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Contents

Volume 66 Issue No. 2 2009

189 > The postcranial anatomy of two Middle Devonian lungfishes (Osteichthyes, Dipnoi) from Mt. Howitt,

Victoria, Australia

John A. Long and Alice M. Clement

203 > New asterinid species from Africa and Australia (Echinodermata: Asteroidea: Asterinidae)

P. Mark O’Loughlin

215 > Observations of reproductive strategies for some dendrochirotid holothuroids (Echinodermata:

Holothuroidea: Dendrochirotida)

P. Mark O’Loughlin, John Eichler, LeonAltoff, Audrey Falconer, Melanie Mackenzie, Emily

Whitfield and Chris Rowley

221 > Australian Axiidae (Crustacea: Decapoda: Axiidea)

Gary C.B. Poore and David J. Collins

289 > New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

UlfScheller

Memoirs of Museum Victoria 66: 189-202 (2009)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://museumvictoria.com.au/About/Books-and-Journals/Journals/Memoirs-of-Museum-Victoria

The postcranial anatomy of two Middle Devonian lungfishes (Osteichthyes, Dipnoi)

from Mt. Howitt, Victoria, Australia

John A. Long 1, 2,3,4 and Alice M. Clement 1,2

Abstract

Keywords

1 Department of Sciences, Museum Victoria, PO Box 666, Melbourne, Victoria, Australia, 3001. (jlong@ museum.vic.

gov.au, aclement@museum.vic.gov.au):

2 Research School of Earth Sciences, The Australian National University, Canberra, Australia; 0200;

3 School of Geosciences, Monash University, Clayton, Victoria, Australia, 3800.

4 Natural History Museum of Los Angeles County, 900 Exposition Boulevard, California 90007, USA (jlong@nhm.org)

Long, J.A. and Clement, A.M. 2009. The postcranial anatomy of two Middle Devonian lungfishes (Osteichthyes, Dipnoi)

from Mt. Howitt, Victoria, Australia. Memoirs of Museum Victoria 66: 189-202.

The postcranial skeletons of two upper Givetian lungfishes from Mt. Howitt, Victoria, Australia, show remarkable

similarities, despite the fact that one is a tooth-plated form ( Howidipterus Long 1992) whilst the other has a denticulate

dentition ( Barwickia Long 1992). Both genera show identical body shape with a short first dorsal fin and greatly elongated

second dorsal fin, and small anal fin. The cleithra and clavicles are remarkably similar except for Barwickia lacking

external ornament on the lateral lamina of the cleithrum and having a smaller branchial lamina on the clavicle. Both have

paddle-shaped subdermal anocleithra that meet the posterior process of the I bone, approximately the same numbers of

cranial ribs, pleural ribs, supraneural and subhaemal spines, the same expanded dorsal and anal fin basals with similar

number of proximal and middle radials supporting the fins, and approximately the same number of radials supporting the

hypochordal lobe of the caudal fin. These numerous similarities in the postcranial skeletons of the two genera strongly

suggest that their differing feeding mechanisms probably evolved from a shared ancestral form having a similar postcranial

skeleton. Implications for hypotheses of dipnoan phylogeny are discussed.

Pisces, osteichthyes, Dipnomorpha, Devonian, postcranial skeleton, anatomy, evolution, Australia

Introduction

Since the time of Dollo (1895) the significance of postcranial

features in the large scale evolutionary trends of the Dipnoi

has been repeatedly noted (Graham-Smith and Westoll, 1937;

Westoll, 1949; Lehman, 1966; Bemis, 1984; Long, 1990;

Pridmore and Barwick, 1993). However, despite the recent

wealth of new information on the cranial anatomy of early

lungfishes, there is a lack of information on their postcranial

skeletons. Over seventy Devonian genera of lungfish are now

known (Marshall, 1987; Jarvik, 1980; Janvier 1996) yet only

four of these, Fleurantia denticulata (Graham-Smith and

Westoll, 1937), Dipterus valencienessi (Ahlberg and Trewin,

1994) and two genera from the Late Devonian Gogo Formation

of Western Australia, Chirodipterus australis and

Griphognathus whitei (Pridmore and Barwick, 1993; Campbell

and Barwick, 2002), have had the postcranial skeleton

described in detail. Other Devonian dipnoans which have had

aspects of the postcranial skeleton described include

Uranolophus (Denison, 1968; Campbell and Barwick, 1988a),

Dipterus (e.g. Schultze 1970, 1975; Campbell and Barwick,

1988a, Campbell et al. 2006), Rhinodipterus (Schultze, 1975),

Pillar arhynchus (Barwick and Campbell 1996), Adololopas

(Campbell and Barwick 1998), Griphognathus (Schultze,

1969; Campbell and Barwick, 1988a; Pridmore and Barwick,

1993). Isolated vertebral centra of dipnoans from indeterminate

taxa have been figured and described also by several workers

(e.g. Jarvik, 1952). Therefore the complete description of the

postcranial skeleton in two more Devonian genera, presented

in this paper, contributes significant new information to the

subject, and allows discussion of phylogenetic problems

concerning the monophyly of tooth plated versus denticulated

dipnoan lineages.

The Mt. Howitt fauna, of uppermost Givetian age (Young,

1993, 1999), represents one of the best preserved and most

diverse late Middle Devonian freshwater fish assemblages from

any single site in the Southern Hemisphere, and is also

significant in being the key stone for biostratigraphic correlations

throughout eastern Victoria (Long, 1983, 2004; Long and

Werdelin, 1986; Cas et al 2003). There are two genera of

lungfish at Mt. Howitt, regarded by Long (1993) as members of

the Family Fleurantiidae ( contra Long, 1992, in which

Howidipterus was placed provisionally in the Dipteridae). One

has tooth-plates with occasional denticles between the tooth-

190

J.A. Long and A.M. Clement

ridges ( Howidipterus)-, the other has a denticle-covered

dentition, although rows of teeth may be clearly distinguished

on the pterygoids ( Barwickia ). Although Long (1993) suggested

that the fleurantiid dentitions probably evolved by heterochronic

processes (McKinney and McNamara, 1991), namely

paedomorphic retention of tooth-row development in conjuction

with peramorphic development of denticle fields (“dissociated

heterochony”), it is the nature of the postcranial skeletons in

these forms that gives further information on their possible

phylogenetic affinities. The phylogenetic analysis of Devonian

lungfishes by Ahlberg et al. (2006) supported a close relationship

between Howidipterus and Barwickia.

Materials and methods

The Mt. Howitt lungfishes were studied from latex casts of the

natural moulds preserved in black shale. The specimens are

generally preserved as flattened, slightly disrupted carcasses,

but often fine preservation of cartilage bones, such as elements

of the visceral skeleton, are clearly seen from the latex peels.

Photographs are of latex casts dusted with ammonium chloride.

The description of the postcranial skeleton follows terminology

used by Goodrich (1958), Graham-Smith and Westoll (1937),

Long (1987, for the cleithrum) and Cloutier (1996). Figure 1

outlines the terminology used for axial skeleton components

used in this work.

Outline drawings and descriptions of postcranial features

have been made using a camera lucida. Comparative material

examined includes three-dimensional lungfish bodies from the

Gogo Formation of Western Australia held in the W.A.

Museum and in the Geology Department, The Australian

National University, Canberra, and collections of North

American and European Devonian lungfishes held in the

British Museum of Natural History, London, The National

Museum of Scotland, Edinburgh and the Australian Museum,

Sydney. Specimens referred to in this work are housed in the

palaeontological collections of the Museum of Victoria,

Melbourne (MV), The Australian Museum, Sydney (AM), and

the Western Australian Museum (WAM), Perth.

Descriptions of the postcranial skeletons

The two genera show remarkably similar body form and

postcranial skeletal morphology. Both genera are commonly

preserved in size ranges of 10-20 cm, the largest individual

indicating a maximum length estimated at close to 40 cm

( Howidipterus ). Although there are many specimens

representing both forms which show the overall shape and

proportions of the body and fins (e.g. figs. 4, 6), very few

specimens show good preservation of the axial skeletal

elements, and in most specimens the counts of these elements

are based on impressions of ribs and supraneurals that have

been overprinted by the squamation.

Pectoral girdle

The exoskeletal pectoral girdle in both genera consists of a

large cleithrum and clavicle, and a smaller paddle-shaped

subdermal anocleithrum which articulates anterodorsally with

lepidotrichia (segmented)

lepidotrichia (unsegmented)

middle radials

*/**

<*/

proximal radials

expanded fin basal

supraneural spine

neural arch

(basidorsal)

dorsal ligament

spinal chord

notochord

ventral arch

(haemal spine)

subhaemal spine

expanded fin basal

proximal radial

middle radial'

Figure 1. Terminology used for axial skeleton components.

the posterior subdermal process of the I bone. The

scapulocoracoid is not commonly preserved, and was probably

largely cartilaginous, as were the axial mesomeres that

presumably formed the pectoral and pelvic fin skeletons. In

one specimen (Barwickia, MV P198046) there is an impression

of part of the scapulocoracoid showing the exposed portion to

have a similar form as that figured for Chirodipterus (Campbell

and Barwick, 1987). Neither the shape of the glenoid fossa nor

the support buttresses for the scapulocoracoid can be

determined from the latex peel.

Cleithrum. The cleithra in Howidipterus (fig. 3) and Barwickia

(figs. 3, 5) are very similar in overall form and shape. Both are

generally similar to the cleithra of other Late Devonian

dipnoans, especially Eoctenodus microsoma (Long, 1987) and

Scaumenacia (Jarvik, 1980). The cleithrum has an expanded

dorsal end, strong dorsoventral lateral thickening and extensive,

inwardly directed branchial lamina that meets the branchial

lamina of the clavicle along a prominent thickened ridge. They

differ from each other in that the externally exposed region of

The postcranial anatomy of two Middle Devonian lungfishes (Osteichthyes, Dipnoi) from Mt. Howitt, Victoria, Australia

191

Figure 2. Howidipterus donnae: a, photograph of MV P181792; b, interpretive drawing of MV P198045; c, MV P198042, sketch interpretation

of large specimen, slightly disarticulated.

192

J.A. Long and A.M. Clement

lateral

lamina

neural spines

branchial

lamina ^

operculum ^

lateral

thickening

process

cleithrum

anocleithrum

clavicle

head

cranial rib L

lateral * L

thickeninc

branchia|^<

laminaS*'

anocleithrum

ornamented

lateral lamina

cranial

ribs

branchia[

lamina

ridge^^

clavicle,

cleithrum

_ lateral /

thickening J

cleithrum

clavicle

Figure 3. Shoulder girdle: a, Barwickia downunda cleithrum and clavicle, MV P181890; b, Barwickia downunda with anterior ribs and neural

spines, MV P198046; c, Howidipterus donnae exoskeletal shoulder girdle, MV P 181883; and d, also showing anocleithrum, MV P181792.

the cleithrum (lateral lamina) in Howidipterus has weakly

developed surface pitting, indicating it was situated just below

the dermis in life. The cleithrum of Barwickia shows no

external ornament or marking on its lateral lamina, and

appears to have a more strongly developed lateral thickening.

As in Eoctenodus there is a marked anterior angle on the

branchial lamina in both forms, and a roughened mesial pit is

formed where the branchial lamina meets the lateral lamina.

Eoctenodus differs in having a notch present at the ventromesial

corner of the branchial lamina (Long 1987, Fig. 6) which is not

seen in either of the Mt. Howitt forms.

There are some variations seen within the cleithra of

Howidipterus. P181883 (Fig. 3; figured only in part by Long,

1992, Fig. 3G) shows the presence of a distinct mesial lamina

in addition to a branchial lamina. This outer, mesial lamina is

part of the lateral thickening of the cleithrum, and may have

served to separate the overlap area of the operculum from the

gill chamber.

In visceral view there is no indication of the shape or size of

the scapulocoracoid attachment area in either form, as seen in

some other early lungfishes (e.g. Uranolophus, Campbell and

Barwick, 1988b; Chirodipterus, Campbell and Barwick, 1999).

Clavicle. The clavicles are well-preserved in several specimens

from both genera (figs. 2, 3, 5). They are large bones, almost as

long as the cleithrum and smoothly curved throughout their

extent. Overlap between the cleithrum and clavicle in the Mt.

Howitt genera was relatively short and narrow, unlike the

primitive form Uranolophus in which the clavicle had an

elongate, extensive dorsal overlap surface (Campbell Barwick,

1988b). The ventral laminae in both Mt. Howitt forms are of

simple triangular shape, lacking a notch for overlap of the

principal gular plate as seen in some other Devonian lungfish

such as Chirodipterus (e.g. WAM 90.10.8) and Uranolophus

(Campbell and Barwick, 1988b, Figs. 23-25). The clavicles of

both Howidipterus and Barwickia possess a strong lateral

thickening along the outermost edge, which increases in

thickness towards the junction with the cleithrum. The

branchial lamina of the clavicle of Howidipterus is notably

more extensive than that in Barwickia (fig. 3).

Anocleithrum. The anocleithrum is well-preserved and of

similar paddle-shape in several specimens of both forms

(Barwickia, figs. 3, 5; Howidipterus, Long, 1993: Fig. 5). In

Barwickia the anocleithrum is 80% as long as the cleithrum.

The postcranial anatomy of two Middle Devonian lungfishes (Osteichthyes, Dipnoi) from Mt. Howitt, Victoria, Australia

193

second dorsal fin

first dorsal fin

anal fin

cleithrum

1cm

Figure 4. Outline of postcranial body and fins: a, Barwickia downunda\ b, Howidipterus donnae.

In Howidipterus the anocleithrum appears to be slightly

smaller compared with the cleithrum. The anterior end of the

anocleithrum is slender and produced into a strong anterior

spine that remains in contact with the posterior process of the

I bone of the skull in many specimens, suggesting a strong

ligamentous connection in life.

Pectoral and pelvic fins

Pectoral fin. The pectoral fin is well-preserved in many

specimens, although it shows only the outline of the fringing

fin rays and some small scales covering the fin. There is no

preservation of endoskeletal fin bones in either genus. In both

genera the pectoral fin approximates to the same length as the

skull roof, and is approximately four times as long as its

broadest part. The fin rays emerge from the edges of the fin as

long, curved, unbroken elements which then subdivide into

smaller elements close to the margins of the fin. The fin rays

emerge a short distance from the beginning of the fins, and

there are approximately 45-50 rows of lepidotrichia present.

Pelvic fin and girdle. Part of the endoskeletal pelvic girdle is

seen preserved only in one specimen of Barwickia (AM

F98074 part and counterpart, fig. 6 A, B). It shows a large

articulatory facet for the axial mesomeres of the pelvic fin,

and a short process near this facet which might be the

homologue of the dorsomesial process described on the pelvic

girdle of Chirodipterus (Young et al., 1990, Fig. 4). The

overall shape and size of the girdle in Barwickia closely

matches the girdle of Chirodipterus being almost a

parallelogram in shape, not elongated with a long anterior

process as in Griphognathus.

194

J.A. Long and A.M. Clement

middle

radial proximal f irst dorsal fin

axis of vertebral

column

operculum

pleural rib

cranial

ribs

clavicle

cleithrum

Figure 5. Barwickia downunda, features of postcranial skeleton: a, MV P181784; b, interpretive drawing of same.

The pelvic fin is well-preserved in many specimens (e.g.

figs. 4, 6) and is of identical shape and proportions to that of

the pectoral fin in both genera, exhibiting exactly the same

style of fin-ray bifurcation and proportions. The pelvic fin

emerges opposite the first dorsal fin, at the point where the

paired pleural ribs end. Approximately 40-50 rows of

lepidotrichia fringe the dorsal and ventral margins of the fin.

Median fins

Anterior dorsal fin. The anterior, or first dorsal fin, is the

smallest of the median fins, being about one fifth the length of

the second dorsal fin at its base, and slightly smaller than the

anal fin, being approximately 3% of the total length of the fish

in both forms. It originates from approximately the 20th to

22nd myotomal segment, and is supported by a dorsally

expanded racquet-shaped fin basal (radial), which itself is

supported by a shortened supraneural relative to the lengths of

the supraneurals anterior and posterior to it. In some specimens

of Barwickia there is a short median anteriorly directed

process developed on the expanded fin support (fig. 6E), a

feature not seen in any specimen of Howidipterus. The

expanded fin basal is approximately half as broad as the

expanded anterior support bone for the second dorsal fin.

Groups of three or four stiff lepidotrichia attach to

approximately three proximal radials that articulate ventrally

with the anterior dorsal fin support bone. These bunches of

four or more unsegmented lepidotrichia continue for about

half the extent of the fin before giving way to smaller segmented

and bifurcating fin rays for the distal extent of the fin. About

16-18 lepidotrichial rows are present at the insertion of the

anterior dorsal fin of both genera. The area of the fin supported

by unsegmented lepidotrichia was covered by small scales.

Posterior dorsal fin. The posterior, or second dorsal fin, is the

largest median fin and extends for approximately 15% the total

The postcranial anatomy of two Middle Devonian lungfishes (Osteichthyes, Dipnoi) from Mt. Howitt, Victoria, Australia

195

, expanded

proxima! fln basa| .

radiaL

5mm

Figure 6. Barwickia downunda a and b, AM F98074 body flattened showing postcranial skeleton and head: a, dorsal view; b, ventral view; c, MV

P181784, showing details of tail and fins; d, MV P181784, photograph of tail and fins; e, MV P198044, internal support bones for first and second

dorsal fins.

196

J.A. Long and A.M. Clement

length of the fish in both genera (figs. 4, 6). It begins at a point

slightly anterior to the anterior margin of the anal fin, although

the supraneural leading to the fin-support bones of this fin

meets the notochordal axis at the same myotomal segment as

the infrahaemal supporting the expanded anal fin bone. In

both genera the posterior dorsal fin has a gently lobate shape,

and is supported anteriorly by a large expanded radial that

articulates distally with five proximal radials that support four

middle radials (fig. 6 C, D) that each carry the bunches of 3-4

unsegmented lepidotrichia. This expanded radial has a waisted,

stout shaft that expands ventrally to articulate with a thick

supraneural. The five proximal radials that support the anterior

end of the fin increase evenly in size posteriorly. There are

10-11 other proximal radials that follow posteriorly from the

five, articulating with the anterior expanded bone thus totalling

15 or 16 elements. Each of the anterior proximal radials and

the anterior expanded bone are supported by supraneurals

articulating to the vertebral column, although the posteriormost

three or four may articulate directly to the mineralised section

of the vertebral column. Their exact position is not clear from

the preservation of the material. About 60 rows of unsegmented

lepidotrichia support the ventral half of the fin. The expanded

anterior fin basal is approximately as large and of identical

shape to that of the anal fin support bone.

Anal fin. The anal fin in both genera is only slightly broader in

shape than the first dorsal fin and inserts into the same

myotomal segment (c. 24th) as the anterior margin of the

second dorsal fin. It is supported by a stout racquet-shaped fin

basal bone (. Howidipterus , fig. 2; Barwickia, fig. 6C) which

articulates dorsally with a short but thick infrahaemal spine.

Four proximal radials articulate posteroventrally with the

expanded fin basal and these each articulate with a middle

radial that supports bunches of 3-5 stiff lepidotrichia.

Approximately 15-20 lepidotrichial rows support the dorsal

half of the fin.

Caudal fin. The caudal fin is well-preserved in several

specimens of both genera and appears to have exactly the same

outline and development of fin-ray support bones. The tail is

heterocercal with a triangular shape, the axis of the vertebral

column being deflected about 20° from the main axis of the

body (figs. 4, 6, 8). The ventral edge of the hypochordal lobe

begins almost immediately posterior to the anal fin, and

equivalent in position to half-way along the posterior dorsal

fin. The anterior edge of the hypochordal lobe is supported by

three rows of fin support bones: the dorsal series (subhaemals)

articulate with the vertebral axis, and distally these articulate

with a row of proximal radials which articulate with a 1:1 ratio

with middle radials. The middle radials have bunches of

unsegmented lepidotrichia attached to them. There appears to

be only 8-9 rows of middle radials before the tail narrows, and

the proximal radials or subhaemals support the fin directly on

the vertebral axis. At this point the rest of the fin structure is

unclear, and appears to consist largely of bunches of

lepidotrichia inserting directly into the axis of the vertebral

column. A small epichordal lobe of segmented lepidotrichia is

present in both genera (e.g. Barwickia, fig. 6C).

Axial skeleton

The axial skeleton consists of the vertebral column and its

articulating spines and ribs. Paired pleural ribs are present

throughout the anterior half of the fish, articulating with the

first 19-21 vertebral elements within each myoseptum in

Howidipterus, and between the 20-22 myosepta in Barwickia,

thereby being almost identical (exact counts are difficult to

make due to the overprinting of paired ribs in the crushed state

of preservation).

The vertebral column is well-ossified in the tail region of

both species, although individual centra are not clearly

differentiated, instead there is a continuous ossified or

mineralised column. This may represent mineralisation of the

notochord in this region as suggested by Schultze (1970) and

Arratia et al. (2001), or they could be individual ring centra

that are only well-ossified in the caudal part of the vertebral

column. Anteriorly there are poorly preserved remains of

vertebral arches in some specimens (fig. 7). These closely

resemble the dorsal arch elements (basidorsals) described in

Griphognathus by Campbell and Barwick (1988a, Figs. 34,

35). Ventral elements, possibly representing ossified

basiventrals are sometimes seen, and impressions of whole

body specimens suggest that they were present throughout the

vertebral column in younger individuals. The largest specimens

basidorsal?

5mm

basiventral?

Figure 7. Barwickia downunda: a, photograph and b, interpretive

drawing of MV P181868, details of anterior vertebral elements.

The postcranial anatomy of two Middle Devonian lungfishes (Osteichthyes, Dipnoi) from Mt. Howitt, Victoria, Australia

197

show no vertebral ossification at all (e.g. large Howidipterus,

fig. 2). Supraneurals articulate to the vertebral column

throughout its length, but no secondary supraneurals are

present as exists near the first dorsal fin as in Fleurantia

(Graham-Smith and Westoll, 1937).

Howidipterus and Barwickia have approximately 20-22

vertebrae and supraneurals anterior to the first dorsal fin, then

4-5 or so supraneurals before the second dorsal fin support in

Barwickia , and 5-7 supraneurals before the second dorsal fin

support in Howidipterus (these are accurate counts and reflect

individual variations). Both forms then show identical

development of the second dorsal fin shape and the numbers of

supraneurals supporting this fin and subhaemal spines, as

described above.

Cranial ribs. Cranial ribs are present in both forms, and appear

identical in shape (figs. 3, 5). Long (1993) gave a preliminary

description of the cranial ribs in both the Mt. Howitt lungfishes.

The expanded rectangular distal ends of the cranial ribs can be

often recognised in specimens where the squamation has

overprinted the axial skeleton. Each cranial rib has a slightly

expanded flat head, narrow neck, and a flat shaft that broadens

gradually throughout its distal length. Two pairs of cranial ribs

are present in each genus. They are easily identified as being

present in the head region of weakly disarticulated specimens

of Barwickia, being followed by the first pair of pleural ribs.

In no specimens can we see the neurocranium preserved, so

we can only deduce from the anterior extent of the cranial ribs,

moreso than for the pleural ribs (e.g. Long 1993, Fig. 3) that

they did articulate to the ventral suture of the brainacase and

posterior stalk of the parsphenoidid as in other lungfishes. In

AM F89074 (fig. 6) the flat articulatory heads of the cranial

ribs are seen lying adjacent to the posterior end of the ossified

neurocranium. In Neoceratodus forsteri the cranial ribs are

oriented almost horizontally (Goodrich, 1958), and it appears

that in the fossilised forms from Mt. Howitt the orientation of

the cranial ribs was similar as they are commonly observed

lying in a different orientation to the paired pleural ribs.

Pleural ribs. Paired pleural ribs (figs. 2, 3, 5-8) are gently

curved, almost sigmoid shaped elongate rounded elements

which run for most of the length of the trunk, terminating at the

level of origin of the pelvic fin. Anterior pleural ribs are longer

than the posterior elements, and have a more distinct curvature.

Figure 8. Reconstructions of postcranial skeletons: a, Barwickia and b, Howidipterus. Paired elements such as pleural and cranial ribs are drawn

in full.

198

J.A. Long and A.M. Clement

Approximately 18-20 pairs of these ribs are present in both

forms. They articulate dorsally with the basiventral element of

the vertebral column at a slightly expanded head having a flat

articulatory surface meeting the basiventral (fig. 7).

Phylogenetic significance of dipnoan postcranial features

During the Devonian Period lungfishes underwent major

changes in both their cranial and postcranial skeletons, leading

directly to the lineage of tooth-plated forms including the

modern genera. By the Early Carboniferous, forms like

Uronemus (Ganopristodus) had acquired essentially the same

body and fin shape seen in all subsequent lungfishes, including

extant forms: a single continuous dorsal fin that is merged with

the caudal and anal fins to give a diphycercal fin shape. A

transformation series of intermediate morphological stages in

acquiring this pattern can be seen in various Devonian

dipnoans, represented by the few known from complete or

near complete body fossils (e.g. Long 1993, Fig 7).

The series begins with the only Early Devonian genus in

which the approximate form of the body and fins is known,

Uranolophus. It shows the presence of two dorsal fins and a

large, separate anal fin, and a heterocercal caudal fin with high

angle axis of tail to body (although it is incompletely preserved,

Denison, 1968; Campbell and Barwick, 1988b). Even in the

earliest known dipnoan the anterior dorsal fin is slightly smaller

than the posterior fin. Some Late Devonian forms, like

Rhynchodipterus retain this primitive pattern in having two

almost equidimensional dorsal fins, a similarly sized anal fin and

large upturned heterocercal tail (Save-Soderbergh, 1937). In

Griphognathus there are also two widely separated dorsal fins,

with the anterior fin is seen to be slightly smaller than the posterior

dorsal fin (Schultze 1969; Campbell and Barwick 2002).

Dipterus shows a slightly more derived condition than

these forms in that the second dorsal fin is enlarged much

more than the first dorsal fin (Forster-Cooper, 1937 plate 3;

Ahlberg and Trewin, 1994). In Dipterus the first dorsal fin has

about 18 unsegmented lepidotrichia as in Howidipterus and

Barwickia, while the second dorsal fin has about 40 or so

unsegmented lepidotrichia, as compared with approximately

60 or so in the Mt. Howitt forms. Whilst these Australian

genera closely resemble Pentlandia in this respect, the latter,

from the Middle Devonian of Scotland, has several

distinguishing differences in the skull morphology. However

Pentlandia requires a detailed study to determine its exact

affinities and is here regarded as having similar level of

organisation in its postcranial skeleton as the two Mt. Howitt

genera. Pinnalongus from the Eifelian of Scotland shows a

similar condition to the Mt Howitt forms in having a very

small anterior dorsal fin and extensive posterior dorsal fin

(Newman and Den Blaawen 2007).

Fleurantia represents the next stage in the transformation

series from the Mt. Howitt forms (and possibly Pentlandia).

Fleurantia has a much larger second dorsal fin, with many

more proximal radials (16-21 elements, approximately 100

rows of unsegmented lepidotrichia; Cloutier, 1996). The first

dorsal fin is approximately the same size and has a similar

number of unsegmented lepidotrichia as in the Mt. Howitt

forms, but the anal fin in Fleurantia is further reduced in only

having 3 proximal radials articulating with the expanded fin

basal. Unlike the Mt. Howitt forms, Fleurantia lacks an

expanded fin basal supporting the anterior region of the second

dorsal fin and has a few secondary supraneurals present near

the first dorsal fin.

Scaumenacia represents the next stage in the series in

having a greatly expanded, but low first dorsal fin, and a larger

second dorsal fin (supporting approximately 180 long

lepidotrichia; Cloutier, 1996). It is also more derived than

Fleurantia and the Mt. Howitt forms in having lost the ossified

radials supporting the first dorsal fin, and in having the tail

terminate in a long, thin caudal filament.

Phaneropleuron, from the Famennian Rosebrae Beds of

Scotland, shows similar level of organisation to Scaumenacia

but incorporates both dorsal fins with the enlarged epichordal

lobe of the tail, which has now achieved a diphycercal shape,

although the anal fin is still separate. This genus also requires

further study of its postcranial skeleton before it can be

compared in detail with the previous forms.

Finally, merging the anal fin with the diphycercal tail

arrives at the condition seen in all later lungfishes, as typified

in the Lower Carboniferous genus Uronemus (= Ganopristodus,

Schultze, 1992).

From the above descriptions and discussion the following

observations and hypotheses can be suggested regarding the

phylogenetic significance of each character.

Cleithrum. In primitive dipnoans the cleithrum has a weakly

developed branchial lamina ( Uranolophus , Campbell and

Barwick, 1988b), although the feature is subsequently well-

developed in many Devonian forms (e.g. Scaumenacia,

Chirodipterus, Eoctenodus, Barwickia, Howidipterus).

Campbell and Barwick (1988a) pointed out several differences

between the cleithrum of denticulate lungfishes and that of the

presumed monophyletic ‘tooth-plated forms’. Their

comparisons used Griphognathus and Uranolophus as

denticulated forms, and Chirodipterus, Scaumenacia and

Eoctenodus as tooth-plated forms. The new material from Mt.

Howitt shows that unlike the condition described for

Griphognathus and Uranolophus, Barwickia possessed a

cleithrum (and clavicle) that was essentially the same as in

Howidipterus in possessing a large, medially extensive

branchial lamina. The same type of extensive branchial lamina

is also present in Holodipterus, regarded as one of the members

of the denticle-shedding lineage by Campbell and Barwick

(1991) but by Smith (in Campbell and Smith 1987, p.165) as a

form that could have been derived from earlier tooth-plated

forms such as Dipterus or Speonesydrion.

Fin support bones. The development of expanded racquet-

shaped median fin support bones is seen only in the Mt. Howitt

forms and in the anal fin of Fleurantia. In Griphognathus

whitei there are large expanded basal bones, but these do not

taper into thin rods as occurs in Fleurantia, Barwickia and

Howidipterus. Furthermore, Griphognathus whitei has a

unique type of dorsal and anal fin-support bone with enlarged

secondary fin basals supporting several proximal radials, and

can be regarded as specialised in this respect (e.g. WAM 86.9.

The postcranial anatomy of two Middle Devonian lungfishes (Osteichthyes, Dipnoi) from Mt. Howitt, Victoria, Australia

199

645, Pridmore and Barwick, 1993, Fig. 8 shows the fin basal

for the posterior dorsal fin). Thus the expanded racquet-shaped

fin basals could either represent a synapomorphy of Fleurantia

and the Mt. Howitt lungfishes or a homoplasy. As Fleurantia

and the Mt. Howitt forms otherwise show very similar levels

of development and dentition (Long, 1993), and the similar

shaped second dorsal fin of Scaumenacia does not have a

similar enlarged basal, we here consider it to be a derived

feature of the Family Fleurantiidae (defined nodally as the

clade Fleurantia, Howidipterus and Barwickia in Ahlberg et

al. 2006), and possibly also including Jarvikia, based only on

cranial features shared with Fleurantia (Campbell and

Barwick, 1990, Cloutier, 1996); and Andreyevichthys, based

on similar dentition (Smith et al., 1993). The presence of a few

secondary supraneurals near the first dorsal fin of Fleurantia

is here considered an autapomorphy of that genus as such

bones have not been recorded in any other fossil lungfish.

Fin shapes. The stages leading to the acquisition of the modern

dipnoan body and fin shape have been summarised in the

discussion above. The primitive condition is having two

equidimensional or nearly equally sized dorsal fins, separate

anal and heterocercal caudal fins without epichordal lobes as

seen in other sarcopterygians (e.g. Osteolepis, Glyptolepis).

The following characters are therefore seen as derived with

respect to this condition, as outlined in Ahlberg and Trewin

(1994): (a) reduction of first dorsal fin, slight enlargement of

second dorsal fin (e.g. Dipterus); (b) enlargement of second

dorsal fin (Howidipterus, Barwickia, Pentlandia ); (c) greater

enlargement of second dorsal fin ( Fleurantia ); (d) first dorsal

fin elongated but low, greater expansion of second dorsal fin,

long caudal filament developed on main axis of caudal fin

( Scaumenacia ); (e), continuous long dorsal fin axis of tail

horizontal, not inclined (Phaneropleuron); (f) anal fin merged

with continuous dorsal fin (Conchopoma, Uronemus, all Late

Palaeozoic to Recent lungfishes).

Vertebrae. The vertebrae are weakly ossified in primitive

forms, consisting of ossified neural arches that straddle an

unconstricted notochord (Uranolophus, Campbell and

Barwick, 1988b; Dipterus, Ahlberg and Trewin, 1994;

Schultze, 1975). Through the arches passes the spinal chord

and dorsal ligament.

Ossified spool-shaped centra are found only in a few forms

(e.g. Griphognathus), and are considered to be a derived

condition by outgroup comparison with other primitive

sarcopterygians (e.g. separate intercentra and pleurocentra are

primitive for other sarcopterygians; Andrews and Westoll,

1970; Ahlberg, 1989). The presence of vertebrae, as separate

basidorsal and/or basiventral ossifications is observed in

Scaumenacia (Cloutier, 1996) and at various growth stages in

the Mt. Howitt forms. Modern lungfishes have basidorsals and

basiventrals present as cartilaginous units (Goodrich, 1958;

Shute, 1972), possibly a derived condition due to loss of bone

from primitive forms.

Ribs. These have been found in all dipnoans where whole body

features are preserved, and are often referred to as ‘pleural

ribs’ in the thoracic region of the body. It is unknown whether

paired pleural ribs were extensively present in Uranolophus or

other primitive marine dipnoans like Dipnorhynchus,

Speonesydrion, Ichnomylax or Melanognathus. If so, then

this feature would have no special significance for evolution

within the Dipnoi, but otherwise could be a derived condition

within later dipnoans that co-evolved with the development of

larger lungs. The well-developed ribs present in all the marine

dipnoans from the Middle-Late Devonian Gogo (Campbell

and Barwick 2002) and Bergisch-Gladbach faunas (Schultze

1975) do not appear to be strongly curved as in the Mt Howitt

forms, so we assume this kind of ‘pleural’ rib found in the Mt

Howitt species evolved for accommodation of a larger lung for

air-breathing. The pleural ribs in Dipterus appear to be

primitively short compared with the longer elements seen in

Barwickia and Howidipterus.

Cranial ribs. Early reports of cranial ribs in one specimen of

Fleurantia (Graham-Smith and Westoll: 255) and in

Scaumenacia (Goodrich, 1909) have been confirmed by

observation of casts of these species held in the collections of

the Geology Department at the Australian National University,

and of original specimens of Scaumenacia held in the Museum

of Victoria. Aside from Barwickia and Howidipterus, the only

other Devonian dipnoans to have cranial ribs are Rhinodipterus

ulrichi (Schultze, 1975), a marine form, and possibly incipient

cranial ribs in Dipterus (Ahlberg and Trewin, 1994), known

from both freshwater and marine environments. Observation of

the marine Gogo specimens of Chirodipterus, Gogodipterus

and Griphognathus also show that cranial ribs were absent in

these forms (Campbell and Barwick 2002). The presence of

cranial ribs in lungfishes, being absent in plesiomorphic fully

marine forms), would appear to be a good synapomorphy

uniting air-gulping forms (Long, 1993). The actual morphology

of the cranial ribs has not been previously considered, although

some new information is now at hand. In Dipterus (Ahlberg

and Trewin, 1994, Fig. 6) the enlarged ribs identified as possible

cranial ribs are not ventrally expanded, showing the condition

of being enlarged pleural ribs that probably articulated with the

posterior end of the braincase. The cranial ribs in Barwickia

and Howidipterus are here considered to be more specialised

than those of Dipterus in having distally expanded, flat shapes,

allowing for more surface area on the lateral and mesial surfaces

of the ribs for attachment of ligaments to anchor the pectoral

girdle. We note the occipital ribs, that articulate to the

posterodorsal surface of the neurocranium, have not been

observed in the Mt.Howitt forms.

Dipnoan evolution: evidence from the Mt. Howitt dipnoans

The two genera of lungfishes from the Mt. Howitt deposit

exhibitidentical postcranial skeletons, and cranial morphologies

that differ slightly but are still at a similar grade of evolution

with respect to approximate numbers of skull roof bones and

cheek bone patterns (Long, 1992). Barwickia shows a dentition

that was at first thought to be typical of the denticulate feeding

mechanism (Long, 1992) but later shown to be a form of tooth

plate with large denticle fields present (Long, 1993). It should

be pointed out though that the histology of these tooth plates is

not known as the material can only be studied from latex peels.

200

J.A. Long and A.M. Clement

Howidipterus shows more typical dipnoan tooth plates that

closely resemble those of Scaumenacia in overall morphology

but can also be demonstrated to be closely related to those of

Barwickia. Dissociated heterochrony was invoked as a possible

mechanism for the development of the Barwickia type tooth

plate based on the known growth changes that occur during the

ontogeny of Andreyevichthys toothplates (Long, 1993). This

means that different rates of growth apply to the developmental

stages, such as peramorphic development of the denticle field

whilst there is restrained growth of the tooth rows

(paedomorphosis). From these observations, and the overall

nature of the Mt. Howitt fauna, with a high proportion of

endemic fauna, and palaeogeographically representing a

highland intermontane sedimentary basin deposit (Cas et al.

2004), and the recent phylogenetic analysis supporting the two

lungfishes Barwickia and Howidipterus as very closely related

(Ahlberg et al. 2006), we suggest that they may have had a

comparatively recent divergence from a common ancestor. In

overall body form they are identical, so must have had identical

functional morphologies with respect to their mode and speed

of swimming. As they inhabited the same lake system, each

must have occupied a different niche primarily based on

differing food preference in the lacustrine food chain. To date

there are no invertebrate fossils known from the Mt. Howitt

deposit, despite delicate, articulated preservation of both the

fish (in all stages of growth) and plants, thus sources of food

for the lungfish are possibly to be found in the known fossil

record of the site, or alternatively as soft-bodied invertebrates

not preserved in the fossil deposit. The teeth of Barwickia

suggest it fed by a mechanism similar to those of denticulate

forms, like Fleurantia or holodipterids that have predominantly

denticle-covered plates with a few larger cusps set in rows

(Pridmore et al., 1994), possibly being a predator on either

smaller fishes or soft bodied invertebrates. Howidipterus , on

the other hand, had more typical dipnoan crushing tooth plates

suited to triturating food, potentially lycophytous and

psilophytous plant material that grew or fell into in the lake.

Lakes are often highly endemic, closed systems (Day et al.,

2009) and are analogous to islands in their isolated nature

(Danley and Kocher, 2001). Local speciations and adaptive

radiations are often influenced by past environmental factors

such as climate change (Day et al., 2009) and sea level changes

(Beheregaray et al., 2002, Bohlen et al., 2006). Another driving

factor is that of resource availability (Liem, 1974), the evolution

of variation has been demonstrated particularly for fish in low-

resource environments (Schluter, 1995; Roy et al., 2004).

There are many examples of sympatric lacustrine speciations

of fish (Humphries and Miller, 1981; Day et al., 2009) and

invertebrates such as gastropods (Glaubrecht and Kohler,

2004) and shrimp (von Rintelen et al., 2007). The best-known

example is that of the cichlids in the great lakes of East Africa

(Liem, 1974; Schliewen et al., 1994; Danley and Kocher, 2001;

Streelman et al., 2007). These cichlids underwent three major

bursts of cladogenesis; driven by habitat choice, competition

for food resources, and the third burst has been attributed to

sexual selection for male colouration. The secondary radiation

(trophic morphology) was most pronounced in the rock-

dwelling genera (Danley and Kocher, 2001).

This pattern of diversification of body form and trophic

structure is also seen in many other freshwater fishes including

the threespine stickleback Gasterosteus aculeatus (Cresko and

Baker, 1996), the Arctic charr Salvelinus alpinus (Snorrason et

al., 1989) and the Brook charr Salvelinus fontinalis (Dynes et

al., 1999). However the condition exhibited by the Mt. Howitt

fauna of divergent trophic morphologies with limited postcranial

differentiation is much less common. Fish with similar

postcranial morphologies are likely to have comparable

locomotive ability and occupy a common habitat. Trophic

specializations can diverge extremely rapidly (in “contemporary

time”) in response to different resource availability as seen in

the Arctic charr (Adams et al., 2003; Knudsen et al., 2007;

Michaud et al., 2008) and some cichlids (Liem, 1974; Streelman

et al., 2007). This indicates that the two Mt. Howitt species may

have only relatively recently diverged from a common ancestor

into two morphs with radically differing dentition, most likely

as a result of competition in a low -resource environment.

Acknowledgements

Thanks to Professor Jim Warren (Zoology Department, Monash

University) for allowing JLto initially work on the Mt. Howitt

material, and Mr. Ian Stewart for his assistance during the

preparation and casting of the specimens.The following people

are thanked for helpful discussion on the material: Professor

Ken Campbell, Dr. Dick Barwick and Dr. Peter Pridmore

(A.N.U. Geology Dept.), Dr. Gavin Young, (AGSO, Canberra),

Dr. Richard Cloutier (Universite de Sciences et Techniques,

Lille, France), Dr Ken McNamara (Cambridge University) and

Dr. Hans-Peter Schultze (University of Kansas, U.S.A.). Tim

Holland assisted in making latex peels of the Mt Howitt fishes.

This work was supported under ARC Grant DP 0772138.

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Memoirs of Museum Victoria 66: 203-213 (2009)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://museumvictoria.com.au/About/Books-and-Journals/Journals/Memoirs-of-Museum-Victoria

New asterinid species from Africa and Australia (Echinodermata: Asteroidea:

Asterinidae)

P. Mark O’Loughlin

Marine Science Department, Museum Victoria, GPO Box 666, Melbourne 3001, Victoria, Australia (pmo@bigpond.net.au)

Abstract O’Loughlin, P.M. 2009. New asterinid species from Africa and Australia (Echinodermata: Asteroidea: Asterinidae).

Memoirs of Museum Victoria 66: 203-213.

Three new species are described: Aquilonastra shirleyae sp. nov. from central Western Australia; Aster ina hoensonae

sp. nov. from southern Africa; Callopatiria cabrinovici sp. nov. from central east Africa. The variety Disasterina

leptalacantha var africana Mortensen is Tegulaster leptalacantha (H.L. Clark). Asterina gracilispina H.L. Clark is

reviewed systematically.

Keywords Australia, Africa, Echinodermata, Asteroidea, Asterinidae, Aquilonastra , Asterina, Callopatiria, Parvulastra, Tegulaster,

new species, generic reassignment

Introduction

This paper continues a series on family Asterinidae. Genus

Meridiastra O’Loughlin, 2002 was erected to accommodate

some southern Australian and Pacific species that had been

assigned to Asterina Nardo, 1834. O’Loughlin et al. (2002)

reviewed species Patiriella regularis (Verrill, 1867) using

morphological and molecular data, and a new species of

Patiriella Verrill, 1913 was described for New Zealand.

O’Loughlin et al. (2003) reviewed genus Patiriella Verrill, 1913

using morphological and molecular data, and three new species

were described for southern Australia. O’Loughlin and Waters

(2004) revised genera of Asterinidae based on morphological

systematics and a molecular phylogeny, and four new genera

were erected with O’Loughlin as author. O’Loughlin and Rowe

(2005) erected a new genus for the Indo-West Pacific region,

and described five new species. Most recently O’Loughlin and

Rowe (2006) undertook a morphological systematic revision of

genus Aquilonastra O’Loughlin, 2004, and described 13 new

species. In O’Loughlin and Waters (2004) and O’Loughlin and

Rowe (2006) genera Aquilonastra O’Loughlin, Asterina Nardo,

Callopatiria Verrill, 1913, Parvulastra O’Loughlin, 2004 and

Tegulaster Livingston, 1933 were discussed in detail, and are

the subjects of systematic work here.

This work is based on a continuing study of loan material

from the Australian Museum (Sydney; AM with registration

prefix J), Museum National d’Historie Naturelle (Paris; MNHN

with registration prefix EcAs), Museum of the Republic of

Central Africa (Brussels; MRAC), Museum Victoria (Melbourne;

NMV with registration prefix F), Natural History Museum

(London; NHM), South Africa Museum (Cape Town; S.A.M.

with registration prefix A for asteroids) and University of Florida

(UF). Recently included in this study is a Western Australian

Museum (Perth; WAM with registration prefix Z) loan of

specimens from voyage SS 10/05 by the RV Southern Surveyor

for Australia’s national science agency, the Commonwealth

Scientific and Industrial Research Organization (CSIRO) through

the Marine National Facility. New species of Aquilonastra

O’Loughlin from Western Australia, Asterina Nardo from South

Africa, and Callopatiria Verrill from east Africa are described.

Type specimens of species of Asterina and Disasterina Perrier,

1875 that are held in the South Africa Museum are reviewed.

Methods

Skeletal plates were cleared for observation using commercial

bleach. Terminology follows O’Loughlin and Waters (2004).

Photographs for figures 1, 3a, 3c, and 4 were taken by Leon

Altoff and Audrey Falconer using a Pentax K10D camera, with

an Olympus 80 mm f4 macro lens with bellows for large

specimens and Olympus 38 mm f2.8 macro lens with bellows for

small specimens. Photographs for figures 2, 3b, 3d, and 5 were

taken by Chris Rowley using a Leica MZ16 stereomicroscope,

DC300 Leica digital camera, and “Auto-Montage” software.

Figures were prepared by Caroline Harding.

Asterinidae Gray, 1840

Remarks. See O’Loughlin and Waters (2004).

Aquilonastra O’Loughlin, 2004

Remarks. See O’Loughlin and Waters (2004), O’Loughlin and

Rowe (2006).

204

P.M. O’Loughlin

Aquilonastra shirleyae sp. nov.

Figure la-f

Material examined. Holotype: Western Australia, Point Cloates,

22°50'55" S, 113°30'39" E to 22°51'29" S, 113°30'50" E, 100 m.

Southern Surveyor , SS10/2005 stn 135, M.P. Salotti and S. Slack-

Smith, 9 Dec 2005, WAM Z37278.

Paratype: Red Bluff, 24°02'37" S, 113°01'37" E to 24°02'50" S,

113°01'44" E, 100 m. Southern Surveyor, SS10/2005 stn 126, M.P

Salotti and S. Slack-Smith, 8 Dec 2005, WAM Z37279 (1, denuded).

Description. Small, stellate, R - 13 mm, r = 5 mm (holotype;

abnormal abactinal digitiform growth near disc), R = 8 mm, r =

4 mm (paratype; 2 rays regenerating); 5 sub-equal discrete

rays, subdigitiform, interradial margin deeply incurved; rays

tapering to narrow rounded distal end; rays flat actinally, high

domed elevation abactinally; body integument not evident;

single madreporite; not fissiparous; gonopores abactinal; glassy

convexities on plates; superambulacral and superactinal plates

present internally.

Abactinal: disc not discretely defined; rare proximal

doubly or singly papulate carinal plates on holotype, weakly

developed singly papulate carinal series on paratype, remaining

upper ray plates irregular in arrangement; non-carinal plates

crescentiform with single notch for papula; papular spaces

small, single papula per space; 3 longitudinal series of singly

papulate plates along each side of rays; rare secondary plates

except in disc area; large white opaque bi-valved pedicellariae

over papulae on upper sides of rays of holotype, less developed

on small paratype; spinelets glassy, conical or columnar, up to

about 0.15 mm long, tapered or splay -pointed distally, rugose,

in splayed clusters on plates, in 3 small clusters across rare

doubly papulate carinal plates, up to about 25 spinelets per

plate; ends of distal abactinal interradial splayed spinelets

rarely overlap ends of adjacent plate spinelets.

Margin: superomarginal plates about half size of

inferomarginal plates, both in regular series; up to about 16

spinelets on both superomarginal and inferomarginal plates,

thicker on inferomarginals.

Actinal: plates in longitudinal series, parallel to furrow;

complete series of adradial actinal plates and spines. Actinal

spines per plate: oral 8-10, suboral 7-9, ambulacral / furrow

7-8, subambulacral 7-8, proximal actinal 7-10, distal actinal,

7-10; oral spines digitiform, rugose; other actinal spines thin,

glassy, rugose, pointed distally.

Colour. Live (photo of paratype): abactinal pale mottled

with white, pale brown, pale orange; actinal white.

Preserved: white.

Distribution. Western Australia, Point Cloates, Red Bluff,

continental shelf, 22-24° S, 113° E, 100 m.

Etymology. Named for Shirley Slack-Smith of the Western

Australian Museum, with appreciation of her role in collecting

these specimens, and in recognition of four decades of dedicated

contribution to Australian marine mollusc research.

Remarks. The new species has the diagnostic characters of

genus Aquilonastra O’Loughlin as detailed in the emended

diagnosis by O’Loughlin and Rowe (2006): discrete rays,

interradial margin deeply incurved, stellate; high domed

abactinally, flat actinally; abactinal plates predominantly

irregular in arrangement on upper rays; longitudinal series

along sides of rays, not perpendicular to margin; predominantly

single papular notch per plate; predominantly single papula per

papular space; numerous elongate glassy spinelets on each

abactinal plate; superomarginal and inferomarginal plates in

regular series; suboral spines present; adradial actinal spines in

complete series; superambulacral and superactinal plates

present internally.

A key to the 24 species of Aquilonatra is provided by

O’Loughlin and Rowe (2006). Aquilonastra shirleyae sp. nov.

is close diagnostically to Aquilonastra rowleyi O’Loughlin

and Rowe, 2006 (Sodwana Bay, SE Africa) and Aquilonastra

watersi O’Loughlin and Rowe, 2006 (Arabian and Red Seas,

and western Indian Ocean). Aquilonastra shirleyae is

distinguished from Aquilonastra rowleyi (details in brackets)

by: size (up to R = 23 mm); subdigitiform rays (rays short,

wide at base; fig. 6i in O’Loughlin and Rowe, 2006);

pedicellariae squat (elongate; fig. 10c in O’Loughlin and Rowe,

2006); disc not discretely defined (disc clearly bordered; fig.

10b in O’Loughlin and Rowe, 2006); superomarginal plates

half size of inferomarginals (subequal); actinal interradial

spines 7-10 (3-4).

Aquilonastra shirleyae is distinguished from Aquilonastra

watersi (details in bracket) by: size (up to R = 19 mm); rays

that are thinner, with narrower base (rays short with wide base;

figs. 3j, lOe in O’Loughlin and Rowe, 2006); few singly and

doubly papulate carinal plates (up to 10 doubly papulate); up to

about 25 abactinal spinelets per plate (up to about 20);

pedicellariae conspicuous (inconspicuous; fig. 61 in O’Loughlin

and Rowe, 2006); up to about 16 superomarginal spinelets per

plate (up to about 7); up to 10 actinal spines per plate (up to 8);

mottled very pale white, pale brown, pale orange (mottled pale

brown, red-brown, grey-brown, blue-grey; off-white; fig. 3j in

O’Loughlin and Rowe, 2006).

O’Loughlin and Rowe (2006) reported most Aquilonastra

species from the shallow sub-littoral to about 50 m. A. batheri

(from Japan) was reported to 92 m, and A. cepheus (from

southern China to northern Australia) to 70 m. A. rosea (SW

Australia) was reported to 110 m, the only occurrence deeper

than A. shirleyae.

Asterina gracilispina H.L. Clark, 1923

Figure 2a-d

Asterina gracilispina H.L. Clark, 1923: 286-287, pi. 16 figs.

3-4.— A. M. Clark, 1974: 437 (part).— A. M. Clark and Courtman-

Stock, 1976: 77 (part).

Material examined. Holotype (dry): South Africa, East London, SW

of Cove Rock, 40 m, S.A.M. A6421.

Description. Small, subpentagonal, R = 6 mm, r = 4 mm; rays

5, discrete, wide at base, short, rounded distally; low convex

abactinally, flat actinally, sides not steep, margin acute; body

integument not evident; single inconspicuous madreporite; not

fissiparous; gonopores not seen; glassy convexities on plates;

presence or absence of superambulacral and superactinal plates

New asterinid species from Africa and Australia (Echinodermata: Asteroidea: Asterinidae)

205

Figure 1. Aquilonastra shirleyae sp. nov. (photos by L. Altoff and A. Falconer). Holotype, WAM Z37278 (R = 13 mm; all except b); paratype

WAM Z37279 (R = 8 mm; cleared; b only), a, abactinal surface (one ray dissected off; abnormal digitiform growth arrowed), b, disc not

discretely defined; few proximal carinal plates with single papula arrowed, c, abactinal pedicellariae (arrowed) and spinelets on upper side of ray.

d, section through cleared ray showing sup erambul acral plates (right arrow) and superactinal plates (left arrow), e, actinal surface with actinal

plates in longitudinal series, f, oral, suboral, furrow, subambulacral and proximal actinal spines.

206

P.M. O’Loughlin

Figure 2. Asterina gracilispina H.L. Clark, 1923 (photos by C. Rowley). Holotype, S.A.M. A22559 (R = 6 mm), a, abactinal surface, insert

showing spinelets. b, carinal series of doubly papulate plates and spinelets. c, actinal surface with actinal plates in longitudinal series, d, oral,

suboral, furrow, subambualcral and proximal actinal spines.

unknown (small type specimen not dissected); glassy

convexities on plates.

Abactinal: disc not distinctly bordered; plates imbricate,

projecting proximal edge frequently tabular, plates not

notched, slight proximal indentation for papula sometimes

present, papulae emerge from under projecting proximal

raised edge of plates; doubly papulate carinal series of plates

along most of upper ray, rare secondary plates; papulae large,

single per papular space, rarely 2; 8 longitudinal series of

papulae across mid ray; small subsacciform to conical, pointed

glassy spinelets, up to about 0.15 mm long, spread over plates,

up to 20 per plate.

Margin: superomarginal plates longitudinally elongate, in

regular series, up to about 14 spinelets spread over each plate,

subequal with abactinal and inferomarginal spinelets,

projecting inferomarginal plates with up to about 20 spinelets.

Actinal: interradial plates in longitudinal series, not

predominantly oblique; complete series of adradial actinal

plates and spines. Actinal spines per plate: oral 5 (2 long

proximal; gap to 3 short distally, increasing in length to

distalmost longest of 3); suboral 2 (webbed, long); furrow 4-3;

subambulacral 4-3; adradial actinal 3-5; actinal interradial up

to 7 mid ray, webbed transverse series, frequently 5-6; spines

subsacciform to conical.

Distribution. South Africa, East London, 40 m.

Remarks. For O’Loughlin and Waters (2004) I used a specimen

from Cape Agulhas, registered to the Natural History Museum

in London as Asterina gracilispina (NHM 1975.10.29.47), as

evidence for remarks on A. gracilispina. The subsequent

availability of the holotype of A. gracilispina for examination

in this work has made it possible for me to recognize that the

New asterinid species from Africa and Australia (Echinodermata: Asteroidea: Asterinidae)

207

Cape Agulhas specimen is not conspecific. The Cape Agulhas

specimen is similar in form, but is distinguished (R = 12 mm)

by: disc distinctly bordered; conspicuous madreporite; short,

blunt, digitiform to subgranular abactinal spinelets; 9 oral

spines in series tapering evenly from long to very short, spines

slightly swollen distal ly; up to 3 actinal spines mid ray. This

specimen is described below as a new species.

Mortensen (1933) referred “with considerable doubt” a

specimen (R = 10 mm) in the South Africa Museum from

False Bay (26 m) to A. gracilispina. His grounds for doubt

were: dorsal spinelets blunt; distinct madreporite; 2-3 stout

actinal interradial spines. These characters are consistent with

those of the Cape Agulhas specimen, described below as a

new species of Asterina.

A.M. Clark (1974) reported on six specimens from South

Africa determined as A. gracilispina, but her notes indicate to

me that there were two species. Most of the specimens were in

poor condition, but details of spine number for the Mossel Bay

specimen (R = 6 mm) are compatible with the holotype of A.

gracilispina. Notes that the Algoa Bay specimen (R = 10 mm)

had 3-5 actinal spines per plate and an inconspicuous

madreporite indicate that it is also probably A. gracilispina.

But notes of 9 oral spines and only 2-3 actinal spines for the

Cape Agulhas specimen (R = 12 mm; registered to the NHM)

confirm the observations discussed above that it is not

conspecific with A. gracilispina and it is the type for the new

species referred to above and described below.

H.L. Clark (1923) was uncertain about generic assignment

for this species, and chose Asterina. Currently there is

inadequate data to confirm or reassign. However, the atypical

arrangement and form of the oral spines is similar to that in

Parvulastra O’Loughlin, 2004. If superambulacral and

superactinal plates are present, then the species would be more

appropriately assigned to Parvulastra.

Asterina hoensonae sp. nov.

Figure 3a-d

Asterina gracilispina.— Mortensen, 1933: 255-256 (non A.

gracilispina).— O’Loughlin and Waters, 2004: 11, 15-16 (non A.

gracilispina).— A.M. Clark, 1974: 437 (partnon A. gracilispina).— A.M.

Clark and Courtman-Stock, 1976: 77 (part non A. gracilispina).

Material examined. Holotype (in alcohol; part dissected): South

Africa, Cape Agulhas, 34°S, 20°E, C. Griffiths (University of Cape

Town), NHM 1975.10.29.47.

