Memoirs of Museum Victoria 78: 1—21 (2019) Published 2019

1447-2594 (On-line)

htto://museumvictoria.com.au/about/books-and-journals/journals/memoirs-of-museum-victoria/

DOI https://doi.org/10.24199/j.mmv.2019.78.01

Microscopic analysis of the developing dentition in the pouch young of the extinct

marsupial Thylacinus cynocephalus, with an assessment of other developmental

stages and eruption

W. Patrick LUCKETT!*, Nancy HoNG LUCKETT?**, AND TONY HARPER!

! Center for Functional Anatomy and Evolution, Johns Hopkins University School of Medicine, Baltimore, Maryland, USA

* Caribbean Primate Research Center Museum, University of Puerto Rico, Medical Sciences Campus, San Juan,

Puerto Rico, USA

! Center for Functional Anatomy and Evolution, Johns Hopkins University School of Medicine, Baltimore, Maryland, USA

* To whom correspondence should be addressed. E-mail: plucket3 @jhmi.edu

*'*Deceased

Abstract

Luckett, W.P., Hong Luckett, N. and Harper, T. 2019. Microscopic analysis of the developing dentition in the pouch

young of the extinct marsupial Thylacinus cynocephalus, with an assessment of other developmental stages and eruption.

Memoirs of Museum Victoria 78: 1—21.

A pouch young of the extinct dasyuromorphian marsupial Thylacinus cynocephalus was examined histologically to

study the stages of the developing dentition during the pre-eruptive period of development. Both deciduous and successional

stages of tooth development were examined, and these were compared to later stages of development and eruption of the

teeth in Thylacinus and with selected developmental stages of other dasyuromorphians. Our analysis shows that the

development and eruption of teeth in Thylacinus is most similar to that of dasyurids with only two premolars, such as

Dasyurus and Sarcophilus, rather than the dasyurids with three premolars, such as Antechinus and Sminthopsis.

Keywords

Institutional abbreviations

AM, Australian Museum, Sydney, Australia; AMNH, American

Museum of Natural History, New York, New York, USA; CU,

Cambridge University, Museum of Zoology, Cambridge,

England; LAC, Laboratoire d'Anatomie Comparée, Paris,

France; MN, Museum für Naturkunde, Berlin, Germany; NMV,

Museums Victoria, Melbourne, Australia; SAM, South

Australian Museum, Adelaide, Australia; TMAG, Tasmanian

Museum and Art Gallery, Hobart, Tasmania, Australia; USNM,

United States National Museum, Washington DC, USA, WAM,

Western Australian Museum, Perth, Australia

Introduction

Although the last known living specimen of the marsupial

Thylacinus cynocephalus died in the Hobart Zoo in Tasmania,

Australia, on 7 September, 1936, frequent reports of sightings

have continued to occur in Tasmania; (see Paddle, 2012, for

a recent summary). However, no new living or dead specimens

of the thylacine have been found since 1936, and most

authorities accept that the thylacine is now extinct. An

excellent overview on the biology of Thylacinus was provided

by Joan Dixon (1989).

Thylacinus, marsupials, Dasyuridae, deciduous dentition, dental development, pouch young, homologies

The earliest study on the developing dentition of Thylacinus

known to us was conducted by William Flower (1867), who

illustrated a juvenile pouch young with unerupted teeth. In

lateral view, Flower showed three (of the four) developing

incisors, the canine, three deciduous premolars, a developing

successional third premolar (P3), and the developing MI - 2 in

both jaws. A reproduction of his figure is shown in fig. 1. The

juvenile specimen was from the Museum of the Royal College

of Surgeons of England and the Head Length (HL) measured

71.1 mm.

No teeth had yet erupted above the soft tissues of these

jaws; however, Flower’s dissection revealed the apex of a

small tooth in both the maxilla and dentary, barely elevated

above the level of the bone. These small teeth were identified

as the third deciduous premolars (dP3), overlying the deeper

and less developed third successional premolars (P3). In later

stages of all marsupials then known, Flower noted that the

underlying and larger successional third premolars would

later displace and replace these deciduous predecessors In

both jaws.

The tiny dP3 in the upper jaw measured only 2.54 mm in

length and lacked distinct roots. The tiny dp3 in the dentary

was slightly smaller and also lacked distinct roots. Following

Figure 1. Thylacinus (71.1 mm Head Length) pouch young, with

unerupted dentition; redrawn from Flower, 1867. C, upper and lower

canine; dP3, upper and lower deciduous third premolar; M1, upper and

lower first molar; P3, upper and lower successional third premolar.

his examination of Thylacinus and representatives of five

other families of marsupials, Flower emphasised that the

“peculiar condition of dental succession (Flower, 1867: page

637) in marsupials differed greatly from that in eutherian

mammals. This mode of dental development was

characterised by tooth succession at only the third premolar

position in all marsupials examined by him. Flower therefore

considered this condition in marsupials to be a “rudimentary

diphyodont condition (Flower, 1867: page 638), with tooth

replacement being confined to a single tooth position on

each side of both jaws.

As a closing thought, Flower (1867) suggested that earlier

developmental stages of marsupials might show additional

evidence for deciduous predecessors at other tooth positions.

Later studies by Oldfield Thomas (1887) revealed that

variation occurred in dental development within the marsupial

family Dasyuridae, with some genera having three premolars

(Thylacinus, Sminthopsis and Myrmecobius), whereas others

had only two premolars (Dasyurus and Sarcophilus).

Curiously, Thomas called the posterior deciduous premolar

and its successor “Pm4,” rather than dP3 and P3, considering

them to be homologous with the fourth premolars in

eutherians. Numerous studies on the developing dentitions of

marsupials and eutherians continued during the latter part of

the 19th century, although none of these included additional

specimens of Thylacinus.

several authors began to use microscopic studies of the

marsupial and eutherian dentitions 1n the 1890s. A significant

study by Leche (1893) showed histological sections of early

dental development in the dasyurid Myrmecobius, 1n which he

described and illustrated the occurrence of small abnormal

teeth, which he called "prelacteal" or pre-milk teeth, in the

developing incisor and canine regions. Some of these abnormal

W.P. Luckett, N. Hong Luckett & T. Harper

teeth contained a prominent dentinal nodule and occurred

labial to the normal developing first incisor and canine, which

were in the late bell stage of development (his figs 1 and 2).

This and other related studies on the developing dentition of

marsupials were discussed in great detail by Wilson and Hill

(1897), in their microscopic study of extensive samples of the

developing dentition in the Australian peramelid Perameles.

One of their major findings was to note that the third deciduous

premolar in several genera of marsupials differentiated from

the dental lamina contemporaneously with the so-called

^prelacteal" or pre-milk teeth in the incisor and canine regions,

and they concluded that these teeth belong to the same dental

series and were homologous to the deciduous or milk series of

eutherians.

In contrast, Wilson and Hill (1897) considered the two

premolars anterior to dP3 in both jaws of marsupials to be

homologous with the successional third premolars (1.e., to be

Pl and P2), although they presented no developmental data to

support this hypothesis. These anterior premolars develop

later than dP3 in all marsupials studied to date, and they

develop directly from the primary dental lamina, as do dP3.

We are unaware of any developmental studies that show that

these anterior deciduous premolars are replaced by

successional teeth. Nevertheless, most studies of fossil and

extant marsupials have continued to refer to these teeth as PI

and P2 [see Archer et al.(2016), Murray and Megirian (2006),

and Yates (2014, 2015) as recent examples].

In Flowers (1867) study of the juvenile Thylacinus, he

considered that it belonged in the family Dasyuridae, and he

noted that it would be useful to compare its developing

dentition with that of Dasyurus and other dasyurids.

Numerous studies on dental development in dasyurids have

been carried out since then (Archer, 1976; Luckett and

Woolley, 1996), although Thylacinus was later placed in a

separate family Thylacinidae. Today, both morphological and

molecular studies group Thylacinidae, Dasyuridae and

Myrmecobiidae within the marsupial order Dasyuromorphia

(see Archer et al., 2016, and Westerman et al., 2015).

Unfortunately, additional studies on dental development in

Thylacinus were not carried out before the genus became

extinct in 1936.

Material and methods

During a trip by one of us (WPL) to Melbourne, Australia, in

1992, to study dental development in a variety of marsupials,

numerous specimens of juvenile dasyurids and other

Australian marsupials were examined at Museums Victoria.

This included several specimens of juvenile thylacines

preserved in the alcohol collections of the Mammal

Department. One case included a female thylacine and her

four pouch young, which had been collected in Tasmania on

23 June, 1909 by W.M. McGowan. The four pouch young and

the head of the mother were stored in alcohol. Fortunately, a

testis had been sectioned from one of the pouch young by Dr

Patricia Woolley of La Trobe University; this indicated that

tissue preservation was relatively good. Following discussions

between WPL and Joan Dixon, Curator of Mammals at

Dental development in the thylacine pouch young

Figure 2. Sibling pouch young thylacines. A, NMV C5754, male

specimen sectioned for histology images; B, NMV C 5757, female

specimen, used by Feigin et al. (2018) for genomic analysis.

Museums Victoria at the time, it was agreed that one of these

valuable pouch young would be made available for histological

investigation. The pouch young (PY) selected for study (NM V

C 5754) was a male with a crown - rump length (CRL) of 77.8

mm and a head length (HL) of 34 mm (fig. 2a). The age of this

pouch young litter was recently estimated to be 4.5 weeks old

(Newton et al., 2018).

We decided to have this valuable pouch young sectioned

histologically by our colleague Professor Dr. Milan Klima at

the Dr Senckenbergische Anatomie, J.W. Goethe-Universitat in

Frankfurt am Main, Germany, because of his extensive

experience in the preparation and study of histological serial

sections from mammalian foetuses and pouch young, including

marsupials and whales. Dr. Gerhard Storch at the Senckenberg

Natural History Museum in Frankfurt am Main agreed to

assume responsibility for this valuable specimen during its

preparation in Germany. Following photographs and X-rays,

the head of the pouch young was removed and decalcified in

5% HNO,. It was then embedded in celloidin-paraffin and

sectioned serially at 10 um in a transverse (coronal) plane. The

resulting 323 slides were stained alternately with azan

trichrome, or with haematoxylin and eosin. As a consequence

of the unknown fixation of the specimen in 1909, the resulting

tissue preparation is only fair. Nevertheless, the histological

detail is adequate for the recognition of most soft tissues, the

developing dentition and the enamel matrix. There 1s some loss

of bone and of dentin in the sections. In addition to our study of

the developing dentition, we anticipated that other aspects of

the developing cranium and postcranium might be studied by

other collaborators, including Professor Klima. Following our

examination of the slides for dental development, the slide

series was returned to Museums Victoria.

After preparation of the histological series for dental

development, we were assisted by our colleague Dr. Friedemann

Schrenk, at the Landesmuseum, Darmstadt, Germany, in

Figure 3. Histological section of I1 in Thylacinus, in middle bell stage.

dp, dental papilla; en, epithelial nodule; o, oral epithelium; pdl,

primary dental lamina.

photographing the slides.There is some distortion in the

specimens, but most details can be recognised. One of us (NHL)

made some camera lucida reconstructions from the serial

sections to better show the relationships between dP3 and P3 in

both jaws. There was no eruption of any teeth in the developing

pouch young. For comparison, we show X-ray analysis of an

older pouch young (AM P 762) of the thylacine from the

Australian Museum in Sydney, which shows very early signs of

tooth eruption. Developmental stages and dental homologies

follow those described in Luckett (1993a, b) and Luckett and

Woolley (1996).

Results

Upper jaw

The first incisor (I1) is a moderately large tooth in the middle

bell stage of development, with moderate development of the

stellate reticulum (fig. 3). The outer and inner layers of the

middle bell are artifactually separated; this may be due, In part,

to the lack of dentin and enamel. Development of this tooth is

considerably retarded or delayed compared with that of the other

developing incisors. The primary dental lamina stalk Is

relatively intact and attached to the oral epithelium. The

premaxillary alveolus for this tooth is relatively shallow,

compared with that for I2. Bilaterally, there are tiny buccal

epithelial nodules that may represent epithelial remnants of a

rudimentary deciduous Il, as are known to occur in many other

marsupials (see Leche, 1893). However, this is impossible to

corroborate without access to younger developmental stages.

Such rudimentary deciduous incisors, often containing dentin,

are readily seen in the dasyurids Sminthopsis virginiae (Luckett

and Woolley, 1996) and Dasyurus viverrinus (Luckett et al.,

unpublished research). For instance, in a Dasyurus viverrinus

pouch young of 23 mm greatest length, there is a rudimentary

dll] with a distinct tiny epithelial knot associated with a

moderately large successor l! in the middle bell stage.

Figure 4. Longitudinal section through I4 and transverse section

through smaller I3. e, disrupted enamel.

I2. A large tooth with moderately thick enamel. The dentin was

probably moderately thick to thick; however, partial dissolution

of dentin 1n this and other teeth makes this difficult to determine

accurately. There are only scattered epithelial remnants of the

primary dental lamina, with little if any connections to the oral

epithelium. The alveolus for the developing tooth is much

deeper than that for I1. There 1s no evidence of the remains of a

rudimentary dI2.

I3. A large tooth, similar in development to I2, with moderately

thick enamel and disrupted dentin. The tooth is somewhat

procumbent and its enamel is thicker buccally than lingually

(fig. 4). There are prominent buccal epithelial nodules on both

sides of the jaw; these occur mesio-buccal to I3 and they could

be remnants of a rudimentary dl13.

I4. A moderately-sized tooth, smaller than I2 and I3, with

moderately developed to moderately thick enamel on its apex

(fig. 4). There is no trace of a rudimentary dl4.

W.P. Luckett, N. Hong Luckett & T. Harper

Figure 5. Longitudinal section through large successional upper

canine, with disrupted dentin and well -developed enamel. d, dentin;

e, enamel.

Canine. A large vertically 1mplanted tooth, with moderately

thick enamel and disrupted dentin (fig. 5). The tooth lies in a

deep alveolus at the rostral extent of the maxilla. There are no

traces of a rudimentary dC.

dP1. A moderately sized tooth, with relatively thick enamel

and disrupted dentin. Its primary dental lamina stalk Is

detached from the oral epithelium. A short, flattened lingual

successional lamina occurs at the mid-level of the tooth.

dP2. A moderately large tooth, with relatively thick enamel

and disrupted dentin on the apex of the single cusp (fig. 6).

The primary dental lamina stalk is also detached from the

oral epithelium. A short flattened lingual successional lamina

occurs along the mid-level of the tooth. The tooth lies anterior

to the large orbit. A thin distinct layer of connective tissue

separates the detached primary dental lamina stalk from the

oral epithelium for both dP1 and dP2.

dP3. A small tooth, lying in a shallow alveolus beneath the

anterior margins of the orbit. A layer of moderately thick to

Dental development in the thylacine pouch young

w «4 k. o, un

Las

Figure 6. Longitudinal section through dP2, with disrupted dentin and

well - developed enamel. d, dentin; e, enamel.

thick enamel and disrupted dentin overlies the small ovoid

cusp (fig. 7). There are only slender fragmented remnants of

the detached primary dental lamina stalk. A slender epithelial

strand connects the outer enamel epithelium (OEE) of dP3 to

its lingual successional lamina at a level near to the middle of

the paracone elevation. At this level, the successional lamina

appears relatively short in the sections; however, it is

continuous with the lamina extending anteriorly to the

developing successor P3 (see camera lucida reconstructions of

the same relationships between dp3 and p3 In the lower jaw;

fig. 11a, b).

Successor P3. A large late bell stage tooth with moderately

well-developed stellate reticulum lies immediately anterior to

the smaller dP3. There is no distinct evidence for dentin or

odontoblasts on the apex of the cusp (fig. 8). The larger P3,

although less differentiated than its deciduous predecessor,

extends deeper into its developing alveolus than the more

superficial disto-buccal and smaller dP3. Only a short segment

of the lingual successional lamina Is evident in fig. 8. This

successional lamina runs parallel to the oral epithelium but is

. "A

Figure 7. Longitudinal section through the apex of the small dP3, with

its disrupted dentin and thick enamel. A segment of its lingual

successional lamina is evident, but its attachment to the outer enamel

epithelium is not evident in this section. d, dentin; e, enamel; Isl, lingual

successional lamina.

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Figure 8. Late bell stage successor P3. This tooth lies anterior to dP3

and it lacks dentin and enamel. Isl, short segment of lingual

successional lamina; t, tongue.

not attached to it. Instead, it extends posteriorly through the

sections to its origin on the lingual successional lamina of dP3,

in the same manner as it occurs In the lower jaw (see camera

lucida fig. 11b).

MI. A large tooth overlapping the distal end of dP3 extends

deeper into the jaw than dP3. Only isolated remnants of the

primary dental lamina stalk are evident mesially. A

moderately elevated mesio-buccal paracone is in the late bell

stage but lacks dentin and odontoblasts (fig. 9b). A small

spherical epithelial nodule of stratum intermedium cells is

detached over the distal extent of the paracone elevation (fig.

9a), similar to the condition that occurs in some dasyurids and

Didelphis. Detachment of this small epithelial nodule over the

paracone, but not over the metacone or protocone, has been

suggested to be correlated with the earlier development and

W.P. Luckett, N. Hong Luckett & T. Harper

Figure 9. Section of MI, at level of paracone. A, paracone with overlying epithelial nodule; B, paracone at more central level, lacking the

epithelial nodule; en, epithelial nodule; pa, paracone apex.

calcification of the taller metacone, in relation to the shorter

and later developing paracone in didelphids and some

dasyurids (Luckett et al., unpublished research). Presumably,

a similar developing condition is also occurring in Thylacinus.

A short flat stylar shelf occurs buccally at the paracone

level; a broader flat lingual shelf is also present at this level for

the future development of a protocone, which is still lacking at

this stage. A moderately tall disto-buccal metacone shows

greater development, with moderately thick dentin and enamel

(fig. 10). In contrast with the paracone, there is no evidence of

a detached epithelial nodule over the taller metacone.

M2. Moderately large early-middle bell stage with a

moderately developed central epithelial knot developing distal

to MI. This tooth occurs beneath the distal 1/3 of the eye. The

dental lamina ends distal to this tooth with no trace of a

primordium for M3 or M4.

Lower jaw

il. A large tooth, somewhat procumbent, with moderately

thick enamel and dentin. Only isolated remnants of the dental

lamina stalk are evident. There is no trace of rudimentary

elements for a deciduous precursor for this or other lower

incisors.

i2. A large tooth with moderately thick enamel and

disrupted dentin.

i3. A moderately large tooth with relatively thick enamel

and disrupted dentin.

canine. A very large tooth with moderately thick dentin

and enamel, similar to the condition in the upper canine.

dpl.A moderate sized tooth with relatively thick enamel and

disrupted dentin on its apex. There is a short flattened lingual

successional lamina on both dpl and dp2, as occurred in the

upper jaws.

dp2.A larger tooth with moderately thick enamel; this may be

slightly thinner than that on dpl, but this is difficult to determine.

dp3. A small and somewhat elongate tooth, with relatively

thick enamel and disrupted dentin on the apex of the tooth (fig.

lla, b). The mesial end of the tooth overlaps the more lingually

situated, and larger, successor p3. A nearly intact slender strand

of the primary dental lamina stalk extends from the apex of the

tooth near its mesial end towards the oral epithelium. Then,

larger epithelial islands of the fragmented primary dental

lamina stalk continue towards the oral epithelium. There is no

direct connection with the oral epithelium, but the intermittent

epithelial island remnants make this earlier connection evident

(see camera lucida fig. 11b).

A single moderately elevated cusp (probably the protoconid)

is evident on the tooth; this becomes reduced distally. There is

a slight suggestion of two additional cusps distal to the

protoconid, but these are not very distinct. At the level of the

middle of the protoconid, a nearly complete slender epithelial

strand extends between the greatly thinned outer enamel

epithelium of dp3 and the fragmented proximal portion of the

lingual successional lamina (see fig. 11b). At this level there are

fewer remnants of the primary dental lamina stalk attached to

dp3; these are better developed along the mesial third of the

tooth. Epithelial strands of the lingual successional lamina

continue mesially towards the large developing successor p3.

successor p3. A large, late bell stage tooth, with thin to

moderately developed dentin on its apex (fig. 12). The tooth lies

lingual to, and somewhat mesial to, the smaller, more superficial

dp3 and it extends deeper into the jaw. Ameloblasts are polarised

over the cusp, but there is no distinct evidence of enamel.

Epithelium of the lingual successional lamina is attached to the

apex of p3 and the epithelium runs parallel to the oral epithelium

but is not attached to the latter (fig. 11a). The larger p3 extends

somewhat mesial to the smaller dp3. The intermittent, and

nearly complete successional lamina attachment between p3 and

dp3 is best developed on the right side of the jaw (see fig. 11b).

ml. A very large tooth with thick enamel and disrupted

dentin on the apex of the tall protoconid. There is no trace of

a lingual paraconid mesially. While there are other slight

swellings of the epithelium, there is no distinct evidence for

other developing cusps.

m2. A large tooth with moderately thick enamel and dentin on

the tall mesio-buccal protoconid. There is no evidence for a

Dental development in the thylacine pouch young

enamel on its apex and disrupted dentin; d, dentin; e, enamel; me,

metacone apex.

paraconid elevation mesially, or for other developing cusps.

There is a short residual lingual lamina at the level of

the protoconid.

m3. A moderately large tooth in the middle- late bell stage.

The dental lamina disappears distal to this tooth with no

evidence of a developing m4.

Other developing stages of the dentition in Thylacinus

There has been little, 1f any, attempt to describe early or later

developmental stages of the dentition In Thylacinus. One of the

few examples was the study by Heinz Moeller (1968), in which

he described and illustrated the erupting dentition In a Juvenile

skull of Thylacinus. We have included this specimen in a larger

sample of developing dentitions ın juvenile and subadult

thylacines, collected by one of us (WPL) from several

museums in Australia, North America, France and Germany.

The recent discovery of four juvenile thylacine pouch young

in the collections of the Charles University in Prague, Czech

Republic (Sleightholme et al., 2012), that are even younger than

our specimen, raises hopes for the possible further assessment of

early development of the rudimentary deciduous incisors and

canines in Thylacinus, as well as other aspects of their

developmental biology. The specimens shown in their photos

suggest the possibility that these pouch young are in a poorer

state of preservation than our pouch young, and it is unclear as to

whether the authors plan to attempt a histological study of one of

their pouch young. We hope that they will attempt such a project.

During the preparation of our manuscript, a paper

published by Newton et al. (2018) presented X-ray computed

tomography scans of five thylacine pouch young specimens,

representing all of the known PY litters of Thylacinus (see

their fig. 2). The emphasis in their study was on growth

changes 1n the cranial and postcranial skeleton; only minimal

data were provided concerning the developing dentition in

these PY. Their youngest specimen, from the Prague

collection (DZCU 8021), was estimated to be 1.5 weeks old

(10-11 days). For this pouch young , the authors noted that

"two to three tooth sockets are visible in each of the jaw

quadrants’ (Newton et al, 2018: page 9). They made no

attempt to identify or locate the specific developing teeth.

As we will discuss later, Dasyurus viverrinus appears to be

very close in its pattern of dental development to the conditions

in the thylacine, and we have examined two 10-day old PY of

D. viverrinus, 1n order to obtain an estimate of the likely dental

developmental conditions 1n the Prague pouch young of 10 —11

days old (Luckett et al., unpublished research). There is

evidence of two to three developing teeth in the dasyurid PY,

similar to the condition noted for the Prague PY. In the upper

Jaw of the Dasyurus PY, there is a well-developed dP3 in the

early—middle cap stage, and a tiny spherical epithelial knot for

dC, that is associated with a distinct lingual successional bud

for C. There is also a less distinct nodular cap anteriorly for

dI2 or dI3, with a disto-lingual successional early bud for I2 or

I3. Hopefully, these data would be useful when and if it is

possible to examine one of the Prague PY histologically.

The next older pouch young examined by Newton et al.

(2018) was estimated to be 35—37 days old (see their Fig. 5) and

has an 89 mm CRL (TMAG A931). This PY was considered by

the authors to be similar in its development to the Museums

Victoria specimens. This would include our 34 mm HL PY and

its sister (NMV C5755), examined by them (see their Fig. 2b).

Newton et al. chose to examine and describe the TMAG

specimen rather than the Museums Victoria PY, because of its

apparent better state of preservation. The authors noted the

occurrence of several deciduous teeth in both jaws, without

further comment or illustrations. However, we were able to

examine some aspects of this and the older dentitions of the

thylacine PY studied by Newton et al. (2018) due to their

inclusion of extensive electronic supplementary material that

they made available publicly for study of the dentition and the

cranio-skeletal morphology.

Our brief examination of Newton et als (2018)

supplementary material for IMAG A931, which 1s slightly older

than our sectioned thylacine PY, revealed the presence of tiny,

unerupted dP3 in both jaws, and the apparent very early eruption

of the protoconid of ml above the alveolar margins In the lower

Jaw (fig. 13a). There was no distinct evidence for any tooth

eruption in the upper Jaw. The unerupted canine and dP1-3 were

evident in both jaws. However, we were unable to see any

distinct evidence for the successor P3 1n either jaw. We suspect

that this 1s correlated with the development of little or no dentin

on the P3 as yet, similar to the condition In our sectioned PY.

The next older pouch young examined by Newton et al.

(2018) was a male (TMAG A930), estimated to be about 9.5

weeks old (66 - 67 days), and illustrated in their fig. 6. The

authors noted that this specimen contained several unerupted

teeth in both jaws but gave no further description of the

dentition. Our examination of their supplementary material

revealed the presence of tiny dP3 in both jaws; these appear to

be erupted above the alveolar margins of both jaws (fig. 13b).

These images also show the presence of the unerupted

successor P3 in both jaws, in close proximity and immediately

anterior to the smaller erupting dP3. The lower dp3 1s somewhat

more erupted than the upper one, and its successor p3 1s also

anterior to, but not as closely apposed, to dp3 as 1s the upper P3.

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ET /4

Figure 11. Camera lucida drawings of dp3 and p3 in the dentary, at different planes of section. A, The lingual successional lamina is fragmented,

it shows the fragmented lingual successional lamina between the two teeth. A, ameloblasts; AB, alveolar bone; D, dentin; dp3, deciduous third

premolar; E, enamel; O, odontoblasts; OE, oral epithelium; P, dental papilla; p3, successor third premolar; PL, primary dental lamina; SL, lingual

primary dental lamina connection to the oral epithelium is fragmented but still evident. The section through the successor p3 is not central, but

successional lamina; SR, stellate reticulum.

but still largely intact, between the small dp3 and its larger successional p3; B, The small dp3 is evident with its dentin and enamel, and its

Dental development in the thylacine pouch young

» ` ;

- “< 2 - = `

sA I

a 2

Wa, f ~

a n» ` 4 |

cogow CO Wet 6 aem. LU Wyr, ute

P. AE rg tht D REA p,

Figure 12. Section through dp3 and its successor p3. Only a small

fragment of the lingual successional lamina is evident. Compare this

single section with the camera lucida drawings in Figure 11. dp3,

deciduous third premolar; Isl, lingual successional lamina; p3,

successional third premolar.

Clearly, the complete dentitions of these thylacine PYs should

be carefully described and illustrated, and we hope that Newton

et al. (2018) will do so in a future publication.

A later stage of tooth eruption in Thylacinus available to us

is from a pouch young in the Australian Museum in Sydney.

This specimen (AM P 762) measured 80.5 mm HL and was

collected in Tasmania during 1866 (fig. 14). This PY was recently

estimated to be 12 weeks old (Newton et al., 2018). It was not

possible to section this young, but we were able to examine the

head using X-rays with the help of Dr Lucjan Sych, from the

School of Dentistry at the University of Melbourne. The images

of the head made by Dr Sych showed that the small spherical

dp3 was erupted bilaterally above the alveolar margins in the

dentary, with only minimal suggestions of distinct roots (fig.

14a, c). The unerupted, but larger, successor p3 lies immediately

anterior to the smaller dp3, as it was in our younger sectioned

pouch young and in the TMAG young described above. The

apex of dpl was just slightly above the alveolar margins, whereas

the larger dp2 was at or just below the alveolar margins (see fig.

14c). The lower ml protoconid is in an early stage of eruption

above the alveolar margins; m2 1s evident within its alveolus but

is not yet erupting, as is the less developed m3.

The upper jaw was more difficult to interpret in our X-rays,

and it was especially difficult to identify the dP3 and its state of

eruption. Fortunately, the recent publication by Newton et al.

(2018) on the available thylacine pouch young specimens

included X-ray computed tomography scans of AM P 762, and

we were able to examine this specimen in greater detail, thanks

to their inclusion of extensive supplementary material. We

noted that both of the tiny dP3 In the upper and lower jaws were

at least partly erupted, with greater eruption in the lower jaw

(fig. 15). The unerupted but well-developed successor P3 are

also clearly seen immediately anterior and deeper in both jaws,

as they were in our younger developing pouch young.

A later stage of dental eruption in Thylacinus was described

and illustrated by Moeller (1968). We were not able to examine

this specimen (CU A6 7/10) from the Cambridge University,

Museum of Zoology, but instead relied on the careful

description and figures supplied by Moeller (1968). He showed

that, in this specimen, I2 - 3 were erupting in the upper jaw,

whereas I1 and I4 were unerupted, but evident in their alveoli.

This 1s consistent with our observation of a delayed development

of I] in our sectioned pouch young. Moeller also showed that

dPl and dP2 were in early eruption, but there was no sign of

dP3, except for an alveolus that contained the unerupted P3. It

is unclear whether the poorly rooted dP3 (probably erupted)

was lost or damaged during preparation of the skull. The MI

was unerupted.

In the lower jaw, the canine Is unerupted, and dpl and dp2

are partly erupted, with dpl being more erupted. A small

spherical dp3 is erupted anterior to an erupting ml (see our

reproduction of Moeller's fig. 32a in fig. 16a). The apex of the

unerupted p3 1s evident In an alveolus anterior to the small dp3.

Clearly, Moeller's specimen is only slightly more advanced than

our pouch young (AM P 762) from the Australian Museum. A

slightly later stage of Thylacinus (USNM 115365; skull length =

87.75 mm) shows that dp3 has been lost in the dentary and its

successor p3 is in early eruption (fig. 16b). The ml is almost

completely erupted, and m2 is in early eruption. The dpl is

almost completely erupted, whereas the larger dp2 is In an earlier

phase of eruption.

Our stages for these and later development and eruption in

the upper and lower jaws of Thylacinus are presented in Tables

l and 2. Included in specimens from the dentary we have

presented some samples of fossil Thylacinus cynocephalus

from caves In Western Australia. Radiocarbon analysis of

charcoal samples from the Henschke Fossil Cave suggests that

the cave was filled 1n between 32,000 and 40,000 years ago,

trapping the thylacines and many other marsupial species

within (Pledge, 1990). Although these dentitions are 1n most

cases somewhat smaller than the more recent Tasmanian

thylacines, the developmental stages appear to be identical 1n

both groups. For instance, both p3 and m3 are erupting at

about the same time in the lower jaw.

Comparison of the thylacine pouch young with similar

developmental stages in dasyurids

In comparing our single specimen of the thylacine pouch young

with comparable developmental stages of dasyurids, presumably

one of its closest relatives within the order Dasyuromorphia, it

was interesting to note the occurrence of both similarities and

differences within the family Dasyuridae. Unfortunately, we were

unable to find a similar developmental stage of Myrmecobius (the

single representative of Myrmecobiidae) in our studies.

In Table 3, the thylacine pouch young has the successor P3

in the late bell stage, and M2 is in the early - middle bell stage in

the upper jaw. The closest similar developmental stage 1n our

dasyurid sample is Dasyurus viverrinus, a species in which dP2

has been lost In both jaws (as in all species of Dasyurus

examined by us). The Dasyurus PY has a 13.5 mm HL and its

P3 is in the late bell stage, with a thin layer of dentin. Its M2 is

in the middle - late bell stage. Given the slightly older specimen

of Dasyurus, it 1s quite similar In its developmental stage to our

10 W.P. Luckett, N. Hong Luckett & T. Harper

Table 1. Development and eruption of the upper postcanine dentition In Thylacinus cynocephalus, (HL = Head Length).

Moderately Tiny tooth,

thick enamel with

on apex; flat, moderately

short lingual thick enamel;

successional fragmented

lamina lingual

successional

lamina

extends

anteriorly

No trace

NMV C 5754 34 mm

HL pouch young; no

teeth erupting; estimated

31 - 32 days

No trace

Not evident

Not evident |Not evident

Moderately

developed to

moderately

thick enamel

on apex; short

lingual

successional

lamina

Large, late

bell stage, no

odontoblasts;

lingual and

mesial to tiny

dP3

Large tooth,

with

moderately

thick enamel

on tall

metacone

Moderately

large tooth, in

early -middle

bell stage

71.1 mm HL pouch

young (Flower, 1867)

Museum of the Royal

College of Surgeons

Tooth

calcified; not

erupting

Tooth

calcified; not

erupting

Well calcified

tooth; not

erupting

Tiny, rootless |Tooth deep in Not evident

jaw; beneath

and slightly

lingual to tiny

Partly

calcified

tooth; not

erupting

60 mm HL TMAG 930

Estimated 66 —67 days

Calcified

tooth; not

erupting

Calcified

tooth; not

erupting

Tooth not

erupting

Tooth not

erupting;

close to

alveolar

margins

Tooth evident

but not

erupting;

anterior to

dP3

80.5 mm HL AM P 762 Tooth evident

but not

Tooth evident

but not

Apex of tooth | Apex of larger | Tiny tooth Apex of

Haired pouch young;

estimated 84 days

erupting

alveolar

slightly above |below

tooth just

alveolar

unerupted

tooth evident

mesio-lingual

erupting

margins margins; not

erupting

to tiny dP3

Not evident — |Not evident

Not evident |Notevident [Not evident

Not evident Not evident

Skull length = 80 mm

Cambridge Univ.

