Continental United States and Canada

Author(s): Ronald Altig
Reviewed work(s):
Source: Herpetologica, Vol. 26, No. 2 (Jun., 1970), pp. 180-207
Published by: Herpetologists' League
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180 HERPETOLOGICA Vol. 26, No. 2
. 1969. Evolutionary relationships and osteology of the frog
family Leptodactylidae. Unpubl. Ph.D. Diss., Univ. Kansas. 800 p.
SMITH, H. M., AND E. H. TAYLOR. 1948. An annotated checklist and key to
the Amphibia of Mexico. Bull. U. S. Nat. Mus. 194:1-118.
1966. Herpetology of Mexico/annotated checklists and keys to
the amphibians and reptiles/Reprints of Bulletins 187, 194, 199' of United
States National Museum with a list of subsequent taxonomic innovations.
Eric Lundberg.
TAYLOR, E. H. 1941. Some Mexican frogs. Proc. Biol. Soc. Washington
54:87-94.
Received: 24 October 1969
Accepted: 10 December 19,69
Department of Zoology, University of Nebraska, Lincoln, Ne-
braska 68508
A KEY TO THE TADPOLES OF THE CONTINENTAL

UNITED STATES AND CANADA
RONALD ALTIG
ABSTRACT: A key to the tadpoles of the continental United States and

Canada in developmental stages 25 through 40 of Gosner is presented. A
standard terminology is suggested, and methods of preservation and storage
are discussed. Generic synopses and a bibliography concerning identification
of tadpoles are included.
ADULT anurans of the United States and Canada are relatively

well known, yet studies of tadpoles have lagged behind due pri-
marily to the difficulty of correct identification. Of the 72 species
of tadpoles in the United States and Canada, only the more striking
species can be readily identified. Thus, tadpoles are seldom collected
and are generally excluded from research programs; museum mate-
rial rarely receives attention.
Orton (1952) presented a useful key to the genera of tadpoles
in the United States and Canada. Various other keys covered the
species of a restricted area (e.g., Smith, 1934; Walker, 1946; Bragg
et al., 1950), but the keys by Wright (1929), and subsequently by
Wright and Wright (1949), are the only attempts to cover all species
in the United States and Canada. These keys are of limited value
because of the lack of material and the annotated style. Papers by
various authors, especially Gosner and Orton, added necessary
descriptions and other information.
The primary purpose of the following key is to provide a rela-
tively simple means of identifying preserved tadpoles of the United
HERPETOLOGICA 26:180-207. June, 1970

1970 TADPOLES: U. S. AND CANADA 181
TL
BL
MH
SP TH
FIC. 1.-Left lateral view of a typical tadpole. BL, body length; MH,
musculatureheight; SP, spiracle, TH, tail height; TL, total length.
Statesand Canadain developmentalstages 25 (operculumclosure)

through 40 (immediately before front leg eruption) of Gosner
(1960). Gosner'ssystem has been used widely and the use of a
commonstaging system is imperative to detailed comparisons. In
practice,the limits of this developmentalrange will vary (especially
at younger stages) due to characteristicsof the particularspecies.
Specimens near the middle of this range, with mouthparts and
color patternsfully developed, should be available for best results.
Locality data are required for the separation of certain closely
related species that cannot otherwise be separated satisfactorily;
geographiccouplets are used to shorten and simplify the key.
It seems unlikelythat a key can be written to hatchlingtadpoles
with undevelopedmouthparts,althoughwithin a limited area certain
taxa can be identified by color, size, adhesive gland conformation
and ecology (Gosner and Black, 1957); a key to eggs (Livezey and
Wright, 1947; Wright and Wright, 1949) is useful in this regard.
Specimensolder than S-40 often can be identified with a key to
adults. By the correlation of preserved with live specimens, one
can soon learn to sight identify most tadpoles in an area.
A secondarypurpose of this key is to suggest a standardtermi-
nology (see figuresand glossaryfor suggestedtermsand synonyms).
In contrast to the consecutive anterior to posterior num.beringof
Braggand Bragg (1959) and the peripheryto center numberingon
each labiumof Boulenger(1891), the illustratedmethod of number-
ing tooth rows approximatesthe sequence of appearance of the
rows and facilitates interspecific comparisonsregardless of labial
tooth row formulas.
A fractiondesignatesthe number and position of rows of labial
teeth; the numeratorindicates the number of rows on the anterior
(A.) labium,and the denominatorindicatesthe rows on the posterior

A-2 gap
Al~~~~U
p
57
P1 ,,,::::4:,,.P21
Od
FIG.2.-Schematic drawingof tadpolemouthparts.The left side is emar-
ginate,the right not emarginate.Al, anteriorlabium;A-1, first anteriortooth
row;A-2, secondanteriortoothrow;Lj, lowerjaw;Lp, lateralprocessof upper
jaw; Mo, mouth;Mp, marginalpapilla;Od, oral disc; P1, posteriorlabium;
P-1,firstposteriortoothrow;P-2, secondposteriortoothrow;P-3, thirdposterior
toothrow; Sp, submarginal papilla;Uj, upperjaw.
(P) labium. Letter abbreviations plus row numbers (e.g., A-1, P-3)
reduce confusion. The fraction is written in line with the rows with
median gaps in parentheses. A range in the number of rows is
hyphenated, and variability in the presence of a median gap is
indicated by brackets. For exampile, the formula 2(2)/3-4[1] indi-
cates a tadpole with two rows on the anterior labium, the second
with a median gap, and three or four rows on the posterior labium,
the first with or without a median gap. On the anterior labium,
typically the more central rows vary in occurrence, whereas on the
posterior labium the more peripheral rows usually vary in occur-
rence. This generality, especially concerning the posterior labium,
does not always apply to tadpoles with a tooth row formula of %/4
or larger. Extraneous teeth may occur anywhere on the labia.
Although a formula for the actual number of teeth per row
(Noble and Noble, 1923) is essentially useless because of ontoge-
netic and individual variation in row length, the number of teeth
per unit length suggests size and spacing of labial teeth. Gosner
(1959) presented taxonomic characters of individual teeth.
Characteristics of pigmentation are limited primarily to melanic
patterns that persist in preservative, although live color patterns that
disappear soon in preservative are mentioned if important. The
distinction of dermal and subdermal pigmentation is useful because

