D EBATES

CLADISTICS OF THE FAlvIILY TRICHODACrYLIDAE (CRUSTACEA:

DECAPODA): A REAPPRAISAL
Juan J. Morrone' & Estela C. Lopretto'
LaborJtorio de Sistcm;itica y Biologia Evolutiva (LASBE), Musco de La Plat;"!, Pasco del Bosqllc, 1900
• La Pbt;l ARGENTINA I and Citcdrl de Zoologb hwcrtcbrados I, FJcuitad de Cicilcias N:lturab y
Musco, Pasco del Bosque, 1900 La Plat:t, ARGENTINA 2

Abstract. Morrone,} j. & E. C. Loprettll. 1996. Cladistics afthe family Trichodnctylidnc (Cntstnccn: Dccrrpodn).'
A R'£flpprnisaJ.j. Compo BioI. 1(1/2):65-72. A recendy published cladistic analysis arthc family Trichodacrylidac
ｩＮｾ＠ shown to be defective with respect to certain methodological isslles. It... rC:ln:dysis , based on 29 species
and 40 clllf:tcrcrs, produced 612 cl3dograms under equal weight.. , each with 104 S[CPS, CI=O.37 :lt1d
RJ =0.75. Successive weighting reduced thcm to six c1adogralns with 248 Stcps, C I = 0.68 and IU = 0.92.
Both TriciJodnctyllls Latrcillc and Dilocnrcilllls H. Milne-Edwards appear as non -monophyletic groups, and
;uc newly circumscribcd to includc species previously assigned to othcr genera. Following the phyloge-
netic sequencing convention for asymmetrical cl:ldogr:.lms, five genera are recognized: (1) TricJlOdnctyllls
Latreillc (= Mikrotricbodnctylus Prctzman syn.n. , /{odri..qllczia Bottsyn.n. , andAvotrichodnctyl1ls Pretzmann,
syn .n. ), (2) Vnldil'in White, (3) rorstcria Bott, (4) Sylviocnrcinlls H. Milne-Edwards, and (5 ) Dilocarcinus
A. Milnc-Edwards (=Fredilocarcinus Pretzmann syn.n. and Zilchiopsis syn.n.). Three new combinations
are established: Trichocbctylus oaxcnsis (Rodrfguez) comb.n., Dilocarcinus satderi (Bott) comb.n., and
D. chacei (Prc[ZInann ) comb.n.

Keywords: Crustacca, Decapoda, TrichodactyJidae, taxonomy, cladistics.

Introduction methodological Is s ues and hi s

The freshwater crabs of the family
Triehodaetylidae (Cntstaeea: Deeapoda)
presently comprise 10 genera and 43
named species (Rodriguez, 1992 ). Some
species arc important clements of the
freshwater eareinofauna of South
America, and from a biogeographic
viewpoint they may play an important role
for understanding the biotic evolution of
the region (Morrone & Lopretto, 1994,
1995 ).
Generic assignment'i of trichodactylids
are unclear. Based on characters from the
male pJeopod, Lopretto (1976) found
that the classification of the group (c.g.,
Pretzmann, 1968a,b; Bott, 1969) was not
satisfactory, because the four species
groups delimited by her were incongruent
with the current generic assignment of the
species. A sound phylogenetic hypothesis
for this group is needed to clearly elucidate
the placement of some problematical
species.
Recently, Rodriguez (1992) revised the
Trichodaetylidae and proposed a
cladogram for d1eir species. His analysis,
however, is defective wid1 respect to some
classification is not consistendy cladistic.
When the classification is compared wid1
his cJadogram (Fig. 1) it is seen that some
tJ.'(a are not monophyletic and five clades
are not supported by any character.
Furthermore, the strict consensus
c1adogram presented by Rodriguez
(1992) for t1,e 18 cJadograms obtained
by him (Fig. 2) is strongly contradictory
with the chosen cladogram, e.g., certain
para phyletic groups in the c1adogram
appear as monophyletic in d1e consensus.
The aim of this paper is to revise
Rodriguez'S (1992) analysis , in order to
propose a cladistic classification for d1e
group t1ut reflect' the aculal phylogenetic
relationships between the genera. We
intend to show that some of Rodriguez's
(1992) groups are not supported or arc
even counterindicated by his data, and
that a different c1adogram is a better
hypothesis oftrichodaetylid relationships.
Rodriguez'S analysis
Rodrigue z's ( 1992 ) considered 30
terminal units in the group (Table I) and
 Morrone & Loprctto