Description. Small, subpentagonal, R - 12 mm, r = 9 mm; rays

5, discrete, wide at base, short, rounded distally; body

integument not evident; low convex abactinally, sides not steep,

margin acute, single conspicuous madreporite; gonopores not

detected; absence of pedicellariae; absence of superambulacral

and superactinal plates; margin supported by internal

contiguous projections of abactinal and actinal plates; glassy

convexities on plates.

Abactinal: plates imbricate, surface flat, not broken by

raised edges of plates, plates not notched, shallow concave

proximal indentations for papulae; doubly papulate carinal

series of plates along upper ray; papular spaces large, 0-2

secondary plates per space, 0-3 large papulae per space, 10

longitudinal series of papulae across mid ray; disc distinctly

bordered; spinelets digitiform to subgranuliform, short, blunt,

up to about 0.15 mm long, cover projecting abactinal plates, up

to 16 spinelets per plate.

Margin: superomarginal and inferomarginal plates

longitudinally elongate, in regular series, up to about 11

slightly conical subgranuliform spinelets spread over each

superomarginal plate, subequal with inferomarginal spinelets,

projecting inferomarginal plates with up to about 16 spinelets.

Actinal: interradial plates in variably longitudinal and

oblique series; complete series of adradial actinal plates and

spines. Actinal spines per plate: oral 9 (series tapering uniformly

from tall proximally to short distally, tallest spines slightly

swollen distally, smallest pointed distally); suboral 3; furrow 6;

subambulacral 4; adradial actinal 2-3; actinal interradial 2-3

mid ray, 3-5 distally; spines digitiform, webbed.

Distribution. South Africa, Cape Agulhas (E of Cape Town).

Etymology. Named in appreciation of the contribution to this

work by Elizabeth Hoenson of the South Africa Museum, who

went to considerable lengths to make available essential loans

for this work.

Remarks. The new species has the diagnostic characters of

genus Asterina Nardo as detailed in the emended diagnosis by

O’Loughlin and Rowe (2006): 5 discrete rays; not fissiparous;

disc distinctly bordered; carinal series of doubly papulate

plates; extensive papulate areas, numerous papulae and

secondary plates; abactinal spinelets digitiform to

subgranuliform; predominantly 2-3 digitiform actinal spines

per plate; lacking superambulacral and superactinal plates;

margin supported internally by contiguous projections of

abactinal and actinal plates.

Some characters distinguishing Asterina hoensonae sp.

nov. from Asterina gracilispina are listed under A. gracilispina

above. Asterina hoensonae is distinguished from most of the

remaining species of Asterina (A. gibbosa, A. ocellifera, A.

pancerii, A. phylactica and A. stellifera) by lacking

pedicellariae; and from A. fimbriata by having a distinctly

bordered disc.

I discuss this specimen under Asterina gracilispina above. It

is the specimen I wrongly accepted as being Asterina gracilispina

in O’Loughlin and Waters (2004). Another specimen (False Bay,

26 m; R - 10 mm) in the South Africa Museum, referred “with

considerable doubt” to A. gracilispina by Mortensen (1933), is

probably A. hoensonae (see above). Specimens from Algoa Bay

and Mossel Bay referred to A. gracilispina by A.M. Clark (1974)

and A.M. Clark and Courtman-Stock (1976) are probably A.

hoensonae (see above).

Callopatiria cabrinovici sp. nov.

Figure 4a-f

Material examined. Holotype: East Africa, Zanzibar, M. Angel, NHM

1965.6.1.743 (dry).

Paratypes: type series, NHM 1965.6.1.744 (1); East Africa,

Zanzibar, Mazizini, rocky outcrop, M.D. Richmond, 1993, NHM

2004.2833 (1, dry); Zanzibar, C. Crossland, NHM 1903.4.2.61-62 (2);

208

P.M. O’Loughlin

Figure 3. Asterina hoensonae sp. nov. (photos a, c by L. Altoff and A. Falconer; b, d by C. Rowley). Holotype, NHM 1975.10.29.47 (R = 12 mm),

a, abactinal surface with section of abactinal plates removed, b, abactinal subdigitiform to subgranuliform spinelets. c, abactinal view (plates

cleared, section of plates removed), showing doubly papulate carinal series of plates (left arrows) and absence of superambulacral plates (right

arrow), d, oral, suboral, furrow, subambulacral and proximal actinal spines.

Kenya, Watamu, Ras Ngomeni, W.F. Humphreys, 1 Apr 1969, NHM

1979.2.5.147 (1); Watamu, rock platform, sub-littoral, W.F. Humphreys,

9 Sep 1969, NHM 1979.2.5.146 (2).

Other material. Callopatiria cabrinovici sp. nov. Kenya, Shimoni,

J.D. Taylor, NHM 1973.10.4.48 (1); Zanzibar, Dr Kirk, NHM 68.3.6.13

(1); N Oman, Khesab Bay, coral reef, P. Cornelius, 30 Dec 1971, NHM

1972.4.10.57 (1).

Callopatiria granifera (Gray, 1847). South Africa, Western Cape

Province, NMV F98049 (1, donation to NMV by A. Thandar).

Description. Rays 5, discrete, subdigitiform, narrow base,

tapering to point or narrowly rounded end, broadly flat actinally,

acute angular margin, sides steep, close to perpendicular, high

convex abactinally; size large, rays unequal, up to R = 27-35

mm; integument evident; conspicuous single madreporite, not

fissiparous; lacking pedicellariae; complete series of internal

superambulacral plates; interradial margin supported by

numerous internal superactinal plates; superambulacral and

superactinal plates contiguous on actinal internal surface for

most of ray length; inner resinous brown lining to ray; gonopores

not observed.

Abactinal: plates thick, imbricate, angled; disc weakly

delineated in larger specimens, disc boundary typically 5

transversely elongate radials each with narrow band of up to

about 50 spinelets, 5 small interradials; lacking carinal series

of plates; longitudinal band (“field”) of primary and secondary

upper ray plates, irregular in arrangement and form, plates

with shallow concave indentation for single papula,

crescentiform, not sharply notched; papulate areas extensive,

papular spaces large, predominantly 1 large papula per space,

secondary plates numerous, frequently 1 per papular space; up

New asterinid species from Africa and Australia (Echinodermata: Asteroidea: Asterinidae)

209

Figure 4. Callopatiria cabrinovici sp. nov. (photos by L. Altoff and A. Falconer). Holotype, NHM 1965.6.1.743 (R = 27-35 mm), a, abactinal

view, insert showing spinelets. b, cleared proximal upper ray showing “field” of irregular singly papulate primary and few secondary plates, c,

lateral view of cleared ray showing longitudinal series of plates, and superomarginal and inferomarginal series of plates, d, cleared ray with

section of abactinal plates removed, showing superactinal plates (upper right arrow), superambulacral plates (lower right arrow) and ambulacral

plates (left arrow), e, actinal surface showing longitudinal series of actinal plates, f, oral, suboral, furrow, subambulacral and proximal actinal

spines.

210

P.M. O’Loughlin

to 5 longitudinal series of plates and papulae along each side

of rays, frequently in transverse series also; spinelets glassy,

columnar, thick to thin, splay-pointed, frequently widened

terminally, some narrowed terminally; up to about 40 spinelets

in narrow band 2-3 spinelets wide across projecting edge of

proximal ray plates, spinelets not in discrete tufts; proximal

ray spinelets up to about 0.4 mm long; distal interradius with

up to about 16 splayed spinelets per plate, ends rarely

overlapping those of adjacent plates; glassy convexities

sometimes evident around base of plates.

Margin: superomarginal and inferomarginal plates in

regular series; inferomarginals noticeably larger than

superomarginals; inferomarginals project only slightly;

alignment of superomarginals with inferomarginals frequently

broken by presence of additional superomarginal plate;

superomarginals with up to about 16 typical abactinal spinelets;

inferomarginals with up to about 26 spinelets, proximally

similar to superomarginal spinelets, distally stout digitiform.

Actinal: plates in longitudinal series parallel to ambulacrum;

adradial actinal plates in complete series. Actinal spines per

plate: oral 6, thick wedge-like proximally, slight gradation in

size from proximal to distal; suboral 6-9, big gradation in size;

furrow 6; subambulacral 6-8 in curved series, small at edges;

adradial actinal up to about 10; interradial up to about 10 in

clusters over crest of plate. Interradial spines thick digitiform

and thin, opaque, rugose, bluntly rounded to pointed distally.

Lacking actinal gonopores.

Distribution. East African coast, Zanzibar, Kenya, rock

substrate, shallow sub-littoral; Oman, coral reef.

Etymology. Named for Andrew Cabrinovic (Natural History

Museum, London), with appreciation of his gracious assistance

in facilitating loans to Museum Victoria for this and previous

studies.

Remarks. The generic diagnostic characters (in O’Loughlin

and Waters, 2004) of Callopatiria Verrill, 1913 that are shared

by the new species are: 5 discrete rays; rays long, stellate form;

sides of rays close to perpendicular above angular margin; rays

broadly flat actinally, high convex abactinally; irregular

arrangement of upper ray plates; abactinal plates crescentiform;

abactinal plates covered by narrow band of numerous elongate

glassy spinelets; absence of pedicellariae; secondary plates in

papular spaces; inferomarginals project only slightly; numerous

digitiform actinal spines per plate; presence of series of

superambulacral plates; numerous superactinal plates fill the

interradial angular margin of the rays, contiguous with

superambulacral plates for most of ray length; interior resinous

body lining.

Callopatiria cabrinovici sp. nov. differs from C. granifera

(Gray, 1847), the type species for Callopatiria (distribution

South Africa, from Namibia to Natal), by having: small papular

spaces with predominantly single papula and secondary plate

per papular space (not up to about 10); superomarginal plates

smaller than inferomarginals (not subequal); actinal plates in

longitudinal series (not oblique); more numerous suboral

spines per plate (more than up to 6); more numerous furrow

and subambulacral spines per plate (more than 4). C.

cabrinovici sp. nov. differs from C. formosa (Mortensen,

1933) (type locality False Bay, South Africa) by lacking the

enlarged, rounded, distal abactinal plates that are mostly bare

of spinelets; by having more numerous actinal spines (more

than 3-4 furrow and subambulacral spines, more than 7 actinal

interradial spines).

The specimen from Oman is in poor condition, but is

judged with some uncertainty to belong to the new species.

The other two specimens that are not nominated as types are

small, and do not show the diagnostic characters as distinctively

as the larger specimens, but they are also judged with some

uncertainty to belong to the new species.

Rowe and Richmond (2004) discussed the occurrence of

asterinid species from the western Indian Ocean. They recognized

two undescribed species from Rodrigues, and these have

subsequently been described by O’Loughlin and Rowe (2006)

as Aquilonastra conandae and Aquilonastra richmondi.

Reference was made by Rowe and Richmond (2004) to two

specimens from Zanzibar, thought by A.M. Clark to be “possibly

referable to Paranepanthia Fisher” (discussed fully in note 89

on pages 68-7 1 in Clark and Rowe 1971). Three NHM specimens

that were examined in this work (see above) are from Watamu

(Kenya) and had been determined as Paranepanthia. These

specimens, and others from Kenya and Zanzibar (see above)

that had been determined as Asterina burtoni, are referred here

to the new species Callopatiria cabrinovici.

Tegulaster leptalacantha (H.L. Clark, 1916)

Figure 5a-f

Asterina leptalacantha H.L. Clark, 1916: 57-58, pi. 18 figs. 3-4.

Disasterina leptalacantha.— Livingstone, 1933: 6, 8-10, pi. 3 figs.

5-6, pi. 4 figs. 1, 4.— H.L. Clark, 1946: 139. -A.M. Clark and Rowe,

1971: 38-39, 67,-Rowe and Gates, 1995: 36.

Disasterina leptalacantha var africana Mortensen, 1933:

259-260, pl.12 fig. 3.— A.M. Clark and Courtman-Stock, 1976: 78.

Tegulaster leptalacantha.— O’Loughlin and Waters, 2004: 13,

35-36.

Material examined. Disasterina leptalacantha. Holotype: NE

Australia, Queensland, Capricorn Group, Masthead I., littoral, Dec

1913, AMJ3082.

Disasterina leptalacantha var africana. Syntype: South Africa,

Natal, off Tugela River, 366 m, S.A.M. A22559 (1).

Other material. NE Queensland, AM J6097 (1); AM J12488 (1);

Indian Ocean, Mauritius I., Cape Malheureux, down to 24 m, UF

2499 (1); La Reunion I., Saint Leu, C. Conand, 22 Mar 2003, NMV

F109364 (3); NMV F109367 (4); S Madagascar, Mission Decary,

MNHN EcAsll856 (1); E South Africa, Sodwana Bay, 11 m, MRAC

1746 (1); 14 m, MRAC 1744 (2); Trafalgar Marine Reserve, 14 m,

MRAC 1745 (1).

Description. Thick integument body cover; rays 5, discrete,

medium length, wide base, tapered, blunt to narrowly rounded

distally, elevated, steep sides, acute thin margin, up to R = 24.5

mm; single madreporite; abactinal gonopores; superambulacral

and superactinal plates present.

Abactinal: plates predominantly bare, thick, frequently

with raised domes, generally closely imbricate, rarely spaced

creating non-plated areas, upper ray plates irregular in size,

form, arrangement, regular longitudinal series along sides of

New asterinid species from Africa and Australia (Echinodermata: Asteroidea: Asterinidae)

211

Figure 5. Tegulaster leptalacantha (H.L. Clark, 1916) (photos by C. Rowley), a, c, e, Disasterina leptalacantha var africana Mortensen, 1933,

syntype, S.A.M. A22559 (R = 16 mm), a, abactinal surface (not cleared), c, tufts of acicular inferomarginal spinelets. e, oral, suboral, furrow,

subambulacral and proximal actinal spines. Tegulaster leptalacantha, b, d, f, Sodwana Bay specimens, b, disc and proximal abactinal surface

(not cleared; MRAC 1746; R = 17 mm), d, tufts of acicular inferomarginal spinelets (MRAC 1744; R = 18 mm), f, oral, suboral, furrow,

subambulacral and proximal actinal spines (MRAC 1746; R = 17 mm)

212

P.M. O’Loughlin

rays, lower ray plates indented proximally for papula; lacking

distinct secondary plates; papulae large, single, irregular along

upper ray, sometimes doubly papulate carinal plates mid-ray,

up to 3 longitudinal series long each side of ray, up to 8

longitudinal series across mid-ray; disc variably bordered by 5

radial, 5 interradial plates; small subsacciform spinelets

sometimes present perianally, on disc, around madreporite;

long acicular spinelets sometimes on distal interradii; up to

few small granular or subsacciform or conical spinelets on

abactinal plates of small specimens (R = 5 mm); glassy

convexities on plates.

Margin: superomarginal plates of variable size and

regularity as series, bare except small specimens with single,

small, conical glassy spinelets; inferomarginal plates project,

plates sometimes with constricted waist, distal tuft of up to 10

and more glassy, long, acicular subsacciform spinelets, up to

1.5 mm long.

Actinal: interradial plates in oblique series. Actinal spines

per plate: oral 8-9, strongly tapered series; suboral 1

(sometimes small additional distal one); furrow 5, short;

subambulacral 1, long; adradial actinal 1; actinal interradial 1;

spines long, tapering to thin, sacciform.

Distribution. NE Australia, Mauritius I., La Reunion Is.,

Madagascar, E South Africa, 0-366 m.

Remarks. Mortensen (1933) observed only minor morphological

differences between the type and the two South Africa

specimens on which he based his variety. Amongst these

differences he noted that there were only five oral spines in the

type, but seven in the variety. The type has eight oral spines,

and eight and nine were observed on the specimen of the variety

from Natal. Mortensen (1933) erected the variety “mainly for

zoogeographical reasons” since the type locality for Tegulaster

leptalacantha is Queensland (NE Australia). In O’Loughlin

and Waters (2004) I determined material from Mauritius as T.

leptalacantha, and judged that the variety was not justified. I

confirm that opinion here. In O’Loughlin and Waters (2004) I

reassigned Disasterina leptalacantha to Tegulaster lAv'mgsonz,

1933. 1 confirm the morphological grounds for the reassignment

here. The diagnostic characteristics of Tegulaster leptalacantha

are: 5 discrete, high, tapered rays; thick integument evident

over body; predominantly bare abactinal plates; abactinal plates

frequently with rounded domes; actinal plates in oblique series;

single long sacciform spines on each actinal interradial plate;

inferomarginal plates with distal dense tufts of long, glassy,

acicular subsacciform spinelets; superambulacral and

superactinal plates present.

Tegulaster leptalacantha is distinguished from all other

species of Tegulaster by having a tuft of long acicular sacciform

spinelets on each inferomarginal plate; and is also distinguished

in particular from T. emburyi Livingstone, 1933 (type species

for Tegulaster, type locality Queensland) by having single

suboral and actinal interradial spines per plate, from T. alba

(H.L. Clark, 1938) (type locality Lord Howe I.) by having

abactinal gonopores, and from T. praesignis (Livingstone,

1933) (type locality Queensland) by having bare superomarginal

plates (see O’Loughlin and Waters, 2004 for these and other

distinguishing characters).

Acknowledgments

I am most grateful for the generous assistance of: Leon Altoff

and Audrey Lalconer (Marine Research Group of the Lield

Naturalists Club of Victoria, for some photography); Nadia

Ameziane and Marc Eleaume (MNHN, for facilitating loan

material); Andrew Cabrinovic (NHM, for facilitating loan

material); Chantal Conand (Emeritus Professor at La Reunion

University, for donation of specimens to NMV); Caroline

Harding (NMV, for preparation of figures); Karen Gowlett-

Holmes (colour photo of WAM paratype live); Liz Hoenson

(S.A.M., for facilitating loan material); Loisette Marsh and

Mark Salotti (WAM, for facilitating loan material); Gustav

Paulay (UP, for facilitating loan material); Prank Rowe (formerly

of the AM, for collaboration in earlier stages of this on-going

research); Chris Rowley (NMV, for montage photography);

Ahmed Thandar (University of KwaZulu-Natal, for donation of

a specimen to NMV and facilitating communication with the

S.A.M.); Didier VandenSpiegel MRAC, for facilitating loan

material). I am pleased to acknowledge the Coral Reef Research

Poundation, under contract to the US National Cancer Institute

contract no. N01-CM-27704, for sponsorship of fieldwork by

the University of Florida that yielded specimens studied in this

work. I am most appreciative of the critical review of this paper

by Prank Rowe.

References

Clark, A.M. 1974. Notes on some echinoderms from southern Africa.

Bulletin of the British Museum ( Natural History), Zoology 26:

421-487, figs. 1-16, this 1-3.

Clark, A.M. and Courtman- Stock, J. 1976. The echinoderms of

southern Africa. British Museum (Natural History) Publication

No. 776: London. 277 pp.

Clark, A.M. and Rowe, F.W.E. 1971. Monograph of shallow-water

Indo-West Pacific echinoderms. Pp. vii+238, 100 figs., 31 pis.

British Museum (Natural History): London.

Clark, H.L. 1916. Report on the sea-lilies, starfishes, brittle-stars and

sea-urchins obtained by the F.I.S. Endeavour on the coasts of

Queensland, New South Wales, Tasmania, Victoria, South

Australia and Western Australia. Biological Results of the Fishing

Experiments carried on by the F.I.S. “Endeavour”, 1909-1914

4(1): 1-123, 11 figs., 44 pis.

Clark, H.L. 1923. The echinoderm fauna of South Africa. Annals of

the South Africa Museum 13(7) 12: 221-435, pis 8-23.

Clark, H.L. 1938. Echinoderms from Australia. An account of collections

made in 1929 and 1932. Memoirs of the Museum of Comparative

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Gray, J.E. 1840. A synopsis of the genera and species of the class

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History (1) 6: 175-184; 275-290.

Gray, J.E. 1847. Description of some new genera and species of

Asteriadae. Proceedings of the Zoological Society of London 15:

72-83.

Livingstone, A.A. 1933. Some genera and species of the Asterinidae.

Records of the Australian Museum 19: 1-20, pis 1-5.

Mortensen, Th. 1933. Echinoderms of South Africa (Asteroidea and

Ophiuroidea). Videnskabelige Meddelelser fra Dansk

naturhistorisk Forening i K0benhavn 93: 215-400, pis 8-19.

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Nardo, J.D. 1834. De Asteriis. In Oken, L. Isis 7: 716-717.

O’Loughlin, P.M. 2002. New genus and species of southern Australian

and Pacific Asterinidae (Echinodermata, Asteroidea). Memoirs of

Museum Victoria 59(2): 277-296.

O'Loughlin, P.M., Waters, J.M. and Roy, M.S. 2002. Description of a

new species of Patiriella from New Zealand, and review of

Patiriella regularis (Echinodermata, Asteroidea) based on

morphological and molecular data. Journal of The Royal Society

of New Zealand 32(4): 697-711.

O’Loughlin, P.M., Waters, J.M. and Roy, M.S. 2003. A molecular and

morphological review of the asterinid, Patiriella gunnii (Gray)

(Echinodermata: Asteroidea). Memoirs of Museum Victoria 60(2):

181-195.

O’Loughlin, P.M. and Waters, J.M. 2004. A molecular and

morphological revision of genera of Asterinidae (Echinodermata:

Asteroidea). Memoirs of Museum Victoria 61(1): 1-40.

O’Loughlin, P.M. and Rowe, F.W.E. 2005. A new asterinid genus from

the Indo-West Pacific region, including five new species

(Echinodermata: Asteroidea: Asterinidae). Memoirs of Museum

Victoria 62(2): 181-189.

O’Loughlin, P.M. and Rowe, F.W.E. 2006. A systematic revision of

the asterinid genus Aquilonastra O’Loughlin, 2004

(Echinodermata: Asteroidea). Memoirs of Museum Victoria

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Perrier, E. 1875. Revision de la collection de stellerides du Museum

d’Histoire Naturelle de Paris. 384 pp. Paris. [Also in: Archives de

Zoologie Experimental et Generate (1876) 5: 1-104, 209-304]

Rowe, F.W.E. and Gates, J. 1995. Echinodermata. In Wells, A. (ed.).

Zoological Catalogue of Australia 33: i-xiii, 1-510. CSIRO:

Melbourne.

Rowe, F.W.E. and Richmond, M.D. 2004. A preliminary account of

the shallow-water echinoderms of Rodrigues, Mauritius, western

Indian Ocean. Journal of Natural History 38: 3273-3314.

Verrill, A.E. 1867 (republished 1870). Notes on the Radiata in the

Museum of Yale College, with descriptions of new genera and

species. Descriptions of new starfishes from New Zealand.

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1(2)1: 247-251. [Also in: Transactions and Proceedings of the

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Memoirs of Museum Victoria 66: 215-220 (2009)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://museumvictoria.com.au/About/Books-and-Journals/Journals/Memoirs-of-Museum-Victoria

Observations of reproductive strategies for some dendrochirotid holothuroids

(Echinodermata: Holothuroidea: Dendrochirotida)

P. Mark O’Loughlin 1 , John Eichler 2 , Leon Altoff 2 , Audrey Falconer 2 , Melanie Mackenzie 3 , Emily Whitfield 4 ,

Chris Rowley 5

1 Honorary Associate, Marine Science, Museum Victoria, Melbourne, GPO Box 666, Australia (e-mail: pmo@bigpond.

net.au)

2 Marine Research Group of The Field Naturalists Club of Victoria, 1 Gardenia Street, Blackburn 3130, Australia (email:

mailto:fncv@vicnet.net.au)

3 Research Associate, Marine Science, Museum Victoria

4 Volunteer, Marine Science, Museum Victoria

5 Marine Science, Museum Victoria (e-mail: crowley@museum.vic.gov.au)

Abstract O’Loughlin, P. M., Eichler, J., Altoff, L., Falconer, A., Mackenzie, M., Whitfield, E., Rowley, C. 2009. Observations of

reproductive strategies for some dendrochirotid holothuroids (Echinodermata: Holothuroidea: Dendrochirotida). Memoirs

of Museum Victoria 66: 215-220.

Some recently observed dendrochirotid holothuroid reproductive strategies are reported for the first time: fissiparity

by Cucuvitrum rowei O’Loughlin and O’Hara; probable intra-coelomic brood fissiparity by Staurothyone inconspicua

(Bell); intra-coelomic brood-protection by a species of Parathyonidium Heding; intra-coelomic brood auto-ingestion by

Neoamphicyclus materiae O’Loughlin; brood-protection in anterior interradial marsupia by Psolidiella mollis (Ludwig

and Heding). Some analysis is reported of marsupial brood-protection by Antarctic dendrochirotid holothuroids: the

“ Cucumaria georgiana group”; Echinopsolus acanthocola Gutt; Echinopsolus parvipes Massin; Microchoerus splendidus

Gutt; Psolidiella mollis (Ludwig and Heding); P solus charcoti Vaney. Confusion in the literature about brood-protection

by the Subantarctic dendrochirotid Cladodactyla crocea (Lesson) is clarified.

Keywords Echinodermata, Holothuroidea, Dendrochirotida, auto-ingestion, brood-protection, fissiparity, intra-coelomic, marsupium.

Introduction

Large collections of holothuroids from coastal southern

Australia and from Antarctica (Prydz Bay, the Bellingshausen

Sea, the Antarctic Peninsula and the South Atlantic), most

specimens held by Museum Victoria (NMV), have been

studied by Mark O’Loughlin. This has resulted in many of the

observations reported here. Fieldwork by John Eichler of the

Marine Research Group (MRG) in Victoria has resulted in the

recognition of fissiparity by a dendrochirotid species.

Laboratory research on specimens by NMV volunteers

Melanie Mackenzie and Emily Whitfield has contributed to

observations reported in this work. John Eichler (MRG; JE),

Leon Altoff and Audrey Falconer (MRG; LA), and Chris

Rowley (NMV; CR) have provided photographs.

Fissiparity by Cucuvitrum rowei O’Loughlin and O’Hara

John Eichler collected a live specimen of Cucuvitrum rowei

O’Loughlin and O’Hara, 1992 from Port Phillip Bay in SE

Australia on 20 April 2008 (NMV F157401; variably 19 to 30

mm long live). During subsequent days peristaltic-like body

contractions were observed and photographed (fig. la).

Between April 22 and 25 the specimen divided transversely

into two individuals (smaller 7 mm long). Peristaltic-like body

contractions continued in both post-fissiparity individuals.

Overnight on May 1 to 2 the larger individual divided

transversely again (fig. lb). Preservation and dissection of

these individuals revealed that the larger post-fissiparity oral

end individual had fully developed tentacles and calcareous

ring, but lacked internal organs. Detached internal organ

remnants were present in the smaller anal end individuals that

lacked tentacles and ring. One apparent purpose of the

peristaltic movements was to push the internal organs to the

anal end of the coelom to provide a nutrient source for the

subsequent regeneration of tentacles, ring and internal

organs.

On 3 August 2008, seven smaller specimens were collected

from Port Phillip Bay (NMV F161549; up to 12 mm long).

After nine days none had undergone fissiparity. On 30 August

2008, five specimens were collected from Port Phillip Bay

(NMV F161500; up to 16 mm long). Overnight on September

216

P.M. O’Loughlin, J. Eichler, L. Altoff, A. Falconer, M. Mackenzie, E. Whitfield, C. Rowley

Figure 1. Fissiparity by Cucuvitrum rowei O’Loughlin and O’Hara, a, peristaltic contractions in live specimen (F157401; about 25 mm long;

photo by JE). b, 3 live individuals resulting from fissiparity (F157401; JE). c, live specimen showing regenerating anal end (F157419; 4 mm long;

LA), d, preserved specimens showing evidence of fissiparity, with fully developed tentacles and ring and lacking internal soft organs (upper),

with developing tentacles and ring (lower) (F161501; LA).

6 to 7 one of the larger individuals divided transversely. During

his fieldwork John Eichler frequently noticed individuals in

close proximity on the undersurface of rocks. This clustering

may be a consequence of fissiparity.

Leon Altoff and Audrey Falconer photographed a live

specimen in the field that showed regeneration of the anal end

(fig. lc; 4 mm long; NMV F157419). Dissection by Emily

Whitfield of a large collection of NMV preserved specimens

of Cucuvitrum rowei revealed rare individuals that showed

evidence of fissiparity. Post-fissiparity oral ends lacked internal

organs but had withdrawn fully developed tentacles and

calcareous ring (fig. Id; NMV F161501); and post-fissiparity

anal ends showed a reduced developing calcareous ring and

small tentacles (fig. Id; NMV F161501), or lacked a calcareous

ring and tentacles.

O’Loughlin (1991, 1994) reported fissiparity by similar

mid-body transverse constriction and division in the

dendrochirotid Squamocnus aureoruber O’Loughlin and

O’Hara, 1992 from the rocky shallows of southern Australia.

This is the first record of fissiparity by the dendrochirotid

Cucuvitrum rowei , and the first record of peristaltic body

movements in a dendrochirotid holothuroid.

Coelomic fissiparity by Staurothyone inconspicua (Bell)

Brood juveniles (45) removed from the coelom of a female

Staurothyone inconspicua (Bell, 1887) from Opossum Bay in

SE Tasmania are of different sizes and many show mid-body

constrictions (fig. 2a; NMV F58613). One coelomic juvenile

from another specimen from Opossum Bay shows a deep mid-

Observations of reproductive strategies for some dendrochirotid holothuroids (Echinodermata: Holothuroidea: Dendrochirotida)

217

body constriction (fig. 2b; NMV F58456). For this seasonally

reproducing and coelomic brood-protecting species (see

Materia et al. 1991), and with an assumption of a single

fertilization event, these observations suggest intra-coelomic

brood fissiparity and cloning. However, dissection of coelomic

juveniles has to date failed to reveal confirming evidence of a

coelomic juvenile that lacks a calcareous ring.

Balser (2004) reported on cloning by larvae of echinoderms,

including holothuroids. The evidence here indicates probable

intra-coelomic cloning by a holothuroid.

Coelomic brood protection by a species of Parathyonidium

Heding

A female specimen of an undescribed species of Parathyonidum

Heding, 1954 from Eastern Antarctica (17 mm long, preserved;

NMV F84983) has 39 differentiating coelomic juveniles of

uniform size (2-3 mm long) (fig. 2c). Materia et al. (1991)

reported coelomic brood-protection for Neoamphicyclus

materiae O’Loughlin, 2007 (as Neoamphicyclus lividus

Hickman, 1962) and Staurothyone inconspicua (Bell, 1887)

from SE Australia. This is a third case of intra-coelomic

brood-protection by a dendrochirotid holothuroid species.

Brood auto- ingestion by Neoamphicyclus materiae

O’Loughlin

A 20 mm long female Neoamphicyclus materiae O’Loughlin,

2007 from Kitty Miller Bay on the coast of Victoria, collected

on 25 October 1987, has 528 small coelomic brood juveniles

(fig. 2d; NMV F58592). A 30 mm long female from Cape

Otway on the coast of Victoria, collected on 29 December

1985, has one large coelomic brood juvenile (fig. 2d; NMV

F76371). Typically this species has small coelomic brood

juveniles in October (see Materia et al. 1991), and one or a few

large coelomic juveniles are present in December or January

(NMV F58606; F58720; fig. 2e). These observations indicate

that intra-coelomic brood auto-ingestion occurs in

Neoamphicyclus materiae.

Byrne (1996) reported intragonadal cannibalism in the

small simultaneous hermaphrodite asterinids Parvulastra

vivipara (Dartnall, 1969) and Parvulastra parvivipara

(Keough and Dartnall, 1978) from southern Australia. This is

the first report of intra-coelomic cannibalism in a holothuroid

species.

Brood protection by Psolidiella mollis (Ludwig and

Heding)

Psolidiella mollis (Ludwig & Heding, 1935) is an additional

species of Antarctic dendrochirotid holothuroid that brood-

protects in marsupia (see Table 1). Males have a long genital

papilla (NMV F157414), and females have up to five anterior

interradial internal marsupia (fig. 3a; NMV F104865). One

female from Bouvet Island (fig. 3b; NMV F104882) has 46 and

60 differentiated embryos (3-4 mm long) in each of two

marsupia, and 52, 53, and 60 undifferentiated eggs or embryos

(1.3 mm long) in each of three marsupia, evidence of two

fertilization events.

Table 1. Antarctic species with 5 anterior interradial marsupia.

“ Cucumaria georgiana (Lampert, 1886) group”

Cucumaria acuta Massin, 1992

Cucumaria analis Vaney, 1908

Cucumaria aspera Vaney, 1908

Cucumaria attenuata Vaney, 1906

Cucumaria georgiana (Lampert, 1886)

Cucumaria joubini Vaney, 1914

Cucumaria lateralis Vaney, 1906

Cucumaria perfida Vaney, 1908

Cucumaria periprocta Vaney, 1908

Cucumaria secunda Vaney, 1908

Cucumaria vaneyi Cherbonnier, 1949

First reported: Vaney, 1925; Ekman, 1925

Echinopsolus acanthocola Gutt, 1990

First reported: De Ridder et al., 2005

Echinopsolus parvipes Massin 1992

First reported: Heterier et al., 2004

Microchoerus splendidus Gutt, 1990

First reported: O’Loughlin, 1994

Psolidiella mollis (Ludwig and Heding, 1935)

First reported: this work

Psolus charcoti Vaney, 1914

First reported: O’Loughlin, 2001

Broe4 protection in anterior interradial marsupia

A “ Cucumaria georgiana (Lampert, 1886) group” was created

by Gutt (1988) and adopted by Massin (1992) because of the

systematic confusion resulting from many Antarctic cucumariid

species having similar morphological characters. O’Loughlin

(in O’Loughlin et al. 2009) included Cucumaria aspera Vaney

in this group, but removed Cucumaria armata Vaney. A revised

list is included within Table 1. All species within this “Group”,

and the five other species in Table 1, have the same brood-

protecting habit that is unique to Antarctic holothuroids: males

have a long genital papilla between the dorsal tentacle pair

(fig. 3c), and do not have marsupia; females have a short

genital papilla between the dorsal tentacle pair (fig. 3c), and

have up to five anterior interradial marsupia with external

pores (fig. 3d). Females in all the species in Table 1 have up to

five marsupia, but all five may not be present in an individual.

A “ Cucumaria georgiana group” female specimen from

Casey Station in Eastern Antarctica, collected on 3 November

1997 (fig. 3e; NMV F85853), has 41 and 66 undifferentiated

eggs or embryos (1.3 mm long) in each of two marsupia, and

25, 25 and 56 differentiated embryos (3-4 mm long) in each of

three marsupia, evidence of two fertilization events.

A female specimen of Psolus charcoti Vaney, 1914 from

Prydz Bay, collected on 21 February 1987 (fig. 3f; NMV

F86009) has one and seven undifferentiated eggs or embryos

(1.7 to 1.8 mm long) in each of two marsupia; seven

differentiated embryos (3.5 mm long) in one marsupium; and

one and four marsupial juveniles (4.0 mm long) in each of two

marsupia. The data are evidence of three fertilization events.

Two female specimens from amongst many (up to 35 mm

long, tentacles included) belonging to the “ Cucumaria

218

P.M. O’Loughlin, J. Eichler, L. Altoff, A. Falconer, M. Mackenzie, E. Whitfield, C. Rowley

Figure 2. a, coelomic brood juveniles from a female Staurothyone inconspicua (Bell) showing mid-body constrictions and variable sizes (F58613;

photo by CR). b, coelomic juvenile of S. inconspicua showing a deep mid-body constriction (F58456; CR). c, coelomic juveniles from a female

Parathyonidium Heding species (F84983; CR). d, many small (F58592, late October; CR) and one large (F76371, late December; CR) coelomic

brood juveniles of Neoamphicyclus materiae O’Loughlin. e, single large coelomic juvenile of N. materiae (F58606; late December; LA), f,

invaginated body wall marsupium of Cladodactyla crocea (Lesson) with single remaining egg/embryo (F106967; 27 mm long; LA).

Observations of reproductive strategies for some dendrochirotid holothuroids (Echinodermata: Holothuroidea: Dendrochirotida)

219

Figure 3. a, transverse section through anterior marsupia of Psolidiella mollis (Ludwig and Heding) (F104865; diameter about 15 mm; photo by

LA), b, female P. mollis with differentiated embryos (3-4 mm long) from each of 2 marsupia (left), and undifferentiated eggs or embryos (1.3

mm long) from each of 3 marsupia (right) (F104882; specimen 45 mm long; CR). c, long male genital papilla of Cucumaria acuta Massin

(F160042; left; CR); short female genital papilla of C. acuta (F160020; right; CR). d, internal marsupium of C. acuta (F160038; specimen 37 mm

long; CR). e, C. georgina group species with differentiated embryos (3-4 mm long) from each of 3 marsupia (left), and undifferentiated eggs or

embryos (1.3 mm long) from each of 2 marsupia (right) (F85853; 33 mm long; CR). f, Psolus charcoti Vaney with 1 and 7 undifferentiated eggs

or embryos (1.7 to 1.8 mm long) from each of 2 marsupia; 7 differentiated embryos (3.5 mm long) from 1 marsupium (left); and 4 and 1 marsupial

juveniles (4.0 mm long) from each of 2 marsupia (middle and right) (F86009; specimen 55 mm long; LA).

220

P.M. O’Loughlin, J. Eichler, L. Altoff, A. Falconer, M. Mackenzie, E. Whitfield, C. Rowley

georgiana group”, from Peter I Island in the Bellingshausen

Sea taken at 124 m (Spanish BENTART 2003 collection; stn

A5, 7-10 February), were examined. The gonad tubules had

eggs 1.5 mm long. Marsupial juveniles were uniform in size

(2-3 mm long). Numbers of juveniles in the marsupia were: 5,

0, 0, 0, 0 and 15, 13, 11, 0, 0. The data indicate two fertilization

events.

Two female specimens from amongst many (up to 27 mm

long, tentacles included) belonging to the “Cucumaria

georgiana group”, from Low Island near the Antarctic

Peninsula taken at 86 m (Spanish BENTART 2006 collection;

stn Low 45, 7-10 February), were examined. The gonad

tubules lacked eggs. Marsupial juveniles were uniform in size

(2-3 mm long). Numbers of juveniles in the marsupia were:

63, 39, 27, 0, 0 and 30, 27, 23, 17, 9. These data indicate one

fertilization event.

The differing gonad and marsupial data for the same time

of the year, and the significantly different sizes of the specimens

from very large samples, are evidence of two different species

within the specimens of the “ Cucumaria georgiana group”

from Peter I Island and Low Island.

Heterier et al. (2004) reported that preliminary observations

indicated that Echinopsolus acanthocola Gutt, 1990 and

Echinopsolus parvipes Massin, 1992 “could brood”. De Ridder

et al. (2005) confirmed internal brood-protection for

Echinopsolus acanthocola. Brood-protection in anterior

interradial marsupia is confirmed here for both species of

Echinopsolus.

Marsupium in Cladodactyla crocea (Lesson)

For the subantarctic dendrochirotid species Cladodactyla

crocea (Lesson, 1830), Wyville-Thomson (1878) reported “no

special marsupium”; Bell (1908) reported “brood pouches”;

Vaney (1925) reported “two brood pouches”; and Ekman

(1925) reported “sexual maturity comes late” and “no brood

pouches”. Museum Victoria specimens of Cladodactyla

crocea from the Burdwood Bank (NMV F160031) and

Falkland Islands (NMV F106967) that were studied in this

work have a single dorsal longitudinal invaginated body wall

marsupium (fig. 2f). A small and presumably young specimen

from the Burdwood Bank, collected on 25 January 2004

(NMV F160031), is 8 mm long, has eggs in gonad tubules, and

a dorsal invaginated marsupium with eggs/embryos.

Presumably the authors referred to above were observing

specimens that were not Cladodactyla crocea.

Acknowledgments

We are grateful to: Ben Boonen for the preparation of the

figures; Susie Lockhart for the donation to Museum Victoria

of AMLR R/V Yuzhmorgeologiya (2003) and Icefish (2004)

holothuroid specimens from the southern Atlantic that were

studied in this work; Eugenia Manjon-Cabeza for the

opportunity to study the Spanish BENTART 2003 and 2006

R/V Hesperides collections held by the University of Malaga;

Museum Victoria for the use of the facilities of the Marine

Science Department, and the opportunity to undertake

research on the holothuroid collections.

References

Balser, E.J. 2004. And then there were more: cloning by larvae of

echinoderms. Pp. 3-9, 4 figs in: Heinzeller and Nebelsick (eds).

Echinoderms: Miinchen; Proceedings of the Eleventh

International Echinoderm Conference, Munich, Germany, 6-10

October, 2003. Balkema: London.

Bell, F.J. 1908. Echinoderma. Pp. 1-16, 5 pis in: National Antarctic

Expedition 1901-1904, Natural History, 4 Zoology. British

Museum of Natural History: London.

Byrne, M. 1996. Viviparity and intragonadal cannibalism in the

diminutive sea stars Patiriella vivipara and P. parvivipara (family

Asterinidae). Marine Biology 125: 551-567.

De Ridder, C., David, B., Heterier, V. and Massin, C. 2005. A new case

of brooding in an Antarctic holothuroid. Page 127 in: Evolution

and Biodiversity in Antarctica; Abstract book; IXth International

Antarctic Biology Symposium, Curitiba, Brazil, 20-23 July, 2005.

Ekman, S. 1925. Holothurien. Further zoological results of the

Swedish Antarctic Expedition 1901-1903 1(6): 1-194.

Gutt, J. 1988. Zur Verbreitung und Okologie der Seegurken

(Holothuroidea, Echinodermata) im Weddellmeer (Antarktis).

Berichte zur Polarforschung 41: 1-87.

Heterier, V., De Ridder, C., David, B. and Rigaud T. 2004. Comparative

biodiversity of ectosymbionts in two Antarctic cidaroid echinoids,

Ctenocidaris spinosa and Rhynchocidaris triplopora. Pp. 201-205,

4 tables in: Heinzeller and Nebelsick (eds). Echinoderms: Miinchen;

Proceedings of the Eleventh International Echinoderm Conference,

Munich, Germany, 6-10 October, 2003. Balkema: London.

Massin, Cl. 1992. Three new species of Dendrochirotida

(Holothuroidea, Echinodermata) from the Weddell Sea

(Antarctica). Bulletin de Vlnstitut Royal des Sciences Naturelles

de Belgique, Biologie 62: 179-191.

Materia, C.J., Monagle, J.F. and O’Loughlin, P.M. 1991. Seasonal

coelomic brooding in southern Australian cucumariids

(Echinodermata, Holothurioidea). Pp. 301-307, 5 figs, 5 tables in:

Yanagisawa, T., Yasumasu, I., Oguro, C., Suzuki, N. and

Motokawa, T. (eds). Biology of Echinodermata; Proceedings of

the Seventh International Echinoderm Conference, Atami, 9-14

September, 1990. Balkema: Rotterdam.

O’Loughlin, P.M. 1991. Brooding and fission in shallow water

echinoderms of southern Australia. Pp. 223-228, 5 figs, 1 table in:

Yanagisawa, T., Yasumasu, I., Oguro, C., Suzuki, N. and

Motokawa, T. (eds). Biology of Echinodermata; Proceedings of

the Seventh International Echinoderm Conference, Atami, 9-14

September, 1990. Balkema: Rotterdam.

O’Loughlin, P.M. 1994. Brood-protecting and fissiparous cucumariids

(Echinodermata, Holothurioidea). Pp. 539-547, 1 table, 6 figs in:

David, B., Guille, A., Feral, J-P. and Roux, M. (eds). Echinoderms

through Time; Proceedings of the Eighth International

Echinoderm Conference, Dijon, France, 6-10 September, 1993.

Balkema: Rotterdam.

O’Loughlin, P.M. 2001. The occurrence and role of a digitate genital

papilla in holothurian reproduction. Pp. 363-368 in: M. Barker

(ed.). Echinoderms 2000; Proceedings of the Tenth International

Conference, Dunedin. Swets and Zeitlinger: Lisse.

O’Loughlin, P.M., Manjon-Cabeza, M.E. and Ruiz, F.M. 2009.

Antarctic holothuroids from the Bellingshausen Sea, with

descriptions of new species (Echinodermata: Holothuroidea).

Zootaxa 2016: 1-16.

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Station Zoologique de Wimereux 9: 254-274.

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of propagation of certain echinoderms of the Southern Sea.

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Memoirs of Museum Victoria 66: 221-287 (2009)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://museumvictoria.com.au/About/Books-and-Journals/Journals/Memoirs-of-Museum-Victoria

Australian Axiidae (Crustacea: Decapoda: Axiidea)

Gary C. B. Poore 1 and David J. Collins

Museum Victoria, GPO Box 666, Melbourne, Vic. 3001, Australia ( 1 gpoore@museum.vic.gov.au)

Abstract Poore, G.C.B. and Collins, D. J. 2009. Australian Axiidae (Crustacea: Decapoda: Axiidea). Memoirs of Museum Victoria

66: 221-287.

Three new genera and nine new species of Axiidae are described from shelf and continental margins of Australia:

Acanthaxius garawa sp. nov., Acanthaxius gathaagudu sp. nov., Acanthaxius ningaloo sp. nov., Australocaris pinjarup

gen. and sp. nov., Calastacus myalup sp. nov., Eiconaxius mallacoota sp. nov., Pilbaraxius kariyarra gen. and sp. nov.,

Platyaxius bardi sp. nov.; Michelaxiopsis gen. nov. is erected for Axiopsis australiensis De Man, 1925 and Michelaxiopsis

nauo sp. nov. New records of Indo-West Pacific species for Australia are: Acanthaxius clevai Ngoc-Ho, 2006, Allaxius

clypeatus (De Man, 1888), Axiopsis serratifrons A. Milne-Edwards, 1873, Axiopsis tsushimaensis Sakai, 1992,

Bouvieraxius keiensis Sakai, 1992 and Planaxius brevifrons Komai and Tachikawa, 2008. New distributional records are

given for species previously recorded from Australia: Calaxius acutirostris Sakai and de Saint Laurent, 1989, Dorphinaxius

kermadecensis (Chilton, 1911), and Paraxiopsis pumilus (Sakai, 1994). Ten genera are rediagnosed. Undescribed species

of Axius, Axiopsis and Ambiaxius are mentioned. Photomicrographs of maxillae 1 and 2 and maxillipeds 1 and 2 of 1 1

species are appended. We could find very little of taxonomic value at the genus or species level in these appendages and

question the value of illustrating them in future. A key is provided for the identification of all 30 Australian species.

Keywords Crustacea; Decapoda; Axiidea; Thalassinidea; Axiidae; Acanthaxius ; Allaxius ; Ambiaxius ; Australocaris ; Axiopsis ;

Bouvieraxius ; Calastacus ; Calaxius ; Dorphinaxius ; Eiconaxius ; Michelaxiopsis ; Oxyrhynchaxius\ Pilbaraxius ;

Paraxiopsis ; Planaxius ; Platyaxius ; Scytoleptus\ taxonomy; new genus; new species

Introduction

The Axiidae, a family of burrowing lobsters, have traditionally

been placed in the decapod infraorder Thalassinidea (e.g.,

Poore, 1994) but more recent molecular and morphological

evidence suggests that Thalassinidea is not a monophyletic

taxon. Instead, this family falls within a smaller group, Axiidea

de Saint Laurent, 1973 (Robles et al., 2009).

The Australian axiid fauna comprises 18 recognised

species (Davie, 2002; Poore, 2008). Here we diagnose three

new genera, add nine new species, record six Indo-West Pacific

species for the first time from Australia, reidentify or

synonymise material of three species and provide additional

records for three species previously recorded from Australia.

The taxonomy of some other known species is reassessed and

updated and in the process it was necessary to redescribe some

species and rediagnose ten other genera. We take the

opportunity to list and provide a key to all 30 known Australian

Axiidae and comment on biogeography.

Photomicrographs of maxillae 1 and 2 and maxillipeds 1

and 2 of 11 species were prepared using a Leica DM5000B

microscope and Leica DC500 camera after clearing in lactic

acid and staining with chlorozol black. Between 15 and 25

images were taken in different planes and merged using

Automontage® software. Mandibles were also dissected and

examined stereoscopically. We could find very little of

taxonomic value at the genus or species level in these

appendages and do not present descriptions in words. The only

exceptional morphology is the absence in Acanthaxius clevai

Ngoc-Ho, 2006 of a posterior long seta on the epipod of

maxilla 2 (scaphognathite). Mouthparts of axiids have been

illustrated with or without the complex of setae that ornament

them by many authors. The level of detail has varied

considerably probably largely because of the difficulty of

rendering such complex setose structures as line drawings. For

this reason comparison between species has been rarely

attempted and species have never been differentiated using

these characters. In spite of the detail made possible by our

photographic tools, we are still unable to discover useful

characters. In our view, our own and earlier efforts have not

been informative and we question the value of illustrating

them in future.

Much of the new material arose from recent sampling

(cruises SS10-2005 and SS05-2007) by CSIRO Marine and

Atmospheric Research (CMAR) and Museum Victoria along

the Western Australian continental margin (Poore et al., 2008).

This collection is supplemented by material from the Northern

Territory, tropical Queensland, the south-eastern Australian

slope, and South Australia.

222

G.C.B. Poore and D.J. Collins

Material is deposited in the Australian Museum, Sydney

(AM), Museum Victoria, Melbourne (NMV), South Australian

Museum, Adelaide (SAM), Museum and Art Gallery of the

Northern Territory, Darwin (NTM) and Natural History

Museum, London (BMNH). Measurements are of carapace

length including rostrum (cl.) and total length (tl.). Most

illustrations were made by the second author and prepared for

publication using Adobe Illustrator and many of the

recommendations of Coleman (2003).

Family Axiidae Huxley, 1879

Restricted synonymy.

Axiidae Huxley, 1879: 785.— Sakai and de Saint Laurent, 1989:

4-5.— Poore, 1994: 96.

Calocarididae Ortmann, 1891: 47.— Kensley, 1989: 960.— Poore,

1994: 98

Eiconaxiidae Sakai and Ohta, 2005: 69.

Diagnosis. Carapace more or less laterally compressed,

moderately to well calcified; cervical groove distinct at least

dorsally; lineathalassinica absent; posterior margin of carapace

with lateral lobes interacting with abdominal somite 1. Rostrum

present, acute. Pleuron of abdominal somite 1 more or less

produced. Eyestalks cylindrical. Antenna 1 with article 3 about

as long as article 2. Antenna 2 with scale-like or well produced

scaphocerite. Maxilla 2 scaphognathite with a posterior whip

(rarely without). Pereopods 1 chelate, equal or unequal;

pereopod 2 chelate; pereopod 3 simple; pereopod 4 simple or

subchelate; pereopod 5 chelate. Pleopod 1 present or absent.

Pleopod 2 similar in dimensions to pleopods 3-5, biramous;

pleopods 3-5 endopodal rami elongated, not laterally expanded,

with or without appendix interna. Uropodal rami lamellate,

endopod more or less oval.

Remarks. Axiid lobsters are recognised by the combination of

a prominent rostrum ending in an acute tip, similar laminar

rami on pleopods 2-5, and oval uropodal endopod. They may

be confused with Strahlaxiidae which have a rostrum with an

apical notch and a triangular uropodal endopod. The family

Calocarididae was revived and defined by Kensley (1989) and

included in the key to thalassinidean families by Poore (1994).

Poore’s key to genera included six genera and Poore (2008)

listed two more. Kensley (1989) recognised the family on its

being hermaphroditic, having unpigmented eyes and highly

modified pleopod 2; in his view, axiids are gonochoristic (that

is, the sexes are separate with gonopores on either coxae 3

[female] or coxae 5 [male] and pleopods 1 and 2 usually sexually

dimorphic). In Kensley’s cladogram (1989: fig. 1) four

calocaridid genera form a clade separated from a paraphyletic

clade of three representative “axiid” genera. Tsang et al.’s

(2008) molecular analysis of relationships between two

calocaridid species ( Calastacus crosnieri and Paracalocaris

sagamiensis ) and three axiids found the calocaridids were

sister taxa and the axiids paraphyletic. Robles et al. (2009) used

eight species in another molecular study and found that another

calocarid, Calocaris caribbaeus, was embedded within the

seven species of Axiidae. In their analysis the species labelled

as belonging to the calocaridid genus, Calaxiopis, is here

reidentified as an axiid, Pilbaraxius kariyarra sp. nov. Cladistic

support for the family is currently weak in spite of its strong

morphological unity. The problem lies more with the apparent

paraphyly of Axiidae than with Calocarididae itself. Similarly,

Sakai and Ohta (2005) isolated Eiconaxius in its own family,

Eiconaxiidae, on the basis of rounded spinose dactyli on

pereopods 3-5. This supposed unique feature defines the genus

while nothing except its absence defines remaining axiids.

However, this characteristic is shown in part in another genus

(see discussion of Platyaxius below), weakening support for

this family. We prefer to synonymise Calocarididae and

Eiconaxiidae with Axiidae.

We note a structure hitherto unreported in many of the

new species described below. Attached to the anterolateral

margin of sternite 8, at the base of the socket of pereopod 5, is

an articulating flap, oval-semicircular in shape with a setose

margin (e.g. figs. 7d, e). In other species the structure is reduced

to a setose ridge. Its homology is unclear but it appears to be a

sternal scerite that may act as a valve at the posterior of the

branchial cavity.

Key to Australian species of Axiidae

This key is designed to be as practical as possible and

concentrates on characters most easily determined from what

can sometimes be fragmentary specimens. It does not reflect

relationship nor necessarily use generic characters.

1. Pleopod 2 endopod with dominating terminal appendix

masculina and digitiform appendix interna; pleopods 3-5

rami extremely thin; eyes without pigment; usually

hermaphroditic (with gonopores on both coxae 3 and 5,

pleopods 1 and 2 not sexually dimorphic) 2

- Pleopods 2-5 endopods laminar, similar; pleopod 2 with

appendices masculina and interna (if present in male)

linear and attached mesially; eyes usually pigmented;

gonochoristic (sexes separate, gonopores on either coxae

3 [female] or coxae 5 [male]; pleopods 1 and 2 sexually

dimorphic) 3

2. Rostrum directed upwards, about half length of postorbital

carapace length; pleopod 2 article 2 elongate -triangular,

mesially curved Ambiaxius franklinae Sakai, 1994

- Rostrum directed anteriorly, about quarter length of

postorbital carapace length; pleopod 2 article 2 linear,

2-segmented Calastacus myalup sp. nov.

3. Uropodal exopod without transverse suture; telson

tapering to rounded apex 4

- Uropodal exopod with transverse suture; telson more or

less rectangular, apex truncate or rounded 8

4. Rostrum significantly depressed below level of gastric

region of carapace; gastric region almost vertical anteriorly

Scytoleptus serripes Gerstaecker, 1856

- Rostrum level with or slightly below gastric region of

carapace; gastric region horizontal or gently sloped^,.... 5

Australian Axiidae (Crustacea: Decapoda: Axiidea)

223

5. Rostrum depressed between smooth marginal rim; median

gastric carina smooth; submedian gastric carinae absent ...

- Rostrum scarcely depressed between dentate marginal

rim; median gastric carina dentate; submedian gastric

carinae present, usually toothed 7

6. Telson 1.1 times as long as wide; uropodal endopod with

oblique apex Eiconaxius kimbla Kensley, 1996

- Telson 1.5 times as long as wide; uropodal endopod with

rounded apex Eiconaxius mallacoota sp. nov.

7. Rostrum longer than eyestalks; scaphocerite strongly

curved; lateral gastric carina with supraocular spine plus

1 spine Platyaxius bardi sp. nov.

- Rostrum shorter than eyestalks; scaphocerite straight;

lateral gastric carina unarmed

Platyaxius brevirostris Sakai, 1994

8. Pleopods 2-5 without appendix interna 9

- Pleopods 2-5 each with appendix interna wg. 12

9. Scaphocerite slender, reaching more than half length of

antennal article 4; rostrum with 4 or 5 lateral spines along

length; lateral gastric carina unarmed; submedian gastric

carina with 3 or 4 spines

Bouvieraxius keiensis Sakai, 1992

- Scaphocerite plate-like, triangular or bifurcate, barely

overlapping base of antennal article 4; rostrum with 1 or 2

lateral spines near base; lateral gastric carina unarmed or

with 2 teeth; submedian gastric carina unarmed or with

6-8 spines 10

10. Submedian gastric carina with 5-8 spines

Paraxiopsis austrinus (Sakai, 1994)

- Submedian gastric carina absent or unarmed 11

11. Lateral gastric carina with 1 or 2 spines posterior to

supraocular spine; submedian gastric carina ending

anteriorly as definite tooth

Paraxiopsis brocki (De Man, 1888)

- Lateral gastric carina unarmed posterior to supraocular

spine; submedian gastric carina weak, not well defined

Paraxiopsis pumilus (Sakai, 1994)

12. Uropodal exopod with 5 prominent lateral spines, second

article oval, longer than wide; uropodal endopod with

broad shoulder proximally

Allaxius clypeatus (De Man, 1888)

- Uropodal exopod with small lateral spines or unarmed,

second article short, much wider than long; uropodal

endopod evenly expanding 13

13. Eyestalks longer than spike -like rostrum n,.

Oxyrhynchaxius manningi Lin, Kensley and Chan, 2000

- Eyestalks shorter than rostrum or if longer, rostrum

broadly triangular 14

14. Cheliped dactyli with row of at least 2 erect spines on

upper margin (as well as on propodus upper margin) .... 15

- Cheliped dactyli without erect spines on upper margin

(propodus upper margin may have erect spines) 18

15. Carapace densely spinose 16

- Carapace smooth 17

16. Eyestalks longer than rostrum; major cheliped propodus

with tubercles and blunt spines laterally, gross tubercles

mesial ly Acanthaxius garawa sp. nov.