differentpartsof the color patternmay be formedfrom one or both

layers.
Absolute measurementsare nearly useless as taxonomiccharac-
ters because so many factors affect size of tadpoles. Proportional
measurementsoften follow predictablepatterns (Gosner and Black,
1957; Limbaughand Volpe, 1957) and sometimesare used.
Method of preservationand storage of tadpoles are important,
for they can drastically affect color patterns, measurementsand
configuration. Specimens should be preserved immediatelyin for-
malin mixed to be 10% after dilution due to the tadpoles. Plant
material should be excluded, and the specimens should not be
crowded. Specimenskept alive for even a short time often become
damaged, and the keratinized mouthparts are often shed. After
24 hours the fluid should be changed to fresh neutral 10%formalin
for storage. Tags should not be tied to specimens,even if legs are
present.
Some people store tadpoles in ethyl or isopropyl alcohol. Al-
though alcohol renders the non-keratinizedoral structures more
easily visible, tadpoles in alcohol often become soft and distorted.
Pigmentpatternsare alteredmore than in formalin,since iridophore
patternsare lost sooner and melanophorepatternsare lightened or
lost. Possibly new preservatives that preserve like formalin but
have a more agreeable odor will be a compromise. Antioxidants
aid in color preservation. Specimensdistortedby alcohol dehydra-
tion often can be salvaged by placing them in water overnight,
transferringto 20%formalinfor 2-3 hours and then storing in 10%
neutral formalin.
Notes on color in life and ecological notationsare importantand
likely will become more so. Besides the usual collection data, the
time, amount of shade, water turbidity,bottom color, and amount
and type of aquatic vegetation should be recorded since some of
these factors may affect pattern and color intensity.
Several hylid genera which cannot be separated satisfactorily
and the morphotypes of Spea associated with cannibalism key
directly to species. Pertinent range limitationsare mentioned, but
range maps in Conant (1958), Stebbins (1966) and the Catalogue
of AmericanAmphibiansand Reptiles (American Society of Ich-
thyologists and Herpetologists) will prove useful. Introduced
species were excluded, except for Ranacatesbeianain the western
states and Bufomarinusin southernFlorida.
Phenotypicplasticitymakes tadpoles difficult organismsto cate-
gorize into a key. There is usually considerable individual, geo-
graphic and ontogenetic variation,while social and environmental
factors often impress further alterations. Anomalies of oral disc
componentsare commonand laboratoryspecimensoften show addi-
tional variationsof color patterns, mouthparts,size and conforma-

tion. In any group there are individuals that will not key correctly,
and tadpoles will certainly be no exception; subjectivity cannot be
avoided. Bufo and Pseudacris are the most difficult.
Specimens of all but 6 species were examined; specimens of 40
species were seen alive. Generic synopses and a bibliography con-
cerning identification of tadpoles in the United States and Canada
follow the key.
ACKNOWLEDGMENTS
I thank the following persons and institutions for the loan or
donation of specimens: R. G. Zweifel, American Museum of Natural
History; R. H. Mount and G. W. Folkerts, Auburn University; A. E.
Sanders, Charleston Museum; J. H. Harrison, College of Charleston;
F. H. Pough, Cornell University; E. D. Brodie, Jr., Clemson Univer-
sity; H. Marx, Field Museum of Natural History; H. M. Stevenson,
Florida State University; P. W. Smith, Illinois Natural History
Survey; C. E. Nelson, Indiana University; J. W. Wright, Los Angeles
County Museum of Natural History; R. C. Stebbins, Museum of
Vertebrate Zoology; G. R. Zug, United States National Museum;
K. L. Gosner, Newark Museum; R. S. Funk, Northern Arizona Uni-
versity; R. M. Storm, Oregon State University; J. W. Gibbons,
Savannah River Ecology Laboratory; R. A. Brandon, Southern
Illinois University; C. J. May, Tucson, Arizona; H. A. Dundee,
Tulane University; H. Harima, University of Alabama; T. P. Maslin
and H. M. Smith, University of Colorado; K. R. Porter, University
of Denver; W. J. Auffenberg, University of Florida; C. F. Walker,
University of Michigan; D. E. Metter, University of Missouri; J. H.
Black, University of Oklahoma; C. R. Shoop, University of Rhode
Island; D. B. Ralin, University of Texas; J. M. Legler, University of
Utah; S. Hedeen, Xavier University.
Numerous specimens were collected while the author was a
visiting summer faculty member at the Savannah River Ecology
Laboratory under Division of Nuclear Education and Training funds
[AT(38-1)-310] to the University of Georgia.
Also, I thank E. D. Brodie, Jr., G. H. Clemmer and J. L. Wolfe
for reading parts of the manuscript. Dianne M. Brodie prepared
the figures and Waanda B. Lee typed the manuscript.
GLOSSARY
A-1-an abbreviation for first anteriortooth row; A (anterior)and a number
designatea row on the anteriorlabium. (= upperfringe, outer row on
upperlabium,first upperor anteriorrow.)
A-2 gap-median gap in row A-2. (= medianinterval,medianspace.)
A-2 gap ratio-the A-2 gap ratioequalsthe lengthof one sectionof A-2/width
of the gap betweenthe two sections,regardlesswhichis larger. A number
of 1 or largerindicatesa gap that is equal to or smallerthan the lateral
sections,and a numbersmallerthan1 indicatesa gap widerthanthe lateral
sections. Used in contrastto Wrightand Wright'smethodof alternately
dividingor multiplyingdependingon the widthof the gap.
Amphigyrinid-condition of having dual, lateralspiracles.

Anal flap-small flap present in Ascaphus tadpoles, ventral to the anus.

Biserial-describes a tooth ridge bearing two rows of labial teeth, as in Ascaphus.
Body length-see Fig. 1. (= head-body length.)
Body ratio-total length/body length.
Emarginate-refers to an indentation (lateral in U. S. tadpoles) in the oral disc
margin, as in Rawa (Fig. 2). (= lips infolded, indented laterally.)
Fin-membranous flap along the dorsal and ventral margins of the tail mus-
culature, used for propulsion; dorsal and ventral fins. (= crests, caudal
crests.)
Flagellum-an attenuate posterior portion of the tail capable of undulating
independently of the remainder of the tail, sometimes not pigmented;
present in some hylid tadpoles.
Infralabial rows-(= posterior tooth rows).
Internarial distance-transverse distance between the medial borders of the
external nares.
Interorbital distance-transverse distance between the medial margins of the
pupils. (= interocular distance, interpupillary distance.)
Jaw-narrow-heavily keratinized for less than half the width (longitudinal
dimension) of the jaw.
medium-heavily keratinizedfor about half the width of the jaw.
wide-heavily keratinized for nearly entire width of jaw.
upper-anterior. (= maxilla, beak, mandible, homy beak, labial mandible,
horny jaw. )
lower-posterior. ( mandible, beak, horny beak, labial mandible, horny
jaw, maxilla.)
Labial flap-membranous flap suspended in front of the mouth of microhylid
tadpoles, may or may not bear papillae, but labial teeth are absent.
Labial teeth-keratinized structuresoccurringin transverserows on each labium,
used for scraping food from substrate, not true teeth. A group of replace-
ment teeth (series) occurs above each visible tooth; denticulate or not,
depending on taxon. (= denticles, dermal denticles; labial denticles.)
Labium-half of the oral disc. An anterior and posterior labium, limited by
the lateral emarginationor a similar point, form the oral disc. Each labium
supports labial teeth and papillae. [= lip (part)] See lip and labial flap.
Laevogyrinid-condition of having a single, lateral spiracle on the left side.
Lateral line pores-sites of pressuresensitive neuromasts. See muciferous crypts.
Lateral process-a poorly delimited posterolateral extension of the upper jaw,
often non-serrate;short, extending to or slightly beyond lower jaw; medium,
extending definitely beyond the lower jaw; long, extending well beyond the
lower jaw.
Lip-freshly folds around the mouth of a tadpole, used in contrast to labia
since they support no labial teeth or labial flaps since they are not pendant.
See labium and labial flap.
Longitudinal axis-horizontal axis projecting from the tip of the snout to the
tip of the tail; used in contrast to lateral axis. (= body axis.)
Marginalpapillae-papillae attached to the margin of the oral disc, collectively
forming the papillary border. ( = dermal papillae.)
Mediogyrinid-condition of having a single, medioventral spiracle.
Metamorphic-stage of development between the time the front legs erupt and
metamorphosisis complete.
Muciferous crypts-lateral line pores (part); gland openings (part).
Musculature height-vertical height of tail musculature measured at the base
(Fig. 1) unless stated otherwise.
Oral disc-collective term for combination of anterior and posterior labia sup-
porting labial teeth and papillae. [= mouth disc, oral pad, oral apparatus
(or this may include jaws), mouth pad.]

