Memoirs of Museum Victoria 78: 73–146 (2019) Published 2019
1447-2554 (On-line)
https://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/
DOI https://doi.org/10.24199/j.mmv.2019.78.05
A new classification of Callianassidae and related families (Crustacea: Decapoda:
Axiidea) derived from a molecular phylogeny with morphological support
(http://zoobank.org/urn:lsid:zoobank.org:pub:263C1363-0ADA-4972-9224-AC690A1FD238)
Gary C. B. Poore1 (http://zoobank.org/urn:lsid:zoobank.org:author:c004d784-e842-42b3-bfd3-317d359f8975),
Peter C. DworsChak 2 (http://zoobank.org/urn:lsid:zoobank.org:author:4BCD9429-46AF-4BDA-BE4B-439EE6ADC657),
rafael roBles 3, 4, 5 (http://zoobank.org/urn:lsid:zoobank.org:author:7F54A0F4-892F-407A-AED4-4E65EA8B9360)
fernanDo l. Mantelatto 4 (http://zoobank.org/urn:lsid:zoobank.org:author:882CB189-DEFC-4CFA-8E04-2923C89B1B25) and
Darryl l. felDer 3 (http://zoobank.org/urn:lsid:zoobank.org:author:7A4A3683-FCED-431A-A685-1C68E7CFD8AB)
1
Museums Victoria, PO Box 666, Melbourne, Vic. 3001, Australia gpoore@museum.vic.gov.au
Dritte Zoologische Abteilung, Naturhistorisches Museum, Burgring 7, A 1010 Vienna, Austria
Peter.Dworschak@nhm-wien.ac.at
3
Department of Biology, University of Louisiana—Lafayette, Lafayette, LA 70504, USA dlf4517@louisiana.edu
4
Laboratory of Bioecology and Crustacean Systematics (LBSC), Department of Biology, Faculty of Philosophy, Science
and Letters at Ribeirão Preto (FFCLRP), University of São Paulo (USP), Av. Bandeirantes 3900, 14040-901, Ribeirão
Preto, São Paulo, Brazil flmantel@usp.br
5
Facultad de Ciencias Químico Biológicas, Universidad Autónoma de Campeche, Campus V. Predio s/n por Avenida Ing.
Humberto Lanz Cárdenas y Fracc. Ecológico Ambiental Siglo XXIII, Colonia Ex Hacienda Kalá, San Francisco de
Campeche, Camp., 24085, México
2
Abstract
Poore, G.C.B., Dworschak, P.C., Robles, R., Mantelatto, F., and Felder, D.L. 2019. A new classification of Callianassidae
and related families (Crustacea: Decapoda: Axiidea) derived from a molecular phylogeny with morphological support.
Memoirs of Museum Victoria 78: 73–146.
The classification of the families and genera of Callianassidae and related families (Crustacea: Decapoda: Axiidea) is
significantly revised based on the results of a separately published molecular phylogeny with morphological support.
Seven families are recognised: Anacalliacidae Manning and Felder, 1991; Callianassidae Dana, 1852; Callianopsidae
Manning and Felder, 1991; Callichiridae Manning and Felder, 1991; Ctenochelidae Manning and Felder, 1991;
Eucalliacidae Manning and Felder, 1991; and Paracalliacidae Sakai, 2005.
The families comprise 53 genera, 17 new:
Anacalliacidae—Anacalliax de Saint Laurent, 1973.
Callianassidae—Aqaballianassa gen. nov.; Arenallianassa gen. nov.; Biffarius Manning and Felder, 1991;
Callianassa Leach, 1814; Caviallianassa gen. nov.; Cheramoides Sakai, 2011; Cheramus Bate, 1888; Coriollianassa gen.
nov.; Filhollianassa gen. nov.; Fragillianassa gen. nov.; Gilvossius Manning and Felder, 1992; Jocullianassa gen. nov.;
Lipkecallianassa Sakai, 2002; Necallianassa Heard and Manning, 1998; Neotrypaea Manning and Felder, 1991; Notiax
Manning and Felder, 1991; Paratrypaea Komai and Tachikawa, 2008; Poti Rodrigues and Manning, 1992; Praedatrypaea
gen. nov.; Pugnatrypaea gen. nov.; Rayllianassa Komai and Tachikawa, 2008; Rudisullianassa gen. nov.; Scallasis Bate,
1888; Spinicallianassa gen. nov.; Tastrypaea gen. nov. and Trypaea Dana, 1852.
Callianopsidae—Bathycalliax Sakai and Türkay, 1999; Callianopsis de Saint Laurent, 1973; and Vulcanocalliax
Dworschak and Cunha, 2007.
Callichiridae—Audacallichirus gen. nov.; Balsscallichirus Sakai, 2011; Calliapagurops de Saint Laurent, 1973;
Callichirus Stimpson, 1866; Corallianassa Manning, 1987; Glypturoides Sakai, 2011; Glypturus Stimpson, 1866;
Grynaminna Poore, 2000; Karumballichirus gen. nov.; Kraussillichirus gen. nov.; Laticallichirus Komai, Yokooka,
Henmi and Itani, 2019; Lepidophthalmus Holmes, 1904; Michaelcallianassa Sakai, 2002; Mocallichirus gen. nov.;
Mucrollichirus gen. nov.; Neocallichirus Sakai, 1988; and Thailandcallichirus Sakai, 2011.
Ctenochelidae—Ctenocheles Kishinouye, 1926; Ctenocheloides Anker, 2010; Dawsonius Manning and Felder,
1991; Kiictenocheloides Sakai, 2013; Gourretia de Saint Laurent, 1973; Laurentgourretia Sakai, 2004; and Paragourretia
Sakai, 2004.
Eucalliacidae—Andamancalliax Sakai, 2011; Calliax de Saint Laurent, 1973; Calliaxina Ngoc-Ho, 2003; Eucalliax
Manning and Felder, 1991; Eucalliaxiopsis Sakai, 2011; Pseudocalliax Sakai, 2011; and Paraglypturus Türkay and Sakai,
1995.
74
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Paracalliacidae—Paracalliax de Saint Laurent, 1979.
Of 19 available family-level names that have accumulated since 1852, ten have been previously synonymised or are
synonymised in this work. Of 74 available genus-level names of extant species, 43 have been previously synonymised. The
following are synonymised in this work. Anacalliaopsis Sakai, 2011, is synonymised with Anacalliax de Saint Laurent,
1973. Nihonotrypaea Manning and Tamaki, 1998, and Pseudobiffarius Heard and Manning, 2000, are synonymised with
Neotrypaea Manning and Felder, 1991. Calliaxiopsis Sakai and Türkay, 2014; Bakercalliax Sakai, 2018; Heardcalliax
Sakai, 2018; and Manningcalliax Sakai, 2018, are synoymised with Eucalliaxiopsis Sakai, 2011. Forestcallichirus Sakai,
2011, and Capecalliax Sakai, 2011, are synonymised with Balsscallichirus Sakai, 2011. Podocallichirus Sakai, 1999;
Lepidophthalmoides Sakai, 2011; and Lepidophthalminus Sakai, 2015, are synonymised with Lepidophthalmus Holmes,
1904. Sergio Manning and Lemaitre, 1994, and Callichiropsis Sakai, 2010, are synonymised with Neocallichirus Sakai,
1988. Ivorygourretia Sakai, 2017; Plantesgourretia Sakai, 2017; and Ruiyuliugourretia Sakai, 2017, are synonymised with
Gourretia de Saint Laurent, 1973. Heterogourretia Sakai, 2017, and Tuerkaygourretia Sakai, 2017, are synoymised with
Paragourretia Sakai, 2004.
Keys are presented to families and to genera within families. All available species names are tabulated within the
new family and genus arrangement. Some remain incertae sedis because they have been only partially described.
Accepted species of Callianassidae and related families number 265, excluding junior synonyms, of which one third
(87) are placed in new genus–species combinations. These are tabulated alphabetically by species and in systematic order.
Keywords
Crustacea, Decapoda, Axiidea, Anacalliacidae, Callianassidae, Callianopsidae, Callichiridae, Ctenochelidae,
Eucalliacidae, Paracalliacidae, taxonomy, new genera
Contents
Introduction ........................................................................ 75
Methods .............................................................................. 75
Infraorder Axiidea de Saint Laurent, 1979 ........................ 76
Rayllianassa Komai and Tachikawa, 2008 ..................... 98
Rudisullianassa gen. nov. ................................................ 98
Scallasis Bate, 1888 ......................................................... 99
Spinicallianassa gen. nov. ............................................... 99
Tastrypaea gen. nov. ...................................................... 100
Trypaea Dana, 1852 ....................................................... 100
Key to families of Axiidea ................................................. 77
Callianopsidae Manning and Felder, 1991 ....................... 101
Anacalliacidae Manning and Felder, 1991 ......................... 81
Anacalliax de Saint Laurent, 1973 .................................. 82
Callianassidae Dana, 1852 ................................................. 82
Key to genera of Callianassidae .........................................
Aqaballianassa gen. nov. .................................................
Arenallianassa gen. nov. .................................................
Biffarius Manning and Felder, 1991 ................................
Callianassa Leach, 1814 ..................................................
Caviallianassa gen. nov. ..................................................
Cheramoides Sakai, 2011 ................................................
Cheramus Bate, 1888 .......................................................
Coriollianassa gen. nov. ..................................................
Filhollianassa gen. nov. ...................................................
Fragillianassa gen. nov. ..................................................
Gilvossius Manning and Felder, 1992 .............................
Jocullianassa gen. nov. ....................................................
Lipkecallianassa Sakai, 2002 ..........................................
Necallianassa Heard and Manning, 1998 .......................
Neotrypaea Manning and Felder, 1991 ...........................
Notiax Manning and Felder, 1991 ...................................
Paratrypaea Komai and Tachikawa, 2008 ......................
Poti Rodrigues and Manning, 1992 .................................
Praedatrypaea gen. nov. ..................................................
Pugnatrypaea gen. nov. ...................................................
87
90
91
91
91
92
92
93
93
93
94
94
95
95
95
96
96
96
97
97
97
Key to genera of Callianopsidae ......................................
Bathycalliax Sakai and Türkay, 1999 ............................
Callianopsis de Saint Laurent, 1973 ..............................
Vulcanocalliax Dworschak and Cunha, 2007 ...............
102
102
102
102
Callichiridae Manning and Felder, 1991 .......................... 102
Key to genera of Callichiridae ......................................... 106
Audacallichirus gen. nov. .............................................. 107
Balsscallichirus Sakai, 2011 .......................................... 108
Calliapagurops de Saint Laurent, 1973 ......................... 108
Callichirus Stimpson, 1866 ........................................... 109
Corallianassa Manning, 1987 ....................................... 109
Glypturoides Sakai, 2011 ............................................... 110
Glypturus Stimpson, 1866 ............................................. 110
Grynaminna Poore, 2000 .............................................. 110
Karumballichirus gen. nov. ............................................ 111
Kraussillichirus gen. nov. ............................................... 111
Laticallichirus Komai, Yokooka, Henmi and Itani, 2019 .. 111
Lepidophthalmus Holmes, 1904 .................................... 112
Michaelcallianassa Sakai, 2002 ................................... 113
Mocallichirus gen. nov. ................................................. 113
Mucrollichirus gen. nov. ................................................ 113
Neocallichirus Sakai, 1988 ............................................. 114
Thailandcallichirus Sakai, 2011 ..................................... 114
75
Callianassidae and related families
Ctenochelidae Manning and Felder, 1991 ........................ 115
Key to genera of Ctenochelidae ........................................ 118
Ctenocheles Kishinouye, 1926 ....................................... 118
Ctenocheloides Anker, 2010 .......................................... 119
Dawsonius Manning and Felder, 1991 .......................... 119
Gourretia de Saint Laurent, 1973 .................................. 119
Kiictenocheloides Sakai, 2013 ....................................... 120
Laurentgourretia Sakai, 2004 ....................................... 120
Paragourretia Sakai, 2004 ............................................ 121
Eucalliacidae Manning and Felder, 1991 ......................... 122
Key to genera of Eucalliacidae ........................................
Andamancalliax Sakai, 2011 .........................................
Calliax de Saint Laurent, 1973 ......................................
Calliaxina Ngoc-Ho, 2003 ............................................
Eucalliax Manning and Felder, 1991 .............................
Eucalliaxiopsis Sakai, 2011 ...........................................
Paraglypturus Türkay and Sakai, 1995 .........................
Pseudocalliax Sakai, 2011 .............................................
125
126
126
126
127
127
128
128
Paracalliacidae Sakai, 2005 ............................................. 128
Paracalliax de Saint Laurent, 1979 ............................... 129
Acknowledgements .......................................................... 129
References ........................................................................ 130
Introduction
The earliest published descriptions of callianassids were at
the end of the 18th century and the beginning of the 19th
century (fig. 1). The genus name Callianassa Leach, 1814,
was introduced shortly thereafter. The number of available
species names now totals 305, with 262 accepted by WoRMS
(2019) at the time of writing, twice that number if fossils are
included. However, despite 72 more generic names having
been erected over the last 200 years, authors remain undecided
about the generic affinities of new species and Callianassa
continues to be used as a catch-all genus. The affinities of
many species have changed over time, some often, such that
about 940 generic recombinations now exist (Sakai, 2011, and
later papers), which is more than three times the number of
accepted species.
The unsatisfactory state of ghost shrimp systematics has
been recently outlined by us in a linked study on which this
paper depends (Robles et al., in press). Here, we present a
classification of the family and genera of Callianassidae and
related families based on Robles et al.’s (in press) phylograms
that were derived by multigene analysis of two mitochondrial
(16S, 12S) and two nuclear (histone 3, 18S) markers from 123
named species, one half of all extant described species (265
species; WoRMS, 2019), including 24 as yet undescribed or
not confidently identified to species. The present phylogeny
was supported by a parsimonious analysis of morphological
data from 195 species that recovered terminal clades
compatible with those of the genetic analyses, though not
always with the same deep relationships between terminal
taxa. Fitting the morphological data to the molecular
phylogram discovered characters that could be viewed as
synapomorphies of terminal clades that we treated as families
and genera.
As in Robles et al. (in press), we use “callianassoid” as a
short-hand term to refer to a monophyletic group of taxa that
includes Ctenochelidae and Callianassidae (sensu Dworschak
et al., 2012; Poore et al., 2014) in a well-supported clade found
in the most recent molecular treatment of “Thalassinidea”
(Robles et al., 2009) and subclades Eucalliacinae,
Ctenochelidae and Callianassidae (Callichirinae and
Callianassinae) in another molecular treatment of
Callianassidae and related families (Felder and Robles, 2009).
The complex taxonomy of Callianassoidea is explained below.
Here, seven callianassoid families of Axiidea and their
genera are diagnosed. For diagnoses of the other families, see
Sakai (2011) for Axiidae and Strahlaxiidae, see Poore (2015a)
for Callianideidae, and see Poore and Collins (2015) for
Micheleidae. A key to all families of Axiidea and keys to all
callianassoid genera within the seven families are offered.
Tables 1 and 2 list all 265 accepted species, synonyms excepted,
alphabetically by species and by family and genus, respectively.
One third of all species, 87, are in new combinations. Species
authorities are given in these tables and are not repeated for the
species mentioned by name in this text.
Methods
Representatives of numerous species were examined in
museum collections: University of Louisiana, Lafayette
(ULLZ); US National Museum of Natural History, Washington
(USNM); Florida Museum of Natural History, Gainsville
(UF); Naturhistorisches Museum, Vienna (NHMW); Muséum
national d’Histoire naturelle, Paris (MNHN); Senckenberg
Museum, Frankfurt (SMF); Zoological Museum, Hamburg
(ZMH); Phuket Marine Biological Center, Phuket (PMBC);
Museums Victoria, Melbourne (NMV); Northern Territory
Museum and Art Gallery, Darwin (NTMAG); Australian
Museum, Sydney (AM); and Queensland Museum, Brisbane
(QM). The morphological data assembled during examination
of the literature and examination of specimens representing
about 200 of these species were the basis of the phylogenetic
analysis (Robles et al., in press). The same data, stored in a
DELTA database (Dallwitz, 2010), were used to generate
diagnoses of genera. Another DELTA database was assembled
for families of Axiidea. The Diagnose facility in the interactive
key program Intkey was used to discover a combination of
three characters that would differentiate families or genera (in
those families with three or more genera). Single characters,
or more if necessary, that uniquely diagnose a family or genus
are in bold italics. Only the aggregate of these characters was
used to diagnose families or genera, fewer than were assembled
to build the phylograms. The diagnoses generated by DELTA
were edited for sense and additional characters were
highlighted if these were felt to more readily diagnose genera.
76
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Figure 1. Earliest published figures of callianassids. Clockwise: Cancer candidus Olivi, 1792, pl. 3 fig. 3; Astacus tyrrhenus Petagna, 1792, pl. 5
fig. 3; Cancer (Astacus) subterraneus Montagu, 1808, pl. 3 figs 1, 2.
To shorten diagnoses, character attributes common to the vast
majority of genera in the larger families are not repeated for
these genera. These are listed after the keys under the heading
Implicit attributes, and as a corollary, only the exceptions to
these attributes appear in generic diagnoses. For example, a
triangular sclerite is absent from the anterior branchiostegite
except in Aqaballianassa. Intkey was used as an aid to
generate dichotomous keys to the families and genera within
families, much as explained by Coleman et al. (2010).
Characters used to differentiate families and genera are
illustrated by line drawings (figs 2–20) accompanying the keys.
Most of these were prepared by tracing published illustrations
in Adobe Illustrator but confirmed by our own observations.
Illustrations are simplified, rescaled and reoriented for better
comparison. The mesial margin of pleopods is on the left. Setae
are omitted from these diagnostic drawings unless they are
critical characters. Original illustrations, most by GCBP, are
identified in figure captions.
Infraorder Axiidea de Saint Laurent, 1979
Axiidea de Saint Laurent, 1979b: 19, 28.—Robles et al., 2009:
310–314.—Dworschak et al., 2012: 187.
Callianassoidea.—Sakai, 2005a: 1125.
Callianassidea.—Sakai and Sawada, 2006: 1357–1358.
Callianassida.—Sakai, 2011: 3.
Remarks. The Axiidea have been diagnosed simply as decapods
having pereopods 1 and 2 chelate. The name Axiidea, rather
than others suggested by Sakai (2005a, 2011) and Sakai and
Sawada (2006), has become almost universally adopted (Poore
et al., 2014).
Poore’s (1994) Callianassoidea included Laomediidae
Borradaile,
1903;
Upogebiidae
Borradaile,
1903;
Callianideidae Kossmann, 1880; Thomassiniidae de Saint
Laurent, 1979a; Ctenochelidae Manning and Felder, 1991;
and Callianassidae Dana, 1852. This concept was first doubted
by a reappraisal of morphology (Sakai, 2005a; Sakai and
Sawada, 2006) and later by molecular data (Robles et al.,
2009; Tsang et al., 2008). Sakai (2005a) included in
Callianassoidea, families Callianassidae; Axiidae Huxley,
1879; Callianideidae; Ctenochelidae and Gourretiidae Sakai,
77
Callianassidae and related families
1999, in fact, all Axiidea. But in a later synthesis, Sakai
(2005b) omitted Axiidae and Callianideidae from
Callianassoidea. Sakai and Sawada (2006) expanded
Callianassoidea to five families. Sakai (2011) included ten
families in Callianassoidea, four with two subfamilies each,
and included nine families in Axioidea. This dichotomy does
not reflect the two clades discovered by investigation of
molecular relationships (Robles et al., 2009), where Axiidae
are sister to a clade with all other families. This history was
summarised by Robles et al. (2009) and Dworschak et al.
(2012). The relationships in the phylograms of both Tsang et
al. (2008) and Robles et al. (2009) led to our expanded study
(Robles et al., in press).
We do not recognise Callianassoidea at the formal
superfamily level within Axiidea because, firstly, it has such a
convoluted history (outlined above), secondly, doing so leaves
other axiidean families hanging, and thirdly, it is impossible to
diagnose unambiguously with a unique synapomorphy.
Axiidae have been shown to belong to one of two axiidean
clades (Robles et al., 2009; Tsang et al., 2008), while the
“callianassoid” subclade plus Strahlaxiidae, Micheleidae and
Callianideidae belong to the other in a paraphyletic relationship
(Robles et al., 2009; Tsang et al., 2008). All callianassoids
have a lobster-like form with a flaccid pleon longer than the
carapace (but so do most callianideids); all lack a long seta on
the triangular posterior lobe of the scaphognathite (but so do
some Strahlaxiidae); all have a complete linea thalassinica, a
hinge separating the dorsal regions of the carapace from the
branchiostegite (but this is partially evident in some
Callianideidae); all except one species have flat contiguous
eyestalks (but so do callianideids). Pleopods 3–5 of
callianassoids have triangular or subtriangular endopods with
straight mesial margins that are closely connected to their
pairs by short or moderately long appendices internae and
exopods that are usually longer than and enclosing the
endopods. This condition is less pronounced in some
ctenochelids and callianopsids, which approach axiids,
strahlaxiids and micheleids in having the endopods of
pleopods 3–5 linear or oval, weakly connected to their pairs
by long appendices internae, and the exopods shorter than or
as long as the endopods, but not enclosing the endopods.
Species of Callianidea have pleopods 3–5 as in most
callianassoids. The uropodal exopod of most callianassoids
has an elevated dorsal plate, a region at the end of the anterior
margin defined by a transverse row of setae but, again, it is not
true of all because a dorsal plate is absent in Callianopsidae,
Ctenochelidae and Paracalliax.
The number of well-defined clades with consistent
molecular support prompted us to recognise seven
“callianassoid” families. All had been previously recognised
at least as subfamilies. Four other families of Axiidea are
Axiidae, Callianideidae, Micheleidae and Strahlaxiidae. The
following family diagnoses use 21 characters adequate to
distinguish all 11 families. The bold italic parts in diagnoses
are character states, generated with the aid of Intkey (Dallwitz,
2010), that distinguish each family from every other family in
at least one respect.
Key to families of Axiidea
Figures 2–4
1.
Rostrum prominent, often with erect lateral spines, carapace
with lateral gastric carinae originating from lateral margins
of rostrum, often with submedian and median gastric
carinae (figs 2a, b, c); linea thalassinica absent (figs 2e–g);
eyestalks cylindrical (figs 2a, b, c)
2
–
Rostrum triangular-flat or reduced to short spine shorter
than eyestalk, carapace without median and lateral gastric
carinae (fig. 2f), or rostrum flat-unornamented, longer than
eyestalk, carapace with lateral gastric carinae (fig. 2e);
linea thalassinica present over all or part of carapace length
(figs 2h–o), or absent, or short; eyestalks contiguous, flat or
cylindrical (figs 2d, f, g)
3
2.
Rostrum apex bifid, with lateral teeth (fig. 2c); propodi of
pereopods 3 and 4 without lateral spiniform setae (fig.
3q); pleopods 3–5 with oblique peduncles meeting
mesially, endopods oval, exopods attached laterally,
triangular, shorter than endopods, wider proximally than
distally (fig. 4p); maxilla scaphognathite with (fig. 3a) or
without (fig. 3b) long setae extending from posterior lobe
into branchial chamber
Strahlaxiidae
–
Rostrum with acute or rounded apex (figs 2a, b); propodi
of pereopods 3 and 4 with lateral spiniform setae (fig. 3p);
pleopods 3–5 with linear peduncles not meeting mesially,
endopods linear to elongate-oval, exopods linear-oval,
attached subdistally, shorter than or as long as endopod,
not overlapping endopods (fig. 4o); maxilla scaphognathite
with long setae extending from posterior lobe into
branchial chamber (fig. 3a)
Axiidae
3.
Maxilla scaphognathite with long setae extending from
posterior lobe into branchial chamber (fig. 3a); linea
thalassinica complete, partial or absent
4
–
Maxilla scaphognathite without long setae extending from
posterior lobe into branchial chamber (fig. 3b); linea
thalassinica complete over full carapace length (fig. 2h)
5
4.
Posterior margin of carapace evenly curved, not
interacting with anterolateral lobes on pleomere 1 (fig.
2f); eyestalk flat, contiguous (fig. 2f); chelipeds flattened,
asymmetrical (fig. 3j)
Callianideidae
–
Posterior margin of carapace with lateral lobes interacting
with anterolateral lobes on pleomere 1 (fig. 2g); eyestalks
cylindrical even if continuous (fig. 2g); chelipeds
cylindrical, symmetrical (fig. 3i)
Micheleidae
5.
Pleomere 1 with dorsal pair of lobes interacting with
posterior margin of carapace (fig. 2i); female pleopod 1 with
single broad expanded ramus (fig. 4b); pleopod 2 (at least of
female) similar to pleopods 3–5 (fig. 4c); epipods present
above maxilliped 3 to pereopod 4
Paracalliacidae
(1 species, Paracalliax bollorei de Saint Laurent, 1979)
78
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Figure 2. Diagnostic characters for families of Axiidea. Anterior carapace, dorsal: a, Axiidae, Pillsburyaxius; b, Axiidae, Eiconaxius; c,
Strahlaxiidae, Neaxius; d, Eucalliacidae, Calliaxina; e, Micheleidae, Tethisea; f, Callianeidae, Callianidea. Carapace, lateral: g, Micheleidae,
Michelea; h, Eucalliacidae, Calliaxina. Posterior carapace, pleomeres 1, 2: i, Paracalliacidae; j, Callianassidae. Anterior carapace, branchiostegite,
epistome, basal antenna and eyestalk: k, Eucalliacidae, Calliaxina; l, Callichiridae, Callichirus; m, Callichiridae, Lepidophthalmus; n,
Callianassidae, Coriollianassa; o, Callianassidae, Biffarius.
Original illustrations: i, Paracalliax bollorei, MNHN Th1517; k, Calliaxina sakaii, ULLZ; l, Callichirus islagrande, ULLZ; m,
Lepidophthalmus richardi, ULLZ; n, Coriollianassa coriolisae, MNHN-IU-2014-18276; o, Biffarius biformis.
–
Pleomere 1 with evenly curved dorsal margin, not
interacting with posterior margin of carapace (fig. 2j);
female pleopod 1 with reduced ramus (fig. 4a); pleopod 2
in both sexes (figs 5d–g) smaller than pleopods 3–5 (fig.
5q); epipods absent above maxilliped 3 to pereopod 4
(exception, 2 species of Callianopsidae)
6
6.
Maxilliped 3 dactylus ovate, distally truncate, with dense
distal field of setae (fig. 3e)
7
–
Maxilliped 3 dactylus linear, with scattered groups of
setae (fig. 3f), or densely setose on upper or lower margin
(figs 3g, h)
8
Callianassidae and related families
79
Figure 3. Diagnostic characters for families of Axiidea. Maxilla: a, Axiidae, Eutrichocheles; b, Callianassidae, Arenallianassa. Maxilliped 1: c,
Callichiridae; d, Callianassidae. Maxilliped 3: e, Callianopsidae; f, Anacalliacidae; g, Callianassidae, Callianassa; h, Callianassidae,
Caviallianassa. Major cheliped: i, Micheleidae, Tethisea; j, Callianideidae, Callianidea; k, Ctenochelidae, Ctenocheles; l, Ctenochelidae,
Gourretia; m, Anacalliacidae. Minor cheliped: n, Ctenochelidae, Ctenocheles; o, Ctenochelidae, Gourretia. Pereopod 3, propodus, dactylus: p,
Axiidae, Acanthaxius; q, Strahlaxiidae, Neaxius. Pereopod 5, fingers: r, Ctenochelidae, Ctenocheloides; s, Anacalliacidae. The arrows indicate
features of interest.
Original illustrations: d, Callianassa subterranea, NMV J16779; h, Caviallianassa FP-11, UF 29204.
80
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Figure 4. Diagnostic characters for families of Axiidea. Female pleopod 1: a, Eucalliacidae, Paraglypturus; b, Paracalliacidae. Female pleopod
2: c, Paracalliacidae; d, Eucalliacidae, Calliax; e, Eucalliacidae, Calliaxina; f, Callichiridae; g, Callianassidae. Male pleopod 1: h, i, Callichiridae;
j, Callianassidae. Male pleopod 2: k, Ctenochelidae, Ctenocheles; l, Eucalliacidae, Eucalliax; m, Eucalliacidae, Calliaxina; n, Eucalliacidae,
Paraglypturus. Pleopod 3: o, Axiidae, Marianaxius; p, Strahlaxiidae, Neaxiopsis; q, Callichiridae. Pleomere 6, uropod, telson: r, Ctenochelidae,
Gourretia; s, Ctenochelidae, Paragourretia; t, Eucalliacidae, Calliaxina; u, Eucalliacidae, Eucalliax; v, Callianassidae; w, Anacalliacidae; x,
Callianopsidae. Arrows indicate features of interest.
Original illustrations: s, Eucalliax quadracuta, NHMW 25916; w, Anacalliax agassizi MNHN Th1206.
81
Callianassidae and related families
7.
–
Uropodal exopod oval, without dorsal plate (fig. 4x);
telson parallel-sided over proximal half, then tapering to
evenly rounded apex, or widest basally over anterior
third, sharp lateral step, then tapering posteriorly to
broadly rounded apex; maxilliped 3 propodus longer than
wide, not prominently lobed on lower margin (fig. 3e);
male pleopod 1 with article 2 triangular, with lobed
mesial margin, or flagellate
Callianopsidae
Uropodal exopod with dorsal plate, indicated by secondary
row of setae diverging on upper surface from anterior
margin (figs 4t, u); telson convex-sided, widest near
midpoint, or semicircular, or curving to broad convex
apex (figs 4t, u); maxilliped 3 propodus about as wide as
long; male pleopod 1 with article 2 linear, or blade like,
with bifid or simple apex, or digitiform (sometimes fused)
Eucalliacidae
8.
Uropodal exopod without dorsal plate (fig. 4r), or with
distal margin interrupted by weak notch (fig. 4s)
9
–
Uropodal exopod with dorsal plate, secondary row of
setae on upper surface branching subdistally from anterior
margin (fig. 4v)
10
9.
Pereopod 5 semichelate (fixed finger closing on proximal
part of dactylus), dactylus a twisted plate longer than fixed
finger (fig. 3r); major cheliped merus lower margin with
weakly toothed squarish blade plus proximal erect spine
(fig. 3l), or simple proximal spine, or oblique blade or
spine near midpoint; propodus evenly tapering or
cylindrical, with fingers tapering and irregularly toothed
or thin and pectinate (fig. 3k); minor cheliped merus with
spine on lower margin (figs 3n, o); telson mostly parallelsided, with rounded posterolateral corners (fig. 4r), or
parallel-sided over proximal half, then tapering to evenly
rounded apex (fig. 4s)
Ctenochelidae
–
Pereopod 5 chelate (fixed finger closing complete length
of dactylus; fig. 3s); major cheliped merus lower margin
with 1 or 2 small proximal teeth, propodus parallel-sided,
fixed finger as long as dactylus (fig. 3m); minor cheliped
merus lower margin smooth; uropodal endopod with
small distal notch (fig. 4w); maxilliped 3 ischium linear,
with curved lower margin, palp narrow, dactylus linear,
generally setose (fig. 3f); telson tapering from greatest
width near base (fig. 4w)
Anacalliacidae
10. Anterior branchiostegal lobe well-produced anteriorly
beyond junction with oblique branchiostegal ridge with
which it articulates by means of a virtual condyle (figs 2l,
m); male pleopod 1 article 2 usually of 2 articles, article 2
simple or apically notched (figs 4h, i); male and female
pleopod 2 rami narrower and with less setation than
pleopods 3–5, endopod flattened and 2–5 times as long as
wide (fig. 4f); maxilliped 1 epipod with acute anterior
lobe lying obliquely and along oblique suture of exopod
(fig. 3c)
Callichiridae
–
Anterior branchiostegal lobe merging smoothly with
anterodorsal branchiostegal angle (figs 2n, o), or with
small independent triangular sclerite; male pleopod 1
absent, or if present, uniarticulate or with second simple
article (fig. 4j); male pleopod 2 absent or reduced; female
pleopod 2 rami styliform, endopod much longer than wide
(fig. 4g); maxilliped 1 epipod truncate, without anterior
lobe (fig. 3d)
Callianassidae
Anacalliacidae Manning and Felder, 1991
Figure 5
Anacalliinae Manning and Felder, 1991: 786.
Anacalliacinae.—Sakai, 1999a: 126.—Sakai, 2005b: 208–210.
Anacalliacidae.—Sakai, 2011: 341.
Type genus. Anacalliax de Saint Laurent, 1973.
Diagnosis. Rostrum flat, short, triangular, shorter than
eyestalks; median carina on rostrum only; gastric carinae
absent; cervical groove well defined; suture between ocular
lobe and end of linea thalassinica horizontal in lateral view;
anterior branchiostegal margin sinusoidal or semicircular;
anterior branchiostegal lobe simple, scarcely calcified, merging
smoothly with anterodorsal branchiostegal angle and
anterolateral margin of carapace; posterior margin of carapace
without lateral lobes, pleomere 1 without anterolateral lobes,
weakly chitinised. Eyestalks flattened, contiguous, with
subdistal dorsal cornea. Antennal scaphocerite elongate.
Maxilla scaphognathite without long seta on posterior lobe
extending into branchial chamber. Maxilliped 1 epipod with
acute anterior lobe lying alongside exopod. Maxilliped 3
propodus longer than wide, not prominently lobed on lower
margin; dactylus slender, digitiform, with setae irregularly
spaced along all margins. Cheliped merus lower margin
spinose; major cheliped palm oval in cross-section, barely
crested above or below. Pereopod 3 propodus broad, with
proximal lobe on lower margin, without distal spiniform setae
on lateral face (often with 1 distal spiniform seta on lower
margin). Pereopod 5 minutely chelate or subchelate. Female
pleopod 2 rami narrower and with more reduced setation than
pleopods 3–5; endopod 2–5 times as long as wide. Pleopods
3–5 with oblique peduncles, endopods oval, exopods attached
laterally, not proximally lobed, shorter than and barely
overlapping endopods; appendices internae reduced and almost
embedded in mesial margin of endopod. Uropodal exopod
with margin divided by notch.
Remarks. The only genus, Anacalliax de Saint Laurent, 1973,
is recognised by the unique combination of a short flat rostrum
and the uropodal exopod having a marginal notch.
The subfamily Anacalliacinae (misspelled Anacalliinae)
was erected by Manning and Felder (1991) as one of three
subfamilies of Ctenochelidae, the others being Ctenochelinae
and Callianopsinae. Manning and Felder (1991) included
Paracalliax, Gourretia and Dawsonius in the nominate
subfamily and only their type genus in the other two. This
arrangement reflects in a single family, with all four basally
derived families recognised in Robles et al.’s (in press) analysis
as a paraphyletic grouping. Sakai (2005b) treated the taxon as
a subfamily of Callianassidae while retaining Gourretiidae for
82
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Figure 5. Diagnostic characters of Anacalliacidae, Anacalliax: a, dorsal anterior carapace, eyestalks, antennules, antennae; b, lateral carapace;
c, maxilliped 3; d, male pleopod 1; e, female pleopod 2; f, telson, uropod.
other subfamilies. Sakai (2011) elevated the taxon to family
level. The family comprises a single genus. Two genera erected
by Sakai (2011) are herein synonymised, Anacalliaopsis with
Anacalliax, and Capecalliax with Balsscallichirus
(Callichiridae; see below).
The similarity of the “dorsal oval” to that of Callianassa
s.s. was highlighted in Sakai’s (2011) discussion of this group,
but this character grades to such an extent throughout all
families that we were unable to use it at any level. The broad
(almost axiid-like) article 2 of the male pleopod 1, presence of
an appendix masculina on the male pleopod 2, absence of any
tooth on the merus of the cheliped, absence of a dorsal plate on
the uropodal exopod and absence of a distal lobe on the epipod
of maxilliped 1 suggest a basal position for this family, as
realised by Manning and Felder (1991).
The family is known only from the type material of its two
species. No specimen was available for inclusion in the
molecular analysis.
Anacalliax de Saint Laurent, 1973
Anacalliax de Saint Laurent, 1973: 515.—Manning and Felder,
1991: 786–787.—Sakai, 1999a: 126.—Sakai, 2005b: 210.—Sakai,
2011: 343.
Anacalliaopsis Sakai, 2011: 342 (type species: Callianassa
agassizi Biffar, 1971, by original designation and monotypy) syn. nov.
Type species. Callianassa argentinensis Biffar, 1971b, by
original designation and monotypy.
Diagnosis. With characters of the family.
Remarks. The genus is the sole member of the family. Sakai
(2011) erected a new genus Anacalliaopsis for a single species
Callianassa agassizi. He did not explain how the species
differed from Anacalliax argentinensis except in his key where
the only difference was in the posterior margin of the telson
(other characters used are identical or not compared).
Anacalliax argentinensis has a concave margin whereas A.
agassizi has a convex margin, differences that, if substantiated,
would be best treated at species level. Otherwise, the two share
similar maxillipeds, chelipeds and pleopods.
