Notes on the taxonomy of Frevillea A. Milne-Edwards, 1880 (Crustacea, Brachyura, Goneplacoidea), with the description of a new genus for F. sigsbei A. Milne-Edwards, 1880

Di ec eu De la publica ion / Publica ion di ec o : Gilles Bloch
P ésiden du Muséum na ional d’His oi e na u elle
éDac ice en che / edi o -in-chie : Lau e Desu e -G andcolas
assis an e De éDac ion / assis an edi o : Anne Mabille ([email p o ec ed])
Mise en page / Page layou : Anne Mabille
coMi é scien i ique / scien i ic boa d :
Nes ine Akka i (Na u his o isches Museum, Vienne, Au iche)
Ma ia Ma a Cigliano (Museo de La Pla a, La Pla a, A gen ine)
Se ge Go as (Uni e sidad de Málaga, Málaga, Espagne)
Syl ain Hugel (CNRS, Uni e si é de S asbou g, F ance)
Ma co Isaia (Uni e si à degli S udi di To ino, Tu in, I alie)
Ra ael Ma quez (CSIC, Mad id, Espagne)
Jose Ch is ophe E. Mendoza (Lee Kong Chian Na u al His o y Museum, Singapou )
Annema ie Ohle (MNHN, Pa is, F ance)
Jean-Y es Rasplus (INRA, Mon e ie -su -Lez, F ance)
Wanda M. Weine (Polish Academy o Sciences, C aco ie, Pologne)
cou e u e / co e :
Diuplax sigsbei (A. Milne-Edwa ds, 1880) a e A. Milne-Edwa ds & Bou ie (1923: pl. 6, ig. 2).
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Ng P. K. L., Cas o P. & Rod íguez Mo eno P. A. 2024. — No es on he axonomy o F e illea A. Milne-Edwa ds, 1880
(C us acea, B achyu a, Goneplacoidea), wi h he desc ip ion o a new genus o F. sigsbei A. Milne-Edwa ds, 1880.
Zoosys ema 46 (29): 733-748. h ps://doi.o g/10.5252/zoosys ema2024 46a29. h p://zoosys ema.com/46/29
ABSTRACT
The iden i ies o he Wes e n A lan ic b achyu an c abs F e illea ba ba a A. Milne-Edwa ds, 1880,
F. osaea A. Milne-Edwa ds, 1880, F. iden a a A. Milne-Edwa ds, 1880, and F.sigsbei A. Milne-
Edwa ds, 1880, o iginally placed in Goneplacidae MacLeay, 1838, a e cla i ied on he basis o he
examina ion o ype ma e ial as well as o addi ional specimens. The co ec iden i y o he ype ma-
e ial o he ou species is esol ed. A new genus, Diuplax n.gen., is desc ibed o F.sigsbei, which
had p e iously been ans e ed o Goneplax Leach, 1814 ( amily Goneplacidae). The new genus is
dis inguished om Goneplax s.s . by ha ing a sho cheliped me us ha possesses a low submedian
ube cle on i s do sal ma gin (elonga ed me us wi h a dis inc sho submedian spine on i s do sal
ma gin in Goneplax s.s .) and he una med do sal ma gin o he me i o he ambula o y legs (wi h
dis inc subdis al spine in Goneplax s.s .). F e illea ba ba a and F. osaea a e now in he Eu yplaci-
dae S impson, 1871, F.sigsbei is in Diuplax n.gen. (Goneplacidae, Goneplacoidea MacLeay, 1838),
whe eas F. iden a a is in T apezioplax Guino , 1969 in Pseudo hombilidae Alcock, 1900 (Xan hoidea
MacLeay, 1838).
Pe e K. L. NG
Lee Kong Chian Na u al His o y Museum,
Conse a o y D i e, Na ional Uni e si y o Singapo e,
Singapo e 117377 (Republic o Singapo e)
Pe e CASTRO
Biological Sciences Depa men , Cali o nia S a e Poly echnic Uni e si y,
Pomona, CA 91768 (Uni ed S a es)
Paula A. RODRÍGUEZ MORENO
Di ec ion géné ale déléguée aux collec ions,
case pos ale 53, 57 ue Cu ie , F-75231 Pa is cedex 05 (F ance)
[email p o ec ed] (co esponding au ho )
Submi ed on 15 Decembe 2023 | Accep ed on 15 Ap il 2024 | Published on 12 No embe 2024
No es on he axonomy o F e illea A. Milne-Edwa ds, 1880
(C us acea, B achyu a, Goneplacoidea), wi h he desc ip ion
o a new genus o F. sigsbei A. Milne-Edwa ds, 1880
KEY WORDS
Wes e n A lan ic egion,
Goneplacoidea,
nomencla u e,
new combina ion,
new genus.
734 ZOOSYSTEMA • 2024 • 46 (29)
Ng P. K. L. e al.
RÉSUMÉ
No es su la axonomie de F e illea A. Milne-Edwa ds, 1880 (C us acea, B achyu a, Goneplacoidea),
a ec la desc ip ion d’un nou eau gen e pou F. sigsbei A. Milne-Edwa ds, 1880.
L’iden i é des c abes de la égion a lan ique occiden ale F e illea ba ba a A. Milne-Edwa ds, 1880,
F. osaea A. Milne-Edwa ds, 1880, F. iden a a A. Milne-Edwa ds, 1880, e de F.sigsbei A. Milne-
Edwa ds, 1880, o iginalemen placés dans les Goneplacidae MacLeay, 1838, es cla i iée su la base
de l’examen des spécimens ypes ainsi que de spécimens addi ionnels. L’iden i é co ec e du ma é iel
ype des qua e espèces es ésolue. Un nou eau gen e, Diuplax n.gen., es déc i pou F.sigsbei,
an é ieu emen ans é é dans Goneplax Leach, 1814 (Goneplacidae). Le nou eau gen e se dis ingue
de Goneplax s.s . pa le cou mé us du chélipède do é d’un aible ube cule submédian su le bo d
do sal (mé us ne emen plus allongé, a ec une cou e épine su le bo d do sal chez Goneplax s.s .) e
le bo d do sal du mé us des pa es ambula oi es sans épine (une épine subdis ale chez Goneplax s.s .).
F e illea ba ba a e F. osaea on main enan pa ie des Eu yplacidae S impson, 1871 e F.sigsbei es
dans Diuplax n. gen. (Goneplacidae, Goneplacoidea MacLeay, 1838), andis que F. iden a a es dans
T apezioplax Guino , 1969 dans les Pseudo hombilidae Alcock, 1900 (Xan hoidea MacLeay, 1838).
INTRODUCTION
As pa o his epo on he B achyu a collec ed in he Gul o
Mexico, Ca ibbean Sea, and he A lan ic coas o he Uni ed
S a es by he Uni ed S a es Coas Su ey and Geological Se -
ice (USCSGS) S eame Geo ge S.Blake be ween 1877 and
1879, Alphonse Milne-Edwa ds (1880) es ablished he genus
F e illea o ou species o ca ome opous c abs: F.ba ba a
A.Milne-Edwa ds, 1880, F. osaea A. Milne-Edwa ds, 1880,
F.sigsbei A. Milne-Edwa ds, 1880, and F. iden a a A. Milne-
Edwa ds, 1880. Much la e , A. Milne-Edwa ds& Bou ie
(1923) ea ed he genus and species in mo e de ail, adding
no es on he ma e ial hey had a ailable as well as p o iding
o he i s ime, igu es o he ou species. No ype species
was o iginally designa ed o F e illea, bu Ra hbun (1918:
25) selec ed F.ba ba a as he ype species.