P-1-abbreviationfor first posteriortooth row; P (posterior)and a number

designatea row on the posteriorlabium. (= innerrow on lower labium,
first posterioror lowertooth row.)
Papillaryborder-collective term for marginalpapillaesurroundingthe oral
disc, may be complete,with a dorsalgap or with dorsaland ventralgaps.
(= oralpapillae,papillaryfringe,labialpapillae,oralfringe,labialfringe.)
Premetamorphic-stage of developmentbetweenthe time the operculumcloses
and the rearlimb buds appear.
Prometamorphic-stageof developmentbetweenthe time the rearlimb buds
appearand the front legs erupt.
Sinistral-conditionof having a single spiraclesituatedon the left side. (
laevogyrinid.)
Spiracle-externalopeningfor the exit of respiratorywater.
Submarginal papillae-papillaeattachedto the oraldisc apartfromthe margin
and marginalpapillae. ( = innerpapillae.)
Supralabialrows-(= anteriortooth rows).
Tail height-Fig. 1. (= tail depth.)
Tail length-Fig. 1.
Total length-Fig. 1.
Uniserial-refersto a tooth ridge bearingone row of labial teeth, as in most
tadpoles.
Upperfringe-( = A-1).
KEY
Section A: Key to families of tadpoles plus genera and species of
Rhinophrynidae, Microhylidae, Ascaphidae, Pelobatidae and
Leptodactylidae.
1. Jaws without keratinized sheaths; oral disc and labial teeth
absent - --------------------------------------------------------------- 2
Jaws with keratinized sheaths; oral disc and labial teeth
present ----------------------------------------------------------------------5
2. Spiracles dual and lateral (amphigyrinid); oral barbels
present; upper lip without a median notch; southern
Texas -Rhinophrynidae, Rhinophrynus dorsalis
Spiracle single and ventromedial (mediogyrinid); oral bar-
bels absent; labial flaps with a median notch; east of
Continental Divide and extreme southcentral Arizona
------------------------------------ Microhylidae-3
3. Edge of labial flaps scalloped or with distinct papillae;
medial margins of labial flaps convergent; extreme south-
ern Texas-Hypopachus cuneus
Edge of labial flaps smooth; medial margins of labial flaps
parallel or divergent -Gastrophryne-4
4. Dorsum grayish-brown; tail stripe indistinct to absent;
underside of labial flap sometimes with small black ex-
crescences; total length/body length 2.0 or less; light
ventral markings small; west of Mississippi River
-------------------------------------Gastrophryne olivacea
Dorsum dark brown to black; tail stripe usually distinct;
underside of labial flap without excrescences; total
length/body length 2.1 or more; light ventral markings

*
*/ 4fX\
FIG. 3.-Eye and anus positions: (A) Hyki versicolor, lateral eyes, (B)
Scaphsopus holbrooki dorsal eyes, (C) Leptodactylus labwlis, medial anus,
( D) Rana catesb na, dextral anus.
generally large; primarily east of Mississip!pi River

-~~~~~~~~Gastrophryne carolinensis
5. Spiracle ventromedial (mediogyrinid), nearer mouth than
anus; mouth large and ventral; labial tooth row formula
2-3/10-12(1); some tooth rows biserial; mountain streams

of northwestern United States and adjacent Canada

--------------------------------Ascaphidae, Ascaphus trutei
Spiracle on left side (sinistral, laevogyrinid); labial tooth
row formula 7/6 or less, commonly 2/3; all tooth rows
uniserial; widespread- 6
6. Anus medial (Fig. 3C); eyes dorsal (Fig. 3B) -7
Anus dextral (Fig. 3D); eyes dorsal or lateral (Fig. 3A) 12
7. Papillary border with a wide dorsal gap about equal to A-1
and a ventral gap equal to or larger than P-3; oral disc
emargina.te; labial tooth row formula 1-2(2) /3[1]
-------------------------------Bufonidae (part )-Section B
Papillary border without a ventral gap, dorsal gap present
or not; oral disc not emarginate; labial tooth row formula
2/3 or more - 8
8. Tooth row formula 2/4 or more; papillary border complete
or with a narrow dorsal gap; spiracle well below longi-
tudinal axis; body somewhat depressed; darkly or lightly
pigmented; widespread -Pelobatidae-9
Tooth row formula 2/3; papillary border with wide dorsal
gap; spiracle at or near longitudinal axis; body not de-
pressed; darkly pigmented; extreme southern Texas
--------------Leptodactylidae (part), Leptodactylus labialis
9. Jaws narrow to medium, never cuspate; lower jaw striated;
keratinized area on roof of mouth absent; dorsum usually
dark brown to black; body typically wider posteriorly
than anteriorly; to 35 mm total length; eastern and south-
western states -Scaphiopus-10
Jaws wide, frequently cuspate; lower jaw not striated; fre-
quently a small keratinized area on roof of mouth; dorsum
typically lightly pigmented and somewhat transparent;
body often depressed and wider anteriorly than poste-
riorly; to 75 mm total length; widespread from Missouri
westward -Spea-11
10. Jaws medium; labial tooth row formula 2-5(3-5)/4-5(1-3);
spiracle nearer eye than vent; interorbital distance/inter-
narial distance 1.4 or more; tail height/musculature height
2.2 or more; last posterior row subequal to upper jaw and
.50 or less times next anterior row; A-2 normally without
a median gap; central Texas, westward Scaphiopus couchi
Jaws narrow; labial tooth row formula 4-6 (2-6) / 3-6 ( 1-3);
spiracle equidistant between eye and vent; interorbital
distance/internarial distance 1.8 or less; tail height/mus-
culature height 2.5 or less; last posterior tooth row longer
than upper jaw and .50 or more times next anterior row;
A-2 normally with a median gap; central Texas eastward
-----------------------------------------Scaphiopus holbrooki

11. Tail length/tail height 1.9 or less; tail height/musculature

height 3.1 or more; dorsal fin arises abruptly from body;
body globular; primarily east of Rocky Mountains
----------------------------------------Spea bombifrons
Tail length/tail height 2.0 or more; tail height/musculature
height 2.9 or less; dorsal fin does not arise abruptly from
body; body globular to depressed; primarily west of
Rocky Mountains -Spea hammondi,S. intermontanius
12. Eyes dorsal; oral disc emarginate; papillary border does not
or barely reaches above lateral tips of A-1; labial tooth
row formula 1-7(2-7) /2-6[1], commonly 2-3/34 - 13
Eyes lateral or dorsal; oral disc not emarginate; papillary
border commonly reaches considerable distance above
lateral tips of A-1; labial tooth row formula 2(2) /2-4[l]
-------------------------------------Hylidae-Section C
13. Labial tooth row formula 2/2; papillary border with a pos-
terior gap equal to length of P-2; marginal papillae few;
central Texas to Arizona - Bufonidae (part) -Section B
Labial tooth row formula rarely 2/2; papillary border with-
out a posterior gap; marginal papillae common; wide-
spread -Ranidae-Section D
Section B: Key to species of the genus Bufo.