A6 7/10

Tooth not

erupting

Tooth evident

in alveolus,

but not

erupting

No trace Large tooth;

very early

erupting, just

above alveolar

margins

Tooth erupting| Tooth erupting | Tiny tooth not

evident;

probably lost

USNM 115365 Juvenile

female (skull length —

87.75 mm)

Tooth almost |Moderately

completely X |large; about

erupted 2/3 erupted

Large tooth;

early erupting

(about 1/4)

(less erupted

than dP1)

Tooth erupted | Tooth erupted

Erupted Erupted No trace Erupted Erupted Erupted Evident In

alveolus; not

Erupted Erupted No trace Erupted Erupted Erupted

above alveolar

margins

MN Berlin An 13914

Juvenile (skull length —

129.4 mm)

Not evident

Not evident |Not evident

Large tooth;

erupted

SAM M1958 Juvenile

male (skull length =

130 mm)

Large tooth;

erupted

NMV C 5744 Juvenile

male (skull length =

157 mm)

Not evident

AM P 778 Juvenile

(skull length —

159.4 mm)

Not evident

Dental development in the thylacine pouch young

SAM M1956 Juvenile

(skull length = 149 mm)

NMV C 5743 Juvenile

(skull length =

153.5 mm)

NMV C 5600 Juvenile

female (skull length =

153.8mm); born In zoo;

18 months old

AM P778 Juvenile (skull

length = 159.4 mm)

MN Berlin A 1745

Juvenile (skull length =

168 mm)

AMNH 77701 Juvenile

(skull length =

191.2 mm)

WA F6358 Subadult

fossil (skull length =

138.2 mm)

AM S1180

Subadult; skull length =

180 mm

AM 775

Subadult; skull length =

182 mm

(collected 1866)

LAC A 3298

Adult; skull length =

192 mm

Other adult skulls, with

all teeth erupted; with

skull lengths from

192 mm to 253mm

Erupted Erupted No trace Erupted Erupted Erupted Early Not evident

emergence

above alveolar

margins

Erupted Early erupting | Not evident

(less than 1/3)

Erupted Early erupting | Not evident

(about 1/4)

Erupted Early Not evident

emergence

above alveolar

margins

Erupted Not distinct;

alveolus

only

Erupted Erupted Region

damaged;

uncertain

Erupted Erupted About 1/3

erupted

No trace Erupted Erupted

lš

Erupted

No trace Erupted Erupted

TI T e T

= = = =

£= a= "3 I

ct = — =

C C C C

aN O CL Le

12 W.P. Luckett, N. Hong Luckett & T. Harper

Table 2. Development and eruption of the lower postcanine dentition in Thylacinus cynocephalus, (HL = Head Length).

Stage dp, jae, an — |» m m m [m —

MV C 5754 Moderately |Moderately Large tooth, |Large tooth, No trace

with thick with

enamel on tall |moderately ` 'middle-late

protoconid thick enamel |bell stage;

dental lamina

ends distal to

thick enamel (thick enamel

Tooth deep in | Well calcified |Partly Not evident

jaw; beneath |tooth; not calcified l

and slightly — |erupting tooth; not

lingual to tiny erupting

dp3

on apex; short [on apex; short

Tiny tooth Unerupted ‘Tooth at Tooth evident |Apex evident |Not evident

erupted tooth apex alveolar but not but not

evident mesial margins, not |erupting erupting

to tiny dp3 erupting

flat lingual flat lingual

Tiny spherical | Apex of Protoconid of |Apex of tooth |Partly Not evident

tooth; erupted |unerupted large tooth in jevident just — |calcified tooth

above the tooth evident [early eruption |below deep in Jaws,

alveolar mesio-lingual alveolar not erupting

margins to tiny dp3 margins, but

Tiny, elongate |Large, late

tooth, with bell, with

moderately ` (moderately

thick enamel; |developed

lingual dentin; lingual

successional and mesial to

lamina smaller dp3

extends

mesial to

larger P3

34 mm HL pouch

young; no teeth

erupting; sectioned

histologically

estimated 31 - 32 days

old

71.1 mm HL pouch Calcified Calcified

young (Flower, 1867) |tooth; not tooth; not

erupting erupting

Tiny, rootless

tooth; apex

just above

alveolar

margins

TMAG A930 Probably not |Apex of large

erupting; tooth near

region alveolar

damaged margins

60 mm HL; estimated

66—67 days old

AM P 762 Apex of tooth | Apex of larger

erupting just [tooth just

above alveolar|above alveolar

margins margins

80.5 mm HL Haired

pouch young;

successional |successional

estimated 84 days old

not erupting

lamina lamina

Tooth partially | Tiny tooth Apex of Protoconid of |Protoconid ` [Tooth not Not evident

erupted (about |erupted above |unerupted large tooth in apex just erupting

1/3) alveolar tooth evident [early eruption |below

margins mesio-lingual alveolar

to tiny dp3 margins; not

erupting

Early erupting | No trace Alveolus only |Early Not evident — |Not evident

(about 1/3) (probably erupting;

| erupted) slightly less

than that of

dp2

Tooth erupting} No trace; Large tooth; [Large tooth; |Unerupted Not evident

possible erupted early eruption |tooth in

(1/3) alveolar crypt

Erupting tooth | Alveolus only |Alveolus only |Erupting tooth | Not erupting

for erupting |(about 1/3)

(or erupted)

tooth

shallow

buccal

for erupted

tooth

Erupting tooth | Not distinct

(about 1/2)

Cambridge University | Tooth almost

A6 7/10 completely

ted

80 mm skull length |)

(Moeller, 1968)

WAM 72.1.1149 Alveolus

only; at least

partially

erupted

Juvenile fossil right

dentary, Murray Cave,

Western Australia

USNM 115365 Tooth erupted

Juvenile female; skull

length = 87.75 mm

alveolus for

lost tooth

SAM P 33482 Partial

alveolus only

Double rooted | No trace

alveolus only

for erupting

(or erupted)

tooth

WAM 61-2-26 Tooth erupted |No trace

Juvenile fossil dentary,

Henschke Fossil Cave,

South Australia

Tooth erupting] Alveolus only

for erupted

tooth

Alveolus only

for erupted

Juvenile fossil left tooth

dentary, Mammoth

Cave, Western

Australia

Dental development in the thylacine pouch young 13

Roaches Rest Cave, Tooth erupted |No trace No trace;

fossil 67-3-21 probably

erupted

Berlin An 13914 Erupted Erupted No trace

Tooth erupted | Tooth erupted |Erupting tooth | Unerupted, in

deep alveolus

Erupted Erupted

Juvenile skull and

dentary (skull length =

129.4 mm)

SAM 1958 Erupted Erupted No trace Completely

Juvenile skull and erupted

dentary (skull length =

130 mm)

MV C 5744 Erupted Erupted No trace Erupted Erupted

Juvenile skull and

dentary (skull length =

157.1 mm) male

SAM M1956 Erupted Erupted No trace Erupted Erupted

Juvenile skull and

dentary (skull length =

149 mm)

LAC, Paris 1883-352 |Erupted No trace Erupted Erupted

Juvenile skull and

dentary damaged

(Collected in 1844)

MV C5743 Erupted Erupted No trace Erupted Erupted

Juvenile female (skull

length =153.5 mm)

AM P 778 Erupted Erupted No trace Erupted Erupted

Juvenile (skull length

= 159.4mm)

MN Berlin A 1745 Erupted No trace Erupted Erupted

Subadult (skull length

= 168 mm)

WAM F6358 Erupted Erupted

Subadult fossil (skull

length = 138.2 mm)

Nullabor Plain,

lli

ii i

Erupted Erupted Not evident

erupted (about

1/2)

Erupted

Early

emergence of

Western Australia

Adult female (skull

length = 192mm)

a m E

= = =

"a E= E=

= = =

C C C

14 W.P. Luckett, N. Hong Luckett & T. Harper

a P3

Figure 13. Computed tomography images from the supplementary data of Newton et al. (2018). A, section of the skull and dentition from TMAG

A931, a thylacine pouch young of 35 - 37 days old; B, Section of the skull and dentition from TMAG A930, a thylacine pouch young of 66 - 67

days old. Scale bars are 5 mm. C, canine; dP1, deciduous first premolar; dP2, deciduous second premolar; dP3, deciduous third premolar; M1,

first molar; P3, successional third premolar.

Dental development in the thylacine pouch young 15

Figure 14. Images of the head, skull, and dentition of the thylacine pouch young from the Australian Museum (AM P 762). A, X-ray of the skull,

showing deciduous and successional teeth in varying stages of development and early eruption; B, Head and upper body of the pouch young, prior

to X-ray analysis; C, Higher magnification of a portion of the X-ray shown in figure A, with emphasis on the erupted lower dp3, and the unerupted

but larger successional p3 immediately anterior to it.

W.P. Luckett, N. Hong Luckett & T. Harper

Figure 15. Computed tomography image of the skull and dentition of AM P 762, taken from the supplementary data of Newton et al. (2018). The

figure clearly shows the erupted dP3 in both jaws, and the unerupted successor P3 immediately anterior to their predecessors. Scale bar equals

5 mm. Other early erupting and unerupted teeth are clearly labeled. C, successional canine; dP1, deciduous first premolar; dP2, deciduous second

premolar; dP3, deciduous third premolar; M1, first molar; P3, successional third premolar.

Thylacinus PY. In the lower jaw, Dasyurus viverrinus is also

more similar in its developmental stages of the dentition with

Thylacinus than with the dasyurids with three premolars,

Antechinus and Sminthopsis (see Table 4). Note that there 1s an

accelerated phase of dental development in the lower jaw,

compared with that in the upper jaw for both Thylacinus and

Dasyurus. Thus, the lower p3 Is most similar with m3 in its

developmental stage, rather than with m2.

In contrast, the two dasyurid genera with three premolars

that we examined (Antechinus and Sminthopsis). are quite

different and considerably delayed or retarded In the

development of their successional P3, compared with Thylacinus

and Dasyurus. Thus, in Antechinus, the successor P3 Is only

suggested by the slight swelling of the lingual successional

lamina of dP3, and Sminthopsis has P3 In a very early bud stage

in the upper jaw (Table 3). However, the M2 of both genera with

three premolars are in a similar middle - late bell stage. Note

that the differences occur only in the development of the

premolars, and not the molars.

Although early developmental stages of Myrmecobius were

not available to us for comparison with the thylacine 34 mm

HL PY, we were able to compare later stages of development

of Myrmecobius with comparable stages of Thylacinus from

our Tables 1 and 2. In the upper jaw of a juvenile Myrmecobius

(WAM M 19214, with skull length 2 41.66 mm), P3 was about

half erupted and M2 was almost completely erupted. In the

lower Jaw, p3 was about 3/4 erupted and m3 was erupting.

These relationships are most similar to those of a juvenile

Thylacinus from the Berlin Museum (MN 13914; skull length

= |29.4 mm). In the upper jaw of the Berlin thylacine, P3 is

almost completely erupted and M2 is about 3/4 erupted (see

Table 1). In the lower jaw, p3 is almost completely erupted and

m3 is about 2/3 erupted (see Table 2). These relationships

suggest that Myrmecobius is most similar in its dental

development to the dasyurids with two premolars and with

Thylacinus, rather than with the dasyurids with three

premolars, despite the fact that Myrmecobius has three

premolars in both jaws.

Dental development in the thylacine pouch young

p3 In crypt

p3 erupting

te 9 '

`? w tst

Figure 16. Later stages of early eruption in Thylacinus showing the presence and early loss of dp3 in the dentary. A, Part of the dentary (CU A6

7/10), redrawn from Moeller (1968), showing the erupted dp3, the unerupted successor p3 in its alveolar crypt, immediately anterior to dp3, and

the erupting ml. The erupting dpl and dp2 are also labeled; B, A slightly later stage of eruption In the dentary (USNM 115365) shows that the

dp3 has been lost, and successor p3 1s in early eruption. The ml is now almost completely erupted. c indicates lower successional canine in B.

We were also able to examine a similar later stage of

development in Sarcophilus harrisii (AMNH 65674), a dasyurid

with two premolars, for comparison with Thylacinus. As In

other dasyurids with two premolars, and in the thylacine, dental

development is accelerated, with dPl, P3, and MI - 2 erupted in

both jaws. In the lower Jaw, m3 is almost completely erupted.

Although most authors have concluded that it is dP3 (and P3)

that is the missing tooth in Sarcophilus (Archer, 1976; Tate,

1947; Thomas, 1877; Wroe, 1999), our developmental studies

have demonstrated the presence of dP3 and P3 in both jaws of

Sarcophilus, as well as the absence of a developing dP2 in both

early and later stages, similar to the condition in Dasyurus

(Luckett et al., unpublished research).

For an additional comparison, we examined one specimen of

Peramelidae (Perameles sp.) and one Didelphidae (Monodelphis

domestica). Both were similar in developmental stages to the

dasyurids with three premolars (see Tables 3, 4). The successor

P3 for Perameles sp. (16 mm HL) was in the early bud stage, and

M2 was in the late bell stage with early odontoblasts in the upper

jaw. In the lower jaw, Perameles sp. was in the early bud stage

for p3, and m3 was in the late bell stage, with early odontoblasts.

In Monodelphis domestica (14.5 mm HL), P3 was in the early -

middle bud stage and M2 was in the late bell stage in the upper

Jaw. In the lower jaw, p3 was in the late bud - early cap stage and

m3 was in the middle bell stage. Note that the molars are in

similar developmental stages for all taxa examined, whereas it IS

the successional P3 that varies in both jaws.

18 W.P. Luckett, N. Hong Luckett & T. Harper

position of p3 < SH

s tro "UR Fš "ee ae

. < See, ARTT STY

Figure 17. Selected dentaries of Thylacinus showing differences in the diastemata between the premolars as the effect of increasing age. A,

subadult (in labial view) with m3 erupted, but not m4. Only slight suggestions of diastemata are evident between the premolars; B (lingual view)

and C (labial view), showing later stages of m4 eruption and the increase of diastemata in adults.

Relationship of paracone and metacone in thylacines and including Nimbacinus (Muirhead and Archer, 1990), Badjcinus

other marsupials (Muirhead and Wroe, 1998), Wabulacinus (Muirhead, 1997),

In Thylacinus cynocephalus, the metacone is larger than the Ngamalacinus (Muirhead, 1997) and Mutpuracinus (Murray

paracone on MI - 3. A similar condition has been found alsoin and Megirian, 2006). It is likely that a similar condition to that

numerous Late Oligocene-Miocene fossil thylacinids, discovered in Thylacinus and didelphids, in which the apical

Dental development in the thylacine pouch young

Table 3. Comparison of development In Thylacinus and Dasyuridae upper postcanine dentitions, (HL = Head Length).

Thylacinus (34mm Moderately

HL) estimated 31 - 32 |thick enamel

days

Moderately — |Tiny tooth;

moderately

thick to thick

Moderately

thick dentin

and enamel

Dasyurus viverrinus

(13.5 mm HL) About

35 days

shallow bony

alveolus

Middle - late

bell stage

Moderately

thick dentin;

Moderately

thick dentin

and enamel

Antechinus stuartii

(9.5 mm HL) 34 days

successional

lamina

Late bell stage | Moderately

thick dentin

and enamel

Sminthopsis virginiae

(8.8 mm HL) 30 days

dentin; thin

enamel

Moderate Moderate

sized, late bell |sized, late

stage; thin

dentin

Perameles sp. 16 mm Moderate

HL sized, late

bell; possible

early

odontoblasts

Monodelphis Well-

domestica (14.5 mm |developed

HL) dentin and

enamel

Well-

developed

dentin and

enamel enamel

epithelial nodule is detached from the developing paracone but

not from the metacone, 1s a causative factor in this relationship.

It would be interesting to know whether some Oligocene -

Miocene thylacinids also had a more molariform and

functional dP3, with a tall metacone, than in Thylacinus

cynocephalus. Unfortunately, we are unaware of any findings

of a dP3 In these earlier fossil thylacinids .

Some comments and criticisms concerning similarities and

differences between thylacines and other dasyuromorphians

and with marsupials in general

It 1s often noted that Thylacinus cynocephalus differs from

other thylacinids because of the pronounced diastema between

the premolars in its adult dentitions. Indeed, 1n some cases this

has been used in character analyses within thylacinids and

other dasyuromorphians (see Muirhead and Wroe, 1998;

Yates, 2014). This, however, is simply a factor that increases

Late bell

stage; thin

dentin on apex|thick dentin,

Small early

bud stage

Small early

welldeveloped |-middle bud

Large, late

bell stage; no

odontoblasts;

deeper bony

alveolus

Large tooth; |Moderately — [No trace No trace

moderately ` |large, early

thick dentin |- middle bell

and enamel on stage

tall metacone

Moderately ` |No trace No trace

large, middle

-Jate bell stage

bell stage

Early bud

Stage

Large tooth;

moderately

thin enamel

on tall

metacone

Moderately

thick dentin;

moderately

developed

enamel

Late bell

stage

Moderately

thick dentin;

moderately

developed

enamel

Large tooth, |Large, late

with

moderately

thick dentin

and

moderately

developed

Small, early

- middle bud

bell with thin

dentin

enamel

Large tooth; Moderately |Early bud

moderately {large late bell, |stage

with

moderately

thick dentin

and enamel

developed

dentin

with age in Thylacinus cynocephalus. If we examine some

younger juvenile to subadult stages, as shown in Figure 17a,

there are few, 1f any, spaces separating dpl dp2, and p3. In a

juvenile (NM V C 5744; skull length = 157.1 mm) in which dPl

- 2, P3 and MI - 2 are erupted In both jaws, and m3 is erupted

in the lower Jaw, the diastema in the upper jaw measured 2.0

mm between dP1 and dP2, and also between dP2 and P3. In

the lower jaw, dpl and dp2 are separated by a diastema of 1.5

mm, as are dp2 and p3. With increasing age and completion of

eruption of M4 in both jaws, there is an increase in the length

of the diastemata (Fig. 17b, c). We have examined and

measured the diastema In a large number of subadult and adult

thylacines (measuring192 mm to 253 mm skull length for the

adults with all teeth erupted), and this clearly demonstrates

that these diastemata increase with age, between the premolars,

but not between the molars. As an example, in the oldest (1.e,

largest) adult skull examined by us (AM P 767, measuring

W.P. Luckett, N. Hong Luckett & T. Harper

Table 4. Development of Thylacinus cynocephalus and dasyurid lower postcanine dentitions, (HL = Head Length).

Tan | dp | dp | dp | p | m | m | m | m _

Moderate

sized, thick

enamel and

disrupted

dentin

Tiny tooth,

with thick

Moderately

large, with

thick enamel

Thylacinus

cynocephalus

NMV C 5754 34 mm

HL; estimated 31 - 32

days

enamel and

disrupted

and disrupted

dentin

distinct cusp

Moderately

developed

dentin, thin

enamel

Dasyurus viverrinus

23 - 25 mm GL (35

days)

Thin dentin;

no enamel

Moderately

thick dentin,

thin enamel

Sminthopsis virginiae

thick dentin,

10 mm HL (35 days) thin enamel

Middle-late

bell stage

Moderately

thick dentin,

Moderately

thick dentin,

moderately

developed

enamel

Antechinus stuartii

9.5 mm HL (34 days)

Small tooth;

late bell; thin

dentin

Moderately

large, late

bell; early

odontoblasts

Moderately

large, late

bell; very thin

dentin

Perameles nasuta

bud

35 mm GL (16 mm

Large tooth;

thick dentin

Moderately

sized tooth;

moderately

thick dentin

and enamel

Large

caniniform

tooth; thick

dentin and

enamel

about 253 mm skull length), the teeth were heavily worn, and

the diastema in the upper jaw measured 3.3 mm between dPl

and dP2, and 4.9 mm between dP2 and P3. In the lower jaw,

the diastema was 5.2mm between dpl and dp2, and 5.8mm

between dp2 and p3. Considerable variation occurred between

the sizes of the diastemata in both jaws of all adults examined.

We suggest that the presence, absence and size of diastemata

between premolars should be used with caution in assessing the

phylogenetic relationships among fossil thylacinids, especially

when only one or two specimens are known for any fossil species.

Another comment refers to the continued use of the terms

PI and P2 in the adult jaws of fossil and extant thylacinids, and

in other marsupials in general. In all marsupials that have been

examined histologically from early developmental stages up to

the eruption of all teeth, there is no evidence known to us for

the replacement of dP1 or dP2 by a successor P1 or P2 in the

upper or lower jaws of any fossil or extant marsupial. This has

been emphasised previously for both fossil (Cifelli et al., 1996)

and extant (Luckett, 1993a, b; Luckett and Woolley, 1996)

marsupials. Authors who continue to use the terms “P1 and

P2” should present evidence for the replacement of their

Large, late

bell stage; thin thick dentin

to moderately land enamel on [thick dentin

developed

dentin; single |dentin

Tiny abnormal | Early - middle | Moderately

slight

swelling of

lingual

successional

lamina

Slight

swelling of

lingual

successional

lamina

Lingual early

Moderately

large, middle

- late bell

Stage

Large tooth;

moderately

and enamel on

protoconid

Not developed

deciduous predecessors in marsupials, as is the case for dP3

and P3. Similar usage of the term "PI" instead of dP] within

fossil and extant eutherians has been discussed recently in a

study on Eocene juvenile perissodactyls (Rose et al., 2018).

A final, very positive, observation on thylacine biology and

evolution. We were pleased to see the publication on development

of the immune system (Old, 2015) in the pouch young of

Thylacinus that was based on the histological sections of NM V C

5754 from Museums Victoria. Hopefully, other investigators will

study additional aspects of the cranial and postcranial biology in

this specimen. We are also happy to note the recent publication

of the thylacine genome (Feigin et al., 2018), thanks to the use of

material from the sister (NMV C 5757) of our sectioned pouch

young in the Museums Victoria collection (see fig. 2b).

Middle - late

bell stage

Moderately

thick dentin;

thin enamel

Middle - late

bell stage

Moderately

thick dentin;

thin enamel

thick dentin;

thin enamel

Middle - late

bell stage

Thick dentin,

moderately

thick enamel

Moderately

thick dentin,

moderately

developed

enamel

Moderately

large, late

bell, with

early

odontoblasts

Large tooth;

dentin and

thin enamel

on protoconid

Large tooth;

Moderately

large tooth, in

Large tooth;

thick dentin

Acknowledgements

This research was carried out in great part due to the financial

support provided to WPL by the Visiting Scholar Program at

the University of Melbourne, through the efforts and support of

Dr Marilyn Renfree, and by a similar program at La Trobe

Dental development in the thylacine pouch young

University, Bundoora, Victoria, through the support of Dr

Patricia Woolley during the years 1992-1996. In addition to the

generous help provided by Joan Dixon and Lina Frigo at

Museums Victoria, many additional people provided great

assistance at the other museums visited in Australia. These

included Michael Archer, Tim Flannery and Linda Gibson at

the Australian Museum in Sydney; Peter Baverstock and

Neville Pledge at the South Australian Museum, Adelaide; and

Ken Aplin at the Western Australian Museum, Perth, among

many others.

Special thanks of appreciation go to Axel Newton and his

colleagues for providing unlimited access to their thylacine

pouch young specimens; these helped greatly in our

understanding of the later stages of unerupted and early erupting

phases of the thylacine pouch young. Ken Rose provided

numerous constructive criticisms that greatly 1mproved our

manuscript. Adam Sylvester helped considerably in the

preparation of the figures.

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Memoirs of Museum Victoria 78: 23—55 (2019) Published 2019

1447-2554 (On-line)

htto://museumvictoria.com.au/about/books-and-journals/journals/memoirs-of-museum-victoria/

DOI https://doi.org/10.24199/).mmv.2019.78.02

Taxonomic revision of dragon lizards in the genus Diporiphora (Reptilia: Agamidae)

from the Australian monsoonal tropics

(http://zoobank.org/urn:|sid:zoobank.org:pub:08B3925A-6720-44E4-BF1C-EED106581DD4)

JANE MELVILLE'*, KATIE SMITH Date!, PAUL HORNER’ AND PAUL DOUGHTY?

! Department of Sciences, GPO Box 666, Museum Victoria, Melbourne, Victoria, 3001, Australia.

* Museum and Art Gallery of the Northern Territory, GPO Box 4646, Darwin, Northern Territory, 0801, Australia (current address:

Cannonvale, Queensland, 4802, Australia).

* Department of Terrestrial Zoology, Western Australian Museum, 49 Kew St, Welshpool, Western Australia, 6016, Australia.

* Corresponding author: jmelv@museum.vic.gov.au

Melville, J., Smith Date, K.L., Horner, P., and Doughty, P. 2019. Taxonomic revision of dragon lizards in the genus

Diporiphora (Reptilia: Agamidae) from the Australian monsoonal tropics. Memoirs of Museum Victoria 78: 23—55.

The Australian dragon lizard genus Diporiphora currently comprises 21 species based on genetic and morphological

evidence, with 11 of these species occurring in the monsoonal tropics of northern Australia. Diporiphora are climbing

lizards that are found on either trees, grasses or rocks, with usually only subtle morphological differences to distinguish

between species. Since the last taxonomic treatment of this genus 1n northern Australia over 40 years ago, species delimitation

using genetic techniques has clarified the number of lineages and increased collections from recent surveys have significantly

broadened the distributions of these taxa. However, no formal taxonomic assessments have been undertaken to redefine

species, including the many lineages that represent undescribed species. Currently, there are seven species of Diporiphora

with vast distributions across northern Australia and a broad and variable set of morphological characteristics that make

species identification challenging, even for experienced field workers. Here, we provide a comprehensive taxonomic

treatment of Diporiphora species across northern Australia based on previously published genetic data and morphological

examination of voucher specimens. Our analyses demonstrate that these broadly distributed taxa actually comprise multiple,

often allopatric, species, with especially high diversity in the Kimberley region of Western Australia.

We redescribe nine previously described species and describe five new species of Diporiphora based on historical

types, newly collected material and older museum vouchers. In the D. australis species group, we resurrect D. jugularis

Macleay from synonomy. In the D. bennettii species group, we synonomise D. arnhemica Storr with D. albilabris Storr,

and raise to full species the latter and D. sobria Storr. In addition, we describe as new a wide-ranging saxicoline species

previously attributed to D. bennettii Gray. In the D. bilineata species group, we resurrect D. margaretae Storr from

synonomy with D. magna Storr and describe three new species. Lastly, we describe a species from the northwest Kimberley

The revision of the northern Diporiphora dragons here stabilises the taxonomy, redefines many species distributions

and reveals many new species. Further work on Diporiphora includes further surveys to better understand distributions

and habitat preferences and continue to refine their evolutionary history and biogeography in northern Australia.

Agamid lizards; Australia; Cape York Peninsula; Diporiphora carpentariensis sp. nov.; Diporiphora gracilis sp. nov.;

Diporiphora granulifera sp. nov.; Diporiphora pallida sp. nov.; Diporiphora perplexa sp. nov.; Kimberley; monsoonal

Abstract

that is more closely related to an arid zone radiation.

Keywords

tropics; taxonomy; Top End.

Introduction

The Australian monsoonal tropics (AMT) constitute a

significant element of the Australian landmass, spanning the

northern quarter of the continent from the far west coast of the

Kimberley region in Western Australia (WA), across the Top

End of the Northern Territory (NT) and around the Gulf of

Carpentaria to the Cape York Peninsula in Queensland.

Tropical savannah woodlands and grasslands are the region’s

dominant vegetation types and are characterised by a dense

grass understorey and an over-storey of widely scattered trees

(Bowman et al., 2010). These woodlands are home to a major

component of Australian biodiversity, with sandstone

escarpments possibly having similar levels of diversity to the

rainforests (Bowman et al., 2010). Despite the importance and

size of this region In terms of phylogenetic diversity, there are

many vertebrate groups for which there remain significant

taxonomic problems, particularly in terms of under-

representing the true species diversity (e.g. Doughty 2011;

Laver et al., 2018; Moritz et al., 2018; Pepper et al., 2011).

One group with such taxonomic problems is the dragon

lizard genus Diporiphora (family Agamidae, sub-family

Amphibolurinae), which is a diverse group of slender small-

bodied dragons, including some of the most common and

widespread diurnal lizards across the tropical savannah

woodlands. This genus presents particular difficulties in

species-level identification, with many of the morphological

characters used in keys and diagnoses being as variable within

species as between species. The last significant taxonomic

treatment of Diporiphora in this region of northern Australia

was published more than 40 years ago, describing eight species

and subspecies (Storr, 1974). More recently, genetic work has

shown that the current taxonomy significant under-represents

— (b) D. bennett) species group

(a) D. australis species group

D. amphiboluroides

D. pindan

_* |

| |o. valens Arid Zone

t D. adductus Species group

D. paraconvergens

D. winneckeí/ D. ameliae / D. linga

— D. reginae

D. superba

Pogona

Tympanocryptis * | l

- Tropicagama

Rankinia

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

the true species diversity (Couper et al., 2012; Edwards and

Melville, 2011; Smith et al., 2011).

We undertook a comprehensive taxonomic review of

Diporiphora species across northern Australia, with particular

focus on the taxa that have been identified as species complexes

In previously published and unpublished genetic work (fig. 1).

Although Storr’s (1974) work provides a good framework

towards resolving the taxonomy of northern Diporiphora, many

issues remain. As was typical of Storr’s work, his revision was

based on collections at the Western Australian Museum

(WAM); he did not address Queensland, despite Diporiphora

occurring across the Gulf of Carpentaria region including the

Cape York Peninsula.

Storr (1974) recognised considerable diversity within

several species and described multiple subspecies to

accommodate this variation: D. albilaris (D. a. albilaris, D. a.

sobria), D. bennettii (D. b. bennettii, D. b. arnhemica) and D.

(a) D. australis species group

D. australis

D. phaeospinesa

D. jugularis

: jug

(b) D. bennettii species group

D. perplexa sp. nov.

D. sobria

D. albilabris

D. bennettii

— D. bilineata

D. magna

D. falling

D. gracilis sp. nov.

| D. margaretae

D. granulifera sp. now.

D. campentanensis sp. rav.

[ | "o [ | Available names raised to

Mew species |

species

[ | Change in species | | Unchanged, taxonomic

| distribution | | treatment in this work

Figure 1. Bayesian 50% majority-rules phylogenetic tree for Diporiphora based on mtDNA on ~1200 bp mitochondrial DNA (ND2). Asterisks

on branches represent >99% posterior probability support. Clades highlighted 1n green are expanded, with phylogenetic relationships within each

of the species groups reviewed in the current paper: a, D. australis; b, D. bennettii; c, D. bilineata. Species reviewed in the current paper are

coloured to represent the taxonomic revision that is undertaken.

Taxonomic revision of dragon lizards

Table 1. Meristic characters scored and morphological characters measured in this study

Meristic characters

Pre-cloacal pores

Femoral pores

Canine teeth

Guar fold

Post-auricular fold

Scapular fold

Dorsal scales

Throat stripes

Position of lateral dark spot

Paravertebral and dorsolateral scales

Vertebral scales

Ath toe lamellae (4TL)

Infralabials (ILB)

supralabials (SLB)

Measurements

SVL

Tail (TL)

Head length (HL)

Head width (HW)

Interlimb length (IL)

Forelimb (FLL)

Hindlimb (HLL)

Number of pre-cloacal pores in total

Number of femoral pores in total

Number of canine teeth on each side of upper Jaw

Absent/weak/present

Absent/weak/present/strong

Absent/weak/ present/strong

Homogenous/weakly heterogeneous/strongly heterogeneous

Absent/weak/strong

On side of neck/axilla (arm pit)

Enlarged or homogenous with surrounding scales

Number of enlarged subdigital lamellae under fourth toe, counted from toe junction to base of claw

Number of infralabial scales, ending with the last small scale in contact with the posterior margin of the

last upper labial

Number of supralabial scales

Snout-vent length

Tail length from cloaca to tip (unbroken tails only)

Head length, measured obliquely from tip of snout to anterior margin of tympanum

Head width, measured at the widest point

Length from the axilla to the groin

Length, measured from the elbow bend to the tip of the 4th finger including claw

Length, measured from the knee bend to the tip of the 4th toe including claw

bilineata (D. b. bilineata, D. b. margaretae). At the time of

Storr’s revision, subspecies were fashionable to employ to deal

with the variation observed and to hedge against taxonomic

uncertainty, especially with the relatively small number of

specimens he had to work with. Since Storr's revision, the

concept of these species has changed according to various

authors and field guides but without any formal taxonomic

treatment. For example, the subspecies D. b. arnhemica 1s

usually recognised as a full species (e.g. Cogger et al., 2014;

Wilson and Swan, 2017). Additionally, the distributions of

these subspecies have significantly expanded since Storr's

(1974) original descriptions. For example, D. b. arnhemica

was originally limited to the type location, D. a. sobria was

limited to two locations in the central NT and D. a. albilabris

was limited to four locations In the northwest Kimberley.

subsequently, the distributions of these taxa have expanded to

encompass vast areas of northern Australia (e.g. Cogger, 2014;

Wilson and Swan, 2017).

Taxonomic problems, such as those outlined above, extend

across all northern Diporiphora species. Storr (1974) described

D. b. margaretae as a new subspecies of D. bilineata, with

distributions of the two subspecies (D. b. bilineata and D. b.

margaretae) defined as being disjunct in the NT and northern

Kimberley, respectively, with the intervening savannah

woodlands occupied by D. magna (also described in the same

work). Finally, D. lalliae was described as a generalist species

with a broad east-west distribution across the southern

portions of the AMT (Storr, 1974). Since the description of

these species and subspecies, many of the distributions of

these lineages have expanded to be widespread and broadly

sympatric across the AMT and even into the arid zone (D.

lalliae) without further taxonomic treatment. Storr apparently

changed his mind about the legitimacy of D. b. margaretae,

omitting it from Storr et al. (1963) without comment 10 years

later. This change has been largely followed since (e.g. Cogger,

2014; Wilson and Swan, 2017) despite being listed as a full

species by Cogger et al. (1983). Thus, the currently used

taxonomy of northern Australian Diporiphora does not reflect

storr's original work and a full taxonomic assessment of these

lizards 1s long overdue.

Table 2. Diagnostic characters to distinguish between lineages

(a) D. australis

species group

l. No. of canine teeth on each One Two

side of upper Jaw

2. Axilla granular scales Absent Absent

On side of neck

Absent

3. Lateral dark spot

4. Femoral pores

(b) D. bennettii

species group

Axial if present

Present (D. albilabris,

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

species group species group

One One

Present Absent

Exception: absent in

D. lalliae

Axial Absent

Absent Absent

D. sobria) or absent

Here, we provide the first comprehensive taxonomic

treatment of Diporiphora species across northern Australia

since Storr’s work, using an understanding of relationships based

on genetic data and an examination of newly collected and

historical collections, including type specimens.

Methods

Specimens examined and external morphology

Specimens were examined from the collections of the Western

Australian Museum (WAM), Queensland Museum, Brisbane

(QM), Museum Victoria (NMV), Museum and Art Gallery of

the Northern Territory (MAGNT), Darwin, Australia Museum,

Sydney (AMS), the Natural History Museum, London (NHMUK,

registration prefix — BMNH), the Naturhistorisches Museum,

Vienna (NHMW) and the Swedish Museum of Natural History

(NHRM). New type material is deposited in the WAM and QM.

A set of 13 meristic and seven morphometric characters

(Table 1) were recorded for all primary types. Measurements

were made with electronic callipers to the nearest 0.1 mm,

except for snout-vent length (SVL) and tail length (broken tails

were not measured), which were measured with a ruler to the

nearest 0.5 mm. Individuals were sexed by observations of

everted hemipenes in males or heavily gravid females, or by

direct examination of the gonads. Aside from the type series,

an additional >10 well-preserved adult specimens (where

available) from each taxon were examined (listed in Appendix

1). On the dorsum, we refer to the vertebral scale row at the

midline, with the paravertebral scale row adjacent and with

the 2nd paravertebral scale row (often enlarged and hence

referred to in the text) to the outside of the paravertebral row.

We detail both meristic and morphological characters to

diagnose species and to identify the species groups from each

other. In the Comparisons with other species sections, we

compare the focal species with other species that have

overlapping distributions.