Guino (1969a-c, 1971) discussed he axonomy o a ious
goneplacids a leng h and es ic ed F e illea o F.ba ba a
and F. osaea, ans e ing, wi h doub , F.sigsbei o Goneplax
Leach, 1814, and e e ed F. iden a a o a new genus, T a-
pezioplax Guino , 1969 (c . Guino 1969c). In his e ision
o he Goneplacidae MacLeay, 1838 s.s ., Cas o (2007)
ecognised F.sigsbei as a alid species o Goneplax (see also
Ng e al. 2008); T apezioplax is now in Pseudo hombilidae
Alcock, 1900 (see Guino 1971; Ng e al. 2008; Thoma e al.
2014), and F e illea s.s . is in Eu yplacidae S impson, 1871
(Guino 1971; Ng e al. 2008; Cas o& Ng 2010).
As pa o collec ion managemen in he Muséum na ional
d’His oi e na u elle, Pa is (MNHN), a echnician, Lau a
Flamme, ecen ly highligh ed o he second au ho ha he
ype specimens o F.ba ba a and F. osaea as eco ded we e
p obably mislabelled and/o mixed. Taking ad an age o a isi
o he MNHN in 2023, he i s au ho checked he MNHN’s
holdings o F e illea wi h he hi d au ho a he eques o
he second au ho , who had examined and made no es o
his ma e ial in 2006 as pa o his s udies on Goneplacoidea
(see Cas o 2007; Cas o& Ng 2010). The i s au ho also
sough assis ance om s a in he Museum o Compa a i e
Zoology, Ha a d Uni e si y, Camb idge, MA, Uni ed S a es
(MCZ), whe e a pa o A. Milne-Edwa ds’ (1880) o iginal
ma e ial is housed. One o he esul s o ou in es iga ion
was he es ablishmen o a new genus, Diuplax n.gen., o
F e illea sigsbei A. Milne-Edwa ds, 1880.
MATERIAL AND METHODS
Measu eMen s
Measu emen s (in millime es) a e o he maximum ca apace
leng h ollowed by wid h (including ee h), espec i ely. The
o iginal ype ma e ial examined was e-measu ed using he
same me hod used o he mo e ecen ma e ial. The e mi-
nology used ollows Da ie e al. (2015). Specimens examined
a e deposi ed in MCZ, ZRC and MNHN.
abb e ia ions
Ins i u ions
MCZ Museum o Compa a i e Zoology, Ha a d Uni e si y,
Camb idge, MA;
MNHN Muséum na ional d’His oi e na u elle, Pa is;
ZRC Zoological Re e ence Collec ion, Lee Kong Chian Na u al
His o y Museum, Na ional Uni e si y o Singapo e.
Mo phology
G1 male i s gonopod;
G2 male second gonopod.
noMencla u e
The specimens o A. Milne-Edwa ds (1880) and A. Milne-
Edwa ds& Bou ie (1923) ha e been a iously lis ed, labelled,
and/o da abased as “ ype”, “co ypes”, “holo ype”, “lec o ype”
o “pa alec o ype” by a ious au ho s and cu a o s. This con-
usion s ems om he manne hese au ho s iden i ied which
is he “ ype” (in he sense o he name-bea ing specimen) and
how hey lis ed hei ma e ial. They also did no s a e whe e
each specimen was deposi ed (i.e., MCZ o MNHN). Speci-
mens we e p obably only sha ed be ween he wo museums
MOTS CLÉS
Région a lan ique
occiden ale,
Goneplacoidea,
nomencla u e,
combinaison nou elle,
gen e nou eau.
735
Taxonomy o F e illea and desc ip ion o Diuplax n. gen.
ZOOSYSTEMA • 2024 • 46 (29)
la e on. The si ua ion was compounded by Ra hbun’s (1918)
nomencla u al ac ions (see below). The p esen decisions on
which specimens belongs o he syn ype se ies and whe he
hey a e holo ypes o lec o ypes ollow a s ic in e p e a ion
o he cu en zoological code (ICZN 1999).
Alphonse Milne-Edwa ds (1880) p o ided collec ion da a,
measu emen s and sex o only one specimen o each o he
ou species he desc ibed (F.ba ba a, F. osaea, F.sigsbei, and
F. iden a a). We ne e heless canno assume ha he ou
specimens in ques ion a e he holo ypes o he ou species,
because he ne e s a ed anywhe e in he ex which we e he
ype specimens. Fu he mo e, he did no say i hese ou
specimens we e he ones he had. I is wo h no ing, howe e ,
ha when A. Milne-Edwa ds (1880: 15-16) ga e he measu e-
men s o he specimen o each o he ou species o F e illea,
he w i es “La geu de la ca apace d’un mâle” [Ca apace leng h
o one male] ( o F.ba ba a), “La geu de la ca apace d’un
[sic] emelle” [Ca apace leng h o one emale] ( o F. osaea),
“La geu de la ca apace d’une emelle” [Ca apace leng h o
one emale] ( o F. iden a a), and “La geu de la ca apace
d’une emelle cha gée d’œu s” [Ca apace leng h o one emale
wi h eggs] ( o F.sigsbei). The wo ding he used sugges s ha
he had mo e han one specimen o each species. A icle 73.1.2
o he Code also s a es ha o a specimen o be a holo ype
i should be s a ed as such o in an equi alen manne in he
o iginal publica ion. Recommenda ion 73F o he code ad ises
ha “Whe e no holo ype o syn ype was ixed o a nominal
species-g oup axon es ablished be o e 2000, and when i is
possible ha he nominal species-g oup axon was based on
mo e han one specimen, an au ho should p oceed as hough
syn ypes may exis and, whe e app op ia e, should designa e
a lec o ype a he han assume a holo ype.” Ra hbun (1918)
subsequen ly “men ioned” he ype ma e ial o he ou species
as ollows: F.ba ba a, “ ype in Pa is Mus.” and “male holo ype
(Pa is Mus.)” (Ra hbun 1918: 26); F. osaea, “holo ype in Pa is
Mus.” (Ra hbun 1918: 27); F. iden a a, “holo ype in Pa is
Mus.” (Ra hbun 1918: 29); and F.sigsbei, “holo ype in Pa is
Mus.” (Ra hbun 1918: 26). She appa en ly did no examine
hese “holo ypes” as she lis ed hem unde he “o he eco ds”
o “ ype locali y” and no as “ma e ial examined”, and all he
in o ma ion (loca ion and sex) o hese was exac ly as s a ed
in A. Milne-Edwa ds (1880). While she did no indica e how
many specimens he e we e, he ac ion ne e heless cons i u es
a alid lec o ype designa ion unde A icles 74.5 and 74.6,
e en i he eposi o y, as we now know, is inco ec . I is no
known why she assumed he “holo ypes” we e in MNHN and
no in MCZ. Ra hbun had isi ed MNHN a he beginning o
he 20 h cen u y o mainly wo k on eshwa e c abs in Pa is
(see Bou ie , in Ra hbun 1904: 225-226) bu she appa en ly
did no examine F e illea du ing ha ime. While we canno
be ce ain as he e a e no eco ds o his, i is possible ha
when Ra hbun (1918) published he wo k, all he syn ypes
o he ou species we e s ill a Pa is, and some o hem we e
only sen back o MCZ du ing o a e 1918, o mo e likely,
a e he pape o A. Milne-Edwa ds& Bou ie (1923). This
would explain why Ra hbun (1918) did no men ion he
MCZ ype ma e ial o he ou species.