ROOT ROOT
SIDE BIDE
BORE eORE
PANO PANO
FlUV "lUV
VlLC YILL
PETR PETR
KENS KENS
QUIN QUIN
CONS CONS
OAXE OAXE
DEVI BULB /
PIRI cENT
PleT PAGE
SATT MEOE
SPEC Nice
MALO TRUN
BULB MUSM
TRUN EMJU
DENT EMAR
MEOE CAME
PAGE HART
NICE SERR
MUSM GILA
ENIAR OEVl
EMJU PIRI
CAME PleT
HART SPEC
SERR SATT
GILA MALO
1 VENE 2 VENE

ROOT ROOT
BIDE BIDE
BORE BORE
PANC PANQ
FLUV FLUV
VILL VILL
PETR PETR
KENS KENS
QUIN
C1UIN
CONS
CONS
QAXE
OAxe
BULB
BULB
DENT
TRUN
MEOE
NICE
DENT
PAGE MEOE
NICE
SATT
TRUN PAGE
MUSM EMAR
EMAR MUSM
EMJU SATT
CAME SPEC
HART Pier
SERR DElli
GILA PIRI
DEVI MALO
PIRI VEHE
MALO CAME
PleT GILA
SPEC HART
3 VENE 4 SERR

Figures 1-4. I . Original c1adogram presented by Rodriguez (1992). 2. Strict consensus c1:ldogram
of the 18 c1adograms obtained by ROdriguez (1992). 3 . Strict consenSllS c1adogram of the 15
dadograms found for Rodriguez's (1992) original data. 4. Consensus c1adogram based on six
c1adograms obtained by us.

66
 Cladistics of the fami ly Trichodactylidae

Table r. Species of Trichodactylidae treated as units of the analysis. A=Avotrichodactylus

D=Dilocarcinusj Of=Forsteria j Fr=Fredilocarcinus; H=Holthus i:lj M=Mikrotrichodactylus
P = Poppi:ma j Ro = Rodriguezia j Rv=Rotundovaldivi:l j S=Sylviocarcinusj T =Trichodactylus
V=Valdi"iaj Z= Zilchiopsis.

, Prcvious gcncric and subgcncric assignmcnu

Acronyms Species Rathbun Prerzm:ulIl Bott Rodrfgue·z;

( 1906) (1 96S',b) ( 1969) ( 1992)

BIDE A. hidnu T(V) T(M) T(T) A

BORE M . bore/Jirmus T(Ro) M
BULB D. btllbiftr D
CAME V. cnmcrrmi T(V) V(Rv) T(T) V
CONS A. c01lStri,,1IS T(A) T(Ro) A
DENT D. dwtntm T(D) D P D
DEVI S. dCJII'//ci T(V) V(V) S S
EMAR Z. emnrgi,m!lIS T(D) D Z Z
EMjU Z. emnrgj'lntl/s jllv. Z
FLUV Tflm1intilu T(T) T(T) T(T) T
GILA v.giIa V
HART V. bami; T(V) V(Rv) V(V) V
KENS T. kemleyi l'
MALD S. maJdonndoensis S
MEDE D. medemi D
MUSM F lII/mll/lSchiae Fr
NICE D. lIiceforei V( Rv) D
OAXE A.ofJXmsis A
PAGE D.pngei T(D) D(D) D
PANO M. panopllls T(V) T(M) T(T) M
PETR T. petropolitfl'J/ff T(V) T(T) T(T) T
PIer S. pictttS T(D) H S S
PlRI S. pirifonllis V(V) S
QUIN T . fJtlillfJlICdmtntllS T(1') T(T) T(Ro) T
SATf Z. sattleri Z Z
SERR v.scrrata T(V) V(V) V(V) S
SPEC S. spec. V
TRUN D. tnmentlls D
VENE F l'etJezllefmsis T(V) H V(Fo) 1'0
VILL It l,i/lnlobosi T(Ro) Ro

