Memoirs of Museum Victoria 69:1-235 (2012)

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

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

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the

Bass Strait, Victoria, Australia (other than the Tanaidae)

Magdalena Blazewicz-Paszkowycz 1 ’ 2 And Roger N. Bamber 3

Abstract

Keywords

1 Laboratory of Polar Biology and Oceanobiology, University of Lodz, Banacha 12/16, PL-90-237 Lodz, Poland,

(magdab@biol.uni.lodz.pl)

2 Museum Victoria, PO Box 666, Melbourne, Victoria 3001, Australia

3 ARTOO Marine Biology Consultants, Ocean Quay Marina, Belvidere Road, Southampton S014 5QY, United Kingdom,

(roger.bamber @ artoo.co.uk)

Blazewicz-Paszkowycz, M. and Bamber, R.N. 2012. The Shallow-water Tanaidacea (Arthropoda: Malacostraca:

Peracarida) of the Bass Strait, Victoria, Australia (other than the Tanaidae). Memoirs of Museum Victoria 69: 1-235.

All of the shallow-water tanaidacean taxa (except for species of the family Tanaidae) from material collected

between 1964 and 1999 within the Bass Strait, Victoria, Australia, have been analyzed. The material had been collected

predominantly by staff from the Museum Victoria, Melbourne. The species treated here are those occurring at depths <200

m; substrata were predominantly sands. A total of 65 species in 43 genera is discussed, of which 44 species, five genera

and one subgenus are described as new, although two of the species are not named owing to inadequacy of the material.

Only nine of the species are known from elsewhere in Australia, and none from outside Australia. In addition, after

examination of more material of its type- (and only) species, the genus Annexos is synonymized with Apseudes, as is

Xanthapseudes ; Apseudes tuski is moved to Apseudopsis; subgenera of Bunakenia are rejected; the “tribes” Parapseudini

and Pakistanapseudini are raised to Subfamily rank; Magniaculeus is synonymized with Saltipedis: intraspecific variation

in Kalliapseudes obtusifrons is discussed; the genera of the Pagurapseudinae are resolved, and Pagurapseudes abrucei is

transferred to Macrolabrum\ the first male for the genus Bathytanais is described; the validity of the genus Araphuroides

is discussed, and A. io is returned to Araphura\ Protanaissus makrotrichos, P. alvesi and P.floridensis are moved to new

genera; the family Tanaopsidae is erected to accommodate the genus Tanaopsis (at least).

Tanaidacea; Apseudomorpha; Tanaidomorpha; Australia; Tasman Sea; Bass Strait

Introduction

The Tanaidacea is a group of generally small, mainly marine,

peracarid crustaceans found in benthic habitats from the

shore to the deep sea. A comprehensive review of their

biology and ecology has been presented by Larsen (2005). As

a rule, they are outnumbered in these habitats by the two

dominant peracarid groups, the Amphipoda (in shallower

waters) and the Isopoda (in deeper waters), although they

appear to be a dominant macrofaunal group on abyssal plains

where they potentially rival polychaetes in ecological

importance (Blazewicz-Paszkowycz et al. 2012).

Despite this, the group has been understudied historically.

Most species are small in the size-spectrum of macrofauna

(commonly only a few mm long, and less than 1 mm wide);

they also have a hydrophobic cuticle, which results in their

adhering to surface water-films during sample-collection. As

a result, they have been undersampled in general macrofaunal

collection, and their identification even to genus has posed

problems for the non-specialist. Fortunately, an increase in

the number of specialists studying the Tanaidacea since the

1980s, coinciding with more rigorous investigations of the

smaller taxa in the deep sea, the tropics and the Antarctic,

has begun to improve the understanding of, and the

availability of identification texts for, the group. Valuable

baselines now exist regionally (e.g. Bird & Holdich, 1989;

Gufu, 1997; Poore, 2002, 2005; Larsen, 2005; Larsen &

Shimomura, 2007b; Bamber, 2008; Edgar, 2008) and globally

(e.g. Sieg, 1980b; Sieg, 1983a; Blazewicz-Paszkowycz, 2007;

Drumm et al., 2008).

In shallow waters (<500 m), the regions where studies on

the Tanaidacea have been most comprehensive have been the

northeast Atlantic (Europe) and eastern coasts of the USA,

simply because of the history of effort in these areas. There

has also been a long history of study in the Mediterranean,

although over the last century that has concentrated on

apseudomorph species (e.g. Gufu, 2002), and more recently in

the Antarctic (e.g. Blazewicz-Paszkowycz & Sekulska-

Nalewajko, 2004; Jozwiak & Blazewicz-Paszkowycz, 2007).

As a general rule, the more comprehensive studies have

found that in shallow waters tanaidaceans are less diverse,

M. Biazewicz-Paszkowycz & R.N. Bamber

but the species have denser populations, when compared

with deep waters where diversity can be surprisingly high,

but species are often represented by only a few (or one)

individuals. It was therefore entirely unexpected when

Bamber (2005), surveying the shallow-water (0 to 40 m

depth) tanaidacean fauna of one bay in southwestern

Australia over three weeks, found 26 species in 21 genera;

further, that in quantitative sandy-beach samples of an

admittedly low-diversity community, the tanaidaceans were

the dominant peracarid group, representing 80% of the

peracarid fauna numerically.

Most recent studies in Australian waters (Larsen, 2001;

Gufu, 2006; Biazewicz-Paszkowycz & Bamber, 2007a, b;

Bamber, 2008; Edgar, 2008) have found an extraordinary

shallow-water diversity and dominance of the Tanaidacea

when compared with other parts of the world. For example,

Holdich & Jones (1983), listed 28 species in 16 genera for the

long-studied British waters, this total having risen only to 33

as a result of subsequent publications (Bird & Holdich, 1989;

Bird, 2002, 2004; Bamber, 2011); Larsen (2005) listed a total

of 58 shallow-water (undefined) species from the Gulf of

Mexico/ western Caribbean Sea. Furthermore, the large

majority of the taxa found in Australian waters was new to

science, and showed both high local diversity and high regional

distinctness. Bamber (2005) listed 28 species in 21 genera

recorded from Australian waters previously to that study

(although two species were missed from that list); a short time

later, Bamber (2008) listed a new total of 113 species in 61

genera from Australian waters. Of this impressive number,

only six species have also been recorded outside Australia

(and four of those are recorded as “cf.” in Australian waters,

and may yet prove to be distinct). Seventeen genera are

currently endemic to Australia. One entire family, the

Whiteleggiidae Gufu, 1972, is also endemic [Biazewicz-

Paszkowycz & Bamber (2007b) showed that the only other

record, of Whiteleggia multicarinata (Whitelegge, 1901) off

South Africa, was in fact a lapsus calami ]. At the regional

level, Bamber (2008) found that, of 29 species discovered in

Moreton Bay, Queensland, only six species also occurred

elsewhere in Australia.

The present paper brings together the results of four

decades of sampling in the Bass Strait, southeastern

Australia. This region lies between the Great Australian

Bight to the west and the Tasman Sea to the east, and between

Victoria in the north and Tasmania in the south, extending

for some 400 km east-to-west and 250 to 300 km north to

south (Fig. 1). It is the widest area of continental shelf of

temperate Australia (Wilson & Poore, 1987), and forms part

of the Southeast Australia Large Marine Ecosystem (LME),

considered notable for its biodiversity (e.g. Morgan, 1989).

The prevailing strong current runs from east to west. Most of

the Strait is around 50 m depth, and the substrata are

predominantly sands.

This work describes all the shallow-water tanaidacean

taxa analysed from material collected between 1964 and

1999 within the Bass Strait (see Poore, 1986; Wilson and

Poore, 1987), except for taxa in the Family Tanaidae Dana,

1849, which are being treated elsewhere. Species analyzed

are those occurring at depths <200 m (although some of

these extend to greater depths). Species occurring in adjacent

waters exclusively below 200 m will also be treated

elsewhere.

From the foregoing, it is perhaps now no surprise that

analysis of the hundreds of samples collected over that period

revealed a total of 65 species in 43 genera, of which 57 species

and eight genera were new to science. Some of this material

has already been described (Biazewicz-Paszkowycz &

Bamber, 2007a, b). The following description of all taxa from

the Bass Strait describes 44 new species, five new genera, one

new subgenus and designates one new family. The material is

held in the collections of Museum Victoria, Melbourne.

Methods

A map of the Bass Strait region, showing principal sampling

areas mentioned in the text, is given as Figure 1.

Sampling techniques, including trawls, dredges, epibenthic

sled and Smith-Mclntyre grab, are listed in Poore (1986) and

Wilson & Poore (1987); these papers also give sample-station

details, including sediments where known. Techniques for

more recent samples are not known, but none are treated

herein as quantitative.

The type material and other studied materials are

deposited at Museum Victoria (Melbourne, Australia). In the

case of very numerous species, not all the material is listed

below even when studied for confirmation of identification; in

these cases, primary types are only those specimens

designated as such herein. In addition, some previously

described material held at the Museum was re-examined to

resolve issues of identity, and in one case to designate new

type material (neotype).

Dissected material was stained with chlorazol black and

mounted in glycerine for microscopic examination. Drawings

were done with the aid of a camera lucida. Measurements are

made axially, dorsally on the body and antennae, laterally on

pereopods. Body-proportions are based on length (from

anterior of rostrum to tip of telson) versus width of pereonite

2. Morphological terminology is as in Biazewicz-Paszkowycz

and Bamber (2007b).

Systematics

Order Tanaidacea Dana, 1849

Suborder Apseudomorpha Sieg, 1980

Superfamily Apseudoidea Leach, 1814

Family Apseudidae Leach, 1814

Subfamily Apseudinae Leach, 1814

Genus Apseudes Leach, 1814

Xanthapseudes Gufu, 2008, new synonymy.

Apseudes abditospina (Biazewicz-Paszkowycz & Bamber,

2007) comb. nov.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 1. Map of the Bass Strait, showing main place names mentioned in the text and 100 m depth contour (dotted line) (partly redrawn after

Wilson & Poore, 1987).

Figure 2

Annexos abditospina Blazewicz-Paszkowycz & Bamber, 2007b,

111-116, figs 1-3.

Remarks. On its original description, Annexos abditospina

was attributed to a new genus of the Apseudinae owing to its

not having exopodites on the cheliped nor on pereopod 1. Re¬

examination of paratypes of this species has found that it does

indeed have exopodites on both of these appendages (Fig. 2),

although they readily break off. Both are of three articles, that

on the cheliped has seven marginal plumose setae on the distal

article, while the exopodite on pereopod 1 has five.

The species is therefore transferred to the genus

Apseudes, of which Annexos becomes a junior synonym.

A. abditospina was compared with, and distinguished from,

other Australian species of Apseudes during its original

description (Blazewicz-Paszkowycz & Bamber, 2007b).

With the discovery of exopodites on the cheliped and

pereopod 1, A. abditospina sits comfortably with that group

of typical Apseudes which includes the generotype, A. talpa

(Montagu, 1808).

There are distinct proximal hook-like apophyses on the

bases of the first three pereopods of this species, a feature

known elsewhere only in A. atuini (Bamber 2005), from

Western Australia, although in that species they are also

present on the posterior pereopods. Like Apseudes poorei

(Blazewicz-Paszkowycz & Bamber, 2007) (see below), this

species has distinct articulated inner-distal spines on the first

and second maxilliped palp articles. For discussion of the

relevance of this feature to Gufu’s (2008a) suggested genus

Xanthapseudes see under remarks for A. poorei below.

This species was found throughout the Bass Strait, between

38°43' and 40°22'S and 144°18' and 148°24'E, and from 22 to

79 m depth on sandy to coarse shell substrata.

Apseudes poorei Blazewicz-Paszkowycz & Bamber, 2007

A. poorei Blazewicz-Paszkowycz & Bamber, 2007b, 120-125, figs 7-9.

Xanthapseudes poorei G 11 ( 11 , 2008a, 39.

Remarks. Apseudes poorei is very similar to A. bucospinosus

Gufu, 2006, a species from Heron Island on the Great Barrier

Reef (depth not recorded), although the pereopods of the latter

species are not fully described. An unusual distinguishing

feature of A. poorei is the presence of plumose setae on the

bases of pereopods 2 and 4; the two species are distinguished

further on the setation of the mouthparts and the morphology

of the cheliped, inter alia. A. bucospinosus has strong inner-

distal “spiniform processes” rather than spines on the first and

second maxilliped palp articles. Gufu (2008a) assigned these

M. Blazewicz-Paszkowycz & R.N. Bamber

two species to a new genus, Xanthapseudes, on the basis of an

assumed similarity in the character of spines or spine-like

apophyses on the proximal maxilliped palp articles. In fact, in

this character, A. poorei is much closer to A. abditospina, yet

quite distinct on a number of other characters (for example, the

anterolateral spiniform apophyses on the pereonites and the

hook-like apophyses on the anterior pereopod bases of the

latter species). It is therefore evident that this character of one

article of the maxilliped palp alone is not a sufficient basis on

which to distinguish a separate genus.

Apseudes poorei was found throughout the Bass Strait,

between 38°00' and 40°23'S and 144°05' and 148°37'E, and

from 13 to 82 m depth on sandy substrata.

Apseudes quasimodo sp. nov.

Figures 3-5

Material examined. 1 ? with oostegites (Registration no, J58462),

holotype. Eastern Bass Strait, Stn MSL-EG 95, 37°51.70'S 148°14.60'E,

37 m depth, February 1991, coarse sand, coll. N. Coleman, Smith-

Mclntyre grab; 2 $$ with oostegites, 6 subadults, 1 juvenile (J28515),

paratypes, same sample as Holotype. 1 ? with oostegites and penial

tubercle, 3 juveniles (J28513), paratypes, Stn MSL-EG 69, 37°51.70'S

148°14.6'E, 37 m depth, 4 June 1991, coarse sand; 2 $2 with oostegites, 3

brooding ??, 23 juveniles (J28514), paratypes, Stn MSL-EG 77,

37°49.89'S 148°30.13'E, 27 m depth, 4 June 1991, coarse sand; 3 subadults

(J28512), paratypes, Stn MSL-EG 44, 37°53.18'S 148°28.96'E, 45 m

depth, 26 September 1990, sand and shell; 1 brooding 2 , 8 subadults

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 3 .Apseudes quasimodo sp. nov., holotype female. A, dorsal view; B, lateral view. Scale = 1 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 4 .Apseudes quasimodo sp. nov., female paratype (J56389). A, antennule; B, antenna; C, left mandible; C', mandible molar; D, right mandible;

E, maxillule; E', maxillule palp; F, maxilla; G, labium; H, maxilliped; H', maxilliped endite; I, epignath. Scale: A = 0.6 mm; B, I = 1 mm;

C-H = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 5. Apseudes quasimodo sp. nov., female paratype. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5;

G, pereopod 6; H, pleopod; I, uropod. Scale = 0.1 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

(J28516), paratypes, Stn MSL-EG 96, 37°51.70'S 148°14.60'E, 37 m

depth, February 1991, coarse sand; 6 99 with oostegites, 8 subadults, 7

juveniles (J28517), paratypes, Stn MSL-EG 99,37°53.39'S 148°15.40'E,

43 m depth, February 1991, coarse sand; 1 subadult (J28518), paratypes,

Stn MSL-EG 103, 37°49.89’S 148°30.13'E, 27 m depth, February 1991,

coarse sand; 1 subadult (J28519), paratypes, Stn MSL-EG 104,

37°49.89’S 148°30.13'E, 27 m depth, February 1991, coarse sand; 2 $9

with oostegites (J28520), paratypes, Stn MSL-EG 30, 37°5L77'S

148°13.63'E, 40 m depth, 25 September 1990, sand with shell; 1 9 with

oostegites (J28521), paratypes, Stn MSL-EG44,37°53.18'S 148°28.96'E,

45 m depth, 26 September 1990, sand with shell; 1 9 with oostegites

(J28522), paratypes, Stn MSL-EG 55, 37°50.63'S 148°43.47'E, 49 m

depth, 28 September 1990, sand with shell; 1 9 with oostegites (J51300),

paratypes, Stn MSL-EG VC-41-C3, 37°32.95'S 148°03.78'E, 40 m

depth. May 1998; all Eastern Bass Strait, coll. N. Coleman, Smith-

Mclntyre grab. 1 9 with oostegites (J56335), paratypes, steel wharf Stn

MSL ref C27 grab nr4, 12 m depth, 5 March 1997; 1 9 with oostegites

(J57555), paratypes. Western Bass Strait, Stn CR 89-K-5 Stn 52,38°57'S

143°27'E, 49 m depth, 08 October 1980, coarse sand, Smith-Mclntyre

grab. 2 juveniles (J57644), 1 9 with oostegites, 1 juvenile (J57665), 1 9

with oostegites (J57671), paratypes, Stn CPBS 23N, 38°20.29'S

145°14.18'E, 10 m depth, 10 March 1965, sandy gravel; 3 99 , 2 subadults

(J57668), paratypes, Stn CPBS 33S, 38°22.06'S 145°14.10'E, 13 m

depth, 5 March 1965, reef, sponge; 2 99 with oostegites (J57680),

paratypes, CPBS 23S/1 1973, ca38°21'S 145°14'E, 10 m depth; all Crib

Point Benthic Survey, Western Port, Smith-Mclntyre grab.

Description of female/hermaphrodite. Body (Fig. 3),

dorsoventrally flattened, holotype 12.7 mm long, 5.6 times as

long as wide, narrower posteriorly. Cephalothorax subrectangular,

as long as wide, anterior margin with conspicuous pointed

rostrum with “shoulders” at base. Eyes present, eyelobes with

small spine-like apophyses directed anteriorly; lateral spiniform

apophyses at anterior margin of branchial chambers. Pereonites

1,3, 5 and 6 subequal, about 0.4 times as long as cephalothorax,

pereonite 2 just shorter, pereonite 4 longest, half length of

cephalothorax; lateral margins of pereonites 1 and 2 uniformly

convex, pereonites 3 to 6 with small anterolateral spine-like

apophyses and expanded posterolaterally at attachment of coxae

(all pereonites respectively 2.6, 2.8, 2.1, 1.6, 2.0 and 1.9 times as

wide as long); ventral pointed, forwardly-curved hyposphenia on

pereonites 2, 4 and 5, but variable - rarely also on pereonite 1,

sometimes absent on pereonites 2 and 4; penial tubercle mid-

ventrally on pereonite 6. Pleon just longer than last three

pereonites together, with five free subequal pleonites bearing

pleopods; pleonites dorsally convex, over three times as wide as

long, not bearing lateral spiniform apophyses. Pleotelson less

than half-length of whole pleon, twice as long as wide, with

conspicuous lateral setae, and with pronounced mid-dorsal boss

towards anterior margin.

Antennule (Fig. 4A). Peduncle proximal article three times

as long as wide, inner margin without rugosity, with three

shorter setae in proximal half and mid-length and subdistal

tufts of four and two simple setae, outer margin with three

proximal penicillate setae and three pairs of simple setae as

figured; second article slightly longer than wide, 0.25 times as

long as article 1, with four outer distal setae, four inner distal

setae and two inner proximal setae; third article about half as

long as wide, 0.25 times as long as second, with single inner

and outer distal setae; fourth article as long as third, with

single inner distal seta. Main flagellum of 12 segments,

segments 6, 8, 10 and 12 each bearing 1 aesthetasc; accessory

flagellum of five segments.

Antenna (Fig. 4B). Proximal peduncle article simple;

article 2 with inner rugosity, single outer and inner setae at

mid-length, and bearing elongate squama with 18 simple

marginal setae; peduncle article 3 as long as wide, with one

seta; article 4 0.8 times as long as article 2, with two inner

setae; article 5 0.8 times as long as article 4, with three

penicillate setae and four longer outer simple setae. Flagellum

of 13 segments.

Labrum (not figured) rounded, distally finely setulose;

sharp epistome present. Left mandible (Fig. 4C) outer margin

rugose, bearing strong, denticulate pars incisiva, robust,

denticulate lacinia mobilis, setiferous lobe with two trifurcate,

three bifurcate and one simple setae, pars molaris (Fig. 4C')

robust, distally concave, with fine marginal spinules. Right

mandible (Fig. 4D) as left but without lacinia mobilis;

mandibular palp of three articles, proximal article longer than

wide with five inner setae, article 2 twice as long as article 1

with three longer proximal setae, two longer distal setae, and

row of about 14 shorter setae in distal half; article 3 two-

thirds length of article 2, densely setose along inner margin

and distally. Labium (Fig. 4G) with small mid-distal tuft of

setules, palp with fine lateral setules and three simple distal

setae. Maxillule (Fig. 4E, E') inner endite with finely setose

outer margin and five finely setulate distal setae; outer endite

with eleven distal spines and two subdistal setae, outer and

inner margins finely setose; palp of two articles, distally with

six setae increasing in length towards tip of article. Maxilla

(Fig. 4F) with smooth outer margin; outer lobe of moveable

endite with two finely plumose subdistal setae and six

distally-denticulate distal setae; inner lobe of moveable endite

with three distally-denticulate setae, four simple setae and

two subdistal setulose setae; outer lobe of inner endite with

three stout trifurcate distal spines, and three distal and one

subdistal setulose setae; inner lobe of fixed endite with rostral

row of over 40 setae guarding seven longer finely denticulate

setae. Maxilliped (Fig. 4H) basis naked; palp article 1 with

two inner distal setae and five setae on slight outer-distal

apophysis; palp article 2 longer than wide, with dense rows of

numerous filtering setae on inner margin, outer margin with

one slender distal spine and adjacent subdistal short, simple

setae; palp article 3 longer than wide, with six shorter and

thirteen longer simple setae in two rows along expanded inner

margin; palp article 4 with twelve distal setae. Endite (Fig.

4FT) with simple inner caudodistal seta, plumose outer

subdistal seta, outer fine distal setae and inner rod-like distal

spines. Epignath (Fig. 41) large, cup-shaped, with distally-

plumose distal seta.

Cheliped (Fig. 5A) robust. Basis 1.8 times as long as

wide, dorsally naked, ventrally with two smaller and two

longer proximal seta, mid-ventral spine-like apophysis and

tuft of four distal setae; exopodite present, 3-articled, second

article naked, elongate, distal article with nine plumose

setae. Merus narrowing proximally, with three longer simple

setae and paired short spines on ventrodistal “shoulder”.

Carpus subtriangular, widest distally (here 0.7 times as wide

as carpus length), with row of simple setae along entire free

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

ventral margin, dorsodistal and ventrodistal shorter setae.

Chela stout, propodus 1.3 times as long as wide, fixed finger;

dense row of setae along majority of ventral margin; cutting

edge of fixed finger with row of fine setules and proximal

tooth-like apophysis; dactylus with fine setae but no

apophyses on cutting edge, distal claw pointed, meeting claw

of fixed finger.

Pereopod 1 (Fig. 5B) with coxal spine-like apophysis

pronounced. Basis stout, 1.9 times as long as wide, with two

proximal dorsal setae, sparse small ventral setae, small

ventrodistal spine and adjacent setae; exopodite present,

3-articled, article 3 with four distal plumose setae. Ischium

with dense tuft of ventrodistal setae. Merus widening distally,

0.56 times as long as basis, with row of longer mesial setae,

ventral marginal setae in distal half, stout ventrodistal spine,

five dorsodistal simple setae but no dorsodistal spine. Carpus

three-quarters as long as merus, with dorsodistal stout spine

surrounded by tuft of setae, two ventral stout spines. Propodus

just shorter than carpus and articulating slightly ventral of

carpus midline, with three ventral stout spines, two dorsal

stout spines surrounded by setae. Dactylus stout, with mid¬

dorsal fine seta and fine inner denticulation, unguis short, both

together 0.85 times as long as propodus.

Pereopod 2 (Fig. 5C) more slender. Coxa without apophysis.

Basis 4.1 times as long as wide with longer dorsal setae in the

proximal half and tufts of ventral setae. Merus 0.7 times as

long as carpus, with ventrodistal slender spine. Carpus

elongate, with ventrodistal slender spine. Propodus articulating

subdistally on ventrodistal corner of carpus, just longer than

carpus, densely setose on both margins, with mid-ventral and

ventrodistal spines. Dactylus with paired mid-dorsal setae and

fine ventral denticulation, unguis short, the two together 0.63

times as long as propodus. Pereopod 3 (Fig. 5D) similar to

pereopod 2, but propodus with outer mesial and subdistal

dorsal spines.

Pereopod 4 (Fig. 5E) similar to pereopod 2 but basis with

plumose sensory setae, merus only half length of carpus and

with two ventral spines, carpus with four ventral, two distal

and one slender dorsodistal spines; propodus as long as carpus,

with dorsodistal tuft of four short and two long finely

denticulate setae, and adjacent spinulation; dactylus plus claw

0.6 times as long as propodus and shorter than longest

dorsodistal propodal setae. Pereopod 5 (Fig. 5F) similar to but

larger than pereopod 4, carpus without dorsodistal spine,

propodus with two long, lender and one short dorsodistal

spines, and with ventral row of 12 short spinules bounded

proximally, distally and mesially by small spines; dactylus

ventrally denticulate, together with claw almost as long as

propodus. Pereopod 6 (Fig. 5G) basis with both dorsal and

ventral marginal plumose setae, merus with one plumose and

three simple setae all longer than article, carpus densely setose

on all margins, with subdistal and ventrodistal spines,

propodus with tapering row of fine spines along most of

ventral margin and around distal margin; dactylus together

with claw almost as long as propodus.

Pleopods (Fig. 5H) all alike. Basis elongate, with four

inner but no outer plumose setae. Endopod and exopod

subequal, linguiform, each with about 30 plumose setae.

Uropod (Fig. 51) biramous, both rami filiform, multi-

segmented. Basis with five setae distally; exopod one-quarter

as long as endopod, with five segments; endopod elongate,

with about 22 segments.

Description of younger stages. Juveniles with slender cheliped,

exopodite with only 5 setae; hyposphenia sparse, one on pereonite

6; subadults with robust cheliped similar to that of adult, fewer

hyposphenia than adult, one on pereonite 6; no oostegites.

Etymology. Named after Quasimodo, a central character from

French author Victor Hugo’s 1831 novel Notre Dame de Paris,

who also had a distinctive dorsal hump.

Remarks. Apseudes quasimodo sp. nov. is unique amongst the

Apseudidae in having a pronounced mid-dorsal boss towards the

anterior margin of the pleotelson, as well as the anaxial

articulation of the propodus on the anterior pereopods. In the

presence of a row of small spinules on the ventral margin of the

propodus of pereopod 5 (as well as of pereopod 6), it resembles

only Apseudes sensu stricto and Paradoxapseudes (see below)

in the Apseudidae, but in the conformation of the cheliped, the

pereonites, and with spine-like apophyses at the anterior margin

of branchial chambers, inter alia, shows similarities with

Spinosapseudes and Tuberapseudes, as well as such taxa as

Apseudes grossimanus (which also has suggestions of an anaxial

articulation of the pereopod propodus) and A. tenuimanus.

Apseudes quasimodo was found in Western Port at

10-13 m depth, and in the Eastern Bass Strait off the Metung

to Mario coast (to the east of Gippsland Lakes) from 27 to 49 m

depth on coarse sandy substrata.

Genus Apseudopsis Norman, 1899

Apseudopsis tuski (Blazewicz-Paszkowycz & Bamber, 2007)

comb. nov.

Apseudes tuski Blazewicz-Paszkowycz & Bamber, 2007b, 1 lb-

120, figs 4-6.

Remarks. This species lacks anterolateral spiniform apophyses on

the pereonites, a comb of spinules on the pereopod 5 propodus,

and a dorsodistal spine on the merus of pereopod 1, so is clearly a

member of Apseudopsis Norman 1899 sensu Gu(u (2006), indeed

close in overall morphology to A. latreilli (Milne-Edwards, 1828).

Apseudopsis tuski also has unusual spination on the maxilliped

palp, as is the case for two of the species of Apseudes discussed

above, but in this case there are three spines on the outer margin

of the second article, the most distal of which is large and robust.

This species was found throughout the Bass Strait, between

37°50' and 40°07'S and 143°14' and 148°30'E, and from 18 to

84 m depth on sandy to coarse shell substrata.

Genus Spinosapseudes Gufu, 1996

Spinosapseudes colobus Blazewicz-Paszkowycz & Bamber,

2007

S. colobus Blazewicz-Paszkowycz & Bamber, 2007b, 126-127, figs

10-13.

Remarks. Spinosapseudes colobus was the second species of

the genus to be described after S. setosus (Lang, 1968), recorded

from the other side of the Tasman Sea off New Zealand at 610 m

M. Biazewicz-Paszkowycz & R.N. Bamber

depth. S. colobus is a species more compact in its pereopods, and

of a shallower distribution. This species was found throughout

the Bass Strait, between 38°39' and 40°23'S and 144°18' and

148°40'E, and from 22 to 124 m depth.

Genus Bunakenia Gufu, 1995

Bunakenia labanticheiros sp. nov.

Figures 6-9

Material examined. 1 brooding 9(J50813), holotype. Central Bass Strait

Stn VC 31 C2, 39°02.52'S 146°10.47'E, 40 m depth, 14 May 1999, coll.

N. Coleman; 1 <3, 7 99, 2 juveniles (J50814), paratypes, Eastern Bass

Strait Stn VC 31 Cl, 38°18.3’S 147°15.25’E, 40 m depth, 10 May 1998,

coll. N. Coleman; 115 specimens (J23633), paratypes. East Gippsland

Survey Stn MSL-EG 32, 37°54.07'S 148°12.09’E, 42 m depth, 25

September 1990, sand with shell; 27 specimens (J23634), paratypes.

East Gippsland Survey Stn MSL-EG 33, 37°53.42'S 148°11.87'E, 43 m

depth, 25 September 1990, sand with shell; 20 99. 5 SS (J23637),

paratypes. East Gippsland Survey Stn MSL-EG55, 37°50.63’S

148°43.47’E, 49 m depth, 28 September 1990, sand with shell; 3 99,1 <3

(J23639), paratypes. East Gippsland Survey Stn MSL-EG57,37°51.29'S

148°43.73'E, 50 m depth, 28 September 1990, sand with shell; 27

specimens (J29174), paratypes. East Gippsland Survey Stn MSL-EG71,

37°53.39'S 148°15.40'E, 43 m depth, 4 June 1991, coarse sand; 45

specimens (J28619), paratypes. East Gippsland Survey Stn MSL-EG97,

37°53.39'S 148°15.40’E, 43 m depth, February 1991, coarse sand.

Description of female. Body (Fig. 6) dorsoventrally flattened,

slender, holotype 2.4 mm long, 5.6 times as long as wide,

tapering towards posterior, glabrous. Cephalothorax

subrectangular, as long as wide, with uniform triangular

rostrum; eyelobes and eyes present. Six free subequal

pereonites, each glabrous and without lateral apophyses;

pereonite 1 trapezoidal, wider anteriorly, just less than half as

long as cephalothorax; pereonites 2 and 6 subequal,

subrectangular, half as long as cephalothorax; pereonites 3 and

5 subequal, subrectangular, two-thirds as long as cephalothorax;

pereonite 4 longest, 0.8 times as long as cephalothorax (all

pereonites respectively 2.3, 2.0, 1.5, 1.25, 1.5 and 1.8 times as

wide as long), lateral margins smooth, without apophyses.

Pleon 2.4 times as long as pereonite 6, tapering posteriorly, of

five free subequal pleonites bearing pleopods plus pleotelson;

each pleonite about five times as wide as long, without

dorsolateral rows of plumose setae. Pleotelson subrectangular,

O. 7 times as long as all pleonites together, as long as wide.

Antennule (Fig. 7A) proximal peduncle article 3.2 times as

long as wide, outer margin with numerous penicillate setae

and one subdistal and two distal simple setae, inner margin

centrally rugose, one simple proximal seta and five plumose

setae; second peduncle article 1.5 times as long as wide, 0.3

times as long as first, with penicillate outer seta and numerous

simple inner setae; third article half length of second, 1.5

times as long as wide, distally with inner and outer setae, the

former longer than article; fourth peduncle article about half

length of third. Main flagellum of 9 segments, single

aesthetascs present on fifth, eighth and ninth segments;

accessory flagellum of three segments.

Antenna (Fig. 7B), proximal peduncle article without

apophysis. Second article with linguiform squama bearing 11

marginal setae, two setae next to base of squama. Third

peduncle article twice as long as wide and two-thirds as long

as second, fourth article as long as second, fifth article half

length of fourth. Flagellum of five segments.

Labrum rounded, simple, distally setose. Right mandible

(Fig. 7C) with strong, crenulate pars incisiva, setiferous lobe

with six forked setae, outer margin finely spinose. Left mandible

(Fig. 7D) similar but with crenulate lacinia mobilis. Pars molaris

robust, with posterodistal denticulation; palp of three articles,

proximal article with five simple setae, second article longest

with two medial simple setae and, distal to these, three rows of

two, three and four simple, third article with ten setae in a single

row. Labium (Fig. 7G) with denticulate outer margin, palp with

fine lateral setules and three simple distal setae. Maxillule (Fig.

7E) inner endite with outer apophysis, finely setose outer margin

and five setulose distal setae, outer endite with eleven distal

spines and two subdistal setae (not seen on figure), outer and

inner margins finely setose, palp of two articles, distally with

three setae. Maxilla (Fig. 7F) typical of the genus, with a rostral

row of 25 setae, compound setae on the fixed endite and serrate

sickle-like setae on the moveable endite inside five compound

outer setae on the outer lobe. Maxilliped (Fig. 7H) basis with

medial inner seta; first palp article with one very long inner setae

and small, naked outer apophysis; second palp article just longer

than wide, with one row of inner simple setae, parallel row of

shorter setulose setae, and outer distal spine; third palp article

wider than long with inner distal group of thirteen simple setae;

fourth palp article mounted anaxially, with eight distal setae.

Endites (Fig. 7H') with three coupling hooks, distally with four

outer setae and numerous inner slender, blunt spines.

Cheliped (Fig. 8A) slender, smaller than pereopod 1, basis

twice as long as wide, ventrally with long proximal seta,

central sharp spine and distal group of four simple setae;

three-articled exopodite present, slender, distal article with

three plumose setae. Merus subtriangular, twice as long as

wide, ventral margin with eight setae; carpus 4.7 times as long

as wide, with longer ventral marginal setae and shorter dorsal

marginal setae. Chela slender, fixed finger just shorter than

palm with dense distal setation; dactylus and claw slightly

overreaching fixed finger, both fingers without apophyses on

cutting edge.

Pereopod 1 (Fig. 8B) with conspicuous sharp setose

apophysis on coxa; stout basis less than twice as long as wide,

dorsal margin bearing only simple setae in proximal half,

single ventrodistal spine; exopodite large, three-articled, distal

article with six plumose setae. Ischium with simple ventrodistal

setae. Merus widening distally, with single dorsodistal and

ventrodistal spines and associated simple setae. Carpus

compact, wider than long, with fan of dorsodistal setae, one

dorsodistal and two ventrodistal blunt spines. Propodus with

two dorsodistal spines and four ventral blunt spines interspersed

with single fine setae. Dactylus slender, with paired mid-dorsal

and ventral fine setae; claw slender, finely denticulate.

Pereopods 2 and 3 (Fig. 8C, D) basis 3.2 times as long as

wide with sparse penicillate setae, one (pereopod 2) or two

(pereopod 3) ventromedial setae and tuft of longer ventrodistal

setae; ischium half as long as wide; merus as long as carpus,

widening distally and with long ventral setae and single

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 6. Bunakenia labanticheiros sp. nov., holotype female. A, lateral view; B, dorsal view. Scale = 1 mm.

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 7. Bunakenia labanticheiros sp. nov., female paratype. A, antennule; B, antenna; C, right mandible; D, left mandible; E, maxillule;

E', maxillule palp; F, maxilla; G, labium; H, maxilliped; FT, maxilliped endite. Scale: A, B = 0.1 mm; C-H = 0.01 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 8. Bunakenia labanticheiros sp. nov., female paratype. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 6.

Scale = 0.1 mm.

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 9. Bunakenia labanticheiros sp. nov., male. A, antennule; B, antenna; C, juvenile male cheliped; D, adult male cheliped; E, pereopod 1;

F, pereopod 2; G, pleopod; H, uropod. Scale = 0.1mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

ventrodistal slender spine; carpus with slender ventrodistal

spine I tuft of simple setae and crown of dorsodistal simple

setae; propodus with two or three ventral and one dorsodistal

slender spines; dactylus slender, claw not denticulate.

Pereopods 4 (Fig. 8E) and 5 with slender basis with

penicillate seta and fine ventrodistal simple setae; carpus

slightly longer than merus; propodus of pereopod 4 with mid¬

dorsal penicillate seta; dactylus and claw less slender than

those of anterior pereopods.

Pereopod 6 (Fig. 8F) proportionately as pereopod 5, but

basis with ventral marginal row of five simple and three

plumose setae; merus shorter than carpus and with three

dorsal plumose setae; carpus with four dorsal plumose setae;

propodus with 20 small compound spines ventrally and

distally on each side in tapering row; dactylus plus claw

slender, simple.

Pleopods as those of male (Fig. 9G) all alike, basis with

two inner but no outer plumose setae, endopod and exopod

subequal, slender, each with 11 plumose setae.

Uropod (Fig. 9H) biramous, basis with three distal simple

and two penicillate setae; exopod just less than three times as

long as basis and of six segments; endopod elongate, filiform,

multisegmented.

Description of male, generally similar to female. Antennule

(Fig. 9A) main flagellum of 11 segments with single aesthetascs

on segments 3, 5, 7 and 9, accessory flagellum of 5 segments.

Antenna (Fig. 9B) flagellum of eight segments.

Conspicuous dimorphism of cheliped (Fig. 9D): basis 1.5

times as long as wide, ventrally with central spine and paired

distal plumose setae; three-articled exopodite present, stout,

distal article with four plumose setae. Carpus stouter, widening

distally, twice as long as wide. Chela robust, highly modified,

fixed finger flexed back along distal margin of propodus (palm),

distal tip truncate with rugose cutting edge, tooth-like apophysis

in angle between fixed finger and palm; dactylus narrowing

rapidly from base, distally truncate, with five spines but no

apophysis on cutting edge. Chela of subadult male (Fig. 9C)

intermediate between that of mature male and that of female,

carpus slender, fixed finger longer than palm, but not reflexed.

Pereopod 1 (Fig. 9E) with stout basis twice as long as wide,

dorsal margin bearing nine plumose setae in proximal half.

Ischium dorsally wide. Merus dorsal margin with flattened,

flange-like apophysis and dorsodistal spine. Pereopods 2 (Fig.

9F) and 3, merus shorter than carpus.

Etymology. From the Greek - labe - something for grasping,

and anticheiros - thumb, in reference to the extra tooth on the

fixed finger of the chela of the male.

Remarks. Gufu (1996c) described two subgenera of Bunakenia.

The nominate B. (Bunakenia), distinguished only by having

rows of plumose setae on the basis of pereopod 1, includes three

species, B. (B.) indonesiana Gufu, (1995a) from Sulawesi, at

4-5 m depth, B. (B.) tanzaniana Gufu, 1996(d) from the Indian

Ocean coast of Africa at 20 m depth, and B. (B.) salzella

Bamber, 2005 from the littoral to 30 m depth in southwestern

Australia. The subgenus B. (Extensibasella), without plumose

setae on the basis of pereopod 1, includes B. (E.) sudvestatlantica

Gufu, 1996(c) from Brazil at 31 m depth, B. (E.) aspalieus

Bamber, Bird and Angsupanich, 2003, from Thailand in littoral-

infralittoral seagrass beds, B.(E.) kadazan Bamber and Sheader,

2005 from Sabah in sand at 23-35 m depth, and B. (E.) anomala

Gufu, 2006 from Moreton Bay, Australia (depth unspecified).

The present species has plumose setae on the basis of

pereopod 1 in the male, but not in the female, and thus falls

quite between the two subgenera; Bamber & Sheader (2005)

questioned the validity of the subgenera, their species B.

kadazan showing little affinity to B. (E.) sudvestatlantica, and

the zoogeography of these two “groups” is inconsistent. We

therefore choose to dispense with those subgenera.

In the conformation of the male chela, with extreme

reflexion of the fixed finger, Bunakenia labanticheiros sp. nov.

is similar only to the other Australian species, B. salzella,

from which it can be distinguished by the presence of the

tooth-like apophysis in the angle between the fixed finger and

the palm of the male chela (absent in B. salzella ), the absence

of plumose setae on the basis of pereopod 1 in the female

(present in B. salzella), the mid-ventral spine on the basis of

the cheliped (a plumose seta in B. salzella ), fewer ventral

spines on the propodus of pereopod 1, fewer plumose setae on

the basis of pereopod 6, and details of the setation of the

mouthparts and pleopods, inter alia.

Unlike the present species, B. kadazan has a thin, pointed

rostrum; B. sudvestatlantica has a more elaborate cheliped

basis in the male, and is without the tooth-like apophysis in

angle between fixed finger and palm of the male chela, as also

are B. tanzaniana and B. anomala', B. aspalieus (male

unknown) has posterolateral hook-like apophyses on the

pereonites; all four have plumose setae on both margins of the

pleopod; B. indonesiana has only one basis seta on the pleopod,

but dense rows of plumose setae on the pereopod 1 basis of

both genders, and a more pronounced rostrum. All of these

species have an inner apophysis on the proximal peduncle

article of the antenna, unlike Bunakenia labanticheiros.

All specimens were taken from sandy substrata at between

40 and 50 m depth in the Central and Eastern Bass Strait.

Genus Paradoxapseudes Gufu, 1991

Gollumudes Gufu, 1991

Remarks, in a reanalysis of material from Cuba, the type

locality for the then monotypic genus Paradoxapseudes, Gufu

(2008a) revised the morphology of the type species, P. cubensis

Gufu 1991, and realised that Gollumudes Bamber 2000 is a

junior synonym of Paradoxapseudes. As a result, he was able to

assign 12 species to the genus, which now showed a worldwide

distribution. The genus is partly characterized by the row of

leaf-like propodal spines on pereopod 5, as well as the row on

pereopod 6 found in other apseudomorphs (but see also

Apseudes). Owing to this new resource of information on the

morphological variation within Paradoxapseudes (including

Gollumudes ), the Bass-Strait material, including that attributed

to “G.” larakia Edgar (1997) by Bfazewicz-Paszkowycz &

Bamber (2007b) was re-examined, and found to be of two

distinct species, which are described below.

M. Biazewicz-Paszkowycz & R.N. Bamber

During the analysis of the morphology of all the taxa now

included in Paradoxapseudes, it also became apparent that the

material from Tanzania mentioned by Gufu (2007), and from

the Strait of Malacca and the South China Sea mentioned by

Gufu (2008a), all attributed to the Japanese species P. littoralis

(Shiino, 1952), was in fact not of that species (unlike P. littoralis,

they have serrations on the antennule peduncle article 1,

significantly more segments in the main flagellum of the

antennule, and plumose, not simple, dorsal setae on the basis of

pereopod 1, and the last two have far fewer leaf-like spines on

the propodus of pereopod 5, inter alia). It is certainly quite

unlikely that the material from Tanzania would be conspecific

with a species from Japan. This material is considered to

represent at least two further taxa, for which more detailed

description is required before diagnosis and naming.

Paradoxapseudes paneacis sp. nov.

Figures 10-12

Gollumudes larakia Biazewicz-Paszkowycz & Bamber, 2007b

(partim - shallow water specimens), non -Apseudes larakia Edgar, 1997.

Material examined. 1 9 (J58580), holotype, 1 <3 (J58581), allotype, 155

further specimens (J57662), paratypes. Crib Point Benthic Survey Stn

CPBS 33S/2, Western Port, 38°21.60'S 145°13.67'E, 13 m depth, 12

March 1965, muddy sand. Smith McIntyre Grab. 21 specimens

(J57649), Stn CPBS 33S, same data as holotype; 1 9 with oostegites

(J55880), 3 specimens (J57659), paratypes, Stn CPBS 23N, 38°20.29’S

145°14.18’E, 10 m depth, 10 March 1965, sandy gravel; 7 specimens

(J57672), paratypes, Stn CPBS 23S, 38°21.69'S 145 0 13.51'E, 11 m

depth, 9 March 1965, muddy sand; 1 brooding 9 (J56169), 20 specimens

including 33 and brooding 99, 1 brooding 9 (J57674), paratypes, Stn

CPBS 41N, 38°20.81'S 145°13.85’E, 13 m depth, 30 March 1965, gravel

and sand; all Western Port, Crib Point Benthic Survey, Smith McIntyre

Grab. 1 9 (J56292), paratype. Western Port, “sublittoral”, 25 November

1971.

Other material (as in Biazewicz-Paszkowycz & Bamber, 2007b). 1

individual (J53143), 50 m south of Twin Reefs, Venus Bay (38°41'S,

145°39'E), 9 m, 07 March 1982, coll. M. McDonald; 1 individual

(J55759), 50 m east of Petrel Rock, Venus Bay (38°39'S, 145°42'E), 8 m,

05 March 1982, (CPA 1) coll. M. McDonald and M.F. Gomon; 5

individuals (J55761), 1 km east of Harmers Haven, 500 m offshore

(38°34'S, 145°40'E), 11 m, 06 March 1982, (CPA 14), coll. C. Larsen

and G. Barber; 1 individual (J55762), 1 km east of Harmers Haven,

300 m offshore (38°34'S, 145°40'E), 6 m, 06 March 1982 (CPA 15),

coll. R.S. Wilson and C. Larsen; 1 individual (J55763), east side of

Cape Paterson (38°41'S, 145°36’E), 6 m, 05 March 1982, (CPA 12), coll.

R.S. Wilson, G. Barber, etal:, 1 individual (J55765) Bennison Channel

1.0 km south of Granite Island (38°49'S, 146°23'E), 6.0 m, 23 November

1983, (CIN 28), coll. G.J. Morgan.

Description of female. Body (Fig. 10) dorsoventrally flattened,

holotype 2.9 mm long, 5.8 times as long as wide, tapering

towards posterior. Cephalothorax subrectangular, 1.4 times as

long as wide, with large triangular rostrum; eyelobes and eyes

present. Pereonites 1 and 2 subequal in length, 0.28 times as

long as cephalothorax, with convex lateral margins, paired

anterodorsal setae and posterolateral plumose setae; pereonite

3 longest, 1.5 times as long as pereonite 2, with anterolateral

pointed apophysis, midlateral invagination and posterolateral

rounded apophysis above pereopod attachment, and with

lateral plumose setae and paired anterodorsal setae; pereonites

4 to 6 similar to, but progressively shorter than, pereonite 3,

pereonite 6 being 1.26 times as long as pereonite 2 (all

pereonites respectively 2.7, 2.5, 1.7, 1.8, 1.9 and 1.9 times as

wide as long). Pleon three times as long as pereonite 6, narrower

than pereon, with five free subequal pleonites bearing pleopods;

each pleonite about three times as wide as long and extended

laterally into sharp, triangular apophysis bearing plumose

setae. Pleotelson subpentangular, with two rounded apophyses

bearing plumose setae on each side, as long as last three

pleonites together, just longer than wide.

Antennule (Fig. 11 A) proximal peduncle article 3.6 times

as long as wide, outer margin with penicillate setae and sparse

simple setae in distal half, inner margin with sparse simple

setae and proximal serration; second peduncle article wider

distally, 1.5 times as long as wide, 0.3 times as long as first,

with three penicillate and five simple distal setae; third article

0.7 times length of second, about twice as long as wide; fourth

peduncle article slender, half length of third. Main flagellum

of 7 segments, aesthetascs present on second, third and fifth

segments, seventh segment anaxial on sixth; accessory

flagellum of three segments.

Antenna (Fig. 11B), proximal peduncle article with

subrectangular inner apophysis bearing two plumose setae.

Second peduncle article twice as long as first, twice as long as

wide, margins sinuous, with medial and distal simple setae on

outer margin, medial and distal tooth-like apophyses on inner

margin, the former with an adjacent plumose seta; elongate

linguiform squama bearing two longer distal and two shorter

subdistal setae. Third peduncle article as long as wide and

one-quarter as long as second, with inner-distal spine-like

apophysis; fourth and fifth articles 0.7 times as long as second.

Flagellum of four segments.

Labrum (not figured) rounded, distally finely setulose. Left

mandible (Fig. 11C) with strong, crenulate pars incisiva, robust

lacinia mobilis with three distal crenulations, setiferous lobe

with two compound and three simple setae; outer margin finely

denticulate; pars molaris (Fig. 11C") robust, with radial rows of

distal rugosity; palp (Fig. 11C') of three articles, proximal article

shortest with five simple setae, second article longest with single

row of two longer and eight shorter finely setulose setae along

distal half of ventral margin, third article with 11 finely setulose

setae, distal setae much longer than more proximal setae; right

mandible (Fig. 11D) similar but without lacinia mobilis.

Maxillule (Fig. 11E) inner endite with slight outer apophysis,

finely setose outer margin and four setulose distal setae, outer

endite with eleven distal spines and two subdistal setae, outer

margin finely setose, palp (Fig. 11E') of two articles, distally

with four setae. Maxilla (Fig. 11F) outer lobe of moveable endite

with two subdistal setulose sickle-like setae and five distal

setulose setae, inner lobe with eight simple curved setae and five

stouter plumose setae; outer lobe of fixed endite with six

compound distal spines and subdistal biserrate spine, inner lobe

with three stout, proximally setulose setae and rostral row of 26

setae. Labium (Fig. 11G) with setulose outer margin, palp with

dense tufts of fine lateral setules and three simple distal seta.

Maxilliped (Fig. 11H) basis naked; first palp article with one

very long plumose inner seta and adjacent fine simple seta, and

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 10. Paradoxapseudes paneacis sp. nov., holotype, adult female. A, dorsal view; B, lateral view. Scale = 0.1 mm.

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 11. Paradoxapseudes paneacis sp. nov., female paratype. A, antennule; B, antenna; C, left mandible; C', mandible palp; C”, mandibular

molar; D, right mandible; E, maxillule, with E', detail of palp; F, maxilla; G, labium; H, maxilliped; H', maxilliped endite; I, epignath. Scale:

A, B = 0.1 mm; C-I = 0.01 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 12. Paradoxapseudes paneacis sp. nov., female paratype. A, cheliped; A', cheliped male; B, pereopod 1; C, pereopod 2; D, pereopod 3;

E, pereopod 4; F, pereopod 5; G, pereopod 6; H, pleopod; I, uropod. Scale = 0.1 mm.

M. Biazewicz-Paszkowycz & R.N. Bamber

small plumose outer seta; second palp article longer than wide,

with inner rows of five ventral plumose setae and numerous

distally-curved simple; third palp article nearly twice as long as

wide, with inner group of nine simple curved setae; fourth palp

article with one subdistal and seven distal simple setae. Endites

(Fig. 11H') with three coupling hooks, outer margin densely

setulose, distally with one simple seta and numerous slender,

blunt, bifurcate spines, outer subdistal plumose seta. Epignath

(Fig. Ill) oval, with setulose distal spine.

Cheliped (Fig. 12A) basis 1.6 times as long as wide, with

two ventroproximal setae on small tubercles, mid-ventral

curved spine, two ventrodistal plumose setae; exopodite

present, distal article with 4 plumose setae (as on male, Fig.

12A); merus with one longer and two shorter setae on

ventrodistal shoulder; carpus widening distally, subpentangular,

with stout, curved spine and three simple setae mid-ventrally;

chela robust, palm of propodus just longer than wide, fixed

finger two-thirds length of palm, ventral margin regularly

setose, cutting edge of fixed finger with crenulations and central

rugose tooth-like apophysis, numerous submarginal setae;

dactylus stout, with crenulated cutting edge.

Pereopod 1 (Fig. 12B) with conspicuous setose apophysis

on coxa; basis 2.9 times as long as wide, dorsal margin bearing

four plumose setae in proximal half, ventral margin with

single proximal plumose seta, single mid-ventral and paired

distal simple setae; exopodite three-articled, distal article with

four plumose setae. Ischium with three simple ventrodistal

setae. Merus widening distally, with ventrodistal row of simple

marginal setae, and single long dorsodistal and shorter

ventrodistal finely denticulate spines. Carpus just shorter than

merus, with setose margins including paired dorsodistal setae

longer than propodus, one dorsodistal and two ventrodistal

finely denticulate spines. Propodus 0.6 times as long as carpus,

with two dorsodistal and four ventral finely denticulate spines

interspersed with single fine setae. Dactylus slender, claw

short, together longer than propodus.

Pereopods 2 and 3 (Fig. 12C, D) coxa with rounded, setose

apophysis, basis four times as long as wide, sparsely setose,

ventrodistal seta reaching past half length of merus; ischium

half as long as wide, with fine dorsal seta, one short ventral

seta and one ventral seta longer than merus; merus shorter

than carpus, with two to four ventral setae, pair of ventrodistal

spines, one very short, and single strong dorsodistal seta;

carpus with ventrodistal spine with adjacent simple setae,

smaller submarginal ventral spines, inner distal finely-

denticulate spine and long dorsodistal simple setae exceeding

tip of propodus; propodus longer than carpus, with three or

four finely-denticulate ventral spines, one (P2) or two (P3)

dorsodistal finely-denticulate spines, dorsodistal setae

exceeding tip of claw; dactylus with ventrodistal seta, dactylus

and claw curved, together as long as carpus.

Pereopod 4 (Fig. 12E) basis three times as long as wide,

ventrodistal seta reaching past half length of merus; ischium

half as long as wide, with fine dorsal seta, two shorter ventral

setae and one ventral seta longer than merus; merus 0.6 times

as long as carpus, with one ventral seta and pair of ventrodistal

finely-denticulate spines; carpus with paired mid-distal spines

and ventrodistal row of five finely denticulate spines

interspersed with simple setae; propodus 0.8 times as long as

carpus, with two finely-denticulate ventral spines and group of

numerous finely-setulose dorsodistal setae and single

dorsodistal seta exceeding tip of claw; dactylus with

ventrodistal seta, dactylus and claw curved, together just

shorter than propodus.

Pereopod 5 (Fig. 12F) similar to pereopod 4, but merus

with single ventral spine, long dorsodistal and ventrodistal

setae reaching or exceeding tip of carpus, carpus with two

ventral spines and dorsodistal seta exceeding tip of propodus,

propodus with one short and two longer setae, one of which

longer than dactylus plus claw, and with ventral comb of 10

leaf-like spines in the distal half.

Pereopod 6 (Fig. 12G) proportionately similar to pereopod

5, basis with seven plumose dorsal setae; ischium with single

dorsal plumose seta; merus with two dorsal plumose setae;

carpus with dorsal and ventral simple setae; propodus with

small leaf-like spines ventrally and distally; dactylus plus claw

slender, curved, together as long as propodus

Pleopods typical for genus (Fig. 12H), basis with two inner

but no outer plumose setae, endopod with eight plumose marginal

setae, exopod shorter with seven plumose marginal setae.

Uropod (Fig. 121) biramous, basis with four distal simple

setae; exopod broken, of at least four segments; endopod

elongate, filiform, with about 14 segments.

Description of male. Generally similar to female, but with

dimorphic cheliped (Fig. 12A), basis stout, 1.5 times as long as

wide, with ventroproximal setae on small tubercles, mid-

ventral curved spine, ventrodistal plumose setae; exopodite

present, distal article with 4 plumose setae; merus sparsely

setose; carpus stouter than that of female, widening distally,

almost triangular, with stout, curved ventral spine; chela robust,

stouter than that of female, palm of propodus as long as wide,

fixed finger almost half length of palm, ventral margin regularly

setose, cutting edge of fixed finger with crenulations and sub-

proximal rugose tooth-like apophysis, numerous submarginal

setae; dactylus stout, with crenulated cutting edge. Setae on

most articles longer than those of female.

Etymology. Named after Crib Point, the type locality, contrived

from the Greek pahnee - a crib, and akis - a point.

Remarks. The shallow-water material described as Gollumudes

larakia in Biazewicz-Paszkowycz & Bamber (2007) is in fact

of this species, to which are added numerous further

specimens, principally collected during the Crib Point Benthic

Survey (see Poore, 1986). Paradoxapseudes paneacis sp. nov.

is unusual in the genus in having long ventrodistal and

dorsodistal setae on merus, carpus, ischium and basis of each

pereopod. It is the only Australian species with a spine on the

cheliped carpus, a feature also present in P. littoralis (from

Japan), P. garthi (Menzies, 1956) from the Gulf of California,

P. heroae(Sieg, 1986)fromtheSubantarctic,and P. intermedius

(Hansen, 1895) from the Mediterranean. P. paneacis is also

the only Australian species to have inner proximal serration

on the antennule peduncle (like only P. intermedius of the four

species listed above); as well as the long distal setae on the

pereopod article, the present species differs from P. intermedius

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

in having more mandibular palp setae, fewer basis setae on

pereopods 1 and 6, and in the conformation of the rostrum,

inter alia.

All specimens were collected in Western Port on shallow

sands at depths between 6 and 13 m.

Paradoxapseudes attenuata sp. nov.

Figures 13-15

Gollumudes larakia Blazewicz-Paszkowycz & Bamber, 2007b

(partim - deeper-water specimens), non -Apseudes larakia Edgar, 1997.

Material : 1 9 (J47130), holotype, Stn BSS109, Central Bass Strait,

40°30.9'S 144°56'E, 27 m depth, 2 November 1980, very coarse sand,

coll. M. Gomon & G.C.B. Poore; 2 99 with oostegites (J55843),

paratypes, Stn BSS117, Central Bass Strait, 40°38.0'S 145°23'E, 36 m

depth, 4 November 1980, muddy shell and grit, coll. M. Gomon &

G.C.B. Poore; 1 9 (J58464), paratype, Stn BSS161, Central Bass Strait,

39°48.3'S 147°19.2'E, 60 m depth, 14 November 1981, muddy sand,

coll. R. Wilson.

Other material: 2 99 (J55842), Stn BSS119, western Bass Strait,

39°06.7'S 143°28.7'E, 92 m depth, 31 January 1981, fine sand, coll. M.

Gomon et al .; 1 9 (J57559), Stn VC 18 C2, Central Bass Strait,

38°30.2'S 144°15.0'E, 40 m depth, 30 May 1998, coll. N. Coleman,

Smith McIntyre grab. Material in Btatewicz-Paszkowycz & Bamber

(2007b): 4 individuals (J47131), Australia, Tasmania, eastern Bass

Strait, 37 km NNE of Eddystone Point (40°43.48'S, 148°37.12'E),

67 m, 14/11/1981, (BSS 164), coll. R.S. Wilson; 1 individual (J55756),

western Bass Strait, 30 km SSW of Warrnambool (38°38.12'S, 142

35.00'E), 59 m, 20/11/1981, (BSS 188), coll. R.S. Wilson; 3 individuals

(J55757), western Bass Strait, 15 km S of Port Fairy (38°32.00'S, 142

28.36'E), 52 m, 20/11/1981, (BSS 187), coll. R.S. Wilson; 1 individual

(J55760), western Bass Strait, 15 km south of Port Fairy (38°32.00'S,

142 28.36'E), 52 m, 20/11/1981, (BSS 187), coll. R.S. Wilson; 2

individuals (J55766), Victoria, western Bass Strait, 5 km south of

Point Reginald (38°48.00'S, 143°14.30'E), 47 m, 20/11/1981, (BSS

185), coll. R.S. Wilson; 1 individual (J55758), western Bass Strait, 5

km southwest of Bluff Point (40°48.06’S, 144°38.00'E), 42 m,

02/02/1981, (BSS 126 G), coll. M.F. Gomon.

Description of female with oostegites. Body (Fig. 13)

dorsoventrally flattened, elongate, holotype 2.75 mm long,

seven times as long as wide, tapering towards posterior.

Cephalothorax subrectangular, 1.5 times as long as wide, with

triangular rostrum; eyelobes and eyes present. Pereonite 1

laterally convex, 0.23 times as long as cephalothorax; pereonite

2 1.2 times as long as pereonite 1, with slight anterolateral

apophysis bearing plumose seta, smaller posterolateral plumose

seta; pereonites 3 to 6 with anterolateral pointed apophyses and

posterolateral rounded apophyses, each bearing plumose setae,

pereonites 3 and 5 subequal, 1.8 times as long as pereonite 1,

pereonite 4 longest, twice as long as pereonite 1, pereonite 6 as

long as pereonite 2 (all pereonites respectively 2.9, 2.3, 1.5, 1.2,

1.4 and 1.9 times as wide as long). Pleon narrower than pereon,

just longer than cephalothorax, tapering posteriorly, with five

free subequal pleonites bearing pleopods; each pleonite about

3.3 times as wide as long and extended laterally into sharp,

triangular apophysis bearing plumose setae. Elongate

pleotelson subpentangular, with two rounded apophyses

bearing plumose setae on each side, as long as last four pleonites

together, 1.75 times as long as wide.

Antennule (Fig. 14A) proximal peduncle article 4.2 times

as long as wide, outer margin with slight proximal rounded

apophysis, three penicillate setae and three simple setae in

distal half, inner margin with sparse simple setae, no proximal

corrugation; second peduncle article twice as long as wide, 0.4

times as long as first, with two penicillate and five simple

distal setae; third article half length of second, about twice as

long as wide; fourth peduncle article slender, slightly shorter

than third. Main flagellum of 7 segments, single aesthetasc

present on sixth segments, seventh segment anaxial on sixth;

accessory flagellum of three segments.

Antenna (Fig. 14B), proximal peduncle article with inner

apophysis bearing small seta and two teeth. Second peduncle

article twice as long as first, twice as long as wide, margins

sinuous with mid-inner setae on apophysis, with elongate

linguiform squama bearing 6 marginal setae. Third peduncle

article as long as wide and one-quarter as long as second,

fourth article 0.8 times as long as second, fifth article half

length of second. Flagellum of five segments.

Labrum (Fig. 14C) rounded, simple, distally with rows of

setules. Left mandible (Fig. 14D) with strong, crenulate pars

incisiva, robust lacinia mobilis with five distal crenulations,

setiferous lobe with four slender setae; pars molaris robust,

with radial rows of distal rugosity; palp (Fig. 14D') of three

articles, proximal article shortest with six simple setae, second

article longest with four distomedial simple setae, third article

with six distal setae. Right mandible (Fig. 14E) similar but

without lacinia mobilis. Labium (Fig. 14H) with denticulate

outer margin, setulose distal margin, palp with fine lateral

setules and one simple distal seta. Maxillule (Fig. 14F) inner

endite with slight outer apophysis, finely setose outer margin

and four setulose distal setae, outer endite with eleven distal

spines and two subdistal setae, outer and inner margins finely

setose, palp of two articles, distally with one short and one

longer setae. Maxilla (Fig. 14G) outer lobe of moveable endite

with two simple subdistal sickle-like setae and five distal

setulose setae, inner lobe with eight simple curved setae and

five stouter plumose setae; outer lobe of fixed endite with six

compound distal spines and subdistal biserrate spine, inner

lobe with three stout, serrate and proximally setulose setae

and rostral row of 15 setae. Maxilliped (Fig. 141) basis naked;

first palp article with one very long plumose inner seta and

small outer seta; second palp article with artefactual suggestion

of proximal articulation, longer than wide, with four proximal

inner plumose setae and rows of inner curved setae in distal

half, mostly plumose; third palp article nearly twice as long as

wide, with inner group of six simple curved setae; fourth palp

article with one subdistal and seven distal simple setae. Endites

(Fig. 14F) with two coupling hooks, outer margin densely

setulose, distally with one simple seta and numerous slender,

blunt, bifurcate spines. Epignath (Fig. 14J) oval, with setose

distal spine.

Cheliped (Fig. 15A) basis twice as long as wide, ventrally

with central sharp seta and distal pair of simple setae; three-

articled exopodite present, slender, distal article with four

plumose setae. Merus with small spine and two simple setae

on ventrodistal “shoulder”; carpus 2.4 times as long as wide,

with ventral marginal setae and single dorsodistal seta. Chela

stout, fixed finger as long as palm with six ventral setae, cutting

edge setose and with proximal tooth-like apophysis; dactylus

and claw slightly overreaching fixed finger, with fine setules

on cutting edge.

Pereopod 1 (Fig. 15B) with conspicuous setose apophysis

on coxa; basis three times as long as wide, dorsal margin

bearing four simple setae in proximal half, ventral margin

with single proximal, single mid-ventral and paired distal

simple setae; exopodite three-articled, distal article with four

plumose setae. Ischium with three simple ventrodistal setae.

Merus widening distally, with ventroproximal row of simple

setae, and single long dorsodistal and shorter ventrodistal

finely-denticulate spines and associated simple setae. Carpus

shorter than merus, with setose margins including paired

dorsodistal setae longer than propodus, one dorsodistal and

two ventrodistal finely-denticulate spines. Propodus just

shorter than carpus, with two dorsodistal and four ventral

finely-denticulate spines interspersed with single fine setae.

Dactylus slender, claw short.

Pereopods 2 and 3 (Fig. 15C, D) basis four times as long as

wide, sparsely setose, ventrodistal setae reaching to half length

of merus; ischium half as long as wide, with fine dorsal seta,

one short ventral seta and one ventral seta longer than merus;

merus shorter than carpus, with four ventral setae and single

dorsodistal seta; carpus with slender ventrodistal spine with

adjacent simple setae and long dorsodistal simple setae as long

as article; propodus longer than carpus, with two or three

slender ventral spines, dorsodistal setae as long as article;

dactylus with ventrodistal seta, dactylus and claw slender,

curved, together as long as propodus.

Pereopod 4 (Fig. 15E) similar to pereopod 3 but with

stouter basis just less than 3 times as long as wide, with

penicillate seta and fine ventrodistal simple seta as long as

ischium and merus combined; dactylus and claw less curved

than those of anterior pereopods, one of dorsodistal setae

longer than dactylus and claw combined.

Pereopod 5 (Fig. 15G) similar to pereopod 4, but basis with

three penicillate setae and three ventrodistal simple setae,

propodus with one longer seta shorter than dactylus plus claw,

and with ventral comb of 11 leaf-like spines in the distal half.

Pereopod 6 (Fig. 15F) basis with four plumose dorsal

setae; merus shorter than carpus but proportionately longer

than on pereopod 5, with two dorsal plumose setae; carpus

with one dorsal plumose seta; propodus with small leaf-like

spines ventrally and distally; dactylus plus claw slender,

curved, together as long as propodus

Pleopods typical for genus (Fig. 15H), basis with two inner

but no outer plumose setae, endopod with 12 plumose marginal

setae, exopod shorter with eight plumose marginal setae.

Uropod (Fig. 151) biramous, basis with five distal simple

setae; exopod about twice as long as basis and of four segments;

endopod elongate, filiform, with about 17 segments.

Male unknown.

Etymology. From the Latin - attenuatus : long, drawn out, thin.

Remarks. Paradoxapseudes attenuata sp. nov. is one of only

three species of the genus without plumose setae on the basis

of pereopod 1 or proximal serration on the antennal peduncle,

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 13. Paradoxapseudes attenuata sp. nov., holotype, adult female,

dorsal view. Scale = 1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 14. Paradoxapseudes attenuata sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, left mandible; D' mandible palp; E, right

mandible; F, maxillule; G, maxilla; H, labium; I, maxilliped; I', maxilliped endite; J, epignath Scale: A, B. J = 0.1 mm; C-I = 0.01 mm.

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 15. Paradoxapseudes attenuata sp. nov., female paratype. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4;

F, pereopod 5; G, pereopod 6; H, pleopod; I, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

and with only two maxillule palp setae; of the other two,

P. littoralis is distinct in having only two segments in the

accessory flagellum and four in the main flagellum of the

antennule (three and seven respectively in the other two),

while P. mortoni (Bamber, 1997) has one fewer ventral

propodus spine on pereopod 1, one fewer setae on the pleopod

basis, one more distal seta on the labial palp, fewer setae on the

proximal mandibular palp article, and ventral tubercles on the

carpus of the female cheliped. The second antennal-peduncle

article in the present species is unusual in widening at its

midpoint where the setae attach, and in only having four

ventral setae on the second article of the mandibular palp, all

other species having at least 6. P. attenuata is one of the most

elongate species in the genus, particularly in the proportions

of its cephalothorax.

Paradoxapseudes attenuata was collected throughout the

Bass Strait on sandy substrata at depths between 27 and 92 m.

Subfamily Pugiodactylinae Gufu, 1995

Genus Pugiodactylus Gufu, 1995

Pugiodactylus syntomos Blazewicz-Paszkowycz & Bamber, 2007

P. syntomos Bfazewicz-Paszkowycz & Bamber, 2007b, 131-132,

figs 14-16.

Material examined. 1 brooding ? (J57921), Slope Stn. 49, 41°56.50'S

148°37.90'E, coarse bryozoan mud, 200 m depth, 27 July 1986, coll.