We provide a taxonomic revision of each of the species

groups outlined in fig. 1. This encompasses most of the

Diporiphora that occur across the AMT and a species

description for a taxon that falls outside these species groups

but occurs within the AMT:

(a) D. australis species group

(b) D. bennettii species group

(d) arid zone species group.

Five Diporiphora species that occur in the northern tropical

savannahs are not reviewed in the current study. Diporiphora

superba Storr is extremely elongate and green or yellow, and is

easily distinguished from all other Australian dragon lizards

(Storr et al., 1983; Wilson and Swan, 2017). The type of D.

convergens Storr was examined as part of this study but an

account is not included because this specimen has the keels of

the dorsal scales converging towards the midline, which is

unique in AMT Diporiphora. This species 1s only known from

the type specimen, which was collected from Crystal Creek in

the northern tip of the Kimberley (Storr, 1974) and has not

been collected again. Taxonomic treatments of D. nobbi

(Witten), D. phaeospinosa Edwards and Melville or D. pindan

storr are also not provided because they have been recently

treated elsewhere (Doughty et al, 2012a; Edwards and

Melville, 2011).

Phylogenetic analyses

The phylogenetic tree (fig. 1), based on a region of ~1200 bp

mtDNA (including the protein coding gene ND2 and five

tRNAs) across 283 Diporiphora samples, provides strong

support for all lineages covered in this taxonomic review. The

ND2 coding region and flanking tRNA regions were found to

follow the GT R+I+G model of substitution with no partitioning

schemes using the corrected Akaike information criterion on

PartitionFinder2 on the CIPRES Science Gateway (Lanfear et

al., 2016). Bayesian analysis was performed using MrBayes

(Huelsenbeck and Ronquist, 2001) on the CIPRES Science

Gateway, with two runs of four independent Markov chain

Monte Carlo analyses (each 50,000,000 generations long and

sampled every 1,000 generations) under a GTR+I+G model

with flat priors. Tracer v1.6 was used to check for stationarity

and convergence of the chain outputs. The trees were subject

to a 25% burn-in in MrBayes, summarised and posterior

probabilities obtained.

Taxonomic revision of dragon lizards

Diporiphora Gray, 1842

Type species. Diporiphora bilineata Gray, 1842, by monotypy.

A diverse genus, distributed across all states and territories,

except Tasmania, predominately in semi-arid, arid and AMT

biomes. Small- to medium-sized lizards that are mostly terrestrial

or semi-arboreal. Moderately long snout with relatively long legs

and tail. All species have exposed tympanum, and most species

have pre-cloacal pores with femoral pores occasionally present.

Generally lacking prominent crests, large spinose scales or

dermal appendages, although present In some species (e.g. D.

amphiboluroides). Colour patterning variable within species,

particularly between breeding males and females or juveniles. In

general, most species have pale dorsolateral stripes, dark

transverse dorsal bars, a black circular blotch on shoulders or

neck, and a pink to mauve flush on hips and tail base (particularly

in adult males).

(a) D. australis species group (fig. 1a)

Content:

Diporiphora australis (Steindachner, 1867)

Diporiphora jugularis (Macleay, 1877)

Diporiphora nobbi (Witten, 1972)

Diporiphora phaeospinosa Edwards and Melville, 2011

Diagnostic characters for group (Table 2):

° one canine tooth on each side of upper jaw (see fig. 2 for

illustration of this character)

* scales in axillary region not granular

* lateral dark spot on side of neck.

(a) D. bennett! species group

Figure 2. Images from micro X-ray computed tomography scans

showing the differences in pleurodont (canine) tooth number in the

upper Jaw: a, arrangement in D. bennettii species group; b, other

species groups included in the current study.

Remarks

This species group Is restricted to the eastern portion of the

AMT (fig. 3), occurring along the east coast, adjacent inland

areas and throughout the Cape York Peninsula. Relative to other

species groups in the AMT, they tend to be large bodied (68—75

mm SVL) with generalist habits. We do not include D. nobbi and

D. phaeospinosa below because Edwards and Melville (2011)

recently reviewed these species. Our analyses of all Diporiphora

species (fig. la) recovered the same relationships among species

within this group as the mtDNA phylogeny in Edwards and

Melville (2011), with D. australis and D. jugularis being highly

supported as sister species, but the relationships between this

lineage and D. phaeospinosa and D. nobbi are not well resolved.

We provide an account of D. australis and D. jugularis, raising

the latter species from a junior synonym of D. bilineata (Cogger

et al., 1983) based on genetic results in Edwards and Melville

(2011) and examination of types and specimens herein.

Diporiphora australis (Steindachner, 1867)

Common name. Tommy roundhead.

Figure 4, Tables 3, 4

Calotella australis Steindachner, F. 1867. Reptilien. pp. 1—98 1n,

Reise der Osterreichischen Fregatte Novara um die Erde in den Jahren

18657, 1858, 1559 unter den Befehlen des Commodore B. von

Wüllerstorff- Urbair. Zoologie 1(3). State Printer: Vienna. [1869 on title

page| [29, pl. 1 fig. 9]. Type data: lectotype - NHMW 19821.1 Australia

("Cape York, QLD” on type label), taxonomic designation, this work.

Grammatophora macrolepis Günther, A. 1867. Additions to the

knowledge of Australian reptiles and fishes. Annals and Magazine of

Natural History 20: 45—68 [51]. Type data: holotype - BMNH

1946.8.12.74, Australia (by implication).

@ D. jugularis

@ D. australis

Figure 3. Distributions of D. australis and D. jugularis based on

specimens examined and collection records.

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

Table 3. Measurements and sex of types. All measurements are in millimetres (mm) and key to abbreviations can be found in Table 1. Museum

abbreviations are as follows: Western Australian Museum (WAM); Australian Museum, Sydney (AMS); Queensland Museum (QM); Natural

History Museum, United Kingdom (BMNH); Naturhistorisches Museum, Vienna, Austria (NHMW)

Species Accession No. Sex

D. australis NHMW 19821:1 female

D. jugularis AMS R40672 Juvenile

D. jugularis AMS R40673 juvenile

D. jugularis AMS R40674 juvenile

D. bennettii BMNH 1946.8.12.77 female

D. albilabris WAM R435 17 female

D. sobria WAM R23180 subadult

D. perplexa sp. nov. WAM R177290 male

D. bilineata BMNH 1946.8.12.75 male

D. bilineata BMNH 1946.8.12.76 male

D. lalliae WAM R23020 male

D. magna WAM R42786 female

D. margaretae WAM R27648 female (gravid)

D. gracilis sp. nov. WAM R177291 male

D. granulifera sp. nov. QM J96362 male

D. carpentariensis sp. nov. QM J88197 male

D. pallida sp. nov. WAM R177292 male

* = tail broken

Diporiphora nuchalis De Vis, C.W. 1884. On new species of

Australian lizards. Proceedings of the Royal Society of Queensland |:

97-100 [98]. Type data: syntype(s) — whereabouts unknown, central

and south-coast district, Queensland.

Diporiphora ornata De Vis, C.W. 1884. On new species of

Australian lizards. Proceedings of the Royal Society of Queensland |:

97-100 [99]. Type data: holotype — whereabouts unknown, locality

unknown.

Physignathus nigricollis Lonnberg, E., and Andersson, L.G. 1915.

Results of Dr. E. Mjóberg's Swedish Scientific Expeditions to

Australia 1910-1913. VII. Reptiles collected in northern Queensland.

Kongliga Svenska Vetenskaps-Academiens Nya Handlingar,

Stockholm 52: 1-9 [4] [incorrect spelling as Physignatus nigricollis].

Type data: holotype - NHRM 3209, Cooktown, Queensland.

Diagnosis. Body size moderately large (to 70 mm SVL) with

long tail (2.3-2.7 x SVL). Gular, post-auricular and scapular

folds present. Lacks spinose scales on thighs or neck. Pre-

cloacal pores 4; femoral pores 0.

Description of lectotype. Female; 69 mm SVL; 146 mm tail

length. Medium-sized dragon, moderately robust with

moderately long limbs and tail. Single canines on each side of

upper jaw. Gular and post-auricular folds present and scapular

fold weak. Dorsal scales homogenous but prominent keels form

longitudinal ridges along dorsum along midline with

dorsolateral ridges on each side; these raised vertebral and

dorsolateral scale rows extend anteriorly onto nape to head and

posteriorly onto base of tail, not extending down tail. Scales on

SVL TL HL HW IL HLL FLL 4TL SLB ILB

69 146 18 12 33 49 27 23 11 12

38 9 l2 7 l 35 22 23 ll 12

35 82 12 8 l 34 22 23 11 12

30 65 11 8 l 30 19 22 10 11

5 68 15 13 24 30 23 17 ll 10

44 94 l4 9 20 40 24 18 10 10

34 «68 12 8 14 25 19 18 11 12

63 140 22 11 24 52 31 21 11 11

58 | 56* 18 13 24 46 26 23 11 10

57 131 17 le 4 42 24 22 10 11

62 152 18 11 3M 44 27 B 9 9

47 141 15 9 20 42 25 23 ll 11

55 128 16 9 24 4l 21 19 11 13

57 165 18 10 20 49 27 23 11 9

65 176 21 12 29 46 3l 23 11 12

62 185 20 12 28 55 33 22 11 11

46 95 15 10 24 40 24 16 10 11

flanks heterogeneous with scattered slightly enlarged scales.

Scales on neck, limbs and tail not spinose. Scales in axilla

small but not granular. Ventral scales strongly keeled. Pre-

cloacal pores 4; femoral pores 0.

Dorsum strongly patterned. Vertebral and dorsolateral

lines very faint, almost undiscernible with colouring a slightly

paler replication of dorsum patterning, running from back of

head to pelvis. Five wide dark transverse bands across dorsum

between head and pelvis. Dark bands are a similar width to the

pale background. Head relatively plain with little patterning,

labials similar colour to rest of head and lacking pale line

between eye and ear. Lacks dark spot on posterior of

tympanum. Flanks a similar colour to dorsum. Lacks lateral

stripe between axilla and groin. Dark patch on sides of neck at

anterior edge of shoulder. Arms weakly banded with dark

bands slightly narrower than light. Legs and tail strongly

banded with dark bands narrower than light. Ventral surface

cream with no patterning.

Variation. 46—10 mm SVL; 105—174 mm tail length. Tail long,

ranging from 2.3—2.7 x SVL. Strong scapular fold present, post-

auricular fold usually absent although present but weak In a few

individuals. Dorsal scales homogenous but prominent keels form

longitudinal ridges. There are often five longitudinal ridges: one

along the vertebral midline, one each on the 2nd paravertebral

scale rows and one on each of the dorsolateral stripes. Some

individuals only have these longitudinal ridges running along the

Taxonomic revision of dragon lizards 29

Table 4. Diagnostic characters distinguishing species within species groups

Tail Dorsolateral Gular Post- Scapular Dark spot Extent of axial Dorsal scales Pre- Femoral

length stripes fold auricular fold on dark granular between cloacal pores

(x SVL) fold tympanum scales dorsolateral pores (total)

edge stripes (total)

(a) D. australis

species group

D. australis Long Present Present — Weak/ Present Absent Absent Homogeneous 4 0

(2.3—2.7) absent

D. jugularis Lone Usually | Absent Absent Absent Absent Absent Homogeneous 4 O

(1.8—2.7)

(b) D. bennettii

species group

D. albilabris Long Present Present Strong Absent Absent Absent Strongly 4 2

(1.8—2.5) heterogeneous

D. bennettii Short Absent Present Strong Weak Absent/ Absent Homogeneous 2 0

(<2.0) weak

D. perplexa sp. nov. Long Present Weak Strong Weak Strong Absent Homogeneous 2-4 0

(1.9-2.5)

D. sobria Long Present Present Weak/ Weak/ Absent/ Absent Homogeneous in 4 2

(2.3—2.6) present present weak Western Australia

(some Northern

Territory

populations

heterogeneous)

species group

D. bilineata Long Present Absent Absent Present Absent Over arm onto Moderately 2 0

(2.2—2.7) shoulder and heterogeneous

posteriorly with outer row of

along flanks, dorsolateral stripe

with pale having weakly

flecks raised trailing edge

D. gracilis sp. nov. Very Sometimes Absent Weak/ Weak Absent Overarmto Homogeneous 4 O

lone absent scapular fold lacking raised

(2.7—3.0) scales In

dorsolateral stripes

D. lalliae Very Usually Present Strong strong Absent Absent Homogeneous 4 O

long lacking raised

scales in

Ur dd) dorsolateral stripes

D. magna Very Present Absent Strong Strong Absent Overarm but Homogeneous 4 0

long not to sides of lacking raised

2630 neck scales in

(5:099) dorsolateral stripes

D. margaretae Long Present Absent Weak/ Weak/ Absent |Ontop of arm Homogeneous but 4 0

25.27 present present and posteriorly with outer row of

Gc T] along flanks, dorsolateral stripe

with pale having raised

flecks trailing edge

D. granulifera sp. | Very Present Absent Weak— Strong Absent Overarmand Homogeneous but 4-6 O)

nov. lone strone along the full with outer row of I!

27.30 length of the dorsolateral stripe pu: >;

s scapular fold: having raised )

anterior to trailing edge

scapular fold

small, slightly

eranular scales

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

Tail Dorsolateral Gular Post- Scapular Dark spot Extent of axial Dorsal scales Pre- Femoral

length stripes fold auricular fold on dark granular between cloacal pores

(x SVL) fold tympanum scales dorsolateral pores (total)

edge stripes (total)

D. carpentariensis Very Present Absent Weak- Strong Absent Over armand Homogeneous 4-5 V

Sp. nov. long strong along the full lacking raised "

21-30 length of the scales in ms M

(2 scapular fold dorsolateral stripes )

(d) Arid zone

species group

D. pallida sp. nov. Short Absent Present Absent Present Absent Absent Homogeneous 2 O

(2.0)

second scales out from the midline, giving a smoother appearance

to the dorsum. Some individuals have weakly heterogeneous

dorsal scales with the 2nd paravertebral scale rows slightly

enlarged. In individuals with weak post-auricular fold, a row of

4—6 enlarged but not spinose scales may be present.

Dorsal pattern variable from strong, complex patterning to

unpatterned. Dorsolateral stripes always present, wide cream to

grey vertebral stripe present in all but weakly patterned

individuals. In strongly patterned individuals, usually six dark

transverse bands across dorsum between head and pelvis but

ranges from 4—7. Transverse bands similar in size or slightly

narrower than pale background. Flanks in these individuals

have similar patterning to dorsum and commonly have wide

pale lateral stripe between axilla and groin. In weakly patterned

individuals, transverse bands are either absent or are narrow

dark bands contacting the dorsolateral stripes and extending

towards, but not contacting, the vertebral stripe. In these plain

animals, the bands on the tail also contact the dorsolateral

stripes but do not meet at the midline. Flanks in these individuals

lack patterning, including a lateral stripe between axilla and

eroin. Dark patch on sides of neck often present but variable

from being absent to running along scapular fold and extending

anteriorly onto the sides of neck from the dorsal extent of the

fold, with the dark pigmentation contacting the pale dorsolateral

stripe. This dark spot on the neck never extends posteriorly

from the scapular fold towards the axial region or onto the

ventral surface. Ventral surface usually unpatterned and cream

in colour, but a few individuals, usually females, have pale grey

colouring on gular region.

Distribution and ecology. Widespread along the north-eastern

coast of Australia, as far south as the Coffs Harbour area in

northern New South Wales, extending north along the coast

into the south-eastern portions of Cape York Peninsula as far

north as the Cooktown area (fig. 3). It extends inland

approximately 600 km from the coast, across the base of Cape

York Peninsula to the Normanton area. It extends south along

the Great Dividing Range, into northern New South Wales.

A common and abundant species that is an ecological

generalist, occurring in most dry forests and woodlands within

its distribution (fig. 5c and e). Individuals use low vegetation,

fallen timber and termite mounds as perches. This species 1s a

common sight in some of the northern cities and appears to

have adapted well to bushy suburban areas (Kutt et al., 2011).

Although D. australis is one of the most common species

encountered in suburban areas of eastern Queensland,

relatively little is known about its biology.

(b)

Figure 4. Diporiphora australis: a, adult, Karawatha, south-eastern

Queensland (photo: S. Wilson); b, lectotype NHMW 19821.1,

Australia (“Cape York, QLD” on type label); c, ventral view of head

showing gular fold.

Taxonomic revision of dragon lizards

Comparison with other species. Overlaps extensively with the

distribution of D. nobbi and overlaps broadly with D. jugularis

at the northern edge of its range (fig. 3) and with D.

carpentariensis sp. nov. in the north-western extent of its range.

It differs from D. nobbi in having smaller body size and lacking

spinose scales on the thighs or neck. It can be distinguished

morphologically from D. jugularis and D. carpentariensis sp.

nov. by possessing a gular fold.

Remarks. As part of this study we examined all available types,

including those of synonymised species, held in Australian and

European collections. In addition, we examined a specimen

Figure 5. Examples of the range of habitats in which the Diporiphora species of the Australian monsoon tropics occur: a, sandstone escarpment,

Mitchell Plateau, Kimberley region, Western Australia; b, rocky outcrops in savannah woodlands, western Arnhem Land, Northern Territory; c,

savannah woodlands, Kimberley region, Western Australia; d, savannah grasslands on cracking clay soils, floodplain of the Lennard River,

Kimberley Region, Western Australia; e, savannah woodlands, western Arnhem Land, Northern Territory; f, arid spinifex grasses with scattered

trees on stony ground, Tennant Creek, Northern Territory (photos: J. Melville).

held in the NHMUK that is housed in the type collection based

on an account by Günther (1867). This account named

Grammatophora calotella as a replacement name for a species

described earlier in the same year by Steindachner, who

described Calotella australis, basing his name on specimens in

the Vienna collection. Günther renamed Calotella australis as

the species Grammatophora calotella and referred to the two

specimens in the NHMUK from Cape York. However,

examination of the Grammatophora calotella specimen

revealed that it lacks a gular fold, indicating that it is D.

jugularis from Cape York, and not C. australis. In the same

paper, Günther described Grammatophora macrolepis as a

new species, which examination confirms is D. australis.

Cogger et al. (1983) suggested that the two specimens with

the single registration number of NHMW 19821 were likely to

be the syntypes and that these bore the locality “Cape York”,

suggesting that they were collected by Daemel. As such, they

would have been collected by Daemel at the same time as the

Grammatophora calotella specimens in the NHMUK (BMNH

66.12.28.22—23). However, it is probable that the location of

these specimens is incorrect and they were not from Cape

York but were from Rockhampton. Comparison of the

lectotype NHMW19821:1 to the image provided in the original

description (fig. 6) suggests that this specimen was the one

described by Steindachner but with incorrect locality data.

There are several species with types collected by Daemel

where there is a mix-up between specimens collected in Cape

York and Rockhampton (G. Shea, personal communication).

The type of Physignathus nigricollis Lönnberg and

Andersson was also examined as part of this study and was

confirmed to be a specimen of D. australis.

(a)

Figure 6. Lateral images of the Diporiphora australis syntype NHMW

19821:1: a, Naturhistorisches Museum, Vienna (photo: J. Melville); b,

taken from the original species description (Steindachner, 1867).

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

Diporiphora jugularis (Macleay, 1877)

Common name. Black-throated two-pored dragon.

Figure 7, Tables 3, 4

Grammatophora jugularis Macleay, W. 1877. The lizards of the

Chevert Expedition. Second paper. Proceedings of the Linnean

Society of New South Wales 2: 97—104 [1878 on title page] [104]. Type

data: syntype(s) — AMS R40672-4, Cape Grenville, Cape York

Peninsula, Queensland.

Diporiphora brevicauda De Vis, C.W. 1884. On new species of

Australian lizards. Proceedings of the Royal Society of Queensland 1:

97—100 [99]. Type data: holotype — whereabouts unknown, Cape York

Peninsula, Queensland. E

Diporiphora pentalineata De Vis, C.W. 1884. On new species of

Australian lizards. Proceedings of the Royal Society of Queensland 1:

97—100 [99]. Type data: holotype — whereabouts unknown, Cape York

Peninsula, Queensland.

Diagnosis. Body size moderately large (to 68 mm SVL) with

long tail (1.8—2.7 x SVL). Gular and scapular folds absent;

post-auricular fold absent or weak. Scales in axilla not granular.

Lacks spinose scales on thighs or neck. Pre-cloacal pores 4;

femoral pores 0.

(a)

(b)

10 mm

Figure 7. Diporiphora jugularis: a, adult male with breeding colouration,

Iron Range, Queensland (photo: S. Wilson); b, syntype(s) AMS R40672—

4, juveniles, Cape Grenville, Cape York Peninsula, Queensland.

Taxonomic revision of dragon lizards

Description of syntypes. Juveniles; 30—3 mm SVL; 65—96 mm

tail lengths. Small dragons, moderately robust with moderately

long limbs and tail. Single canines on each side of upper Jaw.

Gular, scapular and post-auricular folds absent. Dorsal scales

homogenous but raised keels form four prominent longitudinal

ridges along dorsum, with a ridge on either side of midline

running along the 2nd paravertebral scale row, and a

dorsolateral ridge on along each side. These raised vertebral

and dorsolateral scale rows extend anteriorly to neck,

terminating at head and extending posteriorly onto base of tail.

Scales on flanks strongly heterogeneous with scattered scales

that are distinctly larger than rest, with keels angling away

from dorsum towards ventral surface from midbody. Scales on

neck, limbs and tail not spinose. Scales in axilla small but not

eranular. Ventral scales strongly keeled, keels forming

longitudinal ridges running length of body. No pre-cloacal

pores visible in these juvenile specimens (usually 4); femoral

pores 0.

Dorsum not patterned, which may be due to preservation.

Lack vertebral and dorsolateral lines. Dark transverse bands

across dorsum absent. Heads plain without patterning, labials

same colour to rest of head and lacking pale line between eye

and ear. Lacking dark spot on posterior of tympanum. Flanks

same colour as dorsum. No lateral stripe between axilla and

eroin. Faint dark pigmented patch on either side of neck at

anterior edge of shoulder. Arms, legs and tail plain. Ventral

surface cream with no patterning.

Variation. 54—68 mm SVL; 99-178 mm tail length. Tail long,

ranging from 1.8—2.7 x SVL. Gular, scapular and post-auricular

folds absent in all individuals examined. Dorsal scales

homogenous but prominent keels form longitudinal ridges.

There are often five longitudinal ridges: one along the vertebral

midline, one on either side midline running along the 2nd

paravertebral scale row and one on each of the dorsolateral

stripes. However, some individuals only have these longitudinal

ridges running along the 2nd paravertebral scale row, giving a

smoother appearance to the dorsum. Weakly heterogeneous

scales In some individuals, with the scale rows second out from

the midline being slightly enlarged. Spinose scales absent, but

some individuals have 4—6 enlarged (not spinose) scales where

a post-auricular fold would occur. Pre-cloacal pores 4; femoral

pores 0.

Dorsal patterning variable from plain to strongly patterned.

Adult males often have little patterning, but with strongly

keeled dorsolateral ridge of scales with a pale cream colour

forming prominent dorsolateral stripes. Flanks on these

individuals are dark brown with the scattered enlarged pale

brown or cream scales giving a speckled or flecked appearance,

lacking lateral stripe. Patterned individuals, typically adult

females and juveniles, have approximately 4—7 broken dark

transverse bands across dorsum on a pale background. Dark

bands are of similar width or slightly broader than pale

background. These bands are broken by a pale grey or light

brown vertebral stripe. The dark transverse bands continue

laterally beyond the pale cream dorsolateral stripes, becoming

diffuse and terminating at a poorly defined lateral stripe. Wide

dark band on sides of neck mostly present, extending from the

dorsolateral stripe to the gular ventral surface and anteriorly

from where a post-auricular fold would be to the anterior of the

shoulder. The dark band is prominent in males and smaller or

occasionally absent in females. This dark gular band does not

extend onto chest but forms a well-defined posterior edge

across the gular region. In adult males, the dark gular band

extends anteriorly to the labials, while in individuals with

smaller gular bands, the apex of the small central triangle

extends anteriorly. Individuals without the dark gular band may

have a dark spot on either side of the neck or a narrow grey

band across gular. Legs and tail banded in strongly patterned

individuals, otherwise little patterning on legs and tail.

Distribution and ecology. Restricted to the eastern edge of the

Cape York Peninsula, extending as far south as Mount Misery,

southwest of Cairns (fig. 3). Widespread along the eastern

extent of Cape York Peninsula, extending inland 150 km from

the coast. Little is known about the ecology of this species, but

presumed to be similar to that of its sister species D. australis

in the NT. If so, it would be a generalist species that occurs In

dry open forests, woodlands and shrublands.

Comparison with other species. Overlaps geographically with

D. australis, D. nobbi and D. carpentariensis sp. nov. It differs

morphologically from both D. australis and D. nobbi in lacking

a gular fold. Also differs from D. nobbi in being smaller in

body size (65 mm vs. 75 mm SVL), lacking spinose scales on

the thighs or neck, having fewer pre-cloacal pores and lacking

femoral pores. It differs from D. carpentariensis sp. nov. In

that it lacks a scapular fold, scales in axilla are not reduced in

size and not granular, scales on flanks strongly heterogeneous

with scattered scales that are distinctly larger than surrounding

scales, and a black gular band or black spots on sides of neck.

Remarks. Diporiphora jugularis 1s restricted to the Cape York

Peninsula and has been referred to as D. bilineata owing to the

lack of a gular fold (e.g. Cogger, 2014; Wilson and Swan, 2017).

The distribution maps for D. bilineata usually show a

distribution (e.g. Cogger, 2014; Wilson and Swan, 2017), with

most of the range In the NT extending across the Gulf of

Carpentaria region of Queensland and onto the Cape York

Peninsula. However, genetic work has shown that the Cape

York Peninsula population is not closely related to D. bilineata

and 1s instead the sister lineage to D. australis (Edwards and

Melville, 2010), from which it is distinguished by lacking a

eular fold (as opposed to a gular fold being present in D.

australis).

There are a number of names available for Diporiphora on

the Cape York Peninsula. Examination of specimens held in

the type collection at the NHMUK showed that the earliest

known treatment of this species was in 1867 by Günther. As

detailed above for D. australis, Grammatophora calotella 1s a

replacement name created by Günther In 1867 for a species

described earlier in the same year by Steindachner. The

Grammatophora calotella treatment was based on BMNH

66.12.28.22—23, but these specimens are the taxon from Cape

York Peninsula (rie. D. jugularis). However, because

Grammatophora calotella Günther, 1867 is a replacement

name for Calotella australis Steindachner, 1867 and the

account contains no description of that species, these BMNH

specimens have no type status. Thus, the first valid description

of this species was in 1877: Grammatophora jugularis

Macleay.

(b) D. bennettii species group (fig. 1)

Content:

D. albilabris Storr, 19774

D. bennettii (Gray, 1845)

D. perplexa sp. nov.

D. sobria Storr, 19774

Diagnostic characters for group (Table 2):

° two canine teeth on each side of upper jaw (fig. 2)

e scales in axillary region not granular

e lateral dark spot in axilla, if present.

Remarks

The D. bennettii species group is widely distributed across

northern Australia from the Kimberley to north-western

Queensland (fig. 8). This species group has moderate size overall,

with D. sobria and D. perplexa sp. nov. attaining maximum sizes

of «70 mm SVL, and with the two Kimberley endemics reaching

only 61 mm SVL (D. albilabris) or 55 mm SVL (D. bennettii).

A key character in distinguishing the D. bennettii species

group from the other species groups In northern Australia, and in

particular the broadly sympatric D. bilineata species group, 1s

the number of canine teeth on each side of the upper jaw (fig. 2).

Agamid lizards have two kinds of teeth: acrodont, which are

fused to the jaw and constitute most of the teeth running along

the extent of the jaw, and pleurodont, which sit in sockets

(Richman and Hadrigan, 2011). The canines in Diporiphora

are pleurodont teeth. In all members of the D. bennettii species

group, there are two canines on each side of the upper jaw,

where the posterior pleurodont tooth is usually slightly larger

than the anterior tooth, a possible consequence of the tooth

replacement process. In contrast, all of the species in the D.

bilineata and D. australis species groups have a single canine

on each side of the upper jaw. This variation in tooth number

250

O D. albilabris

^ D. bennettii

O D. perplexa

* D. sobria

Figure 8. Distributions of D. albilabris, D. bennettii, D. sobria and D.

perplexa sp. nov. based on specimens examined and collection records.

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

IS consistent within these species groups. However, in some

Juvenile and hatchling lizards, the pleurodont teeth may be

absent or not fully enlarged, so this character 1s best used to

distinguish adult lizards from each species group.

Owing to the structure of the tree 1n this group (fig. 2c),

major changes to the current taxonomy are required. From the

molecular data, we recovered a sister relationship between true

D. bennettii (1.e. a small-bodied north-west Kimberley form)

and D. albilabris albilabris. Sister to this pair was a monotypic

lineage with specimens assigned to either D. albilabris sobria

or D. bennettii arnhemica. which indicated that the latter two

subspecies were a single species. Lastly, an unnamed species

that was assigned to D. bennettii but lacked the short body and

tail of the holotype was sister to these three lineages.

To resolve conflicts with subspecies and stabilise the

taxonomy, several changes were necessary. First, D. albilabris

albilabris was raised to full species because its sister taxon ts the

distinctive true D. bennettii. Next, we raised D. albilabris sobria

to full species and synonomised D. bennettii arnhemica with D.

sobria owing to page precedence in Storr’s (1974) original

descriptions (sobria — p. 135; arnhemica — p. 137). With the

description of the widespread taxon usually attributed to D.

bennettii (1.e. D. perplexa sp. nov.), we feel stability has been

achieved in this species group from these taxonomic changes.

Should further work show structure in the D. sobria lineage,

then the name D. arnhemica would be available 1f it corresponded

to an unnamed lineage. But at this point, there is insufficient

evidence from genetic work and morphology to warrant

recognition of two taxa within this group.

Diporiphora albilabris Storr, 1974

Common name. White-lipped two-lined dragon.

Figure 9, Tables 3, 4

Diporiphora albilabris albilabris Storr, G.M. 1974. Agamid

lizards of the genera Caimanops, Physignathus and Diporiphora in

Western Australia and Northern Territory. Records of the Western

Australian Museum 3: 121—146 [133]. Type data: holotype - WAM

R43517, Mitchell Plateau, WA [14° 48'S, 125° 50' E].

Diagnosis. Body size moderate (to 61 mm SVL), tail moderately

long (from 1.8-2.5 x SVL). Gular and post-auricular folds

present; scapular fold absent. White dorsolateral stripes on

raised scale rows. Lacks dark smudge on posterior of tympanum.

Scales between dorsolateral stripes are heterogeneous (fig. 10a),

with paravertebral row reduced, 2nd paravertebral row enlarged.

Pre-cloacal pores 4; femoral pores 2.

Description of holotype. Female. 44 mm SVL, 94 mm tail length.

Small body size, moderately robust with long limbs and tail. ‘Two

canines on either side of upper jaw. Gular and post-auricular

folds present, scapular fold absent. Dorsal scales heterogeneous,

with dorsolateral row of enlarged, strongly keeled white scales

running from back of head to pelvis and forming well-defined

white dorsolateral stripes. Paravertebral rows, either side of the

vertebral scale row, are reduced in size compared with adjacent

scale rows, leading to the scales of the paravertebral and adjacent

scale rows being strongly heterogeneous. To the outer edge of the

white dorsolateral stripes, scales are relatively small and keeled,

Taxonomic revision of dragon lizards

with keels angling toward ventral surface at midbody. Scales on

flanks are heterogeneous with scattered slightly enlarged scales.

Scales on limbs and tail not spinose, a row of enlarged pale

spines on the post-auricular fold. Scales in axilla not granular.

Ventral scales strongly keeled. Pre-cloacal pores 3 (2 right; |

left); femoral pores 2.

Dorsum pattern complex. Wide (~3 scales) poorly defined

grey vertebral stipe and prominent white dorsolateral lines from

back of head to base of tail. Five wide dark transverse bands

between dorsolateral stripes between head and pelvis. Dark

bands extend irregularly from the outer side of the white stripes,

fading into the colour patterns on the sides. Dark bands are

narrower, approximately half the width, than the pale interspaces

with background colour. Head strongly patterned: poorly defined

line of pale scales from back of eye, over top of ear towards white

dorsolateral stripes, although not contacting stripes; labials pale

cream, with a few darker flecks, extending as a broad pale band

along jaw to back of head and ending at the enlarged spinose

scales on the post-auricular fold. Lacks dark spot on posterior of

tympanum. Lateral surfaces distinct from dorsum, with sides

dark brown with light brown spots, the spots consisting of

(D)

Figure 9. Diporiphora albilabris: a, adult male in breeding colouration

(registered specimen NMV D73860) from King Edward River

crossing, Western Australia (photo: J. Melville); b, dorsal view of

holotype WAM R43517, Mitchell Plateau, Western Australia.

clusters of 2—4 pale scales on the dark brown scales. Lateral

stripe between axilla and groin absent. Lateral dark spot absent

Arms weakly banded with dark bands approximately the same

width as intervening pale colour. Banding on legs not well

defined, dark bands on anterior two-thirds of tail, with dark

bands wider than the light bands, fading to unbanded for

remainder of tail. Gular region with six grey longitudinal stripes

ending at gular fold, ventral surface of torso and tail with cream

colouration and unpatterned.

Variation. 43—61 mm SVL; 92-125 mm tail length. Tail long,

ranging from 1.8—2.5 x SVL. Usually a spine above tympanum,

with several spines along post-auricular fold. A short row of

spines extends from post-auricular fold to above tympanum,

variably expressed with most reaching less than half way to

tympanum. Nuchal crest absent to very low in profile. On

dorsum, paravertebral row sometimes approaches the size of

vertebral row, but is typically reduced in size. The enlarged

dorsolateral row decreases in size posteriorly, reaching normal

size at the base of the tail.

Dorsal patterning variable and complex with 5—6 dark brown

bands from nape to legs between dorsolateral white stripes,

intersected by a poorly defined vertebral stripe of grey,

approximately 3 scales wide. These dark bands extend irregularly

to the outer side of the white stripes, fading into the colour

patterns on the sides, which are dark brown with light brown

spots. Dorsal patterning tends to be obscured in males in full

breeding colouration, where they have strongly contrasting

charcoal black, white and chestnut or orange-red colouring on

head and upper body, with ventral surface of head, chest and tail

also having an orange-red flush in some individuals. Labial

scales pale cream with a few darker flecks, extending as a broad

pale band along jaw to back of head ending at the enlarged

spinose scales on the post-auricular fold. Ventral surface of head

often has grey longitudinal stripes ending at gular fold. Ventral

surface of body and tail unpatterned and cream in colour.

Distribution and ecology. Restricted to the central and northern

Kimberley region (fig. 8). Most records are from the Mitchell

Plateau and Prince Regent River National Park, with other

records from Kalumburu, Theda and Doongan stations. No

specimens known from islands.