The la e publica ion by A. Milne-Edwa ds& Bou ie
(1923) showed ha he e we e mo e specimens o each o
he ou species o F e illea, and iden i ied he specimen
men ioned by A. Milne-Edwa ds (1880) o be he “ ype”
(in he sense o he name-bea ing specimen) o each axon.
The designa ed “ ype” o each species is he same as hose
selec ed by Ra hbun (1918). As a as he ype s a us o he
addi ional specimens lis ed by A. Milne-Edwa ds& Bou ie
(1923), he bes solu ion is o s ic ly ollow A icles 72.4.1
and 72.4.1.1 o he ICZN code, which s a e “The ype se ies
o a nominal species-g oup axon consis s o all he specimens
included by he au ho in he new nominal axon (whe he
di ec ly o by bibliog aphic e e ence), excep any ha he
au ho exp essly excludes om he ype se ies [A . 72.4.6], o
e e s o as dis inc a ian s (e.g., by name, le e o numbe ),
o doub ully a ibu es o he axon.” and “Fo a nominal
species o subspecies es ablished be o e 2000, any e idence,
published o unpublished, may be aken in o accoun o de-
e mine wha specimens cons i u e he ype se ies”. I is clea
hen ha A. Milne-Edwa ds (1880) had mo e han one speci-
men when he made he desc ip ion e en hough he did no
men ion he o he s, and he e o e mus ha e used hem when
making his decision. In all cases, hese addi ional specimens
we e om he same Blake s a ion as he “ ype.” In addi ion,
while A. Milne-Edwa ds& Bou ie (1923) did no iden i y
he addi ional specimens as “co ypes” o he equi alen , hey
also do no explici ly exclude hem as syn ypes. As such, i
is bes o ea all he ma e ial o he species in A. Milne-Ed-
wa ds& Bou ie (1923) as syn ypes. Wi h he iden i ica ion
o a name-bea ing specimen o lec o ype o each species by
Ra hbun (1918) and A. Milne-Edwa ds& Bou ie (1923),
he emaining specimens a e de ac o pa alec o ypes.
SYSTEMATICS
Supe amily Goneplacoidea MacLeay, 1838
Family eu yplacidae S impson, 1871
Genus F e illea A. Milne-Edwa ds, 1880
F e illea A. Milne-Edwa ds, 1880: 15.
ype
species
. — F e illea ba ba a A. Milne-Edwa ds, 1880 (by
designa ion, Ra hbun 1918: 25; gende eminine).
F e illea ba ba a A. Milne-Edwa ds, 1880
(Figs1; 2)
F e illea ba ba a A. Milne-Edwa ds, 1880: 15.— Ra hbun 1898:
287.— A. Milne-Edwa ds& Bou ie 1923: 336, ig.2, pl.2, igs10,
11, pl.7, igs1, 2.— Guino 1969b: 513, pl.2, ig.2; 1971: 1080
[in lis ]; 1984: pl.1, ig.H.— Powe s 1977: 114.— So o 1985:
484, 487, 488; 1986: 3, 4, 36; 1991: 627.— Abele& Kim 1986:
54, 593, ig.a.— Boschi 2000: 82.— McLaughlin e al. 2005: 257
[in lis ].— Ng e al. 2008: 78 [in lis ].— Felde e al. 2009: 1081
[in lis ].— Thoma e al. 2009: 553, 563, igs1, 2.— Cas o&
Ng 2010: 53 (pa ).

736 ZOOSYSTEMA • 2024 • 46 (29)
Ng P. K. L. e al.
Goneplax ba ba a – Ra hbun 1918: 25, 26, pl.4, igs1, 3, pl.5.
F e illea osaea – Cas o& Ng 2010: 56 (pa ) (no F. osaea
A.Milne-Edwa ds, 1880).
No F e illea ba ba a – Poupin 1994: 48, pl.5e (= F.hi su a (Bo -
adaile, 1916)).
H
is o y
.— F e illea ba ba a was desc ibed om Blake’s s a-
ion36 in he Gul o Mexico (23°13’N, 89°16’W) om a dep h
o 84 a homs (= 154m) (A. Milne-Edwa ds 1880: 15). Measu e-
men s we e p o ided o one male (26.0 × 17.0mm), bu no ype
was indica ed (see abo e). Fo he species, Ra hbun (1918: 26)
s a ed: “ ype-locali y, la . 23° 13’ N.; long. 89° 16’ W., 84 a homs,
A
B
C
DE
FG
H
ig. 1
. — F e illea ba ba a A. Milne-Edwa ds, 1880: A-D, lec o ype ♂ (29.5 × 18.4 mm) (MCZ-IZ-CRU-9155); E, pa alec o ype ♀ (27.3 × 17.2 mm) (MCZ-IZ-
CRU-9155); F, pa alec o ype ♂ (16.5 × 10.0 mm) (MNHN-IU-2014-7505); G, pa alec o ype ♀ (22.7 × 15.7 mm) (MNHN-IU-2014-7506); H, ju enile pa alec o ype
♀ (7.9 × 5.0 mm) (MNHN-IU-2014-7506): A, D-H, o e all do sal iews; B, igh chela; C, pleon. C edi s: A-C a e A. Milne-Edwa ds & Bou ie (1923: ig. 2; pl. 7,
igs 1, 2); D, E, pho og aphs by Alana Ri e a, MCZ; , F-H, pho og aphs by P. K. L. Ng.
737
Taxonomy o F e illea and desc ip ion o Diuplax n. gen.
ZOOSYSTEMA • 2024 • 46 (29)
s a ion 36, Blake; ype in Pa is Mus.” and “Yuca an Channel; la .
23° 13’ 00”N, long. 89° 16’ 00” W.; 84 a homs; emp. 60° F.;
s a ion 36, U. S. C. S. S , Blake; male holo ype (Pa is Mus.).”;
and al hough she did no examine he specimen, he ac ion con-
s i u es he i s alid designa ion o a lec o ype o he species.
Ra hbun (1918: 26), howe e , did examine ano he Blake speci-
men in MCZ: “O G enada; la . 12° 01’ 45” N.; long. 61°47’
25” W.; 92 a homs; ne. S.; Ma . 1, 1879; s a ion 262, Blake;
1male (Ca . No. 4116. M. C. Z.).” Alphonse Milne-Edwa ds&
Bou ie (1923: 336) elabo a ed ha hey s udied one “ ype” male,
one imma u e male, wo adul emales, and wo ju eniles. This
a i ied he name-bea ing designa ion made by Ra hbun (1918).