Callianassidae Dana, 1852
Figures 6–11
Callianassidae Dana, 1852a: 12, 14.—Dana, 1852b: 508.—
Bouvier, 1940: 100.—Balss, 1957: 1581.—de Saint Laurent, 1973:
513.—de Saint Laurent, 1979a: 1395.—Manning and Felder, 1991:
766.—Poore, 1994: 101—Sakai, 1999a: 7.—Sakai, 2005b: 9–11.—
Sakai and Sawada, 2006: 1357.—Sakai, 2011: 353–357 partim.
Callianassinae.—Bouvier, 1940: 100.—Balss, 1957: 1582.—de
Saint Laurent, 1973: 514.—de Saint Laurent, 1979a: 1395–1396.—
Manning and Felder, 1991: 767.—Sakai, 1999a: 10.—Sakai, 2005b:
11–25.—Sakai, 2011: 357–358.
Callianassidae and related families
83
Figure 6. Diagnostic characters for genera of Callianassidae. Anterior carapace, eyestalks, antennules, antennae: a, Aqaballianassa; b,
Arenallianassa; c, Caviallianassa; d, Coriollianassa; e, f, Filhollianassa; g, Jocullianassa; h, Notiax i, Praedatrypaea; j, Pugnatrypaea; k, l,
Rayllianassa; m, Rudisullianassa; n, Trypaea. Mandible, mesial and lower views: o, Callianassa; p, Rayllianassa.
Original illustrations: a, Aqaballianassa lewtonae, MNHN-IU-2016-8152; c, Caviallianassa FP-11, UF 29204; j, Pugnatrypaea GMX,
USNM 1559553 (ULLZ 17962); o, Callianassa subterranea, NMV J16779; p, Rayllianassa amboinensis, MNHN-IU-2014-2778.
Cheraminae Manning and Felder, 1991: 780.—Tudge et al., 2000:
136.
Lipkecallianassinae Sakai, 2005: 212.
Lipkecallianassidae Sakai, 2011: 521.
Diagnosis. Rostrum flat, short, triangular, shorter than eyestalks,
or spike-like, longer than wide; gastric carinae absent; cervical
groove well defined; suture between ocular lobe and end of
linea thalassinica oblique in lateral view; anterior branchiostegal
margin sinusoidal or semicircular; anterior branchiostegal
lobe sclerotised, merging smoothly with anterodorsal
branchiostegal angle; posterior margin of carapace without
lateral lobes, pleomere 1 without anterolateral lobes, weakly
chitinised. Eyestalks flattened, contiguous, with subdistal dorsal
cornea. Antennal scaphocerite rudimentary. Maxilla
scaphognathite without long seta on posterior lobe extending
into branchial chamber. Maxilliped 1 epipod truncate, without
anterior lobe. Maxilliped 3 dactylus slender, digitiform, with
setae irregularly spaced along all margins. Cheliped merus
84
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Figure 7. Diagnostic characters for genera of Callianassidae. Maxilliped 3: a, Callianassa; b, Caviallianassa; c, Cheramoides; d, Lipkecallianassa;
e, Necallianassa; f, Neotrypaea; g, Praedatrypaea; h, Pugnatrypaea; i, Scallasis; j, Spinicallianassa; k, Trypaea; l, Arenallianassa; m, Biffarius.
Original illustrations: a, Callianassa; b, Caviallianassa; c, Cheramoides; d, Lipkecallianassa; e, Necallianassa; m, Biffarius delicatulus,
NHMW 25542.
lower margin smooth, or with prominent proximal tooth; major
cheliped with distinctively flattened palm, sometimes with
strong crest above and below. Pereopod 3 propodus wide, with
proximal lobe on lower margin, without distal spiniform setae
on lateral face (often with 1 distal spiniform seta on lower
margin). Pereopod 5 minutely chelate or subchelate. Female
pleopod 2 rami styliform; endopod much longer than wide, or
absent. Pleopods 3–5 with oblique peduncles meeting mesially,
endopods triangular, with straight mesial margin, exopods
attached laterally, proximally lobed, longer than and enclosing
endopods; appendices internae elongate, much longer than
wide, or reduced and almost embedded in mesial margin of
endopod. Uropodal exopod with elevated dorsal plate.
Remarks. Callianassidae differs from its sister taxon
Callichiridae in having the rami of the female pleopod 2
styliform and lacking an appendix interna, often absent in the
male, rather than broad and often with an appendix interna,
and the truncate epipod on maxilliped 1, whereas an anterior
lobe runs alongside the exopod in Callichiridae.
The clades resolved in the molecular analysis of over 50
species (Robles et al., in press) bore little resemblance to
existing classifications. This necessitated the acceptance of 14
existing genera, the creation of 12 new genera and the
synonymy of others. All 26 can be differentiated using
morphological features, some of which had been overlooked in
earlier accounts. The following key does not reflect phylogeny
but attempts to eliminate the most distinctive genera first.
Callianassidae and related families
85
Figure 8. Diagnostic characters for genera of Callianassidae. Male major cheliped: a, Cheramus; b, Fragillianassa; c, Coriollianassa; d, e,
Biffarius; f, Arenallianassa; g, Caviallianassa; h, Callianassa; i, Cheramoides; j, Filhollianassa; k, Gilvossius. Minor cheliped: l, Cheramoides;
m, Neotrypaea.
Original illustrations: c, Coriollianassa MOZ-33, MNHN-IU-2008-10314; e, Biffarius biformis, NMV J20793; g, Caviallianassa FP-11, UF
29204; j, Filhollianassa filholi, NMV J62111.
86
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Figure 9. Diagnostic characters for genera of Callianassidae. Male major cheliped: a, Jocullianassa; b, Notiax; c, Necallianassa; d, Neotrypaea;
e, f, Paratrypaea; g, Rayllianassa; h, Rudisullianassa; i, Scallasis; j, Spinicallianassa; k, Tastrypaea. Minor cheliped: l, Jocullianassa; m,
Rudisullianassa.
87
Callianassidae and related families
Figure 10. Diagnostic characters for genera of Callianassidae. Pereopod 3: a, Lipkecallianassa; b, Scallasis; c, Spinicallianassa; d, Jocullianassa.
Male pleopod 1: e, Caviallianassa. Male pleopod 2: f, Caviallianassa; g, Poti.
Original illustration: e, f, Caviallianassa FP-11, UF 29204.
Key to genera of Callianassidae
1.
Antennular peduncle about 4 times the width of both
eyestalks, twice as long as antennal peduncle, with 2
dense rows of adjacent long setae on lower surface (fig.
6n); maxilliped 3 merus produced as massive triangular
lobe alongside palp (fig. 7k)
Trypaea
–
Antennular peduncle at most 3 times the width of both
eyestalks, with at most bands of scattered long setae on
lower surface (fig. 6a); maxilliped 3 merus distally convex
at most
2
–
Maxilliped 3 merus distally oblique with obtuse angle
between distal and lower margins (fig. 7h); pereopod 3
propodus oval, lower margin slightly convex, leading to
narrow, distinctly rounded proximal lobe; uropodal
exopod distal margin clearly differentiated from anterior
margin, anterodistal corner right-angled; telson tapering
over distal third to pair of posterior lobes separated by
deep notch (fig. 11o)
Pugnatrypaea
5.
Antennular peduncle exceeded by all or most of antennal
peduncular article 5 (figs 6d, i)
6
2.
Small triangular sclerite present on branchiostegite at
anterior end of linea thalassinica (fig. 6a) Aqaballianassa
–
Antennular peduncle as long as or longer than antennal
peduncle (figs 6b, c, e, l)
10
–
Without triangular sclerite on branchiostegite at anterior
end of linea thalassinica (figs 6f, k, h)
3
6.
3.
Telson tapering, posterior margin with medial spine in
deep notch (figs 11g, o)
4
Maxilliped 3 merus with spine on distal free margin (fig.
7g); proximolateral lobe of telson prominent, defined
posteriorly by clear unchitinised region (fig. 11n)
Praedatrypaea
–
Telson posterior margin convex, truncate or at most with
shallow medial concavity, sometimes with small medial
spine
5
–
Maxilliped 3 merus without spine on distal free margin;
proximolateral lobe of telson prominent or not, indefinitely
defined posteriorly
7
4.
Maxilliped 3 merus almost rectangular, distally truncate
with squarish angle between distal and lower margins (fig.
7d); pereopod 3 propodus linear, without lobe on lower
margin (fig. 10a); uropodal exopod distal margin with
anterodistal angle obtuse; telson with pair of broad
posterior lobes, widely excavate at midpoint (fig. 11g)
Lipkecallianassa
7.
Major cheliped carpus articulating by means of a short
neck with merus, merus with bifid proximal spine (fig.
8c); scaphocerite bifid (fig. 6d)
Coriollianassa
–
Major cheliped carpus without neck, lower margin evenly
curved, merus lower margin simple, with simple spine or
blade; scaphocerite simple
8
88
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Figure 11. Diagnostic characters for genera of Callianassidae. Pleomere 6, telson, uropod: a, Arenallianassa; b, c, Caviallianassa (uropod in
detail); d, e, Cheramoides (with pleomere 6 lateral); f, Gilvossius; g, Lipkecallianassa; h, Necallianassa; i, Neotrypaea californiensis; j, N.
petalura; k, Notiax; l, Poti; m, Paratrypaea; n, Praedatrypaea; o, Pugnatrypaea; p, Tastrypaea.
Original illustrations: a, Arenallianassa arenosa, NMV J31887; b, c, Caviallianassa FP-11, UF 29204; d, e, Cheramoides marginata,
MNHN-IU-2016-2462; i, Neotrypaea californiensis, NMV J20600; i, N. petalura, NMV J59981; k, Notiax brachyophthalma, NMV J58880; m,
Paratrypaea maldivensis, UF 28781.
89
Callianassidae and related families
8.
Major cheliped upper margins of ischium and merus,
lower margins of merus and carpus beaded, with dense
row of fine setae, propodus with deep notch at base of
fingers (figs 8d, e)
Biffarius
–
Major cheliped upper margins of ischium and merus,
lower margins of merus and carpus smooth, with few
well-spaced setae, propodus without deep notch at base of
fingers
9
9.
–
Rostrum acute, as long as eyestalk (fig. 6i); cornea diffuse;
maxilliped 3 longer than wide at ischium-merus suture
(fig. 7c); major and minor chelipeds similar, with sawtooth blade on lower margin of merus, upper margin
concave (fig. 8a); pleomere with sublateral ventral sharp
ridge, flared posteriorly (figs 11d, e)
Cheramus
Rostrum obtusely triangular, not reaching cornea (fig.
6m); cornea well defined; maxilliped 3 wider at ischiummerus suture than long (fig. 7m); major and minor
chelipeds dissimilar, merus unarmed or with small teeth
(figs 8h, m); pleomere without sublateral ventral sharp
ridge
Rudisullianassa
10. Rostrum acute, reaching beyond cornea (figs 6g, i)
–
11
Rostrum obsolete or triangular, not reaching cornea (figs
6b–f, h, l, m)
15
11. Pereopod 3 propodus bean-shaped, lower margin concave,
with broadly rounded proximal lobe (fig. 10d); major
cheliped merus with simple perpendicular proximal spine
(fig. 9a)
Jocullianassa
–
–
15. Major cheliped with dorsodistolateral propodus and
lateral dactylus with dense brush of setae (figs 9e, f);
uropodal endopod with facial distal transverse row of
short spiniform setae (fig. 11m)
Paratrypaea
–
–
13. Uropodal exopod at least twice as long as wide (fig. 11d);
pleopod 2 absent in male
Cheramoides
–
Uropodal exopod about as long as wide (fig. 11l); pleopod
2 present in male (fig. 10g)
Poti
14. Maxilliped 3 merus wider at ischium-merus suture than
long, distally convex, extending only slightly beyond
articulation of carpus (fig. 7j); male major cheliped merus
with oblique spine about one third or half-way along lower
margin, propodus swollen, longer than carpus (fig. 9j);
pereopod 3 propodus subrectangular (fig. 10c)
Spinicallianassa
Anterior carapace moderately or not depressed in lateral
view (fig. 6h)
17
17. Antennular peduncle exceeding antennal peduncle by
about half length of article 3 (fig. 6l); major and minor
chelipeds similar, merus without prominent hook or spine
on lower margin, carpus and propodus ovoid in crosssection, upper and lower margins not carinate, carpus of
both shorter than upper margin of propodus (fig. 9g);
mandibular molar dominating, calcified, swollen projection
without sharp edge, incisor without teeth (fig. 6p)
Rayllianassa
–
Pereopod 3 propodus oval or with straight lower margin
(Figs 10b, c); major cheliped merus smooth (fig. 8i), or
with spine (fig. 9h), or with blade (figs 9i, j)
12
Maxilliped 3 distally convex, extending beyond
articulation of carpus (fig. 7j), or distally oblique with
obtuse angle between distal and lower margins (fig. 7i);
major cheliped merus with prominent hooked blade (figs
9i, k)
14
Major cheliped with dorsodistolateral propodus and
lateral dactylus sparsely setose; uropodal endopod
sometimes with facial spiniform setae but not as obvious
transverse row
16
16. Anterior carapace strongly depressed by about 45° in
lateral view, rostrum blunt (figs 6e, f)
Filhollianassa
12. Maxilliped 3 almost rectangular, distally truncate with
squarish angle between distal and lower margins (fig. 7c);
major cheliped merus without prominent hooked blade
(unknown in Poti; fig. 8i)
13
–
Maxilliped 3 merus longer than wide at ischium-merus
suture, distally oblique with obtuse angle between distal
and lower margins (fig. 7i); male major cheliped merus
with 1 or 2 proximal similar teeth and distal denticles on
lower margin, propodus narrow, shorter than carpus (fig.
9i); pereopod 3 propodus oval (fig. 10b)
Scallasis
Antennular peduncle about as long as or a little longer
than antennal peduncle (figs 6b, c); chelipeds dissimilar,
major cheliped merus with prominent hook or spine on
lower margin, carpus and propodus flattened, upper and
especially lower margins carinate, submarginal mesial
face slightly concave (figs 8b, g); minor cheliped carpus
longer than upper margin of propodus (fig. 8m);
mandibular molar with simple sharp margin, incisor
toothed (fig. 6o)
18
18. Maxilliped 3 dactylus with dense brush of long setae over
most of upper-distal margin, few setae along lower margin
(fig. 7b)
19
–
Maxilliped 3 dactylus with scattered setae over upper
margin, dense brush of short setae distally on lower
margin (fig. 7a)
20
19. Male and female major cheliped merus with prominent
truncate hook armed with serrations along lower margin,
excavate laterally at base, with deep notch at base of
fingers (fig. 8b); pleopod 1 present in male, pleopod 2
absent in male; uropodal endopod without facial spiniform
setae
Fragillianassa
–
Male major cheliped merus with 3 oblique similar short
proximal spines on lower margin, beaded beyond (fig. 8g),
female with simple hook; pleopods 1 and 2 present in
male (figs 10e, f); uropodal endopod with at least 1 facial
spiniform seta (figs 11b, c)
Caviallianassa
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G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
20. Maxilliped 3 merus longer than width at ischium-merus
suture, distally oblique with obtuse angle between distal
and lower margins (see above; fig. 7a); male major cheliped
merus with simple proximal hook on lower margin (fig.
8h)
Callianassa
–
Maxilliped 3 merus wider at ischium-merus suture than
long, distally convex, extending beyond articulation of
carpus (figs 7e, f); male major cheliped merus with
prominent complex truncate hook (figs 8f, k, 9c)
21
21. Uropodal endopod anterior margin with distal spine (fig.
11h); telson usually with 1 or 2 pairs of lateral spines
Necallianassa
–
Uropodal endopod anterior margin and telson lateral
margin without spines (figs 11f, i, j)
22
22. Telson wider than long, posterior margin semicircular
(fig. 11f) or subtruncate; uropodal endopod without facial
spiniform setae
Gilvossius
–
Telson as wide as long as or longer than wide, posterior
margin truncate, slightly concave or slightly convex
between posterolateral angles; uropodal endopod with
short distal transverse row of facial spiniform setae (figs
11i–k, p)
23
23. Male major cheliped merus with dentate blade at midpoint
(fig. 9k); uropodal exopod posterodistal margin with row
of 6–8 long blade-like setae proximal to long setae on
distal margin (fig. 11c)
Tastrypaea
–
Male major cheliped merus with prominent basal truncate
tooth (figs 8f, 9b, d); uropodal exopod distal and posterior
margins densely setose, without blade-like setae
24
24. Rostrum acute, anteriorly directed, with ventral broad
swelling (fig. 6h); maxilliped 3 crista dentata absent or
comprising few proximal denticles; male major cheliped
with distal margin of propodus bearing tooth at base of
dactylus (fig. 9b); uropodal endopod distal margin well
differentiated from anterior margin, distal margin
truncate; telson longer than wide, tapering from base (fig.
11k)
Notiax
–
Rostrum flat against eyestalks (fig. 6b); maxilliped 3 crista
dentata consisting of a row of denticles (fig. 7f); male major
cheliped with distal margin of propodus having deep notch
at base of fingers (fig. 9d); uropodal exopod distal margin
well or poorly differentiated from anterior margin; telson
wider than or about as wide as long
25
25. Maxilliped 3 merus distally convex, extending only
slightly beyond articulation of carpus (fig. 7l); telson
wider than long, posterior margin as wide as base, weakly
convex between rounded posterolateral angles (fig. 11a)
Arenallianassa
–
Maxilliped 3 merus expanded distally as rounded lobe
beyond articulation with ischium (fig. 7f); telson at least
as wide as long, usually wider than long, posterior margin
narrower than base, often with median spine (figs 11i, j)
Neotrypaea
Implicit attributes. Unless indicated otherwise, the following
attributes are implicit throughout the generic diagnoses.
Gonochoristic. Triangular sclerite absent from branchiostegite
at anterior end of linea thalassinica; anterior carapace almost
flat in lateral view. Rostrum flat against eyestalk. Pleomere 6
without sublateral ventral sharp ridge. Cornea well defined,
pigmented, eyestalk distal lobes rounded, largely contiguous.
Antennular peduncle length about twice width of both
eyestalks, about as long as or a little longer than antennal
peduncle; articles 2 and 3 with lateral band (3–5 setae wide) of
20–30 irregularly placed long setae along lower margin, and
mesial row of shorter setae. Antennal scaphocerite simple,
about as long as wide, apically rounded. Mandibular molar flat,
with sharp margin; incisor dentate. Maxilliped 3 merus distally
convex, extending only slightly beyond articulation of carpus,
without distal spine on mesial margin; crista dentata consisting
of a row of numerous denticles; dactylus tapering, with
scattered setae over upper margin, dense brush of short setae
distally on lower margin. Male major cheliped merus with
spine or tooth or variously denticulate along lower margin;
upper margins of ischium and merus, and lower margins of
merus and carpus, smooth with few well-spaced setae; carpus
proximal and lower margins evenly convex; carpus and
propodus flattened, upper and especially lower margins
carinate, submarginal mesial face slightly concave; propodus
distal margin unornamented, oblique; upper distal margin of
propodus and dactylus with scattered lateral clusters of setae.
Minor cheliped slender, considerably narrower than major
cheliped; carpus upper margin longer than propodus. Male
pleopod 2 present. Uropodal endopod anterior margin
unarmed, without facial setae. Uropodal exopod about as long
as wide, distal margin poorly differentiated from anterior
margin, anterodistal corner rounded, posterodistal margin
densely setose, with short spiniform setae along upper margin.
Aqaballianassa gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:27E2FC97-B39348F5-B69F-83AFD8FF49D7
Type species. Callianassa aqabaensis Dworschak, 2003, by
present designation.
Diagnosis. Hermaphrodite. Triangular sclerite separate from
branchiostegite at anterior end of linea thalassinica.
Rostrum obsolete or obtusely triangular, flat, not reaching
cornea. Pleomere 1 tergite undivided, or with weak transverse
ridge. Pleomere 6 with sublateral ventral sharp ridge, flared
posteriorly. Antennular peduncle articles 2 and 3 with single
lateral row of 6–10 well-spaced long setae along lower margin.
Mandibular molar calcified, swollen projection without sharp
edge; incisor with few teeth. Maxilliped 3 merus wider at
ischium-merus suture than long. Male major cheliped merus
with oblique spine about one third to half-way along lower
margin; propodus distal margin with small lateral tooth;
dactylus with dense setae along upper margin. Minor cheliped
two-thirds width of major cheliped, both flattened. Pereopod 3
propodus rectangular, lower margin deeply convex, leading to
broadly rounded free proximal lobe. Male pleopod 2 present or
absent. Uropodal endopod ovoid, usually longer than wide,
91
Callianassidae and related families
anterior margin straight or slightly convex, posterodistal
margin evenly convex, with spiniform setae near anterior and
distal margins, or with facial spiniform setae on rib. Uropodal
exopod 1.0–1.8 times as long as wide, distal margin clearly
differentiated from anterior margin, anterodistal corner rightangled, posterodistal margin with row of 6–8 long blade-like
setae proximal to long setae on distal margin. Telson
anterolateral lobe obsolete, undefined; truncate or slightly
convex between posterolateral angles, or slightly concave,
sometimes with medial spine.
Etymology. An alliteration of the name of the type species and
Callianassa, type genus of the family.
Remarks. The genus is recognisable by the unique triangular
sclerite separated by a clear suture or discontinuity from the
branchiostegite at the anterior end of the linea thalassinica. All
specimens of A. aqabaensis have male and female gonopores,
but pleopods 1 and 2 are sexually dimorphic (Dworschak,
2003; Markham and Dworschak, 2005). Two species are
included besides the type species, A. brevirostris from Thailand
and A. lewtonae from Queensland, plus an undescribed species
from Papua New Guinea.
Arenallianassa gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:7A9C78DE-06CA4407-BA4A-0A2BA6258BDE
Type species. Callianassa arenosa Poore, 1975, by present
designation.
Diagnosis. Rostrum obsolete or obtusely triangular, flat, not
reaching cornea. Pleomere 1 tergite undivided or with weak
transverse ridge. Maxilliped 3 merus wider at ischium-merus
suture than long. Male major cheliped merus with prominent
truncate hook armed with serrations along lower margin,
excavate laterally at base; carpus and propodus flattened, upper
and lower margins carinate, blade-like, submarginal mesial
face especially of carpus deeply concave; propodus distal
margin with deep notch at base of fixed finger. Pereopod 3
propodus rectangular, lower margin deeply convex, leading to
broadly rounded free proximal lobe. Male pleopod 2 absent.
Uropodal endopod ovoid, usually longer than wide, anterior
margin straight or slightly convex, posterodistal margin
evenly convex, with facial distal transverse row of short
spiniform setae (reduced). Uropodal exopod distal margin
clearly differentiated from anterior margin, anterodistal corner
right-angled. Telson lateral margins convex.
Etymology. An alliteration of the name of the type species and
Callianassa, type genus of the family.
Remarks. Only the type species is known, a common shallow
subtidal species in south-eastern Australia. Arenallianassa
arenosa resembles Paratrypaea but lacks the dense setation on
the major cheliped of the latter genus and has a prominent
truncate meral hook on the major cheliped, whereas
Paratrypaea has a dentate blade, widest proximally instead.
The chelipeds resemble those of species of Neotrypaea, a
genus defined by a prominent distal lobe on the merus of
maxilliped 3, but this is not especially more developed in some
species than in A. arenosa. The telson is widest at its midpoint,
whereas species of Paratrypaea and Neotrypaea have a
tapering telson. Arenallianassa differs from Trypaea in lacking
an extremely long and setose antennule, and differs from
Filhollianassa in the anterior carapace being flat in profile and
the uropodal endopod being ovoid rather than asymmetrical.
Biffarius Manning and Felder, 1991
Biffarius Manning and Felder, 1991: 769–771.—Poore, 2004: 181
(partim).
Trypaea.—Sakai, 2011: 385–387 (partim; not Trypaea Dana,
1952).
Type species. Callianassa biformis Biffar, 1971, by original
designation.
Diagnosis. Rostrum obsolete or obtusely triangular, flat, not
reaching cornea. Pleomere 1 tergite undivided or with weak
transverse ridge. Antennular peduncle exceeded by most to all
of antennal peduncular article 5; articles 2 and 3 with single
lateral row of 6–10 well-spaced long setae along the lower
margin. Maxilliped 3 merus wider at ischium-merus suture
than long; crista dentata absent (or few proximal spines only).
Male major cheliped merus with prominent truncate hook
armed with serrations along lower margin, excavate laterally at
base; upper margins of ischium and merus, and lower margins
of merus and carpus beaded, with dense row of long fine
setae; propodus distal margin with deep notch at base of fixed
finger. Pereopod 3 propodus rectangular, lower margin deeply
convex, leading to broadly rounded free proximal lobe. Male
pleopod 2 present or absent. Uropodal endopod ovoid, usually
longer than wide, anterior margin straight or slightly convex,
posterodistal margin evenly convex. Uropodal exopod distal
margin clearly differentiated from anterior margin, anterodistal
corner right-angled, posterodistal margin with row of 6–8 long
blade-like setae proximal to long setae on distal margin. Telson
longer than wide, tapering evenly from near base; anterolateral
lobe defined posteriorly by short transverse slit; transverse
ridge with only fine setae; truncate or slightly convex between
posterolateral angles.
Remarks. Biffarius has been widely misused by several authors,
with up to a dozen species allocated to it, possibly because it
was a genus of last resort in Poore’s (1994) key. In fact, the type
species shares with one other species, B. limosus, and possibly
B. delicatulus, a long antennal peduncle and the lower margins
of the major cheliped merus and carpus being beaded and with
a dense row of long fine setae. Biffarius biformis occurs from
the North Atlantic to Caribbean waters, B. delicatulus occurs
in the South Atlantic and B. limosus is known from southeastern Australia, a somewhat enigmatic generic distribution.
Callianassa Leach, 1814
Callianassa Leach, 1814:
(comprehensive synonymy).
386,
400.—Sakai,
2011:
359
Type species. Cancer (Astacus) subterraneus Montagu, 1808,
by original designation and monotypy.
92
Diagnosis. Rostrum obsolete or obtusely triangular, flat, not
reaching cornea. Pleomere 1 tergite undivided or with weak
transverse ridge. Maxilliped 3 merus distally oblique with
obtuse angle between distal and lower margins, longer than
wide at ischium-merus suture. Male major cheliped merus with
simple proximal hook on lower margin. Pereopod 3 propodus
rectangular, lower margin deeply convex, leading to broadly
rounded free proximal lobe. Male pleopod 2 rarely present.
Uropodal endopod ovoid, usually longer than wide, anterior
margin straight or slightly convex, posterodistal margin evenly
convex or asymmetrical, at least as wide as long, distal margin
truncate-convex, at right angles to straight anterior margin.
Uropodal exopod distal margin clearly differentiated from
anterior margin, anterodistal corner right-angled.
Remarks.
Callianassa
differs
from
Paratrypaea,
Arenallianassa, Filhollianassa and Trypaea, genera it most
resembles in these analyses, in having the telson tapering
evenly from near base, the merus of maxilliped 3 longer than
wide and tapering, the major cheliped with simple proximal
hook on its lower margin and absence of facial setae on the
uropodal endopod.
Callianassa has been the common catch-all genus for
many species whose systematic position was uncertain. Sakai
(2011) restricted the genus to a single species but his diagnosis
of few characters could refer to any of numerous callianassid
genera. The name Montagua Leach, 1814, is sometimes listed
as a synonym. Montagua appears only in the index of Leach
(1814: 436) referring to “Genus 44”, Callianassa in the main
text. Gebios Risso, 1822, was treated as a junior synonym of
Callianassa by Sakai (2011) and in many earlier works. If
anything, Gebios Risso, 1822, is a synonym of Gilvossius and
is discussed below.
Caviallianassa gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:9819F120-10C6496D-80E2-BDDAD3FBAE07
Type species. Cheramus cavifrons Komai and Fujiwara, 2012,
by present designation.
Diagnosis. Rostrum obsolete or obtusely triangular, flat, not
reaching cornea. Pleomere 1 tergite undivided or with weak
transverse ridge. Antennular peduncle articles 2 and 3 with
single lateral row of 6–10 well-spaced long setae along lower
margin. Maxilliped 3 merus wider at ischium-merus suture
than long; dactylus ovate, with dense brush of long setae over
most of upper-distal margin, few setae along lower margin.
Male major cheliped merus with 1–3 oblique similar short
proximal spines on lower margin, beaded beyond; propodus
distal margin unornamented, oblique, or with small lateral
tooth. Minor cheliped two-thirds width of major cheliped,
both flattened. Pereopod 3 propodus rectangular, lower margin
deeply convex, leading to broadly rounded free proximal lobe.
Uropodal endopod ovoid, usually longer than wide, anterior
margin straight or slightly convex, posterodistal margin evenly
convex, with facial spiniform setae on rib. Uropodal exopod
posterodistal margin with row of 6–8 long blade-like setae
proximal to long setae on distal margin. Telson about as wide
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
as long, tapering from anterolateral lobe; anterolateral lobe
obsolete, undefined.
Etymology. An alliteration of the name of the type species and
Callianassa, type genus of the family.
Remarks. A group of two, possibly three, undescribed species
from Papua New Guinea and French Polynesia was revealed by
our molecular treatment (Robles et al., in press). Morphological
examination revealed these to be similar to Cheramus cavifrons,
of which only the female is known. The chelipeds are similarly
compact in all species and have a row of tubercles on the lower
margin of the merus beyond a proximal tooth or series of small
short spines. Komai and Fujiwara (2012) did not illustrate the
unusual setation of the dactylus of maxilliped 3 but its
proportions are otherwise similar to the undescribed species,
nor did they illustrate the spiniform setae on the face of the
uropodal endopod; both features are generally overlooked.
Caviallianassa shares with two genera related on genetic
and molecular evidence (Robles et al., in press) a maxilliped 3
dactylus with a dense brush of long setae over most of the
upper-distal margin and few setae along the lower margin.
Caviallianassa differs from the first, Rudisullianassa, in
having the antennular peduncle about as long as or a little
longer than the antennal peduncle (vs exceeded by all or most
of the antennal peduncular article 5) and having a small spine
on the merus of the major cheliped (vs smooth). Males and
females of Caviallianassa are known but only females with
male gonopores of Rudisullianassa are known. Caviallianassa
differs from Fragillianassa in having a wider minor cheliped,
lacking a prominent hook on the merus of the major cheliped
and having facial setae on the uropodal endopod.
Cheramoides Sakai, 2011
Cheramoides Sakai, 2011: 362.
Cheramus.—Sakai, 2011: 363–365 (partim).
Type species. Callianassa marginata Rathbun, 1901, by
original designation and monotypy.
Diagnosis. Rostrum acute, anteriorly directed, as long as
eyestalks. Pleomere 1 tergite fused, divided into 2 sections by
transverse step. Pleomere 6 with sublateral ventral sharp ridge,
flared posteriorly. Cornea with scattered reduced pigmentation.
Antennular peduncle articles 2 and 3 with single lateral row of
6–10 well-spaced long setae along lower margin. Antennal
scaphocerite simple, longer than wide, acute. Maxilliped 3
merus almost rectangular, distally truncate with squarish angle
between distal and lower margins, longer than wide at ischiummerus suture; crista dentata a prominent toothed ridge
extending beyond proximal margin of merus. Male major
cheliped merus without prominent hook or spine on lower
margin. Minor cheliped slender, narrower than major cheliped,
with attenuated curved dactylus, longer than palm. Pereopod 3
propodus oval, lower margin slightly convex, leading to narrow
sharply rounded proximal lobe. Male pleopod 2 absent.
Uropodal endopod ovoid, usually longer than wide, anterior
margin straight or slightly convex, posterodistal margin evenly
convex, with facial spiniform setae on rib. Uropodal exopod
93
Callianassidae and related families
twice as long as wide, distal margin clearly differentiated from
anterior margin, anterodistal corner right-angled, posterodistal
margin with row of 6–8 long blade-like setae proximal to long
setae on distal margin. Telson anterolateral lobe obsolete,
undefined; slightly concave, sometimes with medial spine.
Remarks. Cheramoides is unique in that the uropodal exopod
is twice as long as wide and much longer than the endopod. The
maxilliped 3 merus is rectangular, distally truncate with a
squarish angle between the distal and lower margins. Only C.
oblonga from West Africa and C. brachytelson from the
Andaman Sea are similar to the type species from the
Caribbean deep sea.
Cheramus Bate, 1888
Cheramus Bate, 1888: 30.—Manning and Felder, 1991: 91.—
Poore, 1994: 101.—Davie, 2002: 459.—Sakai, 2011: 363–366
(partim).—Komai et al., 2014b: 504–505 (partim).
Callianassa (Cheramus).—Borradaile, 1903: 545–546.—De Man,
1928: 26, 95.—Gurney, 1944: 8.
Type species. Cheramus occidentalis Bate, 1888, preoccupied,
replaced by Callianassa profunda Biffar, 1973, by subsequent
designation.
Diagnosis. Rostrum acute, anteriorly directed, as long as
eyestalks. Pleomere 1 tergite fused, divided into 2 sections by
transverse step. Pleomere 6 with sublateral ventral sharp ridge,
flared posteriorly. Cornea with scattered reduced pigmentation.
Antennular peduncle exceeded by all or most of antennal
peduncular article 5. Maxilliped 3 merus almost rectangular,
distally truncate with squarish angle between distal and lower
margins, longer than wide at ischium-merus suture; crista
dentata a prominent toothed ridge extending beyond proximal
margin of merus. Male major cheliped merus with serrate
blade over lower margin, upper margin concave. Minor and
major chelipeds similar, both attenuated, with swollen
palms; carpus upper margin as long as or shorter than propodus.
Pereopod 3 propodus oval, lower margin slightly convex,
leading to narrow sharply rounded proximal lobe. Male
pleopod 2 present. Uropodal endopod ovoid, usually longer
than wide, anterior margin straight or slightly convex,
posterodistal margin evenly convex. Uropodal exopod about
1.5–1.8 times as long as wide, distal margin clearly
differentiated from anterior margin, anterodistal corner rightangled. Telson anterolateral lobe obsolete, undefined.
Remarks. Cheramus is similar to Cheramoides but differs in
having uniquely similar minor and major chelipeds, both
attenuated and with swollen palms. Contrary to the views in
recent accounts, only the type species agrees with this
characterisation. Komai et al. (2014b) explained the complex
taxonomic history of the type species but followed Manning
and Felder (1991) in including other species, C. marginatus
(type species of Cheramoides), C. orientalis and C. oblonga.
Several other species have been included since 1991 (Komai et
al., 2014b), but it has to be concluded from Robles et al.’s (in
press) phylogeny, supported by morphology, that these and the
16 species included by Sakai (2011) are a diverse assemblage.
Komai et al. (2014b) could list only “possibly diagnostic
characters”, most of which, in our analyses, are features of
more than one genus.
Coriollianassa gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:A26A8A5B-426A4110-9945-31A8B1F5831B
Type species. Callianassa coriolisae Ngoc-Ho, 2014, by
present designation.
Diagnosis. Rostrum acute, anteriorly directed, as long as
eyestalks, with a ventral keel. Pleomere 1 tergite fused, divided
into 2 sections by transverse step. Cornea with scattered
reduced pigmentation. Antennular peduncle exceeded by all or
most of antennal peduncular article 5; articles 2 and 3 with
single lateral row of 6–10 well-spaced long setae along lower
margin. Antennal scaphocerite bifid. Maxilliped 3 merus
distally oblique with obtuse angle between distal and lower
margins, longer than wide at ischium-merus suture; crista
dentata a prominent toothed ridge extending beyond proximal
margin of merus. Male major cheliped merus with proximal
curved spine on lower margin; carpus articulating by means
of a short neck with merus. Pereopod 3 propodus oval, lower
margin slightly convex, leading to narrow sharply rounded
proximal lobe. Male pleopod 2 present or absent. Uropodal
endopod ovoid, usually longer than wide, anterior margin
straight or slightly convex, posterodistal margin evenly convex,
with facial spiniform setae on rib. Uropodal exopod about 1.5–
1.8 times as long as wide, posterodistal margin with row of 6–8
long blade-like setae proximal to long setae on distal margin.
Telson anterolateral lobe obsolete, undefined.
Etymology. An alliteration of the name of the type species and
Callianassa, type genus of the family.
Remarks. Robles et al. (in press) recognised a well-defined
genus on molecular and morphological grounds. Two of the
four species included were undescribed. The genus is defined
by the sharp anteriorly directed rostrum with a ventral keel, the
long antennal peduncle, bifid scaphocerite, the unusual neck at
the base of the carpus of the chelipeds and the usually bifid
meral tooth on the major cheliped. Ngoc-Ho (2014) recognised
the similarity between C. coriolisae and C. sibogae, notably
the characteristic chelipeds, and realised their differences from
Cheramus where they had been placed.
Filhollianassa gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:9A34F27E-ABF546ED-AFE8-BA399225F2C6
Type species. Callianassa filholi A. Milne-Edwards, 1878, by
present designation.
Diagnosis. Anterior carapace strongly domed, depressed
over anterior quarter. Rostrum obsolete or obtusely triangular,
flat, not reaching cornea. Pleomere 1 tergite undivided or with
weak transverse step. Antennular peduncle length about 2.5–3
times the width of both eyestalks. Maxilliped 3 merus wider at
ischium-merus suture than long. Male major cheliped merus
with prominent truncate hook armed with serrations along
94
lower margin, excavate laterally at base; carpus and propodus
flattened, upper and lower margins carinate, blade-like,
submarginal mesial face especially of carpus deeply concave;
propodus distal margin with deep notch at base of fixed finger.