- Eyestalks shorter than rostrum; major cheliped propodus

with sharp spines laterally, small tubercles and few spines

mesially Acanthaxius ningaloo sp. nov.

17. Lateral gastric carina with 1 prominent tooth posterior to

supraocular spine; major cheliped dactylus with 2 erect

spines on upper margin; telson significantly longer than

wide Acanthaxius clevai Ngoc-Ho, 2006

- Lateral gastric carina with 2 prominent teeth posterior to

supraocular spine; major cheliped dactylus with 5 erect

spines on upper margin; telson as wide as long

Acanthaxius gathaagudu sp. nov.

18. Chelipeds densely setose, setae completely obscuring

tuberculation; rostrum extremely narrow, with about 6

small lateral teeth

Acanthaxius polychaetes Sakai, 1994

- Chelipeds sparsely setose, ornamentation visible through

setae; rostrum broadly triangular or with fewer than 6

lateral teeth if narrow 19

19. Rostrum strongly depressed below median gastric carina;

gastric carinae with only supraocular spines and minute

anterior median tooth above steeply sloping anterior ridge

Dorphinaxius kermadecensis (Chilton, 1911)

- Rostrum level with or slightly below median gastric carina;

gastric carinae more or less dentate or spinose 20

20. Abdominal pleura 2-5 acutely produced ventrally to sharp

point 21

- Abdominal pleura 2-5 rounded ventrally 22

21. Carapace tuberculate; rostrum with 1 pair of lateral

spines; chelipeds without spines on upper margin of

carpus and propodus; uropodal endopod apically rounded,

without spines Pilbaraxius kariyarra gen. and sp. nov.

- Carapace smooth; rostrum with 2 or more pairs of lateral

spines; chelipeds with strong curved spines on upper

margin of carpus and propodus; uropodal endopod

apically truncate, with 2 distal spines ^

Calaxius acutirostris Sakai and de Saint Laurent, 1989

22. Rostrum a triangular projection well separated from

gastric region, with 10 spines each side; without

supraocular spines; cheliped dactyli massive, with convex

upper margin Spongiaxius brucei (Sakai, 1986)

224

G.C.B. Poore and D.J. Collins

- Rostrum triangular, not well separated from gastric

region, variously dentate; with supraocular spines clearly

or weakly differentiated; cheliped dactyli tapering | ; jj. 23

23. Scaphocerite longer than antennal peduncle, with 3 spines

on lower margin and 1 spine on mesial margin; eyes

unpigmented

. . Australocaris pinjarup gen. and sp. nov.

- Scaphocerite shorter than antennal peduncle, unarmed,

minutely bifurcate or with basiomesial spine; eyes

pigmented ^ 24

24. Lateral gastric carina unarmed posterior to supraocular

spine; submedian gastric carina unarmed or absent 25

- Submedian and usually lateral gastric carinae denticulate

posterior to supraocular spine 26

25. Median gastric carina with 2 teeth anteriorly; submedian

gastric carina a simple ridge; cheliped merus upper margin

unarmed Axius werribee (Poore and Griffin, 1979)

- Median gastric carina unarmed; submedian gastric carina

absent; cheliped merus upper margin with 2 small teeth ....

Planaxius brevifrons Komai and Tachikawa, 2008

26. Lateral gastric carina unarmed; submedian gastric carina

a hair-pin shaped double ridge of small beads; median

gastric carina with few spines and/or beads 27

- Lateral gastric carina with several teeth; submedian

gastric carina of numerous teeth; median gastric carina

with several teeth 28

27. Carapace smooth; telson without lateral teeth; inner row

of tubercles of submedian gastric carina with wide hiatus

Michelaxiopsis australiensis (De Man, 1925)

- Carapace covered with numerous short stiff setae; telson

with 3 lateral teeth; inner row of tubercles of submedian

gastric carina continuous

Michelaxiopsis nauo gen. and sp. nov.

28. Gastric region with numerous tubercles between median

and submedian rows of denticles; cheliped merus without

tooth on upper margin

Axiopsis serratifrons (A. Milne-Edwards, 1873)

- Gastric region with single or duplicated submedian rows

of denticles; cheliped merus with subdistal tooth on upper

margin 29

29. Gastric region with duplicated submedian rows of

denticles; larger cheliped propodus tuberculate over distal

half of lateral face Axiopsis tsushimaensis Sakai, 1992

- Gastric region with single submedian row of denticles;

larger cheliped propodus smooth over lateral face . ■ .

Axiopsis consobrina De Man, 1905

Acanthaxius Sakai and de Saint Laurent, 1989

Acanthaxius Sakai and de Saint Laurent, 1989: 66 (diagnosis, list

of species).— Kensley, 1996d: 70-71 (diagnosis, list of species).

Type species. Axiopsis ( Axiopsis ) pilocheira Sakai, 1987 by

original designation.

Diagnosis. Carapace generally smooth or spinose; cervical

groove visible laterally over half distance to anterolateral

margin. Rostrum spine -like, narrow, laterally obscurely

denticulate, about as long as eyestalks, not depressed below

level of carapace, continuous with definite lateral carinae;

supraocular spines prominent (or not); lateral carina spinose;

submedian carina present, spinose; median carina a spinose

ridge; postcervical carina absent. Abdominal somite 1 pleuron

acute; pleuron 2 broad, anteriorly rounded, posteriorly rounded;

pleura 3-5 posteriorly rounded. Eyestalk cylindrical,

articulating; cornea pigmented. Antenna, scaphocerite short,

curved; distal spine on antenna article 2 anteromesially angled

and acute. Maxilliped 3 exopod not clearly bent at base of

flagellum. Pleurobranchs absent; podobranchs and

arthrobranchs well developed; epipods present on maxilliped 2

to pereopod 4. Pereopods 1 asymmetrical, with propodus

cylindrical; carpus-dactylus upper margins prominently

spinose (or not). Pereopods 3-5 propodi with transverse rows

of robust setae; dactyli tapering, with longitudinal row of robust

setae. Pleopods 3-5, appendix interna present. Pleopod 1 of

male absent (or present). Pleopod 2 of male with appendix

masculina. Uropodal exopod with transverse suture. Telson

with lateral fixed spines and posterolateral robust setae; apex

truncate -rounded.

Remarks. Acanthaxius is a genus of 13 nominal species, from

the Pacific Ocean and Caribbean Sea. Kensley (1996d) stated

that “the definition of this genus contains some uncertainties”

and was supported in this by Ngoc-Ho (2006). Most of its

members are unambiguously recognised by the combination of

spinose rostrum, supraocular spine, spine-bearing median,

submedian and lateral gastric carinae on the gastric region of

the carapace, narrow chelipeds (pereopods 1) with prominently

spinose upper margin and elongate fingers, absence of

pleurobranchs, absence of the male pleopod 1, and

anteromesially angled acute distal spine on antenna article 2.

Some species, including the type species, are known only

from females and the absence of the male pleopod 1 would

appear to have been assumed by the authors of the genus from

the condition in A. miyazakiensis Yokoya, 1933 that they

redescribed from abundant material. The male pleopod 1 is

absent also in species described subsequently, A. formosa

Kensley and Chan, 1998, A. grandis Kensley and Chan, 1998,

A. gadaletae Ngoc-Ho, 2006, A. clevai Ngoc-Ho, 2006 and

one of the new species described here.

Two species, Calocaris ( Calastacus ) hirsutimana Boesch

and Smalley, 1972 and Axiopsis ( Axiopsis ) caespitosa Squires,

1979, both referred to Acanthaxius by Sakai and de Saint

Laurent (1989), and A. polychaetes Sakai, 1994 do possess a

male pleopod 1 and also differ from typical Acanthaxius in

absence of a prominent supraocular spine, more compact

propodus on the major cheliped with few lateral spines, and a

prominent lateral spine on the telson. Acanthaxius spinulicauda

(Rathbun, 1902) is also less spinose than the type species.

These species may well deserve another genus whose

exceptions are given in parentheses in the diagnosis above.

Australian Axiidae (Crustacea: Decapoda: Axiidea)

225

Acanthaxius clevai Ngoc-Ho, 2006

Figures 1-3, 37

Acanthaxius clevai Ngoc-Ho, 2006: 59-62, figs. 1, 2,

3A-C).

Material examined. WA, off Mermaid Reef, 17°11.83'S,

119°34.8rE-17°12.37'S, 119°35.00'E (stn SS05-2007 079), 435-438

m, 18 Jun 2007, NMV J55706 (female, cl. 28.8 mm, tl. 64 mm). WA,

off Point Leveque, 14°58.22'S, 121°38.56'E-14°57.76'S, 121°38.26’E

(stn SS05-2007 143), 232-228 m, 02 Jun 2007, NMV J55705 (juvenile

male, cl. 8.3 mm, tl. 20 mm).

Diagnosis. Carapace generally smooth, with scattered setae

and small tubercles dorsally. Rostrum 0.4 times length of front-

to-cervical groove, acute, with 2 lateral spines anterior to

supraocular spine, continuous with definite lateral gastric

carinae (connecting by diverging curved ridges). Supraocular

spines prominent. Lateral gastric carina with 1 spine, diverging

anteriorly. Submedian gastric carina with 4 spines (last small,

flattened). Median gastric carina with 3 spines. Abdominal

pleuron 1 ventrally acute; pleura 2-5 posteroventrally rounded;

pleuron 6 of male with small spine on anteroventral margin.

Antennal article 1 with 1 spine on lower distal margin;

article 2 distal spine slender, directed inwards, reaching distally

to middle of antennal article 4; scaphocerite slender, straight,

reaching distally almost to end of article 4; article 3 with 1

spine on mesial lower margin. Maxilliped 3 ischium with 2

spines on lower margin; merus with 3 spines on lower margin.

Pereopods 1 well differentiated, of similar length, propodus

of major cheliped more swollen than minor cheliped. Major

pereopod 1 merus upper margin convex, with 2 spines, lower

margin with 4 spines, lateral face spinose distally, mesial face

spinose distally; carpus upper margin with 4 midline spines

and 1 spine on each side, lower margin with 3 spines, mesial

face tuberculate, with 1 spine; propodus upper margin with 3

midline spines and 2 spines each side, lower margin with 7

spines in lateral row and 3 spines in mesial row, lateral face

tuberculate, with 5 scattered spines, mesial face tuberculate,

with 4 scattered spines; fixed finger 1.2 times length of upper

palm, cutting edge with large rounded teeth; dactylus upper

margin with 2 erect spines, 1 submarginal spines on each side,

tuberculate, cutting edge with large rounded teeth.

Minor pereopod 1 merus, carpus as in larger cheliped;

propodus as in larger cheliped except narrower, lower margin

with 7-9 spines in lateral row and 3 spines in mesial row; fixed

finger 1.7 times length of upper palm, cutting edge straight

with numerous small sharp teeth; dactylus armature as in

major pereopod.

Telson rectangular, approximately parallel-sided, 1.3 times

as long as wide, lateral margin with 3 spines, distal margin

convex with posteromedian spine, posterolateral angle with 2

robust setae; dorsal face with 2 spines in each oblique row.

Uropodal endopod 1.4 times as long as wide, with 1 lateral

spine, longitudinal ridge with 4 spines (including marginal).

Uropodal exopod 1.6 times as long as wide, with 7 lateral

spines, 2 longitudinal ribs (outer rib with 4 spines),

posterolateral angle with 1 fixed spine and 1 robust seta;

transverse suture with 10 spines.

Distribution. Solomon Islands; WA, North-West continental

margin, 15°-17°S, 119°-122°E, 228-438 m depth.

Remarks. The new material, a female and juvenile male from

north-western Australia, are barely distinguishable from Ngoc-

Ho’s (2006) figures and description of a similarly-sized male

specimen from the Solomon Islands. We illustrate them fully

and present a short diagnosis. The species is recognisable by

the pattern of spination on the rostrum and carapace (two lateral

rostral spines, median gastric carina with four spines,

submedian with five spines including the supraorbital, and a

single spine on the curved lateral gastric carina). The complex

patterns of spination on the major and minor chelipeds are also

virtually identical to those of the holotype. N. Ngoc-Ho

illustrated the minor cheliped of a female paratype (MNHN

Thl492) for us and we detect no significant differences from

our male. Slight differences in proportions (the chelipeds of the

Australian adult female are slightly more slender than in the

male holotype) can be attributed to sexual dimorphism. We

identify the small male (about one -third as long as the adult) as

the same species in spite of substantial differences in the

chelipeds. The chelipeds of the small juvenile male are more

elongate than those of the adult female and with more prominent

spination (cf. figs, li-1 with 3c, d). The upper margin of the

dactylus possesses three spines while the Australian female

and male holotype have only only two. The juvenile male

possesses spinules on the anteroventral corners of abdominal

pleura 3-6, not present in the adult female nor in the male

holotype.

Ngoc-Ho (2006) described a second similar species,

Acanthaxius gadaletae, from the Solomon Islands and New

Caledonia, based on type material about half the size of

specimens of A. clevai. The species, apparently adult males

and females, differs from A. clevai only in better defined

gastric carinae, maxilliped 3 basis with a spine, more stout

pereopod 1 with four propodal and dactylar spines on the

upper margins, and abdominal pleura 3-5 slightly angled

posteriorly. The cheliped spination alone (more spines than in

adult and juveniles of A. clevai) is sufficient to differentiate A.

gadaletae. The maxilliped 3 basis of the type of A. clevai was

said to lack a distal spine and to differentiate the species from

A. gadaletae ; both of the Australian specimens possess a

strong spine but we don’t consider this difference alone

diagnostic. This small species possesses spinules on the

anteroventral corners of abdominal pleura 3-6 as in the small

Australian male.

Acanthaxius gadaletae differs from the type species, A.

pilocheira (Sakai, 1987) most obviously in not having a spinose

carapace and in having one, not two, spines on each lateral

gastric carina.

Both specimens of A. clevai from WA lack a long straight

seta on the posterior lobe of the scaphognathite (maxilla 2

epipod; fig. 37) as was reported for this species and the similar

A. gadaletae (Ngoc-Ho, 2006). The seta is typical of Axiidae

and is found in all other species of Acanthaxius (Squires,

1979; Sakai, 1987; Sakai and de Saint Laurent, 1989; own

observations of four species). Only A. hirsutimana is

unknown.

226

G.C.B. Poore and D.J. Collins

Figure 1. Acanthaxius clevai Ngoc-Ho, 2006. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c, telson and right

uropod. d, sternites 7 and 8. e, right sternite 8 (lateral), f, left antenna peduncle, g, epistome, left antennule and antenna (ventral), h, maxilliped

3 (anterior and posterior views); i, major pereopod 1 (left, lateral), j, same (propodus-dactylus, without setae, mesial), k, minor pereopod 1 (right,

lateral). 1, same (carpus-dactylus, without setae, mesial). All figures from NMV J55706.

Australian Axiidae (Crustacea: Decapoda: Axiidea)

227

Figure 2. Acanthaxius clevai Ngoc-Ho, 2006. a-d, pereopods 2-5 (right except d, with detail of dactylus of pereopod 3). e, f, female pleopods 1,

2. All figures from NMV J55706.

Acanthaxius garawa sp. nov.

Figures 4, 5

Material examined. Holotype. Qld, Gulf of Carpentaria, 12°10.5'S,

139°56.7'E (stn SS05-1991 040, A J. Bruce), 59 m, 25 Nov 1991, NTM

Cr008808 (male, cl. 9.9 mm, tl. 28 mm).

Paratype. Qld, Gulf of Carpentaria, 11°25.2'S, 139°25.2'E (stn

SS05-1991 054, A.J. Bruce), 49 m, 28 Nov 1991, NTM Cr009067

(male, cl. 10.0 mm, tl. 28 mm).

Description of male holotype. Carapace heavily spinulose.

Rostrum 0.2 times length of front-to-cervical, acute, with 4

lateral spines anterior to supraocular spine, weakly continuous

with lateral gastric carinae. Supraocular spines moderately

prominent. Lateral gastric carina with 10 spines. Submedian

gastric carina duplicated, external row with 10-12 spines,

internal with 10-12 spines. Median gastric carina with about

20 spines. Sternite 7 (pereopod 4) deeply divided in midline

over posterior two-thirds and with sharp oblique lateral ridge.

Sternite 8 (pereopod 5) with setose semicircular flap on anterior

face at base of leg. Abdominal pleuron 1, 1.4 times as deep as

middorsal length, ventrally obtuse; pleuron 2 broad, lateral

length as long as dorsal length, anteroventrally rounded; pleura

3-5 becoming more ventrally quadrate, each with small

anteroventral tooth; pleuron 6 rounded, with small anteroventral

tooth; pleura 1-5 without lateral crease, slightly flared

laterally.

228

G.C.B. Poore and D.J. Collins

Figure 3. Acanthaxius clevai Ngoc-Ho, 2006. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c, major pereopod

1 (left, lateral), d, minor pereopod 1 (right, lateral). All figures from NMV J55705, without setae.

Eyestalk, 1.6 length of rostrum; cornea weakly pigmented.

Antennular peduncle reaching to middle of antennal article 5.

Antennal article 1 with 1 spine and 3 spinules on distoventral

margin; article 2 distal spine slender, directed slightly inwards,

reaching distally to middle of antennal article 4; scaphocerite

slightly curved, reaching distally to just beyond midpoint of

article 4; article 3 with with 4 spines on mesial lower margin;

article 4 about two-thirds length of article 2 (excluding distal

spine), with distoventral spine; article 5 about half length of

article 4. Maxilliped 3 basis with 1 spine; ischium with 2

spines on lower margin; crista dentata with ~24 teeth; merus

with 3 spines; carpus with 1 spine.

Pereopods 1 asymmetrical, propodus of major more

swollen, 1.15 times as long as minor. Major pereopod 1 (left)

coxa lower margin with 2 spines; basis lower margin without

spine; ischium lower margin with 3 spines; merus upper

margin convex, with 4 spines, lower margin with 10 spines,

lateral face spinose distally near upper margin, mesial face

rugose distally; carpus upper margin with 6 spines, lower

margin with 3 spines laterally, 1 spine mesially, lateral face

tubercular, mesial face with 3 spines plus others smaller;

propodus upper margin with row of 7 spines, lower margin

with 12 spines in lateral row and obsolete mesial row, lateral

face covered with dome-like tubercles, mesial face covered

with larger distinct tubercles, with 1 spine near gape; fixed

finger 1.2 times length of upper palm, cutting edge straight,

with c. 20 irregular rounded teeth; dactylus upper margin with

8 spines, lateral face with row of blunt spines, mesial face with

row of 6 spines and denticles, cutting edge as in fixed finger;

both fingers bearing setae.

Minor pereopod 1 coxa, ischium as in larger cheliped;

merus upper margin with 5 spines, lower margin with 7 spines;

carpus upper margin with 6 spines, lower margin with 4 spines

laterally, 1 spine mesially, lateral face weakly spinose, mesial

face with 3 spines; propodus upper margin with 5 spines, lower

margin with 7 spines in lateral row and 7 spines in mesial row,

lateral face weakly tuberculate, mesial face grossly tuberculate,

with 2 spines near gape; fixed finger 2.5 times length of upper

palm, cutting edge with c. 40 irregular teeth; dactylus upper

margin with 6 spines, lateral face with proximal row of 5

spines, mesial face with row of 4 denticles, cutting edge as in

fixed finger; both fingers setose as in major pereopod.

Pereopod 2 ischium lower margin with 1 distal spine;

merus lower margin with 4 spines; carpus about as long as

chela; propodus upper margin 0.8 length of dactylus. Pereopod

3 merus lower margin with 5 strong spines; carpus with 1

spine; propodus 1.9 times as long as dactylus, with 7 marginal

robust setae (some duplicated). Pereopod 4 missing. Pereopod

5 propodus 2.5 times as long as dactylus, subchelate, with

short fixed finger; dactylus with broad blade on cutting edge.

Pleopods 2-5 appendix interna one quarter length of

endopod; appendix masculina just exceeding appendix interna.

Telson about as long as broad, lateral margin with proximal

tooth, distal margin broadly convex, with posteromedian

spine, posterolateral angle with 1 minute tooth and 2 robust

setae; dorsal face with 2 spines in each oblique row. Uropodal

Australian Axiidae (Crustacea: Decapoda: Axiidea)

229

Figure 4. Acanthaxius garawa sp. nov. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c, telson and left uropod.

d, major pereopod 1 (left, lateral), e, same (propodus-dactylus, without setae, mesial), f, same (carpus-propodus, without setae, lower), g, minor

pereopod 1 (right, lateral), h, same (propodus-dactylus, without setae, mesial), i, same (carpus-propodus, without setae, lower). All figures from

holotype.

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G.C.B. Poore and D.J. Collins

Figure 5. Acanthaxius garawa sp. nov. a, maxilliped 3 (without setae), b, male pleopod 2. c, pereopod 2 (right), d, pereopod 3 (right), e, pereopod

5 (right). All figures from holotype.

endopod 2.0 times as long as wide, with 2 lateral and 1

distolateral spines, longitudinal ridge with 5 spines (including

marginal). Uropodal exopod 1.8 times as long as wide, with 4

lateral spines, 2 longitudinal ribs (outer rib with 1 spine),

posterolateral angle with 1 fixed spine and 1 robust seta;

transverse suture with 6 spines.

Etymology. Garawa is an Australian Aboriginal language

which was spoken in the Gulf of Carpentaria region close to the

Northern Territory-Queensland border.

Distribution. Qld, Gulf of Carpentaria, 12°S, 140°E, 49-59 m

depth (known only from the type locality).

Remarks. With its spinose carapace, Acanthaxius garawa is

closest to A. ningaloo sp. nov. described below. It differs in

having tubercles rather than sharp spines on lateral and mesial

faces of the chelipeds and the rostrum being shorter than the

eyes. The tubercles on the mesial face of the major cheliped are

strikingly prominent. See the Remarks following A. ningaloo

for differences from other similar Indo-West Pacific species.

Acanthaxius gathaagudu sp. nov.

Figures 6-8

Material examined. Holotype. WA, Shark Bay, 25°55.40'S,

112° 14.35'E-25°56. 17'S, fl2°i4.46'E (stn SS10-2005 113), 404-407

m, 06 Dec 2005, NMV J53448 (female, cl. 36 mm, tl. 85 mm).

Paratype. WA, off Mermaid Reef, 17°01.09'S,

119 o 35.46'E-17°01.81'S, 119°35.00'E (stn SS05-2007 080), 451-440

m, 18 Jun 2007, NMV J55704 (male, cl. 22 mm, tl. 55 mm).

Description of female holotype. Carapace smooth, with few

setae. Rostrum (broken in holotype, 0.4 times length of front-

to-cervical groove in paratype), acute, with 3-4 lateral spines

anterior to supraocular spine, weakly continuous with lateral

gastric carinae. Supraocular spines prominent. Lateral gastric

carina with 2 spines. Submedian gastric carina with 4 spines.

Median gastric carina with 3 anterior spines, 1 tubercle, 1

posterior spine. Sternite 7 (pereopod 4) deeply divided in

midline over posterior two-thirds and with sharp oblique lateral

ridge. Sternite 8 (pereopod 5) with setose semicircular flap on

anterior face at base of leg. Abdominal pleuron 1 twice as deep

as middorsal length, ventrally acute; pleuron 2 broad, lateral

length 1.2 times dorsal length, anteroventrally rounded; pleura

3-5 becoming more ventrally quadrate; pleuron 6 rounded;

pleura 1-5 without lateral crease, slightly flared laterally.

Eyestalk (broken in holotype, 0.5 length of rostrum in

paratype); cornea pigmented. Antennular peduncle reaching

to proximal part of antennal article 5. Antennal article 1 with

1 spine on distoventral margin; article 2 distal spine slender,

directed slightly inwards, reaching distally to middle of

antennal article 4; scaphocerite slightly curved, reaching

distally to just beyond midpoint of article 4; article 3 with

broad mesiodistal spine on lower margin; article 4 about two-

Australian Axiidae (Crustacea: Decapoda: Axiidea)

231

Figure 6. Acanthaxius gathaagudu sp. nov. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c, telson and left

uropod. d, sternites 7 and 8. e, left sternite 8 (lateral), f, left antenna peduncle (lateral), g, epistome, left antennule and antenna (ventral), h,

maxilliped 3. i, major pereopod 1 (left, without setae, lateral), j, same (propodus, dactylus, without setae), mesial, k, same (propodus, without

setae, lower). All figures from holotype.

232

G.C.B. Poore and D.J. Collins

Figure 7. Acanthaxius gathaagudu sp. nov. a, minor pereopod 1 (right, without setae, lateral), b, same (propodus, dactylus, without setae, mesial),

c, same (propodus, lower), d-g, pereopods 2-5 (left except f). h, pereopod 5 fingers, i, j, female pleopods 1, 2. All figures from holotype.

thirds length of article 2 (excluding distal spine); article 5

about half length of article 4. Maxilliped 3 basis with 1 spine;

ischium crista dentata with 15-20 teeth; merus with 4 spines;

carpus unarmed.

Pereopods 1 scarcely differentiated, of similar length,

propodus of major more swollen. Major pereopod 1 (right)

coxa lower margin with 1 spine; basis lower margin without

spine; ischium lower margin with 4 spines; merus upper

margin convex, with 4 spines, lower margin with 7 spines,

lateral face spinose distally, mesial face smooth; carpus upper

margin with 5 spines, lower margin spinulose, lateral face

spinose, mesial face with 1 spine; propodus upper margin with

1 row of 4 spines, lower margin with 18 spines in lateral row,

5 spines in proximal mesial row and 7 smaller intermediate

spines, lateral face spinose, mesial face tuberculate, with 5

spines concentrated near gape; fixed finger 1.5 times as long as

upper palm, cutting edge straight with c. 25 irregular rounded

teeth; dactylus upper margin with (est. 5) spines, lateral face

with 1 small spine, mesial face unarmed, cutting edge as in

fixed finger; both fingers bearing setae.

Australian Axiidae (Crustacea: Decapoda: Axiidea)

233

Figure 8. Acanthaxius gathaagudu sp. nov. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c, major pereopod

(right, lateral, without setae), d, minor pereopod (left, lateral, without setae), e, male pleopod 2. All figures from paratype, NMV J55704.

Minor pereopod 1 coxa, ischium and merus as in larger

cheliped; carpus upper margin with 3 spines, lower margin with

2 spines, lateral face spinose, mesial face with 1 spine; propodus

upper margin with 1 row of 4 spines, lower margin with 16

spines in lateral row and 4 spines in mesial row, lateral face

spinose, mesial face tuberculate, with 2 spines near gape; fixed

finger 2.8 times length of upper palm, cutting edge with c. 20

oblique sharp teeth; dactylus upper margin with 8 spines, lateral

face with obsolete spine, mesial face unarmed, cutting edge as

in fixed finger; both fingers setose as in major pereopod.

Pereopod 2 ischium lower margin with 1 distal spine;

merus lower margin with 3 spines; carpus slightly shorter than

chela; propodus upper margin 0.8 length of dactylus. Pereopod

3 merus lower margin with 3 spines; propodus 2.2 times as

long as dactylus, with 7 marginal robust setae (some

duplicated). Pereopod 4 merus lower margin with 3 spines;

propodus 2.7 times as long as dactylus, with 6 marginal robust

setae (some duplicated). Pereopod 5 propodus 4 times as long

as dactylus, subchelate, with short fixed finger; dactylus with

broad blade on cutting edge.

Pleopods 2-5 appendix interna one quarter length of

endopod.

Telson about as long as broad, lateral margin with 2 spines,

distal margin broadly convex, with posteromedian spine,

posterolateral angle with 1 minute tooth and 2 robust setae;

dorsal face with 2 spines in each oblique row. Uropodal endopod

1.6 times as long as wide, with 3 lateral spines, longitudinal

ridge with 3 spines (including marginal). Uropodal exopod 1.7

times as long as wide, with 9 lateral spines, 2 longitudinal ribs

(outer rib with 4 spines), posterolateral angle with 1 fixed spine

and 1 robust seta; transverse suture with 7 spines.

Male paratype. Essentially indistinguishable from female

except for: maxilliped 3 merus with 3 spines, carpus with 1

spine; details of spination of chelipeds; more slender minor

cheliped; pleuron 1 having a small ventral spine, a slightly

more rectangular pleuron 5 and a small spine on pleuron 6.

Pleopod 2 with appendix masculina about half length of distal

endopod; appendix interna slightly shorter.

Etymology. Gathaagudu is the name for Shark Bay in the local

Malgana language.

234

G.C.B. Poore and D.J. Collins

Distribution. WA, North West Shelf to Shark Bay, 17°-26°S,

112°-119°E, 400-450 m depth.

Remarks. Comparison of the figures of the holotype (figs. 6, 7)

and paratype (fig. 8) illustrates variability in spination of the

carapace, maxillipeds and chelipeds. Acanthaxius gathaagudu

is most similar to two other species of Acanthaxius with two

spines on the lateral gastric carina. The most significant

difference between them is in the number of spines on the upper

margin of the dactylus of the major cheliped, five (in both sexes)

in the new species, 11 in A. grandis and 8-10 in A. pilocheira.

The number of dactylar spines on the minor cheliped are similar

in the three species, 8-9 in A. gathaagudu, 10-11 in the other

two species. In A. gathaagudu the minor cheliped of the male is

slightly more slender than in the female so any comparisons

must be made between individuals of the same sex. The holotype

of A. grandis from Taiwan is a female of carapace length 33.5

mm whose chelipeds were figured by Kensley and Chan (1998:

fig. 4). The figured holotype of A. pilocheira is also a female of

a similar size. The female major and minor chelipeds of A.

gathaagudu are narrower (depth: dorsal length of propodus

0.85) and with more prominent marginal spines than in A.

grandis (depth : dorsal length of propodus 1.25) whereas A.

pilocheira is intermediate between them. Acanthaxius

gathaagudu has narrower uropodal rami than the other species

and, like A. pilocheira, has 8-9 marginal spines on the uropodal

exopod compared to only five in A. grandis. Kensley and Chan

(1998) remarked on the abdominal pleura, especially pleuron 5,

which is posteriorly rectangular in A. grandis, rather than

rounded as in A. pilocheira. We suspect that this may be a sexual

difference-their figures of a paratype may be of a male whereas

those of A. pilocheira are of female. The male of A. gathaagudu

has a slightly more rectangular pleuron 5 than the female and a

small spine on pleuron 6, absent in the female (cf. figs. 6a, 8a).

Acanthaxius ningaloo sp. nov.

Figures 9, 10, 38

Material examined. Holotype. WA, off Ningaloo North, 21°59.10'S,

113 0 49.12'E-21°59.47'S, 113°49.08'E (stn SS10-2005 153), 165 m, 11

Dec 2005, NMV J53446 (female, cl. 22.2 mm, tl. 56.7 mm).

Paratype. Collected with holotype, NMV J53447 (male, cl. 16

mm, complete but fragmented).

Description of female holotype. Carapace heavily spinulose.

Rostrum 0.4 times length of front-to-cervical groove, acute,

with 3-4 lateral spines anterior to supraocular spine, continuous

with definite lateral gastric carinae. Supraocular spines

prominent. Lateral gastric carina with 9 spines. Submedian

gastric carina duplicated, external row with 7-10 spines,

internal with 12-13 spines. Median gastric carina with 21

spines. Sternite 7 (pereopod 4) deeply divided in midline over

posterior two-thirds and with sharp oblique lateral ridge.

Sternite 8 (pereopod 5) with setose semicircular flap on anterior

face at base of leg. Abdominal pleuron 1 twice as deep as

middorsal length, ventrally acute; pleuron 2 broad, lateral

length 1.2 times dorsal length, anteroventrally rounded; pleura

3-5 becoming more ventrally acute; pleuron 6 with small spine

on ventral margin; pleura 1-5 with lateral crease.

Eyestalk 0.8 length of rostrum; cornea weakly pigmented.

Antennular peduncle reaching to proximal part of antennal

article 5. Antennal article 1 with 1 spine and 2 spinules on

lower distal margin; article 2 distal spine slender, directed

slightly inwards, reaching distally to middle of antennal article

4; scaphocerite slender, straight, reaching distally almost to

end of article 4; article 3 with 4 spines on mesial lower margin;

article 4 about half length of article 2 (excluding distal spine),

with distoventral spine; article 5 about two-thirds length of

article 4. Maxilliped 3 basis with 1 spine; ischium with 2

spines on lower margin; crista dentata with 16 teeth; merus

with 3 spines on lower margin; carpus unarmed.

Pereopods 1 scarcely differentiated, of similar length,

propodus of major more swollen. Major pereopod 1 (right)

coxa lower margin with 2 spines; basis lower margin with 1

spine; ischium lower margin with 4 spines; merus upper

margin convex, with 4 spines, lower margin with 9 spines,

lateral face spinose distally, mesial face spinose distally;

carpus upper margin with 6 spines, lower margin with 4 spines

laterally, 1 spine mesially, lateral face spinose, mesial face

with 1 spine; propodus upper margin with 2 rows each of 4

spines, lower margin with 21 spines in lateral row and 8 spines

in mesial row, lateral face spinose, mesial face tuberculate,

with 8 spines concentrated near gape; fixed finger twice length

of upper palm, cutting edge straight, with c. 20 irregular

rounded teeth; dactylus upper margin with 9 spines, lateral

face with proximal row of 5 submarginal spines, mesial face

with proximal spine and row of denticles, cutting edge as in

fixed finger; both fingers bearing setae.

Minor pereopod 1 coxa, ischium and merus as in larger

cheliped; carpus upper margin with 4 spines, lower margin with

1 spine, lateral face spinose, mesial face with 1 spine; propodus

upper margin with 2 rows each of 4 spines, lower margin with

23 spines in lateral row and 5 spines in mesial row, lateral face

spinose, mesial face tuberculate, with 6 spines concentrated

near gape; fixed finger 2.5 times length of upper palm, cutting

edge with c. 30 oblique sharp teeth; dactylus upper margin with

10 spines, lateral face with proximal row of 5 submarginal

spines, mesial face with row of denticles, cutting edge as in

fixed finger; both fingers setose as in major pereopod.

Pereopod 2 ischium lower margin with 1 distal spine;

merus lower margin with 3 spines; carpus slightly shorter than

chela; propodus upper margin 0.8 length of dactylus. Pereopod

3 merus lower margin with 3 spines; propodus 2.8 times as

long as dactylus, with 6 marginal robust setae (some

duplicated). Pereopod 4 merus lower margin with 4 spines;

propodus 2.6 times as long as dactylus, with 7 marginal robust

setae (some duplicated). Pereopod 5 propodus 2.9 times as

long as dactylus, subchelate, with short fixed finger; dactylus

with broad blade on cutting edge.

Pleopods 2-5 appendix interna one third length of

endopod.

Telson about as long as broad, lateral margin with 1 spine,

distal margin convex with posteromedian spine, posterolateral

angle with 1 or 2 minute teeth and 2 robust setae; dorsal face

with 2 spines in each oblique row. Uropodal endopod 1.6 times

as long as wide, with 3 lateral spines, longitudinal ridge with 3

spines (including marginal). Uropodal exopod 1.6 times as

Australian Axiidae (Crustacea: Decapoda: Axiidea)

235

Figure 9. Acanthaxius ningaloo sp. nov. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c, telson and right

uropod. d, sternites 7 and 8. e, left sternite 8 (lateral), f, left antenna peduncle (lateral), g, epistome, left antennule and antenna (ventral), h, female

major pereopod 1 (right, dislocated, lateral), i, same (carpus-dactylus, without setae, mesial), j, female minor pereopod 1 (left, dislocated,

without setae, lateral), k, same (carpus-dactylus, without setae, mesial). All figures from holotype.

236

G.C.B. Poore and D.J. Collins

Figure 10. Acanthaxius ningaloo sp. nov. a, maxilliped 3. b-e, pereopods 2-5 (right except d). f, female pleopod 1. g, male pleopod 2 (rami

without setae), h, male major pereopod 1 (right, dislocated, lateral), i, same (propodus-dactylus, without setae, mesial), j, male minor pereopod

1 (left, dislocated, without setae, lateral), k, same (propodus-dactylus [broken], without setae, mesial). Figs a-f from holotype; figs, g-k from

paratype NMV J53447.

long as wide, with 4 lateral spines, 2 longitudinal ribs (outer

rib with 2 spines), posterolateral angle with 1 fixed spine and 1

robust seta; transverse suture with 7 spines.

Male. Carapace spination and abdomen indistinguishable

from female. Pereopods 1 grossly differentiated, of similar

length, propodus of major considerably more swollen than

minor. Major pereopod 1 (right) coxa lower margin with 2

small spines; basis lower margin with 1 spine; ischium lower

margin with 5 spines; merus upper margin convex, with 4

spines, lower margin with 10 spines, lateral face spinose

distally, mesial face spinose distally; carpus upper margin

with 6 spines, lower margin with 3 spines laterally, 1 spine

mesially, lateral face spinose and densely setose in upper half,

mesial face with 1 spine; propodus upper margin with 2 rows

each of 4 spines (mesial row obsolete), lower margin with 19

spines in lateral row and 14 spines in mesial row, diminishing,

lateral face spinose and densely setose in upper half, mesial

face tuberculate, with 8 spines concentrated near gape; fixed

finger as long as upper palm, cutting edge straight, with c. 20

irregular rounded teeth; dactylus upper margin with 6 spines,

lateral face with proximal row of 5 small submarginal spines,

densely setose, mesial face with proximal spine and row of

denticles, cutting edge as in fixed finger.

Minor pereopod 1 coxa, ischium and merus as in larger

cheliped; carpus upper margin with 4 spines, lower margin

with 3 lateral spines, 1 mesial spine, lateral face spinose and

setose, mesial face with 1 spine; propodus upper margin with

2 rows of 4 lateral and 3 small mesial spines, lower margin

Australian Axiidae (Crustacea: Decapoda: Axiidea)

237

with 20 spines in lateral row and 5 spines in mesial row, lateral

face spinose and setose, mesial face tuberculate, with 6 spines

concentrated near gape; fixed finger 4 times length of upper

palm, cutting edge with c. 30 oblique sharp teeth; dactylus

upper margin with spinose (broken).

Pleopod 2 with appendix masculina about third length of

endopod, attached one-third along; appendix interna slightly

shorter than appendix masculina.

Etymology. Ningaloo, the name of a major WA fringing reef, is

from the local Gnulli language.

Distribution. WA, off Ningaloo North, 22°S, 114°E, 165 m

depth (known only from the type locality).

Remarks. Acanthaxius ningaloo is most similar to A.

polyacantha (Miyake and Sakai, 1967) described from the East

China Sea, also recorded from the Solomon Islands by Ngoc-

Ho (2006), and to A. formosa from off Taiwan (Kensley and

Chan, 1998). The type specimen is a female of similar size to

the holotype of the other three species; the Solomon Islands

specimen is much smaller. All species share a carapace covered

with spinules and duplicated submedian gastric carinae.

Acanthaxius ningaloo has a longer rostrum with four (as in A.

formosa, rather than two as in A. polyacantha) pairs of lateral

spines, antenna article 3 with four spines (rather than one as in

the other two species), relatively shorter telson, more pronounced

spines along the margin of the cervical groove and more definite

spines on the postcervical carapace. The spines along the

margins of the fingers of the chelipeds are apparently longer

and more erect. Further, A. formosa lacks denticles anterolateral

to the cervical groove. Acanthaxius miyazakiensis, reported

from Japan, Philippines and New Caledonia (Sakai and de

Saint Laurent, 1989), is similar to all three but has a more

elongate minor first cheliped and fewer carapace spinules.

Acanthaxius ningaloo has strongly sexually dimorphic

chelipeds, the major one of the male being more setose, more

swollen than the minor, and with shorter marginal spines than

the female. In no species described to date has both sexes been

illustrated. Most figures are of females but the male of A.

grandis, a species with a similar cheliped but smooth carapace,

would appear to be of the grossly swollen form (Kensley and

Chan, 1998).

Acanthaxius polychaetes Sakai, 1994

Acanthaxius polychaetes Sakai, 1994: 193-198, figs.

11-13.— Davie, 2002: 450.

Distribution. Qld, continental slope, 260 m depth.

Remarks. Acanthaxius polychaetes is remarkable for the

absence of spines on the upper margins of the carpus-dactylus

and abundance of long setae on the chelipeds obscuring any

ornamentation. Like American species of this genus and unlike

the Indo-West Pacific species, the male possesses a pleopod 1.

The distal spine on article 2 of the antenna is, however,

characteristically anteromesially directed.

Allaxius Sakai and de Saint Laurent, 1989

Allaxius Sakai and de Saint Laurent, 1989: 73-74.

Remarks. Allaxius, with five species from the Indian Ocean,

Indonesia and Papua New Guinea, is recognised by the uropodal

exopod being spined laterally with its second article at least as

long as wide and usually longer than wide, and the uropodal

endopod having broadly shoulders proximolaterally. The

rostrum is short, about the same length as the eyestalks, with

short lateral teeth.

Allaxius clypeatus (De Man, 1888)

Axius clypeatus De Man, 1888: 470, pi. 20 fig. 2.

Axiopsis ( Axiopsis ) clypeata. — De Man, 1925b: 70.

Allaxius clypeatus.— Sakai and de Saint Laurent, 1989: 73-74.

Material examined. Qld, Yonge Reef, Lizard Island (14°38'S,

145°38'E), 8 Nov 1975, AM P25014 (male, cl. 4.5 mm).

Remarks. Allaxius clypeatus is known from reefs in Indonesia

and is here reported from Lizard Island, northern Great Barrier

Reef. The rostrum is narrow and as long as the eyestalks, with

two pairs of lateral teeth. The median gastric carina has a blunt

tooth anteriorly and a broad triangular plate posteriorly. The

submedian carina comprises two blunt teeth and the lateral

gastric carina just one tooth. The tapering telson has four lateral

spines and a minute posterolateral one. The uropodal exopod

has three spines in a longitudinal ridge, four lateral spines,

three small spines along the transverse suture and a larger

mesiodistal spine. A long moveable spine lies lateral to the oval

second article of the exopod. The uropodal endopod has a

strong lateral shoulder, four spines in a longitudinal ridge, and

one lateral and two distolateral spines.

Ambiaxius Sakai and de Saint Laurent, 1989

Ambiaxius Sakai and de Saint Laurent, 1989: 54.— Poore, 1994:

99 (key).— Sakai and Ohta, 2005: 82.

Callistocaris Kensley, 1989: 961 (objective synonym: same type

species).

Diagnosis. Carapace smooth; cervical groove visible laterally

over most of distance to anterolateral margin. Rostrum triangular

or spine-like, dentate or not, longer than eyestalks, depressed

below level of carapace, continuous with lateral carinae;

supraocular spines prominent; lateral carina weak; submedian

carina absent; median carina smooth; postcervical carina absent.

Abdominal somite 1 pleuron triangular; pleuron 2 posteriorly

rectangular; pleura 3-5 posteriorly rounded. Eyestalk cylindrical,

articulating; cornea unpigmented. Antenna, scaphocerite short.

Maxilliped 3 exopod not clearly bent at base of flagellum.

Pleurobranchs absent above pereopods 2^4; podobranchs and

arthrobranchs present; epipods present on maxilliped 2 to

pereopod 4. Pereopods 1 asymmetrical, with propodus

cylindrical; propodus with spine on upper margin. Pereopods

3-5 propodi with transverse rows of robust setae; dactyli 3 and

4 elongate, with scattered robust setae; dactylus 5 elongate.

Pleopods 3-5, appendix interna present. Hermaphroditic.

Pleopod 1 article 1 flattened; article 2 subtriangular, one-third

length of first, folded obliquely, posterior part distally lobed,

anterior part smaller, narrower, with distal margin oblique, with

short appendix interna. Pleopod 2 endopod article 2 elongate-

triangular, as long as article 1, mesial margin concave, with

238

G.C.B. Poore and D.J. Collins

basal thumb-like appendix interna, appendix masculina with a

row of strong spines followed by a short apical setose appendage.

Uropodal exopod with transverse suture. Telson without lateral

teeth, without posterolateral robust setae; apex deeply rounded

and continuous with lateral margins.

Remarks. Species of Ambiaxius are recognised by the prominent

supraocular spines, unarmed and obsolete or absent gastric

carina, being hermaphroditic, triangular short second article

on pleopod 1, and pleopod 2 endopod elongate -triangular, with

concave mesial margin and short setose end. The genus is

closest to Calastacus Faxon, 1893 (see below). Sakai and Ohta

(2005) distinguished their new genus Briancaris from

Ambiaxius on the shape of the rostrum, the former having a

broad triangular toothed rostrum while the latter has a styliform

rostrum that is barely toothed if at all. These differences are

slight and otherwise the two genera, each of three species, have

virtually identical pleopods.

Ambiaxius franklinae, from the Coral Sea, is the only

Australian species known so far. A specimen of another

species, too incomplete to describe, is recorded below.

Ambiaxius franklinae Sakai, 1994

Ambiaxius franklinae Sakai, 1994: 177-180, figs. 1, 2.— Davie,

2002: 451.

Distribution. WA, continental slope, 1300 m depth

Remarks. The species is distinguished from others in the genus

and from all other Australian axiids in the possession of a

narrow styliform upturned rostrum.

Ambiaxius sp.

Material examined. WA, off Cape Leveque, 14°36.89'S,

121°19.65'E-14°36.25'S, 121°20.74'E (stn SS05-2007 147), 700-698

m, 02 Jun 2007, NMV J54313 (hermaphrodite, cl. 10 mm, tl. 31 mm).

Remarks. The single individual lacks chelipeds and has a neat

U-shaped excision at the anterior of the gastric region where the

rostrum and supraocular spines, typical of species of Ambiaxius,

might attach. Pleopods 1 and 2 are typical of the genus. Ambiaxius

franklinae, from the Coral Sea, has similar maxilliped 3 , pleopods

1 and 2, fused triangular eyestalks, and telson and uropods, but

has a shorter scaphocerite, quite linear in this specimen.

Australocaris gen. nov.

Type species. Australocaris pinjarup sp. nov., here designated.

Diagnosis. Carapace smooth; with weakly defined short

cervical groove. Rostrum styliform, elongate, laterally

denticulate, 3 times as long as eyestalks, level with carapace,

continuous with definite lateral gastric carinae; supraocular

spines (spine at anterior end of lateral gastric carina and base of

rostrum) prominent; lateral gastric carina unarmed except for

anterior supraocular spine; submedian gastric carina present,

obsolete; median gastric carina as weak ridge; postcervical

carina absent. Abdominal somite 1 pleuron produced; pleuron

2 broad, anterioirly rounded, ventrally fiat, posteriorly rounded;

pleura 3-5 posteriorly rounded. Eyestalk cylindrical,

articulating; cornea unpigmented. Antenna, scaphocerite

extending beyond antennal peduncle, with basal mesial spine

and spines on lower margin. Maxilliped 3 exopod not clearly

bent at base of flagellum. Pleurobranchs absent above pereopods

2-4; podobranchs and arthrobranchs well developed; epipods

present on maxilliped 2 to pereopod 4. Pereopods 1 symmetrical,

with propodus laterally flattened, broad, carinate on upper and

lower margins; carpus-dactylus upper and lower margins with

strong spines. Pleopods 3-5, appendix interna present. Pleopod

1 of male minute. Pleopod 2 of male without appendix

masculina. Uropodal exopod with transverse suture.

Etymology. A combination derived from Australia and karis

(Greek), a shrimp (feminine).

Remarks. It is unfortunate that the only individual of the type

species is a male of uncertain development. It has a pair of

simple pleopods 1 and lacks an appendix masculina on pleopod

2. These limbs may develop at a later instar. Nevertheless, the

unique form of the rostrum and associated gastric carinae, the

long spinose scaphocerite, and the form of the chelipeds

distinguish the species from all other axiids and a new genus is

justified. Poore’s (1994) key leads this species to Calocarides

Wollebaek, 1908 (uropodal exopod with suture, epipods present,

appendix interna present, pleurobranchs absent, scaphocerite

well developed, carapace smooth, eyestalks not more than half

length of rostrum, eyes weakly pigmented). Differences are in

the absence of the male pleopod 1 (minute in the new genus but

not considered of generic importance— see discussion of

Paraxiopsis below) and presence of the appendix masculina

(absent in the new species). Calocarides was reviewed by

Kensley (1996c). The gastric carinae of all its 11 species are

more or less armed; apart from the supraocular spine and rostral

dentition, gastric carinae in the new species are obsolete and

unarmed. The first pereopods of species of Calocarides are

asymmetrical, narrow, cylindrical and linear with teeth

concentrated on the upper margins of the propodus and dactylus

while in the new species the chelipeds are symmetrical, broad,

flattened, with strong spines on the upper and lower margins of

the carpus-dactylus. These chelipeds set the new species apart

from all other axiid genera. Genera without an appendix

masculina are Axiorygma Kensley and Simmons, 1988,

Bouvieraxius Sakai and de Saint Laurent, 1989 (some species),

Parascytoleptus Sakai and de Saint Laurent, 1989, and Paraxius

Bate, 1888 but the new genus bears little resemblance to these

genera. All except Bouvieraxius are monotypic.

The spinose scaphocerite is unique to axiids but similar

spination is seen in the unrelated Neaxius acanthus Milne-

Edwards, 1878 (Strahlaxiidae).

Australocaris pinjarup sp. nov.

Figures 11, 12, 39

Material examined. Holotype. WA, off Bunbury, 33°00.35’S,

114 0 34.12'E-32°59.37'S, 114°34.55'E (stn SS10-2005 067), 423-397

m, 29 Nov 2005, NMV J53443 (male, cl. 10.1 mm, tl. 27 mm,

damaged).

Description of holotype. Carapace smooth except for slight

rugosity at base rostrum, with few setae on gastric region and

Australian Axiidae (Crustacea: Decapoda: Axiidea)

239

Figure 11. Australocaris pinjarup gen. and sp. nov. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna and antennae,

c, telson and right uropod. d, sternites 7 and 8, abdominal somite 1 and pleopods 1 (ventral), e, pereopod 4-5 coxa (left, lateral), f, epistome, left

antennule and antenna (ventral), g, pereopod 1 (left, lateral), h, same (carpus-dactylus, without setae, upper). All figures from holotype.

240

G.C.B. Poore and D.J. Collins

Figure 12. Australocaris pinjarup gen. and sp. nov. a, maxilliped 3. b, c, pereopods 2 (right), 3 or 4 (left), e, pleopod 2. All figures from

holotype.

rostrum. Rostrum 0.75 times length of front-to-cervical groove,

narrow, with 5-6 short oblique lateral spines anterior to

supraocular spine, continuous with lateral gastric carinae.

Supraocular spines prominent. Lateral gastric carina unarmed.

Submedian gastric carina obsolete, curved mesially between

supraocular spines. Median gastric carina sharp on rostrum,

unarmed. Sternite 7 (pereopod 4) deeply divided in midline

over posterior two-thirds and with sharp oblique lateral ridge.

Sternite 8 (pereopod 5) with setose ridge on anterior face at

base of leg. Abdominal somite 1 pleuron ventrally truncate;

pleuron 2 asymmetrical, posteriorly rounded; pleura 3-5

rounded; pleura 6 rounded.

Eyestalk, 0.3 length of rostrum; cornea unpigmented.

Antennular peduncle reaching to midpoint of antennal article 4;

article 1 swollen proximally, with small lateral spinule. Antennal

article 1 unarmed; article 2 distal spine slender, directed slightly

upwards, reaching distally to middle of antennal article 4;

scaphocerite reaching distally beyond distal margin of article 5,

with 3 strong spines on lower margin, 1 spine on mesial margin;

article 3 with sharp mesiodistal spine on lower margin; article 4

about as long as article 2 (excluding distal spine), with mesial

distal spine (left side only); article 5 about half length of article

4. Maxilliped 3 coxa and basis lower margin each with distal

spine; ischium unarmed; crista dentata of about 20 teeth; merus

with 2 spines; carpus with 1 spine.

Pereopods 1 symmetrical, flattened, carinate; coxa lower

margin with 2 spines; basis lower margin with 1 spine; ischium

lower margin with 2 spines; merus upper margin barely convex,

with 3 distal spines, lower margin with 7 spines, lateral face

smooth, mesial face smooth; carpus upper margin with 6 spines,

lower margin with 3 spines laterally, mesial face smooth; propodus

upper margin with 1 row of 4 spines, lower margin convex, with

9 spines, lateral face tuberculate, mesial face smooth; fixed finger

1.2 times as long as upper palm, cutting edge convex, with c. 15

irregular triangular teeth; dactylus upper margin with 4 spines

along proximal half, lateral face smooth, mesial face smmoth,

cutting edge denticulate; both fingers bearing setae.

Pereopod 2 ischium lower margin unarmed; merus lower

margin unarmed; carpus 0.7 length of chela; propodus upper

margin 0.5 length of dactylus. Pereopod 3 or 4 merus lower

Australian Axiidae (Crustacea: Decapoda: Axiidea)

241

margin with 1 spine; propodus 2.3 times as long as dactylus,

with oblique rows of simple setae but without robust setae

except for one distally. Pereopod 5 missing.

Pleopod 1 minute, cylindrical. Pleopod 2 without appendix

masculina; appendix interna third length of endopod.

Telson 1.2 times as long as wide, widest proximally, then

approximately parallel-sided, lateral margin unarmed, distal

margin truncate-convex, with posteromedian spine, posterolateral

angle rounded, unarmed; dorsal face without oblique ridges or

spines. Uropodal endopod 1.5 times as long as wide, without

lateral spines, longitudinal ridge with 4 spines (none marginal).

Uropodal exopod 1.6 times as long as wide, with 6 obscure

lateral spines, no longitudinal ribs, posterolateral angle with 1

fixed spine and 1 robust seta; transverse suture unarmed.

Etymology. Pinjarup is the name of the Australian Aboriginal

people inhabiting the coast close to the type locality (noun in

apposition).

Distribution. WA, off Bunbury, 33°S, 114°E, c. 400 m depth

(known only from type locality).

Remarks. The type specimen is in poor condition but sufficient

features can be ascertained to enable the species to be described

as a new genus and species. The single article of the male

pleopod 1 is enigmatic and may not reflect the adult state.

Axiopsis Borradaile, 1903

Axiopsis Borradaile, 1903: 538.— Sakai and de Saint Laurent,

1989: 76,-Komai et al„ 2002: 29-30.

Type species. Axius affinis De Man, 1888.

Diagnosis. Carapace smooth; cervical groove visible laterally

almost to anterolateral margin. Rostrum triangular, broad, laterally

denticulate, longer than eyestalks, not depressed below level of

carapace, continuous with definite lateral carinae; supraocular

spines not differentiated from other spines; lateral carina spinose;

submedian carina present, spinose; median carina a spinose ridge;

postcervical carina absent. Abdominal somite 1 pleuron acute;

pleuron 2 broad, anteriorly rounded, posteriorly rounded; pleura

3-5 posteriorly rounded. Eyestalk cylindrical, articulating; cornea

pigmented. Antenna, scaphocerite long, acute. Maxilliped 3

exopod not clearly bent at base of flagellum. Pleurobranchs

absent; podobranchs and arthrobranchs well developed; epipods

present on maxilliped 2 to pereopod 4. Pereopods 1 asymmetrical,

with propodus cylindrical; carpus-dactylus upper margins

smooth. Pereopods 3-5 propodi with transverse rows of robust

setae; dactyli tapering, with longitudinal row of robust setae.

Pleopods 3-5, appendix interna present. Pleopod 1 of male

absent. Pleopod 2 of male with appendix masculina. Uropodal

exopod with transverse suture. Telson with lateral fixed spines

and posterolateral robust setae; apex truncate-rounded.

Remarks. Axiopsis is defined by the absence of a male pleopod

1, a triangular rostrum with marginal teeth running uninterupted

on to lateral gastric carinae, similar even denticles along the

lateral, submedian and median gastric carinae, absence of

pleurobranchs and the presence of the appendix masculina on

the male pleopod 2 and appendices internae (Komai et al.,

2002). We rediagnose Axiopsis here so as to differentiate it

from the new genus, Michelaxiopsis gen. nov., below.

Axiopsis tsushimaensis Sakai, 1992 differs from others in

that the submedian gastric carinae are each duplicated but in

A. serratifrons (Milne-Edwards, 1873) irregular intermediate

tubercles occupy the space between median and submedian

gastric carinae.

Axiopsis consobrina De Man, 1905

Axiopsis consobrina De Man, 1905: 595-596.— Sakai and de

Saint Laurent, 1989: 77-78.— Sakai, 1994: 198-201, fig. 14.— Davie,

2002: 451.