Alphonse Milne-Edwa ds& Bou ie (1923) did no s a e he
size o he ype male (only one adul male is men ioned), bu we
AB
C
D
E
F
ig. 2
. — F e illea ba ba a A. Milne-Edwa ds, 1880: A, B, F, pa alec o ype ♂ (16.5 × 10.0 mm) (MNHN-IU-2014-7505); C, pa alec o ype ♀ (27.3 × 17.2 mm)
(MCZ-IZ-CRU-9155); D, lec o ype ♂ (29.5 × 18.4 mm) (MCZ-IZ-CRU-9155); E, pa alec o ype ♀ (22.7 × 15.7 mm) (MNHN-IU-2014-7506): A, le side o ca a-
pace and le cheliped; B, igh chela; C-E, en al iew o cephalo ho ax and pleon; F, ♂ s e nopleonal ca i y. C edi s: A, B, E, F pho og aphs by P. K. L. Ng;
C, D, pho og aphs by Alana Ri e a, MCZ.
738 ZOOSYSTEMA • 2024 • 46 (29)
Ng P. K. L. e al.
p esume i is he one A. Milne-Edwa ds o iginally measu ed, i.e.,
26.0 × 17.0mm. They igu ed he male pleon o he ype male
(A. Milne-Edwa ds& Bou ie 1923: ig.2), as well as he ca a-
pace and cheliped (pl.7, igs1, 2) and he ou h ambula o y leg
(pl.2, igs10, 11) (Fig.1A-C). In any case, appa en ly Ra hbun
(1918) and A. Milne-Edwa ds& Bou ie (1923) selec ed he
same specimen as he lec o ype.
Alphonse Milne-Edwa ds& Bou ie (1923: 337) also commen ha
“En deho s de la locali é p écéden e qui se ou e dans le Gol e du
Mexique au N.du Canal du Yuca an, l’espèce a é é signalée, a ec
dou e, au la ge de La Ha ane pa Mlle Ra hbun.” [Apa om he
p e ious locali y, which is in he Gul o Mexico, N. o he Yuca an
Channel, he species has been epo ed, wi h doub , o Ha ana by
Miss Ra hbun.] (see Ra hbun 1898). Guino (1969b: 512, pl.12
ig.2) nex eco ded and igu ed a “co ype” emale measu ing 16.5 ×
10.0mm om s a ion 36. Cas o& Ng (2010: 53) lis ed one ju-
enile male (ca apace wid h 5.9mm, MNHN-B8733), and one
male, one emale and one ju enile (MCZ-IZ-CRU-9155). Guino ’s
(1969b) “co ype” emale measu ing 16.5 × 10.0mm om s a ion
36 is ac ually a male (see Ma e ial Examined below), al hough he
gonopods a e missing.
Du ing he examina ion o ma e ial by he i s au ho in May 2023
in he MNHN bo le labelled as “F e illea ba ba a” (MNHN-
B8733), he ound only one small ju enile male measu ing 7.8 ×
5.9mm and labelled as he lec o ype. In he ca d ca alogues in
MNHN (unde MP B8733), only one co ype emale measu ing
16.5 × 10.0mm om he Gul o Mexico is lis ed. The locali y
and s a ion labels in he bo le ma ch hose gi en by A. Milne-
Edwa ds (1880), and hey include he o iginal hand-w i en labels
a A. Milne-Edwa ds ime om he Blake c uise. We a e ce ain
his specimen is no F.ba ba a, bu F. osaea (see below in Tax-
onomy sec ion). The ma e ial in MCZ consis s o one ca alogued
lo (MCZ-IZ-CRU-9155) wi h h ee specimens: one adul male
(29.5 × 18.4mm), one badly damaged ju enile male (ca apace
wid h c. 9.0mm), and one adul emale (27.3 × 17.2mm), all
being lis ed in he labels as “co ypes” and in he da abase as “syn-
ypes” (Figs1D, E; 2D).
ype Ma e ial.— The 26.0 × 17.0mm male o A. Milne-Edwa ds
(1880) is he lec o ype o F e illea ba ba a because he did no iden i y
a ype specimen and did no indica e how many specimens he had
(see abo e). This specimen is clea ly he p esen MCZ male speci-
men measu ing 29.5 × 18.4mm (MCZ-IZ-CRU-9155); i ag ees
e y well wi h he o iginal igu es (A. Milne-Edwa ds& Bou ie
1923: pl.7, igs1, 2) (Figs1D; 2D). The p esen disc epancy in
measu emen s can be explained by whe he he ca apace wid h
was measu ed a he wides poin o he ex e nal o bi al ee h o
ac oss hei bases; and i he leng h was aken medially, whe e he
pos e io ma gin is dis inc ly conca e, o a i s la e al pa which
is mo e conca e. As no ed abo e, he subsequen i e specimens
epo ed by A. Milne-Edwa ds& Bou ie (1923) should be now
ega ded as pa alec o ypes.
Ma e ial exaMined.— Lec o ype (by in e ence o holo ype by
Ra hbun 1918: 26, as Goneplax ba ba a; and A. Milne-Edwa ds&
Bou ie 1923: 336 as F.ba ba a). Mexico • ♂ (29.5 × 18.4mm); s a-
ion 36, no h o Alac an Ree , Campeche Bay, Yuca an; 84 a homs
(= 154m); 23°13’0”N, 89°16’0”W; coll. USCGS; Geo ge S.Blake
c uise; 19-24.I.1878; MCZ-IZ-CRU-9155.
Pa alec o ypes. Mexico • 1 ♀ (27.3 × 17.2mm), 1 badly damaged
ju enile ♂ (c. 9.0mm ca apace wid h); same da a as o lec o ype;
MCZ-IZ-CRU-9155) • 1 ♂ (16.5 × 10.0mm); same da a as o
lec o ype; MNHN-IU-2014-7505) (= MNHN-B8733) • 1 ♀ (22.7×
15.7mm), 1 ju enile ♀ (7.9 × 5.0mm); same da a as o lec o ype;
MNHN-IU-2014-7506) (= MNHN-B8733).
G
eoG apHical
dis ibu ion
.— Wes e n A lan ic egion: G enada
and Gul o Mexico coas s o Flo ida and Mexico.
axonoMy
When he i s au ho compa ed he specimens in he MNHN
bo les now labelled as “F e illea ba ba a” and “F e illea
osaea” wi h he desc ip ions and he o iginal igu es o he
species in A. Milne-Edwa ds (1880) and A. Milne-Edwa ds&
Bou ie (1923), i was ob ious ha he specimens had been
swi ched. In any case, F. osaea was desc ibed only om wo
specimens, so he h ee specimens in he bo le clea ly do
no ma ch.