M. Gomon et al., WHOI Epibenthic sled.

Remarks. P. syntomos is distinct from the other four species,

which have been described in this genus owing to its rounded

rostrum, short cheliped carpus and more compact antenna,

inter alia. It was distributed sparsely throughout the Bass

Strait at between 9 and 200 m depth (the present specimen

extending the deeper end of the range marginally), on muddy

to coarse sand substrata and once on a predominantly rocky

bottom. The genus is found from the Antarctic through

Australia (Victoria and Queensland) to Malaysia and the

South Pacific, usually in shallow waters.

Family Whiteleggiidae Gufu, 1972

Genus Whiteleggia Lang, 1972

Whiteleggia multicarinata (Whitelegge, 1901)

W. multicarinata, Lang, 1970, 605-615, figs 3-8; - Bfazewicz-

Paszkowycz & Bamber, 2007, 132 (Bass Strait material).

Additional material examined. 2 specimens, Stn. BSS 206, Eastern

Bass Strait, 19 km E of Lake Tyers Entrance, 37°50.5'S, 148°16.0'E,

26 m depth, coarse sand, 30 July 1983, coll. M. Gomon & R. Wilson,

FV Silver Gull.

Remarks. This endemic south-east-Australian species

occurred across the Bass Strait, on heterogeneous sandy

substrata from depths between 26 and 124 m. Previous records

were of the type material, off New South Wales, Australia, at

37 to 108 m depth, and further specimens in 1914 on sand and

mud in depths of 70 to 100 m off Merimbula, New South

Wales (not off South Africa - see Bfazewicz-Paszkowycz &

Bamber, 2007, p. 132).

Genus Pseudowhiteleggia Lang, 1970

Pseudowhiteleggia typica Lang, 1970

P. typica, Lang, 1970,616-626, figs 9-15; - Bfazewicz-Paszkowycz

& Bamber, 2007, 132-136 (Bass Strait material).

Remarks. Also endemic to southeast Australia, this species

occurred in the Central and Western Bass Strait at depths

between 39 and 84 m, on coarse to heterogeneous sands,

commonly sympatric with Whiteleggia multicarinata. The only

previous record was of the types at 50 m depth off northern New

South Wales, Australia.

Family Kalliapseudidae Lang, 1956

Subfamily Kalliapseudinae Gufu, 1972

Genus Kalliapseudes Stebbing, 1910

Kalliapseudes obtusifrons (Haswell, 1882)

Figures 16-21

Apseudes obstusifrons Haswell, 1882; -Kalliapseudes obtusifrons

Drumm & Heard, 2006, 29-38, figs 1-4 (redescription, literature).

Material examined. A total of 684 individuals were examined from 65

samples, including 361 females (71 with oostegites, 75 with brood

pouches), 141 males, 100 juveniles and 82 mancae. Samples were from

West of Cape Otway to Port Philip Bay (depths from 9 to 124 m). Port

Phillip Bay itself (15 to 20 m). Western Port (2 to 19 m). East of Wilson’s

Promontory (11 to 40 m), and the Gippsland coast (22 to 29 m), thus

ranging from 142.03°E to 148.7°E, and variously between 38.24°S to

40.4°S. Substrata included mud, but mainly fine to coarse sands, to

sandy gravel and shell. Numbers per sample ranged from 1 to 141

specimens. A selection of the samples examined is listed in Appendix

1. Numerous further samples held in the collections of Museum

Victoria and confirmed as this species were not examined in detail.

Remarks. Kalliapseudes obtusifrons was originally known

from Port Jackson, New South Wales (33.85°S 151.27°E), and

the species remained somewhat enigmatic until Drumm &

Heard (2006) rediscovered a syntype (designating it the

lectotype) and valuably redescribed a further specimen - an

ovigerous female collected from Cabbage Tree Island, New

South Wales (31.95°S 152.59°E) (these authors also gave an

identification key to the Australian species of Kalliapseudes ).

These had been the only known specimens of this species. It

was therefore of some surprise to find K. obtusifrons common

throughout the Bass Strait.

The large amount of material available has enabled us to

confirm the description of the female given by Drumm & Heard

(2006) and to supplement that description where their material

was damaged, to describe the dimorphism of the male, and to

examine intraspecific variation in some meristic characters

across the width of the Bass Strait.

Supplementary description of female (Fig. 16). Antennule (Fig.

17A) main flagellum with 7 to 10 segments, accessory flagellum

with 3 or 4 segments, both with three distal setae; antenna (Fig.

17B) first article with large apophysis with 4 to 6 plumose setae,

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 16. Kalliapseudes obtusifrons , adult female. A, dorsal view; B, lateral view. Scale = 1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 17. Kalliapseudes obtusifrons, female. A, antennule; B, antenna; C, labrum; D, left mandible; E, right mandible; F, maxillule; G, maxilla;

H, labium; I, maxilliped; J, maxilliped endite.

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 18 . Kalliapseudes obtusifrons.A, female cheliped; A', female cheliped, details; B, male cheliped; C, male antennule; D, epignath. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 19. Kalliapseudes obtusifrons. A, pereopod 1, female; B, details of pereopod 1, female; C, details of pereopod 1, male; D, details of pereopod

1, brooding female; E, details of pereopod 1, manca. Scale: A = 0.2 mm; B-E = 0.1 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 20. Kalliapseudes obtusifrons, female. A, pereopod 2; B, pereopod 3; C, pereopod 4; D, pereopod 5; E, pereopod 6; F, pleopod; G, uropod.

Scale = 0.1 mm.

no. no.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

-«*- ♦

44 ♦ <

►

♦

TM'

♦

ll'r 'I' ^

4 ^

- -4

iXu

142 143 144

145 146 147

Easting, degrees

148 149 150

Fig. 21. Kalliapseudes obtusifrons : graphs of A, number of ventral spines on the propodus of pereopod 1 and B, number of squama setae, both

against longitude across the Bass Strait (means and ranges where available), with linear trend-lines.

M. Blazewicz-Paszkowycz & R.N. Bamber

squama with 4 to 7 simple setae, third article with two inner

plumose setae, fourth article fused to fifth; flagellum of 6

segments, distal segment with four distal setae.

Cheliped (Fig. 18A) basis with two longer and one shorter

ventrodistal setae. Pereopod 1 (Fig. 19) basis with group of

ventrodistal simple setae, exopodite with three distal setae;

ischium with ventral seta; propodus with 4 to 6 ventral

propodal spines. Pereopod 2 (Fig. 20A) with group of long

subdistal ventral setae on basis. Pereopod 6 (Fig. 20E) basis

with 4 to 6 plumose setae on ventral margin.

Pleopod (Fig. 20F) basis with 3 to 5 plumose seta on outer

margin. Uropod endopod filiform, 2.7 times as long as

pleotelson, of about 19 segments.

Dimorphism of male. Antennule (Fig. 18C) peduncle article 3

and 4 shorter than wide, main flagellum with 8 segments,

segments 2 to 5 each bearing distal row of four or five

aesthetascs. Cheliped (Fig. 18B) basis swollen, as long as wide,

with corrugated ventrodistal margin; merus with ventral and

distal margins corrugated; carpus twice as long as wide, with

parallel ventral rows of filtering setae increasing in length

distally; propodus robust, ventral margin corrugated in distal

two-thirds, palm of chela as long as wide, with row of five

plumose filtering setae as long as width of palm. Fixed finger

about half length of palm, cutting edge with corrugated

triangular distal apophysis and smooth triangular tooth-like

proximal apophysis; dactylus cutting edge with proximal

triangular tooth-like apophysis, distally corrugated.

Note: Drumm & Heard (2006) describe a “distinct line of

fusion” on the cheliped carpus; this would be a stress line in the

cuticle related to the internal proximal attachment of the caudal

carpus muscle, as seen elsewhere in Apseudes bruneinigma

Bamber, 1998 (q.v.) and Pakistanapseudes goof Bamber &

Sheader, 2003 (Bamber & Sheader, 2003, fig. 4A). It was not

observed in any of the Bass Strait K. obtusifrons material.

Morphometric variation. As mentioned above, the large amount

of material from the Bass Strait has allowed observation of

variation in certain meristic characters. Variations were found in

the numbers of segments in the antennule flagella (the NSW

specimen of Drumm & Heard, 2006, had 10 segments in the

main flagellum, 4 in the accessory flagellum; Fig. 17A shows an

example of a main flagellum of 7, accessory flagellum of 3), in

the number of squama setae (the NSW specimen had 7, the

example in Fig. 17B shows 6), in the number of pereopod 1

propodus ventral spines (Fig. 19 shows an adult range of 4 to 6,

and a manca with 3), in the number of pereopod 1 exopodite

setae (the NSW specimen had 2, most Bass Strait specimens had

3, as in Fig. 19A), and in the number of pleopod basis setae (the

NSW specimen had 4, Bass Strait juveniles had 1 to 2, Bass

Strait adults had 3 to 4 or once 5, e.g. Fig. 20F).

When those features with sufficient variation are plotted

across the east-west range of the Bass Strait material, a distinct

trend is revealed: the number of ventral propodal spines in

adults (Fig. 21A) and the number of squama setae (Fig. 21B)

show either a decline from east to west, or a step somewhere

around 147°E (east of Wilson’s Promontory). An identical

trend is shown in the number of ventral propodal spines in

juveniles (not figured). A step change around 147°E may

represent a general separation of two populations, possibly

partially isolated by hydrographic conditions, as has been

shown in other crustaceans with intraspecific meristic

variation between populations (e.g. Henderson et al., 1990);

there would then be no reason why the New South Wales

specimens should continue this trend directly, as they would

again be a semi-isolated population. Intraspecific ranges in

meristics in other species of Kalliapseudes were discussed by

Bamber et al. (2003), who found increases in the number of

accessory flagellum segments and the number of squama setae

in Kalliapseudes makrothrix Stebbing, 1910, K. gobinae

Bamber, 1999 and K. tomiokaensis Shiino, 1966, but in relation

to size (larger in larger individuals), and based on much

smaller samples. The patterns found here for K. obtusifrons

are not size-related.

The adult female to male sex ratio of all the material

examined was 2.6:1.

Family Metapseudidae Lang, 1970

Subfamily Metapseudinae Lang, 1970

Genus Cyclopoapseudes Menzies, 1953

Subgenus Exopoapseudes subgen. nov.

Diagnosis. Cyclopoapseudes with exopodites on cheliped and

pereopod 1.

Type species. Cyclopoapseudes diceneon Gardiner, 1973.

Other species C. (E.) plumosa sp. nov.

Etymology. Combined from Exo - from “exopod”, and

“- poapseudes ” from the last part of the name of the genus

Cyclopoapseudes (female).

Remarks. With the following new species, there are now four

species of Cyclopoapseudes known. The Pacific species

C. indecorus Menzies, 1953 (the generotype), from Ecuador, and

the Indian Ocean species C. estafricana Bacescu, 1975 (Tanzania)

are both without exopodites on the cheliped and pereopod 1, while

C. diceneon Gardiner, 1973 (New Zealand) and the new species

described below from the Bass Strait, both from Antipodean

waters, have these exopodites. In other apseudomorph taxa, this

difference has been considered sufficient to distinguish separate

genera (rightly or wrongly); here we distinguish two subgenera,

the nominate Cyclopoapseudes and the presently Antipodean

Exopoapseudes , the latter being the more plesiomorphic.

Cyclopoapseudes (Exopoapseudes) plumosa sp. nov.

Figures 22-25

Material examined. 1 $ (J60994), holotype, 1 (5 (3.5 mm long)

(J60993), allotype, 2 females with empty brood pouch (3 mm long), 1

brooding 9 (3 mm long), 7 SS, 8 subadult <5S (1 dissected), 47 other

specimens (J57560), paratypes, CPBS 33S, Western Port off Crib

Point, 38°22.06'S 145°14.10’E, 13 m depth, reef with sponges, 5 March

1965, coll. AJ. Gilmour.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 22. Cyclopoapseudes (Exopoapseudes)plumosa sp. nov. A, holotype female dorsal view; B, male lateral view. Scale = 1 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 23. Cyclopoapseudes (Exopoapseud.es) plumosa sp. nov., female paratype. A, antennule; B, antenna; C, left mandible; D, right mandible D',

mandibular molar; E, maxillule; F, maxilla; G, labium; H, maxilliped; H', maxilliped endite; I, epignath Scale: A-D, G, I = 0.1 mm; E-F, H = 0.01 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 24. Cyclopoapseudes (Exopoapseudes)plumosa sp. nov. A, cheliped male; B, cheliped female. Scale = 0.1 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 25. Cyclopoapseudes (Exopoapseudes) plumosa sp. nov., female paratype. A, pereopod 1; B, pereopod 2; C, pereopod 3; D, pereopod 4;

E, pereopod 5; F, pereopod 6; G, pleopod; H, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Description of female with oostegites. Body (Fig. 22A) compact,

grossly similar to that of C. diceneon, holotype 3.7 mm long (tip

of rostrum to posterior of pleotelson), 3.5 times as long as wide,

narrower posteriorly. Cephalothorax subrectangular, wider than

long (1.5 times as wide as long without rostrum), anterior margin

with conspicuous rounded rostrum with smooth anterior margin.

Eyes present on robust eyelobes; paired dorsal plumose setae,

lateral plumose and simple setae as figured. Pereonites all with

lateral margins expanded and uniformly convex, each with

anterior row of 6 to 8 plumose setae, posterior pair of plumose

setae, and numerous lateral marginal plumose and simple setae;

pereonites 1 to 5 subequal in length (pereonite 1 just shortest),

about 0.4 times as long as cephalothorax; pereonite 6 shortest,

0.6 times as long as pereonite 2 (all pereonites respectively 3.25,

3.0, 3.0, 2.6, 2.6 and 3.8 times as wide as long). Pleon three

times as long as pereonite 2, of five free subequal pleonites

bearing pleopods; pleonites dorsally with paired low posterior

tubercles, about seven times as wide as long, with paired mid¬

dorsal plumose setae, laterally expanded by spiniform apophyses

each bearing three or four plumose setae. Pleotelson distally

with truncated protuberance, slightly longer than wide and half

as long as whole pleon, with midlateral indentation; antero-

dorsal row of four plumose setae, postero-dorsal triad of

plumose setae, laterally with seven marginal plumose setae.

Antennule (Fig. 23A). Peduncle proximal article compact,

twice as long as wide, inner margin with paired mesial and

three subdistal plumose setae, outer margin with entire row of

plumose setae; second article one-third as long as first, with

inner and distal groups of plumose setae; third article two-

thirds length of second, with outer distal seta and inner

marginal plumose setae; fourth just shorter than third, with

distal simple setae. Main flagellum of 7 segments, segments 2,

3 and 5 each bearing single aesthetasc; accessory flagellum of

4 segments.

Antenna (Fig. 23B). Proximal peduncle article with inner

distal pair of plumose setae; article 2 just longer than article 1,

inner margin bearing pair of plumose setae and sub-proximal

rounded apophysis, outer margin with three plumose setae and

subdistal squama with seven simple marginal setae; peduncle

article 3 shorter than wide, one-quarter the length of article 2,

with one simple and one plumose inner setae; article 4 twice as

long as article 3, with inner distal pairs of simple and

penicillate setae; article 5 slightly longer than article 4, with

inner and outer distal simple setae and paired inner penicillate

setae. Flagellum of seven segments.

Labrum (not figured) rounded, distally finely setulose. Left

mandible (Fig. 23D) bearing strong, pointed and crenulated

pars incisiva, lacinia mobilis slender with fine denticulations,

setiferous lobe with one stout and five finer compound setae,

pars molaris (Fig. 23D') robust, blunt, margin with row of

rounded tubercles and fine teeth; mandibular palp of three

articles, proximal article with single subdistal seta, article 2

more than half as long as whole palp with four simple distal

setae, article 3 shorter than article 1 with four subdistal and

two distal simple setae. Right mandible (Fig. 23C) as left, but

lacinia mobilis with more robust dentition, setiferous lobe

with one simple and seven compound setae. Maxillule (Fig.

23E) inner endite with five finely setulate distal setae, inner

margin finely setulose; outer endite with nine distal spines and

two subdistal setulose setae, outer margin finely setose, inner

margin with fine rows of setules. Palp of two articles, distal

article with distal row of five simple setae Maxilla (Fig. 23F)

with naked outer margin; outer lobe of moveable endite with

two simple subdistal setae and eight simple distal setae; inner

lobe of moveable endite with four simple and two setulose

setae; outer lobe of inner endite with six outer simple setae

interspersed with two bidenticulate spines, inner half with two

bidenticulate spines and three distally compound spines; inner

lobe of fixed endite with rostral row of 19 setae guarding five

longer setulose setae. Labium (Fig. 23G) with smooth outer

margin, palp with fine lateral setules and two simple distal

spines. Maxilliped (Fig. 23H) basis with finely setose outer

margin, no distal setae; palp article 1 with outer distal simple

seta and inner distal finely setulose seta; palp article 2 longer

than wide, inner margin with rows of numerous short setae,

and two simple and two setulose longer setae in proximal half,

outer margin finely denticulate with slender distal spine

reaching tip of third article; palp article 3 with 10 simple setae

along inner margin; palp article 4 with eleven setae along

broad distal margin. Endite (Fig. 23H') with paired, slender,

simple inner caudodistal setae, linguiform inner distal spines

and simple outer distal setae, setulose outer margin, two

coupling-hooks. Epignath (Fig. 231) slender, linguiform, with

distally setulose distal spine.

Cheliped (Fig. 24B) basis 2.8 times as long as wide, narrow

proximally, with short dorsodistal simple setae and four longer

ventrodistal setae; exopodite with four plumose setae. Merus

subrectangular, with four distal simple setae. Carpus 2.5 times

as long as wide, with row of long simple setae along ventral

margin. Chela stout, palm just longer than wide and fixed

finger 0.6 times as long as palm, ventral margin densely setose;

slight setose apophysis between fixed finger and articulation of

dactylus; cutting edge of fixed finger serrated, distal claw

stout; dactylus with proximal tooth-like apophysis on cutting

edge, distal claw pointed.

Pereopod 1 (Fig. 25A) basis three times as long as wide,

dorsal margin with five simple setae on a slightly convoluted

margin, ventral margin with paired proximal plumose setae, two

mid-ventral setae and four ventrodistal setae; exopodite present,

3-articled, article 3 with five distal plumose setae. Ischium with

four simple ventrodistal setae. Merus just under half as long as

basis, expanded distally, with numerous ventral simple setae,

ventrodistal spine, and curved, slender dorsodistal spine with

longer adjacent simple setae. Carpus 0.8 times as long as merus,

with two ventral spines and intervening simple setae, and single

dorsodistal spine amongst tuft of longer setae. Propodus slightly

anaxial on carpus, longer than merus, with four ventral spines

alternating with simple setae, one ventrodistal spine adjacent to

dactylus, and five dorsal spines alternating with simple setae.

Dactylus more than half as long as propodus, with mid-dorsal

fine setae, ventrodistal seta; unguis mounted subdistally, short.

Pereopod 2 (Fig. 25B) more slender but similar to pereopod

1. Coxa with plumose setae. Basis three times as long as wide

with ventral and ventrodistal plumose setae. Merus 1.1 times

as long as carpus, with ventrodistal spine but no dorsodistal

spine. Propodus with four ventral and three dorsal spines with

M. Biazewicz-Paszkowycz & R.N. Bamber

interspersed setae. Pereopod 3 (Fig. 25C) similar to pereopod

2, but basis with longer and dorsal plumose seta, merus with

two ventral spines, carpus with two dorsodistal spines,

propodus with two dorsal spines.

Pereopod 4 (Fig. 25D) basis 5 times as long as wide, naked

other than three ventrodistal plumose setae; merus with two

ventrodistal spines and inner-distal group of setae as long as

carpus; carpus twice as long as merus, distally with slender,

curved spines and simple setae; propodus as long as carpus,

articulated anaxially on carpus, widening distally to square

end, with dense group of finely denticulate shorter and longer

setae; dactylus plus unguis as long as propodus. Pereopod 5

(Fig. 25E) similar to pereopod 2, basis with ventral simple

setae; merus shorter than carpus, with two ventrodistal spines;

carpus with two ventrodistal and one dorsodistal spines;

propodus with three ventral and two dorsodistal spines.

Pereopod 6 (Fig. 25F) basis arrayed with proximal, dorsal

subdistal, and ventral plumose setae as figured; ischium with

four ventrodistal plumose setae; merus and carpus with

plumose setae along dorsal and ventral margins, carpus with

single dorsodistal spine; propodus with comb of fine leaf-like

spines along distal half of ventral margin and around dactylus,

dactylus with adjacent slender, curved spine.

Pleopods (Fig. 25G) all alike. Basis with two dorsal

plumose setae, ventral margin naked. Endopod shorter than

exopod, linguiform, with nine ventral and distal plumose

setae; exopod subovate, with seven plumose setae around

distal margin.

Uropod (Fig. 25H) biramous, exopod just shorter than

pleotelson, of six segments, endopod 3.75 times as long as

exopod, of fifteen segments.

Distinctions of male. Sexual dimorphism minimal (Fig. 22B),

cheliped (Fig. 24A) more robust, dorsal margin of basis

expanded and bearing row of 11 simple setae; merus narrow

proximally, with two distal setae; carpus only 1.7 times as long

as wide, with three mid-ventral setae; propodus as long as wide,

with dorsal marginal rows of microtrichia, chela fingers as

those of female but more robust.

Etymology, named for the plumose dorsal setae on the carapace.

Remarks. The morphology of the body of the present species is

typical for the genus, as are the conformation of the mandible

palp, and of the fourth pereopod, both being unusual features

characteristic of the genus. As commented above,

Cyclopoapseudes (Exopoapseudes) plumosa sp. nov. shares the

possession of exopodites on the cheliped and pereopod 1 only

with C. (E.) diceneon. It is distinguished from that species

particularly in the form of the pleotelson, that of the present

species being almost square in outline (although with mid-lateral

indentation), that of C. (E.) diceneon narrowing at half its length

to give a T-shape when viewed dorsally, with a conical posterior

protuberance bearing long paired distal setae. In addition, C. (E.)

plumosa has plumose setae dorsally on the carapace (that of C.

(E.) diceneon being naked), a somewhat more slender and more

setose antennule, more setose pereopods, inter alia.

Gardiner (1973) distinguished brooding females and

subadult males (with a presumed male genital cone “anlage”).

and only the latter had a cheliped with a stout propodus and a

tooth-like apophysis on the cutting edge of the dactylus. The

cheliped of the female figured here (Fig. 24B) is even more

robust than that of Gardiner’s male, but has an oostegite,

contrary to Gardiner’s (ibid.) contention that all metapseudids

are without an oostegites on the cheliped. The male cheliped

of the present species also demonstrates sexual dimorphism.

Menzies (1953) based C. indecorus on what he referred to as a

male (without specifying why), and his relatively incomplete

description shows a chela without tooth-like apophysis; with

only one specimen, no sexual dimorphism is known for this

species. Finally, Bacescu (1975) gave an even less complete

description of C. estafricana, but based on a brooding female

and a juvenile, so again no information is available on dimorphism

or hermaphroditism. This genus unfortunately is generally of

sparse occurrence.

Genus Labraxeudes Biazewicz-Paszkowycz & Bamber, 2007

Labraxeudes heliodiscus Biazewicz-Paszkowycz & Bamber,

2007

Figure 26

L. heliodiscus Biazewicz-Paszkowycz & Bamber, 2007b, 136-141,

figs 17-19.

Material examined. 6 99 (3 brooding, 3 with oostegites), 3 juveniles

(J55848), Stn WBES 1746, Western Port, 38°29.78'S 145°06.28’E, sand,

24 m depth, 25 November 1974, Smith-Mclntyre Grab; 14 99 (6 with

oostegites, 5 brooding), 2 juveniles (J56346), Stn CRUST 21,

Whaleback Rock, 0.5 km south of Point Hicks, 37°48.30’S 149°16.48'E,

ca 30 m depth; 22 99 (5 brooding, 15 with oostegites) (J57612), Stn

CPBS 33S, Crib Point, Western Port, 38°22.04'S 145°14.06’E, “reef/

sponge”, 13 m depth, 5 March 1965; 7 99 (1 brooding, 3 with oostegites)

(J57679), Stn CPBS 23N, Crib Point, Western Port, 38°20.17'S

145 0 14.11'E, sandy gravel, 10 m depth, 10 March 1965.

Remarks, the types (four females) of this species were described

from a sample off Phillip Island, at the mouth of Western Port.

Examination of the further material listed above has allowed

description of some ontogenic variation, and the correction of

some misinterpretations of the type material.

Contrary to the type description, all setae on the

cephalothorax, pereonites, pleonites and pleotelson are

plumose. The pleotelson has a rounded posterior protuberance

bearing plumose setae (Fig. 26F).

The uropods of the figured type now appear to be from a

damaged specimen: large-adult uropods (Fig. 26F) have 7

segments in the endopod and 5 segments in the exopod. The

juveniles (post-manca) have shorter uropods (Fig. 26D), but

already 7 articles in the endopod and 4 in the exopod, as do

brooding females with a simple cheliped (Fig. 26E); the

juveniles also have a simpler, semicircular rostrum (Fig. 26C).

The chelipeds show progressive development: those of

juveniles (Fig. 26A) are simple, smaller versions of those of

the smaller brooding females (Fig. 26B): these have a basis

twice as long as wide, with one proximal and two distal ventral

setae and a mid-ventral spine; exopodite as in the type; merus

with mesial and subdistal setae and two ventrodistal spines;

carpus 2.25 times as long as wide, with paired proximodorsal

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 26. Labraxeudes heliodiscus. A, cheliped of 2.53 mm brooding female; B, cheliped of 1.2 mm juvenile; C, cephalothorax of 1.2 mm juvenile;

D, pleotelson and right uropod of 1.2 mm juvenile (plumose nature of uropod setae not shown); E, left uropod of 2.53 mm brooding female

(plumose nature of all setae not shown); F, left uropod of 3.74 mm female with oostegites. Scale line = 0.5 mm.

M. Biazewicz-Paszkowycz & R.N. Bamber

setae and three ventral setae; chela palm 1.4 times as long as

wide, fixed finger with three ventral setae, dactylus with no

tooth-like apophysis on the cutting edge. The larger specimens

have the cheliped as shown for the type (Biazewicz-

Paszkowycz & Bamber, 2007b, fig. 19A), showing a

dimorphism suggestive of a male, yet these specimens also

have oostegites, and no penial tubercle was found on any

specimen. This may imply progynous, or later simultaneous,

hermaphroditism in this species, but until a specimen with a

penial tubercle is discovered, no conclusion can be drawn

about this.

All the known material of this species was collected in

Western Port except for the specimens from off Point Hicks, at

depths from between 10 and 30 m and on a range of substrata.

Genus Metapseudes Stephensen, 1927

Metapseudes wilsoni Biazewicz-Paszkowycz & Bamber, 2007

M. wilsoni Biazewicz-Paszkowycz & Bamber, 2007b, 141-146, figs

20-23.

Material examined. 1 specimen, Stn. BSS 198, Western Bass Strait,

36 km SSW of Stokes Point, King Island, 40°26.7'S, 143°41.4'E, 85 m

depth, medium sand, 22 November 1981, coll. R. Wilson, RV Tangaroa;

1 brooding ?, Stn. BSS 203, Central Bass Strait, 44 km NE of Cape

Wickham, King Island, 39°22.0’S, 144°18.3’E, 60 m depth, coarse

sand, 23 November 1981, coll. R. Wilson, RV Tangaroa. 1 9 (J57543),

Stn SA62, Flinders Island, “The Hotspot” reef, 5 n miles W of N end

of Flinders Island, South Australia, 33°40.30'S 132°22.00'E, 17 m

depth, tufted bryozoans on rock face, exposed, 19 April 1985, SCUBA,

coll. G.C.B. Poore.

Remarks. The large type collection of this species was taken in

the Eastern Bass Strait at 32 m depth; the present specimens

extend the range through the Bass Strait, as well as to South

Australia, and to a depth of 85 m. This, the second species of the

genus, is most easily distinguished from the generally similar

generotype, M. aucklandiae Stephensen, 1927, described from

New Zealand in shallow waters (Stephensen, 1927, Gardiner,

1973) (depth range 0-113 m), by the more slender antennules

and antennae, and the distinct form of the rostrum, inter alia.

Family Parapseudidae Gufu, 1981

Subfamily Pakistanapseudinae Gufu, 2008 new rank

Remarks. Gu(u (2008b) separated the family Parapseudidae into

two groups, separating the species of the Pakistanapseudes-

group discussed by Biazewicz-Paszkowycz & Bamber (2007a)

from the remaining parapseudids. Gufu (ibid.) erected two tribes

(Parapseudini and Pakistanapseudini), but it is entirely more

appropriate to consider these as subfamilies, so we have given

them that new rank herein.

Genus Pakistanapseudes Bacescu, 1978

Remarks, in describing his new genus, Bacescu (1978) did not

designate a type-species. By inference, and as stated by Bamber

& Sheader (2003), it should be Pakistanapseudes leptochelatus

Bacescu 1978, herein so designated (not Pakistanapseudes

leptodactylus Bacescu 1978 as cited by Gufu, 2008, a lapsus

calami ). It should be noted that members of this genus have a

great propensity for autotomizing their appendages on fixation

unless relaxed first, so in many species not all of the pereopods,

chelipeds, antennules or uropods are known. In both of the new

species described below, chelipeds and first pereonites were so

rare that no complete exopodite was found.

Pakistanapseudes bassi Biazewicz-Paszkowycz & Bamber, 2007

P. bassi Biazewicz-Paszkowycz & Bamber, 2007a, 14-19, figs 7-9.

Remarks. This species was described originally from numerous

specimens collected throughout the Bass Strait, on sandy

substrata from depths between 60 and 293 m. Numerous further

samples of this species exist in the collections of Museum

Victoria, including some from water as shallow as 2 m (Port

Phillip Bay), over 100 specimens having been examined in the

course of this study in addition to the type-collection, and it

appears clearly to be the commonest Pakistanapseudes species

in the Bass Strait region. It is morphologically similar to

P. perulpa (rounded rostrum, no bifurcate claws), but unlike that

species it has no ventral setae on the pleopod basis, and pereonites

5 and 6 are wide than long (longer than wide in P. perulpa ).

Pakistanapseudes lucifer sp. nov.

Figures 27-29

Material examined. 1 ovigerous 9 (J28617), holotype, Stn MSL-EG

117, Eastern Bass Strait, 37°52.65'S 148°42.15’E, 49 m depth, February

1991, coarse sand; 4 99 (J28627), paratypes, Stn MSL-EG 78, Eastern

Bass Strait, 37°43.89'S 148°30.13’E, 27 m depth, 4 June 1991, coarse

sand; 1 brooding 9, 2 99 with oostegites (J28611), paratypes, Stn MSL-

EG 99, Eastern Bass Strait, 37°53.29’S 148°15.40'E, 43 m depth,

February 1991, coarse sand; 1 3 (J28630), paratype, Stn MSL-EG 108,

Eastern Bass Strait, 37°53.14’S 148°28.94’E, 45 m depth, February

1991, medium sand; 1 $ (J28627), paratype, Stn MSL-EG 72, Eastern

Bass Strait, 37°53.39’S 148°15.40’E, 43 m depth, 4 June 1991, coarse

sand; 3 33, 1 brooding 9 (J28629), paratypes, Stn MSL-EG 104,

Eastern Bass Strait, 37°49.89’S 148°30.13'E, 27 m depth, February

1991, coarse sand; 2 99 (J57574), paratypes, Stn BSS170, Eastern Bass

Strait, 39°51.8'S 148°26.5'E, 130 m depth, 15 November 1981, fine

sand, coll. R.S. Wilson; 1 3 (J57574), paratype, Stn BSS169, Eastern

Bass Strait, 39°02.4'S 148°30.6'E, 120 m depth, 15 November 1981,

muddy sand, coll. R.S. Wilson; 1 $ (J57574), paratype, Stn BSS188,

Western Bass Strait, 38°38.2’S 142°35.0’E, 59 m depth, 20 November

1981, coll. R.S. Wilson; 1 9 (J51790), paratype, Stn VC 27 Cl, Western

Bass Strait, 38°23.92'S 145°18.43’E, 40 m depth, 11 May 1998, fine

sand; 2 99 (J51317), paratypes, Stn VC 18 C3, Western Bass Strait,

38°30.2'S 144°15.00E, 40 m depth, 13 May 1998; 2 brooding 99

(J57655), 1 $ with oostegites (J57646), 1 brooding $ (J57653), 1 3

(J57654), paratypes, Stn CPBS 41N, Western Port, 38°20.81'S

145°13.85’E, 13 m depth, 30March 1965, gravel and sand; 1 9 (J55925),

12 March 1965, 1 9 (J55902), 20 March 1967, 2 33 and 4 99 (1 with

oostegites, 2 brooding) (J55885), paratypes, Stn CPBS 32S, Western

Port, 38°22.06'S 145°14.10'E, 13 m depth, reef with sponge.

Description of female. Body (Fig. 27A), dorsoventrally flattened,

elongate, holotype 3.2 mm long (tip of rostrum to posterior of

pleotelson), six times as long as wide, tapering towards

posterior. Cephalothorax subrectangular, just longer than wide,

with pronounced pointed rostrum curving downward, laterally

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 27. Pakistanapseudes lucifer sp. nov. A, holotype ovigerous female, dorsal. B, male, dorsal; C, cephalothorax of female, lateral; D, male

antennule; E, male pleopod; F, female pleopod. Scale = 0.1 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 28. Pakistanapseudes lucifer sp. nov., female paratype. A, antennule; B, antenna; C, left mandible; D, right mandible; E, mandibular molar; E',

mandibular palp; F, maxillule;.; F', maxillule palp; G, maxilla; H, labium; I, maxilliped; J, maxilliped endite. Scale A, B = 0.1 mm; C -1 = 0.01 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 29. Pakistanapseudes lucifer sp. nov., female paratype. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 4; E, pereopod 5; F, pereopod

6; G, uropod. Scale = 0.1 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

indented at anterior of branchial chambers; eyelobes distinct,

dark ocelli present. Conspicuous forward-pointing spine-like

hyposphenium mid-ventrally between chelipeds (Fig. 27C).

Each pereonite laterally convex, with anterolateral seta

(posterolateral seta on pereonite 1); pereonites 1 and 2 subequal,

shortest, about 0.35 times as long as cephalothorax; pereonite 3

one-and-a-half times length of pereonite 1; pereonite 4 longest,

nearly twice as long as pereonite 1, pereonite 5 just shorter than

pereonite 4, pereonite 6 just shorter than pereonite 3 and

narrowest (all pereonites respectively 2.7, 2.7, 1.8, 1.3, 1.3 and

1.3 times as wide as long). Pleon three times as long as pereonite

5, with five free pleonites bearing pleopods; pleonites with

single midlateral seta on each side; pleonite 1- 0.8 times as long

as wide, posterior pleonites progressively shorter. Pleotelson

rectangular, wider posteriorly, 0.3 times length of pleon, 1.4

times as long as wide, with paired lateral setae.

Antennule (Fig. 28A) proximal peduncle article 2.7 times

as long as wide, with inner and outer subdistal tufts of setae and

midlateral outer group of three penicillate setae and single

simple seta. Article 2 about 1.4 times as long as wide, 0.34

times length of first, with inner and outer distal setae exceeding

distal edge of third article, single outer distal penicillate seta.

Article 3 just under half-length of article 2, as long as wide.

Peduncle article 4 half-length of article 3, wider than long.

Main flagellum sparsely setose, of twelve segments, three

aesthetascs present on segment 4, four on segment 6, one on

each of segments 8 and 10; accessory flagellum of five segments.

Antenna (Fig. 28B) with naked proximal peduncle article

(not figured) expanded on inner margin. Article 2 longer than

first, with two simple setae adjacent to elongate squama

bearing seven inner marginal and distal setae. Peduncle article

3 shorter than wide with inner seta. Article 4 half as long as

second with inner penicillate setae; article 5 twice as long as

article 4. Flagellum of six segments, first and second segments

with setae longer than three flagellar segments.

Labrum rounded, simple, naked; epistome not obvious.

Right mandible (Fig. 28D) with five cusps on pars incisiva;

setiferous lobe with three trifurcate, two bifurcate and one

simple setae, pars molaris (Fig. 28E) stout, blunt with distal

rugosity and denticulate margin; palp (Fig. 28E') of three

articles, proximal article with one distal seta, second article

twice as long as first and naked, third article just longer than

second and with two longer stout distal setae and one shorter

subdistal seta. Left mandible (Fig. 28C) as right but with

dentate lacinia mobilis; outer margin finely denticulate.

Labium with outer serrations, distally finely setose, palp (Fig.

28H) with inner and outer fine lateral setules and two longer

and two shorter distal spines, and conspicuous rounded inner

apophysis. Maxillule (Fig. 28F) inner endite with outer

apophysis and finely-setose distal margin, and five plumose

distal setae; outer endite with ten distal spines and three

distally denticulate subdistal setae, outer margin finely setose;

palp of two articles, distally with three setae graduated in

length. Maxilla (Fig. 28G) typical of genus, outer margin

finely setose; moveable endite outer lobe with two subdistal

and five distal finely denticulate setae, inner lobe with six

plumose/denticulate setae; fixed endite outer lobe with simple,

trifurcate and bilaterally denticulate distal spines, inner lobe

with five longer plumose setae and rostral row of 24 setae.

Maxilliped (Fig. 281) with simple setae; first palp article with

short outer seta and long inner distal seta; second palp article

with inner margin bearing proximal serrations and numerous

setae largely in two rows, longest inner seta not reaching distal

margin of article, single outer distal setae; third palp article

with nine recurved inner setae; fourth palp article with eleven

setae around distal margin. Endite (Fig. 28J) distal margin

with outer simple setae and inner blunt compound spines, two

coupling hooks. Epignath not recovered.

Cheliped (Fig. 29A) slender. Basis four times as long as

wide, ventrally without spine but with single ventrodistal seta.

Exopodite damaged. Merus with single ventrodistal seta.

Carpus slender, four times as long as wide, naked. Chela not

slender, palm (propodus) 1.6 times as long as wide with single

inner and outer distal setae at base of dactylus and small

dorsodistal seta; ventral margin of fixed finger with two setae;

cutting edge of fixed finger without apophyses but with three

setae. Dactylus shorter than palm, naked, with no apophyses

on cutting edge.

Pereopod 1 (Fig. 29B) generally bearing simple (not

tapering) longer setae. Basis stout, 1.4 times as long as wide,

with small ventroproximal and mid-ventral spine and larger

ventrodistal spine, dorsal margin with four simple, fine setae.

Exopodite damaged. Ischium with two shorter and one longer

ventrodistal setae. Merus more than half length of basis, wider

distally, with single dorsodistal seta and curved dorsodistal

spine, row of mid-ventral setae, stout ventrodistal spine

without adjacent setae. Carpus compact, as long as merus, 1.2

times as long as wide, with two short ventral and one longer

dorsodistal blunt spines, sparse ventral and dorsodistal setae

as figured. Propodus shorter than carpus, with four ventral

blunt spines increasing in length towards distal margin

interspersed with single, simple setae, two dorsodistal slender

blunt spines and few setae along dorsal margin. Dactylus stout,

with two ventral denticulations and two dorsal setae. Unguis

distinct, pointed.

Pereopods 2 (Fig. 29C) and 3 similar to each other. Basis

four times as long as wide, with two dorsoproximal penicillate

setae and two shorter and one elongate ventrodistal setae,

longest seta exceeding tip of merus. Ischium as long as wide

with one ventrodistal seta; merus short, 0.2 times as long as

basis, much shorter than carpus, with three ventral setae, dorsal

margin naked. Carpus elongate, three times as long as merus,

with inner and ventral rows of setae, single dorsodistal seta.

Propodus 0.8 times as long as carpus, with row of elongate,

simple ventral setae, paired mesial setae, dorsoproximal

penicillate seta and three dorsodistal setae. Dactylus slender, as

long as propodus, with subdistal unguis forming bifurcate tip.

Pereopod 4 (Fig. 29D) slender, similar to pereopod 2, basis

4.7 times as long as wide, without ventrodistal setae. Carpus 3.2

times as long as merus. Propodus with dorsodistal group of one

short spine, and one longer slender spine and two setae all as long

as dactylus; dactylus elongate, 0.8 times as long as propodus,

bifurcate as pereopod 2. Pereopod 5 (Fig. 29E) similar to

pereopod 4 but propodus with ventral row of fine spinules and

three dorsodistal setae; dactylus with subdistal unguis forming

bifurcate tip. Pereopod 6 (Fig. 29F) similar to pereopod 5 but

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

propodus ventral and distal margin with row of some 19 small

leaf-like spinules, dorsodistal group of four spines and single

seta. Dactylus with subdistal unguis forming bifurcate tip.

Pleopods (Fig. 27F) all alike. Basis naked; rami linguiform.

Endopod longer than exopod, respectively with 12 and 10

marginal plumose setae.

Uropod (Fig. 29G) biramous, basis with single inner and

outer distal setae; exopod of nine segments; endopod over four

times as long as exopod, 0.6 times as long as body length, of

about 35 segments.

Description of male, similar to female (figured male 3.7 mm

long, Fig. 27B), cephalothorax proportionately slightly larger,

rostrum more pronounced. Antennule and antenna (Fig. 27D)

with multisegmented flagella bearing dense aesthetascs;

squama of antenna fused to second peduncle article, with small

distal spinule, fourth peduncle article short, fifth article with

two simple distal setae. Cheliped not recovered. Pleopods with

more elongate basis, with articulation (Fig. 27E).

Etymology, named after the Devil, owing to its having bifurcated

claws (“cloven hooves”) on all pereopods other than pereopod 1.

Remarks, bifurcation of the pereopod claws (used in this context

to mean the combination of dactylus and unguis, whether fused

or not) in Pakistanapseudes is a variable feature, ranging from

no bifurcation (e.g. P. leptochelatus, P. bassi ), to a bifurcation

resulting from the subdistal attachment of the unguis on the

dactylus on pereopods 2 and 3, as, for example, in P. goofi

Bamber & Sheader, 2003, and finally to a subdistal fusion of the

unguis to the dactylus on pereopods 2 and 3 as, for example, in

P. tenuicorporeus (Shiino, 1963); apart from the present species,

the only other species of the Pakistanapseudinae to have

bifurcating claws (with an unfused dactylus) on pereopods 5 and

6 as well as 2 and 3 is Swireapseudes birdi Gufu & Iliffe, 2008,

but in that species the claw of pereopod 4 is not bifurcate, whereas

the claw of pereopod 4 in P. lucifer sp. nov. is bifurcate, identical

to those of the other pereopods.

Other distinctions of P. lucifer include the complete absence

of setae on the basis of the pleopod: many other species lack

outer (ventral) setae, but all appear to have inner (dorsal) setae,

although the setation of the basis in P. leptochelatus is unclear.

In addition, the basis of pereopod 1 is surprisingly short

compared with other species, and all other species show some

modification of the dactylus and unguis of pereopod 4 (where

known), normally a marked reduction in size compared with the

other pereopods (although it is larger in P. perulpa Bfazewicz-

Paszkowycz & Bamber, 2007 and P. ridculli Bamber, 2005): in

the present species, it shows no difference.

In the very sparse setation of the mandibular palp, only P.

brasiliensis Gufu 1996, from Brazil, approaches P. lucifer, but

that species has a naked proximal article and eight distal/subdistal

setae on the distal article. The sparsity of setation on the cheliped

and pereopods is more extreme than any other pakistanapseudid,

while the presence of a ventral row of fine spinules on the

propodus of pereopod 5 is unique in this group, and the

presence of three subdistal setae on the outer endite of the

maxillule is unknown in the Apseudomorpha, as far as we are

aware, all other species having two when they are present.

With regard to the identification key to Australian

Pakistanapseudes species given by Bfazewicz-Paszkowycz &

Bamber (2007), the only other Australian species with a pointed

rostrum is P. australianus Gufu, 2006, from Queensland, but

that species has two setae on the pleopod basis, far more

segments in the antennular and antennal flagella, and is without

bifurcated claws on pereopods 2,4, 5 and 6 (pereopod 3 has not

been described).

Pakistanapseudes lucifer occurred throughout the Bass

Strait, at depths from 13 to 130 m on sandy substrata.

Pakistanapseudes perulpa Bfazewicz-Paszkowycz & Bamber,

2007

P. perulpa Bfazewicz-Paszkowycz & Bamber, 2007a, 3-8, figs 1-3.

Material examined. 1 9 with oostegites, 1 subadult (J30441), La Trobe

Valley Ocean Outfall Survey Stn MSL LV 4 T5, Eastern Bass Strait, 1

km off Delray Beach, Victoria, 38°14'S 147°22'E, 15-16 m depth, 24

October 1989; 2 99 (1 with oostegites) (J30444), La Trobe Valley

Ocean Outfall Survey Stn MSL LV 6 T8, Eastern Bass Strait, 1 km off

Delray Beach, Victoria, 38°14’S 147°22'E, 15-16 m depth, 29 August

1990; 4 $9 (2 with oostegites) (J30405), La Trobe Valley Ocean Outfall

Survey Stn MSL LV 6 SI, Eastern Bass Strait, 1 km off The

Honeysuckles, Victoria, 38°22'S 147°12'E, 15-16 m depth, 28 August

1990; 6 99 (5 with oostegites) (J30409), La Trobe Valley Ocean Outfall

Survey Stn MSL LV 6 S4, Eastern Bass Strait, 1 km off The

Honeysuckles, Victoria, 38°22'S 147°12'E, 15-16 m depth, 28 August

1990, SCUBA Airlift, coll. EPA & Marine Science Laboratory.

Remarks : this species was described originally from Moreton

Bay, Queensland, from clean sandy substrata at depths between

7 and 28 m. The present material is within this depth range, but

extends the distribution much further south. It has a rounded

rostrum, setae on both margins of the pleopod basis, an overlong

dactylus to pereopod 4, and is without bifurcating claws.

Pakistanapseudes taylorae sp. nov.

Figures 30-32

Material examined. 1 $ with oostegites (J57588), holotype, Stn BSS76,

Western Bass Strait, 39°19'S 143°38'E, 95 m depth, 10 October 1980,

coarse sand, carbonate, coll. G C B Poore; 1 juvenile without appendages

(J57596), paratype, Stn BSS112, Central Bass Strait, 40°22.2'S 145°17'E,

40 m depth, 3 November 1980, mainly sand, coll. M.F. Gomon and G C

B Poore; 3 SS, 3 9$ (J55840), paratypes, Stn BSS180, Central Bass

Strait, 39°12.9'S 146°27.3'E, 65 m depth, 18 November 1981, medium

sand, coll. R.S. Wilson; 1 brooding 9 (J55865), paratype, Stn CPBS 32N,

Western Port, 38°20.83'S 145°13.49'E, 13 m depth, 21 February 1969,

sandy gravel. 1 9 (not registered), Stn BSS209, Eastern Bass Strait,

38°18.0'S 147°37.0'E, 55 m depth, 31 July 1983, muddy with fine shell,

coll. M. Gomon & R.S. Wilson.

Description of female. Body (Fig. 30A), dorsoventrally

flattened, elongate, holotype 4 mm long (tip of rostrum to

posterior of pleotelson), 6.5 times as long as wide, tapering

towards posterior. Cephalothorax subrectangular, just wider

than long, with pronounced apparently rounded rostrum but

with small distal point (Fig. 31C), single lateral setae in front of

branchial chamber on each side; eyelobes distinct, dark ocelli

present. Forward or backward pointing hyposphenium mid-

ventrally between chelipeds. Pereonite 1 naked, pereonites 2

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 30. Pakistanapseudes taylorae sp. nov., holotype female. A, dorsal view; B, pleopod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 31 . Pakistanapseudes taylorae sp. nov., female paratype. A, antennule; B, antenna; C, rostrum; D, labrum; E, left mandible; F, right mandible;

F' mandible palp; G, maxillule; H, maxilla; I, labium; J, maxilliped; J', maxilliped endite; K, epignath. Scale A, B, D = 0.1 mm; C, E-K= 0.01 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 32. Pakistanapseudes taylorae sp. nov. A, antennule, male; B, cheliped (fragment), female; C-H, pereopods 1-6 respectively, female.

Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

and 3 with anterolateral seta, pereonites 4 to 6 with anterolateral

seta and posterolateral seta; pereonite 1 shortest, about 0.4

times as long as cephalothorax, pereonites 2 and 3 subequal,

half length of cephalothorax; pereonite 4 longest, 1.3 times as

long as pereonite 1, pereonite 5 just shorter than pereonite 4,

pereonite 6 just longer than pereonite 1 (all pereonites

respectively 2.6, 1.8, 1.7, 1.1, 1.2 and 1.7 times as wide as long);

pereonites 1 and 2 with or without hyposphenia. Pleon three

times as long as pereonite 4, with five free pleonites bearing

pleopods; pleonite 1- 0.4 times as long as wide, posterior

pleonites progressively shorter, each with four or five lateral

setae. Pleotelson rectangular, one-third length of pleon, twice

as long as wide, with paired lateral setae.

Antennule (Fig. 31A) proximal peduncle article 3.2 times

as long as wide, outer margin with few shorter simple setae,

inner margin with numerous penicillate setae and more sparse

longer simple setae, longest seta reaching distal edge of second

peduncle article. Second article 1.5 times as long as wide, half

length of first, with distal crown of simple and penicillate

setae. Article 3 about 0.2 times length of article 2, half as long

as wide, with long distal setae all round. Peduncle article 4 as

long as article 3, wider than long. Main flagellum regularly

setose, of eight segments, single aesthetascs present on

segments 6 and 8; accessory flagellum of five segments.

Antenna (Fig. 31B) with naked proximal peduncle article

expanded on inner margin. Article 2 just longer than first, with

one simple seta adjacent to elongate squama bearing two distal

setae. Peduncle article 3 shorter than wide with inner spine-like

apophysis. Article 4 one-and-a-half times as long as second

with crown of penicillate setae; article 5 just longer than article

4, with single distal seta. Flagellum of four segments.

Labrum (Fig. 31D) truncate, simple, marginally setose.

Pointed epistome obvious. Right mandible (Fig. 31F) with five

rounded “teeth” on pars incisiva; setiferous lobe with one

bifurcate and three trifurcate setae, pars molaris (Fig. 31F")

stout, blunt with crenulate distal margin; palp (Fig. 31F') of

three articles, proximal article with one inner seta; second

article nearly three times as long as first, naked; third article as

long as second, with two longer and two shorter simple distal

setae. Left mandible (Fig. 31E) as right but with dentate lacinia

mobilis and five setae on setiferous lobe. Labium (Fig. 311)

without serrations, not setose, palp with inner and outer fine

lateral setules and two longer and one minute simple distal

setae, and widely rounded inner apophysis. Maxillule (Fig.

31G) inner endite with outer finely-setose margin distal of

apophysis, and four plumose and one simple distal setae; outer

endite with ten distal spines and three subdistal setae, outer

margin finely setose; palp of two articles. Maxilla (Fig. 31H)

typical of genus, outer margin naked, outer lobe of moveable

endite with two subdistal and seven distal finely denticulate

setae, inner lobe with nine plumose/denticulate setae; fixed

endite outer lobe with simple, trifurcate, plumose and

bilaterally denticulate distal spines, inner lobe with two longer

plumose setae and rostral row of 13 setae. Maxilliped (Fig.

31J) mostly with simple setae; first palp article with short outer

distal seta and longer inner distal seta almost reaching tip of

article 2; second palp article with outer distal spine, inner

margin bearing 16 setae largely in two rows, longest inner seta

reaching fourth article, inner proximal margin with about

three thorn-like apophyses; third palp article with five recurved

inner setae; fourth palp article with seven setae around distal

margin, inner five with finely denticulate inner margins. Endite

(Fig. 31J') distal margin with outer simple setae and gradation

of inner blunt spines, caudal seta leaf-like. Epignath (Fig. 31K)

large, cup-shaped, with large, setose proximal lobe and distally

setulose distal spine.

Cheliped (Fig. 32B) only one available on one specimen,

damaged; basis slender, 2.6 times as long as wide, ventrally

with small proximal and longer distal setae, without spine.

Exopodite damaged. Merus with three ventrodistal setae.

Carpus slender, 3.3 times as long as wide, with one mid-ventral

and two ventrodistal setae. Chela badly damaged, fixed finger

with 3 ventral setae, and tuft of distal curled setae, and row of

spatulate setae on cutting edge.

Pereopod 1 (Fig. 32C) basis three times as long as wide,

with small ventroproximal and mid-ventral spine and seta

between these two, longer, pointed ventrodistal spine with

adjacent seta exceeding distal margin of ischium, dorsal

margin with two proximal simple setae and two mid-dorsal

penicillate setae. Exopodite damaged. Ischium with three

ventrodistal setae. Merus half as long as basis, wider distally,

with slender dorsodistal spine and long dorsodistal seta as

long as carpus, row of five ventral setae and ventrodistal spine.

Carpus just shorter than merus, with three ventral spines

interspersed with three setae, and one elongate dorsodistal

spine with adjacent row of four setae. Propodus with five

ventral spines increasing in length towards distal margin

interspersed with single simple setae, two dorsodistal slender

blunt spines and row of setae along dorsal margin. Dactylus

stout, with three ventral denticulations, two small mid-dorsal

setae. Unguis distinct, pointed.

Pereopod 2 (Fig. 32D) basis 2.7 times as long as wide, with

single mid-ventral spine and three elongate ventrodistal setae.

Ischium as long as wide with row of three ventrodistal setae;

merus shorter than carpus, with single long, slender dorsodistal

and ventrodistal spines. Ventral margin with row of six setae,

one ventrodistal seta. Carpus 1.5 times as long as merus, with

row of three slender spines interspersed with single setae

along ventral margin, and diagonal row across inner face of

two spines and six setae, one longer, slender dorsodistal spine;

propodus just longer than carpus, ventrodistal margin with

four slender spines interspersed with single simple setae,

dorsally with three slender spines and five setae in distal half.

Dactylus slender, together with distinct unguis 0.8 times as

long as propodus, not bifurcate. All spines with fine

denticulation in distal half.

Pereopod 3 (Fig. 32E) similar to pereopod 2, but basis

more slender, merus with only one ventral spine and without

dorsal spine, carpus and propodus with fewer spines and setae,

dactylus 1.24 times as long as propodus, very small unguis

subdistal, giving bifurcation.

Pereopod 4 (Fig. 32F) basis 4.3 times as long as wide, with

ventrodistal and dorsoproximal penicillate setae and single,

long ventrodistal simple seta exceeding tip of merus. Ischium

with single ventrodistal seta. Merus with three ventral setae.

Carpus 1.5 times as long as merus, ventrally with four slender,

M. Biazewicz-Paszkowycz & R.N. Bamber

blunt, finely denticulate spines interspersed with nine setae.

Propodus with distal crown of ten setae, single dorsal

penicillate seta in proximal half; dactylus elongate, sinuous,

pointed, 1.6 times as long as propodus.

Pereopod 5 (Fig. 32B) similar to pereopod 4 but basis

without ventrodistal seta, ventral setae on ischium and merus

shorter, unguis attached subdistally, shorter than extension of

dactylus, giving bifurcation.

Pereopod 6 (Fig. 32H) basis naked, ischium with two

ventrodistal setae and single smaller dorsodistal seta. Carpus

with eight setae and five denticulate spines ventrally. Propodus

ventral and distal margin with row of some 26 small leaf-like

spines, ventrally with slender subdistal spine, dorsodistal group

of two setae. Dactylus as long as propodus, unguis attached

subdistally, shorter than extension of dactylus, giving bifurcation.

Pleopods (Fig. 30B) all alike. Basis with two outer plumose

setae and three inner plumose setae; rami linguiform. Endopod

longer than exopod, respectively with 17 and 13 marginal

plumose setae.

Uropods missing on all specimens.

Description of male. Generally similar to female, available

specimens missing most appendages (cheliped unknown);

antennule missing, antenna (Fig. 32A) with multi segmented

flagellum with array of aesthetascs over whole surface; squama

with three distal setae.

Etymology, named after Dr Joanne Taylor, Collection Manager

at Museum Victoria, in gratitude for all her diligent efforts and

her tolerance of our interference with her collections.

Remarks. Pakistanapseudes taylorae sp. nov. is the seventh

species of this genus recorded from Australian waters (see

Biazewicz-Paszkowycz & Bamber, 2007, and above). The only

other species to have a pointed rostrum and bifurcate claws is

P. lucifer (described above); the present species is distinguished

from P. lucifer by having basal setae on the pleopod, a very short

third peduncle article on the antennule, a non-bifurcate claw on

pereopods 2 and 4, far more and longer setae and spines on the

cheliped and pereopods, fewer segments in the flagellum of the

antenna and the main flagellum of the antennule, fewer setae on

the distal maxilliped palp article, and the thorn-like apophyses

on the second palp article, inter alia. Interestingly, both of these

species have the unusual attribute of three subdistal setae on the

outer endite of the maxillule.

Elsewhere, the only other species of the Pakistanapseudinae

with a pointed (if only slightly) rostrum, eyes, and bifurcate

pereopod claws is Swireapseudes birdi from the Bahamas, but

that species has far more segments in the antennular and

antennal flagella, far fewer spines on its much more slender

pereopod 1, a proportionately longer third peduncle article on

the antennule, and distinct setation of its mouthparts.

P. taylorae 'vs particularly unusual in the Pakistanapseudinae

in the reduced setation of the antennal squama: only the deep-

sea species Leptolicoa thokozele (Bamber & Sheader, 2003),

very distinct from the present species in a number of features,

has as few as two squama setae.

P. taylorae was collected sparsely throughout Bass Strait, on

coarse to medium sands, and in depths between 13 and 95 m.

Pakistanapseudes C sp. nov.

Material examined. 1 damaged female (J58891), Stn MSL EG 49,

Eastern Bass Strait, 15.1 km WSW OF Pt Ricardo, 37°51.38'S

148°28.14'E, 34 m depth, 26 September 1990, Smith-Mclntyre Grab.

Remarks. The body of this taxon is very elongate, much more

so than, and thus quite different from, all the other described

Australian species of Pakistanapseudes. Unfortunately, the

single specimen is in very poor condition, with the only

appendages being the cheliped and pereopods 1 and 2, and the

pleon in particular is substantially damaged, so it does not

warrant a proper description, nor naming, despite its being

clearly a distinct and new species.

Subfamily Parapseudinae Gufu, 1981 new rank

Genus Parapseudes Sars, 1882

Parapseudes blandowskii sp. nov.

Figures 33-36

Material examined. 1 brooding 9, holotype (J24152), Stn CRUST 153,

Cappers Camp, west end of Nelson Bay, off rock platform, 38°24'S

141°34'E, 5 m depth, 29 February 1992, coll. B.F. Cohen & R.S. Wilson,

SCUBA airlift. 4 (5(5, 22 99 (10 brooding, 7 with oostegites), 5 subadults

(J57615), 1 $ (J57795), 1 6' (J58574), paratypes, Stn CPBS 41N, Western

Port, 38°20.81'S 145°13.85'E, 13 m depth, 30 March 1965, sandy gravel;

2 6(5, 8 99 (6 brooding) (J57634), paratypes, Stn CPBS 23S, Western

Port, 38°21.69'S 145 0 13.5PE, 11 m depth, 9 March 1965, muddy sand; 1

subadult (J57630), paratype, Stn CPBS 23N, Western Port, 38°20.29'S

145°14.18'E, 10 m depth, 10 March 1965, sandy gravel; 33 specimens

(including 6(5 and brooding 9$) (J57622), paratypes, Stn CPBS 33S,

Western Port, 38°22.06'S 145°14.10'E, 13 m depth, 5 March 1965, reef

with sponges; 3 99 (2 brooding, 1 with oostegites) (J56359), paratypes,

Stn CPBS 41N, Western Port, 38°20.81'S 145°13.85'E, 13 m depth, 30

March 1965, sandy gravel; 1 9 (J56199), paratype, Stn CPBS 32N,

Western Port, 38 0 19.71'S 145°13.82'E, 14 m depth, 25 Agust 1966, sand;

2 (56', 4 99 (1 brooding) (J57627), paratypes, Stn CPBS 33S, Western

Port, 38°22.06’S 145°14.10'E, 13 m depth, 5 March 1965, reef with

sponges; 32 specimens (including (5(5 and brooding 99) (J57645),

paratypes, Stn CPBS 41N, Western Port, 38°20.81'S 145°13.85'E, 13 m

depth, 30 March 1965, sandy gravel, coll. A.J. Gilmour; 4 99 (3

brooding), 1 subadult (J56363), paratypes. Red Rock Point Island,

sublittoral, 23 July 1974; 1 (5, 1 9 (J56361), paratypes. Cruise 81-T-l Stn

BSS 185, Western Bass Strait, 38°48.0'S 143°14.5’E, 47 m depth, 20

November 1981, rocky bottom, coll, R. Wilson; 1 (5, 1 brooding 9

(J56291), paratypes. Western Port sublittoral, 25 November 1971, coll J

E Verse, H F Seed.

Description of female. Body (Fig. 33) typical of the genus,

holotype 4.0 mm long (tip of rostrum to posterior of pleotelson),

five times as long as wide, narrower posteriorly. Cephalothorax

pentangular, as long as wide including rostrum, anterior margin

produced into convex, triangular rostrum with smooth anterior

margin; lateral indentation anterior to branchial chambers. Eyes

present on rounded eyelobes. All pereonites with lateral margins

uniformly convex, appearing as posterolateral rounded

apophyses on pereonites 3 to 6 owing to anterior (3 and 4) or

posterior (5 and 6) lateral indentations, each with two to four

conspicuous simple lateral setae; pereonite 1 shortest, about one-

quarter as long as cephalothorax; pereonites 2 to 5 progressively

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

longer, pereonite 6 as long as pereonite 3 (all pereonites

respectively 3.5, 2.8, 1.7, 1.6, 1.5 and 1.7 times as wide as long).

Pleon twice as long as pereonite 5, of five free subequal pleonites,

the first four only bearing pleopods, and rectangular pleotelson;

pleonites more than four times as wide as long, laterally expanded

by spiniform apophyses each bearing two or three simple setae

distally, pleonite 1 with dorsal row of setae. Pleotelson distally

extended and rounded, half as long as whole pleon, 1.3 times as

wide as long, bearing lateral and dorsal simple setae.

Antennule (Fig. 34A). Peduncle proximal article compact,

widest at mid-length, 2.3 times as long as wide, inner margin

with row of four simple setae just distal of mid-length and

subdistal seta, outer margin with penicillate setae in proximal

third and four simple setae in distal two-thirds; second article

one-third as long as article 1, 1.5 times as long as wide, with

outer and inner subdistal tufts of three simple setae; third

article 0.4 times as long as second and wider than long; fourth

article just shorter than third, naked. Main flagellum of 10

segments, segments 6 and 8 each bearing single aesthetasc;

accessory flagellum of 4 segments.

Antenna (Fig. 34B). Proximal peduncle article with outer

rounded apophysis, naked; article 2 0.8 times as long as article

1, outer margin with one simple marginal seta, linguiform

squama with six simple setae around distal margin; peduncle

article 3 shorter than wide, one-third as long as article 2, with

one distal seta; article 4 as long as article 1, with one distal

seta; article 5 one-third as long as article 4, with one distal

seta. Flagellum of five segments.

Labrum (Fig. 34C) rounded, distally finely setulose. Left

mandible (Fig. 34D) bearing strong, crenulated pars incisiva,

lacinia mobilis robust with five strong denticulations, setiferous

lobe with three trifurcate and one bifurcate setae, pars molaris

robust, blunt, margin with anterodistal row of finely denticulate

teeth (as shown for right mandible, Fig. 34E); mandibular palp

of three articles, proximal article longer than wide with four

setae on inner margin, article 2 twice as long as article 1 with

two longer and three shorter setae in distal half, article 3 0.8

times as long as article 2 with eight inner finely denticulate

setae in distal two-thirds and six longer subdistal to distal

curved simple setae. Right mandible (Fig. 34E) as left but

without lacinia mobilis. Maxillule (Fig. 34F) inner endite with

five setulate distal setae, inner and outer margins setulose;

outer endite with ten distal spines and two subdistal setae,

outer margin finely setulose; palp of two articles, distally with

four setae. Maxilla (Fig. 34G) with sparse fine setae on outer

margin; outer lobe of moveable endite with three simple

subdistal setae and seven simple distal setae; inner lobe of

moveable endite with five simple and seven setulose distal

setae, inner margin with three subdistal simple setae; outer

lobe of inner endite distally with four outer simple setae, one

distally-bilaterally-setulose spine, two mid-distal simple

spines, three stout trifurcate spines, and one inner stout spine

distally setulose on outer margin, subdistally with one distally-

bilaterally-setulose spine; inner lobe of fixed endite with

rostral row of 18 setae guarding six longer finely-denticulate

setae, inner margin finely denticulate. Labium (Fig. 341) with

microtrichia along outer margin, palp with fine lateral setules

and three simple distal spines. Maxilliped (Fig. 34H) basis

Fig. 33 . Parapseudes blandowskii sp. nov., holotype female, dorsal view.

Scale = 1 mm.

Fig. 34. Parapseudes blandowskii sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, left mandible; E, right mandible; F, maxillule;

F', maxillule palp; G, maxilla; H, maxilliped; H', maxilliped endite; I, labium. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 35. Parapseudes blandowskii sp. nov. A, female cheliped; B, male cheliped (exopodite not shown); C, uropod. Scale = 0.1 mm.

naked; palp article 1 with single fine distal spine on outer

margin and three fine simple inner proximal setae; palp article

2 longer than wide, with rows of numerous short setae and two

longer simple setae along inner margin, outer margin with four

distal setae; palp article 3 as long as wide, with 14 simple setae

along inner margin, in two rows; palp article 4 with seven

distal setae and one subdistal seta. Endite (Fig. 34H') with

bilaterally-setulose inner caudodistal seta, distal margin with

simple outer setae and inner half bearing slender, distally

rugose spines.

Cheliped (Fig. 35A) slender, basis 2.7 times as long as wide,

dorsally naked, ventrally with one subdistal and paired distal

fine setae; exopodite present, 3-articled, distal article with four

plumose setae. Merus lozenge-shaped, with five ventrodistal

simple setae. Carpus 2.6 times as long as wide, with two mid-

ventral and two ventrodistal setae. Chela palm (propodus)

longer than wide, with ventral submarginal group of three

setae, dorsal submarginal row of three shorter setae, comb of

four longer setae adjacent to dactylus articulation. Chela fingers

shorter than palm, ventral margin of fixed finger with five

setae; two setae near inner base of fixed finger; cutting edge

with fine spinules and row of eight setae but no apophyses,

distal claw slender, curved; dactylus with three subdistal setae,

row of stout setae along cutting edge, distal claw pointed.

Pereopod 1 (Fig. 36A, B) basis 3.7 times as long as wide,

dorsally with two proximal spines and adjacent seta, and one

subdistal spine, ventrally with proximal and mid-ventral setae,

ventrodistally with small spine and tuft or two shorter and three

longer setae; exopodite present, 3-articled, distal article with six

plumose setae. Ischium with single dorsodistal and tuft of longer

ventrodistal setae. Merus half as long as basis, expanded distally,

with entire row of ventral simple setae, submarginal spinules,

and ventrodistal spine, dorsally with slender, curved dorsodistal

spine and long adjacent simple setae almost as long as carpus.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 36. Parapseudes blandowskii sp. nov. A, pereopod 1; B, pereopod 1 basis details; C, pereopod 2; D, pereopod 3; E, pereopod 4; E', distal

articles of pereopod 4; F, pereopod 5; G, pereopod 6. H, pleopod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Carpus as long as merus, with six or seven ventral spines and

intervening simple seta, dorsal margin with numerous simple

setae and slender, curved dorsodistal spine. Propodus as long as

carpus, with nine ventral spines alternating with simple setae,

four ventral submarginal spinules, two distal spinules, simple

dorsal setae in proximal half and two dorsodistal spines.

Dactylus half as long as propodus, with mid-dorsal fine setae;

unguis half length of dactylus.

Pereopod 2 (Fig. 36C) more slender. Basis 3.7 times as

long as wide with small dorsal spine in distal half and

ventrodistal tuft of setae mostly twice as long as ischium.

Ischium with ventrodistal tuft of setae as long as merus. Merus

0.7 times as long as carpus, with curved dorsodistal spine, row

of ventral simple marginal setae and straight ventrodistal

spine. Carpus with five ventral spines interspersed with setae,

groups of inner mesial and dorsodistal simple setae, and one

shorter straight subdistal spine and one longer curved

dorsodistal spine. Propodus articulating anaxially on merus,

just longer than merus, with five ventral and two dorsodistal

spines with interspersed setae. Dactylus curved, with fine mid¬

dorsal seta, unguis slender, together 0.8 times as long as

propodus. Pereopod 3 (Fig. 36D) similar to pereopod 2, but

basis with dorsal seta rather than spine, merus with two

ventrodistal spines, carpus with longer marginal and shorter

submarginal ventral spines and three dorsodistal spines,

propodus with dorsal penicillate seta.