A habitat generalist occurring in tropical savannah

woodlands and grasslands. Although it occurs in savannah

woodlands, it appears to be found 1n rockier areas within these

habitats, including laterite, basalt and sandstone. Observed to

perch on rocks or termite mounds.

Comparison with other species. The distribution overlaps a

number of other Diporiphora species in the Kimberley. From

D. sobria it can be distinguished in lacking a scapular fold and

having strongly heterogeneous scales between the pale

dorsolateral stripes. It differs from D. perplexa sp. nov. In

having heterogeneous dorsal scales, 2 (vs. 0) femoral pores and

no dark markings on the tympanum. It differs from D. magna

in having a gular fold, femoral pores, double canine teeth in

upper jaw and white labial scales. It differs from D. margaretae

in having a gular fold and double canine teeth on each side of

upper jaw.

Figure 10. Images of dorsal scales of D. albilabris and D. sobria,

depicting: a, heterogeneous dorsal scales in D. albilabris; b,

homogeneous dorsal scales in D. sobria from Western Australia; c,

moderately heterogeneous dorsal scales in D. sobria from northern

and eastern Northern Territory.

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

Remarks. Originally a nominate subspecies, elevated here to full

species (see account above). Although, based on mtDNA, this

species IS sister to D. bennettii, with which it overlaps in

distribution, it Is quite different in appearance. Diporiphora

albilabris has a more typical Diporiphora appearance of slender

body shape and contrasting markings on the dorsum, whereas D.

bennettii has a more derived morphology of small body size,

short tail length and lack of highly contrasting dorsolateral

stripes. Diporiphora albilabris has a relatively small distribution

in the northern central Kimberley, yet it is reasonably abundant

where it occurs and not under immediate threat, making its

conservation status likely to be of least concern.

Diporiphora bennettii (Gray, 1845)

Common name. Kimberley sandstone dragon.

Figure 11, Tables 3, 4

Gindalia bennettii Gray, J.E. 1845. Catalogue of the specimens of

lizards in the collection of the British Museum. British Museum:

London. xxvii4289 pp. [247]. Type data: holotype — BMNH

1946.8.12.77, “N.W. Coast of Australia”.

Diagnosis. Small body size (to 55 mm SVL) and short tail («2.0

x SVL), with robust habitus and wide head with short snout.

Gular fold present, post-auricular fold strong, scapular fold

weak. Dorsals homogeneous, scales in axilla not granular. Lacks

Figure 11. Diporiphora bennettii: a, adult from Little Mertens Falls

area, Mitchell Plateau, Western Australia. (photo: S. Wilson); b,

holotype BMNH 1946.8.12.77, from the “NW coast of Australia”,

showing dorsal view. x

Taxonomic revision of dragon lizards

dorsolateral stripes. Usually lacks dark smudge on tympanum,

although some individuals have a faint dark spot on posterior

edge of tympanum. Pre-cloacal pores 2; femoral pores 0.

Description of holotype. Female; 50 mm SVL; 68 mm tail length.

small body size with wide head and short wide neck. Two

canines on either side of upper jaw. Gular fold present. Strong

post-auricular fold, weak scapular fold. Post-auricular fold with

1-3 spines. Dorsal scales mostly homogeneous, strongly keeled

with keels parallel to midline forming lines running

longitudinally down dorsum from shoulders to one third of the

way down the tail; enlarged scales associated with vertebral or

dorsolateral stripes are absent. Scales on flanks homogeneous,

with keels angled towards dorsum. Lacks spinose scales on

limbs or tail. Scales In axilla not granular. Weakly keeled scales

on ventral gular region and strongly keeled scales on ventral

torso. Pre-cloacal pores 2; femoral pores 0.

Dorsum patterning faint and diffuse. Lacks vertebral or

dorsolateral stripes. Lacks visible dark transverse bands

between head and pelvis. Head virtually unpatterned, without

pale labials or pale line between eye and ear. Faint dark spot

on posterior of tympanum. Flanks similar to dorsum, with

sides of torso being slightly darker than dorsum with a few

light brown spots. Lateral stripe between axilla and groin

absent. Lateral dark spot absent. Very faint banding on lower

hind limbs, dark banding on tail, with light bands much wider

than darker bands. Arms lack dark banding. Ventral surface of

head, torso and tail unpatterned and cream colouration.

Variation. 41—55 mm SVL, 53—83 mm tail length. Tail short, less

than 2.0 x SVL. Gular fold ranges from weak to strong, always

with reduced scales along fold, even if skin does not fold over;

post-auricular fold strong and scapular weak. Head triangular

with usually pronounced adductor muscles, snout short. Typically,

one or two enlarged spinose scale above tympanum and on post-

auricular fold, with 0—3 adjacent slightly enlarged scales. No

conspicuous nuchal crest, but occasionally slightly raised scales

present. Dorsal scales homogeneous with no abrupt demarcation

of dorsal and lateral scale sizes. Keels of scales on sides angled

dorsally and posteriorly. Scales above cloaca on sides of tail at

base with raised keels. Tail short, terminating 1n a blunt tip.

Background colour an admixture of dark brown, tan and

pale white scales generating a rather diffuse background pattern

with no highly contrasting dorsolateral stripes. Dark brown

cross-bands obscure, but in some heavily marked individuals

they can form loose networks that enclose pale spots. In some

individuals, especially juveniles, there are alternating wide dark

brown blotches and thinner pale lines along the dorsum,

although these are usually interrupted by variable admixture of

different-coloured scales. Lateral surfaces tend to be darker

with pale stippling. Tails with thin dark bands with wide pale

interspaces. Usually lacks dark spot on tympanum. Ventral

surface pale with occasional dark stippling on chin, sometimes

extending to ventrum.

Distribution and ecology. Restricted to the northern Kimberley

(fig. 8), with many records from the Mitchell Plateau and Prince

Regent River National Park, extending east to Drysdale River

National Park and including a few neighbouring offshore

islands: Augustus, Darcy and Unwins.

Collection records (over 40) are nearly all from sandstone

rock outcrops and escarpments with spinifex cover.

Comparison to other species. This species Is similar to D.

perplexa sp. nov., with extensive distributional overlap and

occurring in similar habitats, but differs in having a much

smaller body size, a short tail and no dorsolateral stripes. It

differs from D. albilabris 1n having a smaller body size, a short

tail, no femoral pores, homogeneous dorsal scales and in pattern

by lacking any prominent vertebral or dorsolateral stripes. It

differs from D. magna and D. margaretae in having a smaller

body size, a short tail, a gular fold and a pair of enlarged canines

on each side of upper jaw. It differs from D. pallida sp. nov. 1n

having two canine teeth on each side of upper jaw.

Remarks. The species was previously believed to include what

we describe below as D. perplexa sp. nov. Diporiphora

bennettii, however, 1s the sister species to D. albilabris (fig. 1b;

also restricted to the northern Kimberley), with these two being

sister species to D. sobria, which 1s now understood to be

widely distributed In the southern AMT. D. perplexa sp. nov.

occurs over a broad area In the Kimberley and extends to the

NT, and accordingly, D. bennettii is now known to be restricted

to the north-west Kimberley. Diporiphora bennettii has an

unusual morphology for a Diporiphora, owing to its small

body size, short tail, robust habitus, obscure dorsal patterning

and close association with sandstone outcrops in the northern

Kimberley. The redefinition of this species and reduction of its

range adds to the large number of endemic reptiles and frogs

from this region (e.g. Powney et al., 2010; Palmer et al., 2013).

Diporiphora perplexa sp. nov.

ZooBank LSID: http://zoobank.org/urn:Isid:zoobank.org:act:

4A6A3ASB-5FBI-40D5-843B-258D095238F3

Common name. Kimberley rock dragon.

Figure 12, Tables 3, 4

Holotype. WAM R177290 (formerly NMV D73819) (adult male),

Gibb River Rodd, 20 km west of Ellenbrae Station, WA (15° 57.31' S,

126° 52.9' E). Collected by J. Melville on 9 September 2005.

Paratypes. NMV D73805 (adult female), Home Valley Station,

WA (15° 44.39' S, 127° 49.83' E); NMV D73841 (adult female), King

Edward River Campground, Mitchell Plateau Road, WA

(14° 56' 57.1" S, 126° 12' 10.4" E; NMV D73978 (adult male),

Buchanan Highway, south of Jasper Creek, NT (16° 02' 46.8" S.

130° 51' 49.3" E); NMV D73980 (adult female), Bullo Road off

Victoria Highway, NT (15° 48' 39.2" S, 129° 40' 13.5" E); WAM

119719 (male), Emma Gorge, Cockburn Range, WA (15° 50' S,

128° 02' E); WAM R162517 (female), 25 km S Wyndham, WA

(15.7154° S, 128.2684" E); WAM R171418 (male), Prince Regent River

National Park, WA; WAM R175785 (female), Waterfall Yard, 15 km N

Mt Elizabeth Homestead, WA (16.2822° S, 126.1059° E).

Diagnosis. Body size moderately large (to 68 mm SVL) with

long tail (1.9—2.5 x SVL). Gular and scapular folds present but

weak; post-auricular fold strong. Homogeneous dorsal scales.

Pale dorsolateral stripes from back of head to one-third down

torso. Black smudge on posterior edge of tympanum, extending

on to scales posterior to tympanum. Pre-cloacal pores 2—4;

femoral pores 0.

Figure 12. Holotype of Diporiphora perplexa sp. nov. (WAM R177290,

formerly NMV D73819): a, in life — adult male In breeding colouration

from Gibb River Road, west of Ellenbrae Station, Western Australia;

b, c, d, preserved specimen in dorsal, ventral and lateral (head) views.

Yellow arrow highlights a key diagnostic character: dark pigment

“smear” on posterior of tympanum spreading onto neighbouring head

scales (photos: J. Melville).

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

Description of holotype. Male; 63 mm SVL; 140 mm tail length.

Moderately stout with long tail and limbs. Two canines on

either side of upper jaw, with posterior canines extremely

enlarged relative to anterior canines. Gular fold present but

weak. Strong post-auricular fold, weak scapular fold. Dorsal

scales homogeneous, strongly keeled with keels parallel to

midline forming lines running longitudinally down dorsum

from shoulders to base of the tail; enlarged scales associated

with vertebral or dorsolateral stripes not conspicuous. Scales

on flanks homogeneous, with keels on posterior flanks angled

towards dorsum. Lacks spinose scales on head, limbs and tail.

Scales in axilla small but not granular. Ventral scales weakly

keeled in gular region and strongly keeled on body. Pre-cloacal

pores 2; femoral pores 0.

Dorsum patterning faint. Lacks vertebral stripe and faint

dorsolateral stripes from back of head to one-third down torso.

Lacks visible dark transverse bands between head and pelvis.

Head relatively unpatterned: labials same colour as rest of head;

and faint, narrow pale line from posterior of eye to above ear.

Prominent black smudge on posterior edge of tympanum,

extending onto scales behind ear. Flanks have large dark patches

in axillary region extending up onto shoulder, then extending

posteriorly and fading to dark speckled appearance on a pale

background. Lateral stripe between axilla and groin absent.

Large lateral dark patch in axilla, extending up onto shoulder.

Arms lack dark banding. Very faint banding on hind limbs and

tail, with light bands much wider than darker bands. Ventral

surface plain cream to white.

Variation. 48—76 mm SVL; 101—183 mm tail length. Tail long,

ranging from 1.9-2.5 x SVL. In some individuals keeling on

dorsal scales converge towards the midline approximately

halfway down torso but run parallel to midline from mid-

dorsum to pelvis. Usually a single spinose scale above

tympanum, with 1—3 spines on post-auricular fold, occasionally

with a short row of spines extending from post-auricular fold to

above tympanum. Dorsolateral scale rows raised on neck. [n

some individuals, a slightly raised nuchal crest (males) and the

scales of dorsolateral rows on body appear to be enlarged with

slightly thickened keels, but this 1s barely perceptible (hence,

we have not included it as a diagnostic character). Pre-cloacal

pores were usually 2, but many had 4; no specimens with

femoral pores.

Dorsal colouration variable from strongly patterned

(mainly smaller 1ndividuals and females) to unpatterned (adult

males). Most individuals have pale dorsolateral stripes running

from the back of the head to at least a third of the way down the

torso. [n strongly patterned individuals, these stripes continue

onto the tail where they converge approximately halfway down.

More patterned individuals have 5—7 dark cross-bands between

the pale dorsolateral stripes, continuing onto the tail. These

cross-bands are intersected by a narrow pale white, cream or

erey vertebral stripe. Also, these individuals will often have a

black patch on shoulder, into the axilla, with pale flecks.

Individuals with little patterning are usually adult males with

breeding colours, which includes a large black patch in axillary

region extending up onto shoulder, then posteriorly fading to

dark speckled appearance with a bright yellow background,

Taxonomic revision of dragon lizards

and a pink flush on tail and rear legs. No distinct patterning on

head, upper labials flecked with light brown and cream, with no

pale labial stripe. Ventral surfaces plain cream to white. Gular

region plain or with diffuse brown speckling (no lines).

Distribution and ecology. Widespread in the Kimberley and

extreme western Top End of the NT. They occur from the

Yampi Peninsula in the south-western Kimberley, extending

eastwards to the Kununurra area and to the western NT in the

Jasper Gorge area.

This species 1s almost always associated with rocks. It will

also climb onto vegetation, such as cane grass, small shrubs

and trees, spinifex and even pandanus and mangroves, but

with rocks or creek lines nearby.

Etymology. Named from the Latin for confused or cryptic, in

reference to Allan Greer’s (former curator at the AMS) thoughts

on this species when carrying out earlier work on the group in

the 1990s. This species remained hidden until a genetic analysis

and consultation of the D. bennettii type with its small body

size, short tail and diffuse pale patterning.

Comparison with other species. Similar to D. bennettii,

occurring in similar rocky habitats, but differs in having a long

tail and limbs and dorsolateral stripes. Can be distinguished

from D. albilabris and D. sobria in mostly lacking femoral

pores, having no white or pale stripes on the head (on upper

labials or between eye and ear), no stripes under chin and

homogenous dorsal scales. Differs from D. magna, D.

margaretae and D. pindan in having a gular fold, a black spot

on tympanum and two canine teeth on each side of upper jaw.

Diporiphora lalliae differs from D. perplexa sp. nov. in lacking

both a black spot on tympanum and double canine teeth on

each side of upper jaw.

Remarks. This species has previously been confused with D.

bennettii (see account above), and this is the species usually

depicted in field guides as D. bennettii. However, both genetic

work and examination of the original specimens have now

distinguished these two species. Interestingly, D. perplexa sp.

nov. is the only member of the D. bennettii species group that

Is sympatric with the three other species (which are generally

allopatric). This distributional pattern suggests that D. perplexa

sp. nov. may have different habitat preferences and ecology to

the other species that allows for sympatry.

Recent unpublished phylogenomic research using single-

nucleotide polymorphisms shows that D. perplexa sp. nov. 1s

highly divergent and genetically distinguishable from the D.

sobria (as defined below), even in areas of syntopy (J. Fenker,

unpublished data).

Diporiphora sobria Storr, 1974

Common name. Northern savannah two-pored dragon.

Figure 13, Tables 3, 4

Diporiphora albilabris sobria Storr, G.M. 1974. Agamid lizards

of the genera Caimanops, Physignathus and Diporiphora in Western

Australia and Northern Territory. Records of the Western Australian

Museum 3: 121—146 [135]. Type data: holotype - WAM R23180, Pine

Creek, NT [14° 04' S, 131° 58' E].

Diporiphora bennettii arnhemica Storr, G.M. 1974. Agamid

lizards of the genera Caimanops, Physignathus and Diporiphora in

Western Australia and Northern Territory. Records of the Western

Australian Museum 3: 121—146 [137]. Type data — holotype ANWC

R740, near upper Katherine River, NT [14° 13' S, 132° 36' E].

Synonymy decision of current work.

Diagnosis. Body size moderately large (to 69 mm SVL) with

moderately long tail (2.3—2.6 x SVL). Gular fold present, post-

auricular fold strong and scapular fold present but often weak.

Dorsolateral stripes present. Lacks dark smudge on posterior

edge of tympanum. In WA, the scales between dorsolateral

stripes are homogeneous (fig. 10b); in the NT, heterogeneous

(fig. 10c). Pre-cloacal pores 4; femoral pores 2.

Description of holotype. Subadult, unknown sex; 34 mm SVL;

68 mm tail length. Moderately robust with long limbs and tail.

Two canines on either side of upper jaw. Gular fold present,

scapular and post-auricular folds present but weak. Dorsal scales

homogeneous between faint pale dorsolateral stripes. These

homogeneous scales are relatively large and strongly keeled. On

the outer sides of the faint dorsolateral stripes are smaller keeled

scales. Scales on flanks homogeneous, with keels running

parallel to dorsum. Lacks spinose scales on head, limbs or tail.

Lacks granular scales in axilla. Ventral scales homogenous and

strongly keeled. Pre-cloacal pores 4; femoral pores 2.

(b)

5 mm

Figure 13. Diporiphora sobria: a, adult male with breeding colouration,

Halls Creek, Western Australia (photo: S. Wilson); b, holotype WAM

R23180 from Pine Creek, Northern Terrotory.

Dorsal colour uniform brown and grey tones, without

apparent patterning. Lacks vertebral stripe but has faint

dorsolateral stripes on anterior half of body, fading to

background colour midway down torso. Lacks visible dark

transverse bands between head and pelvis. Head relatively

unpatterned: labials pale; and very faint, narrow pale line from

posterior of eye to above ear. Dark smudge on posterior of

tympanum absent. Flanks unpatterned and consistent with

dorsal colouration. Lateral stripe between axilla and groin

absent. Lacks lateral dark patch. Arms lack dark banding. Very

faint banding on hind limbs; dark bands on anterior two-thirds

of tail, with dark bands being narrower than the light, fading to

unbanded for remainder of tail. Ventral surface of head, torso

and tail unpatterned and white or cream colouration.

Variation. 46—69 mm SVL; 114—162 mm tail length. Tail long,

ranging from 2.3-2.6 x SVL. Gular fold always present and

often strong, post-auricular fold from weak to strong and scapular

fold present but variable. No low nuchal crest. Most individuals

have homogeneous dorsal scales between pale dorsolateral

stripes, with all individuals examined from WA having

homogeneous scales. However, animals examined from the

eastern NT, on the Barkly Tablelands and the Gulf of Carpentaria

region, and from north-central regions, such as the Mt Wells and

Pine Creek area, had somewhat heterogeneous scales between

pale dorsolateral stripes. In these animals, paravertebral rows on

either side of the vertebral scale row are not reduced in size In

comparison to the vertebral scale row, but the 2nd paravertebral

scale row is slightly enlarged, with a strong central keel that is

aligned to the scales immediately anterior and posterior. These

strongly keeled scales that are aligned form a ridge running

along the dorsum either side of the midline. Animals with these

dorsal scales include NMV D7/2666—68, D72706. D72707,

D72715, D72720, D73995, D74001, D74003, D74022, D74024

and D74262. Scales on the sides are similar to typically sized

dorsal scales and are angled up towards midline. Ventral scales

homogenous and strongly keeled. Pre-cloacal pores 4

(occasionally up to 5); femoral pores 2 (occasionally 3 or 4).

Dorsal patterning variable from plain to strongly patterned.

Strongly patterned individuals, typically adult females and

juveniles, have approximately 7—9 irregular dark transverse

bands from nape to legs across dorsum on a pale background.

Dark bands are of similar width or narrower than pale

background. These bands are separated by an indistinct faint

erey or light brown vertebral stripe. The dark transverse bands

continue laterally beyond the pale cream dorsolateral stripes,

becoming diffuse and terminating on sides, with defined lateral

stripe absent. Labial scales pale cream, with a few darker flecks,

extending as a broad pale band along jaw to back of head ending

at the post-auricular fold. In these animals, the lateral surfaces

of the neck, axilla and flanks lack a defined dark patch. Some

individuals are less strongly patterned, particularly in the NT,

and lack the pale dorsolateral stripes or the dorsolateral stripes

fade midway down dorsum. Adult males in breeding colouration

often have little patterning, having strongly contrasting charcoal

black, white and chestnut or orange-red colouring on head,

flanks and upper body, with tail also having a pink flush in some

individuals. Arms and legs banded. Ventral surface of head,

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

torso and tail usually unpatterned and white or cream

colouration. However, adult males may have up to three pairs of

erey stripes in gular region, terminating at gular fold.

Distribution and ecology. Widely distributed from the southern

Kimberley region (ie. south of the barrier ranges: King

Leopold and Durack), Top End of the NT (but with no records

from Arnhem Land) and just extending into western Queensland

(fig. 8). Extends south to the Katherine area.

A habitat generalist occurring in tropical savannah

woodlands and grasslands, within these habitats it appears to be

associated with rockier areas. Similar habitats to D. albilabris.

Seen to perch on low vegetation, rocks or termite mounds.

Comparison with other species. This species has a very wide

distribution and overlaps with numerous other Diporiphora

species. In WA, it can be distinguished from D. albilabris in

having a scapular fold, fewer spines around tympanum and

post-auricular fold and having homogenous scales between the

dorsolateral stripes. It differs from D. perplexa sp. nov. In

having pale rows of dorsolateral scales, usually a strong post-

auricular fold, two femoral pores and lacks dark markings on

the posterior of the tympanum. Differs from D. bilineata, D.

magna and D. lalliae in having femoral pores, two pairs of

canines in upper jaw and white labial scales. It also differs from

D. bilineata, D. magna and D. margaretae in having a gular

fold and two pairs of canine teeth on each side of upper Jaw.

Remarks. This species incorporates what was D. arnhemica

and much of the distribution of D. albilabris albilabris sensu

lato. Despite the original name, this species does not appear to

occur in Arnhem Land from our observations, although

collections from this region are scarce. This taxon is diverse

and exhibits morphological variation In dorsal scalation. Where

it occurs near the morphologically similar D. albilabris in the

Kimberley region, the homogeneous dorsal scalation differs

and separates the two species.

Content:

. bilineata Gray, 1842

. lalliae Storr, 1974

magna Storr, 1974

. margaretae Storr, 1974

. gracilis sp. nov.

. granulifera sp. nov.

. carpentariensis sp. nov.

TO OS Me ior

Diagnostic characters for group (Table 2):

* one canine tooth on each side of upper jaw

e granular scales in axilla present, with the exception of D.

lalliae

e lateral dark spot in axilla

* femoral pores absent.

Remarks

The D. bilineata species group Is widely distributed across

northern Australia (fig. 14), from the Kimberley to the Cape York

Taxonomic revision of dragon lizards

Peninsula. Almost all species appear to be generalists, with the

possible exception of D. gracilis sp. nov. being specialised for

erasses. Body size ranges from small (D. margaretae, to 55 mm

SVL) to large (D. magna, to 77 mm SVL). This is a genetically

diverse species group (Smith et al., 2011) for which there has

been significant difficulty in species delimitation based on

morphology. We provide species accounts for D. bilineata, D.

lalliae and D. magna, with key morphological characters for

species identification and a revision of their distributions. We

also raise D. margaretae from synonymy of D. bilineata (Cogger

et al., 1983), based on genetic results and examination of types.

We also describe three new species: two from Queensland and

one from the southern Kimberley.

Diporiphora bilineata Gray, 1842

Common name. Two-lined dragon.

Figure 15, Tables 3, 4

Diporiphora bilineata Gray, J.E. 1842. Description of some

hitherto unrecorded species of Australian reptiles and batrachians. Pp.

51—57 in: Gray, J.E. (ed.). The zoological miscellany. Treuttel, Würz &

Co: London. [54]. Type data: syntype(s) - BMNH 1946.8.12.75—76,

Port Essington, NT.

Diagnosis. Body size moderate (to 58 mm SVL) with long tail

(2.2—2.7 x SVL). Gular and post-auricular folds absent, scapular

fold present but weak. Granular scales In axilla, extending over

arm to neck. Dorsal scales moderately heterogeneous: scales on

2nd paravertebral row and the two rows of pale dorsolateral

scales slightly enlarged and raised. Flanks dark in colour with

dark colour of granular scales extending posteriorly onto flanks,

which have a speckled appearance due to scattered white scales

on the dark background. Pre-cloacal pores 2; femoral pores 0.

Description of syntypes. Males; 57 and 58 mm SVL, 151 and 56

(broken) mm tail length. Medium-sized Diporiphora,

moderately gracile with long limbs and long tail. One canine on

either side of upper jaw. Gular and post-auricular folds absent,

scapular fold present. Dorsal scales heterogeneous; vertebral

row of scales, plus the 3—4 rows immediately adjacent on either

side, are enlarged and strongly keeled. The vertebral row and

the fourth longitudinal scale row from the vertebral are raised.

Beyond these enlarged vertebral scales are four rows of small

homogenous scales and then a dorsolateral longitudinal row of

@ D. bilineata

D. magna

D. margaretae

O D. gracilis sp. nov.

€ D. lalliae

@ D. granulifera sp. nov.

D. carpentariensis sp. nov.

Figure 14. Distributions of D. magna, D. bilineata, D. lalliae, D.

margaretae, D. gracilis sp. nov. D. granulifera sp. nov. and D.

carpentariensis sp. nov. based on specimens examined and collection

records.

enlarged scales, with scales on each side strongly keeled. Raised

vertebral and dorsolateral scale rows extend up onto neck onto

the back of head and posteriorly onto tail. Scales on flanks

homogeneous, although changing from small granular scales in

0 M

TOCE SSA >

. LE u seh 1

vd ista.

k. , 4

ee H»

LAS

. o

* Ld

°. @

° ‘> "

> = m

Figure 15. Diporiphora bilineata: a, adult In non-breeding colours,

Casuarina, Northern Territory (photo: S. Wilson); b, syntypes BMNH

1946.8.12.75—76, Port Essington, Northern Territory.

axilla to small but non-granular scales on the posterior two-

thirds of flanks. A few small pale spinose scales at back of head

on each side, lacks spinose scales on limbs or tail. Granular

scales in axilla, extending over arm onto neck. Ventral scales

strongly keeled. Pre-cloacal pores 2; femoral pores 0.

Dorsum strongly patterned. Wide pale, poorly defined

vertebral stripe associated with enlarged vertebral scale rows

and narrow pale dorsolateral stripes associated with row of

enlarged scales, extending from back of head onto base of tail.

Six dark transverse bands between head and pelvis, narrower

than the pale background. Head relatively plain with little

patterning, labials similar colour to rest of head and lacking

pale line between eye and ear. Dark smudge on posterior of

tympanum absent. Flanks dark in colour with dark colour of

eranular scales extending posteriorly onto flanks, which have

a speckled appearance due scattered white scales on the dark

background. Lateral stripe between axilla and groin absent.

Dark patch in axilla extending up onto shoulder. Arms and

legs lack dark banding. Faint banding on hind limbs; dark

bands down length of tail, with dark bands being wider than

the light bands. Ventral surface cream with no patterning.

Variation. 46—58 mm SVL; 113-156 mm tail length. Tail long,

ranging from 2.2—2.7 x SVL. Gular fold always absent and

scapular fold present in all animals examined. In a few

Specimens a small post-auricular fold present. Outer raised

trailing edge of scales on outer row of dorsolateral stripes

usually present but often trailing edge of scales are only weakly

raised, providing demarcation between dorsal and lateral

surface. Pre-cloacal pores 2; femoral pores 0.

Variable patterning from strongly patterned to plain

individuals. In patterned individuals, 5—7 dark brown transverse

bands ranging from narrower to wider than pale background.

Dark bands are offset to each other on either side of a narrow

ereyish or cream vertebral stripe and intersected two pale

dorsolateral stripes. In plain individuals, dark dorsal transverse

bands are faint or absent, vertebral stripe may be absent but

there are usually still the pale dorsolateral stripes from neck to

at least level of mid-dorsum. However, dorsolateral stripes are

also absent in a few animals. Granular scales on flanks around

arm are usually dark brown, extending posteriorly onto flanks

that have speckled appearance due to scattered white scales on

a dark background. These scattered pale scales sometimes

form vertical lines. No white markings on face, labial scales

speckled with light brown flecks. Usually faint or no banding

on legs but in some more patterned animals banding present.

Ventral surface cream, usually plain but some individuals have

faint longitudinal stripes on ventral surface of head. Males with

breeding colouration tend to lose some of their dorsal

patterning, having a yellow wash over the head and upper body

with a large black patch in axilla that extends onto shoulder.

Distribution and ecology. Diporiphora bilineata is widely

distributed across the Top End of NT. It occurs sympatrically

with D. magna in the Pine Creek area. It is a common generalist

that occurs throughout the tropical savannah woodlands and

grasslands of this region, and is often seen perching on low

vegetation, rocks or termite mounds.

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

Comparison with other species. The distribution of D. bilineata

overlaps with two other Diporiphora species. Diporiphora

bilineata can be distinguished from D. magna in usually

having fewer pre-cloacal pores (2 vs. 4), lacking a post-auricular

fold and by possessing heterogeneous dorsal scales, rows of

enlarged vertebral scales and dark flanks with scattered white

scales. Diporiphora bilineata can be distinguished from D.

sobria in lacking a gular fold and femoral pores, and having

single canines on either side of the upper jaw.

Remarks. This taxonomic revision has reduced the distribution

of this species to the Top End of the NT, with overlap with D.

magna in the Pine Creek area. Previously, adult males of D.

bilineata in breeding colouration may have been mistaken for

D. magna, and D. jugularis has been resurrected that was often

attributed to D. bilineata, further reducing this species’ range.

Genetic work has shown that these two species do not overlap

extensively (Smith et al., 2011).

Diporiphora lalliae Storr, 1974

Common name. Northern deserts dragon.

Figure 16, Tables 3, 4

Diporiphora lalliae Storr, G.M. 1974. Agamid lizards of the

genera Caimanops, Physignathus and Diporiphora in Western

Australia and Northern Territory. Records of the Western Australian

Museum 3: 121-146 [138]. Type data: holotype — WAM R23020,

Langey Crossing, WA [17° 39'S, 123° 34' E].

Diagnosis. Body size moderately large (to 62 mm SVL) with

very long tail (2.6—3.4 x SVL). Gular, post-auricular and

scapular folds present. Small scales in axilla but usually not

eranular. Homogeneous dorsal scales between pale dorsolateral

lines that usually lack raised scales In outer row, providing

little demarcation between dorsal and lateral scales. Pre-

cloacal pores 4; femoral pores 0.

Description of holotype. Male; 62 mm SVL; 152 mm tail length.

Medium-large Diporiphora with long tail and long limbs. One

canine on either side of upper jaw. Gular fold present. Strong

scapular and post-auricular folds. Dorsal scales homogeneous,

relatively large and strongly keeled. At the shoulder, dorsal

scales are raised and strongly keeled (but not enlarged) In a

longitudinal series of paravertebral and dorsolateral scales that

fade by midbody. Scales on flanks homogeneous. Single white

spinose scale at back of head sitting on ventral end of post-

auricular fold, lacks spinose scales limbs or tail. Small scales

in axilla but not granular. Ventral scales strongly keeled. Pre-

cloacal pores 4; femoral pores 0.

Dorsum strongly patterned. Wide grey vertebral stripe (~4

scales wide at middle of dorsum), extending from back of head

onto base of tail. Prominent pale dorsolateral stripes from

shoulder to mid-dorsum, fading into background patterning;

these scales form the enlarged longitudinal row of dorsolateral

scales at the shoulder. Six dark transverse bands between head

and pelvis, approximately the same width as the pale

background. Head relatively plain with little patterning, labials

similar colour to rest of head but have faint pale line between

eye and ear. Dark smudge on posterior of tympanum absent.

Taxonomic revision of dragon lizards

Flanks pale in colour, similar to dorsum. Lateral stripe

between axilla and groin absent. Dark patch in axilla absent.

Arms with faint dark banding. Strong well-defined alternating

light and dark bands on legs; dark bands on tail, continuing to

near end of tail, with dark bands being a similar width to the

light bands. Ventral surface white and unpatterned.

Variation. 49-62 mm SVL; 130-194 mm tail length. Very long

tail, ranging from 2.7—3.4 x SVL. Gular fold always present,

although weak In some individuals. Scapular fold strong and post-

auricular fold usually strong, although weak in some individuals.

No enlarged spinose scales above tympanum or along post-

auricular fold, although the latter may have several slightly

enlarged scales. Very low nuchal crest. Outer raised trailing edge

of scales on outer row of dorsolateral stripes usually absent, but in

some individuals, the trailing edge of scales are weakly raised,

particularly over shoulder, providing weak demarcation between

dorsal and lateral surface. Pre-cloacal pores 4; femoral pores 0.

Dorsal patterning variable from strongly patterned to little

patterning. In strongly patterned individuals, there are 6—8

wide, dark brown transverse bars between shoulders and pelvis,

intersected by a wide grey vertebral stripe; pale dorsolateral

10 mm

Figure 16. Diporiphora lalliae: a, adult male, Three Ways, Northern

Territory (photo: S. Wilson); b, holotype - WAM R23020, Langley

Crossing, Western Australia.

stripes from neck and usually extending onto tail. At the

shoulder, these dorsolateral stripes consist of an enlarged

longitudinal row of scales. Usually no patterning on the head,

but in some individuals a white stripe between the eye and ear.

In more plain individuals, dark transverse bars faint or absent,

and vertebral stripe often absent. Pale dorsolateral stripes from

neck to tail usually present, even in unpatterned animals. Dark

bands on tail, continuing to near tip. Usually well-defined

alternating light and dark bands on upper and lower legs. Dark

spot in axillary region absent and flanks usually pale, but some

individuals have dark lateral spot above and slightly posterior to

shoulder. Ventrum plain.

Distribution and ecology. The distribution of D. lalliae spans

the northern extent of the arid zone along the western deserts,

from the southern Kimberley In WA to the far west of

Queensland. It extends Into the southern reaches of the AMT

but does not occur in the central arid zone as occasionally

depicted (e.g. Cogger, 2014).

This species occurs In a variety of habitats from savannah

woodlands and grasslands to arid habitats. It is a generalist

species that is found in many habitats, often seen perching on

small rocks, termite mounds or clumps of earth.

Comparison to other species. Diporiphora lalliae 1s sympatric

with D. magna, D. gracilis sp. nov. and D. granulifera sp. nov.

in the northern parts of its range, occurring in similar habitats

and is superficially similar in appearance. However, D. lalliae

can be distinguished from these species by the presence of a

cular fold, which is unique in the D. bilineata species group

(Table 2). The distribution of D. lalliae also overlaps with D.

sobria, from which it can be distinguished in having single

canine teeth on each side of upper jaw and lacking femoral

pores. In the southern Kimberley region, D. lalliae can be

distinguished from D. pindan in having a gular fold and strong

post-auricular and scapular folds.

Remarks. This species has previously been confused with

numerous other species owing to its generalised appearance.

Phylogenetic work has confirmed that it is a member of the D.

bilineata species group (Smith et al., 2011) but is unique in this

group due to the presence of a gular fold.

Diporiphora magna Storr, 1974

Common name. Yellow-sided two-lined dragon.