In sepa a ing F. osaea om F.ba ba a, A. Milne-Edwa ds
(1880: 15) no ed ha “Ce e espèce se dis ingue de la p é-
céden e pa sa ca apace plus épaisse e moins éla gie en
a an ; les bo ds la é aux é an p esque pa allèles. Le on
es plus la ge e à bo d plus d oi . Les pédoncules oculai es
son plus g os e plus cou s. L’angle pos o bi ai e es o mé
pa une den poin ue, en a iè e de laquelle exis e un pe i
en lemen ube culi o me puis une épine hépa ique cou e
mais acé ée. Les pinces e les pa es ambula oi es son dis-
posées comme chez le F e illea ba ba a.” [This species is
dis inguished om he p eceding by i s hicke ca apace
and less widened (na owe ) in on ; he side edges being
nea ly pa allel. The on is wide and wi h a s aigh e edge.
The eyes alks a e la ge and sho e . The pos o bi al angle
is o med by a poin ed oo h, behind which he e is a small
ube culi o m swelling and hen a sho bu sha p hepa ic
spine. Claws and walking legs a e a anged as in F e illea
ba ba a.]. Alphonse Milne-Edwa ds& Bou ie (1923: 337)
added ha while F.ba ba a and F. osaea we e close, “Aux
obse a ions p écéden es j’ajou e ai que les pédoncules
oculai es de ce e espèce son ès dila és dans leu égion
co néenne, que les deux échanc u es du bo d o bi ai e su-
pé ieu son ès accen uées, su ou la plus in e ne e que
les pa es ambula oi es semblen un peu moins o es que
celles du F e illea ba ba a. A pa quelques di é ences ès
légè es, ous les au es ca ac è es son iden iques à ceux de
ce e de niè e espèce, du moins dans le ype cha gé d’œu s e
dans le ès jeune exemplai e qui m’on é é soumis.” [To he
p eceding obse a ions I will add ha he ocula peduncles
o his species a e e y dila ed in hei co neal egion, ha
he wo inden a ions o he supe io o bi al edge a e e y
accen ua ed, especially he mos in e nal one and ha he
ambula o y legs seem a li le less s ong han hose o he
F e illea ba ba a. Apa om a ew e y sligh di e ences,
all o he cha ac e s a e iden ical wi h hose o he la e
species, a leas in he egg-laden ype and in he e y young
specimen p o ided o me.].
The cha ac e s o he p esen MNHN ju enile male la-
belled as “F e illea ba ba a” (7.8 × 5.9mm, MNHN-B8733)
(Fig.3E-G) ac ually ma ch hose desc ibed and igu ed
as F. osaea by A. Milne-Edwa ds (1880) and A. Milne-
Edwa ds& Bou ie (1923) exac ly. Con e sely, he cha -
ac e s o he h ee MNHN specimens labelled as “F e illea
osaea” (MNHN-B10152), ma ch hose o F e illea ba ba a.
How he specimens became swi ched in MNHN emains
unknown and could ha e occu ed du ing examina ion by
esea che s o o he cu a o ial/digi isa ion p ocesses. The
second au ho had examined he MNHN ma e ial be ween
739
Taxonomy o F e illea and desc ip ion o Diuplax n. gen.
ZOOSYSTEMA • 2024 • 46 (29)
Ma ch and June 2006, and he ma e ial had al eady been
mixed, as con i med by no es and ske ches o he wo spe-
cies made a he ime. As such, he eco d o F.ba ba a (and
F. osaea) mus be ega ded as swi ched as hey e e ed o
bo h he ypes as well as he ma e ial hey examined a ha
ime. The e a e no p oblems wi h he MCZ specimens,
wi h all he specimens o he wo species ag eeing wi h he
o iginal desc ip ions.
The MNHN male specimen (16.5 × 10.0mm) now labelled
as “F e illea osaea” (MNHN-B10152) ac ually closely ma ches
wha Guino (1969b: pl.12 ig.2) igu ed as he MNHN
“co ype” emale o F.ba ba a om s a ion 36 (no ca alogue
numbe indica ed). They ag ee in measu emen s and he
specimen ag ees well in all physical ea u es. We belie e hey
a e he same specimen. As such, a e he p esen s udy o he
li e a u e and analysis o mo phological cha ac e s o MNHN
specimens o he wo species in ques ion, he specimens we e
e u ned o hei co ec ja s.
F e illea ba ba a A. Milne-Edwa ds, 1880 is he ype spe-
cies o F e illea (c . Cas o& Ng 2010), amily Eu yplacidae
S impson, 1871 (see Ra hbun 1918: 25).
F e illea osaea A. Milne-Edwa ds, 1880
(Fig.3)
F e illea osaea A. Milne-Edwa ds, 1880: 15.— A. Milne-Edwa ds&
Bou ie 1923: 337, pl.6, ig.1.— Chace 1940: 47.— Guino
1969b: 513; 1971: 1080 [lis ].— McLaughlin e al. 2005: 257.—Ng
e al. 2008: 78.— Cas o& Ng 2010: 56 (pa ).
F e illea osae [sic] – So o 1986: 3, 4, 33; 1991: 627.
F e illea osea [sic] – Felde e al. 2009: 1081.
Goneplax osaea – Ra hbun 1918: 25, 27.— Chace 1940: 47.
F e illea ba ba a – Cas o& Ng 2010: 53 (pa ). (no F.ba ba a
A. Milne-Edwa ds, 1880).
His o y.— F e illea osaea was desc ibed om Blake’s s a ion232
o S . Vincen , Lesse An illes, om a dep h o 88 a homs (161m)
(A. Milne-Edwa ds, 1880: 15). He p o ided measu emen s o one
emale, 20.0 × 15.0mm, bu no ype was indica ed. Fo he species,
Ra hbun (1918: 27) s a ed: “ ype-locali y, S . Vincen , 88, co -
ec ly 87, a homs, s a ion 232, Blake; holo ype in Pa is Mus.” and
al hough she did no examine he specimen, he ac ion cons i u es
he i s alid designa ion o a name-bea ing specimen o lec o ype
o he species. Alphonse Milne-Edwa ds& Bou ie (1923: 337)
la e commen ed ha hey had one ype o ige ous emale (has o be
he one wi h he measu emen s 20.0 × 15.0mm) and one ju enile.
They igu ed he do sal ea u es, including he ca apace o he ype
emale (A. Milne-Edwa ds& Bou ie 1923: pl.6, ig.1) (Fig.3A).
In addi ion, hey w o e: “A pa quelques di é ences ès légè es,
ous les au es ca ac è es son iden iques à ceux de ce e de niè e
espèce, du moins dans le ype cha gé d’œu s e dans le ès jeune
exemplai e qui m’on é é soumis.” [Apa om a ew e y sligh
di e ences, all he o he cha ac e s a e iden ical o hose o he la -
e species, a leas in he egg-bea ing ype and in he e y young
specimen passed o me.] (A. Milne-Edwa ds& Bou ie 1923: 337).
These commen s also cons i u e a name-bea ing o lec o ype desig-
na ion, e en i done a e Ra hbun (1918). Bo h Ra hbun (1918)
and A. Milne-Edwa ds& Bou ie (1923) appa en ly selec ed he
same specimen o be he lec o ype.
Chace (1940: 47) subsequen ly eco ded he species om Cuba.
Cas o& Ng (2010: 56) lis ed one emale “lec o ype” (22.7 ×
15.7mm), one “pa alec o ype” male (16.3 × 10.1mm) and one
p e-adul emale “pa alec o ype” (7.9 × 5.0mm) (MNHN-B10152)
o he species.