Pereopod 4 (Fig. 36E) similar to pereopod 3 but basis stouter,

twice as long as wide; merus with six ventral and one dorsodistal

setae; carpus with numerous ventral and distal setae; propodus

with dorsoproximal penicillate seta; dactylus with claw (Fig.

36E') half length of adjacent setae, half length of propodus.

Pereopod 5 (Fig. 36F) similar to pereopod 4, but basis with fine

dorsoproximal setae; carpus with curved dorsodistal spine,

ventral margin densely setose and spinose; dactylus plus unguis

0.8 times as long as propodus. Pereopod 6 (Fig. 36G) similar to

pereopod 5 but basis with plumose setae along entire dorsal and

ventral margins, two ventrodistal setae plumose; merus with

single long dorsal plumose seta; carpus with three dorsal plumose

setae in proximal half; propodus with ventrodistal submarginal

row of 30 spinules, fine distal compound spinules, dactylus plus

unguis 0.8 times as long as propodus.

Pleopods (Fig. 36H) in four pairs all alike. Basis elongate,

with four dorsal and three ventral plumose setae. Endopod

shorter than exopod without proximal articulation; both rami

slender, with 12 to 14 marginal plumose setae.

Uropod (Fig. 35C) biramous; basis with distal crown of

about 12 simple and two penicillate setae; exopod 2.75 times

as long as basis with seven elongate segments; endopod nearly

five times as long as exopod, with about 27 segments.

Distinctions of male. Penial tubercle conspicuous. Flagella of

antennule and antenna with numerous aesthetascs.

Cheliped (Fig. 35B) robust and highly dimorphic; basis 2.4

times as long as wide, dorsally with conspicuous paired tooth¬

like apophyses in proximal half, ventrally with mid-ventral

spine and two fine distal setae; exopodite present, 3-articled,

distal article with four plumose setae. Merus stout with

ventrodistal shoulder bearing five setae. Carpus as long as wide,

with paired mid-dorsal setae, dorsal subdistal tooth-like

apophysis, ventroproximal hooked apophysis with three

adjacent setae, ventrodistal corner finely rugose with row of five

setae. Chela palm (propodus) as long as wide with mid-distal

triangular apophysis with tuft of numerous setae; fixed finger

distally squared with conspicuous proximal invagination,

ventral margin with five longer setae and distal comb of seven

shorter setae; cutting edge with small apophyses; dactylus with

two larger apophyses on cutting edge, distal claw overreaching

fixed finger.

Basis of pereopod 1 (Fig. 36B) dorsally with two proximal

spines and adjacent seta, and one subdistal spine as female.

Etymology, named after Wilhelm Blandowski (1822-1878), a

founder of the Geological Society of Victoria, and the first

scientist appointed to the then new Victorian Museum,

Melbourne, on 1 April 1854.

Remarks : Lang (1965) synonymized all Parapseudes material

worldwide into P. latifrons (Grube, 1864), with a putative

distribution from the Yugoslavian Adriatic (type locality), the

Mediterranean, the Atlantic Ocean, the Caribbean, Pacific

Central and South America through Hawaii to Japan. His

decision was based on observing variation in the number of

uropod segments, the number of ventral spines on the distal

articles of pereopod 1, and the number of segments in the

antennule flagella, all characters on which earlier species had

been distinguished. From our present knowledge of sibling

species in Tanaidacea, such a synonymy is no longer tenable.

Both Gut,u (1998a), in his preliminary reassessment of the genus,

and Larsen & Shimomura (2008) in their sensible discussion of

Parapseudes, point out that the many described species require

detailed re-examination in order to determine their validity, and

indeed to understand the world-wide diversity of this genus.

Gufu (1998a; 1998b; 2001) distinguished four species in

the genus based, inter alia, on the number of dorsal proximal

spines on the basis of pereopod 1. To extend this concept, P.

latifrons sensu Sars (1882) (Mediterranean), P. algicola

(Shiino, 1952) (Japan) and P. francispori (Bacescu, 1980)

(Mediterranean) have one proximal spine, P. latifrons sensu

Gufu (1998b) (Tanzania) and P. latifrons sensu Lang (1965)

(Japan) (both non Rho'ea latifrons Grube, 1864) have two

proximal spines, and P. inermis (Silva Brum, 1974) (Brazil)

and P. trispinosus Gufu, 1998(a) (Indonesia) have three; none

of these have a subdistal spine. P. pedispinis (Boone, 1923)

(California) has one proximal spine and one subdistal spine on

the pereopod 1 basis. P. neglectus Miller, 1940 (Hawaii),

P. similis Vanhoffen, 1914 (Cape Verde) and P. spongicola

Brown, 1958 (South Africa) apparently have no such spines,

although the original (and only) descriptions of these three

species are somewhat wanting. P. arenamans Larsen &

Shimomura, 2008 (Japan) definitely has no such spines.

These last authors suggested that the appearance of such

spines may be an artefact based on setae embedded in mucus:

this is not the case for the present material, nor for that of

P. latifrons agg. sensu Bamber (2005) from S.W. Australia

(see below). The spination of the pereopod 1 basis in P. goodei

Richardson, 1905 (Bermuda) and P. hirsutus Stebbing, 1910

(Chagos) is not known.

M. Biazewicz-Paszkowycz & R.N. Bamber

Parapseudes blandowskii sp. nov. has two proximal spines

on the basis preceded by a seta and one subdistal spine dorsally

on the basis of pereopod 1. This basis spination most closely

resembles that of P. latifrons agg. of Bamber (2005), from

Esperance, southwest Australia, (based only on females) which

has three proximal and one subdistal basis spines, and of P.

pedispinis, redescribed from California by Menzies (1953),

although that species has only one proximal spine and is without

the adjacent seta. Further, the male cheliped of P. pedispinis (as

that of all other described males) is without the dorsal and ventral

tooth-like apophyses on the carpus shown by P. blandowskii.

The Esperance species, currently being redescribed elsewhere,

has a distinctly different dactylus on pereopod 4.

From species where the spination of the pereopod 1 basis

is unknown, P. blandowskii differs in the ventral spination of

the merus, carpus and propodus of pereopod 1, in the number

of segments in the antennule flagella (notwithstanding the

variation inferred by Lang, 1965), in the proportions of the

pereonites, the plumose setation of pereopod 6, and particularly

in the conformation of the male cheliped carpus, inter alia.

Both Larsen & Shimomura (2008) and Gufu (1998a; 1998b)

found in their species and in P. francispori (see Gufu, 2001)

that the dactylus of pereopod 4 was reduced to a small tubercle¬

like structure with the unguis reduced to a seta, and both

speculated that this might be the norm in the genus. The present

species shows a normal (although reduced in size) dactylus plus

unguis on pereopod 4 (Fig. 36E, detail). A normal dactylus and

unguis were also shown by Sars (1886) for what must be taken

as P. latifrons sensu stricto, and by Shiino (1952) in P. algicola,

while that of P. latifrons agg. from Esperance has a reduced but

not tubercle-like dactylus and unguis.

Parapseudes blandowskii is only the second Parapseudes

species presently known from Australasia, and occurred on

sandy substrata at depths of 10 to 15 m in Western Port,

occasionally deeper outside that embayment. P. latifrons agg.

of Bamber (2005) was found on sandy substrata with rhodoliths

and on the red alga Osmundaria prolifera at depths from 18 to

40 m in Esperance Bay, southwest Australia.

Gufu (1998a) gave a revised diagnosis for the genus, but

failed to include therein two significant characterizing

features, viz the dorsal row of setae on pleonite 1 and the

presence of only four pairs of pleopods. This last characteristic

is particularly diagnostic for Parapseudes.

Genus Saltipedis Gufu, 1995

Saltipedis nugoris Biazewicz-Paszkowycz & Bamber, 2007

Figure 37

S. nugoris Biazewicz-Paszkowycz & Bamber, 2007a, 26-31, figs

13-15.

Magniaculeus nugoris Gufu, 2008, p. 58.

Description of male cheliped (Fig. 37). Much more robust than

that of female. Basis stout, twice as long as wide, with

dorsoproximal two-humped apophysis and dorsodistal

expansion into which proximal part of merus fits; with mid-

ventral spine and ventrodistal seta. Exopodite with three

articles, distal article with six marginal plumose setae. Merus

almost rectangular, with mid-ventral seta and tuft of eight

simple distal setae. Carpus stout, 1.5 times as long as wide, with

paired small dorsodistal setae, ventrally with three submarginal

and two marginal setae. Chela stout, propodus with tufts of

setae dorsodistally and on mid-distal apophysis, fixed finger

with array of distal setae as figured, conspicuous tooth-like

apophysis on cutting edge; dactylus cutting edge with fine

crenulation and setules distally, tooth-like apophysis proximally.

Remarks. This species was described originally from numerous

specimens collected throughout the Bass Strait, on muddy- to

coarse sandy substrata from depths between 12 and 293 m.

Numerous further samples of this species exist in the collections

of Museum Victoria, over 100 specimens having been examined

in the course of this study in addition to the type-collection, and

conforming to the same habitat range as the type collection.

This re-examination of material has enabled us to expand on

the original description, to include the dimorphic male cheliped.

While this species was originally confused with Saltipedis forex

Bamber, 2005, until the distinctions from morphological detail

were determined, the male cheliped reinforces the differences:

that of S. forex is without the proximal apophysis on the basis,

has a spine rather than a seta mid-ventrally on the basis (as does

the female), no mid-ventral seta on the merus, and a truncated

fixed finger to the chela (thus without tooth-like apophysis),

more reminiscent of the male chela of the unrelated apseudid

Mendamanus ailurostoma Bamber, 1999.

Saltipedis floccus sp. nov.

Figures 38-40

Material examined. 1 brooding 9 (J55832), holotype, Stn BSS159,

Central Bass Strait, 39°43.5'S 146°18.8'E, 80 m depth, 13 November

1981, muddy shell, coll. R. Wilson; 1 9 (J57600), paratype, Stn BSS155,

Central Bass Strait, 38°55.5'S 145°17.0'E, 70 m depth, 12 November

1981, fine sand, coll. R. Wilson. 2 99 (J57731), paratypes, Stn BSS117,

Central Bass Strait, 40°38.0'S 145°23E, 36 m depth, 4 November 1980,

muddy shell with grit, coll. M. Gomon & G C B Poore. 1 9 (J57721),

paratype, Stn BSS68, Western Bass Strait, 39°27'S 142°55'E, 180 m

depth, 10 October 1980, coarse sand, carbonate, coll. G C B Poore. 1 ( 5 , 6

subadults (J57718), paratypes, Stn BSS 133T, Central Bass Strait, 30 km

N of Wynyard, Tasmania, 40°33.07'S 145°44.68'E to 40°36.22'S

145°48.68’E, 68 m depth, mud, 04 February 1981, coll. M.F. Gomon et

al., 20 m otter trawl. 3 9 $ (J57737), paratypes, Stn BSS 158, Central Bass

Strait, 66 km S of Rodondo Island, Victoria, 39°4836'S 146°18.48'E,

82 m depth, sand with silt and mud, 13 November 1981, coll. R.S. Wilson.

1 9 (J57725), paratype, Stn BSS158, Central Bass Strait, 39°49.5'S

146°18.5’E, 82 m depth, 13 November 1981, sand-silt-mud, coll. R.

Wilson. 1 9 (J57725), paratype, Stn BSS158, Central Bass Strait,

39°49.5'S 146°18.5E, 82 m depth, 13 November 1981, sand-silt-mud,

coll. R. Wilson. 1 S without cephalothorax (J55850), paratype, Stn WBES

1746, Western Port, 38°29.47'S 145°06.17'E, 79 m depth, 25 November

1974, Smith-Mclntyre grab, coll. N. Coleman. 1 9 (J55864), paratype, Stn

HP BES 1747/2, Western Port, Victoria, 38°27.32'S 145°0835'E, 18 m

depth, 25 November 1974, Smith-Mclntyre grab, coll. N. Coleman. 1 9

(J55900), paratype, Stn CPBS 100, Western Port, 38°21.15'S 145°13 23%

4 m depth, 24 March 1965, mud and Zostera; 2 99 (J57663), paratype, Stn

CPBS 01S, Western Port, 38°21.73'S 145°13.23'E, 3 m depth, 1 April

1965.1 9 (J17162),l 9 (J46538), paratypes, Stn MSL EG6, Eastern Bass

Strait, 19 km S of Lakes Entrance, Victoria, 38°04'S 148°00'E, 92 m

depth, 12 August 1989, Smith-Mclntyre grab, coll. G.D. Parry.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 37. Saltipedis nugoris, male cheliped. Scale = 0.1 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 38. Saltipedis floccus sp. nov. A, holotype female, dorsal; B, male, dorsal; C, cephalothorax, lateral; C', pleon, lateral. Scale = 1.0 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 39. Saltipedis floccus sp. nov., female. A, proximal articles of antennular peduncle; B, antenna; C, right mandible incisor; D, left mandible

incisor; D', mandible molar; D”, mandible palp; E, maxillule; F, maxilla; G, labium; H, maxilliped; H', maxilliped endite; I, epignath.

Scale A, B = 0.1 mm; C-I = 0.01 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 40. Saltipedis floccus sp. nov., female paratype. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5;

G, pereopod 6; H, pleopod; I, uropod. Scale = A-F= 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Description of female. Body (Fig. 38A) dorsoventrally flattened,

holotype 9.2 mm long (tip of rostrum to posterior of pleotelson),

five times as long as wide, tapering towards posterior.

Cephalothorax subrectangular, as long as wide, with pronounced

triangular rostrum; eyelobes distinct, dark ocelli present. Spine¬

like hyposphenium present mid-ventrally between chelipeds,

but no hyposphenium on pereonite 2. Pereonites 1 to 3 with

tufts of four to six anterolateral setae, pereonites 4 to 6 with

setae dispersed along lateral margins; pereonites 1 and 2

subequal, shortest, about one-third as long as cephalothorax,

pereonite 3 1.2 times as long as pereonite 2; pereonites 4 and 6

subequal, 1.7 times length of pereonite 2; pereonite 5 longest,

2.3 times as long as pereonite 2 (all pereonites respectively 3.3,

3.2, 2.6, 1.8, 1.2 and 1.4 times as wide as long). Pleon one-

quarter as long as whole body, with five free, pleonites bearing

pleopods and backwardly-directed hyposphenia (Fig. 38C');

pleonite 1 with row of setae around anterior margin, posterior

pleonites naked; pleonites 4.7 times as wide as long. Pleotelson

(Fig. IB) rectangular, more than half-length of pleon, twice as

long as wide, with sparse lateral setae.

Antennule: only basal peduncle articles present on any

specimen (Fig. 39A); proximal peduncle article 1.7 times as

long as wide, inner margin with row of five setae at mid-length

and distal tuft of 7 or 8 setae, outer margin with four groups of

setae, a proximal row of penicillate setae, a subproximal tuft

of simple and penicillate setae, a subdistal row of simple setae

and a distal tuft of six simple setae. Second article 1.5 times as

long as wide, 0.5 times length of first, with simple inner distal

setae, simple outer proximal setae and an outer distal tuft of

simple and penicillate setae.

Antenna (Fig. 39B) with proximal peduncle article

expanded on inner margin as a rounded apophysis with three

spinules. Article 2 as long as first, with four simple setae

adjacent to elongate squama bearing seventeen marginal setae.

Peduncle article 3 shorter than wide with inner seta. Fourth

article just longer than third, as long as wide, with two inner

setae; fifth article twice as long as third, with long inner seta.

Flagellum of thirteen segments, mostly with setae longer than

two flagellar segments.

Epistome conspicuous (Fig. 38C). Labrum rounded,

distally setulose. Right mandible (Fig. 39C) with three rounded

“teeth” on pars incisiva; setiferous lobe with four trifurcate

and one simple setae; left mandible (Fig. 39D) as right but with

dentate lacinia mobilis, and six trifurcate setae on setiferous

lobe; pars molaris (Fig. 39D') stout, blunt with distal rugosity;

palp (Fig. 39D") of three articles, proximal article with field of

twelve inner setae; second article twice as long as first with

three shorter and one longer simple inner setae at mid-length;

third article as long as first, with eleven shorter inner subdistal

setae, two longer distal setae and three shorter submarginal

dorsal setae. Labium (Fig. 39G) with outer rows of microtrichia,

inner distal margin finely setose, palp with inner fine lateral

setules, three simple distal setae, outer proximal microtrichia

and small, rounded, setose inner apophysis. Maxillule (Fig.

39E) inner endite with outer apophysis, and four compound

distal setae; outer endite with twelve distal spines and two

subdistal setae, inner and outer margins finely setose; palp of

two articles, distally with six graduated setae. Maxilla (Fig.

39F) typical of genus, outer margin denticulate, moveable

endite outer lobe with two subdistal and five distal finely

denticulate setae, inner lobe with twelve plumose/denticulate

setae; fixed endite outer lobe with simple, trifurcate, plumose

and bilaterally denticulate distal spines, inner lobe with nine

longer plumose setae and rostral row of 37 setae. Maxilliped

(Fig. 39H) with simple setae; basis with inner and outer simple

setae and outer distal spine-like apophysis; first palp article

with paired inner and outer setae and outer distal spine-like

apophysis; second palp article with inner margin bearing

numerous setae largely in two rows, longest two inner setae

exceeding fourth article, and tuft of nine outer distal setae;

third palp article with about 13 recurved inner setae; fourth

palp article with seven setae around distal margin and one

outer subdistal seta. Endite (Fig. 39H') distal margin with

outer simple setae, central rod-pike setae and inner blunt

spines, inner margin with plumose setae and four coupling

hooks. Epignath (Fig. 391) large, cup-shaped, with setose outer

margin and finely plumose distal spine.

Cheliped (Fig. 40A) slender. Basis 2.7 times as long as

wide, with mid-ventral spine and two subdistal penicillate

setae; dorsally naked. Exopodite 3-articled; article naked,

article 3 with four plumose setae. Merus subrectangular, with

ventroproximal, mesial and ventrodistal groups of setae.

Carpus very slender, four times as long as wide, two simple

setae along ventral margin, five setae along inner midline, and

tufts of shorter dorsodistal and ventrodistal setae. Chela

slender, palm (propodus) 1.2 times as long as wide with

numerous setae on inner face; ventral margin setose along

fixed finger; cutting edge of fixed finger without apophyses but

with small curved setae. Dactylus longer than palm, with no

apophyses on cutting edge.

Pereopod 1 (Fig. 40B) basis 2.35 times as long as wide,

with small ventral spinules, ventrodistal spine and long

ventrodistal setae exceeding distal margin of ischium, dorsal

margin with four setae in proximal half. Exopodite

conspicuous, 3-articled, article 2 with one seta, article 3 with

five plumose setae. Ischium with three ventrodistal setae.

Merus wider distally, with slender dorsodistal spine and

shorter, stouter ventrodistal spine, tufts of simple setae

ventrally, mesially and dorsodistally. Carpus compact, as long

as merus, with two elongate dorsodistal spines, two shorter

ventrodistal spines, setae along entire dorsal margin,

dorsodistal, ventral and ventrodistal tufts of setae. Propodus

as long as carpus, with six ventral spines increasing in length

towards distal margin interspersed with setae, two dorsodistal

slender spines and row of setae along dorsal margin, mesial

field of sparse setules. Dactylus stout, with two ventral

denticulations and dorsal seta, unguis distinct, pointed, both

together 0.8 times as long as propodus.

Pereopod 2 (Fig. 40C) basis 1.6 times as long as wide, with

three ventral spinules, and tuft of long ventrodistal setae

exceeding distal margin of ischium. Ischium shorter than wide

with tuft of seven ventrodistal setae; merus shorter than

carpus, with dense field of ventral setae, two ventrodistal

slender spines, dorsal margin naked. Carpus with diagonal

row of setae along inner face, two dorsodistal slender spines,

ventral margin with five fine spines interspersed with setae;

M. Biazewicz-Paszkowycz & R.N. Bamber

propodus just longer than carpus, ventrodistal margin with

five fine spines interspersed with setae, dorsally with three fine

spines interspersed with setae. Dactylus compact, unguis

shorter, distinct, both together 0.6 times as long as propodus.

Pereopod 3 (Fig. 40D) similar to pereopod 2, but basis

armed only with one mid-ventral and three dorsoproximal

penicillate setae.

Pereopod 4 (Fig. 40E) basis slender, with sparse ventral

spinules and tuft of ventrodistal setae. Ischium with

ventrodistal row of six setae. Merus with numerous ventral

setae and one ventrodistal slender spine. Carpus 1.3 times as

long as merus, ventrally with five fine spines interspersed with

long setae, diagonal row of setae along inner face distally

including five fine spines. Propodus just longer than carpus,

with dorsoproximal penicillate seta, ventrally with five fine

spines interspersed with long setae, diagonal row of setae

along inner face distally including two fine spines; dactylus

and claw elongate, 0.75 times as long as propodus.

Pereopod 5 (Fig. 40F) basis stouter with single ventrodistal

seta, ischium with slender dorsodistal spine, spines on merus,

carpus and propodus longer than those on pereopod 4, distal

propodal spines finely denticulate.

Pereopod 6 (Fig. 40G) similar to pereopod 5 but basis

without complete dorsal marginal row of plumose setae, five

ventral plumose setae in distal half; ischium with two

dorsodistal spines; propodus ventral and distal margin with

row of some 26 small leaf-like spines.

Pleopods (Fig. 40H) all alike. Basis with five ventral

(inner) plumose setae and six dorsal (outer) plumose setae;

rami linguiform. Endopod longer than exopod, respectively

with 24 and 22 marginal plumose setae.

Uropod (Fig. 401) biramous. Basis with two tufts each of

four distal setae. Exopod five times as long as basis and of

about ten poorly-distinguished segments; endopod damaged

on all specimens, but at least twice as long as exopod.

Description of male. Only one male found (Fig. 38B), with

damaged antennae and antennules, without chelipeds. Body

similar to that of female, but pereonites 4 and 5 proportionately

shorter; cephalothorax with three lateral setae posterior to each

eyelobe; all pereonites and pleonites with more dorsal setae.

Etymology. From the Latin, floccus, a tuft or lock of hair, with

reference to the distinctive tufts of setae on the lateral margins

of the pereonites.

Remarks. Saltipedis floccus sp. nov. is the fourth species of the

genus to be described from Australian waters after S. forex,

S. incognita Bamber, 2005 (both from southwestern Australia),

and S. nugoris from the Bass Strait (see above). All four are

generally similar in their habitus (as are all species of the genus

other than S. achondroplasia Bamber, Bird & Angsupanich,

2003), and they share the unusual feature, not found in any other

species of the genus, of outer spine-like apophyses on the

maxilliped basis and palp-article-1. They are, however, readily

distinguished: S, nugoris and S. forex are without an epistome,

and have a triangular or blunt rostrum respectively, while

S. floccus and S. incognita have a conspicuous epistome and a

pointed rostrum, but no hyposphenia on pereonites 1 or 2

(present in S, nugoris)-, S. forex is the only one of these species

without a cephalothoracic hyposphenium between the chelipeds,

or a dorsodistal spine on the merus of pereopod 1.

S. forex is notably different from the other Australian

species owing to the tufts of setae on the lateral margins of its

pereonites; the mouthparts are also much more densely setose

(notably the mandibular and maxilliped palps), and the present

species is the only one to have ventral (as well as dorsal)

plumose setae on the basis of pereopod 6.

Despite all of these differences, the four Australian species

do broadly seem to be closely related. Gufu (2008b) moved the

previously-described three species into a separate genus,

Magniaculeus, simply on the feature of the spine-like maxilliped

apophyses, ignoring the numerous features by which they differ.

In fact, the present species disagrees with a number of the

diagnostic characters of Magniaculeus (many of which were

vague or not distinct to the newly-described genus), e.g. the labial

palp is not ovate, and the propodus of pereopod 1 is not longer

than the carpus. Both features are in fact more like that of another

genus - Brachylicoa - which Gufu (2006) also somewhat

tenuously separated from Saltipedis. Equally, the fine spines on

the basis of pereopod 1 in S, nugoris and S. forex are diagnostic

features of another of GufiTs (2006) new genera derived from

Saltipedis, Podictenius. Until a more comprehensive and rational

analysis of the genus Saltipedis (including Brachylicoa,

Magniaculeus and Podictenius) is undertaken, separating these

four species into a separate genus is premature.

Saltipedis floccus was collected throughout the Bass Strait,

at depths between 3 and 180 m, normally on heterogeneous

substrata, and it was often sympatric with S, nugoris.

Genus Remexudes Biazewicz-Paszkowycz & Bamber, 2007

Remexudes toompani Biazewicz-Paszkowycz & Bamber, 2007

R. toompani Biazewicz-Paszkowycz & Bamber, 2007a, 19-25,

figs 10-12.

Remarks. The presently monotypic genus Remexudes shows

affinities to both Saltipedis and Pakistanapseudes, but is quite

distinct owing to the elongate pereopod 1 propodus and the

flattened distal articles of pereopod 2 which are of the fossorial

form more typical of first pereopods in the Apseudomorpha.

With a dorsal row of plumose setae on pleonite 1, it accords

with the Parapseudinae. The outer spine-like apophysis on the

basis of the maxilliped is also found in Australian species of

Saltipedis, although they also have a similar apophysis on the

first palp article.

Remexudes toompani occured throughout the Bass Strait,

on sandy substrata from depths of 11 to 630 m.

Family Pagurapseudidae Lang, 1970

Subfamily Hodometricinae Gufu, 1981

Genus Indoapseudes Bacescu, 1976

Indopaseudes macabre Bamber, 2005

I. macabre Bamber, 2005, 650-654, figs 17-18.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Material examined. 1 ? with oostegites, 1 brooding ? (J46401), Cruise

81-T-l Stn 196 DP, Western Bass Strait, 6km W of Currie, King Island,

38°54.7'S 143°43.4'E, 49 m depth, coarse sand, 21 November 1981,

coll. R.S. Wilson, Smith McIntyre grab, RV Tangaroa.

Remarks. Of the three described species of Indoapseudes,

I. macabre is the only species found in Australia (so far). The

type (and only other) material was of 11 specimens collected

in Esperance, southwestern Australia, in association with

macroalgae, from 18 to 26 m depth: the present specimens

extend the distribution to southeastern Australia, and the lower

end of the depth range to 49 m.

Genus Similipedia Gufu, 1989

Similipedia diarris Blazewicz-Paszkowycz & Bamber, 2007

S. diarris Blazewicz-Paszkowycz & Bamber, 2007b, 146-147, figs

24-26.

Material examined. 5 specimens, Stn. BSS 158, Central Bass Strait,

66 km S of Rodondo Island, 39°49.5'S, 146°18.5'E, 82 m depth, “sand-

silt-mud”, 13 November 1981, coll. R. Wilson, RV Tangaroa.

Remark. Similipedia diarris was originally described from 45

specimens collected off Wilson’s Promontory at 65 m depth.

The present specimens are from slightly further south, and

slightly deeper water. The only other species of the genus,

S. eminescui Gufu, 1989, was recorded from the north-east

Mozambique Channel.

Subfamily Pagurapseudinae Lang, 1970

The Pagurapseudinae incorporates species which are

obligately adapted to living within gastropod shells, and show

extreme morphological adaptations, often convergent with

those of pagurid decapods. In particular, the pereon and pleon

are twisted, the pleotelson consequently asymmetrical, the

number of pleopods is usually reduced, the chelipeds are

robust and often asymmetrical, the first pereopods are

proportionately large (long), and the second to sixth pereopods

bear rows of short, cylindrical spines or tubercles on the

merus, carpus and propodus, used for gripping the inside of

the empty snail-shell.

There are three genera described within this subfamily,

Pagurapseudes Whitelegge 1901, Pagurotanais Bouvier,

1918, and Macrolabrum Bacescu, 1976(b), but, as more species

have been discovered over the years, there has been some

confusion over the features which distinguish or characterize

them (see Gufu, 1996b; Bamber, 2007; 2008). Pagurotanais is

distinguished in having an exopodite present on the cheliped

(absent in one species) but absent on pereopod 1 (these being

respectively absent and present in the other two genera).

Macrolabrum was distinguished by, and named for, an

unusually long epistome exceeding the tip of the rostrum

(anterior margin of the carapace) when viewed from above. In

comparison with Pagurapseudes, this genus usually also has

pronounced cheliped dimorphism in the male and robust distal

setae or spines on the uropod endopod; other features which

have been cited are the basis of pereopod 1 being conspicuously

wider than subsequent articles, and the presence of large

plumose setae on the maxilliped palp. However, some of these

features are subjective, and some overlap these two genera as

defined by the other characters.

A further character, disregarded before but confirmed in

the present material, which does serve to distinguish these

genera consistently is the conformation of the pleopods.

These are best developed in Pagurapseudes species, with two

equal linguiform rami, each as long as the basis (protopod)

and with a few setae on all margins (e.g. Fig. 44H), and

present on at least 1 and up to 5 pereonites (juveniles have

fewer pairs). In Macrolabrum species, the pleopods are

present as only two pairs in adults, again well developed and

biramous, but, while the exopod is similar to that of

Pagurapseudes species, the endopod is characteristically

shorter and almost circular (e.g. Fig. 50H). In Pagurotanais

species, the pleopods are either absent entirely (including the

generotype, see Bouvier, 1918), or present as a single pair, in

the male only in one species, and of highly reduced form with

the rami bearing 1 to 3 setae, and shorter than the basis, (e.g.

McSweeney, 1982, figs 4G, 6C, unfortunately described

as Pagurapseudes).

As the other features have not been found to be entirely

consistent (e.g. the epistome of Macrolabrum distonyx

Bamber, 2007 does not exceed the anterior margin of the

carapace), this pleopod character is most stable in

distinguishing the genera. As a result, Pagurapseudes abrucei

Bacescu 1981, incidentally a species with a notably wide

basis to pereopod 1 (less than twice as long as wide), is moved

to Macrolabrum. The three genera may thus be keyed out

as follows:

1. Pereopod 1 without exopodite; pleopods in the adult

absent, or present as a single pair with reduced,

unequal rami shorter than basis and bearing 3 or

fewer setae; epistome not exceeding anterior margin

of carapace ... Pagurotanais

Pereopod 1 with conspicuous exopodite; pleopods

present in the adult, with well-developed rami, at

least one of which is linguiform and subequal in

length to the basis ... 2

2. Rami of pleopods subequal in length, linguiform;

epistome not exceeding anterior margin of carapace

... Pagurapseudes

Endopod of pleopods circular and shorter than

exopod; epistome usually (but not always) exceeding

anterior margin of carapace ... Macrolabrum.

The original material of the generotype Pagurapseudes

spinipes, from New South Wales, probably included more than

one species. The type-description clearly accords with a

Pagurapseudes, and is good enough to recognize as a species.

Those specimens which Whitelegge (1901) mentions as

females having no pleopods may well have been Pagurotanais

koonungai Bamber, 2008 (recorded from Brisbane), while his

other specimens with less than three pleopods were possibly

juveniles or other species.

Five distinct species of the Pagurapseudinae were found in

the Bass Strait material, all new, doubling the species

complement for this subfamily in Australia.

M. Blazewicz-Paszkowycz & R.N. Bamber

Genus Pagurapseudes Whitelegge, 1901

Pagurapseudes victoriae sp.nov.

Figures 41-44

Material examined. 1 brooding 9 (J57789), holotype, Stn CPBS 03N,

Western Port, 38°20.56'S 145°15.08'E, 2 m depth, 5 April 1965; 3 99 (1

brooding), 1 S (J57790), paratypes, same sample as holotype; 2 99

(J48009), paratypes, Stn CPBS 25S, Western Port, 38°21.63'S

145°15.08'E, 9 m depth, 23 February 1965, sand; 1 9 with oostegites, 2

66 (J48004), paratypes, Stn CPBS 11S, Western Port, 38°22.00'S

145°13.38’E, 3 m depth, 17 March 1965, shelly gravel; 3 99 (J56614), 2

99 (56612), paratypes, 95 further specimens (unregistered), Stn PPBES

985, Port Phillip Bay, 38°21.00'S 144°41.5'E, 9 m depth, 9 December

1971, sand; 3 99 (J43098), paratypes, Port Phillip Bay “wet sandy

region”, 38°10.51'S 144°43.9’E, 7.5 m depth, 17 October 1994, sand

(labelled “P. spinipes ”); 2 99 (J56618), paratypes, 1 8, 4 99

(unregistered), Stn BSS180, Central Bass Strait, 39°12.9'S 146°27.3'E,

65 m depth, 18 November 1981, medium sand, coll. R.S. Wilson; 5 99

(J56617), paratypes, 10 99 (unregistered), Stn BSS170, Eastern Bass

Strait, 31°51.8S 148°26.5'E, 130 m depth, 15 November 1981, fine

sand, coll. R.S. Wilson; 1 brooding 9 , 1 6 (J56613), paratypes, Stn

BSS209, Eastern Bass Strait, 38°18.0’S 147°37.0’E, 55 m depth, 31 July

1983, muddy fine shell, coll. M. Gomon & R.S. Wilson.

Other material. A further 447 specimens from the Tasmanian

Coast, Flinders Island, Western Port, Port Phillip Bay and throughout

the Bass Strait, at depths from 5 to 69 m.

Description of female. Body (Fig. 41A) typical of a

pagurapseudid, pleon skewed to the right and curved under

pereon; small, holotype about 5 mm long. Cephalothorax (Fig.

41B) subrectangular, as long as wide, rostrum variable (Fig.

42): often trilobed, anterior margin either smooth (Fig. 42A), or

with fine (Fig. 42B) or coarse (Fig. 42C) denticulation, this

variation irrespective of gender or maturity; lateral margins of

branchial chamber with 8 or 9 plumose setae, sparse plumose

setae scattered over dorsal surface of branchial chambers.

Eyelobes distinguished with anterior pointed apophysis, eyes

present as group of black-pigmented ocelli. Epistome not

visible dorsally. Each pleonite with anterolateral and

posterolateral tufts of plumose setae; pereonite 1 shortest, 0.3

times as long as cephalothorax; pereonite 2 1.4 times as long as

pereonite 1; pereonite 3 longer, pereonite 4 longest, nearly

twice as long as pereonite 1; pereonites 5 and 6 progressively

shorter, pereonite 6 just longer than pereonite 2 (all pereonites

respectively 3.0, 2.0,1.4, 1.3,1.4 and 1.5 times as wide as long).

Pleon with five free subequal, asymmetrical pleonites, each

pleonite about one-third as long as pereonite 6, with sparse

lateral and occasional dorsal plumose setae. Pleonites 1 to 3

only bearing pleopods. Pleotelson almost semicircular, longer

than last two pleonites together, just shorter than wide, with

sparse plumose lateral setae and simple distal setae.

Antennule (Fig. 43A) proximal peduncle article 3.25 times

as long as wide, with conspicuous inner-distal tridentate

apophysis and inner-medial expansion bearing tooth-like

apophyses and two plumose setae; outer margin also

denticulate, each “tooth” with an adjacent plumose seta.

Second peduncle article 0.3 times as long as first, expanded

distally to 1.5 times as long as wide, with plumose distal setae;

third article as long as second, fourth article one-third length

of third. Main flagellum of five (rarely six) segments, with

single aesthetascs on each segment; accessory flagellum of

two (rarely three) segments, distally not quite reaching distal

edge of third segment of main flagellum.

Antenna (Fig. 43B) with two basal articles fused into wide

proximal peduncle article inner margin bearing denticulation

and distal apophysis and two plumose and three penicillate

setae, outer margin with blunt apophysis; third article as long

as wide, 0.4 times as long as fused basal articles, with single

simple seta; fourth peduncle article half as long as fused basal

articles, naked; fifth article just longer than fourth, with one

simple and four penicillate distal setae. Flagellum of two

segments, distal segment with three distal setae.

Labrum (Fig. 43D) bilobed, rounded, sparsely setose, small

pointed epistome present (Fig. 43C). Left mandible (Fig. 43E)

with finely denticulate outer margin, quadricuspid pars incisiva,

tricuspid lacinia mobilis, setiferous lobe with simple and

bifurcate setae, pars molaris round, blunt, with ventrodistal

spinules on grinding surface; palp of three articles, proximal

article with robust, plumose inner seta, second article longest,

twice as long as proximal article, with 13 inner distally-

denticulate setae in distal half; third article two-thirds as long

as second, with nine progressively longer distally-denticulate

setae along inner margin, distal seta longer than article. Right

mandible as left but without lacinia mobilis (Fig. 43F). Labium

(Fig. 431) typically marginally setose, palp with two strong

distal setae, fine outer setules and longer inner setae. Maxillule

(Fig. 43G) inner endite with five plumose distal setae outer

apophysis and row of simple setae, inner margin with fine

setules; outer endite with ten distal spines, inner and outer

margins setose; palp (Fig. 43G') of two articles with distinct

articulation, distally with three setae each bearing rounded

setulose tips. Maxilla (Fig. 43H) outer margin setulose, outer

lobe of moveable endite with two subdistal and seven distal

finely setulose setae, inner lobe with four simple and four

setulose setae; fixed endite outer lobe with two bifurcate, two

trifurcate, two setulose and three bilaterally denticulate distal

spines, inner lobe with four longer plumose setae and rostral

row of 20 setae. Maxilliped (Fig. 43K) basis with seven distal

plumose setae, inner margin with two rows of denticulation and

plumose seta; proximal palp article with simple outer margin

with one plumose and one simple setae, and short spine, inner

margin naked; second article with coarsely denticulate inner

and outer margins, three plumose seta on outer margin, two

shorter simple setae and five plumose setae along inner margin;

third article widening distally, with coarsely denticulate outer

margin, six inner simple setae; fourth article with ten distal and

outer sub-distal setae each with fine denticulation in distal half;

endite (not figured) with finely setose outer margin, simple

distal spines, three coupling-hooks. Epignath (Fig. 43J) large,

inner lobes conspicuous, distal spine proximally setose.

Chelipeds (Fig. 44A) showing no conspicuous dimorphism.

Compact basis 1.3 times as long as wide, with complex

proximal surface denticulations dorsally and ventrally, three

mid-ventral simple setae, one simple and two plumose

subdistal ventral setae; exopodite absent. Merus quadrate,

distal half of ventral margin with coarse denticulation, three

inner and one ventral subdistal plumose setae and four mid-

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 41. Pagurapseudes victoriae sp. nov., female. A, dorsal; B, cephalothorax. Scale = 1 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 42. Pagurapseudes victoriae sp. nov., anterior of carapace of A-E, females (lengths of carapaces 1.2 mm - 0.9 mm); F, ovigerous female

(length of carapace 1.3 mm); G, ovigerous female (length of carapace 1.7 mm); H, manca.

ventral simple setae. Carpus elongate, twice as long as wide,

widening distally, with denticulate inner margin, sparse dorsal

and ventral marginal fine setae, two plumose and one simple

inner proximal setae. Propodus elongate, 2.1 times as long as

wide, with few ventral setae; fixed finger with four ventral

setae, three setae adjacent to cutting edge, saw-like row of

small tooth-like spines distally on cutting edge (Fig. 44A');

dactylus curved, cutting-edge with fine denticulations and

three distal tooth-like spines (Fig. 44A').

Pereopod 1 (Fig. 44B) longest pereopod, coxa with slight

apophysis having denticulate margin and one simple and one

plumose setae; basis 2.3 times as long as wide, dorsal margin

bearing seven plumose setae interspersed amongst triangular

tooth-like apophyses with further four submarginal plumose

setae, ventral margin with four ventral and one distal plumose

setae, two simple ventrodistal setae; exopodite present (Fig.

44B'), large, distal article with seventeen plumose setae.

Ischium one-quarter as long as basis, with mid-dorsal simple

seta, ventrally with one plumose and two simple distal setae.

Merus 0.8 times as long as basis, dorsally with two distal

setae, ventral margin with plumose setae and distal spine.

Carpus shorter than merus, with four ventral spines, each with

crenulate anterior face, interspersed with setae. Propodus 1.4

times as long as carpus, with five ventral spines. Dactylus

curved, 0.8 times as long as propodus, with fine ventral setae,

unguis slender, sharp, 0.4 times as long as dactylus.

Pereopods 2 to 6 similar to each other, each about one-half

to one-third as long as pereopod 1. Pereopod 2 (Fig. 44C)

<r C

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 43. Pagurapseudes victoriae sp. nov., female paratype. A, antennule; B, antenna; C, epistome; D, labrum; E, left mandible; F, right mandible;

G, maxillule endites; G', maxillule palp; H, maxilla; I, labium; J, epignath; K, maxilliped. Scale A-B = 0.1 mm; C-K = 0.01 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 44. Pagurapseudes victoriae sp. nov., female paratype. A, cheliped; A', detail of chela; ; B, pereopod 1; B', exopod; C, pereopod 2;

D, pereopod 3; E, pereopod 4; F, pereopod 5; G, pereopod 6; H, pleopod; I, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

basis stout, 2.2 times as long as wide, with one plumose and

two penicillate dorsoproximal setae, simple ventrodistal seta;

ischium with three ventrodistal setae. Merus, carpus and

propodus bearing “sucker-like” spines, generally in two

ventral rows, and plumose setae as figured. Merus longer than

carpus; propodus 0.8 times as long as carpus, distal propodal

spine simple, stout; dactylus and unguis not fused into hook¬

like claw, dactylus with minute inner distal spine. Pereopod 3

(Fig. 44D) with more plumose setae on basis, no dorsal seta on

merus. Pereopod 4(Fig. 44E) basis stouter, 1.7 times as long as

wide, fewer “sucker-like” spines on merus, without stout distal

propodal spine. Pereopod 5 (Fig. 44F) as pereopod 4. Pereopod

6 (Fig. 44G) basis with only one plumose dorsal seta, propodus

with distal denticulate spine adjacent to dactylus.

Pleopods (Fig. 44H) only present on pleonites 1 to 3,

biramous, reduced; basis with single dorsal and ventral

plumose setae; exopod with outer proximal and three distal

plumose setae, endopod with four distal plumose setae, inner

margin with three simple setae and proximal plumose seta.

Uropod (Fig. 441) biramous, basis with two plumose distal

setae; endopod longer than basis, of three segments increasing

in length, second segment distally with two setae, third

segment twice as long as first with one stout and two more

slender distal setae; exopod of one segment, just shorter than

proximal endopod segment, with two distal setae.

Description of male. Male closely similar to female, chelipeds

not significantly dimorphic, but antennule with main flagellum

of six segments. Penial tubercle conspicuous ventrally on

pereonite 6.

Etymology. Named after the State of Victoria (and thus

indirectly Queen Victoria), off which this species is by far the

commonest pagurapseudid.

Remarks. P. inquilinus Bamber 2007, from 440-450 m depth off

New Caledonia, is the only previously-described species of

Pagurapseudes to have two segments in the accessory flagellum

of the antennule (all others having only one), and shows many

similarities to the present species in the morphology of the

antennule, antenna and pereopods, but has seven segments in the

main flagellum (in both sexes). P. victoriae sp. nov. is also

distinguished in having the complex denticulation on the

maxilliped basis and palp (absent in P. inquilinus), and only

three setae on the maxillule palp (six in P. inquilinus)', conversely,

the New Caledonia species has ventral spine-like apophyses

rather than simple setae on the basis of the cheliped, and simple

spines rather than plumose setae proximally on the cheliped

carpus, and is without the dorsal denticulations on the basis of

pereopod 1, has fewer lateral but more dorsal plumose setae on

the carapace, and the uropod exopod is longer than the proximal

endopod segment (shorter in P. victoriae, indeed, notably small

for the genus).

The only species of Macrolabrum to have two segments in

the accessory flagellum of the antennule are M. aenigmaticus

Gufu, 1997, (from Bali), M. boeri Bacescu 1981 and M. abrucei

(Bacescu, 1981) comb. nov. (both from the Great Barrier Reef),

but those species have only four segments in the main flagellum

(the distal segment being comparatively minute in all three),

and no complex apophyses on the proximal article of the

antennule peduncle (as well as typical Macrolabrum pleopods).

The variation in the numbers of antennular flagella articles,

although consistent in their distinction from other species, and

the variation in denticulation of the rostrum in the present

species are notable, as these characters have been used (albeit

not in isolation) in distinguishing between other pagurapseudid

species, which are rarely taken in such profusion as was

P. victoriae.

Pagurapseudes victoriae was collected throughout the

Bass Strait, at depths from 2 to 130 m, on sandy substrata.

Pagurapseudes kimbla sp. nov.

Figures 45-47

Material examined. 1 2 with oostegites (56368), holotype, Stn BSS185,

Western Bass Strait, 38°48.0'S 143°14.5'E, 47 m depth, 20 November

1981, hard rock, coll. R.S. Wilson; 1 juvenile (J56613), paratype, Stn

BSS68, Western Bass Strait, 39°27'S 142°55'E, 183 m depth, 10

October 1980, bryozoan mud, coll. G C B Poore; 1 ? (J56370),

paratype, Stn BSS171, Eastern Bass Strait, 38°53.7’S 147°55.2'E, 71 m

depth, 18 November 1981, medium sand, coll. R.S. Wilson; 1 ? with

oostegites (J55834), paratype, Stn BSS162, Central Bass Strait,

40°09.2'S 147°31.9'E, 51 m depth, 14 November 1981, shelly sand, coll.

R.S. Wilson; 1 9 (J56369), paratype, Stn BSS203, Central Bass Strait,

39°22.0'S 144°18.3'E, 60 m depth, 23 November 1981, coarse sand,

coll. R.S. Wilson; 1 subadult (J29171), paratype, Stn MSL EG88,

Eastern Bass Strait, 37°52.65’S 148°42.15’E, 49 m depth, 4 June 1991,

coarse sand, coll. N Coleman; 1 2 (J29172), paratype, Stn MSL EG115,

Eastern Bass Strait, 37°52.65’S 148°42.15'E, 49 m depth, February

1991, coarse sand, coll. N Coleman; 1 brooding $ (J29173), paratype,

Stn MSL EG111, Eastern Bass Strait, 37°52.65'S 148°42.15’E, 49 m

depth, February 1991, coarse sand, coll. N Coleman; 1 subadult

(J51377), paratype, Stn VC 27 Cl, Central Bass Strait, 36°23.92’S

145°18.43’E, 40 m depth, 11 May 1998, fine sand, coll. N Coleman.

Description of female. Body (Fig. 45A) typical of a

pagurapseudid, small, holotype about 1.7 mm long.

Cephalothorax (Fig. 45B) slightly rounded, just longer than

wide, with convex anterior margin, rostrum a finely-denticulate

semicircle; lateral carapace without denticulations, sparse,

irregular plumose setae. Eyelobes rounded, eyes present as

group of black-pigmented ocelli. Epistome not apparent.

Pereonites 1 and 3 subequal, two-thirds as long as cephalothorax,

pereonite 1 with paired anterolateral plumose or simple setae

and single posterolateral simple setae; pereonite 2 shortest, 0.6

times as long as cephalothorax; pereonites 4 to 6 subequal, 0.9

times as long as cephalothorax; pereonite 5 longest, as long as

cephalothorax. Pleon of five free subequal pleonites, each

pleonite about half as long as cephalothorax. Pleonites 1, 2 and

3 only bearing pleopods. Pleotelson semicircular, 1.5 times as

long as last pleonites, with plumose lateral seta on each side,

paired longer and single shorter simple setae above uropod

attachment, and two small posterior spines (Fig. 471).

Antennule (Fig. 46A) proximal peduncle article 3.3 times

as long as wide, with three larger and one smaller inner spine¬

like apophyses accompanied by plumose setae, no distal

apophysis; outer margin with two proximal tooth-like

apophyses and numerous penicillate and fewer plumose setae.

Second peduncle article 0.35 times as long as first, with array

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 45. Pagurapseudes kimbla sp. nov., holotype female. A, lateral view; B, cephalothorax. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 46. Pagurapseudes kimbla sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, left mandible; D', mandible palp; D” mandible

molar; E, maxillule; F, maxilla; G, labium; H, maxilliped; H', maxilliped endite; I, epignath. Scale = 0.1 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 47. Pagurapseudes kimbla sp. nov., female paratype. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5;

G, pereopod 6; H, pleopod; I, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

of plumose and penicillate distal setae; third article 0.9 times as

long as second, fourth article 0.4 times as long as third. Main

flagellum of three segments, with single aesthetascs on each

segment; accessory flagellum of two segments, distally not

quite reaching distal edge of second segment of main flagellum.

Antenna (Fig. 46B) with two basal articles fused into wide

proximal peduncle article with complex inner denticulation,

outer tooth-like apophysis and paired inner and single outer

plumose setae; third article 1.3 times as long as wide with

outer plumose seta; fourth peduncle article just longer than

third, with two outer penicillate setae; fifth article twice as

long as third with distal array of penicillate setae and one

simple seta. Flagellum of two segments, distal segment with

four distal setae.

Labrum (Fig. 46C) bilobed, rounded, sparsely setose.

Right mandible (Fig. 46D) outer margin denticulate, with

quadricuspid pars incisiva, setiferous lobe with four bifurcate

setae, pars molaris (Fig. 46D') round, blunt, with marginal

crenulations; palp (Fig. 46D") of three articles, proximal

article with single long, plumose inner seta (broken on figure),

second article longest, 1.8 times as long as proximal article,

with four inner distally setulose setae in distal half; third

article 0.8 times as long as second, with five progressively

longer distally setulose setae in distal third, distal seta longer

than article. Left mandible as right but with narrow, tricuspid

lacinia mobilis (not figured). Labium (Fig. 46G) outer margin

denticulate, palp with two distal setae and setulose margins.

Maxillule (Fig. 46E) inner endite with four plumose distal

setae (one broken on figure), outer apophysis and setulose

margin; outer endite with 10 distal spines, outer margin finely

setose; palp of two articles with distinct articulation, distally

with four simple setae with rounded setulose tips. Maxilla

(Fig. 46F) moveable endite damaged in preparation; fixed

endite outer lobe with four simple, one sabre-like, three

trifurcate, one setulose and three bilaterally denticulate distal

spines, inner lobe with two longer plumose setae and rostral

row of 13 setae. Maxilliped (Fig. 46H) basis with three outer

plumose setae; proximal palp article with tridenticulate outer

margin with one plumose seta, inner margin naked; second

article with coarse denticulations along inner and outer

margins, one long and one short plumose setae on outer

margin, seven setulose setae and four plumose setae along

inner margin; third article with bidenticulate outer margin, six

inner marginal simple setae each with fine denticulation in

distal half; distal article with seven inner-marginal and three

distal setae, each with fine denticulation in distal half; endite

(Fig. 46H') with naked outer margin, seven compound distal

spines decreasing in size inwards and two subdistal plumose

setae. Epignath (Fig. 461) large, inner lobes conspicuous, distal

spine with setae surrounding tip.

Chelipeds (Fig. 47A) showing no conspicuous dimorphism.

Compact basis 1.6 times as long as wide, with three

dorsoproximal setae, ventroproximal penicillate seta, mid-

ventral spine and ventrodistal seta, ventrodistal margin densely

setulose; exopodite absent. Merus quadrangular, two ventral

simple setae and two tooth-like apophyses on ventrodistal

margin. Carpus elongate, twice as long as wide, widening

distally, with sparse dorsal and ventral fine setae and two tooth¬

like apophyses on mid-ventral margin. Propodus robust, as

long as wide, fixed finger with three ventral, three distal and

three dorsal setae, row of small rounded teeth along cutting

edge; moveable finger stout, curved, with two distal setae.

Pereopod 1 (Fig. 47B) longest pereopod, with basis slender,

4.7 times as long as wide, dorsal margin bearing five plumose

setae interspersed amongst six triangular tooth-like apophyses,

ventral margin with two spines in distal half and distal simple

seta; exopodite present, large, distal article with thirteen

plumose setae. Ischium one-quarter as long as basis, with

naked dorsal margin, three ventrodistal plumose setae. Merus

0.6 times as long as basis, with two simple dorsodistal setae,

ventral margin with nine plumose setae and two subdistal

spines. Carpus shorter than merus, with one ventral simple

and two ventrodistal plumose setae and three ventral spines.

Propodus 1.3 times as long as carpus, with four slender ventral

spines. Dactylus curved, as long as propodus, with fine ventral

setae, unguis slender, sharp, 0.4 times as long as dactylus.

Pereopods 2 to 6 similar to each other, each about one-half

to one-third as long as pereopod 1. Pereopod 2 (Fig. 47C) basis

stout, twice as long as wide, with plumose ventrodistal seta;

ischium with paired ventrodistal setae. Merus, carpus and

propodus bearing “sucker-like” spines, generally in two

ventral rows, and plumose setae as figured. Merus 1.2 times as

long as carpus; propodus 1.1 times as long as carpus, without

distal spine; dactylus and unguis not fused, with minute inner

seta. Pereopod 3 (Fig. 47D) similar but carpus as long as

propodus. Pereopod 4 (Fig. 47E) slightly more compact, basis

1.7 times as long as wide, fewer “sucker-like” spines on merus,

carpus longer than merus or propodus. Pereopod 5 (Fig. 47F)

as pereopod 4. Pereopod 6 (Fig. 47G) propodus with dorsodistal

denticulate spine.

Pleopods (Fig. 47H) only present on pleonites 1, 2 and 3,

biramous, reduced; basis naked; exopod with outer proximal

and three distal plumose setae, endopod with four distal

plumose setae. Uropod (Fig. 471) biramous, basis with two

plumose distal setae; endopod longer than basis, of three

segments, first distally naked, second segment as long as first

with simple distal seta, third segment with three distal setae

and one penicillate seta; exopod of one segment, subequal in

length to proximal endopod segment, with two distal setae.

Male. Unknown.

Etymology. The HMAS Kimbla was one of the vessels used on

the Bass Strait Survey between 1979 and 1984 (noun in

apposition).

Remarks. Pagurapseudes kimbla sp. nov. is the only species of

the genus to have two segments in the accessory flagellum and

three in the main flagellum of the antennule. Equally, no

previously described species of the related genus Macrolabrum

has this combination of antennular flagellar segments. The

present species is also unusual in having the propodus of

pereopod 2 longer than the carpus (in other species it is

conspicuously shorter). The only other species of Pagurapseudes

to have only two segments in the antennular accessory flagellum

are P. inquilinus Bamber (2007) from 440-450 m depth off

New Caledonia, which has seven segments in the main

M. Biazewicz-Paszkowycz & R.N. Bamber

flagellum (Bamber, 2007), and P. victoriae (see above) which

has 5 or 6 segments in the main flagellum. Both of those species

have one dorsal and one ventral seta on the pleopod basis,

whereas P. kimbla has none.

Pagurapseudes kimbla was taken only occasionally, from

throughout the Bass Strait at depths from 40 to 183 m, and on

varied substrata.

Genus Macrolabrum Bacescu, 1976

Macrolabrum tangaroa sp. nov.

Figures 48-50

Material examined. 1 S (J56366), holotype, Stn BSS202, Western

Bass Strait, 39°00.2'S 144°33.9'E, 74 m depth, 23 November 1981,

sandy shell, coll. R.S. Wilson.

Description of male (limited by dissection of half of only

available specimen). Body (Fig. 48A) typical of a pagurapseudid,

small, holotype about 1.85 mm long. Cephalothorax apparently

naked, rostrum (Fig. 49A) convex, smooth. Eyelobes

distinguished with anterior pointed apophysis, eyes present as

group of black-pigmented ocelli. Epistome not visible dorsally.

Pereonites 1, 3 and 5 subequal, about 0.6 times as long as

cephalothorax; pereonites 2 and 4 subequal, 1.15 times as long

as pereonite 3; pereonite 6 longest, 1.45 times as long as

pereonite 3. Pleon with five free subequal pleonites, each

pleonite about half as long as pereonite 6. Pleonites 1 and 2

only bearing pleopods. Pleotelson semicircular, about as long

as last two pleonites together, with plumose lateral seta and

simple posterior seta on each side (Fig. 501).

Antennule (Fig. 49B) proximal peduncle article 2.25 times

as long as wide, margins without denticulation or apophyses

inner margin with simple setae, outer margin with simple

setae and two penicillate setae in distal half; second peduncle

article 0.44 times as long as first with simple mesial and distal

setae; third article 0.6 times as long as second, fourth article

half length of second. Main flagellum of two segments, with

single aesthetascs on each segment; accessory flagellum of

one segment, distally just reaching distal edge of second

segment of main flagellum.

Antenna (Fig. 49C) with two basal articles fused into wide

proximal peduncle article with complex inner denticulation

and single inner and outer-distal plumose setae; second article

not as long as wide, with small inner distal spine; third

peduncle article 1.5 times as long as second, fourth 2.5 times

as long as second, both with distal penicillate setae. Flagellum

of two very short segments, distal segment with one short and

one very long distal setae, longer seta 1.5 times as long as

distal three peduncle articles together.

Labrum (not figured) bilobed, rounded, sparsely setose,

epistome present, not reaching anterior margin of carapace. Left

mandible (Fig. 49D) with quadricuspid pars incisiva, narrow,

bicuspid lacinia mobilis, setiferous lobe with four variously

crenulate setae, pars molaris round, blunt, with distal marginal

crenulations; palp (Fig. 49D') of three articles, basal article with

single ventrodistal plumose seta, second article with five ventral

plumose setae in distal half, third article with six plumose setae

in distal third, these setae progressively longer distally such that

proximal seta half length of article, distal seta more than twice

as long as article; right mandible not seen. Labium (Fig. 49G)

typically marginally setose, palp with two distal setae. Maxillule

(Fig. 49E) inner endite with four distally-setulose distal setae,

no outer apophysis; outer endite with 9 distal spines, outer and

inner margins densely setose; palp (Fig. 49E') of two articles

with distinct articulation, distally with four simple setae.

Maxilla (Fig. 49F) outer margin naked, outer lobe of moveable

endite with two subdistal and five distal simple setae, inner lobe

with five simple setae; fixed endite outer lobe with five simple,

one sabre-like, one trifurcate and two bilaterally denticulate

distal spines and subdistal bilaterally denticulate spine, inner

lobe with two longer distally denticulate setae and rostral row of

15 setae; one large inner distally denticulate seta. Maxilliped

(Fig. 49H) basis with two inner plumose setae, three inner

marginal denticulations, outer margin with small setose

apophysis and two fine setae; proximal palp article with coarsely

denticulate inner and outer margins and with one inner and one

outer plumose seta; second article with coarsely denticulate

inner and outer margins extended into large teeth inner-distally,

and with six submarginal plumose setae and four plumose setae

along inner margin, single outer plumose seta; third article with

four setae on slight inner apophysis, each with fine denticulation

in distal half; fourth article with six distal and two outer

subdistal simple setae, each with fine denticulation in distal

half; endite not seen. Epignath (Fig. 491) large, marginally

densely setose, inner lobes inconspicuous, distal spine setose.

Cheliped (Fig. 50A) with compact basis 1.2 times as long

as wide, dorsally naked, ventrally with proximal seta, mud-

ventral spine and six distal setae; exopodite absent. Merus

subtriangular, with ventral plumose setae and ventrodistal

denticulate triangular apophysis. Carpus unusually wide, 1.8

times as long as wide, widening distally to form cuff into

which reflexed propodus could sit, cuff lined with crenulations;

dorsally with sparse proximal plumose and simple setae,

ventrally with sparse simple setae. Propodus robust, as long as

wide with four ventral setae; fixed finger only half length of

body of propodus (“palm”), with three ventrodistal and five

dorsal setae, saw-like row of small teeth distally on cutting

edge; moveable finger stout, curved, with three distal setae and

one seta on cutting edge.

Pereopod 1 (Fig. 50B) longest pereopod, with stout basis

2.2 times as long as wide, dorsal margin bearing nine plumose

setae but no apophyses, two subdistal submarginal outer

plumose setae, ventral margin with three simple setae and

plumose ventrodistal seta; exopodite present, large, distal

article with fifteen plumose setae. Ischium 0.2 times as long as

basis, with naked dorsal margin, simple ventral seta and single

ventrodistal plumose seta. Merus 0.6 times as long as basis,

naked, ventral margin with six plumose setae and nine shorter

denticulate setae. Carpus just shorter than merus, with two

ventral and three ventrodistal denticulate setae and longer

ventrodistal simple seta, dorsodistally with two simple and

two denticulate setae. Propodus as long as merus, with three

ventral spines and two ventrodistal spines with adjacent simple

setae. Dactylus curved, longer than propodus, with fine ventral

setae, unguis slender, sharp, 0.6 times as long as dactylus.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 48. Macrolabrum tangaroa sp. nov., holotype female. A, lateral view. Scale = 0.1 mm.

Pereopods 2 to 6 similar to each other, each about one-half as

long as pereopod 1. Pereopod 2 (Fig. 50C) basis stout, 1.75 times

as long as wide, with two plumose ventral seta; ischium with four

plumose ventrodistal setae. Merus, carpus and propodus bearing

“sucker-like” spines, generally in three ventral rows, and plumose

setae as figured. Merus just shorter than carpus; propodus 0.3

times as long as carpus, with minutely denticulate distal spine;

dactylus and unguis not fused, together longer than propodus.

Pereopod 3 (Fig. 50D) with more setae on basis, only three

plumose setae on ischium, carpus proportionately longer.

Pereopod 4 (Fig. 50E) with only two plumose setae on ischium,

carpus shorter than merus. Pereopod 5 (Fig. 50F) similar to

pereopod 3, fewer sucker-like spines on merus. Pereopod 6 (Fig.

50G) with only one plumose seta on ischium.

Pleopods (Fig. 50H) only present on pleonites 1 and 2,

biramous, reduced; basis with two dorsal but no ventral

plumose setae; exopod with five distal plumose setae,

endopod almost circular, with nine distal and inner plumose

setae. Uropod (Fig. 501) biramous, basis with one simple

and one plumose distal setae; endopod longer than basis, of

two segments, first segment shorter than basis, naked,

second segment with three robust distal setae and one

penicillate seta; exopod of two segments, first segment

shorter than proximal endopod segment, second segment

reaching half length of distal endopod segment, with two

distal setae.

Female. Unknown.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 49. Macrolabrum tangaroa sp. nov., female paratype. A, rostrum; B, antennule; C, antenna; D, left mandible; D', mandible palp; E, maxillule;

E', maxillule palp; F, maxilla; G, labium; H, maxilliped; I, epignath. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 50. Macrolabrum tangaroa sp. nov., female paratype. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod

5; G, pereopod 6; H, pleopod; I, uropod. Scale = 0.1 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

Etymology. The RV Tangaroa was one of the vessels used on

the Bass Strait Survey between 1979 and 1984 (noun in

apposition).

Remarks. Unusually for a Macrolabrum species, the epistome

of M. tangaroa sp. nov. does not exceed the anterior margin of

the carapace (a condition also found in M. distonyx, a species

with a three-segmented uropod endopod). The only other

Macrolabrum species to have two segments in both uropod

rami is M. aenigmaticus (known only from a juvenile), but that

species differs from M. tangaroa in having a four-segmented

main flagellum and a two-segmented accessory flagellum on

the antennule, a propodus on pereopod 2 not shorter than the

carpus, and distinct setation on the pereopod 1 basis, inter alia.

The extremely wide, almost oval, cheliped carpus of

M. tangaroa appears to be unique in the genus, but may

represent a sexual dimorphism (males are not known for all

species). The very short propodus on pereopod 2 is also an

unusual and characterizing feature of this species.

The single specimen of Macrolabrum tangaroa was taken

at 74 m depth in the western Bass Strait.

Macrolabrum sarda sp. nov.

Figures 51-54

Material examined. 1 9 (J57788), holotype, Stn SA63, Flinders Island,

South Australia, “The Hotspot” reef, 5 n miles W of N end of Flinders

Island, 33°40.30'S 134°22.00'E, 17 m depth, 19 April 1995, SCUBA,

coll. G.C.B. Poore; 1 9 , 1 <5 (J56372), paratypes, same sample as

Holotype; 1 9 (J56373), paratype, Stn SA59, Flinders Island, South

Australia, bay on NW coast of Flinders Island, 33°41.42'S 134°28.30’E,

3 m depth, 23 November 1981, hand dredge, coll. G.C.B. Poore; 1 9

(J56366), paratype, Stn BSS180, Central Bass Strait, 8 km south of

South East Point, Wilsons Promontory, Victoria, 39°12.9’S 146°27.3’E,

65 m depth, 18 November 1981, medium sand, coll. R.S. Wilson.

Description of female. Body (Fig. 51A) typical of a

pagurapseudid, small, holotype about 3.5 mm long.

Cephalothorax (Fig. 51A, B) subrectangular, slightly narrower

anteriorly, almost as long as wide, with denticulate rostrum;

paired plumose setae behind ocular lobe, lateral margins with

six hooked spine-like apophyses and seven plumose setae.

Eyelobes distinct, eyes present as group of black-pigmented

ocelli. Epistome conspicuous, exceeding anterior margin of

carapace, visible dorsally. Pereonite 1 with undulating anterior

and posterior margins, 0.4 times as long as cephalothorax and

2.8 times as wide as long, laterally with seven or eight plumose

setae on each side; pereonite 2 similar to but just longer than

pereonite 1, twice as wide as long, laterally with two plumose

setae on each side; pereonites 3 to 6 naked. Pleon 0.4 times as

long as whole body, with five free subequal pleonites, each

pleonite nearly as long as pereonite 6 and slightly wider than

long; pleonites 1 and 2 only bearing pleopods. Pleotelson

subrectangular, about twice as long as last pleonite, 1.5 times as

long as wide, with three lateral setae on each side.

Antennule (Fig. 52A) proximal peduncle article 3.5 times

as long as wide, inner margin with four proximal spine-like

apophyses accompanied by simple setae, further simple setae

in distal half, outer margin with proximal penicillate setae,

distal simple setae and distal spine; second peduncle article

0.5 times as long as first, inner spine-like apophysis at mid¬

length, both margins with single mid-length plumose seta and

tuft of distal plumose setae; third article 0.85 times as long as

second, with short inner and outer plumose setae; fourth

article one-third length of third. Main flagellum of four

segments including minute distal segment, first, second and

third segments with 3, 2 and 1 aesthetascs respectively;

accessory flagellum of three segments, first segment short,

second segment longest and distally exceeding distal edge of

first segment of main flagellum, third segment minute.

Antenna (Fig. 52B) with two basal articles fused into wide

proximal peduncle article bearing complex denticulation

along inner margin, one plumose and one simple inner setae;

third article one-third length of fused proximal articles, as

long as wide with inner distal spine-like apophysis and outer

distal spine and adjacent simple seta; fourth peduncle article

twice as long as third with inner distal spine-like apophysis

and outer distal penicillate seta; fifth article 2.5 times as long

as third with distal crown of penicillate setae and long outer

simple seta. Flagellum of two unequal segments, distal

segment with three distal setae.

Labrum (not figured) bilobed rounded, sparsely setose,

epistome large, exceeding rostrum. Left mandible (Fig. 52C)

with heavily denticulate outer margin, quadricuspid pars

incisiva, tricuspid lacinia mobilis, setiferous lobe with four

distally denticulate setae, pars molaris round, blunt, crushing

face with marginal crenulations; palp of three articles, proximal

article with inner distal crenulations and long, plumose inner

seta, second article longest, 1.7 times as long as proximal article,

inner margin with five plumose and two simple marginal setae,

and 23 shorter simple setae essentially in two rows; third article

half as long as second, with six progressively-longer inner setae

and two outer setulose subdistal setae. Right mandible (Fig.

52D) as left but without lacinia mobilis. Labium (Fig. 52G) with

hook-like denticulation on outer margin, palp elongate, setose,

with two distal setae. Maxillule (Fig. 52E) inner endite with four

distally-denticulate distal setae and outer apophysis below

setose margin, outer endite with 9 distal spines, outer margin

setose; palp of two articles with indistinct articulation, distally

with six setae each minutely denticulate in its distal half.

Maxilla (Fig. 52F) outer margin setulose, outer lobe of moveable

endite with two subdistal and five distal simple setae, inner lobe

with seven simple setae and one plumose seta; fixed endite outer

lobe with fpur simple, three trifurcate and one bilaterally

denticulate distal spines, inner lobe with one longer distally

denticulate seta and rostral row of 16 setae. Maxilliped (Fig.

52H) basis with two inner and three distal setae and small outer

setulose apophysis; proximal palp article with denticulate inner

and outer margins, and one inner and one outer plumose setae;

second article with denticulate inner and outer margins, outer

margin with one distal and three marginal plumose setae, inner

margin with two distal plumose setae and two more in proximal

half, and sparse simple setae; third article with five simple and

four finely-denticulate inner marginal setae; distal article with

six finely-denticulate inner-marginal and distal setae, paired

outer subdistal plumose setae; endite (Fig. 52FT) with finely

setose outer margin, distally with two plumose setae and five

bi- or trifurcate spines progressively smaller towards inner

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

margin, and two subdistal plumose setae; three coupling-hooks.