Axiopsis (Axiopsis) consobrina.— De Man, 1925b: 69, 80-84, pi.

6 figs. 13-13c.— Ngoc-Ho, 2005: 55-57, fig. 4.

Distribution. Indo-West Pacific, WA, North West Shelf.

Remarks. Axiopsis consobrina possesses a distal spine on the

upper margin of the merus of the major cheliped; the lateral

face of the propodus is tuberculate only distally. The gastric

submedian gastric carina is a single row of teeth without

surrounding tubercles. Poore and Griffin’s (1979) Queensland

record of this species is of A. serratifrons. Sakai (1994)

recorded A. consobrina from the North West Shelf.

Axiopsis serratifrons (A. Milne-Edwards, 1873)

Limited synonymy. Axia [sic.] serratifrons A. Milne-Edwards, 1873:

263, pi. 13.

Axiopsis (Axiopsis) serratifrons.— De Man, 1925b: 68, 72-80, pi.

6 figs. 12— 12i.

Axiopsis serratifrons.— Sakai and de Saint Laurent, 1989:

76-77.— Ngoc-Ho, 2005: 53, fig. 3.— Komai and Tachikawa, 2008:

20-22, fig. 1 (synonyms).

? Axiopsis (Axiopsis) consobrina. — Poore and Griffin, 1979:

230-232, fig. 4.

Distribution. Indo-West Pacific, Caribbean, Brazil; Qld, Gulf

of Carpentaria.

Remarks. Milne-Edwards (1873) reported his species from

Upolu, Samoa (specimen now in Zoological Museum,

Hamburg) and Hawaii (specimen now in Museum national

d’Histoire naturelle, Paris), the latter being assumed by some to

be the type locality (Kensley, 1981; Sakai and de Saint Laurent,

1989) . De Man (1925b) described the “cotype” from Upolu and

illustrated marked tuberculation on the lateral face of the

propodus of the cheliped. Ngoc-Ho (2005) redescribed what

she called the “holotype” from Hawaii with this face being

smooth. It is possible that more than one species is involved

which is significant because the species has been reported

many times and has at least two synonyms (Kensley, 1981;

Sakai and de Saint Laurent, 1989; Komai and Tachikawa,

2008). No lectotype has been selected and Ngoc-Ho’s use of

“holotype” does not constitute lectotype designation (ICZN

74.7).

The Australian record of A. consobrina (Poore and Griffin,

1979) is another species (M. de Saint Laurent, pers. comm.,

1990) , potentially A. serratifrons. The material has not been

re-examined for this study. Until the identity (or identities) of

this widespread species is decided, A. serratifrons is best

242

G.C.B. Poore and D.J. Collins

distinguished from other Australian species of Axiopsis by the

absence of a spine on the upper margin of the merus and the

numerous tubercles that ornament the space between the

denticulate median and submedian gastric carinae.

Axiopsis tsushimaensis Sakai, 1992

Figures 13, 40

Axiopsis tsushimaensis Sakai, 1992: 173-175, figs. 14, 15.— Sakai,

1994: 198.— Komai et al„ 2002: 19-30, figs. 1-3.

Material examined. WA, off Bald Island, 35°11.26'S,

118 0 38.42'E-35°11.15'S, 118°39.00'E (stn SS10-2005 035), 157-157 m,

24 Nov 2005, NMV J55437 (male, cl. 6.8 mm, tl. 19 mm). WA, Jurien

Bay, 29°48.25'S, 114 0 25.52'E-29°48.33'S, 114°25.55'E (stn SS10-2005

083), 113-114 m, 02 Dec 2005, NMV J55444 (juvenile, cl. 3.0 mm).

Diagnosis. Carapace generally smooth except in gastric region.

Rostrum 0.4 times length of front-to -cervical groove, broadly

acute, with 4-5 lateral spines anterior to supraocular spine,

continuous with definite lateral gastric carinae. Supraocular

spines not differentiated from others in row. Lateral gastric

carina with 9-10 low spines. Submedian gastric carina

duplicated, each row of 6-8 low spines. Median gastric carina

with 9-10 low spines.

Antennal article 2 distal spine slender, directed slightly

upwards, reaching distally to quarter of antennal article 4;

scaphocerite reaching distally to end of article 4, simple;

article 3 with sharp mesiodistal spine on lower margin.

Pereopods 1 significantly differentiated. Major pereopod 1

ischium lower margin with 5 denticles and 1 spine; merus

upper margin convex, with 1 spine, lower margin with 7 spines,

last longest, and sublateral proximal row of denticles, lateral

face smooth, mesial face smooth; carpus upper margin

unarmed, lower margin unarmed; propodus upper margin with

ridge of 11 low blunt teeth, lower margin with ridge of 11 low

blunt teeth, lateral face with low tubercles concentrated near

margins on distal two-thirds (absent proximally and from

fixed finger), mesial face with low tubercles concentrated near

margins on distal two-thirds (absent proximally and from

fixed finger); fixed finger 0.5 length of upper palm, cutting

edge with irregular rounded teeth, largest at midpoint; dactylus

upper margin tuberculate, lateral face tuberculate proximally,

mesial face tuberculate proximally, cutting edge with major

tooth at midpoint and notch proximally; both fingers bearing

setae.

Minor pereopod 1 coxa, ischium, merus and carpus as in

major cheliped but narrower; propodus upper margin with sharp

ridge, lower margin ridge of 11 low blunt teeth, lateral face

mostly smooth, mesial face with few tubercles; fixed finger as

long as upper palm, cutting edge with c. 25 short sharp teeth;

dactylus upper margin unarmed, lateral face smooth, mesial

face with few denticles, cutting edge obscurely denticulate.

Telson 1.1 times as long as wide, lateral margin with 3

spines, distal margin convex, with posteromedian spine,

posterolateral angle with 4 robust setae well separated; dorsal

face with 2 spines in each oblique row. Uropodal endopod 1.5

times as long as wide, with 2 lateral spines, longitudinal ridge

with 5 spines (including marginal). Uropodal exopod 1.4 times

as long as wide, with 6 lateral spines, 2 longitudinal ribs

unarmed, posterolateral angle with 1 fixed spine and 1 robust

seta; transverse suture with 10 spines.

Male. Pleopod 2 appendix interna 0.3 length of endopod;

appendix interna 0.7 length of appendix interna.

Distribution. Tsushima Strait (Korea Strait), southern Japan;

Sulu Sea, Philippines; WA, south-western coast; 34°N-35°S,

114°-130°E, 102-157 m depth.

Remarks. Sakai (1992) based his description of Axiopsis

tsushimaensis on a juvenile female (tl. 18 mm) without a larger

cheliped. Komai et al.’s (2002) detailed description and

illustrations were likewise based on a small female. We

illustrate for the first time an adult male and show in particular

the strongly dimorphic chelipeds and pleopod 2. We could not

detect differences in the carapace armature, abdomen, tail fan

or antennae between our material and the type. The species is

similar to A. consobrina De Man, 1905 in having a spine on

the upper margin of the merus of the chelipeds and the

proximal lateral surface of the larger cheliped smooth. Axiopsis

tsushimaensis differs principally in having a double (rather

than single) gastric submedian gastric carina and scaphocerite

without a basal mesial spine (obvious to De Man, 1925b).

While A. consobrina was described from slightly larger

specimens (tl. 21, 29.5 mm), these differences would not

appear to be size related as evidenced by our examination of a

size range of the similar A. serratifrons (Milne-Edwards,

1873) in NTM collections (Cr0009886, Cr010271).

Axiopsis sp. aff. serratifrons (A. Milne-Edwards, 1873)

Material examined. Vic., eastern Bass Strait, 11.7 km W of Pt Ricardo

(37°49.53'S, 148°30.08'E), 27 m (stn MSL-EG), NMV J31777 (female,

tl. 28 mm)

Remarks. The single female without pereopods has four spines

on the rostrum, 13 teeth on the lateral gastric carina, numerous

setae in the submedian region, some in longitudinal rows, and

15 teeth on the median gastric carina. A description awaits the

discovery of more complete specimens.

Axius Leach, 1815

Axius Leach, 1815: 343.— Sakai and de Saint Laurent, 1989:

26.— Ngoc-Ho, 2003: 447.

Remarks. Axius was diagnosed most recently by Ngoc-Ho (2003).

Three species are known, the type species, A. stirhynchus Leach,

1815, from the eastern North Atlantic (Ngoc-Ho, 2003) and two

from the western North Atlantic, A. serratus Stimpson, 1852, and

A. armatus Smith, 1881 (Kensley, 2001). Species of Axius are

recognised by having a triangular, laterally armed rostrum,

undifferentiated supraocular spine, unarmed submedian gastric

carina anteriorly flexed towards the midline, uropodal exopod

with a transverse suture, pleurobranchs present above pereopods

2-4 and sometimes 5, male pleopod 1 present, and pleopods 2-5

with an appendix interna. We confirmed Ngoc-Ho’s (2003)

observation that the male pleopod 1 of A. stirhynchus is simple

but with an obsolete suture between two articles and a ridge that

hints at an obsolete appendix interna (fig. 14a). Kensley reported

Australian Axiidae (Crustacea: Decapoda: Axiidea)

243

Figure 13. Axiopsis tsushimaensis Sakai, 1992. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c, right antenna

peduncle (lateral), d, telson and left uropod. e, major pereopod 1 (left), f, minor pereopod 1 (right), g, h, i, pereopods 2, 4, 5 (left except g, with

detail of perepod 4). j, male pleopod 2, with detail of appendices interna and masculina. All figures from NMV J55437.

244

for A. serratus the “pleopod 1 in both sexes consisting of single

slender setose ramus.” We include Axiopsis ( Axiopsis ) werribee

Poore and Griffin, 1979 (and possibly another species, see below)

in this genus, the only ones outside the North Atlantic.

Sakai and de Saint Laurent (1989) listed Axiopsis

australiensis as a member of Axius but this species lacks a

male pleopod 1 and appendix masculina and possesses more

complex gastric armature. It is here redescribed in a new

genus, Michelaxiopsis gen. nov.

Axius werribee (Poore and Griffin, 1979)

Figure 14b

Axiopsis ( Axiopsis ) werribee Poore and Griffin, 1979: 232-235,

figs. 5, 6.— Gowlett-Holmes, 2008: 217 (colour photo).

Calocarides werribee.— Sakai and de Saint Laurent, 1989:

84.— Sakai, 1994: 175, 201,-Davie, 2002: 452.

Axiopsis werribee Poore, 2004: 174, figs. 45c, d, 46b, pi. llh.

Distribution. Tas., Vic. SA, 2-25 m depth.

Remarks. Axius werribee has five gastric carinae unomamented

except for two teeth on the median gastric carina at the base of

the rostrum. The triangular rostrum has five pairs of lateral teeth

from which the supraocular tooth is not differentiated. Poore and

Griffin (1979) misinterpreted the pleopods. Reexamination of

type material in Museum Victoria has revealed that the male

pleopod is present, two-articled, with the second article triangular

and bearing a thumb-like appendix interna (Poore and Griffin,

1979: fig. 6h). The male pleopod 2 has both appendices interna

and masculina. These characters together place the species

clearly in Axius rather than Axiopsis or Calocarides. Both have

spinose lateral carinae and lack pleurobranchs. The presence of

this species in SA relies on Gowlett-Holmes (2008).

Of the three species of Axius, A. werribee most closely

resembles A. armatus redescribed by Kensley (2001). These two

species alone share two teeth on the median gastric carina and

two pairs of spines dorsally on the telson and lack a pleurobranch

over pereopod 5. The chelipeds of A. werribee lack a meral spine

(present in A. armatus ) and are more compact. The male pleopod

1 (fig. 14b) is more complex than in A. stirhynchus (fig. 14a; see

too Ngoc-Ho, 2003: fig. 2E); that of A. armatus is unknown and

of A. serratus slender and setose (Kensley, 2001).

Axius sp. aff. werribee (Poore and Griffin, 1979)

Material examined. NSW, 44 km E of Nowra (34°55.47'S-34°56.04'S,

151°08.04'E-151°07.52'E), 429 m, (stn SLOPE 56), G.C.B. Poore on

RV Franklin, 22 Oct 1988, NMV J16793 (juvenile male, cl. 4.0 mm;

juvenile, cl. 2.9 mm)

Remarks. These two specimens from the continental slope lack

all pereopods except one pereopod 3 on the smaller one. The

rostrum and gastric carinae are exactly as in Axius werribee

but article 4 and the scaphocerite of the antenna are significantly

more elongate than in the subtidal species. They were compared

with similarly sized individuals of A. werribee from Port

Phillip Bay in NMV collections whose antenna is as described

for the larger adults. The larger specimen has visible male

gonopores but lacks pleopod 1 and appendix masculina as do

G.C.B. Poore and D.J. Collins

Figure 14. Male pleopods 1. a , Axius stirhynchus Leach, 1815 (France,

NMV J34093). b, Axius werribee (Poore and Griffin, 1979) (holotype,

NMV J280).

small males of A. werribee. The more elongate antenna and the

ecological separation (slope depths vs subtidal) suggest a

second Australian species of Axius similar to A. werribee (and

to A. armatus ). A description awaits the discovery of adults.

Bouvieraxius Sakai and de Saint Laurent, 1989

Bouvieraxius Sakai and de Saint Laurent, 1989: 45.— Sakai, 1992:

165-166.— Poore, 2008: 161-162.

Posthonocaris Kensley, 1989: 964.

Type species. Axius longipes Bouvier, 1905, by original

designation.

Diagnosis. Carapace smooth; cervical groove visible laterally

over almost half distance to anterolateral margin. Rostrum

Australian Axiidae (Crustacea: Decapoda: Axiidea)

245

triangular, acute, laterally denticulate, longer than eyestalks, not

depressed below level of carapace, continuous with definite

lateral carinae; supraocular spines barely differentiated from

other spines; lateral carina smooth; submedian carina present,

spinose; median carina smooth; postcervical carina absent.

Abdominal somite 1 pleuron acute; pleuron 2 broad, anteriorly

rounded, posteriorly rounded; pleura 3-5 posteriorly rounded.

Eyestalk cylindrical, articulating; cornea pigmented. Antenna,

scaphocerite long, acute. Maxilliped 3 exopod not clearly bent at

base of flagellum. Pleurobranchs present over pereopods 2^4;

podobranchs and arthrobranchs well developed; epipods present

on maxilliped 2 to pereopod 4. Pereopods 1 subequal, with

propodus cylindrical; carpus-dactylus upper margins smooth,

with distal tooth on propodus. Pereopods 3-5 propodi with

transverse rows of robust setae; dactyli tapering, with longitudinal

row of robust setae. Pleopods 3-5, appendix interna absent.

Pleopod 1 of male present, of 3 articles, second lobed and with

field of hooks (appendix interna remnant). Pleopod 2 of male

with appendix masculina elongate and with appendix interna.

Uropodal exopod with transverse suture. Telson with lateral fixed

spines and posterolateral robust setae; apex truncate-rounded.

Remarks. The genus has been discussed but not recently

redefined. It is defined by the combination of uropodal exopod

with a transverse suture, three pairs of pleurobranchs, toothed

triangular rostrum, male pleopods 1 and 2 present and,

importantly, pleopods 3-5 without an appendix interna (not

‘with’ as in the key of Poore, 2008).

Bouvieraxius keiensis Sakai, 1992

Bouvieraxius keiensis Sakai, 1992: 166-168, figs. 8, 9.

Bouvieraxius rudis.— Sakai, 1994: 177 (not Axius rudis Rathbun,

1906).

Bouvieraxius michelae Poore, 2008: 162-164, fig. 1. (syn. nov.)

Figures 15, 16, 41

Material examined. WA, off Kalbarri, 27°48.48'S,

113°18.40'E-27°49.06'S, 1 13° 18.43 'E (stn SS10-2005 102), 96-98 m,

05 Dec 2005, NMV J55441 (male, cl. 9.8 mm; female, cl. 10.0 mm).

Off Zuytdorp, 27°03.06'S, 113°13.19'E-27°02.56'S, 113°06.00'E (stn

SSI 0-2005 104), 97 m, 05 Dec 2005, NMV J53445 (damaged carapace

with chelipeds, abdomen missing). Off Shark Bay, 25°54.27’S,

112°49.23'E-25°54.26'S, 112°49.44'E (stn SS10-2005 035), 100 m, 06

Dec 2005, NMV J53444 (2 ovigerous females, cl. 10.0 mm; male with

bopyrid parasite, cl. 8.3 mm). S of Shark Bay, (SSI 0-2005 stn not

recorded), NMV J55471 (male, cl. 10.8mm). OffBarrow I., 21°02.15'S,

114°53.28'E-21°01.99'S, 114°53.14'E (stn SS05-2007 008), 90-100

m, 10 Jun 2007, NMV J55709 (damaged male, cl. 6.0 mm). North West

Shelf, 19°50.0’S 115°34.0'E, 80 m, NTM Cr000886 (ovigerous female,

cl. 8.0 mm, tl. 23.5 mm, determined as Bouvieraxius rudis by Sakai,

1994).

Diagnosis. Carapace smooth. Rostrum 0.3 times length of front-

to-cervical groove, narrowly triangular, with 3 (sometimes fourth

obsolete) lateral spines anterior to supraocular spine, continuous

with definite lateral gastric carinae. Supraocular spines similar in

size to other rostrum spines. Lateral gastric carina unarmed behind

supraocular spine. Submedian gastric carina with 3 or 4 spines.

Median gastric carina unarmed. Abdominal pleura 3-6 each with

small anteroventral tooth in male only (female pleura rounded).

Eyestalk 0.5 length of rostrum; cornea pigmented.

Antennular peduncle reaching to proximal part of antennal

article 5. Antennal article 1 with 1 spine and 2 spinules on

lower distal margin; article 2 distal spine slender, directed

distally, reaching distally to one third of antennal article 4;

scaphocerite slender, straight, reaching distally to 0.7 length

of article 4; article 3 with 1 or 2 spines on mesial lower margin;

article 4 about half length of article 2 (excluding distal spine),

without spine, article 5 about two-thirds length of article 4.

Maxilliped 3 basis with 1 spine; ischium with 3 spines on

lower margin; crista dentata with 18 teeth; merus with 4 spines

on lower margin; carpus with 1 spine on lower margin.

Pereopods 1 unequal, of similar length, propodus of major

cheliped more swollen than in minor. Major pereopod 1 coxa

lower margin with 1 spine; basis lower margin with 1 spine;

ischium lower margin with 2^4 spines; merus upper margin

convex, with 2 or 3 spines, lower margin with 5-9 spines, lateral

face smooth, mesial face smooth; carpus smooth and unarmed;

propodus upper margin with 1 row of c. 27 blunt truncate teeth,

lower margin with obsolete scale-like teeth, lateral face

tuberculate, especially distally near upper and lower margins,

mesial face tuberculate, especially near upper margin; fixed

finger 1.4— 1.6 length of upper palm, cutting edge weakly toothed;

dactylus upper margin smooth— weakly tuberculate, lateral face

smooth, mesial face smooth, cutting edge irregularlly toothed.

Minor pereopod 1 spination and tuberculation as in larger

cheliped (numbers of spines on merus show same range of

values but pair often asymmetrical); propodus less swollen,

slightly narrower, fixed finger as long as upper palm.

Male pleopod 1 article 2 with distal lobe bearing hooks

proximally, mesial lobe reaching halfway along article 3. Male

pleopod 2 appendix masculina about as long as endopod,

extending beyond endopod by half its length. Pleopods 2-5

appendix interna one third length of endopod.

Telson 1.3 times as long as broad, lateral margin with 3

spines, distal margin convex with posteromedian spine,

posterolateral angle with 1 robust seta; dorsal face with 2 spines

in each oblique row. Uropodal endopod 1.3 times as long as

wide, with 2 lateral spines, longitudinal ridge with 5 spines

(including marginal). Uropodal exopod 1.4 times as long as

wide, with 1 or 2 lateral spines, 2 longitudinal ribs (ribs with 7

scattered denticles), posterolateral angle with 1 fixed spine and

1 robust seta; transverse suture with 16-20 spinules.

Distribution. Indonesia (Kei Island), 245 m; Timor Sea, 18 m;

Mauritius, 73 m; WA, North West Shelf to Shark Bay, 20°-27°S,

113°-115°E, 80-100 m depth.

Remarks. Bouvieraxius keiensis was described from material

from Kei Island, Indonesia (type locality) and Mauritius. The

new material from the region of Shark Bay, central WA, totals

nine specimens that vary in the number of lateral spines on the

rostrum anterior to the supraocular spine (two or three, the last

often obsolete), teeth on the submedian gastric carina (usually

four, three in two individuals, five on one side in one individual),

spines on the merus of the cheliped (two or three on the upper

margin, 6-9 on the lower margin, usually asymmetrical), and

extent of tuberculation on the propodus of the cheliped (from

one third to two thirds of the lateral surface). Sakai’s (1994)

246

G.C.B. Poore and D.J. Collins

Figure 15. Bouvieraxius keiensis Sakai, 1992. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c, telson and left

uropod. d, epistome, left antennule and antenna (ventral), e, major pereopod 1 (right), f, minor pereopod 1 (left), g, maxilliped 3. h, i, male

pleopods 1, 2. j, k, female pleopods 1, 2. 1, left pleura of abdominal somites 3-6. Figs a-e, j, k from NMV J55441 female; figs, h, i, 1 from male,

NMV J55441.

Australian Axiidae (Crustacea: Decapoda: Axiidea)

247

Figure 16. Bouvieraxius keiensis Sakai, 1992. a-d, pereopods 2-5 (right except d). All figures from female, NMV J55441.

illustrations (taken from three individuals) show two lateral

spines on the rostrum, four teeth on the submedian gastric

carina, four upper and five lower spines on the merus of the

cheliped, and extensive tuberculation on the propodus of the

cheliped. While the spination of the cheliped meri of the

holotype falls outside the range of the Australian material we

can see no other differences; otherwise, the tail fan and male

pleopods seem identical. We illustrate an ovigerous female of a

size similar to Sakai’s specimens; both show the inequality in

the female chelipeds that is mirrored in males.

Bouvieraxius michelae was described on the basis of a

small male (cl. 4.2 mm) from the Timor Sea. The holotype

lacks chelipeds. Dorsal spination is essentially the same as in

the figures of B. keiensis-, its more elongate telson could be

attributed to its small size. The male pleopod 1 of B. michelae

lacks a mesial lobe overlapping an apical third article,

apparently less developed than that figured for the new material

(cf. fig. 15h with Poore, 2008: fig. 1). The pleopod 2 of the

small male of B. michelae has a shorter and less setose

appendix masculina and rami than those of the larger specimen

figured here but male pleopods 1 and 2 of both specimens are

consistent with those figured for B. keiensis (Sakai, 1992: fig.

9). We conclude that B. michelae is a junior synonym of B.

keiensis.

Sakai (1994) identified a small female from the North West

Shelf, WA, as the Hawaiian species, Bouvieraxius rudis

(Rathbun, 1906). We examined his specimen (NTM Cr000886)

and noted three teeth on the submedian gastric carina as

reported for B. rudis (the types of B. keiensis have four). Two

males (J55471, J55709) from our new collections also have

only three pairs of submedian teeth but are otherwise identical.

Sakai noted that the rostrum has three rostral spines (including

the supraocular) while B. rudis has two (Rathbun said “three

or four” and Sakai and de Saint Laurent illustrated two or

three). The cheliped merus of the NTM specimen has 2+6

spines, different from the types of both species but within the

range of our new Australian material. Sakai (1994)

differentiated B. keiensis from B. rudis on the number of teeth

on the submedian gastric carina (4 or 5 vs 3) and the number

of mesiodistal spinules on article 1 of the antenna (2 vs 4 or 5

in B. rudis from New Caledonia). The identity of the New

Caledonian specimens (Sakai and de Saint Laurent, 1989) is

uncertain (M. de Saint Laurent, pers. comm., 1990). The

mesiodistal angle of article 1 of the antenna in our material

varied from barely angular to two or three spinules.

The synonymy or otherwise of B. rudis and B. keiensis

remains uncertain and until this is resolved we prefer to use

the name associated with a species from the region.

Calastacus Faxon, 1893

Calastacus Faxon, 1893: 194. —Schmitt, 1921: 112. — de Saint

Laurent, 1972: 353. —Sakai and de Saint Laurent, 1989: 59. — Kensley,

1989: 961. -Kensley, 1996: 158, 159.

Calocaris ( Calastacus ). — Alcock, 1901: 191. — Borradaile, 1903:

539. -De Man, 1925b: 115.

Type species. Calaxius stilirostris Faxon, 1893, by monotypy.

Diagnosis. Carapace smooth; cervical groove visible laterally

over most of distance to anterolateral margin. Rostrum spine-

248

G.C.B. Poore and D.J. Collins

like, laterally weakly dentate, longer than eyestalks, depressed

below level of carapace, continuous with lateral carinae;

supraocular spines prominent; lateral carina weak; submedian

carina absent; median carina smooth; postcervical carina absent.

Abdominal somite 1 pleuron triangular; pleuron 2 posteriorly

rectangular; pleura3-5 posteriorly rounded. Eyestalk cylindrical,

articulating; cornea unpigmented. Antenna, scaphocerite short.

Maxilliped 3 exopod not clearly bent at base of flagellum.

Pleurobranchs absent above pereopods 2^4; podobranchs

rudimentary on maxilliped 3-pereopod 3, arthrobranchs present;

epipods present on maxilliped 2 to pereopod 4. Pereopods 1

asymmetrical, with propodus cylindrical; propodus with spine

on upper margin. Pereopods 3-5 propodi with transverse rows

of robust setae; dactyli 3 and 4 elongate; dactylus 5 elongate.

Pleopods 3-5, appendix interna present. Hermaphroditic.

Pleopod 1 article 1 flattened; article 2 subtriangular, two-thirds

length of first, folded longitudinally, posterior part distally lobed,

anterior part larger, broader, with short appendix interna. Pleopod

2 endopod article 2 elongate, twice length of article 1, mesial

margin straight, with basal digitiform appendix interna, appendix

masculina divided into 2 equal parts bearing rows of strong

setae. Uropodal exopod with transverse suture. Telson without

lateral teeth, without posterolateral robust setae; apex deeply

rounded and continuous with lateral margins.

Remarks. Calastacus is a genus of six species, C. stilirostris

Faxon, 1893 from the central eastern Pacific, C. laevis de Saint

Laurent, 1972 from the eastern Atlantic and Mediterranean, C.

colpos Kensley, 1996 and C. mexicanus Kensley, 1996 from the

Gulf of Mexico (Kensley, 1996e), C. crosnieri Kensley and

Chan, 1998 and C. inflatus Komai, Lin and Chan, 2009 from

the north-western Pacific. The genus was diagnosed by de

Saint Laurent (1972) whose concept has been followed by later

authors. The genus differs from Ambiaxius only in the

characteristic pleopods 1 and 2. Pleopod 1 is more elongate

than in Ambiaxius and pleopod 2 has a long second segment on

the endopod. A seventh species, first from the Indian Ocean, is

described below.

Calastacus myalup sp. nov.

Figures 17, 42

Material examined. Holotype. WA, off Bunbury, 33°00.35'S,

1 14°34.12'E-32°59.37'S, 114°34.55'E (stn SS10-2005 067), 423-397

m, 29 Nov 2005, NMV J53460 (hermaphrodite, cl. 8.0 mm, tl. 19 mm;

carapace damaged, pereopods 1 missing, 1 egg case on pleopod 3).

Paratype. Collected with holotype, NMV J58382 (hermaphrodite,

cl. 8.0 mm, tl. 19 mm; most pereopods missing).

Description of hermaphrodite holotype. Carapace smooth except

for slight rugosity between supraocular spines. Rostrum 0.5

times length of front-to-cervical groove, acute, unarmed anterior

to supraocular spine, not continuous with lateral gastric carinae.

Supraocular spines prominent. Lateral gastric carina absent.

Submedian gastric carina absent. Median gastric carina unarmed.

Abdominal pleuron 1 twice as deep as middorsal length, short,

ventrally rounded; pleuron 2 broad, lateral length 1.3 times

dorsal length, anteroventrally rounded; pleura 3-5 rounded;

pleura 6 rounded.

Eyestalk 0.4 length of rostrum; cornea unpigmented.

Antennular peduncle reaching two-thirds along antennal

article 4. Antennal article 1 with 1 sharp spine and spinule on

lower distal margin; article 2 distal spine slender, directed

slightly inwards, reaching distally to middle of antennal article

4; scaphocerite slender, directed slightly upwards, reaching

distally to middle of antennal article 4; article 3 with sharp

distal spine on lower margin; article 4 little longer than article

2 (excluding distal spine); article 5 about half length of article

4. Maxilliped 3 basis unarmed; ischium unarmed on lower

margin; crista dentata with 20 teeth; merus with 2 long spines

on lower margin; carpus unarmed.

Pereopods 1 coxa with 1 spine; basis unarmed; remainder

missing.

Pereopod 2 ischium unarmed; merus unarmed; carpus 0.7

length of chela. Pereopod 3 merus unarmed; propodus 3 times

as long as dactylus, with oblique rows of simple setae but

without robust setae. Pereopod 4 unarmed; propodus 3 times

as long as dactylus, with oblique rows of simple setae distally

but without robust setae. Pereopod 5 unarmed; propodus with

oblique rows of simple setae distally but without robust setae,

with short rounded fixed finger; dactylus broken.

Pleopod 1 article 2 0.8 length of article 1, leaf-like, concave

posteromesially, hooks representing the appendix interna on a

blunt lobe on broad triangular anterior fold, lateral fold almost

semicircular, the pair together forming a broad open tube joined

anteriorly by the hooks. Pleopod 2 endopod of 2 articles (third

weakly differentiated), all articles more or less in line with

peduncle, with thumb-like appendix interna at base of article 2,

setae of appendix masculina not well differentiated from more

distal setae (as in other species); exopod half length of endopod.

Telson 1.6 times as long as wide, widest proximally, then

approximately parallel-sided, lateral margin unarmed, distal

margin semicircular without posteromedian spine, posterolateral

region rounded; dorsal face without spines on obsolete oblique

row. Uropodal endopod 2.0 times as long as wide, without

lateral spines, longitudinal ridge unarmed. Uropodal exopod

2.0 times as long as wide, without lateral spines, 2 obsolete

longitudinal ribs unarmed, posterolateral angle with 1 fixed

spine and 1 robust seta; transverse suture with 11 robust setae

of varying lengths.

Etymology. Myalup is a settlement on the coast close to the

type locality (noun in apposition).

Distribution. WA, south-western coast, 400 m depth.

Remarks. The new species differs from C. crosnieri and C.

inflatus, the two West Pacific species, in having narrower

uropods.

Calaxius Sakai and de Saint Laurent, 1989

Calaxius Sakai and de Saint Laurent, 1989: 84 (diagnosis, list of

species).— Sakai, 1994: 192 (diagnosis).— Clark et al., 2007: 64

(diagnosis).

Manaxius Kensley, 2003: 367-368.

Type species. Calaxius acutirostris Sakai and de Saint Laurent,

1989, by original designation.

Australian Axiidae (Crustacea: Decapoda: Axiidea)

249

Figure 17. Calastacus myalup sp. nov. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c, telson and right uropod.

d, maxilliped 3. e-h, pereopods 2-4 (right except e). i, j, right and left pleopod 1 (right from anterolateral view, left from mesial view), k, pleopod

2. All figures from holotype.

250

G.C.B. Poore and D.J. Collins

Diagnosis. Carapace smooth or tuberculate; cervical groove

visible laterally over third distance to anterolateral margin.

Rostrum acutely triangular, with 2 or 3 pairs of lateral spines,

longer than eyestalks, not depressed below level of carapace,

continuous with definite lateral carinae; supraocular spines

prominent; lateral carina with at least 1 spine; submedian carina

present, with 1 or few spines; median carina a weak ridge, with

1 or few spines; postcervical carina absent. Abdominal somite

1 pleuron acute; pleuron 2 acute or rounded, pleura 3-5 acute,

sometimes with anteroventral tooth, or rounded. Eyestalk

cylindrical, articulating; cornea pigmented. Antenna,

scaphocerite long. Maxilliped 3 exopod not clearly bent at base

of flagellum. Pleurobranchs present above pereopods 2-A\

podobranchs and arthrobranchs well developed; epipods present

on maxilliped 2 to pereopod 4. Pereopods 1 slightly

asymmetrical, with propodus flattened; carpus-propodus upper

margin with strong curved spines (rarely obsolete), densely

covered with long setae. Pereopods 3-5 propodi with transverse

rows of robust setae; dactyli tapering, with longitudinal row of

robust setae. Pleopods 3-5, appendix interna present. Pleopod 1

of male absent. Pleopod 2 of male with appendix masculina.

Uropodal endopod with lateral and distolateral spines; exopod

with transverse suture. Telson with lateral fixed spines and

posterolateral robust setae; apex rounded.

Remarks. Sakai and de Saint Laurent (1989) listed eight species.

Subsequently, six species have been described, including one

originally in the synonymised genus Manaxius Kensley, 2003

(Clark et ah, 2007). While Clark et al.'s (2007) diagnosis serves

to enable species to be recognised, it should be added that

pleopods 3-5 each possess an appendix interna. Most species

can be recognised by the long rostrum, pattern of gastric spines

and chelipeds with long spines and dense mat of long setae.

The chelipeds of Calaxius euophthalmus (De Man, 1905)

would appear to lack strong spines and setae. Here, we

redescribe the type species from Australian material and erect

a new genus, Pilbaraxius, for a similar axiid with affinities to

Calaxius (see below).

Calaxius acutirostris Sakai and de Saint Laurent, 1989

Ligures 18-20, 43

Calaxius acutirostris Sakai and de Saint Laurent, 1989: 86-92,

fig. 25 (probably not figs. 23, 24).— Sakai, 192-193, fig. 10.

Material examined. WA, off Zuytdorp, 27°08.01'S,

112°45.04'E-27°08.48'S, 112°45.43'E (stn SS10-2005 105), 414-405

m, 05 Dec 2005, NMV J53450 (female, cl. 20 mm, tl. 48 mm). WA,

north-west slope, 16°45.3S, 119°46.4'E, 502-504 m, NTM Cr004234

(male, cl. 22.2, tl. 56.0). Qld, off Mackay, 22°55.1’S, 153°00.5’E,

338-325 m, NTM Cr007158 (male, cl. 26.5, tl. 65.0).

Description of female NMV J53450. Carapace smooth (with

scattered setae). Rostrum 0.5 times length of front-to-cervical

groove, acute, with 2 lateral spines anterior to supraocular

spine, continuous with definite lateral gastric carinae.

Supraocular spines prominent. Lateral gastric carina with 1

prominent spine posterior to supraocular spine. Submedian

gastric carina with 2 spines. Median gastric carina present as a

ridge with 3 small spines. Laint postcervical carina present,

becoming stronger posteriorly. Sternite 7 (pereopod 4) deeply

divided in midline over posterior two-thirds and with sharp

oblique lateral ridge. Sternite 8 (pereopod 5) with setose

semicircular flap on anterior face at base of leg. Abdominal

pleuron 1 2.5 times as deep as middorsal length, ventrally

acute; pleura 2-5 broad, lateral length 1.3 times dorsal length,

tapering posteroventrally to a point, pleura 4 and 5 with small

anteroventral tooth; pleuron 6 rounded.

Eyestalk 0.3 length of rostrum; cornea pigmented.

Antennular peduncle reaching to end of antennal article 4.

Antennal article 1 with 2 spines on lower distal margin; article

2 distal spine slender, reaching distally to middle of antennal

article 4; scaphocerite slender, straight, reaching distally 0.8

length of article 4; article 3 with 1 spine on mesial lower

margin; article 4 about 1.3 length of article 2 (excluding distal

spine), article 5 about two-thirds length of article 4. Maxilliped

3 basis with 1 spine; ischium with 2 spines on lower margin;

crista dentata with 18 teeth; merus with 3 spines on lower

margin; carpus unarmed.

Pereopods 1 scarcely differentiated, of similar length, major

slightly more swollen than minor, merus of major slightly

broader. Major pereopod 1 (right) coxa lower margin with 1

spine; basis lower margin with 1 spine; ischium lower margin

with 1 spine; merus upper margin convex, with 2 spines, lower

margin with 3 spines, lateral face with 1 distal spine, mesial

face smooth; carpus upper margin with 3 spines, lower margin

with 2 spines laterally, lateral face smooth, mesial face smooth;

propodus upper margin with 4 spines (2 in midline, 1 on each

side), lower margin with 8 spines in lateral row, lateral face

with row of 4 spines, mesial face with 1 spine and few tubercles

near lower margin; fixed finger 1.5 times upper palm, cutting

edge with 5 sharp spine-like teeth; dactylus unarmed, cutting

edge denticulate; distal articles bearing clusters of stiff setae.

Minor pereopod 1 coxa, ischium, merus and carpus as in

larger cheliped; propodus as in larger cheliped except mesial

face with 2 spines and few tubercles near lower margin; fixed

finger 1.8 times length of upper palm, cutting edge with uneven

blunt teeth; dactylus unarmed, cutting edge denticulate; distal

articles bearing clusters of stiff setae.

Pereopod 2 ischium lower margin unarmed; merus lower

margin with 3 spines; carpus slightly shorter than chela;

propodus upper margin 0.8 length of dactylus. Pereopod 3

merus lower margin with 2 spines; propodus 2.7 times as long

as dactylus, with 4 marginal robust setal rows. Pereopod 4

merus lower margin with 1 spine; propodus 2.8 times as long

as dactylus, with 3 marginal robust setal rows. Pereopod 5

propodus 5 times as long as dactylus, subchelate, with short

fixed finger; dactylus twisted.

Pleopod 1 article 2, 3 times article 1. Pleopods 2-5

appendix interna 0.3 length of endopod.

Telson 1.3 times as long as broad, lateral margin with 1

robust seta on proximal lobe, distal margin convex without

posteromedian spine, posterolateral angle with 2 robust setae;

dorsal face with 2 small spines in each oblique row. Uropodal

endopod 1.6 times as long as wide, with 1 distolateral spine,

longitudinal ridge with 2 spines (including marginal). Uropodal

exopod 1.7 times as long as wide, with 2 obsolete lateral spines,

2 longitudinal ribs unarmed, posterolateral angle with minute

Australian Axiidae (Crustacea: Decapoda: Axiidea)

251

a-c,h,i 5 mm

Figure 18. Calaxius acutirostris Sakai and de Saint Laurent, 1989. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna,

c, telson and left uropod. d, sternites 7 and 8. e, right sternites 7 and 8 (lateral), f, epistome, left antennule and antenna (ventral), g, maxilliped 3.

h, major pereopod 1 (right, lateral), i, same (propodus-dactylus, without setae, mesial). All figures from NMV J53540.

252

G.C.B. Poore and D.J. Collins

Figure 19. Calaxius acutirostris Sakai and de Saint Laurent, 1989. a, minor pereopod 1 (left, lateral), b, same (carpus-dactylus, without setae,

mesial), c-f, pereopods 2-5 (right), g, female pleopod 1. h, male pleopod 2. Figs a-g from NMV J53540, fig. h from NTM Cr007158.

fixed spine (robust seta probably lost); transverse suture with

6-10 uneven spines.

Description of male NTM Cr007158. Essentially as female

except: rostrum with 3 and 4 lateral spines anterior to

supraocular; major cheliped more swollen, propodus mesial

face more obviously tuberculate; pleopod 1 absent; pleopod 2

with appendices interna and masculina of similar lengths,

each about third length of endopod; telson lateral margin with

fixed spine (in place of robust seta) on proximal lobe and

robust seta at midpoint; uropodal exopod posterolateral angle

with minute fixed spine and robust seta.

Distribution. Madagascar (type locality); Australia, central

and northern WA, central Qld, to 27°S, 325-505 m depth.

Remarks. De Saint Laurent (pers. comm., 1990) informed GCBP

that, in her view, only figure 25 of C. acutirostris in the original

description (Sakai and de Saint Laurent, 1989) refers to this

species whose holotype is from Madagascar; the other figures and

part of the description refer to another species from the Philippines.

This would appear to differ only in the degree of spination of the

chelipeds. Sakai (1994: 192, fig. 10) recorded Calaxius acutirostris

from off Queensland and Western Australia, and illustrated and

Australian Axiidae (Crustacea: Decapoda: Axiidea)

253

Figure 20. Calaxius acutirostris Sakai and de Saint Laurent, 1989. Detail of rostum (and left antennae) of three specimens, a, female, cl. 20 mm,

NMV J53540. b, male, cl. 22.2 mm, NTM Cr004234. c, male, cl. 26.5 mm, NTM Cr007158.

described maxilla 2 and maxillipeds 1-3. We examined his

material and found it, like our own, more resembled figure 25 of

the type specimen than figure 24 of the Philippines specimen. We

figure dorsal views of all Australian specimens to illustrate the

elongation of the rostrum with increased size (fig. 20).

The original diagnosis and description were extensive and

covered all limbs (Sakai and de Saint Laurent, 1989) but apart

from pereopod 1 no limbs were illustrated. Because of the

confusion about which species was referred to, we provide

figures and another description based on the Australian

material. We note differences in the number of rostral spines

(between individuals and between left and right sides)— two to

four spines possible, in degree of spination and tuberculation

of chelipeds, and in armature of the telson.

Calaxius acutirostris , is similar to C. euophthalma (De

Man, 1905), C. manningi Kensley et al., 2000 and C. tungi

Zhong, 2000 in the possession of two well defined spines on

each lateral and sublateral gastric carina. All possess two to

four asymmetrical spines on each lateral margin of the rostrum.

Calaxius acutirostris and C. manningi also have a similar

spination pattern on the chelipeds while C. tungi has fewer and

weaker spines. Calaxius euophthalma lacks prominent cheliped

spines. Also belonging to this group of Indo-West Pacific

species with two or three asymmetrical rostral spines are C.

mimasensis (Sakai, 1967) and C. sibogae (De Man, 1925b) but

these species have less spinose chelipeds than C. acutirostris.

Dorphinaxius Sakai and de Saint Laurent, 1989

Dorphinaxius Sakai and de Saint Laurent, 1989: 33-34.— Poore,

2004: 175.

Type species. Axiopsis ( Paraxiopsis ) appendiculis Poore and

Griffin, 1979 by original designation.

Diagnosis. Carapace smooth; cervical groove visible laterally

over half distance to anterolateral margin. Rostrum triangular,

broad, laterally obscurely denticulate, about as long as eyestalks,

depressed below level of carapace, continuous with definite

lateral carinae; supraocular spines prominent; lateral carina

unarmed; submedian carina present, with blunt anterior tooth;

median carina a weak ridge, with one blunt tubercle; postcervical

carina absent. Abdominal somite 1 pleuron rounded; pleuron 2

broad, anteriorly rounded, ventrally flat, posteriorly rounded;

pleura 3-5 posteriorly rounded. Eyestalk cylindrical,

articulating; cornea pigmented. Antenna, scaphocerite short,

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G.C.B. Poore and D.J. Collins

curved. Maxilliped 3 exopod not clearly bent at base of

flagellum. Pleurobranchs present above pereopods 2-4;

podobranchs and arthrobranchs well developed; epipods

present on maxilliped 2 to pereopod 4. Pereopods 1 slightly

asymmetrical, with propodus cylindrical; carpus-dactylus

upper and lower margins unarmed. Pereopods 3-5 propodi

with transverse rows of robust setae; dactyli tapering, with

longitudinal row or robust setae. Pleopods 3-5, appendix

interna present. Pleopod 1 of male absent. Pleopod 2 of male

with appendix masculina. Uropodal exopod with transverse

suture. Telson with lateral fixed spines and posterolateral robust

setae; apex rounded.

Remarks. Sakai and de Saint Laurent (1989) likened their new

genus to Scytoleptus Gerstaecker, 1856, the two sharing a

depressed rostrum, broad foliaceous epipods on pereopods

1- 4, and sexually dimorphic pleopods 2-5. The epipods of

these two genera seem not different from those of many other

axiids and pleopods are scarcely sexually dimorphic (cf. figs.

23g, h). According to their account, Scytoleptus has a shorter

cervical groove than Dorphinaxius. They erroneously

differentiated the two genera on pleurobranchs above pereopods

2- 4 (present in both genera). Here, we rediagnose the genus

and confirm the taxonomy of the type and only species.

Dorphinaxius kermadecensis (Chilton, 1911)

Figures 21-23

Iconaxiopsis kermadecensis Chilton, 1911: 550-551, figs. 1, 2.

Axius ( Eiconaxius ) kermadecensis.— De Man, 1925b: 4, 10, 15.

Axiopsis ( Paraxiopsis ) appendiculis Poore and Griffin, 1979:

224-226, fig. 1.

Eiconaxius kermadecensis.— Sakai and de Saint Laurent, 1989: 23.

Dorphinaxius appendiculis.— Sakai and de Saint Laurent, 1989:

34.— Sakai, 1994: 200.

Dorphinaxius kermadecensis.— Davie, 2002: 452.— Poore, 2004:

175, figs. 45e, f, 46c.

Material examined. Iconaxiopisis kermadecensis Chilton, 1911.

Syntypes. New Zealand, Kermadec Islands, Meyer I. and Coral Bay,

rock pools at Sunday I. [= Raoul I.] (29°16’S, 177°55’W), Captain

Bollons, BMNH 1912.5.25.44-46 (fragments of 5 chelipeds, 1 body

with only abdominal segments identifiable, 1 body with anterior

carapace, 1 right uropod; annotated “Cotypes Pres. Prof. Chilton. The

specimens came back from Godstowe very macerated— only fragments

left. I. G Lord an j v/46.”).

Axiopsis ( Paraxiopsis ) appendiculis Poore and Griffin, 1979.

Holotype. Australia, NSW, Shellharbour, under stones between tide

marks, G. McAndrew, 1926, AM P9359 (ovigerous female, cl. 14 mm).

Paratypes. Australia, central NSW: Grafton, Collaroy, Port Jackson

and Shellharbour (29 o 47.0’S-34 o 35’S, 151°12’E-153°18’E), AM, 12 of

19 paratype lots (6 males, cl. 10-19 mm; 14 females, 6 ovigerous, cl.

6.5-21 mm).

Other material. Australia, Norfolk I., Emily Bay, Point Hunter

(29°03.8’S, 167°57.3’E), 0-0.25 m, H. Larsen, 18 Apr 1984, NTM

Cr001666 (juvenile, cl. 4.0 mm).

Description. Carapace smooth except for slight rugosity at base

rostrum, with few setae on gastric region and rostrum. Rostrum

0.25 times length of front-to-cervical groove, broadly triangular,

without or with 2 or 3 lateral tubercles anterior to supraocular

spine, depressed below level of median carina, anteriorly

directed or directed slightly upwards, continuous with lateral

carinae. Supraocular spines prominent, broadly triangular.

Lateral carina unarmed. Submedian carina smooth, with 1

anterior blunt tooth. Median carina obsolete, with single tooth,

sloping down at 20-40°. Abdominal somite 1 pleuron ventrally

rounded; pleuron 2 asymmetrical, posteriorly rounded; pleura

3-5 rounded; pleura 6 rounded.

Eyestalk, 0.5 length of rostrum; cornea pigmented.

Antennular peduncle reaching to midpoint of antennal article

4; article 1 with mesial and distal spinule. Antennal article 1

unarmed; article 2 stylocerite obsolete; scaphocerite one third

length of article 4, curved downwards; article 4, 1.5 times

length of article 2; article 5 about half length of article 4.

Maxilliped 3 coxa and basis lower margin each with distal

spine; ischium unarmed; crista dentata of about 17 teeth;

merus with 3 spines; carpus with 1 spine.

Pereopods 1 asymmetrical, robust, not sexually dimorphic.

Major cheliped coxa to ischium unarmed; merus upper margin

strongly convex, unarmed, lower margin with 1 small spine;

carpus unarmed; propodus unarmed; fixed finger 0.5 times as

long as upper palm, cutting edge with 1 triangular tooth;

dactylus distally curved, cutting edge unevenly toothed.

Minor cheliped similar to major, propodus narrower, fixed

finger 0.6 times as long as upper palm, cutting edge with 2

blunt teeth.

Pereopod 2 ischium lower margin unarmed; merus lower

margin unarmed; carpus slightly shorter than chela; propodus

upper margin 1.1 times length of dactylus. Pereopod 3 merus

unarmed; propodus 2.0 times as long as dactylus, with 5

transverse rows each of 3-9 robust setae; dactylus tapering, with

8 large robust setae plus longitudinal row of 7 smaller robust

setae on inner face, plus unguis. Pereopod 4 propodus 2.4 times

as long as dactylus, distally densely setose, with 5 transverse

rows each of 3-10 robust setae; dactylus tapering, with 11 large

robust setae plus longitudinal row of 7 smaller robust setae on

inner face, plus unguis. Pereopod 5 propodus 2.5 times as long

as dactylus, weakly subchelate, distally densely setose, with

distal transverse row of 7 robust setae; dactylus spoon-shaped,

with 2 robust setae mesially, plus unguis.

Pleopods 2-5 each with appendix interna 0.2 length of

endopod. Pleopod 1 of male absent. Pleopod 2 of male

appendix masculina 1.1 times as long as appendix interna.

Pleopod 1 of female a single narrow article.

Telson 1.2 times as long as wide, widest proximally, then

tapering, lateral margin with 4 teeth and 2 robust setae, distal

margin convex between distolateral spines and robust setae,

without posteromedian spine; dorsal face with oblique ridges

bearing 2 spines. Uropodal endopod 1.7 times as long as wide,

with 3 or 4 lateral spines, longitudinal ridge with 2^4 spines (last

submarginal). Uropodal exopod 1.5 times as long as wide, with 3

or 4 lateral spines, longitudinal rib with 3 or 4 spines, posterolateral

angle with 1 fixed spine and 1 robust seta; transverse suture 3-5

spines, two most medial strongest, adjacent and diverging.

Distribution. New Zealand: Kermadec Is; Australia: Norfolk

I., central NSW. Intertidal to 8 m depth.

Remarks. Chilton (1911) based his new species on “several

specimens,” the largest with a carapace length of 17 mm and

Australian Axiidae (Crustacea: Decapoda: Axiidea)

255

Figure 21. Dorphinaxius kermadecensis (Chilton, 1911). Fragments of 5 chelipeds, abdominal segments, anterior carapace, rami of right uropod.

All from syntypes of Iconaxiopisis kermadecensis Chilton, 1911. BMNH 1912.5.25.44-46. Top scale bar refers to carapace, abdomen and

uropod; bottom scale bar refers to chelipeds.

abdomen to end of telson, 29 mm. The “cotypes” at the BMNH

are the remains of only three specimens; no material could be

found at the Canterbury Museum, Christchurch, New Zealand,

where Chilton placed other species, nor at the National Museum

of New Zealand (Te Papa). His description was brief, mentioning

only the rostrum, the gastric region of the carapace and the

chelipeds; he illustrated only the left and right chelipeds of one

individual. He likened the species to “Iconaxiopsis” andamanensis

Alcock, 1901 now placed in Eiconaxius Bate, 1888.

In his key, De Man (1925b) correctly assumed that the

species possessed pleurobranchs and placed it in the subgenus

Axius (. Eiconaxius ). Sakai and de Saint Laurent (1989) failed

to report the pleurobranchs above pereopods 3-4 of the species

they called D. appendiculis in spite of having examined

material and noting their presence in the generic diagnosis.

The synonymy of I. kermadecensis and A. appendiculis

was first proposed by Davie (2002) on the advice of the late

Brian Kensley but we are unaware of how he reached this

conclusion. We have carefully examined what remains of the

types of 7. kermadecensis and compared them with types of A.

appendiculis , both reillustrated here. We can detect no

differences and agree with Kensley. The attitude of the rostrum

varies remarkably (cf. figs. 22a, g), as does the number of

lateral spines on the uropodal rami. A submedian tubercle sits

between those on the median and lateral gastric carinae on

some specimens, for example, on three of six individuals in

one sample. The difference is not of taxonomic significance.

Although the species occurs at the Kermadec Islands north

of New Zealand, at Norfolk Island in the Tasman Sea, and in the

Sydney region, Australia, it has not been recorded from the main

islands of New Zealand (S. Ahyong, R. Webber, pers. comm.).

Eiconaxius Bate, 1880

Eiconaxius Bate, 1888: 40.— Sakai and de Saint Laurent, 1989:

15-16.— Sakai and Ohta, 2005: 69-70 (list of species).

256

G.C.B. Poore and D.J. Collins

Figure 22. Dorphinaxius kermadecensis (Chilton, 1911). a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c, telson

and right uropod. d, epistome, bases of right antennule and antenna (ventral), e, major pereopod 1 (right, lateral), f, minor pereopod 1 (left), g,

lateral anterior carapace and rostrum, a-f from holotype of Axiopsis ( Paraxiopsis ) appendiculis Poore and Griffin, 1979 (AM P9359, ovigerous

female, cl. 14 mm); g from female, cl. 21 mm (AM P1511).

Australian Axiidae (Crustacea: Decapoda: Axiidea)

257

Figure 23. Dorphinaxius kermadecensis (Chilton, 1911). a, maxilliped 3. b-e, pereopods 2-5. f, g, female pleopods 1, 2. h, male pleopod 2. a-g

from holotype of Axiopsis ( Paraxiopsis ) appendiculis Poore and Griffin, 1979 (AM P9359, ovigerous female, cl. 14 mm), h from male, cl. 19 mm

(AM P18557).

258

G.C.B. Poore and D.J. Collins

Iconaxiopsis Alcock, 1901: 193-195.— Borradaile, 1903:

537.— Balss, 1925: 210 (type species: Eiconaxius kermadeci

laccadivensis Alcock and Anderson, 1894, subsequent designation by

Borradaile, 1903.

Axius (Eiconaxius).— Borradaile, 1903: 537-538.— De Man,

1925b: 8-9 (synonymy of Iconaxiopsis), 14.

Type species. Eiconaxius acutifrons Bate, 1888, subsequent

designation by Borradaile, 1903.

Diagnosis. Carapace smooth; cervical groove weak to

inconspicuous. Rostrum triangular, broad, laterally smooth or

obscurely denticulate, longer than eyestalks, depressed below

level of carapace, continuous with definite lateral carinae;

supraocular spines absent; lateral carina unarmed; submedian

carina present, converging anteriorly and joining median carina;

median carina a weak ridge on rostrum only, armed or not;

postcervical carina absent. Abdominal somite 1 pleuron rounded;

pleuron 2 posteriorly acute; pleura 3-5 posteriorly rounded.

Eyestalk cylindrical, articulating; cornea pigmented or not.

Antenna, scaphocerite blade-like. Maxilliped 3 exopod not

clearly bent at base of flagellum. Pleurobranchs present above

pereopods 2^4; podobranchs and arthrobranchs well developed;

epipods present on maxilliped 2 to pereopod 4. Pereopods 1

asymmetrical, with propodus cylindrical; carpus-dactylus upper

and lower margins smooth or toothed, propodus at least with

distal tooth on upper margin. Pereopods 3-5 propodi with

transverse rows of robust setae; dactyli spatulate, with row of

robust setae along oblique margin. Pleopods 3-5, appendix

interna present. Pleopod 1 of male absent. Pleopod 2 of male

with appendix interna and appendix masculina. Uropodal

exopod without transverse suture. Telson with lateral teeth,

without posterolateral robust setae; apex rounded or truncate.

Remarks. Eiconaxius is a well-characterised genus of 25 similar

species from deep waters in the Indo-Pacific, Caribbean and

Gulf of Mexico. Sakai and Ohta (2005) erected a family,

Eiconaxiidae, for the genus defining it mainly using typical

generic characters. They believed the family “is conspicuously

different from all other genera of the family Axiidae” citing the

chelate pereopod 2, rounded dactyli with robust marginal setae

on pereopods 3-5, indistinct cervical groove and absence of the

male pleopod 1. Chelate pereopods 2 are found in all axiideans.

A similar “rounded” or spatulate dactylus is found on pereopod

5 of Platyaxius Sakai, 1994 (see below) but not on pereopods 3

and 4. Platyaxius also has an oval telson and uropodal rami, the

exopod without a transverse suture, similar to those of species

of Eiconaxius. In the telson and uropod (but not the male

pleopod 1) these two genera are similar to Scytoleptus

Gerstaecker, 1856. Several axiid genera lack a male pleopod E

Many genera lack a pleopod 1 and have an indistinct cervical

groove. These similarities suggest a more complex relationship

between the genera than proposed by Sakai and Ohta (2005)

and we include the genus in Axiidae for now.

The median carina has been described as bifurcating

posteriorly in species of Eiconaxius. We interpret the

bifurcation as the two submedian gastric carinae converging

anteriorly on the median carina and look to axiid genera such

as Axiopsis Borradaile, 1903 for homology (see for example

Ngoc-Ho, 2005).

Eiconaxius kimbla Kensley, 1996

Eiconaxius kimbla Kensley, 1996b: 481-483, fig. 8.— Davie, 2002:

453.

Distribution. Qld, c. 150 m depth.

Remarks. In this species the rostral rim is unarmed and

uropodal rami are obliquely truncated. No material was

examined.

Eiconaxius mallacoota sp. nov.