Du ing he examina ion o ma e ial by he i s au ho in May 2023,
in he MNHN bo le labelled as “F e illea osaea” (MNHN-B10152),
he ound h ee specimens: a male (16.5 × 10.0mm, wi hou pleon
and gonopods), a emale (22.7 × 15.7mm), and a ju enile emale
(7.9 × 5.0mm) (Figs1F-H; 2A, B, E, F). The adul emale (22.7×
15.7mm) (Figs1G; 2E) is labelled as he “lec o ype” o he spe-
cies, wi h he o he wo “pa alec o ypes.”. In he ca d ca alogues in
MNHN, h ee specimens a e lis ed om S . Vincen , o which one
ju enile is lis ed as he co ype (MP B10152). The locali y labels
in he bo le ma ch hose gi en by A. Milne-Edwa ds (1880), bu
he e a e no o iginal labels om he Blake c uise. As discussed ea lie
unde F.ba ba a, we a e ce ain ha hese h ee specimens do no
belong o F. osaea bu o F.ba ba a ins ead. The ma e ial in MCZ
consis s o only one o ige ous emale specimen (22.1 × 15.4mm)
(MCZ-IZ-CRU-9167) (Fig.3A-D) and is labelled as a “syn ype”
in he da abase.
ype Ma e ial.— On he basis o he published in o ma ion, he
20.0 × 15.0mm emale epo ed by A. Milne-Edwa ds (1880) is he
lec o ype o F. osaea. This specimen is clea ly he MCZ o ige ous
emale specimen measu ing 22.1 × 15.4mm (MCZ-IZ-CRU-9167)
(Fig.3A-D). The disc epancies in measu emen s a e mino and can
easily be accoun ed o by di e ences in how he measu emen s we e
made. Mo phologically, i ag ees e y well wi h he o iginal igu es
in A. Milne-Edwa ds& Bou ie (1923: pl.6 ig.1). The ju enile
specimen epo ed by A. Milne-Edwa ds& Bou ie (1923) should
be ega ded as a pa alec o ype.
Ma e ial exaMined.— Lec o ype (by in e ence o holo ype by
Ra hbun 1918: 27, as Goneplax osaea; A. Milne-Edwa ds& Bou ie
1923: 337 as F. osaea). Sain Vincen and he G enadines • o ige -
ous ♀ (22.1 × 15.4mm); s a ion232; o S . Vincen ; 88 a homs
(=161m); 13°06’45”N, 61°06’55”W; coll. USCGS; Geo ge S. Blake
c uise; 21.II.1879; MCZ-IZ-CRU-9167.
Pa alec o ype. Sain Vincen and he G enadines • 1 ju enile ♂
(7.8 × 5.9mm); same da a as lec o ype; MNHN-IU-2014-19822
(= MNHN-B10152).
GeoG apHical dis ibu ion.— Wes e n A lan ic egion: Cuba
and S . Vincen .
axonoMy
See he ea lie discussion on he con usion o he ypes o
F.ba ba a and F. osaea.
F e illea osaea A. Milne-Edwa ds, 1880 is included in
F e illea s.s . as ede ined by Cas o& Ng (2010), amily
Eu yplacidae S impson, 1871.
Family pseudo HoMbilidae Alcock, 1900
Genus T apezioplax Guino , 1969
T apezioplax Guino , 1969: 712.
ype species. — F e illea iden a a A. Milne-Edwa ds, 1880, by
o iginal designa ion; gende eminine.
746 ZOOSYSTEMA • 2024 • 46 (29)
Ng P. K. L. e al.
male pl 2 ( ig.72), longe han he pl 1. In sigsbei ( ig.63),
as in homboides ( ig.64), he male o i ice opens on he coxa
o p5 bu , as a esul o he beginning o he union o s e -
ni es 8 and 7, he penis is in an impe ec ly closed gu e ,
o med be ween hese wo egions. I should be no ed ha
in homboides, he la e al po ion o s e ni e 8 no co e ed
by he abdomen is impo an and is widely inse ed be ween
he en i e basila , na ow pa o he abdomen (segmen s 1
and 2) and he coxae o p5; on he o he hand, in sigsbei, he
i s abdominal segmen s, mo e ex ended, occupy p ac ically
all he space be ween he coxae o p5, wi h he excep ion o
a small pa o s e ni e 8 which is le unco e ed. Mo eo e ,
i does no seem ha in sigsbei he lagellum o pl 2, which is
cu ed, is bi id as in G. homboides.]
F e illea sigsbei A. Milne-Edwa ds, 1880 is clea ly a mem-
be o Goneplacidae MacLeay, 1838, s.s . (c . Cas o 2007).
Compa ing he specimens o F.sigsbei also shows ha Gui-
no ’s (1969b) doub s abou ans e ing i o Goneplax a e
jus i ied. The di e ences in he pleon, s e nal and gonopodal
s uc u es, as well as di e ences in he ca apace and chelipeds
wi h Goneplax s.s . a gue o ea ing F.sigsbei as a sepa a e
new genus. I is he e made he ype species o a new genus,
Diuplax n.gen.
Acknowledgemen s
We a e g a e ul o museum echnician Lau a Flamme (un-
de MNHN P oje : RECOLNAT (ANR-11-INBS-0004)
who i s highligh ed he p oblems o he ypes o F.ba ba a
and F. osaea o he second au ho . Thanks a e due o Adam
Baldinge (MCZ) o checking on he specimens in MCZ,
and o Alana Ri e a (Cu a o ial Assis an ) o aking he pho-
og aphs o he ma e ial. We would also like o hank Lau e
Co ba i (Cu a o o C us acea, MNHN) o access o he
ABC
DE
FG
ig. 7. — Goneplax homboides (Linnaeus, 1758), ♂ (36.5 × 22.6 mm) (ZRC 2010.0069), Sco land. A, do sal iew o ca apace; B, on al iew o cephalo ho ax;
C, le hi d maxilliped; D, an e io ho acic s e num and pleon; E, pos e io ho acic s e num and pleon; F, igh me us o cheliped (inne iew); G, le ou h
ambula o y leg. C edi s: pho og aphs by P. K. L. Ng.

747
Taxonomy o F e illea and desc ip ion o Diuplax n. gen.
ZOOSYSTEMA • 2024 • 46 (29)
collec ions. The i s and second au ho s also hank Danièle
Guino o he hospi ali y and ad ice on goneplacoids du ing
ou isi s o MNHN, as well as he help wi h he manusc ip .
She and Shane Ahyong kindly e iewed he manusc ip , o
which we a e mos g a e ul.
REFERENCES
abele l. G. & kiM W. 1986.— An Illus a ed Guide o he Ma ine
Decapod C us aceans o Flo ida. Depa men o En i onmen al
Regula ion S a e o Flo ida and Flo ida S a e Uni e si y Tal-
lahassee Technical Se ies 8 (1) Pa s 1, 2, 760p.
Alcock a. 1900.— Ma e ials o a ca cinological auna o India.