Epignath (Fig. 521) large, oval, distal spine coarsely setose.

Chelipeds showing dimorphism. Right cheliped (Fig. 53B)

with compact basis 1.3 times as long as wide, with three dorsal

setae on slight apophysis, mid-ventral finely-denticulate spine,

two ventrodistal finely-denticulate spines and ventrodistal

plumose seta; exopodite absent. Merus subtriangular with

pronounced distal triangular extension, five simple and three

plumose ventral setae. Carpus elongate, 2.3 times as long as

wide, dorsally with sparse fine setae and mid-dorsal hook-like

apophysis, ventrally with denticulate margin in distal half,

four simple and one plumose marginal setae. Propodus robust,

1.35 times as long as wide, with three ventral and one

dorsoproximal short setae; fixed finger wide, blunt, distally

rounded, claw nor evident, with row of crenulations distally

and tuft of three proximal setae on cutting edge, two ventral

setae; moveable finger stout, strongly curved, cutting edge

with rounded crenulations. Left chela (not figured) more

slender, fixed finger pointed with distal claw.

Pereopod 1 (Fig. 54A) longest pereopod, coxa with triangular

apophysis bearing two plumose setae; basis stout, 2.3 times as

long as wide, with mid-proximal spine, dorsal margin expanded,

expansion incomplete distally, bearing 19 plumose setae

interspersed with eleven triangular spinules, ventral margin with

three proximal simple setae, three central plumose setae and one

distal plumose seta; exopodite present (Fig. 54A'), large, second

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 52. Macrolabrum sarda sp. nov., female paratype. A, antennule; B, antenna; C, left mandible; D, right mandible; E, maxillule; F, maxilla;

G, labium; H, maxilliped; H', maxilliped endite; I, epignath. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 53 . Macrolabrum sarda sp. nov. A, pleopod; B, cheliped, female. Scale = 0.1mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 54. Macrolabrum sarda sp. nov., male paratype. A, pereopod 1; A', pereopod 1, exopod; B, pereopod 2; C, pereopod 3; D, pereopod 4;

E, pereopod 5; F, pereopod 6; G, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

article with two dorsal setules, distal article with 18 plumose

setae. Ischium as long as wide, with naked dorsal margin, three

plumose ventral setae. Merus 0.8 times as long as basis, dorsally

with two plumose and one simple distal setae, ventrally with 10

marginal plumose setae, 17 submarginal denticulate setae in two

rows. Carpus short, half length of merus, ventrally with three

spines and subdistal simple setae, and with two mid dorsal

simple setae and dorsodistal tuft of three simple setae and one

spine. Propodus 1.7 times as long as carpus, with four ventral

spines in distal half and sparse distal setae. Dactylus stout,

curved, 0.8 times as long as propodus, with three ventral and one

dorsal setae, unguis slender, sharp, 0.3 times as long as dactylus.

Pereopods 2 to 6 similar to each other, each about one-half to

one-third as long as pereopod 1. Pereopod 2 (Fig. 54B) coxa with

simple seta; basis stout, 1.9 times as long as wide, dorsally with

plumose proximal seta and four simple and one penicillate setae

in distal half, ventrally with two distal plumose setae; ischium

with paired ventrodistal setae. Merus, carpus and propodus

bearing “sucker-like” spines, generally in three ventral rows, and

sparse plumose setae as figured. Merus 1.5 times as long as basis;

carpus 0.7 times as long as merus; propodus 0.75 times as long as

carpus; dactylus and unguis not fused, with minute inner seta,

together 0.8 times as long as propodus; adjacent distal propodal,

spine simple but with inner distal fine denticulation. Pereopod 3

(Fig. 54C) with only plumose setae on basis, three ventrodistal

plumose setae on ischium, dorsal penicillate seta on propodus.

Pereopod 4 (Fig. 54D) slightly more compact, basis 1.5 times as

long as wide; fewer sucker-like spines on merus; carpus just

longer than merus; propodus short, 0.3 times as long as carpus,

with two denticulate distal spines; dactylus plus unguis 1.5 times

as long as propodus. Pereopod 5 (Fig. 54E) as pereopod 4 but

with more plumose setae on basis and ischium. Pereopod 6 (Fig.

54F) carpus twice as long as merus, only one sucker-like spine

on merus.

Pleopods (Fig. 53A) only present on pleonites 1 and 2,

biramous, reduced; basis with single dorsal and no ventral

plumose seta, and suggestion of proximal articulation; exopod

with six outer and distal plumose setae, endopod almost

circular, with eight marginal plumose setae.

Uropod (Fig. 54G) biramous, basis outer margin with two

plumose and one penicillate setae, inner margin with one

subdistal simple seta; endopod longer than basis, of three

segments increasing in length, first and second segments each

with inner distal simple seta, third segment with three stout

distal setae each with fine serrations in distal half and one

penicillate seta; exopod of one segment, subequal in length to

proximal two endopod segments together, with three distal

setae each with fine serrations in distal half.

Male. Closely similar to female, chelipeds not significantly

sexually-dimorphic.

Etymology. The SV Sarda was one of the vessels used on the

Bass Strait Survey between 1979 and 1984 (noun in apposition).

Remarks. The only other species of Macrolabrum to show

minute distal segments on both flagella of the antennule are

M. aenigmaticus, M. boeri and M. abrucei, but these have only

two segments in the accessory flagellum; further,

M. aenigmaticus has no pleopod basis setae and only two

uropod endopod segments, while the other two have only one

uropod exopod segment, and M. boeri has six ventral setae on

the pleopod basis.

In fact, the only other species of the Pagurapseudinae to

have a three-segmented accessory flagellum on the antennule

is Pagurapseudes victoriae (see above), and then only in rare,

larger individuals, while the complex setation of the basis and

merus of pereopod 1 and of the second mandible palp article

are unique to Macrolabrum sarda sp. nov..

Macrolabrum sarda sp. nov. was recorded from the

Central Bass Strait, at 65 m on medium sand, as well as from

South Australia at 3 to 17 m depth.

Macrolabrum haikung sp. nov.

Figures 55-57

Material examined. 1 brooding 9 (J57787), holotype, Stn CRUST 23,

“The Whaleback”, Bommie, 0.5 km S of Point Hicks, Victoria,

37°48.30'S 149°16.48'E, 13 m depth, 08 April 1989, SCUBA, coll.

G.C.B. Poore; 14 99 (J56374), paratypes, same sample as Holotype; 1

9 (27697), paratype, Stn BSS175, Eastern Bass Strait, 40 km north of

Deal Island, Tasmania, 39°05.8'S 147°26.2'E, 59 m depth, 18 November

1981, medium sand, coll. R.S. Wilson.

Description of female. Body (Fig. 55A) typical of a

pagurapseudid, pleon skewed to the right and curved under

pereon; small, holotype about 2.4 mm long. Cephalothorax

slightly longer than wide, rostrum rounded, finely denticulate

(Fig. 55B). Eyelobes distinguished with anterior pointed

apophysis, eyes present as group of black-pigmented ocelli.

Epistome not conspicuous. Pereonites 1, 2 and 3, 4 subequal,

0.3 times as long as cephalothorax; pereonites 4 and 5 subequal,

1.4 times as long as pereonite 1; pereonite 6 longest, 1.7 times

as long as pereonite 1. Pleon of five free subequal pleonites,

each pleonite about 0.7 times as long as pereonite 6; pleonites 1

and 2 only bearing pleopods. Pleotelson (Fig. 571)

subrectangular, about as long as last pleonite, 1.4 times as wide

as long, with single plumose lateral seta on each side, and

paired simple posterior setae.

Antennule (Fig. 56A) compact, proximal peduncle article

2.2 times as long as wide, without apophyses, inner margin

sparsely setose, inner margin with three penicillate and one

simple setae distally; second peduncle article 0.4 times as long

as first with simple distal setae; third article 0.7 times as long as

second, with simple distal setae; fourth article half length of

second, with two simple and one penicillate distal setae. Main

flagellum of two segments, with simple setae and single

aesthetasc on each segment; accessory flagellum of one segment,

distally exceeding distal edge of first segment of main flagellum.

Antenna (Fig. 56B) with two basal articles fused into wide

proximal peduncle article bearing inner denticulation and plumose

seta, outer margin expanded into a flange with two simple setae;

third article shorter than wide, one-fifth length of combined

proximal articles, with simple inner seta; fourth peduncle article

1.7 times as long as third, with inner penicillate seta; fifth article

three-times as long as third, with penicillate and simple distal

setae. Flagellum of two minute segments, proximal segment with

penicillate seta, distal segment with two distal setae.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 55. Macrolabrum haikung sp. nov., holotype female. A, lateral view; B, rostrum. Scale = 0.1 mm.

Labrum (not figured) bilobed, rounded, sparsely setose.

Right mandible (Fig. 56C) with tricuspid pars incisiva, setiferous

lobe with three trifid setae, pars molaris slender, round, blunt,

simple; palp of three articles, proximal article with long,

plumose inner seta, second article longest, twice as long as first

article, naked; third article as long as first, with four progressively

longer distal setae and one outer subdistal seta, each seta finely

denticulate. Left mandible (not figured) as right but with narrow,

bicuspid lacinia mobilis. Labium typically marginally setose,

palp (Fig. 56F) with setulose margins and two distal setae.

Maxillule (Fig. 56D) inner endite with four finely serrate distal

setae, margins naked, no outer apophysis; outer endite with 9

serrate distal spines, outer margin setose; palp of two articles,

distally with outer setules and three simple setae. Maxilla (Fig.

56E) outer margin naked, outer lobe of moveable endite with

two subdistal and four distal simple setae, inner lobe with six

simple setae; fixed endite outer lobe with three simple, three

trifurcate and one bilaterally denticulate distal spines, inner

lobe with one longer distally denticulate seta and rostral row of

10 setae. Maxilliped (Fig. 56G) basis with two inner plumose

setae, outer margin denticulate and with one short plumose seta;

proximal palp article with two denticulations and one plumose

seta on inner and outer margins; second article with denticulate

inner and outer margins, outer margin with two plumose setae,

inner margin with four plumose and two simple setae; third

article with three inner marginal simple setae; distal article with

seven finely-denticulate and one simple distal setae, paired outer

subdistal setae each with setules at mid-length and finely-

denticulate distal half; endite (Fig. 56G') with naked outer

margin, five bifurcate distal spines, and two coupling-hooks.

Epignath (Fig. 56H) narrow, inner lobes conspicuous, distal

spine with short marginal setules in distal half.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 56. Macrolabrum haikung sp. nov., female paratype. A, antennule; B, antenna; C, left mandible; D, maxillule; E, maxilla; F, labial palp;

G, maxilliped; G', maxilliped endite; H, epignath. Scale = 0.1 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 57. Macrolabrum haikung sp. nov., female paratype. A, right cheliped; B, left cheliped; C, pereopod 1; C', exopod; D, pereopod 2; E, pereopod 3;

F, pereopod 5; G, pereopod 6; H, pleopod; I, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Chelipeds showing slight dimorphism. Right cheliped (Fig.

57A) with compact basis 1.1 times as long as wide, with mid-

ventral spine, two simple setae proximal to this, and two

plumose setae and one spine ventrodistally; stout proximal

spine dorsally; exopodite absent. Merus subtriangular, with

complex, denticulate triangular distal apophysis, two mid-

ventral, two inner-proximal and three outer-proximal plumose

setae. Carpus elongate, 1.65 times as long as wide, widening

distally, with six teeth in ventrodistal denticulation, four ventral

and four dorsal simple setae. Propodus 1.3 times as long as

wide, single dorsodistal seta, ventrally with two simple setae

and expanded into thin flange; fixed finger with three ventral,

one distal and five dorsal marginal setae, crenulate cutting edge;

moveable finger stout, curved, naked. Left cheliped (Fig. 57B)

similar, ventral denticulations on carpus limited to three on

distal apophysis, chela slightly more slender (propodus 1.2

times as long as wide), with fewer setae and denticulation

restricted to distal half of cutting edge of fixed finger.

Pereopod 1 (Fig. 57C) longest pereopod, with stout basis

twice as long as wide, dorsal margin bearing seven plumose

setae interspersed amongst triangular tooth-like apophyses, and

three rounded paddle-like apophyses proximally; ventral

margin with simple proximal seta and distal plumose seta;

exopodite present (Fig. 57C'), large, second article naked, distal

article with thirteen plumose setae. Ischium 0.3 times as long as

basis, with single ventral plumose seta. Merus relatively compact

for the genus, twice as long as wide, 0.75 times as long as basis,

with two denticulate dorsodistal setae, ventral margin with five

longer plumose setae and four shorter denticulate spines. Carpus

shorter than merus, with denticulate spine and simple seta

dorsodistally and ventrodistally. Propodus as long as merus,

with dorsodistal group of one simple seta, one penicillate seta

and one curved spine, ventrodistally with two simple setae and

one denticulate spine. Dactylus almost straight, just longer then

propodus, with single dorsal but no ventral setae, unguis slender,

curved, blunt, half as long as dactylus.

Pereopods 2 to 6 similar to each other, each about one-half to

one-third as long as pereopod 1. Pereopod 2 (Fig. 57D) basis

stout, 1.7 times as long as wide, naked; ischium with one shorter

and one longer ventrodistal setae. Merus, carpus and propodus

bearing “sucker-like” spines, generally in three ventral rows, and

sparse plumose setae as figured. Merus longer carpus; propodus

0.6 times as long as carpus, both with stout, finely denticulate

distal spine; dactylus and unguis not fused, with minute inner

seta. Pereopod 3 (Fig. 57E) with very short basis, shorter than

wide, ischium with three ventrodistal plumose setae, merus with

fewer sucker-like spines, merus and carpus subequal in length,

carpus and propodus both with stout, finely denticulate distal

spine. Pereopod 4 as pereopod 5. Pereopod 5 (Fig. 57F) with one

dorsal and one ventrodistal setae on basis, one longer and one

shorter ventrodistal setae on ischium, merus without setae,

shorter than carpus; stout, finely denticulate distal spine on

propodus only. Pereopod 6 (Fig. 57G) basis with two plumose

dorsal setae, mid-ventrally with plumose seta and penicillate

seta; ischium with one ventrodistal seta; merus with only one

sucker-like spine; unguis mounted subdistally on dactylus.

Pleopods (Fig. 57H) only present on pleonites 1 and 2,

biramous, reduced; basis with two ventral plumose setae; exopod

with three distal plumose setae, endopod almost circular with

seven marginal plumose setae. Uropod (Fig. 571) biramous, basis

with one simple and one plumose distal setae; endopod longer

than basis, of three segments, first and second segments subequal,

distally naked, third segment longer than first two together, with

three stout distal setae and one penicillate seta; exopod of two

segments, together subequal in length to proximal two endopod

segments together, with two distal setae.

Male. Unknown.

Etymology. The FRV Hai Rung was one of the vessels used on

the Bass Strait Survey between 1979 and 1984 (noun in

apposition).

Remarks. The only other species of Macrolabrum with three and

two segments in the uropod endopod and exopod respectively,

and two and one segments in the antennular main and accessory

flagellum respectively is M. distonyx, from New Caledonia (see

Bamber, 2007), which also has a proximal spine on the cheliped

basis like M. haikung sp. nov. and an epistome not exceeding the

anterior margin of the carapace, but that species has a far more

elongate antennule with denticulation on the proximal peduncle

article, a triangular, denticulate rostrum, four spines on the

mandible palp second article, more slender pereopod bases, and

a huge chela on the right cheliped, inter alia.

Indeed, the present species is unique in its extremely compact

pereopod bases, the relatively short articles of pereopod 1 with

Table 1. Numbers of segments in the antennule flagella and uropod rami, for all described Australian species of Pagurapseudinae.

Species

Main flagellum

Accessory flagellum

Uropod exopod

Uropod endopod

Pagurotanais koonungai Bamber, 2008

Macrolabrum abrucei (Bacescu 1981)

Macrolabrum boeri Bacescu 1981

Macrolabrum impedimenta Bamber, 2005

Macrolabrum tangaroa sp. nov.

Macrolabrum sarda sp. nov.

Macrolabrum haikung sp. nov.

Pagurapseudes spinipes Whitelegge 1901

Pagurapseudes victoriae sp. nov.

5(6)

2(3)

Pagurapseudes kimbla sp. nov.

M. Blazewicz-Paszkowycz & R.N. Bamber

their sparse setation, the lack of ventral setae on the dactylus of

pereopod 1, the subdistal attachment of the unguis on pereopod

6, the serration of the distal spines on the maxillule outer endite,

and in particular the inner flange on the antenna proximal article

and the ventral flange on the cheliped propodus.

Macrolabrum haikung was taken from the Eastern Bass

Strait at 13 to 59 m depth on medium sand.

Comment on the Pagurapseudinae of Australia

The known Pagurapseudinae of Australia are largely

easily distinguishable by the numbers of segments in the

antennule flagella and in the uropod rami, as shown in Table 1.

Suborder Tanaidomorpha Sieg, 1980

Superfamily Paratanaoidea Lang, 1949

Family Paratanaidae Lang, 1949

Subfamily Paratanaidinae Lang, 1949

Genus Paratanais Dana, 1952

Paratanais malignus Larsen, 2001

Figures 58-59

Paratanais malignus Larsen, 2001, 368-372, figs 11-13, 17.

Material examined. 1 9 (J56667), Stn BUN2, off Honeysuckle Hill,

Bunurong, Victoria, 38°40.32'S 145°37.47'E, 10 to 11 m depth, 1 April

1997, SCUBA, coll. T,D. O’Hara. 2 99 (J56668), 1 9 (J56669), Stn

WV11, Beware Reef, near Cape Conran, Victoria, 37°49.21'S

148°47.23'E, 5 to 6 m depth, 15 April 1998, SCUBA, coll. T,D. O’Hara.

3 99 (J56670), Stn WV13, Sailor’s Grave, off East Cape Conran,

Victoria, 37°48.13’S 148°44.41'E, 4 to 5 m depth, 15 April 1998,

SCUBA, coll. T,D. O’Hara. 1 9 (J56671), Stn WV5, Cheviot Beach,

Point Nepean, Victoria, 38°18'S 144°40’E, 1.9 to 3.5 m depth, 31 March

1998, SCUBA, coll. T,D. O’Hara. 1 9 (J56694), Stn BSS 213T, Eastern

Bass Strait, 24 km SW of Lakes Entrance, Victoria, 38°03’S 147°50'E,

45 m depth, 01 October 1983, otter trawl, coll. M. Gomon & R.S.

Wilson. 1 9 (J56734), Stn BSS 181S, Central Bass Strait, 26 km SE of

Aireys Inlet, Victoria, 38°39.48'S 144°18.12'E, 79 m depth, very fine

sand, 19 November 1981, WHOI epibenthic sledge, coll. R.S. Wilson.

1 9 (J56738), Stn BSS 188, Western Bass Strait, 30 km SSW of

Warrnambool, Victoria, 38°38.12'S 142°35.00'E, 59 m depth, 20

November 1981, coll. R.S. Wilson. 1 9 (J56766), Stn PPBES 1218, Port

Phillip Bay, off Brighton, Victoria, 37°54.45’S 144°58.30'E, 4 m depth,

coll. Department of Fisheries & Wildlife Marine Pollution Studies. 3

juveniles (J57546), Stn BSS 158, Central Bass Strait, 66 km S of

Rodondo Island, Victoria, 39°48.36’S 146°18.48'E, 82 m depth, sand

with silt and mud, 13 November 1981, coll. R.S. Wilson. Numerous

other lots in the collections of Museum Victoria.

This species is particularly recognizable owing to its having a

leaf-shaped spine at junction of the chela fingers; other

characterizing features are the rugose lacinia mobilis of the left

mandible, and the dorsal field of marginal setules on article 2 of

the antenna peduncle. The additional material found here allows

some additions to the type-description of Larsen (2001). The

original type-material included only females and mancae; the

present material also includes the male, which is described below.

Supplementary description of female. Body-length up to

3.7 mm. Distal spine on article three of antennal peduncle often

not showing articulation; labrum densely setose (Fig. 58A).

Pars molaris of left mandible with elaborate distal spination

(Fig. 58B). Maxilla (Fig. 58C) subtriangular, naked. Maxilliped

(Fig. 58D) endites with anteromedial seta (originally described

as absent in the more limited type-material), second palp article

with long simple seta in addition to serrated spine and finely

shorter setulose seta.

Cheliped (Fig. 58E) carpus with long inner seta; fixed

finger of propodus with two ventral setae. Pereopods 1 to 3

(Figs 58F, G) with seta on coxa.

Description of male. Body (Fig. 59A) shorter than that of

female, 2.6 mm long, 4.8 times as long as wide. Cephalothorax

subtriangular, as long as wide, longer than pereonites 1 to 3

together; large eyes present, with large pigmented ocelli.

Pereonite 1 shortest, pereonites 2 and 3 progressively longer,

pereonite 3 twice as long as pereonite 1, pereonites 4 to 6 equal

in length, 2.7 times as long as pereonite 1. Pleon with five free

subequal pleonites bearing pleopods.

Antennule (Fig. 59B) of seven articles, proximal article 1.2

times as long as wide, with inner tuft of penicillate setae;

second article 0.7 times as long as wide, about half length of

first, with inner seta longer than article width; third article less

than half length of second with longer inner and shorter outer

distal setae. Flagellum of four segments; first segment very

short, with proximal and distal rows of aesthetascs; second

segment 2.5 times as long as wide, with distal row of

aesthetascs; third segment as long as second, with simple

distal seta; fourth segment shorter than third, distally with two

penicillate and five simple setae, and single aesthetasc.

Antenna similar to that of female.

Mouthparts (Fig. 59C) largely reduced. Maxilliped basis

with single distal seta, endites naked, palp with simple setae

longer than those of female.

Cheliped (Fig. 59D) compact, more robust than that of

female; carpus just longer than wide, with longer ventral and

shorter dorsal single setae; propodus with inner comb-row of

14 setae; fixed finger with numerous fine spinules (“teeth”)

along cutting edge; dactylus strongly curved, with two setae

on inner margin.

Pereopods (Fig. 59E, F, G) more slender than those of

female, pereopods 2 and 3 similar to pereopod 1; posterior

pereopods with bases about 2.5 times as long as wide, propodi

nearly five times as long as wide and as long as carpus and

merus together.

Pleopods (Fig. 59H) similar to those of female, but setae

longer.

Uropod (Fig. 591) rami more slender than those of female;

exopod almost as long as proximal endopod segment; proximal

endopod segment with additional array of penicillate setae in

proximal half.

Remarks. The type material of Paratanais malignus was

collected in kelp epifauna at 4 to 4.5 m depth in Botany Bay,

New South Wales. The present material extends the known

distribution to throughout the Bass Strait in depths from 2 to

82 m on sandy to muddy substrata.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 58. Paratanais malignus, female. A, labrum; B, left mandible; C, maxilla and maxillule endite; D, maxilliped; E, cheliped with detail of chela

fingers; F, pereopod 1; G, pereopod 3; H, pleopod. Scale = 0.01 mm for A to D, 0.1 mm for E to H.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 59. Paratanais malignus, male. A, dorsal; B, antennule; C, mouthparts; D, cheliped; E, pereopod 1; F, pereopod 2; G, pereopod 4; H, pleopod;

I, uropod. Scale = 1 mm for A, 0.1 mm for B to I

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Paratanais tanyherpes sp. nov.

Figures 60-62

Material examined. 1 $ (J56762), holotype, Stn CPBS-N 32, Western

Port, off Crib Point, 30°20.83'S 145°13.48'E, 13 m depth, sandy gravel,

23 March 1965, coll. A J. Gilmour, Smith-Mclntyre Grab. 1 9 (J56763),

paratype, Stn CPBS-A 4, Western Port, off Crib Point, 38°21'S

145°14'E, 9 m depth, sand, 12 October 1964, coll. A.J. Gilmour, Smith-

Mclntyre Grab. 4 99 (J56755), paratypes, Stn CPBS-N 31, Western

Port, off Crib Point, 38°20.93'S 145°13.62'E, 15 m depth, fine sand and

mud, 29 March 1965, coll. A.J. Gilmour, Smith-Mclntyre Grab. 1 9

(J56757), paratype, Stn CPA 22, Cape Paterson, Victoria, 38°41'S

145°36'E, 0 m depth, 07 March 1982, coll. R.S. Wilson & H.M. Lew-

Ton. 2 99 (J56754), paratypes, Stn BSS 174 S, Eastern Bass Strait,

25 km NE of Deal Island, Tasmania, 39°16.8'S 147°33.2'E, 57 m depth,

medium sand, 18 November 1981, coll. R.S. Wilson, WHOI epibenthic

sled. 2 99 (J56761), paratypes, Stn BSS 155, Central Bass Strait, 38 km

SW of Cape Paterson, 38°55.5’S 147°17.0’E, 70 m depth, fine sand, 12

October 1964, coll. R.S. Wilson. 1 9 (J57533), paratype, Stn BSS 203,

Central Bass Strait, 44 km NE of Cape Wickham, King Island, 39°22'S

144°18.3'E, 60 m depth, coarse sand, 23 November 1981, coll. R.S.

Wilson. 1 9 (J56689), paratype, Stn BSS 199, Western Bass Strait, 20

km SSW of Stokes Point, King Island, 40°19.5'S 143°48.8'E, 71 m

depth, fine mud, 22 November 1981, coll. R.S. Wilson. 3 99 (J56722),

paratypes, Stn BSS 192 DRC, Western Bass Strait, 44 km SW of Cape

Otway, 39°06.7'S 143°07.4'E, 81 m depth, medium sand, 21 November

1981, coll. R.S. Wilson. 499(156705), paratypes, Stn BSS 209, Eastern

Bass Strait, 40 km SSW of Lakes Entrance, 38°18.0'S 147°37.0'E, 55 m

depth, muddy fine shell, 31 July 1983, coll. M.F. Gomon & R.S.

Wilson. 3 99 (J56764), paratypes, Stn BSS 173, Eastern Bass Strait, 30

km north of North Point Flinders Island, 39°26.3'S 147°48.7'E, 49 m

depth, medium sand, 17 November 1981, coll. R.S. Wilson. 1 9

(J56756), paratype, Stn CPBS-S 32, Western Port, off Crib Point,

38°21.6'S 145°13.7'E, 13 m depth, muddy sand, 12 March 1965, coll.

A.J. Gilmour. 1 9 (J56765), 2 99 (J56758), 1 9 (J56759), paratypes, Stn

CPBS-N 32, Western Port, off Crib Point, 38°20.83'S 145°13.48'E,

13 m depth, sandy gravel, 23 March 1965, coll. A.J. Gilmour, Smith-

Mclntyre Grab. 1 9 (J57542), paratype, Stn CPBS-S 26, Western Port,

off Crib Point, 38°22.18'S 145°15.22’E, 10 m depth, sand, 26 February

1965, coll. A.J. Gilmour, Smith-Mclntyre Grab. 1 9 (J57563), paratype,

Stn SPPS 5, Southern Port Phillip Bay, 38°17.3'S 144°41.4'E to

38°16.4'S 144°41.8'E, 7 m depth, coll. Marine Research Group of

Victoria, dredge.

Description of female. Body (Fig. 60A) elongate, 8.7 times as

long as wide, holotype 3 mm long. Cephalothorax subrectangular,

1.4 times as long as wide, twice as long as pereonite 1, with slight

triangular rostrum, naked. Pereonite 1 shortest, pereonites 2 and

3 about 1.5 times as long as pereonite 1; pereonite 4 longest, 1.9

times as long as pereonite 1; pereonites 5 just shorter than

pereonite 4, pereonite 6 just shorter than pereonite 2 (all

pereonites respectively 1.5,1.1, 1.0,0.9,0.9 and 1.1 times as wide

as long). Pleonites 3.6 times as wide as long, pleonites 1 to 4 with

one plumose, articulating lateral seta on each side. Pleotelson

semicircular, short, twice as long as pleonite 5, 1.7 times as wide

as long, distally with four posterolateral setae, marginal single

simple setae either side of each uropod attachment, and marginal

penicillate seta distal of uropod attachment (Fig. 621).

Antennule (Fig. 61A) of five articles, shorter than

cephalothorax; proximal article 1.6 times as long as wide, with

inner tufts of penicillate setae and short outer-distal simple seta;

second article wider than long, about one-third length of first,

with inner distal tuft of penicillate setae and simple seta longer

than article width; third article two-thirds length of second with

inner and outer distal setae; fourth article slender, tapering, almost

as long as second and third articles together, with one distal seta;

distal article minute, with four distal setae and single aesthetasc.

Antenna (Fig. 61B) proximal article compact, naked;

second article just shorter than wide, ventral margin produced,

without distal apophyses, with mid-ventral seta, shorter

laterodistal and dorsodistal seta, dorsal margin densely

setulose; third article two-thirds as long as wide, shorter than

second article, dorsal margin convex, with stout dorsodistal

spine; fourth article just longer than second, with two distal

simple setae and mesial and distal penicillate setae; fifth

article half as long as fourth with one distal seta; sixth article

minute with one very short and four longer distal setae.

Labrum (Fig. 61C) hood-shaped, apically rounded, setose.

Left mandible (Fig. 61D) with wide, crenulate lacinia mobilis,

right mandible (Fig. 61E) pars incisiva bilobate; left pars

molaris (Fig. 61D') robust distally with both sharp, slender and

short, rounded “teeth”. Labium (Fig. 61G) simple, densely and

finely setose, without palp. Maxillule (Fig. 61F) with nine

distal spines, rows of outer and inner setae on endite, palp not

retrieved. Maxilla (Fig. 61F) linguiform, naked. Maxilliped

(Fig. 61H) endites characteristic of genus, with denticulate

outer margin, two ovate spines and long single inner seta; palp

first article with outer-distal seta; second article with two

simple setae and one shorter denticulate, reflexed spine, distal

rows of microtrichia; third article with three marginal and one

submarginal inner, finely denticulate setae; fourth article with

five distal finely-denticulate setae and fine inner setules; single

distal seta on basis comfortably exceeding distal margin of

first palp article but not exceeding distal margin of endites.

Cheliped (Fig. 62A) compact, basis 2.1 times as long as

wide, with dorsodistal seta; merus triangular, occupying

almost all of ventral margin of carpus, with single mid-ventral

seta; carpus 1.75 times as long as wide, with proximal and

distal dorsal setae and two longer ventrodistal setae; propodus

1.25 times as long as wide, fixed finger short, half as long as

body of propodus (“palm”), with two ventral setae and three

setae alongside cutting edge; with lamellate apophyses on

cutting edge, terminal spine robust; dactylus with outer margin

smooth, two proximal simple setae on cutting edge.

Pereopod 1 (Fig. 62B) longer than others, coxa simple with

long seta; basis slender, arcuate, 4.15 times as long as wide,

with dorsal seta in proximal third; ischium compact with

single seta; merus slender, with single ventrodistal seta; carpus

0.8 times as long as merus, with one longer and two shorter

dorsodistal setae; propodus 1.75 times as long as carpus, with

two subdistal setae and fine dorsodistal sharp apophysis;

dactylus with dorsoproximal seta exceeding tip of dactylus;

unguis slender, curved, twice as long as dactylus, both together

1.1 times as long as propodus. Pereopod 2 (Fig. 62C) more

compact than pereopod 1, basis 2.9 times as long as wide;

merus 0.7 times as long as carpus, merus with two ventrodistal

setae; carpus with one dorsodistal seta, and one dorsodistal

and two ventrodistal curved, finely denticulate spines;

propodus twice as long as carpus, and just longer than dactylus

plus unguis. Pereopod 3 (Fig. 62D) similar to pereopod 2.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 60. Paratanais tanyherpes sp. nov., female holotype. A, dorsal view; B, lateral view. Scale = 1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 61 .Paratanais tanyherpes sp. nov., female paratype. A, antennule; B, antenna (proximal article not shown); C, labrum; D, left mandible, with D',

detail of molar process; E, right mandible; F, maxillule endite and maxilla; G, labium; H, maxilliped. Scale = 0.2 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 62. Paratanais tanyherpes sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5;

G, pereopod 6; H, pleopod; I, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Pereopod 4 (Fig. 62E) slightly more robust than anterior

pereopods, basis 2.8 times as long as wide with two mid-

ventral penicillate setae; merus and carpus subequal in length,

each with two ventrodistal curved, finely-denticulate spines,

carpus also with two dorsodistal curved, finely-denticulate

spines, ventrodistal surface of carpus with rows of microtrichia;

propodus 1.4 times as long as carpus, with simple dorsodistal

seta, mid-dorsal penicillate seta, two ventrodistal curved,

finely-denticulate spines, and row of fine setules along distal

margin; dactylus stout, with microtrichia; unguis distinct,

curved, with minute ventrodistal denticulation, half length of

dactylus, both together 0.6 times as long as propodus. Pereopod

5 (Fig. 62F) as pereopod 4. Pereopod 6 (Fig. 62G) as pereopod

4, but propodus with three dorsodistal serrate spines and no

dorsal penicillate seta.

Pleopods (Fig. 62H) all alike, with naked basis, endopod

with single inner subdistal plumose seta; exopod without setae

on inner margin, proximal seta on outer margin of both rami

separated from remaining setae.

Uropod (Fig. 621) basis naked, exopod of one segment,

over half as long as endopod, with one mesial and two unequal

distal setae; endopod of one segment, with mesial and distal

tufts of penicillate and simple setae; rami slender.

Male. Unknown.

Etymology. From the Greek tanaos - long and herpes - a

creeping thing, the present species being the most elongate yet

known in the genus.

Remarks. There are only two described species of Paratanais

with a single-segmented uropod endopod, viz. P. intermedius

Dojiri & Sieg, 1997 (q.v.) from 98 to 591 m depth in the Santa

Maria Basin, California, and P. wanga Bamber, 2008 from 4 to

26 m in Moreton Bay, Queensland. Although the description

and figures of P. intermedius are poor, it is distinct from

Paratanais tanyherpes sp. nov. in having a more elongate

antenna (peduncle articles 2 and 3 clearly longer than wide), the

distal seta on the maxilliped basis not reaching the distal margin

of first palp article, the merus of the cheliped covering only

some 75% of the ventral margin of the carpus, the merus on

pereopod 1 shorter than the carpus, and the uropod rami much

less elongate. P. wanga is distinct from P. tanyherpes in having

the second antennal peduncle article 1.4 times as long as wide,

and again the uropod rami much less elongate. In many respects,

P. tanyherpes is very close in morphology to the New Zealand

species P. tara Bird, 2011, with which it shares the proportions

of the antennule, the setation/spination of the antennule, of

many details of the maxilliped, but that species differs in having

a two-articled uropod endopod ( inter alia).

Paratanais tanyherpes is further distinguished from these

three species, and indeed all others in the genus, in being very

elongate, the body being nearly nine times as long as wide

(“about 6” times in P. intermedius ; 6.4 times in P. wanga ; up

to 7.5 times in P. tara', eight times as long as wide in the

previously most-elongate species, P. martinsi Bamber &

Costa, 2009, q.v.). P. tanyherpes was found throughout the

Bass Strait, including in Western Port and Port Phillip Bay, at

depths between 0 to 81 m.

Paratanais vetinari Bamber, 2005

Figures 63-64

Paratanais vetinari Bamber, 2005, 712-716, figs 51-52.

Material examined. 1 8 (J58466), 5 99 (J56735), Stn 190, 81-T-l,

Western Bass Strait, 50 km SSW of Warrnambool, Victoria, 38°49.5'S

142°35.4'E, 89 m depth, coarse sand, 21 November 1981, coll. R.S.

Wilson. 1 9 (J56666), Stn WV5, Cheviot Beach, Point Nepean,

Victoria, 38°18'S 144°40'E, 3.5 to 5 m depth, 31 March 1998, SCUBA,

coll. T,D. O’Hara. 1 9 (J56663), Stn BUN4, East of Eagles Nest,

Victoria, 38°40.46’S 145°39.14'E, 5 to 11 m depth, 01 April 1997,

SCUBA, coll. coll. T,D. O’Hara et al.. 1 9 (J46369), Stn WV2,

Schomberg Reef, near Peterborough, Victoria, 38°36.49'S 142°53.19'E,

3.5 to 5 m depth, 19 May 1998, SCUBA, coll. T,D. O’Hara. 3 99

(J51620), Stn VC 08 Cl, Western Bass Strait, 38°14.36'S 142 o 10.H'E,

40 m depth, 14 May 1998, Smith-Mclntyre Grab, coll. N. Coleman. 1

9 (J46370), Stn CRUST 21, “The Whaleback”, bommie 0.5 km S of

Point Hicks, 37°48.30'S 149°16.48'E, 13 m depth, 08 April 1989,

SCUBA, coll. G.C.B. Poore. 1 9 (J56665), 1 9 (J56662), Stn WV13,

Sailor’s Grave, off East Cape Conran, Victoria, 37°48.13'S 148 0 44.41'E,

4 to 5 m depth, 15 April 1998, SCUBA, coll. T,D. O’Hara. 19 (J56664),

Stn WV8, Nepean Bay, Point Nepean, Victoria, 38°18.24'S 144°39.28'E,

5 to 6 m depth, 08 April 1998, SCUBA, coll. T.D. O’Hara. Numerous

other lots in the collections of Museum Victoria.

Paratanais vetinari was described initially from Esperance,

southern Western Australia, based on four females. The present

females seem to agree with the type-description; the presence

of the male in the Bass Strait material allows its first description.

Description of male. Body (Fig. 63A, B) smaller and more

compact than that of female, 1.8 mm long, 5 times as long as

wide. Cephalothorax narrow in anterior half, 1.15 times as long

as wide, longer than pereonites 1, 2 and 3 together, with

conspicuous triangular rostrum; eyes present, large, pigmented.

Six free dorsoventrally-flattened pereonites; pereonite 1

shortest, pereonites 2 just longer than pereonite 1, both short

and with single anterolateral seta on each side; pereonites 3 to

6 subequal in length (all pereonites respectively 4.4, 3.7, 2.4,

2.4, 2.2 and 2.1 times as wide as long). Pleon laterally convex,

with five free subequal pleonites bearing pleopods; pleonites

4.6 times as wide as long. Pleotelson semicircular, short, les

than twice as long as pleonite 5, 2.2 times as wide as long.

Antennule (Fig. 63C) peduncle of three articles, proximal

article twice as long as wide, second article 0.6 times as long

as wide, about one-third length of first, with inner simple seta

longer than article width; third article half length of second

with inner and outer distal setae; flagellum of seven segments,

segments progressively longer distally, segments 1 to 6 with

subdistal rows of 8 or 9 aesthetascs, distal segment with one

penicillate and four simple distal setae.

Antenna (Fig. 63D) of six articles, proximal article

compact, naked; second article shorter than wide with single

simple distal and mesial setae; third article as long as wide,

about half as long as second article, naked; fourth article 1.25

times as long as second; fifth article longer than fourth,

slender, tapering; sixth article minute with five distal setae.

Mouthparts largely atrophied; maxilla (Fig. 63F)

subtriangular, naked; maxilliped (Fig. 63E) endites narrow,

with single distal seta, basis with long distal seta, palp with

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 63. Paratanais vetinari, male. A, lateral view; B, dorsal view; C, antennule; D, antenna; E, maxilliped; F, maxilla; G, epignath. Scale A-B = 0.5 mm;

C-G = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 64. Paratanais vetinari, male. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5; G, pereopod 6;

H, pleopod; I, uropod. Scale = 0.1 mm.

M. Blazewicz-Paszkowycz & R.N. Bamber

simple setae much longer than those of female; epignath (Fig.

63 G) narrow, linguiform.

Cheliped (Fig. 64A) similar to that of female, merus

shorter, carpus 1.3 times as long as wide; propodus

proportionately longer, fixed finger with triangular tooth-like

apophysis proximally on cutting edge; dactylus with three

simple setae on cutting edge.

Pereopods 1 to 3 (Fig. 64B, C, D) similar to those of

female, merus of pereopod 1 1.3 times as long as carpus, distal

spines on carpi of pereopods 2 and 3 simple and more slender.

Pereopods 4 to 6 (Fig. 64E, F, G) proximally similar to

those of female, but propodus 6 times as long as wide with

dense ventral fields of microtrichia, dactylus and unguis two-

thirds as long as propodus, articulation between the two

indistinct, dactylus with fields of microtrichia.

Pleopods (Fig. 64H) all alike, similar to those of female

but setae proportionately longer.

Uropod (Fig. 641) similar to but more elongate than that of

female, distal exopod segment longer than proximal segment;

proximal endopod segment with inner array of penicillate setae.

Remarks. Of the seven previously-described species of

Paratanais from Australasia (including P. tanyherpes above),

only P. vetinari and P. maleficus Larsen, 2001 have a two-

segmented uropod exopod. Females of these two are most easily

distinguished by the uropod segments being more than twice as

long as wide in P. vetinari, but less than 1.5 times as long as

wide in P. maleficus. Most features of the appendages of the

male of P. maleficus as described and figured by Larsen (2001)

are different from those shown here. Another characteristic

feature of the present species is the subdistal ventral seta on the

propodus of pereopods 2 and 3 being longer than the dactylus

(shorter in P. maleficus ). The only other species described with

a two-segmented uropod exopod is P. hessleri Kudinova-

Pasternak, 1985 (q.v.), but, despite the relatively poor description,

that species has the subdistal ventral propodus seta on pereopods

2 and 3 shorter than the dactylus, and it clearly has a more

elongate body, antennules and antennae, inter alia.

The type-collection of Paratanais vetinari was from 20 to

30 m depth on gravelly sand with rhodoliths and sparse

macroalgae. In the Bass Strait, this species was collected from

between 3.5 and 89 m depth on sandy substrata, including

sympatrically with P. malignus.

Subfamily Bathytanaidinae Larsen & Heard, 2001

Genus Bathytanais Beddard, 1886

Bathytanais bathybrotes (Beddard, 1886)

Figures 65-67

Paratanais bathybrotes Beddard, 1886(a), 119 - Bathytanais

bathybrotes Bamber, 2008, 175-176, literature.

Material examined. 12 specimens (including 2 brooding $$) (J56271),

1 3 (J56272), from sand wall at front of Pope’s Eye, Port Philip Bay,

7 m depth, 28 February 1982, coll. R Lipson.

The female of this species was comprehensively redescribed by

Lang (1972); however, the male was previously unknown -

indeed, no male of any Bathytanais species has been recorded

before, making the following description particularly important.

Description of male. Body (Fig. 65A, B) with typical gross

appearance of a Paratanais male, 2.1 mm long, 5.3 times as

long as wide. Cephalothorax subrectangular, as long as wide, as

long as pereonites 1 to 3 together, with conspicuous

subtriangular rostrum; eyes large, one-third as long as

cephalothorax, with black ommatidia. Pereonite 1 shortest,

pereonites 2, and 3 subequal in length, 1.4 times as long as

pereonite 1, pereonites 4 and 5 subequal, 1.8 times as long as

pereonite 1, pereonite 6 just shorter than pereonite 5 (all

pereonites respectively 3.6, 2.7, 2.8, 2, 2 and 2.3 times as wide

as long). Pleon of five free subequal pleonites bearing pleopods,

ventrally with blunt keel; pleonites 2.7 times as wide as long

and 1.2 times as long as pereonite 1. Pleotelson (Fig. 47K)

semicircular, as long as pereonite 6, 1.5 times as wide as long.

Antennule (Fig. 66A) of three peduncular and four flagellar

articles, proximal peduncle article 1.9 times as long as wide, with

outer mesial tufts of penicillate setae, inner distal simple seta;

second article just shorter than wide, 0.4 times as long as first

article, with outer-distal tuft of penicillate setae; third article

one-third length of second, with two simple distal setae; first

flagellar article half as long as third peduncle article, with

proximal and distal rows of six or seven aesthetascs and outer-

distal simple seta; second and third flagellar articles subequal in

length, four times as long as first, with distal rows of six

aesthetascs; fourth flagellar article just shorter than third, distally

with four simple setae, one penicillate seta and one aesthetasc.

Antenna (Fig. 66B) of six articles, proximal article

compact, naked; second article with ventrodistal tuft of three

penicillate setae and one ventrodistal and one dorsodistal

simple setae; third article as long as wide, 0.6 times as long as

second article, with dorsodistal seta; fourth article twice as

long as third, with one mid-dorsal and three dorsodistal

penicillate setae, single dorsal and ventral subdistal simple

setae; fifth article 0.9 times as long as fourth with one dorsal

subdistal seta; sixth article minute with five simple and one

penicillate distal setae.

Mouth parts underdeveloped in comparison with those of

female. Labrum (Fig. 66C) apically rounded, naked. Mandibles

absent. Maxillule (Fig. 66D) with naked endite and simple

palp bearing distal setule; maxilla (Fig. 66D) ovoid, naked.

Maxilliped (Fig. 66E) endites relatively wide, with two minute

ovate tubercles and single inner seta; palp first article with

outer-distal seta; second article with three inner-distal simple

setae; third article with three inner setae in proximal half;

fourth article with five inner to distal setae and one outer

subdistal seta; single inner seta on basis not reaching distal

margin of endites.

Cheliped (Fig. 66F) compact, basis twice as long as wide,

naked; merus subtriangular, ventrally convex, with one mid-

ventral seta; carpus 1.3 times as long as wide with two mid-

ventral setae, one dorsoproximal and one dorsodistal fine

setae; propodus as long as wide, with inner comb-row of 12

setae; fixed finger just shorter than palm, with two ventral

setae, three setae adjacent to cutting edge; dactylus with

dorsoproximal seta, two proximal setae on cutting edge.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 65. Bathytanais bathybrotes, male. A, lateral; B, dorsal. Scale = 0.2 mm.

100

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 66. Bathytanais bathybrotes , male. A, antennule; B, antenna; C, labrum; D, maxillule and maxilla; E, maxilliped; F, cheliped. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

101

Fig. 67. Bathytanais bathybrotes, male. A, pereopod 1; B, pereopod 2; C, pereopod 3; D, pereopod 4; E, pereopod 5; F, pereopod 6; G, pleopod;

H, uropod. Scale = 0.1 mm.

102

M. Biazewicz-Paszkowycz & R.N. Bamber

Pereopod 1 (Fig. 67A) longer than others, coxa simple with

seta; basis slender, 4.25 times as long as wide with dorsoproximal

penicillate seta; ischium compact with single seta; merus one-

third as long as basis, with fine ventrodistal seta; carpus 0.9 times

as long as merus, with fine ventrodistal seta and two dorsodistal

setae; propodus 1.8 times as long as carpus, with one dorsal and

two ventral subdistal setae, and dorsal and ventral marginal

microtrichia; dactylus half as long as slender unguis and with

distal seta, dactylus and unguis together 0.9 times as long as

propodus. Pereopod 2 (Fig. 67B) similar to pereopod 1, but more

compact, basis 3.4 times as long as wide; merus and carpus equal

in length, carpus without dorsodistal setae; propodus with two

dorsal and one ventral subdistal setae, dactylus with proximal

seta. Pereopod 3 (Fig. 67C) similar to pereopod 2.

Pereopod 4 (Fig. 67D) basis stout, 2.6 times as long as wide,

with three ventral subdistal penicillate setae; ischium with two

ventral setae; merus one-third as long as basis, naked; carpus as

long as merus, with two ventrodistal and one dorsodistal tooth¬

like spines; propodus 1.7 times as long as carpus with dorsodistal

seta, two ventrodistal spines, mid-distal penicillate seta and

marginal rows of penicillate setae; dactylus long and slender,

four times as long as distinct unguis, both together 0.8 times as

long as propodus. Pereopod 5 (Fig. 67E) similar to pereopod 4,

but basis with dorsoproximal pair of penicillate setae, merus

with two ventrodistal tooth-like spines, carpus with additional

larger outer distal spine and dorsodistal seta, dactylus and unguis

distinctly more curved. Pereopod 6 (Fig. 67F) as pereopod 5.

Pleopods (Fig. 67G) all alike, with naked basis, rami

subequal; endopod with inner subdistal plumose seta and 13

plumose setae around outer margin; exopod without setae on

inner margin, 19 plumose setae around outer margin.

Uropod (Fig. 67H) basis naked; exopod of two subequal

segments, together more than half as long as endopod, first

segment with simple outer-distal seta, second segment with two

unequal distal simple setae; endopod of three segments,

proximal segment short, naked, second segment twice as long as

first, with proximal array of seven penicillate setae, distally with

one simple and one penicillate setae; third segment as long as

second, distally with five simple and one penicillate setae.

Remarks. The history of this species was discussed by Bamber

(2008), from which we may disregard the type-locality as being

a lapsus calami. Bathytanais bathybrotes was previously

known from New South Wales at depths of 6 to 50 m depth

(Beddard, 1886b; Lang, 1972) and from Moreton Bay,

Queensland at depths between 8 and 42 m (Bamber, 2008) on

clean sand. The present material extends its distribution further

west to Victoria.

The Bathytanais male is very similar to males of

Paratanais species, and does not show the expanded antenna

peduncle articles of the females, but appears to differ in the

presence of a proximal seta on the dactyli of pereopods 2 and

3. Discovery of the males of other species is necessary to

confirm this as a generic character.

Bathytanais fragilis Larsen & Heard, 2001

Bathytanais fragilis Larsen & Heard, 2001, 13-16, figs 5-7

(partim: deeper-water specimens only).

Material examined. 1 ? (J37854), holotype. Slope 21, 36°57.40'S

150°18.80'E, 220 m depth, muddy shell, 20 July 1986. 3 $? (J58583),

paratypes (“other material” in Larsen & Heard, 2001), same data as

holotype. 1 ? (J39282), idiotype. Slope 32, 38°21.90'S 149°20.00'E,

1000 m depth, fine mud, 23 July 1986. All coll. G.C. Poore et al .,

WHOI suprabenthic sled, R.V. Franklin.

Remarks. In the light of other Bathytanais material found in

the collections, the type-material of B. fragilis was re¬

examined, and some specimens (from shallower samples) were

separated as belonging to a distinct species, described below.

Significantly, we were able to confirm the shape and, in

particular, the transparency of the ventral expansion on the

second article of the antenna peduncle in B. fragilis, as well as

other features of its morphology.

Bathytanais fragilis is recorded from the Eastern Bass

Strait, from depths between 200 and 1000 m, mainly on muddy

substrata.

Bathytanais parageios sp. nov.

Figures 68-70

Bathytanais fragilis Larsen & Heard, 2001 partim (shallow-water

specimens only).

Material examined. 1 9 (J58584), holotype, MAFRI-H28 GI, Long

Island Hastings, 38°19'S 145°14'E, 17 m depth, 4 March 1997; 6 99

(J66538), paratypes, MAFRI-H27 G4, Crib Point, 38°21'S 145°14'E,

12 m depth, 4 March 1997; 4 99 (J66579), paratypes, MAFRI-H28 GI,

Long Island Hastings, 38°19’S 145°14’E, 17 m depth, 4 March 1997; all

coll. DPI Victoria, Smith McIntyre grab. 1 $ (J56781), paratype, WBES

stn 1723, Western Port, 38°17.07’S 145°14.86'E, 14 m depth, sand, 22

November 1973, coll. Marine Studies group. Smith McIntyre grab. 3 9

9 (J51621), paratypes, VC-34-C1, 38°32.42'S 146°29.68'E, 40 m depth,

fine mud, 11 May 1998, coll. N Coleman, Smith McIntyre grab. 22

specimens (incl. 1 brooding 9 ), MSL-EG 70, 37°53.39’S 148°15.40'E,

43 m, coarse sand, 4 June 1991, coll. N Coleman, Smith McIntyre grab

(cited as paratypes of B. fragilis under a different registration no. in

Larsen & Heard, 2001). 1 $ (J58465), paratype (dissected and figured),

CR 81-T-l stn 155, 38°55.5’S 145°17.0’E, 70 m depth, fine sand, 12

November 1981, coll. R.S. Wilson, Smith McIntyre grab. 1 9 (J55908),

paratype, CPBS 31N/3, Crib Point, 38°20.94'S 145°13.62’E, 15 m depth,

fine sand and mud, 29 March 1965, coll. Marine Studies group. Smith

McIntyre grab. 2 99 (J56676), paratype, CPBS 31N/4, same data as

previous. 3 99 (J23557), paratypes, MSL-EG 58, Eastern Bass Strait,

37°51.19'S 148°38.53'E, 51 m depth, mud and shell, 29 September 1990,

coll. Marine Studies group. Smith McIntyre grab, RV Sarda. 1 9

(J17283) and 1 9 (J17281), paratypes, MSL-EG 16, Eastern Bass Strait,

38°04'S 148°25.7'E, 28 m depth, sand and shell, 12 August 1989, coll. G

Parry, Smith McIntyre grab, RV Sarda.

Description of female. Body (Fig. 68) elongate, holotype

3.8 mm long, eight times as long as wide. Cephalothorax

subrectangular, as long as wide, shorter than pereonites 1 and

2 together, with rounded triangular rostrum, naked; eyes

present, pigmented. Pereonites cylindrical; pereonite 1

shortest, half length of cephalothorax; pereonite 2 longer, 0.75

times as long as cephalothorax; pereonite 3 longest, as long as

cephalothorax; pereonites 4 and 5 subequal, longer than

pereonite 2, pereonite 6 just shorter than pereonite 2 (all

pereonites respectively 1.9, 1.1, 0.8, 1.0, 0.9 and 1.1 times as

wide as long). Pleon of five free subequal pleonites bearing

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 68. Bathytanais parageios sp. nov., holotype female. A, dorsal view; B, lateral view. Scale = 0.1 mm.

104

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 69. Bathytanais parageios sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, left mandible; E, right mandible; F, maxillule

and maxilla G, maxilliped; G', maxilliped endite. Scale A-C = 0.1 mm; D-H= 0.01 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

105

Fig. 70. Bathytanais parageios sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 6;

G, pleopod; H, uropod. Scale = 0.1 mm.

106

M. Biazewicz-Paszkowycz & R.N. Bamber

pleopods plus pleotelson, the whole shorter than pereonites 5

and 6 together; pleonites 5 times as wide as long; each pleonite

with one plumose, articulating lateral seta on each side.

Pleotelson semicircular, short, 1.6 times as long as pleonite 5,

twice as wide as long, distally with two posterolateral simple

setae on each side.

Antennule (Fig. 69A) of four articles, proximal article

twice as long as wide, naked; second article as long as wide,

about one-third length of first, distally with two simple and

three penicillate setae; third article two-thirds length of

second with two simple distal setae; fourth article slender, as

long as second article, with five setulose distal setae, longest

three longer than cephalothorax and antennule together, but

without aesthetasc (possibly a generic character: G Bird,

pers. comm.).

Antenna (Fig. 69B) of six articles, proximal article

compact, naked; second article dorsally with slight flange-like

expansion bearing single simple distal seta, ventral margin

with three distal setae and laminar apophysis (flange)

exceeding distal edge of third peduncle article, distally

rounded, not transparent, with long mid-distal seta; third

article just shorter than wide, half as long as second article,

expanded mid-dorsally with mid-distal spine; fourth article

half as long as second, distally with one simple seta, two

penicillate setae and long dorsal setulose seta 3.5 times as long

as article; fifth article half as long as fourth with two long

setulose distal setae; sixth article minute with two long

setulose distal setae.

Labrum (Fig. 69C) apically rounded, setose. Left mandible

(Fig. 69D) with wide, curved, crenulate lacinia mobilis, right

mandible (Fig. 69E) without lacinia mobilis; pars incisiva

bilobate, pars molaris robust, distally flat with marginal

rugosity extended as short, rounded “teeth” ventrally. Labium

(Fig. 47G) rounded, finely setose with microtrichia on inner

face, outer margins finely denticulate. Maxillule (Fig. 69F)

with seven distal spines and sparse outer setules, palp slender

with two distal setae. Maxilla (Fig. 69F) oval, naked.

Maxilliped (Fig. 69H) with single inner spine on basis reaching

distal margin of second palp article; palp first article naked,

second article with two simple setae and shorter denticulate

seta on inner margin, outer margin with one simple seta; third

article with three denticulate setae on inner margin; fourth

article with five submarginal simple ventral setae, four distal

denticulate setae and one mid-dorsal denticulate seta; endites

(Fig. 69H') characteristic of genus, wider than maxilliped

bases, with denticulate outer corner, two elongate ovate spines

and single inner slender spine. Epignath (Fig. 691) slender,

curved, distally setose.

Cheliped (Fig. 70A) basis 1.8 times as long as wide, naked;

merus with one mid-ventral seta; carpus 1.6 times as long as

wide, ventral margin convex with two mid-ventral setae,

dorsally with proximal and distal simple setae; propodus

longer than wide, distally with comb of five shorter setae on

inner face and three longer setulose setae, fixed finger short

with lamellate apophyses on cutting edge, one longer and one

shorter ventral setae, one longer and two shorter simple setae

distally below cutting edge; dactylus with outer margin smooth

and bearing one setulose seta.

Pereopod 1 (Fig. 70B) longer than others, coxa simple with

seta; basis slender, arcuate, 4.4 times as long as wide with

three dorsal setae in proximal half; ischium compact with

single seta; merus just over half length of basis, with

ventrodistal seta; carpus 0.6 times as long as merus with one

ventrodistal and two dorsodistal setae; propodus as long as

merus, with two distal setae; dactylus curved, half as long as

propodus, unguis slender, twice as long as dactylus. Pereopod

2 (Fig. 70C) more compact, basis 4 times as long as wide;

merus 0.8 times as long as basis, with two ventrodistal setae;

carpus 1.3 times as long as merus, with slight dorsal expansion

and single dorsal and ventral distal setae; propodus twice as

long as carpus, with two distal setae; dactylus with long

dorsoproximal seta, one-third length of propodus and half

length of slender, curved unguis. Pereopod 3 (Fig. 70D) similar

to pereopod 2, but dactylus without dorsal seta.

Pereopod 4 (Fig. 70E) coxa not evident; basis stouter, 2.3

times as long as wide, with three simple dorsal setae and two

ventrodistal penicillate setae; ischium with two ventrodistal

setae; merus half as long as carpus, each with ventrodistal

microtrichia and rugosity and molar spines, carpus with

dorsodistal seta; propodus as long as carpus, with mid-dorsal

penicillate seta, dorsodistal spine exceeding claw, inner and

outer ventrodistal molar spines; dactylus and claw fused into

unguis, curved, bearing microtrichia, half as long as propodus.

Pereopod 5 (not figured) as pereopod 4 but without dorsal

penicillate seta on propodus. Pereopod 6 (Fig. 70F) similar to

pereopod 4, but basis without penicillate setae, merus with

triangular dorsodistal spine, propodus with three distal serrate

spines adjacent to unguis.

Pleopods (Fig. 70G) all alike, with naked basis; endopod

shorter than exopod, with single inner-subdistal plumose seta,

exopod without setae on inner margin; outer margins of

endopod and exopod with 16 and 23 plumose setae respectively,

on both rami proximal seta on outer margin separated from

remaining setae.

Uropod (Fig. 70H) basis with one outer simple seta. Rami

slender, exopod of one segment exceeding proximal segment

of endopod, with one mesial and two distal simple setae,

endopod of two segments, proximal segment with two

penicillate and one simple distal setae, distal segment with

two penicillate and five simple distal setae.

Male. Unknown.

Etymology. From the Greek parageios - pertaining to shallow

water.

Remarks. Bathytanaisparageios sp. nov. is close in morphology

and locality to B. fragilis, but there are consistent differences.

B. fragilis is characterized, inter alia, by the expansion on the

second peduncle article of the antenna being transparent

(confirmed in the examination of the types, see above), and

somewhat wider proximally than distally, with two medial

setae and no dorsal seta on the article; in the present species,

this article has a slight dorsal expansion with a seta, and the

ventral expansion is of uniform width throughout, with three

medial setae, and quite opaque. The second peduncle article is

also subtly different, that of B. fragilis having a forward-

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

107

pointing spine-like apophysis bearing a seta, while that of

B. parageios has a stout spine. Further, the third article of the

antennule of B. fragilis has a seta some five times as long as the

fourth antennular article, this seta being less than twice as long

in B. parageios.

While these differences are sufficiently subtle to suggest

variability with depth in one species, other notable differences

are found in the uropod segmentation, B. fragilis having a one-

segmented endopod some three times as long as wide, while

that of B. parageios is two-segmented, and conspicuously

more slender (as is the exopod) at five times as long as wide.

Further differences are found in the spination of the posterior

pereopods (especially pereopod 6), the setation of the

maxilliped basis and endite, the proportionately shorter second

and third articles of the antennular peduncle, the

proportionately shorter fifth article of the peduncle of the

antenna (half as long as the fourth in B. parageios , subequal in

length in B. fragilis), and the more numerous plumose setae on

the rami of the pleopods, and the more gracile posterior

pereopods in B. parageios. No variation between these

morphologies was observed within the material examined.

These two taxa are thus considered sibling species, close

in morphology but showing a number of distinctions, and

separated by depth and habitat ( B. parageios on shallower

sandy sediments, B. fragilis on deeper muds), this niche-

specificity presumably having lead to allopatric speciation.

Their antennal morphology distinguishes them entirely from

all other described Bathytanais species.

Bathytanais parageios was collected from throughout the

Bass Strait, from depths between 12 and 70 m, mainly on

sandy substrata.

Family Leptocheliidae Lang, 1973

Genus Leptochelia Dana, 1849

Leptochelia billambi sp. nov.

Figures 71-74

Material examined. 1 ? (J58472), holotype, 1 3 (J58474), allotype, 162

$? (J58473), paratypes, sample Stn CPBS 000/4, 38°21.17'S

145°13.15’E, 2 m depth, sandy mud, 6 April 1965, coll. MSG, Smith

McIntyre grab.

Portland Exotic Species survey material: 31 $? (J66139) at 3 m

depth, 9 9 (J66142) at 0.5 m depth, 16 specimens (J66139) at 7 m depth,

paratypes, MAFRI-P17 SCR, Portland No. 6 berth, 38°21'S 147°37'E,

3 May 1996. 2 99 (J66126) at 7 m depth, 4 99 (J66123) at 3 m depth,

paratypes, MAFRI-P16 SCR, Portland No. 2 berth, 38°20'S 147°37’E,

3 May 1996. 1 9 (J66094) at 3 m depth, paratypes, MAFRI-P15 SCR,

Portland SL Pattersons Berth, 38°21'S 147°37'E, 2 May 1996. 3 99

(J66156) at 3 m depth, 2 99 (J66158) at 0.5 m depth, paratypes,

MAFRI-P18 SCR, Portland No. 1 berth, 38°21’S 147°37'E, 4 May

1996. 1 <3 (J66386), paratype, MAFRI-M47-G47, Port of Melbourne,

37°50'S 144°53'E, 14 m depth, 7 December 1999.

Other material examined. A further 521 idiotypic individuals

from the Bass Strait, Western Port and Port Philip Bay, many without

registration numbers, and including 303 99 (9 brooding), 3 <3(3 and 38

juveniles. Numerous further specimens exist in the collections of

Museum Victoria.

Description of female. Body (Fig. 71A, B) slender, holotype

2.4 mm long, 6.4 times as long as wide. Cephalothorax

subrectangular, tapering towards anterior, 1.4 times as long as

wide, longer than pereonites 1 and 2 together, with slight

rostrum, eyelobes rounded, eyes present and black, single setae

at posterior of eyelobes and mid-laterally. Pereonites 1 and 6

subequal and shortest, pereonites 2 to 5 subequal, progressively

longer, pereonite 5 longest and 1.4 times as long as pereonite 1

(all pereonites respectively 1.9, 1.7,1.5, 1.5,1.4 and 1.9 times as

wide as long). Pleon with five free subequal pleonites bearing

pleopods; each pleonite about 5.6 times as wide as long, with 2

or 3 lateral setae. Pleotelson pentangular, as long as last two

pleonites together, twice as wide as long, with one anterolateral

and posterolateral seta on each side and two distal setae.

Antennule (Fig. 72A) of three longer and one minute distal

articles, proximal article 4 times as long as wide, 1.6 times as

long as distal three articles together, with proximal, mesial and

distal inner groups of penicillate setae, inner and outer simple

mesial setae and one long inner distal seta longer than second

article; second article twice as long as wide, one-third as long

as first article, distal seta just longer than article; third article

just shorter than second, with one distal seta; fourth article

minute, with three distal setae and one aesthetasc.

Antenna (Fig. 72B) of six articles, proximal article compact,

with fine distal seta; second article as long as wide, with single

inner distal and dorsodistal slender spines; third article just

longer than wide, with dorsodistal slender spine; fourth article

longest, 3.2 times as long as wide and twice as long as third,

with mid-length seta reaching two-thirds of length to distal

margin, and distal tufts of penicillate and simple setae; fifth

article 0.6 times as long as fourth; sixth article minute, with

four simple and one penicillate distal setae.

Labrum (Fig. 72C) hood-shaped, setose, typical of genus.

Left mandible (Fig. 72D) with crenulate lacinia mobilis wider

than distal end of mandible, distal crenulation on pars incisiva,

pars molaris with strong rugosity; right mandible (Fig. 72E)

similar but without lacinia mobilis, pars incisiva distally

bifurcate. Labium (Fig. 72G) wide, bilobed, distally finely

setose, without palp. Maxillule (Fig. 72F) with ten distal

spines and setose margins, rows of setules on inner distal face;

palp distinct, with two distal setae. Maxilliped (Fig. 72H) palp

first article naked, second article with finely setose inner

margin, and with one outer, one ventral and three inner setae,

distal-most inner seta not reaching distal margin of third palp

article; third article with nine inner marginal and one distal

submarginal finely denticulate setae; fourth article with nine

inner/distal finely denticulate setae and one outer subdistal

seta; basis with five long setae extending to third palp article;

endites distally with fine outer setules and three robust

spatulate spines, inner spine shorter than others.

Cheliped (Fig. 73A) with rounded, comparatively slender

basis 2.3 times as long as wide; merus triangular with three

ungrouped ventral setae; carpus twice as long as wide, with

two midventral setae and two shorter marginal dorsal setae;

propodus slightly longer than wide, with inner distal comb of

seven setae and longer seta at base of dactylus; fixed finger

with four ventral and three inner setae, cutting edge crenulate;

dactylus naked.

108

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. ll.Leptochelia billambi sp. nov., female holotype. A, dorsal view; B, lateral view; C, male lateral view. Scale = 0.2 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

109

Fig. 72. Leptochelia billambi sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, left mandible; E, right mandible; F, maxillule;

G, labium; H, maxilliped. Scale = 0.2 mm.

110

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 73. Leptochelia billambi sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5;

G, pereopod 6; H, pleopod; I, uropod. Scale = 0.2 mm.

* ttt

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

111

Fig. 74. Leptochelia billambi sp. nov., male paratype. A, antennule; B, antenna; C, cheliped; D, mouth parts; E, pereopod 1; F, pereopod 2;

G, pereopod 3; H, pereopod 4; I, pereopod 5; J, pereopod 6; K, pleopod; L, uropod. Scale = 0.2 mm.

112

M. Biazewicz-Paszkowycz & R.N. Bamber

Pereopod 1 (Fig. 73B) longer than other pereopods, coxa

with seta; basis slender, 4.4 times as long as wide, with single

dorsoproximal simple and penicillate setae; ischium compact

with one ventral seta; merus as long as carpus, with one

dorsodistal and two ventrodistal setae; carpus with two

ventrodistal setae, one medial distal seta and three dorsodistal

setae, longest of which is half length of propodus; propodus

1.8 times as long as carpus, with three setae on subdistal dorsal

mound and one subdistal ventral seta; dactylus slender,

extending into shorter slender unguis 0.7 times as long as

dactylus, the two together as long as propodus. Pereopod 2

(Fig. 73C) more compact than pereopod 1; coxa with longer

seta; basis 3.1 times as long as wide; ischium with 2 setae;

merus as long as carpus and 1.5 times as long as wide, with

strong ventrodistal spine and ventral rows of microtrichia;

carpus with single dorsodistal and ventrodistal setae and two

ventrodistal spines, and ventral rows of microtrichia; propodus

1.6 times as long as carpus, with three distal setae and ventral

rows of microtrichia; dactylus and short unguis curved,

together 0.7 times as long as propodus. Pereopod 3 (Fig. 73D)

similar to pereopod 2, but carpus shorter than merus and with

two dorsodistal setae.

Pereopod 4 (Fig. 73E) basis stout, 2.23 times as long as

wide with two ventroproximal penicillate setae; ischium with

two ventrodistal setae; merus with paired ventrodistal spines

and ventral rows of microtrichia; carpus just shorter than

merus, with outer, ventral and inner distal spines each with

fine subdistal setules, and ventral rows of microtrichia;

propodus 1.2 times as long as carpus, with two ventrodistal

spines, five dorsodistal setae mostly as long as dactylus, and

ventral rows of microtrichia; dactylus and distinct unguis

curved, 0.6 times as long as propodus. Pereopod 5 (Fig. 73F)

as pereopod 4, but carpus with additional simple distal seta,

propodus as long as carpus. Pereopod 6 (Fig. 73G) as pereopod

5, but propodus with seven finely denticulate and one simple

distal setae.