Pereopod 3 propodus rectangular, lower margin deeply convex,
leading to broadly rounded free proximal lobe. Male pleopod 2
absent. Uropodal endopod asymmetrical, at least as wide as
long, distal margin truncate-convex, at right angles to straight
anterior margin, with facial distal transverse row of short
spiniform setae. Uropodal exopod posterodistal margin with
row of 6–8 long blade-like setae proximal to long setae on
distal margin. Telson lateral margins convex.
Etymology. An alliteration of the name of the type species and
Callianassa, type genus of the family.
Remarks. The two species of Filhollianassa from south-eastern
Australia and New Zealand are immediately recognisable from
the strongly domed anterior carapace and strongly inwardly
curved carinate upper and lower margins of the carpus and
propodus of the major chelipeds. The major cheliped of Trypaea
australiensis, also in south-eastern Australia but ecologically
separate, is similar but this species has a flat dorsum, much
longer and more setose antennule, considerably expanded merus
of maxilliped 3 and narrower uropodal endopod.
Fragillianassa gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:CFA11DDC-A9C54ACC-AC90-E2C6A5456E82
Type species. Callianassa fragilis Biffar, 1970, by present
designation.
Diagnosis. Rostrum obsolete or obtusely triangular, flat, not
reaching cornea. Pleomere 1 tergite undivided or with weak
transverse step. Antennular peduncle length about 2.5–3
times the width of both eyestalks. Antennal scaphocerite
reduced to small floating disc. Maxilliped 3 merus wider at
ischium-merus suture than long; dactylus ovate, with dense
brush of long setae over most of upper-distal margin, few
setae along lower margin. Male major cheliped merus with
prominent truncate hook armed with serrations along lower
margin, excavate laterally at base; propodus distal margin with
deep notch at base of fixed finger. Pereopod 3 propodus
rectangular, lower margin deeply convex, leading to broadly
rounded free proximal lobe. Male pleopod 2 absent. Uropodal
endopod ovoid, usually longer than wide, anterior margin
straight or slightly convex, posterodistal margin evenly convex.
Uropodal exopod posterodistal margin with row of 6–8 long
blade-like setae proximal to long setae on distal margin. Telson
truncate, or slightly convex between posterolateral angles, or
slightly concave, sometimes with medial spine.
Etymology. An alliteration of the name of the type species and
Callianassa, type genus of the family.
Remarks. Fragillianassa, with two species recognised so far, is
most similar to Necallianassa and Neotrypaea, with which it
shares a similar eyestalk, antennule and maxilliped 3 merus.
Fragillianassa differs from both in having an extremely
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
reduced scaphocerite, a dense brush of long setae over most of
the upper-distal margin of the dactylus of maxilliped 3 and an
ovoid uropodal endopod longer than wide, more symmetrical
than in the other two. Fragillianassa differs from Necallianassa
most obviously in lacking spines on the sides of the telson and
on the anterior margin of the uropodal endopod, and it differs
from Neotrypaea in lacking a transverse facial row of setae on
the uropodal endopod.
Gilvossius Manning and Felder, 1992
Gebios Risso, 1822: 243 (type species, Gebios davianus Risso,
1822, junior subjective synonym of Cancer candidus Olivi, 1792, by
monotypy) nomen oblitum.
Gebius Agassiz, 1846: 160 (unjustified emendation of Gebios
Risso, 1822).
Gilvossius Manning and Felder, 1992: 558.—Sakai, 2011: 372
(partim).
Pestarella Ngoc-Ho, 2003: 475 (type species: Astacus tyrrhenus
Petagna, 1792, by original designation).
Type species. Gonodactylus setimanus DeKay, 1844, by
original designation and monotypy.
Diagnosis. Rostrum obsolete or obtusely triangular, flat, not
reaching cornea. Pleomere 1 tergite fused, divided into 2
sections by transverse step. Antennular peduncle length about
2.5–3 times the width of both eyestalks, length exceeding
narrower antennal peduncle. Maxilliped 3 merus wider at
ischium-merus suture than long. Male major cheliped merus
with prominent truncate hook armed with serrations along
lower margin, excavate laterally at base. Pereopod 3 propodus
rectangular, lower margin deeply convex, leading to broadly
rounded free proximal lobe. Male pleopod 2 present or absent.
Uropodal endopod ovoid, usually longer than wide, anterior
margin straight or slightly convex, posterodistal margin evenly
convex. Telson wider than long, almost semicircular; or
rarely posterior margin subtruncate.
Remarks. The genus is recognised by the telson being wider than
long, almost semicircular in most species, but rarely subtruncate
posteriorly. Gilvossius differs from Trypaea, Arenallianassa
and Paratrypaea, which share uropodal exopods with densely
setose margins, in lacking a deep notch at the base of the cheliped
fingers and lacking a facial distal transverse row of short
spiniform setae on the uropodal endopod.
Manning and Felder (1992) and Ngoc-Ho (2003)
compared Gilvossius and Pestarella respectively with
Callianassa, but this latter genus is only remotely similar in
Robles et al.’s (in press) analyses. Both Pestarella and
Gilvossius are in the recent literature. Gilvossius was erected
for a single species, Gonodactylus setimanus, shown by
molecular and morphological evidence (Robles et al., in
press) to be congeneric with Gilvossius tyrrhenus, G.
candidus and G. whitei (Sakai, 1999a). Ngoc-Ho (2003)
erected Pestarella for the type species (Astacus tyrrhenus),
G. candidus and three others; Pestarella is here treated as a
subjective junior synonym of Gilvossius.
Gebios davianus Risso, 1822, is generally agreed to be a
junior subjective synonym of Cancer candidus Olivi, 1792.
Gebios, therefore, could be invoked as a senior subjective
95
Callianassidae and related families
synonym of both Pestarella Ngoc-Ho, 2003, and Gilvossius
Manning and Felder, 1992. However, Gebios has never been
used as a generic name. Ngoc-Ho (2003) provided a
comprehensive synonymy of Pestarella, a junior synonym of
Gilvossius, and its included species. The conditions of the
International Code of Zoological Nomenclature Article 23.9.1
required for prevailing usage are met. We therefore invoke
International Code of Zoological Nomenclature Article 23.9
and designate Gebios Risso, 1822, as nomen oblitum and both
Pestarella Ngoc-Ho, 2003, and Gilvossius Manning and
Felder, 1992, as nomina protecta.
Jocullianassa gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:1AC3D163-889545EF-8EC6-E87B435B9FDF
Type species. Callianassa joculatrix De Man, 1905, by present
designation.
Diagnosis. Rostrum acute, anteriorly directed, as long as
eyestalks. Pleomere 1 tergite fused, divided into 2 sections by
transverse step. Cornea with scattered reduced pigmentation.
Antennular peduncle articles 2 and 3 with single lateral row of
6–10 well-spaced long setae along lower margin. Antennal
scaphocerite simple, longer than wide, acute. Maxilliped 3
merus tapering, not mesially produced, longer than wide at
ischium-merus suture. Male major cheliped merus with
proximal perpendicular spine on lower margin; propodus distal
margin with deep notch at base of fixed finger. Pereopod 3
propodus bean-shaped, lower margin concave, leading to
broadly rounded proximal lobe. Male pleopod 2 absent.
Uropodal endopod ovoid, usually longer than wide, anterior
margin straight or slightly convex, posterodistal margin evenly
convex. Uropodal exopod about 1.5–1.8 times as long as wide,
distal margin clearly differentiated from anterior margin,
anterodistal corner right-angled, posterodistal margin with row
of 6–8 long blade-like setae proximal to long setae on distal
margin. Telson anterolateral lobe obsolete, undefined.
Etymology. An alliteration of the name of the type species and
Callianassa, type genus of the family.
Remarks. Jocullianassa joculatrix is relatively easily
recognised by the combination of its acute rostrum, sharp basal
spine on the merus of the major cheliped and the excavate
lower margin of the propodus on pereopod 3 (Komai and
Tachikawa, 2008). Jocullianassa joculatrix is far removed
from the type species of Trypaea, the genus in which it was
included by Sakai (2011). The species is common in collections
from the Indo-West Pacific.
Lipkecallianassa Sakai, 2002
Lipkecallianassa Sakai, 2002: 477.—Sakai, 2005b: 212.—Sakai,
2011: 522.
Type species. Lipkecallianassa abyssa Sakai, 2002, by original
designation and monotypy.
Diagnosis. Rostrum acute, anteriorly directed, as long as
eyestalks. Cornea with scattered reduced pigmentation.
Maxilliped 3 merus almost rectangular, distally truncate with
squarish angle between distal and lower margins, longer than
wide at ischium-merus suture, with acute distomesial angle.
Pereopod 3 propodus linear, without lobe on lower margin.
Uropodal endopod ovoid, usually longer than wide, anterior
margin straight or slightly convex, posterodistal margin evenly
convex. Uropodal exopod about 1.5–1.8 times as long as wide.
Telson anterolateral lobe obsolete, undefined; posterior margin
with pair of broad posterolateral lobes, widely excavate at
midpoint, with mesial spine.
Remarks. The monotypic genus Lipkecallianassa is known
from a single damaged incomplete specimen of the type
species. But based on Sakai’s (2002) short description and
figures, the species has a linear propodus of pereopod 3, seen
elsewhere only in Praedatrypaea longicauda and P. modesta
but neither of these species has a strongly excavate posterior
telsonic margin. The telson resembles that of species of
Pugnatrypaea in having an excavate posterior margin; both
genera have a narrow ischium-merus of maxilliped 3. All of
these three genera have a tooth or spine on the distal margin of
the merus of maxilliped 3 but are not allied in the molecular or
morphological trees of Robles et al. (in press).
Necallianassa Heard and Manning, 1998
Necallianassa Heard and Manning, 1998: 883–884.
Trypaea.—Sakai, 2011: 385–387 (partim) (not Trypaea Dana, 1952).
Type species. Necallianassa berylae Heard and Manning,
1998, by original designation.
Diagnosis. Rostrum obsolete or obtusely triangular, flat, not
reaching cornea, or acute, anteriorly directed, as long as
eyestalks. Pleomere 1 tergite undivided or with weak transverse
step. Antennular peduncle length about 2.5–3 times the width
of both eyestalks. Antennal scaphocerite simple, longer than
wide, acute. Maxilliped 3 merus wider at ischium-merus suture
than long. Male major cheliped merus with prominent truncate
hook armed with serrations along lower margin, excavate
laterally at base; propodus distal margin with deep notch at
base of fixed finger. Pereopod 3 propodus rectangular, lower
margin deeply convex, leading to broadly rounded free
proximal lobe. Male pleopod 2 absent. Uropodal endopod
asymmetrical, at least as wide as long, distal margin truncateconvex, at right angles to straight anterior margin, anterior
margin with distal spine. Uropodal exopod posterodistal
margin with row of 6–8 long blade-like setae proximal to long
setae on distal margin.
Remarks. The sharp spine on the anterior margin of the
uropodal endopod and usually one or two spines on the lateral
margins of the telson uniquely differentiate Necallianassa
from all other callianassids (Heard and Manning, 1998).
Necallianassa truncata lacks lateral spines on the telson but
the male major cheliped has the same setose propodus and
dactylus as N. acanthura (cf. Ngoc-Ho, 2003). The genus was
synonymised with the very different Trypaea by Sakai (2011)
without explanation.
96
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Neotrypaea Manning and Felder, 1991
Notiax Manning and Felder, 1991
Neotrypaea Manning and Felder, 1991: 711–712.
Nihonotrypaea Manning and Tamaki, 1998: 889–891 (type
species, Callianassa japonica Ortmann, 1891, by original designation)
syn. nov.
Pseudobiffarius Heard and Manning, 2000: 70.—Sakai, 2005b:
26 (as synonym of Callianassa).—Sakai, 2011: 286 (as synonym of
Trypaea) (type species, Pseudobiffarius caesari Heard and Manning,
2000, by original designation and monotypy) syn. nov.
Trypaea.—Sakai, 2011: 385–387 (partim; not Trypaea Dana, 1852).
Notiax Manning and Felder, 1991: 772–773.—Sakai, 2005b: 18
(as synonym of Callianassa).
Not Notiax.—Sakai, 2011: 381–382.
Type species. Callianassa californiensis Dana, 1854, by
original designation.
Diagnosis. Rostrum obsolete or obtusely triangular, flat, not
reaching cornea. Pleomere 1 tergite undivided or with weak
transverse step. Eyestalk distal lobes triangular, apices
separate or contiguous, produced and tapering over distal
half. Antennular peduncle length about 2.5–3 times the width
of both eyestalks. Maxilliped 3 merus expanded distomesially
as rounded lobe beyond articulation with ischium, wider at
ischium-merus suture than long. Male major cheliped merus
with prominent truncate hook armed with serrations along
lower margin, excavate laterally at base; propodus distal
margin with deep notch at base of fixed finger. Pereopod 3
propodus rectangular, lower margin deeply convex, leading to
broadly rounded free proximal lobe. Male pleopod 2 absent.
Uropodal endopod asymmetrical, at least as wide as long,
distal margin truncate-convex, at right angles to straight
anterior margin, with facial distal transverse row of short
spiniform setae. Uropodal exopod distal margin clearly
differentiated from anterior margin, anterodistal corner rightangled. Telson lateral margins convex; posterior margin
slightly concave, sometimes with medial spine.
Remarks. Neotrypaea probably includes more species than any
genus of Callianassidae. Species have a triangular distal lobe
on the eyestalk, the appendix interna of pleopods 3–5 embedded
in the endopod, or only slightly protruding, and the merus of
maxilliped 3 projecting as a rounded lobe beyond the
articulation of the carpus (Manning and Felder, 1991) but not
as much as in Trypaea. The uropodal endopod is asymmetrical,
at least as wide as long, with the distal margin truncate-convex,
at right angles to straight anterior margin.
Manning and Tamaki (1998) differentiated Nihonotrypaea
with the appendix interna of pleopods 3–5 projecting from the
endopod margin from Neotrypaea, in which the appendix
interna is completely embedded in the margin. The same
difference was used for erection of Pseudobiffarius by Heard
and Manning (2000) but this difference is slight. Two clades
were evident in the molecular analyses of Robles et al. (in
press) but not in the morphological treatment. Species assigned
to Nihonotrypaea were reported to differ from Neotrypaea in
having the antennular peduncle shorter than the antennal
peduncle (Lin et al., 2007a) but this is true, or they are of
similar lengths, in all species from both genera. Both
Nihonotrypaea and Pseudobiffarius were synonymised with
Trypaea by Sakai (2011).
Type species. Callianassa brachyophthalma A. MilneEdwards, 1870, by original designation and monotypy.
Diagnosis. Rostrum acute, reaching middle of cornea, with
ventral broad swelling. Pleomere 1 tergite undivided or with
weak transverse step. Maxilliped 3 merus wider at ischiummerus suture than long; crista dentata absent (or few proximal
spines only). Male major cheliped merus with prominent
truncate hook armed with serrations along lower margin,
excavate laterally at base; carpus and propodus flattened, upper
and lower margins carinate, blade-like, submarginal mesial
face especially of carpus deeply concave. Pereopod 3 propodus
rectangular, lower margin deeply convex, leading to broadly
rounded free proximal lobe. Male pleopod 2 present or absent.
Uropodal endopod asymmetrical, at least as wide as long,
distal margin truncate-convex, at right angles to straight
anterior margin, with facial distal transverse row of short
spiniform setae. Telson longer than wide, tapering evenly from
near base; anterolateral lobe defined posteriorly by short
transverse slit; transverse ridge with only fine setae; truncate or
slightly convex between posterolateral angles.
Remarks. Manning and Felder (1991) differentiated Notiax
from other callianassids because of its “rostral spine” but it is
not the only genus with this feature. Notiax differs, at least as
adults, in having a broad swelling under the rostrum not seen in
others. The telson is longer than wide, tapering from its base,
whereas it is more rectangular in similar genera such as
Neotrypaea and Arenallianassa. The maxilliped 3 merus is
not lobed as in Neotrypaea.
Sakai (2011) redefined Notiax to include, besides the type
species, five species here included in Rayllianassa, Filhollianassa
and Biffarius. His diagnosis included alternate states for the
shapes of the maxilliped 3, male pleopods 1 and 2 and the telson.
Nothing was given that would unify these five species.
Paratrypaea Komai and Tachikawa, 2008
Paratrypaea Komai and Tachikawa, 2008: 36.—Komai and
Fujita, 2014: 5–6.
Gilvossius.—Sakai, 2011: 372–373 (partim).
Type species. Callianassa (Trypaea) bouvieri Nobili, 1904, by
original designation.
Diagnosis. Rostrum broadly or sharply triangular, almost or as
long as eyestalks. Pleomere 1 tergite fused, divided into 2
sections by transverse step. Antennular peduncle length about
2.5–3 times the width of both eyestalks. Maxilliped 3 merus
wider at ischium-merus suture than long. Male major cheliped
merus with blade dentate over lower margin, widest proximally;
carpus and propodus flattened, upper and lower margins
carinate, blade-like, submarginal mesial face especially of
carpus deeply concave; propodus distal margin with deep notch
at base of fixed finger; upper distal margin of propodus and
dactylus with dense brush of setae. Pereopod 3 propodus
97
Callianassidae and related families
rectangular, lower margin deeply convex, leading to broadly
rounded free proximal lobe. Male pleopod 2 absent. Uropodal
endopod ovoid, usually longer than wide, anterior margin
straight or slightly convex, posterodistal margin evenly convex,
with facial distal transverse row of short spiniform setae.
Uropodal exopod distal margin differentiated from anterior
margin, anterodistal corner rounded or right-angled. Telson
longer than wide, tapering evenly from near base; anterolateral
lobe defined posteriorly by short transverse slit; transverse
ridge with only fine setae; truncate or slightly convex between
posterolateral angles.
Remarks. Species of Paratrypaea are immediately recognisable
by the dense brush of setae on the propodus and lateral dactylus
of the major cheliped, and the merus of the male major cheliped
having a dentate blade on its lower margin, rather than a hook
as in many other genera.
Paratrypaea was synonymised with Gilvossius by Sakai
(2011) because the two genera share the same type of male
pleopods 1 and 2, but the two genera differ in many ways
(Komai and Fujita, 2014). Gilvossius chichijimaensis Sakai,
2015, was shown to be a junior synonym of Paratrypaea
bouvieri (see Komai, 2017).
Poti Rodrigues and Manning, 1992
Poti Rodrigues and Manning, 1992a: 9–10.
Callianassa.—Sakai, 1999a: 11–13 (partim).
Cheramus.—Sakai, 2011: 363–365 (partim).
Type species. Poti gaucho Rodrigues and Manning, 1992, by
original designation and monotypy.
Diagnosis. Rostrum acute, anteriorly directed, as long as
eyestalks. Pleomere 1 tergite fused, divided into 2 sections by
transverse step. Cornea with scattered reduced pigmentation.
Antennular peduncle articles 2 and 3 with single lateral row of
6–10 well-spaced long setae along lower margin. Maxilliped 3
merus almost rectangular, distally truncate with squarish
angle between distal and lower margins, longer than wide at
ischium-merus suture; crista dentata a prominent toothed ridge
extending beyond proximal margin of merus. Pereopod 3
propodus oval, lower margin slightly convex, leading to narrow
sharply rounded proximal lobe. Male pleopod 2 present.
Uropodal endopod ovoid, usually longer than wide, anterior
margin straight or slightly convex, posterodistal margin evenly
convex. Uropodal exopod about 1.5–1.8 times as long as wide,
distal margin clearly differentiated from anterior margin,
anterodistal corner right-angled, posterodistal margin with row
of 6–8 long blade-like setae proximal to long setae on distal
margin. Telson anterolateral lobe obsolete, undefined.
Remarks. Poti was said to differ from all other callianassid
genera in having an incomplete linea thalassinica (Rodrigues
and Manning, 1992a), a condition we were unable to confirm.
Otherwise, the maxilliped 3 and minor cheliped of the only
species resembles those of species of Cheramoides. The
uropodal exopod is broad, while it is exceptionally long and
narrow in Cheramoides.
Poti was synonymised with Callianassa by Sakai (1999a),
along with seven other genera, and with a redefined Cheramus
by Sakai (2011) who took a broad view of both genera.
Praedatrypaea gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:CDBA883E-288C4CDA-80F4-1BEE665FA3D0
Type species. Callianassa praedatrix De Man, 1905, by present
designation.
Diagnosis. Rostrum acute, anteriorly directed, as long as
eyestalks. Pleomere 1 tergite fused, divided into 2 sections
by transverse step. Pleomere 6 with sublateral ventral sharp
ridge, flared posteriorly. Antennular peduncle exceeded by
all or most of antennal peduncular article 5; articles 2 and 3
with single lateral row of 6–10 well-spaced long setae along
lower margin. Antennal scaphocerite simple, longer than
wide, acute. Maxilliped 3 merus wider at ischium-merus
suture than long, with distal spine on distal free margin.
Male major cheliped merus with blade dentate over lower
margin, widest proximally. Pereopod 3 propodus oval, lower
margin slightly convex, leading to narrow sharply rounded
proximal lobe. Uropodal endopod ovoid, usually longer than
wide, anterior margin straight or slightly convex, posterodistal
margin evenly convex, with spiniform setae near anterior
and distal margins. Uropodal exopod about 1.5–1.8 times as
long as wide, distal margin clearly differentiated from
anterior margin, anterodistal corner right-angled,
posterodistal margin with row of 6–8 long blade-like setae
proximal to long setae on distal margin. Telson anterolateral
lobe prominent, defined posteriorly by clear unchitinised
region; truncate or slightly convex between posterolateral
angles, or slightly concave, posterior margin sometimes with
medial spine.
Etymology. An alliteration of the name of the type species and
Trypaea, a genus of the family.
Remarks. The hooked spine on the distal margin of the merus
of maxilliped 3 immediately defines Praedatrypaea. The
genus is also unusual in that the proximolateral lobes at the
base of the telson are prominent, especially ventrally, and are
defined by a weakly chitinised suture dorsally and laterally. In
other genera, these lobes may be evident and defined at most by
a narrow slit.
Pugnatrypaea gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:0F41845B-F5264AB8-80B7-A60B6538919E
Type species. Callianassa pugnatrix De Man, 1905, by present
designation.
Diagnosis. Rostrum acute, anteriorly directed, as long as
eyestalks. Pleomere 1 tergite fused, divided into 2 sections by
transverse step. Cornea with scattered reduced pigmentation.
Antennular peduncle from about as long exceeded by all of
antennal peduncular article 5; articles 2 and 3 with single
lateral row of 6–10 well-spaced long setae along lower margin.
98
Maxilliped 3 merus distally oblique with obtuse angle between
distal and lower margins, without distal spine on mesial
margin. Male major cheliped merus with simple proximal hook
on lower margin; propodus distal margin with deep notch at
base of fixed finger. Pereopod 3 propodus oval, lower margin
slightly convex, leading to narrow sharply rounded proximal
lobe. Male pleopod 2 present or absent. Uropodal endopod
ovoid, usually longer than wide, anterior margin straight or
slightly convex, posterodistal margin evenly convex, with
spiniform setae near anterior and distal margins, or with facial
spiniform setae on rib. Uropodal exopod about 1.5–1.8 times as
long as wide, distal margin clearly differentiated from anterior
margin, anterodistal corner right-angled, posterodistal margin
with row of 6–8 long blade-like setae proximal to long setae on
distal margin. Telson anterolateral lobe obsolete, undefined;
tapering over distal third to pair of posterior lobes separated
by deep notch, with medial spine.
Etymology. An alliteration of the name of the type species and
Trypaea, a genus of the family.
Remarks. The telson tapers over its distal third to a pair of
lobes separated by a deep notch with a medial spine. The two
pairs of spiniform setae typically found on the posterolateral
angles of the telson of callianassids are prominently enlarged
and displaced anteriorly. Pugnatrypaea pugnatrix and an
unidentified species Pugnatrypaea GMX lie on the same
molecular clade (Robles et al. 2019). Both have a simple curved
spine on the lower margin of the merus of the major cheliped.
The antennular peduncle is shorter than the antennal peduncle
in Pugnatrypaea GMX (fig. 6j) while they appear to be of
similar lengths in Pugnatrypaea pugnatrix (De Man, 1928: fig.
23). Species with similar maxillipeds 3, telsons and short
antennular peduncles are P. intermedia and P. lobetobensis but
these differ in having a truncate, instead of rounded, uropodal
exopod and the merus of the major cheliped dentate along the
lower margin and with a spine on the upper margin.
Rayllianassa Komai and Tachikawa, 2008
Rayllianassa Komai and Tachikawa, 2008: 42–43.—Komai and
Fujita, 2014: 551–552.
Notiax.—Sakai, 2011: 381 (partim).
Type species. Callianassa amboinensis De Man, 1888, by
original designation and monotypy.
Diagnosis. Hermaphrodite. Rostrum obsolete or obtusely
triangular, flat, not reaching cornea. Pleomere 1 tergite
undivided or with weak transverse step. Eyestalk distal lobes
obliquely truncated, apices diverging. Antennular peduncle
exceeding antennal peduncle by about half length of article 3;
articles 2 and 3 with single lateral row of 6–10 well-spaced long
setae along lower margin. Antennal scaphocerite simple, longer
than wide, acute. Mandibular molar calcified, swollen projection
without sharp edge, incisor with few teeth. Maxilliped 3 merus
wider at ischium-merus suture than long. Male major cheliped
merus without prominent hook or spine on lower margin;
carpus and propodus ovoid in cross-section, upper and lower
margins not markedly carinate; propodus distal margin with
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
small lateral tooth. Minor cheliped about two-thirds width of
major cheliped, both swollen; carpus upper margin as long as or
shorter than propodus. Pereopod 3 propodus rectangular, lower
margin deeply convex, leading to broadly rounded free proximal
lobe. Uropodal endopod ovoid, usually longer than wide,
anterior margin straight or slightly convex, posterodistal margin
evenly convex, with facial spiniform setae on rib. Uropodal
exopod posterodistal margin with row of 6–8 long blade-like
setae proximal to long setae on distal margin. Telson lateral
margins convex; anterolateral lobe obsolete, undefined; slightly
concave, sometimes with medial spine.
Remarks. Rayllianassa amboinensis is recognisable by the
absence of a hook on the merus of the major cheliped, the
propodi being swollen and the pair being little different in size.
The lobes on the eyestalks are truncate-oblique and the
maxilliped 3 particularly broad. Komai et al. (2014a) added a
second species R. rudisulcus and emended the generic
diagnosis to accommodate the presence or absence of a dorsal
oval, but the authors were ambivalent about the value of this
character in callianassid systematics. This species and another
were shown by Robles et al. (in press) to differ on both
molecular and morphological criteria (see Rudisullianassa
below where the two genera are compared).
Rayllianassa amboinensis has been reported from
throughout the Indo-West Pacific and illustrated several times
since its discovery in Ambon, Indonesia (Komai et al., 2014a;
Komai and Tachikawa, 2008; Ngoc-Ho, 2005; Poore and Griffin,
1979; Sakai, 1984, 1988, 1999a). There is considerable
discrepancy between the illustrations. Robles et al. (in press)
found substantial genetic difference between eight individuals
from Papua New Guinea, Philippines and the Line Islands
pointing to the probability of more than one species in this genus.
All authors cited above have reported only females with
the exception of Sakai (1999a) who reported two males and a
female from Ambon, all twice as long as the “female” holotype
and other subsequent records. Examination of all the material
available to us shows them to be hermaphrodites with both
male and female gonopores. Ngoc-Ho (1991) illustrated minute
pleopods 1 and 2 on a single male that was renamed
Callianassa ngochoae by Sakai (1999a) and later synonymised
by Komai et al. (2014a). The generic position of Callianassa
ngochoae is uncertain. Males of species of Rudisullianassa
and Spinicallianassa have also never been reported. All
“females” with typical pleopods 1 and 2, including ovigerous
individuals, have both female gonopores on coxae of pereopods
3 and male gonopores on coxae of pereopods 5 and are
probable hermaphrodites.
Rudisullianassa gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:C2EFC849-6C6B4466-8B47-9DF66EBF5D46
Type species. Rayllianassa rudisulcus Komai, Fujita and
Maenosono, 2014, by present designation.
Diagnosis. Hermaphrodite. Rostrum obsolete or obtusely
triangular, flat, not reaching cornea. Pleomere 1 tergite
undivided or with weak transverse step. Antennular peduncle
99
Callianassidae and related families
exceeded by all or most of antennal peduncular article 5;
articles 2 and 3 with single lateral row of 6–10 well-spaced
long setae along lower margin. Maxilliped 3 merus wider at
ischium-merus suture than long; dactylus ovate, with dense
brush of long setae over most of upper-distal margin, few
setae along lower margin. Male major cheliped merus without
prominent hook or spine on lower margin; propodus distal
margin with small lateral tooth. Minor cheliped about twothirds width of major cheliped, both swollen; carpus upper
margin as long as or shorter than propodus. Pereopod 3
propodus oval, lower margin slightly convex, leading to narrow
sharply rounded proximal lobe. Uropodal endopod ovoid,
usually longer than wide, anterior margin straight or slightly
convex, posterodistal margin evenly convex, with or without
facial spiniform setae on rib. Uropodal exopod 1.0–1.8 times as
long as wide, posterodistal margin with row of 6–8 long bladelike setae proximal to long setae on distal margin. Telson about
as wide as long, tapering from anterolateral lobe; anterolateral
lobe obsolete, undefined; posterior margin slightly concave,
sometimes with medial spine.
Etymology. An alliteration of the name of the type species and
Callianassa, type genus of the family.
Remarks. Komai et al. (2014a) described Rayllianassa
rudisulcus based on one female; they did not note how the
setation on the dactylus of maxilliped 3, which they correctly
described as “stout”, differed from that of R. amboinensis. The
form found in Rayllianassa, tapering with few long setae on
the upper surface and a dense brush of short setae below, is
typical of most callianassids. Rudisullianassa differs further
from Rayllianassa in having the antennular peduncle shorter
than the antennal peduncle, whereas the opposite is true in
Rayllianassa, and the minor cheliped is relatively smaller. The
atypical maxilliped 3 dactylus seen in R. rudisulcus is also
seen in Biffarius, Fragillianassa and Caviallianassa but these
are gonochoristic genera. Rudisullianassa rudisulcus and a
second undescribed species were common in collections from
Papua New Guinea (Robles et al. 2019). All individuals had
both male and female gonopores. Both species were associated
with submerged wood.
The unusual maxilliped 3 was initially mistaken by one of
us (GCBP) for the form seen in eucalliacids, also with swollen
similar chelipeds. In eucalliacids, the dactylus is also
extremely setose but in this case the apex is decidedly truncate.
Scallasis Bate, 1888
Scallasis Bate, 1888: 34.—Manning and Felder, 1991: 780.
Callianassa (Scallasis).—Borradaile, 1903: 547–548 (partim).
Cheramus.—Sakai, 2011: 363–365 (partim).
Type species. Scallasis amboinae Bate, 1888, by monotypy.
Diagnosis. Rostrum acute, anteriorly directed, as long as
eyestalks. Pleomere 1 tergite fused, divided into 2 sections by
transverse step. Pleomere 6 with sublateral ventral sharp ridge,
flared posteriorly. Eyestalk distal lobes acute, apices separate.
Antennular peduncle articles 2 and 3 with single lateral row of
6–10 well-spaced long setae along lower margin. Antennal
scaphocerite simple, longer than wide, acute. Maxilliped 3
merus distally oblique with obtuse angle between distal and
lower margins, longer than wide at ischium-merus suture;
crista dentata a prominent toothed ridge extending beyond
proximal margin of merus. Male major cheliped merus with 2
or 3 proximal similar teeth and distal denticles on lower
margin; propodus distal margin with small lateral tooth.
Pereopod 3 propodus oval, lower margin slightly convex,
leading to narrow sharply rounded proximal lobe. Uropodal
endopod ovoid, usually longer than wide, anterior margin
straight or slightly convex, posterodistal margin evenly convex,
anterior margin unarmed or with spine at midpoint, with facial
spiniform setae on rib. Uropodal exopod 1.0–1.8 times as long
as wide, posterodistal margin with row of 6–8 long blade-like
setae proximal to long setae on distal margin. Telson lateral
margins convex; transverse ridge with fine setae and spiniform
setae; posterior margin truncate, or slightly convex between
posterolateral angles, or slightly concave, sometimes with
medial spine.
Remarks. We now follow Clark (2018) in attributing this genus
to Bate rather than Spence Bate. The holotype of Scallasis
amboinae was examined (by GCBP) and additional specimens
were found in Papua New Guinea, enabling the species to be
well characterised. The genus is recognised by the combination
of a narrow rostrum, eyestalks with acute apices, a narrow
maxilliped 3, a simple or bifid spine on the merus of the major
cheliped, and exceptionally strong spiniform setae on the face
of the uropodal endopod and on the face of the telson.
Spinicallianassa gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:DCD1DFBE-9DFF481E-89CA-483A0AC6DDD8
Type species. Cheramus spinicauda Komai, Maenosono and
Fujita, 2014, by present designation.
Diagnosis. Hermaphrodite. Rostrum acute, anteriorly directed,
as long as eyestalks. Pleomere 1 tergite undivided or with weak
transverse step. Pleomere 6 with or without sublateral ventral
sharp ridge, flared posteriorly. Antennular peduncle articles 2
and 3 with single lateral row of 6–10 well-spaced long setae
along lower margin. Antennal scaphocerite simple, longer than
wide, acute. Mandibular molar calcified, swollen projection
without sharp edge, incisor with few teeth. Maxilliped 3 merus
wider at ischium-merus suture than long. Male major cheliped
merus with oblique spine one third to half-way along lower
margin; dactylus with dense setae along upper margin. Minor
cheliped two-thirds width of major cheliped, both flattened.
Pereopod 3 propodus rectangular, lower margin deeply convex,
leading to broadly rounded free proximal lobe. Uropodal
endopod ovoid, usually longer than wide, anterior margin
straight or slightly convex, posterodistal margin evenly convex,
with spiniform setae near anterior and distal margins, or on rib.
Uropodal exopod about 1.5–1.8 times as long as wide, distal
margin clearly differentiated from anterior margin, anterodistal
corner right-angled, posterodistal margin with row of 6–8 long
blade-like setae proximal to long setae on distal margin. Telson
about as wide as long, tapering from anterolateral lobe;
100
anterolateral lobe obsolete, undefined; transverse ridge
sometimes with spiniform setae; posterior margin slightly
concave, sometimes with medial spine.
Etymology. An alliteration of the name of the type species and
Callianassa, type genus of the family.
Remarks. Species of Spinicallianassa have dense setae on the
upper margin of the dactylus of the male major cheliped,
approaching that of Paratrypaea but differing from this genus
in many ways, notably having a small meral spine on the major
cheliped rather than a prominent toothed blade. The row of
short spiniform setae along the posterior margin of the telson
may be characteristic of this genus only. Spinicallianassa
shares with Aqaballianassa and Rayllianassa a dominating
calcified mandibular molar without a sharp edge and an incisor
without teeth.
Tastrypaea gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:B5299239-C36C4B78-8A7A-BB82710B67B5
Type species. Callianassa poorei Sakai, 1999, herein
designated.
Diagnosis. Rostrum obsolete or obtusely triangular, flat, not
reaching cornea. Pleomere 1 tergite undivided or with weak
transverse step. Maxilliped 3 merus wider at ischium-merus
suture than long. Male major cheliped merus with 1 or 2
small teeth just before midpoint; carpus and propodus
flattened, upper and lower margins carinate, blade-like,
submarginal mesial face especially of carpus deeply concave;
propodus distal margin with deep notch at base of fixed finger.
Minor cheliped two-thirds width of major cheliped, both
flattened. Pereopod 3 propodus rectangular, lower margin
deeply convex, leading to broadly rounded free proximal lobe.
Male pleopod 2 absent. Uropodal endopod asymmetrical, at
least as wide as long, distal margin truncate-convex, at right
angles to straight anterior margin, with facial distal transverse
row of short spiniform setae. Uropodal exopod distal margin
clearly differentiated from anterior margin, anterodistal corner
right-angled, posterodistal margin with row of 6–8 long bladelike setae proximal to long setae on distal margin. Telson
lateral margins convex.
Etymology. An alliteration of Tasmania, type locality of the
type species, and Trypaea, a genus of the family.
Remarks. We have no molecular data to support this genus but
Callianassa poorei displays several significant morphological
differences from its nearest neighbours to justify a new genus
(Sakai, 1999b). Tastrypaea resembles species of Paratrypaea
in having a well-developed transverse row of spiniform setae
near the anterodistal angle of the uropodal endopod but differs
in not having especially setose fingers on the major cheliped.
The species also differs from Paratrypaea and from
Arenallianassa arenosa, which has an obsolete row of
spiniform setae on the uropodal endopod, in that the chelipeds
are not extremely dissimilar, in not having a well-developed
blade on the major cheliped, the uropodal endopod being
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
asymmetrical, as wide as long, with its distal margin truncateconvex, at right angles to the straight anterior margin (rather
than ovoid and longer than wide) and the uropodal exopod
having a row of long blade-like setae proximal to long setae on
the distal margin (rather than having densely setose margins).