40 characters (Table II), whose polarity c1arustic analysis) is as follows:

was determined by outgroup comparison
with the subfamily Carcininae. The data
matrix (T able III ) was analyzed by
Rodriguez (1992 ) with programs
PHYLIP 3.0 (Felsenstein, 1986) and
PAUP 2.2 (Swofford, 1985), generating
five and 18 c1adograms, respectively, all
of them with 108 steps and cr of 0.37.
From them, he chose one c1adogram (Fig.
1) and presented a consensus c1adogram
(Fig. 2 ) ( note t he in co ns iste ncies
commented above .)
R o drigu ez's ( 1992) proposed
classification for d1C whole family (i.c.]
including species not considered in his
1.0. Subfami ly T ri chodactylinac H . Mi lne-
Edwards, 1853
1.1. Genus TricbodnctylllS Larreille, 1828
T. fiuvinti/is Latreille, 1828
T. maytni Pret7.mann, 1978
T. kensleyi Rodrfguez, 1992
T. petropolitnlllls (Gocldi, 1886 )
T. qtlillqlled£1lfatlls Rathbu n, 1893
T. ehrhnrdti ( Bott, 1969)
1.2. Genus Mikrotrjcbodn czylus Preczmann ,
1968
M. borel/i{l1lJ1s (Nobili , 1896)
M. pmlOpilu (von Martcns, 1869)
1.3. Genus Rndrigllezin. BoCt, 1969
lL mmsnbnk (Cottarclli & Argano, 1977)
R. l>illnlobosi ( Rcxl.dgucz & Manriquc, 1967)
1.4. Genus Avotrichodnctyills Prcrzmann. 1968
A. bidem ( Bort, 1969)