Pleopods (Fig. 73H) all alike, typical for the genus but

basis naked, endopod with single inner plumose seta and

proximal outer seta separated from remainder.

Uropod (Fig. 731) biramous, basis naked; exopod of one

segment, 0.8 times as long as proximal endopod segment,

outer distal seta longer than inner distal seta; endopod of five

segments, distal segments slender.

Description of male. Larger than female (allotype length

4.2 mm), body (Fig. 71C) more compact, cephalon as long as

pereonites 2 to 4 together, with large eyelobes bearing large

black eyes; pereonite 1 shortest, pereonites 2,3 and 6 subequal,

twice as long as pereonite 1, pereonite 4 longest, three times as

long as pereonite 1, pereonite 5 just shorter than pereonite 4.

Five free pleonites, subequal in length, each as long as pereonite

1, pleotelson as long as pleonites 5 and 6 together.

Antennule (Fig. 74A) elongate, first peduncle article 6.4

times as long as wide, curved, with single dorsodistal

penicillate and simple setae; second article 0.43 times as long

as first with single ventral and dorsal distal simple setae; third

article half as long as second and twice as long as wide with

single ventral and dorsal distal simple setae; flagellum of 9

segments, segments 1 to 7 bearing proximal row of 6 or 7

aesthetascs, segment 8 naked, segment 9 minute, distally with

six simple and one penicillate setae.

Antenna (Fig. 74B) proximal article compact; second

article as long as wide, with single ventrodistal seta; third

article longer than second, twice as long as wide, with single

simple dorsodistal seta; fourth article 3.3 times as long as

third, with incipient secondary articulation at mid-length

bearing single simple and penicillate setae, distally ventral

pairs of penicillate and simple setae, and stronger dorsal seta

reaching half length of fifth article; fifth article 0.9 times as

long as fourth with two longer and one shorter distal setae;

sixth article minute, with four simple distal setae.

Mouthparts (Fig. 74D) atrophied, naked maxilliped and

maxillule palp with two distal setae distinguishable.

Cheliped (Fig. 74C) larger and more slender than that of

female; basis 1.7 times as long as wide; merus short, with two

ventral setae; carpus three times as long as wide, with three

midventral setae and four dorsal marginal setae; propodus 1.4

times as long as wide, fixed finger 1.25 times as long as palm,

with six ventral setae, three setae adjacent to cutting edge,

two inner tooth-like apophyses on cutting edge; dactylus

slender, curved, with proximal seta and row of 13 setae along

cutting edge.

Pereopods (Fig. 74E to J) similar to but more slender than

those of female, merus shorter than carpus on pereopods 1 to

3; on pereopods 4 to 6 propodus much longer than carpus,

distal carpal spines more elongate than those of female,

microtrichia restricted to dactylus, dactylus proportionately

longer than in female.

Pleopods (Fig. 74K) with longer setae than on those of

female. Uropod (Fig. 74L) basis with inner distal row of setae;

exopod two-segmented, longer than proximal endopod

segment; endopod of five segments, more heavily setose than

that of female.

Etymology. Named after William Lamb, 2nd Viscount

Melbourne, after whom the city of Melbourne was named in

1837 (the original Melbourne being a village in Derbyshire,

England).

Remarks. Bamber (2008) gave an identification key to the

species of the Leptocheliidae then known from Australian

waters, through which the present species would key out to

Leptochelia opteros Bamber, 2008, the commonest species in

Moreton Bay, Queensland: it is indeed close to that species, and

shares with it a distal seta on the proximal article of the antenna;

however, L. billambi sp. nov. is a larger species, its cheliped and

antennule are more elongate than those of L. opteros, the fixed

finger of the chela has 4, not 3, ventral setae, the chela comb-

row has more setae, and the maxilliped basis has 5 distal setae

(4 in L. opteros). The male appears generally as a larger version

of that of L. opteros, but that male is characterized by having a

dorsodistal flange on the basis of pereopod 6, absent in the

present species. Finally, L. billambi was recorded on sandy

mud substrata, while L. opteros lives amongst sublittoral algae

and epifaunal communities. The incipient secondary

articulation of the fourth antennal peduncle article in the male

is so far unique to this species in the Leptocheliidae.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

113

A range of morphometric and meristic characters in

females has been analyzed for some 22 taxa attributable to the

Leptochelia savignyi (Kr0yer, 1842)-complex (see Bamber,

2008; Bamber, 2010), with which the present species has been

compared. The only other species which has the proximal

article of the antennule approaching as slender as 4 times as

long as wide is L. savignyi sensu stricto, ranging from 3.5 to

3.8 times as long as wide (Bamber, 2010), while in the figure

of L. savignyi sensu Sars (1886) from the Mediterranean this

article is 4 times as long as wide. However, that species has

only 4 maxilliped-basis setae (five in L. billambi ) and a

proportionately shorter cheliped basis and carpus in the adult,

while the distal seta on article 2 of the antennule is not longer

than the article itself (longer in L. billambi ), the uropod

exopod is only just longer than half of the proximal endopod-

segment length (0.8 times as long in L. billambi ), and the

dactylus plus unguis of pereopod 1 are shorter than the

propodus (equal in length in L. billambi)', further, the male of

L. savignyi sensu stricto also has a dorsodistal flange on the

basis of pereopod 6 (Bamber, 2010), unlike that of L. billambi.

Other than the present species, there are only four described

species which have the distal seta on article 2 of the antennule

as long as or longer than the article itself (L. itoi Ishimaru

1985, L. daggi Bamber, 2005, L. opteros Bamber, 2008 and

L. guduroo Bamber, 2008), but all of these have the first

peduncle article of the antennule less than three times as long

as wide.

Leptochelia billambi was recorded commonly at between 0

and 14 m depth in Port Phillip Bay and Western Port, Victoria.

Genus Araleptochelia gen. nov.

Diagnosis. Female with 4-articled antennule, third article

longer than second; antenna 6-articled, articles 2 and 3 with

slender distal spines, article 2 without ventral spine; maxilliped

basis with four long setae extending to half length of second

palp article, endites distally with single seta and three slender

spatulate spines; cheliped relatively slender, merus covering

less than half of ventral margin of carpus, propodus (palm of

chela) nearly twice as long as wide; dactylus plus unguis on

first pereopod 1.5 times as long as propodus; merus of

pereopods 2 and 3 with ventrodistal spine about two-thirds as

long as carpus; pleopod without inner plumose seta on the

pleopod endopod; uropod exopod 1-segmented, endopod

5-segmented, with all segments elongate (more than four times

as long as wide). Otherwise typical of the family. Male

showing dimorphism in the antennule with secondary

segmentation of the flagellum to more than 17 segments,

proximal flagellum segment bearing large dorsodistal spine

almost as long as flagellum; cheliped slender, sinuous;

posterior pereopods without flange on basis. Sub-adult male

with five-articled antennule.

Type species: Araleptochelia macrostonyx sp. nov. by

monotypy,

Etymology, from the Greek araeos - thin, as is the chela of the

female (and the proximal uropod endopod segments) - and

ii Leptochelia , \ to which genus it is closest (female).

Remarks. Ostensibly, the species described below has the gross

appearance of a typical Leptochelia', females of that genus are

remarkably conservative in their morphology, a factor which in

the past has lead to over-synonymization. The present species

has a sufficient number of distinguishing characters, notably

the slender chela (somewhat reminiscent of that of a

typhlotanaid); the very slender distal spines on the second and

third peduncle articles of the antenna; the proportionately very

long dactylus and unguis of pereopod 1; the long ventrodistal

spines on the merus of pereopods 2 and 3; the lack of an inner

plumose seta on the pleopod endopod; and the elongate

proximal uropod segments, that it warrants separation into a

distinct genus.

The spination/setation of the antennal peduncle articles is

more typical of such genera as Konarus Bamber, 2006,

Pseudonototanais Lang, 1973 and Parakonarus Bird, 2011

rather than of Leptochelia, while the proportions of the dactylus

and unguis of pereopod 1 are more typical of Leptochelia.

The mouthparts are typical of Leptochelia, inter alia,

and while the maxilliped basis has four distal setae in the only

known species (thus distinguishing it from such other

leptocheliid genera as Heterotanais and Pseudoleptochelia ),

the number of these setae is known to be variable in, for

example, Leptochelia (e.g. Bamber, 2008), so cannot as yet

be regarded as diagnostic. While the chela of the male may

be regarded as somewhat intermediate between that of

the L. savignyi- type and that of the L. minuta- type, the extreme

secondary segmentation of the antennule flagellum, with an

extraordinarily long spine on the first flagellum segment,

reinforces the difference between this taxon and species

of Leptochelia.

Araleptochelia macrostonyx sp. nov.

Figures 75-78

Material examined. 1 brooding $ (J58470), holotype, 11 99 (J56622),

paratypes, Stn 81-HK-l 134, 40°56.0'S 146°05.40'E, 68 m depth, mud,

4 February 1981, coll. MF Gamon et al., FRV Hai Kung, pipe-dredge.

3 99 (J56629), paratypes, Stn 80-Sa-l 113, 40°23.8'S 146°32'E, 65 m

depth, muddy sand, 3 November 1980, coll. MF Gamon et al., FRV

Sarda. 14 99 (J56630), paratypes, Stn 83-SG-l 209, 38°18.0'S

147°37.0'E, 55 m depth, muddy fine shell, 31 July 1983, coll. MF

Gamon et al., FV Silver Gull.

13 99 , 2 88 (J56636), 5 99 , 1 juvenile (J56635), paratypes, Stn 81-T-

1 158, 39°49.5'S 146°18.5'E, 82 m depth, sand-silt-mud, 13 November

1981. 1 $ (J56634), paratype, Stn 81-HK-l 189, 38°42.8'S 142°35.6'E,

69 m depth, coarse sand, 20 November 1981. 1 9 (J56626), paratype,

Stn 81-HK-l 201, 39°08.3'S 144°43.9'E, 66 m depth, coarse sand, 23

November 1981. 9 99 , 1 8 (J56627), 1 preparatory 8 (J56639), 8

preparatory 88 (J56640), paratypes, Stn 81-HK-l 159, 39°43.5’S

146°18.8'E, 80 m depth, muddy shell, 13 November 1981.1 9 (J56631),

1 $ (J56633), 1 preparatory 8 (J56638), paratypes, Stn 81-HK-l 161,

39°48.3'S 147°19.2'E, 60 m depth, muddy sand, 14 November 1981.1 $

(J56624), paratype, Stn 81-HK-l 173,39°26.3'S 147°48.7'E, 49 m depth,

medium sand, 17 November 1981. 2 99 (J56628), paratypes, Stn 81-T-l

169, 39°02.4'S 148°30.6'E, 120 m depth, muddy sand, 15 November

1981. All coll. R.S. Wilson, RV Tangaroa.

1 preparatory 8 (J56637), paratype, Stn 83-SG-l 209, 38°18.0’S

147°37.0'E, 55 m depth, muddy fine shell, 31 July 1983, coll. M.F.

Gomon & R.S. Wilson, FV Silver Gull.

-rfftfr

114

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 7 5. Araleptochelia macrostonyx sp. nov., female holotype. A, dorsal view; B, lateral view; C, male lateral view. Scale = 1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

115

Fig. 76. Araleptochelia macrostonyx sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, right mandible; E, E' left mandible;

F, maxillule endite; G, maxilla; H, labium; I, maxilliped. Scale = 0.1 mm.

116

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 77 Araleptochelia macrostonyx sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 6;

G, pleopod; H, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

117

Fig. 78. Araleptochelia macrostonyx sp. nov., male. A, antennule; B, antenna; C, cheliped; D, pereopod 1; E, pereopod 2; F, pereopod 3;

G, pereopod 6. Scale = 0. 1 mm.

118

M. Biazewicz-Paszkowycz & R.N. Bamber

Description of female. Body (Fig. 75A, B) slender, holotype

2.3 mm long, 7.4 times as long as wide. Cephalothorax

subrectangular, 1.3 times as long as wide, as long as pereonites

1 and 2 together, with slight rostrum, eyelobes and black eyes

present, single setae at posterior of eyelobes, midlaterally and

posterolaterally. Pereonites 1 and 2 subequal, shortest;

pereonites 3 and 6 subequal, slightly longer than pereonite 1;

pereonites 4 and 5 subequal, longest and almost twice as long

as pereonite 1 (all pereonites respectively 1.5, 1.6, 1.5, 0.9, 0.9

and 1.4 times as wide as long). Pleon with five free subequal

pleonites bearing pleopods, each pleonite about 6 times as wide

as long, with paired lateral setae. Pleotelson semicircular,

longer than last two pleonites together, 1.7 times as wide as

long, on each side bearing one posterolateral seta and a pair of

one simple and one penicillate setae posterior to uropod

attachment, and two distal setae.

Antennule (Fig. 76A) of four tapering articles, as long as

cephalothorax, proximal article 3.1 times as long as wide;

second article almost twice as long as wide, one-quarter as

long as first article, longest distal outer only just longer than

article; third article 1.3 times as long as second; fourth article

minute, eccentric, with four distal setae and one aesthetasc.

Antenna (Fig. 76B) of six articles, proximal three articles

subequal in length; proximal article naked; second article with

single dorsodistal slender spine; third article as long as wide,

with slender dorsodistal spine; fourth article longest, 2.7 times

as long as third article and nearly 5 times as long as wide; fifth

article 0.75 times as long as fourth; sixth article minute.

Labrum (Fig. 76C) hood-shaped, setose, typical of genus.

Left mandible (Fig. 35E, E') with crenulate lacinia mobilis

narrower than distal end of mandible, proximal crenulation on

pars incisiva, pars molaris (Fig. 76E) robust; right mandible

(Fig. 76D) similar but without lacinia mobilis. Labium (Fig.

76H) wide, bilobed, distally finely setose, without palp.

Maxillule (Fig. 76F) with ten distal spines and setose margins,

setules on inner distal face paired. Maxilla (Fig. 76G) simple,

linguiform, naked. Maxilliped (Fig. 761) palp first article naked,

second article with one outer and three inner setae in distal half,

distal-most inner seta only half as long as third palp article;

third and fourth articles with filtering rows of six and five setae

respectively, fourth article with outer seta; basis with four long

setae extending to half length of second palp article; endites

distally with single seta and three slender spatulate spines.

Cheliped (Fig. 77A) basis nearly twice as long as wide;

merus subtriangular with two ventral setae; carpus 2.8 times

as long as wide, with three ventral setae in distal half, one

proximal and one distal dorsal setae; propodus elongate for the

genus, palm 1.8 times as long as wide, fixed finger 0.45 times

as long as palm with two ventral and three inner setae alongside

cutting edge, cutting edge crenulate, comb-row at base of

dactylus of four setae with adjacent microtrichia; dactylus

with outer proximal seta.

Pereopod 1 (Fig. 77B) longer than other pereopods, coxa

with seta; basis slender, five times as long as wide; ischium

compact with one ventral seta; merus 0.6 times as long as

carpus, with single distal seta; merus-carpus articulation

strongly oblique; carpus with one dorsal and three ventral

distal setae, longest of which is less than one-third as long as

propodus; propodus twice as long as carpus, with three setae

on subdistal dorsal mound and one subdistal ventral seta;

dactylus slender, with proximal seta, extending into slightly

longer slender unguis, the two together 1.5 times as long as

propodus. Pereopod 2 (Fig. 77C) more compact than pereopod

1; basis 3.3 times as long as wide; ischium with 2 setae; merus

as long as carpus, with slender ventrodistal spine more than

half as long as carpus; carpus with dorsodistal seta and short

ventrodistal spine; propodus 1.75 times as long as carpus, with

two dorsodistal and one ventrodistal setae, and dorsodistal

sharp apophysis; curved dactylus and slightly longer unguis

together 0.7 times as long as propodus. Pereopod 3 (Fig. 77D)

similar to pereopod 2, but dactylus and unguis subequal.

Pereopod 4 (Fig. 77E) basis stouter than those of anterior

pereopods, twice as long as wide; ischium with two seta; merus

and carpus subequal, merus with two short, ventrodistal spines

and microtrichia, carpus with one outer, one ventral and one

inner distal spines each with serrate ventral margin, single inner

and outer simple dorsodistal setae and ventral microtrichia;

propodus 1.2 times as long as carpus, with ventral microtrichia,

two ventrodistal serrate spines, and three dorsodistal finely-

denticulate setae almost as long as dactylus; dactylus and short

unguis distinct, curved, together two-thirds as long as propodus.

Pereopod 5 as pereopod 4. Pereopod 6 (Fig. 77F) similar to but

stouter than pereopod 4, but distal carpal spines larger, propodus

with two pectinate and three finely-denticulate distal setae.

Pleopods (Fig. 77G) all alike, with no inner plumose seta

on endopod; proximal outer seta separated from others on

both rami.

Uropod (Fig. 77H) biramous, basis naked; exopod of one

slender segment as long as proximal endopod segment, outer

distal seta longer than inner distal seta; endopod of five

elongate segments.

Description of male. Typical primary male, smaller than female

(allotype length 1.6 mm), body (Fig. 75C) more compact,

cephalon as long as pereonites 1 to 3, with large eyelobes

bearing large black eyes; pereonite 5 longest, other pereonites

subequal in length. Five free pleonites, subequal in length and

about as long as pereonite 1, pleotelson nearly twice as long as

pleonite 5. Sexual dimorphism as follows.

Antennule (Fig. 78A) elongate, first peduncle article

curved, 2.8 times as long as wide with one dorsodistal seta;

second article compact, 0.25 times as long as first with long

outer distal seta; third article compact with dorsodistal spine

and ventrodistal penicillate and simple setae; flagellum of 19

segments, first 18 each bearing distal row of aesthetascs,

proximal flagellum article with extraordinarily-long

dorsodistal spine extending almost full length of flagellum;

distal article short and with four simple setae. Antenna (Fig.

78B) with slender distal setae rather than spines on articles 2

and 3, articles 4 and 5 apparently fused. Mouthparts atrophied.

Cheliped (Fig. 78C) very slender, three-quarters as long as

body; basis arcuate, about 3 times as long as wide; carpus slender

and sinuous, about 5 times as long as wide with two ventral setae

in proximal half; palm of propodus twice as long as wide, fingers

of chela 0.8 times length of palm; fixed finger longer than palm,

with two ventral and three inner setae alongside cutting edge,

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

119

cutting edge with sharp denticulations but no large tooth-like

apophyses, distal spine rugose; dactylus with sharp tooth-like

denticulations along cutting edge, unguis rugose.

Pereopods (Fig. 78D to G) more elongate than those of

female; unguis of pereopod 1 shorter than dactylus; on

pereopods 4 to 6 carpal spines more slender, distal articles

without dense fields of microtrichia, unguis longer and more

slender, together with dactylus >0.8 times as long as propodus.

Preparatory male. As female, but antennule with four longer

articles as well as minute distal article, from division of third

peduncle article of neuter.

Etymology. From the Greek macro - long and stonyx - a sharp

point, in reference to both the extremely long dactylus and

unguis on pereopod 1, the ventrodistal spine on the merus of

pereopods 2 and 3, and the spine on the fourth antennal article

of the male, all characterizing features of this species.

Remarks. The numerous distinctions of this species from those

most similar, i.e. species of Leptochelia, are described under

the remarks for the genus. Araleptochelia macro stonyx sp. nov.

contributes further to the great diversity of the Leptocheliidae

in Australian waters and is readily distinguished from other

species without dissection owing to its unique cheliped

morphology and the extremely long dactylus plus unguis on

pereopod 1. The antennular morphology of four longer

segments in the subadult male was demonstrated for the related

Leptochelia savignyi (Kr0yer, 1842) by Bamber (2010).

A. macrostonyx was found on muddy to coarse sand

substrata at depths from 49 to 120 m right across the northern

half of the Bass Strait.

Genus Pseudoleptochelia Lang, 1973

Pseudoleptochelia occiporta sp. nov.

Figures 79-82

Material examined. 1 9 (J58467), holotype, 1 6 (J58468), allotype, 48 9

9 , 2 juveniles (J55820), paratypes, Stn 81-T-l 162,40°09.2'S 147°31.9'E,

51 m depth, shelly sand, 14 November 1981, coll. R.S. Wilson, RV

Tangaroa. 4 99 , 2 juveniles (J50804), paratypes, Stn VC 23 Cl,

38°19.18'S 144°37.62E, 40 m deep, reef, 12 May 1998. 10 specimens

(J48183), paratypes, CPBS 01S/4,38°21.73'S 145°13.23'E, 3 m depth, 1

April 1965. 6 specimens, (J48188), paratypes, CPBS 03S/2, 38°21.65'S

145°15.21'E, 2 m depth, sandy-mud, 13 April 1965. 25 specimens,

(J48175), paratypes, CPBS 11N/4, 38°23.23'S 145°13.28'E, 5 m depth,

fine sand, 21 March 1965. 10 specimens, (J48178), paratypes, CPBS

11S/2, 1 specimen, (J48751), paratypes, CPBS 11S/4, both 38°22.00’S

145°13.38'E, 3 m depth, shelly gravel, 17 March 1965. 5 specimens,

(J48755), paratypes, CPBS 22N/4,38°20.60'S 145°13.46'E, 13 m depth,

shelly sand, 18 March 1965. 66 99 , 26 juveniles, (J48967), paratypes,

CPBS 32N/367, 38°20.83’S 145°13.49’E, 13 m depth, sandy gravel, 20

March 1967. 1 9 (J48972), 3 specimens (J48970), paratypes, CPBS

32N/769, 38°20.83'S 145°13.49'E, 13 m depth, sandy gravel, 15 July

1969. 12 specimens, (J48971), paratypes, CPBS 32N/870, 38°20.83'S

145°13.49'E, 13 m depth, sandy gravel, 12 August 1970. 20 99 , 10

juveniles, (J48980), paratypes, CPBS 32S/770,38°21.60'S 145°13.67'E,

13 m depth, muddy sand, 6 July 1970.8 specimens, (J48974), paratypes,

CPBS 32S/866, 38°21.60’S 145°13.67'E, 13 m depth, muddy sand, 26

August 1966. 1 9 . (J48955), paratypes, CPBS 400/3, 38°21.17'S

145°14.00'E, 15 m depth, sand, 1 April 1965. 3 specimens, (J48958),

paratypes, CPBS 41N/2, 38°20.81'S 145°13.85'E, 13 m depth, gravelly

sand, 30 March 1965.

Description of female. Body (Fig. 79A, B, C, D) relatively small

for the genus, holotype 2.7 mm long, 6.3 times as long as wide.

Cephalothorax subrectangular, tapering towards anterior, 1.4

times as long as wide, 1.5 times as long as pereonites 1 and 2

together, with slight rostrum, eyelobes rounded, eyes present

and black, paired setae at posterior of eyelobes and single setae

midlaterally. Pereonite 1 shortest, pereonite 2 and 6 subequal

and 1.5 times as long as pereonite 1, pereonites 3 to 5 subequal

and 1.9 times as long as pereonite 1 (all pereonites respectively

2.6, 1.8, 1.4, 1.4, 1.5 and 1.7 times as wide as long). Pleon with

five free subequal pleonites bearing pleopods; each pleonite

about 5.3 times as wide as long, with single midlateral setae.

Pleotelson pentangular, as long as last two pleonites together,

1.6 times as wide as long, with one anterolateral and two

posterolateral setae on each side and two distal setae.

Antennule (Fig. 80A) of three longer and one minute distal

articles, proximal article 3.5 times as long as wide, 1.5 times

as long as distal three articles together, with proximal, mesial

and distal inner groups of penicillate setae, inner simple

mesial seta, and one outer and one long inner distal seta longer

than second article; second article twice as long as wide, one-

third as long as first article, longer distal seta just longer than

article; third article just shorter than second, distally with one

simple and one penicillate seta; fourth article minute, distally

with three simples and two penicillate setae and one aesthetasc.

Antenna (Fig. 80B) of six articles, proximal article compact,

naked; second article as long as wide, with single inner distal

and dorsodistal slender spines; third article as long as second

and just longer than wide, with dorsodistal stouter spine (Fig.

80b'); fourth article longest, 3.8 times as long as wide and twice

as long as third, with short mesial setae and distal tufts of

penicillate and simple setae; fifth article 0.7 times as long as

fourth; sixth article minute with six distal setae.

Labrum (Fig. 80C) hood-shaped, setose. Left mandible

(Fig. 80D) with crenulate lacinia mobilis wider than distal end

of mandible, distal crenulation on pars incisiva, pars molaris

with strong rugosity; right mandible (Fig. 80E) similar but

without lacinia mobilis, crenulation of pars incisiva extending

down inner margin. Labium (Fig. 80G) wide, bilobed, distally

finely setose, without palp. Maxillule (Fig. 80F) with nine

distally bifurcate spines and setose margins, rows of setules on

inner distal face; palp distinct, with two distal setae. Maxilliped

(Fig. 80H) palp first article naked, second article with finely

setose inner margin, and with one outer and two inner setae,

distal-most inner seta not reaching distal margin of third palp

article; third and fourth articles with 11 inner marginal setae

in two rows of 7 (dorsal) and 4 (ventrodistal); fourth article

with one outer subdistal seta; basis with 4 to 6 long setae

(sometimes asymmetrical - Fig. 80H') extending to distal edge

of second palp article; endites (Fig. 80FI") distally with fine

outer setules, long outer seta and three robust spatulate spines,

with additional subdistal pair of spatulate spines.

Cheliped (Fig. 81A) with rounded, comparatively stout

basis 1.6 times as long as wide, with subdistal dorsal seta;

merus triangular with three ventral setae; carpus 1.7 times as

120

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 79. Pseudoleptochelia occiporta sp. nov., holotype female. A, lateral view; B. manca, dorsal view; C, 4 mm female paratype; D, 3 mm female

paratype; E, male lateral. Scale: A, B, D = 0.2 mm; E = 0.1 mm (scale for C as for D).

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

121

Fig. 80. Pseudoleptochelia occiporta sp. nov., female paratype. A, antennule; B, antenna (proximal article not shown); B', spine on antennal

article 3; C, labrum; D, left mandible; E, right mandible; F, maxillule; F', maxillule endite details; G, labium; H, maxilliped; H', maxilliped bases

of smaller female; H”, endite of H. Scale = 0.2 mm.

122

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 81. Pseudoleptochelia occiporta sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 6;

G, pleopod; H, uropod H', uropod details. Scale = 0. 2 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

123

Fig. 82. Pseudoleptochelia occiporta sp. nov., male. A, cheliped; B, antennule; C, antenna; D, pereopod 1; E, pereopod 2; F, pereopod 3;

G, pereopod 4; H, pereopod 5; I, pereopod 6; J, pleopod; K, uropod. Scale = 0.1 mm.

124

M. Biazewicz-Paszkowycz & R.N. Bamber

long as wide, with three midventral setae and three shorter

dorsal marginal setae; propodus as long as wide, with

dorsodistal seta and one seta at base of dactylus; fixed finger

with five ventral and three inner setae, cutting edge crenulate,

subdistal claw; dactylus with finely-crenulate cutting edge.

Pereopod 1 (Fig. 81B) longer than other pereopods, coxa

with long seta; basis slender, 4.4 times as long as wide, with

single dorsoproximal simple and penicillate setae; ischium

compact with one ventral seta; merus 0.9 times as long as carpus,

with one dorsodistal and two ventrodistal setae; carpus with

distal crown of two dorsodistal, one inner and two outer medial,

and one ventrodistal setae, longest of which is much less than

half length of propodus; propodus 1.5 times as long as carpus,

with three setae on subdistal dorsal mound and one subdistal

ventral seta; dactylus slender, curved, extending into shorter

slender unguis 0.7 times as long as dactylus, the two together

some 1.4 times as long as propodus. Pereopod 2 (Fig. 81C) more

compact than pereopod 1; basis 2.45 times as long as wide;

ischium with 2 setae; merus just longer than carpus and 1.1 times

as long as wide, with single dorsodistal and ventrodistal setae

and tooth-like ventrodistal spine and ventral rows of microtrichia;

carpus with two dorsodistal and single ventrodistal setae and two

small ventrodistal spines, and ventral rows of microtrichia;

propodus 1.9 times as long as carpus, with three distal setae and

dorsodistal sharp apophysis; dactylus and shorter unguis curved,

together 0.6 times as long as propodus. Pereopod 3 (Fig. 81D)

similar to pereopod 2, but merus and carpus with no and one

dorsodistal setae, propodus with two distal setae.

Pereopod 4 (Fig. 81E) basis stout, 1.7 times as long as wide

with two ventroproximal penicillate setae; ischium with two

ventrodistal setae; merus with paired ventrodistal spines and

ventral rows of microtrichia; carpus just shorter than merus,

with one dorsodistal seta, outer, ventral and inner distal spines

each with fine subdistal setules, and ventral rows of microtrichia;

propodus 1.1 times as long as carpus, with two ventrodistal

spines, three dorsodistal setae mostly as long as dactylus,

serrated spines adjacent to dactylus articulation, and ventral

rows of microtrichia; dactylus and distinct unguis curved, 0.6

times as long as propodus, dactylus with microtrichia. Pereopod

5 as pereopod 4. Pereopod 6 (Fig. 81F) as pereopod 4, but

propodus with four shorter and one longer distal setae.

Pleopods (Fig. 81G) all alike, typical for the genus, basis

naked, endopod with single inner plumose seta and proximal

outer seta separated from remainder.

Uropod (Fig. 81H) biramous, basis naked; exopod of two

segments, as long as proximal endopod segment, outer distal

seta longer than inner distal seta; endopod of five or six

segments, distal segments slender.

Description of male. Smaller than female (allotype length

1.5 mm), body (Fig. 79E) more compact, cephalon 1.25 times

as long as pereonites 1 to 3 together, with large eyelobes

bearing large black eyes; pereonite 1 shortest, pereonite 2 just

longer, pereonites 3 and 6 subequal, 1.8 times as long as

pereonite 1, pereonites 4 and 5 twice as long as pereonite 1.

Five free pleonites, subequal in length, as long as pereonite 1,

pleotelson as long as pleonites 4 and 5 together. Pleotelson with

much longer distal setal pair (Fig. 82K).

Antennule (Fig. 82B) first peduncle article 2.4 times as

long as wide, with paired dorsomedial penicillate setae and

single dorsodistal penicillate and simple setae; second article

0.65 times as long as first with single ventroproximal and

dorsal subdistal simple setae; third article 0.3 times as long as

second with single ventrodistal simple seta; flagellum of 8

segments, segment 1 with proximal and distal rows of

aesthetascs, segments 2 to 7 bearing distal row of 5 or 6

aesthetascs, segment 8 minute, distally with four simple setae.

Antenna (Fig. 82C) of six articles, proximal article

compact, naked; second article longer than wide, longer than

first article, with single dorsoproximal and three distal seta;

third article shorter than second, 1.6 times as long as wide,

with single simple distal seta; fourth article nearly twice as

long as third, with mesial seta and distal tufts of penicillate

and simple setae; fifth article as long as fourth with two long

distal setae; sixth article minute, with three simple distal setae.

Mouthparts atrophied.

Cheliped (Fig. 82A) with subchelate chela; basis twice as

long as wide; merus short, with seven ventral setae; carpus

1.9 times as long as wide but with large subtriangular ventral

apophysis bearing the three midventral setae, dorsal margin

with continuous row of nine setae; propodus twice as long as

wide with dorsodistal seta, fixed finger reduced to small

apophysis with claw, with three ventral setae, three setae

adjacent to remains of cutting edge, inner diagonal comb-

row of ten shorter and one longer setae; dactylus slender,

curved, as long as propodus, with row of four setae along

cutting edge.

Pereopods (Fig. 82D to I) similar to but more slender than

those of female, with in particular more elongate propodi;

merus shorter than carpus on pereopods 1 to 3; on pereopods

4 to 6 propodus much longer than carpus, ventrodistal merus

spines more elongate than those of female, dactylus

proportionately longer than in female.

Pleopods (Fig. 82J) with longer setae than on those of

female. Uropod (Fig. 82K) basis with inner distal row of setae;

exopod two-segmented, longer than proximal endopod

segment; endopod of five segments, less-elongate and more

heavily setose than that of female.

Distinctions ofmanca. Smaller than female (Fig. 79B), similar

in morphology and proportions, uropod exopod of one segment.

Etymology. From the Latin occidentalis - western, and portus

- a port, the species mainly occurring in, and common in,

Western Port, Victoria.

Remarks. The genus Pseudoleptochelia is in need of review.

Bird & Bamber (2000) listed twelve species in the genus, since

when three further species have been described, P. fairgo

Bamber, 2005, and P. straddi Bamber, 2008, both from

Australia, and P. bulbus Bamber, 2006 from New Caledonia.

However, the male of P. bulbus , and P. straddi (known only

from the male) are currently suspected to be species of a

different genus (Bamber, in prep.). Many of these

Pseudoleptochelia species are poorly described, particularly

their females and their mouthparts, some being known only

from males.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

125

Lang (1973) diagnosed the genus Pseudoleptochelia as

having stout posterior pereopod bases, with spines on the

second and third antennal peduncle articles (as in Leptochelia),

the uropod exopod with one or two segments, the endopod

with at least three segments, and the maxilliped bases with

two distal setae. Unfortunately, Lang (ibid.) based his genus

on Heterotanais anomalus Sars, 1882, the female of which is

not fully described, and on his own new species,

Pseudoleptochelia mortenseni, which, judging from his

figure 16, is clearly a chimaera, as the antennule of “a small

female” (Lang, 1873: fig. 16g) is typical of a species of

Konarus Bamber, 2006 or Parakonarus Bird, 2011, and not

Pseudoleptochelia sens, auctt., casting doubt on with which

female his subchelate male should be associated (see Bird,

2011). In practice, the maxilliped-basis setation and female

antennal spination are unknown in at least half of the

Pseudoleptochelia species.

Pseudoleptochelia occiporta sp. nov. is consistent with the

generic diagnosis of Lang (loc. cit.) except for its having 4 to 6

maxilliped-basis setae, a feature of three other leptocheliid

genera, Leptochelia, Konarus and Parakonarus. It shares this

feature, as well as the five distal spatulate spines on the

maxilliped endite, with P. bulbus Bamber, 2006 from New

Caledonia, a species differing owing to its one-segmented

uropod exopod and distinct setation of the pereopod 1 carpus,

inter alia.

The genera Konarus and Parakonarus both have setae

rather than spines on the second and third antennal peduncle

articles, and an unguis distinctly longer than the dactylus on

pereopod 1, unlike the present species. We therefore choose to

place the present species in Pseudoleptochelia until the genus

has been properly resolved.

The only currently accepted species of Pseudoleptochelia

recorded previously from Australia is P. fairgo, known from

Esperance, Western Australia, and Brisbane, Queensland

(Bamber, 2005; 2008). The female of P. fairgo, which also

has four maxilliped-basis setae, is unique in the genus in

having setose tubercles on the merus of pereopods 4 to 6, and

a tuft of elongate setae on the cheliped carpus pointing

proximally in life, neither feature being present in

P. occiporta.

The male of the present species has a rounded apophysis

on the carpus of the cheliped like Pseudoleptochelia fairgo,

but differs from that species in that P. fairgo has a

ventrodistal apophysis on the propodus of the cheliped

which is wider and rounded (slender and pointed in

P. occiporta ) and a strong dorsodistal seta on the third

antennular peduncle article as long as the proximal four

flagellum segments (only a very small dorsodistal seta in

P. occiporta ). In P. occiporta the uropod exopod is two-

segmented in both sexes, but the exopod in P. fairgo is one-

segmented with only an incipient fusion line. Of the four

Pseudoleptochelia species having a two-segmented uropod

exopod, none have a ventral carpal apophysis on the male

cheliped as found in P. occiporta.

Pseudoleptochelia occiporta was taken at depths between

3 and 51 m, on gravelly to muddy sands, mainly in Western

Port.

Genus Bassoleptochelia gen. nov.

Diagnosis. Female with 4-articled antennule, third article longer

than second; antenna with setae rather than spines on articles 2

and 3; mandibles with relatively simple lacinia mobilis and pars

molaris, maxilliped with three basal setae, elongate distal

spatulate spines on endites, setae on palp article 2 simple;

cheliped very slender, merus covering less than half of ventral

margin of carpus and with ventral tuft of numerous long setae,

propodus (palm of chela) longer than wide, fixed finger with two

ventral setae; pereopod 1 with elongate merus and distal crown

of setae on carpus, dactylus longer than unguis; merus, carpus,

propodus and dactylus of pereopods 4 to 6 with fields of

microtrichia, carpus of pereopod 6 with dense brush of

microtrichia (prickly tubercle); uropod exopod 1-segmented,

endopod 4-segmented. Otherwise typical of the family. Male

showing dimorphism in the antennule with secondary

segmentation of the flagellum to more than 5 segments; cheliped

slender, chela almost subchelate, fixed finger one-third as long as

dactylus, distal edge of propodus with triangular tooth-like

apophysis, comb-row vertical; pereopods more slender than

those of female, posterior pereopods ambulatory.

Type species. Bassoleptochelia verro sp. nov. by monotypy.

Etymology. Named for the Bass Strait, plus -Leptochelia (female).

Remarks. The species described below has the gross appearance

of a typical leptocheliid, but the tufts of microtrichia on the

posterior pereopods are not found in any other species of the

family (being more like those found on some typhlotanaid

species - see Blazewicz-Paszkowycz, 2007). Among Australian

leptocheliids, the “brush” of long setae on the cheliped merus is

also found in Pseudoleptochelia fairgo, although in that species

they point proximally in life. Equally, the male is unusual for the

family in having ambulatory posterior pereopods, which must

develop secondarily from those of the subadult form which are

like those of the female, and in the unique cheliped chela, which

falls somewhere between the normal chelate form of most genera

and the subchelate form found in some species of

Pseudoleptochelia and Parakonarus. The mouthparts are also

atypical of the family, in having simple inner setae on the second

palp article of the maxilliped, while the mandibular lacinia

mobilis and pars molaris are both simple, rather than crenulate

or rugose, respectively.

Bassoleptochelia verro sp. nov.

Figures 83-86

Material examined. 1 9 (J58469), holotype, 5 99 (J55822), paratypes, Stn

81-T-l 177,38°53.7'S 147°06.5'E, 58 m depth, coarse shell, 18 November

1981. 1 8 (J58471), allotype, 1 $ (J56650), paratype, Stn 81-T-l 199,

40°19.5'S 143°48.8'E, 71 m depth, sandy shell, 22 November 1981.4 99

(J56657), 1 $ (J56652), paratypes, Stn 81-T-l 203,39°22.0'S 144°18.3'E,

60 m depth, coarse sand, 23 November 1981.13 99 (J56646), paratypes,

Stn 81-T-l 180, 39°12.9'S 146°27.3’E, 65 m depth, medium sand, 18

November 1981. 1 9 (J56643), paratype, Stn 81-T-l 187, 38°32.0'S

147°28.6’E, 52 m depth, medium sand, 20 November 1981. All coll.

R.S. Wilson, RV Tangaroa. 8 99 , 1 S' (J56651), paratypes, Stn 81-Sa-l

116,40°32.0’S 145°23'E, 43 m depth, muddy shell and grit, 4 November

1980, coll. M.F. Gomon & G.C.B. Poore, FRY Sarda.

126

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 83. Bassoleptochelia verro sp. nov. A, female holotype, dorsal; B, neuter; C, juvenile; D, manca II; E, male, lateral. Scale = 1mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

127

Fig. 84. Bassoleptochelia verro sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, right mandible; E, incisor and E', molar of left

mandible; F, maxillule; G, labium; H, maxilliped and FI' endite. Scale = 0.1mm.

128

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 85. Bassoleptochelia verro sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 6 with F'

distal detail; G, pleopod; H, uropod. Scale = 0.1mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

129

Fig. 86. Bassoleptochelia verro sp. nov., male. A, antennule; B, antenna; C, mouthparts; D, cheliped; E, pereopod 1; F, pereopod 2; G, pereopod

4; H, pereopod 5; I, uropod. Scale = 0.1mm.

130

M. Biazewicz-Paszkowycz & R.N. Bamber

Description of female. Body (Fig. 83A) narrowing anteriorly,

holotype 3.8 mm long, 6.4 times as long as wide. Cephalothorax

subrectangular, laterally convex, tapering towards anterior, 1.4

times as long as wide, longer than pereonites 1 and 2 together,

with slight triangular rostrum, eyelobes extended into spine-like

apophysis (Fig. 84A), eyes present and black, single setae at

posterior of eyelobes. Pereonites 1 to 4 with single anterolateral

setae, pereonites 5 and 6 with paired lateral setae on each side;

pereonites 1 and 2 subequal and shortest, pereonites 3, 4 and 6

subequal, 1.6 times as long as pereonite 1, pereonite 5 longest

and twice as long as pereonite 1 (all pereonites respectively 2.0,

2.0, 1.3, 1.5, 1.2 and 1.5 times as wide as long). Pleon with five

free subequal pleonites bearing pleopods; each pleonite about

5.3 times as wide as long, with single midlateral seta. Pleotelson

pentangular, as long as last two pleonites together, twice as wide

as long, with one simple and one penicillate posterolateral seta

on each side and two distal setae.

Antennule (Fig. 84A) of three longer and one minute distal

articles, proximal article 2.4 times as long as wide, 1.6 times

as long as distal three articles together, with mesial and distal

inner groups of penicillate setae, inner and outer simple mesial

setae, outer distal seta and one long inner distal seta longer

than tip of antennule, outer margin of article finely setulose in

proximal half; second article as long as wide, one-quarter as

long as first article, inner and outer distal setae three times as

long as article; third article 1.3 times as long as second, with

two distal setae; fourth article minute, distally with three

simple setae, one penicillate seta and one aesthetasc.

Antenna (Fig. 84B) of six articles, proximal article longer

than wide, naked; second article as long as wide, with single

inner distal and dorsodistal strong setae; third article half as

long as wide, one-third as long as second article, with

dorsodistal slender seta; fourth article longest, 3.2 times as

long as wide and 4.5 times as long as third, with distal crown

of penicillate and simple setae; fifth article 0.6 times as long as

fourth; sixth article minute, with three simple distal setae.

Labrum (Fig. 84C) hood-shaped, finely setose. Left

mandible (Fig. 84E) with simple subrectangular lacinia

mobilis, pars incisiva distally bilobed with crenulation on

proximal inner margin, pars molaris narrow with slight

marginal crenulation distally; right mandible (Fig. 84D)

similar but without lacinia mobilis, pars incisiva distally

crenulate. Labium (Fig. 84G) wide, bilobed, distally finely

setose, outer lobe with single outer distal seta, without palp.

Maxillule (Fig. 84F) with eight distal spines, outer distal tuft

of setules, rows of setules on inner face; palp with distinct

articulation, with two distal setae. Maxilla not recovered.

Maxilliped (Fig. 84H) palp first article elongate, naked; second

article with four simple inner setae; third article with five

inner marginal finely-denticulate setae; fourth article with six

longer and one shorter distal finely denticulate setae and one

outer subdistal seta; basis with three long setae extending to

third palp article; endites distally with fine outer setules and

three elongate spatulate spines and long inner simple seta.

Epignath not recovered.

Cheliped (Fig. 85A) with elongate slender basis 3.5 times

as long as wide; merus subtriangular with brush of nine ventral

setae and one outer dorsoproximal seta; carpus twice as long

as wide, with three midventral setae on slightly flattened edge

and four shorter dorsal marginal setae, dorsoproximal margin

of carpus slightly crenulate; propodus relatively slender, 1.3

times as long as wide, with inner distal comb of three setae

and longer seta at base of dactylus; fixed finger with two

ventral and three inner setae, cutting edge crenulate; dactylus

with proximal seta, cutting edge simple.

Pereopod 1 (Fig. 85B) slender, longer than other pereopods,

coxa with seta; basis 3.5 times as long as wide, with single

dorsoproximal simple seta; ischium compact with one ventral

seta; merus slender, six times as long as wide and as long as

carpus, with one dorsodistal seta; carpus widening distally,

with single ventrodistal, inner and outer distal and dorsodistal

setae, longest of which is less than half length of propodus;

propodus 1.5 times as long as carpus, with three setae on

subdistal dorsal mound and one subdistal ventral seta; dactylus

slender, extending into longer slender unguis 1.4 times as long

as dactylus, the two together 0.85 times as long as propodus.

Pereopod 2 (Fig. 85C) more compact than pereopod 1; basis

2.2 times as long as wide, with dorsoproximal penicillate seta;

ischium with two ventral setae; merus as long as carpus and as

long as wide, with small ventrodistal spine, ventrodistal seta

and ventral rows of microtrichia; carpus with single dorsodistal

and two ventrodistal setae and ventral rows of microtrichia;

propodus 1.6 times as long as carpus, with two distal setae,

dorsodistal sharp apophysis and ventral rows of microtrichia;

dactylus and subequal unguis only slightly curved, together

0.9 times as long as propodus, proximal seta on dactylus.

Pereopod 3 (Fig. 85D) similar to pereopod 2, but basis with

fields of microtrichia, merus without ventrodistal spine.

Pereopod 4 (Fig. 85E) basis stout, twice as long as wide

with midventral penicillate seta; ischium with two ventrodistal

setae; merus with paired ventrodistal slender spines and

ventral rows of microtrichia; carpus just longer than merus,

with dorsodistal seta and ventral rows of microtrichia;

propodus as long as carpus, with two ventrodistal spines, three

dorsodistal setae longer than dactylus, and ventral rows of

microtrichia; dactylus and distinct, short unguis curved, 0.9

times as long as propodus, with lateral rows of microtrichia.

Pereopod 5 as pereopod 4. Pereopod 6 (Fig. 85F) as pereopod

4, but carpus with dense field of microtrichia, propodus with

four distal setae.

Pleopods (Fig. 85G) all alike, typical for the genus, basis

naked, endopod with single inner plumose seta and proximal

outer seta separated from remainder.

Uropod (Fig. 85H) biramous, basis with outer distal seta;

exopod of one segment, as long as proximal endopod segment,

outer distal seta longer than inner distal seta; endopod of four

segments, distal segment slender.

Subadults. Neuter, juvenile and manca essentially smaller

versions of adult female (Figs 83B, C, D).

Description of male. Smaller than female (allotype length

1.5 mm), body (Fig. 83E) more compact, cephalon as long as

pereonites 1 to 3 together, with large eyelobes bearing large

black eyes; pereonite 1 shortest, pereonites 2 and 3 respectively

1.3 and 1.5 times as long as pereonite 1, pereonite 4 longest, 2.7

times as long as pereonite 1, pereonites 5 and 6 progressively

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

131

shorter, respectively 2.2 and 1.8 times as long as pereonite 1.

Five free pleonites, subequal in length, each as long as pereonite

2, pleotelson 1.4 times as long as pleonite 5.

Antennule (Fig. 86A) peduncle compact, first peduncle

article twice as long as wide, with ventrodistal tuft of penicillate

and simple setae; second article 0.4 times as long as first and as

long as wide, with ventrodistal tuft of penicillate and simple

setae; third article 0.25 times as long as first and slightly shorter

than wide with single ventral and long dorsal distal simple

setae; flagellum of 6 segments, segment 1 with proximal and

distal rows of aesthetascs, segments 2 to 5 bearing distal row of

3 to 5 aesthetascs, segment 6 with four distal setae.

Antenna (Fig. 86B) of six articles, proximal article longer

than wide, naked; second article as long as first, with three

distal setae; third article shorter than second, with two distal

setae; fourth article longest, 1.6 times as long as second and 4

times as long as wide, with mesial simple and penicillate setae

and distally one penicillate and three long simple setae; fifth

article 0.8 times as long as fourth; sixth article minute, with

four simple distal setae.

Mouthparts (Fig. 86C) atrophied, naked maxilliped and

maxillule palp with two distal setae distinguishable.

Cheliped (Fig. 86D) proportionately larger than that of

female; basis three times as long as wide; merus short, with

brush of seven ventral setae; carpus 1.8 times as long as wide,

with twp midventral setae. Propodus almost square, 1.3 times

as long as wide, fixed finger short with slender distal claw, with

two ventral setae and three setae adjacent to naked, simple

cutting edge; large, triangular tooth-like apophyses on distal

face of propodus, and inner vertical comb-row of 8 shorter and

one longer setae; dactylus slender, curved, nearly three times

as long as propodal fixed finger, with proximal seta and three

spinules on cutting edge, articulation of unguis obscure.

Pereopod 1 (Fig. 86E) similar to that of female but

propodus proportionately longer (1.8 times as long as carpus)

and more elongate; pereopods 2 (Fig. 86F) and 3 similar to but

shorter than pereopod 1, carpus with distal crown of setae.

Pereopods 4 to 6 (e.g. Fig. 86G, H) ambulatory, merus, carpus

and propodus slender, all distal spines more elongate than

those of female but distal setae proportionately shorter,

microtrichia restricted to carpus and propodus without dense

brush on pereopod 6, dactylus long and slender, naked, 0.7

times as long as propodus, unguis just less than half as long as

dactylus.

Pleopods with longer setae than on those of female. Uropod

(Fig. 861) similar to that of female.

Etymology. From the Latin - verro - a brush, in reference to the

tuft of setae on the cheliped merus and the dense fields of

microtrichia on the posterior pereopods, especially the carpus of

pereopod 6, analogous to the “prickly tubercles” found in some

typhlotanaids (noun in apposition).

Remarks. The numerous distinctions of this species from other

leptocheliids are described under the remarks for the genus,

and again it contributes to the great diversity of the

Leptocheliidae in Australian waters. Bassoleptochelia verro

sp. nov. was found on coarser sandy substrata at depths from 43

to 71 m right across the Bass Strait.

Family Tanaellidae Larsen & Wilson, 2002

Remarks. The family Tanaellidae was erected to include four

genera, Araphura Bird & Holdich, 1984; Araphuroides Sieg,

1986(a); Arthrura Kudinova-Pasternak, 1966, and Tanaella

Norman & Stebbing, 1886.

In particular, Araphuroides was split from Araphura by

Sieg (1096a) for two species, Araphura brevispina Bird &

Holdich, 1984 and Araphuroides parabreviremis Sieg, 1986.

Features distinguishing these two genera have been discussed

subsequently and inconsistently (e.g. Sieg & Dojiri, 1989;

Larsen, 2005; Larsen et al., 2009). The tanaellid taxa described

herein confound this issue further, and their generic attribution

is discussed below after their description.

Genus Araphura Bird & Holdich, 1984

Araphura pygmothymos sp. nov.

Figures 87-89

Material examined. 1 9 (J58833), holotype. Central Bass Strait, 66 km

S of Rodondo Island, Stn BSS 158, 39°48.6'S 146°18.8'E, 82 m depth,

sand with silt and mud, 13 November 1981; 1 9 (J58834), paratype,

same data as holotype; 1 9 (J56692), paratype. Central Bass Strait, 100

km SSE of Cape Liptrap, Victoria, Stn BSS 156,39°45.90’S 145°33.3'E,

74 m depth, muddy fine sand, 13 November 1981; 1 9 (J58835),

paratype. Central Bass Strait, 38 km SW of Cape Paterson, Stn BSS

155, 38°55.5'S 145°17.00'E, 70 m depth, fine sand, 12 November 1981;

all coll. R.S. Wilson.

Description of female. Body (Fig. 87A, B) slender, holotype

2 mm long, 9.7 times as long as wide. Cephalothorax

subrectangular, narrowing anteriorly with slight triangular

rostrum, 1.6 times as long as wide, twice as long as pereonite 1,

naked, eyes absent. Pereonites all naked and rectangular;

pereonites 1 and 5 subequal in length; pereonites 2 to 4

subequal, 1.2 times as long as pereonite 1; pereonite 6 shortest,

0.8 times as long as pereonite 5 (all pereonites respectively 1.3,

0.9, 0.9, 0.9,1.0 and 1.3 times as wide as long). Pleon of five free

subequal pleonites bearing pleopods plus pleotelson; each

pleonite 4.2 times as wide as long. Pleotelson subrectangular,

as long as all pleonites together, 1.25 times as long as wide.

Antennule (Fig. 88A) of four articles, proximal article

nearly 2.6 times as long as wide, as long as distal three articles

together, with single outer distal simple seta surrounded by

four penicillate setae; second article longer than wide, 0.43

times as long as first article, with three outer distal penicillate

setae; third article compact, 0.6 times as long as second article,

naked; fourth article tapering, with six simple and one

penicillate distal setae, and one aesthetasc.

Antenna (Fig. 88B) of six articles, proximal article

compact, fused to cephalothorax; second article 1.3 times as

long as wide, with dorsodistal seta; third article as long as

wide, 0.8 times as long as second article, with dorsodistal seta;

fourth article longest, 4.5 times as long as wide, nearly three

times as long as second article, with penicillate seta in

proximal half and crown of one simple and four penicillate

distal setae; fifth article half as long as second, with one distal

simple seta; sixth article minute with four distal setae.

132

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 87. Araphurapygmothymos sp. nov., female holotype. A, dorsal view; B, lateral view. Scale =1.0 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

133

Fig. 88 .Araphura pygmothymos sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, left mandible; E, right mandible; F, maxillule;

F' maxillule endite spines; G, maxilla; H, labium; I, maxilliped. Scale = 0.1 mm.

134

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 89. Araphura pygmothymos sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5;

G, pereopod 6; H, pleopod; I, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

135

Labrum (Fig. 88C) rounded, hood-shaped, setose. Left

mandible (Fig. 88D) with wide, spade-like crenulate pars incisiva

and linguiform lacinia mobilis, right mandible (Fig. 88E) with

lanceolate pars incisiva and without lacinia mobilis; pars molaris

of both mandibles tapering, with fine distal denticulations.

Labium (Fig. 88H) simple, outer distal corner with unarticulated

setose palp-like projection. Maxillule (Fig. 88F) with nine finely-

denticulate distal spines. Maxilla (Fig. 88G) simple, linguiform,

naked. Maxilliped (Fig. 881) palp first article naked, second

article with one outer and three distal inner setae, third article

with two longer mesial and two shorter distal inner setae, fourth

article with four longer and one shorter distal setae and one small

subdistal outer seta; basis naked; endites distally naked, with

outer-distal microtrichia and paired inner setae.

Cheliped (Fig. 89A) with rounded, naked basis 1.5 times as

long as wide, merus subtriangular with single ventral seta

shorter than width of merus, and covering about half of ventral

margin of carpus; carpus 1.7 times as long as wide, with two

midventral setae, one dorsodistal seta; propodus 1.2 times as

long as wide, with two ventral setae, outer face with curving

ridge of rounded tubercles from mid proximal to dorsodistal,

inner comb-row of two setae; fixed finger with three setae

below cutting edge; dactylus with dorsal rounded tubercles in

proximal half.

Pereopod 1 (Fig. 89B) longer than others, coxa without

apophysis, with seta; basis slender, 4.2 times as long as wide;

ischium compact, with one ventrodistal seta; merus just shorter

than carpus, ventrodistally with seta and longer distally-

denticulate spine exceeding half length of carpus; carpus

distally with ventral seta and single distally-denticulate spines

dorsally and ventrally; propodus 1.5 times as long as carpus,

with ventrodistal spine and dorsodistal spinous apophysis;

dactylus with proximal seta, unguis 1.4 times as long as

dactylus, both together 0.9 times as long as propodus. Pereopod

2 (Fig. 89C) similar to pereopod 1, basis 3.5 times as long as

wide with dorsal penicillate seta; merus longer than carpus;

propodus 1.8 times as long as carpus and with ventral fields of

microtrichia; dactylus and unguis together 0.8 times as long as

propodus. Pereopod 3 (Fig. 89D) similar to pereopod 2.

Pereopod 4 (Fig. 89E) somewhat more compact, basis 3.25

times as long as wide; ischium with two ventrodistal setae;

merus 0.7 times as long as carpus, with two ventrodistal

spines; carpus with four ventrodistal spines; propodus just

longer than carpus, with ventral fields of microtrichia, two

ventrodistal spines and one dorsodistal spine; dactylus about

twice as long as unguis, and with fields of microtrichia,

dactylus and unguis not fused into a claw, the two together 1.2

times as long as propodus. Pereopod 5 (Fig. 89F) as pereopod

4, but with ventral penicillate seta on basis, carpus with

dorsodistal seta. Pereopod 6 (Fig. 89G) as pereopod 5, but

propodus with two ventral and three dorsal distal spines.

Pleopods (Fig. 89H) all alike, with naked basis, endopod

and exopod without setae on inner margin, outer margins with

respectively 8 and 12 plumose setae.

Uropod (Fig. 891) basis naked, exopod 0.6 times as long as

proximal endopod segment, with one mesial and two distal

setae; endopod of two segments, distal segment 0.6 times as

long as proximal segment, setose as figured.

Male. Unknown.

Etymology. From the Greek pygme - a fist, and thymos - a

warty excrescence, referring to the tubercles on the chela (noun

in apposition).

Remarks. The characteristics of the genus Araphura and its

distinctions from the closely related genera Araphuroides Sieg,

1886 and Tanaella Norman & Stebbing, 1886 are discussed by

Sieg (1986a) and Larsen et al. (2009) (but see below): the latter

give a key to the genus Araphura, in which the present species

keys out to A. parabrevimanus. That species, found at >3200 m

off Panama (the record from 720 m in the Subantarctic by

Kudinova-Pasternak, 1975, is highly unlikely), is well figured by

Lang (1968), from which the distinctions of the present shallow-

water Antipodean species, although subtle, can be seen clearly.

In particular, in A. parabrevimanus the dorsal surfaces of

the propodus and dactylus of the chela are smooth, ornamented

with rows of microtrichia, while in A. pygmothymos sp. nov.

these surfaces are highly tuberculate, the line of tubercles on

the propodus extending across the dorsal outer face. Recently,

Bird (2011) has described Araphura whakarakaia from New

Zealand. Also a species with tubercles on the cheliped

propodus and dactylus; as well as the different orientation of

this tuberculation, A. pygmothymos differs from the New

Zealand species in lacking a crenulate cheliped carpus, in its

more elongate pleotelson, and in having stout spines on the

merus of pereopods 1 to 3, inter alia. Other differences

characterizing the present species are the more compact

antennule peduncle articles, the lack of a pseudoarticulation¬

line on the fourth article of the antenna, the stronger distal

spines on the merus and carpus of the pereopods, and the

less-elongate uropodal exopod-process.

Araphura pygmothymos was found in the Central Bass Strait

at depths of 70 to 82 m on fine to muddy sands.

Araphura yarra sp. nov.

Figures 90-92

Material examined. 1 $ (J58836), holotype, Stn BSS 155, Central Bass

Strait, 38 km SW of Cape Paterson, 38°55.5'S 145°17.00'E, 70 m depth,

fine sand, 12 November 1981; coll. R.S. Wilson; 1 9(J58845), paratype,

Stn VC 31 Cl, Central Bass Strait, 38°02.52'S 146°10.47'E, 40 m

depth, 14 May 1999; coll. N. Coleman; 1 9 (J56681), paratype, Stn VC

18 C2, Central Bass Strait, 38°30.2'S 144°15.00'E, 40 m depth, 13 May

1998; coll. N. Coleman. 1 9 (J28487), paratype, Stn MSL-EG 118,

Eastern Bass Strait, 10.8 km E of eastern edge of Lake Tyers,

37°50.92’S 148°12.83'E, 25 September 1990, coll. N. Coleman (depth

not available). 1 9 (J58837), paratype, Stn BSS 197, Western Bass

Strait, 4 km SSW of Currie, King Island, 40°00.7'S 143°49.9’E, 46 m,

21 November 1981, coll. R.S. Wilson,Smith-McIntyre Grab.

Description of female. Body (Fig. 90A, B) slender, holotype

1.9 mm long, 8.8 times as long as wide. Cephalothorax

subrectangular, narrowing anteriorly with slight rounded

rostrum, 1.6 times as long as wide, 2.5 times as long as pereonite

1, naked, eyes absent. Pereonites all naked and rectangular;

pereonites 1 and 6 subequal in length; pereonites 2 to 5

subequal, 1.5 times as long as pereonite 1 (all pereonites

respectively 1.6, 1.0, 1.2,1.0, 0.9 and 1.3 times as wide as long).

136

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 90. Araphura yarra sp. nov., female holotype. A, dorsal; B, lateral; C, cephalothorax, ventral; D, posterior of pleotelson with attachment of

uropods, ventral, showing adjacent tubercles. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

137

Fig. 91. Araphura yarra sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, left mandible; E, right mandible; F, maxillule;

G, maxilliped; H, labium; I, epignath. Scale = 0.1 mm.

138

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 92 .Araphura yarra sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 4; E, pereopod 5; F, pereopod 6; G, pleopod;

H, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

139

Pleon of five free subequal pleonites bearing pleopods, plus

pleotelson; each pleonite 3.6 times as wide as long. Pleotelson

subrectangular, as long as last three pleonites together, 1.2

times as wide as long, without mid-distal process, but with

disto-marginal tubercle adjacent to attachment of each uropod

(Fig. 90D).

Antennule (Fig. 91A) of four articles, proximal article 2.8

times as long as wide, longer than distal three articles together,

with single outer distal simple and penicillate setae; second

article longer than wide, 0.35 times as long as first article,

outer distal margin with one penicillate and one longer simple

setae, latter longer than article; third article as long as wide,

0.6 times as long as second article, with two simple mesial

setae; fourth article tapering, 1.3 times as long as third, with

five simple distal setae.

Antenna (Fig. 91B) of six articles, proximal article

compact, fused to cephalothorax; second article 1.3 times as

long as wide, with two dorsodistal setae; third article as long

as wide, just shorter than second article, with single simple

dorsodistal seta; fourth article longest, 3.6 times as long as

wide, nearly three times as long as third article, with single

distal simple and penicillate seta setae; fifth article as long as

second, with one distal simple seta; sixth article minute with

five distal setae.

Labrum (Fig. 91C) rounded, hood-shaped, naked. Left

mandible (Fig. 91D) with bilobed pars incisiva and tridentate

linguiform lacinia mobilis, right mandible (Fig. 91E) with

tridentate pars incisiva and without lacinia mobilis; pars

molaris of both mandibles stout, with fine distal denticulations,

ventralmost longest. Labium (Fig. 91H) simple, outer distal

corner slightly setose. Maxillule (Fig. 91F) with eight finely-

denticulate distal spines, palp with two relatively short distal

setae. Maxilla not recovered. Maxilliped (Fig. 91G) palp first

article naked, second article with one outer and three distal

inner setae, third article with two longer mesial and two

shorter distal inner setae, fourth article with four distal setae

and one small subdistal outer seta, ventral microtrichia; basis

naked; endites distally with single inner setae but no tubercles.

Epignath (Fig. 911) ribbon-like, naked.

Cheliped (Fig. 92A) sclerite with triangular insertion into

basis; rounded, naked basis 2.2 times as long as wide, with

conspicuous proximal extension but not reaching to pereonite

1 ventrally (Fig. 90C); merus subtriangular with single ventral

seta shorter than width of merus, and covering about one-third

of ventral margin of carpus; carpus nearly twice as long as

wide, dorsal margin convoluted, with one longer and one

shorter midventral setae, one dorsodistal seta and one

dorsoproximal seta; propodus 1.2 times as long as wide, with

two ventral setae, outer face with two submarginal tooth-like

tubercles, inner comb-row of three setae; fixed finger with

three setae below cutting edge, cutting edge with rounded

crenulations; dactylus with dorsal rounded tubercles.

Pereopod 1 (Fig. 92B) not longer than others, coxa without

apophysis, with seta; basis 3.4 times as long as wide, naked;

ischium compact, without seta; merus 1.4 times as long as

carpus, ventrodistally with distally-denticulate spine

exceeding half length of carpus; carpus distally with single

distally-denticulate spines dorsally, mesially and ventrally;

propodus 1.4 times as long as carpus, with subdistal seta,

ventrodistal spine, distal microtrichia and dorsodistal spinous

apophysis; dactylus naked, unguis 1.8 times as long as dactylus,

both together as long as propodus. Pereopod 2 (Fig. 92C)

similar to pereopod 1, basis 3.8 times as long as wide; ischium

with seta; merus 1.2 times as long as carpus; propodus 1.5

times as long as carpus and with ventral fields of microtrichia;

dactylus and unguis together 0.9 times as long as propodus.

Pereopod 3 (not figured) similar to pereopod 2.

Pereopod 4 (Fig. 92D) coxa naked, basis 3.6 times as long

as wide with midventral penicillate seta; ischium with ventral

seta; merus as long as carpus, with two ventrodistal distally-

denticulate spines; carpus with three distal distally-denticulate

spines; propodus 1.3 times as long as carpus, with ventral

fields of microtrichia, three distal distally-denticulate spines,

distal microtrichia and dorsodistal spinous apophysis; dactylus

0.8 times as long as unguis, dactylus and unguis not fused into

a claw, the two together 1.2 times as long as propodus.

Pereopod 5 (Fig. 92E) as pereopod 4, but basis somewhat

stouter, propodus only just longer than carpus. Pereopod 6

(Fig. 92F) as pereopod 5, but basis without penicillate seta,

propodus with two ventral and three dorsal distal spines.

Pleopods (Fig. 92G) all alike, with naked basis, endopod

and exopod without setae on inner margin, outer-distal

margins with respectively 7 and 9 plumose setae, exopod with

additional separated proximal plumose seta.

Uropod (Fig. 92H) basis naked, exopod process half as

long as proximal endopod segment, with three distal setae;

endopod of two segments, distal segment half as long as

proximal segment, setose as figured.

Male .Unknown.

Etymology. Named after the Yarra River which runs through

Melbourne (noun in apposition).

Remarks. Like Araphura pygmothymos (se above), Araphura

yarra sp. nov. keys out to A. brevimanus in the key presented by

Larsen et al. (2009), but is distinguished from that species for

similar reasons, such as the tuberculate dorsal margin of the

cheliped dactylus, the more compact antennule peduncle

articles, the lack of a line of pseudoarticulation on the fourth

article of the antenna, and the stronger distal spines on the

merus and carpus of the pereopods. The present species is

distinguished from the New Zealand species A. whakarakaia by

its different tuberculation of the cheliped, and distinct

spinulation of the pereopods (as in A. pygmothymos ). It is

distinguished from A. pygmothymos by the shorter pereonites,

the more slender cephalothorax, the absence of outer rows of

tubercles on the cheliped propodus, the absence of a seta on the

ischium of pereopod 1, in having only three (rather than four)

spines on the carpi of pereopods 4 to 6, and in the setation of the

uropodal exopod process, inter alia.

Araphura yarra was found in the Central Bass Strait at

depths of 40 to 70 m on fine sand.

Araphura doutagalla sp. nov.

Figures 93-95

140

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 93 .Araphura doutagalla sp. nov., female holotype. A, dorsal; B, lateral. Scale = 0.5 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

141

Fig. 94. Araphura doutagalla sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, right mandible; E, maxillule; F, labium;

G, epignath; H, maxilliped. Scale = 0.1 mm.

142

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 95. Araphura doutagalla sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5;

G, pereopod 6; H, pleopod; I, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

143

Material examined. 1 9 (J58838), holotype, 1 9 (J58848), paratype.

Central Bass Strait, 28 km E of Cape Farewell, King Island, Stn BSS

107, 39°32.8'S 144°16.00'E, 18 m depth, fine sand, 1 November 1980;

coll. M.F. Gomon & G.C.B. Poore.

Description of female. Body (Fig. 93A, B) slender, holotype

1.8 mm long, 7.4 times as long as wide. Cephalothorax pear-

shaped, widest centrally, with distinct rounded rostrum, 1.4

times as long as wide, 2.6 times as long as pereonite 1, naked,

eyes absent. Pereonites rectangular, although pereonites 1,5 and

6 with convex lateral margins, bearing single anterolateral

setae; pereonite 1 short; pereonites 2 to 4 subequal in length, 1.2

times as long as pereonite 1; pereonite 5 shorter, 1.1 times as

long as pereonite 1; pereonite 6 shortest, 0.7 times as long as

pereonite 1 (all pereonites respectively 1.6, 1.4, 1.3, 1.3, 1.4 and

2.3 times as wide as long). Pleon of five free subequal pleonites

bearing pleopods plus pleotelson; each pleonite four times as

wide as long, with single epimeral seta on each side. Pleotelson

subpentangular, twice as long as each pleonite, 1.6 times as wide

as long, with two laterodistal simple and one penicillate setae on

each side of slight rounded mid-distal process (Fig. 951).

Antennule (Fig. 94A) of four articles, proximal article 2.9

times as long as wide, as long as distal three articles together,

with single outer distal simple seta preceded by four penicillate

setae; second article longer than wide, half as long as first article,

with two outer distal penicillate setae and adjacent simple seta

1.4 times as long as article; third article compact, half as long as

second article, with two simple distal setae; fourth article

tapering, with six simple and one penicillate distal setae.