Tastrypaea differs from Filhollianassa, its sister taxon in
Robles et al. (in press), in not having a depressed anterior
carapace and chelipeds without prominent marginal crests.
Trypaea Dana, 1852
Trypaea Dana, 1852a: 14.—Poore, 2004: 184.—Sakai, 2011: 385–
387 (partim).
Callianassa (Trypaea).—Borradaile, 1903: 546 .—De Man, 1928:
27, 96 (partim).
Type species. Trypaea australiensis Dana, 1852, by monotypy.
Diagnosis. Rostrum obsolete or obtusely triangular, flat, not
reaching cornea. Pleomere 1 tergite undivided or with weak
transverse step. Eyestalk distal lobes obsolete, truncate.
Antennular peduncle length about 4 times the width of both
eyestalks, twice as long as antennal peduncle; articles 2 and
3 with 2 single similar rows of closely spaced setae laterally
and mesially along lower margin, extending on to flagellum.
Maxilliped 3 merus grossly expanded distomesially beyond
articulation with ischium, wider at ischium-merus suture than
long; crista dentata absent (or few proximal spines only). Male
major cheliped merus with prominent truncate hook armed with
serrations along lower margin, excavate laterally at base; carpus
and propodus flattened, upper and lower margins carinate,
blade-like, submarginal mesial face especially of carpus deeply
concave; propodus distal margin with deep notch at base of
fixed finger. Pereopod 3 propodus rectangular, lower margin
deeply convex, leading to broadly rounded free proximal lobe.
Male pleopod 2 absent. Uropodal endopod ovoid, usually longer
than wide, anterior margin straight or slightly convex,
posterodistal margin evenly convex, with facial distal transverse
row of short spiniform setae. Telson lateral margins convex;
slightly concave, sometimes with medial spine.
Remarks. Trypaea australiensis is immediately distinguished
from all other callianassids by the massive antennular
peduncles, far exceeding the antennal peduncles, bearing on
their lower margin a double row of adjacent long setae. The
merus of maxilliped 3 is more grossly expanded distally than
in any other genus. In all other genera that appear to have
setose antennae, the setae are scattered in broad bands and not
adjacent. Trypaea has only one species which is genetically
well separated from its sister taxa (Robles et al. 2019). These
are Arenallianassa from south-east Australia, Filhollianassa
from south-east Australia and New Zealand, and the more
widespread Indo-West Pacific Paratrypaea.
Sakai (1999a) synonymised Trypaea and eight other genera
with Callianassa, and later, Sakai (2005b) added a further two
genera to this synonymy. Sakai (2011) revived the genus and
synonymised five genera with Trypaea based on shared
features of the male pleopods 1 and 2, which are poorly
developed or absent in all callianassids. In doing so, he
included 53 species.
Callianassidae and related families
Callianopsidae Manning and Felder, 1991
Figure 12
Callianopsinae Manning and Felder, 1991: 787–789.—Schweitzer
Hopkins and Feldmann, 1997: 237.—Sakai, 2005b: 226–227.—Sakai,
2011: 477–478.—Sakai et al., 2015: 122–124.
Callianopsidae.—Sakai, 2011: 477.—Sakai et al., 2015: 121–
122.—Dworschak and Poore, 2018: 66–67.
Neocallianopsinae Sakai, 2011: 482.
Bathycalliacinae Sakai and Türkay, 1999a: 204.—Sakai, 2005b:
213–214.—Sakai, 2011: 347–348 syn. nov.
Bathycalliacidae Sakai, 2011: 347 syn. nov.
Vulcanocalliacinae Dworschak and Cunha, 2007: 37.—Sakai,
2011: 350 syn. nov.
Diagnosis. Rostrum flat, short, triangular, shorter than eyestalks,
or spike-like; median carina absent, or on rostrum only; gastric
carinae absent; cervical groove well defined; suture between
ocular lobe and end of linea thalassinica horizontal in lateral
view; anterior branchiostegal margin sinusoidal or semicircular;
anterior branchiostegal lobe simple, scarcely calcified, merging
smoothly with anterodorsal branchiostegal angle and anterolateral
margin of carapace; posterior margin of carapace without lateral
lobes; pleomere 1 without anterolateral lobes, weakly chitinised.
Eyestalks flattened, contiguous, with subdistal dorsal cornea.
Antennal scaphocerite usually elongate, rarely rudimentary.
Maxilla scaphognathite without long seta on posterior lobe
extending into branchial chamber. Maxilliped 1 epipod with
acute anterior lobe lying alongside exopod. Maxilliped 3 dactylus
dilating, truncate, with dense field of setae on distal margin.
Cheliped merus lower margin spinose, or with small proximal
tooth; palm oval in cross-section, barely crested above or below.
Pereopod 3 propodus rectangular or oval, up to twice as long as
wide, with proximal lobe on lower margin, without distal
spiniform setae on lateral face (often with 1 distal spiniform seta
on lower margin). Pereopod 5 minutely chelate. Female pleopod
2 rami narrower and with more reduced setation than pleopods
3–5; endopod flattened and 3–5 times as long as wide. Pleopods
3–5 with oblique peduncles meeting mesially, endopods
triangular, with straight mesial margin, exopods attached
laterally, proximally lobed, longer than and enclosing endopods;
appendices internae elongate, much longer than wide. Uropodal
exopod without elevated dorsal plate.
101
Remarks. Callianopsids differ from other callianassoids except
eucalliacids in having the dactylus of maxilliped 3 dilating,
truncate and with a dense field of setae on its distal margin. The
uropodal exopod lacks a dorsal plate, which is present in
eucalliacids. Members of the family have lateral ridges on the
eyestalks and maxilliped 3 ischium with a strong proximal lobe
on the lower margin. These features place the family far removed
from Callianassidae, with which it was compared by Sakai (2011).
Callianopsinae was treated as a subfamily of Ctenochelidae
by Manning and Felder (1991) and by Schweitzer Hopkins and
Feldmann (1997), as a subfamily of Gourretiidae by Sakai
(2005b) and as a full family by Sakai (2011).
Dworschak and Poore (2018) showed that Neocallianopsis is
a synonym of Callianopsis, and therefore, Neocallianopsinae
Sakai, 2011, is a synonym of Callianopsidae. Bathycalliacinae
and Vulcanocalliacinae, both monotypic and originally
subfamilies of Callianassidae, were included in Bathycalliacidae,
newly elevated to family rank by Sakai (2011). Dworschak and
Cunha (2007) noted that the two taxa shared epipods on
maxilliped 3 to pereopod 4 (found also in the only species of
Paracalliacidae), similar maxillipeds 3, propodi on pereopod 3,
telsons and uropods, and blindness. The morphological analysis
found the absence of an appendix masculina on male pleopod 2
to be a synapomorphy.
In his discussion of Bathycalliacinae, Sakai (2011)
contradicted Dworschak and Cunha’s (2007) assertion that the
holotype of Bathycalliax geomar is a male (as originally
stated). Subsequent examination by PCD has shown the
original observation was correct. Sakai (2011) erroneously
stated that the male pleopod 1 is absent and that Vulcanocalliax
arutyunovi possesses a dorsal plate on the uropodal exopod,
again not true.
The molecular analysis of Robles et al. (in press) found one
bathycalliacid representative, Vulcanocalliax arutyunovi, to
be a sister taxon to one example of Callianopsidae, Callianopsis
goniophthalma, a result consistent with the morphological
analysis that found Bathycalliax to be closely allied. On the
basis of this evidence, we synonymise Bathycalliacidae with
Callianopsidae and move Vulcanocalliacinae to Callianopsidae
without recognising subfamilies.
Figure 12. Diagnostic characters for genera of Callianopsidae. Carapace: a, Bathycalliax. Pleomere 6: c, Callianopsis. Maxilliped 3: c, Bathycalliax.
102
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Key to genera of Callianopsidae
1.
Pleomere 6 with prominent lateral projections (fig. 12b);
carapace with longitudinal carina running from rostrum;
epipods absent
Callianopsis
–
Pleomere 6 without prominent lateral projections;
carapace without longitudinal carina; epipods present on
maxilliped 3 to pereopod 4
2
2.
Two cardiac sulci present (fig. 12a); maxilliped 3 with
rudimentary exopod (fig. 12c)
Bathycalliax (1 species, B. geomar)
–
Cardiac sulci absent; maxilliped 3 exopod absent
Vulcanocalliax (1 species, V. arutyunovi)
Bathycalliax Sakai and Türkay, 1999
Bathycalliax Sakai and Türkay, 1999: 204.—Sakai, 2005b: 214.—
Sakai, 2011: 349.
Type species. Bathycalliax geomar Sakai and Türkay, 1999, by
original designation and monotypy.
Diagnosis. Rostrum obsolete or obtusely triangular, flat, not
reaching cornea; carapace without median carina; cardiac
sulci present. Pleomere 6 without lateral projections.
Maxilliped 3 exopod present. Male major cheliped merus lower
margin straight, with proximal row of denticles.
Remarks. Bathycalliax geomar is the sole species, placed by its
authors in its own callianassid subfamily, Bathycalliacinae.
Dworschak and Cunha (2007) confirmed that the holotype
male pleopod 1 has a unique flagellate article 2 but that the
smaller male paratype lacks pleopod 1. Sakai (2011)
contradicted his earlier observations, saying that the male
pleopod 1 is absent.
Callianopsis de Saint Laurent, 1973
Callianopsis de Saint Laurent, 1973: 515.—Schweitzer Hopkins
and Feldmann, 1997: 237–238.—Sakai, 2005b: 226.—Sakai, 2011:
478–479.—Dworschak and Poore, 2018: 67.
Pleurocalliax Sakai, 2011: 480–481 (type species, Callianassa
caecigena Alcock and Anderson, 1894, by original designation and
monotypy).
Neocallianopsis Sakai, 2011: 482 (type species, Callianopsis
anovalis Lin, Komai and Chan, 2007, by original designation and
monotypy).
Phaetoncalliax Sakai, Türkay, Beuck and Freiwald, 2015: 124–
128 (type species, Phaetoncalliax mauritana Sakai, Türkay, Beuck
and Freiwald, 2015, by original designation and monotypy).
Type species. Callianassa goniophthalma Rathbun, 1902, by
original designation and monotypy.
Diagnosis. Rostrum acute, produced, with dorsal carina
leading to gastric region; carapace with median carina strong
on rostrum, weaker more posteriorly; cardiac sulci absent.
Pleomere 6 with lateral projections. Maxilliped 3 exopod
absent. Male major cheliped merus with straight or weakly
convex blade on lower margin bearing a short proximal curved
spine and 1 or more distal spines.
Remarks. Callianopsis is recognised by the combination of a
dorsally carinate rostrum and lateral projections on pleomere 6.
Callianopsis was placed in its own subfamily of the
Ctenochelidae by Manning and Felder (1991). The subfamily
was included as a member of Gourretiidae by Sakai (2005b)
and at family rank by Sakai (2011). Sakai (2005b: 227) treated
Dawsonius, a gourretiid genus, as a junior synonym of
Callianopsis but in a note added in proof to the same paper (p.
245) recognised it as a separate genus. Dworschak and Poore
(2018) synonymised Neocallianopsis Sakai, 2011,
Pleurocalliax Sakai, 2011, and Phaetoncalliax Sakai, Türkay,
Beuck and Freiwald, 2015, with Callianopsis. All had been
inadequately diagnosed or based on errors.
Vulcanocalliax Dworschak and Cunha, 2007
Vulcanocalliax Dworschak and Cunha, 2007: 37.—Sakai, 2011:
350–351.
Type species. Vulcanocalliax arutyunovi Dworschak and
Cunha, 2007, by original designation and monotypy.
Diagnosis. Rostrum obsolete or obtusely triangular, flat, not
reaching cornea; carapace without median carina; cardiac sulci
absent. Pleomere 6 without lateral projections. Maxilliped 3
exopod absent. Male major cheliped merus with 1–2 small
proximal teeth on lower margin.
Remarks. The sole species, Vulcanocalliax arutyunovi
Dworschak and Cunha, 2007, was placed in its own subfamily
of Callianassidae close to Bathycalliacinae by its authors but
the subfamily was transferred to Bathycalliacidae by Sakai
(2011). Contrary to Sakai’s (2011) assertion, the only species
does not have a dorsal plate on the uropodal exopod. It differs
from Bathycalliax geomar in the absence of cardiac sulci and
possession of a maxilliped 3 exopod.
Callichiridae Manning and Felder, 1991
Figures 13–15
Callichirinae Manning and Felder, 1991: 775–776.—Hyžný and
Müller, 2012: 968–969.
Calliapaguropinae Sakai, 1999a: 7.—Sakai, 2005: 205.—Sakai,
2011: 491–492.
Callichiridae.—Sakai, 2011: 418.—Sakai et al., 2014: 490 (lapsus,
Callichirinae on p. 500).—Dworschak, 2018: 21.
Diagnosis. Rostrum flat, short, triangular, shorter than eyestalks,
or spike-like, longer than wide; median carina absent; gastric
carinae absent; cervical groove well defined; suture between
ocular lobe and end of linea thalassinica oblique in lateral view;
anterior branchiostegal lobe sclerotised, well produced
anteriorly beyond junction with oblique branchiostegal ridge
with which it articulates by means of a virtual condyle;
posterior margin of carapace without lateral lobes; pleomere 1
without anterolateral lobes, weakly chitinised. Eyestalks
flattened, contiguous, with subdistal dorsal cornea, or cylindrical,
with terminal subspherical cornea (Calliapagurops only).
Antennal scaphocerite rudimentary. Maxilla scaphognathite
without long seta on posterior lobe extending into branchial
chamber. Maxilliped 1 epipod with acute anterior lobe lying
alongside exopod. Maxilliped 3 dactylus slender, digitiform,
Callianassidae and related families
103
Figure 13. Diagnostic characters for genera of Callichiridae. Rostrum, eyestalks, antennules, antennae: a, Calliapagurops; b, Lepidophthalmus;
c, Mocallichirus; d, Mucrollichirus; e, Karumballichirus; f, Audacallichirus; g, h, Glypturus; i, Corallianassa. Maxilliped 3: j, Mocallichirus;
k, Glypturoides; l, Karumballichirus; m, Thailandcallichirus; n, Calliapagurops. Male major pereopod 1: o, Glypturus; p, Thailandcallichirus;
q, Corallianassa; r, Mucrollichirus; s, Karumballichirus; t, Glypturoides; u, Laticallichirus. Minor pereopod 1: v, Balsscallichirus; w,
Laticallichirus.
Original illustrations: c, Mocallichirus mocambiquensis, UF 13986; d, r, Mucrollichirus mucronatus, MNHN-IU-2013-2777.
104
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Figure 14. Diagnostic characters for genera of Callichiridae. Pereopod 3: a, Audacallichirus; b, Karumballichirus; c, Neocallichirus; d,
Lepidophthalmus; e, Mucrollichirus. Pleon, telson, uropods: f, Callichirus; g, Grynaminna; h, Michaelcallianassa. Telson, uropod: i, Audacallichirus;
j, Balsscallichirus; k, Glypturoides; l, Karumballichirus; m, Lepidophthalmus; n, Mocallichirus; o, Kraussillichirus; p, Neocallichirus.
Original illustrations: n, Mocallichirus mocambiquensis, UF 13986; e, Mucrollichirus mucronatus, MNHN-IU-2013-2777.
Callianassidae and related families
105
Figure 15. Diagnostic characters for genera of Callichiridae. Male pleopod 1: a, Mocallichirus mocambiquensis; b, Balsscallichirus balssi; c, B.
pixii; d, Corallianassa martensi; e, C. xutha; f, Glypturus armatus; g, Lepidophthalmus eiseni; h, L. madagassus; i, Neocallichirus raymanningi;
j, N. vigilax. Male pleopod 2: k, Glypturus armatus; l, Corallianassa coutierei; m, Grynaminna tamakii; n, Michaelcallianassa indica. Female
pleopod 2: o, Balsscallichirus balssi; p, Laticallichirus grandis.
Original illustrations: a, Mocallichirus mocambiquensis, UF 13986; j, Neocallichirus vigilax MNHN-IU-2015-7072.
106
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
with setae irregularly spaced along all margins. Cheliped merus
lower margin smooth or spinose; major cheliped with distinctively
flattened palm, sometimes with strong crest above and below.
Pereopod 3 propodus broad, with proximal lobe on lower margin,
without distal spiniform setae on lateral face (often with 1 distal
spiniform seta on lower margin). Pereopod 5 minutely chelate or
subchelate. Female pleopod 2 rami narrower and with more
reduced setation than pleopods 3–5; endopod flattened and 2–5
times as long as wide. Pleopods 3–5 with oblique peduncles
meeting mesially, endopods triangular, with straight mesial
margin, exopods attached laterally, proximally lobed, longer
than and enclosing endopods; appendices internae reduced and
almost embedded in mesial margin of endopod. Uropodal
exopod with elevated dorsal plate.
Remarks. In callichirids, the endopod of the female pleopod 2
is flattened, variously broadened and more or less parallelsided, unlike in callianassids where it is narrow. The anterior
branchiostegal margin is interrupted by an oblique sclerotised
ridge, sometimes almost articulating on the margin, unlike in
callianassids where this margin is uninterrupted. Pleopod 1 is
always present in the male, often substantial with a distolateral
sharp curved lobe (fig. 15d–j) or variously reduced (fig. 15a–c),
whereas in callianassids it is small and simple or absent. The
epipod of maxilliped 1 takes the plesiomorphic form with a
triangular lobe overlapping the exopod; this lobe does not exist
in callianassids.
The use of the name as a family, rather than subfamily, has
not been previously argued.
Key to genera of Callichiridae
1.
–
2.
–
3.
–
Eyestalk cylindrical, separate, with terminal spherical
cornea, without terminal lobe (fig. 13a); maxilliped 3
merus with 3 spines along distal margin (fig. 13n)
Calliapagurops
Eyestalk flattened, contiguous, about twice as long as wide,
with cornea subterminal and more or less elevated, with
terminal lobe (figs 13b–g); maxilliped 3 merus without
spines along distal margin (figs 13j–m)
2
Rostrum spine-like, anteriorly or upwardly directed;
anterolateral margins of carapace with conical curved spine,
sometimes weakly calcified at base (fig. 13g)
3
Rostrum obsolete or triangular, flat even if sharp;
anterolateral margins of carapace not produced (figs 13b–f)
or rarely sharp (in Neocallichirus vigilax, Lepidophthalmus
tridentatus)
4
Major chelipeds with 3 spines on upper inner margin of
merus and propodus, and 4–6 on lower margin of carpus
(fig. 13o); rostrum with midventral ridge (fig. 13h);
scaphocerite small, discoid; male pleopod 2 with partially
articulating appendix masculina (fig. 15k)
Glypturus
Major chelipeds without spines on upper margin of merus
and propodus or lower margin of carpus (fig. 13q); rostrum
without midventral ridge (fig. 13i); scaphocerite absent;
male pleopod 2 with appendix masculina fused to
endopod (fig. 15l)
Corallianassa
4.
Maxilliped 3 exopod present, minute (figs 13k, l)
5
–
Maxilliped 3 exopod absent
8
5.
Antennular peduncle much longer than antennal peduncle
(fig. 13b); telson posterior margin with medial lobe
(fig. 14m, o)
6
–
Antennular peduncle shorter than antennal peduncle (fig.
13e); telson posterior margin convex or concave
7
6.
Male major cheliped merus with sinuous lower margin,
widest proximally and excavate beyond, or with proximal
tubercle bearing spine(s) with or without more distal teeth
along lower margin; pereopod 3 propodus lower margin
with obtuse notch between distal lobe and heel (fig. 14d);
Lepidophthalmus
–
Male major cheliped merus with denticulate blade, more
prominent over proximal half of lower margin; pereopod
3 propodus lower margin simply concave
Kraussillichirus
7.
Male major cheliped fingers longer than square palm (fig.
13s); telson widest proximally or at midpoint, with convex
posterior margin (fig. 14l)
Karumballichirus
–
Male major cheliped fingers shorter than rectangular
palm (fig. 13t); telson widest at midpoint, posterior margin
concave (fig. 14k)
Glypturoides
8.
Antennular peduncle shorter or as long as antennal
peduncle (figs 13c, d, f)
9
–
Antennular peduncle significantly longer than antennal
peduncle (fig. 13b)
13
9.
Major cheliped merus lower margin with simple proximal
hook, dactylus massive (fig. 13p); maxilliped 3 dactylus
with expanded setose apex (fig. 13m)
Thailandcallichirus
–
Major cheliped merus lower margin denticulate or with
toothed blade, dactylus not significantly broadened (figs
13q, r); maxilliped 3 dactylus tapering
10
10. Rostrum spine-like, thickened; cornea subdistal,
hemispherical, elevated, with minute mesiodistal plate;
female pleopod 2 endopod 3 times as long as wide
Mucrollichirus
–
Rostrum flat, triangular; reaching cornea mid-distal,
scarcely elevated, with obvious mesiodistal plate; female
pleopod 2 endopod 4 times as long as wide
11
11. Telson parallel-sided over most of length, with obtuse
angled apex, domed dorsally (fig. 14n); maxilliped 3
propodus longer than wide, free distal margin oblique;
male pleopod 1 consisting of 1 slender article (fig. 15a)
Mocallichirus
–
Telson widest proximally or at midpoint; maxilliped 3
propodus as long as wide, free distal margin nearly
transverse; male pleopod 1 consisting of 2 articles
12
107
Callianassidae and related families
12. Telson converging over most of length from subproximal
width to about half maximum width, with rounded
posterolateral corners, narrow posterior margin (fig. 14p);
pereopod 3 propodus with proximal lobe on lower margin
slightly overlapping carpus (fig. 14c)
Neocallichirus
–
Telson converging from subproximal width to about
three-quarters maximum width, with strongly convex
lateral margins and rounded distolateral corners, wide
sinuous posterior margin (fig. 14i); pereopod 3 propodus
with prominent proximal lobe on lower margin appearing
to overlap carpus (fig. 14a)
Audacallichirus
13. Uropodal endopod strap-like, curved, longer than exopod;
pleomeres 3–5 with dorsal pattern of symmetrical
grooves; telson almost semicircular, thickened, with
constriction on proximal region, with posterolateral
swellings (fig. 14f)
Callichirus
–
Uropodal endopod ovate; pleomeres 3–5 without pattern
of symmetrical grooves; telson convex-sided, widest
proximally or near midpoint (figs 14g, h)
14
14. Uropodal exopod about as wide as length of anterior
margin; distal margin of telson convex, upper surface
with long setae only (fig. 14g)
Grynaminna
–
Uropodal exopod much wider than length of anterior
margin; distal margin of telson concave, upper surface
usually with transverse row of short spiniform setae (figs
14h, j)
15
15. Minor cheliped palm about one third as wide as major
cheliped palm, carpus longer than wide, fingers straight,
shorter than palm (fig. 13v)
Balsscallichirus
–
Minor cheliped palm at least half as wide as major cheliped
palm, carpus as long as wide, fingers curved, pincer-like,
much longer than palm (figs 13u, w)
16
16. Pleomeres 3–5 with anteriorly converging longitudinal
grooves and oblique–transverse rows of long setae (fig.
14h); male major cheliped merus with lower margin
almost straight; female pleopod 2 uniramous (fig. 15o)
Michaelcallianassa
–
Pleomeres 3–5 with prominent lateral tufts of setae; male
major cheliped merus with convex denticulate blade (fig.
13u); female pleopod 2 biramous (fig. 15p)
Laticallichirus
Implicit attributes. Unless indicated otherwise, the following
attributes are implicit throughout the generic diagnoses.
Rostrum without midventral ridge; anterolateral spines absent,
anterolateral angle obsolete. Pleomere 1 tergite and oblique
narrow lateral pleura fused into a single sclerifed unit, or pleura
separated by triangular uncalcified region from tergite;
rectangular median lateral sternite plates (pleopod attached)
fused to pleuron and median sternite. Pleomeres 3–5 without
symmetrical pattern of deep grooves, with narrow transverse
rows of setae, pleomere 6 without longitudinal grooves on
anterior section, with slight lateral incision. Eyestalk flattened
and contiguous. Antennular peduncle length less than 3 times
as width of both eyestalks, shorter than antennal peduncle.
Antennal scaphocerite vestigial. Maxilliped 3 merus without
distal spine on mesial margin; crista dentata a row of numerous
denticles; propodus longer than wide, lower margin convex;
dactylus tapering, with scattered setae over upper margin,
dense brush of short setae distally on lower margin; exopod
absent. Male major cheliped carpus shorter than palm; without
spines on upper margin of merus and propodus or lower margin
of carpus; male major cheliped dactylus typically tapering.
Pereopod 3 propodus lower margin straight or weakly concave.
Male pleopod 2 biramous. Female pleopod 2 biramous;
endopod about 4 times as long as wide. Telson smooth dorsally.
Audacallichirus gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:F54AD76A-81724D0F-ADCB-2174D5A3FC6B
Type species. Callianassa audax De Man, 1911, by present
designation.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique
branchiostegal ridge with which it articulates by means of a
virtual condyle. Rostrum obsolete or obtusely triangular, flat,
not reaching cornea. Pleomere 1 tergite undivided or with weak
transverse ridge. Maxilliped 3 ischium and merus narrow,
more than twice as long as wide at their articulation; propodus
about as wide as long, free distal margin transverse or nearly
so. Male major cheliped merus lower margin convex or blade
like, especially proximally, with small denticles. Male minor
cheliped much narrower than major, carpus longer than palm,
fingers shorter than palm. Pereopod 3 propodus subpentagonal,
with strong broadly rounded proximal lobe on lower margin,
lower margin straight-concave. Pereopod 4 subchelate. Male
pleopod 1 consisting of 2 articles, article 2 distally expanded,
with shallow apical notch between 2 rounded setose distal
lobes. Male pleopod 2 appendix interna absent or reduced to
obsolete distomesial lobe. Pleopods 3–5 appendices internae
barely emerging from endopod margin. Uropodal endopod
with convex anterior margin, acute-rounded apex, straightslightly curved posterior margin, longer than wide. Telson
convex-sided, widest near midpoint, posterior margin trilobed,
excavate each side of medial lobe.
Etymology. An alliteration of the name of the type species and
Callichirus, type genus of the family.
Remarks. Audacallichirus audax differs from members of
Neocallichirus, the genus in which it was placed by Sakai
(1999a, 2011), in having a telson with strongly convex lateral
margins, uropodal endopod tapering, the propodus of pereopod
3 usually with a strong proximal lobe on the lower margin, and
the male pleopod 1 with a broad second article with an wide
apical notch (Ngoc-Ho, 2014; Rao and Kartha, 1967; Tirmizi,
1967). The molecular analysis of Robles et al. (in press) found
the species similar to Neocallichirus mirim, a species
sometimes included in Sergio. The two species share a similar
male pleopod 1, uropodal endopod, telson and cheliped, but the
propodus of pereopod 3 is less expanded in N. mirim.
108
Balsscallichirus Sakai, 2011
Balsscallichirus Sakai, 2011: 414–415.—Hyžný, 2016: 43–46.
Tirmizicallichirus Sakai, 2011: 474–475 (type species,
Callianassa (Callichirus) masoomi Tirmizi, 1970, by original
designation and monotypy).
Barnardcallichirus Sakai, 2011: 416–417 (type species,
Callichirus tenuimanus de Saint Laurent and Le Loeuff, 1979, by
original designation).
Capecalliax Sakai, 2011: 345 (type species, Callianassa pixii
Kensley, 1976, by original designation and monotypy) syn. nov.
Forestcallichirus Sakai, 2011: 426–427 (type species, Callichirus
foresti Le Loeuff and Intès, 1974, by original designation and
monotypy) syn. nov.
Type species. Callianassa (Callichirus) balssi Monod, 1933, by
original designation and monotypy.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique
branchiostegal ridge with which it articulates by means of a
virtual condyle. Rostrum obsolete or obtusely triangular, flat,
not reaching cornea. Pleomere 1 tergite divided into 2 sections
by unsclerified band; tergite weakly sclerotised, if so only
posteriorly; pleura oblique thin rods independent of tergite
dorsally, separated by extensive flexible region; sternite a thin
transverse plate. Antennular peduncle longer than antennal
peduncle. Maxilliped 3 ischium and merus narrow, at least 3
times as long as wide at their articulation; crista dentata
absent (or few proximal spines only); propodus longer than
wide, lower margin convex or about as wide as long, free distal
margin clearly oblique or free distal margin transverse or
nearly so. Male major cheliped merus with lower margin
almost straight or with denticulate blade, more prominent over
proximal half of lower margin or with prominent proximal
denticulate blade-like tooth on lower margin. Male minor
cheliped much narrower than major, carpus longer than palm,
fingers shorter than palm. Pereopod 3 propodus subpentagonal,
with strong broadly rounded proximal lobe on lower margin,
lower margin straight-concave, lower margin straight or weakly
concave or weakly convex. Pereopod 4 subchelate. Male
pleopod 1 of 2 articles, article 2 longer than wide, with or
without slight apical notch. Male pleopod 2 biramous or
uniramous; appendix interna absent. Pleopods 3–5 appendices
internae barely emerging from endopod margin. Uropodal
endopod with convex anterior margin, acute-rounded apex,
straight-slightly curved posterior margin, longer than wide.
Telson converging to about half basal width from near anterior
width over most of length to rounded posterolateral corners or
with convex lateral margins and rounded posterolateral corners,
posterior margin trilobed, excavate each side of medial lobe or
slightly excavate or obscurely excavate between rounded
posterolateral corners, with transverse row of robust setae.
Remarks. Balsscallichirus is best recognised by its very narrow
maxilliped 3. Sakai (2011) distinguished Balsscallichirus (B.
balssi, B. guineensis), Tirmizicallichirus (T. masoomi),
Barnardcallichirus (B. tenuimanus, B. gilchristi) and
Forestcallichirus (F. foresti) from Podocallichirus (P.
madagassus) on the basis of male pleopods 1 and 2. The
distinction is valid—Podocallichirus is considered here a
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
junior synonym of Lepidophthalmus. He did not compare the
new genera with each other or with others. Hyžný (2016)
synonymised Tirmizicallichirus and Barnardcallichirus with
Balsscallichirus and listed five Recent species and four fossils.
Kensley (1976) recognised that his new species Callianassa
pixii belonged in the “subgenus Callichirus”, likening it to
Callianassa guineensis. He was supported by de Saint Laurent
and Le Loeuff (1979), who grouped this with four of the
species listed above, B. balssi, B. foresti, B. tenuimanus and B.
guineensis (as species of Callianassa), on the similarity of
triangular rostrum, rounded posterior border of the telson,
pediform maxilliped 3 and reduced male pleopod 2. Sakai
(2011) erected another monotypic genus, Capecalliax for C.
pixii, this time in the family Anacalliacidae, to which it clearly
does not belong. Capecalliax and Forestcallichirus, for C.
foresti listed in this group correctly characterised by de Saint
Laurent and Le Loeuff (1979) are also synonyms of
Balsscallichirus. The exceptionally narrow maxilliped 3 was
treated as a symplesiomorphy by the morphological analysis
of Robles et al. (in press).
The seven species are known only from around Africa, as
far north as Mauritania in the west and Pakistan in the east.
Calliapagurops de Saint Laurent, 1973
Calliapagurops de Saint Laurent, 1973: 515.—Sakai, 1999a: 8.—
Ngoc-Ho, 2002: 540–541.—Sakai, 2005b: 207.—Sakai, 2011: 492.
Type species. Calliapagurops charcoti de Saint Laurent, 1973,
by original designation and monotypy.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique branchiostegal
ridge with which it articulates by means of a virtual condyle.
Rostrum spine-like; anterolateral spines prominent. Pleomere 1
tergite undivided or with weak transverse ridge. Eyestalk
cylindrical, with subspherical terminal pigmented cornea.
Antennal scaphocerite elongate, longer than wide, with free
acute tip. Maxilliped 3 ischium and merus less than twice as
long as wide at their articulation; merus with 2 or more distal
spines on distal free margin; crista dentata of few separate
spines proximally and toothed ridge distally overlapping
proximal margin of merus; propodus about as wide as long, free
distal margin clearly oblique. Male major cheliped merus with
row of sharp oblique spines, usually 2 proximally and 1 or more
along length of lower margin. Male minor cheliped half or more
as wide as major, carpus as long as or shorter than palm, fingers
as long as or longer than palm. Pereopod 3 propodus oval, lower
margin convex, not proximally lobed (slightly produced
distally), lower margin weakly convex. Pereopod 4 propodus
minutely chelate, fixed finger as long as dactylus. Male pleopod
1 consisting of 2 articles, article 2 longer than wide, with or
without slight apical notch. Male pleopod 2 appendix interna
free, articulating, distal on mesial endopodal margin. Female
pleopod 2 endopod 2–3 times as long as wide. Pleopods 3–5
appendices internae barely emerging from endopodal margin.
Uropodal endopod with convex anterior margin, acute-rounded
apex, straight-slightly curved posterior margin, longer than
wide. Telson convex-sided, widest near midpoint, posterior
109
Callianassidae and related families
margin concave between rounded posterolateral corners, with
transverse row of robust setae.
Remarks. The two species of Calliapagurops differ from all
other callianassoids in having cylindrical eyestalks with
terminal cornea. Ngoc-Ho (2002) pointed out numerous
similarities between Calliapagurops and Corallianassa,
similarities borne out in the morphological analysis (Robles et
al., in press) but not by the molecular data, where the genus is
closer to Grynaminna. Sakai (2011) argued that Calliapagurops
and its subfamily Calliapaguropinae belong in Eucalliacidae
because of differences from Callichirus and Callichirinae,
namely the absence of the dorsal oval, the unique eyestalks and
a small appendix interna on the male pleopod 2. He listed no
similarities to Eucalliacidae – there are few.
Callichirus Stimpson, 1866
Callichirus Stimpson, 1866: 47.—de Saint Laurent and Le Loeuff,
1979: 55–56.—Manning and Felder, 1991: 775–776.—Sakai, 2011: 418
(abbreviated synonymy).
Callianassa (Callichirus).—Borradaile, 1903: 546–547.—De
Man, 1928: 28, 96 (partim).
Type species. Callianassa major Say, 1818, by original
designation and monotypy.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique branchiostegal
ridge with which it articulates by means of a virtual condyle.
Rostrum obsolete or obtusely triangular, flat, not reaching
cornea. Pleomere 1 tergite divided into 2 sections by unsclerified
band; tergite weakly sclerotised, if so only posteriorly; pleura
oblique thin rods independent of tergite dorsally, separated by
extensive flexible region; sternite a thin transverse plate.
Pleomeres 3–5 with symmetrical pattern of deep curved
transverse and longitudinal grooves plus paired tufts of dense
setae, pleomere 6 with pair of deep longitudinal grooves on
wide swollen anterior section, defined by lateral concavity.
Antennular peduncle length longer than antennal peduncle.
Maxilliped 3 ischium and merus less than twice as long as wide
at their articulation; crista dentata absent (or few proximal
spines only); propodus about as wide as long, free distal margin
clearly oblique or free distal margin transverse or nearly so.
Male major cheliped merus with denticulate blade, more
prominent over proximal half of lower margin or with prominent
proximal denticulate blade-like tooth on lower margin; major
cheliped carpus much longer than palm, especially in male.
Male minor cheliped much narrower than major, carpus longer
than palm, fingers shorter than palm. Pereopod 3 propodus
subpentagonal, with strong broadly rounded proximal lobe on
lower margin, lower margin straight-concave. Pereopod 4
subchelate. Male pleopod 1 consisting of 2 articles, article 2
longer than wide, with or without slight apical notch. Male
pleopod 2 appendix interna absent or reduced to obsolete
distomesial lobe. Pleopods 3–5 appendices internae barely
emerging from endopod margin. Uropodal endopod strap-like,
posterior margin concave. Telson thickened, with constriction
on anterior region, with posterolateral swellings, posterior
margin with medial notch at end of longitudinal groove.
Remarks. Species of Callichirus are recognised by the
symmetrical pattern of deep curved transverse and longitudinal
grooves plus paired tufts of dense setae on pleomeres 3–5, deep
longitudinal grooves on the wide anterior section of pleomere
6, the long antennular peduncle, the strap-like curved uropodal
endopods and the thickened telson, with a proximal constriction.
Callichirus kraussi is here removed to its own genus,
Kraussillichirus, on genetic evidence (Robles et al., in press)
and uropodal endopods more typical of the family. Species of
Callichirus may be functional hermaphrodites with some or all
females having male gonopores (Souza et al., 2017, 2018)
Corallianassa Manning, 1987
Corallianassa Manning, 1987: 392–394.—Poore, 2004: 184.—
Ngoc-Ho, 2005: 71.—Komai et al., 2015: 54–55 (synonymy).
Corallichirus Manning, 1992: 571–574.—Sakai, 2011: 422–423
(type species: Corallianassa xutha Manning, 1988, by original
designation).