Figure 24

Material examined. Holotype. Australia, Victoria, S of Point Hicks

(38°19.36’S, 149°24.18’E-38°19.00’S, 149°27.18’E, 930-951 m (stn

SLOPE 33), M.F. Gomon et al. on RV Franklin, WHOI epibenthic

sled, 23 Jul 1986, NMV J15061 (male, cl. 5.7 mm, tl. 15.0 mm).

Paratypes. Collected with holotype, NMV J53161 (male, cl. 4.1

mm, tl. 11.0 mm); NMV J53162 (male, cl. 6.2 mm, tl. 16.8 mm). S of

Point Hicks (38°21.90’S, 149°20.00’E, 1000 m (stn SLOPE 32), G.C.B.

Poore et al. on RV Franklin, WHOI epibenthic sled, 23 Jul 1986,

NMV J15060 (juvenile, cl. 2.0 mm, abdomen damaged).

Description of male holotype. Carapace smooth. Rostrum

0.25 times length of front-to-posterior margin of carapace,

concave dorsally, parallel-sided over eyes, then tapering to

acute tip, with 10 marginal lateral tubercles on oblique

margins, depressed below level of median carina, anteriorly

directed, continuous with lateral carinae. Supraocular spines

absent. Lateral carina unarmed. Submedian carina smooth,

together semicircular and converging on median carina.

Median carina obsolete, on base of rostrum only. Abdominal

somite 1 pleuron ventrally rounded-truncate; pleuron 2 oblique

angled, posteroventrally acutely produced; pleura 3-4

posteroventrally acutely produced, pleuron 5 less so, all with

anteroventral tooth; pleura 6 subacute; abdominal somite 6

dorsal posterior margin with pair of lateral teeth at base of

telson, with (3 uneven) denticles along dorsal posterior

margin.

Eyestalk 0.5 length of rostrum; cornea unpigmented.

Antennular peduncle reaching to end of antennal article 4;

article 1 unarmed. Antennal article 1 unarmed; article 2

stylocerite a vertical blade, reaching to midpoint of article 5;

scaphocerite a vertical blade, reaching beyond end of article 5;

article 3 lower margin with mesial tooth; article 4 as long as

article 2; article 5 about half length of article 4. Maxilliped 3

coxa-ischium unarmed; crista dentata of about 15 similar

teeth; merus and carpus unarmed.

Pereopods 1 asymmetrical, robust. Major cheliped coxa

with 1 spinule; basis unarmed; ischium lower margin with few

irregular teeth; merus upper margin strongly convex, with 2

small teeth, lower margin with 6 small teeth; carpus lower

margin with 1 distal tooth; propodus greatest depth equal to

upper margin length; upper margin with distal tooth, lower

margin with 5 small teeth on lateral submarginal ridge; fixed

finger 0.7 times as long as upper palm, cutting edge with

irregular teeth in shallow proximal concavity, irregular tooth

distally; dactylus distally curved, cutting edge smooth.

Minor cheliped more slender than major; coxa with 1

spinule; basis unarmed; ischium lower margin with few

Australian Axiidae (Crustacea: Decapoda: Axiidea)

259

Figure 24. Eiconaxius mallacoota sp. nov. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c, telson and right

uropod. d, epistome, right antennule and antenna (ventral), e, maxilliped 3. f, major pereopod 1 (left, lateral), g, minor pereopod 1 (right), h-k,

pereopods 2-5 (with details of dactyli of pereopods 3 and 4). 1, male pleopod 2. All figures from holotype.

260

G.C.B. Poore and D.J. Collins

irregular teeth; merus upper margin strongly convex, with 2

small teeth, lower margin with 5 small teeth; carpus lower

margin with 1 distal tooth; propodus greatest depth 1.1 times

upper margin length; upper margin with distal tooth, lower

margin with 4 small teeth on lateral submarginal ridge; fixed

finger 1.2 times as long as upper palm, with lateral ridge

parallel to cutting edge, cutting edge straight, with irregular

teeth; dactylus tapering, cutting edge smooth.

Pereopod 2 ischium lower margin unarmed; merus lower

margin unarmed; carpus slightly shorter than chela; propodus

upper margin 3 times as long as dactylus. Pereopod 3 merus

unarmed; propodus 2.5 times as long as dactylus, with 6 rows of

robust setae, of 1 or 2 setae; dactylus spatulate, with 13 robust

setae along oblique margin, plus unguis. Pereopod 4 virtually

identical to pereopod 3; propodus 2.5 times as long as dactylus,

with 6 rows of robust setae, of 1-3 setae; dactylus spatulate,

with 1 1 robust setae along oblique margin, plus unguis. Pereopod

5 propodus 2.8 times as long as dactylus, simple, distally densely

setose, without robust setae; dactylus spatulate, with 8 robust

setae along oblique margin, plus unguis.

Pleopods 2-5 each with appendix interna 0.25 length of

endopod. Pleopod 2 of male appendix masculina 0.7 times as

long as appendix interna.

Telson 1.5 times as long as wide, widest proximally, then

tapering more steeply distally, lateral margin with 8-10 small

teeth, distal margin a shallow obtuse angle between weak

distolateral teeth, without posteromedian spine; dorsal face

with obsolete longitudinal ridges. Uropodal endopod 2.35

times as long as wide, oval, with 12 small irregular lateral

teeth, last tooth distal, without longitudinal ridge. Uropodal

exopod 1.6 times as long as wide, oval with 14-20 small

irregular lateral teeth over distal two-thirds, last tooth distal,

without longitudinal rib.

Variation. Paratype male, cl. 6.2 mm, NMV J53162. Abdominal

somite 6 posterodorsal margin with pair of lateral teeth and 2

pairs of submedian denticles. Telson with 7 small lateral teeth.

Paratype male, cl. 4.1 mm, NMV J53161. Rostrum with 5

sharp lateral teeth. Abdominal somite 6 posterodorsal margin

with pair of lateral teeth, 1 pair of submedian denticles, 1

median tooth. Minor cheliped merus upper margin with 3

teeth, lower margin with 2 teeth; propodus upper margin with

2 teeth; dactylus upper margin with 1 tooth. Telson with 7

small lateral teeth.

Paratype juvenile, cl. 2.0 mm, NMV J15060. Major

cheliped merus upper margin with 2 teeth, lower margin with

2 teeth; propodus upper margin with 2 teeth, lower margin

with 4 teeth (all more prominent than in larger specimens).

Minor cheliped merus upper margin with 1 tooth, lower

margin with 1 tooth; propodus upper margin with 4 teeth,

lower margin with 3 teeth; dactylus upper margin unarmed.

Etymology. Mallacoota, a township and estuary not far from

the type locality (noun in apposition).

Distribution. Australia, off eastern Vic., 930-1000 m depth.

Remarks. The new species is represented by three males and

one juvenile. The males vary only slightly but the juvenile has

more prominent spination on the chelipeds than the others.

Eiconaxius mallacoota is similar to E. kermadeci (Bate, 1888)

from a depth of 1100 m at the Kermadec Islands, north of New

Zealand. The latter was redescribed by Sakai and de Saint

Laurent (1989: 16-18, fig. 5) and a lectotype designated. The

acute rostrum and gastric region of E. kermadeci are similarly

weakly ornamented but the rostrum is shorter, not reaching to

article 3 of the antennule, and less acutely tapering. The telson

and uropodal rami of E. kermadeci are relatively broader than

in the new species and the fixed finger of the larger cheliped is

basally strongly toothed and gaping. At 37 mm long (according

to Bate, or 23 mm according to Sakai and de Saint Laurent) E.

kermadeci is larger than the new species but few specimens are

available for either species.

Another similar species is E. parvus (Bate, 1888), known

from a single 12 mm long ovigerous female (holotype, not

lectotype as stated by Sakai and de Saint Laurent) taken at 950

m also near the Kermadec Islands. Bate’s short description

and simple drawing do not allow a comparison but Paul Clark

(Natural History Museum, London) kindly figured the holotype

for us (fig. 25). The rostrum of E. parvus is not so clearly

tapered as in E. mallacoota , the telson is relatively broader

and the chelipeds slightly more elongate.

A third similar species is E. demani Sakai, 1992 from

Indonesia and the Arafura Sea (just outside Australia’s EEZ)

but its rostrum is apically rounded rather than acute. The only

other Australian species is E. kimbla Kensley, 1996b, which

differs from the new species in having an irregularly ornamented

tapering rostrum, shorter telson, asymmetrical uropodal rami,

and more massive elongate chelae with short fingers.

Michelaxiopsis gen. nov.

Type species. Axiopsis ( Axiopsis ) australiensis De Man, 1925,

herein designated.

Diagnosis. Carapace and abdomen smooth or covered with

numerous stiff setae; cervical groove visible laterally over most

of distance to anterolateral margin. Rostrum triangular, broad,

laterally denticulate, longer than eyestalks, slightly depressed

below level of carapace, continuous with definite lateral carinae;

supraocular spines barely differentiated from other spines;

lateral carina beaded; submedian carina present, beaded,

duplicated as hair-pin shape; median carina toothed and

beaded; postcervical carina absent. Abdominal somite 1

pleuron acute; pleuron 2 broad, anteriorly rounded, posteriorly

rounded; pleura 3-5 posteriorly rounded. Eyestalk cylindrical,

articulating; cornea pigmented. Antenna, scaphocerite long,

acute. Maxilliped 3 exopod not clearly bent at base of flagellum.

Pleurobranchs present above pereopod 2-4; podobranchs and

arthrobranchs well developed; epipods present on maxilliped 2

to pereopod 4. Pereopods 1 asymmetrical (in male), with

propodus cylindrical; carpus-dactylus upper margins smooth.

Pereopods 3-4 propodi with transverse rows of robust setae;

dactyli tapering, with longitudinal row of robust setae. Pleopods

3-5, appendix interna present. Pleopod 1 of male absent.

Pleopod 2 of male without appendix masculina. Uropodal

exopod with transverse suture. Telson with lateral fixed spines

and posterolateral robust setae; apex truncate -rounded.

Australian Axiidae (Crustacea: Decapoda: Axiidea)

261

Figure 25. Eiconaxius parvus (Bate, 1888). a, b, dorsal and lateral views of anterior carapace, peduncles of antennule and antenna, c, telson and

right uropod. d, major cheliped (left), e, minor cheliped (right). All figures prepared for publication by DJC from pencil drawings of the holotype

by Paul Clark.

Etymology. This species is dedicated to the late Michele de Saint

Laurent (1926-2003) who in 1990 alerted the first author to the

probability that the type species belonged to a new genus.

Remarks. Although superficially similar to species of Axiopsis,

the type species and the second described below differ in two

important characters. Pleurobranchs are present and well

developed over pereopods 2-4 (absent and probable apomorphy

in Axiopsis ) and the male pleopod 2 lacks an appendix

masculina (present and probable plesiomorphy in Axiopsis ).

Michelaxiopsis australiensis (De Man, 1925) comb. nov.

Figure 26

Axiopsis ( Axiopsis ) australiensis De Man, 1925a: 127-132, fig.

4.— De Man, 1925b: 69,-Poore and Griffin, 1979: 226-228, fig. 2.

Axiopsis australiensis.— Poore, 2004: 174, figs. 45a, b, 46a.

Axius australiensis.— Sakai and de Saint Laurent, 1989: 26,

29.— Sakai, 1992: 165, fig. 7.-Davie, 2002: 451,-Sakai, 1994: 200.

Material examined. NSW, Long Reef, W end of reef (33°44’S,

151°19’E), I. Bennett, Apr 1964, AM P24699 (2 ovigerous females, cl.

20.0, 24.7 mm). Port Jackson, Watsons Bay (33°50.8’S, 151°16.8’E),

McIntosh and Whitelegge, Mar 1908, AM P15036 (1 male, cl. 11.7

mm; 2 females, cl. 13.6, 15.0 mm).

Vic., Shoreham (38°26’S, 145°03’E), AM P1757 (2 juvenile males,

cl. 7.2, 8.5 mm).

Diagnosis. Carapace smooth, with scattered long setae.

Rostrum 0.25 times length of front-to-cervical groove, broadly

acute, rugose dorsobasally, with 5 lateral spines anterior to

supraocular spine, continuous with definite lateral gastric

carinae. Supraocular spines barely differentiated from others

in row. Lateral gastric carina with 3 or 4 obscure low beads

fading posteriorly. Submedian gastric carina duplicated, outer

row of 12-15 tubercles curving inwards at anterior end, beading

fading posteriorly, inner row of 3-5 tubercles anteriorly, 3-4

posteriorly and clear hiatus between. Median gastric carina

with 18-22 tubercles, scarcely taller anteriorly, fading

posteriorly.

Antennule article lwith sharp stylocerite. Antennal article

2 distal spine slender, directed anteriorly, reaching distally to

quarter of antennal article 4; scaphocerite reaching two-thirds

length of article 4, simple; article 3 with short mesiodistal

spine on lower margin.

Major pereopod 1 ischium lower margin with 1 spine;

merus upper margin convex, unarmed, lower margin with 5

spines, last longest, lateral face smooth, mesial face smooth;

carpus upper margin unarmed, lower margin unarmed;

propodus upper margin tuberculate, lower margin smooth,

lateral face with squamous tubercles concentrated on distal

two-thirds along lower margin, few along upper margin,

mesial face with squamous tubercles concentrated on distal

two-thirds along lower margin, few along upper margin; fixed

finger 0.8 length of upper palm, cutting edge with 1 blunt tooth

and 1 triangular tooth; dactylus upper margin smooth, lateral

face smooth, mesial face smooth, cutting edge with blunt tooth

at midpoint and notch proximally, with subapical robust setae

prominent.

Minor pereopod 1 of similar length and ornamentation as

major cheliped but narrower.

Telson 1.2 times as long as wide, lateral margin unarmed

(rarely 1 spine), distal margin convex, with posteromedian

262

G.C.B. Poore and D.J. Collins

Figure 26. Michelaxiopsis australiensis (De Man, 1925). a, b, lateral and dorsal view of anterior carapace, peduncles of antennule and antenna,

c, telson and left uropod. d, cheliped (left). All figures from female (AM P24699).

spine, posterolateral angle with inconspicuous robust setae;

dorsal face with 2 spines in each oblique row. Uropodal

endopod 1.2 times as long as wide, with or without lateral

spine and 1 distolateral spine, longitudinal ridge with 4-5

spines. Uropodal exopod 1.3 times as long as wide, with 5-6

lateral spines, 2 longitudinal ribs with 3-4 on outer rib,

posterolateral angle with 1 fixed spine and 1 robust seta;

transverse suture with 6-10 spines.

Distribution. NSW, Vic., intertidal-subtidal.

Remarks. Michelaxiopsis australiensis is differentiated from

M. nauo by the absence of short stiff setae on the carapace and

abdomen. The submedian gastric carina is duplicated, the outer

row of tubercles continous and the inner one of three anterior

and three or four posterior tubercles separated by a distinct

hiatus. The median gastric carina is composed largely of bead-

like tubercles, the anteriormost ones only slightly more elevated

than posterior ones. The lateral margin of the telson is usually

smooth, and only rarely with a small tooth. The absence of the

Australian Axiidae (Crustacea: Decapoda: Axiidea)

263

male pleopod 1 and gastric ornamentation exclude this species

from Axius to which it has been assigned. Poore and Griffin

(1979) listed numerous specimens from central NSW, a few of

which where re-examined for this study.

The two small specimens from Victoria are in poor

condition. They differ from the others in that the gastric

carinae are almost smooth, with only vestiges of the beads or

tubercles of larger specimens. The hiatus in the inner

submedian row is quite evident.

Michelaxiopsis nauo sp. nov.

Figures 27, 28

Material examined. Holotype. SA, Sir Joseph Banks Group: Roxby I.

(34°35’S, 136°19’E), 6 m, in burrow under rocks, N. Holmes, 9 Jan

1988, SAM C6811 (ovigerous female, cl. 24.5 mm).

Paratypes. SA, Reevesby I. (34°31’S, 136°16’E), offshore from

Northwest Point, 3 m, under rocks, W. Zeidler, 13 Jan 1984, SAM

C6812 (male, 9.0 mm); Marum I., North Point (34°30’S, 136°15’E),

5-6 m, under rocks, K. Gowlett, 22 Jan 1985, SAM C6813 (male, 14.5

mm); between Reevesby and Partney Is., opposite Nicholas Bay, 6 m,

under dead Pinna shells, K. Gowlett and N. Holmes, 23 Jan 1985,

NMV J59765 (female, cl. 11.0 mm).

Description of female holotype. Carapace covered with short

stiff setae, often in small bunches, and scattered longer setae.

Rostrum 0.25 times length of front-to-cervical groove, broadly

acute, with 4 lateral spines anterior to supraocular spine,

continuous with definite lateral gastric carinae. Supraocular

spines barely differentiated from others in row. Lateral gastric

carina with obscure low beads fading posteriorly. Submedian

gastric carina duplicated in form of a hair-pin, of 20-25 beads in

each row, fading posteriorly. Median gastric carina with about 5

erect spines near base of rostrum and about 15 beads fading

posteriorly. Abdominal somite 1 pleuron ventrally rounded;

pleuron 2 rounded anteriorly and posteriorly; pleura 3^4

posteroventrally rounded, pleuron 5 less so; pleuron 6 rounded.

Eyestalk, 0.5 length of rostrum; cornea pigmented.

Antennular peduncle reaching almost to end of antennal article

4; article 1 with sharp stylocerite. Antennal article 2 distal

spine slender, directed anteriorly, reaching distally to quarter

of antennal article 4; scaphocerite reaching two-thirds length

of article 4, simple; article 3 with sharp mesiodistal spine on

lower margin. Maxilliped 3 coxa-ischium unarmed; crista

dentata of about 20 similar teeth; merus with 4 spines, largest

distal; carpus with 1 spine.

Pereopods 1 symmetrical; ischium lower margin with 1

spine; merus upper margin convex, unarmed, lower margin

with 6 spines; propodus upper margin tuberculate, lower

margin smooth, lateral face with squamous tubercles

concentrated on distal two-thirds (absent proximally and from

fixed finger), mesial face with squamous tubercles concentrated

on distal two-thirds (absent proximally and from fixed finger);

fixed finger 0.7 length of upper palm, cutting edge with 1 blunt

bicuspid tooth; dactylus upper margin smooth, lateral face

smooth, mesial face smooth, cutting edge with blunt tooth at

midpoint and notch proximally.

Pereopod 2 ischium lower margin unarmed; merus lower

margin unarmed; carpus slightly longer than chela; propodus

upper margin 1.2 times as long as dactylus. Pereopod 3 merus

lower margin unarmed; propodus 2.5 times as long as dactylus,

with 7 transverse rows of 2 or 3 robust setae; dactylus with 2

longitudinal rows of robust setae. Pereopod 4 merus unarmed;

propodus 2.1 times as long as dactylus, with 8 transverse rows

of 1-4 robust setae; dactylus with 2 longitudinal rows of robust

setae. Pereopod 5 propodus 2.2 times as long as dactylus,

subchelate, with short fixed finger; dactylus with row of 6

robust setae.

Pleopod 1 a simple, setose article. Pleopods 2-5 each with

appendix interna 0.25 length of endopod.

Telson 1.2 times as long as wide, lateral margin with 3

spines, distal margin convex, with posteromedian spine,

posterolateral angle with 2 robust setae; dorsal face with 2

spines in each oblique row. Uropodal endopod 1.5 times as

long as wide, with 1 lateral spine and 1 distolateral spine,

longitudinal ridge with 4-6 spines. Uropodal exopod 1.3 times

as long as wide, with 4 lateral spines, 2 longitudinal ribs with

6 and 2 spines, posterolateral angle with 1 fixed spine and 1

robust seta; transverse suture with 7-11 spines.

Male. Pereopods 1 significantly differentiated. Major

pereopod 1 ischium lower margin with 1 spine; merus upper

margin convex, with 2 spines, lower margin with 5 spines, last

longest, lateral face smooth, mesial face smooth; carpus upper

margin unarmed, lower margin unarmed; propodus upper margin

tuberculate, lower margin smooth, lateral face with squamous

tubercles concentrated on distal two-thirds (absent proximally

and from fixed finger), mesial face with squamous tubercles

concentrated on distal two-thirds (absent proximally and from

fixed finger); fixed finger 0.6 length of upper palm, cutting edge

with 1 blunt bicuspid tooth and 1 triangular tooth; dactylus upper

margin smooth, lateral face smooth, mesial face smooth, cutting

edge with blunt tooth at midpoint and notch proximally.

Minor pereopod 1 of similar length and ornamentation as

major cheliped but narrower (85% of width) and merus upper

margin unarmed.

Etymology. The species is named for the Nauo people of the

southern part of the Eyre Peninsula, South Australia, close to

the Sir Joseph Banks Group of islands (noun in apposition).

Distribution. SA, Sir Joseph Banks Group of islands; 5-6 m

depth.

Remarks. Michelaxiopsis nauo, from South Australia, differs

from M. australiensis, from NSW and Victoria, most obviously

in having a setose carapace and abdomen. The submedian

gastric carina is duplicated, both rows of tubercles continous

and fading posteriorly. The anterior-most tubercles of the

median gastric carina are decidedly more elevated than

posterior ones. The lateral margin of the telson always has

three teeth, absent in M. australiensis.

Oxyrhynchaxius Pari si, 1917

Remarks. Oxyrhynchaxius is characterised by elongate

eyestalks, longer than the spike -like rostrum and a carapace

covered with spinules (Lin et ah, 2000). The Australian species

is only the second known.

264

G.C.B. Poore and D.J. Collins

Figure 27. Michelaxiopsis nauo sp. nov. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c, telson and left uropod.

d, epistome, right antennule and antenna (ventral), e, f, maxilliped 3. g, pereopod 1 (left, lateral), h, i, female pleopods 1, 2. j, lateral anterior

carapace and rostrum, a-i from holtype; j from paratype male (SAM C6813).

Australian Axiidae (Crustacea: Decapoda: Axiidea)

265

Figure 28. Michelaxiopsis nauo sp. nov. a-d, pereopods 2-f (with detail of dactylus of pereopod 5). e, major pereopod 1 (right, lateral), f, minor

pereopod 1 (right, lateral), a-d from holotype; e, f from paratype male (SAM C6813).

266

G.C.B. Poore and D.J. Collins

Oxyrhynchaxius manningi Lin, Kensley and Chan, 2000

Oxyrhynchaxius manningi Lin et al., 2000: 203-205, figs. 3,

4— Davie, 2002: 453.

Distribution. WA, North West Shelf, 134 m depth.

Remarks. The generic characters immediately identify the

species in Australia.

Paraxiopsis De Man, 1905

Axiopsis ( Paraxiopsis ) De Man, 1905: 597.

Paraxiopsis.— Kensley, 1996a: 709-712.

Eutrichocheles.— Sakai and de Saint Laurent, 1989: 51 (part).

Diagnosis. Carapace and abdomen smooth or covered with

numerous stiff setae; cervical groove visible laterally over half

distance to anterolateral margin. Rostrum triangular, broad,

laterally denticulate or unarmed, longer than eyestalks, slightly

depressed below level of carapace, continuous with definite

lateral carinae; supraocular spines barely differentiated from

other spines; lateral carina spinose or smooth; submedian carina

present, dentate or smooth; median carina smooth; postcervical

carina absent. Abdominal somite 1 pleuron acute; pleuron 2

broad, anteriorly rounded, posteriorly rounded; pleura 3-5

posteriorly rounded. Eyestalk cylindrical, articulating; cornea

pigmented. Antenna, scaphocerite short, acute or asymmetrically

bifid. Maxilliped 3 exopod not clearly bent at base of flagellum.

Pleurobranchs absent; podobranchs and arthrobranchs well

developed; epipods present on maxilliped 2 to pereopod 4.

Pereopods 1 asymmetrical, with propodus cylindrical; carpus-

dactylus upper margins smooth. Pereopods 3^4 propodi with

transverse rows of robust setae; dactyli tapering, with longitudinal

row of robust setae. Pleopods 3-5, appendix interna absent.

Pleopod 1 of male absent or minute article. Pleopod 2 of male

without appendix interna, with appendix masculina. Uropodal

exopod with transverse suture. Telson without lateral fixed

spines and with posterolateral robust setae; apex rounded.

Remarks. The subgenus Paraxiopsis De Man, 1905 was

synonymised with Eutrichocheles Wood-Mason, 1876 by Sakai

and de Saint Laurent (1989) but later resurrected at full genus

status (Kensley, 1996a). Kensley (2003) brought the number of

species to fourteen. The two genera share several features, male

pleopod 1 lacking or reduced, absence of appendix interna on

pleopods 2-5, and similar gastric carinae. Kensley (1996a) listed

several characteristics distinguishing Paraxiopsis from

Eutrichocheles , notably the absence of a postcervical carina,

absence of a gape and tubercle on fingers of the cheliped, presence

of a spine and absence of a notch on the telson, and small size

(maximum cl. 8 mm). His differentiation was accepted with

minor discrepancies by Ngoc-Ho et al. (2005) who redescribed

E. modestus Wood-Mason, 1876, type species of Eutrichocheles.

Kensley (1996a) diagnosed Paraxiopsis as lacking male pleopod

1 but one male of P. pumilus displays minute digitiform male

pleopods 1 (see fig. 30e) as in species of Eutrichocheles (Ngoc-

Ho et al., 2005). The female pleopod 1 is typically a uniramous

appendage but in P. pauley i Kensley, 1996a, P. majuro Kensley,

1996a, P. austrinus (fig. 29b) and P. pumilus pleopod lisa minute

conical articulating projection similar to that seen in some males.

Kensley (1996a) noted the presence of an appendix masculina on

the male pleopod 2 (in spite of few species of the genus actually

being documented) and this is confirmed for P. pumilus below.

Paraxiopsis austrinus (Sakai, 1994)

Figure 29

Eutrichocheles austrinus Sakai, 1994: 185, figs. 6, 7.

Paraxiopsis austrinus.— Kensley, 2003: 373

Material examined. NT, Bullocky Point, Darwin, 12°26'S, 130°50'E, low

water rocky outcrops with muddy pools, AJ. Bruce, 3 Dec 1982 (stn

AJB-10), NTM 0003173 (2 females, cl. 8.3 mm, 6.8 mm; juvenile, cl.

3.4 mm). NT, Shell Island, Darwin, 12°30'S, 130°45'E, reef pools, D.

Sachs, 18 Mar 1988 (stn AJB-38), NTM 0006384 (female, cl. 10 mm).

Distribution. NT, Darwin region, 12°S, 130°E, low intertidal.

Remarks. Kensley (2003) differentiated this species from nine

other species of Paraxiopsis. It is recognisable by the presence

of a tomentum of long and short setae over much of the body

and limbs. The rostrum has three close-set spines on the lateral

margins running towards three more on the lateral gastric

carina. The submedian gastric carinae each has usually six

spines but in the female illustrated here there are eight on one

side only. Sakai based his species on two females only. One of

the females in the new collection, with gonopores on coxae 3,

has a minute pair of pleopods 1 while the others do not.

Paraxiopsis brocki (De Man, 1888)

Restricted synonymy.

Axius Brocki De Man, 1888: 475, pi. 20 fig. 3.

Axiopsis ( Paraxiopsis ) brocki.— De Man, 1925b: 101, pi. 8 figs.

19— 19f. — Poore and Griffin, 1979: 228, fig. 3.— Tirmizi, 1983: 88, fig.

3.— Morgan, 1990: 6, 63.

Eutrichocheles brocki.— Sakai and de Saint Laurent, 1989: 52, fig.

4B.— Ngoc-Ho, 1998: 365, fig. 1.

Paraxiopsis brocki.— Kensley, 1996a: 712, figs. 1, 2.— Poore,

2004: 176, figs. 45g, h, 46d.-Ngoc-Ho et al., 2005: 200.

Material examined. WA, near Mermaid Reef, 17°46.10'S,

120°43.15'E-17°45.95'S, 120°42.94'E (stn SS05-2007 097), 97-109 m,

20 Jun 2007, NMV J155708 (ovigerous female lacking most pereopods,

cl. 6 mm).

Distribution. Indo-West Pacific from Japan and Hawaii in

north, Tuamotu in east, through Indonesia, to southwestern

Australia, and to Kenya; to 100 m depth.

Remarks. Paraxiopsis brocki has been redescribed and

illustrated several times recently. It has been recorded from

WA and NT in Australia by Poore and Griffin (1979) and

Morgan (1990). The new specimen has prominent rostral spines

like that figured by Poore and Griffin (1979), more prominent

than on the specimen from Tuamotu figured by Ngoc-Ho

(1998). The female has prominent uniramous first pleopods.

Paraxiopsis pumilus (Sakai, 1994)

Figures 30, 31, 44

Eutrichocheles pumilus Sakai, 1994: 188-192, figs. 8, 9.

Australian Axiidae (Crustacea: Decapoda: Axiidea)

267

Figure 29. Paraxiopsis austrinus (Sakai, 1994). a, dorsal view of

carapace, peduncles of antennule and antenna, b, thoracic sternites 7

and 8, abdominal sternites 1 and 2 showing pleopods 1, 2. c, pleopod

2. All figures from female, cl. 8.3 mm, NTM Cr003173.

Paraxiopsis pumilus.— Kensley, 2003: 373

Paraxiopsis dianae Poore, 2008: 165-168: fig. 2. (syn. nov.)

Material examined. WA, off Barrow I., 20°59.05'S,

1 14°54.25'E-20°59.40'S, 114°54.32'E (stn SS10-2005 170), 101-100

m, 13 Dec 2005, NMV J53449 (male, cl. 8.3 mm, tl. 21.5 mm). WA,

Bonaparte Archipelago, Port George IV (15°23.474’S, 124°37.793’E),

10-16 m, 8 Oct 2007, J. James (stn P23), NMV J59647 (4 juvenile

females, cl. 4.8-6. 0 mm; female, cl. 7.3 mm)

NT. W side of Barrow Bay, Port Essington (11°22.0'S, 132°12.0’E),

low water, J.R.Hanley, 18 Sep 1985 (stn CPV8), NTM Cr013204 (1

female), NTM Cr013205 (1 female). Arafura Sea, 09°36.63'S,

134°10.95'E-09°36.59'S, 134°10.87'E (stn SS05-2005 BS014), 95 m,

25 May 2005, AM P74506 (1 juvenile).

Distribution. NT, Cobourg Penisula, Arafura Sea; WA, Barrow

Island— Dampier Archipelago, c. 9°-21°S, 115°-132°E, 6-100

m depth.

Remarks. The species is distinguished from other species of

Paraxiopsis by the possession (in adults) of two pairs of spines

at the base of the tapering rostrum and none on the submedian

gastric carina (Kensley, 2003). Poore (2008) compared his new

species, Paraxiopsis dianae, with several descriptions of the

similar species, P. brocki De Man, 1888, and concluded that

probably more than one species over a wide geographical range

had been referred to the latter name. He did not compare it with

P. pumilus (Sakai, 1994), described as a species of

Eutrichocheles, from the Northern Territory and north-western

WA. For this paper, topotypic material was compared with the

WA material. Sakai figured the tail fan, gastric region and

rostrum of two individuals of P. pumilus. The number of lateral

spines on the rostrum differed between these two and between

individuals in the new collections from WA and NT. On small

individuals lateral spines are absent, in others there is one spine

on one or both sides, in addition to the supraocular spine (fig.

31). The two individuals of P. dianae fall within this range of

variability and the species must be synonymised with P.

pumilus.

Most species of Paraxiopsis and Eutrichocheles possess a

bifid scaphocerite and illustrations of type material of both

Paraxiopsis pumilus and P. dianae show this. Some specimens

possess a simple comma-shaped scaphocerite on both antennae,

a difference we do not consider of specific importance (cf. figs.

30b, 31g, h). Paraxiopsis johnstoni Edmondson, 1925 from

Hawaii was also illustrated with a simple scaphocerite but has

a different carapace.

Type material of the two nominal Australian species was

collected at 6-40 depth; the new specimen is from 100 m

depth but at a similar latitude.

Pilbaraxius gen. nov.

Type species. Pilbaraxius kariyarra sp. nov., herein

designated.

Diagnosis. Carapace smooth or tuberculate; cervical groove

visible laterally over third distance to anterolateral margin.

Rostrum acutely triangular, with pair of lateral spines, longer

than eyestalks, not depressed below level of carapace, continuous

with definite lateral carinae; supraocular spines prominent;

lateral carina with 1 spine; submedian carina present, with 1

spine; median carina a weak ridge, unarmed; postcervical carina

absent. Abdominal somite 1 pleuron acute; pleuron 2 acute,

pleura 3-5 acute, with antero ventral tooth. Eyestalk cylindrical,

articulating; cornea weakly pigmented. Antenna, scaphocerite

long. Maxilliped 3 exopod not clearly bent at base of flagellum.

Pleurobranchs present above pereopods 2^4; podobranchs and

arthrobranchs well developed; epipods present on maxilliped 2

to pereopod 4. Pereopods 1 slightly asymmetrical, with propodus

flattened; carpus-propodus upper margin unarmed, sparsely

setose. Pereopods 3-5 propodi with transverse rows of robust

setae; dactyli tapering, with longitudinal row of robust setae.

Pleopods 3-5, appendix interna present. Pleopod 1 of male

absent. Pleopod 2 of male without appendix masculina. Uropodal

endopod without lateral and distolateral spines; exopod with

transverse suture. Telson with lateral fixed spines and

posterolateral robust setae; apex rounded.

268

G.C.B. Poore and D.J. Collins

Figure 30. Paraxiopsis pumilus (Sakai, 1994). a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c, telson and right

uropod. d, thoracic sternites 7 and 8, abdominal somite 1 with pleopods 1. e, left thoracic sternites 7 and 8, abdominal somite 1 with pleopod 1

(lateral), f, epistome, left antennule and antenna (ventral), g, maxilliped 3. h, major pereopod 1 (right, lateral), i, male pleopod 2. All figures from

male, cl. 8.3 mm, NMV J53449.

Australian Axiidae (Crustacea: Decapoda: Axiidea)

269

Figure 31. Paraxiopsis pumilus (Sakai, 1994). a-c, pereopods 2-4. d-h, anterior carapace, peduncles of antennule and antenna and rostrum of

five individuals, cl. 4.8, 4.8, 5.2, 6.0, 7.3 mm. Figs a-c from male, cl. 8.3 mm, NMV J53449; figs, d-e from male, 7.3 mm, NMV J59647.

Etymology. Pilbara is the name of the region of North-western

Australia close to the type locality of the type species.

Remarks. The problematic generic placement of the type

species highlights issues with the family Axiidae. The spinose

rostrum, diverging palm and long fingers on the chelipeds and

abdominal pleura with prominent ventral spination closely

resemble those of Calaxiopsis serrata Sakai and de Saint

Laurent, 1989, a species belong to the “calocaridid” group.

The new species differs from all in this group in possessing

pleurobranchs and lacking the modified pleopods 1 and 2. The

new species also resembles species of Calaxius but these

usually have highly setose chelipeds and rows of prominent

spines on the upper margin of the chelipeds (as in C. acutirostris

redescribed above). Neither of these conditions characterises

the new species. In addition, the uropodal endopod is apically

rounded and lacking lateral and distolateral spines typical of

Calaxius and many other axiid genera. Further, pleopod 2

lacks an appendix masculina, a state seen in few axiids. In

summary, a new genus seems warranted.

Pilbaraxius kariyarra sp. nov.

Calaxiopsis sp.— Robles et al., 2009: 314, 316 (molecular

phylogeny, GenBank numbers).

Figures 32, 33, 45

Material examined. Holotype. WA, off Port Hedland, 18°34.19'S,

1 17°27.86'E-18°34.06'S, 117°28.63'E (stn SS05-2007 052), 405-401

m, 14 Jun 2007, NMV J55576 (male, cl. 6.8, tl. 15.7 mm).

Description of male holotype. Carapace covered in small

tubercles. Rostrum 0.4 times length of front-to-cervical

groove, acute, elongate, with 1 long lateral spine anterior to

270

G.C.B. Poore and D.J. Collins

a, b 5 mm

Figure 32. Pilbaraxius kariyarra sp. nov. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c, telson and left uropod.

d, epistome, left antennule and antenna (ventral), e, major pereopod 1 (right, lateral), f, same (propodus-dactylus, mesial, setae not shown), g,

minor pereopod 1 (left, lateral), h, same (propodus-dactylus, mesial, setae not shown). All figures from holotype.

Australian Axiidae (Crustacea: Decapoda: Axiidea)

271

Figure 33. Pilbaraxius kariyarra sp. nov. a, thoracic sternites 7 and 8. b, maxilliped 3. c-f, pereopods 2-5 (c, f right; d, e left), g, pereopod 5 distal

propodus and dactylus. h, female pleopod 2. All figures from holotype.

supraocular spine, continuous with definite lateral gastric

carinae. Supraocular spines prominent. Lateral gastric carina

with 1 spine. Submedian gastric carina with 1 spine. Median

gastric carina present as a ridge extending anteriorly onto

rostrum, with 1 tubercle located midway between rostrum

and cervical groove. Postcervical carina on carapace absent.

Sternite 7 (pereopod 4) deeply divided in midline over

posterior two-thirds and with oblique lateral ridge ending in

sharp spine. Sternite 8 (pereopod 5) with setose semicircular

flap on anterior face at base of leg. Abdominal pleuron 1, 2.6

times as deep as middorsal length, with small ventral spine;

pleuron 2 asymmetrical, lateral length 1.3 times dorsal length,

concave ventrally, with 2 distinct ventral spines; pleura 3 and

4 tapering to a ventral spine, with another spine located

anteroventrally; pleuron 5 tapering to a ventral spine, with

another 2 spines located anteroventrally; pleuron 6 with 1

ventral spine.

Eyestalk 0.25 length of rostrum; cornea weakly pigmented.

Antennular peduncle reaching to end of article 4 of antenna.

Antennal article 1 with 2 small spines on distal margin; article

2 distal spine straight, approximately half length of article 2;

scaphocerite simple, straight, reaching distally almost to end

of article 4; article 3 with 1 spine on lower margin; article 4

about as long as article 2 (excluding distal spine); article 5

about half as long as article 4. Maxilliped 3 basis with 1 spine;

crista dentata with 15 teeth; merus with 2 spines on lower

margin (1 large, 1 small); carpus unarmed.

Pereopods 1 differentiated, propodus of major cheliped

longer and more swollen than minor. Major pereopod 1 (right)

coxa lower margin with 1 spine; basis lower margin unarmed;

ischium lower margin with 1 spine; merus upper margin

convex, with 1 hooked spine, lower margin with 1 spine and

obsolete tubercle laterally, lateral face with broad tubercle

distally, mesial face smooth; carpus upper margin tuberculate,

lower margin unarmed, lateral face tuberculate, mesial face

smooth; propodus upper margin with 1 distal spine, lower

margin with low lateral carina, lateral face tuberculate, mesial

face smooth; fixed finger 1.2 times length of upper palm,

cutting edge with 4 large irregular rounded teeth; dactylus

upper margin smooth, lateral face smooth, mesial face smooth,

cutting edge with narrow proximal notch.

Minor pereopod 1 coxa, ischium, merus and carpus as in

larger cheliped; propodus similar except mesial face with spine

near gape; fixed finger about as long as upper palm, cutting

edge with 2 large triangular teeth and smaller intermediate

denticles; dactylus cutting edge excavate proximally.

Pereopod 2 unarmed; carpus slightly shorter than chela;

propodus upper margin as long as dactylus. Pereopod 3

unarmed; propodus 2.2 times as long as dactylus, with 6

marginal robust setae (some duplicated). Pereopod 4 unarmed;

propodus 2.2 times as long as dactylus, with 7 marginal robust

setae (some duplicated). Pereopod 5 propodus 3.9 times as

long as dactylus, subchelate, with short fixed finger bearing 5

distinct robust setae; dactylus slightly flattened.

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G.C.B. Poore and D.J. Collins

Pleurobranchs present above pereopods 2-4; arthrobranchs

on maxilliped 2 (rudimentary) to pereopod 4; epipods with

well developed podobranchs (with up to 10 gill filaments) on

maxilliped 2 to pereopod 3

Pleopod 1 absent. Pleopod 2 appendix masculina absent;

appendix interna slender, about quarter length of endopod.

Telson 1.3 times as long as broad, lateral margin with 4

spines, distal margin convex without posteromedian spine,

posterolateral angle with 1 robust seta; dorsal face without

spines on each oblique row. Uropodal endopod 1.9 times as

long as wide, without lateral spines, longitudinal ridge

unarmed. Uropodal exopod 1.9 times as long as wide, without

lateral spines, longitudinal ribs unarmed, posterolateral angle

with 1 robust seta; transverse suture unarmed.

Etymology. Kariyarra is the name of the Australian Aboriginal

people inhabiting the coast close to the type locality (noun in

apposition).

Distribution. WA, off Port Hedland, 18°S, 118°E, c. 400 m

depth (known only from type locality).

Remarks. See the discussion for the genus Pilbaraxius.

Unfortunately, the species was placed in a molecular analysis

as Calaxiopsis sp. by Robles et al. (2009).

Planaxius Komai and Tachikawa, 2008

Planaxius Komai and Tachikawa, 2008: 22-24.

Remarks. Planaxius is recognised by the combination of the

absence of submedian gastric carinae, uropodal exopod with a

transverse suture, pereopodal epipods present, three pairs of

pleurobranchs, toothed triangular rostrum, male pleopods 1 and

2 present and pleopods 3-5 with an appendix interna. It is the

absence of submedian gastric carinae and presence of appendices

intemae that distinguishes the genus from Bouvieraxius whose

species have five gastric carinae and lack appendices internae

(Komai and Tachikawa, 2008). These authors described the

type and only species as lacking a median gastric carina. The

single individual from Australia referred to this species below

has a definite median gastric carina at the base of the rostrum

extending as far back as the lateral gastric carinae.

Planaxius brevifrons Komai and Tachikawa, 2008

Planaxius brevifrons Komai and Tachikawa, 2008: 24-29, figs. 2-6.

Figures 34, 46

Material examined. WA, off Jurien Bay, 29°48.33'S,

114 0 25.52'E-29°48.33'S, 114°25.55'E(stn SS10-2005 083), 113-114 m,

02 Dec 2005, NMV J55445 (male, cl. 3.3 mm, tl. 9.5 mm, without

pereopods 3-5).

Description of male. Carapace smooth, cervical groove short.

Rostrum 0.3 times length of front-to-cervical groove, triangular,

with 2 or 3 short oblique lateral spines anterior to supraocular

spine, continuous with definite lateral gastric carinae.

Supraocular spines prominent. Lateral gastric carina unarmed.

Submedian gastric carina absent. Median gastric carina low,

sharp, unarmed. Branchiostegal angle produced, lobe-like,

with 2 minute marginal teeth. Sternite 8 (pereopod 5) with

setal ridge on anterior face at base of leg. Abdominal pleuron 1,

3 times as deep as middorsal length, ventrally acute; pleuron 2

broad, lateral length 1.8 times dorsal length, anteroventrally

rounded; pleura 3-5 becoming more posteroventrally square,

each with anteroventral tooth; pleuron 6 with small tooth on

ventral margin.

Eyestalk 0.5 length of rostrum; cornea pigmented.

Antennular peduncle reaching to proximal part of antennal

article 5. Antennal article 1 with 2 spinules on lower distal

margin; article 2 distal spine slender, directed slightly inwards,

reaching distally one third of antennal article 4; scaphocerite

slender, straight, reaching distally two-thirds of article 4; article

3 with 1 spine on mesial lower margin; article 4 about 1.3 times

length of article 2 (excluding distal spine), article 5 about one-

third length of article 4. Maxilliped 3 basis with 1 spine; ischium

with 3 spines on lower margin; crista dentata with 13 teeth;

merus with 4 spines on lower margin; carpus with 1 spine.

Pereopods 1 asymmetrical, propodus of major longer,

more swollen. Major pereopod 1 (right) coxa lower margin

with 1 spine; basis lower margin unarmed; ischium lower

margin with 3 spines; merus upper margin convex, with 2

spines, lower margin with 6 spines, lateral face smooth, mesial

face smooth; carpus unarmed, smooth; propodus upper margin

carinate, with small distal spine, lower margin smooth, lateral

face smooth, mesial face smooth; fixed finger half length of

upper palm, cutting edge unevenly denticulate; dactylus

smooth, cutting edge as in fixed finger.

Minor pereopod 1 coxa as in larger cheliped; ischium

lower margin with 1 spine; merus upper margin convex, with

2 spines, lower margin with 6 spines, lateral face smooth,

mesial face smooth; carpus unarmed, smooth; propodus upper

margin carinate, with small distal spine, lower margin smooth,

lateral face smooth, mesial face smooth; fixed finger as long as

upper palm, cutting edge unevenly denticulate; dactylus

smooth, cutting edge scarcely denticulate.

Pereopod 2 ischium lower margin unarmed; merus lower

margin unarmed; carpus as long as chela; propodus upper

margin 0.8 length of dactylus. Pereopods 3-5 missing.

Pleopod 1 of 2 fused articles; article 2 blade-like, apex

unevely bilobed, appendix interna represented by few hooks.

Pleopod 2 with appendix masculina as long as remaining

endpod; appendix interna third length of endopod, 0.6 length

of appendix masculina. Pleopods 2-5 appendix interna one

third length of endopod.

Telson 1.5 times as long as wide, lateral margin with 3

spines, distal margin convex without posteromedian spine,

posterolateral angle with 2 robust setae, one much larger; dorsal

face with 2 small spines in each oblique row. Uropodal endopod

1.8 times as long as wide, with 1 lateral spine, longitudinal

ridge with 4 spines (including marginal). Uropodal exopod 1.6

times as long as wide, with 4 lateral spines, 2 longitudinal ribs

(outer rib with 2 spines), posterolateral angle with 1 fixed spine

and 1 robust seta; transverse suture with 4 spines.

Distribution. Japan, Kii Peninsula, Ogasawara Is, 47-100 m;

WA, south-western coast, 114 m depth.

Remarks. The single male is virtually indistinguisable from

slightly larger specimens reported from Japan by Komai and

Australian Axiidae (Crustacea: Decapoda: Axiidea)

273

Figure 34. Planaxius brevifrons Komai and Tachikawa, 2008. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c,

telson and right uropod. d, sternites 7 and 8, pleonite 1 with left pleopod 1 in situ, e, sternites 7 and 8, pleonite 1 with left pleopod 1 in situ

(lateral), f, epistome, right antennule and antenna (ventral), g, maxilliped 3. h, major pereopod 1 (right, lateral), i, minor pereopod 1 (left), j,

pereopod 2. k, 1, male pleopods 1, 2. All figures from NMV J55445.

274

G.C.B. Poore and D.J. Collins

Tachikawa (2008). Dr Komai kindly re-examined the type

material at our request and confirmed the presence of a median

gastric carina, more clear and sharp in the paratypes than in the

illustrated holotype, as in the WA specimen. While the ischium

of maxilliped 3 of the Australian specimen has blunt teeth

along the lower margin some of the Japanese specimens have

one or two minute spinules obscured by stiff setae. Dr Komai

confirmed that the two minute teeth on the anterior

branchiostegal lobe of the Australian specimen are absent in

the Japanese material. The uropodal endopod of the holotype

lacks the lateral and distolateral spines seen in the Australian

specimen. While we have only one specimen, none of these

differences argues strongly for a separate species but the record

is a considerable range extension.

Platyaxius Sakai, 1994

Platyaxius Sakai, 1994: 180-181.

Diagnosis. Carapace smooth, with scattered long setae; cervical

groove visible laterally over half or more of distance to

anterolateral margin. Rostrum triangular, broad, laterally

weakly dentate, as long as or slightly longer than eyestalks,

slightly depressed below level of carapace, continuous with

definite lateral carinae; supraocular spines absent; lateral carina

weakly dentate or unarmed; submedian carina dentate; median

carina, dentate; postcervical carina absent. Abdominal somite

1 pleuron triangular; pleuron 2 posteriorly rectangular; pleura

3-5 posteriorly rounded. Eyestalk cylindrical, articulating;

cornea pigmented. Antenna, scaphocerite short. Maxilliped 3

exopod not clearly bent at base of flagellum. Pleurobranchs

absent above pereopods 2-4; podobranchs and arthrobranchs

well developed; epipods present on maxilliped 2 to pereopod 4.

Pereopods 1 asymmetrical, with propodus cylindrical; carpus-

dactylus upper and lower margins smooth, propodus with

obsolete distal tooth on upper margin. Pereopods 3-5 propodi

with transverse rows of robust setae; dactyli 3 and 4 triangular,

with scattered robust setae; dactylus 5 spatulate, with row of

robust setae along oblique margin. Pleopods 3-5, appendix

interna present. Pleopod 1 of male absent. Pleopod 2 of male

with appendix interna and appendix masculina. Uropodal

exopod without transverse suture. Telson with lateral teeth,

with posterolateral robust setae; apex deeply rounded and

continuous with lateral margins.

Remarks. The type species, Platyaxius brevirostris Sakai, 1994

from the Australian North West Shelf, is recognised by the

absence of a suture on the uropodal exopod, absence of the

male pleopod 1 and lateral teeth on the rostrum. Sakai compared

his new genus to species of Eiconaxius Bate, 1888 (see Sakai,

1992 for descriptions of five species) and the two genera share

similar uropods, pleopods, swollen chelipeds and a rounded

telson, differing largely in the dentition of the rostrum. The

dactyli of pereopods 3-5 of species of Eiconaxius are spatulate,

non-tapering and with a row of robust setae on the oblique

distal margin. See, for example, E. farreae Ortmann, 1891

figured by Sakai and Ohta (2005) and E. mallacoota sp. nov.

(fig. 24). The same unusual form is seen only on pereopod 5 of

the new species described here; dactyli of pereopods 3 and 4

are tapering with several facial robust setae. Sakai (1994)

illustrated only pereopod 3 of P. brevirostris and did not

mention pleurobranchs in his diagnosis. The new species

described below differs from all species of Eiconaxius in

lacking pleurobranchs above pereopods 2-4.

Axius odontorhynchus De Man, 1905 was included in

Spongiaxius by Sakai and de Saint Laurent (1989) but is better

placed in Platyaxius. It too has denticulate rostrum margins,

swollen chelipeds and non-tapering dactylus only on pereopod

5 (De Man, 1925b: pi. 1 figs. 11, m).

Sakai and Ohta (2005) removed Eiconaxius from Axiidae

and placed it in its own family, Eiconaxiidae, on the basis of

‘P3-5 dactyli . . . rounded in shape . . . and their ventral margins

spinulate ...’ and other features that are shared with one or

more other axiid genera. Eiconaxius and Platyaxius are

clearly related but whether or not they belong in a clade

separate from another containing all other axiids remains to

be investigated.

Platyaxius bardi sp. nov.

Figures 35, 36, 47

Material examined. Holotype. WA, near Mermaid Reef, 17°29.23'S,

120 o 27.64'E-17°29.72'S, 120°28.07'E (stn SS05-2007 091), 187-184

m, 20 Jun 2007, NMV J55707 (male, cl. 5.9 mm, tl. 17.2 mm).

Paratypes. WA, North-west Shelf, between Port Hedland and

Dampier, 18°36'S, 118°02'E-18°39'S, 118°04'E (stn NWA-24), 184 m,

06 Jun 1983, NMV J15420 (2 ovigerous females, cl. 5.7 mm).

Description of male holotype. Carapace smooth. Rostrum 0.3

times length of front-to-cervical groove, narrowly triangular,

with 4 weak lateral spines anterior to supraocular spine,

continuous with definite lateral gastric carinae. Supraocular

spines similar size to other rostrum spines. Lateral gastric

carina with 2 prominent spines in addition to supraocular spine.

Submedian gastric carina with 4 spines. Median gastric carina

with 6 spines. Sternite 7 (pereopod 4) deeply divided in midline

over posterior two-thirds and with sharp oblique lateral ridge.

Sternite 8 (pereopod 5) with setose semicircular flap on anterior

face at base of leg. Abdominal pleuron 1, 2.6 times as deep as

middorsal length, ventrally acute; pleuron 2 asymmetrical,

lateral length 1.2 times dorsal length, posteroventrally quadrate;

pleura 3-5 posteroventrally angled, each with small

anteroventral spine; pleuron 6 with small spine on ventral

margin.

Eyestalk 0.8 length of rostrum; cornea pigmented.

Antennular peduncle reaching to distal part of antennal article

4. Antennal article 1 with 2 spinules on lower distal margin;

article 2 distal spine well developed, reaching to proximal part

of antennal article 5; scaphocerite strongly curved downwards,

reaching distally almost to end of article 4; article 3 with spine

on lower margin; article 4 approximately as long as article 2

(excluding distal spine); article 5 about two-thirds length of

article 4. Maxilliped 3 basis with 1 spine; ischium with 1 spine

on lower margin; crista dentata with numerous small, even

teeth; merus with 3 spines on lower margin; carpus unarmed.

Pereopods 1 differentiated, propodus of major cheliped

longer and more swollen than minor. Major pereopod 1 (left)

coxa lower margin with 1 spine; basis lower margin unarmed;

Australian Axiidae (Crustacea: Decapoda: Axiidea)

275

Figure 35. Platyaxius bardi sp. nov. a, lateral view, b, dorsal view of carapace, peduncles of antennule and antenna, c, thoracic sternites 6-8. d,

telson and right uropod. e, major pereopod 1 (left, lateral), f, minor pereopod 1 (right, lateral). All figures from holotype.

276

G.C.B. Poore and D.J. Collins

Figure 36. Platyaxius bardi sp. nov. a, maxilliped 3. b-e, pereopods 2-5 (b, e right; c, d left; each with details of dactylus). f, male pleopod 2.

All figures from holotype.

Australian Axiidae (Crustacea: Decapoda: Axiidea)

277

ischium lower margin with 4 small spines and 1 larger; merus

upper margin strongly convex, unarmed, lower margin with 10

teeth, lateral face smooth, mesial face smooth; carpus upper

margin carinate, lower margin unarmed, lateral face smooth,

mesial face smooth; propodus upper margin carinate, with

blunt distal tooth, lower margin with weak lateral carina,

better defined on finger, lateral face tuberculate over upper and

lower distal third, mesial face tuberculate near base of finger;

fixed finger 0.6 length of upper palm, cutting edge straight,;

dactylus margins and faces smooth, cutting edge with few

blunt teeth.

Minor pereopod 1 coxa, ischium, merus and carpus as in

larger cheliped, narrower; propodus upper margin carinate,

with blunt distal tooth, lower margin with weak lateral carina,

better defined on finger, lateral face with few tubercles, mesial

face with few tubercles near base of finger; fixed finger as long

as upper palm, cutting edge with regularly spaced sharp teeth;

dactylus margins and faces smooth, cutting edge denticulate.

Pereopod 2 ischium lower margin unarmed; merus lower

margin unarmed; carpus slightly longer than chela; propodus

upper margin 0.4 length of dactylus. Pereopod 3 merus lower

margin with 3 weak spines; propodus 2.9 times as long as

dactylus, with 8 transverse rows each of 4-7 robust setae;

dactylus with 13 robust setae on inner face plus unguis.

Pereopod 4 propodus 4.1 times as long as dactylus, with 8

transverse rows each of 3-8 robust setae; dactylus with 12

robust setae on inner face plus unguis. Pereopod 5 propodus

3.5 times as long as dactylus, weakly subchelate, distally with

2 transverse rows each of 2 or 3 robust setae; dactylus with 6

robust setae on distal margin, 3 robust setae on inner face.

Pleopods 2-5 appendix interna one third length of endopod.

Pleopod 2 appendix masculina as long as appendix interna.

Telson ovate, slightly tapering posteriorly, 1.1 times as

long as wide, lateral margin with 3 spines, distal margin

convex with posteromedian spine, posterolateral curve with 1

robust seta; dorsal face with 2 spines in each oblique row.

Uropodal endopod 1.7 times as long as wide, with 7 lateral

spines, longitudinal ridge with 4 spines (including marginal).

Uropodal exopod 1.6 times as long as wide, with 11 lateral

spines, 2 longitudinal ribs (inner rib ending in marginal spine,

outer rib with 7 spines), posterolateral angle with 1 fixed spine

and 1 robust seta; transverse suture absent.

Female (2 paratypes). As male except: rostrum with 4-6

lateral spines anterior to supraocular spine; lateral gastric

carina with 2 or 3 prominent spines in addition to supraocular

spine; submedian gastric carina with 4 or 5 spines; median

gastric carina with 5-7 spines; pleura 3-5 posteroventrally

angled, each with small anteroventral spine (as male); pleuron

6 with small spine on ventral margin; major and minor

chelipeds fixed finger and dactylus with lateral ridge; pleopod

1 uniramous, article 2, 3 times length of article 1; ovigerous

with at least 6 eggs.

Etymology. Bardi is the name of the Aboriginal Australian

people inhabiting Cape Leveque close to the type locality (noun

in apposition).

Distribution. WA, continental slope of North-west Shelf,

17°-19°S, 118°-120°E, 180-184 m depth.