Antenna (Fig. 94B) of six articles, proximal article compact,

fused to cephalothorax; second article as long as wide, with

single dorsodistal and ventrodistal setae; third article as long as

wide, 0.85 times as long as second article, with dorsodistal

seta; fourth article longest, 4.4 times as long as wide, three

times as long as third article, with penicillate seta and two

simple setae distally; fifth article as long as third, with one

distal simple seta; sixth article minute with four distal setae.

Labrum (Fig. 94C) rounded, hood-shaped, naked. Left

mandible not recovered; right mandible (Fig. 94D) with four

rounded “teeth” on pars incisiva, pars molaris stout with fine

distal denticulations, ventral ones elongate. Labium (Fig. 94F)

simple, outer distal corner of both lobes setulate. Maxillule

(Fig. 94E) with eight finely-denticulate distal spines and distal

setules, palp not recovered. Maxilla not recovered. Maxilliped

(Fig. 94H) palp first article naked, second article with one

outer and two distal inner setae, third article with one longer

mesial and three shorter distal inner setae, fourth article with

five distal setae; basis naked; endites distally each with

rounded tubercle and two fine setae, and with outer-distal

microtrichia and longer inner seta. Epignath (Fig. 94G) long,

tapering, ribbon-shaped, naked.

Cheliped (Fig. 95A) sclerite with triangular insertion into

basis, rounded basis 1.6 times as long as wide, with simple

dorsal seta, and with conspicuous proximal extension but not

reaching to pereonite 1 ventrally; merus subtriangular with

single ventral seta longer than width of merus, and covering

about half of ventral margin of carpus; carpus 1.5 times as

long as wide, with two midventral setae, one dorsodistal seta

and one dorsoproximal seta, dorsal margin smooth; propodus

1.2 times as long as wide, with two ventral setae, outer face

with curving ridge of rounded tubercles along fixed finger and

three such tubercles dorsodistally, inner comb-row of three

setae; fixed finger with three setae below cutting edge, cutting

edge with three “teeth”; dactylus with dorsal rounded tubercles

and two fine spinules on cutting edge.

Pereopod 1 (Fig. 95B) not longer than others, coxa without

apophysis, with seta; basis three times as long as wide; ischium

compact, with one ventrodistal seta; merus as long as carpus,

ventrodistally with seta as long as carpus and shorter distally-

denticulate spine exceeding half length of carpus; carpus

distally with mesial seta, distally-denticulate dorsodistal spine

as long as carpus, and shorter simple ventrodistal spine;

propodus 1.3 times as long as carpus, with simple ventrodistal

spine; dactylus naked, unguis 1.8 times as long as dactylus,

both together 1.2 times as long as propodus. Pereopod 2 (Fig.

95C) similar to pereopod 1, basis 3.5 times as long as wide;

carpus with additional distally-denticulate ventrodistal spine;

dactylus and unguis together 0.9 times as long as propodus.

Pereopod 3 (Fig. 95D) similar to pereopod 2.

Pereopod 4 (Fig. 95E) slightly more compact, basis 2.7

times as long as wide; ischium with two ventrodistal setae;

merus as long as carpus, with two ventrodistal distally-

denticulate spines; carpus with four distally-denticulate distal

spines; propodus 1.4 times as long as carpus, with ventral

fields of microtrichia, two ventrodistal distally-denticulate

spines and one dorsodistal distally-denticulate spine; dactylus

1.3 times as long as unguis, dactylus and unguis not fused into

a claw, the two together just shorter than propodus. Pereopod

5 (Fig. 95F) as pereopod 4, but carpus with additional

dorsodistal simple seta, ventrodistal microtrichia evident on

merus and carpus. Pereopod 6 (Fig. 95G) as pereopod 5, but

propodus distally with two ventral and two dorsal distally-

denticulate spines.

Pleopods (Fig. 95H) all alike, with naked basis, endopod

and exopod without setae on inner margin, outer-distal

margins with respectively 7 and 8 plumose setae, exopod with

additional separated proximal plumose seta.

Uropod (Fig. 951) longer than pleotelson, basis naked,

exopod process 0.4 times as long as proximal endopod

segment, with two distal setae; endopod of two segments,

distal segment 0.6 times as long as proximal segment, setose

as figured.

Male. Unknown.

Etymology. “Doutagalla” was used by the European settlers at

Melbourne as one of the early names for the colony: it may have

been a mistranslation of the name of a prominent tribal elder, but

is also said to translate as “treeless plain” (noun in apposition).

Remarks. Araphura doutagalla sp. nov., like the previous two

species, is generally of the A. brevimanus form, but is also

distinguished from that species and from A. whakarakaia as are

those two taxa, and again has tuberculation on the chela. The

patterns of this tuberculation are distinct from those of both A.

pygmothymos and A. yarra\ in addition, A. doutagalla has a

more robust body form (7.4 times as long as wide, compared

with 9.7 or 8.8 times in the other two respectively), and

144

M. Biazewicz-Paszkowycz & R.N. Bamber

conspicuously the uropodal exopod process is less than half the

length of the proximal segment of the endopod (longer than or as

long as half the length respectively). Its relatively stout

mandibular molar process is similar to that of A. yarra (and not

those of A, pygmothymos ), while the proportions of the merus

and carpus of pereopod 1 are like those of A. pygmothymos (and

not those of A, yarra).

Araphura doutagalla was found in the Central Bass Strait

north of Tasmania at a depth of 18 m on fine sand.

Genus Araphuroides Sieg, 1886

Araphuroides stabastris sp. nov.

Figures 96-99

Material examined. 1 9 (J58536), holotype. Central Bass Strait, 33 km

S of Deal Island, Stn BSS 161,39°48.3'S 147°19.2'E, 60 m depth, muddy

sand, 14 November 1981; coll. R.S. Wilson; 1 S (J58544), allotype.

Central Bass Strait, 35 km NNE of Cape Wickham, King Island,

Tasmania, Stn BSS 204 DRC, 39°16.0'S 144°05.4'E, 82 m depth, sandy

shell, 23 November 1981; coll. R.S. Wilson; 2 99 (J58537), paratypes.

Central Bass Strait, 33 km S of Deal Island, Stn BSS 161, 39°48.3'S

147°19.2'E, 60 m depth, muddy sand, 14 November 1981; coll. R.S.

Wilson; 5 99 (J58538), paratypes. Central Bass Strait, 44 km NE of

Cape Wickham, King Island, Stn BSS 203,39°22.0'S 144°18.3'E, 60 m

depth, coarse sand, 23 November 1981; coll. R.S. Wilson; 1 9 (J58541),

paratype. Central Bass Strait, 38 km SW of Cape Paterson, Stn BSS

155, 38°55.5'S 145°17.00'E, 70 m depth, fine sand, 12 November 1981;

coll. R.S. Wilson; 1 9 (J58540), paratype. Eastern Bass Strait, 24 km

NNE of Eddystone Point, Stn BSS 163G, 40°43.9'S 148°32.5'E, 56 m

depth, muddy sand, 14November 1981; coll. R.S. Wilson; 2 99 (J58543),

paratypes. Central Bass Strait, 65 km ENE of Cape Rochon, Three

Hummock Island, Stn BSS 157, 40°10.9'S 145°44.3’E, 75 m depth,

shelly sand, 13 November 1981; coll. R.S. Wilson; 1 9 (J58546),

paratype. Eastern Bass Strait, 100 km NE of North Point, Flinders

Island, Stn BSS 170, 38°52.6’S 148°25.2’E, 130 m depth, fine sand, 15

November 1981; coll. R.S. Wilson; 1 9 (J58548), paratype. Eastern Bass

Strait, 28 km SSW of Mario, Stn BSS 207,37°59.0'S 148°27.0'E, 51 m

depth, muddy sand and fine shell, 30 July 1983; coll. M.F. Gomon; 1 9

(J58539), paratype. Central Bass Strait, 25 km SW of Cape Frankland,

Flinders Island, Stn BSS 162,40°09.4’S 147°32.7'E, 51 m depth, shelly

sand, 14 November 1981; coll. R.S. Wilson; 2 99 (J58542), paratypes.

Western Bass Strait, 40 km SSW of Warrnambool, Victoria, Stn BSS

189,38°42.8'S 142°35.6'E, 69 m depth, coarse sand, 20 November 1981;

coll. R.S. Wilson; 2 99 (J58561), paratypes. Central Bass Strait, 99 km

WSW of Cape Liptrap, Stn BSS 13IT, 39°45.55'S 145°33.82'E to

39°48.03'S 145°32.58'E, 78.7 m depth, very fine sand, 03 February

1981; coll. M.F. Gomon, G.C.B. Poore & C.-C. Lu.

Description of female. Body (Fig. 96A, B) slender, holotype

2.35 mm long, 7 times as long as wide. Cephalothorax

subrectangular, tapering anteriorly with slight triangular

rostrum, 1.3 times as long as wide, longer than pereonites 1 and

2 together, naked, eyes absent. Pereonite 1 shortest, 0.4 times as

long as cephalothorax; pereonites 2 to 6 subequal, half as long

as cephalothorax, with rounded lateral margins and wider than

long (all pereonites respectively 2.0, 1.6, 1.4, 1.3, 1.3 and 1.4

times as wide as long). Pleon of five free subequal pleonites

without pleopods plus pleotelson; each pleonite 3.2 times as

wide as long. Pleotelson pentangular, as long as last pereonite,

with paired posterior setae on each side.

Antennule (Fig. 97A) of four articles, proximal article 1.6

times as long as wide, 1.3 times as long as distal three articles

together, with single outer distal simple seta surrounded by four

penicillate setae; second article just longer than wide, 0.3 times

as long as first article, with outer distal tuft of one simple and

two penicillate setae; third article two-thirds as long as second,

distally with outer simple seta and inner pair of one simple and

one penicillate setae; fourth article tapering, just longer than

second article, with six distal setae and one aesthetasc.

Antenna (Fig. 97B) of six articles, proximal article

compact, fused to cephalothorax; second and third articles as

long as wide, each with dorsodistal seta; fourth article longest,

as long as three proximal articles together and 4.3 times as

long as wide, with two simple and two penicillate distal setae;

fifth article as long as third, with one distal seta; sixth article

minute with four distal setae.

Labrum (Fig. 97C) rounded, hood-shaped, setose. Left

mandible (Fig. 97D) with irregularly denticulate pars incisiva

and triangular, crenulate lacinia mobilis, right mandible (Fig.

97E) with bilobed pars incisiva and without lacinia mobilis;

pars molaris apically tuberculate. Labium (Fig. 97G) simple,

finely setose at outer distal corners, without palp. Maxillule

(Fig. 97F) with eight distal spines, each distally finely-

denticulate, palp (Fig. 97F') with two distal setae. Maxilla

(Fig. 97H) simple, naked. Maxilliped palp (Fig. 971) first

article with single outer distal seta; second article with three

inner setae, two of these distally finely serrated; third article

with two longer and two shorter inner setae; fourth article with

five distal setae, each distally finely serrated, and one outer

subdistal seta; basis with single, long seta almost reaching

distal margin of endites; endites distally with two setae and

rounded tubercle.

Cheliped (Fig. 98A) with rounded, naked basis about twice

as long as wide; merus subtriangular with single ventral seta

and covering more than half of ventral margin of carpus;

carpus 1.5 times as long as wide, with two midventral setae,

one dorsodistal seta; propodus as long as wide, comb-row of

three setae, fixed finger with two ventral and three setae below

cutting edge; dactylus naked.

Pereopod 1 (Fig. 98B) coxa without apophysis, with seta;

basis slender, five times as long as wide, naked; ischium

compact with one ventral seta; merus one-quarter as long as

basis, with slender ventrodistal spine; carpus 1.75 times as

long as merus, distally with inner seta and single distally-

denticulate spine dorsally and two ventrally; propodus 1.25

times as long as carpus, with ventrodistal seta and distal tuft of

setules; dactylus 0.42 times as long as unguis, both together

1.3 times as long as propodus. Pereopod 2 (Fig. 98C) similar

to pereopod 1 although articles proportionately shorter, basis

four times as long as wide with two mid-dorsal penicillate

setae. Pereopod 3 (Fig. 98D) similar to pereopod 2.

Pereopod 4 (Fig. 98E) basis stouter, 3.6 times as long as

wide, with two mid-ventral penicillate setae; ischium with two

ventral setae; merus two-thirds as long as carpus and with two

ventrodistal spines; carpus with three distal spines; propodus

1.14 times as long as carpus, with dorsal penicillate seta and

three distal spines, each with fine serrations; dactylus longer

than distinct unguis, both together just longer than propodus.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 96 . Araphuroides stabastris sp. nov., female holotype. A, dorsal view; B, lateral view. Scale = 0.1 mm.

146

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 97 . Araphuroides stabastris sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, left mandible; E, right mandible; F, maxillule;

F' maxillule palp; G, labium; H, maxilla; I, maxilliped. Scale = 0.2 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

147

Fig. 98. Araphuroides stabastris sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5;

G, pereopod 6; H, uropod. Scale = 0.2 mm.

148

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 99 . Araphuroides stabastris sp. nov., male allotype. A, dorsal view ; B, lateral view ; C, antennule ; D, pleopod. Scale = 1 mm for A (and B),

0.1 mm for C and D.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

149

Pereopod 5 (Fig. 98F) as pereopod 4, but all spines showing

fine serrations, microtrichia present on propodus and dactylus.

Pereopod 6 (Fig. 98G) as pereopod 5, but propodus with four

distal spines, and basis without penicillate setae.

Pleopods absent.

Uropod (Fig. 98H) basis naked, exopod process 0.7 times

as long as endopod, with one mesial and one distal setae;

endopod of one segment, with subdistal tuft of two penicillate

setae and distal group of five simple and one penicillate setae.

Distinctions of male (Fig. 99). Generally similar to female,

allotype body length 2.9 mm; pereonite 2 only half as long as

wide. Antennule with incipient articulation of distal article.

Five pairs of pleopods present, rami subequal in length, exopod

with nine distal setae, endopod with eight distal and one inner

subdistal setae.

Etymology. Stabastris is an anagram of Bass Strait.

Remarks. The genus Araphuroides is distinguished from

Araphura and Tanaella in having pereonites laterally rounded

and wider than long (unlike Araphura ), with a distinct uropodal

exopod-process (unlike Tanaella ), but see discussion below.

Larsen (2005) re-diagnosed the genus and gave a key to the

four species then known, in which A. stabastris sp. nov. is not

resolved. Since that paper, A. io (Bamber, 2005) was transferred

to this genus from Araphura by Larsen et al. (2009), and the

present species is indeed closest to A. io, a species recorded

from south-western Western Australia at depths from 23 to

40 m.

Araphuroides stabastris is distinguished from A. io in

having a shorter cephalothorax; a cheliped with two ventral

carpal setae, and only three setae in the chela comb-row;

pereopod 1 with a more elongate basis (five times as long as

wide, only three times in A. io), a carpus approaching twice as

long as the merus (subequal in A. io), and a proportionately

longer unguis (2.4 times as long as dactylus, only 1.3 times in

A. io); all pereopods with a proportionately longer dactylus

plus unguis, proportionately longer carpus, and longer distal

spines on the merus and carpus. In addition, A. io has pleopods

in the female. Both species are without a line of pseudo¬

articulation on the fourth antennal article found in all other

species apart from A. bombus Larsen, 2005.

Araphuroides stabastris was found at 31 to 71 m depth on

muddy to shelly sands throughout the Bass Strait.

Araphuroides batmania sp. nov.

Figures 100-101

Material examined. 1 ? (J58572), holotype, 4 $9 (J58904), paratypes.

Central Bass Strait, 28 km E of Cape Farewell, King Island, Stn BSS

107S, 39°32.8'S 144°16.0'E, 18 m depth, fine sand, 1 November 1980;

25 99 (J58571), paratypes. Central Bass Strait, 35 km E of Cape

Farewell, King Island, Stn BSS 108G, 39°32.8'S 144°21.0'E, 27 m

depth, fine sand, 01 November 1980; all coll. M.F. Gomon & G.C.B.

Poore.

Description of female. Body (Fig. 100A, B) slender, holotype

3.2 mm long, 6.6 times as long as wide. Cephalothorax

subrectangular, 1.3 times as long as wide, shorter than

pereonites 1 and 2 together, with single anterolateral seta on

each side, eyes absent. Pereonites rectangular, mostly with

convex lateral margins; pereonite 1 longest, 0.6 times as long as

cephalothorax; pereonites 2 to 5 subequal, half as long as

cephalothorax; pereonite 6 shortest, half as long as pereonite 1

(all pereonites respectively 1.25, 1.5, 1.6, 1.5, 1.5 and 2.4 times

as wide as long). Pleon with five free subequal pleonites bearing

pleopods; each pleonite 4.8 times as wide as long. Pleotelson

semicircular, as long as last two pleonites together, with one

posterolateral seta on each side.

Antennule (Fig. 100C) of four articles, proximal article 2.7

times as long as wide, just longer than distal three articles

together; second article 1.5 times as long as wide, 0.4 times as

long as first article; third article shorter than wide, 0.4 times as

long as second article; fourth article tapering, twice as long as

third article, with four distal setae and one aesthetasc.

Antenna (Fig. 100D) of six articles, proximal article

compact, fused to cephalothorax; second article longer than

wide, with dorsodistal seta; third article as long as wide, with

dorsodistal seta; fourth article longest, 4.7 times as long as

wide, with midventral and ventrodistal penicillate setae and

ventrodistal simple seta; fifth article 0.4 times as long as fourth

with one distal seta; sixth article minute with four distal setae.

Labrum (Fig. 100E) hood-shaped, naked. Left mandible

(Fig. 100F) with narrow, crenulate lacinia mobilis, cutting-

edge angled; pars incisiva truncate with prominent

crenulations. Right mandible (Fig. 100G) without lacinia

mobilis, pars incisiva pointed with inner crenulations; pars

molaris of both mandibles distally with strong, rounded tooth¬

like protrusions. Labium not recovered. Maxillule (Fig. 100H,

H') with nine distal spines, at least some of these finely

denticulate, and fine distal setules, palp with two distal setae.

Maxilla (Fig. 100H) ovoid, simple, naked. Maxilliped palp

(Fig. 1001) first article with simple outer seta, second and third

articles with three inner setae, fourth article with four distal,

finely denticulate spines and one outer subdistal seta; basis

naked; endites distally with single seta, outer group of setules

and slight inner tubercle.

Cheliped (Fig. 101A) naked basis attached to substantial

sclerite; merus subtriangular with single ventral seta; carpus

1.5 times as long as wide, with two midventral and one

dorsodistal setae; propodus with row of rounded tubercles

along outer ventral margin including fixed finger, with two

ventral setae, three setae on crenulate cutting edge; dactylus

dorsal margin with rounded tuberculation.

Pereopod 1 (Fig. 101B) coxa with seta; basis 3.9 times as

long as wide, naked; ischium compact with ventral seta as long

as merus; merus just longer than carpus, with fine dorsodistal

seta, ventrodistally with slender spine denticulate in distal two-

thirds and simple seta; carpus distally with strong dorsodistal

spine, smaller ventrodistal spine and mid-distal seta; propodus

1.4 times as long as carpus, with short ventrodistal spine; dactylus

half as long as slender unguis, both together as long as propodus.

Pereopod 2 (Fig. 101C) similar to pereopod 1, but basis with

proximal seta, merus with stouter ventrodistal spine but without

dorsodistal seta, carpus with longer ventrodistal spine, propodus

with dorsodistal seta. Pereopod 3 (Fig. 101D) similar to pereopod

2, but basis naked.

150

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 100. Araphuroides batmania sp. nov. A, holotype female dorsal; B, holotype female lateral; C, antennule; D, antenna; E, labrum; F, left

mandible; G, right mandible; H, maxillule and maxilla; H', maxillule palp; I, maxilliped. Scale: A-B = 1 mm, C-H = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

151

Fig. 101. Araphuroides batmania sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5;

G, pereopod 6; H, pleopod; I, uropod; I', uropod basis, lateral.

152

M. Biazewicz-Paszkowycz & R.N. Bamber

Pereopod 4 (Fig. 101E) basis stouter than those of anterior

pereopods, 3.2 times as long as wide, with ventral penicillate

seta; ischium with two ventral setae; merus and carpus

subequal, merus 0.6 times as long as carpus and with two

ventrodistal spines, carpus with distal crown of four denticulate

spines; propodus 1.3 times as long as carpus, with mid-dorsal

penicillate seta, one dorsodistal and two ventrodistal spines;

dactylus longer than unguis. Pereopod 5 (Fig. 101F) as

pereopod 4, but basis naked. Pereopod 6 (Fig. 101G) as

pereopod 5, but propodus with three dorsodistal setae.

Pleopods (Fig. 101H) all alike, with naked basis, endopod

and exopod without setae on inner margin, outer distal margins

with respectively 7 and 12 plumose setae, additional proximal

exopod seta separated from others.

Uropod (Fig. 1011) exopod process 0.4 times as long as

proximal endopod segment, with three distal setae; endopod of

two segments, proximal segment with one simple and two

penicillate distal setae, distal segment half as long as proximal

segment and with four simple and three penicillate distal setae.

Male. Unknown.

Etymology. Batmania was one of the names of the early

settlement at Melbourne (ca. 1835), named after John Batman,

a leading member of the Tasmanian Port Phillip Association.

Remarks. For reasons given above under Araphuroides

stabastris, the present species is again closest only to A. io;

unlike the previous species, A. batmania sp. nov. also has on

pereopod 1 the shorter carpus, shorter distal merus and carpus

setae, and proportionately shorter dactylus plus claw of A. io,

and again shares the absence of a line of pseudo-articulation on

the fourth antennal article. Like A. io, but unlike A. stabastris,

the female of the present species has pleopods.

Araphuroides batmania is distinguished from A. io by its

longer first pereonite on a generally stouter body, fewer setae

on the maxilliped palp, and a mandible with more complex

crenulation on the pars incisiva and rounded distal tubercles

on the pars molaris, and is distinguished from A. io, and all

other described species in the genus, in having rounded

tubercles on both fingers of the chela.

Araphuroides batmania was taken in the Central Bass

strait east of Cape Farewell, King Island at depths of 18 to

27 m in fine sand.

Araphuroides sala sp. nov.

Figures 102-104

Material examined. 1 9 (J58841), holotype. Western Port, off Crib

Point, Stn CPBS-N 52/272, 38°19.92'S 145°13.95'E, 19 m depth, sand

and gravel, 31 March 1965; coll. A.J. Gilmour. 1 9 (J58843), paratype.

Western Port, off Crib Point, Stn CPBS-N 25/1, 38°20.25’S

145°14.68'E, 11 m depth, sand, 10 March 1965; coll. A.J. Gilmour. 1 9

(J58839), Stn CPBS-N 32/1, 1 9 (J58840), Stn CPBS-N 32/2, and 4 99

(J58842), Stn CPBS-N 32/3, paratypes, all Western Port, off Crib

Point, 38°20.83'S 145°13.48'E, 13 m depth, sandy gravel, 23 March

1965; coll. A.J. Gilmour. 2 99 (J58534, J58535), paratypes. Eastern

Bass Strait, 63 km E of North Point, Flinders Island, Stn BSS 167,

39°44.8'S 148°40.6'E, 124 m depth, muddy sand, 14 November 1981,

coll. R.S. Wilson.

Description of female. Body (Fig. 102A, B) slender, holotype

1.4 mm long, 8.1 times as long as wide. Cephalothorax pear-

shaped, widest centrally, with distinct rounded rostrum, 1.6

times as long as wide, 2.7 times as long as pereonite 1, naked,

eyes absent. Pereonites all naked, lateral margins convex;

pereonite 1 short, 0.4 times as long as cephalothorax; pereonites

2 to 5 subequal, 1.25 times as long as pereonite 1; pereonite 6

shortest, 0.8 times as long as pereonite 1 (all pereonites

respectively 1.5, 1.25, 1.25, 1.5, 1.3 and 2.0 times as wide as

long). Pleon of five free subequal pleonites bearing pleopods plus

pleotelson; each pleonite 4.4 times as wide as long, with single

lateral epimeral seta on each side. Pleotelson subpentangular, 0.4

times as long as whole pleon, as long as wide, paired laterodistal

setae either side of rounded mid-distal margin.

Antennule (Fig. 103A) of four articles, proximal article 3.6

times as long as wide, longer than distal three articles together,

distally with two tufts of penicillate setae and single outer

simple seta; second article longer than wide, 0.4 times as long

as first article, with four inner distal penicillate setae and one

outer simple seta; third article compact, half as long as second

article, with two simple distal setae; fourth article tapering, 1.4

times as long as third article, with six simple and one

penicillate distal setae.

Antenna (Fig. 103B) of six articles, proximal article

compact, shorter than wide, fused to cephalothorax; second

article as long as wide, with two distal setae; third article

shorter than wide, 0.7 times as long as second article, with

dorsodistal seta; fourth article longest, five times as long as

wide, more than three times as long as second article, with

penicillate seta in proximal half, crown of one simple and five

penicillate distal setae and dorsal rows of microtrichia; fifth

article as long as second, with one distal simple seta; sixth

article minute with four distal setae.

Labrum (Fig. 103C) rounded, hood-shaped, distally setose.

Left mandible (Fig. 103D) with wide, spade-like crenulate pars

incisiva and triangular, crenulate lacinia mobilis, right

mandible (Fig. 103E) with crenulate and distally bifid pars

incisiva and without lacinia mobilis; pars molaris of both

mandibles stout, with fine, elongate distal denticulations.

Labium (Fig. 103H) simple, outer distal corner of both lobes

setulose. Maxillule (Fig. 103F) with nine finely-denticulate

distal spines and few distal setules, palp with two distal setae.

Maxilla (Fig. 103G) simple, triangular, naked. Maxilliped

(Fig. 1031) palp first article naked, second article with one

outer and three distal inner setae, two of these finely

denticulate; third article with two longer mesial and two

shorter distal inner setae, one of each of these finely denticulate;

fourth article with four longer and one shorter distal finely-

denticulate setae and one small subdistal outer seta; basis

naked; endites distally each with two rounded tubercles, with

outer-distal microtrichia and inner seta. Epignath (Fig. 103J)

slender, ribbon-like, distally pointed, naked.

Cheliped (Fig. 103K) with rounded, naked basis 2.3 times

as long as wide, merus subtriangular with single ventral seta,

and covering about half of ventral margin of carpus; carpus 1.5

times as long as wide, with one longer and one much shorter

midventral setae, one dorsodistal seta and one dorsoproximal

seta, dorsal margin smooth; propodus just longer than wide,

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 102 . Araphuroides sala sp. nov., female holotype, dorsal. Scale = 0.5 mm.

154

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 103. Araphuroides sala sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, left mandible; E, right mandible; F, maxillule;

G, maxilla; H, labium; I, maxilliped; J, epignath; K, cheliped. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

155

Fig. 104. Araphuroides sala sp. nov., female paratype. A to F, pereopods 1 to 6 respectively; G, pleopod; H, uropod; H' pleotelson, dorsal (distal

setae not shown). Scale = 0.1 mm.

156

M. Biazewicz-Paszkowycz & R.N. Bamber

with two ventral setae, dorsally with two rows of rounded

tubercles in distal half, outer face with row of rounded tubercles

along ventral margin of fixed finger, inner comb-row of four

setae; fixed finger with three setae below cutting edge and two

or three small tooth-like apophyses on cutting edge; dactylus

with rounded tubercles along dorsal margin.

Pereopod 1 (Fig. 104A) not longer than others, coxa

without apophysis, with seta; basis 2.9 times as long as wide,

with dorsoproximal penicillate seta; ischium compact, with

one ventrodistal seta almost as long as merus; merus 1.2 times

as long as carpus, ventrodistally with seta and longer distally-

denticulate spine just exceeding half length of carpus; carpus

distally with shorter ventral spine and single distally-

denticulate dorsodistal spines anteriorly and posteriorly;

propodus 1.25 times as long as carpus, with ventrodistal spine

and dorsodistal and ventral microtrichia; dactylus naked,

unguis 1.5 times as long as dactylus, both together 1.4 times as

long as propodus. Pereopod 2 (Fig. 104B), similar to pereopod

1, basis with two dorsoproximal penicillate setae; merus and

carpus subequal in length; propodus 1.5 times as long as

carpus; dactylus and unguis together 1.2 times as long as

propodus. Pereopod 3 (Fig. 104C) similar to pereopod 2.

Pereopod 4 (Fig. 104D) not more compact, coxa naked,

basis three times as long as wide, with two dorsoproximal and

two midventral penicillate setae; ischium with one shorter and

one longer ventrodistal setae, latter as long as merus; merus just

shorter than carpus, with two finely-denticulate ventrodistal

spines; carpus with single dorsodistal and ventrodistal setae and

three finely-denticulate ventrodistal spines, distally with

microtrichia; propodus 1.2 times as long as carpus, with ventral

fields of microtrichia, dorsal penicillate seta, two ventrodistal

and one dorsodistal finely-denticulate spines; dactylus 0.8 times

as long as unguis, both with fields of microtrichia, dactylus and

unguis not fused into a claw, the two together 1.1 times as long

as propodus. Pereopod 5 (Fig. 104E) as pereopod 4, but basis

without penicillate setae. Pereopod 6 (Fig. 104F) as pereopod 5,

but propodus with two ventral and three dorsal distal spines.

Pleopods (Fig. 104G) all alike, with naked basis, endopod

with subdistal inner plumose seta, exopod without setae on

inner margin, outer margins with respectively 5 and 12 plumose

setae, additional proximal exopod seta separated from others.

Uropod (Fig. 104H, H') half as long as pleotelson, basis

naked, exopod process half as long as endopod, with three

distal setae; endopod of one segment, setose as figured.

Male. Unknown.

Etymology. Named after the English journalist George

Augustus Henry Sala who, during a visit to Victoria in 1885,

coined the phrase “Marvellous Melbourne”, which stuck long

into the twentieth century and is apparently still used today by

Melburnians (noun in apposition).

Remarks. Within the genus Araphuroides, only the three species

described herein and A. bombus Larsen, 2005 are without a

pseudo-articulation line on the fourth antennal article, and only

the present species and A. batmania have rounded tubercles on

the chela; these two also share the elongate setae on the ischia of

the pereopods. A. batmania differs from A. sala sp. nov. in

being without the dorsodistal tubercles on the chela, as well as

in having only rounded distal tubercles on the mandibular molar

process, and in having a uropod with a longer exopod process

and a two-segmented endopod.

Araphuroides sala was found in Western Port and off

Flinders Island in coarse to muddy sands at depths between 11

and 124 m.

Discussion of the genera Araphura and Araphuroides.

Sieg (1986a) originally distinguished Araphuroides from

Araphura by their “body shape”, the pereonites of Araphura

having parallel margins while those of the two species he

attributed to Araphuroides (see above) having “gently rounded”

margins; additionally, pereonite 2 in Araphura is “normally” as

long as wide or slightly wider than long, while it is “distinctly

broader than long” in Araphuroides ; further, he maintained that

the merus on pereopod 1 is short, “only slightly longer than

broad”, in Araphura, but “distinctly longer than broad” in

Araphuroides. Finally, the pars molaris of the mandible is broad,

with “at least one longer and several small toothlike processes”

in Araphuroides, but is pointed “ending in three or four tiny

tips” in Araphura.

Sieg & Dojiri (1989) expanded on these distinctions, citing

pereonite 2 as being “at least as long as broad, but mostly

longer than broad” in Araphura; strangely, these authors

showed the features of the ratio of length to width of pereonite

2, and of parallel- or convex-sided pereonites to be

ontogenically variable. Larsen (2005) and Larsen et al. (2009)

added the body length to width proportions as distinguishing

these two genera ( Araphura 9 to 13 times as long as wide,

Araphuroides less than nine times), even though this feature was

not cited by Sieg (locc. cit.).

These various features for the Australian species of these

two genera are shown in Table 2. None of the species has a

pereopod 1 merus “only slightly longer than broad”. Other than

this, it is apparent that the only species agreeing wholly with

Sieg’s concept of Araphura is A. pygmothymus, and it is the

only one with a “pointed” mandibular molar process. Yet, from

their remaining morphology, A. yarra is clearly a species close

to A. pygmothymus. Indeed, a number of these species appear

to be siblings. Considering Larsen’s (2005) concept of body

length to width, these Australian species show a gradation

suggesting that any such distinction is entirely arbitrary.

Indeed, it is apparent that the “characters” diagnosing these

genera according to Sieg (locc. cit.) are not consistent, and

consideration of the features shown in table 2 as being diagnostic

is to fall into the error or classifying characters rather than

animals. To quote Linnaeus, “Characters come from the genus,

not the genus from the characters. Characters are not there so

that there should be a genus but in order that the genus should be

recognized.” (Linne, 1751; see also Mayr, 1969). One might be as

justified in using the tuberculation on the chela found here (and

in A. whakarakaia) in all three species named in Araphura plus

two of the Araphuroides species; however, this can hardly be a

generic character, as it is also present in another and quite distinct

tanaellid genus, described below, as well as some species of the

unrelated genera Chauliopleona Dojiri & Sieg, 1997 and

Akanthophoreus Sieg, 1986.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

157

Table 2. Characters of the Australian species of Araphura and Araphuroides, together with those defined as characterizing the genera by Sieg &

Dojiri (1989). *data from Larsen (2005), who further distinguished these genera on body proportion, but note that Sieg (1986a) and Sieg & Dojiri

(1989) did not.

Body

length:

width

pereonite

2 width:

length

pereopod

1 merus

length:

width

pereopod

1 merus:

carpus

mandible

molar

process

mandibular

molar

ventrodistal

slender

spines

lateral

borders of

pereonites

Tuberculation

on the chela

Araphura pygmothymus

9.7

0.9

1.7

-1.0

tapering

absent

parallel

present

Araphura yarra

8.8

2.1

1.4

stout

present

~ parallel

present

Araphura doutagalla

7.4

1.4

1.7

-1.0

stout

present

~ parallel

present

Araphuroides stabastris

1.6

1.3

0.6

stout

absent

convex

absent

Araphuroides batmania

6.6

1.5

1.7

-1.0

stout

absent

~convex

present

Araphuroides sala

8.1

1.25

1.7

1.2

stout

present

convex

present

Araphura io Bamber, 2005

7.3

1.5

1.6

-1.0

stout

present

~ parallel

absent

Araphura sensu Sieg &

Dojiri, 1989

9 to 13*

< 1.0

-1.0

tapering

absent

parallel

Araphuroides sensu Sieg &

Dojiri, 1989

<9.0*

>1.0

stout

present

convex

The right conclusion is probably to dismiss Araphuroides

as a distinct genus, but that must necessitate a reanalysis of all

24 species attributed to these two genera (including those

described herein). At present, the three species above attributed

to Araphuroides are distinguished simply on their convex (or

relatively convex) lateral pereonite margins. On that basis,

despite its being apparently close to the Araphuroides species

described above, Araphura io is returned to its original genus.

Genus Inconnivus gen. nov.

Diagnosis, female. Tanaellid with cephalothorax showing

lateral concavity towards anterior. Eyelobes present.

Antennule with four articles, antenna with six articles,

proximal article fused to cephalothorax; second and third

articles with slender dorsodistal spines. Mandibular pars

molaris longer than pars incisiva, with distal ring or spines,

longer ventrally. Pereopods, chelipeds, maxillipeds with

microtrichia; merus and carpus of all pereopods with spines.

Dactylus and unguis of pereopods distinct, as long as or

longer than propodus; distal propodal spine of pereopods 2

and 3 coaxial with dactylus. Pleopods present. Uropods stout,

exopod present as small process fused to basis; endopods

short, the two not configured in the form of “pincers”.

Type species. Inconnivus billibunteri sp. nov. by monotypy.

Etymology, from the Latin “that does not close the eyes”,

alluding to the presence of distinct eyelobes in a taxon otherwise

hardly distinct from the eyelobe-less genus Tanaella Norman

and Stebbing, 1886; noun derived from the adjective, male.

Remarks. With the anterolateral concavity to the cephalothorax,

the conformation of the antennules, antennae, pereopods,

pleopods, uropods and mouthparts, the species described

below shows a very close affinity to Tanaella. The uropods are

very short for a Tanaella, and clearly not “pincer-like” (see

diagnosis of Tanaella by Larsen & Heard, 2004b), although

this configuration is also approached by, for example, T. kroyeri

Larsen et al., 2009. However, the present species takes this

reduction in the uropods much further, and, most distinctly,

has evident eyelobes, although no ocelli were observed in the

preserved material: Larsen and Heard (2004b) included a lack

of eyelobes in their generic diagnosis for Tanaella, and Larsen

(2005) considered it one diagnostic feature of the Tanaellidae.

The species below shows some superficial similarities to

the Cryptocopinae, but the conformation of the uropods,

antennae, maxilliped endites and chelipeds, and the pereopod

spination all suggest otherwise.

This species is therefore attributed to a separate genus,

closely related to Tanaella, but with the presence of eyelobes,

probably more plesiomorphic. It is not possible at present to

say whether the additional features of the tuberculate rugosity

on the cheliped or the microtrichia on the pereopods are

generic or specific characters.

Inconnivus billibunteri sp. nov.

Figures 105-107

Material examined. 1 ? (J37873), holotype, off Nowra, New South

Wales, Stn SLOPE 1, 34°59.52'S 151°05.93'E, 204 m depth, 14 July

1986; coll. G.C.B. Poore; 1 ? (J58564), paratype. Central Bass Strait, 25

km SW of Cape Frankland, Hinders Island, Stn BSS 162, 40°09.4'S

147°32.6’E, 51 m depth, shelly sand, 14 November 1981; coll. R.S.

Wilson; 1 $ (J58903), paratype, Eastern Bass Strait, 100 km NE of North

Point, Hinders Island, Stn BSS 170,38°52.6'S 148°25.2'E, 130 m depth,

fine sand, 15 November 1981; coll. R.S. Wilson.

Description of female. Body (Fig. 105) compact, holotype

2.9 mm long, 4.8 times as long as wide. Cephalothorax narrowing

towards anterior, as long as wide, about as long as pereonites 1 to

158

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 105. Inconnivus billibunteri sp. nov. A, holotype female dorsal view. Scale = 1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

159

Fig. 106 . Inconnivus billibunteri sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, left mandible; E, right mandible; F, maxillule

endite; G, maxilliped (distal palp article damaged); H, labium. Scale = 0.1 mm.

160

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 107. Inconnivus billibunteri sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5;

G, pereopod 6; H, pleopod; I, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

161

4 together, naked; eyelobes present, eyes absent. All pereonites

laterally convex; pereonites 1 to 3 subequal in length, narrow,

less than one-quarter as long as cephalothorax; pereonite 4

longest, 1.5 times as long as pereonite 3; pereonites 5 and 6

subequal, 0.9 times as long as pereonite 4 (all pereonites

respectively 4, 4.4, 3.8, 2.3, 2.7 and 2.9 times as wide as long).

Five free subequal pleonites bearing pleopods; each pleonite 6.4

times as wide as long. Pleotelson semicircular, as long as

maximum width, longer than last three pleonites together, naked.

Antennule (Fig. 106A) of four articles, proximal article

2.4 times as long as wide, 1.3 times as long as distal three

articles together, with mid-dorsal row of two simple and about

eight penicillate setae; second article slightly longer than

wide, one-third as long as first article, with distal tuft of one

simple and four penicillate setae; third article compact, one-

third as long as second article, with two simple distal setae;

fourth article 2.5 times as long as third with six simple distal

setae and one aesthetasc.

Antenna (Fig. 106B) proximal article compact, fused to

cephalothorax; second article as long as wide, with distal seta

and dorsodistal spinule; third article as long as wide, with

dorsodistal seta; fourth article longest, 3.9 times as long as wide,

with suggestion of secondary articulation just proximal of mid¬

length, and with one penicillate seta just proximal of mid-length

and group of three simple and three penicillate distal setae; fifth

article 0.3 times as long as fourth with one simple distal seta;

sixth article minute with four distal setae.

Labrum (Fig. 106C) rounded, distally setose. Left mandible

(Fig. 106D) with narrow, irregularly-crenulate pars incisiva,

lacinia mobilis triangular with fine outer denticulations, right

mandible (Fig. 106E) as left but without lacinia mobilis; pars

molaris of both mandibles robust with slender ventrodistal

spines. Labium (Fig. 106H) simple, elongate, with outer-distal

spinule. Maxillule (Fig. 106F) with nine distal spines, six of

these slender, scythe-like with fine denticulations; palp not

recovered. Maxilliped palp (Fig. 106G) with groups of

microtrichia; first article naked; second article with outer distal

seta and three longer inner-distal setae; third article with four

inner setae, longest seta much longer than article; fourth article

damaged in preparation; basis with single, long seta not reaching

distal margin of endites; endites distally with two oval tubercles.

Epignath not recovered.

Cheliped (Fig. 107A) sclerite with triangular attachment to

basis; basis just longer than wide and bearing microtrichia;

merus subtriangular with single ventral seta; carpus rounded,

compact, 1.1 times as long as wide, with two midventral setae,

one dorsoproximal and one dorsodistal setae; propodus as

long as carpus, with comb-row of four setae, fixed finger with

two ventral and one inner setae, three setae on cutting edge,

cutting edge crenulate; dactylus with robust row of rounded

crenulations along outer dorsal margin, cutting edge with

three coarser crenulations.

Pereopod 1 (Fig. 107B) coxa naked; basis slender, 4.5 times

as long as wide, with mid-ventral microtrichia and ventrodistal

seta; ischium compact with long ventral seta 0.85 times as long

as merus; merus, carpus and propodus subequal in length; merus

with midventral microtrichia and ventrodistal spine; carpus with

microtrichia, ventrodistal spine, two inner-distal setae and long

dorsodistal spine, 0.75 times as long as propodus and with inner

denticulation; propodus with ventral microtrichia, subdistal

dorsal seta and ventrodistal spine longer than dactylus; dactylus

and claw subequal in length, both together 1.15 times as long as

propodus. Pereopod 2 (Fig. 107C) similar to pereopod 1, basis

four times as long as wide, without ventrodistal seta; carpus with

two ventrodistal spines, dorsodistal spine half as long as

propodus; propodus 1.3 times as long as carpus, without dorsal

seta; dactylus with proximal seta. Pereopod 3 (Fig. 107D) similar

to pereopod 2 but carpus and propodus subequal in length and

1.4 times as long as merus.

Pereopod 4 (Fig. 107E) basis stout, 2.5 times as long as wide,

with two penicillate setae; ischium with ventrodistal seta as long

as merus; merus shorter than carpus, with two finely-denticulate

ventrodistal spines; carpus with three finely-denticulate

ventrodistal spines and slender dorsodistal blunt spine; propodus

1.2 times as long as carpus, with dorsal penicillate seta, ventral

rows of microtrichia, and three distal spines almost as long as

dactylus; dactylus and claw subequal in length, curved, both

together 1.3 times as long as propodus. Pereopod 5 (Fig. 107F) as

pereopod 4, but seta on ischium longer than merus, merus and

carpus subequal in length, distal setae on propodus longer than

dactylus. Pereopod 6 (Fig. 107G) as pereopod 4, but basis more

slender (3.25 times as long as wide), propodus with four distal

finely-denticulate spines and one simple seta.

Pleopods (Fig. 107H) all alike, with naked basis, endopod

shorter than exopod, both without setae on inner margin, outer

margins with respectively 7 and 14 plumose setae.

Uropod (Fig. 1071) short; basis naked and with minute

rounded exopodal process bearing two long and one short

distal setae; endopod of one segment just longer than basis,

with two penicillate setae at mid-length and array of five simple

distal setae.

Male. Unknown.

Etymology, named after William George (“Billy”) Bunter, a

proportionately-fat schoolboy character in books written by

Charles Hamilton using the pen-name Frank Richards.

Remarks. The characterizing features of this species, which

distinguish it from the related genus Tanaella, particularly the

presence of eyelobes, are described above under the generic

remarks. Further, it is much less slender than species of Tanaella.

In comparison with those species, Inconnivus billibunteri sp.

nov. would key out to T. mclellandi Larsen and Heard, 2004b, in

the key to the genus given by Guerrero-Kommritz & Blazewicz-

Paszkowycz (2004) but the antennule is distinct, the uropods

too short, the chela smaller, the propodi of pereopods 1 and 2

proportionately shorter, and pleopods are present.

Since that publication, two further species of Tanaella

have been described. T. kommritzia Larsen & Shimomura,

2007(a) has much longer (and incurved) uropods, inter alia,

and is distinctly more slender. The only species of Tanaella

recorded from Australia, T. dongo Bamber, 2005, is the

slenderest of the genus, and has a 2-segmented uropod.

Neither, of course, has eyelobes.

Inconnivus billibunteri was collected sporadically at

depths from 51 to 204 m on shelly to fine sand substrata.

162

M. Biazewicz-Paszkowycz & R.N. Bamber

Family Mirandotanaidae Biazewicz-Paszkowycz & Bamber,

2009

Genus Pooreotanais Biazewicz-Paszkowycz & Bamber, 2009

Pooreotanais gari Biazewicz-Paszkowycz & Bamber, 2009

Pooreotanais gari Biazewicz-Paszkowycz & Bamber, 2009,7-11,

figs 1-3.

Remarks. Pooreotanais gari was described from a male and a

number of females and subadults collected from Western Port,

Victoria, at depths between 13 and 18 m on a variety of

substrata. The genus, and indeed the family Mirandotanaidae,

is characterized by having a grossly inflated posterior half (or

more) of the body, including the pleon and at least pereonite 6

(pereonites 4 to 6 in the present species). The function of this

inflated posterior is unknown, but is unlikely to be related to

reproduction as it is also shown by the male.

The only other species of Pooreotanais, P. ningaloo, is

from Western Australia, and is distinct in having only

pereonite 6 inflated, and the pleotelson longer than any

pleonites (shorter in P. gari), inter alia.

Family Typhlotanaidae Sieg, 1984

Genus Typhlotanais Sars, 1882 sensu lato

Typhlotanais herthio sp. nov.

Figures 108-110

Material examined. 1 9 (J58514), holotype. Eastern Bass Strait, 60 km

E of North Point, Flinders Island, Stn BSS 32, 39°41.7'S 148°39.5'E,

115 m depth, muddy sand, 27 March 1979; coll. G.C.B. Poore; 45 99

and neuters (J58515), paratypes. Central Bass Strait, 32 km SE of Cape

Otway, Stn BSS 48DN, 39°01'S 143°49’E, 81 m depth, coarse sand, 07

October 1980; coll. G.C.B. Poore; 41 9? and neuters (J58518),

paratypes. Central Bass Strait, 66 km S of Rodondo Island, Stn BSS

158S, 39°48.6'S 146°18.8'E, 82 m depth, sand with silt and mud, 13

November 1981; coll. R.S. Wilson.

Description of female. Body (Fig. 108A, B) slender, holotype

2.7 mm long, 6.4 times as long as wide. Cephalothorax

subrectangular, tapering towards anterior with slight triangular

rostrum, 1.2 times as long as wide, about as long as pereonites

1 and 2 together, naked, eyes absent. Pereonite margins parallel,

pereonite 1 shortest, 0.4 times as long as cephalothorax;

pereonites 2 to 5 subequal, 0.8 times as long as cephalothorax,

pereonite 6 shorter, 0.6 times as long as pereonite 5 (all

pereonites respectively 2.4, 1.4, 1.4, 1.3, 1.4 and 1.7 times as

wide as long). Pleon with five free subequal pleonites bearing

pleopods; each pleonite 5.8 times as wide as long. Pleotelson

pentangular, one-third length of pleon and twice as wide as

long, with four small distal setae (Fig. 1101).

Antennule (Fig. 109A) of three articles, proximal article

5.2 times as long as wide, 1.8 times as long as distal two

articles together, with row of three strong inner-dorsal setae,

outer margin with tufts of one simple and two or three

penicillate setae at mid-length and distally; second article

nearly twice as long as wide, 0.4 times as long as third article,

with single inner distal penicillate and longer simple setae;

third article tapering, 0.4 times as long as first article, with five

simple and one penicillate distal setae.

Antenna (Fig. 109B) of six articles, proximal article

compact, naked; second article stout, as long as wide, with

dorsodistal seta longer than article; third article shorter than

wide, with fine dorsodistal seta; fourth article longest, ten times

as long as wide, curved, with one simple and one penicillate

distal setae; fifth article one-quarter as long as fourth with one

distal seta; sixth article minute with four distal setae.

Labrum (Fig. 109C) rounded, hood-shaped, distally setose.

Left mandible (Fig. 109D) with subtriangular pars incisiva and

wide, crenulate lacinia mobilis, right mandible (Fig. 109E)

without lacinia mobilis; pars molaris of both mandibles with

strong, rounded tooth-like protrusions around distal margin.

Labium (Fig. 109H) simple, finely setose on outer margins.

Maxillule (Fig. 109F) with eight distal spines, palp (Fig. 109F')

with two distal setae. Maxilla (Fig. 109G) ovoid, naked.

Maxilliped palp (Fig. 1091) first article naked, second article

with one outer and three inner setae, distal of these finely

denticulate in distal half; third article with four inner setae in

distal half of article, two of these finely denticulate in distal

half; fourth article with five inner to distal setae, four of these

finely denticulate in distal half, and one outer subdistal seta;

basis with single, long seta reaching distal margin of endites;

endites distally with two setae and two slight tubercles, outer

distal margin slightly denticulate. Epignath (Fig. 109J)

elongate, linguiform, naked.

Cheliped (Fig. 110A) with rounded basis reaching pereonite

1 ventrally, 1.2 times as long as wide, with single dorsodistal

seta; merus subtriangular with single ventral seta; carpus

elongate, three times as long as wide, with two midventral

setae of markedly unequal length, one fine ventrodistal seta,

and row of eight setae along dorsal margin; propodus slender,

curved, twice as long as wide, fixed finger 0.73 times as long

as palm, with two ventral setae, three setae on cutting edge;

dactylus with fine proximal seta.

Pereopod 1 (Fig. 110B) longer than others, coxal apophysis

large, triangular, pointed, with seta; basis arcuate, slender,

nearly six times as long as wide, with six simple setae along

dorsal margin; ischium compact, with ventral seta two-thirds

as long as merus; merus 0.4 times as long as basis, with three

simple distal setae; carpus just shorter than merus with distal

crown of eight simple setae; propodus 1.5 times as long as

carpus, with three dorsal subdistal setae, longer ventral

subdistal seta; short, stout dactylus with proximal seta longer

than dactylus, slender unguis 2.6 times as long as dactylus,

both together 0.9 times as long as propodus. Pereopod 2 (Fig.

110C), coxa similar to that of pereopod 1, basis 3.3 times as

long as wide, with midventral seta and eight setae along dorsal

margin; ischium with seta only half as long as merus; merus

0.25 times as long as basis, with single dorsal and two ventral

distal simple setae, and dense field of microtrichia across

ventral and ventrolateral surfaces in distal two-thirds; carpus

1.4 times as long as merus, with distal crown of eight setae and

dense field of microtrichia across ventral and ventrolateral

surfaces in distal two-thirds; propodus 1.6 times as long as

carpus, with two dorsal subdistal setae, longer ventral subdistal

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 108. Typhlotanais herthio sp. nov., female holotype. A, dorsal view; B, lateral view. Scale = 1.0 mm.

164

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 109. Typhlotanais herthio sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, left mandible; E, right mandible; F, maxillule

endite; F' maxillule palp; G, maxilla; H, labium; I, maxilliped; J, epignath. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

165

Fig. 110. Typhlotanais herthio sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5;

G, pereopod 6; H, pleopod; I, uropod-6. Scale = 0.1 mm.

166

M. Biazewicz-Paszkowycz & R.N. Bamber

seta, and fields of microtrichia; short, stout dactylus with

proximal seta longer than dactylus, slender unguis longer than

dactylus, both together 0.4 times as long as propodus. Pereopod

3 compact (Fig. 110D), similar to pereopod 2, basis with six

dorsal marginal setae; merus with three ventrodistal setae.

Pereopod 4 (Fig. 110E) basis stout, 2.1 times as long as

wide, with simple mid-dorsal seta and two penicillate setae

near ventrodistal corner; ischium with two ventrodistal setae;

merus 0.8 times as long as carpus, with field of microtrichia

across ventral and ventrolateral surfaces in distal two-thirds,

and two small ventrodistal spines; carpus with robust distal

molar spine, two simple mid-dorsal and one dorsodistal setae,

and “prickly tubercle” ( sensu Biazewicz-Paszkowycz, 2007)

surrounded by minute spines in ventrodistal half; propodus

1.5 times as long as carpus, with fields of microtrichia, mid¬

dorsal penicillate seta, strong dorsodistal seta, and two

ventrodistal dentiform spines; dactylus slender, with fields of

microtrichia, three times as long as curved unguis, both

together longer than propodus. Pereopod 5 (Fig. 110F) as

pereopod 4, but carpus with mid-dorsal and dorsodistal spines.

Pereopod 6 (Fig. 110G) as pereopod 5, but basis without

penicillate setae, propodus with three dorsodistal setae.

Pleopods (Fig. 110H) all alike, with naked basis, exopod

shorter than endopod; endopod and exopod without setae on

inner margin, outer margins with respectively 15 and 21 plumose

setae, proximal seta on both rami separated from others.

Uropod (Fig. 1101) biramous, basis naked; exopod and

endopod of one segment, exopod shorter than endopod, with

one fine proximal, one slender and one stouter distal setae;

endopod with four slender, one stouter and one penicillate

distal setae.

Male. Unknown.

Etymology. From the Anglo-Saxon haer - hairy, and thioh - the

thigh, alluding to the density of dorsal marginal setae on the

bases of the anterior pereopods, which distinguish this species

most evidently from the other species of the greenwichensis-

group of Typhlotanais sensu lato\ noun in apposition

Remarks. With the pronounced coxal spurs on the anterior

pereopods, the curving carpus-propodus and the dorsal

marginal spines on the carpus of the cheliped, and the prickly

tubercles on the posterior carpi, this species fits into the

“greenwichensis- group” of Biazewicz-Paszkowycz (2007). The

two described species of this group are T. greenwichensis

Shiino, 1970, from the Antarctic-Subantarctic, and

T. messinensis Sars, 1882 from the Mediterranean.

Typhlotanais greenwichensis differs from T. herthio sp.

nov. in being more elongate (nearly seven times as long as

wide), with a less-slender proximal article to the antennule

(four times as long as wide), and, most obviously, has only a few

dorsal marginal setae on the pereopods 1 to 3 (4, 4 and 3

respectively). T. messinensis is quite distinct in having a more

compact proximal peduncle article to the antennule (three

times as long as wide), and two-segmented rami on the uropods.

Typhlotanais herthio was collected sporadically through

the Bass Strait, from sandy substrata at depths between 81 and

115 m.

Genus Antiplotanais Bamber, 2008

Antiplotanais actuarius sp. nov.

Figures 111-113

Material examined. 1 9 (J58529), holotype. Western Bass Strait, 26

km SW of Cape Otway, Stn BSS 120, 39°01.0'S 143°22.1'E, 84 m

depth, medium sand, 31 January 1981; coll. M.F. Gomon; 1 $ (J56616),

paratype. Southern Port Phillip Bay, Stn PPBES 985, 38°21.0'S

144°51.5'E, 9 m depth, sand, 09 December 1971; coll. G.C.B. Poore &

S.F. Rainer.

Description of female. Body (Fig. 111A, B) compact, holotype

1.7 mm long, 4.5 times as long as wide. Cephalothorax

subrectangular, tapering towards anterior with slight triangular

rostrum, as long as wide, longer than pereonites 1 to 3 together,

naked, eyes absent. Pereonite 1 shortest, 0.2 times as long as

cephalothorax; pereonites 2 to 5 progressively longer, pereonite

2 being 1.4 times as long as pereonite 1, pereonite 5 being twice

as long as pereonite 1; pereonite 6 as long as pereonite 4 (all

pereonites respectively 4.5, 3.3, 2.8, 2.5, 2.1 and 2.4 times as

wide as long). Pleon with five free subequal pleonites bearing

pleopods; each pleonite 3.7 times as wide as long. Pleotelson

pentangular, one-third length of pleon and 1.6 times as wide as

long, with four small dorso-distal setae.

Antennule (Fig. 112A) of three articles, proximal article 1.9

times as long as wide, 2.4 times as long as distal two articles

together, with row of five inner setae, outer margin with

proximal and mid-length tufts penicillate setae, one simple seta

at mid-length and two distally; second article nearly twice as

wide as long, 0.6 times as long as third article, with single inner

distal penicillate and longer simple setae and outer distal

simple seta; third article tapering, 0.25 times as long as first

article, with six simple and one penicillate distal setae.

Antenna (Fig. 112B) of six articles, proximal article

compact, naked; second article stout, as long as wide, with

dorsodistal seta much shorter than article; third article shorter

than wide, with fine dorsodistal seta; fourth article longest,

nearly five times as long as wide, with one simple and three

penicillate distal setae; fifth article 0.4 times as long as fourth

with one distal seta; sixth article minute with five distal setae.

Labrum (not figured) rounded, hood-shaped, distally

setose. Left mandible (Fig. 112C) with subtriangular pars

incisiva and linguiform, crenulate lacinia mobilis, right

mandible (Fig. 112D) without lacinia mobilis but with wider,

rounded pars incisiva; pars molaris of both mandibles stout

with few (two or three) large, pointed, tooth-like protrusions on

distal margin. Labium (Fig. 112G) simple, finely setose on

outer and distal margins. Maxillule (Fig. 112E) with eight

distal spines, palp with two distal setae. Maxilla not recovered.

Maxilliped palp (Fig. 112F) first article naked, second article

with one outer and three inner setae, at least one of these finely

denticulate in distal half; third article with four inner setae in

distal half of article, two of these finely denticulate in distal

half; fourth article with five inner to distal setae, four of these

finely denticulate in distal half, and one outer subdistal seta;

basis with single, long seta exceeding distal margin of endites;

endites distally with two setae and two distinct tubercles, and

microtrichia. Epignath (Fig. 112H) elongate, linguiform, naked.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. Ill . Antiplotanais actuarius sp. nov., female holotype. A, dorsal view; B, lateral view. Scale = 1 mm.

168

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 112. Antiplotanais actuarius sp. nov., female paratype. A, antennule; B, antenna; C, left mandible; D, right mandible; E, maxillule;

F, maxilliped; G, labium; H, epignath. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

169

Fig. 113. Antiplotanais actuarius sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 6;

G, pleopod; H, uropod. Scale = 0.1 mm.

170

M. Biazewicz-Paszkowycz & R.N. Bamber

Cheliped (Fig. 113A) with rounded basis reaching pereonite

1 ventrally, 1.65 times as long as wide, with single dorsodistal

seta; merus subtriangular with single ventral seta; carpus

elongate, 2.5 times as long as wide, with three midventral

setae very unequal in length, and row of six minute setae along

dorsal margin; propodus slender, curved, 1.5 times as long as

wide, fixed finger 0.7 times as long as palm, with two ventral

setae, three setae on cutting edge; dactylus naked.

Pereopod 1 (Fig. 110B) longer than others, coxal apophysis

large (evident dorsally on whole animal, see Fig. 108A),

triangular, pointed, with seta; basis curved, 4.6 times as long

as wide, with three simple setae along dorsal margin; ischium

compact, with short ventral seta; merus 0.4 times as long as

basis, with one dorsal and one ventral simple distal setae;

carpus as long as merus with three dorsal and two ventral

simple distal setae; propodus 1.4 times as long as carpus, with

three distal setae; short, naked dactylus half as long as slender

unguis, both together 1.2 times as long as propodus. Pereopod

2 (Fig. 113C), coxa similar to that of pereopod 1, basis 3.7

times as long as wide, with two setae on dorsal margin;

ischium with single seta; merus 0.4 times as long as basis,

ventrodistally with two simple setae; carpus 1.1 times as long

as merus, with one dorsodistal seta, ventrodistally with two

simple setae and field of microtrichia; propodus 1.6 times as

long as carpus, with two dorsal subdistal setae, shorter ventral

subdistal seta, and fields of microtrichia; short, stout dactylus

with proximal seta longer than dactylus, slender unguis longer

than dactylus, both together 0.4 times as long as propodus.

Pereopod 3 compact (Fig. 113D), similar to pereopod 2, basis

with two simple setae on dorsal margin; ischium with single

seta; merus ventrodistally with two simple setae and field of

microtrichia; carpus ventrodistally with two simple setae and

field of microtrichia; propodus with two dorsal subdistal setae,

shorter ventral subdistal seta, and fields of microtrichia; short,

stout dactylus with proximal seta longer than dactylus, slender

unguis longer than dactylus.

Pereopod 4 (Fig. 113E) basis stout, 2.2 times as long as

wide, with two penicillate setae mid-ventrally; ischium with two

ventrodistal setae; merus 1.1 times as long as carpus, with field

of microtrichia across ventral and ventrolateral surfaces in

distal two-thirds, and two small ventrodistal spines; carpus with

robust distal molar spine, one dorsodistal blunt seta, and “prickly

tubercles” ( sensu Biazewicz-Paszkowycz, 2007) surrounded by

minute spines in ventrodistal half; propodus as long as carpus,

with fields of microtrichia, mid-dorsal penicillate seta, simple

dorsodistal seta, and two small ventrodistal spines; dactylus

slender, with fields of microtrichia, three times as long as curved

unguis, both together 0.9 times as long as propodus. Pereopod 5

(not figured) as pereopod 4. Pereopod 6 (Fig. 113F) as pereopod

4, but basis with simple setae, propodus with three dorsodistal

setae finely denticulate in their distal half.

Pleopods (Fig. 113G) all alike, with naked basis, exopod

shorter than endopod; endopod and exopod without setae on

inner margin, outer margins with respectively 9 and 16 plumose

setae, proximal seta on both rami separated from others.

Uropod (Fig. 113H) biramous, basis naked; exopod half as

long as endopod, with one fine proximal, one shorter and one

longer distal setae; endopod with residual (atrophied)

articulation line, with two penicillate setae just proximal to

this line, one subdistal and four distal simple setae and two

subdistal penicillate.

Male. Unknown.

Etymology. From the Latin actuarius - a shorthand writer, a

pun referring to the shorter chela (“hand”) in proportion to the

cheliped carpus in this species when compared with the other

two described species of Antiplotanais.

Remarks. The genus Antiplotanais shares with the Typhlotanais

greenwichensis- group (see above) the conspicuous coxal

apophyses on the anterior pereopods, and prickly tubercles on

the carpi of the posterior pereopods. It differs in the much more

compact antennule and antenna, the proportionately shorter

habitus (all less than 6 times as long as wide with all pereonites

at least twice as wide as long), the relatively long pleonites, the

presence of dorsal or dorsodistal setae on the pleotelson, the

stouter dactyli on the posterior pereopods and the presence of

distal lobes on the mandibular pars molaris.

There were two described species of Antiplotanais,

A. coochimudlo Bamber, 2008, from off Brisbane, Queensland,

and A. lutze (Bamber, 2005), from Esperance Bay, Western

Australia. Antiplotanais actuarius sp. nov. differs from the

other two particularly in the presence of dorsal setae on the

bases of the anterior pereopods, in the cephalothorax being no

longer than wide, and in the proportionately-shorter cheliped

propodus (including the fixed-finger), being 0.64 times as long

as the carpus compared with 0.85 times in A. coochimudlo and

the same length in A. lutze. All three species are from sandy

substrata in shallow waters off Australia.

Bamber (2008) remarked that the dactyli and ungues of the

posterior pereopods appeared to be fused; it is evident from

the present material that this is not the case. The posterior

ungues are distinct but very short, and confirmed by

re-examination of paratypic material of A. lutze, so evidently

missed in the examination of the previous material of this

genus of very small animals.

Antiplotanais actuarius was recorded only twice, from

Port Phillip Bay and the Western Bass Strait at 9 and 24 m

respectively, on sandy substrata.

Genus Hamatipeda Biazewicz-Paszkowycz, 2007

Hamatipeda sima sp. nov.

Figures 114-116

Material examined. 1 9 (J58901), holotype, 4 99 (J58902), paratypes.

Eastern Bass Strait, 85 km NE of North Point, Flinders Island, Stn

BSS 169S, 39°02.4'S 148°30.6'E, 120 m depth, sandy-mud, 15

November 1981; coll. R.S. Wilson.

Description of female. Body (Fig. 114A, B) elongate, slender,

holotype 3.9 mm long, 13 times as long as wide. Cephalothorax

subrectangular, tapering towards anterior with slight triangular

rostrum, 1.4 times as long as wide, naked, eyes absent. All

pereonites with parallel sides, all but the sixth longer than wide:

pereonite 1 as long as cephalothorax; pereonites 2 and 4

subequal, 1.5 times as long as cephalothorax; pereonite 3 longest,

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 114. Hamatipeda sima sp. nov. female holotype. A, dorsal view; B, lateral view. Scale = 1 mm.

172

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 115. Hamatipeda sima sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, left mandible; E, right mandible; F, maxillule;

G, maxilla; H, maxilliped; I, labium; J, epignath. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

173

Fig. 116. Hamatipeda sima sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5; G, pereopod 6;

H, pleopod; I, uropod. Scale = 0.1 mm.

174

M. Biazewicz-Paszkowycz & R.N. Bamber

1.75 times as long as cephalothorax; pereonite 5 shorter than

pereonite 4, 1.4 times as long as cephalothorax; pereonite 6

shortest, 0.6 times as long as cephalothorax (all pereonites

respectively 0.8, 0.5, 0.4, 0.5, 0.6 and 1.1 times as wide as long).

Pleon with five free subequal pleonites bearing pleopods; each

pleonite 4.8 times as wide as long. Pleotelson semicircular, twice

as long as each pleonite and 1.9 times as wide as long, with four

small distal setae.

Antennule (Fig. 115A) of three articles, proximal article

stout, 2.7 times as long as wide, five times as long as distal two

articles together, with one simple inner seta, outer margin with

tufts of two or three penicillate setae proximally, at mid-length

and distally, the last two tufts also with one simple seta; second

article nearly as long as wide, attached within invagination of

proximal article but as long as third article, with single outer and

inner simple setae; third article tapering, 1.4 times as long as

wide, with six simple and one penicillate distal setae.

Antenna (Fig. 115B) of six articles, proximal article compact,

naked; second article twice as long as wide and twice as long as

first article, with dorsodistal seta; third article as long as wide, as

long as first article, with dorsodistal seta and ventral microtrichia;

fourth article longest, five times as long as wide, with fields of

microtrichia, and distal crown of three simple and three

penicillate setae; fifth article one-quarter as long as fourth with

one distal seta; sixth article minute with four distal setae.

Labrum (Fig. 115C) rounded, hood-shaped, distally very-

finely setose. Left mandible (Fig. 115D) with subtriangular pars

incisiva and linguiform, bilobed lacinia mobilis, right mandible

(Fig. 115E) without lacinia mobilis but with longer cutting edge

on pars incisiva; pars molaris of both mandibles with strong,

smooth distal margin. Labium (Fig. 1151) simple, finely setose on

outer margins. Maxillule (Fig. 115F) with eight distal spines,

rows of microtrichia on outer margin of endite, palp with two

distal setae. Maxilla (Fig. 115G) ovoid, naked. Maxilliped palp

(Fig. 115H) first article naked, second article with one outer and

three inner setae; third article with one robust and three shorter

inner setae, all finely denticulate in distal two-thirds; fourth

article with five inner to distal setae, proximal of these finely

denticulate in distal half, and one outer subdistal seta; basis with

single, long seta not reaching distal margin of endites; endites

distally with two setae and non-articulate tubercle. Epignath

(Fig. 115J) elongate, linguiform, naked.

Cheliped (Fig. 116A) with rounded basis not reaching

pereonite 1 ventrally, twice as long as wide; merus subtriangular

with single ventral seta; carpus stout, 1.5 times as long as wide,

with two midventral setae, one fine dorsoproximal seta, and one

dorsodistal seta; propodus palm just longer than wide, fixed

finger 0.85 times as long as palm, with two ventral setae, three

setae on cutting edge, cutting edge finely denticulate; dactylus

with fine proximal seta.

Pereopod 1 (Fig. 116B) longer than others, coxal without

apophysis, with seta; basis straight, 4.3 times as long as wide,

with one simple and one penicillate dorsoproximal setae and one

ventrodistal seta; ischium compact, with ventral seta; merus one-

third as long as basis, with two ventrodistal and one dorsodistal

simple setae; carpus just longer than merus with two ventrodistal

and two dorsodistal simple setae; propodus 1.3 times as long as

carpus, with two dorsal subdistal setae and one ventral subdistal

seta; dactylus half as long as slender unguis, both together 0.9

times as long as propodus. Pereopod 2 (Fig. 116C), coxa similar

to pereopod 1, basis 4.1 times as long as wide without

dorsoproximal penicillate seta; merus 0.25 times as long as basis,

without dorsodistal seta; carpus with three dorsodistal and three

ventrodistal setae and small ventrodistal spine; propodus 1.5

times as long as carpus, with two dorsal subdistal setae and one

ventral subdistal seta; short, stout dactylus half as long as unguis,

both together 0.7 times as long as propodus. Pereopod 3 compact

(Fig. 116D), similar to pereopod 2, merus with dorsodistal seta.