1. Two rows of labial teeth on posterior labium -2
Three rows of labial teeth on posterior labium -3
2. Tail musculature unicolored; large specimens with mela-
nophores in ventral fin; collection locale from east of
Tucson, Arizona to central Texas -- Bufo debilis
Tail musculature bicolored; large specimens without mela-
nophores in ventral fin; collection locale west of Tucson,
Arizona -Bufo retiformis
3. Papillary border extends to lateral tips of P-2, although a
single papilla may occur at each end of P-3; P-1 with a
median gap; P-3 short; tail musculature bicolored and
often with dorsal light saddles; Coastal Plain from North
Carolina to Mississippi River -Bufo quercicus
4. P-3 length subequal to P-1 -5
P-3 length considerably shorter than P-1 -7
5. Mlarginalpapillae small and difficult to distinguish; upper
jaw with medium lateral processes; tail musculature dark
with light dorsal saddles; Gulf Coast from Mississippi to
central Texas -Bufo valliceps
Mlarginalpapillae obvious; upper jaw with medium to long
lateral processes; tail musculature without light dorsal
saddles; widespread -6
6. Upper jaw with medium to long lateral processes that

project posterolaterally; submarginal papillae rare to

absent; throat unpigmented; body oval in dorsal view;
central Texas to California -Bufo punctatus
Upper jaw with medium lateral processes that project pos-
teriorly; submarginal papillae common; throat pigmented;
body rounded in dorsal view; southeastern Florida and
southern Texas -Bufo marinus
7. P-1/P-3 more than 2; upper jaw angulate; several to many
submarginal papillae at lateral tips of P-3; A-2 gap wide;
dorsum typically lightly pigmented; Great Plains states
and adjacent Canada southwestward to southern Cali-
fornia - ------------------------------------------------------
Bufo cognatus
P-1/P-3 2 or less; upper jaw not noticeably angulate; sub-
marginal papillae at tips of P-3 not prominent or absent;
A-2 gap narrow; dorsum light or dark; widespread- 8
8. Collection locale east of Mississippi River- 9
Collection locale west of Mississippi River -11
9. Upper and lower fins equal to musculature height; dorsal
fin often higher than ventral; dorsum dark brown to
black, with a light oblique mark behind each eye in life;
Coastal Plain from southeastern Virginia to Mississippi
River - Bufo terrestris
Upper and lower fins lower than musculature height; fins
subequal in height; dorsum dark without light mark
behind each eye in life; widespread -10
10. Dorsum unicolored; snout sloping in lateral view; eyes
small; tail length/tail height 2.7 or less; musculature usu-
ally distinctly bicolored; tail height/musculature height
2.0 or less; widespread above Fall Line, north to Hudson
Bay -Bufo americanus (part)
Dorsum commonly with light mottlings in life; snout
rounded in lateral view; eyes large; tail length/ tail
height 2.8 or more; musculature often not distinctly bi-
colored; tail height/musculature height 2.0 or more;
widespread except in southeast- Btfo woodhousii (part)
11. Collection locale east of Continental Divide -12
Collection locale west of Continental Divide -16
12. Dorsum lightly pigmented; tail musculature light with dark
lateral blotches that tend to form a stripe; Texas, western
Oklahoma, and southeastern New Mexico Bufo speciosus
Dorsum typically darkly pigmented; tail musculature uni-
colored or bicolored without contrasting lateral markings;
widespread -13
13. Tail musculature not bicolored in lateral view, but ventral
surface may be unpigmented; fins pigmented; throat
evenly pigmented with punctate melanophores; to 56 mm

total length; high elevationsnorthwestwardfrom cerntral

Colorado-Bufo boreas (part)
Tail musculaturebicolored,althoughlight ventralarea may
be narrow;fins unpigmentedor with only scatteredstel-
late melanophores;at least part of throat clear of pig-
ment; to 30 mm total length; lower elevations generally
outside above range-14
14. Clear throat patch extends nearly to gut, remainder of
venter unicoloredor with reticulate melanophores;light
ventral part of tail musculaturenarrow;northwestward
from west-central Minnesota to northcentralSaskatch-
ewan plus LaramieHills, Wyoming- Bufo hemiophrys
Throat patch small to absent; remainderof venter evenly
pigmented with punctate melanophores;light ventral
part of tail musculaturewide or narrow;widespread 15
15. Throat largely pigmented; no melanic gaps in dorsal tail
musculaturepigment; snout sloping in lateral view; eye
small;tail height/musculatureheight 2.0 or less; dorsum
dark without contrastingmarkings;widespread east of
100thmeridian- Bufo americanus(part), B. houstonensis
Throatpatch present;often irregularmelanicgaps in dorsal
tail musculaturepigment; light ventral portion of tail
musculaturewide; snout rounded in lateral view; eyes
large;tail height/musculatureheight 2.0 or more;dorsum
dark,often with contrastingmottlingsin life; widespread
-------------------------------------Bufo woodhousii (part)
16. Dorsum and venter black; fins darklypigmented; P-1 gap
commonlypresent; tail musculaturetotally black; mon-
tane meadows above 2,000 meters in central Sierra
Nevada Range-Bufo canorus
Dorsumdarkor light, venter lighter than dorsum;fins clear
to moderately pigmented, dorsal more so than ventral;
P-1 gap absent; tail musculatureunicolored,mottled or
bicolored;widespread-17
17. Fins moderately pigmented, dorsal more so than ventral;
submarginalpapillae common at lateral tips of A-2 and
P-2 and on emarginationfold; tail musculatureunicolored
or with a narrow ventral area unpigmented;to 56 mm
total length;southernCalifornianorthand northeastward
to south Alaska- Bufo boreas (part), B. exsul, B. nelsoni
Fins clear or with few stellate melanophoresmore numer-
ous in dorsalfin than ventral;submarginalpapillae rare
to absent;tail musculaturemottled or bicolored;less than
56 mm total length; widespread- .------------------------------------------
18
18. Body lightly pigmented, brassy in life and darker in pre-
servative due to melanic pigment in deeper layers;