Remarks. The three individuals of Platyaxius bardi vary in the

numbers and size of spines/teeth on the rostrum and gastric

carinae. The species differs from P. brevirostris and P.

odontorhynchus (from Indonesia) in the more pronounced

dentition on the rostrum and gastric carina (obsolete in the other

two). The rostrum exceeds the eyestalks in the new species (does

not in the other two species) and uropodal rami are broader. The

scaphocerite of P. bardi is strongly curved while it is straight in

the other two species. Platyaxius bardi and P. brevirostris have

both been taken from similar limited latitudes and depths on the

north-western Australian shelf. Platyaxius bardi has teeth on the

lateral gastric carina while P. brevirostris does not.

Platyaxius brevirostris Sakai, 1994

Platyaxius brevirostris Sakai, 1994: 181-185, figs. 4, 5.— Davie,

2002: 454.

Distribution. WA, North West Shelf, 141 m depth.

Remarks. See comments under P. bardi above.

Scytoleptus Gerstaecker, 1856

Remarks. The steep margin of the median gastric carina, falling

almost vertically to the short rostrum, and with three small

teeth, immediately differentiate the genus and its only species.

Scytoleptus serripes Gerstaecker, 1856

Scytoleptus serripes Gerstaecker, 1856: 158, pi. 6 figs. 1-4. De

Man, 1925b: 49, pi 4 figs. 9-9h.-Poore and Griffin, 1979: 243-245,

fig. 11.— Sakai and de Saint Laurent, 1989: 37-39, fig. 9.— Sakai, 1994:

200.— Davie, 2002: 454.

Distribution. Indo-West Pacific, NT, WA, northern and central

coast, to 36 m depth

Remarks. Scytoleptus serripes is recognised by the characteristic

median gastric carina that defines the genus.

Spongiaxius Sakai and de Saint Laurent, 1989

Spongiaxius Sakai and de Saint Laurent, 1989: 41.— Sakai and

Ohta, 2005: 88-89.

Sakaiocaris Kensley, 1989: 964 (objective synonym: same type

species).

Remarks. The rostrum with its margin of erect spines, separated

by a constriction from the carapace, serves to recognise species

of Spongiaxius.

Spongiaxius brucei (Sakai, 1986)

Axiopsis brucei Sakai, 1986: 12-20, figs. 1-6.

Spongiaxius brucei.— Sakai and de Saint Laurent, 1989:

44-45.— Davie, 2002: 454.— Sakai and Ohta, 2005: 89-90, fig.

11.— Poore, 2008: 168.

Sakaiocaris brucei.— Kensley, 1989: 964-965.

Distribution. WA, slope of North West Shelf, Sulu Sea,

450-690 m depth.

Remarks. Spongiaxius brucei is a large species with erect

spines on the five gastric carinae, the carinae themselves

278

G.C.B. Poore and D.J. Collins

separated by a constriction from the elevated rostrum

surrounded by about 20 erect spines. The massive dactylus of

the major cheliped is also distinctive. Unlike most axiids from

deep water, numerous specimens have been taken, including

more than those already reported in NTM collections.

Biogeographical commentary

In earlier reviews, Poore and Griffin (1979) recognised only

six species of Axiidae and Sakai (1994) thirteen (plus species

now placed in Strahlaxiidae). Now, 30 are known with others

probable. Eleven of these have also been reported from

elsewhere in the Indo-West Pacific, some as far away as Japan

or Madagascar (Table 1). These include both shallow-water,

shelf and deep-water species whose distribution in WA range

as far south as 35°S. This generalisation is subject to the

proviso that identifications of Australian specimens as

species known from elsewhere depend on subjective

judgements of morphological similarity between material at

hand and published descriptions. Poore (2008) has already

tabulated differences between six published descriptions of

the seemingly widespread Paraxiopsis brocki. A further 12

species are now known only from tropical Australia, most

from WA (Table 1). Nine are known only from the type

locality. It is reasonable to assume that some of these could

subsequently be found north of Australia. Nine species are

found in southern temperate Australia south of 30°S (Table

1). Two of these are Indo-West Pacific species and another

occurs also in New Zealand. Of the southern species six

could be treated as endemic. Three are shallow-water species

from south-eastern Australia reported from narrow

geographic ranges. The other three are species from the

continental slope taken only once. The distribution of these

is unknown until these burrowing cryptic animals from deep

water can be reliably sampled. Three genera, monotypic

Australocaris and Dorphinaxius, and Michelaxiopsis with

two species, are confined to southern Australia-New Zealand.

Most others are widespread in the Indo-West Pacific. Axius

and Calastacus, with species in the North Atlantic and

southern Australia appear to have anomalous global

distributions.

Acknowledgements

Some of the collections on which this project was based were

made during expeditions organised by CSIRO Marine and

Atmospheric Research (CMAR). We thank co-Principal

Investigators, Alan Williams and Rudy Kloser, CMAR, for the

organisation and management of the voyages, and to Karen

Gowlett-Holmes and Mark Lewis for help on board RV

Southern Surveyor. For financial support we acknowledge the

Commonwealth Department of the Environment and the

CSIRO Wealth from Oceans Flagship that provided funds for

the field and laboratory components of the “Voyages of

Discovery” program, and the Commonwealth Environmental

Research Facilities (CERF) Marine Biodiversity Hub. We

thank Jane Jelbart, University of Newcastle, Newcastle, Buz

Wilson and Helen Stoddart, Australian Museum, Sydney,

Suzanne Horner, Museums and Art Gallery of the Northern

Territory, Darwin, Thierry Laperousaz, South Australian

Museum, Adelaide, and Miranda Lowe, Natural History

Museum, London, for the loan of material. At Museum

Victoria, we appreciate the efforts of Joanne Taylor in

organising and registering the material and of Anna McCallum

in providing new animals for study.

We thank too Colin McLay, Christchurch, Simon Pollard,

Canterbury Museum, Christchurch, Rick Webber, National

Museum of New Zealand (Te Papa), and Shane Ahyong,

National Institute of Water and Atmosphere, Wellington, who

attempted to chase down types and other material in New

Zealand.

This paper benefited from discussions between the first

author and the late Michele de Saint Laurent in the 1990s and

some of her thoughts are incorporated here. We are grateful

too for valuable comments on a first draft by Tomoyuki Komai,

Chiba, Japan, and for his willingness to re-examine specimens

in his care.

This paper is a contribution to the Census of Marine Life

project COMARGE (Continental Margins Ecosystems).

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Poore, G.C.B. 2008. Thalassinidean shrimps (Crustacea: Decapoda)

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Decapoda) of Australia. Records of the Australian Museum 32:

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Australia: preliminary identifications of 524 species from FRV

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Robles, R., Tudge, C.C., Dworschak, P.D., Poore, G.C.B., and Felder,

D.L. 2009. Molecular phylogeny of the Thalassinidea based on

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Crandall, K.A., and Felder, D.L. (eds). Crustacean Issues Vol. 18:

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Gascogne, Calastacus laevis sp. nov. Remarques sur le genre

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Museum National d’Histoire Naturelle, Paris (2e serie) 35:

347-356.

de Saint Laurent, M. 1973. Sur la systematique et la phylogenie des

Thalassinidea: definition des families des Callianassidae et des

Upogebiidae et diagnose de cinq genres nouveaux. Comptes Rendus

Hebdomadaires de Seances de TAcademie des Sciences, Paris 277:

513-516.

Sakai, K. 1967. Three new species of Thalassinidea (Decapoda,

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Sakai, K. 1987. Two new Thalassinidea (Crustacea: Decapoda) from

Japan, with the biogeo graphical distribution of the Japanese

Thalassinidea. Bulletin of Marine Science 41: 296-308.

Sakai, K. 1992. Axiid collections of the Zoological Museum, Copenhagen,

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Australian Axiidae (Crustacea: Decapoda: Axiidea)

281

Table 1. Distributions of Australian Axiidae. Species are divided into three groups: (1) those distributed elsewhere in the Indo-West Pacific; (2)

those known only from northern tropical and subtropical Australia; and (3) those known from southern temperate Australia.

Taxa grouped by distribution and ranked Depth range (m)

by depth range

Australian distribution: state and southern

(Indo-West Pacific and subtropical species) or

northern (temperate species) latitudinal limit

Indo-West Pacific

Allaxius clypeatus

Qld, 15°S

Axiopsis serratifrons

0-30

Qld, 15°S

Scytoleptus serripes

0-36

NT, WA, 22°S

Paraxiopsis brocki

0-100

WA, 32°S

Bouvieraxius keiensis

18-245

WA, 27° S

Planaxius brevifrons

47-114

WA, 30°S

Axiopsis consobrina

75-113

WA, 19°S

Axiopsis tsushimaensis

102-157

WA, 35°S

Acanthaxius clevai

228-438

WA, 17° S

Calaxius acutirostris

325-505

Qld, 27° S

Spongiaxius brucei

450-690

WA, 17° S

Tropical and subtropical Australia only

Paraxiopsis austrinus

NT, 12°S

Paraxiopsis pumilus

6-100

NT, WA, 9°-21°S

Acanthaxius gawara

49-59

Qld, 12°S

Oxyrhynchaxius manningi

134

WA, 19°S

Platyaxius brevirostris

141

WA, 19°S

Eiconaxius kimbla

150

Qld, 27° S

Acanthaxius ningaloo

165

WA, 22°S

Platyaxius bardi

180

WA, 19°S

Acanthaxius polychaetes

260

Qld, 18°S

Pilbaraxius kariyarra

400

WA, 18°S

Acanthaxius gathaagudu

400-450

WA, 17-26° S

Ambiaxius franklinae

1300

WA, 17° S

Temperate only

Dorphinaxius kermadecensis

0-8

New Zealand, NSW, 32-34°S

Michelaxiopsis australiensis

0-5

Central NSW, Vic., 29-38°S

Michelaxiopsis nauo

5-6

SA, 34° S

Axius werribee

2-25

Tas., Vic. SA, 35°S

Australocaris pinjarup

400

WA, 33°S

Calastacus myalup

400

WA, 33°S

Eiconaxius mallacoota

930-1000

Vic., 38°S

282

G.C.B. Poore and D.J. Collins

Figure 37. Acanthaxius clevai Ngoc-Ho, 2006. Photomicrographs of (left to right) maxilla 1, maxilla 2, maxilliped 1, maxilliped 2.

Figure 38. Acanthaxius ningaloo sp. nov. Photomicrographs of (left to right) maxilla 1, maxilla 2 (posterior epipod seta missing), maxilliped 1,

maxilliped 2.

Australian Axiidae (Crustacea: Decapoda: Axiidea)

283

Figure 39. Australocaris pinjarup gen. and sp. nov. Photomicrographs of (left to right) maxilla 1, maxilla 2 (posterior epipod seta truncated),

maxilliped 1, maxilliped 2.

Figure 40. Axiopsis tsushimaensis Sakai, 1992. Photomicrographs of (left to right) maxilla 1, maxilla 2 (posterior epipod seta truncated),

maxilliped 1, maxilliped 2.

284

G.C.B. Poore and D.J. Collins

Figure 41 . Bouvieraxius keiensis Sakai, 1992. Photomicrographs of (left to right) maxilla 1, maxilla 2 (posterior epipod seta truncated), maxilliped

1, maxilliped 2.

0,5 mm

Figure 42. Calastacus myalup sp. nov. Photomicrographs of (left to right) maxilla 1, maxilla 2 (posterior epipod seta missing), maxilliped 1,

maxilliped 2.

Australian Axiidae (Crustacea: Decapoda: Axiidea)

285

Figure 43. Calaxius acutirostris Sakai and de Saint Laurent, 1989. Photomicrographs of (left to right) maxilla 1, maxilla 2 (posterior epipod seta

truncated), maxilliped 1, maxilliped 2.

Figure 44. Paraxiopsis pumilus (Sakai, 1994). Photomicrographs of (left to right) maxilla 1, maxilla 2 (posterior epipod seta truncated),

maxilliped 1, maxilliped 2.

286

G.C.B. Poore and D.J. Collins

Figure 45. Pilbaraxius kariyarra sp. nov. Photomicrographs of (left to right) maxilla 1, maxilla 2 (posterior epipod seta truncated), maxilliped

1, maxilliped 2.

Figure 46. Planaxius brevifrons Komai and Tachikawa, 2008. Photomicrographs of (left to right) maxilla 1, maxilla 2 (posterior epipod seta

truncated), maxilliped 1, maxilliped 2.

Australian Axiidae (Crustacea: Decapoda: Axiidea)

287

Figure 47. Platyaxius bardi sp. nov. Photomicrographs of (left to right) maxilla 1, maxilla 2 (posterior epipod seta missing), maxilliped 1,

maxilliped 2.

Memoirs of Museum Victoria 66: 289-329 (2009)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://museumvictoria.com.au/About/Books-and-Journals/Journals/Memoirs-of-Museum-Victoria

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

Ulf Scheller

Haggeboholm, Haggesled, S-53194 Jarpas, Sweden (ulf.scheller@telia.com)

Abstract Scheller, U. 2009. New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests. Memoirs of Museum

Victoria 66: 289-329.

Seventeen species new to science belonging to five genera in two families are described from a collection of

Pauropoda made during an intensive survey of temperate rainforest in Tasmania. The new species are Allopauropus

fraterculus sp. nov., Allopauropus inusitatus sp. nov., Decapauropus heis sp. nov., Decapauropus attenuatus sp. nov.,

Decapauropus ungulatus sp. nov., Decapauropus convexus sp. nov., Decapauropus saltuarius sp. nov., Decapauropus

terrestris sp. nov., Nesopauropus tasmaniensis sp. nov., Stylopauropoides erectus sp. nov., Stylopauropoides rounsevelli

sp. nov., Stylopauropoides quadripartitus sp. nov., Stylopauropoides scissus sp. nov., Stylopauropoides hetaeros sp. nov.,

Stylopauropoides eximius sp. nov., Pauropus vandiemi sp. nov. and Borneopauropus dignus sp. nov. The high level of

local endemism in Tasmanian rainforest is emphasised.

Keywords biodiversity, Allopauropus, Decapauropus, Nesopauropus, Stylopauropoides, Pauropus, Borneopauropus, Australia,

endemism

Introduction

An intensive survey of invertebrates of rainforest in Tasmania

was carried out between 1989 and 1990 funded by the National

Rainforest Conservation Programme (Coy et al., 1993). Several

higher taxa collected during the survey were distributed to

specialist taxonomists for study. A large number of Pauropoda

formed part of this collection. It contained seventeen new

species and two already described species (Greenslade, 2008).

The current paper describes the new species from this collection

and reports the collecting sites for all the species studied. Other

Arthropoda were described or recorded by Clark and

Greenslade (1996) and Greenslade (2008).

The Pauropoda are a little known group of soil organisms

being small, cryptic Myriapoda but they are widespread in

distribution and can be abundant (Scheller, 1990). The fauna

often exhibits a high level of local endemism in forest soils as

is demonstrated by the collection described here. Pauropods

can inhabit strata from litter to the subsoil in a variety of plant

communities and soil types, even agricultural habitats

(Scheller, 1990).

Materials and Methods

Specimens were collected from 19 rainforest sites in different

regions of Tasmania and at different altitudes. The sites

covered four different rainforest types. A number of collection

methods were used to collect specimens. They included funnel

extraction of leaf litter, soil and moss, pitfall trapping and

pyrethrin knockdown from tree trunks. The collecting methods

and sites are described in detail by Coy et al. (1993) and

documented by Greenslade (2008) as localities 1 to 19.

All specimens were preserved in ethanol. In the descriptions

below, individuals have been classified as adults, subadults

and juveniles according to the number of pairs of legs. The sex

of adults and subadults was recorded.

Holotypes have been lodged in the Australian National

Insect Collection, CSIRO, Canberra; paratypes and other

material are deposited in the Queen Victoria Museum,

Launceston, Tasmania.

Abbreviations

Deposition: ANIC, Australian National Insect Collection,

CSIRO, Canberra, ACT, Australia; QVM, Queen Victoria

Museum, Launceston, Tasmania, Australia.

Morphology: ad., adult; subad., subadult; juv., juvenile

with number of pairs of legs as indicated.

Collectors: ATW, A.Trumbull-Ward; DR, D. Rounsevell;

HM, H. Mitchell; JD, J. Diggle; MN, M. Neyland; PG, P.

Greenslade; RC, R. Coy; SS, S. Smith.

GR - grid reference; NRCP, National Rainforest

Conservation Programme; PKD, pyrethrin knock down

collecting method.

Systematics

The characters studied to identify species are those used by

previous workers, in particular Hansen (1902), Remy (1931,

1952a, 1956c) and Scheller (1985, 1988, 1993). The setal

290

U. Scheller

nomenclature used here follows these publications. All setal

names are in italics. The length of body, excluding the

antennae, is given and the range of variation in adult paratypes

(subadult in Decapauropus convexus ) is provided in brackets

in mm and other measurements are given in micrometres. For

most measurements the range for adult paratypes is given in

brackets either before (lower figure) or after (higher figure) the

mean measurement. Lengths and ratios of lengths of setae are

given in the descriptions. Distances between setae are given as

eg. a-a=... if distance between same seta on left and right

side of body is measured and as eg. a-a 2 - if distance between

setae on the same side of the body is measured. The number of

pairs of legs, either nine or less, is given before the specimen

details.

Checklist of Tasmanian Pauropoda

Pauropodidae

Allopauropus Silvestri, 1902

Allopauropus fraterculus sp. nov.

Allopauropus inusitatus sp. nov.

Decapauropus heis sp. nov.

Decapauropus attenuatus sp. nov.

Decapauropus ungulatus sp. nov.

Decapauropus convexus sp. nov.

Decapauropus saltuarius sp. nov.

Decapauropus terrestris sp. nov.

Nesopauropus Scheller, 1997

Nesopauropus tasmaniensis sp. nov.

Stylopauropoides Remy, 1956

Stylopauropoides ringueleti Remy, 1962

Stylopauropoides erectus sp. nov.

Stylopauropoides rounsevelli sp. nov.

Stylopauropoides quadripartitus sp. nov.

Stylopauropoides scissus sp. nov.

Stylopauropoides hetaeros sp. nov.

Stylopauropoides eximius sp. nov.

Pauropus Lubbock, 1867

Pauropus dolosus Remy, 1956

Pauropus vandiemeni sp. nov.

Brachypauropodidae

Borneopauropus, Scheller, 1994

Borneopauropus dignus sp. nov.

Systematics

Genus Allopauropus Silvestri

Type species: Allopauropus brevisetus Silvestri, 1902: Fasc. 95,

no. 12, pi. 5.

Allopauropus fraterculus sp. nov.

(Figs. 1-11)

Material Examined. Holotype. Ad. 9 (female), Bruny Island, Mount

Mangana, Loc. 12 (43°22.1'S, 147°17.0'E), moss on dead log, 4-9.

iv.1989 (JD,PG).

Paratypes. Big Sassy Creek, Loc. 7, (42°08.5'S, 147°54.3'E),

rainforest, rotten log, 6 ad. 9 (2 males, 4 females), 17.V.1989 (JD). Rivaux

Creek, (41°15'S, 146°40'E), litter, 1 ad. 9 (female), 20.xii.1988 (PG).

Other material (185 specimens). Loc. 1, moss on Nothofagus, 1

subad. 8 (female), 1. iv.1989 (JD) and moss on dead trunk, 1 ad. 9

(female), 31. iv. 1989 (PG). Loc. 2, moss on ground, 1 ad. 9 (female), 1

juv. 6, 21. iv.1989 (HM,JD). Loc. 4, litter, 1 ad. 9 (male), (HM) and soil

core, 1 subad. 8 (female), 1 juv. 5, 3-8. xi. 1989 (RC). and moss on log,

2 ad. 9 (female), 18. xi. 1989 (HM). Loc. 6, moss on tree trunk, 1 ad. 9

(male), 1 subad. 8 (female), 3 juv. 6, ll.vi.1990 (ATW). Loc. 7, rotten

log, 119 ad. 9 (27 male, 56 female, 36 sex?), 30 subad. 8 (2 female, 28

sex?), 10 juv. 6, 8 juv. 5, 17.V.1989 (JD). Loc. 12, moss on dead log, 1

ad. 9 (female), 4-9.iv.1989 G) and moss on log, 1 ad. 9 (female), 9.

xi.1989 (PG). Loc. 13, litter, 2 ad. 9 (male, female), 27.ii.1989 (SS).

Diagnosis. The new species is very close to A. maoriorum

Remy described from New Zealand (Remyl956a), in natural

habitats also known from New Caledonia and southern Chile,

but can be distinguished in the following manner. The antennal

globulus g has a thin stalk, only 4-5 bracts and longish capsule

in fraterculus but has a thick stalk, several bracts, and spherical

capsule in maoriorum-, the 4th antennal segment has the seta u

(not mentioned by Remy in maoriorum)-, the anal plate is an

almost regular hexagon in fraterculus, but is 5-sided with

posterior margin in a long curve in maoriorum.

Description. Length. -(0.55-) 0.69(-0.72) mm.

Head. -Tergal setae of medium lengths, sublateral and

lateral ones fairly long, subcylindrical-cylindrical, annulate,

blunt. Relative lengths of setae, 1st row: 10, < 2 gS| 10-)11(-12);

2nd row: a i =(8-)9(-ll), a 2 =14(-16), a^(9-)10(-12); 3rd row:

a i =(9-)ll,a 2 =(14-)16(-17);4throw:a i =(12-)16,a 2 =(20-)21(-26),

a 3 -( 18-)21, a 4 -(18-)19(-20); lateral group setae:

/ ; -/ 2 -(17-)19(-21), l={ 32-) 40(-45). The ratio afa 1 -a 1 in 1st row

0.7(-0.8), 2nd row 0.6(-0.7), 3rd row (1.3 -)1.5(-1.6) and 4th row

1.3(-1.5). Temporal organs small, their length in tergal view

0.6(-0.7) of their shortest distance apart, posterior aperture

absent. Head cuticle almost glabrous.

Antennae. -Segment 4 with 5 subcylindrical annulate blunt

setae; their relative lengths: p= 100, /?-(58-)61(-66),

p"=(54-)56(-59), lr=|22-)23, u=( 3-)4. The p'" rudimentary.

Tergal seta p ( 1 . 1 -) 1 . 2 times as long as tergal branch t. The

latter somewhat clavate, (2.7-)3.0(-3.1) times as long as its

greatest diameter and as long as (-1.1 times as long as) sternal

branch s, that branch (2.2-)2.4 times as long as its greatest

diameter; anterodistal corner of s distinctly truncate. Seta q as

seta p of 4th segment, (as long as-) 1.2 times as long as s.

Relative lengths of flagella (basal segments included) and

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

291

Figs. 1-11. Allopauropus fraterculus sp.nov., holotype 1-6, 8-11; paratype 7: 1, head, median and right part, tergal view; 2, right antenna, sternal

view; 3, collum segment, median and left part, sternal view; 4, tergite VI, posterior part; 5, Tp 6, Tp 7, right genital papilla and seta on coxa of

2nd pair of legs, anterior view; 8, seta on trochanter of 9th pair of legs; 9, tarsus of 9th pair of legs; 10, pygidium, posterior and left part, sternal

view; 11, anal plate, lateral view. Scale line a for figures 5, 6, 8, 9; b for figures 1-3, 4, 7; c for figures 10, 11.

292

U. Scheller

basal segments: 7q=100, bs= 8(-12); 77=42(-54), bs={6-)l{-9)\

7^=85(400), /w 3 =10(-ll). The F 1 (2.7-)3.1 times as long as t, F 2

and F 3 1.4(-1.5) and (2.7-)2.8(-2.9) times as long as 5 respectively.

Distal calyces hemispherical; distal part of flagellar axes

between last lamella and calyx strongly widened, ball-shaped.

Globulus g (1.5-)1.8(-1.9) times as long as wide; (4-)5 bracts,

capsule ovoid, distinctly longer than wide; width of g 0.5 of the

greatest diameter of t. Antennae glabrous.

Trunk.- Setae of collum segment somewhat clavate, dense

but distinctly annulate, blunt, furcate but with rudimentary

glabrous pointed secondary branches; sublateral setae 3.0(-3.1)

times as long as submedian ones; sternite process triangular,

with anterior lengthening narrow and with apical incision;

appendages barrel-shaped, caps with collar; process and basal

segment of appendages with minute pubescence.

Setae on tergites subequal in length; on anterior tergites as

on tergal side of head, on posterior tergites cylindrical and

with oblique pubescence. There are 4+4 setae on tergite I, 6+6

on II-IV, 6+4 on V, 4+2 on VI. Submedian posterior setae on

VI 0.5(-0.6) of their distance apart and (2.4-)2.5 times as long

as pygidial setae a y Tergites almost glabrous.

Relative lengths of bothriotricha: 7^=100, T 2 = (98-)101(-105),

r r 102(406). 7;=(116-)128(-134), 7>(124-)135(-141). They

have simple, straight axes, thin in all but T y the latter with

thicker axes, in proximal half compact and in distal half

annulate, each annulus with a whorl of erect hairs. Pubescence

hairs on T p T 2 , T 4 and T 5 oblique in proximal 1/3, more outwards

erect.

Genital papillae (paratypes) glabrous, conical, with inner

sides only a little curved, outer sides strongly convex, 1.4(4. 8)

times as long as their greatest diameter; seta (0.3-)0.4 of the

length of organ.

Legs. -Setae on coxa and trochanter of leg 9 similar,

simple, subcylindrical, annulate, blunt, those more anteriorly

with glabrous blunt, rudimentary secondary branches. Coxal

seta on leg 2 in male as other coxal setae but somewhat

clavate. Legs short, tarsus of leg 9 subcylindrical, 2.9(-3.4)

times as long as its greatest diameter. Proximal seta thin and

with oblique pubescence, its length (0.3-)0.4 times length of

tarsus and (1.3 -)1.5 times as long as distal seta; the latter

somewhat clavate, annulate, blunt. Cuticle of tarsus almost

glabrous.

Pygidium. Tergum. -Posterior margin between st straight (or

somewhat convex). Relative lengths of setae: a= 10,

a 2 =(25-)27(-30), a=( 56-)73, st=(2-)3. The a p a 2 and a 3 shortly

pubescent, subcylindrical, tapering, pointed, curved inwards, a j

also pointing inwards; st straight and clavate, with distinct

pubescence, pointing inwards. Distance a 1 -a 1 (2.2-)2.7 times as

long as ay. distance a 1 -a 2 (3.5-)5.0 times as long as distance a 2 -a 3 \

distance st-st (6.4-)9.0 times as long as st and (as long as-) 1.1

times as long as distance a-a J Cuticle minutely granular.

Sternum. -Posterior margin between b 1 with broad and low

lobe below anal plate; hind area divided into two rounded

parts by a posteromedian incision. Relative lengths of setae

(a ; =10): b=( 38-)50, A,=(14-)18, b={ 9-)12, all setae tapering,

pointed and with very short oblique pubescence; b 1 about as

long as their distance apart, sometimes with distal swelling;

b 2 (0.8-)0.9 of the length of distance b f b 2 , b 3 0.3 of their

distance apart. Anal plate somewhat longer than broad,

glabrous, hexagonal, with anterior and posterior margins

subequal in length; two cylindrical, blunt appendages with

short oblique pubescence protrude from posterosternal

margin, 0.6(-0.7) of the length of plate, somewhat pointing

outwards.

Subad. 8. -Setae d j on pygidial tergum 0.5 of their distance

apart; setae d 2 0.7 of the length of d } and 0.8 of distance dj-d 2 .

Etymology— From Latin frater, fratris = brother (of A.

maoriorum).

Distribution in Tasmania. Most specimens were collected

from a rotten log at Big Sassy Creek but the species seems to be

very widely distributed in the State.

Allopauropus inusitatus sp.nov.

(Figs. 12-24)

Material Examined. Holotype. Ad. 9 (female), Bruny Island, Mount

Mangana, (43°22.1'S, 147°17.0'E), litter, 9.iv.l989 (PG).

Paratypes. Same data as holotype, 2 ad. 9 (male, female), 1 juv. 6,

2 juv. 3. Loc. 8, Sandspit River, (42°42.1'S, 147°51.5'E), litter, 11 ad. 9

(5 male, 6 female), 1 juv. 6, 1 juv. 3, 22.V.1989 (PG).

Other material (7 specimens). Loc. 7, moss on log, 1 ad. 9 (female),

17.V.1989 (HM). Loc. 12, leaf litter, 1 juv. 3, 9.iv.l989 (PG), and in

moss, 1 ad. 9 (female), 9.iv.l989 (JD), and in moss on dead log, 1 ad. 9

(female), 1 juv. 6, 2 juv. 3, 4.iv.l989 (JD, PG).

Diagnosis. Allopauropus inusitatus is easily recognised and

well delineated by the combination of good characters in the

shape of the temporal organs, anal plate, the 77, the antennae

and legs. It is closest to A. sphaeruliger Remy, described from

the Ivory Coast (Remy, 1948) and later found in Gambia,

Gabon, Angola, Madagascar, Reunion and Mauritius and also

in Asia, Pondichery, Sri Lanka and Japan and in South America

in Brazil. It is distinguished from that species by the shape of

the temporal organs (only slightly visible in tergal view in

inusitatus ; but clearly visible in sphaeruliger ), the shape of the

tergal antennal branch (1.4-)1.6 times as wide as long; not

2.0(-2.8), the number of tergal setae on tergite V (6+6; not 6+4),

trichobothrium T 3 (with simple pubescence and distal ovoid

swelling, not with branched pubescence and distal swelling

absent) and some pygidial characters (setae a 4 clavate, not

cylindrical; the proportion a 2 /a 3 about 0.1, not 0.5-0. 8;

appendages of the anal plate directed posteriorly and with short

pubescence, not club-shaped, strongly pointing outwards, with

long pubescence hairs).

Description. Length. -( 0.96-)l. 23(4.24) mm.

Head. -Tergal setae short to medium length, somewhat

clavate, annulate, blunt; lateral ones subcylindrical, annulate.

Relative lengths of setae, 1st row: a= 10, a 2 -(9-)10(-12); 2nd

row: tf ; -(9-)10(-ll), a 2 -9(-12), a 3 - 6(-9); 3rd row: a= 9(42),

0=9(43); 4th row: <jt 7 =10(-14), a 2 =10(-15), o=(13-)15(-16),

a 4 -l0(-l4y, lateral group setae: / ; =(13-) 14(46), Z 2 =13(-16),

/ 3 =(16-)19(-2 0). The ratio a/a f a 1 in 1st row (0.7-)0.8(-0.9), 2nd

row (0.6-)0.7(-0.9), 3rd row (0.7-)0.8(-0.9) and 4th row 0.6(-0.9).

Temporal organs small and laterosternal with narrow

anterosternal extension the distal part of which raised from

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

293

Figs. 12-24. Allopauropus inusitatus sp.nov., holotype 12-19; 21-24, paratype 20: 12, head, median and right part, tergal view; 13, left temporal

organ, sternal view; 14, temporal organ, posterior part; 15, left antenna, sternal view; 16, collum segment, median and left part, sternal view; 17,

tergite VI, posterior part; 18, T 19, 20, genital papillae and seta on coxa of 2nd pair of legs; 21, seta on coxa of 9th pair of legs; 22, tarsus of

9th pair of legs; 23, pygidium, posteromedian and left part, sternal view, to the right setae a J (above) and V; 24, anal plate, lateral view. Scale line

a for figure 20; b for figures 18,19; c for figures 12-14, 16, 17, 21, 22; d: 15, 23, 24.Figs. 39-43.

294

U. Scheller

head surface; their length in tergal view 0.4(-0.5) of their

shortest distance apart; small aperture at posterior margin

between / ; and l r Head cuticle distinctly granular and with

transverse suture anterior of 3rd row of setae.

Antennae. Segment 4 with 4 subcylindrical annulate blunt

setae; their relative lengths: p= 100, p -(85-)96(-104),

p"-{ 85-)92(-96), r-(48-)53(-62). The p'" seta rudimentary, u

absent. Tergal seta p (0.6-)0.8 of the length of tergal branch t.

The latter fusiform, (1.4-)1.6 times as long as its greatest

diameter and (0.8-)0.9 of the length of sternal branch s, that

branch (1.4-)1.7 times as long as its greatest diameter;

anterodistal corner of s truncate. Seta q subcylindrical,

annulate, blunt, (almost as long as-)1.0(-1.2) times as long as

5. Relative lengths of flagella (basal segments included) and

basal segments: 7^-100, fo ; -9(-10); F 2 -(43-)47(-49),

bs= 7(-8); F=(83-)87(-95), bs= 7(-9). The F, (4.2-)4.6(-4.8)

times as long as t, F 2 and F 3 which are thinner than F } are

1.5(-1.9) and (3.5-)4.2(-5.0) times as long as s respectively.

Distal calyces subhemispherical; distal part of flagellar axes

widened only just below calyces. Globulus g (1.2-)1.3(-1.4)

times as long as wide; 5(-6) bracts, capsule small, bottom

convex; width of g 0.4(-0.5) of the greatest diameter of t.

Antennae almost glabrous, minute pubescence on basal

segments of flagella only.

Trunk.-Setae of collum segment (subcylindrical-) somewhat

clavate, annulate, blunt, furcate but with rudimentary glabrous

blunt secondary branches; sublateral ones (1.2-)1.4 times as

long as submedian ones; sternite process short, with shallow

anterior incision (or almost blunt); appendages barrel-shaped

with flat caps; process and basal segment of appendages with

distinct, almost erect pubescence.

Setae on tergites thin, cylindrical, annulate, blunt

decreasing in length posteriorly. There are 4+4 setae on tergite

I, 6+6 on II-V, 4+2 on VI. Submedian posterior setae on VI

0.1(-0.2) of their distance apart and (0.9-)1.0(-l.l) times as long

as pygidial setae a v Tergites with short dense pubescence.

Relative lengths of bothriotricha: !T ; -100, !T 2 =(82-)?(-118),

7j-(75-)82(-98). 7;=(90-)101(-110), r=(81-)95(-100). They

have thin, simple, straight axes, 7j with distal swelling.

Pubescence hairs on T p T 2 , T 4 and T 5 oblique in proximal 1/3,

more outwards erect; T 3 with oblique hairs, very short on

proximal half, longer and in whorls on nodulated axis below

distal swelling.

Genital papillae (paratypes) glabrous, conical, with convex

inner and outer sides, 1.4-1. 7 times as long as their greatest

diameter; seta 0.4-0. 5 of the length of organ.

Legs.- Setae on coxa and trochanter of leg 9 similar, simple,

subcylindrical, annulate, blunt. More anteriorly, these setae

with glabrous blunt rudiments of secondary branches. Coxal

seta on leg 2 in male not divergent. Tarsus of leg 9 short, thick,

barrel-shaped, (1.9-)2.1(-2.2) times as long as its greatest

diameter. Setae subcylindrical, annulate, proximal seta thin,

its length 0.2 of the length of tarsus and (0.6-)0.7(-0.8) of the

length of distal seta; the latter distinctly thicker than proximal

seta. Cuticle of tarsus with short but distinct pubescence.

Pygidium. Ter gum. -Posterior margin between st rounded

and with shallow median indentation. Relative lengths of setae

a= 10, a 2 =10(-15), a 3 =(106-)107(-129), st=5(-l). The and st

straight and clavate, the former striate and the latter with short

pubescence; a 2 cylindrical, annulate, somewhat curved

inwards; a 3 thin, subcylindrical, tapering, striate-annulate,

pointing outwards and curved inwards. Distance a 1 -a 1

(2.5-)2.7(-4.8) times as long as ay distance a f a 2 (2.2-)2.3(-3.7)

times as long as distance a 2 -ay distance st-st (7.6-)9.5(-11.0)

times as long as st and (1.0-)1.1(-1.5) times as long as distance

a 1 -a 1 . Cuticle glabrous.

Sternum. -Posterior margin between b } with a broad

indentation and a small posteromedian lobe below anal plate.

Relative lengths of setae (a= 10): fc y =(43-)47(-55),

fc 2 =(17-)18(-23), 6=(14-)15(-33). The subcylindrical,

tapering, striate, distally annulate; b 2 and b 3 subcylindrical

annulate. The b } about as long as their distance apart; b 2 (0.4-)0.6

times as long as distance bj-b 2 , b 3 (0.3-)0.4 of their distance

apart. Anal plate (1.1-)1.2 times as broad as long, glabrous,

spatulate, anteriorly constricted and posteriorly protruding into

a median lobe being somewhat longer than broad, lobe with

small posteromedian incision, lateral margins of plate anterior

of posterior lobe strongly convex; two appendages protrude

from sternal side at the base of the posterior lobe, being 0.7 of

the length of plate, cylindrical but with small distal swelling,

curved inwards and shortly pubescent.

Etymology.- From Latin inusitatus = unusual, extraordinary

(shape of the temporal organs).

Distribution in Tasmania. Allopauropus inusitatus was found

at three sites only, indicating a south-eastern range.

Genus Decapauropus Remy

Type species: Decapauropus cuenoti Remy, 1931: 67-83, Figs.

1-3, 6-12.

Decapauropus heis sp.nov.

(Figs. 25-38)

Material Examined. Holotype. Ad. 9 (female), Bruny Island, Mount

Mangana, Loc. 12, (43°22.1'S, 147°17.0'E), litter, 9.iv.l989 (PG).

Paratypes. Same data as holotype, 1 ad. 9 (female). Sandspit

River, Loc. 8, (147°51.5'S, 42°42.1'E), leaf litter, 3 ad. 9 (1 male, 2

female), 22.V.1989 (PG).

Other material. 50 specimens. Loc. 2, soil core, 1 ad. 9 (female),

21.iv.1989 (JD,HM). Loc. 7, leaf litter, 1 ad. 9 (female), 12.V.1989 (PG).

Loc. 8, soil core, 2 ad. 9 (female), 22. v. 1989 (PG). Loc. 11, south track,

leaf litter, 1 ad. 9 (female), 21.iii.1989 (PG,JD). Loc. 12, litter, 16 ad. 9

(7 male, 9 female), 3 subad. 8 (1 male, 2 female), 9.iv.l989, and leaf

litter, 6 ad. 9 (female), 1 subad. 8 (female), 9.iv.l989 (PG), and leaf

litter, 1 ad. 9 (female), 4.iv.l989 (JD,PG), and in moss, 3 ad. 9 (1 male,

2 female), 9.iv.l989 (JD). Loc. 13, litter, 3 ad. 9 (1 male, 2 female), 27.

ii.1989 (SS). Loc. 16, litter, 4 ad. 9 (2 male, 2 female), 1 subad. 8

(female), 1 juv. 6, 20.xii.1988 (PG). Loc. 18, leaf litter, 2 subad. 8

(female), 1 juv. 6, xii.1987 (MN). Loc. 19, leaf litter, 3 ad. 9 (female),

20.vi.1989 (PG).

Diagnosis. The species may be closest to D. acer Scheller from

Central Amazon (Scheller 1994). They are similar in the

antennal morphology, the process of the collum segment, the

bothriotricha, the pygidial chaetotaxy and the singular shape of

the anal plate. Reliable distinguishing characters are the thick

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

295

£>-

0 ~

20 Mm

iS -

i ,

^ <

Figs. 25-38. Decapauropus heis sp.nov., holotype 25-31, 33-38; paratype 32: 25, head, median and right part, tergal view; 26, posterior part of

temporal organ; 27, right antenna, sternal view; 28, collum segment, median and left part, sternal view; 29, tergite VI, posterior part; 30, Tp 31,

77; 32, genital papillae and seta on coxa of 2nd pair of legs, anterior view; 33, seta on coxa of 9th pair of legs, 34, seta on trochanter of 9th pair

of legs; 35, tarsus of 9th pair of legs; 36, pygidium, posterior part, sternal view; 37, anal plate, lateral view; 38, anal plate, tergal view. Scale line

a for figures 30-35; b for figures 25, 28; c for figures 26, 27, 29, 36-38.

296

U. Scheller

annulate setae on the collum segment and the proportionately

longer collum appendages in D. heis, also the posterosternal

lobe of the anal plate, which is absent in D. acer and the shape

of the anal plate appendage.

Description. Length.-(0.5l-)0.6l(-0.65) mm.

Head. -Tergal setae annulate and of different lengths, in 1st

and 2nd rows of medium lengths, in 3rd and 4th rows rather

long; anterior and submedian ones somewhat clavate, sublateral

and lateral ones cylindrical. Relative lengths of setae, 1st row:

a =10, a= 10(11); 2nd row: a= 11(12), a 2 =10(-12), a=l\ 3rd

row: a =17(-20), a 2 =20(-23); 4th row: a 7 =15(-16), a 2 =(23-)24,

a J =25(-26), a={ 13-) 16; lateral group setae: / ; -(17-)25,

Z 2 — ( 12-) 14, l={ 16-) 19. The ratio aja-a 1 in 1st row ( 1 . 1 -) 1 .2,

2nd row 0.6(-0.8), 3rd row 1 ,7(-2. 1 ) and 4th row (3 .4-)3 ,7(-3 .8).

Temporal organs oval in tergal view, their length 0.8(-0.9) of

their shortest distance apart; small aperture near surface in

median part at level of l 2 aperture with thin interior canal

backwards. Head cuticle almost glabrous; temporal organs

with delicate pubescence.

Antennae- Segment 4 with four subcylindrical, annulate,

blunt setae; their relative lengths: p= 100, p -(69-)73(-81),

p M =(40-)41(-52), r=(24-)25(-26). Neither p"\ nor u. Tergal seta

p (as long as-) 1.2 times as long as the length of tergal branch

t. The latter fusiform, (2.4-)2.6(-2.9) times as long as its

greatest diameter and about as long as sternal branch s, that

branch 1.8(-2.3) times as long as its greatest diameter;

anterodistal corner of s truncate. Seta q subcylindrical,

annulate, blunt, 0.8 (-as long as) 5. Relative lengths of flagella

(basal segments included) and basal segments: 7q=100,

bs=(6-)l(-8)-, F 2 =(70-)82(-85), bs=( 7-)8; F=(76-)S6, bs = 8.

The Fj (3.2-)3.6 times as long as t, F 2 and F 3 2.2(-2.5) and

(2.6-)2.8(-2.9) times as long as 5 respectively. Distal calyces

somewhat flattened; distal part of flagella axes fusiform.

Globulus g 1.5(-1.6) times as long as wide; 12(-13) bracts;

capsule spherical; width of g 0.9(-as wide as) the greatest

diameter of t. Antennae almost glabrous.

Trunk.-Setae of collum segment somewhat clavate,

distinctly annulate, blunt, furcate but with rudimentary

glabrous blunt secondary branches; sublateral ones

(1.8-)2.1(-2.5) times as long as submedian ones; sternite

process narrow, with anterior lengthening with apical incision;

appendages conical strongly narrowing distally and with

proportionately very small caps; process and basal segment of

appendages very delicately granular.

Setae of about the same length on all tergites, on anterior

ones subcylindrical, annulate, blunt, on posterior tergites

cylindrical, tapering, pointed, with short oblique pubescence.

There are 4+4 setae on tergite I, 6+6 on II-IV, 6+4 on V, 4+2

on VI. Submedian posterior setae on VI 0.6(-0.7) of their

distance apart and 0.8(-0.9) of the length of pygidial setae a r

Tergites glabrous. Relative lengths of bothriotricha: 7^-100,

r 2 =(103-)109(-121), r-( 97-) 1()()(-1 19). r 4 =(115-)119(-130),

r y -(169-)180(-182) i axes simple, straight, in all but T 3 being

very thin, with proximal 2/3 somewhat fusiform and distal 1/3

thin. Pubescence hairs very short and thin on all but T 3 , on the

latter stronger.

Genital papillae (paratypes) glabrous, fusiform, (1.9-)2.1

times as long as their greatest diameter; seta 0.5(-0.6) the

length of organ.

Legs- Setae on coxa and trochanter of leg 9 similar, furcate,

branches subequal in length, striate, blunt; they are cylindrical

except the main branch of coxal seta somewhat clavate, more

anteriorly, these setae with rudimentary secondary branches,

setae on trochanter longest, those on coxae somewhat clavate.

Coxal seta on leg 2 in male not divergent. Tarsus of leg 9

slender, (3.7-)4.4 times as long as its greatest diameter.

Proximal seta tapering, pointed, with very short oblique

pubescence; distal seta cylindrical striate blunt. The former

0.4 of the length of tarsus and 2.0(-2.2) times as long as the

latter. Cuticle of tarsus with delicate pubescence.

Pygidium. Tergum. -Posterior margin between st somewhat

indented, straight. Relative lengths of setae: a= 100,

a 2 =(86-)93(-98), ^=(136-)137(-157), v/-53(-71 ). These setae

thin, tapering, almost glabrous, a } and a 2 almost straight, a 3

and st curved inwards, the latter one also pointing inwards.

Distance a ] -a ] 0.7(-0.8) of the length of ay, distance a ] -a 2

(1.4-)1.8 times as long as distance a 2 -ay distance st-st

(2.3-)2.5(-2.6) times as long as st and (1.7-)1.8(-2.0) times as

long as distance a 1 -a f Cuticle somewhat granular.

Sternum. -Posterior margin between b } with broad shallow

indentation. Relative lengths of setae (a ; -100):

fc 7 =(335-)338(-381), fc 2 =(87-)90(-108). The thin,

subcylindrical, striate; b 2 as a : and a 2 of pygidial tergum. The

b 3 (1.6-)1.8 times as long as their distance apart; b 2 0.8(-0.9)

times as long as distance b 1 -b 2 . Anal plate broadest anteriorly,

glabrous, linguiform with parallel lateral margins and rounded

posteriorly; about three times longer than broad; from the

thickened median part protrudes posteriorly from sternal side

a narrow, and in the vertical plane undulated appendage about

as long as plate; lateral margins of appendage thickened or

curved, so it looks like two thread-like parallel structures.

Etymology.- From Greek heis = one (the appendage of the anal

plate).

Distribution in Tasmania. The range is wide but it has not been

collected from the central and north-eastern parts.

Decapauropus attenuatus sp.nov.

(Figs. 39-50)

Material Examined. Holotype. Ad. 9 (female), Bruny Island, Mount

Mangana, Loc. 12, (43°22.1'S, 147°17.0'E), litter, 4.iv.l989 (PG).

Paratypes. Locality as for holotype, in moss, 3 ad. 9 (2 male, 1

female), 1 subad. 8 (female), 9.iv.l989 (JD).

Other material. 10 specimens. Loc. 4, moss on log, 1 ad. 9

(female), 1 subad. 8 (female), 18. xi. 1989 (HM). Loc. 7, rotten log, 1 ad.

9 (female), 17.V.1989 (JD). Loc. 11, moss on dead log, 1 ad. 9 (female),

21.iii.1989 (JD). Loc. 12, moss on log, 1 juv. 3, 9.xi.l989, and leaf

litter, 2 ad. 9 (female), 9.iv.l989, and moss on dead log, 2 ad. 9 (male,

female), 9.iv.l989 (PG), and in moss, 1 ad. 9 (female), 9.iv.l989 (JD).

Diagnosis. The affinities are difficult to trace but the species is

well delineated by the combination of the following characters:

rather long tergal head setae, anterior ones clavate; the T 3 with

proximal 2/3 fusiform but very thin distally; the shape of the

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

297

Figs. 39-43. Decapauropus attenuatus sp.n., holotype: 39, head, median and right part, tergal view; 40, right temporal organ, posterior part,

lateral view; 41, left antenna, tergal view; 42, collum segment, median and left part, sternal view; 43, tergite VI, posterior part. Scale line a for

figures 39, 40, 42; b for figures 41, 43.

298

U. Scheller

tfi¥t i (fW f WWWi HWW *

Figs. 44-50. Decapauropus attenuatus sp.nov., holotype 44-45, 47-50; paratype 46: 44, Tp 45, 77; 46, genital papillae and seta on coxa of 2nd

pair of legs; 47, seta on trochanter of 9th pair of legs; 48, tarsus of 9th pair of legs; 49, pygidium, posteromedian and left part, sternal view; 50,

anal plate, lateral view. Scale line a for figures 44-48; b for figures 49, 50.

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

299

anal plate with concave lateral margins, distal incision and two

long appendages which are curved inwards.

Description. Length. -($). 56-) 0.63 (-0.74) mm.

Head- Most tergal setae of medium length, some

posterolateral rather long, anterior and submedian ones

somewhat clavate, others cylindrical, striate, blunt but lateral

group setae pointed. Relative lengths of setae, 1st row: a = 10,

a 2 =10(-12); 2nd row: a 7 =(ll-)13, a 2 =(14-)16(-18), ^=(13-)16;

3rd row: a=ll(-l2), a 2 -13(-15); 4th row: a=( 14-)15(-16),

a 2 -19(-23), a={\l-)\9, a 4 =(13-)15(-16); lateral group setae:

l~(22-)25. / 2 -13(-18), 1-2(22). The ratio a/a-a l in 1st row

l.O(-l.l), 2nd row (0.6-)0.7, 3rd row 1.0(-1.2) and 4th row

1.4(-1.7). Length of temporal organs 0.7(-0.9) of their shortest

distance apart; small aperture in posterior part anterior of l r

Head cuticle somewhat granular, temporal organs glabrous.

Antennae.- Segment 4 with five thin cylindrical striate

setae; their relative lengths: p= 100, p'~ (71-)76(-78),

j p"-(22-)24(-28), /?" -11(-18), r=24(-27), u absent. Tergal seta p

(1.4-)1.5(-1.6) times as long as tergal branch t. The latter

fusiform, 2.8(-3.6) times as long as its greatest diameter and as

long as sternal branch 5, that branch (2.0-)2.2(-2.6) times as

long as its greatest diameter; anterodistal corner of s truncate.

Seta q cylindrical, striate, as long as (-1.2 times as long as) 5.

Antennal flagella often broken; relative lengths (basal segments

included) and basal segments: F 7 =100, bs=( 6-)8; F 2 =(78-81),

bs 2 -l\ F 3 -{ 80-92), bs 3 =( 6-)8. The 7q 2.8(-3.4) times as long as

t, F 2 and F 3 ?( 1. 8-2.7) and (2. 0-2.9) times as long as 5

respectively. (Antennal flagella broken in holotype). Distal

calyces somewhat flattened; distal part of flagellar axes

fusiform. Globulus g (1.2-)1.3 times as long as wide; nine

bracts, capsule spherical; width of g 0.6(-0.8) of the greatest

diameter of t. Antennae with short pubescence.

Trunk.-Setae of collum segment somewhat clavate,

annulate, blunt, furcate, but with rudimentary glabrous blunt

secondary branches; sublateral ones (1.8-)2.1(-2.6) times as

long as submedian ones; sternite process triangular with small

anterior incision; appendages obliquely barrel-shaped with

flattened caps; process and basal segment of appendages with

delicate pubescence.

Setae on anterior tergites cylindrical, annulate, blunt; on

posterior tergites somewhat shorter, subcylindrical, tapering,

with oblique pubescence. 4+4 setae on tergite I, 6+6 on II-IV,

6+4 on V, 4+2 on VI. Submedian posterior setae on VI

(0.6-)0.7 of their distance apart and (0.8-)0.9 of the length of

pygidial setae a r Tergites with short minute pubescence.

Relative lengths of bothriotricha: 7^-100, T 2 -95(-101),

7/r(;97-)112(-113). 7^-( 1 07) 1 08(1 30). 7;=(165-)167(-187),

with simple straight axes, in all but T 3 very thin; the latter with

proximal 2/3 thickened, fusiform, tapering into a thin distal

1/3. Pubescence hairs on T„ 77 and 7/ similar, hairs thin,

straight, rather sparse, on proximal halves short and oblique,

erect and longer distally. The T 5 with very short oblique

pubescence.

Genital papillae (paratypes) subcylindrical, glabrous,

2.2(-2.3) times as long as their greatest diameter; seta 0.3(-0.4)

of the length of organ.

Legs. -Setae on coxa and trochanter of leg 9 similar, furcate;

branches striate, blunt, subequal in length, primary branch

somewhat clavate, secondary branch cylindrical, these seta

more anteriorly with blunt glabrous rudimentary secondary

branches; setae on trochanter somewhat widened distally, those

on coxa clavate. Coxal seta on leg 2 in male clavate too. Tarsus

of leg 9 slender, 3.8(-4.3) times as long as its greatest diameter.

Setae thin, proximal one tapering pointed, with oblique

pubescence; distal one cylindrical, blunt, striate; proximal seta

(0.4-)0.5 of the length of tarsus and (2.2-)2.5(-2.6) times as

long as distal seta. Cuticle of tarsus with distinct pubescence.

Pygidium. Tergum. -Posterior margin between st rounded

and with shallow median indentation. Relative lengths of setae:

iq-100, o 2 -( 100-) 106(-1 13), a~( 1 35- ) 1 37( - 1 60 );.

s?-(60-)62(-80), setae thin, tapering, pointed, indistinctly striate,

somewhat curved inwards, a 1 and a 3 with short pubescence

distally; o ; directed posteriorly, a 2 and st pointing inwards, a 3

somewhat pointing outwards. Distance o ; -o ; 0.9(-1.2) times as

long as ay, distance a 1 -a 2 (1.5-)1.8 times as long as distance

a 2 -ay distance st-st (2. 1 -)2.4 times as long as st and ( 1 .4-) 1 .6(- 1 .7)

times as long as distance a 1 -a y Cuticle glabrous.

Sternum. -Posterior margin between b 1 with a broad shallow

indentation. Relative lengths of setae (o ; -100):

&,-(343-)362(-404), & 2 =(80-)94(-100), setae thin, b }

subcylindrical, tapering, striate, blunt; b 2 tapering, pointed,

striate. The b 1 (1.3-)1.4(-1.5) times as long as their distance

apart; b 2 (0.8-)0.9 of distance b-b 2 . Anal plate as long as (-1.2

times as long as) broad, granular, broadest anteriorly, with

somewhat indented lateral margins and posterior margin

divided into two rounded lobes by a broadly V-shaped

indentation; from postero sternal margin of the lobes protrude

backwards two pointing outwards appendages slightly curving

inwards, being 1.3(-1.6) times as long as the length of plate.

Stage subad. 8. The setae d 2 of pygidial tergum thin,

straight, cylindrical, striate, 0.4 of the length of pygidial setae

ay d 2 -d 2 about 10 times longer than d 2 .

Etymology. -From Latin attenuatus - drawn out, thin (Ty setae

of the pygidial tergum).

Distribution in Tasmania. Seems to be a rare species collected

in the eastern parts only.

Decappauropus ungulatus sp. nov.

(Figs. 51-61)

Material Examined. Holotype. Ad. 9 (female), Rivaux Creek, Loc. 16,

(43°10'S, 146°38.6'E), litter, 20.xii.1988 (PG).

Diagnosis. A single adult specimen is available but the singular

shape of the anal plate in combination with the unusual shape

of the antennal setae, the very small collum process and

appendages and the arrangement and shape of the pubescence

on the bothriotricha, make it well defined. It has some similarity

with D. fibratus Scheller from Sri Lanka (Scheller 1970), but

the T 3 and the anal plate are dissimilar (the former with distal

swelling in D. fibratus, not in D. ungulatus-, the appendages of

the anal plate short, cylindrical and glabrous in D. fibratus,

distinctly longer, clavate and pubescent in D. ungulatus ).

300

U. Scheller

Figs. 51-61. Decapauropus ungulatus sp.nov., holotype: 51, head, median and right part, tergal view; 52, temporal organ, posterolateral part,

lateral view; 53, left antenna, sternal view; 54, collum segment, median and left part, sternal view; 55, T 56, 57, seta on coxa of 9th pair of

legs; 58, seta on trochanter of 9th pair of legs; 59, tarsus of 9th pair of legs; 60, pygidium, posterior and left part, sternal view; 61, anal plate,

lateral view. Scale line a for figures 55, 56; b for figures 51, 52, 54, 57-59; c for figures 60, 61; d: 53.

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

301

Description. Length.-0.66 mm.

Head. -Tergal setae annulate, blunt, submedian ones

somewhat clavate, lateral ones cylindrical. Relative lengths of

setae, 1st row: 10, a 2 - 13; 2nd row: a =8, a 2 -10, a= 15; 3rd

row: a ; -12, a 2 -20; 4th row: a=a= 17, a 2 - 26, a 4 = 19: lateral

group setae: l =30, 1=1=21. The ratio in 1st row 1.0,

2nd row 0.5, 3rd row 0.9, 4th row 1.7. Temporal organs oval in

tergal view, their length 1.1 times as long as their distance

apart; small aperture outside posterior margin anterior of l y

Head cuticle and temporal organs glabrous.

Antennae.- Segment 4 with 4 annulate setae: p, p 1 and p"

somewhat clavate and distinctly annulate, r cylindrical; their

relative lengths: p=\00,p '=44,/?' -30, r= 52. Neither nor u.

Tergal seta p 1.7 times as long as the length of tergal branch t.

The latter somewhat fusiform, 2.1 times as long as its greatest

diameter and as long as sternal branch s, that branch twice as

long as its greatest diameter; anterodistal corner of 5 deeply

truncate. Seta q subcylindrical, annulate, blunt, 1.2 times as

long as 5. Relative lengths of flagella (basal segments included)

and basal segments: 7q=100, bs=l; F = 34, bs 2 = 4; F=l, bs=l.