Figure 17, Tables 3, 4

Diporiphora magna Storr, G.M. 1974. Agamid lizards of the

genera Caimanops, Physignathus and Diporiphora in Western

Australia and Northern Territory. Records of the Western Australian

Museum 3: 121—146 [137]. Type data: holotype - WAM R42786, Old

Lissadell (now submerged by Lake Argyle), WA [16° 30'S, 128° 41' E].

Diagnosis. Body size moderately large (to 77 mm SVL) with

very long tail (to 3 x SVL). Gular fold absent, post-auricular

and scapular folds strong. Granular scales in axilla, extending

over arm but not extending onto sides of neck. Homogeneous

dorsal scales between pale dorsolateral lines that usually lack

raised scales in outer row, providing little demarcation between

dorsal and lateral scales. Pre-cloacal pores 4; femoral pores 0.

Description of holotype. Female; 47 mm SVL; 141 mm tail

length. Medium Diporiphora, moderately gracile with long

limbs and very long tail. One canine on either side of upper Jaw.

Gular fold absent. Post-auricular and scapular folds strong.

Dorsal scales strongly keeled, relatively small and homogeneous.

Scales on flanks homogeneous, although changing from small

eranular scales in axilla to small but non-granular scales on the

posterior two-thirds of flanks. Cluster of spinose scales at back

of head on each side, lacks spinose scales on limbs and tail.

Granular scales in axilla, extending over arm but not extending

onto sides of neck. Ventral scales strongly keeled. Pre-cloacal

pores 3 (2 on right; | on left); femoral pores 0.

Dorsum strongly patterned. Wide grey vertebral stripe (~2

scales wide at mid-dorsum), extending from back of head onto

base of tail, and prominent pale dorsolateral stripes from

shoulder to base of tail, fading into background patterning.

seven dark transverse bands between head and pelvis,

narrower than the pale background and offset across the

vertebral stripe. Head relatively plain with little patterning,

10 mm

Figure 17. Diporiphora magna: a, adult male, Larrimah, Northern

Territory (photo: S. Wilson); b, holotype WAM R42786, Old Lissadell

(now submerged by Lake Argyle), Western Australia.

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

labials similar colour to rest of head and lacking pale line

between eye and ear. Dark smudge on posterior of tympanum

absent. On flanks, granular scales in axilla are dark brown and

strongly defined with an abrupt transition to pale flanks with

little patterning. Lateral stripe between axilla and groin

absent. Dark patch in axilla extending up onto shoulder. Faint

banding on hind limbs; dark bands on anterior two-thirds of

tail that are wider than the light bands, fading to unbanded

light colouration for remainder of tail. Arms and legs with

faint dark banding. Ventral surface cream with a few very

faint longitudinal stripes on ventral surface of head.

Variation. 54—77 mm SVL; 145—209 mm tail length. Very long

tail, ranging from 2.6—3.0 x SVL. Gular fold always absent,

post-auricular and scapular strong. In a few individuals, the

post-auricular fold, although strong, may be short («2 mm).

Low nuchal crest, more prominent 1n males. One spinose scale

above tympanum and one on post-auricular fold with from 0—3

additional spines on post-auricular fold. Scales rows of

dorsolateral stripes on neck and above arms with raised

posterior edges. All individuals examined have pre-cloacal

pores 4 and femoral pores 0, except for one individual: large

male (NM V D73812) from the southern Kimberley has a small

indistinct femoral pore on each side.

Variable patterning from strongly patterned individuals to

plain individuals. In patterned individuals, 5—7 dark brown

transverse bands ranging from narrower to wider than pale

interspaces of background colour. Dark bands are often offset

to each other on either side of a broad greyish or cream

vertebral stripe 3—4 scales wide and two pale dorsolateral

stripes usually present. In more plain individuals, dark dorsal

transverse bands are faint or absent, vertebral stripe may be

absent but with pale dorsolateral stripes from neck to at least

level of mid-dorsum. However, dorsolateral stripes are also

absent In a few individuals, particularly breeding males.

Granular scales In or near the axilla are dark brown, flanks

cream, grey, light brown with little patterning. No white

markings on face, labial scales speckled with light brown

flecks. Faint or no banding on legs. Ventral surface cream,

usually plain but some individuals have faint longitudinal

stripes In gular region. Males with breeding colouration tend

to lose some of their dorsal patterning, having a yellow wash

over the head and upper body with a large back patch in axilla

that extends onto shoulder.

Distribution and ecology. Diporiphora magna is widely

distributed across the tropical savannah region of northern

Australia, from the central Kimberley, WA, through the NT and

Just over the Queensland border at Lawn Hill National Park. It

appears to have been replaced by other D. bilineata species

group members as follows: to the south by D. lalliae, which

occupies the northern deserts, to the west In the Kimberley by

D. margaretae and D. gracilis sp. nov., and 1n the Top End of the

NT by D. bilineata, although there Is a wide (~100 km) area of

sympatry between roughly Pine Creek and Katherine.

A habitat generalist occurring in tropical savannah

woodlands and grasslands, individuals have been observed to

perch on low vegetation, rocks or termite mounds.

Taxonomic revision of dragon lizards

Comparison to other species. The distribution of D. magna

overlaps with numerous other Diporiphora species across the

tropical savannahs of northern Australia. In the Kimberley

region, D. magna differs from D. margaretae in having a

smaller body size (to 77 vs. 55 mm SVL), stronger post-

auricular and scapular folds, a longer tail and flanks lacking

speckled appearance (scattered white scales on a dark

background). In the northern NT, D. magna differs from D.

bilineata in having a post-auricular fold and homogenoous

dorsal scales, lacking rows of enlarged vertebral scales and

lacking dark flanks with scattered white scales. In the southern

part of its range, D. magna can be distinguished from D. lalliae

in lacking a gular fold, a longer tail 1n proportion to body and

eranular scales in axilla. Across its range, D. magna can be

distinguished from D. albilabris, D. bennettii, D. sobria and D.

perplexa sp. nov. in lacking a gular fold, lacking femoral pores

and having single canines on either side of the upper jaw.

Remarks. Phylogenetic work has redefined the distribution of

this species (Smith et al., 2011). Specimens of D. margaretae

were formerly assigned to D. magna, meaning now that D.

magna does not occur in the north-west Kimberley region. This

taxon Is diverse and exhibits morphological variation In colour

patterning and overlaps or contacts a number of phylogenetically

close species. Where it occurs in close proximity to these

species, it can be distinguished on the basis of scalation.

Diporiphora margaretae Storr, 1974

Common name. Northwest Kimberley two-lined Dragon.

Figure 18, Tables 3, 4

Diporiphora bilineata margaretae Storr, G.M. 1974. Agamid

lizards of the genera Caimanops, Physignathus and Diporiphora in

Western Australia and Northern Territory. Records of the Western

Australian Museum 3: 121-146 [143]. Type data: holotype - WAM

R27648, Kalumburu, WA [14° 18'S, 126° 30' E].

Diagnosis. Body size small to moderate (to 55 mm SVL) with

long tail (2.5—2.7 x SVL). Gular fold absent, post-auricular fold

weak to moderate, scapular fold moderate to strong. Granular

scales in axilla, extending over arm onto neck to posterior edge

of the scapular fold. Scales on outer rows of dorsolateral stripes

have raised trailing edge in some individuals, particularly over

shoulder, giving moderate demarcation between dorsal and

lateral surfaces. Enlarged pale scales on sides tending to form

vertical bars. Pre-cloacal pores 4; femoral pores 0.

Description of holotype. Female; 55 mm SVL; 128 mm tail

length. A medium-sized Diporiphora, moderately gracile with

long limbs and tail. One canine on either side of upper Jaw.

Gular, post-auricular and scapular folds absent. Dorsal scales

strongly keeled, relatively large and homogenous, although band

of smaller homogenous scales across back of head and neck.

Longitudinal series of raised, but not enlarged, pale paravertebral

and dorsolateral scales at the shoulder, not extending down

dorsum. Scales on flanks homogeneous, although changing from

small granular scales in axilla to small, non-granular scales on

the posterior two-thirds of flanks. A few small pale spinose

scales at back of head on each side, lacks spinose scales limbs on

tail. Granular scales in axilla, extending over arm onto neck to

posterior edge of the scapular fold. Ventral scales strongly

keeled. Pre-cloacal pores 4; femoral pores 0.

Dorsum strongly patterned. Six irregular dark brown

squares (rather than transverse bands) between head and pelvis

either side of a broad grey vertebral stripe, intersected by

poorly defined, light brown dorsolateral stripes. Dark brown

squares are narrower than the pale background. Head relatively

plain with little patterning, labials similar colour to rest of

head and lacking pale line between eye and ear. Dark smudge

on posterior of tympanum absent. Complex patterning on

flanks; granular scales on flanks around arm are dark brown,

extending posteriorly onto flanks that have speckled

appearance due to seemingly random assortment of scale

colours from white, cream, grey, light brown and dark brown;

dark dorsal squares also extend as dark patches onto flanks.

Lateral stripe between axilla and groin absent. Darker scales

in axilla not extending up onto shoulder. Arms with faint dark

banding. Banding on legs not well defined, with light bands

narrower than dark bands. Dark bands on anterior third of tail,

fading to unbanded light colouration for remainder of tail.

Ventral surface cream with a few scattered fleck of light brown

on ventrum, throat and head.

(a)

(b)

Figure 18. Diporiphora margaretae: a, adult male (NMV D73834),

King Edward River, Kimberley, Western Australia (photo: J. Melville);

b, holotype WAM R27648, Kalumburu, Western Australia.

Variation. 44—55 mm SVL; 110—135 mm tail length. Tail long,

ranging from 2.5-2.7 x SVL. Gular fold always absent, post-

auricular and scapular vary from weak to moderate. Low

nuchal crest present. Spinose scale above tympanum, with 1—4

low spines on post-auricular fold. On dorsolateral row on neck,

scales with posterior edges raised. Dorsal scales relatively

homogeneous; an exception is the 2nd paravertebral row

tending to be slightly enlarged. On flanks, keels of scales

angled dorsally and posteriorly; flanks with scattered enlarged

scales, tending to align vertically. Pre-cloacal pores 4; femoral

pores 0.

Dorsal colour pattern variable. In patterned individuals,

cream, grey, light brown to dark brown complex pattern with

white dorsolateral stripes. Five or six irregular dark brown

bands either side of a broad greyish vertebral stripe 3—5 scales

wide. Granular scales on flanks around arm are usually dark

brown, extending posteriorly onto flanks. Sides with speckled

appearance due to scattered pale scales on enlarged scales on

a dark background, often forming vertical rows. The dark

bands on the dorsal surface do not extend laterally on the body

and in most individuals the bands do not extend beyond the

white dorsolateral stripes. In small heavily marked individuals,

there is often a dark-edged pale lateral stripe. In plainer

individuals, often adult males, dark transverse bands across

the dorsum are either absent or faint. Lateral surfaces not as

dark and speckled as for patterned individuals. No white

markings on face, labial scales speckled with dark brown

flecks. Banding on limbs not well defined, with light bands

narrower than dark bands. Dark bands on anterior third of tail,

fading to unbanded light colouration for remainder of tail.

Ventral surface cream sometimes with a few scattered flecks

of light brown on gular region and ventrum; occasionally

several pairs of lines 1n gular region.

Distribution and ecology. Restricted to the far north Kimberley

region, with records from the Anjo Peninsula and Sir Graham

Moore Island in the extreme north, Kalumburu, Mitchell

Plateau, Prince Regent River and Drysdale River National

Parks, and on Mary Island.

This species has been recorded from Eucalyptus woodland,

cane grass, triodia on sandstone and other rocky areas.

Comparison to other species. The distribution of D. margaretae

overlaps with a number of other Diporiphora species, including

D. albilabris, D. bennettii, D. perplexa sp. nov., D. magna and

D. pallida sp. nov. Diporiphora maragetae differs from D.

magna in having weak or absent post-auricular and scapular

folds (as opposed to consistently strong folds), and having

flanks that have a speckled or barred appearance due to

scattered pale scales on a dark background. Diporiphora

margaretae can be distinguished from D. albilabris, D.

bennettii and D. perplexa sp. nov. in lacking a gular fold and

femoral pores, and having single canines on either side of the

upper Jaw. Diporiphora margaretae differs from D. pallida sp.

nov. in possessing a more gracile habitus with longer limbs and

tail, lacking a gular fold and having granular scales in axilla.

Remarks. This species was originally described as a subspecies

on D. bilineata (Storr, 1974) but was subsumed into D. magna

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

without comment in Storr et al. (1983), despite having the

largest difference in body size of any species pair within the D.

bilineata species group. Genetic work, across multiple genes

(Smith et al. 2011), demonstrated the distinctiveness of this

species relative to D. bilineata and D. magna. Phylogenetic

analyses support it being the sister species to D. gracilis sp.

nov. (fig. Ic).

Diporiphora gracilis sp. nov.

ZooBank LSID: http://zoobank.org/urn:lsid:zoobank.oreg:act:

F3BBI6E0-259E-4396-A D68-FFE06AO0FE624

Common name. Gracile two-lined dragon.

Figure 19, Tables 3, 4

Holotype. WAM R177291 (formerly NMV D75540) (adult male),

Fairfield-Leopold Downs Road, south of Gibb River Road, WA

(17° 29' 37.0" S, 125° 2' 17.7" E). Collected by P. Oliver on 2 November

2013.

Paratypes. NM V D75541 (adult females), as for holotype; NM V

D73901 (adult male), as for holotype; WAM R163503 (female) and

WAM R163504 (male), Mornington Station, WA (17° 30' 23" S,

126° 02' 07" E); WAM R177952 (formerly NMV D75542) (adult

female), as for holotype.

Diagnosis. Body size moderately long (to 61 mm SVL) with

elongate and gracile body shape and very long tail (to 3 x SVL).

Gular fold absent, post-auricular and scapular folds weak.

Granular scales in axilla, extending over arm to scapular fold.

Homogeneous dorsal scales between pale dorsolateral lines

that usually lack raised scales 1n outer row, providing little

demarcation between dorsal and lateral scales. Pre-cloacal

pores 4; femoral pores 0.

Description of holotype. Male; 57 mm SVL; tail length 165

mm. A medium-sized gracile Diporiphora, with long limbs

and a very long tail. One canine on either side of upper jaw.

Gular fold absent; post-auricular and scapular folds weak.

Dorsal scales on torso strongly keeled, parallel to midline and

homogenous; heterogeneous scales on head with wide band of

smaller scales across back of head and neck. Low nuchal crest

of ~8 scales. Granular scales in axilla, extending over arm to

the weak scapular fold. Scales on flanks homogeneous,

although changing from small granular scales in axilla to small

but non-granular scales on the posterior two-thirds of flanks.

Lacks spinose scales on limbs or tail. Ventral scales weakly

keeled on head and throat, strongly keeled on body. Pre-cloacal

pores 4; femoral pores 0.

Dorsal colour greyish-brown without patterning.

Dorsolateral and vertebral stripes absent. Dark transverse

bands between head and pelvis absent. Head relatively plain,

labials similar colour to rest of head and lacking pale line

between eye and ear; lateral portions of snout paler than dorsal

surface of snout. Dark smudge on posterior of tympanum

absent. Granular scales on axilla dark brown, posterior two-

thirds of flanks same colour as dorsal surface. Lateral stripe

between axilla and groin absent. Dark patch in axilla, not

extending up onto shoulder. Faint banding on hind limbs; tail

plain without banding. Arms lack dark banding. Ventral

surface cream with no patterning, lower labials faintly stippled.

Taxonomic revision of dragon lizards 4/

Figure 19. Diporiphora gracilis sp. nov. Images of holotype (WAM R177291, formerly NMV D77540), Fairfield-Leopold Downs Road, south of

Gibb River Road, Western Australia: a, in life (photo: J. Sumner); b, c, d, dorsal, ventral and, lateral (head) views. Pattern variation (individuals

from Mornington Station, Western Australia: e, adult male with breeding colour; f, gravid female (plain); g, gravid female (patterned) (photos:

Melissa Bruton, Australian Wildlife Conservancy).

Variation. 52-61 mm SVL; 140—185 mm tail length. Very long

tail, ranging from 2.7—3.0 x SVL. Gular fold always absent,

post-auricular fold weak or absent, and scapular fold weak. At

most, a single small spinose scale above tympanum and one on

post-auricular fold. Dorsal scales homogeneous. Presence of

outer raised trailing edge of scales on outer row of dorsolateral

stripes variable — usually absent in more plain individuals but

present in those with strong patterning, providing weak

demarcation between dorsal and lateral surface. Gulars smooth.

Pre-cloacal pores 4; femoral pores 0.

Variable patterning from strongly patterned individuals to

plain individuals. In patterned individuals, dark brown bands

offset each other to either side of a broad greyish vertebral

stripe, pale well-defined dorsolateral stripes (approximately

two scales wide) that extend from back of head to hind legs,

where stripes continue as broken stripes down a third of the

tail. Dark dorsal cross-bands continue beyond dorsolateral

stripes to a narrow pale lateral stripe. Below lateral stripes 1s

plain with start of cream ventral colour. No patterning on

head, lacks pale stripe between eye and ear. Faint banding on

limbs and banding on anterior third of tail. In plain individuals,

there is no patterning on body or head, including no dorsolateral

or vertebral stripes. In these plain individuals, granular scales

in axilla are dark brown and in life have a well-defined black

spot on sides and a greenish-yellow hue to body. No white

markings on face, labial scales speckled with light brown

flecks. Faint or no banding on limbs. Ventral surface cream,

without markings. In life, males with breeding colouration

have a well-defined black spot on sides, a greenish-yellow hue

to body and a pink flush on the tail.

Etymology. Named for the gracile body shape, with noticeably

long and slender body, limbs and tail. Used as a noun in apposition.

Distribution and ecology. Restricted to the south-western

Kimberley region (fig. 14). Currently only known from two

locations, the type location on the Fairfield-Leopold Downs

Road and further east on Mornington Station, approximately

80 km apart.

Diporiphora gracilis sp. nov. appears to be a grassland

specialist, occupying savannah grasslands on clay soils

associated with the floodplain of the Lennard River. More

collecting 1s required to determine how far the distribution of

this species extends and whether it 1s only associated with

erasslands on clay soils.

Comparison with other species. The distribution of D. gracilis

sp. nov. overlaps numerous other Diporiphora species. The

distribution of D. pindan overlaps with D. gracilis sp. nov. but

they appear to occupy different habitats (D. gracilis sp. nov. in

erasslands on floodplains; D. pindan 1n shrubs and spinifex).

Diporiphora pindan can be distinguished from D. gracilis sp.

nov., with the latter lacking a well-defined white stripe between

eye and ear, having strongly keeled dorsal scales where keels

form longitudinal ridges running along torso and having a very

long tail In proportion to body size. Differs from D. magna in

lacking strong post-auricular and scapular folds, and having a

more gracile habitus. Differs from D. lalliae in lacking gular

fold and possessing granular scales in axilla. Differs from D.

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

albilabris, D. bennettii, D. sobria and D. perplexa sp. nov. 1n

lacking femoral pores, lacking a gular fold and having single

canines on each side of the upper jaw.

Remarks. This species is sister to D. margaretae from the

northern Kimberley, both of which form a Kimberley endemic

lineage compared to the next closely related species, D. lalliae

and D. magna, which both extend across the Kimberley and

NT to just Inside the Queensland border.

Diporiphora gracilis sp. nov. was first collected on

Mornington Station in 2004. The collection of further

individuals by J. Melville that had tissues taken establishes

this species' distinctiveness, and the morphology of these

specimens Is shared by the two Mornington specimens.

Diporiphora granulifera sp. nov.

ZooBank LSID: http://zoobank.org/urn:lsid:zoobank.oreg:act:

B6B3DAEB-9DSE-4D5A-A53B-04BF6962C3BlI

Common name. Granulated two-lined dragon.

Figure 20, Tables 5, 4

Holotype. QM J96362 (formerly NMV D74060) (adult male),

Downs Road, 2 km from Barkly Highway, Queensland (20.3714? S,

139.1529" E). Collected by K. Smith in October 2005.

Paratypes. NMV D74047 (adult female), Lawn Hill Station,

Queensland (18.6536? S, 138.5653? E); NMV D74054 (adult male),

Carpentaria Highway, west of Burketown, Queensland (18.0242* S,

139.0077? E); NMV D74062 (adult male), Downs Road, 2 km from

Barkly Highway, Queensland (20.3717? S, 139.1525" E).

Diagnosis. Medium-large species (to 68 mm SVL) with a very

long tail (22.5 x SVL). Gular fold absent, post-auricular fold

weak and scapular fold strong. Granular scales in axilla,

extending over arm and along the full length of the scapular

fold. Scales on neck anterior to scapular fold small and slightly

eranular. Outer scale row in dorsolateral stripes have raised

posterior edge, particularly over shoulder, giving strong

demarcation between dorsal and lateral surfaces. Pre-cloacal

pores 4—6 (usually 4); femoral pores 0.

Description of holotype. Male; 65 mm SVL; 176 mm tail length.

Medium-large Diporiphora, moderately gracile with long

limbs and very long tail. One canine on either side of upper

Jaw. Gular fold absent. Post-auricular and scapular folds strong.

Dorsal scales strongly keeled, relatively homogenous.

Longitudinal series of raised but unenlarged pale dorsolateral

scales at the shoulder, extending down posteriorly along

dorsum to base of tail. Scales on flanks homogeneous, although

changing from small granular scales in axilla to small but not

eranular scales on the posterior two-thirds of flanks. Lacking

cluster of spinose scales at rear of head on the post-auricular

fold, lacks spinose scales limbs or tail. Small granular scales in

axilla, extending up over shoulder and along under full extent

of the scapular fold. Scales on sides of neck anterior to scapular

fold small and slightly granular. Ventral scales strongly keeled.

Pre-cloacal pores 6; femoral pores 0.

Dorsum light brown with little patterning. Faint broad

grey vertebral stripe; well-defined cream dorsolateral stripes

running from back of head to base of tail and associated with

Taxonomic revision of dragon lizards

raised but not enlarged longitudinal scales. Dark transverse

bands absent. Head relatively plain with little patterning;

labials similar colour to rest of head, although posterior third

paler than anterior; poorly defined, faint pale stripe from eye

(b)

Figure 20. Diporiphora granulifera sp. nov.: a, in life, Lawn Hill,

Queensland (photo: S. Wilson). b, c, d, dorsal, ventral and lateral

(head) views of holotype QM J96362 (formerly NMV D74060) Downs

Road, 2 km from Barkly Highway, Queensland.

to top of ear. Dark smudge on posterior of tympanum absent.

On flanks, granular scales in axilla are dark brown then

transition posteriorly into cream, grey and light brown with

little patterning; flanks have darker dorsal patterning above

and pale cream ventrally; flecked with a few light brown

scattered scales. Lateral stripe between axilla and groin

absent. Dark patch in axilla extending up over shoulder and

down full length of scapular fold. Arms lack dark banding. No

patterning on legs, faint banding on anterior third of tail with

dark bands wider than pale bands. Ventral surface cream with

no patterning.

Variation. 44—68 mm SVL; 121—179 mm tail length. Very long

tail, ranging from 2.7—3.0 x SVL. Gular fold always absent,

post-auricular fold weak to present, scapular fold strong,

extending onto edges of ventrum. In some animals, enlarged

scales on post-auricular fold has an enlarged cluster of scales at

the ventral extent but without a single spinous scale. Pre-cloacal

pores variable from 4—6 (usually 4); femoral pores always 0.

Variable patterning from strongly patterned individuals to

plain individuals. In patterned individuals, approximately 6—8

dark brown bands slightly offset to each other on either side of

a broad undefined greyish vertebral stripe. Banding extends

onto tail, fading out half way down tail. Well-defined pale

dorsolateral stripes, running from back of head to base of tail

and becoming interrupted by banding pattern of tail. Flanks

have a faint, poorly defined light brown lateral stripe, with

darker dorsal patterning above and pale cream below. Flanks

flecked with a few light brown scattered scales. In more plain

individuals, pale dorsolateral stripes from neck onto base of

tail. Granular scales in axilla are dark brown; flanks cream,

grey or light brown with little patterning. No white markings

on face, labial scales speckled with light brown flecks. Faint or

no banding on limbs. Ventral surface cream, usually plain but

some individuals have faint dark flecking on ventral surface of

head. Males with breeding colouration tend to lose some of

their dorsal patterning, having a large black patch in axilla

extending on to shoulder and some having a pink flush on base

of tail.

Etymology. Named for the extensive granulated scales on sides

of body.

Distribution and ecology. Restricted to the far north-west Gulf of

Carpentaria region of Queensland. More collecting is required to

determine how far east this species extends and whether it

extends west into the NT. A habitat generalist occurring In

tropical savannah woodlands and grasslands. Has been observed

perching on low vegetation, rocks or termite mounds.

Comparison to other species. The distribution of D. granulifera

sp. nov. contacts a number of other Diporiphora species in the

western Gulf of Carpentaria region. Very similar

morphologically to D. carpentariensis sp. nov. and it remains

unclear whether they come into contact in the central Gulf of

Carpentaria region, but can be distinguished from this species

in lacking or having weak spinose scales on the post-auricular

fold and in having granular scales extending anteriorly to the

scapular fold, with scales on the sides of neck more homogenous

and outer scale row In dorsolateral stripes having raised trailing

edge, particularly over shoulder, giving strong definition

between dorsal and lateral surface. Differs from D. magna in

having granular scales extending over shoulder and along

scapular fold, slightly onto ventral surface of neck. Differs from

D. lalliae in lacking a gular fold. Can be distinguished from D.

sobria in lacking a gular fold and femoral pores, and having

single canines on either side of the upper Jaw.

Remarks. Although very similar morphologically to D.

carpentariensis sp. nov., the genetic analysis of Smith et al.

(2011) across multiple genes distinguished these two species as

divergent lineages that are not each other's closest relatives. It

remains to be determined if these two species contact In the

central Gulf of Carpentaria region.

Recent unpublished phylogenomic research using single-

nucleotide polymorphisms shows that D. granulifera sp. nov. 1s

highly divergent and genetically distinguishable from D. magna

(as defined above) (J. Fenker, unpublished data). However, there

is evidence of gene flow and mtDNA introgression between

these two species in areas of contact, including along the Barkly

Highway, Queensland. The holotype (QM J96362), which was

included in this genomic analysis, was confirmed as being D.

granulifera sp. nov. and is not a hybrid animal, with no evidence

of genomic contribution from D. magna.

Diporiphora granulifera sp. nov. contributes to a small

number of endemic lizard and frog species from the Gulf of

Carpentaria region, including Cryptoblepharus zoticus Horner,

Oedura bella Oliver and Doughty and Litoria electrica Ingram

and Corben. Further genetic and morphological work on species

that span the Gulf of Carpentaria area may reveal further

interesting phylogenetic patterns in this poorly sampled region.

Diporiphora carpentariensis sp. nov.

ZooBank LSID: http://zoobank.org/urn:lsid:zoobank.oreg:act:

6795AD24-03D8-4E4E-9FD8-4757A EE964C5

Common name. Gulf two-lined dragon.

Figure 21, Tables 3, 4

Holotype. QM 37388197 (adult male), Littleton National Park,

northern Queensland (18.2243°S, 142.7489 E). Collected by E.

Vanderduys on 20 November 2008.

Paratypes. NMV D74080 (adult female), Mt Turner Road, 5 km

north of Gulf Development Road, Queensland (18.2692? S,

143.3647? E); NMV D74079 (adult male), Mt Turner Road, 20 km

north of Gulf Development Road, Queensland (18.2733* 5,

143.3656? E); NM V D74076 (adult female), road to Strathmore, 3 km

north of Gulf Development Road, Queensland (18.1783? S,

142.8844" E); NM V D74068 (adult male), Bourke Development Road,

25 km NE of Karumba turnoff, Queensland (17.3931? S, 141.3989? E).

Diagnosis. Moderately large body size (to 65 mm SVL), with

adult males with very long tails (to 3 x SVL) and females with

shorter tails (to 2.3 x SVL). Gular fold absent, post-auricular fold

weak to strong, scapular fold strong. Granular scales in axilla,

extending over arm and along the full length of the scapular fold.

Scales on neck in anterior to scapular fold small but not granular.

Outer scale row in dorsolateral stripes lack raised trailing edge,

without strong delimitation between dorsal and lateral surface.

Pre-cloacal pores 4 or 5 (usually 4); femoral pores 0.

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

Description of holotype. Male; 62 mm SVL; 185 mm tail length.

Medium-large Diporiphora, gracile with long limbs and very

long tail. One canine on either side of upper jaw. Gular fold

absent, post-auricular fold weak and scapular fold strong,

extending onto edges of ventrum. Dorsal scales strongly keeled,

relatively homogenous. Longitudinal series of raised but not

enlarged pale paravertebral and dorsolateral scales at the

(b)

J88197), Littleton National Park, northern Queensland, a, in life (photo:

E. Vanderduys); b, c, d, dorsal, ventral and lateral (head) view of

paratype NM V D74068.

Taxonomic revision of dragon lizards

shoulder, extending down back to base of tail. Scales on flanks

homogeneous, although changing from small granular scales

in axilla to small but non-granular scales on the posterior two-

thirds of flanks. Cluster of small spinose scales on post-

auricular fold with one cream-coloured spine being clearly

larger than the others. Lacks spinose scales on limbs or tail.

Granular scales In axilla, extending over arm and along the full

length of the scapular fold. Scales on neck, anterior to scapular

fold small but not granular. Ventral scales strongly keeled. Pre-

cloacal pores 5 (2 on right; 3 on left); femoral pores 0.

Dorsum light brown with little patterning. Faint, broad

grey vertebral stripe; well-defined cream dorsolateral stripes

running from back of head to base of tail, associated with

raised but not enlarged longitudinal scales. Dark transverse

bands absent. Head relatively plain with little patterning,

labials similar colour to rest of head and lacking pale line

between eye and ear. Dark smudge on posterior of tympanum

absent. On flanks, granular scales in axilla are dark brown

then transition posteriorly into cream, grey and light brown

without patterning; flanks have darker dorsal patterning above

and pale cream ventrally; flecked with a few light brown

scattered scales. Lateral stripe between axilla and groin

absent. Dark patch in axilla, associated with granular scales,

continues up over shoulder and down full length of scapular

fold. Banding on limbs absent; faint banding on anterior third

of tail, with dark bands narrower than the light bands. Ventral

surface cream with no patterning.

Variation. 52—65 mm SVL; 118—187 mm tail length. Very long

tail, ranging from 2.1—3.0 x SVL. Tail length appears to be

sexually dimorphic with adult male tails ranging from 2.7—3.0

x SVL, while adult female tails ranging from 2.1—2.3 x SVL.

Gular fold always absent, post-auricular fold weak to present,

scapular fold strong, extending onto edges of ventrum. In some

animals, a cluster of small spinose scales on post-auricular fold

has an enlarged scales that is not obviously spinous. Pre-cloacal

pores variable from 4—5 (usually 4); femoral pores always 0.

Variable patterning from strongly patterned individuals to

plain individuals. In patterned individuals, eight or nine dark

brown bands slightly offset to each other on either side of a

broad undefined greyish vertebral stripe. Banding extends

onto tail, fading out halfway down tail. Well-defined pale

dorsolateral stripes, running from back of head to base of tail

and becoming interrupted by banding pattern of tail. Flanks

have faint poorly defined light brown lateral stripe, with darker

dorsal patterning above and dark background colour extending

slightly below stripe. Flanks flecked with a few light brown

scattered scales. In more plain individuals, pale dorsolateral

stripes from neck onto base of tail. Granular scales in axilla

are dark brown or black, flanks cream, grey, light brown with

little patterning. No white markings on head, labial scales

speckled with light brown flecks. Faint or no banding on legs

and tail. Ventral surface cream, usually plain but some

individuals have faint dark flecking on ventral surface of head.

Males with breeding colouration tend to lose some of their

dorsal patterning, having a large back patch in axilla extending

onto shoulder but not anteriorly of the scapular fold, and some

males have a pink flush on base of tail.

Etymology. Named for the region in which this two-lined

dragon occurs on the Gulf of Carpentaria.

Distribution and ecology. Diporiphora carpentariensis sp. nov.

is restricted to the far north-east Gulf of Carpentaria region of

Queensland at the western extent of Cape York Peninsula.

More collecting is required to determine whether this species

extends further west into the Gulf of Carpentaria region. It

appears to be habitat generalist, occurring in tropical savannah

woodlands and grasslands. Little is known about this species

and further field-based work is needed to determine the extent

of its distribution and its ecological requirements, habitat

preferences and behaviour.

Comparison to other species. The distribution of D.

carpentariensis sp. nov. overlaps a number of other Diporiphora

species in the Gulf of Carpentaria region. Very similar

morphologically to D. granulifera sp. nov. and remains unclear

whether they contact in the central Gulf of Carpentaria region but

can be distinguished from this species in having spinose scales

on the post-auricular fold, with a single spine clearly larger than

the others, and in lacking granular scales extending anteriorly

from the scapular fold, and outer scale row in dorsolateral stripes

lack raised trailing edge without a strong demarcation from the

dorsal to lateral surface. Diporiphora carpentariensis sp. nov.

differs from D. jugularis in having a strong scapular fold,

eranular scales in axilla that extend over shoulder and along

scapular fold, scales on flanks relatively homogeneous and

lacking a black gular band or black spot on sides of neck.

Diporiphora carpentariensis sp. nov. can be distinguished from

D. australis in lacking a gular fold and having granular scales in

axilla that extend over shoulder and along scapular fold.

Remarks. This species has previously been identified as

Diporiphora bilineata. It is probable that animals previously

identified as D. bilineata on Cape York Peninsula actually

comprise two species: D. carpentariensis sp. nov. and D.

jugularis. Phylogenetic work (Smith et al., 2011) clearly shows

that this species in unrelated to either D. bilineata or D.

jugularis (fig. 1).

(d) Arid zone species group

Content:

. adductus Doughty, Kealley and Melville, 2012

. ameliae Emmott, Couper, Melville and Chapple, 2012

. linga Houston, 1977

. paraconvergens Doughty, Kealley and Melville, 2012

. pindan Storr, 1980

. valens Storr, 1980

. vescus Doughty, Kealley and Melville, 2012

. winneckei Lucas and Frost, 1896

D. pallida sp. nov.

TO OO SS D T

Diagnostic characters for group in north-western Kimberley:

° one canine tooth on each side of upper Jaw

e axilla granular scales absent

e lateral dark spot absent

* femoral pores absent.

Remarks

The arid zone species group is widely distributed across arid

WA (Couper et al., 2012; Doughty et al., 2012a), with highest

species diversity in the Pilbara region but extending north to

the southern Kimberley (D. pindan) and east into the arid

interior (D. paraconvergens), crossing into South Australia

and the NT. However, D. pallida sp. nov. is from the Mitchell

Plateau (fig. 22), north-western Kimberley, more than 450 km

north of the nearest member of this species group (D. pindan).

Diporiphora pallida sp. nov. expands the bioclimatic and

distributional limits of the arid zone species group. Key

characters in distinguishing the arid zone species group from

the other species groups in northern Australia is the number of

canine teeth on each side of the upper jaw combined with the

absence of granular scales in the axilla. In addition to these

characters, in the Kimberley region this species group can be

distinguished from D. lalliae (a member of the D. bilineata

species group) by species-specific characters: the lack of a

cular fold for D. pindan and a short tail for D. pallida sp. nov.