67

A . constricttts (Pc:trse, 1911 )
A. oaxC1lsis (Rodriguez, 1992)
2.0. Sub6m ily Dilocarcinin:tc Pretzmann, 1978
2.1. T ribe Holthuisiini Pretzmallll , 1978
2 .1.1. Genu s Sy ll'iocarcilills H . Mi lnc-
Edwards, 1853
S. del,ille; H . ｍｩｬｮ･ Ｍ ｅ､ｷＺｴｲＮｾ Ｌ＠ 1853
s. m a/dolladoem;s (Pretzmann, 1978)
S. pimlS (H . Milne-Edwards, 1853 )
S. pinfonnis (PretzIlUI111, 1968)
S. spec.
2.2. Tri be V:tldiviini PrCtzlll:tll ll, 1978
2. 2.1. Genus Valdil'ia White, 11)47
V. (flIllCYfmi (Nobil i, 1896)
J1.11ila (PrCtzIlUnn, 1978)
V. bnrttii (R:lthbull, 1906)
V. lntidcns (A . Milne- Edwards, 1869 )
V. scrra ta White, 1847
2.2.2 Genus Forrtcria Bott, 1969
F I'cnezlIeietlsis (lbthbull, 1906)
2.3. T ribe Dilocarcinini Pretzl1l:lllll, 1978
2.3 .1. Genus Z ilclJiopsis Bott, 1969
Z. cbllee; (Prctznunn, 1968 )
Z. cryptodus (Onnunn , 1893)
Z . CIIuu'Jl; nntw (H . Mi lnc- Edw:lrds,
1853)
z. sllttleri Bott, 1969
2 .3.2. Gen us Dilt)CIll·cillttS H . Mi lne -
ｅ､ｷＺｴ ｲ ､ＮｾＬ＠ 11)53
D . nrqcmillilllllls (R:tthbull, 1906)
D. tr;/1lcn tlls Rodrfguez, 1992
D. bulbiftr Rod riguez, 1992
D . eflstc/wlU; H. Milne-Edw:trds, 1853
D. detl tllt llS (Randall , 1839)
D ./ncl'ifm1lS Morcir:l, 190 1
D . mcdcmi Smalley & Rodrfguez, 1972
D . lIieeforci (Schmin & Pretzl1l:lnn, 1968 )
D. ｰ ｮ ｾｱ｣ ｩ＠ Stimpson, 186 1
D. septcmdcntntllS (H erbst, 1783 )
D. spill ifer H . Milne-Edwards, 1853
2.3 .3. Genm Frcdilo((lrcilltls Pretznunll ,
1978
1-: m ddn i (Pretznun n, 1978)
r: m lwmlSc/Jillc (Pretznunll & M:l)'ta,
1980)
Species incertae ｳ･ ､ｩ Ｎｾ＠
Tric/;odncty/w (D i/()(flrci1/1/ s) ｾ ｱＱｴｙｬｰｷｳｪ
Rathbun, 1906
Tn·chodnctylllJ petropolitn",ls pnrnllnmsis Bott,
1969
Tricbndnetyllls (VnldiJ'ia) fo.wmi R:lthbllJl , 1906
W hen this classification is compared
with Rodriguez's (1992) c1adogram (Fig.
1 ) it re sul ts t h at t he s ubfa mil y
Diiocarcininac, the tribe H olthuisiini,
Trichodactylus, A potrichodactyltfs, Sylvio-
carcinus, and Zilchiopsis a re 110n -
monophyletic
Platni ek (1987 , 1989 ) fo und that
PHYLIP and o lde r ve rsions ofPAUP are
less rel iab le tha n F arris' s ( 19 88 )
Morrone & Lopretto
Table III. Data matrix analyzed by Rodriguez
( 1992 ) . Acronym s :lS in Table l.
O= plesiomorphicj l=:lpomorphic.
ROOT 00000 00000 00000 00000 00000 00000 00000 00000
BIDE 10100 00011 00001 00010 10110 00001 01100 00101
BORE 1 0100 00011 00011 00011 11110 00001 01111 000 1 0
BULB 11110 10000 00101100 11100 11 00010 0 1 010 11000
CAME 00001 0 1000000000011010001 10000 0 1 000 01001
CONS 101 0000001 0000 1 000 1 0 10 110 00001 01100 00100
DENY' 11110 1001 000101 1101110011 10100 01010 11000
DEVI 11110010000 1 000 1 000 1 0001110100 00000 01000
£MAR 0 1 000 00000 0000 110011 1 0011 00 1 00 010 1 0 11000
FUN 10100 000011100 1 000 11 1 0 110 1 0001 0 110 1 00011
GILA 0000111000 00000 000 10 100 11 1 0100 0 1 000 11000
.'
HART 00001 0 1000000000011010001 101 00 0 1 000 11000
KENS 11000 100 11 00001 00010 111 00 00001 01101 00011
MALO 11110 100000000000000100 11 10000 01000 01000
MEDE 11110 100 1 000101 110 11 100 11 101 00 010 1 0 11000
MUSH 01110 1000000001100 11 100 11 00000 01010 11000
NIC E 11110 1 0000001 01 110 11 10010 10100 01010 11000
OAXA 1 010000001 00011 000 1 0 10110 00000 01100 00101
PAGE 11110 100 1 00010110011 1 00 11 11100 01010 11001
PANO 1 0 1 00000 11 00011 000 1111110 00001 01111 00010
PIITR 1 0 1 00000 1 0 00001 0001110110 1000101101 00011
PICT 11110010000000 1 00 110100 11 10100 01000 01000
PIR I 1111001100 1 0000 00000 000 11 100000000001000
QUIN 1 11 10000 11 000 11 00 110 1 0100 00001 01101 00000
SA"IT1111 00001 0 1 0001 10 1111 0001101000 101011001
SERR 00001 0 1 000 00000 0001 0 1 0001 1000001000 11000
SPEC 111 100 1010 00001 00 11 110001 101000 1000 01000
TRUN 11110 10010001 011001110001 000 1 0 01010 11000
VENE 0110001000 10000 00 110 100 11 10001 01000 11000
VILL 101000000111001 00011 10110 100010110 1 001 11
H ennig86 ve rsion 1.5. Therefore, we
reanalyzed R odriguez (1992) data with
the heuristic algorithms mhelUlig* and
b b * o f Henni g 86 , obta ining 15
c1adograms, with 107 steps, C1= 0 .37,
and lU = 0. 75.
This anal ys is is equivalent to that of
R o drig u ez ( 1992 ), althou g h th e
c1adograms obtained by us are one step
shorrer than his c1adogram. If we compare
the consensus cladogram of these trees
(Fig . 3) with that of R odriguez (1992)
(Fig. 2), several difrerences appear.
ｳ＠
Reanalysis
From the te rminal units considered by
Rodriguez ( 1992 ), we exeluded the
juvenile o f Z ilchiopsis emarg inat tts
(EMJU ), because comparison should
involve only comparable semaphoronts
(Hennig, 1966).
The data matrix (T able III) was analyzed
with H ennig86, applying mhennig* and
bb * algo ri t hms. W e obt ain ed 612
c1adograms, with 104 steps, CI = 0.37 and
R1 = 0 .75. The successive weighting
proce dur e reduce d them to S I X
c1adograms, with 248 steps, CI = 0 .68 and
68
 Cladistics of the family Trichodactylidae