melanophores absent in center of belly; jaws coarsely

serrate; tail musculature pigmented evenly with large
punctate melanophores; southern Arizona and adjacent
New Mexico and California -Bufo alvarius
Body lightly or darkly pigmented, brassy or not in life;
melanic pigment present in outer layers of tissue; punc-
tate melanophores present over gut; jaws finely serrate;
widespread -19
19. Tail musculature bicolored, although melanic gaps may
occur anywhere on the tail musculature; dorsum typically
dark; widespread -Bufo woodhousii (part)
Tail musculature light with dark blotches; dorsum light;
scattered localities in southwestern states
--------------------------------------Bufo microscaphus
Section C: Key to species of the family Hylidae.
1. Two rows of labial teeth on posterior labium -2
Three or four rows of labial teeth on posterior labium 6
2. A-2 gap wide; spiracular tube at least partially free from
body wall; tail tip often black; dorsum of tail musculature
frequently banded; body slightly depressed; eyes dorso-
lateral; nostrils large; fins without bold markings; east
of Continental Divide -Acris-3
A-2 gap narrow; spiracular tube almost fully attached to
body wall; tail tip never black; dorsum of tail muscula-
ture banded or not; body globular; eyes lateral; nostrils
large or small; fins with or without bold markings;
widespread -4
3. Free section of spiracular tube long, almost entire length
of tube; throat dark; tail musculature finely flecked; pri-
marily Coastal Plain from Virginia to Mississippi River
----------------------------------------------------Acris gryllus
Free section of spiracular tube short, one-half or less of the
length of the tube; throat light; tail musculature mottled
or reticulated; widespread outside of Florida and Atlantic
Coastal Plain -Acris crepitans
4. Tail musculature striped in lateral view; light stripe extends
forward to eye from dorsal musculature stripe; throat and
chest rnottled; dorsum of tail musculature often banded;
small size; Coastal Plain from Virginia to Florida pan-
handle -Limnoedus ocularis (part)
Tail musculature not striped in lateral view; light stripes
from tail to eye absent; throat and chest light; dorsum of
tail not banded; larger; widespread -5
5. Tail musculature unicolored or bicolored; one row of mar-
ginal papillae; fins clear with some stellate melanophores;
A-2 subequal to A-1 -Pseudacris triseriata (part)

Tail musculature mottled; two rows of marginal papillae;

fins clear with or without large black blotches, if present,
a clear area near musculature; A-2 longer than A-1
-------------------------------------------Hyla crucifer (part)
6. Three rows of labial teeth on posterior labium; widespread 7
Four rows of labial teeth on posterior labium; extreme
southern Florida -Hyla septentrionalis
7. Eyes dorsal; primarily west of Continental Divide-- 8
Eyes lateral; east or west of Continental Divide-- 9
8. Dorsum of tail musculature typically with light saddles;
body depressed; jaws medium to wide; dorsal fin ter-
minates near body-tail junction; P-1 sometimes with a
median gap; P-2/P-3 2 or more; rocky streams of south-
western California -Hyla cadaverina
Dorsum of tail musculature never with light saddles; body
globular; jaws wide; dorsal fin terminates at level of
spiracle; P-1 without a median gap; P-2/P-3 2 or less;
streams in Arizona, southern Utah, and Mexico
--------------------------------------------Hyla arenicolor
9. Collection locale west of Continental Divide -10
Collection locale east of Continental Divide -13
10. Upper jaw without or with short lateral processes; P-3 long;
jaws narrow to medium; A-2 gap ratio 2 or more; extreme
southcentral Arizona -Pternohyla fodiens
Upper jaw with short to long lateral processes; P-3 short or
long; jaws wide or not; widespread- 11
11. P-1 with a median gap; jaws wide, upper jaw with short
lateral processes; one row of marginal papillae; tail pat-
terned with dendritic melanophores to varying degree;
widespread from western Montana -Hyla regilla
P-1 without a median gap; upper jaw wide or not, lower
jaw narrow; two rows of marginal papillae; widespread 12
12. Upper jaw wide; lower jaw narrow and shallow V-shaped;
dorsal fin low, terminating between spiracle and vent;
numerous submarginal papillae; central and southern
Arizona and adjacent New Mexico - Hyla w-rightorum
Upper jaw medium; lower jaw narrow and wide U-shaped;
dorsal fin high, terminating at or anterior to the spiracle;
few submarginal papillae; Rocky Mountain states north
to Yukon Territory -Pseudacris triseriata (part)
13. Papillary border with a posterior gap -14
Papillary border without a posterior gap -15
14. Tail musculature striped in lateral view; light stripes extend
to eye from dorsal light stripe on tail; dorsum of tail
musculature often banded; to 25 mm total length; fins

clear or with few melanophores; Coastal Plain from

Virginia to Florida panhandle Limnoedus ocular'is (part)
Tail musculature not striped, mottled in lateral view; light
stripes from tail to eyes absent; dorsum of tail muscula-
ture not banded; to 35 mm total length; fins often with
large dark blotches and a clear area near musculature
15. Tail musculature dark with light (reddish in life) dorsal
saddles; fins dark without bold markings; light inter-
orbital and orbitonasal bands present in life; gut not
visible, belly black; swamp areas primarily in Mississippi
embayment area -Hyla avivoca
Tail musculature dark or light without dorsal saddles; tail
fins variously patterned; interorbital bars absent, orbito-
nasal bars may or may not be present; gut visible or not;
widespread -16
16. P-3 long, .65 or more times P-2 and longer than upper jaw;
submarginal papillae well developed -17
P-3 short, less than .65 times P-2 and equal to or shorter
than upper jaw; submarginal papillae poorly developed
to absent ------------------------------------------------------------------------
20
17. Upper jaw with long lateral processes; body brownish with
pale crescent mark on posterior part; prominent fin mark-
ings absent; extreme southern Texas - Smilisca baudini
Upper jaw with short to medium lateral processes; body
without crescent mark on posterior part; prominent fin
markings typically present; widespread elsewhere than
above -18
18. Tail musculature distinctly striped; fins flecked or blotched
with clear area remaining near musculature; clear part
of fin reddish in large specimens in life; flagellum well
developed and clear of pigment; Coastal Plain from
Maryland to Mississippi River - Hyla femoralis
Tail musculature not distinctly striped; fins blotched or not,
with or without clear area near musculature; clear part
of fin reddish or not; flagellum less developed; wide-
spread ------------------------------------------------------------------ 19
19. Tail height/body height at eye level 1.5 or less; dorsal fin
equal to or greater than musculature height; tail length/
tail height 3.1 or more; greatest body width/basal width
of tail musculature 2.7 or more; large specimens often
with reddish fins; throat seldom pigmented; widespread
-------------------------------Hyla versicolor, H. chrysoscelis
Tail height/body height at eye level 1.5 or more; dorsal fin
less than musculature height; tail length/tail height 3.3
or less; greatest body width/basal width of tail muscula-