Diporiphora pallida sp. nov.

ZooBank LSID: http://zoobank.org/urn:Isid:zoobank.org:act:

7967A 170-9260-40F9-A895-D5C5041 E4729

Common name. Pale two-pored dragon.

Figure 22, Tables 3, 4

Holotype. WAM R177292 (formerly NMV D73853) (adult male),

Mitchell Plateau, WA (14° 49.45' S, 125° 42.12' E). Collected by J.

Melville on 12 September 2005.

Diagnosis. Small body size (to 46 mm SVL), short tail (2.0 x

SVL), robust head and prominent brow above eye. Gular fold

strong, post-auricular fold weak, scapular fold present. Single

canine In upper jaw, no granular scales in axilla. Pre-cloacal

pores 2; femoral pores 0.

Description. Male; 46 mm SVL; 91 mm tail length. Small

Diporiphora, robust head with relatively short tail (~2.0 x SVL)

and limbs. One canine on either side of upper jaw. Gular fold

strong, post-auricular fold weak and scapular fold present.

Prominent ridges above eye, extending along canthal ridge to

dorsal corner of nasal scale, with short tapered snout. Dorsal

scales strongly keeled and homogenous, running parallel to the

midline. Scales on flanks homogeneous, keels angled posteriorly

and dorsally. One small white spinose scale at back of head

sitting on ventral end of post-auricular fold. Lacks spinose scales

on limbs or tail. Small scales in axilla but not granular. Ventral

scales strongly keeled. Pre-cloacal pores 2; femoral pores 0.

Dorsum relatively plain with little patterning. Broad (~4

scales wide) greyish indistinct vertebral stripe and broad

yellow-cream dorsolateral stripes (~3 scales wide) running

from back of head to base of tail; dorsolateral stripes continuous

from head to mid-dorsum, and from there broken into sections

by background colour. Dark transverse bands absent. Head

relatively plain with little patterning, labials similar colour to

rest of head and lacking pale line between eye and ear. Dark

smudge on posterior of tympanum absent. Flanks pale without

patterning. Lateral stripe between axilla and groin absent.

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

Dark patch in axilla absent. Banding on limbs and tail absent.

Ventral surface cream with no patterning.

Variation. This description is based on a single specimen

collected on the Mitchell Plateau, thus the variation in this

Species 1s unknown.

Etymology. Named for the pale appearance of this species in

life. Used as an adjective.

Distribution and ecology. Based on a single animal, D. pallida

sp. nov. occurs on the Mitchell Plateau in the northwest

Kimberley. This animal was found perched in spinifex grass on

a rocky outcrop (pictured in fig. 22). Little is known of this

species but it appears to be associated with spinifex grasses on

rocky substrates.

Comparison to other species. Unlike other dragons found in the

northwest Kimberley, D. pallida sp. nov. appears to be a habitat

specialist In spinifex grasses and has a distinctive morphology.

Its distribution overlaps with D. margaretae, but differs in

having a gular fold. It also differs from D. albilabris, D.

bennettii and D. perplexa sp. nov. in having a single canine on

either side of upper jaw and further differs from D. albilabris

in lacking femoral pores.

Remarks. Genetic analyses indicate that this species is not

related to other Diporiphora species in the Kimberley but

instead belonging to the arid zone species group containing

other spinifex specialist species (e.g. D. linga and D. winneckei),

which occur in arid central Australia. In mtDNA phylogenetic

analysis, this species was resolved as the sister to D.

paraconvergens, but this relationship was not strongly supported

(fig. 1). Diporiphora pallida sp. nov. can be readily distinguished

from D. paraconvergens by dorsal scales that run parallel to the

midline (vs. converging) and by having a short tail (vs. very

long). Based on genetic results and morphological distinctiveness

we feel confident that this a new species for the Kimberley

region, joining D. convergens as another Diporiphora species

known only from the holotype. Further sampling is required to

gain a better understanding of the ecology, biology and

evolutionary relationships of this little-known species.

Discussion

Our study provides the first comprehensive taxonomic treatment

of Diporiphora species across northern Australia since Storr’s

revision in 1974. Using an understanding of relationships based

on genetic data and a re-examination of newly collected and

historical specimens, we describe five new species, raise a

further two taxa to full species status and redefine six existing

species. This work significantly increases the species diversity

of Diporiphora, and agamid lizards, in the AMT.

Diporiphora species occur in most habitats in the AMT and

are one of the most common and abundant lizards in the tropical

savannah woodlands and grasslands. The Australian tropical

savannah is one of the largest and most intact in the world

(Bowman et al., 2010; Laver et al., 2018), and is globally

significant. Despite this importance, we are only beginning to

document the true species diversity of many vertebrate groups

(e.g. Afonso Silva et al., 2017; Doughty 2011; Doughty et al.,

Taxonomic revision of dragon lizards

2012b, 2018; Laver et al., 2018; Melville et al., 2018; Pepper et

al., 2011; Potter et al., 2012). With these recent studies and our

taxonomic treatment documenting lizard diversity patterns In

geckos, skinks and agamids, we can start to look for concordant

patterns of diversity between these lineages. Three clear

patterns are apparent across these major lineages: (1) greater

species diversity In the northern extent of the AMT, particularly

the Kimberley region; (2) fewer and more widely distributed

species in southerly portions of the AMT; and (3) consistent

major biogeographic breaks across terrestrial vertebrate groups.

A consistent pattern In studies investigating

phylogeographic structuring and species diversity across the

AMT is a north-south gradient of diversity In lizard lineages.

In the northern parts of the AMT, species diversity is greatest,

with particular diversity hotspots in the north-western

Kimberley and Arnhem Land regions (e.g. Powney et al.,

2010). In addition to higher species diversity, there are higher

numbers of short-range endemic species in these northern

hotspots. For example, in the Oedura geckos of the AMT,

genetic evidence suggests that microendemism and diversity

is highest in the northern AMT high-rainfall regions (Laver

et al., 2018), while the most widespread lineages occur in the

more southerly boundary between the AMT and Australian

arid zone. Similarly, in Diporiphora the highest species

diversity is in the Kimberley, with three lineages (D. bilineata,

D. bennettii and arid zone species groups) occurring in the

region. Each of these species groups contain more than one

species in the Kimberley, with all four species from the D.

bennettii species group occurring in the region, including two

locally endemic (D. bennettii and D. albilaris). In contrast,

the southern region of the AMT has fewer Diporiphora

species but they have broad east—west distributions (D. sobria,

D. magna and D. lalliae), with the later of these encroaching

into the Australian arid zone. This north-south pattern of

Species diversity, which appears across multiple groups, has

been attributed to an aridity gradient, climate stability or

instability, and historical refugia in mesic areas (Afonso Silva

et al., 2017; Laver et al., 2018; Palmer et al., 2013). The

hypothesis that the presence of rocky refugia drives patterns

of diversity 1s supported by higher diversity along the barrier

ranges in the southern Kimberley (Doughty et al., 2018;

Oliver et al., 2014, 2016).

In addition to documenting greater species diversity in

Diporiphora, our work also demonstrates that there is high

diversity In body size and ecology within the Kimberley

region, with large and small sympatric species within each

Species group and with ecological variation. In the D. bennetti

species group, there are two rock-specialist species, one large

(D. perplexa sp. nov.) and one small (D. bennettii), and a

smaller generalist woodland species (D. albilabris). Similar

body size divergence in sympatric rock-dwelling Gehyra

species has been documented in the Kimberley (Moritz et al.,

2018; Oliver et al., 2016). In the D. bilineata species group,

there is a generalist woodland species (D. margaretae) with

smaller body size in the northern Kimberley and a gracile

species found In grasslands on river floodplains in the southern

Kimberley (D. gracilis sp. nov.), although it 1s unlikely these

species are sympatric. These patterns suggest that more than

10 mm

Figure 22. Diporiphora pallida sp. nov. Images of holotype (WAM

R177292, formerly MNV D73853), Mitchell Plateau, Western

Australia: a, in life (photo: J. Melville); b, c, dorsal and ventral views;

d, collection location.

mesic refugia and aridity gradients are shaping diversity in

Kimberley Diporiphora species, and that ecological and

morphological diversification through processes other than

evolutionary drift may drive diversity patterns. Regardless of

the mechanisms, there is a clear pattern for high species

diversity in the Kimberley, with more widespread species

within the AMT appearing to arise within these Kimberley

species groups.

In the more southerly regions of the AMT, this diversity in

body size is not apparent In Diporiphora. Across the east—west

span of the AMT, past climatic fluctuations and aridity

eradients have probably shaped species diversity. A number of

major biogeographic breaks have been identified 1n different

lizard lineages, several which appear to be relatively consistent

across groups (e.g. Noble et al., 2018). Genetic differentiation

across two biogeographic barriers — the Carpentaria Gap in

the Gulf of Carpentaria region of Queensland and the Ord

River region between the Kimberley region and the Top End of

the NT — has been well documented across numerous taxa

(Catullo et al., 2014; Noble et al., 2018; Potter et al., 2012),

including agamid lizards (see Pepper et al., 2017). In

Diporiphora, the divergence across the Carpentaria Gap

appears to be most dramatic. For example, in the D. bilineata

species group, the distributions of D. bilineata, D. magna and

D. granulifera sp. nov. appear to truncate at the western

portions of the Carpentaria Gap, while D. carpentariensis sp.

nov. 1s to the east of this biogeographic barrier. In addition, the

D. australis species group is only found to the east of the

Carpentaria Gap (see Edwards and Melville, 2010, 2011). The

divergence of species across the Gulf of Carpentaria region

has been attributed to increased aridity in these regions

compared to adjacent topographic uplands (Pepper et al.,

2017). It remains unclear as to whether AMT Diporiphora

species groups have a distributional gap in the Gulf of

Carpentaria region of Queensland, as is currently suggested

by maps, or is this due to a lack of sampling and records in this

region. Further field-based research 1s needed to fully elucidate

the distributions of Diporiphora species across this

biogeographic barrier.

Acknowledgements

JM, PD and PH examined specimens at the following

institutions: Australian Museum, Sydney; Western Australian

Museum; Museums Victoria; Queensland Museum; Museum

and Art Gallery of the Northern Territory; Natural History

Museum, United Kingdom, Naturhistorisches Museum,

Vienna and Naturhistoriska Riksmuseet, Stockholm. We

would like to thank staff and researchers from these institutions

for their help and assistance, in particular, P. Wagner (Bonn,

Germany) for organising type loans to be examined. We thank

S. Wilson, E. Vanderduys, M. Bruton and the Australian

Wildlife Conservancy for the use of images. We thank G. Shea

for advice on nomenclature and discussions regarding

synonymies and we thank A. Bauer for providing a high-

resolution scan of the D. australis syntype from the original

species description. Research funding provided to JM by the

Australian Research Council.

J. Melville, K.L. Smith Date, P. Horner & P. Doughty

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Memoirs of Museum Victoria 78: 57—64 (2019) Published 2019

1447-2554 (On-line)

htto://museumvictoria.com.au/about/books-and-journals/journals/memoirs-of-museum-victoria/

DOI https://doi.org/10.24199/j.mmv.2019.78.03

A redescription of Eulimnadia rivolensis (Brady, 1886) (Branchiopoda:

Spinicaudata: Limnadiiidae), and its transfer to Paralimnadia

BRIAN V. TIMMS

Honorary Research Associate, Australian Museum, | William St, Sydney, New South Wales, 2010 and Centre for

Ecosystem Science, School of Biological, Earth and Environmental Sciences, University of New South Wales, Kensington,

New South Wales, 2052 (brian.timms@unsw.edu.au)

Eulimnadia rivolensis occurs across the southern Australian mainland and Tasmania but has not been collected in

Abstract Timms, B.V. 2019. A redescription of Eulimnadia rivolensis (Brady, 1886) (Branchiopoda: Spinicaudata: Limnadiiidae),

and its transfer to Paralimnadia. Memoirs of Museum Victoria 78: 51—64.

Victoria since 1910 and in south-east South Australia since 1975, where its former habitat has been destroyed. E. rivolensis

is redescribed from syntype material and transferred to Paralimnadia. This species lacks a subcercopod spine and has

other less characteristic features of Paralimnadia. Eulimnadia palustera Timms, 2015 1s a junior synonym based on egg

morphology and some characteristics of the telson.

Keywords Subcercopod spine, cercopods, resting eges, Eulimnadia palustera

Introduction

The taxon Eulimnadia rivolensis Brady, 1886, of southern

Australia has had a chequered history. It was first applied to

specimens from the Rivoli Bay environs in south-eastern

south Australia, but the description 1s only of the carapace

shape, which Is now known to vary with age and habitat

(Rogers et al., 2012; StraSkraba, 1965), and the illustration

could apply to many limnadiid species. Next, the name was

used without any justification by Spencer and Hall (1896) for

specimens supposedly from Central Australia. Sayce (1903)

published some details of the morphology of E. rivolensis and

provided more accurate drawings, basing his observations on

specimens from Victoria and South Australia. Sayce's (1903)

Onkaringa (Onkaparinga) Creek site was erroneously recorded

as being In Central Australia, but it drains the eastern Adelaide

Hills, and so this error introduces possible inaccuracies in

distribution. Sayce (1903) suggested, without any evidence,

that E. rivolensis may be synonymous with Limnadia sordida,

which at that time was also poorly defined. Thus, New South

Wales, the habitat of L. sordida, was added to the supposed

distribution. This synonymy was perpetuated by Dakin (1914),

who noted its occurrence 1n south-western Western Australia,

and Henry (1924), except that L. sordida was moved to

Eulimnadia. To add to the confusion, Daday (1925) retained

the specific epithet rivolensis but transferred it to Limnadia.

This was followed by Richter and Timms (2005), based on

figures in Sayce (1903), and Gurney (1927) because neither

recorded a subcercopod spine, a defining feature of Eulimnadia

(Martin, 1989). Importantly, the species epithet was hidden in

the synonymy, so its possible existence was not acknowledged

in a recent review of Australian Eulimnadia (Timms, 2016a).

In 2015, I described Eulimnadia palustera from south-

west Western Australia, which shares some features with

sayce's version of E. rivolensis, then thought to be L. sordida.

Given that egg morphology in limnadiids is useful in

distinguishing species (Belk, 1898; Rabet, 2010; Rogers et al.,

2012; Timms, 2016a, 2016b), a comparison of the eggs from

the few collections labelled E. rivolensis in the NMV and AM

suggest a close similarity between the two species. In

summary, there is uncertainty over the validity of E. rivolensis

and E. palustera, to what species they are related and, indeed,

to which genus they belong. Fortunately, there is enough

material in the Australian Museum and National Museum

Victoria to find solutions to these uncertainties. One subsidiary

aim is to accurately plot the distribution of these species.

Material and methods

Drawings were made using a Wild M5 dissection microscope

equipped with a camera lucida. Body measurements were

made by placing a template marked in 0.5 mm spacings

underneath the specimen at magnifications of 10—40 x and

distance was estimated to the nearest half division. Accuracy

is deemed to be + 0.25 mm.

Eggs were prepared as detailed in Timms and Lindsay

(2011) and studied on a Zeiss Evo LS15 Scanning Electron

Microscope using a Robinson Backscatter Detector.

lerminology of the claspers of the Diplostraca follows

Kaji et al. (2014): the hand (or palm) is composed of endites I V

and V; its thumb (or gripping knob) is derived from endite IV,

the small palp from endite IV and the large palp from endite

V; and the finger is derived from endite VI (or endopod).

Other abbreviations used in the text: AM = Australian

Museum; BMNH = British Museum of Natural History; NMV

= National Museum Victoria; SAM = South Australian

Museum; WAM = Western Australian Museum.

Results

Taxonomy

Diplostraca Gerstaecker, 1866

Spinicaudata Linder, 1945

Limnadiidae Baird, 1849

Paralimnadia Sars, 1896, emend Rogers et al., 2012.

Paralimnadia rivolensis Brady, 1886

Figures 1—4

Eulimnadia rivolensis Brady, 1886: 86—67, fig. D. — Simon,

1886: 456 (list); Spencer and Hall, 1896: 238; Sayce, 1903: 245-246

(text), 248 (synopsis), pl. 32; Wolf, 1911 (list); Dakin, 1914: 295 (list),

300 (text); Gurney, 1927: 60—61, fig. 1A.

Limnadia rivolensis — Daday, 1925: 150 (key), 173—175, fig. 121;

Webb and Bell, 1979: 243 (text), table 1; Richter and Timms (text): 348.

Eulimnadia palustera Timms, 2015: 447—449, fig. 6. New synonym

Lectotype. South Australia, hinterland of Rivoli Bay, R. Tate,

date unknown but before 1886, BMNH 1890.2.1.9. Male 9.0

mm long and 6.0 mm high.

Paralectotypes. South Australia, hinterland of Rivoli Bay, R.

Tate, date unknown but before 1886, BMNH 1890.2.1.10. Male

8.9 mm long and 5.5 mm high; Rivoli Bay, freshwater swamps,

11 November 1882, collector unknown, 4 males, 1 female,

NMV J14426.

Comment. Because the Brady collection in the National Museum

Victoria 1s labelled from Rivoli Bay and has a date that aligns

with the approximate date of collection of the lectotype, I

believe the two are the contemporaneous. Hence, the collection

NMV J14426 are herein designated as paralectotypes, which 1s

convenient given there are no females or eggs among the

original syntypes In the British Museum of Natural History.

Other material. Northern Territory: Central Australia, 5 males,

5 females, from Sayce collection but no further data, NMV

J54016; South Australia; Lake Robe, nearby puddle, Margaret

Brock, 23 September 1975, 3 males, 1 female, SAM C12297;

snake Lagoon, Kangaroo Island, South Australia, 24 August

1981, D.J. Williams, 4 males, 2 females, SAM C12296;

Tasmania: no site recorded, R.W. Davis, 18 October 1969, 4

males, 2 females, NMV J46599; 4 km north of Campbelltown,

41.93? S, 147.5? E , 24 November 1963, J. Wilson, 1 male, 2

females, AM P55663; 4 km north of Campbelltown, 41.95? S,

147.5? E, 20 March 1964, J. Wilson, 7 males, 10 females, AM

P55640; 1 female, AM P98988; 1 male, AM P99519; 1 female,

AM P99520; Coles Bay, 31 December 1964, no collector

recorded, 30 males, 42 females and 28 sex uncertain, NMV

54005; Campbelltown, 23 October 1965, no collector recorded,

B.V. Timms

] female, NMV J46622; Bruny Island, between Big Lagoon

and Little Lagoon, 21 September 1975, R.B. Manning, 3 males,

4 females, NMV J46600; Victoria: Elwood Swamp, 18 July

1899, collector unrecorded, 23 males, 24 females, NMV

J53989; Elwood, from Sayce collection but no further data but

co-types for E. victoriensis Sayce, 4 individuals sex uncertain,

NMV J68583; Mordialloc, 25 October 1902, collector

unrecorded, | male, NMV J46622; Cheltenham, 22 October

1910, collector unrecorded, 17 males, 25 females, NMV

J53987; Cheltenham, from Sayce collection but no further data,

4 individuals sex uncertain, NMV J54049;

Diagnosis. Egg astroform, projections grooved. First antenna

with about 11 lobes, second antenna of about 12 antennomeres.

Trunk 18-20 segmented, long palps of claspers with 2—3

palpomeres and palpomere junctions generally inerm. Telson

with about 20 dorsal spines, first 5 usually larger and more

spaced than others. Cercopod basal section about 60% of total

length and bearing about 8 setae of medium length.

Description. Male: Head (fig Ib) with ocular tubercle prominent,

the compound eye occupying most (~80%) of it. Rostrum

slightly more prominent than ocular tubercle, also slightly

asymmetrical and with a rounded apex. Ocellus triangular

dorsobasially in rostrum. Frons-rostrum angle about 90*.

Dorsal organ posterior to eye by about its half its height,

pedunculate about height of ocular tubercle.

First antenna (fig. 1b) distinctly longer than peduncle of second

antennae, with 11 lobes, each with numerous short sensory

setae. Second antenna (fig. ld) with a spinose peduncle

subequal to length to the rostrum, each flagellum with 11

antennomeres dorsally with 1—2 spines and ventrally with 1—7

longer setae. Basal and distal antennomeres with minimal

spines, setae maximal on antennomeres 4—9 and only 1—3 setae

on basal 3 antennomeres.

Carapace (fig. la) elongated oval, pellucid and with weakly

expressed growth lines, numbering about 9. Older growth lines

well spaced compared with closer-spaced newer growth lines

near carapace margin. Both anterior and posterior angles

hardly noticeable.

Twenty pairs of thoracopods, the first two modified as

claspers. Claspers (fig. If) with palm (endites IV and V)

trapezoidal with a slight rounded protrusion distomedially.

Apical club (endite I V) rounded with thick denticles distomedially

and many spines apicolaterally, moveable finger (endite VI) of

normal curved structure and palps of typical structure. Moveable

finger terminating in a suctorial disc and distoventrally with

many small pits. Long palp (endite V) subequal in length to the

palm in the first clasper and about 1.5x longer in second clasper.

short and long palps, both with three palpomeres with junctions

between them inerm. Last palpomere the longest, particularly 1n

the second-longest palp. Other thoracopods of typical structure

for Eulimnadia, decreasing In size and complexity after 10th

thoracopod. Dorsal surface of trunk (fig.le) with a short spine

posteriormedially on each of the 12 posterior trunk segments.

Telson (fig. 1c) with about 20 pairs of dorsal spines, with

the first three larger than the next 17, although these generally

increase in length posteriorly. Most spines inerm. Caudal

On Paralimnadia rivolensis (Branchiopoda: Limnadiidae)

filaments originating from a mound a little higher than the

dorsal telsonic floor and between the 4th and 5th spine. This

dorsal floor posterior to the mound with a moderate declivity

then an even slope to cercopod posterior. Cercopod almost as

long as the telson dorsum, the basal 60% hardly thinning to a

small naked spine, then rapidly thinning to an acute apex. The

basal 60% with about 8 short setae dorsolaterally; length of

most about basal cercopod diameter, with setae 5thto 7th

longest and the last one the shortest). Many tiny denticles

dorsolaterally on apical 40% of cercopod. All setae geniculate.

Triangular projection beneath the cercopods at the

ventroposterior corner of the telson.

Comments

Three previous authors have commented on aspects of the

morphology of this species. Brady’s (1886) original description

is ambiguous and could apply to many limnadlids. Sayce (1903)

notes the 20 trunk segments, which are unusual among

Eulimnadia and Paralimnadia (Timms, 2016a, b), and comments

on 20 telsonic spines and proximal half of cercopod bearing

about 10 shortish seta Brady’s (1886) illustration (Plate X X XII)

confirms a rounded protruding rostrum in the male, many

erowth lines similar to that described presently from the syntypes

(Brady, 1886) and a similar clasper also as described presently

but with 1—2 spines at the palpomere junctions. Sayce (1903)

Figure 1. Drawings of types of P. rivolensis, male lectotype BMNH 1890.2.1.9. A, carapace; B head, C, telson and cercopod; D, an antennal

flagellum; E trunk segments XIV to XV VII dorsa; F. clasper I with insert of long palp of clasper II; female from paralectotype NM V J14426; G.

head; H, telson and cercopod. Scale bars | mm.

60 B.V. Timms

, " d

P d

Figure 2. Digital images of male P. rivolensis from NMV J55640. A, male carapace; B, male head, C, male telson; D, male claspers,

E, female head.

On Paralimnadia rivolensis (Branchiopoda: Limnadiidae)

illustrates a third trunk segment with a long palp of endite V, as

is typical of Eulimnadia and Paralimnadia (Timms, 2016a,

2016b). Gurney (1927) illustrates a male telson, which besides

showing some variation in size and spacing of 21 telsonic spines,

clearly shows a basal cylindrical 45% of the cercopod with 7

setae of moderate length and the telsonic base under the cercopod

insertion with a mild triangular protrusion. Gurney (1927) could

not find any growth lines. Nowhere in any of these three early

descriptions Is a subcercopod spine mentioned or illustrated.

This subcercopod spine is also absent in all of the material

seen in the Australia Museum and National Museum Victoria,

in all cases being replaced by a triangular protrusion of various

sizes. Also, given that all specimens examined have 11-12

antennomeres and cercopods with basal 45-60% bearing

setae (Timms 2016b), the conclusion is inescapable that this

species belongs to Paralimnadia, not Eulimnadia. Further

indication that it Is a Paralimnadia and not a Eulimnadia,

although not absolute (Timms, 20162), is that the sex ratios are

Figure 3. Drawings of male and female of P. rivolensis from NM V J55640. A, male carapace; B, male head; C, male antennal flagellum; D, male

clasper I, E, male telson; F, female carapace; G, female head; H. female telson. Scale bars 1 mm.

broadly 1:1 and not female or hermaphrodite dominated. This

indicates gonochoristic reproduction and not the androdioceous

reproduction that is characteristic of Eulimnadia (Timms,

2016a, Weeks et al., 2008).

While the lectotype has 20 trunk segments, all other

material seen has 18 trunk segments, including the paralectotypes

in NMV J14426.

There are no females in among the original syntypes (now

lectotype and a paralectotype), so the single female in NMV

J14426 was studied (fig. 3).

Head (fig. 3g) with ocular tubercle prominent with a

compound eye occupying much of it (SO—/0% in preserved

material). Rostrum a smooth bulge about as prominent as the

ocular tubercle and at an angle of about 120° to the frons.

Ocellus not visible and dorsal organ apparently missing.

First antenna (fig. 3g) a little shorter than peduncle of the

second antenna, and with five small lobes with short sensory

setae. Second antenna as 1n male.

Carapace (fig. 3f) as in male, although dorsum more vaulted.

Nineteen thoracopods of typical Eulimnadia structure.

Trunk dorsum with 3—9 setae terminally, these setae few, short

and stout on posterior few segments, numerous and longer on

B.V. Timms

segments 8-15, and hardly any setae on anterior trunk

segments l-7.

Telson (fig. 3h) dorsally on each side with 4 larger and more

robust spines anteriorly followed by 21 small spines slightly

increasing in length posteriorly and terminating in a large spine.

Most spines inerm. Telsonic filaments inserted on a mound

between the 4th and 5th spines. Cercopod subequal in length to

the telson with a cylindrical basal section about 60% of its length

followed by a rapidly thinning apical section with many denticles

dorsally, the two sections separated by a spine. About 7 setae on

the basal section, all a little longer than the diameter of the

cercopod, but with the 4th to 6th a little longer again. A blunt

triangular projection posteriorly ventral to the cercopod base.

Egg (fig. 4) astroform with 14—20, mean 16.8 + 2.6 (n = 10)

projections, each subtended by 3—8 sharp-edged grooves in

different planes arranged radically around its base. One to three

of these grooves reach the projection apices on any one aspect

of the projection. Projections often bent, length—base ratio

varying from 1-2.5 (n= 10). Egg diameter 325 + 31 um (n= 10).

Variability. Only the lectotype and paralectotypes have 19—20

trunk segments; all other specimens examined had the usual 18

segments of Paralimnadia and Eulimnadia (Timms, 20162,

Figure 4. SEM images of eggs of P. rivolensis. A, from type locality Rivoli Bay, SA, freshwater swamps, NMV J14426; B, from Campbelltown

Tas, AM P55640; C, from Bruny Island, Tas, NMV J46600; D, from Central Australia, NMV J54016; E, from Flinders Island, Tas, AM P97387;

F, swamps from near Lake Muir, WA, WAM C57251

On Paralimnadia rivolensis (Branchiopoda: Limnadiidae)

2016b). Male antennomeres ranged from 11—13, and lobes on

the first antennae perhaps varied by one unit. The number of

telsonic spines was more variable (20—26, but typically 21),

while the palpomeres and cercopod setae were also variable.

Palpomere numbers ranged from 2-3, again the lectotype and

paralectotype the only collections with 3 + 3. The cercopod

setae ranged from 7-10 and their lengths varied a little from

being uniform and of moderate length (1.e. ca. 1.5x cercopod

diameter) to being of variable length, some being subequal to

cercopod diameter. The lack of a dorsal organ on the female

paralectotype is most unusual.

Synonymy of E. palustera

This species was originally assigned to Eulimnadia on the sole

criterion of an apparent spine beneath the cercopod base

(Timms, 2015). However, this spine is not atypical subcercopod

spine of most Eulimnadia but a rather sharp triangular

ventroposterior corner of the telson. Hence, an assignment to

Paralimnadia is necessary. Furthermore, three other features

suggest placement in Paralimnadia: a cercopod with a spine

approximately midlength and not at about 80% of its length, 13

antennomeres rather than about 8, and a sex ratio approximating

1:1, all generally (but not absolutely) indicating Paralimnadia

(Timms, 2016a, 2016b).

Given the placement of P. palustera within Paralimnadia,

its eggs are identical with those of P. rivolensis being astroform

with 14—20 projections subtended by 3—8 sharp-edged grooves

(fig. 4). Egg morphology has proved to be the most reliable

character separating species within Eulimnadia (Belk, 1998;

Rabet, 2010; Rogers et al, 2012; Timms, 20163) and

Paralimnadia (Timms, 2016b). The next most reliable species

indicator in both genera is the nature of the cercopod setae.

Both P. palustera and P. rivolensis have about 8 medium length

(1.e. 1—2x cercopod diameter) setae (cf. fig. 6 in Timms, 2015

and figs 1—3). Again, both species have about 21 telsonic spines,

although spacing 1s different In the two species. [n P. rivolensis,

all are evenly sized and spaced, except for the first three, which

are larger and more spaced. In P. palustera, the telsonic spines

are mixed in size (cf. fig 6 in Timms 2015 and figs 1—3). Two

characters generally of poor differentiating ability are the first

antennae and rostrum, although in these two species, there are

only minor differences (cf. fig 6 in Timms and figs 1—3).

The claspers are somewhat different between the two

species. P. palustera has a distinct hamulus medially on the

hand (endite IV), while P. rivolensis has Just a slight swelling

there. The palps are variable, with 3 palpomeres in the

paralectotype of P. rivolensis, but only 2 indistinct ones in

most other specimens examined. P. palustera generally has 3

palpomeres but may have the second division indistinct or

incomplete. Sometimes there are spines at palpomere junction

1-2 in P. rivolensis. Similar variability has sometimes been

observed in a few other Paralimnadia species (Timms 2016b).

Distribution. South-western Western Australia, south-eastern

South Australia, southern Victoria and Tasmania. There is a

single record from central Australia, which is difficult to accept

considering the prominent maritime distribution across southern

Australia. It has not been collected in Victoria since 1910, its

habitat in the swamps of eastern Port Philip Bay being drained

and urbanised in the early 1900s. Widespread drainage in the

south-east of South Australia seems to have denied it habitat

there. The most recent collection from near the type locality is

dated 1975, and my expeditions there in the spring of 2010 and

winter of 2016 were unsuccessful. Sites In central Tasmania

seem (as of March 2018) also to be drained, so that perhaps it

now only occurs in refuges of Flinders Island, Kangaroo Island

and south-western Western Australia.

Acknowledgements

I thank Michael Geddes for passing on to me two collections of

P. rivolensis from South Australia; the curators and collection

managers of the Australian Museum, British Museum of Natural

History, National Museum of Victoria, South Australian

Museum, for facilitating of loans of collections, Ron Lovett

for digital images and D. Christopher Rogers for discussions

and criticism of the manuscript.

References

Belk, D. 1989. Identification of species in the conchostracan genus

Eulimnadia by egg shell morphology. Journal of Crustacean

Biology 9: 115—125.

Brady, G.S. 1886. Notes on freshwater Entomostraca from South

Australia. Proceedings of the Zoological Society of London 1886:

82—93, pl 8-10.

Daday, E. 1925. Monographie systématique des Phyllopodes

Conchostracés. Troisiéme partie. Annales des Sciences Naturelles,

Zoologie 10e série 8: 143—184 (453—504).

Dakin, W.J. 1914. Fauna of Western Australia. II. The Phyllopoda of

Western Australia. Proceedings of the Zoological Society of

London 1914: 293—305.

Gurney, R. 1927. Some Australian freshwater Entomostraca reared

from dried mud. Proceedings of the Zoological Society of London

1927: 59-79.

Henry, M. 1924. A monograph of the freshwater Entomostraca of New

South Wales. Part IV Phyllopoda. Proceedings of the Linnean

Society of New South Wales 1924: 120—137.

Kap, T., Frisch, M, Schwentner, M., Olesen, J., and Richter, S. 2014.

Male claspers in clam shrimps (Crustacea, Branchiopoda) in the

light of evolution: A case study of homology versus analogy.

Journal of Experimental Zoology (Molecular and Developmental

Evolution) 322B: 269—280.

Martin, J.W. 1989. Eulimnadia belki, a new clam shrimp from

Cozumel, Mexico (Conchostraca: Limnadiidae), with a review of

Central and South American species of the genus Eulimnadia.

Journal of Crustacean Biology 9: 104—114.

Olesen, J., Martin, J.W, and Roessker, E.W. 1996. External morphology

of the male of Cyclestheria hislopi (Baird, 1850) (Crustacea,

Branchiopoda, Spinicaudata), with a comparison of male claspers

among the Conchostraca and Cladocera and its bearing on

phylogeny of the ‘bivalved’ Branchiopoda. Zoologica Scripta 25:

291-316.

Rabet, N. 2010. Revision of the egg morphology of Eulimnadia

(Crustacea, Branchiopoda, Spinicaudata). Zoosystema 32: 373—

390.

Richter, S., and Timms, B.V. 2005. A list of the recent clam shrimps

(Crustacea: Laevicaudata, Spinicaudata, Cyclestherida) of

Australia, including a description of a new species of Eocyzicus.

Records of the Australian Museum 57: 341—354.

Roeers, D.C., Rabet, N, and Weeks, S.C. 2012. Revision of the extant

genera of Limnadiidae (Branchiopoda: Spinicaudata). Journal of

Crustacean Biology 32: 827-842.

Sayce, O.A. 1903. The Phyllopoda of Australia, including descriptions

of new genera and species. Proceedings of the Royal Society of

Victoria 15: 224—261, pl. 27-36.

Spencer, W.B., and Hall. T.S. 1896. Crustacea. Pp 227—248 in: Report

on the Work of the Horn Scientific Expedition to Central Australia.

II Zoology. Dulau and Co: London.

stra$Skraba, M. (1965) Taxonomic studies on Czechoslovak

Conchostraca. I. Family Limnadiidae. Crustaceana 9: 263—273.

Timms, B.V. 2015. Eulimnadia (Branchiopoda: Spinicaudata) in

Western Australia: Three new species and a description of a

rediscovered species. Journal of Crustacean Biology 35: 441—453.

Timms, B.V. 2016a. A partial revision of the Australian Eulimnadia

Packard, 1874 (Branchiopoda: Spinicaudata: Limnadiidae). Zootaxa

4066: 351—389.