No. 6. The B achyu a Ca ome opa, o G apsoidea. Jou nal on he
Asia ic Socie y o Bengal 69 (2): 279-456. h ps://doi.o g/10.5962/
bhl. i le.15344
bo adaile l. a. 1916.— Decapoda C us acea pa 1, in B i -
ish An a c ic (“Te a No a”) Expedi ion 1910. Na u al His o y
Repo s Zoology B i ish Museum (Na u al His o y) London ol.3
A h opoda (2): 75-110. h ps://www.biodi e si ylib a y.o g/
bibliog aphy/111654
boscHi e. 2000.— Species o decapod c us aceans and hei dis i-
bu ion in he Ame ican ma ine zoogeog aphic p o inces. Re is a
de In es igación y Desa ollo Pesque o 13: 1-136.
cas o p. 2007.— A eapp aisal o he amily Goneplacidae
MacLeay 1838 (C us acea Decapoda B achyu a) and e i-
sion o he sub amily Goneplacinae wi h he desc ip ion o
10 new gene a and 18 new species. Zoosys ema 29 (4): 609-
774. h ps://www.biodi e si ylib a y.o g/page/7637388
cas o p. & nG p. k. l. 2010.— Re ision o he amily Eu yplacidae
S impson 1871 (C us acea: Decapoda: B achyu a: Goneplacoidea).
Zoo axa 2375: 1-130. h ps://doi.o g/10.11646/zoo axa.2375.1.1
cHace F. a. 1940.— The b achyu an c abs in Repo s on he
scien i ic esul s o he A lan is Expedi ions o he Wes Indies
unde he join auspices o he Uni e si y o Ha ana and Ha a d
Uni e si y. To eia 4: 1-67.
da ie p. J. F., Guino d. & nG p. k. l. 2015.— Ana omy
and unc ional mo phology o B achyu a (Chap e 71-2), in
c
as o
p., d
a ie
p. J. F., G
uino
d., s
cH aM
F. . &
on
aupel klein J. C. (eds), Decapoda: B achyu a, T ea ise on
Zoology– Ana omy, Taxonomy, Biology. Vol. 9C-I. B ill, Leiden
and Bos on: 11-163.
Felde d. l., Ál a ez F., Goy J. W. & leMai e . 2009.—
Decapoda (C us acea) o he Gul o Mexico wi h commen s
on he Amphionidacea. Chap e 59, in
unnell
J. W. J
.
Felde d. l. & ea le s. a. (eds), Gul o Mexico O igin Wa e s
and Bio a ol. 1 Biodi e si y Ha e Resea ch Ins i u e o Gul
o Mexico S udies Se ies Texas A&M Uni e si y P ess Texas
USA: 1019-1104.
Guino d. 1969a.— Reche ches p éliminai es su les g oupemen s
na u els chez les C us acés Décapodes B achyou es. VII Les Gon-
eplacidae. Bulle in du Muséum na ional d’His oi e na u elle Pa is
sé . 2 41(1): 241-265, pl.2. h ps://www.biodi e si ylib a y.
o g/page/55599183
G
uino
d. 1969b.— Reche ches p éliminai es su les g oupe-
men s na u els chez les C us acés Décapodes B achyou es VII.
Les Goneplacidae (sui e). Bulle in du Muséum na ional d’His oi e
na u elle sé . 2, 41 (2): 507-528, pl.2. h ps://www.biodi e si-
ylib a y.o g/page/55599454
G
uino
d. 1969c.— Reche ches p éliminai es su les g oupe-
men s na u els chez les C us acés Décapodes B achyou es VII.
Les Goneplacidae (sui e e in). Bulle in du Muséum na ional
d’His oi e na u elle sé . 2, 41: 688-724, pls3-5. h ps://www.
biodi e si ylib a y.o g/page/55599642
Guino d. 1971.— Reche ches p éliminai es su les g oupemen s
na u els chez les C us acés Décapodes B achyou es VIII. Syn hèse
e bibliog aphie. Bulle in du Muséum na ional d’His oi e na u elle
sé . 2, 42(5) [1970]: 1063-1090. h ps://www.biodi e si ylib a y.
o g/page/56079257
Guino d. & cas o p. 2007.— A new species o Goneplax Leach
1814 (C us acea Decapoda B achyu a Goneplacidae) om he
sou h A lan ic and wes e n limi s o he Indo-Wes Paci ic egion
long con used wi h G. homboides (Linnaeus 1758). Zoo axa 1577:
17-31. h ps://doi.o g/10.11646/zoo axa.1577.1.3
G
uino
d.,
a a es
M. & c
as o
p. 2013.— Signi icance o he
sexual openings and supplemen a y s uc u es on he phylogeny
o b achyu an c abs (C us acea Decapoda B achyu a) wi h new
nomina o highe - anked podo eme axa. Zoo axa 3665 (1):
1-414. h ps://doi.o g/10.11646/zoo axa.3665.1.1
He Moso-salaza M. & a izu-coyo zi k. 2015.— C us áceos
del Sis ema A eci al Ve ac uzano, in G anados-ba ba a., o iz-
l
ozano
l. d., s
alas
-M
on eal
d. & G
onzalez
-G
Ánda a
c.
(eds), Apo es al conocimien o del Sis ema A eci al Ve ac uzano:
Hacia el Co edo A eci al del Su oes e del Gol o de México. Uni-
e sidad Ve ac uzana Ve a C uz Mexico: 47-74.
iczn, i
n e na ional
c
oMMission
on
z
ooloGical
n
oMen
-
cla u e. 1999.— In e na ional Code o Zoological Nomencla-
u e. In e na ional Commission o Zoological Nomencla u e.
Fou h Edi ion. Adop ed by he XXI Gene al Assembly o he
In e na ional Union o Biological Sciences. In e na ional T us
o Zoological Nomencla u e in associa ion wi h he B i ish
Museum (Na u al His o y) London 338p. h ps://www.iczn.
o g/ he-code/ he-code-online/
l
eacH
W. e. 1814.— C us aceology, in The Edinbu gh Encyclopaedia
conduc ed by Da id B ews e LLD ol. 7 (2) pp. 385-487, 765-
766 (pla e legend) (pla e CCXXI issued wi h ol. 9 (1) in 1815).
Edinbu gh. h ps://www.biodi e si ylib a y.o g/page/37187274
l
innaeus
c. 1758.— Sys ema Na u ae pe Regna T ia Na u ae
Secundum Classes O dines Gene a Species cum Cha ac e ibus Di -
e en iis Synonymis Locis Tomus I. Lau en ii Sal ii, Holmi., [i ]
+ 824p. h ps://doi.o g/10.5962/bhl. i le.542
Macleay W. s. 1838.— On he B achyu ous Decapod C us acea.