ture 2.7 or less; large specimens without red fins; throat

typically pigmented; Coastal Plain from Virginia to
central Texas --- Hyla squirella
20. Tail musculature striped in lateral view -21
Tail musculature bicolored, unicolored or mottled in lateral
view -22
21. Wide lateral tail stripe bordered dorsally and ventrally by
light stripes; dorsal light stripe extends forward to eye;
dorsum of tail musculature often banded; to 25 mm total
length; Coastal Plain from Virginia to Florida panhandle
----------------------------------Limnoedus ocularis (part)
Dark stripe with indistinct edges projects obliquely from
center of tail musculature to dorsal margin of muscula-
ture at about midlength; stripe to eye absent; dorsum of
tail not banded; to 35 mm total length; scattered localities
from northern South Carolina to New Jersey Hyla andersoni --
22. Jaws wide; upper jaw angulate; dorsal fin high, terminating
anterior to spiracle; body compressed slightly; to 55 mm
total length; tail clear, black or mottled; Coastal Plain
from North Carolina to Mississippi River -23
Jaws narrow to medium; upper jaw not angulate; dorsal fin
high or not; tail clear or blotched; never to 55 mm total
length; widespread -24
23. Total length less than 30 mm.;fins clear; light stripe extends
from tail musculature forward to eye; dorsum of tail
musculature with a black saddle slightly anterior to mid-
length -Hyla gratiosa (part)
Total length more than 30 mm; fins clear, black or mottled;
stripe from tail to eye absent; black saddle on tail mus-
culature absent -Hyla gratiosa (part)
24. Fins and tail musculature typically mottled or reticulated
without a clear area near musculature; A-2 gap ratio 3 or
more; light orbitonasal stripe present at least in life;
small specimens with two light body blotches that form
an incomplete transverse body band (usually lost in
preservative); dorsal fin equals ventral; Coastal Plain
from Delaware to central Texas and north along the
Mississippi River to southern Illinois -Hyla cinerea
Fins not mottled or reticulated, sometimes blotched; if
blotched, a clear area present near musculature; A-2 gap
ratio 3 or less; light orbitonasal stripe absent; light body
blotches absent; dorsal fin variable; widespread -25
25. Fins commonly blotched with a clear area remaining near
musculature; tail musculature mottled; dorsal fin higher
than ventral; P-3 very short; widespread

Fins not blotched, either clear or with a few stellate mela-

nophores; tail musculature mottled, unicolored, bicolored
or striped; dorsal fin variable; widespread -26
26. Collection locale east of the Mississippi River -27
Collection locale west of the Mississippi River -32
27. Throat pigmented; A-2 gap relatively wide; tail musculature
striped and dorsal light stripe extends forward to eye;
P-1 with a median gap; to 21 mm; Coastal Plain from
Virginia to Georgia -Pseudacris brimleyi
Throat not pigmented; A-2 gap relatively narrow; tail mus-
culature not striped; no light stripe from tail to eye; P-1
with or without median gap; to 35 mm total length;
widespread -28
28. Dorsal fin high, extending anterior of spiracle; tail mus-
culature distinctly bicolored; P-1 indented or with a
narrow median gap; tail height/musculature height 3.5
or more; Coastal Plain from North Carolina to Mississippi
River-- Pseudacrisornata
Dorsal fin high or low, usually extending no further than
to spiracle; tail musculature unicolored or indistinctly
bicolored, i.e., not for full length of tail or with consider-
able pigment in lower half; P-1 with or without a median
gap; tail height/musculature height 3.2 or less; wide-
spread -29
29. Body dark brassy in life, subdermal layers dark and dermal
layers transparent in preservative; A-2 gap ratio 2 or
more; P-1 and P-2 equal and sometimes united laterally;
dorsal fin low, terminates posterior of spiracle; eastern
Pennsylvania southwestward to northeastern Mississippi
---------------------------------Pseudacris brachyphona
Body not uniformly brassy in life; dermal layers of body
darkly pigmented in preservative; A-2 gap ratio 2 or
less; P-1 and P-2 equal or not; dorsal fin terminates at
spiracle or anterior; widespread -30
30. P-1 often with a median gap; tail height/musculature height
2.9 or more; upper jaw wide; fins clear, musculature
lightly pigmented; snout sloping in lateral view; dorsal
fin terminates anterior of spiracle; dorsum brownish-
gray; scattered localities in southern and northcentral
Illinois -Pseudacris streckeri (part)
P-1 without a median gap; tail height/musculature height
2.8 or less; upper jaw medium; fins blotched or mottled,
musculature heavily pigmented; snout rounded in lateral
view; dorsal fin extends to spiracle; dorsum dark; wide-
spread -31
31. Chest pigmented; dorsum uniform black to dark brown

without small black dots; fins subequal and both lower

than tail musculature;CoastalPlain from North Carolina
to MississippiRiver-Pseudacris nigrita
Chest not pigmented; dorsum black to dark brown and
typically with small black dots; dorsal fin higher than
ventral and about equal to tail musculature;widespread
primarilyoutside of Coastal Plain-
-----------------------------------Pseudacris triseriata (part)
32. P-1 often with a median gap; musculature lightly pig-
mented; snout sloping in lateral view; dorsal fin ter-
minates anterior of spiracle; fins unpigmented; central
Texas to northeasternArkansas Pseudacrisstreckeri(part)
P-1 without a median gap; musculaturedarklypigmented;
snout rounded in lateral view; dorsal fin terminatesat
spiracleor posterior;fins often pigmented;widespread-- 33
33. Submarginalpapillae few to absent; upper jaw medium;
dorsumdark gray to black; A-2 gap ratio 2 or more; tail
length/tail height 2.5 or more; tail height/musculature
height 2.5 or less; widespread Pseudacristriseriata(part)
Submarginalpapillae common; upper jaw wide; dorsum
grayish-olive;A-2 gap ratio 2.0 or less; tail length/tail
height 2.5 or less; tail height/musculatureheight 2.5 or
more;southernTexasto centralNebraska Pseudacrisclarki
Section D: Key to species of the genus Rana.
1. Four or more rows of labial teeth on anteriorlabium- 2
One to three rows of labial teeth on anteriorlabium - 5
2. Tail mottled or spotted; collection locale in California,
Oregon or Arizona -3
Tail not mottledor spotted;collectionlocale elsewherethan
above, widespread in Canada, Alaska and eastern and
northcentralUnited States -----------
-------------------Rana sylvatica (part), Rana maslini (part)
3. Jaws medium;tail lightly spotted or mottled; four or more
rows of labial teeth on posteriorlabium;collection locale
in Oregonor California -4
Jaws wide; tail and body boldly spotted; three rows of
labial teeth on posterior labium; collection locale in
southcentralArizona-Rana tarahumarae
4. Labial tooth row formula 6-7(2-5)/6(1); submarginal
papillae few to absent; streamsof Cascade-SierraRange
and Coast Range of Oregon and Califomia- Rana boylii
Labial tooth row formula4(2-4) /4(1); submarginalpapil-
lae definitely present; central SierraNevada Range and
mountainsof southwesternCalifornia Ranamuscosa(part)
5. Four rows of labial teeth on posteriorlabium -6
Two or three rows of labial teeth on posteriorlabiurn- 9