B.V. Timms

Timms, B.V. 2016b. A review of the Australian endemic clam shrimp,

Paralimnadia Sars 1896 (Crustacea: Branchiopoda: Spinicaudata).

Zootaxa 4161: 451—508.

Timms, B.V., and Lindsay, S. 2011. Morphometrics of the resting eggs

of the fairy shrimp Branchinella in Australia (Anostraca:

Thamnocephalidae). Proceedings of the Linnean Society of New

South Wales 133: 51—68.

Webb, J.A., and Bell, G.D. 1979. A new species of Limnadia

(Crustacea: Conchostraca) from the granite belt in southern

Queensland and north New South Wales. Proceedings of the

Linnean Society of New South Wales 103: 237—245.

Weeks, S.C., Sanderson, T.F., Zofkova, M., and Knott, B. 2008.

Breeding systems in the clam shrimp family Limnadiidae

(Branchiopoda, Spinicaudata). Invertebrate Biology 127: 336—349.

Wolf, E., 1911. Phyllopoda.Pp. 353-276 in: Michaelsen, W. and

Hartmeyer, R. (eds), Die Fauna Südwest-Australiens. Ergebnisse

der Hamburger südwest-australischen Forschungsriese 1905. G.

Fischer: Jena.

Memoirs of Museum Victoria 78: 65-72 (2019) Published 2019

1447-2554 (On-line)

htto://museumvictoria.com.au/about/books-and-journals/journals/memoirs-of-museum-victoria/

DOI https://doi.org/10.24199/j.mmv.2019.78.04

Bathyal and abyssal hydroids (Hydrozoa, Leptothecata) from southeastern Australia

(http://zoobank.org/urn:|sid:zoobank.org:pub: D3BA513B-E/7D6-41C5-92E4-E643ACA586E])

JEANETTE E. WATSON (http://zoobank.org/urn:lsid:zoobank.org:author:B439E439-8E2A-4F96-8417-E371A40C4B72)

Honorary Research Associate, Museum Victoria. GPO Box 666 Melbourne VIC 3001, Australia.

Email: (hydroidw @ gmail.com)

Abstract

Museum Victoria 78: 65—72.

Watson, J.E. 2019. Bathyal and abyssal hydroids (Hydrozoa, Leptothecata) from southeastern Australia. Memoirs of

A biological survey of the zone extending from ‘Tasmania (40 S) to southern Queensland (25 S) and into the Coral

Sea (23 S) was carried out along the south-eastern continental margin of Australia in 2017. Hydroids collected included

three known species (Acryptolaria angulata, Cryptolarella abyssicola and Zygophylax concinna), three new species

(Amphisbetia ramifera, Hebella macroplana and Lytocarpia parvispiralis) and two genera (Hebella and Halecium)

not identified to species. C. abyssicola was the predominant species in terms of abundance and geographical range.

Keywords

Introduction

A biological survey of the bathyal to abyssal zone along the

south-eastern continental margin of Australia was carried out in

2017 under the auspices of the Commonwealth Scientific

Industrial Research Organisation (CSIRO), the Museum of

Victoria (MV) and the Queensland Museum (QM). The objective

of the survey was to determine the biodiversity of the lower

bathyal (to 2500 m) and abyssal (to 4000 m) seafloor habitats off

south-eastern Australia and the deep-water ecosystems of seven

Commonwealth Marine Reserves. The survey was undertaken

by the Australian research vessel RV /nvestigator (Cruise V

IN2017-V03) and comprised 60 benthic stations sampled from

May to June from Tasmania (40 S) to southern Queensland (25

S) and the Coral Sea (23 S).

Field. Thecate hydroids were recovered from stations sampled

by various trawling methods at depths from 1151 m to 4173 m.

Preliminary sorting of invertebrate material was undertaken by

scientific staff on board the ship. Samples were preserved in

bulk in 95% ethanol.

Laboratory. Bulk collections were sorted in more detail at the

Queensland Museum and specimens were provided to the

author by the Queensland Museum. Representative specimens

were first stained In an aqueous solution of lignin pink and

were then dehydrated over three days by passing through three

increasingly concentrated washes in isopropanol (30%, 60%

and 100%) followed by three days of washes in xylene (30%,

60% and 100%) to harden the material. Specimens were then

permanently mounted on microslides in Malinol mountant and

heated for one week at 40° C to harden the mount.

South-eastern Australia; six bathyal to abyssal species; Cryptolarella abyssicola.

Family Lafoeidae A. Agassiz, 1865

Acryptolaria angulata (Bale, 1914)

Figure | a, b

Cryptolaria angulata Bale, 1914: 166, pl. 35, fig. 1.— Bale, 1915:

251.— Stranks 1993: 7.

Acryptolaria angulata.— Blackburn 1942: 111.— Vervoort and

Watson 2003: 41 (synonymy).

Record. QM G337451, microslide. Coll: off Fraser Island,

Queensland, 25.3253 S, 154.0683 E to 25.3513 S, 154.076 E, 2350-

2342 m, beam trawl, 11/06/2017.

Description. A small infertile colony 30 mm long with remnant

hydrorhizal stolons. Stem fascicled, polysiphonic tubes thin,

irregularly parallel, of same diameter as stolon.

Hydrothecae given off all around stem, tubular, a sharp

outward bend in hydrotheca at junction of abcauline wall with

stem; some hydrothecae widening almost imperceptibly to

margin. Margin circular, everted, rim minutely outrolled;

margin usually with many replications.

Perisarc throughout (preserved material) thin and lax.

Table 1. Measurements (in um) of Acryptolaria angulata

Distance between hydrothecae 900-1140

Hydrotheca | —]

length from abcauline bend, incl. replications 800-1020

264-272

Remarks. The strengthening buttresses reported in Acryptolaria

angulata by Vervoort and Watson (2003) are actually abcauline

intrathecal septa; these do not occur in the present specimen.

Septae probably develop to strengthen the hydrotheca in strong

water movement, not encountered by specimens in quieter deep-

water conditions. The marginal replications of the hydrothecae

considerably extends their length.

Distribution. A widespread deep-water species recorded from

the Indian Ocean, New Caledonia and rarely, New Zealand.

The deepest previous record for the species is 913 m at the

Kermadec Ridge.

Cryptolarella abyssicola (Allman, 1888)

Figure Ic—e

Cryptolaria abyssicola Allman, 1888: 40, pl. 18, fig. 2, 2a.

Cryptolarella abyssicola.— Marques et al. 2005: 711, fig. 1, Table

1, (Synonymy, discussion).

Records. QM G337422, microslide. Coll: off Freycinet, Tasmania,

41.626 S, 149.5515 E to 41.6892 S 149.5843 E, 4022—4052 m, beam

trawl, 18/05/2017. QM G337426, microslide. Coll: off Flinders Island

Tasmania, 40.386 S, 148.928 E to 40.383 S 148.951 E, 932-1151 m,

beam trawl, 20/05/2017. QM G337427, microslide. Coll: off Flinders

Island Tasmania, 40.464 S. 149.3967 E to 40.464 S. 149.4255 E. 4114—

4139 m, beam trawl, 20/05/2017. QM G337438, microslide. Coll: off

Jervis Bay, New South Wales, 35.333 S, 151.258 E to 35.332 S, 151.214

E, 2650-2636 m, beam trawl, 29/05/2017. QM G337439, microslide.

Coll: off Newcastle, New South Wales, 33.435 S, 152.702 E to 33.435

S. 152.665 E, 4280-4173 m, beam trawl, 30/05/2017. QM G337443,

microslide. Coll: off central New South Wales coast, 30.099 S,

153.596 E to 30.128 S, 153.571 E, 1257-1194 m, beam trawl, 5/06/2017.

QM G337448, microslide. Coll: off Byron Bay, New South Wales,

28.0544 S, 154.083 E to 28.097 S, 154.081 E, 999-1013 m, beam

trawl, 9/06/2017. QM G337452, microslide. Coll: Coral Sea,

Queensland, 23.587 S, 154.194 E to 23.617 S, 154.1947 E, 1013—1093

m, beam trawl, 13/06/2017. QM G337421, Coll: off Freycinet

Tasmania, 41.7305 S, 140.1197 E, to 41.7913 S. 149.1558 E, 2751— 2820

m, beam trawl, 18/05/2017. QM G337431, Coll: Bass Strait, 39.552 S,

149.553 E, to 39.496 S. 149.598 E, 4197-4133 m, beam trawl,

23/05/2017. QM G337435, Coll: off Bermagui, New South Wales,

36.418 S, 150.8 E, 3980 m, beam trawl, 26/05/2017. QM G337440.

Coll: off Newcastle, New South Wales, 32.985 S. 152.952 E, to 33.015

S, 152.913 E, 2704—2902 m, beam trawl, 31/05/2017.

Description. Small lax colonies to several centimetres high;

some colonies fertile.

Colonies — fasciculated basally, ultimate branches

monosiphonic. Hydrothecae numerous, tubular, given off from

around branches in fasciculated sections, more or less subalternate

on monosiphonic branches. Hydrotheca adherent to branch for

more than half of length, abcauline wall variably concave,

adcauline wall convex, curving smoothly outwards, free wall

shorter than adnate wall. Hydrotheca narrowing basally but

without floor. Margin circular, not everted, without replications.

Gonothecae large, sausage-shaped, adnate to outer stem

tubes, body narrow proximally, becoming tubular, abcauline

wall minutely wrinkled, orifice wide, circular, upturned to

varying degrees.

Perisarc soft and thin throughout, colour (preserved

material) grey.

J.E. Watson

Table 2. Measurements (in um) of Cryptolarella abyssicola

Hydrtheca ——— |

800-1800

136-192

Gonotheca

1700-2200

320-600

width of orifice 336—404

Remarks. Without a discernible floor, the length of the

hydrothecae is highly variable, making it difficult to provide a

precise estimate of length.

Cryptolarella abyssicola was fist described from

Challenger Station 160 from a depth of 4755 m south of

Australia (42°42 S, 134°10 E) (Allman 1888). In his description

Allman commented on the “vast depth" from which the

Species came, and the height of the colony, about 2 inches

(=4.5 cm). C. abyssicola has since been recorded world-wide

from Sierra Leone, the Azores, Tierra del Fuego, Peru and the

Antarctic (Marques et al. 2005) and from abyssal depths under

several specific names. The present specimens conform well

to the redescription and dimensions of the holotype given by

Marques et al. (2005). This survey indicates that C. abyssicola

is a dominant abyssal species around southern Australia.

Distribution. Australia (type locality), Sierra Leone, Azores,

Tierra del Fuego, Peru, Antarctic.

Family Hebellidae Fraser, 1912

Hebella macroplana sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:CC386770-19A6-

4ACE-SF5C-91 F9963120B8

Figure If

Record. QM G337336. Holotype, one microslide. Coll: eastern

Bass Strait, Victoria, 39.552 S, 149.553 E to 39.496 S, 149.598 E,

4197—4133 m, beam trawl, 23/05/2017.

Description. Five stolonal hydrothecae creeping on stem of an

antipatharian. Colony entangled with remnants of an

unidentifiable anthoathecate hydroid.

Hydrorhizal stolon smooth, thin. Pedicel of hydrotheca

short, smooth, curved, passing upwards to diaphragm.

Hydrothecae tubular, very large, slightly asymmetrical or

symmetrical, adcauline side convex to above diaphragm, walls

smooth to margin. Diaphragm a barely discernible transverse

or Slightly oblique ring. Margin circular, slightly everted, one

hydrotheca with two widely separated marginal replications.

Perisarc very thin, smooth.

Table 3. Measurements (in um) of Hebella macroplana

Hydrorhizal stolon, width 60—64

Hydrotheca ooo o

length, diaphragm to margin 740-1920

520-584

160-192

length of pedicel 120-184

Bathyal and abyssal hydroids (Hydrozoa, Leptothecata) from southeastern Australia 67

Remarks. One hydrotheca contains approximately 10 degenerated

tentacles indicating that the structure it is not an empty gonotheca.

The unsegmented hydrothecal pedicel is very short, and in some

hydrothecae it 1s slightly curved to accommodate the asymmetry

of the hydrotheca. The diaphragm varies from a membranous to

a thin perisarcal ring.

Three genera considered were Hebella, Halisiphonia and

Scandia; the latter two genera were rejected because they have

long hydrothecal pedicels.

Hebella macroplana most resembles the Antarctic species

Hebella plana Ritchie, 1907; however, the hydrotheca of H. plana

is much smaller and has a longer and straighter pedicel [see

Totton (1930); Briggs (1938); Boero et al. (1997)]. Although the

present material is meagre and without gonothecae, the hydrotheca

is extremely large and much bigger than any known species.

Etymology. The name alludes to the large hydrotheca compared

with that of H. plana.

Hebella sp.

Figure lg

Record. QM G337403, one microslide. Coll: off Bermagui, New

South Wales, 36.418 S, 150.8 E, 3980 m, beam trawl, 26/5/2017.

Description. Three damaged stolonal hydrothecae creeping on

stem of Zygophylax concinna. Stolon fragmented, very thin,

tubular. Hydrothecal pedicel long, unsegmented. Hydrotheca

long, expanding from a narrow conical base to diaphragm then

eradually becoming tubular. Diaphragm a distinct perisarcal

ring. Margin circular, rims fragmented.

Perisarc thin and fragile.

Table 4. Measurements (in um) of Hebella sp.

Hydrorhizal stolon width

Hvdrotheca

length, diaphragm to margin 1400

diameter of diaphragm

length of pedicel 240—320

Remarks. Although there are no hydrothecae with intact margins,

sufficient remains to provide an estimate of marginal diameter.

Although morphologically similar to Hebella ritchiei Vervoort,

1966 (2Lafoea tenellula Ritchie, 1911) from coastal New South

Wales, the hydrothecae of the present material are much larger.

The material is inadequate to ascribe it to a new species.

Family Zygophylactidae Quelch, 1885

Zygophylax concinna (Ritchie, 1911)

Figure 2 a, b

Zygophylax concinna Ritchie, 1911: 823, pl. 88, figs 3, 4.

Record. QM G337986, one microslide. Coll: off Bermagui, New

South Wales, 36.418 S, 150.8 E, 3980 m, beam trawl, 26/5/2017. QM

G337446, one microslide. Coll: off Byron Bay, New South Wales,

28.371 S, 154.6487 E to 28.3875 S, 154.617 E, 3825-3754 m, beam

trawl, 9/06/2017.

Description. A broken, heavily fascicled and twisted stem

originally about 15 mm long and one stem fragment 8 mm long

with one undamaged hydrotheca.

Polysiphonic stem tubes parallel, giving off flaccid

monosiphonic branches. Branch internodes long, thin,

cylindrical, nodes transverse, narrow, a tumescence above and

below node; one or two alternate hydrothecae on internode.

Hydrotheca about halfway along internode, inserted on an

inflated apophysis, distal node of apophysis transverse.

Pedicel of hydrotheca of one long, rarely two or three

cylindrical segments expanding distally to diaphragm.

Diaphragm a thin perisarcal ring situated high in hydrotheca;

walls of hydrotheca above diaphragm more or less cylindrical or

expanding a little to margin. Margin circular, transverse to

hydrothecal axis, some slightly inclined, rim everted, often with

several strong replications.

Perisarc of polysiphonic tubes thick, hydrocladia thinner,

hydrothecae fragile, mostly broken.

Table 5. Measurements (in um) of Zygophylax concinna

700-780

40-52

adcauline length of apophysis 40—52

leneth, diaphragm to marei

diameter of margin 136-

Remarks. Zygophylax concinna was first recorded from a fine

sandy bottom at a depth of 100 m off Sydney, New South Wales

(Ritchie 1911). Ritchie’s small colony (Ritchie 1911, pl. 88, fig. 3)

was probably young. The present specimens are probably parts of

much larger complexly branched colonies; otherwise the material

generally conforms to Ritchie’s description of Z. concinna.

Distribution. New South Wales, Australia. This is the second

record of the species.

Family Haleciidae Hincks, 1868

Halecium sp.

Record. QM G337429, one microslide. Coll: near Flinders Island

Tasmania, 39.462 S, 149.276 E to 39.465 S, 149.242 E, 2760—2692 m,

beam trawl, 22/05/2017.

Comment. A large lax tangled, fascicled colony with two

hydrothecae. Specimen too badly damaged for description.

Family Sertulariidae Lamouroux, 1812

Amphisbetia ramifera sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:0OBB245B8-E3F4-

421C-928A-CA2CIFBS4A30

Figure2c,d

68 J.E. Watson

0.5mm

1.0mm

Figure 1. a—g. a, b, Acryptolaria angulata. a, distal monosiphonic stem. b, hydrotheca. c—e, Cryptolarella abyssicola. c, monosiphonic branch

with subalternate hydrothecae. d, gonotheca with wrinkled abcauline wall and upturned margin. e, gonotheca with oblique margin. f, Hebella

macroplana sp. nov., stolonal hydrotheca. g, Hebella sp., stolonal colony and pedicellate hydrotheca.

Bathyal and abyssal hydroids (Hydrozoa, Leptothecata) from southeastern Australia 69

Record. QM G337425. Holotype, one microslide. Coll: near

Flinders Island, eastern Bass Strait, Victoria, 40.386 S. 148.928 E to

40.383 S, 148.951 E, 932-1151 m, beam trawl, 21/05/2017.

Description. A branched stem fragment 5 mm long with four

alternate branches on each side; stem and branches monosiphonic.

Two tubular subopposite hydrothecae on stem internode,

adcauline walls separated, nodes strong, transverse, deeply

indented. Apophysis long, narrowing distally to transverse node,

an axillar hydrotheca pointing along hydrocladium.

First branch internode long, athecate, expanding slightly to a

strong opposed V-shaped joint. Branch internodes same as stem,

nodes may be absent but where present transverse to slightly

oblique, strongly contracted. Hydrothecae opposite, tubular, base

of one hydrotheca usually slightly downwardly displaced with

respect to that opposite. Lower adcauline wall of each pair adnate,

wall straight to weakly convex basally, the convexity increasing

towards free wall; free wall weakly convex or concave to margin.

Abcauline wall smoothly concave, some walls slightly bulging

Just above base. Floor transverse to internode, a small downward

septum from adnate wall passing into internode. Margin deep

saddle-shaped, flanked by a pair of long, sharp lateral cusps.

Perisarc thick, colour (preserved material) shining

golden brown.

Table 6. Measurements (in um) of Amphisbetia ramifera

distance between marginal cusps

width of floor 112-120

Remarks. The fragment 1s probably an apical branch of a larger

colony. The hydrothecae closely resemble Amphisbetia minima

(Thompson, 1879), a common shallow water species in Australia

and New Zealand. A. minima invariably has short unbranched

stems unlike the branching habit of A. ramifera. In colony size

and branching habit, A. ramifera resembles Amphisbetia

maplestonei (Bale, 1884) but in contrast to A. maplestonei the

hydrocladial hydrothecae of A. ramifera are in contact with each

other, do not have an abcauline intrathecal septum and the

marginal cusps are much more prominent. No other Australian

species of Amphisbetia has the smoothly outward-curved

hydrothecae and such prominent marginal cusps as A. ramifera.

Family Aglaopheniidae Marktanner-Turneretscher, 1890

Lytocarpia parvispiralis sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act: DD8BF2A6-A3A6-

4156-AB1E-1F03B1D2C408

Figure 2 e, f

Record. QM G337453. Holotype, one microslide. Coll: Coral Sea,

Queensland, 23.7503 S, 154.5718 E to 23.7739 S, 154.5464 E 2093-

2156 m, Brenke epibenthic sled, 14/06/2017. QM G337985. Paratype,

one microslide from holotype colony.

Description. Infertile colony 90 mm long, broken in two.

Hydrorhiza comprising a group of smooth tubular stolons

coalescing to form parallel polysiphonic tubes of lower stem.

Colony with three primary branches on upper stem section.

Branches monosiphonic, cylindrical, with a row of up to 15

nematothecae below first hydrocladium.

Hydrocladia with many hydrothecae. Hydrocladium long,

lax, apophysis large, distal node oblique, two nematothecae in

a line below hydrocladium and one beside axil. Hydrocladial

internode narrow, node distinct, slightly oblique, two partial

septa passing into internode from base of hydrotheca, one

below hydranth, the other about halfway along internode.

Hydrotheca occupying much of internode, slipper-shaped,

abcauline and adcauline hydrothecal walls gently convex,

adcauline wall fully adnate to internode. Margin slightly

oblique to internode axis, anterior cusp tongue-shaped,

followed by two moderately pointed cusps then three indefinite

low cusps, interspaces between very shallow.

Median nematotheca digitate, very short, almost entirely

adnate to hydrotheca, terminal orifice small, pointing upwards,

open down to hydrotheca. Lateral nematotheca tubular, just

reaching margin of hydrotheca, orifice sinusoidal down to

internode. Cauline nematothecae the same as laterals. Hydranth

with approximately 10 tentacles, hypostome mound-shaped.

Perisarc moderately thick throughout colony.

Table 7. Measurements (in um) of Lytocarpia parvispiralis

Branch — ^ 0 0 |

leneth of internode 568—648

144—152

Hydrocladium

leneth of internode 608—032

width of node

Hydrthea o 0 |

Remarks. Unfortunately the preserved colony was not examined

in detail prior to mounting. Its position on the microslide now

prevents determination of whether the hydrocladia were

spirally arranged. Its close resemblance to Lytocarpia spiralis

(Totton, 1930) suggests that the hydrocladia may be spirally

arranged. The hydrothecae are much smaller and the marginal

cusps less prominent than in L. spiralis (see Vervoort and

Watson 2003). Lytocarpia parvispiralis 1s clearly an abyssal

congener of L. spiralis, a species common around New Zealand

to depths of 1126 m.

Etymology. The name alludes to the smaller size of

L. parvispiralis compared to its close congener L. spiralis.

70 J.E. Watson

Figure 2. a-f. a, b, Zygophylax concinna. a, branch internodes with hydrothecae. b, hydrotheca. c, d, Amphisbetia ramifera sp. nov. c, stem with

subopposite hydrothecae and axillar hydrotheca. d, branch with opposite hydrothecae. e, f. Lytocarpia parvispiralis sp. nov. e, holotype colony.

f, hydrothecae, lateral view.

Discussion Zygophylax concinna), three newly described species (Hebella

Eight species were recovered from depths of 1151-4173 m. OTOP lana, Amphisbetia ramifera and Lytocarpia

Species and their general localities are listed for comparison Parvispiralis) and two species (Hebella sp. and Halecium sp.)

in Table 8. The list includes three previously known species which were too inadequate or in too poor a condition for

(Acryptolaria angulata, Cryptolarella abyssicola and identification.

Bathyal and abyssal hydroids (Hydrozoa, Leptothecata) from southeastern Australia 11

Table 8. Species and their general locations.

Hebellasp FF Bermagui, New SouthWales —

Zygophylax concinna (Ritchie, 191) |2 OFF Bermagui, New SouthWales —

Nei 2 — —

Halecium sp

ooo Off Flinders iona. Tasmania

Amphisbetia ramifera sp. nov. E Off Flinders Island, Tasmania

Lytocarpia parvispiralis sp. nov.

Abyssal hydroids first recorded from the Australian region

were Cryptolarella abyssicola and Halisiphponia megalotheca

from Challenger Station 160 from a depth of 4755 m south of

Australia (Allman 1888). There are no other published reports

of the abyssal hydroid fauna of Australia. Previous moderately

deep-water surveys from which hydroids have been reported

are from the Great Australian Bight carried out by F.I.S.

Endeavour (Bale 1914a; 1914b; 1915), the Thetis expedition

off the coast of New South Wales (Ritchie 1911) and a recent

survey of a marine protected area in the eastern Great

Australian Bight (Watson 2018). None of these surveys were to

depths greater than 100 m.

Cryptolarella abyssicola was by far the most abundant

species with 12 records. The species has previously been

recorded under various names from Sierra Leone, the Azores,

Tierra del Fuego, Peru and the Antarctic (see Marques et al.

2005). It is considered a wide-ranging “true abyssal hydroid"

(Vervoort 1985) recorded from 4600 m in the Southern Ocean

(Allman 1888), from 2470 m in the Kermadec Trench ( Vervoort

1966), from 6328 m from Peru (Vervoort 1972), and 4578 m

from the mid-Atlantic Ridge (Calder and Vervoort 1998). The

Australian range of C. abyssicola 1s now extended along the

south-eastern Australian coast from cool temperate Tasmania

in the south to the subtropical Coral Sea in the north.

Acryptolaria angulata is known from deep water in the

Indian Ocean, Indo-west Pacific, New Caledonia and New

Zealand (Vervoort and Watson 2003). Zygophylax concinna

has been recorded only once previously from off Sydney

(Ritchie 1911); the two new records extend its range south

along the New South Wales coast and its depth range from 100

m to 3754 m.

Two newly described species, Hebella macroplana and

Amphisbetia ramifera, were recovered from depths of 4133 m

and 4131 m respectively in adjacent localities in eastern Bass

Strait, Victoria. The record of A. ramifera at such depth is worthy

of comment: branched, golden-brown species of Amphisbetia

are a moderately common component of the shallow coastal

water hydroid fauna of southern Australia (Watson 1973; pers.

obs.). The present specimen may be a floating fragment from a

shallow water colony entrained in the trawl.

The third newly described species, Lytocarpia parvispiralis

from the Coral Sea, may, when more material 1s found, prove

to be a diminutive subspecies of Lytocarpia spiralis, acommon

deep-water species from around New Zealand (Vervoort and

Watson 2003).

There were surprisingly few species and few locality

records in the collection considering the extensive latitudinal

range of the survey. This may be an artefact of decrease in

hydroid diversity with depth, the sampling gear or sampling of

predominantly sedimentary substrates.

Acknowledgements

I thank Dr Merrick Ekins of the Queensland Museum and Dr

Tim O’Hara of Museum Victoria for the opportunity to

examine the hydroid collection. I also thank Josh Hatton of the

Queensland Museum for sorting of hydroids and the scientific

staff and crew of R.V. /nvestigator for their work during the

Investigator IN2017-V B03 Cruise.

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Memoirs of Museum Victoria 78: 73-146 (2019) Published 2019

1447-2554 (On-line)

httos://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:|sid:zoobank.org:pub:263C1363-0A DA-4972-9224-AC690A 1 FD238)

Gary C. B. Poors! (http://zoobank.org/urn:lsid:zoobank.org:author:c004d784-e842-42b3-bfd3-317d35918975),

PETER C. DWoRSCHAK ° (http://zoobank.org/urn:|sid:zoobank.org:author:4BCD9429-46A F-4BDA-BE4B-439EE6A DC657),

RAFAEL ROBLES > ^? (http://zoobank.org/urn:lsid:zoobank.org:author:7 F54A0F4-892F-407A-A ED4-4E65EA8B9360)

FERNANDO L. MANTELATTO * (http://zoobank.org/urn:|sid:zoobank.org:author:882CB189-DEFC-4CFA-8E04-2923C89B1B25) and

DARRYL L. FELDER ` (http://zoobank.org/urn:|sid:zoobank.org:author:7A4A 3683-FCED-431A-A685-1C68E7CFD8AB)

Abstract

! 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

* Department of Biology, University of Louisiana— Lafayette, Lafayette, LA 70504, USA dlf4517@louisiana.edu

* Laboratory of Bioecology and Crustacean Systematics (LBSC), Department of Biology, Faculty of Philosophy, Science

and Letters at Ribeirao Preto (FFCLRP), University of Sao Paulo (USP), Av. Bandeirantes 3900, 14040-901, Ribeirão

Preto, Sao Paulo, Brazil flmantel@usp.br

` Facultad de Ciencias Químico Biológicas, Universidad Autónoma de Campeche, Campus V. Predio s/n por Avenida Ing.

Humberto Lanz Cardenas y Fracc. Ecológico Ambiental Siglo XXIII, Colonia Ex Hacienda Kala, San Francisco de

Campeche, Camp., 24085, México

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: 13—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—Agaballianassa 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.

/4 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 Rayllianassa Komai and Tachikawa, 2008 ..................... 98

TERCER TROT iL ns Aretes ceva soe sssi CREE 75 nya nasa Serer a D 18 Ta kuu a sm i uki i"

SCallasis Bates TBR aae ebbe ekino ieaiaia 99

IRS CCR paso A SNL dede dede de odo eM ES EES edet 75 Spinicallianassa POT. TOV, seraya rra Re Er Rh der 09

Infraorder Axiidea de Saint Laurent, [0:70.79 ARUARURR UE UE 76 lastrypaea ZCM: DION ceeececeeeesru reve viene sieue sie webs tals a n o daa a te 100

Irypded- Danas 162 ted uuu u uu ren eorr 100

Key to families of Axiidea ......... ccc ccccccecccseecceeeeeeeeeeeeeneeeees 71

Callianopsidae Manning and Felder, 1991 ....................... 101

Anacalliacidae Manning and Felder, 199] ......................... 8 |

Anacalliax de Saint Laurent, 1O73 sese g2 Key to genera of Callianopsidae ...................................... 102

| | Bathycalliax Sakai and Türkay, 1999 ............................ 102

Callianassidae Dana. 1852 ................................................. 82 Callianopsis de Saint Laurent, 1973 ......... e 102

Key to genera of Callianassidae ................... eene 87 Vulcanocalliax Dworschak and Cunha, 2007 ............... 102

Aqaballianassa gen. nov. ................................................. 90 Callichiridae Manning and Felder, 1991 _......... aaa. 102

Arendllidnassd een. OV. [uu eeu as sidia RR a ca 9]

Biffarius Manning and Felder, 1991 ................. sess 9] Key to genera of Callichiridae ......................................... 106

Calliamássa-Leacbh IS Tl n aeter tetti 9] Audacallichirus gen. NOV, 2... euet tenens 107

Cavyialtiamassa Sex NOV ¿L eet er en U SH Re REI 02 Balsscallichirus Sakai, 2011 ................. eee 108

Cheramoides Sakai, 2011 ................................................ 92 Calliapagurops de Saint Laurent, 1975 ......................... 108

Cheramus Bate, 1888 .......... sse 93 Callichirus Stimpson, 1866 .................. sss 109

Coriollianassa gen. NOV. ..................... eene 93 Corallianassa Manning, 1987 ....................................... 109

Filhollianassa gen. nov. ............................ nes 93 Giypturotdes Sakal, 201 T rrenen en errierta 110

Fragillianassa gen. nov... 94 Glypturus Stimpson, 1866 ............ sse 110

Gilvossius Mannine and Felder, 1992 ............................. 04 Grynaminna Poore, POO ES rutuna u au ee et 110

Jocullianassa gen. NOV. ...cccccccseccccsesecesssesesssessssssseseseseseeens 95 Karumballichirus gen. nov. ............................................ 111

Lipkecallianassa Sakal 2002 Lr nee rrr rA 05 Kraussillichirus BOIL HOW. Leacesenettittrtt tttm trennen tee 111

Necallianassa Heard and Manning, 1998 ....................... 95 Laticallichirus Komai, Yokooka, Henmi and Itani, 2019 .. 111

Neotrypaea Manning and Felder, 1991 ........................... 96 Lepidophthalmus Holmes, 1904 .................................... 112

Notiax Manning and Felder, 1991 ................................... 96 Michaelcallianassa Sakai, 2002 ................................... 113

Paratrypaea Komali and Tachikawa, 2008 ...................... 96 Mocallichirus gen; DOV, icieescee cce deveined eese ees vue rop tenus 113

Poti Rodrigues and Manning, 1992 ................................. 97 Mucrollichirus gen. nov. ................................................ 113

Praedatrypaea gen. nov. .................................................. 97 Neocallichirus Sakai, 1988 ............................................. 114

Pugnatrypaea gen. nov. ................................................... 97 Thailandcallichirus Sakai, 2011 ..................................... 114

Callianassidae and related families

Ctenochelidae Manning and Felder, 199] ........................ 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, 199] ......................... 122

Key to genera of Eucalliacidae ........................................ 125

Andamancalliax Sakai, 2011 ......................................... 126

Calliax de Saint Laurent, 1973 ...................................... 126

Calliaxina Ngoc-Ho, 2003 ............................................ 126

Eucalliax Manning and Felder, 1991 ............................. 127

Eucalliaxiopsis Sakai, 2011 ................ eere 127

Paraglypturus Türkay and Sakai, 1995 ......................... 128

Pseudocalliax Sakai, 2011 ............................................. 128

Paracalliacidae Sakai, 2005 ............................................. 128

Paracalliax de Saint Laurent, 1979 ............................... 129

Acknowledgements .......................................................... 129

RESTENE S A. us donk oed tooth took l ama A ede es e a tees 130

Introduction

The earliest published descriptions of callianassids were at

the end of the 18th century and the beginning of the 19th

century (fig. l). The genus name Callianassa Leach, 1814,

was introduced shortly thereafter. The number of available

Species names now totals 305, with 262 accepted by WoR MS

(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 1s 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) 1n 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 (20152)

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 | 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 1n 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 (NHM W); 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 (NT MAG); 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.

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 Agaballianassa. 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.— D worschak 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, 1905;

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 (20053) included in

Callianassoidea, families Callianassidae; Axiidae Huxley,

1879; Callianideidae; Ctenochelidae and Gourretiidae Sakai,

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 1n the phylograms of both Tsang et

al. (2008) and Robles et al. (2009) led to our expanded study

(Robles et al., 1n 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. 40); maxilla scaphognathite

with long setae extending from posterior lobe into

branchial chamber (fig. 3a) Axiidae

3. Maxilla scaphognathıte with long setae extending from

posterior lobe into branchial chamber (fig. 3a); linea

thalassınıca complete, partial or absent 4

— Maxilla scaphognathıte 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 | (fig.

21); eyestalk flat, contiguous (fig. 21); chelipeds flattened,

asymmetrical (fig. 3J) Callianideidae

— Posterior margin of carapace with lateral lobes interacting

with anterolateral lobes on pleomere | (fig. 2g); eyestalks

cylindrical even if continuous (fig. 2g); chelipeds

cylindrical, symmetrical (fig. 31) Micheleidae

5. Pleomere 1 with dorsal pair of lobes interacting with

posterior margin of carapace (fig. 21); female pleopod | 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)

18 G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder

eet

—h^

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: 1, Paracalliacidae; j, Callianassidae. Anterior carapace, branchiostegite,

epistome, basal antenna and eyestalk: k, Eucalliacidae, Calliaxina; 1, Callichiridae, Callichirus; m, Callichiridae, Lepidophthalmus; n,

Callianassidae, Coriollianassa; o, Callianassidae, Biffarius.

Original illustrations: 1, Paracalliax bollorei, MNHN Thl517; 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 6. Maxilliped 3 dactylus ovate, distally truncate, with dense

interacting with posterior margin of carapace (fig. 2j); distal field of setae (fig. 3e... e e etm 7

female pleopod 1 with reduced ramus (fig. 4a); pleopod 2

in both sexes (figs 5d—g) smaller than pleopods 3-5 (fig.