B ough om he Cape by D . Smi h, in s
Mi H
a., Illus a ions o
he Zoology o Sou h A ica; consis ing chie ly o igu es and desc ip-
ions o he objec s o na u al his o y collec ed du ing an expedi ion
in o he in e io o Sou h A ica in he yea s 1834 1835 and 1836;
i ed ou by ‘The Cape o Good Hope Associa ion o Explo ing
Cen al A ica’: oge he wi h a summa y o A ican Zoology and
an inqui y in o he geog aphical anges o species in ha qua e o
he globe published unde he Au ho i y o he Lo ds Commissione s
o He Majes y’s T easu y In e eb a ae. IV [1849], Smi h Elde &
Co. London: 53-71, pls2, 3. h ps://www.biodi e si ylib a y.
o g/page/11059737
MclauGHlin p. a., caMp d. k., anGel M. ., bousField e. l.,
b
unel
p., b
usca
. c., c
adien
d., c
oHen
a. c., c
onlan
k.,
eld edGe l. G., Felde d. l., Goy J. W., Haney ., Hann b.,
Hea d . W., Hend ickx e. a., Hobbs H. H., HolsinGe
J. ., kensley b., laubi z d. ., lec oy s. e., leMai e .,
Maddocks . F., Ma in J. W., nikkelsen p., nelson e.,
neWMan W. a., o e s ee . M., poly W. J., p ice W. W.,
eid J. W., obe son a., oGe s d. c., oss a., scHo e
M., s
cH aM
F. ., s
HiH
c.- ., W
a linG
l., W
ilson
G. d. F. &
u Geon d. d. 2005.— Common and scien i ic names o aqua ic
in e eb a es om he Uni ed S a es and Canada: C us aceans.
Ame ican Fishe ies Socie y, Be hesda, Ma yland, USA, Special
Publica ion 31: xiii + 545. a k:/13960/ 7wn09 95
M
ilne
-e
dWa ds
a. 1880.— É udes p éliminai es su les C us-
acés 1e e pa ie, in Repo s on he esul s o d edging unde
he supe ision o Alexande Agassiz in he Gul o Mexico and
in he Ca ibbean Sea 1877 ’78 ‘79 by he U.S. Coas Su ey
S eame “Blake” Lieu .-Commande C.D. Sigsbee U.S.N. and
Commande J.R. Ba le U.S.N. commanding VIII. Bulle in o
he Museum o Compa a i e Zoology a Ha a d College 8: 1-68,
pls1, 2. h ps://www.biodi e si ylib a y.o g/page/30184918
748 ZOOSYSTEMA • 2024 • 46 (29)
Ng P. K. L. e al.
M
ilne
-e
dWa ds
a. & b
ou ie
e.-l. 1923.— Les Po cellanides e
des B achyou es, in Repo s on he esul s o d edging unde he
supe ision o Alexande Agassiz in he Gul o Mexico (1877-
78) in he Ca ibbean Sea (1878-79) and along he A lan ic coas
o he Uni ed S a es (1880) by he U.S. Coas Su ey S eame
“Blake” Lieu .-Comm. C.D. Sigsbee U.S.N. and Commande
J.R. Ba le U.S.N. commanding XLVII. Memoi s o he Museum
o Compa a i e Zoology a Ha a d College 47 (4): 281-395,
pls1-12. h ps://www.biodi e si ylib a y.o g/page/4368457
nG p. k. l., Guino d. & da ie p. J. F. 2008.— Sys ema B achyu-
o um: Pa I. An anno a ed checklis o ex an b achyu an c abs
o he wo ld. Ra les Bulle in o Zoology Supplemen 17: 1-286.
nizinski M. 2003.— Anno a ed checklis o decapod c us aceans o
A lan ic coas al and con inen al shel wa e s o he U.S. P oceed-
ings o he Biological Socie y o Washing on 116: 96-157. h ps://
www.biodi e si ylib a y.o g/page/34564867
poupin J. 1994.— Faune ma ine p o onde des An illes ançaises.
Récol es du na i e Polka ai es en 1993. É udes e Thèses 1-79, pls1-5.
poWe s l. W. 1977.— A ca alogue and bibliog aphy o he c abs
(B achyu a) o he Gul o Mexico. Con ibu ions in Ma ine
Science [Uni e si y o Texas Ma ine Science Ins i u e] Supple-
men 20: 1-190.
a Hbun M. J. 1898.— The B achyu a o he biological expedi-
ion o he Flo ida Keys and he Bahamas in 1893. Bulle in o
he Labo a o ies o Na u al His o y o he S a e Uni e si y o Iowa
4 (3): 250-294, pls1-9. h ps://www.biodi e si ylib a y.o g/
page/15405204
a Hbun M. J. 1904.— Les c abs d’eau douce, (Po amonidae).
Nou elles A chi es du Muséum na ional d’His oi e na u elle, Sé ie4,
6: 225-312, pls 9-18.
a Hbun M. J. 1918.— The g apsoid c abs o Ame ica. Bulle in
o he Uni ed S a es Na ional Museum 97: xxii + 461, pls1-161.
h ps://www.biodi e si ylib a y.o g/page/7637388
so o l. a. 1985.— Dis ibu ional pa e ns o deep-wa e b achy-
u an c abs in he S ai s o Flo ida. Jou nal o C us acean Biology
5 (3): 480-499. h ps://doi.o g/10.2307/1547920
so o l. a. 1986.— Deep-wa e b achyu an c abs o he S ai s
o Flo ida (C us acea Decapoda). Anales del Ins i u o de Ciencias
del Ma y Limnología, Uni e sidad Nacional Au ónoma de México
13 (1): 1-68.
so o l. a. 1991.— Faunal zona ion o he deep-wa e b achy-
u an c abs in he S ai s o Flo ida. Bulle in o Ma ine Science
49 (1-2): 623-637.
s
iMpson
W. 1871.— B achyu a. P elimina y epo on he C us-
acea d edged in he Gul S eam in he S ai s o Flo ida, by L.
F. Pou ales, Assis . U.S. Coas Su eys, Pa 1. Bulle in o he
Museum o Compa a i e Zoology a Ha a d College 2: 109-160.
h ps://www.biodi e si ylib a y.o g/page/6313618
HoMa b. p., scHuba c. d. & Felde d. l. 2009.— Molecu-
la phylogeny o Wes e n A lan ic ep esen a i es o he genus
Hexapanopeus (Decapoda: B achyu a: Panopeidae), in Ma in J.,
c andall J. W. k. & Felde d. l. (eds). Decapod C us acean
Phylogene ics. C us acean Issues 18: 551-565.
HoMa b. p., Guino d. & Felde d. l. 2014.— E olu iona y
ela ionships among Ame ican mud c abs (C us acea: Decapoda:
B achyu a: Xan hoidea) in e ed om nuclea and mi ochond ial
ma ke s. Zoological Jou nal o he Linnean Socie y 170: 86-109.
h ps://doi.o g/10.1111/zoj12093
WilliaMs a. b. 1984.— Sh imps Lobs e s, and C abs o he A lan ic
Coas o he Eas e n Uni ed S a es Maine o Flo ida. Smi hsonian
Ins i u ion P ess Washing on D. C. x iii + 550p. h ps://doi.
o g/10.2307/1352125
WilliaMs a. b., Mccloskey l. . & G ay i. e. 1968.— New
eco ds o b achyu an decapod c us aceans om he con inen-
al shel o No h Ca olina U.S.A. C us aceana 15 (1): 41-66.
h ps://doi.o g/10.1163/156854068X00755
Submi ed on15Decembe 2023;
accep ed on 15Ap il 2024;
published on 12No embe 2024.