Table II. Characters and character s tates analyzed by Rodriguez ( 1992) . O=plesiomorph ic;
l=apomorphic.

l. Carapace outline: hexagonal (0); suborbicul:lr (I ).

2. Carapace upper surf:lcc: moderately arched (0); strongly arched ( I ).
3. Carapace: uneven (0); smooth ( 1).
4. Carapace tr:msbranchial ridge: prcscnc (0); absent (1).
5. Pair oflunubrcd protrogastric ｬｯ｢｣Ｎｾ Ｚ＠ absent (0); present (1) .
• 6. Carapace median grooves (H depression): deep (0); shallow or obsolcccnr (l).
7. Carapace posterior margin : with a high cari na (0); without a high carina ( 1).
, 8. Cara.pacc: always wider anteriorly (0); becoming wider posteriori), with age (1).
9. Carapace (at /C:lSt in immanlfc specimens): wider on anterior third (0); wider ncar middle (1).
10. Posterol:lteral margin: straight or slighdy arqu:lte (0); strongly arquate (1).
II. Later.ll teeth of carapace: not obsoleccllt in large males (0); presenc in young, obsolecellt in large malc...
(l ).
12. Lueral teeth of carapace: present at least in juveniles (0); absent o r on ly represented by small notches
even in young specimens ( 1) .
13. Lateral teeth in young specimens: 0-5 (0); >5 (1).
14. Lateral teeth: not beyond middle of carapace (0); beyond middle of carapace (I).
15. Front: straight (0); bilobcd ( 1).
16. Front: very advanced, concealing epistoll1e (0); p:miall)' or completely retracted, exposing epistome
( l ).
17. Margin of front : unarmed (0); armed with spines or large granules ( 1).
18. Lower orbit:l l border: without a recess (0); with a recess next to external orbital angle ( I ).
19. Orbit occlusive tooth: cominuou .. (0) ; distinct from inner orbita l :mgk ( I ).
20. Lower orbical margin: directed upward ... at inner orbital angle (0); directed downwards ( I).
21. Middle guner of epistoll1e: ending in a single point (0); ending in twO separate IXlints ( 1).
22. Apcrmre of efierem channels, lateral yugallobe: absent (0); present ( 1).
23. Postgastric pits : present (0); absent ( I ).
24. Sternum: with a deep depression on somite 1 and one on each side of somites 2+3 (0); without
cOnspiCUOll" depressions (1).
25. Abdominal segment... : all sumrcs visible (0); at lea... t sumrcs 3/ 4 and 4/5 obsolete (1).
26. Abdomen: trianhJ"tli:lr (0); tr.lpczoidaJ ( 1).
27. 1l1ird abdominal segment: withoue a carina (0); with a distal carina ( I).
28. 1l1ird maxilliped ments: trapezoidal (0); conspicuously narrow ( I ).
29. "lbird ll1axilliped :Ulteromcsia[ angle of ments: rounded (0); produced into a ttiangular tooth located
ncar articulation ofpa lp ( 1).
30. Third maxilliped distal external spine: triangular, acute (0); unusually wide (1).
31. Third maxillipcd ischium: not unusually wide (0); unusually wide ( 1).
32. Endopodite of 1st maxilliped : with a notch on mesial border (0); without Stich notch ( 1).
33. Penial groove: open cephalad (0); overlapped by 8th tergite and sterna llobc ( 1).
34. Eigth episternite: n:lrrow (0); expandcd laccr.llly ( 1).
35. First gonopod: conica l-elongated (0); ｦｬ｡Ｎｾｫ Ｍｳ ｨ｡ｰ･､＠ (1) .
36. Gonopore: V-shaped, open caudal (0) ; slit-like, open cephalic ( I ).
37. First gonopod apica l setae: if prcsent, small, located on cephalic surface (0); vel)' long, on lacera[
surface ( I ).
38. Second gOlloped: of equal length or longer than first (O)j con... idcfably shorter (1).
39. Pereiopods 2-5: lower margin of propodous and dactylus with rows of long setae (0); lacking long
setae ( 1).
40. Fifth pcteiopods propodous: wide, length/width < 1.8 (0); slender, length/width 2.0 (1).