Pereopod 4 (Fig. 116E) basis stout, 1.8 times as long as wide,

with two penicillate setae near ventrodistal corner; ischium with

two ventrodistal setae; merus 1.4 times as long as carpus, with

field of microtrichia across ventral and ventrolateral surfaces in

distal two-thirds, and two dentiform ventrodistal spines; carpus

with three dentiform hook-like ventrodistal spines and

dorsodistal seta; propodus 1.25 times as long as carpus, with

mid-dorsal penicillate seta, strong dorsodistal seta, and two

ventrodistal dentiform spines; dactylus slender, with fields of

microtrichia, four times as long as curved unguis, unguis distally

trifurcate, both together 0.65 times as long as propodus. Pereopod

5 (Fig. 116F) as pereopod 4. Pereopod 6 (Fig. 116G) as pereopod

4, but basis without penicillate setae, propodus with three

dorsodistal setae.

Pleopods (Fig. 116H) all alike, with naked basis, exopod

shorter than endopod; endopod and exopod without setae on

inner margin, outer margins with respectively 11 and 20 plumose

setae, proximal seta on both rami separated from others.

Uropod (Fig. 1161) biramous, basis naked; exopod and

endopod of one segment, exopod shorter than endopod, with one

fine proximal and two distal setae; endopod with one simple and

two penicillate setae just distal of mid-length, distally with four

simple and one penicillate setae.

Male. Unknown.

Etymology. From the Greek simos, meaning “snub-nosed”,

alluding to the characteristic very short distal antennular articles

of the present species.

Remarks. The distinctive genus Hamatipeda is characterized by

the very elongate body, with pereonites 1 to 5 longer than wide

and as long as (pereonite 1) or longer than the cephalothorax, and

the specialized dentiform hook-like spines on the carpus of the

posterior pereopods, inter alia. H. sima sp. nov. is entirely typical

of the genus.

Of the two previously described species, Hamatipedia

trapezoida Biazewicz-Paszkowycz, 2007, is distinct in having

trapezoidal pereonites which are narrower posteriorly than

anteriorly, while H. longa (Kudinova-Pasternak, 1975) is distinct

in having a two-segmented uropod endopod. Both have a more

slender cheliped carpus, more slender posterior pereopod bases,

lack distinctive tubercles on the maxilliped endites, and have a

much longer distal antennule article than the present species,

more than twice (H. trapezoida) or three times (H. longa ) as long

as wide, compared with less than 1.5 times as long as wide in

H. sima.

Hamatipeda sima is known only from the type-locality off

Tasmania (see above).

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

175

Genus Paratyphlotanais Kudinova-Pasternak & Pasternak,

1978

Paratyphlotanais colouros sp. nov.

Figures 117-119

Material examined. 1 9 (J58551), holotype, 3 99 (J58552), paratypes.

Eastern Bass Strait, 82 km ENE of North Point, Flinders Island, Stn

BSS36, 39°27,7'S 148°51.4'E, 293 m depth, coarse sand, 28 March

1979; coll. G.C.B. Poore; 1 9 (J58553), paratype. Eastern Bass Strait,

67 km ENE of North Point, Flinders Island, Stn BSS38, 39°22.4’S

148°38.7’E, 73 m depth, coarse sand, 29 March 1979; coll. G.C.B.

Poore; 2 9? (J58554), paratypes. Central Bass Strait, 44 km NE of

Cape Wickham, King Island, Stn BSS203,39°22.0’S 144°18.3'E, 60 m

depth, coarse sand, 23 November 1981; coll. R.S. Wilson.

Description of female. Body (Fig. 117A, B) relatively slender,

holotype 1.6 mm long, 6.4 times as long as wide. Cephalothorax

subrectangular, tapering towards anterior with slight triangular

rostrum, 1.5 times as long as wide, naked, eyes absent.

Pereonites wider than long, ventrally with anteriorly-pointed

hyposphenia on pereonites 1 to 3; pereonite 1 shortest, 0.12

times as long as cephalothorax; pereonite 2 parallel-sided, 2.4

times as long as pereonite 1; pereonite 3 with convex margins,

1.3 times as long as pereonite 2; pereonites 4 and 5 subequal,

twice as long as pereonite 2; pereonite 6 as long as pereonite 3

(all pereonites respectively 5.8, 2.5,1.9,1.3, 1.3 and 1.7 times as

wide as long). Pleon with five free subequal pleonites bearing

pleopods; each pleonite four times as wide as long. Pleotelson

pentangular, as long as last two pleonites together and 1.6 times

as wide as long, with four small distal setae.

Antennule (Fig. 118A) stout, proximal article 1.7 times as

long as wide, 1.2 times as long as distal two articles together,

with four simple setae along inner margin, outer margin with

one simple seta at mid-length and two penicillate and one

simple setae distally; second article shorter than wide, one-

quarter as long as first article, with one outer simple seta and

single inner penicillate and longer simple distal setae; third

article tapering, 2.5 times as long as second article, with five

simple and one penicillate subdistal setae adjacent to apical

spur sensu Bird (2004).

Antenna (Fig. 118B) of six articles, proximal article

compact, naked; second article as long as wide, with

dorsodistal seta; third article shorter than wide, 0.6 times as

long as second article, with dorsodistal seta; fourth article

longest, four times as long as third article and four times as

long as wide, curved, with three simple and one penicillate

distal setae; fifth article half as long as fourth article with one

distal seta; sixth article minute with four distal setae.

Labrum (Fig. 118C) rounded, hood-shaped, marginally

setose. Left mandible (Fig. 118D) with bilobed, crenulate pars

incisiva and wide, crenulate lacinia mobilis, right mandible

(Fig. 118E) similar but without lacinia mobilis; pars molaris of

both mandibles with strong, rounded marginal tubercles.

Labium (Fig. 118H) simple, finely setose on outer and inner-

distal margins. Maxillule (Fig. 118F) with eight distal spines,

palp not recovered. Maxilla not recovered. Maxilliped palp

(Fig. 118G) first article naked, second article with one outer

and three inner setae; third article with four inner setae in

distal half of article; fourth article with five inner to distal

setae and one outer subdistal seta; basis with single, long seta

reaching past distal margin of endites; endites distally with

one seta and slight tubercle, outer distal margin denticulate.

Epignath (Fig. 1181) elongate, linguiform, naked.

Cheliped (Fig. 119A) basis not reaching back to anterior of

pereonite 1 ventrally, 1.7 times as long as wide, with single

dorsodistal seta; merus subtriangular with three ventral setae;

carpus 2.2 times as long as wide, with two longer and one

shorter mid-ventral setae, one mid-dorsal and one dorsodistal

setae; propodus slender, palm 1.25 times as long as wide, fixed

finger as long as palm, with two ventral setae, three setae on

cutting edge; dactylus with fine proximal seta.

Pereopod 1 (Fig. 119B) longer than others, coxa without

apophysis, with seta; basis straight, slender, 4.5 times as long as

wide, with two dorsal and one ventral simple setae in proximal

third; ischium compact, with ventral seta; merus 0.3 times as

long as basis, with two ventral and one dorsal simple distal

setae; carpus 1.5 times as long as merus with distal crown of

six simple setae; propodus 1.4 times as long as carpus, with two

dorsal subdistal setae, shorter ventral subdistal seta; slender

dactylus with proximal seta exceeding tip of dactylus, slender

unguis 1.6 times as long as dactylus, both together as long as

propodus. Pereopod 2 (Fig. 119C) similar to pereopod 1, basis

4.8 times as long as wide, with three dorsal but no ventral setae;

ischium with seta; merus with single dorsal and ventral distal

simple setae, and ventrodistal spine; carpus 1.1 times as long as

merus, with dorsodistal and mesiodistal simple setae, and two

unequal ventrodistal spines, longer spine denticulate; propodus

with one dorsal subdistal seta; dactylus and unguis together 0.9

times as long as propodus. Pereopod 3 (Fig. 119D) similar to

pereopod 2, basis with two dorsal and one ventral setae; carpus

with dorsodistal spine half as long as carpus.

Pereopod 4 (Fig. 119E) basis stouter than that of anterior

pereopods, three times as long as wide; ischium with two

ventrodistal setae; merus as long as carpus, with one

denticulate slender ventrodistal spine and one seta; carpus

with one dorsodistal setae and four ventrodistal denticulate

spines; propodus 1.3 times as long as carpus, with one

dorsodistal and two ventral subdistal denticulate spines;

dactylus twice as long as curved unguis, both together 0.6

times as long as propodus. Pereopod 5 (not figured) as

pereopod 4, but basis with ventral penicillate seta. Pereopod 6

(Fig. 119F) basis stout, 2.4 times as long as wide; ischium with

two ventrodistal setae; merus 0.9 times as long as carpus, with

two denticulate slender ventrodistal spines; carpus with one

dorsodistal setae and apparently three ventrodistal slender

denticulate spines; propodus as long as carpus, with three

dorsodistal setae and two ventral subdistal slender spines;

dactylus slender, with ventral microtrichia, twice as long as

curved unguis, both together 0.7 times as long as propodus.

Pleopods (Fig. 119G) all alike, with naked basis, exopod

shorter than endopod; endopod with subdistal inner seta,

exopod without setae on inner margin, outer margins with

respectively 10 and 15 plumose setae, proximal seta on both

rami separated from others.

Uropod (Fig. 119H) biramous, basis naked; exopod of one

segment, less than half as long as proximal endopod segment,

176

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 117. Paratyphlotanais colouros sp. nov., female holotype. A, dorsal view; B, lateral view. Scale = 1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

177

Fig. 118. Paratyphlotanais colouros sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, left mandible; E, right mandible;

F, maxillule endite; G, maxilliped; H, labium; I, epignath. Scale = 0.1 mm.

178

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 119. Paratyphlotanais colouros sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 6;

G, pleopod; H, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

179

with one shorter and one longer distal setae; endopod of two

segments, distal segment 0.6 times as long as proximal

segment; proximal segment with one simple and two

penicillate distal setae, distal segment with one subdistal and

three distal simple setae.

Male. Unknown.

Etymology. From the Greek kolouros - “bobtailed”, alluding to

the characterizing short uropod exopod of this species.

Remarks. The present species, with its pereopod spination,

elongate and tapering cephalothorax, hyposphenia (sternal

spurs) on the anterior pereonites and apical spur on the

antennule, accords comfortably with the genus

Paratyphlotanais, as most recently competently analyzed by

Bird (2004), since when only one further species, P. alveolus

Blazewicz-Paszkowycz, 2007, has been described. P. colouros

sp. nov. has a unique conformation for the genus being without

coxal apophyses (spurs) and with a narrow pereonite 1 (less

than half the length of pereonite 2). Of the nine species

previously attributed to this genus (see Blazewicz-Paszkowycz,

2007) only P. gracilipes (Hansen 1913) from the Northeast

Atlantic south of Iceland, P. pectinatus Bird, 2004, from the

Northeast Atlantic Margin and P. armatus (Vanhoffen, 1914)

from the Antarctic share the short pereonite 1, but the first two

have coxal spurs, while the last has a cephalothorax shorter than

pereonites 1 to 3 together.

Furthermore, none of those species has such a short uropod

exopod in proportion to the endopod, being much longer than

half the length of the proximal segment of the endopod compared

with 0.4 times that length in P. colouros (note that the endopod

of P. pectinatus is only known from the proximal segment, but

the exopod is much longer than that segment and two-segmented:

see Bird, 2004, fig.7f).

P. colouros was collected from the Central and Eastern

Bass Strait at 60 to 293 m in coarse sand.

Genus Peraeospinosus Sieg, 1986

Peraeospinosus tanytrix sp. nov.

Figures 120-122

Material examined. 1 9 (J58534), holotype, and 107 99 , 36 neuters

(J58535), paratypes. Eastern Bass Strait, 63 km E of North Point,

Flinders Island, Stn BSS 167, 39°44.8'S 148°40.6'E, 124 m depth, fine

sand and mud, 14 November 1981; coll. R.S. Wilson; 5 99 and 1 neuter

(J58896), paratypes, Stn BSS 167G, 1 9 (J58898), paratype, Stn BSS

167S, same data as holotype; 1 9 (J58895), 7 99 in tubes with numerous

mancae (J58897), paratypes. Eastern Bass Strait, 60 km E of North

Point, Flinders Island, Stn BSS 32, 39°41.7'S 148°39.5'E, 115 m depth,

muddy sand, 27 March 1979; coll. G.C.B. Poore; 1 9 (J58899), paratype.

Eastern Bass Strait, 100 km NE of North Point, Flinders Island, Stn

BSS 170, 38°52.6'S 148°25.2'E, 130m depth, fine sand, 15 November

1981; coll. R.S. Wilson; 1 9 (J58900), paratype. Eastern Bass Strait, 28

km SSW of Mario, Stn BSS 207,37°59’S 148°27'E, 51 m depth, muddy

sand and fine shell, 30 July 1983; coll. M.F. Gomon; 4 99 in tubes with

mancae (J57817), paratypes, off Nowra, New South Wales, Stn SLOPE

1, 34°59.52'S 151°05.93'E, 204 m depth, 14 July 1986; coll. G.C.B.

Poore; 7 99 with tubes (J37858), paratypes, off Nowra, New South

Wales, Stn SLOPE 2, 34°57.9'S 151°08.0'E, 503 m depth, 14 July 1986;

coll. G.C.B. Poore; 2 99 (J37883), paratypes, off Nowra, New South

Wales, Stn SLOPE 7,34°52.28’S 151°15.02'Eto 34°51.13'S 151°15.13E,

1096 m depth, 15 July 1986; coll. G.C.B. Poore & C.-C. Lu.

Description of female. Body (Fig. 120A, B) slender, holotype

4.7 mm long, seven times as long as wide. Cephalothorax

rounded but tapering towards anterior with triangular rostrum,

as long as wide, naked, eyes absent. Pereonite 1 wider anteriorly,

just over half as long as cephalothorax; pereonites 2 and 3

narrowed at mid-length, subequal in length, 1.6 times as long as

pereonite 1; pereonites 4 and 5 subrectangular, subequal in

length, 1.1 times as long as pereonite 1; pereonite 6 with mid¬

lateral creases, shortest, 0.8 times as long as pereonite 1 (all

pereonites respectively 1.7, 0.9, 0.8, 1.2, 1.2 and 1.5 times as

wide as long). Pleon with five free subequal pleonites bearing

pleopods; each pleonite six times as wide as long. Pleotelson

semicircular, one-third length of pleon and twice as wide as

long, with two small distal setae (Fig. 122H).

Antennule (Fig. 121A) of three articles, proximal article

clavate, twice as wide proximally as distally, 3.3 times as long

as wide, 1.6 times as long as distal two articles together, with

row of three fine inner-dorsal setae, outer margin with tufts of

three, four and three penicillate setae proximally, at mid¬

length and distally, last two tufts with accompanying simple

seta; second article 1.3 times as long as wide, one-third as long

as third article, with two unequal dorsodistal simple setae;

third article tapering, almost half as long as first article, with

five simple distal setae.

Antenna (Fig. 121B) of six articles, proximal article

compact, as long as second article, with ventral microtrichia;

second article swollen, as long as wide, with ventral

microtrichia; third article shorter than wide, 0.7 times as long

as second article, with fine dorsodistal seta; fourth article

longest, 8.4 times as long as wide, four times as long as second

article, curved, with two simple and two penicillate distal

setae; fifth article half as long as fourth with one distal seta;

sixth article minute with five distal setae.

Labrum (Fig. 121C) rounded, hood-shaped, distally setose.

Left mandible (Fig. 121D) with subtriangular, crenulate pars

incisiva and wide, crenulate lacinia mobilis, right mandible

(Fig. 121E) with rounded, smooth cutting edge on pars incisiva,

without lacinia mobilis; pars molaris of both mandibles with

fine denticulations around distal margin. Labium (Fig. 121H)

simple, finely setose on distal margin and with rows of

microtrichia on outer margins. Maxillule (Fig. 121F) with nine

distal spines and sparse microtrichia, palp not recovered.

Maxilla (Fig. 121G) ovoid, naked. Maxilliped (Fig. 1211) palp

first article naked, remaining articles with microtrichia; second

article with one outer simple seta and three inner setae finely

denticulate in distal half; third article with four inner setae in

distal half of article, two of these finely denticulate in distal

half; fourth with five inner to distal setae finely denticulate in

distal half; basis with single seta about half as long as endites;

endites distally with two setae. Epignath not recovered.

Cheliped (Fig. 122A) basis not quite reaching anterior

margin of pereonite 1 ventrally, 1.4 times as long as wide, with

single dorsodistal seta; merus subtriangular with single ventral

seta; carpus 1.5 times as long as wide, with two midventral

setae, one fine dorsodistal seta, and row of five setae along

180

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 120. Peraeospinosus tanytrix sp. nov., female holotype. A, dorsal view; B, lateral view. Scale = 1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

181

Fig. 121 . Peraeospinosus tanytrix sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, left mandible; E, right mandible; F, maxillule

endite; G, maxilla; H, labium; I, maxilliped. Scale = 0.1 mm.

182

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 122. Peraeospinosus tanytrix sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 6;

G, pleopod; H, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

183

dorsal margin, ventral margin distally invaginated to

accommodate propodus on reflexion; propodus about as long

as wide, fixed finger as long as palm, with two ventral setae,

three setae on cutting edge; dactylus naked.

Pereopod 1 (Fig. 122B) coxa without apophysis, naked;

basis arcuate, slender, 5.7 times as long as wide, with seven fine

setae along dorsal margin and three more along ventral margin;

ischium compact, with fine ventral seta; merus elongate, 0.6

times as long as basis and 4.7 times as long as wide, with mid¬

dorsal fine seta and three simple distal setae; carpus 0.6 times as

long as merus, 2.5 times as long as wide, with distal crown of

five simple setae; propodus 1.5 times as long as carpus, six times

as long as wide, with two dorsal subdistal setae; short, stout

dactylus with proximal seta longer than dactylus, slender unguis

1.4 times as long as dactylus, both together 0.4 times as long as

propodus. Pereopod 2 (Fig. 122C), coxa similar to that of

pereopod 1, basis 4.9 times as long as wide, with one penicillate

and four fine simple setae along dorsal margin and three simple

setae along ventral margin; ischium with ventral seta; merus

0.25 times as long as basis, with single dorsodistal seta, short

ventrodistal spine, and dense field of microtrichia across ventral

and ventrolateral surfaces in distal two-thirds; carpus 0.9 times

as long as merus, with dorsal, mesial and ventral distal setae,

short ventrodistal spine, and dense field of microtrichia across

ventral and ventrolateral surfaces; propodus 2.6 times as long as

carpus, with ventral subdistal seta, one shorter dorsodistal seta,

one very long dorsodistal seta 1.3 times as long as propodus;

short, stout dactylus with proximal seta exceeding tip of

subequal slender unguis, both together one-third as long as

propodus. Pereopod 3 (Fig. 122D) similar to pereopod 2, basis

without penicillate seta; merus with additional ventrodistal seta;

dactylus and unguis together 0.2 times as long as propodus, long

distal seta 1.6 times as long as propodus.

Pereopod 4 (Fig. 122E) basis stout, 1.5 times as long as

wide, with two simple mid-dorsal seta and two dorsoproximal

and two ventrodistal penicillate setae; ischium with two

ventrodistal setae; merus 1.2 times as long as carpus, with field

of microtrichia across ventral and ventrolateral surfaces in

distal half, and two small ventrodistal spines; carpus with two

hooked distal spines, fine dorsodistal seta, and prickly tubercle

surrounded by minute spines and microtrichia in ventrodistal

half; propodus 1.3 times as long as carpus, with mid-dorsal

penicillate seta, strong dorsodistal seta, and two short

ventrodistal spines; dactylus slender, nearly three times as

long as denticulate unguis, both together half as long as

propodus. Pereopod 5 (not figured) as pereopod 4. Pereopod 6

(Fig. 122F) similar to pereopod 4, but basis without penicillate

setae, propodus with three dorsodistal setae.

Pleopods (Fig. 122G) all alike, with naked basis, exopod

shorter than endopod; endopod and exopod without setae on

inner margin, outer margins with respectively 19 and 31

plumose setae, proximal seta on both rami slightly separated

from others.

Uropod (Fig. 122H) biramous, basis naked; exopod and

endopod of one segment, subequal in length; exopod with one

fine proximal, one stout distal setae; endopod with four simple

and three penicillate distal setae.

Male. Unknown.

Etymology. From the Greek tany - long, and thrix - a hair, with

reference to the exceedingly long distal seta on the propodi of

pereopods 2 and 3.

Remarks. The genus Peraeospinosus was reviewed most

recently by Blazewicz-Paszkowycz (2005), who presented an

identification key to the ten species then described; in that key,

P. tanytrix sp. nov. identifies as P. emergensis Blazewicz-

Paszkowycz, 2005, sharing the rounded cephalothorax not

longer than wide and the elongate pereonites 1 to 3, but does

not share with that species the elongate merus, carpus and

propodus of pereopod 2, nor the elongate dorso-distal “rod-

seta” on the carpus of pereopod 1. The only other species

described since that key is P. acruxi Blazewicz-Paszkowycz,

2007, which shares with P. emergensis the pereopod 1 rod seta

and elongate anterior pereonites, but has a cephalothorax

longer than wide. Most distinctively, none of the eleven

previously described species have such an extraordinarily long

distal seta on the propodi of pereopods 2 and 3 as found in

P. tanytrix, although this condition is approached in some

species of Torquella Blazewicz-Paszkowycz, 2007.

Peraeospinosus tanytrix was collected from Eastern Bass

Strait at depths between 51 and 1096 m. The tubes, apparently

used for brooding, were of agglomerated sediment particles

and fibrous material.

Genus Meromonakantha Sieg, 1986

Meromonakantha anarsios sp. nov.

Figures 123-125

Material examined. 1 ? (J62058), holotype; 1 ? (J62059), paratype, Stn

BSS 36 (CR 79-K-l), Eastern Bass Strait, 82 km ENE of North Point,

Flinders Island, 293 m, 28 March 1979, 39°27.7'S 148°41.4'E, 293 m

depth, coarse sand, 28 March 1979, coll. G.C.B. Poore.

Description of female. Body (Fig. 123A, B) slender, holotype

1.4 mm long, 7.5 times as long as wide. Cephalothorax

subrectangular, narrowing anteriorly (conical) with slight

triangular rostrum, 1.2 times as long as wide, twice as long as

pereonite 1, naked, eyes absent. Pereonites all naked and

rectangular; pereonites 1 and 6 subequal in length; pereonites

2 and 3 subequal, 1.3 times as long as pereonite 1; pereonites 4

and 5 subequal, 1.5 times as long as pereonite 1 (all pereonites

respectively 1.6, 1.2, 1.1, 1.0, 1.0 and 1.3 times as wide as long).

Pleon of five free subequal pleonites bearing pleopods plus

pleotelson; each pleonite 4.2 times as wide as long. Pleotelson

subpentangular, as long as last two pleonites together, 1.75

times as wide as long.

Antennule (Fig. 124A) of three articles, proximal article

three times as long as wide, 1.7 times as long as distal two

articles together, outer margin with three pairs of penicillate

setae, one central and one distal simple setae each as long as

distal two articles together; second article longer than wide,

0.25 times as long as first article, with two distal simple setae;

third article tapering, 1.4 times as long as second article, with

five simple and one penicillate distal setae.

184

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 123. Meromonakantha anarsios sp. nov., female holotype. A, dorsal view; B, lateral view. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

185

Fig. 124. Meromonakantha anarsios sp. nov., female paratype. A, antennule; B, antenna; C, right mandible; D, maxillule endite; E, maxilla;

F, labium; G, maxilliped; H, epignath; I, cheliped. Scale = 0.1 mm.

186

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 125. Meromonakantha anarsios sp. nov., female paratype. A to F, pereopods 1 to 6 respectively; G, pleopod; H, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

187

Antenna (Fig. 124B) of six articles, proximal article

compact, fused to cephalothorax; second article just longer

than wide, with dorsodistal seta; third article as long as wide

and as long as second article, with strong dorsodistal seta;

fourth article longest, four times as long as wide, 3.6 times as

long as third article, with three simple and two penicillate

distal setae; fifth article 1.5 times as long as third, with one

distal simple seta; sixth article minute with four distal setae.

Labrum and left mandible not recovered. Right mandible

(Fig. 124C) with wide, spade-like pars incisiva bearing upper

and lower marginal rounded “teeth” and submarginal

denticulations; pars molaris stout, blunt, with rounded distal

tubercles. Labium (Fig. 124F) simple, outer distal corner with

unarticulated setulose projection. Maxillule (Fig. 124D) with

eight distal spines and small seta. Maxilla (Fig. 124E) simple,

linguiform, naked. Maxilliped (Fig. 124G) palp first article

naked, second article with one outer and three distal inner

setae, one of inner setae finely plumose; third article with

three longer and one shorter distal inner setae, fourth article

elongate, with four distal setae and one subdistal outer seta;

basis naked; endites distally naked, with outer-distal

microtrichia and paired inner setae. Epignath (Fig. 124H)

elongate, ribbon-like, distally pointed.

Cheliped (Fig. 1241) sclerite dorsally inserted, basis not

reaching anterior of pereonite 1 ventrally, 1.6 times as long as

wide with outer dorsodistal seta; merus subtriangular with

single ventral seta, and covering about half of ventral margin

of carpus; carpus 1.9 times as long as wide, with two midventral

setae, one dorsodistal seta; propodus elongate, 1.5 times as

long as wide, with two ventral setae, inner comb-row of three

setae; fixed finger slender, two-thirds as long as palm, with

three setae below cutting edge; dactylus with proximal seta.

Pereopod 1 (Fig. 125A) coxa without apophysis; basis

slender, 3.8 times as long as wide, with two dorsal penicillate

setae in proximal half; ischium compact, with one ventrodistal

seta; merus 0.7 times as long as carpus, ventrodistally with

two slender spines; carpus distally with slender spines dorsally,

mesially and ventrally; propodus slightly curved, 1.3 times as

long as carpus, with ventral subdistal seta; dactylus naked,

unguis 1.4 times as long as dactylus, both together 1.2 times as

long as propodus. Pereopod 2 (Fig. 125B), similar to pereopod

1, basis with mid-dorsal simple seta but no penicillate setae;

carpus with additional distal seta; propodus straight, 1.9 times

as long as carpus; dactylus and unguis together 0.9 times as

long as propodus. Pereopod 3 (Fig. 125C) similar to pereopod

2, basis naked.

Pereopod 4 (Fig. 125D) somewhat more compact, basis

three times as long as wide with two penicillate setae; ischium

with two ventrodistal setae; merus as long as carpus, with two

stout ventrodistal spines; carpus with four curved distal spines

and dorsodistal seta; propodus 1.2 times as long as carpus,

with two curved ventrodistal spines and one dorsodistal seta;

dactylus with fine ventral denticulation, about twice as long as

unguis, the two together as long as propodus. Pereopod 5 (Fig.

125E) as pereopod 4, but without penicillate seta on basis,

propodus with dorsodistal spine-like apophysis but no seta.

Pereopod 6 (Fig. 125F) as pereopod 5, but propodus with three

dorsodistal setae.

Pleopods (Fig. 125G) all alike, with naked basis, endopod

and exopod elongate, linguiform, without setae on inner or

outer margins, exopod wider and slightly longer than endopod,

each respectively with seven and three distal plumose setae.

Uropod (Fig. 125H) basis naked, twice as long as wide;

exopod of two subequal segments, just longer than proximal

endopod segment; endopod of two subequal segments, setose

as figured.

Male. Unknown.

Etymology. From the Greek anarsios - strange, incongruous,

as this species diverges on a number of characters from the

current diagnosis for the genus (see below).

Remarks. With the long seta on the third article of the antenna,

the simple setation/spination of the pereopods (lacking fields of

microtrichia or prickly-tubercles), the conformation of the

cephalothorax, of the mandibular molar process and of the

uropods, inter alia, the present species accords with

Meromonakantha rather than any other typhlotanaid genus. It

diverges from the diagnosis given by Bfazewicz-Paszkowycz

(2007) in that neither the cephalothorax nor the pleon are wider

than the pereon, and the dactyli and ungues of the posterior

pereopods are not “semi-fused”.

Although, as pointed out by Bfazewicz-Paszkowycz

(2007), the genus is in need of revision once sufficient material

of a number of its less-well described species becomes

available, Meromonakantha anarsios sp. nov. is also

distinguished from all of the other species in having relatively

long curved spines on the posterior pereopods, a more slender

proximal article to the antennule, and the pereonites mostly

parallel-sided.

While these differences may be considered sufficient to

distinguish Meromonakantha anarsios as a separate genus, we

choose at present to maintain its affiliation with other members

of Meromonakantha rather than erect a monotypic genus.

Family Tanaissuidae Bird & Larsen, 2009

Genus Tanaissus Norman & Scott, 1906

Tanaissus giraffa sp. nov.

Figures 126-129

Material examined. 1 9 (J58475), holotype; 1 9 (J23599), paratype, stn

MSL-EG 45, Eastern Bass Strait, 13.5 km E of eastern edge of Lake

Tyers, 37°51.74'S 148°14.77'E, 37 m depth, sand-shell, 25 September

1990, R.V. Sarda, Smith-Mclntyre Grab. 1 S (J28482), dissected, stn

MSL-EG 67, Eastern Bass Strait, 13.3 km E of eastern edge of Lake

Tyers, 37°51.42'S 148°14.36'E, 37 m depth, sand-shell, 4 June 1991,

coll. N. Coleman, Smith-Mclntyre Grab.

Description of female. Body (Fig. 126A, B) slender, ten times

as long as wide, 1.4 mm long. Cephalothorax as long as

pereonites 2 and 3 combined, twice as long as wide, rostral

half much narrower than posterior, with finely-rugose rounded

rostral margin (Fig. 127A). Pereonites all rectangular,

pereonitel shortest, 0.2 times as long as cephalothorax;

pereonite 2 twice as long as pereonite 1; pereonites 3 to 6

188

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 126. Tanaissus giraffa sp. nov. A, holotype female lateral view; B, holotype female dorsal view; C, male lateral. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

189

Fig. 127. Tanaissus giraffa sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, left mandible; D', mandible molar; E, right

mandible; F, maxillule; G, maxilla; H, labium; I, maxilliped; J, epignath. Scale = 0.1 mm.

190

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 128. Tanaissus giraffa sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5; G. pereopod 6;

H, pleopod; I, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

191

Fig. 129. Tanaissus giraffa sp. nov., male. A, antennule; B, antenna; C, cheliped; D, pereopod 1; D' pereopod 1 dactylus; E, pereopod 2;

E', pereopod 2 dactylus; F, pereopod 4; G, pereopod 6; H, pleopod. Scale = 0.1 mm.

192

M. Biazewicz-Paszkowycz & R.N. Bamber

subequal, 2.8 times as long as pereonite 1 (all pereonites

respectively 2.5, 1.2, 0.9, 0.9, 0.9 and 1.0 times as wide as

long). Pleon nearly twice as long as pereonite 6; pleonites

subequal in length, three times as wide as long, all bearing

pleopods; pleotelson subrectangular, as long as two preceding

pleonites, 1.4 times as wide as long, with two setae above each

uropod attachment and two fine distal setae.

Antennule (Fig. 127A) slender, as long as cephalothorax,

three-articled; article 1 four times as long as wide, 1.6 times

as long as articles 2 and 3 combined, with proximal and distal

outer clusters of four penicillate setae, and single inner and

outer simple distal setae; article 2 longer than wide with single

inner and outer simple distal setae; article 3 three times as

long as wide, 1.5 times as long as article 2, with six simple

distal setae and one penicillate seta.

Antenna (Fig. 127B) six-articled, article 1 short and

annular, article 2 twice as long as article 1, 1.5 times as long as

wide, with dorsal seta; article 3 as long as article 1, with dorsal

seta; article 4 longer than articles 1 to 3 combined, five times

as long as wide, curved, with six distal and subdistal penicillate

setae; article 5 as long as article 1, with long seta; article 6 very

small, with four setae.

Labrum (Fig. 127C) rounded, hood-shaped, naked.

Mandibles stout; left mandible (Fig. 127D) with triangular

incisor and broader denticulate lacinia mobilis, molar gently

curved, weak and acutely pointed; right mandible (Fig. 127E)

with finely denticulate distal margin and bifid incisor; molar as

on left mandible. Labium (Fig. 127H) two-lobed with slightly

notched distal processes. Maxillule (Fig. 127F) endite sigmoid,

with two distal setae and eight terminal spines; palp with two

distal setae. Maxilla (Fig. 127G) triangular, naked. Maxilliped

(Fig. 1271) basis with short seta near articulation with palp;

palp article 1 naked, article 2 with three inner weak setae,

article3 with three unequal inner setae, and article 4 with two

longer and three shorter simple setae; endites almost-

completely fused. Epignath (Fig. 127J) with distinct basal

lobe, distally slender and finely setulose.

Cheliped (Fig. 128A) attachment posterior on

cephalothorax, basis extending past anterior margin of

pereonite 1 ventrally, with rounded posterior free margin, 1.4

times as long as wide, with small laterodistal seta; merus

subtriangular, with ventral seta; carpus 1.3 times as long as

broad, with one dorsal and two ventral setae; chela 1.4 times as

long as carpus, propodus without bifid dorsodistal crest,

anterior comb-row of five dendritic setae; fixed finger robust,

with convex cutting edge, two ventral setae and three setae

near cutting edge; dactylus narrow, curved and acute, with

several dorsal nodules and one small anterior seta.

Pereopod 1 (Fig. 128B) longer and more slender than

pereopods 2-3; coxa with seta; basis 7.3 times as long as wide;

ischium with small seta; merus three times as long as wide, half

as long as basis, with ventrodistal seta; carpus as long as merus,

with two distal setae, both much shorter than propodus;

propodus 0.8 times as long as carpus, with small dorsodistal

spine-like apophysis; dactylus half as long as unguis , both

together 0.7 times as long as propodus. Pereopod 2 (Fig. 128C)

stouter than pereopod 1, coxa with seta; basis narrow proximally,

3.5 times as long as wide, with one mid-dorsal and two ventral

penicillate setae; ischium with one small seta; merus distally

expanded, 0.4 times as long as basis, with one seta and one spine

ventrodistally; carpus 0.7 times as long as merus, rectangular,

with one dorsodistal and one ventrodistal spines; propodus 1.2

times as long as carpus, with two ventrodistal spines; dactylus

and unguis together 0.8 times as long as propodus. Pereopod 3

(Fig. 128D) similar to pereopod 2.

Pereopod 4 (Fig. 128E) basis 3.8 times as long as wide

with two mid-ventral penicillate setae; ischium with two setae;

merus 0.8 times as long as carpus, with two ventrodistal

spines; carpus nearly three times as long as wide, with outer

and inner dorsal spines longer than outer and inner ventral

spines, simple dorsodistal seta; propodus 1.25 times as long as

carpus, 2.4 times as long as wide, with one dorsodistal seta

and two ventrodistal spines; dactylus and very short bifurcate

unguis apparently distinct, together 0.9 times as long as

propodus. Pereopod 5 (Fig. 128F) similar to pereopod 4,

dactylus with microtrichia, together with unguis as long as

propodus. Pereopod 6 (Fig. 128G) similar to pereopods 4 and

5 but basis without penicillate setae, propodus with three

dorsodistal setae, dactylus plus unguis shorter than propodus.

Pleopods all similar (Fig. 128H), with basal article about

as long as wide, naked; endopod and exopod rami similar, but

endopod somewhat wider, with eight terminal setae, exopod

with nine terminal setae; all setae plumose and longest barely

as long as rami.

Uropod (Fig. 1281) slender, twice as long as pleotelson;

basal article twice as long as wide, naked; exopod two-

segmented, just longer than proximal article of endopod, with

one distal seta on longer proximal segment, two unequal distal

setae on distal segment; endopod two-segmented, proximal

segment shorter and with one simple and two penicillate distal

setae, distal article with one subdistal and three distal simple

setae and one distal penicillate seta.

Distinctions of male. Body (Fig. 126C) slightly larger than

female, length 1.5 mm. Cephalothorax similar to that of

female. Pereonites more or less rectangular, or with slightly

concave lateral margins; pereonite 1 not shortest, 0.7 times as

long as each of cephalothorax and pereonite 2; pereonites 2

and 3 subequal, pereonites 4 to 6 progressively shorter,

pereonite 6 half as long as pereonite 1. Pleon as long as

pereonites 4 to 6 inclusive.

Antennule (Fig. 129A) seven-articled, shorter than

cephalothorax; peduncle article 1 stout, 3.3 times as long as

wide, with array of nine dorsal penicillate setae in proximal

half, mid-inner simple seta, one longer and one shorter outer

distal simple setae with adjacent penicillate seta, and one

dorsodistal seta; peduncle article 2 compact, 0.2 times as long

as article 1, with two inner setae; flagellum of five segments,

proximal four segments with group of five or six outer distal

aesthetascs; distal segment with four simple distal setae.

Mouthparts absent apart from large maxillipeds.

Maxilliped (Fig. 129B) basis with seta near articulation with

palp; palp articles 1 and 2 similar in length, article 2 with two

small and one long setae; article 3 longer than articles 1 and 2

combined, with three inner setae; article 4 small, with two

short, tow medium and two very long terminal setae.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

193

Cheliped (Fig. 129C) similar to that of female, but carpus

more slender, 1.7 times as long as wide, chela more slender and

longer than carpus, fixed finger narrower; comb-row of 20 setae.

Pereopods (Fig. 129D to G) generally similar to those of

female; pereopod 2 (Fig. 129E) propodus with microtrichia;

pereopods 4 to 6 (Fig. 129F, G) with more slender propodi (about

5 times as long as wide), dactyli slender and severely curved.

Pleopods all similar (Fig. 129H), proportionately larger

than in female and rami more elongate; endopod with one

distal and eleven outer plumose setae, exopod with fourteen

plumose setae, longest setae more than twice as long as rami.

Uropod similar to that of female.

Etymology. With reference to the unusually long pereonite 1 of

the male, Giraffa (from the Arabic - zirafah) is the genus of

the African camelopard ruminant with an extraordinarily long

neck (noun in apposition).

Remarks. There are four described species of Tanaissus (see

Bird, 2002: Bamber et al., 2009), all from the Northern

Hemisphere, three from the North Atlantic and one from the

Mediterranean. Bamber et al. (2009) give a key to three of

these, in which the female of T. giraffa sp. nov. fails at couplet

2, owing to its having a single-pointed, acuminate molar

process on the mandible but no bifid dorsal crest on the chela;

the male of the present species fails at couplet 4 owing to its

having a pleon longer than pereonites 4 and 5 together, but no

ventral apophysis on the cheliped carpus; in addition, its

antennular segmentation is distinct from all of these species,

as is the very long pereonite 1. The fourth species,

T. psammophilous (Wallace, 1919, q.v.), known only from the

female, is incompletely described, but differs from T. giraffa

in being less elongate (about 7 times as long as wide), in the

proportions of the pereonites and of the antennule, and in the

structure of the chela.

Tanaissus giraffa was recorded from the Eastern Bass

Strait on shelly sand at 37 m depth.

Genus Protanaissus Sieg, 1983

Diagnosis (after Shiino, 1970 and Sieg, 1983b). Antennule of

three articles; antenna of six articles. Labrum naked. Mandible

molar process tapering to a point, without grinding surface;

incisor process of right mandible triangular, notched, and

serrated on anterior border; incisor process of left mandible

triangular, serrated on anterior border, lacinia mobilis of

similar shape. Labium simple. Maxillule apically curved

inwards. Maxilliped bases fused; endites flared, short and

broad, fused proximally, not fused distally; palp article 1

naked; article 2 with three distal/inner setae, longest of which

exceeds tip of palp. Chela of cheliped rugose; fixed finger

cutting-edge serrated. Pereopod 1 dactylus plus unguis as long

as or longer than propodus, each of these longer than merus and

carpus combined. Pereopods 2 and 3 carpus with slender

ventrodistal spines; pereopods 4 to 6 with three spines and one

seta on carpus, and short, stout dactylus armed with small

unguis, propodi with dorsodistal seta/setae exceeding length of

dactylus. Pleopods with subequal rami, endopod with subdistal

inner plumose seta. Uropod biramous, both rami with two

segments, exopod shorter than endopod, endopod proximal

segment with conspicuous distal penicillate setae.

Type species. Typhlotanais longidactylus Shiino, 1970 by

monotypy.

Remarks. Sieg (1983b) erected the genus Protanaissus for

Typhlotanais longidactylus Shiino, 1970, a species from the

Antarctic, recognizing both that this species was not a

typhlotanaid, and that it had affinities with the genus Tanaissus.

As the diagnosis given by Sieg (ibid.) was very brief, this has

been expanded above, based also on the original description of

the type species by Shiino (1970). Particular characterizing

features, especially as cited by Shiino (ibid.) are the elongate

propodus and dactylus-plus-unguis of pereopod 1, and the wide

maxilliped endites, while the longer inner seta on maxilliped

palp article 2, the spination of the merus of pereopods 2 and 3,

and the long distal propodal seta of pereopods 4 to 6 are further

features distinguishing Protanaissus from Tanaissus.

Subsequent species attributed to this genus are discussed

below, after the description of a new species of Protanaissus

from the Bass Strait.

Protanaissus huberti sp. nov.

Figures 130-132

Material examined. 1 9 (J50812), holotype, Stn VC 41 C3, Eastern Bass

Strait, 37°32.95'S 148°03.78'E, 40 m depth, 08 May 1998, coll. N.

Coleman, Smith-Mclntyre Grab. 1 9 (J51796), paratype dissected, Stn

VC 40 Cl, Eastern Bass Strait, 37°35.42’S 147°31.88’E, 40 m depth, 08

May 1998, coll. N. Coleman, Smith-Mclntyre Grab. 1 $ (J48971),

paratype, CPBS 32N/810, Western Port off Crib Point, 38°20.83'S

145°13.49'E, 13 m depth, sandy gravel, 12 August 1970, coll. A.J.

Gilmour. 1 9 (J55829), paratype dissected, Stn VC 18 C2, Central Bass

Strait, 38°30.2'S 144°15.0'E, 40 m depth, 13 May 1998, coll. N.

Coleman, Smith-Mclntyre Grab. 19 (J48980), paratype, CPBS 32S/770,

Western Port off Crib Point, 38°21.6'S 145°13.67'E, 13 m depth, muddy

sand, 06 July 1970; coll. A.J. Gilmour. 1 9 (J48919), paratype, CPBS

300/770, Western Port off Crib Point, 38°21.15'S 145 0 13.51'E, 15 m

depth, fine sand with mud, 06 July 1970, coll. A.J. Gilmour.

Description of female. Body (Fig. 130) slender, parallel-sided,

seven times as long as wide, holotype 1.3 mm long.

Cephalothorax longer than pereonites 2 and 3 combined, 1.3

times as long as wide, rostral half narrower than posterior.

Pereonites all rectangular with convex lateral margins,

pereonitel shortest, 0.25 times as long as cephalothorax;

pereonites 2,3 and 6 subequal, 1.7 times as long as pereonite 1;

pereonites 4 and 5 subequal, 2.4 times as long as pereonite 1

(all pereonites respectively 2.5, 1.5, 1.5, 1.1,1.1 and 1.6 times as

wide as long). Pleon 2.5 times as long as pereonite 6; pleonites

subequal in length, four times as wide as long, all bearing

pleopods; pleotelson semicircular, as long as two preceding

pleonites, 1.8 times as wide as long.

Antennule (Fig. 131 A) slender, as long as cephalothorax,

three-articled; article 1 four times as long as wide, 1.5 times

as long as articles 2 and 3 combined, with mid-dorsal and

dorsodistal tufts of penicillate setae, and single mid-length

and distal simple inner setae; article 2 twice as long as wide,

0.4 times a slong as article 1, with single inner and outer simple

194

M. Biazewicz-Paszkowycz & R.N. Bamber

distal setae; article 3 twice as long as wide, 0.7 times as long

as article 2, with five simple distal setae and one aesthetasc.

Antenna (Fig. 131B) six-articled, article 1 short and

annular, article 2 more than twice as long as article 1,1.5 times

as long as wide, with dorsal seta; article 3 half as long as

article 2, with dorsal seta; article 4 longer than articles 1 to 3

combined, 5.6 times as long as wide, curved, with two simple

and three penicillate distal setae; article 5 as long as article 1,

1.5 times as long as wide, with long distal seta; article 6 very

small, with two setae.

Labrum (Fig. 131C) rounded, hood-shaped, naked. Left

mandible (Fig. 131D) with triangular incisor serrated on

anterior border, and broader denticulate lacinia mobilis, molar

gently curved, tapering to narrow tip with spinules but no

grinding surface; right mandible (Fig. 131E) with finely

denticulate distal margin and bifid incisor; molar as on left

mandible. Labium not recovered. Maxillule (Fig. 131F) endite

sigmoid, with two distal setules and six longer and two shorter

terminal spines; palp not recovered. Maxilla (Fig. 131G) wide,

rounded, naked. Maxilliped (Fig. 131G) basis fused, with long

seta near articulation with palp almost reaching distal margin

of palp article 3; palp setae simple, article 1 naked; article 2

with outer distal seta, three inner-distal setae, longest of which

exceeds tip of palp; article 3 with four unequal inner distal

setae; article 4 with two subdistal and four distal setae; endites

basally fused, distally wide, inner distal margin with one short

and one linguiform tubercles. Epignath not recovered.

Cheliped (Fig. 131H) basis with rounded posterior free

margin, 1.6 times as long as wide, with small laterodistal seta;

merus subtriangular, with ventral seta; carpus 1.8 times as

long as broad, with one dorsoproximal, one dorsodistal and

two mid-ventral setae; chela about as long as carpus, stout,

resembling that of Tanaissus spp., propodus wider than long

with dorsodistal rugosity; fixed finger robust, proximally wide,

with convex denticulate cutting edge, one ventral seta and

three setae near cutting edge; dactylus with dorsal rugosity in

proximal half.

Pereopod 1 (Fig. 132A) longer and more slender than

pereopods 2-3; coxa without seta; basis five times as long as

wide, sinuous; ischium with small seta; merus 0.3 times as

long as basis, naked; carpus 1.7 times as long as merus, naked;

propodus 1.2 times as long as merus and carpus combined,

with small dorsodistal spine-like apophysis and fine ventral

subdistal seta; curved dactylus 0.8 times as long as curved

unguis, both together as long as propodus. Pereopod 2 (Fig.

132B) stouter than pereopod 1, coxa with fine seta; basis 3.6

times as long as wide, with one dorsoproximal simple seta;

ischium with one seta; merus distally wider, one-quarter as

long as basis, with one slender ventrodistal spine; carpus 1.8

times as long as merus, with one slender ventrodistal spine;

propodus as long as carpus, with one ventrodistal seta;

dactylus shorter than unguis, both together as long as

propodus. Pereopod 3 (Fig. 132C) similar to pereopod 2, but

coxal seta longer, basis naked, ventrodistal spine on carpus

larger (more than half as long as carpus), propodus with some

distal microtrichia.

Pereopod 4 (Fig. 132D) basis 2.9 times as long as wide;

ischium with two setae; merus 0.4 times as long as basis, with

Fig. 130. Protanaissus huberti sp. nov. female holotype. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

195

Fig. 131 . Protanaissus huberti sp. nov. A, antennule; B, antenna; C, labrum; D, left mandible ; E, right mandible; F, maxillule ; G, maxilliped and

maxilla ; H, cheliped. Scale = 0.01 mm.

196

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 132. Protanaissus huberti sp. nov. A, pereopod 1; B, pereopod 2; C, pereopod 3; D, pereopod 4; E, pereopod 5; F, pereopod 6; G, pleopod;

H, uropod. Scale = 0.01 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

197

two ventrodistal spines; carpus as long as merus, with single

ventral, outer and inner distal spines and dorsodistal spinule;

propodus 1.1 times as long as carpus, with dorsal microtrichia

in distal half, one dorsodistal seta exceeding length of dactylus

plus unguis, and ventrodistal spine almost as long as dactylus;

dactylus and very short bifurcate unguis together half as long

as propodus. Pereopod 5 (Fig. 132E) similar to pereopod 4,

merus somewhat shorter. Pereopod 6 (Fig. 132F) similar to

pereopods 4 and 5 but propodus with two shorter and one

longer dorsodistal setae, no ventrodistal seta or spine.

Pleopods all similar (Fig. 132G), basal article about as

long as wide, naked; endopod and exopod rami similar, but

endopod somewhat shorter, with five outer-distal and one

inner-subdistal plumose setae; exopod with seven outer-distal

and one separated outer proximal plumose setae.

Uropod (Fig. 132H) slender; basal article naked; exopod

two-segmented, just longer than proximal article of endopod,

segments subequal in length, with one distal seta on proximal

segment, two unequal distal setae on distal segment; endopod

two-segmented, segments subequal in length, proximal

segment with two inner penicillate distal setae, distal article

with one subdistal and three distal simple setae.

Male. Unknown.

Etymology. This species is dedicated to the first author’s son

and the second author’s good friend, Hubert.

Remarks. Protanaissus huberti sp. nov. shows many similarities

to P. longidactylus, including the elongate distal articles of

pereopod 1, the rugose cheliped, the conformation of the

mouthparts, particularly the wide maxilliped endites, and the

long dorsodistal setae on the propodi of the posterior pereopods.

It is distinguished from P. longidactylus in having the distal

antennular article shorter than the second article (approaching

twice as long in P. longidactylus ), the more compact fifth

article of the antenna, the presence of distal spinules on the

mandibular molar process, the presence of linguiform distal

tubercles on the maxilliped endites, in having only one

ventrodistal spines on the merus and carpus of pereopods 2 and

3, only one ventral seta on the cheliped propodus (two in

P. longidactylus ), serrations along the whole cutting edge of the

fixed finger of the cheliped propodus (only distally serrate in

P. longidactylus ) and in having only one ventrodistal seta on

the propodus of the posterior three pairs of pereopods (or none

on pereopod 6) compared with two in P. longidactylus.

Protanaissus huberti was taken occasionally throughout the

eastern and central Bass Strait at depths between 13 and 40 m.

The second species to have been attributed to Protanaissus

was P. makrotrichos Sieg 1986, from the shelf off Argentina,

which showed many similarities to P. longidactylus, but had a

distal grinding (“triturating”) surface on the mandibular molar

process, no rugosity on the cheliped, and a maxilliped endite

distally folded to fuse with the basis giving the appearance of

two narrow lobes; rather than having a long dorsodistal seta on

the propodus of pereopod 6, P. makrotrichos has a long

ventrodistal seta. Gufu (1996c) described P. alvesi from Brazil,

which, while again similar to P. longidactylus, also had a distal

grinding surface on a stout (not tapering) mandibular molar

process, no rugosity on the cheliped, no very long seta on

maxilliped palp article 2, a maxilliped endite apparently

distally narrow, and a long ventrodistal seta on the propodus of

pereopod 6, thus more similar to P. makrotrichos. Finally,

Larsen and Heard (2004a) described P. floridensis from

Florida, a species with a uniformly narrow mandibular molar

process, rugosity on the cheliped, and distally narrow (but not

infolded or fused) maxilliped endites, as well as a one-segmented

uropod exopod, a uropod endopod without conspicuous pair of

penicillate setae on the proximal segment, no very long seta on

maxilliped palp article 2, the dactylus plus unguis of pereopods

4 to 6 fused into a claw, no long distal seta on the propodus of

pereopod 6, and a quite distinct pereopod 1, with the propodus

shorter than the merus and carpus combined, and a dactylus-

plus-claw about half as long as the propodus.

That these three additional species have a distally-narrowed

maxilliped endite, that two of them have a grinding mandibular

molar and no cheliped rugosity, while the third has a quite

distinct pereopod 1 and apparently fused claws on pereopods 4

to 6, inter alia, puts them in conflict with the generic diagnosis

given above. Conversely, the new Australian species,

Protanaissus huberti, described above agrees with the generic

diagnosis in all respects.

We therefore remove Protanaissus makrotrichos and

P. alvesi to a separate genus, clearly close to Protanaissus,

while P. floridensis is moved to yet another distinct genus, no

closer to Protanaissus than it is to Tanaissus. These genera are

defined below.

Genus Molotanaissus gen. nov.

Diagnosis of female. Antennule of three articles; antenna of six

articles. Labrum naked. Mandible molar process with distal

grinding or crushing surface; incisor process of right mandible

triangular, notched, and serrated on anterior border; incisor

process of left mandible triangular, serrated on anterior border,

lacinia mobilis of similar shape. Maxillule apically curved

inwards. Maxilliped bases fused, endites short and distally

narrowed and infolded, fused proximally; palp article 1 naked;

article 2 with three distal/inner setae, longest of which does not

exceed tip of palp. Chela of cheliped not rugose; fixed finger

cutting-edge not serrated. Pereopod 1 dactylus plus unguis as

long as or longer than propodus, each of these longer than

merus and carpus combined. Pereopods 2 and 3 carpus with or

without seta and with relatively stout ventrodistal spine;

pereopods 4 to 6 with four distal spines on carpus, and with

short, stout dactylus armed with small unguis, propodi with

long ventrodistal seta exceeding length of dactylus. Pleopods

with subequal rami, endopod with subdistal inner plumose

seta. Uropod biramous, both rami with two segments, exopod

shorter than endopod, endopod proximal segment with

conspicuous distal penicillate setae.

Male. Unknown.

Etymology. From the Latin molo - “grind”, pertaining to the

grinding surface on the molar processes of the mandible (unlike

the condition found in Tanaissus or Protanaissus ), and

Tanaissus ; masculine.

M. Blazewicz-Paszkowycz & R.N. Bamber

Type species. Protanaissus makrotrichos Sieg, 1986 by original

designation.

Species included : Molotanaissus makrotrichos (Sieg, 1986)

comb, nov.; M. alvesi (Gufu, 1996) comb. nov. (see Gufu, 1996c,

for distinctions between these two species).

Distribution. Patagonian Shelf off Argentina (20-50 m depth)

and off Brazil (58=60 m).

Genus Unitanaissus gen. nov.

Diagnosis of female. Antennule of three articles; antenna of six

articles. Labrum naked. Mandible molar process uniformly

narrow, with distal bifurcation but no grinding surface; incisor

process of left mandible triangular, serrated on anterior border,

slightly notched, lacinia mobilis narrower with three rounded

distal crenulations. Maxillule apically curved inwards.

Maxilliped bases fused, endites short, parallel-sided and

entirely narrowed; palp article 1 naked; article 2 with three

distal/inner setae, longest of which does not exceed tip of

article 3. Sclerite triangular, inserted dorsally to cheliped basis.

Chela of cheliped rugose; fixed finger cutting-edge slightly

serrated distally. Pereopod 1 dactylus plus unguis half as long

as propodus, each of these shorter than merus and carpus

combined. Pereopods 2 and 3 carpus with relatively stout

ventrodistal spine; pereopods 4 to 6 with dactylus and unguis

fused into short, stout claw, propodi without long distal seta.

Pleopods with subequal rami, endopod with subdistal inner

plumose seta. Uropod biramous, exopod with one segment and

shorter than endopod, endopod with two segments, proximal

segment without conspicuous distal penicillate setae.

Male. Unknown.

Etymology. From the Latin unus - “one”, and Tanaissus, alluding

to the single-segmented uropod exopod, a condition unlike that

found in Tanaissus , Protanaissus or Molotanaissus ; masculine.

Type species. Protanaissus floridensis Larsen & Heard, 2004

by monotypy.

Species included. Unitanaissus floridensis (Larsen & Heard,

2004) comb. nov.

Distribution. Atlantic coast of Florida, 7 m depth.

Family Agathotanaidae Lang, 1971

Genus Paragathotanais Lang, 1971

Paragathotanais wurundjeri sp. nov.

Figures 133-134

Material examined. 1 ? (J58566), holotype. Eastern Bass Strait, 60 km E

of North Point, Flinders Island, Stn BSS 32,39°41.7'S 148°39.5'E, 115 m

depth, muddy sand, 27 March 1979, coll. G.C.B. Poore; 1 ? (J58567),

paratype. Eastern Bass Strait, 24 km NNE of Eddystone Point, Stn BSS

163,40°43.9’S 148°32.5’E, 56 m depth, muddy sand, 14 November 1981;

coll. R.S. Wilson; 1 ? (J58568), paratype. Eastern Bass Strait, 85 km NE

of North Point, Flinders Island, Stn BSS 169, 39°02.4’S 148°30.6'E,

120 m depth, muddy sand, 15 November 1981; coll. R.S. Wilson.

Description of female. Body (Fig. 133A) slender, holotype

3.7 mm long, 8.5 times as long as wide. Cephalothorax pear-

shaped, widest and laterally-rounded posteriorly, tapering

towards anterior with slight rounded rostrum, 1.5 times as long

as wide, as long as pereonites 1 and 2 together, naked; eyelobes

and eyes absent. Pereonites hexagonal, pereonites 1 to 3 widest

anteriorly, pereonites 4 to 6 centrally; pereonites 1 and 6

subequal in length, shortest, half as long as cephalothorax;

pereonite 2 just longer than pereonite 1; pereonites 3 to 5

subequal in length, longer than wide and about 1.25 times as

long as pereonite 1 (all pereonites respectively 1.4, 1.1, 0.9, 0.8,

0.9 and 1.0 times as wide as long). Pleon narrower than pereon,

pleonites without pleopods, each five times as wide as long.

Pleotelson pentangular (Fig. 134H), one-half length of pleon

and 1.4 times as wide as long.

Antennule (Fig. 133B) of four articles, proximal article

2.9 times as long as wide, as long as distal three articles

together, outer margin with four penicillate setae in distal

half, and simple seta distally as long as second peduncle

article; second article twice as long as wide, 0.4 times as long

as first article, with two outer distal penicillate and single

longer simple setae, one inner distal seta; third article 0.3

times as long as second article, with single inner and outer

distal setae; fourth article as long as second, with five simple

and one penicillate distal setae.

Antenna (Fig. 133C) of six articles, proximal article

compact, naked; second article nearly twice as long as wide,

naked; third article just shorter than wide, half as long as

second article, with fine dorsodistal seta; fourth article

longest, twice as long as second article and four times as long

as wide, with penicillate seta in proximal half and one simple

and two penicillate distal setae; fifth article half as long as

fourth with one distal seta; sixth article minute with five

distal setae.

Labrum not recovered. Left mandible (Fig. 133D) with

rounded “teeth” on pars incisiva, slender hook-like lacinia

mobilis, pars molaris flaccid, lanceolate, directed proximally.

Labium (Fig. 133H) with prominent setose mediodistal

processes. Maxillule (Fig. 133E) with ten distal spines and

proximal tufts of setae, palp not recovered. Maxilla (Fig.

133F) linguiform, naked. Maxilliped palp (Fig. 133G) first

article naked, second article with three inner distal setae; third

article with three inner setae; fourth article distally with two

shorter and two longer setae, longer setae finely denticulate in

distal half; basis naked; endites distally with two setae and

outer rounded tubercle. Epignath (Fig. 1331) elongate, distally

pointed, naked.

Cheliped (Fig. 134A) basis compact, 0.7 times as long as

wide, naked; merus subtriangular with single ventral seta;

carpus 1.5 times as long as wide, with two unequal midventral

setae, one fine dorsodistal seta and one fine mid-dorsal seta;

propodus as long as wide, fixed finger 0.9 times as long as

palm, with one ventral seta, three setae adjacent to cutting

edge, tooth-like apophyses centrally and distally on cutting

edge; dactylus stout, naked.

Pereopod 1 (Fig. 134B) coxa with seta; basis slightly

arcuate, slender, 5.2 times as long as wide, naked; ischium

compact, with ventral seta; merus 0.4 times as long as basis,

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

199

Fig. 133. Paragathotanais wurundjeri sp. nov., female paratype. A, whole body dorsally; B, antennule; C, antenna; D, left mandible; E, maxillule

endite; F, maxilla; G, maxilliped; H, labium; I, epignath. Scale A = 1 mm, B-I = 0.1 mm.

200

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 134. Paragathotanais wurundjeri sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5;

G, pereopod 6; H, pleotelson and uropods, ventral. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

201

ventrodistally with seta and elongate slender spine 0.6 times as

long as merus; carpus as long as merus, distally with dorsal

finely-denticulate spine longer than carpus and one simple

shorter and one finely-denticulate longer ventral spines;

propodus 1.6 times as long as carpus, with two subdistal setae,

shorter ventral spine; dactylus with proximal seta, slender

unguis 1.8 times as long as dactylus, both together 0.8 times as

long as propodus. Pereopod 2 (Fig. 134C) similar to pereopod

1, but merus 0.85 times as long as carpus, propodus without

subdistal setae. Pereopod 3 compact (Fig. 134D), similar to

pereopod 2, basis with ventral penicillate seta.

Pereopod 4 (Fig. 134E) coxa naked; basis slightly stouter

than those of anterior pereopods, 4.7 times as long as wide,

with two midventral penicillate setae; ischium with two

ventrodistal setae; merus one-quarter as long as basis, with

two finely-denticulate ventrodistal spines; carpus 1.2 times as

long as merus, distally with two ventral and one dorsal finely-

denticulate spines and fine mesial seta; propodus as long as

carpus, distally with two ventral finely-denticulate spines, and

one longer and one shorter dorsal setae; dactylus sinuous, with

fields of microtrichia, 1.6 times as long as curved unguis, both

together 1.5 times as long as propodus. Pereopod 5 (Fig. 134F)

as pereopod 4, but carpus with lateral rather than dorsodistal

spine; propodus with three short dorsodistal spines and no

setae. Pereopod 6 (Fig. 134G) as pereopod 5, but propodus

with microtrichia and four dorsodistal spines.

Pleopods absent.

Uropods (Fig. 134H) held ventrally beneath anterior of

pleotelson; basis naked; exopodal process shorter than

endopod, with one distal seta; endopod of one segment, with

one subdistal penicillate seta, four simple and one or two

penicillate distal setae.

Male. Unknown.

Etymology. The Wurundjeri were one of the indigenous hunter-

gatherer tribes of the Melbourne region, from whom John

Batman, in 1835, negotiated a “purchase” of 2,400 km 2 of land

which became the site of the original settlement which

developed into the city of Melbourne (noun in apposition).

Remarks. Larsen (2005) gave a key to the females of

Paragathotanais, in which the present species fails at couplet 5

in having a relatively stout cheliped, but “shoulders” (super-

coxal processes) on the pereonites. In addition to the species

listed that publication, Guerrero-Kommritz (2003) described

P. insolitus from the Angola basin, but that species is distinct in

having only four articles in the antenna, and very reduced

spination of the anterior pereopods; P. abyssorum Larsen,

2007 is distinguished from P. wurundjeri sp. nov. by the much

more compact antennule, the more slender cheliped and the

smaller spines on the anterior pereopods, inter alia (Larsen,

2007); P. vikingus Bird, 2010 has far more compact pereonites,

a characteristically elongate and parallel-sided cephalothorax,

and a distinct mandibular morphology (Bird, 2010); P. zevinae

(Kudinova-Pasternak, 1970), moved to this genus from

Paranarthrura by Larsen (2007), has a more compact antennule

and a more slender cheliped (Kudinova-Pasternak, 1970). All

of these species have a smaller exopodal process on the uropod.

Finally, P. ipy Jozwiak & Bfazewicz-Paszkowycz, 2011, differs

from P. wurundjeri in having no ventrodistal spine on the

merus of pereopods 1 to 3.

Paragathotanais wurundjeri is the shallowest-recorded

species of the genus discovered so far, having been collected

on muddy sand at 56 to 120 m depth in the Eastern Bass Strait.

All of the other species occur at depths greater than 200 m, and

mostly at depths greater than 2000 m.

Genus Ozagathus gen. nov.

Diagnosis. Agathotanaid with three-articled antennule,

apparent five-articled antenna; pereonites wider medially,

pleon slightly narrower than pereon; mandibular molar

membranous and directed proximally, lacinia mobilis reduced

to apophysis; labium with prominent rounded mediodistal

processes; maxilliped bases and endites naked; cheliped

without pseudocoxa; pereopod coxae unfused; anterior

pereopods with carpal spines; dactyli and ungues of posterior

pereopods not fused, their carpi with two spines; uropod

exopod a fused process on basis, endopod one-segmented,

uropods held beneath pleotelson. Female without pleopods.

Male with slightly more robust antennule and with poorly-

developed pleopods.

Type species. Ozagathus watharongus sp. nov. by monotypy.

Etymology. From “Oz”, colloquial slang for “Australia”, and

“agathus” derived from the prefix to the Family name (male).

Remarks. The new species described below shows all the

general features of an agathonataid, but is somewhat

intermediate between the previously recognized genera, having

the three-articled antennule and reduced uropods typical of

Agathotanais Hansen, 1913, but an apparently five-articled

antenna more typical of Paragathotanais. The mouthparts are

generally within the range of morphology shown by the Family,

although the setulose rounded distal processes on the labium

are presently characteristic of Ozagathus gen. nov. Whether the

tuberculation of the cheliped, a feature not previously described

for an agathotanaid, is a generic character is impossible to say

at present, particularly in the light of the variable presence of

this feature in some genera of the Tanaellidae (see above).

Ozagathus watharongus sp. nov.

Figures 135-137

Material examined. 1 9 (J58854), holotype, MSL EG120, Eastern Bass

Strait, 11.7 km Wof Pt Ricardo, 37°49.54'S 148°30.01'E, 29 m depth, 28

September 1990, coll. Marine Science Laboratories. 1 9 (J56381),

paratype, MSL-EG 68, Eastern Bass Strait, 13.3 km E of eastern edge

of Lake Tyers, 37°51.42’S 148°14.36'E, 121 m depth, 04 June 1991, coll.

N. Coleman; 1 9 (J23653), paratype, MSL-EG 41, Eastern Bass Strait,

11.7 km W of Pt Ricardo, 37°49.54'S 148°30.01'E, 29 m depth, sand and

shell, 28 September 1990, Smith-Mclntyre Grab, coll. Marine Science

Laboratories. 1 9 (J50793), paratype, VC 48 Cl, Eastern Bass Strait,

37°21.43'S 149°29.57'E, 40 m depth, 09 May 1998, coll. N. Coleman,

Smith-Mclntyre Grab. 1 9 (J28426), paratype, MSL-EG 121, Eastern

Bass Strait, 11.7 km W of Pt Ricardo, 37°49.54’S 148°30.0PE, 29 m

depth, sand and shell, 28 September 1990, Smith-Mclntyre Grab, coll.

Marine Science Laboratories. 17 99 (J28413), paratypes, MSL-EG 86,

202

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 135. Ozagathus watharongus sp. nov., female holotype. A, dorsal view; B, lateral view. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

203

Fig. 136. Ozagathus watharongus sp. nov. A, female antennule; B, female antenna; C, male antennule; D, male antenna; E, labrum; F, left

mandible; G, maxillule; G‘, maxillule palp; H, labium; I, maxilliped; J, epignath. Scale = 0.01 mm.

204

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 137. Ozagathus watharongus sp. nov. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5; G, pereopod 6;

H, male pleopod; I, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

205

Eastern Bass Strait, 2.9 km SE of Cape Conran, 37°50.00'S 148°38.54'E,

95 m depth, coarse sand, 04 June 1991, Smith-Mclntyre grab, coll. N.

Coleman. 30 $$ (J28416), paratypes, MSL-EG 90, Eastern Bass Strait,

7.3 km SSW of Cape Conran, 37°52.39'S 148°42.09'E, 161 m depth,

coarse sand, 04 June 1991, Smith-Mclntyre grab, coll. N. Coleman. 2 9

9 (J50792), paratypes, Stn VC 37 Cl, Eastern Bass Strait, 38°18.3’S

147°15.25’E, 40 m depth, 10 May 1999, Smith-Mclntyre grab, coll. N.

Coleman. 19 99 (J28420), paratypes, MSL-EG 114, Eastern Bass Strait,

2.9 km SE of Cape Conran, 37°50.00’S 148°38.54’E, 29 m depth, coarse

sand, February 1991, Smith-Mclntyre grab, coll. N. Coleman. 4 99

(J23525), paratypes, MSL-EG 45, Eastern Bass Strait, 13.3 km E of

eastern edge of Lake Tyers, 37°51.44'S 148°14.46'E, 37 m depth, sand

and shell, 25 September 1990, Smith-Mclntyre grab, coll. Marine

Science Laboratories. 22 99 (J23532), paratypes, MSL-EG 59, Eastern

Bass Strait, 9.5 km SW of Cape Conran, 37°52.53'S 148°39.29'E, 48 m

depth, sand and shell, 28 September 1990, Smith-Mclntyre grab, coll.