Type species. Callianassa longiventris A. Milne-Edwards,
1870, by original designation.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique branchiostegal
ridge with which it articulates by means of a virtual condyle.
Rostrum spine-like; anterolateral spines prominent. Pleomere
1 tergite undivided or with weak transverse ridge. Maxilliped 3
ischium and merus linear, around twice as long as wide at their
articulation; crista dentata of few separate spines proximally
and toothed ridge distally overlapping proximal margin of
merus; propodus about as wide as long, free distal margin
transverse or nearly so. Male major cheliped merus with row of
sharp oblique spines, usually 2 proximally and 1 or more along
length of lower margin. Male minor cheliped half or more as
wide as major, carpus as long as or shorter than palm, fingers as
long as or longer than palm. Pereopod 3 propodus subpentagonal,
with strong broadly rounded proximal lobe on lower margin,
lower margin straight-concave, lower margin straight or weakly
concave or weakly convex. Pereopod 4 subchelate. Male
pleopod 1 consisting of 2 articles, article 2 ovoid, with shallow
apical notch between rounded subdistal lobe and curved acute
apex. Male pleopod 2 appendix interna free, articulating, distal
on mesial endopod margin. Pleopods 3–5 appendices internae
longer than wide, clearly emerging from margin of endopod.
Uropodal endopod with convex anterior margin, acute-rounded
apex, straight-slightly curved posterior margin, longer than
wide. Telson converging to about half basal width from near
anterior width over most of length to rounded posterolateral
corners, posterior margin straight or with slight medial lobe.
Remarks. Corallianassa is best recognised by the prominent
anterolateral spines, the hemispherical, elevated, distal cornea
(distinguishing it from Calliapagurops) and the absence of
marginal teeth on the chelipeds (distinguishing it from Glypturus;
see Komai et al., 2015). Komai et al. (2015) reviewed the complex
taxonomic history of Corallianassa, Corallichirus and
Glypturus, and the unjustified confusion introduced by Sakai’s
(2011) revival of Corallichirus, regarded as a synonym of
Corallianassa since this was first proposed by Ngoc-Ho (2005).
110
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Glypturoides Sakai, 2011
Glypturoides Sakai, 2011: 428.
Type species. Callianassa trilobata Biffar, 1970, by original
designation and monotypy.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique
branchiostegal ridge with which it articulates by means of a
virtual condyle. Rostrum obsolete or obtusely triangular, flat,
not reaching cornea. Pleomere 1 tergite undivided or with
weak transverse ridge. Maxilliped 3 ischium and merus less
than twice as long as wide at their articulation; propodus
about as wide as long, free distal margin transverse or
nearly so; exopod present. Male major cheliped merus lower
margin convex or blade like, especially proximally, with
small denticles. Male minor cheliped much narrower than
major, carpus longer than palm, fingers shorter than palm.
Pereopod 3 propodus subpentagonal, with strong broadly
rounded proximal lobe on lower margin, lower margin
straight-concave or weakly convex. Pereopod 4 subchelate.
Male pleopod 1 consisting of 2 articles, article 2 ovoid, with
shallow apical notch between rounded subdistal lobe and
curved acute apex. Male pleopod 2 appendix interna absent or
reduced to obsolete distomesial lobe. Pleopods 3–5 appendices
internae barely emerging from endopod margin. Uropodal
endopod with convex anterior margin, acute-rounded apex,
straight-slightly curved posterior margin, longer than wide.
Telson with convex lateral margins and rounded posterolateral
corners, posterior margin concave between rounded
posterolateral corners.
Remarks. Glypturoides trilobata, the only species, is like
species of Neocallichirus but with an exopod on maxilliped 3.
In addition, the maxilliped 3 propodus has the free distal
margin more oblique than in typical Neocallichirus, and the
telson is short with convex lateral margins and concave
posterior margin.
Glypturus Stimpson, 1866
Glypturus Stimpson, 1866: 46.—Manning, 1987: 390.—Manning
and Felder, 1991: 778.—Sakai, 1999a: 72.—Sakai, 2005b: 130–132.—
Sakai, 2011: 429–430 (partim).—Hyžný and Müller, 2012: 969–971.—
Hyžný et al., 2013: 133.—Klompmaker et al., 2015: 11.—Komai et al.,
2015: 53–54 (complete synonymy).
Callianassa (Glypturus).—Borradaile, 1903: 548.
Type species. Glypturus acanthochirus Stimpson, 1866: 46, by
original designation.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique
branchiostegal ridge with which it articulates by means of a
virtual condyle. Rostrum spine-like, with midventral ridge;
anterolateral spines prominent. Pleomere 1 tergite fused,
divided into 2 sections by transverse groove or shoulder.
Antennal scaphocerite small, discoid. Maxilliped 3 ischium
and merus narrow, more than twice as long as wide at their
articulation; crista dentata of few separate spines proximally
and toothed ridge distally overlapping proximal margin of
merus; propodus about as wide as long, free distal margin
transverse or nearly so. Major cheliped merus with row of
sharp oblique spines, usually 2 proximally and 1 or more
along length of lower margin; with 3 spines on upper inner
margin of merus and propodus, and 4–6 on lower margin
of carpus. Minor cheliped half or more as wide as major,
carpus as long as or shorter than palm, fingers as long as or
longer than palm. Pereopod 3 propodus subpentagonal, with
strong broadly rounded proximal lobe on lower margin, lower
margin straight-concave. Pereopod 4 subchelate. Male
pleopod 1 consisting of 2 articles, article 2 ovoid, with shallow
apical notch between rounded subdistal lobe and curved acute
apex. Male pleopod 2 appendix interna free, articulating,
distal on mesial endopod margin. Pleopods 3–5 appendices
internae longer than wide, clearly emerging from margin of
endopod. Uropodal endopod with convex anterior margin,
acute-rounded apex, straight-slightly curved posterior
margin, longer than wide. Telson strongly diverging to
beyond midpoint and semicircular distally, posterior margin
broadly semicircular.
Remarks. Glypturus is best recognised by the prominent
anterolateral spines and the presence of marginal teeth on the
chelipeds (Komai et al., 2015). Komai et al. (2015) clarified
the confusion around Glypturus introduced by Sakai’s
(2011) revision.
Grynaminna Poore, 2000
Grynaminna Poore, 2000: 150–151.—Sakai, 2011: 438–439.—
Hyžný and Karasawa, 2012: 63–65.
Type species. Grynaminna tamakii Poore, 2000, by original
designation and monotypy.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique
branchiostegal ridge with which it articulates by means of a
virtual condyle. Rostrum obsolete or obtusely triangular, flat,
not reaching cornea. Pleomere 1 tergite undivided or with
weak transverse ridge. Antennular peduncle length more than
3 times the width of both eyestalks, longer than antennal
peduncle. Maxilliped 3 ischium and merus narrow, more than
twice as long as wide at their articulation; crista dentata
consisting of a row of numerous denticles; propodus about as
wide as long, free distal margin transverse or nearly so. Male
major cheliped merus with convex tuberculate blade on
most of lower margin. Male minor cheliped much narrower
than major, carpus longer than palm, fingers shorter than
palm. Pereopod 3 propodus subpentagonal, with strong
broadly rounded proximal lobe on lower margin, lower
margin straight-concave. Pereopod 4 propodus minutely
chelate, fixed finger as long as dactylus. Male pleopod 1
consisting of 2 articles, article 2 ovoid, with shallow apical
notch between rounded subdistal lobe and curved acute apex.
Male pleopod 2 appendix interna free, articulating, distal on
mesial endopod margin (fig. 15m). Pleopods 3–5 appendices
internae barely emerging from endopodal margin. Uropodal
endopod with convex anterior margin, acute-rounded apex,
straight-slightly curved posterior margin, longer than wide.
111
Callianassidae and related families
Telson converging to about half basal width from near anterior
width over most of length to rounded posterolateral corners,
posterior margin straight or convex between tapering
posterolateral margins.
Remarks. Only one Recent species is known. The antennular
peduncle is particularly long, the rostrum particularly short and
the merus of the major cheliped has a convex tuberculate blade
along most of its lower margin.
Karumballichirus gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:E21E0313-39894E03-BBB6-9F9844028DC0
Type species. Callianassa karumba Poore and Griffin, 1979,
by present designation and monotypy.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique
branchiostegal ridge with which it articulates by means of a
virtual condyle. Rostrum spine-like. Pleomere 1 tergite fused,
divided into 2 sections by transverse groove or shoulder.
Maxilliped 3 ischium and merus narrow, more than twice as
long as wide at their articulation; propodus about as wide as
long, free distal margin clearly oblique; exopod present.
Male major cheliped merus lower margin straight,
irregularly dentate, usually with 1 more prominent
proximal denticulate tooth. Male minor cheliped much
narrower than major, carpus longer than palm, fingers shorter
than palm. Pereopod 3 propodus subpentagonal, with strong
broadly rounded proximal lobe on lower margin, lower
margin straight-concave or weakly convex. Pereopod 4
propodus simple, distally rounded on lower margin. Male
pleopod 1 consisting of 2 articles, article 2 ovoid, with shallow
apical notch between rounded subdistal lobe and curved acute
apex. Male pleopod 2 appendix interna free, articulating,
distal on mesial endopodal margin. Pleopods 3–5 appendices
internae barely emerging from endopodal margin. Uropodal
endopod with convex anterior margin, acute-rounded apex,
straight-slightly curved posterior margin, longer than wide.
Telson with convex lateral margins and rounded posterolateral
corners, posterior margin straight or convex between tapering
posterolateral margins.
Etymology. An alliteration of the name of the type species and
Callichirus, type genus of the family.
Remarks. Karumballichirus differs from the most similar
callichirid genera, Neocallichirus, in which it was most recently
placed (see Sakai, 1988) and Corallianassa (see Komai et al.,
2015) in possession of a short maxillipedal 3 exopod, a feature
shared with Lepidophthalmus, which is its sister taxon on the
molecular phylogram (Robles et al., in press). The type species
was described in detail by Dworschak (2008). The type species
was recognised as the extant representative of a group of similar
fossil species by Hyžný et al. (2017).
Kraussillichirus gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:07353B50-6C2E4CD3-A3DF-F37D1497A8C3
Type species. Callianassa kraussi Stebbing, 1900, by original
designation and monotypy.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique
branchiostegal ridge with which it articulates by means of a
virtual condyle. Rostrum obsolete or obtusely triangular, flat,
not reaching cornea. Pleomere 1 tergite divided into 2 sections
by unsclerified band; tergite weakly sclerotised, if so only
posteriorly; pleura oblique thin rods independent of tergite
dorsally, separated by extensive flexible region; sternite a thin
transverse plate. Antennular peduncle length more than 3
times the width of both eyestalks, longer than antennal
peduncle. Maxilliped 3 ischium and merus less than twice as
long as wide at their articulation; crista dentata absent (or few
proximal spines only); propodus about as wide as long, free
distal margin transverse or nearly so; exopod present. Male
major cheliped merus with denticulate blade, more prominent
over proximal half of lower margin. Male minor cheliped much
narrower than major, carpus longer than palm, fingers shorter
than palm. Pereopod 3 propodus subpentagonal, with strong
broadly rounded proximal lobe on lower margin, lower margin
straight-concave or weakly convex. Pereopod 4 subchelate.
Male pleopod 1 consisting of 2 articles, article 2 longer than
wide, with or without slight apical notch. Male pleopod 2
appendix interna absent. Pleopods 3–5 appendices internae
barely emerging from endopod margin. Uropodal endopod
with convex anterior margin, acute-rounded apex, straightslightly curved posterior margin, longer than wide. Telson with
convex lateral margins and rounded posterolateral corners,
posterior margin convex, with medial lobe.
Etymology. An alliteration of the name of the type species and
Callichirus, type genus of the family.
Remarks. Callianassa kraussi was included in the subgenus
Callichirus by De Man (1928) and the genus Callichirus by
Sakai (2005b and later). Kraussillichirus resembles Callichirus
but differs in lacking its ornamented pleomeres 3–6 and having
an ovoid uropodal endopod (fig. 14o). The only species appears
to be estuarine (Hanekom and Russell, 2015; Siebert and
Branch, 2005) as are species of Lepidophthalmus, which it also
somewhat resembles.
Laticallichirus Komai, Yokooka, Henmi and Itani, 2019
Laticallichirus Komai et al., 2019: 463–466.
Type species. “Neocallichirus” grandis Karasawa and Goda,
1996, by original designation and monotypy.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique branchiostegal
ridge with which it articulates by means of a virtual condyle.
Rostrum spine-like. Pleomere 1 tergite divided into 2 sections
by unsclerotised band; tergite weakly sclerotised, if so only
posteriorly; pleura oblique thin rods independent of tergite
112
dorsally, separated by extensive flexible region; sternite a thin
transverse plate. Antennular peduncle length more than 3 times
the width of both eyestalks, longer than antennal peduncle.
Maxilliped 3 ischium and merus broad, less than twice as long
as wide at their articulation; crista dentata absent; propodus
about as wide as long, free distal margin clearly oblique. Male
major cheliped merus with convex denticulate blade. Male
minor cheliped much narrower than major, carpus longer than
palm, fingers shorter than palm. Pereopod 3 propodus
subpentagonal, with strong broadly rounded proximal lobe on
lower margin, lower margin straight-concave. Pereopod 4
subchelate. Male pleopod 1 of 2 articles, article 2 longer than
wide, with or without slight apical notch. Male pleopod 2
appendix interna absent. Pleopods 3–5 appendices internae
barely emerging from endopod margin. Uropodal endopod with
convex anterior margin, acute-rounded apex, straight-slightly
curved posterior margin, longer than wide. Telson with convex
lateral margins and rounded posterolateral corners, posterior
margin concave between rounded posterolateral corners, with
transverse row of robust setae.
Remarks. The type species was described from Japanese
Middle Pleistocene fossil chelipeds but extant specimens
enabled the species to be described more completely. Komai et
al. (2019) compared the new genus with several others. Their
phylogram based on the 16S rRNA gene found the species
closest to Callichirus and more remote from a clade including
Neocallichirus, Corallianassa, Glypturus, Glypturoides and
Lepidophthalmus. The genus appears closest to
Michaelcallianassa, as our key indicates.
Lepidophthalmus Holmes, 1904
Lepidophthalmus Holmes, 1904: 310.—Manning and Felder,
1991: 778–779.—Sakai, 1999a: 64–65.—Sakai and Apel, 2002: 278.—
Sakai, 2005b: 143–144.—Felder, 2001: 440.—Sakai, 2011: 446.—
Robles and Felder, 2015: 462–464, 467–468, fig. 1.—Komai et al.,
2018: 23–24.
Podocallichirus Sakai, 1999a: 53–54.—Sakai, 2005b: 187–189
(partim).—Sakai, 2011: 466–467.—Hyžný and Muñiz, 2012: 619–
621.—Hyžný and Karasawa, 2012: 62–63.—Hyžný, 2016: 44–45.—
Komai et al., 2018: 24 (type species: Callianassa madagassa Lenz
and Richters, 1881, by original designation and monotypy) syn. nov.
Lepidophthalmoides Sakai, 2011: 440–441 (type species:
Lepidophthalmus eiseni Holmes, 1904, by original designation)
objective synonym.
Lepidophthalminus Sakai, 2015: 433 (replacement name for
Lepidophthalmus sensu Sakai, 2011; type species, Callianassa
bocourti A. Milne-Edwards, 1870, by original designation) syn. nov.
Type species. Lepidophthalmus eiseni Holmes, 1904, by
monotypy.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique
branchiostegal ridge which it meets level with linea thalassinica.
Rostrum spine-like; anterolateral angles obsolete or spinous.
Pleomere 1 tergite undivided or with weak transverse step.
Antennular peduncle length less than 3 times as width of both
eyestalks or more than 3 times the width of both eyestalks
(rare), longer than antennal peduncle. Maxilliped 3 ischium
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
and merus narrow, more than twice as long as wide at their
articulation, or rarely less than twice as long as wide at their
articulation; crista dentata absent (or few proximal spines
only); propodus about as wide as long, free distal margin
clearly oblique, or free distal margin transverse or nearly so;
dactylus curved, 3 times as long as wide, concave below;
exopod present. Male major cheliped merus with sinuous
lower margin, widest proximally and excavate beyond, or with
proximal tubercle bearing spine(s) with or without more distal
teeth along lower margin; dactylus typically tapering, or
massive, upper margin expanded, strongly convex, dentate.
Male minor cheliped half or more as wide as major, carpus as
long as or shorter than palm, fingers as long as or longer than
palm. Pereopod 3 propodus subpentagonal, with strong broadly
rounded proximal lobe on lower margin, lower margin straightconcave, lower margin with broad deep notch between distal
lobe and heel. Pereopod 4 subchelate. Male pleopod 1 consisting
of 2 articles, article 2 tapering, with small acute subdistal lobe,
acute distal apex. Male pleopod 2 appendix interna absent, or
reduced to obsolete distomesial lobe, or apparent only as field
of hooks on side of appendix masculina. Pleopods 3–5
appendices internae barely emerging from endopod margin.
Uropodal endopod with convex anterior margin, acute-rounded
apex, straight-slightly curved posterior margin, longer than
wide. Telson with convex lateral margins and rounded
posterolateral corners, posterior margin trilobed, excavate
each side of medial lobe, or convex, with medial lobe.
Remarks. Lepidophthalmus differs from other callichirids in
the combination of short antennal peduncle, telson with convex
lateral margins and quasi-trilobed posterior margin, a series of
concavities along the lower margin of the propodus of pereopod
3, and a curved narrow dactylus on maxilliped 3. Some species
have particularly setose fingers on the minor cheliped or with
long setae on the mesial face of the major cheliped extending
into the gape between the fingers. Lepidophthalmus shares an
exopod on maxilliped 3 with Karumballichirus,
Kraussillichirus and Glypturoides.
Sakai (2011) placed eight species in his new genus
Lepidophthalmoides Sakai, 2011, an objective synonym of
Lepidophthalmus Holmes, 1904. Both have the same type
species. Sakai (2015) realised the error and erected
Lepidophthalminus Sakai, 2015, to replace Lepidophthalmus
sensu Sakai, 2011, this time with Callianassa bocourti A.
Milne-Edwards, 1870, as type species, with only five included
species and referring to his 2011 diagnosis of Lepidophthalmus
to differentiate it from Lepidophthalmus Holmes, 1904. Sakai
(2011) separated the two “genera” in the male pleopod 1 being
“chelate” in the former and simple in the latter. Incidentally, the
adjective “chelate” is inappropriate in this context—the second
article has a mesiodistal notch, not formed by two distal
opposing articles. A review of all 15 known species reveals that
one pleopod form grades into the other and may reflect
developmental stages. Komai et al. (2018) has already pointed
out that the generic division of Sakai (2015) has no merit.
Lepidophthalminus is here synonymised with Lepidophthalmus.
No consistent morphological difference was found between
American and Indo-West Pacific species (Komai et al., 2018;
113
Callianassidae and related families
Robles et al., in press; Robles and Felder, 2015). Most species
of Lepidophthalmus are found in estuarine habitats where
their ecology has been studied (Felder, 2001; Filho et al., 2013;
Hernáez et al., 2012).
Michaelcallianassa Sakai, 2002
Michaelcallianassa Sakai, 2002: 480–481.—Sakai, 2005b: 156–
157.—Sakai, 2011: 450.
Type species. Michaelcallianassa indica Sakai, 2002, by
original designation and monotypy.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique
branchiostegal ridge with which it articulates by means of a
virtual condyle. Rostrum obsolete or obtusely triangular, flat,
not reaching cornea. Pleomere 1 tergite undivided or with weak
transverse step. Pleomeres 3–5 with anteriorly converging
longitudinal grooves and oblique-transverse rows of long
setae. Antennular peduncle longer than antennal peduncle.
Maxilliped 3 ischium and merus narrow, more than twice as
long as wide at their articulation; propodus about as wide as
long, free distal margin clearly oblique. Male major cheliped
merus with lower margin almost straight. Male minor
cheliped half or more as wide as major, carpus as long as or
shorter than palm, fingers as long as or longer than palm.
Pereopod 3 propodus subpentagonal, with strong broadly
rounded proximal lobe on lower margin, lower margin straightconcave. Pereopod 4 subchelate. Male pleopod 1 of 2 articles,
article 2 longer than wide, with or without slight apical notch.
Male pleopod 2 uniramous; appendix interna absent or
reduced to obsolete distomesial lobe. Female pleopod 2
uniramous. Pleopods 3–5 appendices internae barely emerging
from endopod margin. Uropodal endopod with convex anterior
margin, acute-rounded apex, straight-slightly curved posterior
margin, longer than wide. Telson converging to about half
basal width from near anterior width over most of length to
rounded posterolateral corners, posterior margin concave
between rounded posterolateral corners, with transverse row of
robust setae.
Remarks. The two species of Michaelcallianassa have
uniramous pleopods 2 in both sexes distinguishing the genus
from all other callichirids (fig. 15n). The telson has a transverse
row of robust setae. Sakai (2002) noted that the type species
was similar to species of Callichirus but has a row of setae
between the two dorsolateral tufts on pleomeres 4 and 5. This
row is not present on the second species, M. sinica (see Liu and
Liu, 2009).
Mocallichirus gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:8E845452-37C74271-B787-E28D6285AE8F
Type species. Callianassa mocambiquensis Sakai, 2004, by
present designation and monotypy.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique
branchiostegal ridge with which it articulates by means of a
virtual condyle. Rostrum obsolete or obtusely triangular, flat,
not reaching cornea. Pleomere 1 tergite undivided or with weak
transverse step. Maxilliped 3 ischium and merus narrow, more
than twice as long as wide at their articulation; crista dentata a
row of numerous denticles; propodus free distal margin clearly
oblique. Male major cheliped merus with denticulate blade,
more prominent over proximal half of lower margin. Male
minor cheliped half or more as wide as major, carpus as long as
or shorter than palm, fingers as long as or longer than palm.
Pereopod 3 propodus subpentagonal, with strong broadly
rounded proximal lobe on lower margin, lower margin straightconcave. Pereopod 4 propodus simple, distally rounded on
lower margin. Male pleopod 1 consisting only of 1 article.
Male pleopod 2 uniramous; appendix interna absent or reduced
to obsolete distomesial lobe. Pleopods 3–5 appendices internae
barely emerging from endopod margin. Uropodal endopod
with convex anterior margin, acute-rounded apex, straightslightly curved posterior margin, longer than wide. Telson
parallel-sided over most of length, with broadly rounded apex,
domed, posterior margin broadly semicircular.
Etymology. An alliteration of the name of the type species and
Callichirus, type genus of the family.
Remarks. Robles et al.’s (in press) molecular analysis placed
Callianassa mocambiquensis sister to Michaelcallianassa
(two species) in Callichiridae. Sakai (2004) thought the species
similar to Callianassa plantei Sakai, 2004 (also found in
Mozambique), C. joculatrix and three others, all members of
Callianassidae s.s. GCBP was able to examine numerous
specimens from Madagascar from the collections of MNHN
and FMNH. The broad female pleopod 2 , the attenuated distal
lobe on the epipod of maxilliped 1 and the apical notch on the
male pleopod 1 confirm the species as a callichirid. The species
is recognisable by the unusual shape of the telson; it is domed,
longer than in most callichirids with a broadly semicircular
posterior half parallel-sided over most of length, with broadly
rounded apex. The antennal peduncle is longer than the
antennular peduncle, the male pleopod 1 is of one article and
the male pleopod 2 uniramous.
Mocallichirus mocambiquensis shares with members of
the callianassid genus Coriollianassa a proximal neck on the
carpus of the chelipeds, a case of remarkable convergence.
Mucrollichirus gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:B57E69B6-D096459C-B4AF-0D278A73B557
Type species. Callianassa mucronata Strahl, 1862, by present
designation and monotypy.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique
branchiostegal ridge with which it articulates by means of a
virtual condyle. Rostrum spine-like. Pleomere 1 tergite
undivided or with weak transverse step. Maxilliped 3 ischium
and merus narrow, more than twice as long as wide at their
articulation; propodus about as wide as long, free distal margin
114
clearly oblique. Male major cheliped merus lower margin
convex, or blade like, especially proximally, with small
denticles. Male minor cheliped half or more as wide as major,
carpus as long as or shorter than palm, fingers as long as or
longer than palm. Pereopod 3 propodus oval, lower margin
convex, not proximally lobed. Pereopod 4 subchelate. Male
pleopod 1 consisting of 2 articles, article 2 ovoid, with shallow
apical notch between rounded subdistal lobe and curved acute
apex. Male pleopod 2 appendix interna absent or reduced to
obsolete distomesial lobe. Female pleopod 2 endopod 2–3
times as long as wide. Pleopods 3–5 appendices internae barely
emerging from endopod margin. Uropodal endopod with
convex anterior margin, acute-rounded apex, straight-slightly
curved posterior margin, longer than wide. Telson converging
to about half basal width from near anterior width over most of
length to rounded posterolateral corners, posterior margin
straight or with slight medial lobe, smooth dorsally.
Etymology. An alliteration of the name of the type species and
Callichirus, type genus of the family.
Remarks. Mucrollichirus has a more weakly lobed propodus
on pereopod 3 than all other genera except Calliapagurops.
The new genus differs from Neocallichirus (see Sakai, 1988),
Corallianassa (see Komai et al., 2015) and other similar genera
in having a shorter telson. Both the morphological and
molecular analyses placed Mucrollichirus mucronatus on a
clade sister to Corallianassa (Robles et al., in press). The
similarity to Neocallichirus, the genus in which the species
was most recently placed (Sakai, 2011), is more remote.
The molecular analysis of Robles et al. (in press) found no
difference between individuals from Papua New Guinea and
the Philippines but illustrations of Mucrollichirus mucronatus
vary (De Man, 1888, 1928; Dworschak, 1992; Poore and
Griffin, 1979; Sakai, 1999a; Sakai et al., 2014; Tirmizi, 1977).
Neocallichirus Sakai, 1988
Neocallichirus Sakai, 1988: 61–62.—Manning and Felder, 1991:
779–780.—Poore, 1994: 102.—Sakai, 1999a: 84–86.—Sakai, 2000:
92.—Davie, 2002: 461.—Poore, 2004: 184.—Sakai, 2005b: 160–
162.—Sakai, 2011: 451–452.—Hyžný and Karasawa, 2012: 60–65.
Sergio Manning and Lemaitre, 1994: 40.—Sakai, 2011: 467–468
(type species, Callianassa guassutinga Rodrigues, 1971, by original
designation).—Hyžný and Karasawa, 2012: 61–62.
Callichiropsis Sakai, 2010: 1453 (type species, Callichiropsis
spiridonovi Sakai, 2010, by original designation and monotypy)
syn. nov.
Type species. Neocallichirus horneri Sakai, 1988, by original
designation.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique branchiostegal
ridge with which it articulates by means of a virtual condyle.
Rostrum obsolete or obtusely triangular, flat, not reaching
cornea; anterolateral spines absent (rarely with small spines).
Pleomere 1 tergite undivided or with weak transverse step.
Maxilliped 3 ischium and merus linear; propodus about as wide
as long, free distal margin transverse or nearly so. Male major
cheliped merus lower margin convex or blade like, especially
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
proximally, with small denticles. Male minor cheliped palm half
or more width of major cheliped palm, carpus as long as or
shorter than palm, fingers as long as or longer than palm.
Pereopod 3 propodus subpentagonal, with strong broadly
rounded proximal lobe on lower margin, lower margin straightconcave. Pereopod 4 subchelate. Male pleopod 1 consisting of 2
articles, article 2 ovoid, with shallow apical notch between
rounded subdistal lobe and curved acute apex. Male pleopod 2
appendix interna reduced to obsolete distomesial lobe, or rarely
free, articulating, distal on mesial endopod margin, or absent.
Pleopods 3–5 appendices internae barely emerging from
endopod margin. Uropodal endopod with straight or slightly
convex anterior margin ending in rounded angle, transverse
or near-transverse distal margin continuous with curved
posterior margin, as wide or wider than long. Telson converging
to about half basal width from near anterior width over most of
length to rounded posterolateral corners, posterior margin
straight or with slight medial lobe.
Remarks. Neocallichirus is the largest callichirid genus. The
uropodal endopod has a straight or slightly convex anterior
margin ending in a rounded angle and continuous distal and
posterior margins. The endopod is often wider than long. The
telson tapers regularly to about half its width. Sakai (2011)
provided a key to 28 species, including N. mucronatus, herein
removed to Mucrollichirus gen. nov., and N. karumba (=N.
kempi), herein removed to Karumballichirus gen. nov. Sakai
(1999a, 2005b) included Sergio in the synonymy of
Neocallichirus, but in 2011 he redefined the former and placed
some of its originally included species in a re-diagnosed genus.
He differentiated Sergio with a notched apex on the male pleopod
1 (adult form) from Neocallichirus with only a slight distal
indentation (juvenile form), despite most species of
Neocallichirus being described as having the adult form. The
molecular analysis (Robles et al., in press) placed three species of
“Sergio” within the Neocallichirus clade, a result anticipated by
Sepahvand et al. (2018).
Sakai (2010) differentiated a new genus Callichiropsis
from Lepidophthalmus on the basis of a different pereopod 3
and from Podocallichirus on the shape of the telson. We treat
Podocallichirus as a synonym of Lepidophthalmus (see
above). He did not compare Callichiropsis spiridonovi Sakai,
2010, with Neocallichirus, from which it does not differ.
Callichiropsis is here synonymised with Neocallichirus.
Thailandcallichirus Sakai, 2011
Thailandcallichirus Sakai, 2011: 473–474.—Komai et al., 2018:
23–24.
Type species. Callianassa ranongensis Sakai, 1983, by original
designation and monotypy.
Diagnosis. Anterior branchiostegal lobe sclerotised, well
produced anteriorly beyond junction with oblique branchiostegal
ridge, which it meets level with linea thalassinica. Rostrum
spine-like. Pleomere 1 tergite undivided or with weak transverse
step. Maxilliped 3 ischium and merus narrow, more than twice
as long as wide at their articulation; crista dentata absent (or few
proximal spines only); propodus about as wide as long, free
Callianassidae and related families
distal margin clearly oblique; dactylus curved, about twice as
long as wide, concave below. Male major cheliped merus with
proximal tubercle bearing spine(s) with or without more distal
teeth along lower margin; dactylus massive, upper margin
expanded, strongly convex, smooth. Male minor cheliped half
or more as wide as major, carpus as long as or shorter than palm,
fingers as long as or longer than palm. Pereopod 3 propodus
subpentagonal, with strong broadly rounded proximal lobe on
lower margin, lower margin straight-concave, lower margin with
broad deep notch between distal lobe and heel. Pereopod 4
subchelate. Male pleopod 1 consisting of 2 articles, article 2
tapering, with small acute subdistal lobe and acute distal apex.
Male pleopod 2 appendix interna absent or reduced to obsolete
distomesial lobe. Pleopods 3–5 appendices internae barely
emerging from endopod margin. Uropodal endopod with convex
anterior margin, acute-rounded apex, straight-slightly curved
posterior margin, longer than wide. Telson tapering from greatest
width near base, posterior margin straight or convex between
tapering posterolateral margins.
Remarks. Thailandcallichirus differs from Lepidophthalmus,
its closest sibling, in having the antennular peduncle shorter
than the antennal, rather than the other way around, a more
compact maxilliped 3 dactylus, lacking a maxillipedal 3
exopod, a convex posterior margin on the telson and in having
a massive dactylus on the major cheliped (Komai et al., 2018).
The minor cheliped is distally setose as in some
Lepidophthalmus species. Sakai (2011) justified his new genus
on differences between the type species and Neocallichirus,
the genus in which he had previously placed it (Sakai, 1999a,
2005b). Tudge et al. (2000) found it was a sister to other species
of Lepidophthalmus and treated it as a species of this genus as
did Robles and Felder (2015). Robles et al. (in press) found this
alliance again but had no molecular data to support it.
Ctenochelidae Manning and Felder, 1991
Figures 16, 17
Ctenochelinae Manning and Felder, 1991: 784.—Poore, 1994:
103.—Sakai, 1999c: 87.—Sakai, 2005b: 235.
Gourretiinae Sakai, 1999c: 95.—Sakai, 2017a: 1124.
Gourretiidae.—Sakai, 2004: 556.—Sakai, 2005b: 217.—Sakai,
2011: 507–508.—Sakai, 2017a: 1124–1125.—Dworschak and Poore,
2018: 68.
Ctenochelidae.—Sakai, 2005b: 235.—Sakai, 2011: 485–486.—
Dworschak et al., 2012: 189–190.—Poore and Dworschak, 2017: 122
(synonymy).
Pseudogourretiinae Sakai, 2005b: 230.
Dawsoniinae Sakai, 2006: 1276–1277.—Sakai, 2011: 507.—
Sakai, 2017a: 1124.
Ctenocheloidae (pro Ctenocheloididae) Sakai, 2011: 595–596.—
Poore, 2015b: 142 syn. nov.
Pseudogourretiidae Sakai, 2011: 507–508, 523.
Tosacallianassidae Sakai, 2016: 813.
Diagnosis. Rostrum spike-like, longer than wide; median
carina absent, or rarely on rostrum only; gastric carinae absent;
cervical groove well defined; suture between ocular lobe and
end of linea thalassinica horizontal or oblique in lateral view;
anterior branchiostegal margin sinusoidal or semicircular, or
115
with deep obtuse notch leading to oblique ridge; anterior
branchiostegal lobe simple, scarcely calcified, merging
smoothly with anterodorsal branchiostegal angle and
anterolateral margin of carapace; posterior margin of
carapace without lateral lobes. Pleomere 1 without anterolateral
lobes, weakly chitinised. Eyestalks flattened, contiguous, with
subdistal dorsal cornea. Antennal scaphocerite simple,
triangular. Maxilla scaphognathite without long seta on
posterior lobe extending into branchial chamber. Maxilliped 1
epipod with acute anterior lobe lying alongside exopod.
Maxilliped 3 propodus longer than wide, not prominently
lobed on lower margin; dactylus slender, digitiform, with setae
irregularly spaced along all margins. Cheliped merus lower
margin smooth or with prominent proximal tooth; major
cheliped with cylindrical palm, fingers elongate and pectinate,
or with distinctively flattened palm, sometimes with strong
crest above and below. Pereopod 3 propodus rectangular or
oval, up to twice as long as wide, with proximal lobe on lower
margin, without distal spiniform setae on lateral face or lower
margin. Pereopod 5 semichelate (fixed finger closing on
proximal part of dactylus), dactylus a twisted plate longer
than finger. Female pleopod 2 rami narrower and with more
reduced setation than pleopods 3–5; endopod flattened and 2–3
times as long as wide. Pleopods 3–5 with oblique peduncles,
endopods oval, exopods attached laterally, not proximally
lobed, shorter than and barely overlapping endopods, or with
oblique peduncles meeting mesially, endopods triangular, with
straight mesial margin, exopods attached laterally, proximally
lobed, longer than and enclosing endopods; appendices internae
elongate, much longer than wide. Uropodal exopod without
elevated dorsal plate, or with margin divided by notch.
Remarks. Robles et al. (in press) found Ctenochelidae to have
strong molecular support and two morphological
synapomorphies. The character states shared by members of
this family are maxilliped 3 with a prominent toothed crista
dentata, usually extending beyond the meral distal margin, and
a semichelate pereopod 5, its dactylus a twisted plate longer
than the finger. Two clades, ctenochelids s.s. and gourretiids,
were supported by morphological characters but not by
molecular relationships.
Manning and Felder (1991) treated Ctenochelidae as one
of two “callianassid-like families”, distinguished from
Callianassidae by the presence of an appendix masculina on
the male pleopod 2, which was incorrect, as realised later by
Felder and Manning (1994: 352), a longitudinally carinate
uropodal exopod, lacking a dorsal plate and a carapace usually
with a cardiac prominence. Their arrangement was followed
by Poore (1994). The genera of Ctenochelidae known to these
authors are treated in three families here. Sakai (1999c)
included only Paracalliax beside the type genus and excluded
Dawsonius and Gourretia in a new subfamily Gourretiinae.
Sakai (2005b) elevated the taxon to family rank, retaining its
single subfamily that he ignored in Sakai (2011), and confined
Ctenochelidae to Ctenocheles.
As pointed out by Poore (2015b), Sakai’s (2011) diagnosis of
his new family Ctenocheloididae (as Ctenocheloidae) is virtually
the same as Anker’s (2010) diagnosis of Ctenocheloides and was
116
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Figure 16. Diagnostic characters for genera of Ctenochelidae. Pleon (suture between pleomeres 2 and 3 indicated): a, Ctenocheles; b,
Ctenocheloides. Pleomere 6, telson, uropod: c, Dawsonius; d, Gourretia; e, Paragourretia; f, Laurentgourretia. Rostrum, eyestalks, antennules,
antennae: g, h, Ctenocheles; i, Ctenocheloides; j, k, Laurentgourretia.
Original illustrations: f, j, k, Laurentgourretia rhopalommata, MNHN-IU-2014-11417.