50); epipods absent above maxilliped 3 to pereopod 4 setae (fig. 5f), or densely setose on upper or lower margin

(exception, 2 species of Callianopsidae) UU. 6 (HOSSSE INR, ar .uu E E A E AE EN EO O dore ll 8

— Maxilliped 3 dactylus linear, with scattered groups of

Callianassidae and related familles 19

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: 1, Micheleidae, Tethisea; j, Callianideidae, Callianidea; k, Ctenochelidae, Ctenocheles; |, 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,1, Callichiridae;

J, Callianassidae. Male pleopod 2: k, Ctenochelidae, Ctenocheles; |, 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.

Callianassidae and related families

10.

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

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

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. 31), 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 parallel-

sided, with rounded posterolateral corners (fig. Ar), 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 | 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

Anterior branchiostegal lobe well-produced anteriorly

beyond junction with oblique branchiostegal ridge with

which it articulates by means of a virtual condyle (figs 21,

m); male pleopod | article 2 usually of 2 articles, article 2

simple or apically notched (figs 4h, 1); 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, 0), 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 | 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 | 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

G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder

d e f |

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 1n 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, 1f 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: 1l01— 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

Pu TFÍE LA

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| N

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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 1, Praedatrypaea; J, Pugnatrypaea; k, |,

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, NM V 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

eroove 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 I epipod truncate, without

anterior lobe. Maxilliped 3 dactylus slender, digitiform, with

setae irregularly spaced along all margins. Cheliped merus

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;1, 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: 1, Cheramoides;

m, Neotrypaea.

Original illustrations: c, Coriollianassa MOZ-33, MNHN-IU-2008-10314; e, Biffarius biformis, NM V 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; 1, Scallasis; J, Spinicallianassa; k, Tastrypaea. Minor cheliped: |, Jocullianassa; m,

Rudisullianassa.

Callianassidae and related families 87

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: t, Caviallianassa; g, Poti.

Original illustration: e, f, Caviallianassa FP-11, UF 29204.

Key to genera of Callianassidae

Maxilliped 3 merus distally oblique with obtuse angle

between distal and lower margins (fig. 7h); pereopod 3

l. Antennular peduncle about 4 times the width of both 4 L1 a slichtl lead}

eyestalks, twice as long as antennal peduncle, with 2 i a aa QUSS aden sae ae eed ET Sanne 19

dense rows of adjacent long setae on lower surface (fig. narrow, distinctly rounded proximal lobe; uropodal

6n); maxilliped 3 merus produced as massive triangular exopod distal margin clearly differentiated from anterior

lobe alongside palp (fig. 7k) UU aaa Trypaea margin, anterodistal corner right-angled; telson tapering

nih — TM —— over distal third to pair of posterior lobes separated by

— ntennular peduncle at mos imes the width of bo d teh (Ge. 11 p ;

bvestalks. with-ar- most bands st scattered Tone Belas an eep notch (fig. 1160) e e ee ugnatrypaea

lower surface (fig. 6a); maxilliped 5 merus distally convex Antennular peduncle exceeded by all or most of antennal

AEOS Y. ae eee TWO EE IEEE RAE AERE RO E oe eee ee 2 peduncular article 5 (figs 6d, i) aaa 6

2. Small triangular sclerite present on branchiostegite at Antennular peduncle as long as or longer than antennal

anterior end of linea thalassinica (fig. 6a) ..... Agaballianassa peduncle (figs-6b,-6,:& D)... eere 10

— Without triangular sclerite on branchiostegite at anterior Maxilliped 3 merus with spine on distal free margin (fig.

end of linea thalassinica (figs 6f, k, h) nnn 3 7e); proximolateral lobe of telson prominent, defined

3. Telson tapering, posterior margin with medial spine in posteriorly by clear unchitinised region (fig. Ln) oo...

deep notch (figs 11g, 0) oo a aaa Be ad BAR Ra scale Mata Si: Praedatrypaea

— Telson posterior margin convex, truncate or at most with Maxilliped 3 merus without spine on distal free margin;

shallow medial concavity, sometimes with small medial proximolateral lobe of telson prominent or not, indefinitely

SUITE = Rm EXER R3 BOYES RIAL ORI RN ENTE ED X M ENTER 5 defined posteriorly |... 7

4. Maxilliped 3 merus almost rectangular, distally truncate Major cheliped carpus articulating by means of a short

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

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 Ó

G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder

ae AN

1 q N b... /

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; 1, Neotrypaea californiensis; j, N.

petalura; k, Notiax; 1, Poti; m, Paratrypaea; n, Praedatrypaea; 0, 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; 1, Neotrypaea californiensis, NM V J20600; 1, N. petalura, NM V J59981; k, Notiax brachyophthalma, NMV J58880; m,

Paratrypaea maldivensis, UF 2878].

Callianassidae and related families

10.

li.

12.

13.

14.

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

Rostrum acute, as long as eyestalk (fig. 61); cornea diffuse;

maxilliped 3 longer than wide at ischium-merus suture

(fig. 7c); major and minor chelipeds similar, with saw-

tooth 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 ischium-

merus suture than long (fig. 7m); major and minor

chelipeds dissimilar, merus unarmed or with small teeth

(figs 8h, m); pleomere without sublateral ventral sharp

Rudisullianassa

Rostrum obsolete or triangular, not reaching cornea (figs

6b-f, h, 1, m) 15

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

Pereopod 3 propodus oval or with straight lower margin

(Figs 10b, c); major cheliped merus smooth (fig. 81), or

with spine (fig. 9h), or with blade (figs 91, J) 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. 81) 13

Maxilliped 3 distally convex, extending beyond

articulation of carpus (fig. 7j), or distally oblique with

obtuse angle between distal and lower margins (fig. 71);

major cheliped merus with prominent hooked blade (figs

91, k) 14

Uropodal exopod at least twice as long as wide (fig. 11d);

pleopod 2 absent in male Cheramoides

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. 9);

pereopod 3 propodus subrectangular (fig. 100) a...

Spinicallianassa

15.

16.

17.

18.

19.

Maxilliped 3 merus longer than wide at ischium-merus

suture, distally oblique with obtuse angle between distal

and lower margins (fig. 71); male major cheliped merus

with | or 2 proximal similar teeth and distal denticles on

lower margin, propodus narrow, shorter than carpus (fig.

91); pereopod 3 propodus oval (fig. 10b) Scallasis

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

Major cheliped with dorsodistolateral propodus and

lateral dactylus sparsely setose; uropodal endopod

sometimes with facial spiniform setae but not as obvious

transverse row 16

Anterior carapace strongly depressed by about 45° in

lateral view, rostrum blunt (figs 6e, f) Filhollianassa

Anterior carapace moderately or not depressed in lateral

view (fig. 6h) 17

Antennular peduncle exceeding antennal peduncle by

about half length of article 3 (fig. 61); major and minor

chelipeds similar, merus without prominent hook or spine

on lower margin, carpus and propodus ovoid in cross-

section, 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) 0...

Rayllianassa

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. 60) 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

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 | and 2 present in

male (figs 10e, f); uropodal endopod with at least ] facial

spiniform seta (figs 11b, c) Caviallianassa

20.

21.

22.

24.

25.

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.

Sh) 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 Sf, k, 9c) 2]

Uropodal endopod anterior margin with distal spine (fig.

I 1h); telson usually with 1 or 2 pairs of lateral spines ,

Necallianassa

Uropodal endopod anterior margin and telson lateral

margin without spines (figs I1f, 1, J) 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

1 li—k, p) 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 Sf, 9b, d); uropodal exopod distal and posterior

margins densely setose, without blade-like setae 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

Maxilliped 3 merus distally convex, extending only

slightly beyond articulation of carpus (fig. 71); 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 111, J)...

Neotrypaea

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.

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 5 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-B393-

48F5-B69F-83A FDSFFA9D7

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 l 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,

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 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; 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 1s 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. agabaensis have male and female gonopores,

but pleopods | 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:|sid:zoobank.org:act:/A9C78DE-06CA-

4407-BA4A-0A2BA6258BDE

Type species. Callianassa arenosa Poore, 1975, by present

designation.

Diagnosis. Rostrum obsolete or obtusely triangular, flat, not

reaching cornea. Pleomere | 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).

Irypaea.—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 | 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, 5. 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 1s known from south-

eastern Australia, a somewhat enigmatic generic distribution.

Callianassa Leach, 1814

Callianassa Leach, 1814: 386, 400.—Sakai, 2011: 359

(comprehensive synonymy).

Type species. Cancer (Astacus) subterraneus Montagu, 1808,

by original designation and monotypy.

Diagnosis. Rostrum obsolete or obtusely triangular, flat, not

reaching cornea. Pleomere | 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, 1s a synonym of Gilvossius and

is discussed below.

Caviallianassa gen. nov.

http://zoobank.org/urn:|sid:zoobank.org:act:9819F120-10C6-

496D-80E2-BDDAD3FBA E07

Type species. Cheramus cavifrons Komai and Fujiwara, 2012,

by present designation.

Diagnosis. Rostrum obsolete or obtusely triangular, flat, not

reaching cornea. Pleomere | 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 ischium-

merus 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

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 right-

angled. Telson anterolateral lobe obsolete, undefined.

Remarks. Cheramus 1s 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 Gn

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-426A-

4110-9945-31AS8BIF5831B

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 5 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-ABFS5-

46ED-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 | 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

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 7rypaea

australiensis, also 1n 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: CFA 11DDC-A9CS5-

4ACC-AC90-E2C6A5456E82

Type species. Callianassa fragilis Biffar, 1970, by present

designation.

Diagnosis. Rostrum obsolete or obtusely triangular, flat, not

reaching cornea. Pleomere | 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, 1s

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 1n 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 | tergite fused, divided into 2

sections by transverse step. Antennular peduncle length about

2.53 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 1s 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 Negoc-Ho (2003)

compared Gilvossius and Pestarella respectively with

Callianassa, but this latter genus 1s 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

Callianassidae and related families

synonym of both Pestarella Ngoc-Ho, 2003, and Gilvossius

Mannine and Felder, 1992. However, Gebios has never been

used as a generic name. Neoc-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: 1 AC3D163-8895-

45EF-8EC6-E87B435B9FDF

Type species. Callianassa joculatrix De Man, 1905, by present

designation.

Diagnosis. Rostrum acute, anteriorly directed, as long as

eyestalks. Pleomere | 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 |.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 1s 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. (1n press).

Necallianassa Heard and Manning, 1998

Necallianassa Heard and Manning, 1998: 883—884.

Irypaea.—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 | 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 truncate-

convex, at right angles to straight anterior margin, anterior

margin with distal spine. Uropodal exopod posterodistal

margin with row of 6—ë 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.

Neotrypaea 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).

Type species. Callianassa californiensis Dana, 1854, by

original designation.

Diagnosis. Rostrum obsolete or obtusely triangular, flat, not

reaching cornea. Pleomere | 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 right-

angled. 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. (1n

press) but not in the morphological treatment. Species assigned

to Nihonotrypaea were reported to differ from Neotrypaea 1n

having the antennular peduncle shorter than the antennal

peduncle (Lin et al., 20072) 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).

G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder

Notiax Manning and Felder, 1991

Notiax Manning and Felder, 1991: 772—773.—Sakai, 2005b: 18

(as synonym of Callianassa).

Not Notiax.— Sakai, 2011: 381—382.

Type species. Callianassa brachyophthalma A. Milne-

Edwards, 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 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. 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 1s

not lobed as 1n 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 | 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

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 | 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 |.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-288C-

4CDA-80F4-1 BEE665FA3D0

Type species. Callianassa praedatrix De Man, 1905, by present

designation.

Diagnosis. Rostrum acute, anteriorly directed, as long as

eyestalks. Pleomere | 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:lIsid:zoobank.org:act:0F41845 B-F526-

4A B8-80B7-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.

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, 1925: 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 1s recognisable by the

absence of a hook on the merus of the major cheliped, the

propodi being swollen and the pair being little different 1n 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, 19993) 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 (19993) 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 ] and 2 on a single male that was renamed

Callianassa ngochoae by Sakai (19992) and later synonymised

by Komai et al. (2014a). The generic position of Callianassa

ngochoae 1s 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: C2EFC849-6Có6B-

4466-8B47-9DF66EBF5D46

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

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 two-

thirds 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 blade-

like 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. (20143) described Kayllianassa

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 1s true in

Rayllianassa, and the minor cheliped is relatively smaller. The

atypical maxilliped 3 dactylus seen in K. rudisulcus 1s 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—ë 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:DCDIDFBE-9DFF-

481E-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 | 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 Agaballianassa 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-C36C-

4B78-8A7A-BB82710B67B5

Type species. Callianassa poorei Sakai, 1999, herein

designated.

Diagnosis. Rostrum obsolete or obtusely triangular, flat, not

reaching cornea. Pleomere | tergite undivided or with weak

transverse step. Maxilliped 3 merus wider at ischium-merus

suture than long. Male major cheliped merus with I 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 blade-

like setae proximal to long setae on distal margin. Telson

lateral margins convex.

Etymology. An alliteration of Tasmania, type locality of the

type species, and 7rypaea, 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 truncate-

convex, 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).

lastrypaea differs from Filhollianassa, its sister taxon in

Robles et al. (in press) 1n 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 | 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 1schium, 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 1n broad bands and not

adjacent. 7rypaea has only one species which 1s 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 7rypaea based on shared

features of the male pleopods | 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.—Sakai1 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 | 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 | 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 1s

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 | 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

Key to genera of Callianopsidae

|. 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. 12€) aaa

Bathycalliax (1 species, B. geomar)

— A 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 1s 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 1s 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, Tiirkay, 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 | or more distal spines.

G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder

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 WNeocallianopsis 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.—Saka1, 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.—Hyzny$ 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 |

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 | 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; 1, Corallianassa. Maxilliped 3: J, Mocallichirus;

k, Glypturoides; 1, 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

o rS CRI

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. lelson, uropod: 1, Audacallichirus;

J. Balsscallichirus; k, Glypturoides; \, Karumballichirus; m, Lepidophthalmus; n, Mocallichirus; o, Kraussillichirus; p, Neocallichirus.

Original illustrations: n, Mocallichirus mocambiquensis, UF 13986; e, Mucrollichirus mucronatus, MNHN-IU-2013-2777.

105

Callianassidae and related families

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;1, Neocallichirus raymanningt;

J, N. vigilax. Male pleopod 2: k, Glypturus armatus; l, Corallianassa coutierei; m, Grynaminna tamakit; 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

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 | 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 parallel-

sided, unlike 1n callianassids where it 1s 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 | 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 1s small and simple or absent. The

epipod of maxilliped | 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

|. 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

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-1)

orrarely sharp (in Neocallichirus vigilax, Lepidophthalmus

tridentatus) 4

3. Major chelipeds with 3 spines on upper inner margin of

merus and propodus, and 4—6 on lower margin of carpus

(fig. 130); 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. 131); scaphocerite absent;

male pleopod 2 with appendix masculina fused to

endopod (fig. 151) Corallianassa

G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder

10.

|].

Maxilliped 3 exopod present, minute (figs 13k, 1) 5

Maxilliped 3 exopod absent Ó

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

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

Male major cheliped fingers longer than square palm (fig.

13s); telson widest proximally or at midpoint, with convex

posterior margin (fig. 141) Karumballichirus

Male major cheliped fingers shorter than rectangular

palm (fig. 13t); telson widest at midpoint, posterior margin

concave (fig. 14k) Glypturoides

Antennular peduncle shorter or as long as antennal

peduncle (figs 13c, d, f) 9

Antennular peduncle significantly longer than antennal

peduncle (fig. 13b) 13

Major cheliped merus lower margin with simple proximal

hook, dactylus massive (fig. 13p); maxilliped 3 dactylus

with expanded setose apex (fig. 13m)

Major cheliped merus lower margin denticulate or with

toothed blade, dactylus not significantly broadened (figs

13q, r); maxilliped 3 dactylus tapering 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 1]

lelson parallel-sided over most of length, with obtuse

angled apex, domed dorsally (fig. l4n); maxilliped 3

propodus longer than wide, free distal margin oblique;

male pleopod 1 consisting of 1 slender article (fig. 153)...

Mocallichirus

Telson widest proximally or at midpoint; maxilliped 3

propodus as long as wide, free distal margin nearly

transverse; male pleopod | consisting of 2 articles 12

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. 141); 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

erooves; 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

erooves and oblique-transverse rows of long setae (fig.

I4h); male major cheliped merus with lower margin

almost straight; female pleopod 2 uniramous (fig. 150) ......

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)

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

107

as width of both eyestalks, shorter than antennal peduncle.

Antennal scaphocerite vestigial. Maxilliped 5 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:Isid:zoobank.org:act: F54A D76A-8172-

4DOF-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 l 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, straight-

slightly 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

(19992, 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. (1n 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 1s less expanded in N. mirim.

108

Balsscallichirus Sakai, 2011

Balsscallichirus Sakai, 2011: 414—415.— HyZny, 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 Intes, 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 | 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 | 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. Hyzny (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 1n 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. (1n 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, 19993: 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 | 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

| 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

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,

Man, 1928: 28, 96 (partim).

1903: 546—547.—De

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 | 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.

109

Remarks. Species of Callichirus are recognised by the

symmetrical pattern of deep curved transverse and longitudinal

erooves 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.—

Ne0oc-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

| 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 | 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

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 | 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, 1s 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 1s 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).—Hyzny and Müller, 2012: 969—97].—

Hyzny 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

G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder

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 I 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.—

Hyzny 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 | 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.

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: E21 E0313-3989-

4E03-BBB6-9F9844028DCO

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 | 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 | 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 1s 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 Hyzny et al. (2017).

111

Kraussillichirus gen. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:07353B50-6C2E-

4CD3-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 | 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, straight-

slightly 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. 140). 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 Komal 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 | 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 | 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.—Hyzn$ and Muñiz, 2012: 619—

621.—Hyzny and Karasawa, 2012: 62—63.—Hyzny, 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, 201l; 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 | 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 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,

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 maxiliped 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 | being

“chelate” in the former and simple in the latter. Incidentally, the

adjective "chelate" 1s 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.

Lepidophthalminusis heresynonymised with Lepidophthalmus.

No consistent morphological difference was found between

American and Indo-West Pacific species (Komai et al., 2018;

Callianassidae and related families

Robles et al., in press; Robles and Felder, 2015). Most spectes

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 l 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 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 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:8E845452-37C7-

4271-B787-E28D6285A E8F

Type species. Callianassa mocambiquensis Sakai, 2004, by

present designation and monotypy.

Diagnosis. Anterior branchiostegal lobe sclerotised, well

produced anteriorly beyond junction with oblique

113

branchiostegal ridge with which it articulates by means of a

virtual condyle. Rostrum obsolete or obtusely triangular, flat,

not reaching cornea. Pleomere | 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 straight-

concave. Pereopod 4 propodus simple, distally rounded on

lower margin. Male pleopod I consisting only of I 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, straight-

slightly 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 | 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:Isid:zoobank.org:act: B57E69B6-D096-

459C-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 | 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. (1n 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.— HyZny and Karasawa, 2012: 60—65.

Sergio Manning and Lemaitre, 1994: 40.— Sakai, 2011: 467—468

(type species, Callianassa guassutinga Rodrigues, 1971, by original

designation).—Hyzny 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 | 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 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 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 1s 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

(19993, 2005b) included Sergio in the synonymy of

Neocallichirus, but 1n 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

| (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. (2015).

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 1s 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 | tergite undivided or with weak transverse

step. Maxilliped 3 1schium 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, 19992,

2005b). Tudge et al. (2000) found it was a sister to other species

of Lepidophthalmus and treated 1t 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 |

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

pae

eem pe^ |

| e

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; 1, Ctenocheloides; j, k, Laurentgourretia.

Original illustrations: f, J, k, Laurentgourretia rhopalommata, MNHN-IU-2014-11417.

LN AAO VN

t ANT [

T 4 LL HE HTT ll | NN NN

| | NN

\\ W NM

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; 1, Gourretia. Maxilliped 3: J, Gourretia. Male coxa 5:

k, Laurentgourretia. Male pleopods 1, 2: |, m, Laurentgourretia.

Original illustrations: k-m, Laurentgourretia rhopalommata, MNHN-IU-2014-11417.

116

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

] 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

(Hyzny 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 1s 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.

Key to genera of Ctenochelidae

l. 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

G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder

2. Rostrum spine-like, dorsally carinate; cornea flat, with

weak pigmentation (fig. l6g); pleomeres 1 and 2 together

half as long as pleomeres 3—5 (fig. 16a) Ctenocheles

— Rostrum obsolete, flat; cornea disc-like, strongly

pigmented (fig. 161); pleomeres | 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. l6f); 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.

17]) 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. 17]) 6

6. Minor cheliped with sharp proximally directed teeth along

cutting edges of fingers (fig. 171); 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 | 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.

Ctenocheles Kishinouye, 1926

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).— HyZny et al., 2014: 249—250.— Poore

and Dworschak, 2017: 122 (synonymy).

Tosacallianassa Sakai and Türkay, 2016: 813 (type species,

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

| 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. [n 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 1s a sister taxon to two species of

Ctenocheles 1n 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

119

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 Linero

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 1s P. biffari.

Sakai (2005b) treated the genus first as a synonym of

Callianopsis and 1n 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. 161), which is typical of the

family, nor with any ctenochelid.

sakai (2017a) synonymised Dawsonius with Gourretia on

the basis of the same male pleopods | and 2. In fact, Dawsonius

is unique In pleopod l possessing a distomesial rounded blade

and acute apex, whereas in Gourretia and Paragourretia the

distomesial lobed 1s acute and smaller than the apical lobe.

Gourretia de Saint Laurent, 1973

Gourretia de Saint Laurent, 1973: 514.—Le Loeuff and Intés,

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 & Intés, 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 | 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 1n

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.

lelson 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"Jateral 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 1s

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 Intés,

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.

lvorygourretia, 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 | and 2. The

outline of pleopod 1 1s typical of species of Gourretia and its

supposed four articles are probable artefacts. Pleopod 2 1s

typical, except for the absence of an appendix interna, as in

the previous species. Ruiyuliugourretia 1s 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 Kictenocheloides 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 | 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 1s a male with male-like pleopods and a gonopore on

the coxa of pereopod 5 (figs I7k-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 JLaurentgourretia,

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 | 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 1s 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 |

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 1s 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. 51 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. —

Hyzny, 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 | 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 (19992, 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 | 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

Turkay (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 | 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, pleopodl, 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: 1, Paraglypturus; j, Calliax; k, Pseudocalliax.

Pereopod 3: k, Calliaxina; |, Calliax. Pereopod 4: m, Paraglypturus.

Original illustrations: a, Pseudocalliax tooradin NM V J303; e, Calliaxina SA-01, UF 36699; h, Eucalliax quadracuta, Panama, NHM W 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; t, Calliax. Male pleopod 1: g, h, Calliaxina bulimba; 1, C. kensleyi; J, C. SA-01; k, C. punica; |, 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, 1, Calliaxina SA-01, UF 36699; e, 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.

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 | 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 | 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 191—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 | of C. bulimba (fig. 19g; Dworschak, 2005: figs 5c, d;

Poore and Griffin, 1979: fig. 21g), E. inaequimana (fig. 190, p;

Dworschak, 2014: figs 4j-1) 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 1n having a bilobed article 2 with a

separate appendix interna (fig. I9h). This would appear to be a

precursor of the more typical eucalliacid form (figs 191-n). A

third form of male pleopod l 1s seen In E. aequimana where

article 2 1s longer than article 1, parallel-sided and setose

(Dworschak, 2014: figs 9h, 1). These forms appear to cross

generic lines. The role of ontogeny in the morphology of

pleopod | 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). Sakars 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.

125

Key to genera of Eucalliacidae

l. Major chela massive; minor chela with fixed finger acute,

shorter than dactylus (fig. 18]); pereopod 3 propodus oval,

little longer than wide, with short rounded proximal lobe on

lower margin, lower margin straight, upper margin strongly

convex (fig. 181); 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. 191); 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. 16J); 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. 181); 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

Implicit attributes. Unless Indicated otherwise, the followine

attributes are implicit throughout the generic diagnoses. Anterior

dorsal carapace smooth; cardiac sulci absent. Sternite 7 and

pereopod 5, coxa-ischium glabrous. Pleomere | 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 | 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:

G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder

109-110.—Ngoc-Ho, 2003: 489—490.— Sakai, 2005b: 196-197.—

Sakai, 2011: 495-496.—Hyzny 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 Ngoc-

Ho’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 Garcia

Raso et al. (2019). Their records are from 622 m and 457—548

m depth respectively 1n 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.—

Hyzny, 2012: 55—56.— Sakai and Türkay, 2014: 191.—Ngoc-Ho, 2014:

549.— Poore and Dworschak, 2017: 120 (partim).— Sakai, 2018: 739—74].

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 | with pair of

sternal plates anterior to pleopods |. Maxilliped 3 ischium and

merus broad, less than twice as long as wide at their articulation.

Both chelipeds with palms as long as wide 1n 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

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 | 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 | 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. (1n 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 (partım).— Ngoc-

Ho, 2003: 489—490 (partim).—Sakai, 2011: 502-503 (partim).—

Hyzny and Hudackova, 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

127

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 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 | 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 |

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 | 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, F. 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 1s

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, 1).

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 | stout, article |

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 (19992: figs 33a—c).

None of his other generic characters is useful. The male

pleopod 1 of P. tooradin 1s not "small and simple" as stated by

Sakai (2011) but stout, with article 1 setose and article 2 with

shallow apical notch (fig. I9r). 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 I. Eyestalks flattened, contiguous, with subdistal

dorsal cornea. Antennal scaphocerite simple, triangular.

Maxilla scaphognathite without long seta on posterior lobe

Callianassidae and related families

129

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., 1n press). Pleopod 2

is similar to pleopods 3-5 (figs 20d, e). The uropodal exopod

lacks a dorsal plate (fig. 20b) and maxilliped 3 1s exceptionally

narrow (fig. 20c). The male 1s not known.

Manning and Felder (1991) included the only genus

Paracalliax 1n 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

cenera 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-Lévre 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 Turkay, 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 Fundacao de

Amparo a Pesquisa do Estado de Sao Paulo (FAPESP; Tematico

Biota 2010/50188-8; PROTAX 2016/50376-5) Conselho

Nacional de Desenvolvimento Científico e Tecnológico (CNPq;

PQ 304968/2014—5; PROTA X 440417/2015-5), and Coordenação

de Aperfeicoamento de Pessoal de Nivel Superior (CAPES;

Ciéncias 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 post-

doctoral 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

erant 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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Morphological study for understanding the sexual pattern in ghost

shrimp Callichirus major (Crustacea: Axiidea). Acta Zoologica.

http://doi.org/10.1111/azo.12272

Stebbing, T.R.R. 1900. South African Crustacea. Marine

Investigations in South Africa. Department of Agriculture, Cape

Town 2: 14—64, pls 1—4.

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Stimpson, W. 1866. Descriptions of new genera and species of

macrurous Crustacea from the coasts of North America.

Proceedings of the Chicago Academy of Science 1: 46—48.

Tirmizi, N.M. 1967. On the occurrence of Callianassa (Callichirus)

audax de Man off West Pakistan (Decapoda, Thalassinidea).

Crustaceana 13: 151-154. https://doi.org/10.1163/156854067X00314

Tirmizi, N.M. 1970. A new species of Callianassa (Decapoda,

Thalassinidea) from West Pakistan. Crustaceana 19: 245—250.

Tirmizi, N.M. 1977. A redescription of the holotype of Callianassa

mucronata Strahl, 1861 (Decapoda, Thalassinidea). Crustaceana

32: 21-26. https://doi.org/10.1163/156854077 X00845

Tsang, L.M., Lin, F.-J., Chu, K.H., and Chan, T.-Y. 2008. Phylogeny of

Thalassinidea (Crustacea, Decapoda) inferred from three rDNA

sequences: implications for morphological evolution and

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Evolutionary Research 46: 216—223. https://doi.

org/10.1111/1.1439-0469.2008.00459.x

Tudge, C.C., Poore, G.C.B., and Lemaitre, R. 2000. Preliminary

phylogenetic analysis of generic relationships within the

Callianassidae and Ctenochelidae (Decapoda: Thalassinidea:

Callianassoidea). Journal of Crustacean Biology 20 (Special

Issue 2): 129—149. https://doi.org/10.1163/1937240X-90000015

Türkay, M., and Sakai, K. 1995. Decapod crustaceans from a volcanic

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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

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

Family allocation

Callianassidae

Callichinidae

Callianassidae

Callichiridae

Eucalliacidae

Anacalliacidae

Ctenochelidae

Callianassidae

Eucalliacidae

Callianassidae

Callichiridae

Callianassidae

Callianopsidae

Callianassidae

Anacalliacidae

Callianassidae

Callichiridae

Callianopsidae

Callichiridae

Ctenochelidae

Callichiridae

Ctenochelidae

Callianassidae

Callichiridae

Ctenochelidae

Callianassidae

Ctenochelidae

Callianassidae

Ctenochelidae

Callianassidae

Callichinidae

Paracalliacidae

Callichiridae

Ctenochelidae

Callianassidae

Current combination, authority

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 (Bitfar, 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 & Intés, 1974

Necallianassa berylae Heard & Manning, 1998

Pugnatrypaea bicauda (Sakai, 2010) comb. nov.

Neotrypaea biffari (Holthuis, 1991)

Paragourretia biffari (Blanco-Rambla & Lifiero- Arana, 1994)

Biffarius biformis (Bitfar, 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.

Callianassidae and related families

Species, Genus

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

chakratonsae, 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

Family allocation

Eucalliacidae

Callichiridae

Callianopsidae

Callianassidae

Eucalliacidae

Callianassidae

Callichiridae

Callianassidae

Eucalliacidae

Callianassidae

Callichiridae

Ctenochelidae

Callianassidae

Ctenochelidae

Callianassidae

Callichiridae

Ctenochelidae

Callichiridae

Callianassidae

Ctenochelidae

Callianassidae

Eucalliacidae

Callianassidae

Callichiridae

Callianassidae

Callichiridae

Callianassidae

Callichiridae

Ctenochelidae

Callichiridae

Callianassidae

Callianopsidae

Callianassidae

Callichiridae

Callianopsidae

Callichiridae

Callianassidae

Callichiridae

Current combination, authority

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-A guilera, 1998) comb. nov.

Biffarius delicatulus Rodrigues & Manning, 1992

Gourretia denticulata (Lutze, 1937)

Callianassa diaphora LeLoeuft & 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 & Intés, 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 (Coutiere, 1899)

Neocallichirus grandimana (Gibbes, 1850)

Laticallichirus grandis (Karasawa & Goda, 1996)

Incertae sedis gruneri Sakai, 1999

137

Neocallichirus guaiqueri (Blanco-Rambla, Linero-Arana & Beltran Lares, 1995)

138

Species, Genus

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

Family allocation

Callichiridae

Callianassidae

Callichiridae

Ctenochelidae

Callichiridae

Eucalliacidae

Callichiridae

Callianassidae

Callichiridae

Callianassidae

Callichiridae

Callianassidae

Eucalliacidae

Callichiridae

Eucalliacidae

Callianassidae

Callichiridae

Ctenochelidae

Callichiridae

Ctenochelidae

Callichiridae

Ctenochelidae

Callichiridae

Ctenochelidae

Callianassidae

Eucalliacidae

Callianassidae

Callichiridae

Eucalliacidae

Callichiridae

Callianassidae

Ctenochelidae

Callichiridae

Ctenochelidae

G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder

Current combination, authority

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 & Liftiero- 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

Callianassidae and related families

Species, Genus

marchali, Callianassa

marginata, Callianassa

martensi, Callianassa

maryae, Neocallichirus

masoomi, Callianassa

matzt, 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

Family allocation

Callianassidae

Callichiridae

Callianassidae

Callianopsidae

Callichiridae

Eucalliacidae

Callianassidae

Callichiridae

Callianassidae

Callichiridae

Callianassidae

Callichiridae

Callianassidae

Ctenochelidae

Eucalliacidae

Callianassidae

Callichiridae

Callianassidae

Callichiridae

Eucalliacidae

Callianassidae

Callichiridae

Callianassidae

Ctenochelidae

Callichiridae

Callianassidae

Ctenochelidae

Callichiridae

Callianassidae

Ctenochelidae

Callianassidae

Current combination, authority

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, Tiirkay, 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 (Rodngues, 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.

139

140

Species, Genus

punica, Calliax

pygmaea, Callianassa

geshmensis, 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

Family allocation

Eucalliacidae

Callianassidae

Ctenochelidae

Eucalliacidae

Callichiridae

Ctenochelidae

Callichiridae

Callianassidae

Callichiridae

Callianassidae

Eucalliacidae

Callianassidae

Callichiridae

Ctenochelidae

Callianassidae

Callichiridae

Ctenochelidae

Callichiridae

Callianassidae

Callichiridae

Callianassidae

Callichiridae

Callianassidae

Callichiridae

Callianassidae

Callichiridae

Callianassidae

Eucalliacidae

Callianassidae

Eucalliacidae

Callichiridae

G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder

Current combination, authority

Calliaxina punica (de Saint Laurent & Manning, 1982)

Scallasis pygmaea (De Man, 1928) comb. nov.

Gourretia geshmensis 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 & Tiirkay, 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 & Intés, 1974)

Callichirus seilacheri (Bott, 1955)

Ctenocheles serrifrons LeLoeuff & Intes, 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)

Callianassidae and related families

Species, Genus

truncata, Callianassa

tulearensis, Calliax

turnerana, Callianassa

tyrrhenus, Astacus

uncinata, Callianassa

variabilis, Callianassa (Cheramus)

vaugelasi, Neocallichirus

vietnamensis, Irypaea

vigilax, Callianassa (Callichirus)

whitei, Callianassa

winslowi, Callianassa (Callichirus)

xishaensis, Calliaxina

xutha, Corallianassa

zarenkovi, Paragourretia

Family allocation

Callianassidae

Eucalliacidae

Callichiridae

Callianassidae

Callichiridae

Callianassidae

Callichiridae

Callianassidae

Callichiridae

Eucalliacidae

Callichiridae

Ctenochelidae

141

Current combination, authority

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 (Biftar, 1971)

Anacalliax argentinensis (Biffar, 1971)

Callianassidae

Agaballianassa 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 & Intés, 1974

Callianassa marchali LeLoeuff & Intés, 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 (Biftar, 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 (Guzman &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 (Komali, 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, 1985) comb. nov.

Spinicallianassa spinicauda (Komai, Maenosono & Fujita, 2014) comb. nov.

Tastrypaea poorei (Sakai, 1999) comb. nov.

Irypaea 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 (Tirmiz, 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 (Coutiére, 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 familles 145

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, Linero-Arana & Beltran 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, 19774)

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 geshmensis Sepahvand, Pouyani & Momtazi, 2016

Gourretia sinica Liu & Liu, 2010

Gourretia zarenkovi (Sakai, 2010) comb. nov.

146 G.C.B. Poore, P.C. Dworschak, R. Robles, F. Mantelatto and D.L. Felder

Kiictenochelodes nomurai (Komali, 2013)

Laurentgourretia rhopalommata Sakai, 2004

Paragourretia aungtonyae (Sakai, 2002)

Paragourretia biffari (Blanco-Rambla & Linero-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 & Intes, 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 (Bittar, 19770)

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