6. Collection locale in western Washington, western Oregon

or California ------------------------------------------------------ 7
Collection locale elsewhere than above; widespread in
Canada, Alaska and eastern and northcentral United
States -Rana sylvatica (part), Rana maslini (part)
7. P-i/P-4 2 or less; A-2 gap ratio 2 or more; mouth ventral;
dorsal fin terminates posterior to spiracle; body somewhat
depressed; tail musculature massive and tapers slowly,
higher at midlength than ventral fin; body somewhat
transparent; central Sierra Nevada Range and mountains
of southwestern California -Rana muscosa (part)
P-1/P-4 more than 2; A-2 gap ratio 2 or less; mouth antero-
ventral; dorsal fin terminates anterior or posterior of
spiracle; body globular; tail musculature not massive,
about equal to or less than ventral fin at midlength; body
not highly transparent; widespread- 8
8. Submarginal papillae well developed on upper labium;
dorsal fin terminates at or anterior of spiracle; tail length/
tail height 4.4 or less; fins usually unicolored or with few
small blotches; venter pink in life; Cascade-Sierra Range
westward, sea level to 2,650 meters - Rana aurora (part)
Submarginal papillae poorly developed to absent on upper
labium; dorsal fin terminates posterior to spiracle; tail
length/tail height 4.4 or more; tail often with small
blotches; venter not pink in life; about 1,000 meters in
Olympic and Cascade Mountains -Rana cascadae
9. Tail and body greenish, unicolored or more commonly pat-
terned with distinct black dots; fins similarly patterned
with more dots in dorsal fin than in ventral; venter clear
to white depending on size, with or without a contrasting
pattern; proximal portion of tail often more opaque than
remainder; live small specimens (less than 25 mm total
length) black with transverse gold bands on snout and
body; tail appears bicolored due to pigment around cau-
dal blood vessels; frequents permanent water at medium
to low altitudes; natural and artificial range includes most
of United States and southern Canada - Rana catesbeiana
Tail and body greenish or not, seldom with black dots; fins
variously patterned; venter clear, white or dark, seldom
with a contrasting pattern; widespread -10
10. Collection locale west of 100th meridian -11
Collection locale east of 100th meridian -14
11. Lower jaw wide; widespread in United States and Canada
primarily outside western tier of states - Rana pipiens
Lower jaw narrow to medium; widespread -12
12. Tail long, total length/body length 2.6 or more; dorsal fin

terminatesposteriorto spiracle; tail height/musculature

height 2.7 or less; dorsal fin equal to or less than mus-
culature;tail with small flecks and blotches; widespread
in northwesternstates and adjacent Canadianprovinces
----------------------------------------------Rana pretiosa
Tail short, total length/body length 2.6 or less; dorsal fin
terminates at or anterior to spiracle; tail height/mus-
culature height 2.7 or more; dorsal fin equal to or more
than musculature; tail darkly pigmented with or without
dark markings -13
13. Cascade-Sierra Mountain Range westward
---------------------------------------------Rana aurora (part)
Northern Idaho, Laramie Hills of Wyoming and Colorado,
north to central Alaska ----------------
-----------------------Rana sylvatica (part), R. maslini (part)
14. Lower jaw wide; nostrils medium-sized; skin thin; gut
usually visible - -15
Lower jaw narrow; nostrils small to medium; skin thick or
not; gut usually not visible - -17
15. A-2 gap ratio 2.0 or more; marginal papillae below P-3
large, 10 or less present; fins usually heavily marked,
often with dark suffusion; P-1/P-3 1.3 or more; gut often
only slightly visible; eastern United States (except south-
east) and adjacent Canada -Rana palustris
A-2 gap ratio less than 2.0; marginal papillae below P-3
small, more than 10 present; fins heavily marked or not,
speckled or spotted; P-1/P-3 1.5 or less, gut visible or
not; widespread -16
16. Gut usually visible; unpigmented throat patch with con-
trasting margins often present; if tail marked, usually not
with large spots; dorsum not stippled; dorsal fin rounded;
keratinized areas at medial tips of P-1 absent; widespread
--------------------------------------------Rana pipiens (part)
Gut visible or not; throat unpigmented without contrasting
margins or evenly pigmented; if tail marked, usually with
large spots; dorsum often appears stippled; dorsal fin
frequently triangular; keratinized areas at medial tips of
P-1 present on large specimens; Coastal Plain from North
Carolina to central Texas and sporadically up Mississippi
River valley to southern Iowa -Rana areolata
17. Tail musculature distinctly bicolored; south Atlantic Coastal
Plain and northern Florida -18
Tail musculature not bicolored; widespread -19
18. Total length 35 mm or less; fins clear; dorsumnblack, in life
with a prominent transverse gold body band; venter
transparent in preservative -Rana heckscheri (part)

Total length 35-100 mm; fins clear with a prominent black

border; dorsum bluish-black with stippled appearance;
lateral line pores prominent; venter brown to blue-black
--------------------------------------Rana heckscheri (part)
19. Stripe or row of dots formed by pigment around lateral line
pores present in dorsal fin and a less prominent stripe
usually present on tail musculature; light spots sur-
rounded by dark pigment present near edge of fins; gut
not or slightly visible; Coastal Plain from New Jersey to
eastern Texas -20
Stripe in dorsal fin and stripe on tail musculature absent;
light spots surrounded by dark pligment at fin edges
absent or indistinct; gut visible or not; widespread - 21
20. Dorsum brown with black dots; venter brown in pre-
servative, yellow to buff in line; tail musculature stripe
typically present; gut slightly or not visible; row of sub-
marginal papillae present between P-3 and marginal
papillae; tail length/tail height 2.0 or less; acid swamps
on -the Coastal Plain from New Jersey to Florida
--------------------------------------------Rana virgatipes
Dorsum brownish to greenish with black dots or mottled;
venter white, often with a contrasting pattern; tail mus-
culature stripe typically indistinct to absent; gut slightly
or not visible; row of submarginal papillae absent be-
tween P-3 and marginal papillae; tail length/tail height
2.2 or more; Coastal Plain from South Carolina to eastern
Texas -Rana grylio (part)
21. Dorsal fin terminates at spiracle; interorbital distance pro-
jected from posterior margin of cornea extends well past
spiracle; A-2 gap ratio 1.8 or less; dorsum brownish with-
out contrasting marks; widespread in central and north-
eastern area to Hudson Bay -Rana sylvatica (part)
Dorsal fin terminates posterior of spiracle; interorbital
distance projected from posterior margin of cornea ex-
tends to or slightly past spiracle; A-2 gap ratio 1.9 or
more; dorsum brownish to green with small black mark-
ings; widespread -22
22. A-2 gap ratio less than 5.0; dorsum brown to greenish; row
of submarginal papillae often present between P-3 and
marginal papillae; tail with pinkish-buff spots in life;
northward from central Wisconsin and New York to
Hudson Bay -Rana septentrionalis
A-2 gap ratio 5 or more; dorsum greenish; row of submar-
ginal papillae absent between P-3 and marginal papillae;
tail without pinkish-buff spots in life; widespread
------------------------------------------Rana clamitans