RI=O.92 (the consensus cladogram is paraphylctic In terms of

presented in Figs. 4 and 5).
J In the cladogram (Fig. 5) there are two
basal clade s. In the former,
Avotrichodnctylus and Trichodactylus arc
I In order to reflect more accmatcl), the phylogenetic
relationships of the species of Di/ocarcitws, it may
be divided in subgenera or species group; however,
we prefer a more conservative approach and keep it
as a single genus.
Mikrotrichodactylus and Rodriguezia. In
thc second c1adc, therc is a scquccc
(Valdivia (Forsteria (Sylviocarci,,,,s +
Diwcarcinus»). Within Diwcarcimls, d,e
spccies of Zilchiopsis 1 Fredilocarcintts,
Sylviocarci""s pictllS and S. spec. are also
included.
In order to keep a minimum of
categories in the cladistic classification of
the group, we followed Nelson's (1972,

69
 Morrone & Loprctro

ROOT

,,
I. 18
!'
QUIN

" KENS
A

19 II l' II

, ,
VILL _ _ _--..I
1140

,"
B

n 16 17

,,
VENE c

DEVI D

SATT
,,
EMAR
•••
. , 1710
MUSM

Figure 5. Consensus c1adogram of the six c1adograms obtained by us, with character st ates
distributions superimposed. Black squares= synapomorphicsj dotted squares= parallelisms and
reversals; A =TriclJodactylus; B = Valdivia ; C=Forrreria ; O =Sylviocnrci1l us; E=Dilocarcj1JUS.