The Fj 4.3 times as long as t, F 2 1.3 times as long as 5. Distal

calyces of F 1 subhemispherical, those of F 2 flattened. Distal

part of flagella axes thickened, cylindrical in F p fusiform in

F 2 . Globulus g proportionally large, 1.5 times as long as wide;

10 bracts; capsule spherical; width of g 0.8 of the greatest

diameter of t. Antennae glabrous.

Trunk. -Setae of collum segment cylindrical, annulate,

blunt; rudiments of secondary branches not clear. Sublateral

setae 2.3 times as long as submedian ones; sternite process

very small and narrow; appendages subcylindrical with small

flattened caps. Process and appendages glabrous.

Setae on tergites not studied.

Relative lengths of bothriotricha: r ; -100, 7\=108, 7^=117,

T 4 ~\ 27, 7^=148; with simple, straight axes, somewhat thickened

in T 3 only. Pubescence on proximal half of T -T 4 simple,

oblique, increasing in length outwards, more distal ly longer,

branched, arranged in whorls; T 5 with short oblique pubescence

of simple hairs.

Legs. -Setae on coxa and trochanter of leg 9 cylindrical,

annulate, pointed; the former simple and the latter furcate with

secondary branch somewhat shorter than primary branch,

these setae of legs 1-8 simple. Tarsus of leg 9 distinctly

tapering, 2.9 times as long as its greatest diameter. Setae

cylindrical, annulate, blunt; proximal one 0.3 of the length of

tarsus and 1.1 times as long as distal seta. The latter more

densely annulate than the former. Cuticle of tarsus with short

pubescence.

Pygidium. Tergum. -Posterior margin between st straight.

Relative lengths of setae: a ; -100, a = 83, a = 126, st- 19, setae

thin, cylindrical, striate, all but st directed posteriorly, pointed;

st blunt, pointing inwards; a 1 curved outwards and somewhat

pointing outwards, a 2 and a 3 curved inwards. Distance a 1 -a 1

0.8 of the length of ay distance a } -a 2 1.3 times as long as

distance a 2 -ay distance st-st 1.6 times as long as st and 1.5

times as long as distance a 1 -a p Cuticle glabrous.

Sternum. -Posterior margin between b 3 with deep broadly

V-shaped indentation. Relative lengths of setae (a ; -100): b=l,

b 2 - 125. The b 2 as pygidial a 3 , 1.9 times as long as distance

bfb 2 . Anal plate 2.4 times as long as broad with concave

lateral margins and decreasing in width in distal 1/3, posterior

margin with a deep median incision with parallel sides; two

clavate pointing outwards appendages protrude posteriorly

from sternal side; their length 0.3 of the length of plate; lateral

margins and appendages with short pubescence.

Etymology- From Latin ungula = hoof (shape of the anal

plate).

Distribution in Tasmania. Probably rare, distribution outside

the type locality unknown.

Decapauropus convexus sp. nov.

(Figs. 62-71)

Material Examined. Holotype. Subad. 8 (female). Savage River, Loc.

1, (41°18.5'S, 145°16.3'E), soil core, 21.iv.1989 (JD).

Paratype. Same data as holotype, 1 subad. 8 (male).

Diagnosis. Decapauropus convexus shows superficial similarities

in the shape of the T 3 and some pygidial characters with three

species described by Remy: D. burghardti (1941) from France,

D. pachypus (1948) from the Ivory Coast and D. zaianus (1952a)

from Spain and Morocco. Good distinguishing characters in

relation to burghardti are the tergal antennal branch 1.7-1. 8

times longer than wide, not long and slender; the distal swelling

of T 3 well delineated from the bothriotrix axis, axis not slowly

becoming thicker distally; the posterior margin of the pygidial

tergum has a shallow median indentation between the st, not a

distinct rounded lobe. Decapauropus convexus can be

distinguished from D. pachypus by the subspherical antennal

globulus, not longish, the proportionately longer tarsi, by the

posteromedian indentation in the pygidial tergum, not a

triangular projection; and by the thin appendages of the anal

plate, not thick and clavate. Distinguishing characters in relation

to D. zaianus are the size of the antennal globulus, the length of

which is about half of the length of sternal branch s, not 0.7 of

that length, the shape of the distal swelling of the T 3 which is

ovoid, not longish and subcylindrical; and the number of

appendages of the anal plate, being two not four. D. convexus

has some similarity too with D. insignis Remy, 1961 from India,

but is distinguished especially by the much smaller antennal

globulus, the proportionately shorter and thicker tergal antennal

branch and by the the shape of the seta st which are curved and

somewhat clavate, rather than straight and lanceolate.

Description. Length.- 0.48(-0.50) mm.

Head- Most tergal setae short, posterolateral ones of

medium length, subcylindrical, densely annulate, blunt;

relative lengths of setae (holotype only), 1st row: a = 10, a = 12;

2nd row: a =10, a =20, a = 12; 3rd row: <^=10, a = 13; 4th row:

fl ; -10, a = 23, a 3 =a 4 - 24; lateral group setae: / ; -30, / 2 -24,

1= 32. The ratio a/a^aj 0.8 in 1st row, 0.6 in 2nd and 4th rows

and 0.7 in 3rd row. Temporal organs large; their length in tergal

view 1.2 times as long as their shortest distance apart; small

aperture in posterior margin on a level with l } . Head cuticle

glabrous.

Antennae.- Segment 4 with four subcylindrical annulate

setae, p,p' and p" blunt, r pointed; their relative lengths: p-100,

302

U. Scheller

Figs. 62-71. Decapauropus convexus sp.nov., holotype: 62, head, median and right part, tergal view; 63, temporal organ, posterior part, lateral

view; 64, left antenna, sternal view; 65, collum segment, median and left part, sternal view; 66, T', 67, 77; 68, seta on trochanter of 9th pair of

legs; 69, tarsus of 9th pair of legs; 70, pygidium, posterior and left part, sternal view; 71, anal plate, lateral view. Scale line a for figures 62, 63,

66, 67; b for figures 64, 65, 68, 69; c for figures 70, 71.

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

303

p'=44(-47) 5 p"=28(-30), r=(72-)75 and 80. The p'" rudimentary,

u absent. Tergal seta p 1.6(-1.8) times as long as tergal branch

t. The latter subcylindrical, (1.7-)1.8 times as long as its greatest

diameter and 0.9 of the length of sternal branch s; that branch

1.7 times as long as its greatest diameter; anterodistal corner

of s truncate. Seta q subcylindrical, annulate, blunt, 0.8(-0.9)

of the length of 5. Relative lengths of flagella (basal segments

included) and basal segments: F ; -100, bs=l-, F 2 -{ 41-)42,

bs 2 - 4; F J -(79-)81, bs 3 =l(-S). The F } 4.9(-5.4) times as long as

t, F 2 and F 3 (1.8-)1.9 and (3.6-)3.7 times as long as 5 respectively.

Distal calyces small subhemi spherical; distal part of flagella

axes somewhat fusiform in F } and F y distinctly fusiform in F r

Globulus g spherical with thin stalk, 1.2 times as long as wide;

about nine bracts, capsule subspherical with somewhat

flattened bottom; width of g 0.5 of the greatest diameter of t.

Antennae glabrous.

Trunk. -Setae of collum segment cylindrical, striate, blunt,

rudiments of secondary branches probably absent; sublateral

setae 2.5 times as long as submedian ones; sternite process

very narrow; appendages small with rounded caps; process

and appendages glabrous.

Setae on tergites thin, cylindrical, annulate, blunt and of

the same length on all tergites; 4+4 setae on tergite I, 6+6 on

II-IV, 4+4 on VI. Relative lengths of bothriotricha (holotype

only): T=T 3 - 100, T 2 -107, 7^-135, with thin, simple, straight

axes, T 3 with distal swelling. Pubescence hairs on T 5 and on

proximal 1/3 of T jy T 2 , T 3 and on T 4 oblique; on T y T 2 and T 4

pubescence erect, sparse and whorled in outer 2/3; also on

outer half of T 3 , pubescence on distal swelling somewhat

shorter than on the axis below it.

Genital papillae (paratype) small, conical, blunt.

Legs. -Setae on coxa and trochanter of leg 8 simple,

cylindrical, striate, blunt; more anteriorly similar but coxal

setae somewhat thicker than those on trochanter. Tarsus of leg

8 short, strongly tapering, (2. 8)3.0 times longer than its greatest

diameter. Setae cylindrical, blunt, proximal one annulate,

distal one striate; proximal seta 0.2 of the length of tarsus and

0.6 of the length of distal seta. Tarsus with very delicate

pubescence.

Pygidium. Tergum. -Posterior margin between st with two

low submedian lobes separated by a shallow median

indentation. Relative lengths of setae: tf ; -100, a 2 -93(96),

^-241(250), V-93(115), setae blunt, a ] and a 2 subcylindrical,

straight, annulate, the former pointing inwards, the latter

pointing outwards; a 3 and st curved inwards, the former

cylindrical and annulate, the latter somewhat clavate, striate,

pointing inwards. Distance (1.4)1. 6 times as long as ap

distance a f a 2 (2.3)2. 6 times as long as distance a 2 -ap, distance

st-st (2. 1)2.2 times as long as st and 1.3(1. 6) times as long as

distance a 1 -a f Cuticle almost glabrous.

Sternum. -Posterior margin between b } somewhat indented.

Relative lengths of setae (a ; -100): ^-463(490), b 2 = 185(192).

The b 1 cylindrical, densely striate, blunt; b 2 as a 3 of pygidial

sternum. The b 1 (1. 1)1.2 times as long as their distance apart;

b 2 0.9 of distance b ] -b 2 . Cuticle sparsely pubescent. Anal plate

subrectangular with slightly concave lateral margins and

rounded posterolateral corners, 1.2 times as long as broad,

glabrous; two parallel cylindrical and blunt appendages

protrude backwards from posterior part of sternal side; length

of appendages 0.5 of the length of the plate.

Etymology. -From Latin convexus - well rounded (at the top,

posterior part of the anal plate).

Distribution in Tasmania. Known from a single site in north-

eastern Tasmania.

Decapauropus saltuarius sp. nov.

(Figs. 72-81)

Material Examined. Holotype. Ad. 9 (male), Bruny Island, Mount

Mangana, Loc. 12, (147°17.0'S, 43°22.1'E), in moss, 9.iv.l989 (JD).

Paratypes. 3 specimens. Cradle Mountain, Loc. 4, (41°35.4’S,

145°55.9'E), in moss on log, 1 ad. 9 (female), 1 juv. 6, 18. xi. 1989

(HM).

Diagnosis. This species is well defined by the shape of the anal

plate and the combination of good characters in the antennae (/

and s of the same length, anterior truncation of s inconsiderable)

and the last pair of legs (tarsus somewhat bow-shaped, pubescent

and with very thin setae). Since many of these characters are

widespread and shared with several other species, the

relationships are difficult to discern.

Description. Length.-0.64 mm.

Head.- Tergal setae annulate blunt, of medium length, a 2 of

4th row long; lateral and sublateral ones subcylindrical,

submedian ones clavate. Relative lengths of setae, 1st row:

a= 10, a 2 =ll; 2nd row: a= 12, a 2 =a 3 - 15; 3rd row: a=ll, a 2 - 12;

4th row: a= 14, a 2 -10, a 3 - 14, a 4 - 10; lateral group setae: 1= 32,

/ 2 — 15, /.j-20. The ratio aja l -a 1 in 1st row 1.4, 2nd row 1.0, 3rd

row 1.4 and 4th row 1.5. Temporal organs oval in tergal view,

as long as their distance apart; small aperture in an anterior

position of l v Head cuticle almost glabrous.

Antennae- Segment 4 with five cylindrical setae, all but p'"

annulate-striate blunt; p'" very thin with delicate pubescence.

Relative lengths of setae: p-l00,p -84,/V -34,/?" -10, r~ 32; u

absent. Tergal seta p 1.2 times as long as tergal branch t. The

latter fusiform, 2.7 times as long as its greatest diameter and as

long as sternal branch 5 ; that branch 1.8 times as long as its

greatest diameter; anterodistal corner of s truncate. Seta q

cylindrical, annulate, somewhat tapering, as long as s. Relative

lengths of flagella (basal segments included) and basal

segments: Tq-100, bs= 8; F 2 -61, bs 2 - 8; F= 86, bs= 9. The F 2

thinnest; F } 3.3 times as long as t, F 2 and F 3 2.0 and 2.8 times

as long as s respectively. Distal calyces subhemispherical;

distal part of flagella axes fusiform. Globulus g 1.4 times as

long as wide; about nine bracts, capsule subspherical; width of

g 0.7 of the greatest diameter of t. Antennae almost glabrous.

Trunk.-Setae of collum segment subcylindrical, annulate,

blunt, furcate, but with rudimentary glabrous blunt secondary

branches; sublateral ones twice as long as submedian ones;

sternite process thin pointed; appendages subconical with

rounded caps with collar; process and basal segment of

appendages with distinct but short, erect pubescence.

Setae on tergites thin, cylindrical, annulate-striate

insignificantly decreasing in length posteriorly; 4+4 setae on

tergite I, 6+6 on II-IV, 6+4 on V, 4+2 on VI. Submedian

304

U. Scheller

Figs. 72-81. Decapauropus saltuariusx sp.nov., holotype: 72, head, median and right part, tergal view; 73, temporal organ, posterior part, lateral

view; 74, left antenna, sternal view; 75, collum segment, median and left part, sternal view; 76, tergite VI, posterior part; 77, 7\; 78, genital

papillae and seta on coxa of 2nd pair of legs; 79, seta on trochanter of 9th pair of legs; 80, tarsus of 9th pair of legs; 81, pygidium, posterior and

left part, sternal view: Scale line a for figures 77, 80; b for figures 72, 73, 75, 76, 78, 79; c for figures 74, 81.

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

305

posterior setae on VI 0.7 of their distance apart and 1.3 times

as long as pygidial setae a y Tergites glabrous. Relative lengths

of bothriotricha: 7^=100, 7;=104, ^=108, 7^=129, 7^=166,

axes thin, simple, straight, thickest in T y Pubescence hairs

short oblique on proximal parts and on almost the whole T p

more outwards longer erect, strongest on T y

Genital papillae oviform, glabrous, 1.7 times as long as

their greatest diameter; seta about 0.5 of the length of organ.

Legs. -Setae on coxa and trochanter of leg 9 similar, furcate,

branches subequal in length; main branch thick, blunt, with

short pubescence in whorls; secondary branch similar but

thinner, somewhat clavate, more anteriorly, with glabrous

blunt rudiments of the secondary branches. Coxal seta on leg

2 in male not deviating. Tarsus of leg 9 somewhat curved, 3.6

times as long as its greatest diameter. Setae thin, proximal one

tapering and pointed with short depressed pubescence, distal

one cylindrical striate; proximal seta 0.4 of the length of tarsus

and 2.5 times as long as distal seta. Cuticle of tarsus with short

pubescence.

Pygidium. Tergum. -Posterior margin between st evenly

rounded. Relative lengths of setae: a=a 2 - 100, a = 222, st=61.

These setae cylindrical, blunt, indistinctly striate; straight

and somewhat pointing outwards, a 2 and a 3 curved inwards,

the former also pointing inwards, st straight and pointing

inwards. Distance a 1 -a 1 1.2 times as long as ay, distance a f a 2

1.2 times as long as distance a 2 -ay distance st-st 3.5 times as

long as st and 1.9 times as long as distance a 2 -a y Cuticle

glabrous.

Sternum. -Posterior margin between b } almost straight.

Relative lengths of setae (a ; -100): b=455, b 2 about 130; setae

cylindrical blunt striate. The Zq 1.4 times as long as their

distance apart; b 2 about 0.5 of distance b-b 2 . Anal plate 1.5

times as long as broad, with convex lateral margins and a

V-shaped posterior incision separating two short, subcylindrical

posterolateral lobes; from distal part of lobes, two straight,

cylindrical, blunt, outwardly pointing appendages, 0.6 of the

length of the plate; anterior part of plate glabrous, posterior

part shortly pubescent, appendages distinctly striate, anterior

of each base of appendage a small sternal knob.

Etymology.- From Latin saltuarius = of the forest.

Distribution in Tasmania. Found on two widely separated

sites.

Decapauropus terrestris sp. nov.

(Figs. 82-93)

Material Examined. Holotype. Ad.9 (female). Savage River Pipeline

Road, Loc. 1, (41°18.5'S, 145°16.3'E), soil core, 21.iv.1989 (JD).

Paratypes. Frodshams Pass, Loc. 9, (42°49.7'S, 146°22.9'E), soil

core, 2 ad. 9 (female), 20.x. 1989 (RC).

Other material. 2 specimens. Loc. 9, soil core, 1 juv. 3, 20.X.1989

(RC). Loc. 5, soil core, 1 ad. 9 (male), 22.xi.1989 (HM).

Diagnosis. The shape of the antennae and the anal plate indicate

that the new species is related to D. vicinus Remy from

Madagascar (Remy 1956c). They can be reliably distinguished

as in A. terrestris there is a large semicircular lobe between the

st (not in vicinus), the st are thin and cylindrical (not broad

distally) and the anal plate narrows anteriorly (not the

opposite).

Description. Length.-( 0.59-)0.61 mm.

Head.- Tergal setae of short to medium length,

subcylindrical, densely annulate, blunt. Relative lengths of

setae, 1st row: a = 10, a 2 -(9-)ll; 2nd row: a=(l-)S, a 2 -(16-)20,

a.=( 13 -) 17: 3rd row: a = 14, a 2 -(13-)18; 4th row: a ; -(7-)10,

a 2 -?(18), a={ 15-)20, a 4 = 18(-19); lateral group setae: 1=1,

Z 2 — 17, ^-18. The ratio aja 1 -a 1 in 1st row 0.9(-1.0), 2nd row

(0.4-)0.5, 3rd row 0.9 and 4th row 1. 0(-1.3). Temporal organs

oval in tergal view, their length 1.5 times as long as their

shortest distance apart; an unusually large aperture inside

posterior margin on a level with l r Head cuticle glabrous.

Antennae. -Segment 4 with 5 cylindrical annulate blunt setae;

their relative lengths: p=100,p'=43(-50),/?"=28(-29), r=(45-)46.

The p'" rudimentary, u absent. Tergal seta p ( 1 .5-) 1 .6 times as

long as tergal branch t. The latter fusiform, 2.0(-2.2) times as

long as its greatest diameter and as long as sternal branch s; that

branch directed downwards and (1.7-)1.8 times as long as its

greatest diameter; anterodistal comer of s only a little more

truncate than posterior one. Seta q cylindrical annulate blunt, 1.2

times as long as s. Relative lengths of flagella (basal segments

included) and basal segments: F ; -100, bs= 5(-6); F=35(-31),

bs= 4(-5); F=( 83-)85, bs= 5. The 7q (5.3 -)5. 6 times as long as t,

F 2 and F 3 1.8(4. 9) and 4.4 times as long as s respectively. Distal

calyces subhemispherical; distal part of flagellar axes fusiform.

Globulus g proportionately large, 1.4 times as long as wide; 15

bracts and capsule bottom flattened; width of ^1.1 times as long

as the greatest diameter of t. Antennae glabrous.

Tnmfc. -Setae of collum segment clavate, densely annulate-

striate, distal segment large, hemispherical, rudiments of

secondary branches probably absent. Sublateral setae 1.8 times

as long as submedian ones; sternite process triangular, blunt;

appendages narrowing distally and with flat caps; process and

basal segment of appendages with distinct, almost erect, short

pubescence.

Setae on tergites thin, cylindrical, on anterior tergites as

on head, on posterior tergites tapering pointed with short

pubescence distally; 4+4 setae on tergite I, 6+6 on II-IV, 6+4

on V, 4+2 on VI. Submedian posterior setae on VI (0.8-)0.9 of

their distance apart and about as long as pygidial setae a r

Tergites glabrous.

Relative lengths of bothriotricha: 7)=100, 7^-106(413),

7 7 i -(127-)135, 7;=(112-)140, 7;=166(-170), all with simple

straight axes, very thin except in proximal 2/3 of Ty T 3 with

whip-like distal half; pubescence thin erect except on T 3 , there

oblique on thickened part, more distally erect and in distinct

whorls, most distal part annulate with a subhemispherical

distal segment. A specimen from Mount Victoria with T 3

thinner, without widened distal segment.

Legs. -Setae on coxa and trochanter of leg 9 similar, furcate

with subcylindrical blunt branches. Tarsus of leg 9 short,

somewhat tapering, 2.8 times as long as its greatest diameter.

Setae subsimilar, thin, cylindrical, striate; their length 0.2 of the

length of tarsus. Cuticle of tarsus with very delicate pubescence.

Pygidium. Tergum. -Posterior margin between st with large

semicircular lobe. Relative lengths of setae: a ; -100, a 2 -73(-80),

306

U. Scheller

Figs. 82-93. Decapauropus terrestris sp.nov., holotype: 82, head, median and right part, tergal view; 83, temporal organ, posterolateral part,

lateral view; 84, left antenna, tergal view; 85, collum segment, median and left part, sternal view; 86, tergite VI, posterior part; 87, Tp 88, 77; 89,

Tp 90, seta on trochanter of 9th pair of legs; 91, tarsus of 9th pair of legs; 92, pygidium, median and left part, sternal view; 93, anal plate, lateral

view. Scale line a for figure 88; b for figures 86, 87; c for figures 82, 83, 85, 90, 91; d for figures: 84, 89, 92, 93.

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

307

a i =127(- 140), V=47(-60); setae almost straight, blunt; a p a 2

and a 3 directed posteriorly, st pointing inwards, almost

glabrous except a } being striate in distal half. Distance a 1 -a 1

1.4 times as long as ay distance a f a 2 (1.3-)1.4 times as long as

distance a 2 -ay distance st-st (1.5-)1.8 times as long as st and

1.2 times as long as distance a 1 -a y Cuticle glabrous.

Sternum. -Posterior margin between with a broad

indentation below anal plate. Relative lengths of setae (a 7 -100):

& ; =(253-)287, A,=93(-107), setae cylindrical, densely striate,

b 2 with short, oblique pubescence. The b } (1.6-)1.8 times as long

as their distance apart; b 2 as long as distance b ] -b r Anal plate

narrowest anteriorly and with somewhat concave lateral margins,

being 1.2 times as long as broad and with two short, rounded,

posterior lobes separated by a V-shaped incision; the plate

glabrous and with two very short, cylindrical, blunt, shortly

pubescent appendages on the sternal side of the posterior lobes.

Etymology.- From Latin terra, terrestris = of the earth (soil

living).

Distribution in Tasmania. Seems to be a rare but very widely

distributed species.

Genus Nesopauropus Scheller

Type species: Nesopauropus ceylonicus (Scheller, 1970): 63-65,

fig. 29.

Nesopauropus tasmaniensis sp.nov.

(Figs. 94-104)

Material examined. Holotype. Ad.9 (female). Mount Michael, Loc. 6,

(41°10.9'S,148°00.4'E), soil core, ll.xi.1989, (RC).

Diagnosis. Nesopauropus tasmaniensis is the sixth species in

the genus. It is easily distinguished from the previously

described species, three from Sri Lanka (Scheller 1970) and

two from the Seychelles (Scheller 1982), by the shape of the

anal plate: very short, blunt, parallel, posterior appendages and

in between a distinct median incision. Disregarding the very

dissimilar anal plate, the new species may have most in common

with two species from Sri Lanka, subtilis Scheller and

unifibratus Scheller (Scheller 1970).

Description. Length.- 0.63 mm.

Head.- Tergal setae annulate, submedian ones rather short,

somewhat clavate, sublateral and lateral ones at least of

medium length, subclavate-cylindrical. Relative lengths of

setae, 1st row: a= 10, a 2 - 14; 2nd row: a= 10, a 2 - 25, fl^-22; 3rd

row: 10, a 2 -ll\ 4th row: a i =ll, a 2 -l, a 3 - ?, a 4 = 18; lateral

group setae: ^-30, l 2 = 28, 1 = 25. The ratio a/a ] -a 1 in 1st row

0.9, 2nd row 0.5, 3rd row 0.8 and 4th row 1.3. Temporal organs

oval in tergal view, 1.2 times as long as shortest distance apart;

small aperture at posterior margin. Head cuticle glabrous.

Antennae.- Segment 4 with four subclavate-subcylindrical

annulate blunt setae; their relative lengths: p= 100, p'= 34,

p"= 18, r about 16. Neither p'" nor u. Tergal seta p 1.8 times as

long as tergal branch t. The latter fusiform, 2.3 times as long

as its greatest diameter and LI times as long as sternal branch

5 which is 1.6 times as long as its greatest diameter; anterodistal

corner of s truncate. Seta q somewhat clavate, annulate, blunt,

about 1.2 times as long as 5. Relative lengths of flagella (basal

segments included) and basal segments: F ; -100, bs= 8; F 2 = 35,

bs 2 - 5; F= 88, bs 3 -l. The F } 4.6 times as long as t, F 2 and F 3 1.8

and 4.4 times as long as 5 respectively. Distal calyces

subhemispherical, on F 2 and F 3 very small; distal part of

flagella axis fusiform. Globulus g 1.3 times as long as wide; 10

bracts and capsule subspherical; width of g 0.9 of the greatest

diameter of t. Antennae glabrous.

Trunk.-Setae of collum segment simple, somewhat clavate,

annulate, blunt. Sublateral ones 1.7 times as long as submedian

ones; sternite process very small pointed; appendages

proportionally large, almost cylindrical with subhemispherical

caps; process and basal segment of appendages with distinct

almost erect pubescence.

Tergites II and III weakly divided transversally, II between

the groups of setae (two groups of setae, 6+6) and III more

posteriorly (two groups of setae, 8+4). Setae on tergites thin,

(sub)cylindrical, annulate, blunt, not decreasing in length

posteriorly; 4+4 setae on tergite I, 6+6 on II-V, 4+2 on VI.

Submedian posterior setae on VI 0.8 of their distance apart

and 0.8 of the length of pygidial setae a r Tergites glabrous.

Relative lengths of bothriotricha: r 7 -100, Ty= 119, 7^-110,

T 4 = 85, T=159; setae with thin, simple, straight axes, the

proximal half of T 3 and T 5 thickest. Pubescence hairs on T 5 and

on proximal 1/4 of the others short oblique, on distal 2/3 of

T-T 4 much longer, branched and arranged in whorls.

Legs- Setae on coxa and trochanter of legs 1-8 and seta on

coxa of leg 9 simple, somewhat clavate, annulate, blunt; seta

on trochanter of leg 9 furcate, branches cylindrical, annulate,

secondary branch somewhat thinner and shorter than primary

branch. Tarsus of leg 9 tapering, 3.2 times as long as its greatest

diameter. Setae subcylindrical, densely annulate, proximal

seta 0.2 of the length of tarsus and 0.7 of the length of distal

seta; the latter thicker than proximal seta. Cuticle of tarsus

with minute pubescence.

Pygidium. Tergum. -Posterior margin between st almost

straight. Relative lengths of setae: a= 10, a 2 -64, a 3 = 129, V-57;

setae tapering, pointed, striate; a 3 almost straight, a 2 and a 3

curved inwards, st somewhat S-shaped and pointing inwards.

Distance a 1 -a ] 0.7 of the length of ay, distance a ] -a 2 1.8 times

as long as distance a 2 -ay distance st-st 1.9 times as long as st

and 1.5 times as long as distance a ; -a r Cuticle glabrous.

Sternum. -Posterior margin between b ; with a broad

indentation. Relative lengths of setae (fl ; -100): b= 293, b 2

about 98. The b 3 cylindrical, striate, distal ly annulate; b 2

tapering, striate, somewhat pointing outwards. The b 1 1.5

times as long as their distance apart; b 2 about 0.9 of distance

bj-b 2 . Anal plate glabrous, as long as broad, narrowest

anteriorly, with convex lateral margins and two distal and two

sternal appendages; distal ones projecting backwards, short

and thick, cylindrical, blunt, somewhat pointing inwards,

length 1/4 of the length of the plate; sternal appendages of the

same length but thinner and with short oblique pubescence.

Etymology- A latinization of Tasmania.

Distribution in Tasmania. Known from a single locality on

north-east Tasmania.

308

U. Scheller

Figs. 94-104. Nesopauropus tasmaniensisx sp.nov., holotype: 94, head, median and right part, tergal view; 95, right antenna, sternal view; 96,

collum segment, median and left part, sternal view; 97, tergite VI, posterior part; 98, Tp 99, 77; 100, seta on coxa of 9th pair of legs; 101, seta on

trochanter of 9th pair of legs; 102, tarsus of 9th pair of legs; 103, pygidium, posterior part, sternal tergal view; 104, anal plate, lateral view. Scale

line a for figures 98, 99; b for figures 100-102; c for figures 94-97, 103, 104.

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

309

Stylopauropoides Remy

Type species: Stylopauropoides tiegsi (Remy, 1949): 54-56, Fig.

1 a-e.

The genus has its main distribution in the Southern Hemisphere,

has twenty-two species and two only have been reported to the

north of the equator, in the Ivory Coast and Guinea. A

provisional list of species with distributions is given below.

Species

Localities

References

1. S. tiegsi (Remy)

Australia (eastern)

Remy 1949

New Zealand

Remy 1952b, 1956a, 1956b

2. S. novaehollandiae

Australia (eastern)

Harrison 1914

3. S. bornemisszai Remy

Australia (western)

Remy 1957, Postle et al. 1991

4. S. ringueleti Remy

Argentina

Remy 1962

Chile

Scheller 1968

Tasmania

New record

5. S. rounsevelli sp.nov.

Tasmania

New record

6. S. lambda Remy

New Zealand

Remy 1956b

7. S. subantarcticus Scheller

Crozet Islands

Scheller 1974

8. S. infidus (Remy)

New Zealand

Remy 1956a

9. S. duplex (Remy)

New Zealand

Remy 1956a

10. S. erectus sp.nov.

Tasmania

New record

11. S. hetaeros sp.nov.

Tasmania

New record

12. S. scissus sp.nov.

Tasmania

New record

13. S. bilobatus Scheller

New Caledonia

Scheller 1993

14. S. hirtus (Remy)

New Zealand

Remy 1952b, 1956a

15. S. quadripartitus sp.nov

Tasmania

New record

16. S. delamarei (Remy)

Ivory Coast

Remy 1948

Guinea

Remy 1959a

17. S. vadoni (Remy)

Madagascar

Remyl956c, Remy & Bello 1960

18. S. eximius sp. nov.

Tasmania

New record

19. S. incisus Remy & Bello

Madagascar

Remy & Bello 1960

20. S.furcillatus (Remy)

New Zealand

Remy 1952b

New Caledonia

Scheller 1993

21. S. dytanekes Scheller

Brazil

Scheller 2000

22. S. salazarae Scheller

Argentina

Scheller et al. 2004

Stylopauropoides ringueleti Remy, 1962

Material Examined. 38 specimens. Loc. 2, in moss on ground, 2 ad. 9

(female), 1 subad. 8 (female), 21.iv.1989 (HM). Loc. 14, leaf litter, 19

ad. 9 (9 male, 10 female), 2 subad. 8 (male, female), 1 juv. 6, 2 juv. 5,

xii.1988 (MN). Loc. 15, litter, 10 ad. 9 (6 male, 4 female), 22.xii.1988

(MN). Loc. 19, leaf litter, 1 ad. 9 (female), 20.vi.1989 (PG).

Taxonomic remarks. The differences between the type material

from southern Argentina and Chile and the Tasmanian

populations are inconsiderable. In the latter the antennal

globulus seems to be less spherical, the st less clavate and the

T 3 have more erect pubescence and longer branches than has

been reported by Remy, but in all important characters, they

are alike.

Distribution in Tasmania. Known from the northern half of the

State only.

General distribution. Previously known from southern

Argentina (Remy 1962) and southern Chile (Scheller 1968).

Stylopauropoides erectus sp.nov.

(Figs. 105-116)

Material Examined. Holotype. Ad. 9 (female). Savage River Pipeline

Road, Loc.l, (41°18.5'S, 145°16.3'E), litter, 21.iv.1989 (PG).

Paratypes. Same data as holotype, 5 ad. 9 (female), 2 juv. 5.

Other material. 160 specimens. Loc. 1, in moss on Nothofagus, 3

ad. 9 (female), 1 juv. 6, l.iv.1989 (JD), and in moss on ground, 4 ad. 9

(female), 2 subad 8 (male, female), 21.iv.1989 (HM), and in litter 2 ad.

9 (female), 21.iv.1989 (JD,HM), and in moss on log, 1 ad. 9 (female).

310

U. Scheller

Figs. 105-116. Stylopauropoides erectus sp.nov., holotype 105-115, paratype 116: 105, head, median and right part, tergal view; 106, temporal

organ, posterior part with pistil, lateral view; 107, left antenna, tergal view; 108, 3rd antennal segment, tergal view; 109, collum segment, median

and left part, sternal view; 110, tergite VI, posterior part; 111, Ty 112, seta on coxa of 9th pair of legs; 113, tarsus of 9th pair of legs; 114,

pygidium, tergal view; 115, anal plate, lateral view; 116, anal plate, sternal view. Scale line a for figures 105, 106, 110-113; b for figures 107-109,

114-116.

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

311

21.iv.1989 (JD,HM), and in moss on ground, 1 ad. 9 (female), 21.

iv.1989 (JD). Loc. 2, in moss on ground, 1 ad. 9 (female), 1 juv. 6, and

in leaf litter, 1 ad. 9 (female), 1 juv. 5, 21.iv.1989 (JD). Loc. 4, in moss

on ground, 3 ad. 9 (2 male, 1 sex?), 1 subad. 8 (female), 1 juv. 6, 17.

xi.1989 (RC,HM), and in moss on log, 23 ad. 9 (female), 16 subad. 8

(female), 8 juv. 6, 3 juv 5, 1 juv. 3, 18.xi.1989 (RC,HM), and in moss

on myrtle, 1 ad. 9 (female), 1 juv. 6, 18.xi.1989 (HM), and in in moss

on ground, 1 ad. 9 (female), 18.xi.1989 (HM). Loc. 5, in moss on log,

1 subad. 8 (female), 29.xi.1989 (RC), and PKD, 1 juv. 3, 25.xi.1989

(HM). Loc. 6, soil core, 5 ad. 9 (4 male,l female), 1 subad. 8 (female),

1 juv. 6, 11. xi.1989 (RC). Loc. 7, in moss on log, 2 subad. 8 (female), 2

juv. 6, 17.V.1989 (HM), and in moss on ground, 1 ad. 9 (female),

12.V.1989 (PG), and without site description and date, 2 ad. 9 (female),

1 subad. 8 (female), 4 juv. 6, 1 juv. 5. Loc. 11, in leaf litter, 1 ad. 9

(female), and in moss on base of myrtle trunk, 2 ad. 9 (female), 2 juv.

6, and in moss on fallen logs, 15 ad. 9 (female), 6 subad. 8 (female), 5

juv. 6, 1 juv. 3, 21.iii.1989 (PG,JD). Loc. 17, in non-myrtle litter, 9 ad.

9 (female), 3 subad. 8 (female), 7 juv. 6, 5 juv. 5, 4 juv. 3, 8.iii.l989

(PG). Loc. 18, leaf litter, 1 ad. 9 (female), xii.1987 (MN).

Diagnosis. Stylopauropoides erectus belongs to a group of

species in the genus having V-shaped anal plates with short-

stalked appendages distally, among them S. bornemisszai

Remy from west Australia (Remy 1957) but is well distinguished

from it by the shape of the posterior setae of tergite VI and the

distal appendages of the anal plate (cylindrical and similar to a

drawing pin respectively in S. erectus , clavate and irregularly

ovoid in S. bornemisszai). There are also distinct similarites

with S. ringueleti from south Argentina and Chile (Remy 1962,

Scheller 1968) diverging by the shape of the bothriotricha T s

and the st (distal half of T 3 densely provided with branched

pubescence hairs in S. erectus, sparsely provided with thin

branches with short pubescence in S. ringueleti ; st cylindrical,

not clavate). A third similar species is S. subantarcticus

Scheller from the Crozet Islands (Scheller 1974) but the shape

of the antennal globulus is a good separating character (with

short thick stalk, not long conical) as is also the shape of the T 3

(proximal half weakly thickened, not thick clavate). S. erectus

has some characters not often met with in the genus. The F 2 are

longer than the F 3 , a character shared with S. subantarcticus,

and the posterodistal corner of the sternal antennal branch is

more truncate than the anterodistal one, also occurring in

S-infidus Remy from New Zealand (Remy 1956a). The new

species is distinguished from S. infidus by the shape of the

posterior part of the pygidial tergum (with broad rounded bulge

in S. erectus, with median indentation in S. infidus ), by the

proportion st-st/st (9-12, not about 4) and by the shape of the

distal part of the pygidial setae b } (distal part undulated, not

evenly curved). There are also similarities in direction S. tiegsi

Remy from mainland Australia and New Zealand (Remy 1949,

1956a) but in that species the antennal flagella F 3 are as long as

or longer than the F 2 , the st proportionately longer and the

distal part of the pygidial setae b / is straight, not undulated.

Description. Length.-(0.10-)0.13(-0.9S) mm.

Head.- Submedian setae on the tergal side of median

length, subclavate, with somewhat uneven pubescence, lateral

setae fairly long, cylindrical, with short pubescence. Relative

lengths of setae, 1st row: a= 10, a 2 -(8-)ll(-12); 2nd row:

a=( 11)12, a=( 14)15, a 3 =(14-)16; 3rd row: z/ ; -( 1 1)14.

a 2 =(15-)18; 4th row: a=( 13 )18, a 2 =(19-)22, a $ =(20-)2%

a 4 -16(-22); lateral group setae: / ; -(24-)31, l={ 26-)33,

l ={21 -32). The ratio aja 1 -a 1 in 1st row 1.0, 2nd row 0.8, 3rd

row ( 1. 1 -) 1.2(- 1 .3) and 4th row (1.241.5. Length of temporal

organs 0.7(-0.8) of their shortest distance apart; in a depression

of the cuticle in posterior half of the organ anterior of / ; and l 2

a clavate curved vesicle almost 0.2 of the length of temporal

organ. Head cuticle glabrous.

Antennae. -Segment 3 with three setae and rudiment of

globular organ. Segment 4 with five cylindrical annulate blunt

setae; their relative lengths: p= 100, p -(53-)64(-75),

J p"=(48-)52(-57), p"'= 32(-40), u={ 8-)9. Tergal seta p

(0.8-)1.0(-l.l) times as long as tergal branch t. The latter

fusiform, (2.4-)2.9(-3.1) times as long as its greatest diameter

and (almost) as long as sternal branch 5, that branch ( 1 .9-)2.2(-2.3)

times as long as its greatest diameter; posterodistal comer of 5

much more truncate than anterodistal one. Seta q cylindrical,

annulate, blunt, (0.8-)0.9 of the length of 5. Relative lengths of

flagella (basal segments included) and basal segments: F ; -100,

bs =(8-)9(-10); F 2 =(76-)87(-89), fes 2 =8(-10); F =(47-)52(-53),

bs=(l-)8(-9). The (2.9-)3.3(-3.6) times as long as t, F 2 and F s

(2.4-)2.8 and (1.5-)1.7(-1.9) times as long as s respectively.

Distal calyces hemispherical, largest on Fp distal part of flagella

axes not widened. Globulus g (1.4-)1.6 times as long as wide; at

least 10 bracts, capsule bottom flattened; width of g 0.6(-0.7) of

the greatest diameter of t. Antennae with faint pubescence.

Trunk. Setae of collum segment furcate; primary branch

folioform with distinct oblique pubescence, secondary branch

rudimentary, cylindrical, glabrous. Sublateral setae as long as

(-somewhat longer than) submedian ones; sternite process

triangular, anterior part narrow and with distinct incision;

appendages subconical, caps flat with collar; process with

faint lateral pubescence, appendages almost glabrous. Setae

on anterior tergites thin, cylindrical, annulate, blunt, more

posteriorly increasing in length and with short pubescence;

4+4 setae on tergite I, 6+6 on II-V, 4+2 on VI. Submedian

posterior setae on VI 0.4 of their distance apart and (as long

as-)1.2 times as long as pygidial setae a r

Relative lengths of bothriotricha: r ; -100, r 2 -(101-)107

and 114(420), ^=106-114, 7^-( 1 17) 1 22(142),

r 5 -(132-)150(-187); axes thin, simple, straight, those in T 3

moderately thickened. Pubescence hairs on T 5 and on proximal

1/4 of the others short, oblique, on distal 2/3 of T-T 4 much

longer, branched distally and arranged in whorls.

Legs- Setae on coxa and trochanter of leg 9 furcate, clavate,

with oblique short pubescence, secondary branch protruding

from the middle of the primary one and reaching 0. 5-0.7 of the

length of primary branch, these setae more anteriorly with

rudimentary glabrous secondary branches. Tarsus of leg 9

(2.6-) 2.9 times as long as its greatest diameter. Proximal seta

tapering, in distal part annulate, pointed; distal seta cylindrical

annnulate blunt. Proximal seta (0.4-)0.5 of the length of tarsus

and (1.5-)1.9 times as long as distal seta. Cuticle of tarsus with

minute pubescence.

Pygidium. Tergum. -Posterior margin between st with low

rounded bulge. Relative lengths of setae: a ; -100,

a 2 =(200-)215(-242), a 3 =(195-)235(-246), V=(12-)15(-18), setae

curved inwards, thin, tapering; a p a 2 and a 3 pointed, st blunt

and also pointing inwards; all setae with short, oblique

312

U. Scheller

pubescence. Distance a 1 -a 1 (2.1-)2.3 times as long as a p

distance a ] -a 2 2.0(-2.5) times as long as distance a 2 -ay distance

st-st (9.1-)11.3(-12.8) times as long as st and 0.7(-0.8) of distance

a 1 -a r Cuticle with distinct pubescence between st.

Sternum. -Posterior margin above and behind b 3 with a

large and broad triangular lobe. Relative lengths of setae

(a ; -100): & ; -(225-)245(-275); setae thin, shortly pubescent,

tapering, undulated distally, (1.1-)1.2(-1.4) times as long as

their distance apart. Anal plate almost upright, V-shaped,

branches somewhat S -shaped, slender, somewhat increasing in

width distally, each branch with a distal appendage in the

shape of the head of a thread-nail; plate and appendages with

very distinct almost erect pubescence, longest on distal part of

the branches.

Stage juv. 5. Pygidial setae d 2 thin, their length somewhat

more than 0.1 of their distance apart; d 3 absent.

Etymology- From Latin erectus = steep, upright (anal plate).

Distribution in Tasmania. Like the preceding species the

known range is restricted to northern Tasmania but its frequency

in moss and litter samples may indicate a wider distribution.

Stylopauropoides rounsevelli sp.nov.

(Figs. 117-127)

Material Examined. Holotype. Ad.9 (female), Bruny Island, Mount

Mangana, Loc. 12, (43°22.1'S, 147°17.0'E), litter, 4.iv.l989 (PG).

Paratypes. Same data as holotype, 2 ad. 9 (male, female). Cradle

Mountain Loc. 4, (41°35.4'S, 145°55.9'E), in moss on ground, 4 ad. 9

(female), 17.xi.1989 (RC).

Other material. 84 specimens. Loc. 1, Fungi sampled ad. 9

(female), 1 juv. 6, 21.iv.1989 (JD,HM). Loc. 5, soil core, 1 ad. 9

(female), 29.xi.1989 (PG). Loc. 6, in moss on log, 4 ad. 9 (female), 1

juv 6, ll.vi.1990 (ATW), and in litter, 3 ad. 9 (female), 1 subad. 8

(female), 29.xi.1989 (HM), and in pitfall trap, 2 ad. 9 (female), 24-29.

xi.1989 (RC), and suction, 1 ad. 9 (female), 28.xi.1989 (RC). Loc. 7,

soil cores, 1 ad. 9 (female), 1 juv. 5, 12.V.1989 (HM,JD). Loc. 9, soil

core, 1 ad. 9 (male), 20.x. 1989 (RC). Loc. 12, in litter, 2 ad. 9 (female),

4.iv.l989 (PG), and in leaf litter, 1 ad. 9 (female), 1 subad. 8 (male), 1

juv. 6, 4.iv.l989 (JD,PG), and in moss, 16 ad. 9 (4 male, 12 female), 6

subad. 8 (2 male, 4 female), 6 juv. 6, 6 juv. 5, 5 juv. 3 (JD). Loc. 13, in

litter, 11 ad. 9 (4 male, 7 female), 7 subad. 8 (2 male, 5 female), 3 juv.

6, 27.ii.1989 (SS). Loc. 16, in litter, 2 ad. 9 (male, female), 20.xii.1988

(PG). Loc. 17, soil core, 1 juv. 6, 5.vi.l989 (PG).

Diagnosis. S. rounsevelli is a well defined species reason of the

posterior vesicle of the temporal organs, the thin setae and the

anal plate with distally thickened branches and appendages

similar to nail-heads. It may be related to the preceding species,

S. erectus, but is distinguished from that species by the

following characters: antennal flagellae F 2 and F 3 subequal in

length in rounsevelli, F 2 1.4-1. 7 times as long as F 3 in erectus-,

the vesicle of the temporal organs straight and attached to the

cuticle very near the posterior margin anterior of / ; and l 2 , not

curved and situated near the middle of the temporal organ; the

posterior margin of the pygidial sternum with low bulge

between the b p not a large triangular one; the st thin and

pointed, not cylindrical and blunt; the branches of the anal

plate straight and with a short but marked distal swelling, not

somewhat curved and subclavate. There is evidence to consider

S. rounsevelli related to S. bornemisszai Remy from Western

Australia (Remy 1957), but the differences are several, the

shape of the tergal head setae, the shape of the st and the

branches of the anal plate and their appendages.

Description. Length.-{ 0.50-)0.60(-0.84) mm.

Head.- Tergal setae annulate, submedian ones of median

length, subcylindrical, blunt, sublateral and lateral ones fairly

long, cylindrical. Relative lengths of setae, 1st row: a= 10,

a 2 =(9-)10(-ll); 2nd row: c/ ; -(9-)l 1(12), a 2 =13(-14), ^-(13-) 17;

3rd row: a^(9-)10, a 2 -10(-13); 4th row: a ; -10(-12), a 2 -17(-18),

a 3 -23(-21), a 4 -ll(-13); lateral group setae: / ; -24(-29),

l~22{-21). / = 24(417). The ratio aja 1 -a 1 in 1st row l.O(-l.l),

2nd row 0.5(-0.7), 3rd row 1.0(-1.2) and 4th row 0.8(-0.9).

Length of temporal organs (0.8-)0.9 of their shortest distance

apart; near posterior margin on a level with 1 ; an aperture in

the cuticle and in a depression anterior of and l 2 an exterior

clavate straight vesicle; length of vesicle 0.2 of the length of

temporal organ. Head cuticle and temporal organs faintly

pubescent.

Antennae.- Segment 4 with six cylindrical annulate setae;

their relative lengths: p=l00,p -(74-)85(-86), -(50-)51(-57),

^> m -(18-)20(-29), r=(26-)31(-33), u=( 6-)8. Tergal seta p

(0.8-)0.9(-1.0) of the length of tergal branch t. The latter

fusiform, (2.5-)2.8 times as long as its greatest diameter and

about as long as sternal branch, that branch 1.8(-2.2) times as

long as its greatest diameter; distal corners of s equally

truncate. Seta q cylindrical, annulate, pointed, 0.6(-0.7) of the

length of Relative lengths of flagella (basal segments

included) and basal segments: F ; -100, &s ; -(9-)10);

F 2 =(77-)79(-81), *w 2 =9(-11); F^(78-)79(-82), bs= 10(12). The

Fj 3.0(-3.5) times as long as t, F 2 and F 3 (2.2-)2.5 and (2.3 -)2. 5

times as long as 5 respectively. Distal calyces hemispherical

with subglobular swelling of the flagellum axis between it and

the most distal lamella, calyces largest on F p subdistal part of

flagella axes not widened. Globulus g 1.4(-1.5) times as long as

wide; about 10 bracts, capsule bottom convex; width of g 0.7 of

the greatest diameter of t. Antennal branches glabrous, but the

basal segments of flagella with delicate pubescence.

Trunk.-Setae of collum segment furcate; primary branch

folioform with short almost erect pubescence, secondary

branch rudimentary, cylindrical, glabrous. Sublateral setae

(1.2-)1.3 times as long as submedian ones; sternite process

triangular, anterior part narrow and with distinct incision;

appendages subconical, caps rather flat and thick; process and

appendages with short pubescence.

Setae on tergites thin, cylindrical, annulate, blunt, more

posteriorly increasing in length, tapering and with short

pubescence; 4+4 setae on tergite I, 6+6 on II-V, 4+2 on VI.

Submedian posterior setae on VI (0.5-)0.6 of their distance

apart and 1.7(-1.8) times as long as pygidial setae a f

Relative lengths of bothriotricha: 7^-100, T 2 =(82-)95(-97),

r=(84-)98(-105), 7;=(85-)102(-114), 7V=fl47-)175(-180), axes

thin, simple, straight, those in proximal 2/3 of T 3 moderately

thickened. Pubescence hairs on T 5 and on proximal 1/3 of T p

T 2 and T 4 and on proximal half of T 3 simple, oblique-erect,

increasing in length outwards, on distal 1/3 of T-T 4 much

longer, branched distally and arranged in whorls.

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

313

Figs. 117-127. Stylopauropoides rounsevelli sp.nov., holotype: 117, head, median and right part, tergal view; 118, temporal organ, posterior part

with pistil, lateral view; 119, right antenna, sternal view; 120, collum segment, median and left part, sternal view; 121, tergite VI, posterior part;

122, Ty 123, T s \ 124, genital papillae and seta on coxa of 2nd pair of legs, anterior view; 125, seta on coxa of 9th pair of legs; 126, tarsus of 9th

pair of legs; 127, pygidium, sternal view. Scale line a for figures 122-126; b for figures 117, 118, 120, 121, 127; c for figure 119.

314

U. Scheller

Genital papillae subcylindrical with outer lateral part

convex, 1.7(-1.8) times as long as their greatest diameter; distal

seta 0.4 of the length of the organ.

Legs.- Setae on coxa and trochanter of leg 9 furcate, clavate,

with oblique short pubescence, secondary branch protruding

from the middle of the primary one and reaching 0.4-0.6 of

the length of primary branch. These setae are more anteriorly

with rudimentary glabrous secondary branches. Tarsus of leg

9 (3.3-)3.5(-3.7) times as long as its greatest diameter. Setae

tapering, proximal one pointed and with very short oblique

pubescence; distal seta subcylindrical and with longer

pubescence. Proximal seta 0.4(-0.5) of the length of tarsus and

1.8(-2.5) times as long as distal seta. Cuticle of tarsus with

minute pubescence.

Pygidium. Tergum. -Posterior margin between st with

rounded bulge. Relative lengths of setae: a= 100,

a 2 =(165-)167(-188), a 3 =(193-)200(-223), s/=(47-)53(-55); setae

thin, tapering, pointed, curved inwards, st also pointing

inwards. Distance 1.4(-2.0) times as long as ay, distance

a 1 -a 2 (1.7-)2.0 times as long as distance a 2 -ay distance st-st

3.0(-3.5) times as long as st and (0.8-)l.l times as long as

distance a-a r Cuticle glabrous.

Sternum. -Posterior margin above b j with a rounded lobe.

Relative lengths of setae (a ; =100): £ 7 -(164-)190(-198); setae

very thin, tapering, with short pubescence only distally,

1 .3(-1.4) times as long as their distance apart. Anal plate

V-shaped, branches slender, cylindrical but with distal

swelling, each branch with a distal appendage in the shape of

the head of a thread-nail; plate and appendages with very

distinct almost erect pubescence, longest on distal part of the

branches, distal swelling glabrous on sternal side.

Etymology. -Dedicated to David Rounsevell who initiated

the National Rainforest Conservation Program.

Distribution in Tasmania. Seems to be very widespread in

Tasmanian rainforests.

Stylopauropoides quadripartitus sp.nov.

(Figs. 128-141)

Material Examined. Holotype. Ad. 9 (female). Savage River Pipeline

Road, Loc. 1, (41°18.5'S, 145°16.3'E), litter, 21.iv.1989 (PG).

Paratypes. Same data as holotype, 1 ad. 9 (female), 1 subad. 8

(female). Sandspit River, Loc. 8, (42°42.1'S, 147°51.5'E), in leaf litter,

3 ad. 9 (2 male, 1 female), 22.V.1989 (PG).

Other material. 60 specimens. Loc. 1, in moss on ground, 1 ad. 9

(female), 21.iv.1989 (HM). Loc. 3, soil core, 1 ad. 9 (female), 9.iii.l989

(PG). Loc. 4, in moss on ground, 1 ad. 9 (female), 17.xi.1989 (RC), and

in litter, 1 juv. 5, 18. xi. 1989 (HM), and in soil core, 2 ad. 9 (male), 2

subad. 8 (female), 3-8.xi.1989 (PG). Loc. 6, suction, 1 juv. 6, 28.xi.1989

(RC). Loc. 7, tree fern crown, 4 ad. 9 (female), 1 juv. 5, 17.V.1989 (HM),

and in leaf litter, 1 ad. 9 (female), 12.V.1989 (PG). Loc. 8, in soil core,

1 ad. 9 (female), 22.V.1989 (PG). Loc. 9, in soil core, 1 ad. 9 (female),

20.X.1989 (RC), and in litter, 1 ad. 9 (female), 18.xi.1988 (PG). Loc. 11,

in moss on fallen log, 2 ad. 9 (male, female), 1 subad. 8 (female), 1 juv.

6, 21.iii.1989 (JD). Loc. 12, litter, 12 ad. 9 (4 male, 8 female), 3 subad.

8 (1 male, 2 female), 9.iv.l989 (PG). Loc. 13, litter, 14 ad. 9 (6 male, 8

female), 4 subad. 8 (1 male, 3 female), 27.ii.1989 (SS). Loc. 17, in non-

myrtle litter, 1 ad. 9 (male), 8.iii.l989 (PG). Loc. 19, leaf litter, 2 ad. 9

(female), 1 subad. 8 (male), 1 juv. 6, 20.vi.1989 (PG).

Diagnosis. Stylopauropoides quadripartitus is a long-legged

species which is clearly distinguished from its congeners by the

shape of its two-part anal plate: each half divided into two

pointed branches, one lateral and the other posterior. It may be

closest to S. hirtus (Remy) from New Zealand (Remy 1952b,

1956a) but, in that species, the posterior branches are cut

squarely and have short cylindrical appendages and the pygidial

setae a } do not reach more than 0.4-0. 5 of the length of the a 2 ,

not 0.7-0.9.

Description. Length.-(0.14-)l.25(-l.2S) mm.

Head- Submedian setae on the tergal side of medium

length, somewhat clavate, striate; sublateral and lateral setae

fairly long, cylindrical, densely striate. Relative lengths of

setae, 1st row: a = 10, a 2 -(10-)ll; 2nd row: a ; -12(-14),

a 2 -14(-15), <2 3 =15(-16); 3rd row: a ; -ll(-13), a 2 -(12-)14; 4th

row: a ; -14(-15), a 2 =a=( 18-)19, a 4 = 13(44); lateral group setae:

1= 26(-27), / 2 -(19-)24, l={ 26-)29. The ratio aja f a 1 in 1st row

1.0(4. 2), 2nd row 0.6(-0.7), 3rd row 0.7(4. 0) and 4th row

1. 1(4.3). Temporal organs in tergal view ovoid, their length

0.7(-0.8) of their shortest distance apart. Just outside the

posterior margin on a level with / ; a small aperture in the

cuticle and inside the posterior margin, in a depression, a

clavate somewhat curved exterior vesicle. The latter almost 0.2

of the length of temporal organ and attached to it with its

narrow anterior end. Head cuticle with indistinct transverse

suture between 2nd and 3rd rows of setae; head cuticle with

delicate granules, temporal organs with short erect

pubescence.

Antennae.- Segment 4 with six subcylindrical annulate-

striate setae; their relative lengths: p= 100, p -(67-)72(-79),

/>"=( 44-)56, />'"=( 21-)28, r=28(-35), u=( 9-)10. Tergal seta p

(0.8-)0.9(-l.l) times as long as tergal branch t. The latter

fusiform, (2.9-)3.2(-3.4) times as long as its greatest diameter

and as long as (-1.1) times as long as sternal branch s , that

branch (2.1-)2.2(-2.3) times as long as its greatest diameter;

posterodistal and anterodistal corners equally truncate. Seta q

subcylindrical, tapering, annulate-striate, pointed, 0.7(-0.9)

times the length of s. Relative lengths of flagella (basal

segments included) and basal segments: /q-100, £s ; -(6-)8(-9);

F 2 =(71-)74(-83), te 2 =(9-)10(41); F=79(-87), ta 3 =(9-)10(41).

The Fj (2.6-)3.0 times as long as t, F 2 and F s (1.9-)2.2(-2.5) and

(2.2-)2.4(-2.7) times as long as s respectively. Distal calyces

hemispherical; distal part of flagella axes widened only

between calyx and the distal lamella. Globulus g (1.2-)1.5

times as long as wide; (12-)13(-14) bracts, capsule subspherical;

width of g (0.7-)0.8 of the greatest diameter of t. Rudimentary

globulus on posterotergal side of 3rd segment. Antennae

glabrous.