Callianassidae and related families
117
Figure 17. Diagnostic characters for genera of Ctenochelidae. Major cheliped: a, Ctenocheles; b, Ctenocheloides; c, Kiictenocheloides; d,
Gourretia. Minor cheliped: e, Ctenocheloides; f, Kiictenocheloides; g, h, Paragourretia; i, Gourretia. Maxilliped 3: j, Gourretia. Male coxa 5:
k, Laurentgourretia. Male pleopods 1, 2: l, m, Laurentgourretia.
Original illustrations: k–m, Laurentgourretia rhopalommata, MNHN-IU-2014-11417.
118
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
presented without a strong argument for why family status is to
be preferred over genus status for its single species. We treat
Ctenocheloididae as a junior synonym. Poore and Dworschak
(2017) synonymised three taxa erected by Sakai and Türkay
(2016), Tosacallianassidae with Ctenochelidae, Tosacallianassa
with Ctenocheles, and T. hatasagaensis with Ctenocheles balssi.
Gourretiinae was established by Sakai (1999c) as a
subfamily of Callianassidae for Gourretia de Saint Laurent,
1973, and Dawsonius Manning and Felder, 1991. Sakai (2004)
elevated the taxon to family level and added Callianopsis de
Saint Laurent, 1973, and two new genera, Laurentgourretia
and Paragourretia. Pseudogourretiinae was then added for
another new genus, Pseudogourretia Sakai, 2005b. Sakai
(2006) recognised Callianopsinae and Pseudogourretiinae as
subfamilies of Gourretiidae and added Dawsoniinae for
Dawsonius. All subfamilies were ignored in Sakai’s (2011)
synthesis where four of the genera discussed earlier were
included; Paracalliax was added although it had earlier been
treated as member of its own family (see discussion under
Paracalliacidae below), and Callianopsis was excluded (see
Callianopsidae discussed above).
Sakai (2017b) radically dissected Gourretiidae, introducing
five new genera. Paragourretia and Dawsonius were treated
as synonyms of Gourretia, Paracalliax was retained and his
earlier genus Pseudogourretia ignored. Justification of the
new genera centred on differences between the male pleopods
1 and 2. He stressed the significance of these sexually
differentiated structures, citing their importance in the
taxonomy of other decapods and their greater significance
than the few cheliped characters available to palaeontologists
(Hyžný and Klompmaker, 2015). However, he took no account
of pleopodal developmental as individuals mature from
juveniles to adults. All five of Sakai’s (2017b) new genera are
herein synonymised with either Gourretia or Paragourretia.
On strictly cladistic criteria, Dawsonius is a senior
synonym of Paragourretia on molecular evidence (Robles et
al., in press). But the genus has unique apomorphies apparent
in the morphological treatment, pleomere 6 with triangular
lateral projections, maxilliped 3 exopod absent and pereopod
3 with a triangular propodus proximally produced on the
lower margin. Otherwise the two genera have the same
uropods, male pleopods and chelipeds.
Pseudogourretia, said by Sakai (2005b) to possess
pleurobranchs, was found by Dworschak and Poore (2018) to not
have pleurobranchs, as all callianassoids, and the genus was
synonymised with Gourretia. The only character placing the
sole species, P. portsudanensis in Gourretia rather than
Paragourretia is the oval pereopod 3 propodus; chelipeds and
uropods are missing from the only specimen. The family erected
for it (Sakai, 2011) likewise is a junior synonym of Ctenochelidae.
2.
Rostrum spine-like, dorsally carinate; cornea flat, with
weak pigmentation (fig. 16g); pleomeres 1 and 2 together
half as long as pleomeres 3–5 (fig. 16a)
Ctenocheles
–
Rostrum obsolete, flat; cornea disc-like, strongly
pigmented (fig. 16i); pleomeres 1 and 2 together as long as
pleomeres 3–5 (fig. 16b)
Ctenocheloides
3.
Telson longer than wide, parallel-sided over most of
length, with curved truncate apex (fig. 16f); carapace with
anterolateral spines (figs 16j, k); maxilliped 3 exopod
absent
Laurentgourretia
–
Telson shorter or longer than wide, tapering over most of
length to rounded apex (figs 16c–e); anterolateral angles
of carapace obsolete; maxilliped 3 exopod present (fig.
17j) or absent
4
4.
Pleomere 6 with triangular lateral projections (fig. 16c);
telson broadly rounded over posterior two-thirds (fig. 16c)
Dawsonius
–
Pleomere 6 without triangular lateral projections (P.
biffari excepted); telson tapering over posterior half to
two-thirds (figs 16d, e)
5
5.
Major cheliped merus lower margin with tubercle at
midpoint (fig. 17c); minor cheliped palm swollen, fingers
pectinate, spines erect (fig. 17f); maxilliped 3 exopod
absent
Kiictenocheloides
–
Major cheliped merus lower margin with straight or weakly
convex blade on lower margin bearing a short proximal
curved spine and 1 or more distal spines (fig. 17d); minor
cheliped palm tapering (fig. 17g), fingers with or without
teeth; maxilliped 3 exopod present (fig. 17j)
6
6.
Minor cheliped with sharp proximally directed teeth along
cutting edges of fingers (fig. 17i); major cheliped carpus
lower proximal margin swollen (fig. 17d); uropodal exopod
without marginal notch (fig. 16d)
Gourretia
–
Minor cheliped without sharp teeth along cutting edges of
fingers (figs 17g, h); major cheliped carpus lower proximal
margin tapering; uropodal exopod usually with marginal
notch, slight dorsal plate (fig. 16e)
Paragourretia
Implicit attributes. Unless indicated otherwise, the following
attributes are implicit throughout the generic diagnoses.
Cardiac prominence absent. Anterolateral angle obsolete,
unarmed. Pleomere 6 without lateral projections. Maxilliped 3
merus with 1 mesiodistal spine; exopod absent. Major cheliped
carpus proximal and lower margins oblique, evenly curved;
propodus rectangular, fingers stout, fixed finger with broad
tooth. Minor cheliped merus lower margin smooth.
Key to genera of Ctenochelidae
Ctenocheles Kishinouye, 1926
1.
Major cheliped with palm swollen, with narrow pectinate
fingers longer than palm (figs 17a, b)
2
–
Major cheliped with palm flattened, upper and lower
margins usually ridged, fingers rarely longer than palm
(figs 17c, d)
3
Ctenocheles Kishinouye, 1926: 36.—de Saint Laurent, 1973:
514.—Poore and Griffin, 1979: 277.—de Saint Laurent and Le Loeuff,
1979: 81–83.—Sakai, 1999c: 88.—Sakai, 2005b: 237–238.—Sakai,
2011: 486–487 (key to species).—Hyžný et al., 2014: 249–250.—Poore
and Dworschak, 2017: 122 (synonymy).
Tosacallianassa Sakai and Türkay, 2016: 813 (type species,
119
Callianassidae and related families
Tosacallianassa hatasagaensis Sakai and Türkay, 2016, by original
designation and monotypy).
Type species. Ctenocheles balssi Kishinouye, 1926, by monotypy.
Diagnosis. Cardiac prominence present. Rostrum blade-like,
produced anteriorly. Male major cheliped merus with oblique
spine near midpoint of lower margin (or absent); propodus
swollen, with fine pectinate fingers at least twice as long as
palm. Minor cheliped merus lower margin smooth, or with
spine at or near midpoint; propodus cylindrical, slightly
swollen at midpoint, about twice as wide as long. Male pleopod
1 article 2 simple, tapering, without coupling hooks. Telson
parallel-sided over anterior half, then tapering to evenly
rounded apex.
Remarks. The blade-like rostrum combined with the fine long
pectinate fingers of the major cheliped of Ctenocheles make its
species easily recognisable. In fact, some species are known
only from these claws (Holthuis, 1967; de Saint Laurent,
1979a). To the six described species listed by Sakai (2005b,
2011) can be added C. plantei (Burukovsky, 2005), described
as a member of the lobster genus Thaumastochelopsis Bruce,
1988 (Poore and Dworschak, 2017).
Ctenocheloides Anker, 2010
Ctenocheloides Anker, 2010: 1790–1791.—Sakai, 2011: 596.—
Poore, 2015b: 142–143.
Type species. Ctenocheloides attenboroughi Anker, 2010, by
original designation and monotypy.
Diagnosis. Cardiac prominence present. Rostrum obsolete or
obtusely triangular, flat, not reaching cornea. Maxilliped 3
merus with mesial spine at midpoint. Male major cheliped
merus with oblique spine near midpoint of lower margin;
propodus swollen, with fine pectinate fingers about as long as
palm. Minor cheliped merus lower margin with spine at or near
midpoint; propodus cylindrical, slightly swollen at midpoint,
about twice as wide as long. Male pleopod 1 article 2 simple,
tapering, without coupling hooks. Telson parallel-sided over
anterior half, then tapering to evenly rounded apex.
Remarks. The pectinate fingers on the major cheliped are a
little longer than the palm, whereas in Ctenocheles the fingers
are at least twice as long as the palm, and the spikes more
pronounced. Three species are known (Poore, 2015b).
Ctenocheloides boucheti is a sister taxon to two species of
Ctenocheles in the molecular analysis (Robles et al., in press).
Dawsonius Manning and Felder, 1991
Dawsonius Manning and Felder, 1991: 785.—Sakai, 2005b:
245.—Sakai, 2011: 508.
Gourretia.—Sakai, 2017a: 1128–1129 (partim).
Type species. Callianassa latispina Dawson, 1967, by original
designation and monotypy.
Diagnosis. Cardiac prominence present. Rostrum sharply
triangular, produced to near apex of ocular lobes. Pleomere 6
with lateral projections. Male major cheliped merus with
straight or weakly convex blade on lower margin bearing a
short proximal curved spine and 1 or more distal spines;
propodus rectangular, fingers fine, fixed finger smooth. Minor
cheliped merus lower margin with erect proximal spine (as in
major cheliped); propodus tapering, about 3 times as long as
wide, with long narrow fingers. Pereopod 3 propodus triangular,
with straight upper margin, expanded proximally on lower
margin. Male pleopod 1 article 2 with lobed distomesial
margin, distal notch, sharp curved distolateral apex. Telson
parallel-sided over anterior half, then tapering to evenly
rounded apex.
Remarks. Dawsonius latispina differs from other ctenochelids
(except Paragourretia biffari Blanco Rambla and Liñero
Arana, 1994) in having lateral projections on pleomere 6.
Manning and Felder (1991) differentiated Dawsonius from
Gourretia on the absence of an exopod on maxilliped 3 and
the sharp lateral projections on pleomere 6. We (DLF) can
confirm that maxillipedal 3 exopod is absent (Dawson, 1967;
Manning and Felder, 1991) and that Biffar’s (1971a) figure and
Sakai’s (2011) observation of an exopod are in error. Biffar’s
figures are a composite from types and Honduran material but
the latter is P. biffari.
Sakai (2005b) treated the genus first as a synonym of
Callianopsis and in an addendum as a separate genus that he
differentiated from Callianopsis. Callianopsis is a member of
another family. He did not explain what he meant by “sensu
Sakai (not Manning and Felder, 1991)” before his new
diagnosis. Sakai (2011) included Dawsonius in Gourretiidae.
Sakai’s (2011) observations of a “paratype” and a “lectotype”
(USNM 103755, 172310) were not made on the holotype and
paratype designated by Dawson (1967); these specimens may
well have been misidentified. Sakai’s (2011: fig. 69E) figure of
the apex of the male pleopod 2 is impossible to reconcile with
Manning and Felder (1991: fig. 16i), which is typical of the
family, nor with any ctenochelid.
Sakai (2017a) synonymised Dawsonius with Gourretia on
the basis of the same male pleopods 1 and 2. In fact, Dawsonius
is unique in pleopod 1 possessing a distomesial rounded blade
and acute apex, whereas in Gourretia and Paragourretia the
distomesial lobed is acute and smaller than the apical lobe.
Gourretia de Saint Laurent, 1973
Gourretia de Saint Laurent, 1973: 514.—Le Loeuff and Intès,
1974: 26.—Poore and Griffin, 1979: 278.—Manning and Felder, 1991:
785.—Sakai, 2002: 468–469.—Ngoc-Ho, 2003: 498–499.—Sakai,
2005b: 218–219 (partim).—Sakai, 2011: 510–511 (partim).
Pseudogourretia Sakai, 2005b: 230–231.—Dworschak and Poore,
2018: 68–70 (type species, Pseudogourretia portsudanensis Sakai,
2005, by original designation and monotypy).
Ivorygourretia Sakai, 2017a: 1126, 1130 (type species, Gourretia
barracuda Le Loeuff & Intès, 1974, by original designation and
monotypy) syn. nov.
Plantesgourretia Sakai, 2017a: 1126, 1131 (type species,
Gourretia nosybeensis Sakai, 2004, by original designation and
monotypy) syn. nov.
Ruiyuliugourretia Sakai, 2017a: 1126, 1131–1133 (type species,
Gourretia sinica Liu and Liu, 2010, by original designation and
monotypy) syn. nov.
120
Type species. Callianassa subterranea var. minor Gourret,
1887, by original designation and monotypy (now Gourretia
denticulata Lutze, 1937 [Lewinsohn and Holthuis, 1986]).
Diagnosis. Rostrum sharply triangular, produced to near
apex of ocular lobes, or obsolete, or obtusely triangular, flat,
not reaching cornea (rarely). Maxilliped 3 with exopod. Male
major cheliped merus with straight or weakly convex blade on
lower margin bearing a short proximal curved spine and 1 or
more distal spines; carpus proximal and lower margins
convex, swollen; propodus broad, with short fingers. Minor
cheliped merus lower margin with erect proximal spine (as in
major cheliped); propodus tapering, about 3 times as long as
wide; fingers evenly tapering, straight, dactylus longer, with
sharp spines along opposing edges. Pereopod 3 propodus
oval, little longer than wide, with short rounded proximal
lobe on lower margin, lower margin straight, upper margin
strongly convex. Male pleopod 1 article 2 with sharp
distomesial apex, distal notch, sharp curved distolateral apex.
Telson parallel-sided over anterior half, then tapering to
evenly rounded apex.
Remarks. In species of Gourretia the minor cheliped propodus
tapers evenly, the fingers are straight, the dactylus being
slightly longer, and both fingers have sharp spines, almost
hook-like in some, along their opposing edges. The fixed
finger of the major cheliped is stout, as is the dactylus, and
blade-like with a broad tooth. The fused appendices on the
male pleopod 2 reach or overlap the end of the endopod. This
contrasts with the minor chelipeds of species of Paragourretia
whose fingers lack the sharp spines and curve distally, the
fixed finger of the major cheliped evenly tapers and curves,
and the appendices on the male pleopod 2 are midmesial, far
from the apex of the endopod.
Sakai (2004) separated Paragourretia from Gourretia for
two species (see discussion below). In Sakai (2011), the two
genera were redefined and four additional species were
reallocated to Paragourretia from Gourretia. The key
character indicated then to separate the two genera was said to
be the absence in Gourretia and presence in Paragourretia of
a “lateral notch” on the uropodal exopod. A cardiac prominence
is absent in the former but present in the latter according to
their diagnoses. The presence or absence of a distal notch is
sometimes difficult to judge—it is not a black and white
character (see comments below under Paragourretia). Our
morphological analysis separates Paragourretia from
Gourretia on the structure of the major and minor chelipeds,
the shape of the propodus of pereopod 3 and the position of the
appendices on the male pleopod 2, the notch being of lesser
importance. None of these characters was included in the
generic diagnoses by Sakai (2005b, 2011). The allocation of
known species between the two genera is the same as that of
Sakai (2011).
Pseudogourretia was synonymised with Gourretia by
Dworschak and Poore (2018).
The male pleopod 1 of G. barracuda (Le Loeuff and Intès,
1974), said by Sakai (2017a) to be “hooked distally”, ends in a
simple apex, a more subdued form of the bifid apex usual in
this genus. The pleopod may be a subadult form. The appendix
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
masculina of the male pleopod 2 is more adpressed to the
endopod apex than typical but the basic structure is the same.
Ivorygourretia, erected for only this species, is here
synonymised with Gourretia.
Plantesgourretia Sakai, 2017, erected for only G.
nosybeensis, was said to have the male pleopod 2 “biramous;
exopod shorter than endopod; endopod bilobed distally, and
inner lobe bearing distally appendix masculina with setae, but
no appendix interna”. The exopod is shorter in most species;
the appendix masculina is more dominant than in other species
but is otherwise similar; we give no importance to the absence
of the appendix interna. G. nosybeensis is otherwise similar to
all species of Gourretia; Plantesgourretia is here synonymised.
Ruiyuliugourretia Sakai, 2017, again erected for a single
species, was said to have special male pleopods 1 and 2. The
outline of pleopod 1 is typical of species of Gourretia and its
supposed four articles are probable artefacts. Pleopod 2 is
typical, except for the absence of an appendix interna, as in
the previous species. Ruiyuliugourretia is here synonymised
with Gourretia.
Gourretia loeuffintesi Sakai, 2005, is a nomen nudum (no
holotype designated).
Kiictenocheloides Sakai, 2013
Kiictenocheloides Sakai, 2013: 1693.
Type species. Ctenocheloides nomurai Komai, 2013, by
original designation and monotypy.
Diagnosis. Rostrum obsolete or obtusely triangular, flat, not
reaching cornea. Male major cheliped merus with oblique
spine near midpoint of lower margin; propodus broad, with
short fingers. Minor cheliped merus lower margin with spine
at or near midpoint; propodus cylindrical, slightly swollen at
midpoint, about twice as wide as long, with long narrow
fingers. Male pleopod 1 article 2 simple, tapering, without
coupling hooks. Telson parallel-sided over anterior half, then
tapering to evenly rounded apex.
Remarks. The swollen propodus and thin pectinate fingers,
typical of Ctenocheles and Ctenocheloides (fig. 17a, b), are not
found in Kiictenocheloides where the major cheliped has a
propodus and fingers (fig. 17c) typical of callianassoids
generally. The female pleopods are unknown.
Laurentgourretia Sakai, 2004
Laurentgourretia Sakai, 2004: 557.—Sakai, 2005b: 224.—Sakai,
2011: 515.—Sakai, 2017a: 1126, 1130.
Type species. Laurentgourretia rhopalommata Sakai, 2004, by
original designation and monotypy.
Diagnosis. Rostrum sharply triangular, produced to near apex of
ocular lobes; anterolateral spines prominent. Maxilliped 3
merus with 2 or more mesiodistal spines; exopod absent. Minor
cheliped merus lower margin smooth; propodus cylindrical,
slightly swollen at midpoint, about twice as wide as long. Male
pleopod 1 article 2 simple, tapering, without coupling hooks.
Telson mostly parallel-sided, with rounded posterolateral corners.
Callianassidae and related families
Remarks. The only species differs from other ctenochelids in
the parallel-sided telson (fig. 18h) and having three meral
spines on the maxilliped 3. Its chelipeds are unknown.
The only specimen (MNHN-IU-2014-11417) of the type
species is a male with male-like pleopods and a gonopore on
the coxa of pereopod 5 (figs 17k–m; GCBP observations), not
a female as stated by Sakai (2004). The male pleopod 1 has
only one weak suture, the second article half as long as the
first and with eight marginal setae; the male pleopod 2 is
essentially as in other ctenochelids. Its major cheliped is
missing. Sakai (2004) diagnosed Laurentgourretia,
distinguishing it from Gourretia, as having a “stick-like”
eyestalk, a remarkably sharp scaphocerite, and maxilliped 3
lacking an exopod. The eyestalks are more wedge-shaped in
our view (figs 16j, k), contiguous as in other ctenochelids but
not as tapering in dorsal view. The scaphocerite is similar to
that of Dawsonius, which also lacks an exopod on maxilliped
3. The anterolateral angles of the dorsal carapace are sharp, as
is the anterodorsal angle of the branchiostegite (fig. 16k). The
uropodal exopod has a distal notch as in Paragourretia but its
margin has stronger marginal spiniform setae (fig. 16f).
The genus is sister to Dawsonius in the morphology
phylogram; no molecular data are available.
Paragourretia Sakai, 2004
Paragourretia Sakai, 2004: 568–569.—Sakai, 2005b: 225.—
Sakai, 2011: 516.
Heterogourretia Sakai, 2017a: 1126, 1129–1130 (type species,
Gourretia aungtonyae Sakai, 2002, by original designation and
monotypy) syn. nov.
Tuerkaygourretia Sakai, 2017a: 1126, 1133 (type species,
Tuerkaygourretia galathea Sakai, 2017, by original designation and
monotypy) syn. nov.
Type species. Gourretia phuketensis Sakai, 2002, by original
designation and monotypy.
Diagnosis. Cardiac prominence absent (rarely), or present.
Rostrum sharply triangular, produced to near apex of ocular
lobes, or obsolete, or obtusely triangular, flat, not reaching
cornea. Pleomere 6 without or with (P. biffari only) lateral
projections. Maxilliped 3 with exopod. Male major cheliped
merus with straight or weakly convex blade on lower margin
bearing a short proximal curved spine and 1 or more distal
spines; propodus rectangular, fingers tapering, fine, fixed
finger smooth. Minor cheliped merus lower margin with erect
proximal spine (as in major cheliped); propodus tapering, about
3 times as long as wide; fingers evenly tapering, curved
distally, of equal lengths, smooth or with obscure denticles
along opposing edges. Pereopod 3 propodus linear, longer
than wide, without proximal lobe on lower margin, lower
margin straight, upper margin moderately convex. Male
pleopod 1 article 2 with sharp distomesial apex, distal notch,
and sharp curved distolateral apex. Telson parallel-sided over
anterior half, then tapering to evenly rounded apex.
Remarks. Paragourretia differs from Gourretia in having fingers
lacking the sharp spines seen in Gourretia and curving distally,
the fixed finger of the major cheliped evenly tapering and curving,
121
and the appendices on the male pleopod 2 being midmesial, far
from the apex of the endopod rather than distal. None of these
characters was mentioned by Sakai (2011) although his allocation
of species between the two genera is identical to ours. None of the
specific diagnoses of members of this genus and of Gourretia in
Sakai (2011) includes other than an idiosyncratic assemblage of
family and generic features and none that tell one species from
another. Paragourretia biffari shares with Dawsonius latispina
lateral projections on pleomere 6.
Sakai (2004) likened Paragourretia to Dawsonius, not
Gourretia, because of a thick carapace and a cardiac
prominence. Re-examination of the type species failed to
convince GCBP that the carapace differed from other species
of Gourretia in this regard. A transverse cardiac sulcus was
difficult to decipher, and a transverse row of setae on the
anterior branchial region said to also characterise the genus is
found in all ctenochelid species examined. Sakai (2011)
changed the emphasis of his re-diagnosis, this time relying on
the presence of a notch on the uropodal exopod and on simple
versus subchelate pereopod 4 (it is simple in both genera). This
notch might be a precursor to a “dorsal plate”, or as in some
species, an elevated anterior margin over the posterior part of
the exopod, the two separated by transverse furrow. While the
notch is clear and unmissable in some species, in others it is
more ambiguous. The notch is a vague concavity in the type
species P. phuketensis and absent in P. lahouensis (which
Sakai curiously included in his key to species, identifiable as
“without lateral notch”). The notch is absent in all species of
Gourretia. Paragourretia was synonymised with Gourretia
by Sakai (2017b) because they have similar male pleopods 1
and 2. In fact, in species of Paragourretia the appendices on
the midmesial margin do not reach the end of the endopod
whereas in Gourretia s.s. they do, as far as is known.
Gourretia aungtonyae was provisionally included in
Paragourretia by Sakai (2005b) and made type species of a
new genus Heterogourretia (Sakai, 2017a) because of the
absence of male pleopod 1 and a distal “segmented” appendix
interna on pleopod 2. The holotype male has carapace length
of 2.9 mm and an ovigerous female 3.6 mm. The small size of
the male suggests it is immature and lacks male pleopod 1 for
this reason. The figured appendix interna is atypical of
axiideans and possibly an artefact. In all other features, the
species is typical of Paragourretia. For these reasons,
Heterogourretia is synonymised with Paragourretia.
The type species of Tuerkaygourretia, T. galathea Sakai,
2017, was based on a holotype female from Tranquebar, SE India
(Galathea stn 290), two paratypes from the same station, two
females from the Bay of Bengal, India (Galathea stn 304), and
one male from Singapore (Galathea stn 357). The figures of the
holotype (Sakai, 2017a: figs 4A, B, C, E, F, 5A–C, E, G) are of a
species of Paragourretia showing the minor cheliped without
teeth, uropodal exopod with a notch and pereopod 3 with a
narrow propodus having the lower margin straight. Other figures
are problematic, somewhat undermining Sakai’s (2011) argument
for new genera based on differences in male pleopods. Fig. 5I of
pleopod 2 (said to be from the type locality) and fig. 5D of a
broad pereopod 3 (from the Thai-Danish Expedition stn 1160 but
not otherwise listed) are of a species of Gourretia. Other figures
122
from the Thai-Danish Expedition stn 1160 are fig. 5H of pleopod
2 of a species of Gourretia (appendices terminal) and figs 5J, K
of a species of Paragourretia (appendices mesial). Sakai (2017a)
attributed differences between these two forms to size. It is
apparent that more than one species, from both Gourretia and
Paragourretia, have been confounded in this description. Sakai
(2017a) did not explain how these pleopodal forms differed from
those of other genera. To add further confusion, the species’
distribution is given as “Off Tranquebar, India (type locality); W.
of Subitana, Port Moresby, Papua New Guinea; West Malay
Peninsula; Singapore Island” but material is listed for only the
first and last of these four; the Thai-Danish Expedition stn 1160
would appear to be on the West Malay Peninsula.
Tuerkaygourretia is synonymised with Paragourretia on the
basis of probable illustrations of the holotype and paratype.
Eucalliacidae Manning and Felder, 1991
Figures 18, 19
Eucalliinae Manning and Felder, 1991: 781 (misspelling).
Eucalliacinae Sakai, 1999a: 108–109.—Ngoc-Ho, 2003: 487–489
(partim).—Sakai, 2005b: 195–196 (partim).—Sakai, 2011: 493–494.—
Hyžný, 2012: 51–52.—Sakai, 2017a: 1122.—Sakai, 2018: 734–738.
Eucalliacidae.—Sakai, 2011: 491.—Sakai and Türkay, 2014: 190
(outdated key to genera).—Sakai, 2017a: 1122.—Dworschak, 2018:
17.—Sakai, 2018: 734–738.
Diagnosis. Rostrum flat, short, triangular, shorter than eyestalks;
median carina absent; gastric carinae absent; cervical groove
well defined; suture between ocular lobe and end of linea
thalassinica horizontal in lateral view; anterior branchiostegal
margin sinusoidal or semicircular; anterior branchiostegal lobe
simple, scarcely calcified, merging smoothly with anterodorsal
branchiostegal angle and anterolateral margin of carapace;
posterior margin of carapace without lateral lobes. Pleomere 1
without anterolateral lobes, weakly chitinised. Eyestalks
flattened, contiguous, with subdistal dorsal cornea. Antennal
scaphocerite rudimentary. Maxilla scaphognathite without long
seta on posterior lobe extending into branchial chamber.
Maxilliped 1 epipod with acute anterior lobe lying alongside
exopod. Maxilliped 3 propodus about as wide as long; dactylus
dilating, truncate, with dense field of setae on distal margin.
Cheliped merus lower margin smooth; major cheliped with
distinctively flattened palm, sometimes with strong crest above
and below. Pereopod 3 propodus broad, with proximal lobe on
lower margin, without distal spiniform setae on lateral face
(often with 1 distal spiniform seta on lower margin). Pereopod 5
minutely chelate or subchelate. Female pleopod 2 rami narrower,
with more reduced setation than pleopods 3–5; endopod flattened
and 2–5 times as long as wide. Pleopods 3–5 with oblique
peduncles meeting mesially, endopods triangular, with straight
mesial margin, exopods attached laterally, proximally lobed,
longer than and enclosing endopods; appendices internae
reduced and almost embedded in mesial margin of endopod.
Uropodal exopod with elevated dorsal plate.
Remarks. Eucalliacidae differ from Callianopsidae, the only
other family with a dense field of setae on the distal margin of
a truncate maxilliped 3 dactylus, in the propodus of maxilliped
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
3 being wider than long (rectangular in callianopsids), lacking
a lateral longitudinal ridge on the eyestalk (present in
callianopsids) and the uropodal exopod having a dorsal plate
(absent in callianopsids).
The family and generic concepts used in redefining
Eucalliacidae with the support of molecular data, morphological
data or both (Robles et al., in press), differ markedly from those
tabulated by previous authors including Sakai and Türkay
(2014) and Sakai (2018 and his earlier works). Some characters
used by previous authors are too variable or not able to be
easily categorised for generic diagnosis: anterolateral
projections on the carapace, structure of the male pleopod 1,
size of appendices internae on pleopods 3–5, relative length of
the uropodal endopod, and concavity or convexity of the telson
margin. Several of the characters used in this work to separate
genera were not considered by these authors.
The family Eucalliacidae has a complex taxonomic history.
Manning and Felder (1991) erected the subfamily for two
genera, Eucalliax and Calliax. Ngoc-Ho (2003) erected a new
genus Calliaxina and differentiated these three genera from
Paraglypturus Türkay and Sakai, 1995. Sakai (1999a, 2005b)
recognised only Calliax and Paraglypturus, synonymising
Eucalliax and Calliaxina with Calliax.
Sakai (2011) divided Eucalliacidae into two subfamilies,
Eucalliacinae and Calliapaguropinae, the latter for a single
genus, Calliapagurops de Saint Laurent, 1973. Sakai’s remarks
on the new subfamily pointed out differences between
Calliapagurops and Callichirus (a member of another
subfamily) but nothing on why the genus was similar to
remaining eucalliacids. It is not at all similar. As stated much
earlier by Ngoc-Ho (2003), Calliapagurops is clearly a
member of Callichirinae, here treated as Callichiridae, with
which it shares many features. As a result, Sakai’s diagnosis of
the family comprised only characters true for all or most
callianassoids (scaphognathite without long distal seta;
pereopod 1 chelate, unequal, dissimilar; uropodal exopod with
dorsal plate but without notch) or ambiguous (carapace with or
without dorsal oval, maxilliped 3 dactylus “subtriangular” or
digitiform). Sakai’s (2011) Eucalliacinae revived the genera he
had earlier dismissed and added three to bring the total to
seven. He re-diagnosed Calliaxina using one character alone,
not one of those proposed by Ngoc-Ho (2003), but his
arguments were not supported by recent molecular or
morphological analysis (Robles et al., in press). In this same
paper, Sakai corrected his earlier statement (2005) that Calliax
was the type genus but his reasoning was false. Sakai and
Türkay (2014) added an eighth genus and tabulated characters
that they believed distinguished them.
Sakai (2018) re-diagnosed Eucalliacinae without
emphasising the unifying propodus and dactylus of maxilliped
3. He erected three new genera, Bakercalliax, Heardcalliax
and Manningcalliax, here treated as junior synonyms of
Eucalliaxiopsis. He argued heavily on separation of the genera
based on “organs involved in reproduction”, especially male
pleopods, whether pleopod 1 was “simple” or “subchelate”,
and on differences in the degree of fusion of the appendices
masculina and interna. Pleopod 1 of the male varies
considerably and erratically among species of Eucalliacidae
Callianassidae and related families
123
Figure 18. Diagnostic characters for genera of Eucalliacidae. Posterior carapace, sternite 7, pleopod1, coxa 4, basis of pereopod 5: a, Pseudocalliax.
Carapace, eyestalks, antennules, antennae: b, Eucalliax. Telson, uropod: c, Calliaxina; d, Eucalliaxiopsis. Pleomere 6, telson, uropod: e,
Paraglypturus; f, Eucalliax. Major cheliped: g, Paraglypturus; h, Eucalliax. Minor cheliped: i, Paraglypturus; j, Calliax; k, Pseudocalliax.
Pereopod 3: k, Calliaxina; l, Calliax. Pereopod 4: m, Paraglypturus.
Original illustrations: a, Pseudocalliax tooradin NMV J303; e, Calliaxina SA-01, UF 36699; h, Eucalliax quadracuta, Panama, NHMW 25916.
124
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Figure 19. Diagnostic characters for genera of Eucalliacidae. Maxilliped 3: a, Calliaxina punica; b, C. SA-01; c, Andamancalliax; d, Eucalliaxiopsis;
e, Pseudocalliax; f, Calliax. Male pleopod 1: g, h, Calliaxina bulimba; i, C. kensleyi; j, C. SA-01; k, C. punica; l, C. sakaii; m, Eucalliaxiopsis
panglaoensis; n, E. mcilhennyi; o, p, E. inaequimana; q, Calliax; r, Pseudocalliax; s, Paraglypturus. Male pleopod 2: t, Calliaxina; u, v,
Paraglypturus; w, Eucalliax; x, Eucalliaxiopsis. Female pleopod 2, y, Paraglypturus: z, Calliax; z’, Calliaxina. Female pleopod 3: z’’, Paraglypturus.
Original illustrations: b, i, Calliaxina SA-01, UF 36699; g, C. bulimba, MNHN-IU-2013-7097; h, C. bulimba, NMV J71686; w, Eucalliax
quadracuta, NHMW 25916; o, Eucalliaxiopsis inaequimana, MNHN-IU-2013-10008; p, E. inaequimana, UF 16512; e, r, Pseudocalliax
tooradin NMV J303; u, v, Paraglypturus calderus, MNHN Th1416.
125
Callianassidae and related families
and appears to have little generic value, contrary to Sakai’s
(2018) view (figs 19g–s).
Eucalliaxiopsis was differentiated from Eucalliax on
account of its unusual pleopods 1 and 2. The male pleopod 1
of the type species, Eucalliax cearaensis, bears long setae on
article 2 (Rodrigues and Manning, 1992), whereas in the type
species of Eucalliax this article is simple. We were unable to
confirm the structure of the male pleopod 1 of Eucalliax
cearaensis (type mislaid) but Rodrigues and Manning (1992b:
fig. 2s) can be variously interpreted. Apart from the long setae,
article 2 is similar to that of, for example, C. jonesi and E.
panglaoensis (cf. Dworschak, 2006). In most species of
Calliaxina and Eucalliaxiopsis, the male pleopod 1 has long
distal setae on article 1 and a broad article 2 and a curved
apex, sometimes with an intermediate lobe. This is seen in C.
novaebritanniae, C. kensleyi, C. punica, C. sakaii, E. jonesi,
E. mcilhennyi and E. panglaoensis (figs 19i–n; see also figures
in Dworschak, 2006, 2014; Heard, 1989; Felder and Manning,
1994; Manning and Lemaitre, 1994; Ngoc-Ho, 2003; de Saint
Laurent and Manning, 1982). In contrast, the illustrated male
pleopod 1 of C. bulimba (fig. 19g; Dworschak, 2005: figs 5c, d;
Poore and Griffin, 1979: fig. 21g), E. inaequimana (fig. 19o, p;
Dworschak, 2014: figs 4j–l) and E. madagassa (Ngoc-Ho,
2014: fig. 2S, as C. thomassini) has article 1 with distal long
setae and a short oblique article 2. However, a newly discovered
male of C. bulimba differs in having a bilobed article 2 with a
separate appendix interna (fig. 19h). This would appear to be a
precursor of the more typical eucalliacid form (figs 19i–n). A
third form of male pleopod 1 is seen in E. aequimana where
article 2 is longer than article 1, parallel-sided and setose
(Dworschak, 2014: figs 9h, i). These forms appear to cross
generic lines. The role of ontogeny in the morphology of
pleopod 1 is poorly understood but it is evident from some
studies that the male pleopod 1 does not appear fully formed
in juveniles (Dworschak, 2005, 2006, 2014). Development of
the male pleopod 1 with age seems probable.
This throws considerable doubt on the validity of three
recently erected genera Bakercalliax Sakai, 2018, Heardcalliax
Sakai, 2018 and Manningcalliax Sakai, 2018, all differentiated
largely on the basis of the male pleopod 1. Sakai (2018) also
noted differences between the relative sizes of the appendices
interna and masculina of pleopod 2 but this would scarcely
seem of generic value. The type species of all three genera
have a transverse ridge on the telson and lack an exopod on
maxilliped 3. As a consequence, all are synonymised with
Eucalliaxiopsis. Sakai (2011) did not compare Eucalliaxiopsis
with Calliaxina, which it resembles more closely, especially in
the male pleopod 2. Calliaxiopsis Sakai and Türkay, 2014,
was synonymised with Calliaxina by Poore and Dworschak
(2017). Sakai’s shuffling of species in his three papers has
resulted in some species being listed as members of as many
as five genera at different times during their history.
Our morphological analysis (Robles et al., in press) recognised
a single clade containing all eleven of the genera included by
Sakai (2018) in Eucalliacinae. This monophyly was not supported
by the molecular phylogram (Robles et al., in press) where three
sequential paraphyletic clades are evident, equivalent to
Eucalliaxiopsis+Calliaxina, Calliax and Eucalliax.
Key to genera of Eucalliacidae
1.