SYNOPSES
Ascaphus.-Oral disc present and emarginate; jaws unequal,
serrate and with keratinized sheaths; upper jaw plate-like, large,
without lateral processes and sometimes divided medially; lower
jaw small to absent, not in occlusion with upper jaw; labial tooth
row formula 2-3/10-12(1); some tooth rows biserial; labial teeth
non-denticulate; numerous submarginal papillae; spiracle single,
midventral and nearer mouth than anus; anus medial; eyes dorsal;
anal flap present; external nares large and tubular; mouth ventral;
tail fin low; body depressed; darkly pigmented; mountain streams
of northwestern United States; 1 species.
Rhinophrynus.--Oral disc absent; jaws without keratinized
sheaths; upper lip without a median notch; oral barbels present;
spiracles dual, lateral, and near longitudinal axis; nostrils present
throughout development; anus medial; eyes lateral; anal flap absent;
external nares not tubular; mouth terminal; tail fin medium; body
depressed; lightly pigmented; southern Texas; 1 species.
Scaphiopus.-Oral disc present and not emarginate; jaws serrate,
subequal and with keratinized sheaths; lateral processes of upper
jaw medium; lower jaw striated; labial tooth row formula 2-6 (2-6) /
3-6(1-3); tooth rows uniserial; labial teeth commonly non-denticu-
late; papillary border complete or with narrow dorsal gap; sub-
marginal papillae present; spiracle single, sinistral and below longi-
tudinal axis; anus medial; eyes dorsal; external nares not tubular;
anal flap absent; body globular or depressed; tail fin moderate;
eastern and southwestern states; 2 species.
Spea.-Oral disc present and not emarginate; jaws serrate, sub-
equal and with keratinized sheaths; lateral processes of upper jaw
short; lower jaw not striated; upper jaw frequently cuspate, if so,
lower jaw deeply notched; labial tooth row formula 2-6(3-6)/4-6
(2-6); tooth rows uniserial;labial teeth commonlynon-denticulate;
papillary border complete or with narrow dorsal gap; submarginal
papillae few to absent; spiracle single, sinistral and below longi-
tudinal axis; body somewhat depressed; anus medial; eyes dorsal;
external nares not tubular; anal flap absent; tail fin moderate;
Missouri and westward; 3 species.
Rana.-Oral disc present and emarginate; jaws serrate, subequal
and with keratinized sheaths; lateral processes of upper jaw short
to medium; labial tooth row formula 1-7(2-7)/2-6[1]; tooth rows
uniserial; labial teeth commonly denticulate; A-2 gap typically wide;
papillary border with wide dorsal gap; submarginal papillae com-
mon; spiracle single, sinistral and near longitudinal axis; body
globular or slightly depressed; anus dextral; eyes dorsal; external
nares not tubular; anal flap absent; tail fin low to high; widespread;
17 species.

Gastrophryne.-Oral disc absent; jaws without keratinized

sheaths; labial flaps with a median notch, without marginal papillae
and with medial margins parallel or divergent; oral barbels absent;
spiracle single, midventral and immediately ventral to anus; anus
medial; eyes lateral; mouth terminal; nostrils absent until late in
development; body depressed; tail fin low; darkly pigmented; east
of Continental Divide plus extreme southcentral Arizona; 2 species.
Hypopachus.-Oral disc absent; jaws without keratinized
sheaths; labial flaps with median notch, scalloped or with marginal
papillae and with medial margins convergent; oral barbels absent;
spiracle single, midventral and near anus; anus medial; eyes lateral;
mouth terminal; nostrils absent until late in development; body
depressed; tail fin low; darkly pigmented; southern Texas; 1 species.
Bufo.-Oral disc present and emarginate; jaws serrate, subequal
and with keratinized sheaths; lateral processes of upper jaw short
or long; labial tooth row formula 1-2(2)/2-3[1]; tooth rows uniserial;
labial teeth commonly denticulate; A-2 gap narrow or wide; tail
fin low; papillary border with dorsal and ventral gaps; submarginal
papillae common; spiracle single, sinistral and near longitudinal
axis; spiracular tube attached; body globular to slightly depressed;
anus medial or dextral; dorsum dark or light; widespread; 19 species.
Acris.-Oral disc present and not emarginate; jaws serrate, sub-
equal and with keratinized sheaths; lateral processes of upper jaw
medium; labial tooth row formula 2(2)/2; tooth rows uniserial;
labial teeth commonly denticulate; papillary border with wide
dorsal gap; A-2 gap wide; spiracle single, sinistral and near longi-
tudinal axis; median wall of spiracular tube at least partly free
from body wall; anus dextral; eyes dorsolateral; external nares not
tubular; anal flap absent; body slightly depressed; tail tip commonly
black; dorsum of tail musculature often with light saddles; tail fin
low; east of the Continental Divide; 2 species.
Hyla, Pseudacris, Limnoedus, Pternohyla and Smilisca.-Oral
disc present and not emarginate; jaws serrate, subequal and with
keratinized sheaths; lateral processes of upper jaw short, long or
absent; labial tooth row formula 2(2)/2-4[1]; A-2 gap narrow;
oral disc often folds into triangular shape; tooth rows uniserial;
labial teeth commonly denticulate; papillary border with moderate
to narrow dorsal gap; submarginal papillae numerous or not; spiracle
single, sinistral and near longitudinal axis; anus dextral; eyes dorsal
or lateral; external nares not tubular; anal flap absent; spiracular
tube fully attached; body globular to depressed; tail fin low or high;
widespread (see key); 14, 7, 1, 1 and 1 species respectively.
BIBLIOGRAPHY
BABBITT, L. H. 1937. The Amphibia of Connecticut. Connecticut State
Geol. Natur. Hist. Sur. Bull. 57:1-50.
BLAIR, W. F., A. P. BLAIR, P. BRODKORB, F. R. CAGLE, AND G. A. MOORE.

1968. Vertebratesof the United States. 2nd ed. McGraw-Hill Book Co.,
New York. 616 p.
BOULENGER, G. A. 1891. A synopsis of the tadpoles of the European Batra-
chians. Proc. Zool. Soc. London 61:593-627.
BRAGG, A. N. 1936. Notes on the breeding habits, eggs, and embryos of
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Received: 11 November 1969
Accepted: 6 March 1970
Department of Zoology, Mississippi State University, State Col-
lege, Mississippi 39762


A Key To The Tadpoles of The Continental

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A Key to the Tadpoles of the Continental United States and Canada

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A KEY TO THE TADPOLES OF THE CONTINENTAL

ABSTRACT: A key to the tadpoles of the continental United States and

ADULT anurans of the United States and Canada are relatively

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Statesand Canadain developmentalstages 25 (operculumclosure)

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differentpartsof the color patternmay be formedfrom one or both

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Anal flap-small flap present in Ascaphus tadpoles, ventral to the anus.

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P-1-abbreviationfor first posteriortooth row; P (posterior)and a number

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generally large; primarily east of Mississip!pi River

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of northwestern United States and adjacent Canada

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11. Tail length/tail height 1.9 or less; tail height/musculature

Section B: Key to species of the genus Bufo.

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project posterolaterally; submarginal papillae rare to

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total length; high elevationsnorthwestwardfrom cerntral

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melanophores absent in center of belly; jaws coarsely

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Tail musculature mottled; two rows of marginal papillae;

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clear or with few melanophores; Coastal Plain from

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ture 2.7 or less; large specimens without red fins; throat

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Fins not blotched, either clear or with a few stellate mela-

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without small black dots; fins subequal and both lower

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6. Collection locale in western Washington, western Oregon

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terminatesposteriorto spiracle; tail height/musculature

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Total length 35-100 mm; fins clear with a prominent black

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Gastrophryne.-Oral disc absent; jaws without keratinized

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DUELLMAN, W. E., AND A. SCHWARTz. 1958. Amphibians and reptiles of

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