70
 Cladistics of the family Trichodactylidae
1973 ) phylogenetic sequencing
convention for asymmetrical cladograms,
and Amorim 's (1 982 ) "group +"
convention to refer to unnamed inclusive
groups in sequenced classifications. The
revised classification is as follows (species
that were not analyzed are treated as
inccrtae scdis) [ :
,, 1.0. Family Trichodactylidac
Trichodacrylidlc inccrrac ｳ｣､ｩＮｾＺ＠ TricIJodncty!m fnv'l:mli
Rathbull, 1906; T. gUn/pWSll Rathbun, 1906; T.
petropolitanlfs parllnflmsis Bort, 1969.
1.1. Genus Tn'chodnctyftts Latreille, 1828
Mikrotrichodactyl1tS Prcrzm:l11l1, 1968 n. syn.
Rf)drigtlezia Bott, 1969 n. syn.
AI'otn'cIJodactylm Prctzmann, 1968 n. syn.
Tricbodactyftts incenac scdis: T. ehrnrdti (Bott,
1969); T. mayta; Prcrzmann, 1978; T. mensabak
Cottarclli & Argano, 1977
T. onxemis (Rodriguez, 1992) n. comb.
T. bidem Bott, 1969, sedi! mtttabi/is
T. constricttJs Pearse, 1911, sedt's 11IIItabilis
group l]uillqllede1ltntlts+, sedt's 1111ltabilis
T quinquede1ltatttS Rathbun , 1893
T hemleyi Rodrfguez, 1992
T borellianllS Nobili, 1896
T. pfwopltts (von Martens, 1869)
T petropolitanus (Gocldi , 1886)
TjItlJliatilis Latreille, 1828
T. villalobosi Rodrfguez & Manrique,
1967
1. 2. Gellus Valdivia White, 1847
Valdivia ineertac ｳ･､ｩＮｾＺ＠ V. iatidens (A. Milne-
Edwards, 1869)
V. eamemn; (Nobili, 1896)
V. serrata White, 1847
V.gila (Pretzlllann, 1978)
V. harnii (Rathbun, 1906)
1.3. Genus Forsteria Batt, 1969
f. J'enezZlCle1lsis (Rathbun, 1906)
1.4. Genus Sylviocarcinm H. Milne-Edwards,
1853
S. rnaldonadoens;s (Pretzmann, 1978)
S. deville; H. Milne-Edwards, 1853
S. pirifonllis (Pretzmann, 1968)
1.5. Dilocarcintts H . ｍｩｬｮ･Ｍｅ､ｷ｡ｲＮｾＬ＠
Fredilocarei1ltts Pretzm:u1l1, 1978 n. syn.
• Zilchiopsis Bott, 1969 Il. syn .
Dilocareintls incertac sedis: D. argmtiniemis
(Rathbun, 1906); D. easteblflui H . ｍｩｬｮ｣Ｍｅ､ｷ｡ｲｾＬ＠
1853; D. elmee; (Pretznunn, 1968) n. comb.; D.
1 cryptodllS Ortmann, 1893; D. lnevifi'ons Moreira,
1901; D. mdda; Prctzmann, 1978;
septemde1ltatus (Herbst, 1783); D. spimfcr H . Milne-
Edwards, 1853
D. picflls H. Milne-Edwards, 1853
D. spec.
D. mttleri (Bott, 1969) n. comb.
D. e11/nrgillnttts H . Milne-Edwards, 1853
D. mlls11luscbiae Pretzmann & Mayra, 1980
D. pagei Stimpson, 1861
D. bulbijer Rodrfguez, 1992
D. tnmcattts Rodrigucz, 1992
D. dentattts (Randall , 1839)
D. medem; Smalley & Rodrfguez, 1972
D. niceforci (Schmitt & Pretzmann, 1968)
Some features of th e c1adogram
obtained can be summarized as fol.lows:
(1) Trichodactyl1ts (eq uivalent to Ro-
driguez's Trichodactylinac) is the sister
ta..xon to the remaining Trichodactylidae;
(2) Valdivia and Forsteria are successive
sister-taxa, not constituting a
monophyletic tribe Valdiviini;
(3) Sylviocarcinus and Diwcarcinm (the
latter including the s pecies of
Fredilocarcinus, Zilchiopsis, Sylviocarcinus
pictHs, and S. spec., equivalent to
Rodriguez's (1992) tribe Dilocarcinini)
are the less inclusive genera of the family.
Discussion
Lopretto ( 1976 ) concluded that
Sylviocarcilluspictus showed affinities with
spcCles of Dilocarcinus, whereas
Trichodactyltts camerani was considerably
different from T. borelliam" and T.
panoplus. These relationships are
corroborated by our cladogram, because
S. pictus is included inDilocarcinus and T.
camerani belongs in Valdivia.
Some of Rodriguez's (1992)
conclusions about trichodactylid
relationships are not warranted by his
data, namely the monophyly of some taxa.
The alternative classification proposed
herein , although different from his
classification , IS far from being
revolutionary: man y of the species
1853 reassigned by us are placed in the genera
were dley were originally de'icribed. When
compared with Rodriguez's ( 1992 )
classification , all the ge nera herein
relimited are monophyle tic. In addition,
D.
all supragenenc categories are
superfluous, five genera being enough to
c1assity all the species ofTrichodactylidae.
Acknowledgements. We thank Dalton de
Souza Amorim and two anonymous
reviewers for their suggestions to the
manuscript. The research of the senior
author was supported by grant 4662-91
71
 Morrone & Lopretto

from the National Geographic Society. America: F;tmily Trichod;tctylidae and

Support of the Consejo Nacional de Supplement to the Family ｐＮｾ･ｵ､ｯｴ｣ｬｰｨｳｩ｡＠
Editions de l'Orstol1l, Collection F;tune
Investigaciones Cicntfficas y Tecnicas Tropic3ic 31.
(CONICET), to which both authors SWOFFORD, D.L. 1985. PAUP : Phylogcnctic
belong, is gratefully acknowledged . :uulysis using p;trsimony. User's m;tnu;ll. Illinois
N;tUlral History Surve)" Champaign.
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