Major chela massive; minor chela with fixed finger acute,
shorter than dactylus (fig. 18j); pereopod 3 propodus oval,
little longer than wide, with short rounded proximal lobe on
lower margin, lower margin straight, upper margin strongly
convex (fig. 18l); female pleopod 2 without appendix
interna (fig. 19z); scaphocerite small, discoid
2
–
Both chelipeds swollen and similar, minor slightly smaller
if at all (figs 18g, h); pereopod 3 propodus pentagonal,
with strong broadly rounded proximal lobe on lower
margin, lower margin concave (fig. 18k); female pleopod
2 with (fig.18z) or without appendix interna; scaphocerite
absent or vestigial
3
2.
Sternite 7 and pereopod 5 coxa-ischium glabrous;
maxilliped 3 ischium linear, exopod absent (fig. 19f); minor
cheliped with tooth in gape between fingers (fig. 18k); male
pleopod 1 article 2 sickle-shaped (fig. 19q)
Calliax
–
Sternite 7 and pereopod 5 coxa-ischium densely setose (fig.
18a); maxilliped 3 ischium with strong proximal lobe on
lower margin, wider proximally than distally, exopod
present (fig. 19e); minor cheliped with fixed finger shorter
than dactylus (fig. 18j); male pleopod 1 stout, with apical
notch (fig. 19r)
Pseudocalliax (1 species, P. tooradin)
3.
Uropodal endopod triangular, anterior margin strongly
convex, posterior margin straight (fig. 18f); cheliped carpus
upper margin with 2 distal sharp spines (fig. 18h); male
pleopod 2 appendix masculina 4 times as long as wide,
attached near apex of endopod, appendix interna minute
(fig. 19w)
Eucalliax (1 species, E. quadracuta)
–
Uropodal endopod more or less ovate (figs 18c–e);
cheliped carpus upper margin without 2 distal sharp
spines; male pleopod 2 appendix masculina longer than
wide, attached mesially to endopod with appendix interna,
up to twice as long as endopod apex (figs 19t, x)
4
4.
Male pleopod 2 appendix masculina a lobe fused with
appendix interna, subdistal on endopod margin, endopod
broadly triangular (fig. 19u)
Paraglypturus
–
Male pleopod 2 appendix masculina a lobe fused with
appendix interna, attached midway on endopod margin,
endopod longer than wide (figs 19t, x)
5
5.
Telson with transverse ridge, lateral margins convex
(fig. 18d)
Eucalliaxiopsis
–
Telson without transverse ridge; lateral margins tapering
(fig. 18c)
6
6.
Maxilliped 3 ischium and merus length less than twice as
long as wide at their articulation, without proximal lobe,
with exopod (figs 19a, b)
Calliaxina
–
Maxilliped 3 ischium and merus length twice as long as
wide at their articulation; ischium with strong proximal lobe
on lower margin, wider proximally than distally (fig. 19c)
Andamancalliax (1 species, A. andamanica)
126
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Implicit attributes. Unless indicated otherwise, the following
attributes are implicit throughout the generic diagnoses. Anterior
dorsal carapace smooth; cardiac sulci absent. Sternite 7 and
pereopod 5, coxa-ischium glabrous. Pleomere 1 without sternal
plates. Maxilliped 3 ischium linear or wider distally than
proximally. Major cheliped broad, minor cheliped narrower, with
long fingers. Major cheliped carpus upper margin apex rounded
or square. Minor cheliped fingers closing along length. Male
pleopod 2 appendix masculina not reaching tip of endopod.
Uropodal endopod ovate, almost symmetrical, anterior margin
more convex than posterior margin. Telson smooth dorsally.
Andamancalliax Sakai, 2011
Andamancalliax Sakai, 2011: 494–495.—Sakai, 2018: 738.
Type species. Calliax andamanica Sakai, 2002, by original
designation and monotypy.
Diagnosis. Maxilliped 3 ischium and merus narrow, more
than twice as long as wide at their articulation; ischium with
strong proximal lobe on lower margin, wider proximally
than distally. Major cheliped broad, minor cheliped
narrower, with long fingers. Male pleopod 1 article 1 with
distal long setae, article 2 blade like; without appendix
interna. Male pleopod 2 appendix interna digitiform;
appendix masculina a lobe fused to appendix interna attached
midway on endopod margin. Telson widest anteriorly, more
or less semicircular.
Remarks. Andamancalliax was erected for a single species,
sharing with species of Calliaxina similar pereopod 3, pleopod
3, telson, uropod, and male and female pleopods 2. We have
assumed that Sakai’s (2002) figure 2B, labelled as from a
female, is of the simple male pleopod 1; Sakai (2002) reported
it absent but later (Sakai, 2005b, 2011) said it was present,
uniramous and with an emarginate tip. It would appear to be a
juvenile form. Sakai’s (2011) generic diagnosis contains
nothing that would distinguish the new genus from any other
eucalliacid—in fact it is wrong in stating that the male pleopod
2 lacks an appendix interna—it is clearly figured in Sakai’s
(2002) mislabelled figure 2A. He provided no justification for
the new genus; his key separated Andamancalliax with
emarginate distal male pleopod 1 article from Eucalliaxiopsis
where this article was said to incurve distally but the male
pleopod 1 of members of the latter varies markedly. Calliax
andamanica has a more acute rostrum than most species of
Calliaxina but among its members the rostrum ranges from
almost non-existent to sharply triangular (in C. sakaii, for
example). However, A. andamanica has a narrow maxilliped 3
with a proximal lobe (similar to that in Paraglypturus spp. and
Pseudocalliax tooradin) and asymmetrical chelipeds, quite
unlike other eucalliacids and the genus is justified. A new
species of this genus possesses a small exopod on maxilliped
3, not reported for A. andamanica (Poore, in press).
Calliax de Saint Laurent, 1973
Calliax de Saint Laurent, 1973: 514.—Manning, 1987: 397.—
Sakai, 1988: 61.—Manning and Felder, 1991: 783.—Sakai, 1999a:
109–110.—Ngoc-Ho, 2003: 489–490.—Sakai, 2005b: 196–197.—
Sakai, 2011: 495–496.—Hyžný and Gasparic, 2014: 42–45.—Sakai,
2018: 738.
Type species. Callianassa (Callichirus) lobata de Gaillande
and Lagardère, 1966, by original designation and monotypy.
Diagnosis. Maxilliped 3 ischium and merus narrow, more than
twice as long as wide at their articulation. Male major chela
with palm about twice as long as carpus, 1.5 times as long as
wide, minor chela half as wide, with short fingers. Minor
cheliped fingers with wide gape, with tooth at base of fingers,
fingers not closing along length. Male pleopod 1 article 2
curved, simply curving to sharp apex; without appendix
interna. Male pleopod 2 appendix interna absent; appendix
masculina a lobe fused to appendix interna attached midway
on endopod margin, reaching or exceeding tip of endopod.
Telson widest anteriorly, more or less semicircular.
Remarks. The wide gape between the fingers of the minor
cheliped, with an intermediate tooth, immediately defines
species of Calliax.
Sakai (1999a) treated Eucalliax as a junior synonym of
Calliax and later (2005b) added Calliaxina to this synonymy.
Ngoc-Ho (2003) clearly differentiated these three genera and
Paraglypturus. Sakai (2011) appears to have accepted NgocHo’s arguments and he confined Calliax to just two species;
one has been added since (Ngoc-Ho, 2014). Sakai’s (2011)
generic diagnosis contained several generalities referrable to
many callianassoids but apart from mention of “P1 unequal in
size and dissimilar in shape” acknowledges none of the
defining generic characters. His key would fail to discriminate
the genus as presently diagnosed. Sakai’s (2011) “diagnoses”
of the two species are largely replications of his inadequate
generic diagnosis, apart from subtle differences in the
description of the shapes of their telsons. The type species has
been illustrated most recently by Sakai (2017b) and García
Raso et al. (2019). Their records are from 622 m and 457–548
m depth respectively in the Mediterranean; previous records
are from only a few metres depth.
Calliaxina Ngoc-Ho, 2003
Calliaxina Ngoc-Ho, 2003: 493–494.—Sakai, 2011: 497–498.—
Hyžný, 2012: 55–56.—Sakai and Türkay, 2014: 191.—Ngoc-Ho, 2014:
549.—Poore and Dworschak, 2017: 120 (partim).—Sakai, 2018: 739–741.
Calliax.—Sakai, 2005b: 196 (partim, not Calliax de Saint
Laurent, 1973).
Calliamina Sakai and Türkay, 2014: 190 (misspelling).
Type species. Calliax punica de Saint Laurent and Manning,
1982, by original designation.
Diagnosis. Cardiac sulci present. Pleomere 1 with pair of
sternal plates anterior to pleopods 1. Maxilliped 3 ischium and
merus broad, less than twice as long as wide at their articulation.
Both chelipeds with palms as long as wide in both sexes, minor
slightly smaller if at all. Male pleopod 1 article 1 with distal
long setae; article 2 blade-like with bifid apex (sometimes
obscure), or article 2 much shorter, digitiform (sometimes
fused); with appendix interna (present at least as hooks on
127
Callianassidae and related families
endopod). Male pleopod 2 appendix interna digitiform;
appendix masculina a lobe fused to appendix interna attached
midway on endopod margin. Telson widest anteriorly, more or
less semicircular, curving to broad convex apex.
Remarks. Calliaxina differs from Eucalliaxiopsis in having a
tapering telson without a transverse ridge, and at least a vestige
of an appendix interna on pleopod 1 of the male. All species
have an exopod on maxilliped 3, sometimes vestigial, whereas
only one species of Eucalliaxiopsis does.
Calliaxina was erected by Ngoc-Ho (2003) for three species
previously included by Sakai (1999a) in Paraglypturus Türkay
and Sakai, 1995 and later (Sakai, 2005) in Calliax. Ngoc-Ho
(2003) tabulated differences between these genera but did not
include Eucalliax Manning and Felder, 1991, which she
diagnosed separately in her table. Her diagnosis relied
importantly on the presence of an exopod on maxilliped 3, equal
and similar chelipeds, appendix interna on pleopods 1 and 2 of
the male. Some of her key features are difficult to quantify,
notably the shape of the front and anterolateral projections of the
carapace, and length of the appendix interna on pleopods 3–5.
Sakai (2005) synonymised Calliaxina with Calliax without
discussing any morphological similarities or differences. Later,
Sakai (2011) revived Calliaxina which he had earlier
synonymised in new sense (sensu nov.) by including other species
with similar maxillipeds 3 and with one or two cardiac sulci,
although some lack maxillipedal 3 exopods. At the same time,
he erected a monotypic genus Eucalliaxiopsis Sakai, 2011,
relying on the possession of unique male pleopods (see below).
As already explained, Robles et al. (in press) divided the
species allocated by Sakai (2011) to Calliaxina and
Eucalliaxiopsis into two groups. Their molecular analysis
grouped C. bulimba, C. kensleyi, C. novaebritanniae and C.
sakaii into a one clade, and C. aequimana, C. inaequimana,
C. panglaoensis, C. jonesi and three others into a second
clade. The morphological treatment includes C. punica, type
species of Calliaxina in the first and C. cearaensis, type
species of Eucalliaxiopsis in the second. These two names are
applied to these clades.
Eucalliax Manning and Felder, 1991
Eucalliax Manning and Felder, 1991: 781–783 (partim).—NgocHo, 2003: 489–490 (partim).—Sakai, 2011: 502–503 (partim).—
Hyžný and Hudáčková, 2012: 12–14 (partim).—Sakai, 2017a: 1122
(partim).—Sakai, 2018: 742.
Type species. Callianassa quadracuta Biffar, 1970, by original
designation.
Diagnosis. Anterior dorsal carapace with median ridge
posterior to rostrum and submedian pair of oblique ridges.
Maxilliped 3 ischium and merus less than twice as long as wide
at their articulation. Male major cheliped with palm about
twice as long as carpus, 1.5 times as long as wide; minor chela
slightly smaller, with elongate fingers; major cheliped carpus
upper margin with 2 distal sharp spines in both sexes. Male
pleopod 1 article 1 with distal long setae; article 2 blade like,
with bifid apex, sometimes obscure; without appendix interna.
Male pleopod 2 appendix interna minute; appendix masculina
4 times as long as wide, attached near apex of endopod
reaching, or exceeding tip of endopod. Uropodal endopod
triangular, anterior margin strongly convex, posterior
margin straight. Telson convex-sided, widest near midpoint,
posterior margin slightly excavate; with transverse crest.
Remarks. The most obvious characters distinguishing the only
species, Eucalliax quadracuta, from other eucalliacids is the
triangular uropodal endopod and the male pleopod 2 with its
appendix masculina well exceeding the endopod. Re-examination
by us (PCD) showed the presence of a minute appendix interna
(fig. 19w); Sakai’s figure (1999a; fig. 30c) is misleading.
Eucalliax was differentiated initially from Calliax simply
“in having the chelipeds equal and similar”, the two genera
being placed in the same new subfamily (Manning and Felder,
1991). The genus was treated as a synonym of Calliax by Sakai
(1999a, 2005b) without justification but revived later (Sakai,
2011). Sakai (2017a) differentiated Eucalliax from
Eucalliaxiopsis Sakai, 2011 on differences in male pleopods 1
and 2 but not from other genera.
Eucalliaxiopsis Sakai, 2011
Calliaxina.—Ngoc-Ho, 2003: 493–494 (partim).—Sakai, 2011:
497–498 (partim).—Sakai and Türkay, 2014: 191 (partim).—Ngoc-Ho,
2014: 549 (partim).—Poore and Dworschak, 2017: 120 (partim).
Eucalliaxiopsis Sakai, 2011: 503–504.—Sakai, 2018: 742–743.
Calliaxiopsis Sakai and Türkay, 2014: 192.—Sakai, 2018: 741–
742 (type species, Calliaxiopsis madagassa Sakai and Türkay, 2014,
by original designation and monotypy) syn. nov.
Bakercalliax Sakai, 2018: 738 (type species, Callianassa
aequimana Baker, 1907, by original designation) syn. nov.
Heardcalliax Sakai, 2018: 743 (type species, Calliax jonesi
Heard, 1989, by original designation and monotypy) syn. nov.
Manningcalliax Sakai, 2018: 743 (type species, Eucalliax
mcilhennyi Felder and Manning, 1994, by original designation)
syn. nov.
Type species. Eucalliax cearaensis Rodrigues and Manning,
1992, by original designation and monotypy.
Diagnosis. Anterior dorsal carapace smooth, or with median
ridge posterior to rostrum and submedian pair of oblique
ridges; cardiac sulci present. Pleomere 1 without sternal
plates, or with pair of sternal plates anterior to pleopods 1.
Maxilliped 3 ischium and merus less than twice as long as wide
at their articulation. Male major cheliped with palm about
twice as long as carpus, 1.5 times as long as wide; minor chela
slightly smaller, with elongate fingers or both chelipeds with
palms as long as wide in both sexes, minor slightly smaller if at
all (rarely). Male pleopod 1 article 2 longer than article 1,
parallel-sided, setose, or article 1 with distal long setae, article
2 blade like, with bifid apex (sometimes obscure), or article 1
with distal long setae, article 2 much shorter, digitiform
(sometimes fused); without appendix interna. Male pleopod 2
appendix interna digitiform; appendix masculina a lobe fused
to appendix interna attached midway on endopodal margin,
reaching or exceeding tip of endopod. Telson convex-sided,
widest near midpoint; posterior margin slightly excavate or
obscurely excavate between rounded posterolateral corners;
with transverse crest on dorsal surface.
128
Remarks. Eucalliaxiopsis is most easily recognised by the
telson having a transverse ridge and convex lateral margins.
The male pleopod 1 lacks an appendix interna. The appendix
masculina reaches or exceeds the tip of the endopod of pleopod
2. Eucalliaxiopsis shares with Eucalliax absence of a
maxilliped 3 exopod (with one exception, E. madagassa) and a
telson widest near its midpoint and having a sharp transverse
crest. Eucalliaxiopsis differs in having an oval rather than
rhomboid uropodal endopod. The presence of cardiac sulci is
assumed for the type species; it is present in all others
illustrated. The generic synonymy is discussed above under the
family heading.
Paraglypturus Türkay and Sakai, 1995
Paraglypturus Türkay and Sakai, 1995: 26–27.—Sakai, 1999a:
122.—Sakai, 2005b: 204–205.—Sakai, 2011: 504–505.—Sakai, 2018:
744.
Type species. Paraglypturus calderus Türkay and Sakai, 1995,
by original designation and monotypy.
Diagnosis. Maxilliped 3 ischium and merus more than twice as
long as wide at their articulation; ischium with strong proximal
lobe on lower margin, wider proximally than distally. Male
major cheliped with palm about twice as long as carpus, 1.5
times as long as wide, minor chela slightly smaller, with
elongate fingers. Male pleopod 1 article 1 with distal long
setae, article 2 blade like, with bifid apex, sometimes obscure;
with appendix interna (at least with hooks). Male pleopod 2
appendix interna digitiform; appendix masculina a lobe fused
with appendix interna, subdistal on endopodal margin.
Telson widest anteriorly, more or less semicircular.
Remarks. Paraglypturus differs from other Eucalliacidae in
having the pleopod 2 endopod almost as wide as that of pleopod
3 (cf. figs 19u, z’’). The second article of the pleopod 1 of the
male has a minute appendix interna with hooks (fig. 19s). The
species share with many eucalliacids two long setae distally on
the upper margin of the propodus of pereopod 4 (fig. 18m) but
differ in the arrangement of the appendices on the male pleopod
2. The chelipeds are swollen and similar, the minor slightly
smaller (figs 18g, i).
Robles et al. (in press) found weak molecular support for
one of its two species to be more closely related to
Callianopsidae than to any other callianassoids despite the
morphological evidence supporting eucalliacid affinities
(Sakai (2005b, 2011, 2018). Paraglypturus was said by Türkay
and Sakai (1995) to be close to Glypturus but this was realised
to be wrong when Sakai (1999a) noted its similarity to Calliax
and placed both genera in Eucalliacinae. He differentiated the
two on the possession in Paraglypturus (in which he included
five species) and absence in Calliax (seven species) of an
exopod on maxilliped 3. Later, Sakai (2005b, 2011) restricted
the genus to its type species because it alone possesses the
“yellow transparent circular structure on the uropodal
endopod”. Türkay and Sakai (1995) described and figured a
“rounded yellow-transparent circular structure centrally” on
the upper surface of the uropodal exopod of P. calderus. This
was confirmed by PCD on a paratype (SMF 22951) but
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
subsequent examination by GCBP in 2008 and in 2018 of
another specimen (MNHN Th1416) failed to distinguish the
structure. Ahn et al. (2017) figured in colour a yellow patch on
the upper surface of the uropodal endopod of P. tonganus.
Pseudocalliax Sakai, 2011
Paraglypturus.—Poore, 2004: 184.
Pseudocalliax Sakai, 2011: 505–506.—Sakai, 2018: 744.
Type species. Callianassa tooradin Poore and Griffin, 1979, by
original designation and monotypy.
Diagnosis. Sternite 7 and pereopod 5 coxa-ischium densely
setose. Maxilliped 3 ischium and merus more than twice as long
as wide at their articulation; ischium with strong proximal lobe
on lower margin, wider proximally than distally. Male major
cheliped with palm about twice as long as carpus, 1.5 times as
long as wide, minor chela half as wide, with short fingers. Minor
cheliped fingers with wide gape, without tooth at base of fingers,
fingers not closing along length. Male pleopod 1 stout, article 1
setose, article 2 with shallow apical notch; without appendix
interna. Male pleopod 2 appendix interna digitiform; appendix
masculina not differentiated from endopod apex. Telson widest
anteriorly, more or less semicircular.
Remarks. The densely setose sternite 7 and coxa-ischium of
pereopod 5 and the stout male pleopod 1 diagnose the only
species, P. tooradin Poore and Griffin, 1979.
Following “examination of the male specimen [of C.
tooradin] preserved in the USNM”, Sakai (2011) diagnosed
Pseudocalliax as a genus of Eucalliacidae possessing a dorsal
oval and cardiac sulcus but we found no material in the USNM.
We have confirmed by examining types and other specimens
(NMV J301–J303, J59670–J59672) that the species has neither
feature, as was confirmed by Sakai earlier (1999a: figs 33a–c).
None of his other generic characters is useful. The male
pleopod 1 of P. tooradin is not “small and simple” as stated by
Sakai (2011) but stout, with article 1 setose and article 2 with
shallow apical notch (fig. 19r). The species was inadequately
described by Poore and Griffin (1979).
Paracalliacidae Sakai, 2005
Figure 20
Paracalliacinae Sakai, 2005b: 215.
Paracalliacidae.—Dworschak and Poore, 2018: 70.
Diagnosis. Rostrum flat, short, triangular, shorter than
eyestalks; median carina on rostrum only; submedian gastric
carinae absent; cervical groove well defined; suture between
ocular lobe and end of linea thalassinica horizontal in lateral
view; anterior branchiostegal margin sinusoidal or semicircular;
anterior branchiostegal lobe simple, scarcely calcified, merging
smoothly with anterodorsal branchiostegal angle and
anterolateral margin of carapace; posterior margin of carapace
with lateral lobes interacting with anterolateral lobes on
pleomere 1. Eyestalks flattened, contiguous, with subdistal
dorsal cornea. Antennal scaphocerite simple, triangular.
Maxilla scaphognathite without long seta on posterior lobe
129
Callianassidae and related families
Figure 20. Diagnostic characters for Paracalliacidae, Paracalliax bollorei: a, posterior carapace, pleomeres 1, 2; b, telson, uropod; c, maxilliped
3; d, e, female pleopods 1, 2.
extending into branchial chamber. Maxilliped 1 epipod with
acute anterior lobe lying alongside exopod. Maxilliped 3
propodus longer than wide, not prominently lobed on lower
margin; dactylus slender, digitiform, with setae irregularly
spaced along all margins. Cheliped merus lower margin
spinose; major cheliped palm oval in cross-section, barely
crested above or below. Pereopod 3 propodus elongate-oval,
tapering, without proximal lobe on lower margin, without
distal spiniform setae on lateral face (often with 1 distal
spiniform seta on lower margin). Pereopod 5 minutely chelate
or subchelate. Female pleopod 2 rami similar to following
pleopods, with regularly setose margins. Pleopods 3–5 with
oblique peduncles meeting mesially, endopods triangular, with
straight mesial margin, exopods attached laterally, proximally
lobed, longer than and enclosing endopods; appendices internae
elongate, much longer than wide. Uropodal exopod without
elevated dorsal plate.
Remarks. The only species differs from all other callianassoids
in having pleomere 1 with dorsolateral lobes interacting and
overlapping the posterolateral margin of the carapace (fig. 20a;
Dworschak and Poore, 2018; Robles et al., in press). Pleopod 2
is similar to pleopods 3–5 (figs 20d, e). The uropodal exopod
lacks a dorsal plate (fig. 20b) and maxilliped 3 is exceptionally
narrow (fig. 20c). The male is not known.
Manning and Felder (1991) included the only genus
Paracalliax in Ctenochelidae. The subfamily was erected as
member of Callianassidae by Sakai (2005b) who was uncertain
about its affinities (not in Gourretiidae as he asserted later
[Sakai, 2011]). The family was ignored by Sakai (2011) and
Sakai et al. (2015) who treated the genus as part of Gourretiidae
despite treating most other basal groups as distinct families.
Sakai (2017a) also included the genus in Gourretiidae but
considered it a possible synonym of one of three new gourretiid
genera erected at the time.
Paracalliax de Saint Laurent, 1979
Paracalliax de Saint Laurent, 1979a: 1396.—de Saint Laurent and
Le Loeuff, 1979: 84–86.—Manning and Felder, 1991: 785.—Sakai,
2005b: 215.—Sakai, 2011: 515.—Sakai et al., 2015: 134.—Sakai,
2017a: 1126, 1131.
Type species. Paracalliax bollorei de Saint Laurent, 1979, by
original designation and monotypy.
Diagnosis. With characters of the family.
Remarks. Sakai et al. (2015) described a second species,
Paracalliax stenophthalma, from the southern Banc d’Arguin,
the same locality as the holotype of the type species. They are
synonymous (Dworschak and Poore, 2018).
Acknowledgements
GCBP thanks coauthor Darryl Felder for inviting his
participation in this endeavour, part of the Assembling the
Tree of Life (AToL) Decapoda research project.
We particularly thank individuals from two institutions for
the loan of material, access to collections: Laure Corbari,
Paula Martin-Lèvre and Anouchka Krygelmans, Muséum
national d’Histoire naturelle, Paris (MNHN), for loan of
numerous specimens from Papua New Guinea and other
locations; and Gustav Paulay and Amanda Bemis, Florida
Museum of Natural History, Gainsville, for loan of numerous
specimens from the Indo-West Pacific. We thank too the
following individuals and institutions for the loan of type
material: Karen Reed, US Museum of Natural History,
Washington; the late Michael Türkay, Senckenberg Museum,
Frankfurt; Charatsee Aungtonya, Phuket Marine Biological
Center, Phuket; Gavin Dally, Northern Territory Museum and
Art Gallery, Darwin; Stephen Keable, Australian Museum,
Sydney; Peter Davie, Queensland Museum, Brisbane.
130
GCBP is grateful to the Australian Biological Resources
Study National Taxonomy Research Grant Program (NTRGP),
to Phillipe Bouchet and the Alain Crosnier Fund (MNHN) for
financial support while in Paris, and the Naturhistorisches
Museum, Vienna, for support in Vienna.
This study is part of the multidisciplinary and collaborative
research project, with financial support by the Fundação de
Amparo a Pesquisa do Estado de São Paulo (FAPESP; Tematico
Biota 2010/50188-8; PROTAX 2016/50376-5), Conselho
Nacional de Desenvolvimento Científico e Tecnológico (CNPq;
PQ 304968/2014–5; PROTAX 440417/2015-5), and Coordenação
de Aperfeiçoamento de Pessoal de Nível Superior (CAPES;
Ciências do Mar II, 2005/2014–23038.004308/2014-14), granted
to FLM that made possible the formation of ideas, analyses of
the material and discussion of the obtained data by our team.
RR thanks the U.S. National Science Foundation for postdoctoral scholarship (DEB/AToL grant no. EF-0531603 under
DLF), and CNPq (500460/2010–8), FAPESP (2013/05663-8),
and CAPES for post-doctoral scholarships (under FLM), RR
also wishes to acknowledge PRODEP-SEP, Mexico, through
the program “Apoyo a la Incorporación de NPTC (Ago/1/2018
–Jul/31/2019)”.
DLF received funding from the U.S. National Science
Foundation (BS&I grant no. DEB-0315995 and DEB/AToL
grant no. EF-0531603), along with small travel grants from the
Smithsonian Marine Station–Fort Pierce, Florida and cruise
support from the Smithsonian Tropical Research Institute,
Panama. This is contribution number 1116 from the Smithsonian
Marine Station—Fort Pierce, Florida and number 119 from the
University of Louisiana Laboratory for Crustacean Research.
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136
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Table 1. Accepted names of 265 species of callianassoids listed alphabetically by species name, with family allocation and accepted generic and
species combination. 87 are in new combinations. Commonly accepted junior synonyms are not included (see WoRMS Editorial Board 2018).
Eleven species of Callianassidae inadequately described or figured are placed Incertae sedis. See Table 2 for a list of the same accepted species
arranged by family and genus and comments on those Incertae sedis.
Species, Genus
Family allocation
Current combination, authority
abyssa, Lipkecallianassa
acanthochirus, Glypturus
acanthura, Callianassa
acutirostella, Callianassa
adamas, Callianassa
aequimana, Callianassa (Callichirus)
agassizi, Callianassa
almeidai, Ctenocheloides
amboinae, Scallasis
amboinensis, Callianassa
amplimaxilla, Callianassa
andamanica, Calliax
andamaniensis, Trypaea
angelikae, Neocallichirus
anoploura, Callianassa
anovalis, Callianopsis
aqabaensis, Callianassa
arenosa, Callianassa
argentinensis, Callianassa
arguinensis, Gilvossius
armata, Callianassa
articulata, Callianassa
arutyunovi, Vulcanocalliax
assimilis, Callianassa (Callichirus)
attenboroughi, Ctenocheloides
auchenorhynchus, Neocallichirus
audax, Callianassa
aungtonyae, Gourretia
australiensis, Trypaea
australis, subterranea, Callianassa
Balssi, Callianassa (Callichirus)
balssi, Ctenocheles
bangensis, Callianassa
barracuda, Gourretia
berylae, Necallianassa
bicauda, Notiax
biffari, Callianassa
biffari, Gourretia
biformis, Callianassa
bocourti, Callianassa
bollorei, Paracalliax
borradailei, longiventris, Callianassa
boucheti, Ctenocheloides
Bouvieri, Callianassa
brachyophthalma, Callianassa
brachytelson, Callianassa
brevirostris, Callianassa
Callianassidae
Callichiridae
Callianassidae
Callianassidae
Callichiridae
Eucalliacidae
Anacalliacidae
Ctenochelidae
Callianassidae
Callianassidae
Callianassidae
Eucalliacidae
Callianassidae
Callichiridae
Callianassidae
Callianopsidae
Callianassidae
Callianassidae
Anacalliacidae
Callianassidae
Callichiridae
Callichiridae
Callianopsidae
Callichiridae
Ctenochelidae
Callichiridae
Callichiridae
Ctenochelidae
Callianassidae
Callianassidae
Callichiridae
Ctenochelidae
Callianassidae
Ctenochelidae
Callianassidae
Callianassidae
Callianassidae
Ctenochelidae
Callianassidae
Callichiridae
Paracalliacidae
Callichiridae
Ctenochelidae
Callianassidae
Callianassidae
Callianassidae
Callianassidae
Lipkecallianassa abyssa Sakai, 2002
Glypturus acanthochirus Stimpson, 1866
Necallianassa acanthura (Caroli, 1946)
Spinicallianassa acutirostella (Sakai, 1988) comb. nov.
Callichirus adamas (Kensley, 1974)
Eucalliaxiopsis aequimana (Baker, 1907) comb. nov.
Anacalliax agassizi (Biffar, 1971)
Ctenocheloides almeidai Anker & Pachelle, 2013
Scallasis amboinae Bate, 1888
Rayllianassa amboinensis (De Man, 1888)
Aqaballianassa amplimaxilla (Sakai, 2002) comb. nov.
Andamancalliax andamanica (Sakai, 2002)
Scallasis andamaniensis (Sakai, 2010) comb. nov.
Neocallichirus angelikae Sakai, 2000
Incertae sedis anoploura Sakai, 2002
Callianopsis anovalis Lin, Komai & Chan, 2007
Aqaballianassa aqabaensis (Dworschak, 2003) comb. nov.
Arenallianassa arenosa (Poore, 1975) comb. nov.
Anacalliax argentinensis (Biffar, 1971)
Gilvossius arguinensis Sakai, Türkay, Beuck & Freiwald, 2015
Glypturus armatus (A.Milne-Edwards, 1870)
Corallianassa articulata (Rathbun, 1906)
Vulcanocalliax arutyunovi Dworschak & Cunha, 2007
Corallianassa assimilis (De Man, 1928)
Ctenocheloides attenboroughi Anker, 2010
Neocallichirus auchenorhynchus Sakai, 2005
Audacallichirus audax (De Man, 1911) comb. nov.
Paragourretia aungtonyae (Sakai, 2002)
Trypaea australiensis Dana, 1852
Callianassa australis Kensley, 1974
Balsscallichirus balssi (Monod, 1933)
Ctenocheles balssi Kishinouye, 1926
Rayllianassa bangensis (Sakai, 2005) comb. nov.
Gourretia barracuda LeLoeuff & Intès, 1974
Necallianassa berylae Heard & Manning, 1998
Pugnatrypaea bicauda (Sakai, 2010) comb. nov.
Neotrypaea biffari (Holthuis, 1991)
Paragourretia biffari (Blanco-Rambla & Liñero-Arana, 1994)
Biffarius biformis (Biffar, 1971)
Lepidophthalmus bocourti (A. Milne-Edwards, 1870)
Paracalliax bollorei de Saint Laurent, 1979
Corallianassa borradailei (De Man, 1928)
Ctenocheloides boucheti Poore, 2015
Paratrypaea bouvieri (Nobili, 1904)
Notiax brachyophthalma (A.Milne-Edwards, 1870)
Cheramoides brachytelson (Sakai, 2002) comb. nov.
Aqaballianassa brevirostris (Sakai, 2002) comb. nov.
137
Callianassidae and related families
Species, Genus
Family allocation
Current combination, authority
bulimba, Callianassa
cacahuate, Neocallichirus
caechabitator, Neocallichirus
caecigena, Callianassa
caesari, Pseudobiffarius
calderus, Paraglypturus
caledonica, Callianassa
californiensis, Callianassa
Calmani, Callianassa
candidus, Cancer
cavifrons, Cheramus
cearaensis, Eucalliax
ceramica, Callianassa
chakratongae, Callianassa
charcoti, Calliapagurops
collaroy, Callianassa
collini, Ctenocheles
contipes, Callianassa
convexa, Callianassa
coolibah, Gourretia
coriolisae, Callianassa
costaricensis, Callianassa
Coutierei, Callianassa
crosnieri, Gourretia
darwinensis, Neocallichirus
darvishi, Neocallichirus
debilis, Biffarius
delicatulus, Biffarius
denticulata, Callianassa
diaphora, Callianassa
doerjesti, Calliax
ehsani, Callianassa
Eiseni, Lepidophthalmus
exilimaxilla, Callianassa
filholi, Callianassa
foresti, Callichirus
foresti, Calliapagurops
fragilis, Callianassa
frouini, Neocallichirus
galathea, Tuerkaygourretia
garthi, Callianassa
gaucho, Poti
geomar, Bathycalliax
gigas, Callianassa
gilchristi, Callianassa
goniophthalma, Callianassa
grandidieri, Callianassa
grandimana, Callianassa
grandis, Neocallichirus
gruneri, Callianassa
guaiqueri, Sergio
Eucalliacidae
Callichiridae
Callichiridae
Callianopsidae
Callianassidae
Eucalliacidae
Callianassidae
Callianassidae
Callichiridae
Callianassidae
Callianassidae
Eucalliacidae
Callianassidae
Callianassidae
Callichiridae
Callichiridae
Ctenochelidae
Callianassidae
Callianassidae
Ctenochelidae
Callianassidae
Callianassidae
Callichiridae
Ctenochelidae
Callichiridae
Callichiridae
Callianassidae
Callianassidae
Ctenochelidae
Callianassidae
Eucalliacidae
Callianassidae
Callichiridae
Callianassidae
Callianassidae
Callichiridae
Callichiridae
Callianassidae
Callichiridae
Ctenochelidae
Callichiridae
Callianassidae
Callianopsidae
Callianassidae
Callichiridae
Callianopsidae
Callichiridae
Callichiridae
Callichiridae
Callianassidae
Callichiridae
Calliaxina bulimba (Poore & Griffin, 1979)
Neocallichirus cacahuate Felder & Manning, 1995
Neocallichirus caechabitator Sakai, 1988
Callianopsis caecigena (Alcock & Anderson, 1894)
Neotrypaea caesari (Heard & Manning, 2000) comb. nov.
Paraglypturus calderus Türkay & Sakai, 1995
Scallasis caledonica (Ngoc-Ho, 1991) comb. nov.
Neotrypaea californiensis (Dana, 1854)
Neocallichirus calmani (Nobili, 1904)
Gilvossius candidus (Olivi, 1792)
Caviallianassa cavifrons (Komai & Fujiwara, 2012) comb. nov.
Eucalliaxiopsis cearaensis (Rodrigues & Manning, 1992)
Filhollianassa ceramica (Fulton & Grant, 1906) comb. nov.
Incertae sedis chakratongae Sakai, 2002
Calliapagurops charcoti de Saint Laurent, 1973
Corallianassa collaroy (Poore & Griffin, 1979)
Ctenocheles collini Ward, 1945
Scallasis contipes (Sakai, 2002) comb. nov.
Gilvossius convexus (de Saint Laurent & LeLoeuff, 1979)
Paragourretia coolibah (Poore & Griffin, 1979)
Coriollianassa coriolisae (Ngoc-Ho, 2014) comb. nov.
Neotrypaea costaricensis (Sakai, 2005) comb. nov.
Corallianassa coutierei (Nobili, 1904)
Paragourretia crosnieri (Ngoc-Ho, 1991)
Neocallichirus darwinensis Sakai, 1988
Neocallichirus darvishi Sepahvand, Komai, Momtazi & Shahabi, 2018
Fragillianassa debilis (Hernandez-Aguilera, 1998) comb. nov.
Biffarius delicatulus Rodrigues & Manning, 1992
Gourretia denticulata (Lutze, 1937)
Callianassa diaphora LeLoeuff & Intes, 1974
Calliax doerjesti Sakai, 1999
Aqaballianassa ehsani (Sepahvand, Tudge & Momtazi, 2018) comb. nov.
Lepidophthalmus eiseni Holmes, 1904
Incertae sedis exilimaxilla Sakai, 2005
Filhollianassa filholi (A.Milne-Edwards, 1878) comb. nov.
Balsscallichirus foresti (LeLoeuff & Intès, 1974) comb. nov.
Calliapagurops foresti Ngoc-Ho, 2002
Fragillianassa fragilis (Biffar, 1970) comb. nov.
Neocallichirus frouini Ngoc-Ho, 2005
Paragourretia galathea (Sakai, 2017) comb. nov.