Trunk.-Setae of collum segment furcate; primary branch

folioform with short oblique pubescence, secondary branch

rudimentary, cylindrical, glabrous. Sublateral setae as long as

(-1.2 times as long as) submedian ones; sternite process short

and broad, anterior part narrow and with very small incision;

appendages low and wide with flat caps; process with short

lateral pubescence, appendages with delicate pubescence.

Setae on anterior tergites as lateral setae on the head,

tapering posteriorly and with short, oblique pubescence; setae

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

315

Figs. 128-141. Stylopauropoides quadripartitus sp.nov., holotype 128-135, 137-141; paratype 136: 128, head, median and right part, tergal view;

129, temporal organ, posterior part with pistil, lateral view; 130, right antenna, tergal view; 131, 3rd antennal segment, tergal view; 132, collum

segment, median and left part, sternal view; 133, tergite VI, posterior part; 134, Tp 135, T 3 ; 136, genital papillae and seta on coxa of 2nd pair of

legs, anterior view; 137, seta on coxa of 9th pair of legs; 138, seta on trochanter of 9th pair of legs; 139, tarsus of 9th pair of legs; 140, pygidium,

median and left part, sternal view; 141, anal plate, lateral view. Scale line a for figures 133-136; b for figures 128, 129, 132, 137-139; c for figures

130, 131, 140, 141.

316

U. Scheller

on posterior tergites not longer than those on anterior tergites;

4+4 setae on tergite I, 6+6 on II-V, 4+2 on VI. Submedian

posterior setae on VI 0.4(-0.5) of their distance apart and

(1.1-)1.2(-1.4) times as long as pygidial setae a y

Relative lengths of bothriotricha: 7^-100, r 2 -(103-)?(-110),

7>95(-108), r 4 =116(-127), T= 149 and 180(496); axes simple,

thin, straight except proximal half of T 3 being moderately

thickened. Pubescence hairs on T 5 very short, on proximal

halves of T-T 4 stronger, oblique, increasing in length outwards

and on distal 1/3 long, ramose, whorled, most distally erect,

for the rest directed obliquely downwards.

Genital papillae (paratype) short, conical, rounded distally,

about as long as wide, glabrous; seta very thin, 0.4 of the length

of the organ.

Legs.- Setae on coxa and trochanter of leg 9 furcate, densely

pubescent, main branch leaf-shaped, broadest in the middle,

secondary branch clavate, protruding from the middle of the

primary one and reaching (0.6-)0.7 of the length of primary

branch. These setae on more anterior legs with proportionately

broader main branch and rudimentary glabrous secondary

branch. Tarsus of leg 9 (4.4-)4.5(-4.7) times as long as its

greatest diameter. Proximal seta tapering, pointed, with very

short oblique pubescence; distal seta subcylindrical, annnulate,

blunt. Proximal seta 0.4 of the length of tarsus and (2.9-)3.0

times as long as distal seta. Cuticle of tarsus with short

pubescence.

Pygidium. Tergum. -Posterior margin between st straight.

Relative lengths of setae: a = 100, <2 2 -(112-) 140(448),

<2^=181(488), V=(38-)52; setae curved inwards, tapering,

pointed, very shortly pubescent; a 1 and a 2 pointing inwards.

Distance a-a 1 (2.3 -)2 .7 times as long as ay, distance a ] -a 2 as

long as (-1.4) times as long as distance a 2 -ay distance st-st

(3.6-)4.0 times as long as st and (0.6-)0.7 of distance a-a r

Sternum. -Posterior margin above b } with a broad triangular

lobe. Relative lengths of setae (<2 ; =100): & i =(204-)214 and 236;

setae thin, shortly pubescent, tapering, 1.3 and 1.4(4. 5) times

as long as their distance apart. Anal plate directed obliquely

upwards, 1.3(4. 5) times as broad as long; divided longitudinally

into two branches each in turn divided into two branches by a

posterolateral incision; the secondary branches tapering,

pointed, the posterior ones longest; lateral branches pointing

outwards, posterior ones pointing inwards.

Stage subad. 8. Pygidial setae d } and d 2 thin pointed, the

former 0.5 of their distance apart, d 2 0.6 of distance d 1 -d 2 .

Etymology. -From Latin quadripartitus = four-parted (anal plate).

Distribution in Tasmania. Probably one of the most widespread

pauropods of the Tasmanian rainforests as it occurs in all major

regions and has been collected at nine of the 12 main collecting

sites and in three of the additional ones.

Stylopauropoides scissus sp.nov.

(Figs. 142-154)

Material Examined. Holotype. Ad. 9 (male). Savage River Pipeline

Road, Loc. 1, (41°18.5'S, 145°16.3'E), litter, 21.iv.1989 (PG).

Paratypes. Same data as holotype, 1 ad. 9 (male), 1 juv. 6. Mount

Michael, Loc. 6, (41°10.9'S, 148°00.4’E), in moss on log, 1 ad. 9 (male).

1 subad. 8 (female), ll.vi.1990 (ATW). Projection Bluff, Loc. 3,

(146°43.5’S, 41°43.1'E), in soil cores, 13 ad. 9 (8 male, 5 female), 2

subad. 8 (female), 2 juv. 6, 2 juv. 5 (PG).

Other material. 172 specimens. Loc. 1, in moss on log, 1 ad. 9

(female), 1 juv. 6, 1 juv. 5, 21.iv.1989 (HM); ibid.. Fungi sample, 2 ad.

9 (female), 1 subad. 8 (female), 21.iv.1989 (JD). Loc. 2, in moss on

dead log, 1 ad. 9 (male), 1 juv. 6, 1 juv. 3, 21. iv. 1989 (JD); in moss on

ground, 2 subad. 8 (female), 1 juv. 6, 21.iv.1989 (HM); in moss on

rocks, 2 ad. 9 (female), 1 subad. 8 (male), 1 juv. 5, 21.iv.1989 (PG). Loc.

4, in moss on logs, 8 ad. 9 (4 male, 4 female), 6 subad. 8 (3 male, 2

female, 1 sex?), 3 juv. 6, 7 juv. 5, 9.xi.l989 (PG), in moss, 7 ad. 9 (2

male, 5 female), 5 subad. 8 (1 male, 4 female), 3 juv. 6, 1 juv. 5, 18.

xi.1989 (HM). Loc. 5, in moss on log, 1 juv. 5, 29.xi.1989 (RC); PKD,

1 juv. 3, 25. xi.1989 (RC); in soil core, 1 juv. 6, 29.xi.1989 (HM). Loc.

6, pitfall trap, 1 juv. 3, 24-29.xi.1989 (RC). Loc. 7, in leaf litter, 1 juv.

6, 12.V.1989 (PG); Fungi sample, 1 ad. 9 (male), 12.V.1989 (DR). Loc.

8, hand collection, 1 ad. 9 (female), 22.V.1989 (HM). Loc. 9, in leaf

litter, 4 ad. 9 (2 male, 2 female), 3 subad. 8 (2 male, 1 female), 1 juv. 6,

2 juv. 3, 18.xi.1988 (PG); in soil cores, 2 ad. (male, female), 1 subad. 8

(female), 20.x. 1989 (RC). Loc. 10, in moss on log, 7 ad. 9 (2 male, 5

female), 3 subad. 8 (1 male, 2 female), 3 juv. 6, 1 juv. 5, 25.viii.1989

(HM); in lichen on trees, 2 ad. 9 (male, female), 1 subad. 8 (male), 1

juv. 6, 3 juv. 5, 25.viii.1989 (HM); in moss on ground, 1 ad. 9 (female),

25.viii.1989 (RC). Loc. 12, in moss on dead log, 11 ad. 9 (7 male, 3

female, 1 sex?), 5 subad. 8 (3 male, 1 female, 1 sex?), 6 juv. 6, 5 juv. 5,

2 juv. 3, 4-9.iv.1989 (JD,PG); in moss on tree, 2 ad. 9 (female), 4.

iv.1989 (JD,PG,HM); in leaf litter, 4 ad. 9 (1 male, 3 female), 3 subad.

8 (male), 4 juv. 6, 2 juv. 5, 4.iv.l989 (PG), and 6 ad. 9 (2 male, 4 female),

1 subad. 8 (female), 2 juv. 5, 2 stad.?, 9.iv.l989 (PG); in moss, 15 ad. 9

(5 male, 10 female), 1 juv. 5, 3 juv. 3, 9.iv.l989 (JD).

Diagnosis. Stylopauropoides scissus appears related to S.

duplex (Remy) from New Zealand (Remy 1956a). It is

distinguished from that species by the long-stalked antennal

globulus g (short- stalked in duplex), the shape of the T 3 (in two

parts, proximal 2/3 thickened and with dense oblique

pubescence, distal 1/3 very thin with branched hairs arranged

in whorls in scissus ; axes thin, pubescence hairs short and thin

in duplex) and the shape of the anal plate (branches

subcylindrical with subapical constriction in scissus, thickest

in distal half, rounded and without constriction in duplex', and

the distal appendages of the anal plate (with subcentral stalk in

scissus, anteroproximal in duplex).

Description. Length.-(0.16-)0.82(-1.03) mm.

Head.- The a 2 and a 3 of 4th row fairly long, other tergal

setae of medium length, submedian and anterior setae clavate,

a 3 of 2nd row and posterolateral setae subcylindrical. Relative

lengths of setae, 1st row: a= 10, « 2 — (10-)1 1(-12); 2nd row:

^=9(41), a 2 =a^l4-)16(-17); 3rd row: ^=12(44), a 2 ~?( 13-16);

4th row: a y =(13 -)14(-17), a 2 =(24-)25(-31), a=( 18-)25(-28),

a 4 =?(15-18); lateral group setae: / ; =(26-)27(-30), / 2 =(24-)25(-31),

^-(28-)34. The ratio aja 1 -a 1 in 1st row 0.8(-0.9), 2nd row

0.5(-0.7), 3rd row 0. 8(4.0) and 4th row 1. 0(4.3). Temporal

organs in tergal view narrowly ovoid, their length 0.7(-0.8) of

their shortest distance apart, posterolateral part with exterior,

clavate, somewhat curved vesicle, length = 0.1 of the length of

temporal organ. Head and temporal organs almost glabrous.

Antennae.- Segment 4 with six subcylindrical annulate-

striate setae; their relative lengths: /?=100, /P=67(-84),

p"=( 42-)43(-48), /F”-(20-)25(-27), r=20(-22), u=(6-)l. Tergal

seta p (0.8-)0.9(-l.l) times as long as tergal branch t. The latter

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

317

u 148

Figs. 142-154. Stylopauropoides scissus sp.nov., holotype: 142, head, median and right part, tergal view; 143, temporal organ, posterior part with

pistil, lateral view; 144, temporal organ, posterior part with pistil, tergal view; 145, right antenna, tergal view; 146, collum segment, submedian

and left part, sternal view; 147, tergite VI, posterior part; 148, T 149, Ty 150, genital papillae and seta on coxa of 2nd pair of legs, anterior view;

151, seta on coxa of 9th pair of legs; 152, tarsus of 9th pair of legs; 153, pygidium, posterior part, sternal view; 154, anal plate, lateral view. Scale

line a for figure 149; b for figures 145-148, 150-152; c for figures 142-144, 153, 154.

318

U. Scheller

fusiform, (3.1-)3.4(-3.7) times as long as its greatest diameter

and (0.9-)l.l times as long as sternal branch s, that branch

(2.1-)2.3(-2.5) times as long as its greatest diameter;

posterodistal and anterodistal corners equally truncate. Seta q

subcylindrical, annulate-striate, tapering distally, (0.7-)0.8 of

the length of Relative lengths of flagella (basal segments

included) and basal segments: F ; -100, bs=( 9-)ll(-12);

F 2 =(71-)87, &s 2 =(7-)9(-11); 7f=(76-)84(-85), bs=( 10-)12. The

Fj (2.3-)2.4(-2.7) times as long as t, F 2 and F 3 (2.0-)2.2(-3.0)

and (2.0-)2.2(-2.9) times as long as 5 respectively. Distal

calyces subhemispherical; distal part of flagella axes widened

only between calyx and the distal lamella. Globulus g (1.7-)1.9

times as long as wide; 9 (-10) bracts, capsule spherical; width

of g 0.6 of the greatest diameter of t. Rudimentary globulus on

posterotergal side of 3rd segment. Antennae glabrous.

Trunk.- Setae of collum segment furcate; primary branch

folioform with short almost erect pubescence, secondary

branch rudimentary, cylindrical, glabrous. Sublateral setae

1.1(-1.3) times as long as submedian ones; sternite process

rounded but with a small anteromedian incision; appendages

low and wide with flat divided caps with collar; process and

appendages with delicate pubescence.

Setae on most anterior and posterior tergites cylindrical,

medial ones clavate; 4+4 setae on tergite I, 6+6 on II-V, 4+2 on

VI. Submedian posterior setae on VI 0.4 of their distance apart

and (1.3-)1.4(-1.7) times as long as pygidial setae a f

Relative lengths of bothriotricha: 7 , / -100,7 , 2 -(98-)110(-112),

7=?(1 14-136), 7)=(140-) 144(455), 7 5 =(160-)168(-210), axes

thin, simple, straight except 7) being moderately thickened in

proximal 2/3. Pubescence hairs on 7). and on thickened part of

T 3 very short, on proximal halves of T p T 2 and T 4 thicker,

oblique, increasing in length outwards and on distal 1/3 of

Tj-T 4 long, erect, ramose, whorled.

Genital papillae short, subcylindrical, rounded distally,

1.2(-1.3) times as long as wide, glabrous; seta very thin and

short, 0.4 of the length of the organ.

Legs.- Setae on coxa and trochanter of leg 9 furcate, densely

pubescent, main branch leaf-shaped, broadest in the middle,

secondary branch clavate, protruding from the middle of the

primary one and reaching 0.6(-0.7) of the length of primary

branch; these setae more anteriorly with proportionately

broader main branch and the secondary branch rudimentary

and glabrous. Tarsus of leg 9 (3.7-)4.2(-4.4) times as long as its

greatest diameter. Proximal seta tapering, pointed, with very

short oblique pubescence; distal seta subcylindrical annnulate

blunt. Proximal seta 0.4(-0.5) of the length of tarsus and

(2.1-)2.3(-2.5) times as long as distal seta. Cuticle of tarsus

with minute pubescence.

Pygidium. Tergum. -Posterior margin between st straight.

Relative lengths of setae: a ; -100, <2 2 -(171-)200(-231),

<2 i -(247-)250(-308), V-(48-)50(-64). All but st curved inwards,

tapering, pointed, very shortly pubescent; a } and a 2 pointing

inwards; st cylindrical, blunt with short pubescence. Distance

a 1 -a 1 3.2(-3.7) times as long as ay distance a 1 -a 2 1.1(-1.4) times

as long as distance a 2 -ay distance st-st (2.7-)3.8(-3.9) times as

long as st and 0.5 of distance a-a f

Sternum. -Posterior margin between b } with deep

indentation and large posteromedian rounded lobe. Relative

lengths of setae (a ; -100): £> ; =(277-)317(-320); setae thin,

somewhat tapering, shortly pubescent in proximal half, striate

distally, (as long as-)l.l times as long as their distance apart.

Anal plate directed somewhat upwards, somewhat broader

than long and divided longitudinally into two branches

separated by a V-shaped posteromedian incision; branches

with subapical constriction; each branch with a distal stalked

subhemispherical appendage; plate and appendages with very

distinct almost erect pubescence.

Etymology.- From Latin scissus = forked, cloven (anal plate).

Distribution in Tasmania. This species is the most widely

distributed of all the Tasmanian species. It was collected at 11

of the 12 main collecting sites.

Stylopauropoides hetaeros sp.nov.

(Figs. 155-168)

Material Examined. Holotype. Ad. 9 (female). Big Sassy Creek, Loc.

7, (42°08.5'S, 147°54.3'E), soil core, 12.V.1989 (PG,DR).

Paratypes. Mount Michael (41°10.9'S, 148°00.4'E), litter, 1 ad. 9

(male), 1 subad. 8 (female), 2 juv. 5, 4juv. 3, 1 stad. ?, 29.xi.1989 (HM).

Mount Victoria, Loc. 5, (41°20.4'S, 147°49.9'E), litter, 2 ad. 9 (female),

1 subad. 8 (male), 1 juv. 5, 29.xi.1989, (RC). Simons Road, Loc. 17,

(41°21.5',147°31.3'), non-myrtle litter, 1 ad. 9 (female), 8.iii.l989

(PG).

Other material. 30 specimens. Loc. 1, in moss on dead trunk, 2

ad. 9 (female), 31.iv.1989 (PG), and in moss on ground, 6 ad. 9 (3 male,

3 female), 2 subad. 8 (male, female), 21.iv.1989 (HM). Loc. 4, litter, 1

juv. 5, 18. xi. 1989 (RC). Loc. 5, PKD, tree, 1 juv. 5, 25.xi.1989 (RC).

Loc. 6, in moss on ground, 6 ad. 9 (2 male, 4 female), ll.vi.1990

(ATW), and in pitfall trap, 1 subad. 8 (female), 24-29.xi.1989 (RC),

and in litter, 2 juv. 5, 4 juv. 3, 1 juv. stad.?, 29.xi.1989 (HM). Loc. 7,

soil core, 1 juv. 6, 1 juv. 3, 8.iii.l989 (PG). Loc. 17, Vilt area, soil core,

1 juv. 5, 5.vi.l989 (PG).

Diagnosis. There are many similarities between S. hetaeros

and S. duplex (Remy) from New Zealand (Remy 1956a)

particularly in the shape of the setae of the tergal side of the

head, the antennal branches, the setae of the pygidial tergum

and the general shape of the anal plate. Good distinguishing

characters are: the distal part of the branches of the anal plate

(with distinct inner process in haeteros, evenly rounded in

duplex)-, the appendages of the anal plate (in the shape of a nail-

head, not clavate); the shape of the setae in the posterior row of

tergite VI (clavate, not subcylindrical).

Description. Length. -(0.82-)1.03(-1.27) mm.

Head- All tergal setae except for a 3 of 2nd row clavate and

densely pubescent with short oblique hairs, submedian setae

distinctly clavate and sublateral ones weakly, a 3 of 2nd row

cylindrical striate; lateral group setae thin cylindrical striate.

Relative lengths of setae, 1st row: a= 10, « 2 — 11(-13); 2nd row:

tf ; -ll(-12), a 2 -ll(-13), <2^=10(42); 3rd row: a ; -ll(-12),

a 2 — ( 1 1 -) 13 (- 14); 4th row: c/,-1 1(13), a=( 12-)14(-15),

c/ 3 — (1 3 -) 1 4(- 1 5), a 4 -(12-)14(-16); lateral group setae: /,-(22-)26,

Z 2 — 18(-22), l 3 = 23(-25). The ratio a ] /a j -a ] in 1st row ( 1.1-)1. 2(4.4).

2nd row (0.7-)0.8, 3rd row 1.1(-1.2) and 4th row 0.8. Temporal

organs in tergal view ovoid, their length (0.6-)0.7 of their

shortest distance apart; in the middle, a probable interior

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

319

Figs. 155-168. Stylopauropoides hetaerosx sp.nov., holotype, 155-161, 164-168, paratype 162, 163: 155, head, median and right part, tergal view;

156, temporal organ, posterior part with pistil, lateral view; 157, left antenna, sternal view; 158, distal part of 3rd antennal segment, tergal view;

159, collum segment, median and left part, sternal view; 160, tergite VI, posterior part; 161, Ty 162, Ty 163, left genital papilla, anterior view;

164, seta on coxa of 9th pair of legs; 165, tarsus of 9th pair of legs; 166, pygidium, posterior part, sternal view; 167, pygidial seta ay, 168, anal

plate, lateral view. Scale line a for figures 160-163, 165; b for figures 155, 156, 159, 164; c for figures 157, 158, 166-168.

320

U. Scheller

vesicle opening outwards with a very small pore in the cuticula;

length of vesicle almost 0.2 of the length of the temporal organ.

Head cuticle with very dense and short pubescence.

Antennae.- Segment 4 with six cylindrical striate-annulate

setae; their relative lengths: p= 100, p -(64-)67(-72),

^>"-(44-)57(-59), p" '=(21-)30, r=17(-19), (5-6). Tergal seta

p (0.7-)0.8(-0.9) of the length of tergal branch t. The latter

fusiform, obliquely truncate distally, branch 2.7(-3.4) times as

long as its greatest diameter and 0.9(-1.0) of the length of

sternal branch s , that branch (1.5-)1.6(-1.7) times as long as its

greatest diameter; posterodistal and anterodistal corners

equally truncate. Seta q cylindrical, annulate-striate, tapering,

pointed, (0.6-)0.7 of the length of 5. Relative lengths of flagella

(basal segments included) and basal segments: 7q=100,

bs=(S-)9; F 2 =(87-)94, bs=( 8-)9; F=(ll-)92, bs = 8. The 7q

(2.3-)2.6(-2.7) times as long as t, F 2 and F 3 (1.8-)2.6 and

(1.9-)2.5 times as long as 5 respectively. Distal calyces

hemispherical; distal part of flagella axes widened only

between calyx and the distal lamella. Globulus g with thick

stalk, (1.7-)2.2 times as long as wide; about nine bracts, capsule

spherical; width of g (0.5-)0.6 of the greatest diameter of t.

Rudimentary globulus on posterotergal side of 3rd segment.

Antennae with short pubescence.

Trunk. -Setae of collum segment furcate; primary branch

folioform with almost erect short pubescence, secondary

branch rudimentary, cylindrical, glabrous. Sublateral setae as

long as (-1.1) times as long as submedian ones; sternite process

with distinct anterior incision; appendages with flat caps;

process and appendages with minute pubescence.

Setae on tergites as submedian setae on tergal side of head

and of about the same length on all tergites; 4+4 setae on

tergite I, 6+6 on II-V, 4+2 on VI. Submedian posterior setae on

VI (0.2-)0.3 of their distance apart and (1.2-)1.6 times as long

as pygidial setae a r

Relativelengthsofbothriotricha: 7^-100, 7 7 2 -(101-)102(-105),

7^=7(81438), 7^-(105-) 107(4 35), ^-( 1 18 )1 20(172); all setae

with simple straight axes, very thin in all but T 3 , the latter

distinctly thickened in the middle. Pubescence short and

oblique on proximal 1/3 of T j and T 2 , proximal 4/5 of 7) and

9/10 of T 4 and Ty pubescence long erect whorled and partly

branched in distal 2/3 of 7) and T 2 , short and erect in most

distal part of T 4 and T y

Genital papillae (paratype) short, subcylindrical, rounded

distally, 1.5 times as wide as long, glabrous; seta thin, 0.2 of

the length of the organ.

Legs. -Setae on coxa and trochanter of leg 9 furcate, densely

pubescent, main branch leaf-shaped, broadest in the middle,

secondary branch clavate, protruding from a point just below

the middle of the primary one and reaching 0.5 of the length of

primary branch; these setae more anteriorly, with rudimentary

cylindrical glabrous secondary branch. Tarsus of leg 9 almost

straight, tapering, (3.0-)3.3(-3.7) times as long as its greatest

diameter. Setae with oblique pubescence, proximal one tapering,

pointed, distal seta subcylindrical annulate blunt. Proximal seta

0.4 of the length of tarsus and (2.1-)2.2(-2.4) times as long as

distal seta. Cuticle of tarsus with very short dense pubescence.

Pygidium. Tergum.- Posterior margin between st almost

straight. Relative lengths of setae: a ; =100, a 2 =(233-)237(-309),

a 3 - 231(-292), V=65(-81); setae curved inwards; a 3 tapering

blunt, a 2 and a 3 tapering pointed, st subcylindrical blunt;

pubescence very short in all but st, on the latter longer and

whorled; all setae pointing inwards, a ; and st distinctly, a 2 and

a 3 only a little, st thin, somewhat clavate, blunt. Distance a 1 -a 1

(3.1-)3.3(-3.7) times as long as ay, distance a ] -a 2 about twice as

long as than distance a 2 -ay distance st-st (3.0-)3.7 times as

long as s and 0.7 of distance a 1 -a 1

Sternum. -Posterior margin with large median semicircular

lobe below anal plate. Relative lengths of setae (a ; =100):

& ; -212(-357); similar to a 3 of pygidial tergum, but thicker at

base and with stronger pubescence, 0.7(-0.9) of their distance

apart. Anal plate directed obliquely upwards, about as broad

as long, divided longitudinally by a V-shaped incision into two

broad fusiform branches, the latter cut squarely and a with

distal short inner process directed obliquely inwards; each

branch with a short-stalked distal appendage in the shape of a

nail-head; pubescence short oblique, most distally and on the

appendages longer, longest on the inner processes.

Stage subad. 8. Genital papillae rather well developed,

ovoid, seta absent.

Etymology.- From Greek hetaeros = companion (to S. duplex).

Distribution in Tasmania. As are S. ringueleti and S. erectus,

this species may be confined to the northern half of the State.

Stylopauropoides eximius sp.nov.

(Figs. 169-179)

Material Examined. Holotype. Ad. 9 (female). Savage River Pipeline

Road, Loc. 1, (41°18.5'S, 145°16.3'E), in moss on log, 21.iv.1989

(JD,HM).

Paratypes. Hibbs Lagoon Loc. 13, (42°34'S, 145°19.5'E), litter, 4

ad. 9 (1 male, 3 female), 1 subad. 8 (female), 27.ii.1989 (SS).

Other material. 1 specimen. Loc. 1, in moss on ground, 1 ad. 9

(female), 21.iv.1989 (HM).

Diagnosis. Stylopauropoides eximius is a close relative of S.

vadoni Remy from Madagascar (Remy 1956c; Remy & Bello

1960). They have great similarities in the antennae, the tergites

and the pygidium, but can be distinguished by differences in

the T 5 (dense minute pubescence in eximius, sparse thorny in

vadoni), the shape of the posteromedian lobe of the pygidial

tergum (linguiform and longer than broad, not subrectangular

and broader than long), the length of the pygidial setae a p a 2

and a 3 (a } and a 2 about half of the length of a 3 , not all subequal)

and by the shape of the anal plate (branches broad and blunt,

not narrow, tapering, pointed; appendages clavate, not

lanceolate-subcylindrical).

Description. Length.-^ 0.60-)0.82 mm.

Head.- Tergal setae of medium length or fairly long with

dense short oblique pubescence, submedian ones somewhat

clavate, sublateral and lateral ones cylindrical. Relative lengths

of setae, 1st row: a= 10, a 2 =(10-)13; 2nd row: a ; =(12-)14,

a 2 -(15-)18, a=7(ll); 3rd row: a={ 9)11, &=(9-)12; 4th row:

a ; -(12-)14, a 2 -(15-)16, a 3 - 15(49), a 4 -{ 10-)12; lateral group

setae: / ; -(20-)23, Z 2 — (18-)20, / 2 -(17-)20. The ratio a ] /a ] -a 1 in

1st row 14(4.3), 2nd row (0.6-)0.7, 3rd row (1.0-)1.5 and 4th

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

321

Figs. 169-179. Stylopauropoides eximiusx sp.nov., holotype: 169, head, median and right part, tergal view; 170, temporal organ, posterior part

with pistil, lateral view; 171, left antenna, sternal view; 172, collum segment, median and left part, sternal view; 173, tergite VI, posterior part;

174, Tp 175, Ty 176, seta on trochanter of 9th pair of legs; 177, tarsus of 9th pair of legs; 178, pygidium, sternal view; 179, anal plate, lateral view.

Scale line a for figures 174-177; b for figures 169, 170, 172, 173, 178, 179; c for figure 171.

322

U. Scheller

row (1.1-)1.3. Temporal organs (0.9-) as long as their shortest

distance apart. A small, exterior, clavate, straight, vesicle

directed posteriorly and 0.2 of the length of temporal organ

just inside the posterior margin on a level with l f Head cuticle

with minute pubescence, temporal organs glabrous.

Antennae.- Segment 4 with five cylindrical striate-annulate

setae; their relative lengths: /?-100, p -69(471), /?' -37(-43),

p" -20(-23), r=22(-23). Tergal seta p 1.1 times as long as tergal

branch t. The latter fusiform, (3.3-)4.0 times as long as its

greatest diameter and ( 1 . 1 -) 1 . 2 times as long as sternal branch

s, that branch being somewhat clavate, (2.1-)2.3 times as long

as its greatest diameter; posterodistal and anterodistal corners

equally truncate. Seta q cylindrical, annulate-striate, 1.1(-1.2)

times as long as 5. Relative lengths of flagella (basal segments

included) and basal segments (one paratype only): F ; -100,

bs= 11; F 2 =F 3 -13, bs 2 =bs= 12. The F } 2.9 times as long as t,

F 2 and F 3 2.4times as long as 5. Distal calyces subhemi spherical;

distal part of flagella axes widened only just below calyx.

Globulus g 1.3(-1.4) times as long as wide; (ll-)about 13 bracts,

capsule subspherical; width of g (0.9-) as long as greatest

diameter of t. Antennae glabrous.

Trunk.-Setae of collum segment furcate; primary branch

folioform with almost erect pubescence, secondary branch

rudimentary, cylindrical, glabrous. Sublateral setae 1.5 times

as long as submedian ones; sternite process blunt with small

anterior incision; appendages with flat caps with distinct

collar; process and appendages with short pubescence.

Setae on anterior tergites as submedian setae on the tergal

side of head, on posterior tergites subcylindrical, tapering,

pointed and with very short pubescence; posterior setae 1.5-2

times longer than anterior ones; 4+4 setae on tergite I, 6+6 on

II-IV, 6+? on V, 4+2 on VI. Submedian posterior setae on VI

(holotype only) 0.9 of their distance apart and 1.2(-1.3) times

as long as pygidial setae a r

Relative lengths of bothriotricha (most of them broken or

lost): r 7 =100, T= 96(-99), 7>(130), ,77— ( 130). 7>(186); axes

simple, thin, straight except in proximal 3/4 of T y Pubescent

hairs simple oblique on T 4 and T p the main part of T 3 and on

proximal halves of 7) and T 2 , very short on T 4 and Ty hairs

long, erect, branched distally and whorled on distal halves of

T 1 and T 2 , short erect and whorled on most distal part of T y

Legs.- Setae on coxa and trochanter of leg 9 furcate, densely

but very shortly pubescent, main branch leaf-shaped, secondary

branch clavate, protruding from the middle of the primary one

and reaching (0.6-)0.7 of the length of primary branch; these

setae on more anterior legs with rudimentary, cylindrical,

glabrous, secondary branch. Tarsus of leg 9 slender, (3.8-)4.1

times as long as its greatest diameter. Proximal seta tapering,

pointed, with short oblique pubescence; distal seta

subcylindrical striate blunt. Proximal seta 0.4(-0.5) of the

length of tarsus and 2.3(-2.9) times as long as distal seta.

Cuticle of tarsus almost glabrous.

Pygidium. Tergum. -Posterior margin evenly rounded. Very

small but distinct semicircular posterior lobe above st. Relative

lengths of setae: a ; -100, a 2 -107-115, a 3 -(185-)207-223,

s/-( 46-)54(-58); setae all curved inwards; a ]y a 2 and a 3 also

tapering pointed and with short pubescence distally; a 2 and a 3

pointing outwards, st pointing inwards, cylindrical, blunt,

striate. Distance a ] -a 1 (l.O-)l.l and 1.2 times as long as ay,

distance a 1 -a 2 (1.6-)1.9 times as long as distance a 2 -ay distance

st-st (2.6-)2.7(-2.9) times as long as st and 1.3(-1.4) times as

long as distance a ] -a ] .

Sternum. -Posterior margin between b } almost straight.

Relative lengths of setae (tf^-lOO): £ ; -(286-)321-346; setae

thin, tapering, striate, ( 1 . 1 -) 1 . 2 times as long as their distance

apart. Anal plate directed obliquely upwards, somewhat longer

than broad, consisting of two subcylindrical somewhat pointing

outwards blunt branches separated by a deep U-shaped

incision, length of branches 3/4 of the length of plate; from the

sternal side of each branch, a clavate, posteriorly directed

appendage 0.5 of the length of branch; plate and appendages

with very short oblique pubescence.

Etymology.- From Latin eximius = exceptional (among the

Tasmanian representatives of the genus).

Distribution in Tasmania. The small number of localities

makes it impossible to determine the range accurately.

Genus Pauropus Lubbock

Type species: Pauropus huxleyi Lubbock, 1867:182-185, pi. 10,

figs. 1-19.

Pauropus dolosus Remy, 1956a

Material Examined. 44 specimens. Loc. 8, upstream site, soil core, 1

ad. 9 (female), 22.V.1989 (PG). Loc. 11, south track, suction, 1 ad. 9

(sex ?), 1 subad. 8 (female), 16.iii.1989 (PG), and in leaf litter, 3 ad. 9

(1 male, 2 female), 4 subad. 8 (1 male, 3 female), 2 juv. 6, 2 juv. 5,

21.iii.1989 (PG,JD), and in moss, base myrtle trunk, 1 ad. 9 (male), 2

subad. 8 (male, female), 21.iii.1989 (JD), and in moss on fallen log, 3

ad. 9 (1 male, 2 female), 5 subad. 8 (2 male, 3 female), 3 juv. 6, 1 juv.

5, 21.iii.1989 (JD), and PKD, 1 ad. 9 (female), 16.iv.1989 (PG,JD), and

north track, in moss on dead log, 8 ad. 9 (4 male, 3 female, 1 (sex?), 2

juv. 6, 3 juv. 3, 21.iii.1989 (JD). Loc. 12, litter, 1 ad. 9 (female), 9.

iv,1989(PG).

Distribution in Tasmania. Pauropus dolosus has been collected

at three sites only, all in south-eastern Tasmania.

General distribution. Known previously only from the holotype

from New Zealand (Remy 1956a).

Pauropus vandiemeni sp.nov.

(Figs. 180-191)

Material Examined. Holotype. Ad. 9 (male), Bradshaw’s Road, Mount

Murchison, Loc 2, (41°49.9'S, 145°37.0'E), leaf litter, 21.iv.1989 (JD).

Paratypes. Same data as holotype, 1 ad. 9 (male). Bruny Island,

Mount Mangana, Loc. 12, (43°22.1'S, 147°17.0’E), leaf litter, 5 ad. 9

(female), 1 subad. 8 (female), 2 juv. 6, 1 juv. 3, 4.iv.l989 (JD,PG).

Other material. 53 specimens. Loc. 1, in moss on ground, 1 ad. 9

(female), 1 juv. 3, 21.iv.1989 (HM), and in leaf litter, 2 ad. 9 (female),

21.iv.1989 (JD,HM), and suction sample from moss on ground, site 1, 1

ad. 9 (female) and site 2, 1 ad. 9 (female), 21.iv.1989 (PG). Loc. 6, hand

collection, 2 ad. 9 (female), 28.xi.1989 (RC). Loc. 8, leaf litter, 1 ad. 9

(male), 2 juv. 3, 22.V.1989 (PG), and bridge site, soil core, 1 subad. 8

(male), 22.V.1989 (JD), and PKD from Sassafras, 1 ad. 9 (sex?), 2.

vi.1989 (JD,PG). Loc. 9, leaf litter, 2 ad. 9 (male, female), 1 subad. 8

(female), 2 juv. 6, 3 juv. 5, 18.xi.1988 (PG). Loc. 10, in moss on ground/

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

323

Figs. 180-191. Pauropus vandiemeni sp.nov., holotype 180-185, 187-191, paratype 186: 180, head, median and right part, tergal view; 181,

temporal organ, posterior part, lateral view; 182, right antenna, sternal view; 183, collum segment, median and left part, sternal view; 184, tergite

VI, posteromedian part; 185, genital papillae and seta on coxa of 2nd pair of legs, anterior view; 186, genital papillae and seta of coxa of 2nd pair

of legs in subad. 8; 187, seta on coxa of 9th pair of legs; 188, seta on trochanter of 9th pair of legs; 189, tarsus of 9th pair of legs; 190, pygidium,

sternal view; 191, anal plate, lateral view. Scale line a for figures 180, 181, 183-191; b for figure 182.

324

U. Scheller

rock, 6 ad. 9 (1 male, 5 female), 2 subad. 8 (female), 25.viii.1989 (RC).

Loc. 12, in moss on dead log on ground, 4 ad. 9 (female), 4.iv.l989 (JD),

and in litter, 4 ad. 9 (female), 1 subad. 8 (male), 4.iv.l989 (PG) and, 2

ad. 9 (male), 4.iv.l989 (PG), and suction, 4 ad. 9 (female), 9.iv.l989

(PG), and in moss, 2 ad. 9 (female), 9.iv.l989 (JD). Loc. 13, litter, 2 ad.

9 (male, female), 27.ii.1989 (SS). Loc. 17, non-myrtle litter, 1 ad. 9

(female), 2 subad. 8 (male, female), 1 juv. 6, 1 juv. 5, 8.iii. 1989 (PG).

Diagnosis. Pauropus vandiemeni forms a group with P.

montanus Scheller from New Caledonia (Scheller 1993), P.

wieheorum Remy from Mauritius (Remy 1959b) and P. difficilis

Remy from Pondichery (Remy 1961) on the basis of the similar

anal plates but they differ considerably in other features. The

new species is distinguished from P. montanus by anteriorly-

incised collum process (blunt in montanus ), 6+4 setae on

tergite V (not 6+6), lanceolate pubescent setae in the posterior

row on tergite VI (not cylindrical glabrous), the tarsus of leg 9

with strong pubescence on the tergal side (not delicate), the

pygidial tergum with large triangular posterior lobe (not a small

one with posteromedian incision) and distinctly pubescent

pygidial setae a } and a 2 (not glabrous). From P. wieheorum it is

distinguished by the shape of the posterior lobe of the pygidial

tergum (large and subtriangular in vandiemeni , low with

posteromedian incision in wieheorum ), the pygidial setae a 1

(thin and pointed, not thick and blunt) and the st (inner margin

evenly curved and outer with bulge, not inner margin with

bulge and outer evenly curved). There are reliable characters to

distinguish this species from P. difficilis : the length of the

posterior setae on tergite VI in relation to their distance apart

(0.6-0.7 in vandiemeni, 1.1 in difficilis ), the pubescence on

setae a p a 2 and a 3 of the pygidial tergum (distinct from base to

apex, compared to somewhat indistinct and present only

distally) and the shape of the st (similar to a knife-blade

compared to not clavate).

Description. Length. -(0. 84-)0.93(-l. 27) mm.

Head. -Tergal setae long, with dense short oblique

pubescence, anterior and submedian ones somewhat clavate,

sublateral and lateral ones cylindrical. Relative lengths of

setae, 1st row: a= 10, <2 2 -(10-)13; 2nd row: #js|iG-)13,

a 2 -(12-)15, a=( 9)11; 3rd row: a ; =(12-)14(-15), a 2 =(15-)18; 4th

row: fl ; -(12-)13(-14), a 2 =(19-)22(-23), ^=(17-)20(-24),

a 4 -(15-)16(-18); lateral group setae: / 7 =(16-)19(-20), / 2 =(12-)14,

/^about 13(-18). The ratio aja 1 -a 1 in 1st row (0.8-)0.9, 2nd

row (0.8-)1.0, 3rd row 2.0(-2.2) and 4th row (2.2-)3.8. Temporal

organs (0.7-)0.8 of their shortest distance apart. Just inside the

posterior margin on a level with l 3 a small exterior pore. Head

cuticle with minute pubescence, temporal organs almost

glabrous.

Antennae- Segment 4 with 5 cylindrical striate-annulate

setae; their relative lengths: p~l00,p -(83-)89(-95),/fl -30(-44),

r_(30-)42(-44), m-(6-)7; p as a rudimentary knob. Tergal seta

p (0.7-)0.9 of the length of tergal branch t. The latter

subcylindrical, 3.9(-7.2) times as long as its greatest diameter

and 1.2(-1.4) times as long as sternal branch 5, that branch

slightly clavate, (3.2-)3.8 times as long as its greatest diameter;

anterodistal corner somewhat more truncate than posterodistal

one. Seta q cylindrical, striate-annulate, (as long as-)l.l times

as long as 5. Relative lengths of flagella (basal segments

included) and basal segments (paratypes only): F ; -100,

bs=(6-l)- F 2 =( 63-82), hs-m F=( 68-86), bs=(6-l). The

2. 1-3.0 times as long as t, F 2 1.8-1. 9 and F 3 1. 8-2.7 times as

long as ^ respectively. Globulus g (1.3-)1.4(-1.5) times as long

as wide, stalk thin; (11-)13 bracts, capsule with flattened

bottom, subhemispherical; width of g (0.7-)0.8 of greatest

diameter of t. Antennae very faintly pubescent.

Trunk.-Setae of collum segment furcate; primary branch

folioform with oblique pubescence, secondary branch

rudimentary, cylindrical, glabrous. Sublateral setae

(1.8-)1.9(-2.0) times as long as submedian ones; sternite process

blunt with small anterior incision; appendages low with flat

caps; process and appendages with distinct pubescence.

Setae on tergite I as submedian setae on the tergal side of

head, on II subcylindrical blunt, on III-V subcylindrical

tapering, on VI even somewhat lanceolate, but blunt. Posterior

setae about 1.5 times as long as anterior ones; 4+4 setae on

tergite I, 6+6 on II-IV, 6+4 on V, 4+2 on VI. Submedian

posterior setae on VI 0.6(-0.7) of their distance apart and

2.0-(2.3) times as long as pygidial setae a y

Relative lengths of bothriotricha (most broken or lost):

Tj-100, T 2 = 96(-99), the following paratypes only, T 3 -(12 1-136),

r 4 -(113-172), ^-(184-261), setae with simple, straight axes

increasing in thickness posteriorly, T } very thin, T 5 thickest.

Pubescence delicate, distally mostly erect.

Genital papillae subcylindrical with somewhat granular

cuticle, 2.5 times as long as their greatest width; distal seta

straight thin and 0.5 of the length of papilla.

Legs.-Posterior legs very long, anterior ones

proportionationally shorter. Setae on coxa and trochanter of

leg 9 with dense oblique pubescence, furcate, main branch

leaf-shaped, secondary branch clavate and protruding from a

point 1/4 from the base of the seta; secondary branch 0.8 of the

length of the primary branch in coxal seta, 0.9 in the seta on

trochanter; these setae more anteriorly with rudimentary,

cylindrical, glabrous secondary branch. Coxal setae in leg 2 in

males spatulate and very shortly pubescent and with a long

thin cylindrical secondary branch (as long as-)1.3 times as

long as primary branch. Tarsus of leg 9 strongly tapering,

slender, 4.3(-5.3) times as long as its greatest diameter.

Proximal seta tapering, pointed, with distinct oblique

pubescence; distal seta cylindrical striate blunt. Proximal seta

0.3(-0.4) of the length of tarsus and 2.5(-3.0) times as long as

distal seta. Cuticle of tarsus with oblique pubescence, strong

and sparse on tergal side, short and denser on ventral side.

Pygidium. Tergum. -Posterior margin with low bulge

between a 3 and a large subtriangular densely pubescent lobe

projecting backwards above st. Relative lengths of setae:

a= 100, a 2 -(lll-)122(-165), ^=(144-)150(-197), V=100(-lll);

setae all pointed, a } , a 2 and a 3 curved inwards, subcylindrical,

tapering, with distinct oblique pubescence; a 2 and a 3 divergent,

st knife-like, somewhat curved inwards, convergent, glabrous.

Distance (1.7-)1.9 times as long as ay distance a } -a 2

(1.5-)1.6(-2.0) times as long as distance a 2 -a 3 \ distance st-st

(1.1-)1.4 times as long as st and 0.7(-0.8) of distance a 1 -a y

Sternum. -Posterior margin between b 1 with low, very

broad, rounded bulge. Relative lengths of setae (a ; -100):

b=(4l0-)4ll(-5ll), 6 2 =(162-) 178(483), ^=(250-)277(-289).

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

325

The bj subcylindrical, blunt, very shortly pubescent; b 2

subcylindrical, somewhat curved inwards, tapering, pointed,

with distinct oblique pubescence; b 3 somewhat clavate, straight,

with short oblique pubescence. The b j 1.3(-1.5) times as long

as their distance apart; b 2 (1.4-)1.5(-1.7) times as long as

distance bj-by, b 3 0.7(-0.8) of distance b 3 -b y

Anal plate broadest anteriorly, directed steeply upwards,

I. 7(-2.2) times as long as broad; with two lateral and two

submedian branches; the former thin, cylindrical, blunt,

pointing outwards, curved inwards; the latter thicker, straight,

tapering, distally cut obliquely, separated by a V-shaped

incision dividing the plate into two almost unconnected halves;

submedian branches with distal, straight, thin pointed

appendages the length being 0.5 of the length of plate. Plate

and appendages with short oblique pubescence, strongest on

distal halves of appendages.

Stage subad. 8. Genital papillae conical with blunt tip; seta

absent; seta on coxa of leg 2 leaf-shaped, secondary branch

rudimentary, cylindrical, blunt and glabrous.

Etymology. -Named after the original name for Tasmania, Van

Diemen's land.

Distribution in Tasmania. Pauropus vandiemeni belongs to

the group of widely distributed species. It occurs in all main

regions of the State except Central Tasmania.

Family Brachypauropodidae

Genus Borneopauropus Scheller

Type species: Borneopauropus penanorum (Scheller, 1994), in:

Scheller etal. 1994: 8-11, figs 31-42.

Borneopauropus dignus sp. nov.

(Figs. 192-206)

Material Examined. Holotype. Ad. 9 (male). Savage River Pipeline

Road, site 2, Loc. 1, (41°18.5’S, 145°16.3'E), in litter, 21.iv.1989 (PG).

Paratypes. Sandspit River, Loc. 8, (42°42.1'S, 147°51.5'E), PKD

from Sassafras , 1 ad. 9 (female), 2.vi.l989 (JD,PG). Bradshaw’s Road,

Mount Murchison, Loc. 2, (41°49.9'S, 145°37.0'E), site 2, in leaf litter,

3 ad. 9 (2 male, 1 female), 1 subad. 8 (female), 21.iv.1989 (JD).

Other material. 55 specimens. Loc. 1, site 1, in moss on ground, 1

ad. 9 (female), 21.iv.1989 (HM), and in leaf litter, 2 ad.9 (male), 21.

iv.1989 (JD,HM), and site 2, in moss on log, 4 ad. 9 (female), 1 subad.

8 (female), 21.iv.1989 (JD,HM), and in moss on Nothofagus, 6 ad. 9 (2

male, 4 female), 1 juv. 5, 21.iv.1989 (JD), and on moss on ground, 1 ad.

9 (female), 21. iv.1989 (PG). Loc. 2, site 1, in moss on ground, 1 juv. 5,

21.iv.1989 (HM), and in leaf litter, 4 ad. (2 male, 2 female), 1 subad. 8

(male), 21.iv.1989 (PG). Loc. 6, in moss on log, 1 juv. 5, and in moss on

tree trunk, 3 ad. 9 (1 male, 2 female), 1 subad. 8 (female), 4 juv. 5,

II. vi.1990 (ATW). Loc.7, site 2, in leaf litter, 6 ad. 9 (female), 10 juv.

5, 12.V.1989 (PG). Loc. 8, in leaf litter, 1 ad. 9 (male), 1 subad. 8 (male),

22.V.1989 (PG) and suction sample, 1 ad. 9 (female), 2.vi.l989 (JD).

Loc. 13, litter, 2 ad. 9 (male, female), 1 subad. 8 (female), 1 juv. 5, 27.

ii.1989 (SS). Loc. 17, soil core, 1 ad. 9 (male), 5.vi.l989 (PG).

Diagnosis. The genus has three species, B. penanorum

(Scheller) and B. prolatus (Scheller), both from Sabah (in

Scheller et al. 1994 and Scheller, 2001 respectively), and B.

curtipes Scheller from Indonesia (Scheller 2009). The

Tasmanian species is well delineated from them all in having

the tergites II-IV transversely divided (II, IV, V in penanorum,

II, IV in prolatus, II-V in curtipes). It can also be distinguished

from them by other good characters. There are three long

uplifted posterior extensions of the temporal organs (in

penanorum and prolatus two, one large and one very small),

the process of the collum segment is very narrow (broader and

rounded in penanorum and prolatus, broad and with distinct

anterior incision in curtipes ), the posterior end of the anal plate

is cut squarely (triangular in penanorum and curtipes,

cylindrical and blunt in prolatus ), and the appendages of the

anal plate have distinct distal swelling (distally cylindrical in

the other species).

Description. Length. (0.58-)0.68(-0.85) mm.

Head. Tergal and lateral sides with 28 setae arranged as in

Figs. 192. and 193; transversal rows difficult to interpret.

Relative lengths of the five submedian setae: 10, 13, 16, 20, 21;

lateral group: / ; large bladder-shaped, relative length = 18; l 2

subclavate = (12-)18, l 3 subcylindrical striate-annulate tapering

pointed = 52. All tergal and lateral setae except l 2 and l 3

bladder-shaped with distinct and proportionally sparse oblique

pubescence. Temporal organs complicated (fig. 193), longish,

running along from mouth to posterior part of head; anterior

end with short uplifted lobe close to the mouth; posterior part

twice furcate ending in three narrow tubes with subcircular

diameter, the upper tube shortest and curved posteriorly, the

median one straight, the sternal one curved upwards; between

the latter two a low cupulate organ with a short inner conical

structure; on the tergal side of the longish median part a small

protuberance from which a wart-like pubescent process

protrudes from the head cuticle. Tergal side of head sparsely

granular, temporal organs glabrous.

Antennae. Segment 3 with a rudimentary globulus and four

setae, one clavate, distinctly pubescent and 3 subcylindrical

annulate setae. Segment 4 with 5 setae, all thin cylindrical

annulate; relative lengths of them: /?-100, p — ( 1 1 8-) 1 50( - 1 68) ,

/V -(100-)130, -(37-)69, r=50. Tergal seta p 0.4(-0.5) of the

length of tergal branch t. The latter subcylindrical, (3.1-)3.6

times as long as wide, (as long as-) 1.1 times as long as sternal

branch .v. The latter thickest in distal third and with anterodistal

comer somewhat more truncated than posterodistal one;

(1.8-)2.3 times as long as its greatest diameter and its seta q

cylindrical annulate, 0.6(-0.7) of the length of 5. Relative lengths

of flagella (basal segments included) and basal segments:

F y =100, bs~f{- 8); F=81(-92), fa 2 =(6-)7(-8); F r 79(-88),

bs= 6(-8). The F ; (3.3-)3.4(-3.6) times as long as t, F 2 and F 3

(2.6-)3.0(-3.2) and (2.6-)2.9(-3.0) times as long as s respectively.

Distal calyces low, helmet-shaped, glabrous. Globulus g with

slender stalk, (2.1-)2.2(-2.4) times as long as its greatest

diameter, the latter (0.8-)0.9(-1.0) of the greatest diameter of t.

Antennae with g and bs f bs 3 with short pubescence.

Trunk. Setae of collum segment furcate, primary branch

cylindrical, blunt, annulate; secondary branch rudimentarily

glabrous; sublateral seta 1.1(-1.3) times as long as submedian

one. Sternite process very narrow in anterior half and with

apical incision; appendages with low caps; process and

appendages shortly pubescent.

326

U. Scheller

Figs. 192-197. Borneopauropus dignus sp.nov., holotype 192, 194-197, paratype 193: 192, head, median and right part, tergal view; 193, head

with temporal organ, lateral view; 194, left antenna, sternal view; 195, collum segment, median and left part, sternal view; 196, tergites I— III;

197, tergite VI, median and right part, and pygidium, tergal view. Scale line a for figures 196, 197; b for figure 195; c for figures 192-194.

New species of Pauropoda (Myriapoda) from Tasmanian temperate rainforests

327

Figs. 198-206. Borneopauropus dignus, Scheller sp.nov., holotype 198-200, 203-206, paratypes 201, 202: 198, Tp 199, 7\; 200, F 5 ; 201, genital

papillae and seta on coxa of 2nd pair of legs, anterior view; 202, genital papillae and seta on coxa of 2nd pair of legs in subad. 8, anterior view;

203, seta on coxa of 9th pair of legs; 204, tarsus of 9th pair of legs; 205, pygidium, sternal view; 206, anal plate, lateral view. Scale line a for

figures 198, 200, 202; b for figures 199, 201; c for figures 203, 204; d: for figures 205, 206.

328

U. Scheller

Tergites I, V, VI entire, II, III, IV transversely divided

weakly in two. Number of setae on tergites (if two groups of

values, they are anterior and posterior groups respectively: I

(29-)31(-37), II 17(-24)+20, III (23-)26(-27)+(18-)24, IV

(21-)28+(14-)17(-28), V 34(-40), VI 6+4. Setae bladder-shaped

with distinct oblique-erect pubescence, stalk inserted

unsymmetrically. Cuticle of tergites pubescent.

Relative lengths of bothriotricha: 7^-100, T= 98(-105),

r ? -(78-)81(-82), T=(99-)110, 7;=(88-)89(-92), axes simple,

most proximally glabrous; proximal half of T 3 strongly clavate,

distal half very thin; other bothriotricha with thin and curved

axes; pubescence short oblique on proximal halves, erect distally;

clavate part of T 3 with pubescence arranged in sparse whorls.

Genital papillae, 1.2 times as long as their greatest

diameter, widest near the middle, glabrous; seta almost 0.5 of

the length of papilla.

Legs. All legs 5-segmented. Setae on coxa and trochanter

of leg 9 similar, simple, cylindrical, densely annulate, blunt,

without traces of secondary branches. More anteriorly these

setae of the same shape, but with short cylindrical glabrous

rudiments of secondary branches. Tarsus of leg 9 short,

tapering, 1.8(-2.7) times as long as its greatest diameter.

Proximal seta tapering, pointed, with a few oblique pubescence

hairs on tergal side; distal seta cylindrical, striate, blunt; the

former 0.2(-0.3) of the length of tarsus and (0.9-) as long as the

latter. Cuticle of tarsus with delicate pubescence.

Pygidium. Tergum. Hind margin with a shallow incision

between a r Relative lengths of setae: a= 10, a 2 -12(-15),

a J -13(-18), st- 2(-4). All but st bladder-shaped, st very short,

(cylindrical-)clavate, straight, pointing inwards; a ] straight,

subspherical, in tergal view broadly spatulate, with very dense

short erect pubescence; a 2 and a 3 somewhat curved inwards,

ovoid, with sparse but distinct oblique pubescence, these setae

very similar to those on the tergites. Distance a 1 -a 1 (2.0-)2.7

times as long as a p distance a 3 -a 2 considerably longer than

distance a 2 -a 3 ; distance st-st (7.2-)9.3 times as long as st and

0.9(-1.0) times as long as distance a ] -a ] .

Sternum. Posterior margin between rounded and with

broad lobe below anal plate; lobe with rounded posterolateral

corners and shallow median indentation. Relative lengths of

setae (pygidial a= 10): & ; =(43-)50(-62), 7? 2 =13(-15), b 3 - 9(-10),

The bj subcylindrical, tapering, striate, blunt, b 2 and b 3

cylindrical, the former tapering distally and with distinct

oblique pubescence, the latter striate blunt. The b / 1.1(-1.2)

times as long as distance b^b^ b 2 0.5(-0.7) of distance b ] -b 2

and b 3 0.2 of distance b 3 -b 3 . Anal plate twice as long as its

greatest width, widening from its base, lateral margins convex,

broadest about at the middle; distal part narrow with parallel

lateral margins and cut squarely at the end; two thin cylindrical

appendages with globular distal enlargements protrude

backwards from the middle of the sternal side, length of

appendages 0.6(-0.7) of the length of plate.

Stage subad. 8. Genital papillae weakly developed and in

the shape of small blunt cones from a low rounded base.

Stage juv. 5. The number of setae on the tergites I 13-14, II

20, III 20, IV 8(6+2) or 16(8+8) indicating an additional moult

in this stage.

Etymology. From Latin dignus = worth.

Distribution in Tasmania. The species is widely distributed

along both the western and eastern regions of the island.

Notes on collecting methods and sites

Pauropods depend upon sustained conditions of moisture and

humidity in their living space and are normally true soil

dwellers adapted to a uniform type of environment. However,

in humid climates, they sometimes, at least temporarily,

inhabit lower litter layers and can be found under moss and

under bark of rotting wood. By using different collecting

techniques in Tasmanian rainforest and by careful handling of

the material, it has appeared that pauropods are unexpectedly

abundant in moss and are probably living in the contact zone

between the moss and underlying soil or log. They were also

found in habitats not previously considered to be inhabited by

pauropods such as on tree trunks (Greenslade, 2008). The

record of both adults and juveniles of Stylopauropoides

quadripartitus sp. nov. in the crown of a tree fern indicates

that reproduction was occurring in this habitat.

Acknowledgements

Special thanks are directed to the collectors and to Penelope

Greenslade who offered the material to me. Thanks are also

due to the Parks and Wildlife Service, Forestry Commission

and the Department of the Arts, Sport, the Environment and

Territories for financially supporting the National Rainforest

Conservation Programme.

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