Callichirus garthi (Retamal, 1975)
Poti gaucho Rodrigues & Manning, 1992
Bathycalliax geomar Sakai & Türkay, 1999
Neotrypaea gigas (Dana, 1852)
Balsscallichirus gilchristi (Barnard, 1947)
Callianopsis goniophthalma (Rathbun, 1902)
Lepidophthalmus grandidieri (Coutière, 1899)
Neocallichirus grandimana (Gibbes, 1850)
Laticallichirus grandis (Karasawa & Goda, 1996)
Incertae sedis gruneri Sakai, 1999
Neocallichirus guaiqueri (Blanco-Rambla, Liñero-Arana & Beltràn Lares, 1995)
138
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Species, Genus
Family allocation
Current combination, authority
guara, Callianassa
guassutinga, Callianassa
guineensis, Callianassa
hainanensis, Nihonotrypaea
harmandi, Callianassa
hartmeyeri, Callianassa
haswelli, Callianassa
holthuisi, Ctenocheles
horneri, Neocallichirus
inaequimana, Eucalliax
indica, Michaelcallianassa
intermedia, Callianassa
intesi, Callichirus
iranicus, Cheramus
islagrande, Callianassa
jamaicense, Callianassa
japonica, subterranea, Callianassa
joculatrix, Callianassa
jonesi, Calliax
Jousseaumei, Callianassa
karumba, Callianassa
kensleyi, Eucalliax
kowalevski, Trypaea
kraussi, Callianassa
laevidactyla, Gourretia
lahouensis, Gourretia
lanceolata, Callianassa (Callichirus)
laresi, Gourretia
latispina, Callianassa
laurae, Callichirus
loeuffintesi, Gourretia
lemaitrei, Neocallichirus
leviceps, Ctenocheles
lewtonae, Callianassa
lignicola, Callianassa
limosa, Callianassa
lobata, Callianassa
lobetobensis, Callianassa
longicauda, Callianassa
longiventris, Callianassa
louisianensis, jamaicense, Callianassa
madagassa, Calliaxiopsis
madagassa, Callianassa
major, Callianassa
makarovi, Nihonotrypaea
malaccaensis, Callianassa
maldivensis, Callianassa
manihinae, Gourretia
manningi, Lepidophthalmus
manningi, Neocallichirus
maorianus, Ctenocheles
Callichiridae
Callichiridae
Callichiridae
Callianassidae
Callianassidae
Callichiridae
Callichiridae
Ctenochelidae
Callichiridae
Eucalliacidae
Callichiridae
Callianassidae
Callichiridae
Callianassidae
Callichiridae
Callichiridae
Callianassidae
Callianassidae
Eucalliacidae
Callichiridae
Callichiridae
Eucalliacidae
Callianassidae
Callichiridae
Ctenochelidae
Ctenochelidae
Callichiridae
Ctenochelidae
Ctenochelidae
Callichiridae
Ctenochelidae
Callichiridae
Ctenochelidae
Callianassidae
Callianassidae
Callianassidae
Eucalliacidae
Callianassidae
Callianassidae
Callichiridae
Callichiridae
Eucalliacidae
Callichiridae
Callichiridae
Callianassidae
Callianassidae
Callianassidae
Ctenochelidae
Callichiridae
Callichiridae
Ctenochelidae
Neocallichirus guara (Rodrigues, 1971)
Neocallichirus guassutinga (Rodrigues, 1971)
Balsscallichirus guineensis (De Man, 1928)
Neotrypaea hainanensis (Liu & Liu, 2014) comb. nov.
Neotrypaea harmandi (Bouvier, 1901) comb. nov.
Corallianassa hartmeyeri (Schmitt, 1935)
Corallianassa haswelli (Poore & Griffin, 1979)
Ctenocheles holthuisi Rodrigues, 1978
Neocallichirus horneri Sakai, 1988
Eucalliaxiopsis inaequimana (Dworschak, 2014) comb. nov.
Michaelcallianassa indica Sakai, 2002
Pugnatrypaea intermedia (De Man, 1905) comb. nov.
Corallianassa intesi (de Saint Laurent & LeLoeuff, 1979)
Pugnatrypaea iranica (Sepahvand, Momtazi & Tudge, 2015) comb. nov.
Callichirus islagrande (Schmitt, 1935)
Lepidophthalmus jamaicense (Schmitt, 1935)
Neotrypaea japonica (Ortmann, 1891) comb. nov.
Jocullianassa joculatrix (De Man, 1905) comb. nov.
Eucalliaxiopsis jonesi (Heard, 1989) comb. nov.
Neocallichirus jousseaumei (Nobili, 1904)
Karumballichirus karumba (Poore & Griffin, 1979) comb. nov.
Calliaxina kensleyi (Dworschak, 2005)
Jocullianassa kowalevski (Sakai, 2010) comb. nov.
Kraussillichirus kraussi (Stebbing, 1900) comb. nov.
Paragourretia laevidactyla (Liu & Liu, 2010) comb. nov.
Paragourretia lahouensis (LeLoeuff & Intes, 1974)
Corallianassa lanceolata (Edmondson, 1944)
Gourretia laresi Blanco-Rambla & Liñero-Arana, 1994
Dawsonius latispina (Dawson, 1967)
Glypturus laurae (de Saint Laurent, 1984)
Gourretia loeuffintesi Sakai, 2011
Neocallichirus lemaitrei Manning, 1993
Ctenocheles leviceps Rabalais, 1979
Aqaballianassa lewtonae (Ngoc-Ho, 1994) comb. nov.
Rayllianassa lignicola (Alcock & Anderson, 1899) comb. nov.
Biffarius limosus (Poore, 1975)
Calliax lobata (de Gaillande & Lagardere, 1966)
Pugnatrypaea lobetobensis (De Man, 1905) comb. nov.
Praedatrypaea longicauda (Sakai, 1967) comb. nov.
Corallianassa longiventris (A.Milne-Edwards, 1870)
Lepidophthalmus louisianensis (Schmitt, 1935)
Eucalliaxiopsis madagassa (Sakai & Türkay, 2014) comb. nov.
Lepidophthalmus madagassus (Lenz & Richters, 1881) comb. nov.
Callichirus major (Say, 1818)
Neotrypaea makarovi (Marin, 2013) comb. nov.
Coriollianassa malaccaensis (Sakai, 2002) comb. nov.
Paratrypaea maldivensis (Borradaile, 1904)
Gourretia manihinae Sakai, 1984
Lepidophthalmus manningi Felder & Staton, 2000
Neocallichirus manningi Kazmi & Kazmi, 1992
Ctenocheles maorianus Powell, 1949
139
Callianassidae and related families
Species, Genus
Family allocation
Current combination, authority
marchali, Callianassa
marginata, Callianassa
martensi, Callianassa
maryae, Neocallichirus
masoomi, Callianassa
matzi, Callianassa
mauritana, Phaetoncalliax
mauritiana, Callianassa
mcilhennyi, Eucalliax
melissae, Biffarius
mericeae, Sergio
mirim, Callianassa
mocambiquensis, Callianassa
modesta, Callianassa (Calliactites)
moluccensis, Callianassa (Cheramus)
monodi, Callichirus
mortenseni, Neocallichirus
mucronata, Callianassa
natalensis, Callianassa
natesi, Lepidophthalmus
ngochoae, Callianassa
nickellae, Neocallichirus
nieli, Callianassa
nigroculata, Callianassa
nomurai, Ctenocheloides
nosybeensis, Gourretia
novae-britanniae, Callianassa
oblonga, Callianassa
ohuranus, Cheramus
orientalis, Cheramus
pachydactyla, Callianassa
pacificus, Biffarius
panamensis, Lepidophthalmus
panglaoensis, Eucalliax
parva, Callianassa (Calliactites)
parvula, Callianassa
pentagonocephala, Callianassa
persica, Callianassa
petalura, Callianassa
phuketensis, Gourretia
pixii, Callianassa
plantei, Callianassa
plantei, Thaumastochelopsis
pola, Neocallichirus
poorei, Callianassa
portsudanensis, Pseudogourretia
praedatrix, Callianassa
profunda, Callianassa
propinqua, Callianassa
propriopedis, Callianassa
pugnatrix, Callianassa
Callianassidae
Callianassidae
Callichiridae
Callichiridae
Callichiridae
Callianassidae
Callianopsidae
Callichiridae
Eucalliacidae
Callianassidae
Callichiridae
Callichiridae
Callichiridae
Callianassidae
Callichiridae
Callichiridae
Callichiridae
Callichiridae
Callichiridae
Callichiridae
Callianassidae
Callichiridae
Callianassidae
Callianassidae
Ctenochelidae
Ctenochelidae
Eucalliacidae
Callianassidae
Callianassidae
Callianassidae
Callichiridae
Callianassidae
Callichiridae
Eucalliacidae
Callianassidae
Callianassidae
Callichiridae
Callianassidae
Callianassidae
Ctenochelidae
Callichiridae
Callianassidae
Ctenochelidae
Callichiridae
Callianassidae
Ctenochelidae
Callianassidae
Callianassidae
Callianassidae
Callianassidae
Callianassidae
Callianassa marchali LeLoeuff & Intes, 1974
Cheramoides marginata (Rathbun, 1901)
Corallianassa martensi (Miers, 1884)
Neocallichirus maryae Karasawa, 2004
Balsscallichirus masoomi (Tirmizi, 1970)
Jocullianassa matzi (Sakai, 2002) comb. nov.
Callianopsis mauritana (Sakai, Türkay, Beuck & Freiwald, 2015)
Neocallichirus mauritianus (Miers, 1882)
Eucalliaxiopsis mcilhennyi (Felder & Manning, 1994) comb. nov.
Neotrypaea melissae (Poore, 2008) comb. nov.
Neocallichirus mericeae (Manning & Felder, 1995) comb. nov.
Audacallichirus mirim (Rodrigues, 1971) comb. nov.
Mocallichirus mocambiquensis (Sakai, 2004) comb. nov.
Praedatrypaea modesta (De Man, 1905) comb. nov.
Neocallichirus moluccensis (De Man, 1905)
Audacallichirus monodi (de Saint Laurent & LeLoeuff, 1979) comb. nov.
Neocallichirus mortenseni Sakai, 2005
Mucrollichirus mucronatus (Strahl, 1862) comb. nov.
Neocallichirus natalensis (Barnard, 1947)
Lepidophthalmus natesi Felder & Robles, 2015
Incertae sedis ngochoae Sakai, 1999
Neocallichirus nickellae Manning, 1993
Aqaballianassa nieli (Sakai, 2002) comb. nov.
Incertae sedis nigroculata Sakai, 200
Kiictenochelodes nomurai (Komai, 2013)
Gourretia nosybeensis Sakai, 2004
Calliaxina novaebritanniae (Borradaile, 1900) comb. nov.
Cheramoides oblonga (LeLoeuff & Intes, 1974) comb. nov.
Spinicallianassa ohurana (Komai, Maenosono & Fujita, 2014) comb. nov.
Pugnatrypaea orientalis (Bate, 1888) comb. nov.
Neocallichirus pachydactyla (A. Milne-Edwards, 1870)
Neotrypaea pacifica (Guzmán &Thatje, 2003) comb. nov.
Lepidophthalmus panamensis Felder & Robles, 2015
Eucalliaxiopsis panglaoensis (Dworschak, 2006) comb. nov.
Rayllianassa parva (Edmondson, 1944) comb. nov.
Spinicallianassa parvula (Sakai, 1988) comb. nov.
Audacallichirus pentagonocephala (Rossignol, 1962) comb. nov.
Callianassa persica Sakai, 2005
Neotrypaea petalura (Stimpson, 1860) comb. nov.
Paragourretia phuketensis (Sakai, 2002)
Balsscallichirus pixii (Kensley, 1976) comb. nov.
Incertae sedis plantei Sakai, 2004
Ctenocheles plantei (Burukovsky, 2005)
Neocallichirus pola Sakai & Türkay, 2014
Tastrypaea poorei (Sakai, 1999) comb. nov.
Paragourretia portsudanensis (Sakai, 2005) comb. nov.
Praedatrypaea praedatrix (De Man, 1905) comb. nov.
Cheramus profundus (Biffar, 1973)
Praedatrypaea propinqua (De Man, 1905) comb. nov.
Incertae sedis propriopedis Sakai, 2002
Pugnatrypaea pugnatrix (De Man, 1905) comb. nov.
140
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Species, Genus
Family allocation
Current combination, authority
punica, Calliax
pygmaea, Callianassa
qeshmensis, Gourretia
quadracuta, Callianassa
rabalaisae, Glypturus
rafai, Lepidophthalmus
ranongensis, Callianassa (Callichirus)
raymanningi, Neocallichirus
rhopalommata, Laurentgourretia
richardi, Lepidophthalmus
Rochei, Callianassa
Rosae, Callianassa (Callichirus)
rotundicaudata, Callianassa
rotundocula, Trypaea
rudisulcus, Rayllianassa
sahul, Callianassa
sakaii, Calliax
santarita, Notiax
santarosaensis, Callichirus
sassandrensis, Callichirus
seilacheri, Callianassa
serrifrons, Ctenocheles
setimanus, Gonodactylus
Sibogae, Callianassa
sinica, Michaelcallianassa
sinica, Gourretia
sinuensis, Lepidophthalmus
siriboia, Lepidophthalmus
spinicauda, Cheramus
spinoculata, Callianassa
spinophthalma, Callianassa
spinorostra, Trypaea
spiridonovi, Callichiropsis
statoni, Lepidophthalmus
stenomastaxa, Callianassa
subterraneus, Cancer (Astacus)
sulfureus, Sergio
tabogensis, Callianassa
tamakii, Grynaminna
tenuimanus, Callichirus
tenuipes, Callianassa
thailandica, Callianassa
thalesapensis, Neocallichirus
thermophila, Nihonotrypaea
thorsoni, Callianassa
timiris, Neocallichirus
tonganus, Paraglypturus
tonkinae, Callianassa (Scallasis)
tooradin, Callianassa
tridentata, Callianassa
trilobata, Callianassa
Eucalliacidae
Callianassidae
Ctenochelidae
Eucalliacidae
Callichiridae
Callichiridae
Callichiridae
Callichiridae
Ctenochelidae
Callichiridae
Callianassidae
Callichiridae
Callianassidae
Callianassidae
Callianassidae
Callianassidae
Eucalliacidae
Callianassidae
Callichiridae
Callichiridae
Callichiridae
Ctenochelidae
Callianassidae
Callianassidae
Callichiridae
Ctenochelidae
Callichiridae
Callichiridae
Callianassidae
Callianassidae
Callianassidae
Callianassidae
Callichiridae
Callichiridae
Callianassidae
Callianassidae
Callichiridae
Callianassidae
Callichiridae
Callichiridae
Callianassidae
Callianassidae
Callichiridae
Callianassidae
Callianassidae
Callianassidae
Eucalliacidae
Callianassidae
Eucalliacidae
Callichiridae
Callichiridae
Calliaxina punica (de Saint Laurent & Manning, 1982)
Scallasis pygmaea (De Man, 1928) comb. nov.
Gourretia qeshmensis Sepahvand, Pouyani & Momtazi, 2016
Eucalliax quadracuta (Biffar, 1970)
Glypturus rabalaisae Sakai, 2011
Lepidophthalmus rafai Felder & Manning, 1998
Thailandcallichirus ranongensis (Sakai, 1983)
Neocallichirus raymanningi Blanco-Rambla & Lemaitre, 1999
Laurentgourretia rhopalommata Sakai, 2004
Lepidophthalmus richardi Felder & Manning, 1997
Neotrypaea rochei (Bouvier, 1895)
Lepidophthalmus rosae (Nobili, 1904)
Gilvossius rotundicaudatus (Stebbing, 1902)
Incertae sedis rotundocula Sakai & Türkay, 2014
Rudisullianassa rudisulcus (Komai, Fujita & Maenosono, 2014) comb. nov.
Rayllianassa sahul (Poore, 2008) comb. nov.
Calliaxina sakaii (de Saint Laurent, 1979)
Notiax santarita Thatje, 2000
Callichirus santarosaensis Sakai & Türkay, 2012
Neocallichirus sassandrensis (LeLoeuff & Intès, 1974)
Callichirus seilacheri (Bott, 1955)
Ctenocheles serrifrons LeLoeuff & Intès, 1974
Gilvossius setimanus (DeKay, 1844)
Coriollianassa sibogae (De Man, 1905) comb. nov.
Michaelcallianassa sinica Liu & Liu, 2009
Gourretia sinica Liu & Liu, 2010
Lepidophthalmus sinuensis Lemaitre & Rodrigues, 1991
Lepidophthalmus siriboia Felder & Rodrigues, 1993
Spinicallianassa spinicauda (Komai, Maenosono & Fujita, 2014) comb. nov.
Aqaballianassa spinoculata (Sakai, 2005) comb. nov.
Scallasis spinophthalma (Sakai, 1970) comb. nov.
Jocullianassa spinorostra (Sakai, 2010) comb. nov.
Neocallichirus spiridonovi (Sakai, 2010) comb. nov.
Lepidophthalmus statoni Felder, 2015
Incertae sedis stenomastaxa Sakai, 2002
Callianassa subterranea (Montagu, 1808)
Neocallichirus sulfureus (Lemaitre & Felder, 1996)
Neotrypaea tabogensis (Sakai, 2005)
Grynaminna tamakii Poore, 2000
Balsscallichirus tenuimanus (de Saint Laurent & LeLoeuff, 1979)
Incertae sedis tenuipes Sakai, 2002
Caviallianassa thailandica (Sakai, 2005) comb. nov.
Karumballichirus thalesapensis (Sakai & Lheknim, 2014) comb. nov.
Neotrypaea thermophila (Lin, Komai & Chan, 2007) comb. nov.
Aqaballianassa thorsoni (Sakai, 2005) comb. nov.
Callianassa timiris (Sakai, Türkay, Beuck & Freiwald, 2015) comb. nov.
Paraglypturus tonganus Ahn, Kim, Ju & Min, 2017
Scallasis tonkinae (Grebenjuk, 1975) comb. nov.
Pseudocalliax tooradin (Poore & Griffin, 1979)
Lepidophthalmus tridentatus (von Martens, 1868)
Glypturoides trilobata (Biffar, 1970)
141
Callianassidae and related families
Species, Genus
Family allocation
Current combination, authority
truncata, Callianassa
tulearensis, Calliax
turnerana, Callianassa
tyrrhenus, Astacus
uncinata, Callianassa
variabilis, Callianassa (Cheramus)
vaugelasi, Neocallichirus
vietnamensis, Trypaea
vigilax, Callianassa (Callichirus)
whitei, Callianassa
winslowi, Callianassa (Callichirus)
xishaensis, Calliaxina
xutha, Corallianassa
zarenkovi, Paragourretia
Callianassidae
Eucalliacidae
Callichiridae
Callianassidae
Callianassidae
Callichiridae
Callichiridae
Callianassidae
Callichiridae
Callianassidae
Callichiridae
Eucalliacidae
Callichiridae
Ctenochelidae
Necallianassa truncata (Giard & Bonnier, 1890)
Calliax tulearensis Ngoc-Ho, 2014
Lepidophthalmus turneranus (White, 1861)
Gilvossius tyrrhenus (Petagna, 1792)
Neotrypaea uncinata (H.Milne Edwards, 1837)
Neocallichirus variabilis (Edmondson, 1944)
Neocallichirus vaugelasi Dworschak, 2011
Jocullianassa vietnamensis (Sakai, 2010) comb. nov.
Neocallichirus vigilax (De Man, 1916)
Gilvossius whitei (Sakai, 1999)
Corallianassa winslowi (Edmondson, 1944)
Calliaxina xishaensis Liu & Liang, 2016
Corallianassa xutha Manning, 1988
Gourretia zarenkovi (Sakai, 2010) comb. nov.
142
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Table 2. Accepted names of 265 species of callianassoids listed alphabetically by family and genus. 87 are in new combinations. Commonly
accepted junior synonyms are not included (see WoRMS Editorial Board 2018). Eleven species of Callianassidae inadequately described or
figured are placed incertae sedis at the end of the table.
Anacalliacidae
Anacalliax agassizi (Biffar, 1971)
Anacalliax argentinensis (Biffar, 1971)
Callianassidae
Aqaballianassa amplimaxilla (Sakai, 2002) comb. nov.
Aqaballianassa aqabaensis (Dworschak, 2003) comb. nov.
Aqaballianassa brevirostris (Sakai, 2002) comb. nov.
Aqaballianassa ehsani (Sepahvand, Tudge & Momtazi, 2018) comb. nov.
Aqaballianassa lewtonae (Ngoc-Ho, 1994) comb. nov.
Aqaballianassa nieli (Sakai, 2002) comb. nov.
Aqaballianassa spinoculata (Sakai, 2005) comb. nov.
Aqaballianassa thorsoni (Sakai, 2005) comb. nov.
Arenallianassa arenosa (Poore, 1975) comb. nov.
Biffarius biformis (Biffar, 1971)
Biffarius delicatulus Rodrigues & Manning, 1992
Biffarius limosus (Poore, 1975)
Callianassa australis Kensley, 1974
Callianassa diaphora LeLoeuff & Intès, 1974
Callianassa marchali LeLoeuff & Intès, 1974
Callianassa persica Sakai, 2005
Callianassa subterranea (Montagu, 1808)
Callianassa timiris (Sakai, Türkay, Beuck & Freiwald, 2015) comb. nov.
Caviallianassa cavifrons (Komai & Fujiwara, 2012) comb. nov.
Caviallianassa thailandica (Sakai, 2005) comb. nov.
Cheramoides brachytelson (Sakai, 2002) comb. nov.
Cheramoides marginata (Rathbun, 1901)
Cheramoides oblonga (LeLoeuff & Intes, 1974) comb. nov.
Cheramus profundus (Biffar, 1973)
Coriollianassa coriolisae (Ngoc-Ho, 2014) comb. nov.
Coriollianassa malaccaensis (Sakai, 2002) comb. nov.
Coriollianassa sibogae (De Man, 1905) comb. nov.
Filhollianassa ceramica (Fulton & Grant, 1906) comb. nov.
Filhollianassa filholi (A.Milne-Edwards, 1878) comb. nov.
Fragillianassa debilis (Hernandez-Aguilera, 1998) comb. nov.
Fragillianassa fragilis (Biffar, 1970) comb. nov.
Gilvossius arguinensis Sakai, Türkay, Beuck & Freiwald, 2015
Gilvossius candidus (Olivi, 1792)
Gilvossius convexus (de Saint Laurent & LeLoeuff, 1979)
Gilvossius rotundicaudatus (Stebbing, 1902)
Gilvossius setimanus (DeKay, 1844)
Gilvossius tyrrhenus (Petagna, 1792)
Gilvossius whitei (Sakai, 1999)
Jocullianassa joculatrix (De Man, 1905) comb. nov.
Jocullianassa matzi (Sakai, 2002) comb. nov.
Jocullianassa spinorostra (Sakai, 2010) comb. nov.
Jocullianassa vietnamensis (Sakai, 2010) comb. nov.
Jocullianassa kowalevski (Sakai, 2010) comb. nov.
Lipkecallianassa abyssa Sakai, 2002
Necallianassa acanthura (Caroli, 1946)
Necallianassa berylae Heard & Manning, 1998
Necallianassa truncata (Giard & Bonnier, 1890)
Neotrypaea biffari (Holthuis, 1991)
Neotrypaea caesari (Heard & Manning, 2000) comb. nov.
Neotrypaea californiensis (Dana, 1854)
Callianassidae and related families
Neotrypaea costaricensis (Sakai, 2005) comb. nov.
Neotrypaea gigas (Dana, 1852)
Neotrypaea hainanensis (Liu & Liu, 2014) comb. nov.
Neotrypaea harmandi (Bouvier, 1901) comb. nov.
Neotrypaea japonica (Ortmann, 1891) comb. nov.
Neotrypaea makarovi (Marin, 2013) comb. nov.
Neotrypaea melissae (Poore, 2008) comb. nov.
Neotrypaea pacifica (Guzmán &Thatje, 2003) comb. nov.
Neotrypaea petalura (Stimpson, 1860) comb. nov.
Neotrypaea rochei (Bouvier, 1895)
Neotrypaea tabogensis (Sakai, 2005)
Neotrypaea thermophila (Lin, Komai & Chan, 2007) comb. nov.
Neotrypaea uncinata (H.Milne Edwards, 1837)
Notiax brachyophthalma (A.Milne-Edwards, 1870)
Notiax santarita Thatje, 2000
Paratrypaea bouvieri (Nobili, 1904)
Paratrypaea maldivensis (Borradaile, 1904)
Poti gaucho Rodrigues & Manning, 1992
Praedatrypaea longicauda (Sakai, 1967) comb. nov.
Praedatrypaea modesta (De Man, 1905) comb. nov.
Praedatrypaea praedatrix (De Man, 1905) comb. nov.
Praedatrypaea propinqua (De Man, 1905) comb. nov.
Pugnatrypaea bicauda (Sakai, 2010) comb. nov.
Pugnatrypaea intermedia (De Man, 1905) comb. nov.
Pugnatrypaea iranica (Sepahvand, Momtazi & Tudge, 2015) comb. nov.
Pugnatrypaea lobetobensis (De Man, 1905) comb. nov.
Pugnatrypaea orientalis (Bate, 1888) comb. nov.
Pugnatrypaea pugnatrix (De Man, 1905) comb. nov.
Rayllianassa amboinensis (De Man, 1888)
Rayllianassa bangensis (Sakai, 2005) comb. nov.
Rayllianassa lignicola (Alcock & Anderson, 1899) comb. nov.
Rayllianassa parva (Edmondson, 1944) comb. nov.
Rayllianassa sahul (Poore, 2008) comb. nov.
Rudisullianassa rudisulcus (Komai, Fujita & Maenosono, 2014) comb. nov.
Scallasis amboinae Bate, 1888
Scallasis andamaniensis (Sakai, 2010) comb. nov.
Scallasis caledonica (Ngoc-Ho, 1991) comb. nov.
Scallasis contipes (Sakai, 2002) comb. nov.
Scallasis pygmaea (De Man, 1928) comb. nov.
Scallasis spinophthalma (Sakai, 1970) comb. nov.
Scallasis tonkinae (Grebenjuk, 1975) comb. nov.
Spinicallianassa acutirostella (Sakai, 1988) comb. nov.
Spinicallianassa ohurana (Komai, Maenosono & Fujita, 2014) comb. nov.
Spinicallianassa parvula (Sakai, 1988) comb. nov.
Spinicallianassa spinicauda (Komai, Maenosono & Fujita, 2014) comb. nov.
Tastrypaea poorei (Sakai, 1999) comb. nov.
Trypaea australiensis Dana, 1852
Callianopsidae
Bathycalliax geomar Sakai & Türkay, 1999
Callianopsis anovalis Lin, Komai & Chan, 2007
Callianopsis caecigena (Alcock & Anderson, 1894)
Callianopsis goniophthalma (Rathbun, 1902)
Callianopsis mauritana (Sakai, Türkay, Beuck & Freiwald, 2015)
Vulcanocalliax arutyunovi Dworschak & Cunha, 2007
Callichiridae
Audacallichirus audax (De Man, 1911) comb. nov.
Audacallichirus mirim (Rodrigues, 1971) comb. nov.
143
144
G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder
Audacallichirus monodi (de Saint Laurent & LeLoeuff, 1979) comb. nov.
Audacallichirus pentagonocephala (Rossignol, 1962) comb. nov.
Balsscallichirus balssi (Monod, 1933)
Balsscallichirus foresti (LeLoeuff & Intes, 1974) comb. nov.
Balsscallichirus gilchristi (Barnard, 1947)
Balsscallichirus guineensis (De Man, 1928)
Balsscallichirus masoomi (Tirmizi, 1970)
Balsscallichirus pixii (Kensley, 1976) comb. nov.
Balsscallichirus tenuimanus (de Saint Laurent & LeLoeuff, 1979)
Calliapagurops charcoti de Saint Laurent, 1973
Calliapagurops foresti Ngoc-Ho, 2002
Callichirus adamas (Kensley, 1974)
Callichirus garthi (Retamal, 1975)
Callichirus islagrande (Schmitt, 1935)
Callichirus major (Say, 1818)
Callichirus santarosaensis Sakai & Türkay, 2012
Callichirus seilacheri (Bott, 1955)
Corallianassa articulata (Rathbun, 1906)
Corallianassa assimilis (De Man, 1928)
Corallianassa borradailei (De Man, 1928)
Corallianassa collaroy (Poore & Griffin, 1979)
Corallianassa coutierei (Nobili, 1904)
Corallianassa hartmeyeri (Schmitt, 1935)
Corallianassa haswelli (Poore & Griffin, 1979)
Corallianassa intesi (de Saint Laurent & LeLoeuff, 1979)
Corallianassa lanceolata (Edmondson, 1944)
Corallianassa longiventris (A.Milne-Edwards, 1870)
Corallianassa martensi (Miers, 1884)
Corallianassa winslowi (Edmondson, 1944)
Corallianassa xutha Manning, 1988
Glypturoides trilobata (Biffar, 1970)
Glypturus acanthochirus Stimpson, 1866
Glypturus armatus (A.Milne-Edwards, 1870)
Glypturus laurae (de Saint Laurent, 1984)
Glypturus rabalaisae Sakai, 2011
Grynaminna tamakii Poore, 2000
Karumballichirus karumba (Poore & Griffin, 1979) comb. nov.
Karumballichirus thalesapensis (Sakai & Lheknim, 2014) comb. nov.
Kraussillichirus kraussi (Stebbing, 1900) comb. nov.
Laticallichirus grandis (Karasawa & Goda, 1996)
Lepidophthalmus bocourti (A. Milne-Edwards, 1870)
Lepidophthalmus eiseni Holmes, 1904
Lepidophthalmus grandidieri (Coutière, 1899)
Lepidophthalmus jamaicense (Schmitt, 1935)
Lepidophthalmus louisianensis (Schmitt, 1935)
Lepidophthalmus madagassus (Lenz & Richters, 1881) comb. nov.
Lepidophthalmus manningi Felder & Staton, 2000
Lepidophthalmus natesi Felder & Robles, 2015
Lepidophthalmus panamensis Felder & Robles, 2015
Lepidophthalmus rafai Felder & Manning, 1998
Lepidophthalmus richardi Felder & Manning, 1997
Lepidophthalmus rosae (Nobili, 1904)
Lepidophthalmus sinuensis Lemaitre & Rodrigues, 1991
Lepidophthalmus siriboia Felder & Rodrigues, 1993
Lepidophthalmus statoni Felder, 2015
Lepidophthalmus tridentatus (von Martens, 1868)
Lepidophthalmus turneranus (White, 1861)
Michaelcallianassa indica Sakai, 2002
Callianassidae and related families
Michaelcallianassa sinica Liu & Liu, 2009
Mocallichirus mocambiquensis (Sakai, 2004) comb. nov.
Mucrollichirus mucronatus (Strahl, 1862) comb. nov.
Neocallichirus angelikae Sakai, 2000
Neocallichirus auchenorhynchus Sakai, 2005
Neocallichirus cacahuate Felder & Manning, 1995
Neocallichirus caechabitator Sakai, 1988
Neocallichirus calmani (Nobili, 1904)
Neocallichirus darwinensis Sakai, 1988
Neocallichirus darvishi Sepahvand, Komai, Momtazi & Shahabi, 2018
Neocallichirus frouini Ngoc-Ho, 2005
Neocallichirus grandimana (Gibbes, 1850)
Neocallichirus guaiqueri (Blanco-Rambla, Liñero-Arana & Beltràn Lares, 1995)
Neocallichirus guara (Rodrigues, 1971)
Neocallichirus guassutinga (Rodrigues, 1971)
Neocallichirus horneri Sakai, 1988
Neocallichirus jousseaumei (Nobili, 1904)
Neocallichirus lemaitrei Manning, 1993
Neocallichirus manningi Kazmi & Kazmi, 1992
Neocallichirus maryae Karasawa, 2004
Neocallichirus mauritianus (Miers, 1882)
Neocallichirus mericeae (Manning & Felder, 1995) comb. nov.
Neocallichirus moluccensis (De Man, 1905)
Neocallichirus mortenseni Sakai, 2005
Neocallichirus natalensis (Barnard, 1947)
Neocallichirus nickellae Manning, 1993
Neocallichirus pachydactyla (A. Milne-Edwards, 1870)
Neocallichirus pola Sakai & Türkay, 2014
Neocallichirus raymanningi Blanco-Rambla & Lemaitre, 1999
Neocallichirus sassandrensis (LeLoeuff & Intes, 1974)
Neocallichirus spiridonovi (Sakai, 2010) comb. nov.
Neocallichirus sulfureus (Lemaitre & Felder, 1996)
Neocallichirus variabilis (Edmondson, 1944)
Neocallichirus vaugelasi Dworschak, 2011
Neocallichirus vigilax (De Man, 1916)
Thailandcallichirus ranongensis (Sakai, 1983)
Ctenochelidae
Ctenocheles balssi Kishinouye, 1926
Ctenocheles collini Ward, 1945
Ctenocheles holthuisi Rodrigues, 1978
Ctenocheles leviceps Rabalais, 1979
Ctenocheles maorianus Powell, 1949
Ctenocheles plantei (Burukovsky, 2005)
Ctenocheles serrifrons LeLoeuff & Intes, 1974
Ctenocheloides almeidai Anker & Pachelle, 2013
Ctenocheloides attenboroughi Anker, 2010
Ctenocheloides boucheti Poore, 2015)
Dawsonius latispina (Dawson, 1967)
Gourretia barracuda LeLoeuff & Intes, 1974
Gourretia denticulata (Lutze, 1937)
Gourretia laresi Blanco-Rambla & Linero-Arana, 1994
Gourretia loeuffintesi Sakai, 2011
Gourretia manihinae Sakai, 1984
Gourretia nosybeensis Sakai, 2004
Gourretia qeshmensis Sepahvand, Pouyani & Momtazi, 2016
Gourretia sinica Liu & Liu, 2010
Gourretia zarenkovi (Sakai, 2010) comb. nov.
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Kiictenochelodes nomurai (Komai, 2013)
Laurentgourretia rhopalommata Sakai, 2004
Paragourretia aungtonyae (Sakai, 2002)
Paragourretia biffari (Blanco-Rambla & Liñero-Arana, 1994)
Paragourretia coolibah (Poore & Griffin, 1979)
Paragourretia crosnieri (Ngoc-Ho, 1991)
Paragourretia galathea (Sakai, 2017) comb. nov.
Paragourretia laevidactyla (Liu & Liu, 2010) comb. nov.
Paragourretia lahouensis (LeLoeuff & Intès, 1974)
Paragourretia phuketensis (Sakai, 2002)
Paragourretia portsudanensis (Sakai, 2005) comb. nov.
Eucalliacidae
Andamancalliax andamanica (Sakai, 2002)
Calliax doerjesti Sakai, 1999
Calliax lobata (de Gaillande & Lagardère, 1966)
Calliax tulearensis Ngoc-Ho, 2014
Calliaxina bulimba (Poore & Griffin, 1979)
Calliaxina kensleyi (Dworschak, 2005)
Calliaxina novaebritanniae (Borradaile, 1900) comb. nov.
Calliaxina punica (de Saint Laurent & Manning, 1982)
Calliaxina sakaii (de Saint Laurent, 1979)
Calliaxina xishaensis Liu & Liang, 2016
Eucalliax quadracuta (Biffar, 1970)
Eucalliaxiopsis aequimana (Baker, 1907) comb. nov.
Eucalliaxiopsis cearaensis (Rodrigues & Manning, 1992)
Eucalliaxiopsis inaequimana (Dworschak, 2014) comb. nov.
Eucalliaxiopsis jonesi (Heard, 1989) comb. nov.
Eucalliaxiopsis madagassa (Sakai & Türkay, 2014) comb. nov.
Eucalliaxiopsis mcilhennyi (Felder & Manning, 1994) comb. nov.
Eucalliaxiopsis panglaoensis (Dworschak, 2006) comb. nov.
Paraglypturus calderus Türkay & Sakai, 1995
Paraglypturus tonganus Ahn, Kim, Ju & Min, 2017
Pseudocalliax tooradin (Poore & Griffin, 1979)
Paracalliacidae
Paracalliax bollorei de Saint Laurent, 1979
Callianassidae incertae sedis
Callianassa tenuipes Sakai, 2002 – few illustrations, possibly Jocullianassa
Callianassa anoploura Sakai, 2002 – few illustrations
Callianassa chakratongae Sakai, 2002 – female only described
Callianassa exilimaxilla Sakai, 2005 – female only illustrated, possibly Jocullianassa
Callianassa gruneri Sakai, 1999 – few illustrations
Callianassa nigroculata Sakai, 2002 – female only illustrated, possibly Scallasis
Callianassa ngochoae Sakai, 1999 – female only described
Callianassa plantei Sakai, 2004 – possibly two species in illustrated material; chelipeds from holotype typical of Paratrypaea
but maxilliped 3 from another specimen of narrow form
Callianassa propriopedis Sakai, 2002 – female only described
Callianassa stenomastaxa Sakai, 2002 – female only described (possibly same species as C. propriopedis)
Trypaea rotundocula Sakai & Türkay, 2014 – female only described
Rafael Robles