Marine Science Laboratories.

Description of female. Body (Fig. 135A, B) slender, holotype

1.9 mm long, 8.7 times as long as wide. Cephalothorax

subrectangular, tapering in anterior third with slight triangular

rostrum, 1.6 times as long as wide, almost as long as pereonites

1 and 2 together, wider than pereon, naked; eyelobes and eyes

absent. Pereonites almost barrel-shaped, pereonite 1 widest

anteriorly, pereonites 2 to 5 centrally, pereonite 6 posteriorly;

pereonites 1 and 5 subequal in length, half as long as

cephalothorax; pereonites 2 and 4 just longer than pereonite 1;

pereonite 3 longest, 1.3 times as long as pereonite 1; pereonite

6 shortest, 0.8 times as long as pereonite 1 (all pereonites

respectively 1.2, 1.0, 0.9, 1.0, 1.1 and 1.3 times as wide as long).

Pleon just narrower than pereon, pleonites without pleopods,

each 4.5 times as wide as long; pleonite 5 with conspicuous

midlateral seta on each side. Pleotelson subpentangular, 0.4

times length of pleon and as wide as long.

Antennule (Fig. 136A) of three articles, proximal article

4.2 times as long as wide, 1.35 times as long as distal two

articles together, outer margin with mid-length and distal tufts

of four penicillate and one simple setae, mid-length seta

exceeding tip of proximal article, distal seta exceeding tip of

antennule; second article as long as wide, 0.2 times as long as

first article, with simple outer distal seta; third article 1.2 times

as long as second article, with five simple and one penicillate

distal setae.

Antenna (Fig. 136B) of five apparent articles, proximal

article totally fused to carapace; second article as long as wide,

naked; third article just longer than wide, as long as second

article, with fine dorsodistal seta; fourth article longest, 2.2

times as long as second article and nearly four times as long as

wide, with subdistal ventral penicillate seta; fifth article one-

third as long as fourth, naked; sixth article half as long as fifth,

distally with three simple and one penicillate setae.

Labrum (Fig. 136E), compact, rounded, distally setulose.

Left mandible (Fig. 136F) with three rounded lobe-like “teeth”

on pars incisiva, lacinia mobilis reduced to a similar rounded

subdistal tubercle, pars molaris flaccid, blunt, directed

proximally. Labium (Fig. 136H) with prominent setose,

rounded mediodistal processes. Maxillule (Fig. 136G) with

eight distal spines and sparse groups of microtrichia, palp

(Fig. 136G') with two distal setae. Maxilla not recovered.

Maxilliped (Fig. 1361) palp first article naked, second article

with two inner distal setae; third article with two inner setae in

distal half; fourth article tapering, with five setae along inner

margin to tip, single outer subdistal seta; all palp setae other

than the last distally finely denticulate; bases fused, naked;

endites distally with rudiment of outer rounded tubercle.

Epignath (Fig. 136J) elongate, distally pointed, naked.

Cheliped (Fig. 137A) basis compact, 0.85 times as long as

wide, naked; merus subtriangular, outer face with strip of six

rounded tubercles and single seta along dorsodistal margin;

carpus 1.6 times as long as wide, with two unequal midventral

setae, one fine dorsodistal seta and one fine mid-dorsal seta,

and strip of five rounded tubercles along ventral margin of

outer face; propodus longer than wide with ventral submarginal

strip of six rounded tubercles and group of smaller tubercles

dorsodistally, fixed finger 0.8 times as long as palm, with one

ventral seta, three setae adjacent to cutting edge, small “teeth”

centrally and distally on cutting edge; dactylus with rounded

tubercles along dorsal margin and proximally on inner face,

two blunt tubercles on cutting edge.

Pereopod 1 (Fig. 137B) coxa with seta; basis slightly arcuate,

slender, five times as long as wide, naked; ischium compact, with

ventral seta; merus 0.3 times as long as basis, ventrodistally with

two setae; carpus 1.3 times as long as merus, distally with one

dorsal, and one shorter and one longer ventral finely-denticulate

spines; propodus 1.5 times as long as carpus, ventrally with

subdistal spine and seta; dactylus short and robust, tapering

unguis 1.7 times as long as dactylus, both together 0.7 times as

long as propodus. Pereopod 2 (Fig. 137C) similar to but somewhat

stouter than pereopod 1, basis 3.7 times as long as wide, propodus

without subdistal seta but with dorsodistal spine-like apophysis.

Pereopod 3 (Fig. 137D), similar to pereopod 2.

Pereopod 4 (Fig. 137E) basis 4.2 times as long as wide;

ischium with two ventrodistal setae; merus 0.4 times as long

as basis, with two finely-denticulate ventrodistal spines; carpus

as long as merus, distally with outer and inner finely-

denticulate spines and fine dorsal seta; propodus 1.3 times as

long as carpus, distally with three finely-denticulate spines

and dorsodistal spine-like apophysis; dactylus about half as

long as curved unguis, unguis slender, finely denticulate, both

together 1.3 times as long as propodus. Pereopod 5 (Fig. 137F)

as pereopod 4, but basis with two penicillate setae. Pereopod 6

(Fig. 137G) as pereopod 4, but basis naked.

Pleopods absent.

Uropods (Fig. 1371) held ventrally beneath pleotelson;

basis naked but with slight fused exopod with two distal setae

exceeding tip of endopod and visually conspicuous in dorsal

view of animal (Fig. 135); endopod of one segment, widest

proximally, with one simple and two penicillate setae in

proximal half, three simple and two penicillate setae distally.

Distinctions of male. Of similar overall appearance to female;

antennule (Fig. 136C) stouter, proximal two articles respectively

3.5 and 0.7 times as long as wide; antenna (Fig. 136D) also

slightly stouter, second and third articles shorter than wide,

fourth article three times as long as wide; pleopods present

(Fig. 137H), somewhat rudimentary, biramous, rami with

incomplete articulation with naked basis, each with distal tuft

of fine setules.

206

M. Biazewicz-Paszkowycz & R.N. Bamber

Etymology. The Wathaurong were another of the indigenous

hunter-gatherer tribes of the Melbourne region in the mid¬

nineteenth century (see under Paragathotanais wurundjeri

above) (noun in apposition).

Remarks. See above under Remarks for the genus. Ozagathus

watharongus sp. nov. was collected frequently in the Eastern

Bass Strait, at depths from 29 to 161 m on coarse or shelly sands.

Family Akanthophoreidae Sieg, 1986

Genus Gejavis gen. nov.

Diagnosis of female. Similar to Akanthophoreus, uropods

biramous, endopod and exopod two-segmented; molar process

tapering with several terminal spines; maxillule endite with

nine distal spines; maxilliped basis naked; maxilliped palp

article 2 with three inner plumose setae, outer margin naked,

article 3 with three plumose and one simple inner setae;

cheliped carpus without ventral shield but with rugose dorsla

margin, propodus with coarse rugosity; pereopod 1 merus

ventrodistally with one seta and one slender spine, carpus with

dorsodistal seta, ventrodistally naked; dactyli without groove

or small spines; pereopods 4 to 6, unguis of anterior pereopods

much longer than dactylus; pereopods 4 to 6, carpus with three

spines and one seta. Pleonites with long setae.

Type species. Gejavis corsotos sp. nov. by monotypy.

Etymology. Named cryptically after Dr Graham Gird (G J

being his initials, and avis being Latin for a bird) for his

invaluable contributions to tanaidacean taxonomy and

phylogeny (female).

Remarks. The new species described below shows close

affinities to Akanthophoreus Sieg, 1986 (see Bird, 2007, for

diagnosis and discussion of that genus) and Chauliopleona

Dojiri and Sieg, 1997, but is distinguished in the comparatively

reduced spination of the merus and carpus of pereopod 1, in the

simpler conformation of the dactyli of all pereopods, in having

the uropod exopod longer than the proximal endopod segment,

in the absence of a seta on the maxilliped basis and in the

maxilliped palp setation. The new genus differs from

Paraleptognathia Kudinova-Pasternak, 1981 in that the

cheliped is rugose trather than setulose, and is without a

ventrodistal carpal shield, pereopod 1 is without the complex

spinulation of that genus, and again the conformation of the

pereopod dactyli and the uropod rami are distinct. There

remain some species currently assigned to Leptognathia Sars,

1882 with two-segmented uropod exopods which will

presumably be reallocated once the confounded classification

of this group of tanaidaceans is resolved better ( Leptognathia

sensu stricto is currently regarded as having a one-segmented

uropod exopod, e.g. Larsen & Shimomura, 2007a); again,

Gejavis gen. nov. is largely distinguished from those species by

its pereopod 1 spination and uropod exopod conformation.

The rugosity of the cheliped in the present genus is also

distinct from those of other akanthophoreids, although it is not

possible at this stage to say whether this feature is a generic or

specific characteristic.

Biazewicz-Paszkowycz and Bamber (2011) elevated Sieg’s

(1986a) subfamily Akanthophoreinae to familial rank to

accommodate the genus Akanthophoreus (at least), as, despite

this genus having its own higher taxon, being the type genus of

that subfamily, it had been left unclassified (“Family incertae

sedis ”) in a number of recent phylogenetic and taxonomic works

on the genus (e.g. Larsen & Wilson, 2002; Bird, 2007). The new

genus described here is clearly close to Akanthophoreus, and so

is placed within the same Family.

Gejavis corsotos sp. nov.

Figures 138-140

Material examined. 1 ? (J58562), holotype, CPBS 03S, Western Port off

Crib Point, 38°21.65'S 145 0 15.21'E, 2 m depth, sandy-mud, 13 April

1965, Smith-Mclntyre grab, coll. A. J. Gilmour; 1 $ (J23596), paratype,

Stn MSL-EG 40, Eastern Bass Strait, 11.7 km W. of Pt Ricardo, 37°49.90'S

148°30.01'E, 29 m depth, sand-shell, 28 September 1990, R.V. Sarda,

Smith-Mclntyre Grab. 1 ? (J62057), paratype, Stn. BSS 31, Eastern Bass

Strait, 22 km NNE of North Point, Flinders Island, 39°34.3'S 148°04.0'E,

37 m depth, coarse sand, 26 March 1979, dredge, coll. G.C.B. Poore. 1 9

(J56376), paratype (dissected), Stn MSL-EG 69, Eastern Bass Strait, 13.3

km E of eastern edge of Lake Tyers, 37°51.7'S 148°14.6'E, 37 m depth, 04

June 1991, coll. N. Coleman, Smith-Mclntyre Grab.

Description of female. Body (Fig. 138A, B) elongate, slender,

2.4 mm long, nine times as long as wide. Cephalothorax

subrectangular, tapering from mid-length towards anterior, 1.4

times as long as wide, with slight rounded rostrum, naked; eyes

absent. Six free cylindrical pereonites; pereonites 1 to 5 about as

long as wide, subequal in length and 0.6 to 0.7 times as long as

cephalothorax; pereonite 6 shortest, less than half as long as

cephalothorax and 1.3 times as wide as long. All pleonites

bearing pleopods, each pleonite with one midlateral seta on each

side. Pleotelson rounded, almost three times as long as pleonite 5,

1.3 times as wide as long, with paired distal and laterodistal setae.

Antennule (Fig. 139A) of four articles, proximal article 1.9

times as long as wide, 0.6 times as long as last three articles

together, with mesial and distal outer tufts of penicillate setae

and single inner and outer simple distal setae; second article

1.6 times as long as wide, 0.7 times as long as first article, with

one outer distal penicillate seta and single inner and outer

simple distal setae; third article half length of second with one

outer distal penicillate seta and single inner and outer simple

distal setae; distal article more slender, 1.6 times as long as

third article, with three simple and one penicillate distal setae

and single aesthetasc.

Antenna (Fig. 139B) of six articles, proximal article naked;

second article slightly inflated with one small dorsodistal seta;

third article half length of second article, with single

dorsodistal seta three-times as long as article; fourth article

longest, as long as articles 1 to 3 together and 3.5 times as long

as wide, with four distal penicillate setae and single simple

distal seta exceeding tip of antennule; fifth article 0.4 times as

long as fourth, with one distal seta; sixth article 0.25 times as

long as fifth, with four distal setae.

Labrum (Fig. 139C) apically blunt, finely setose. Left

mandible not recovered; right mandible (Fig. 139D) with

bilobed pars incisiva, pars molaris basally stout, tapering, with

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 138. Gejavis corsotos sp. nov., female holotype. A, dorsal view; B, lateral view. Scale = 1 mm.

208

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 139. Gejavis corsotos sp. nov., female paratype. A, antennule; B, antenna; C, labrum; D, right mandible; E, maxillule; F, maxilla; G, labium;

H, maxilliped. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

209

Fig. 140. Gejavis corsotos sp. nov., female paratype. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5;

G, pereopod 6; H, pleopod; I, uropod. Scale = 0.1 mm.

210

M. Biazewicz-Paszkowycz & R.N. Bamber

several distal spinules. Labium (Fig. 139G) with outer distal

and subdistal microtrichia. Maxillule (Fig. 139E) with nine

distal spines and outer brush of setules, palp damaged. Maxilla

(Fig. 139F) quadrangular, simple, naked. Maxilliped (Fig.

139H) endites each with inner-distal rounded tubercle and

slightly crenulated distal margin; palp first article with outer

distal seta, second and third articles with three inner plumose

setae, third article with additional inner simple seta; fourth

article with one inner and three distal plumose setae and one

outer simple seta; basis naked. Epignath not recovered.

Cheliped (Fig. 140A) basis with large posterior lobe, wider

proximally and extended past anterior margin of pereonite 1

ventrally, about 1.8 times as long as wide; merus subtriangular

with one mid-ventral seta; carpus twice as long as wide with

paired mid-ventral setae and no shield, proximal and distal

single dorsal setae, proximal half of dorsal margin coarsely

rugose; propodus longer than wide, with coarse rugosity over

outer face and along dorsal margin; inner comb-row of three

setae; fixed finger short with lamellate cutting edge, two

ventral and three inner setae; dactylus with dorsal margin

coarsely rugose, cutting edge naked.

Pereopod 1 (Fig. 140B) basis damaged in preparation,

curved, about 3.5 times as long as wide; ischium compact with

single seta; merus just shorter than carpus with single

ventrodistal seta and spine; carpus with dorsodistal seta;

propodus 1.9 times as long as carpus, with single dorsal and

ventral subdistal setae; dactylus half as long as slender unguis,

both together 1.4 times as long as propodus. Pereopod 2 (Fig.

140C) similar to but shorter than pereopod 1; basis 2.5 times

as long as wide; merus as long as carpus and with two

ventrodistal spines; carpus with one dorsal and two ventral

distal spines; unguis 1.7 times as long as dactylus, both

together 0.7 times as long as propodus. Pereopod 3 (Fig. 140D)

as pereopod 2, but basis with dorsoproximal penicillate seta.

Pereopod 4 (Fig. 140E) basis 2.8 times as long as wide;

ischium with one ventral seta; merus wider distally, 0.4 times

as long as basis, with two ventrodistal spines; carpus 0.9 times

as long as merus, with fine dorsodistal spines and three stouter

mid- and ventrodistal spines; propodus with one longer

dorsodistal and two shorter ventrodistal spines, all shorter than

dactylus; dactylus twice as long as unguis, both together 1.4

times as long as propodus. Pereopod 5 (Fig. 140F) as pereopod

4, but basis broader, 2.4 times as long as wide, and dorsodistal

spine on [propodus as long as dactylus. Pereopod 6 (Fig. 140G)

as pereopod 5, but propodus with three dorsodistal spines.

Pleopods (Fig. 140H) all alike, with naked basis; endopod

shorter than exopod and with seven mainly distal plumose

setae; exopod with ten distal and outer plumose setae, proximal

seta on ventral margin well-separated from remaining setae,

most setae about as long as rami.

Uropod (Fig. 1401) basis naked; exopod of two subequal

segments, longer than proximal endopod segment, proximal

segment with one distal seta, distal segment with two unequal

distal setae; endopod of two subequal segments, proximal

segment with one seta, distal segment with one subdistal and

four distal simple setae and two distal penicillate setae.

Male. Unknown.

Etymology. From the Greek korsotos - “shorn”, referring to the

relative lack of spines or setae on pereopod 1 of the present

species.

Remarks. See above under Remarks for the genus. Gejavis

corsotos sp. nov. was taken from sandy substrata at 2 to 37 m

depth, in Western Port and the Eastern Bass Strait.

Family Tanaopsidae fam. nov.

Diagnosis. Generally leptognathioid ( sensu lato) facies, pleon

laterally convex and wider than pereon, antennule in female of

four longer articles with or without minute distal article, in male

with multisegmented flagellum; mandible pars incisiva distally

rounded with adjacent serrated incisive margin, molar process

either slender and pointed or absent; maxillule endite reflexed

through about 90° and with five or six distal spines, one

significantly more robust than the others; maxilliped endite

flared. Cheliped with triangular, dorsally-inserted sclerite, fixed

finger with two ventral setae, bifid terminal spine and bifid or

trifid distal denticle on incisive margin. Pereopods 1 to 3 with

setae, unguis longer than slender dactylus, both together about

as long as or longer than propodus, dactylus with proximal

seta; pereopods 4 to 6 with spines on merus, carpus (three in

number) and propodus. Pleopods with plumose seta along

entire outer margin of exopod, but restricted to distal half of

outer margin of endopod; uropods biramous, rami with one or

two segments.

Type genus: Tanaopsis Sars, 1896

Remarks. The present suprageneric classification of the

Paratanaoidea is in a state of flux, owing to recent attempts at

phylogenetic resolution involving cladistics, based on meristics

and morphometries (Larsen & Wilson, 2002; Biazewicz-

Paszkowycz & Poore, 2008; Bird & Larsen, 2009). The only of

these studies to consider the genus Tanaopsis were Larsen &

Wilson (2002) and Biazewicz-Paszkowycz & Poore (2008), but

they were unable to resolve it to a family as defined by their

generated clades.

The diagnostic features listed above taken together

distinguish members of the genus Tanaopsis from all other

paratanaoid genera of the leptognathioid sensu lato facies; that

said, they constitute a diagnosis of the genus itself. Equally,

exclusion of the unique features of the cheliped fixed finger

does not allow inclusion of any other genera. Further detailed

cladistic analyses may associate other genera, which would

then require qualifying the familial diagnosis above, removing

characters not consistent across all associated genera into the

diagnosis of the genus Tanaopsis. Bird (2011) suggests possible

affinities with Cristatotanais Kudinova-Pasternak, 1990

(including Spinitanaopsis Larsen, 2005).

It is further the case that Tanaopsis itself may not be

monophyletic. There appear to be two groups of species, one

with a pointed mandibular molar process and two-segmented

uropod rami ( T. antarcticus Lang, 1967; T. cadieni Sieg &

Dojiri, 1991; T. curtus Kudinova-Pasternak 1984; T. gallardoi

(Shiino, 1970); T. profunda Lang, 1967; T. canaipa Bamber,

2008 and one of the two species described below), the other

without a molar process and with one-segmented uropod rami

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

211

( T. chotkarakde Bird & Bamber, 2000; T. kerguelenensis Shiino,

1979 [mandible unknown]; and one of the two species described

below); the latter group also tend to have a more slender

antennule. That said, T. laticaudata Sars, 1882 is described as

being without a mandibular molar (Sars, 1896), but has two-

segmented uropod rami. It is at present not possible to

distinguish such groups as separate genera, as the type species

of Tanaopsis sensu stricto, T. graciloides (Lilljeborg, 1864)

needs proper redescription based on material from the

northwest Atlantic (see Bamber et al., 2009).

Genus Tanaopsis Sars, 1896

Tanaopsis boonwurrungi sp. nov.

Figures 141-143

Material examined. 1 9 (J57793), holotype, 1 9 (J58563), paratype,

dissected. Western Port, off Crib Point, Stn CPBS-N 03, 38°20.57'S

145°15.08'E, 2 m depth, fine sand, 05 April 1965; 1 9 (J57792),

paratype. Western Port, off Crib Point, Stn CPBS 3IN, 38°20.93'S

145°13.62'E, 15 m depth, fine sand with mud, 29 March 1965; all coll.

A J. Gilmour.

Description of female. Body (Fig. 141) slender with widened

pleon, holotype 3 mm long, 8.7 times as long as wide.

Cephalothorax pear-shaped, widest and laterally-rounded

posteriorly, tapering towards anterior with slight rounded

rostrum, as long as maximum width, shorter than pereonites 1

and 2 together, naked; eyelobes present, eyes apparently absent

in preserved material. Pereonite 1 shortest, 0.4 times as long as

cephalothorax, laterally convex; pereonite 2 nearly twice as

long as pereonite 1, laterally convex; pereonite 3 (and

subsequent pereonites) with parallel sides, 2.5 times as long as

pereonite 1; pereonite 4 longest, longer than wide and 3.2

times as long as pereonite 1; pereonite 5 just shorter than

pereonite 4, pereonite 6 just longer than pereonite 2 (all

pereonites respectively 2.3, 1.4, 1.0, 0.8, 0.9 and 1.2 times as

wide as long). Pleon with five free pleonites bearing pleopods;

first pleonite trapezoidal, longest, as long as pereonite 1 and

2.7 times as long as wide; each remaining pleonite 0.8 times as

long as first pleonite and 3.9 times as wide as long. Pleotelson

pentangular, one-quarter length of pleon and 1.7 times as wide

as long, with two slender distal spines.

Antennule (Fig. 142A) of five articles (four longer articles

plus minute distal article), proximal article 2.1 times as long as

wide, just shorter than distal four articles together, outer

margin with tufts of three penicillate setae at mid-length and

two penicillate and one simple setae distally; second article as

long as wide, 0.35 times as long as first article, distally with

single inner and outer simple setae and three penicillate setae;

third article compact, 0.7 times as long as second article,

distally with single inner and outer simple setae; fourth article

tapering, nearly twice as long as third article, with single

simple distal seta; fifth article (Fig. 142A') minute, with

aesthetasc and four simple and one penicillate distal setae.

Antenna (Fig. 142B) of six articles, proximal two articles

not recovered; third article longer than wide, with fine

dorsodistal seta longer than article and distal microtrichia;

fourth article longest, 2.7 times as long as third article, 4.5

times as long as wide, slightly curved, with three simple and

three penicillate distal setae; fifth article as long as third with

one distal seta; sixth article minute with five distal setae.

Labrum not recovered. Mandibles (Fig. 142C) without

lacinia mobilis or pars molaris, pars incisiva with saw-like row

of denticulations and rounded distal apophysis, larger on right

mandible. Labium (Fig. 142F) with prominent, finely setulose

mediodistal processes. Maxillule (Fig. 142D) with five finely-

denticulate distal spines, one stouter than the others, and outer

tufts of setules, palp not recovered. Maxilla (Fig. 142E)

linguiform but basally cupped, with fine marginal setules.

Maxilliped palp (Fig. 142G) first article naked, second article

with one outer and one inner distal setae; third article with

four slender and curved inner setae; fourth with five distal

setae and one outer subdistal seta; basis with single seta

reaching distal margin of proximal palp article; endite distally

naked. Epignath (Fig. 142H) elongate, linguiform, naked.

Cheliped (Fig. 143A) basis 1.6 times as long as wide,

naked; merus subtriangular with single ventral seta; carpus

stout, 1.2 times as long as wide, with two midventral setae, one

mid-dorsal and one dorsodistal setae; propodus stout, as long

as wide, fixed finger 0.6 times as long as palm, with two ventral

setae, three setae on cutting edge, distal claw with typical

inner and outer bifurcate apophyses; dactylus dorsally finely

crenulate, with slender spinules along cutting edge and fine

proximal seta.

Pereopod 1 (Fig. 143B) longer than others, coxal apophysis

(Fig. 143B') large, pointed, with seta; basis slender, 5.7 times

as long as wide, naked; ischium compact, naked; merus 0.7

times as long as carpus, wider distally, naked; carpus with

single dorsal and ventral distal setae; propodus 1.7 times as

long as carpus, with three dorsal subdistal setae, one ventral

subdistal seta; dactylus half as long as unguis, unguis slender

and as long as propodus. Pereopod 2 (Fig. 143C), coxa rounded

with seta; basis 4.3 times as long as wide; ischium with seta;

merus 0.7 times as long as carpus, with single ventrodistal

seta; carpus with two dorsal and one ventral distal setae;

propodus 2.2 times as long as carpus, with two dorsal subdistal

setae, one ventral subdistal seta; slender unguis longer than

dactylus, both together 1.2 times as long as propodus. Pereopod

3 (Fig. 143D) similar to pereopod 2.

Pereopod 4 (Fig. 143E) basis stout, 2.1 times as long as

wide, naked; ischium with two ventrodistal setae; merus as

long as carpus, with ventral field of microtrichia and two small

distally-denticulate ventrodistal spines; carpus with one

dorsodistal seta and three small distally-denticulate

ventrodistal spines; propodus 1.6 times as long as carpus, with

fields of microtrichia, mid-dorsal penicillate seta, one

dorsodistal and two ventrodistal spines all distally finely

denticulate; dactylus slender, with fields of microtrichia, 1.5

times as long as unguis, both together 0.9 times as long as

propodus. Pereopod 5 (Fig. 143F) as pereopod 4, but basis

with two ventral penicillate setae. Pereopod 6 (Fig. 143G) as

pereopod 4, but propodus with three dorsodistal spines.

Pleopods (Fig. 143H) all alike, with naked basis, endopod

shorter than exopod and with rounded proximal apophysis

on inner margin; endopod with inner subdistal plumose seta

and 14 plumose setae along the distal half of the outer

212

margin, exopod without setae on inner margin, outer margin

with 27 plumose setae, proximal setae on rami not separated

from others.

Uropod (Fig. 1431) biramous, basis naked; exopod and

endopod each of one segment, exopod shorter than endopod,

with one fine proximal, one shorter and one longer distal setae;

endopod with distal penicillate seta on first segment and five

simple and one penicillate distal setae on second segment.

Male. Unknown.

Etymology. The Boonwurrung were another of the indigenous

hunter-gatherer tribes of the (now) Melbourne region in the

mid-nineteenth century (see under Paragathotanais

wurundjeri above).

Remarks. Sieg and Dojiri (1991) gave a key to the genus

Tanaopsis for the species then known, in which the present

species keys out to the generotype, T. graciloides with which

they included T. laticaudata as a synonym (as had most

previous authors). Bamber et al. (2009) cast doubt on this

synonymy, maintaining the distinction of Sars’ Mediterranean

species from Lilljeborg’s Northeast Atlantic-Subarctic species

until a proper redescription of the latter was undertaken; all

descriptions and figures referred to in the recent literature for

T. graciloides, including by Sieg and Dojiri (1991) are from

Sars (1882). Bird & Bamber (2000), while describing as new

T. chotkarakde, also added T. gallardoi to the genus. Since

then, the only new Tanaopsis species that have been described

are T. canaipa from Queensland and T. rawhitia Bird, 2011

from New Zealand.

Tanaopsis boonwurrungi sp. nov. shares the lack of a

molar process on the mandible only with T. graciloides/

laticaudata and T. chotkarakde, and possibly T. kerguelenensis

(mandible not described, but a species also with one-segmented

uropodal rami). T. boonwurrungi is immediately distinguished

from all of these taxa by its distinctly more slender habitus,

with pereonites 4 and 5 longer than wide, and a much more

slender cheliped, while T. chotkarakde has a lacinia mobilis on

the left mandible, and the two European taxa have a two-

segmented uropod exopod.

The small distal article on the antennule has only been

reported before for a Tanaopsis species by Bird (2011),

who noted its presence in T. rawhitia. While such an article

does not appear to have been present in some other more

recently or better-described species, viz. T. kerguelenensis,

T. cadieni, T. chotkarakde or T. canaipa, the possibility of its

having been overlooked in some of the earlier descriptions

cannot be dismissed.

Tanaopsis boonwurrungi was taken only in Western Port

at 2 to 15 m depth on fine sand.

Tanaopsis oios sp. nov.

Figures 144-145

Material examined. 1 ? (J58547), holotype (on microscope slide), 1

further ? (lost). Eastern Bass Strait, 28 km SSW of Mario, Stn BSS

207,37°59'S 148°27'E, 51 m depth, muddy sand and fine shell, 30 July

1983; coll. M.F. Gomon & R.S. Wilson.

M. Blazewicz-Paszkowycz & R.N. Bamber

Fig. 141. Tanaopsis boonwurrungi sp. nov., female holotype, dorsal

view. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

213

Fig. 142. Tanaopsis boonwurrungi sp. nov., female. A, antennule, with A', detail of distal article; B, antenna; C, left and right mandibles;

D, maxillule endite; E, maxilla; F, labium; G, maxilliped; H, epignath. Scale line = 0.1 mm.

214

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 143. Tanaopsis boonwurrungi sp. nov., female. A, cheliped; B, pereopod 1; C, pereopod 2; D, pereopod 3; E, pereopod 4; F, pereopod 5;

G, pereopod 6; H, pleopod; I, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

215

Fig. 144. Tanaopsis oios sp.nov. A, antennule; B, antenna; C, labrum; D, maxillule endite; E, maxilla; F, epignath; G, maxilliped. Scale = 0.1 mm.

216

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 145. Tanaopsis oios sp.nov. A, cheliped; B, detail of chela; C, pereopod 1; D, pereopod 2; E, pereopod 5; F, pereopod 6; G, pleopod;

H, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

217

Description of female. Body slender with widened pleon,

typical for the genus but damaged (not figured).

Antennule (Fig. 144A) of four longer articles but with

incipient articulation of minute distal article, proximal article

1.9 times as long as wide, three-quarters as long as distal three

articles together, outer margin with tufts of three penicillate

setae at mid-length and four penicillate and one simple setae

distally, inner margin with simple distal seta; second article

1.25 times as long as wide, 0.6 times as long as first article,

subdistally with single inner and outer simple setae; third

article compact, 0.4 times as long as second article, distally

with single inner and outer simple setae; fourth article tapering,

2.4 times as long as third article, with single simple subdistal

seta and four simple distal setae.

Antenna (Fig. 144B) of six articles, proximal article

compact, naked; second article as long as wide, 2.5 times as

long as first article, with single dorsodistal and ventrodistal

setae and microtrichia; third article shorter than wide, 0.6

times as long as second article, with fine dorsodistal seta

longer than article; fourth article longest, three times as long

as third article, three times as long as wide, slightly curved,

with two simple and five penicillate distal setae; fifth article

just longer than third, naked; sixth article minute with five

distal setae.

Labrum (Fig. 144C) acorn-shaped, naked. Mandibles and

labium not recovered. Maxillule (Fig. 144D) with five distal

spines, one stouter than the others, and outer tufts of setules,

palp not recovered. Maxilla (Fig. 144E) linguiform but

medially expanded, naked. Maxilliped palp (Fig. 144F) first

article naked, second article with one outer and two inner

distal setae; third article with three inner distal setae; fourth

with six inner to distal setae, distal two much longer than the

others; basis seta not seen; endite distally with inner seta and

setules below outer corner. Epignath (Fig. 144G) elongate,

linguiform, distally hooked, naked.

Cheliped (Fig. 145A) with rounded basis 1.9 times as long

as wide, single dorsodistal seta; merus subtriangular with

single ventral seta; carpus stout, 1.4 times as long as wide,

with two midventral setae, one mid-dorsal and one dorsodistal

setae; propodus stout, 1.5 times as long as wide, fixed finger

0.7 times as long as palm, with two ventral setae, three setae

on cutting edge, distal claw with typical inner and outer

bifurcate rounded apophyses (Fig. 145B); dactylus dorsally

coarsely crenulate.

Pereopod 1 (Fig. 145C) without coxal apophysis, coxa with

seta; basis 3.6 times as long as wide, with dorsoproximal simple

seta; ischium compact, with ventral seta; merus one-third as

long as basis, wider distally, with sparse microtrichia; carpus

0.9 times as long as merus, with three dorsal and single ventral

distal setae; propodus 1.9 times as long as carpus, with three

dorsal subdistal setae, one ventral subdistal seta, sparse dorsal

microtrichia and dorsodistal spine-like apophysis; dactylus

with longer proximal seta, half as long as slender unguis, both

together 1.3 times as long as propodus. Pereopod 2 (Fig. 145D)

basis 3.5 times as long as wide, dorsally with one simple seta

and one penicillate seta in proximal half; ischium with seta;

merus as long as carpus, naked; carpus with two dorsal and one

ventral distal setae and sparse microtrichia; propodus 2.4 times

as long as carpus, with two dorsal subdistal setae, one longer

ventral subdistal seta exceeding tip of dactylus, and sparse

microtrichia; dactylus with longer proximal seta, slender

unguis 1.7 times as long as dactylus, both together as long as

propodus. Pereopod 3 (not figured) similar to pereopod 2.

Pereopod 4 similar to pereopod 5, but basis without

penicillate setae. Pereopod 5 (Fig. 145E) basis relatively stout,

2.3 times as long as wide, with two midventral penicillate setae;

ischium with two ventrodistal setae; merus, carpus and propodus

subequal in length; merus with two small distally-denticulate

ventrodistal spines; carpus with one inner distal seta and three

small distally-denticulate ventrodistal spines; propodus with

fields of microtrichia, mid-dorsal penicillate seta, one

dorsodistal and two ventrodistal spines finely denticulate in

distal half; dactylus with fields of microtrichia, 1.5 times as long

as unguis, both together 0.9 times as long as propodus. Pereopod

6 (Fig. 145F) as pereopod 5, but basis without penicillate setae,

propodus with three dorsodistal spines.

Pleopods (Fig. 145G) all alike, with naked basis, endopod

shorter than exopod and with rounded proximal apophysis on

inner margin; endopod with inner subdistal plumose seta and

nine outer plumose setae in distal half; exopod without setae

on inner margin, outer margins with 21 plumose setae,

proximal seta not separated from others.

Uropod (Fig. 145H) biramous, basis naked; exopod and

endopod each of two subequal segments; exopod distinctly

longer than proximal endopod segment, proximal segment

with one and distal segment with one shorter and one longer

distal setae; endopod with two distal penicillate setae on first

segment and five simple and one penicillate distal setae on

second segment.

Male. Unknown.

Etymology. From the Greek oios - alone, singular, as only one

specimen from a remote part of the Bass Strait remains.

Remarks. Of the previously-described species of Tanaopsis with

two clear segments in both uropod rami, only three have the

uropod exopod exceeding the length of the proximal endopod

segment. In comparison with T. oios sp. nov., T. graciloides sensu

Lang, 1967 has a much less slender cheliped basis, the rugosity on

the cheliped dactylus not restricted to the distal half; a more

slender antennal article 4, and fewer distal setae on anterior

pereopod carpi. T. laticaudata has a more compact cheliped

carpus, the rugosity of the cheliped dactylus more extensive,

fewer distal setae on anterior pereopod carpi, and is without the

long ventral seta on the propodus of pereopod 2. T. cadieni has a

more slender antennal article 4, has a triangular coxal apophysis

on pereopod 1, and is without the long ventral seta on P2 propodus.

T. oios was only found in the Eastern Bass Strait to the east

of the Gippsland Lakes, at 51 m depth on shelly muddy sand.

Family Colletteidae Larsen & Wilson, 2002

Genus Parafilitanais Kudinova-Pasternak, 1989

Parafilitanais vadosus sp. nov.

Figures 146-148

218

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 146. Parafilitanais vadosus sp. nov., female holotype. A, dorsal

view; B, lateral view. Scale = 1 mm.

Material examined. 1 9 (J56378), holotype, Stn MSL-EG 41, Eastern

Bass Strait, 11.7 km W of Pt Ricardo, 37°49.90'S 148°30.02'E, 29 m

depth, 28 September 1990, coll. Marine Sciences Laboratory. Smith-

Mclntyre grab. 1 6 (J28486), paratype dissected, Stn MSL-EG 113,

Eastern Bass Strait, 2.9 km SE. of Cape Conran, 37°50.00'S 148°36.90'E,

29 m depth, coarse sand, February 1991, coll. N. Coleman, Smith-

Mclntyre grab. 1 9 (J56375), paratype, Stn MSL-EG 29, Eastern Bass

Strait, 10.4 km ESE of eastern edge of Lake Tyers, 37°52.52'S

148°12.55'E, 38 m depth, 25 September 1990, coll. Marine Sciences

Laboratory. Smith-Mclntyre grab.

Description of female (Fig. 146A, B). Body slender, holotype

1.45 mm long, nearly nine times as long as wide. Cephalothorax

subrectangular, slightly narrower anteriorly with slight

triangular rostrum, as long as wide, about as long as pereonites

1 and 2 together, naked, eyes absent. Pereonites 1 and 6 shortest,

0.43 times as long as cephalothorax; pereonites 2 to 5 subequal,

pereonite 2 slightly longest, 1.4 times as long as cephalothorax,

(all pereonites respectively 1.6,1.2,1.3,1.3,1.3 and 1.6 times as

wide as long). Pleon with five free subequal cylindrical

pleonites without pleopods; each pleonite 2.7 times as wide as

long. Pleotelson pentangular, one-third length of pleon and as

wide as long, with blunt distal apex.

Antennule (Fig. 147A) of four articles, proximal article just

over twice as long as wide, just shorter than distal three articles

together, with simple distal seta; second article about 1.3 times

as long as third article, with one simple and two penicillate

outer distal setae; third article with two simple inner distal

setae; fourth article 1.4 times as long as third, with five simple

and two penicillate distal setae.

Antenna (Fig. 147B) of six articles, proximal article

compact, naked, fused to cephalothorax; second article stout,

as long as wide, with dorsodistal seta; third longer than wide,

with fine dorsodistal seta; fourth article longest, three times as

long as third article, 5.4 times as long as wide, curved, with

two simple and four penicillate distal setae; fifth article 0.3

times as long as fourth with one distal seta; sixth article minute

with four distal setae.

Labrum not recovered. Left mandible (Fig. 147C) with

subtriangular, crenulate pars incisiva and linguiform, crenulate

lacinia mobilis, right mandible (Fig. 147D) without lacinia

mobilis; pars molaris of both mandibles tapering abruptly with

fine tooth-like protrusions around distal margin. Labium (Fig.

147F) simple, distally slender, naked. Maxillule (Fig. 147E)

with nine finely-denticulate distal spines, palp not recovered.

Maxilla not recovered. Maxilliped (Fig. 147G) palp first and

second articles with microtrichia on outer margin, second

article with two shorter and one stout longer inner setae; third

article with two inner setae in distal half of article; fourth with

four inner to distal setae, and one outer subdistal seta; bases

fused, naked; endites distally with outer microtrichia and

inner rounded tubercle. Epignath not recovered.

Cheliped (Fig. 147H) basis twice as long as wide, naked,

posterior lobe small, sclerite large; merus with one ventral

seta; carpus rounded, 1.2 times as long as wide, wider

proximally, with one dorsodistal and two mid-ventral setae;

propodus elongate, 1.4 times as long as wide, with two ventral

setae, one inner and one outer mid-distal setae adjacent to

dactylus articulation; fixed finger with three setae alongside

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

219

Fig. 147. Parafilitanais vadosus sp. nov., female paratype. A, antennule; B, antenna; C, left mandible, distal; D, right mandible; E, maxillule

endite; F, labium; G, maxilliped; H, cheliped with H', detail of tips of chela fingers. Scale = 0.1 mm.

220

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig. 148. Parafilitanais vadosus sp. nov., female paratype. A to F, pereopods 1 to 6 respectively; G, pleopod; H, uropod. Scale = 0.1 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

221

cutting edge, distal spine with two adjacent tooth-like tubercles

(Fig. 147H'); dactylus naked, shorter than fixed finger.

Pereopod 1 (Fig. 148A) coxa with simple seta; basis four

times as long as wide, naked; ischium compact with one

ventral seta; merus, carpus and propodus subequal in length,

merus with one slender ventrodistal spine, carpus with three

fine distal setae, propodus with subdistal dorsal seta,

dorsodistal spine-like apophysis, distal microtrichia and distal

spine with denticulations in distal half; dactylus shorter than

unguis, both together two-thirds as long as propodus.

Pereopods 2 (Fig. 148B) similar to pereopod 1, but basis with

two dorsal penicillate setae, merus just longer than carpus;

pereopod 3 (Fig. 148C) similar to pereopod 2, basis with one

penicillate seta.

Pereopod 4 (Fig. 148D) coxa with seta, basis four times as

long as wide with two penicillate setae; ischium with one ventral

seta; merus just shorter than carpus, with two ventrodistal

distally-denticulate spines; carpus with two fine dorsodistal

setae, and one dorsodistal and two ventrodistal short, curved

spines; propodus 1.3 times as long as carpus, with dorsodistal

seta and spine-like apophysis, and two ventrodistal distally-

denticulate spines; dactylus half as long as laterally-denticulate

unguis, both together 1.2 times as long as propodus. Pereopod 5

(Fig. 148E) as pereopod 4, but carpus with four distal spines and

one dorsodistal seta; pereopod 6 (Fig. 148F) as pereopod 5, but

ischium with two setae, unguis as long as dactylus.

Pleopods absent.

Uropod (Fig. 148H) uniramous, basis 1.5 times as long as

wide, with one shorter and one longer outer distal (“exopodal”)

setae, longer seta exceeding distal tip of endopod; proximal

segment of endopod about twice as long as basis, 2.5 times as

long as wide, with one simple and two penicillate distal setae;

distal segment half as long as proximal segment, with one

subdistal and five distal setae.

Male similar to female, but with pleopods: pleopod (Fig.

148G) basis naked, endopod half as long as exopod, rami with

four and seven distal setae respectively.

Etymology. From the Latin vadosus - shallow, this species

being by far the shallowest recorded for the genus.

Remarks. The morphology of this species accords entirely with

the diagnosis of the genus Parafilitanais as given by Larsen

(2002), including the proportions of the cheliped and the

pereopods, and the structure of the uropod, but excepting his

statement that the fourth to sixth pereopod are without a coxa:

coxae are present and unfused in this genus, as shown by the

present specimen and by Larsen’s (loc. cit.) figures 1A and 1L.

Of the three previously-described species of the genus (see

Larsen, 2005), Parafilitanais vadosus sp. nov. has a

proportionately shorter pereonite 1 than does P. caudatus

Kudinova-Pasternak, 1989, a proportionately longer pleon than

does P. similis Kudinova-Pasternak, 1990, and is without the

distally-flattened pleotelson and the extended cheliped dactylus

of P. mexicana Larsen, 2002. In addition, the present species is

distinct from all of the other three in having the distal segment

of the uropod endopod about half as long as the proximal

segment (>0.7 times in the others), and in the proportionately

shorter dactylus and unguis of the anterior pereopods.

Parafilitanais vadosus was taken at 29 to 38 m depth in the

Eastern Bass Strait. The previous species were all recorded in

deep waters, P. caudatus at 3660 m in the Indian Ocean,

P. similis at 750 m in the Pacific Ocean, and P. mexicana at

625-2030 m in the Gulf of Mexico.

Genus Bascestus gen. nov.

Diagnosis. Female of leptognathioid ( sensu lato ) facies,

elongate. Cephalothorax longer than wide, eyelobes and eyes

absent. Pleonite epimera each with long seta. Antennule four-

articled; antenna six-articled, fourth article longest, curved,

without secondary articulation. Mandibular molar proximally

broad, tapering, distally with numerous fine teeth; right

mandible incisor not crenulate, but distally concave with

adjacent stout teeth; left incisor with few distal crenulations,

lacinia mobilis narrow, linguiform. Maxilliped bases fused,

naked, endites not fused, distally with single rounded tubercle

and small inner seta. Cheliped attached by distally rectangular

sclerite, basis narrower posterior to this attachment; propodus

with two ventral setae. Pereopods 1 to 3 with one seta on

ischium; pereopods 2 to 6 with compound (denticulate in their

distal half) spines on merus, carpus and propodus; pereopods 2

and 3 merus ventrodistally with one seta and one compound

spine, pereopods 4 to 6 carpus with three spines and one seta.

Ungues of anterior pereopods longer than dactyli; ungues of

posterior pereopods distinct, shorter than dactyli, both articles

with fine ventral denticulation. Pleopods present, functional;

rami rounded, with plumose setae extending along outer

margin. Uropod biramous, basis without distal apophyses, rami

slender, endopod and exopod of two segments.

Male much less elongate than female, antennule of five

articles; other appendages similar to those of female,

mouthparts functional.

Etymology. From “Bass” as in Bass Strait, and the Greek (but

with Latin pronunciation) cestus - a reinforced boxing glove,

alluding to the nodulose chela of at least the type-species of

this genus (male).

Type species: Bascestus melmackenziae sp. nov.

Remarks. The species described below shows morphological

affinities with such genera as Leptognathiella Hansen, 1913,

Leptognathioides Bird & Holdich, 1984, Stenotanais Bird &

Holdich, 1984, and Kanikipa Bird, 2011, without conforming to

any single one of these. Thus the mandibular structure,

particularly the right incisor and the tapering molar, resemble

those of the first three of these three genera, the last two of

which also show some ventrodistal carpal expansion to

accommodate the reflexion of the chela; the linguiform lacinia

mobilis of the left mandible is also consistent with most species

of these four genera, while the molar dentition resembles that of

Leptognathiella and Stenotanais. The presence on the merus of

the anterior pereopods of a ventrodistal seta and a ventrodistal

spine is consistent with Leptognathioides and Kanikipa (but

also the nototanaid genus Tanaissus and the agathotanaid genus

Paragathotanais ), while denticulation of the posterior dactyli

accords with Stenotanais and Akanthophoreus inter alia (the

latter also having a similar mandibular molar morphology).

222

M. Blazewicz-Paszkowycz & R.N. Bamber

Thus, there is no particular characteristic of Bascestus

gen. nov. that distinguishes it from a number of others once

allocated to the Leptognathiidae sensu Sieg, 1976b, but their

combination in the present taxon is unique and excludes it

from any of those genera. In having sexual dimorphism

possibly limited to the antennular structure and habitus

proportions, the present genus is further distinct from many

other “leptognathiid” genera (in a number of which males

remain unknown).

Since the insightful work of Bird and Holdich (1984),

which highlighted the distinctions of a number of these genera

previously subsumed into Leptognathia Sars, 1882, and

sensibly discussed the relative merits of different characters

for distinguishing different taxon levels, these taxa have been

moved variously into such families as the Anarthruridae Lang,

1971, thence to the Colletteidae ( Leptognathiella and

Leptognathioides ), the Akanthophoreidae ( Akanthophoreus )

or currently incertae sedis sensu Larsen and Wilson (2002)

0 Stenotanais , Kanikipa ). Owing to its apparent affinities as

described above, we choose at present to place Bascestus in

the Colletteidae sensu Larsen and Wilson (2002).

Bascestus melmackenziae sp. nov.

Figures 149-151

Material. 1 9 (J23600), holotype. Eastern Bass Strait, 3.2 km S of

Cape Conran, Stn MSL-EG 55, 37°50.38'S 148°43.28'E, 49 m depth,

sand and shell, 28 September 1990, coll. Marine Science Laboratories,

Smith-Mclntyre Grab. 2 99 (J56380), paratypes. Eastern Bass Strait,

13.3 km E of eastern edge of Lake Tyers, Stn MSL-EG 68, 37°51.42'S

148°14.36'E, 37 m depth, 04 June 1991, coll. N. Coleman, Smith-

Mclntyre Grab. 1 S (J28484), allotype. Eastern Bass Strait, 13.3 km E

of eastern edge of Lake Tyers, Stn MSL-EG 69,37°51.42'S 148°14.36'E,

37 m depth, 04 June 1991, coll. N. Coleman, Smith-Mclntyre Grab.

Description of female. Body (Fig. 149C) slender, 12 times as

long as wide, holotype 2 mm long. Cephalothorax parallel-sided

in posterior half, narrowing anteriorly, 1.8times as long as wide,

with slight rounded rostrum; eyelobes and eyes absent.

Pereonites all rectangular, pereonitel half as long as

cephalothorax, as long as wide; pereonites 2 to 5 subequal in

length, 1.4 times as long as pereonite 1, longer than wide;

pereonite 6 shortest, 0.9 times as long as pereonite 1 (all

pereonites respectively 1.0, 0.7, 0.8, 0.8, 0.7 and 1.1 times as

wide as long). Pleon over three as long as pereonite 6; pleonites

subequal in length, 2.8 times as wide as long, all bearing

pleopods and single mid-lateral setae; pleotelson subpentangular,

longer than two preceding pleonites, as wide as long, with single

midlateral setae and two pairs of distal setae.

Antennule (Fig. 150A) shorter than cephalothorax, four-

articled; article 1 nearly three times as long as wide, 0.9 times

as long as articles 2 to 4 combined, with subdistal outer pair of

penicillate setae, and single outer simple distal seta; article 2

slightly overlapping proximal part of article 3, twice as long as

wide, less than half as long as article 1, with single inner and

outer simple subdistal setae and tuft of five dorsodistal setae;

article 3 slightly overlapping proximal part of article 4, 1.7

times as long as wide, 0.6 times as long as article 2, with single

inner and outer simple subdistal setae; article 4 as long as

article 2, five times as long as wide, with three simple and two

penicillate subdistal setae and two longer simple distal setae.

Antenna (Fig. 150C), six-articled, article 1 short and

annular, naked; article 2 about as long as wide, distally with

single shorter dorsal and longer lateral setae; article 3 just

shorter than article 2, with dorsodistal seta; article 4 twice as

long as articles 2 and 3 combined, 4.5 times as long as wide,

curved, distally with two simple and three penicillate setae;

article 5 one-third as long as article 4, with long distal seta;

article 6 very small, with five distal setae.

Labrum (Fig. 150D) rounded, hood-shaped, distally setose.

Mandibles stout; left mandible (Fig. 150E) with distally-crenulate

incisor and narrow, simple lacinia mobilis, molar proximally

stout, tapering, tip with six or seven slender spine-like teeth; right

mandible (Fig. 150F) incisor distally concave with few stout

teeth; molar as on left mandible. Labium (Fig. 150H) simple,

each side with single subdistal spinule. Maxillule (Fig. 150G)

endite slightly sigmoid, with one distal seta and five terminal and

two subterminal spines; palp and maxilla not recovered.

Maxilliped (Fig. 1501) bases rounded, fused, naked; palp article

1 naked, article 2 with two inner distal setae and one outer distal

seta; article 3 with three stout inner setae; article 4 with five

distal and one outer subdistal setae; endites not fused, distally

with rounded tubercle and inner seta. Epignath not recovered.

Cheliped (Fig. 149D, 151A) basis in situ proximally well-

anterior of anterior margin of pereonite 1 ventrally, attached by

distally rectangular sclerite; basis narrower posterior to this

attachment, with rounded posterior free margin, twice as long as

wide, with small dorso-subdistal seta; merus subtriangular, with

ventral seta; carpus twice as long as wide, with three

dorsoproximal and two mid-ventral setae, ventrodistally with

slight expansion of margins into which chela can reflect;

propodus longer than wide, with two ventral setae and comb-row

of four simple setae, dorsal margin and mid-lateral face with

rows of rounded nodules in distal half, and with inner ventral

submarginal crenulate ridge; fixed finger cutting edge with three

proximal denticulations and three adjacent setae; dactylus with

several dorsal nodules and one small proximal seta.

Pereopod 1 (Fig. 151B) basis relatively stout, 2.5 times as

long as wide, with one mid-dorsal simple seta; ischium with

one seta about half as long as merus; merus distally expanded,

0.4 times as long as basis, 1.4 times as long as distal width,

ventrodistally with one shorter seta and one compound spine

almost as long as carpus; carpus 1.1 times as long as merus,

rectangular, with two short ventrodistal spinules and inner and

outer compound distal spines; propodus 1.5 times as long as

carpus, with one compound ventrodistal spine, one distal seta

as long as dactylus, and dorsodistal spinulation; dactylus half

as long as unguis, both together as long as propodus. Pereopod

2 as pereopod 3. Pereopod 3 (Fig. 151C) similar to pereopod 1,

but basis with mid-dorsal penicillate seta, carpus with single

ventral, inner and outer compound distal spines.

Pereopod 4 (Fig. 151D) basis 2.4 times as long as wide;

ischium with two setae; merus 0.3 times as long as basis, with

two ventrodistal compound spines; carpus 1.3 times as long as

merus, with with single ventral, inner and outer compound

distal spines and simple dorsodistal seta; propodus 1.1 times

as long as carpus, with one dorsodistal seta and two ventrodistal

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

223

Fig 149. Bascestus melmackenziae sp. nov. A, male, lateral; B, male, dorsal; C, holotype female, lateral; D, cephalothorax, ventral, showing

attachment of chelipeds. Scale = 0.1 mm.

224

M. Biazewicz-Paszkowycz & R.N. Bamber

Fig 150. Bascestus melmackenziae sp. nov. A, female antennule; B, male antennule; C, antenna; D, labrum; E, left mandible; F, right mandible;

G, maxillule endite; H, labium; I, maxilliped. Scale = 0.01 mm.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

225

Fig 151. Bascestus melmackenziae sp. nov. A, cheliped ; B, pereopod 1 ; C, pereopod 3; D, pereopod 4 ; E, pereopod 5 ; F, pereopod 6 ; G, pleopod ;

H, uropod. Scale = 0.01 mm.

226

M. Blazewicz-Paszkowycz & R.N. Bamber

compound spines; dactylus semicircular in cross-section,

ventral edges finely denticulate; unguis half as long as dactylus,

ventrally finely denticulate, dactylus and unguis together 1.2

times as long as propodus. Pereopod 5 (Fig. 151E) similar to

pereopod 4, propodus with microtrichia. Pereopod 6 (Fig.

151F) similar to pereopods 4 and 5 but basis with dorsal

undulation, propodus with two dorsodistal setae.

Pleopods all similar (Fig. 151G), basal article naked;

endopod and exopod rami rounded, all setae plumose; endopod

with eight outer to distal setae and one inner subdistal seta,

proximal outer margin naked; exopod with eight outer to distal

setae and separated outer proximal seta.

Uropod (Fig. 151H) slender, nearly twice as long as

pleotelson; basal article twice as long as wide, naked, without

distal apophyses; exopod two-segmented, shorter than

proximal article of endopod, segments subequal in length,

with one distal seta on proximal segment, two unequal distal

setae on distal segment; endopod two-segmented, proximal

segment shorter and with one penicillate distal seta, distal

article with one subdistal and two distal simple setae and two

distal penicillate setae.

Distinctions of male. Body (Fig. 149A, B) smaller and less

elongate than that of female, length 0.9 mm, 10 times as long

as wide. Cephalothorax similar to that of female, 1.7 times as

long as wide, tapering anteriorly. Pereonites more or less

rectangular; pereonite 1 half as long as cephalothorax;

pereonites 2 to 5 subequal in length although progressively

shorter, about 1.2 times as long as pereonite 1, not longer than

wide; pereonite 6 shortest, 0.9 times as long as pereonite 1;

pereonites respectively 1.3, 1.0, 1.1, 1.0, 1.1 and 1.4 times as

wide as long. Pleon as long as pereonites 4 to 6 inclusive,

pleonites 2.8 times as wide as long.

Antennule (Fig. 150B) five-articled, shorter than

cephalothorax; peduncle article 1 stout, three times as long as

wide, naked; peduncle article 2 compact, 0.4 times as long as

article 1, 1.3 times as long as wide, with three penicillate and

one simple ventrodistal setae; article 3 as long as wide, 0.6

times as long as article 2, with two dorsodistal simple setae;

flagellum of two segments, proximal segment just longer than

wide, 0.75 times as long as peduncle article 3, naked; second

segment twice as long as wide, as long as peduncle article 3,

with four simple and one penicillate subdistal setae and two

simple distal setae.

Mouthparts, chelipeds, pereopods, pleopods and uropods

similar to those of female.

Etymology. Named after Melanie Mackenzie of the Museum

Victoria in gratitude for her diligent and uncomplaining

assistance with the material and data of the collections.

Remarks. The characters of this species are discussed above

under the generic remarks. The rugose tuberculation of the

cheliped may be a specific rather than generic character as it is

known to show intrageneric variation in other paratanaoid

taxa (see discusison under Araphura ). Bascestus

melmackenziae sp. nov. was taken from depths of 37 to 49 m

in the Eastern Bass Strait.

Tanaidomorpha incertae sedis

Species 33

Figure 152

Material examined. 6 mancae (manca 1) (J51791), Stn VC30 Cl,

Victoria, Central Bass Strait, 38°35.53'S, 146°07.51'E, 40 m depth, 11 th

May 1998, Smith-Mclntyre grab, coll. N. Coleman.

Description. Body (Fig. 152) glabrous, small, eight times as

long as wide, length of all specimens 1.4 mm. Cephalothorax

subrectangular, wider than long, with branchial chambers

expanded as lateral rounded flanges over cheliped attachment;

rostrum straight, smooth, with anterior margin of carapace

protruding either side of rostrum as rounded lobes. Pereonite 1

trapezoidal, 0.8 times as long as cephalothorax, anteriorly 1.4

times as wide as pereonite length, posteriorly 0.8 times as wide

as pereonite length; pereonites 2 and 3 rectangular, 0.75 times

as wide as long, pereonite 2 almost as long as cephalothorax,

pereonite 3 just shorter than pereonite 1; pereonites 4 and 5

hexagonal, pereonite 4 as wide as long and as long as pereonite

3, pereonite 5 about 1.5 times as wide as long and about half

length of cephalothorax; pereonite 6 short, one-quarter as long

as pereonite 5 and four times as wide as long. Pleon laterally

convex, first four pleonites 5.2 times as wide as long, fifth

pleonite as long as pereonite 6 and four times as wide as long.

Pleotelson semicircular, twice as long as each anterior pleonite

and twice as wide as long.

Antennule apparently of three articles. Posterior pereopods

with prickly tubercles on carpus. Uropods biramous, each

ramus of one segment.

Remarks. These specimens are all at the manca 1 stage, with a

highly reduced pereonite 6, hardly longer than pleonite 1, and no

pereopod 6. Their morphology shows no similarity to any known

genus; in particular, the lobate structure of the cephalothorax is

highly unusual, yet appears unlikely to be a juvenile character.

The presence of prickly tubercles on the posterior pereopods

suggests that this taxon may belong in the Typhlotanaidae (but

see Bassoleptochelia. above). Without knowledge of the adult, it

is at present not possible to assign this taxon to any category

lower than Suborder, and it may well represent a distinct Family.

“Species 33” is a Museum Victoria collection reference number.

Discussion

While the Bass Strait has been the subject of unusually extensive

and intensive sampling, from which the collections analyzed

here have arisen, it remains remarkable that we now know of 65

species in 43 genera from this area (Appendix 2), of which 57

species and eight genera have proven to be new to science (and at

present are all endemic except for Remexudes toompani). The

extraordinarily high diversity of this assemblage is the more

remarkable because the habitats are not very diverse, being

predominantly sandy substrata and thus with limited niche

diversity. That the Bass Strait is not unique in this respect

(although Australia may be) is demonstrated by the comparatively

similarly high levels of tanaidacean diversity discovered from

much briefer surveys elsewhere in Australia, such as Esperance

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Fig. 152. Tanaidomorpha incertae sedis Species 33, manca. A, dorsal view ; B, lateral view. Scale line = 0.1 mm.

228

M. Blazewicz-Paszkowycz & R.N. Bamber

(Bamber, 2005) and Moreton Bay (Bamber, 2008), again both

areas with predominantly sandy substrata.

Of those nine species from the Bass Strait known from

elsewhere in Australia, two, Indoapseudes macabre and

Paratanais vetinari, are also known from southwestern

Australia, somewhat consistent with the predominant current

flow coming from the west. All but one of the remainder are also

found in the adjacent territory of New South Wales, and two

extend as far as Brisbane (Queensland).

At the generic level, those described as new for the Bass

Strait are currently endemic (other than Remexudes). Whiteleggia

and Pseudowhiteleggia, both with a fairly long history in the

literature, and Remexudes are so far restricted to the southeastern

corner of Australia. The typhlotanaid genus Antiplotanais now

contains three species, all from Australia, from southern Western

Australia, Victoria and southern Queensland. The at-present

more-numerous genus Bathytanais is almost exclusively

Australian, being represented by seven species in Australian

waters (from all coasts) but also, enigmatically, one species in

Hong Kong.

On a somewhat more widespread basis, the genera

Spinosapseudes and Metapseudes are now both known from

two species only, with some zoogeographic consistency, one of

each genus being from the Bass Strait and the other from New

Zealand waters. A similar distributional association appears to

be shown within Araphura (and Araphuroidesl ): while overall

these genera are relatively widespread in their distribution, the

morphology of the species from the Bass Strait, with nodulose

chelipeds, is also found only in species from New Zealand and

the sub-Antarctic South Shetland Islands. A similar Antarctic/

sub-Antarctic link is to be found in the Mirandotanaidae.

The two pagurapseudid genera recorded above, both well-

represented in the eastern half of Australia, show diversity and

similar distributions throughout Australasia to Micronesia and

also with one ( Macrolabrum ) or five ( Pagurapseudes ) species in

the western Indian Ocean. A similar distribution is shown by the

four known species of Indoapseudes , and by the genus

Pakistanapseudes, five species of which have now been found in

the Bass Strait, which contributes to a surprisingly high diversity

in Australia, (belying the derivation of the generic name) with

some nine of the 18 species recorded globally in the

Pakistanapseudinae coming from Australian waters.

There are thus some consistent trends within the zoogeography

of the Bass Strait taxa, at both regional and wider scales.

However, the origins of this Australian diversity are not

easy to explain. A process of original evolution and subsequent

radiation is occasionally suggested as a source of high

diversity. An early stage in such a process may be exhibited by

the presently Australian-endemic family Whiteleggiidae. Yet

speculation over associations with the tanaidacean faunas of

other parts of what was Gondwana must await better study of

the faunas of South Africa and India, for examples.

Interestingly, there appears little association with the

tanaidacean fauna of Antarctica, where such families as the

Leptocheliidae appear to be absent. The converse hypothesis

of colonization and subsequent allopatric speciation and

radiation is equally well-supported by the distributions of such

taxa as the Pakistanapseudinae and the Pagurapseudidae with

their links through Australasia and the Indian Ocean together

with an apparent greatest diversity in Australia itself.

It remains the case that such hypotheses will remain

speculative until more is known about the tanaidacean faunas

from other, as yet understudied (or at least unpublished) regions

of Australia, such as the tropical waters of northern Queensland,

the Gulf of Carpentaria and the north coast of Australia, and

northern Western Australia, with their extensive coral-reef

habitats, as well as the Great Australian Bight in the south.

Fortunately, information on the tanaidaceans of neighbouring

regions is, in some cases, improving markedly, with valuable

recent work from New Zealand waters by Bird (2008; 2011) and

some from Micronesia (Bamber, 2009 and references therein).

Acknowledgements

We are very grateful to Museum Victoria, and to (alphabetically)

Melanie Mackenzie, Gary Poore, David Staples, Jo Taylor and

Robin Wilson of that museum, for access to, discussions about,

assistance with, and of course collection of the material, as well

as tolerance of our endeavours at the Museum and extraordinary

hospitality. We are also indebted to Graham Bird for numerous

valuable discussions on these taxa, and to the comprehensive

and competent suggestions from the anonymous reviewer. The

research was financed by EU Marie Curie Grant, OIF

040613-DiPoT and grant MNiSW7984/B/P01/2011/40.

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Tanaidae)

Appendix 1. Sample registration numbers for Kalliapseudes

obtusifrons material examined.

Crib Point, Western Port, Victoria

J48195 8 $ (2 brooding), 63,1 juvenile

J48207 15 9 (6 brooding), 4 3, 2 juveniles

J48219 5 9(1 brooding, 2 with oostegites), 1 3

J48228 6 9(1 brooding), 2 juveniles

J48375 1 9 with oostegites, 1 3

J48383 1 3

J48388 7 9 (4 brooding), 2 3, 16 juveniles

J48389 5 brooding 9, 10 juveniles

J48393 3 9 (2 brooding), 2 <3, 1 juvenile

J48398 2 brooding 9, 1 <?, 5 juveniles

J48399 2 brooding 9, 3 juveniles

J48402 49 (1 brooding), 2 juveniles

J48409 1 brooding 9, 2 juveniles

J48416 4 9 (2 brooding, 1 with oostegites), 4 3, 2 juveniles

J48418 1 9, 1 3

Hastings, Western Port, Victoria

J66425 79 9 (24 brooding, 28 with oostegites), 41 <3,21 juveniles

J66520 1 brooding 9

J66665 9 9, 2 <3, 2 juveniles

J66918 19

J67235 3 9, 1 6

J67281 1 9, 1 3

Port Phillip Bay, Victoria

J43615 19

J43616 1 brooding 9, 1 juvenile

J43617 1 9

J43618 1 brooding 9, 1 3

J43619 1 9,1 d

J43620 1 brooding 9

J43621 19

J43623 13

J43624 1 9, 1 3

J43662 3 9, 2 3, 13 juveniles

J43076 13

J43080 1 9, 1 6

J43081 \3

Gippsland, E. Bass Strait

J28079 9 9 (3 with oostegites), 5 3, 7 juveniles

J28114 18 9 (4 brooding, 7 with oostegites), 6 3

J28102 8 9 (2 brooding), 6 3, 2 juveniles

J28110 28 9 (4 with oostegites), 3 3

J28098 1 9 with oostegites

East of Wilson’s Promontory

J50781 2 9(1 brooding), 1 3

J50782 2 juveniles

J50783 1 9, 4 juveniles

J50784 1 9, 1 juvenile

J50785 44 9 (6 brooding), 9 3, 6 juveniles

J50786 3 9, 2 3

J50787 9 9 (2 brooding, 2 with oostegites), 4 3, 3 juveniles

J50788 2 9(1 brooding), 1 3

J50789 9 9, 4 3, 1 juvenile

J50790 13 9, 3 juveniles

J50791 3 9

234

M. Biazewicz-Paszkowycz & R.N. Bamber

Appendix 2. Full species list of Tanaidacea from Bass

Strait, with provenance elsewhere in Australia, if any.

ORDER TANAIDACEA DANA, 1849

Suborder Apseudomorpha Sieg, 1980(a)

Superfamily Apseudoidea Leach 1814

Family Apseudidae Leach 1814

Subfamily Apseudinae Leach 1814

Genus Apseudes Leach 1814

Apseudes abditospina (Biazewicz-Paszkowycz & Bamber,

2007) comb. nov.

Apseudes poorei Biazewicz-Paszkowycz & Bamber, 2007

Apseudes quasimodo sp. nov.

Genus Apseudopsis Norman 1899

Apseudopsis tuski (Biazewicz-Paszkowycz & Bamber, 2007)

comb. nov.

Genus Spinosapseudes Gufu 1996(a)

Spinosapseudes colobus Biazewicz-Paszkowycz & Bamber,

2007

Genus Bunakenia Gufu 1995(a)

Bunakenia labanticheiros sp. nov.

Genus Paradoxapseudes Gufu 1991

Paradoxapseudes paneacis sp. nov.

Paradoxapseudes attenuata sp. nov.

Subfamily Pugiodactylinae Gufu 1995(b)

Genus Pugiodactylus Gufu 1995(b)

Pugiodactylus syntomos Biazewicz-Paszkowycz & Bamber,

2007

Family Whiteleggiidae Gutu 1972

Genus Whiteleggia Lang 1970

Whiteleggia multicarinata (Whitelegge, 1901) [also New

South Wales 1 ]

Genus Pseudowliiteleggia Lang, 1970

Pseudowhiteleggia typica Lang, 1970 [also New South

Wales 1 ]

Family Kalliapseudidae Lang 1956(b)

Subfamily Kalliapseudinae Gufu 1972

Genus Kalliapseudes Stebbing 1910

Kalliapseudes obtusifrons (Haswell 1882) [also New South

Wales 2 ]

Family Metapseudidae Lang 1970

Subfamily Metapseudinae Lang 1970

Genus Cyclopoapseudes Menzies, 1953

Subgenus Exopoapseudes subgen. nov.

Cyclopoapseudes (Exopoapseudes) plumosa sp. nov.

Genus Labraxeudes Biazewicz-Paszkowycz & Bamber, 2007(b)

Labraxeudes heliodiscus Biazewicz-Paszkowycz & Bamber,

2007

Genus Metapseudes Stephensen 1927

Metapseudes wilsoni Biazewicz-Paszkowycz & Bamber,

2007

Family Parapseudidae Gufu 1981

Subfamily Pakistanapseudinae Gufu 2008 new rank

Genus Pakistanapseudes Bacescu 1978

Pakistanapseudes bassi Biazewicz-Paszkowycz & Bamber,

2007

Pakistanapseudes lucifer sp. nov.

Pakistanapseudes perulpa Biazewicz-Paszkowycz &

Bamber, 2007 [also Queensland 7 ]

Pakistanapseudes taylorae sp. nov.

Pakistanapseudes sp. nov. C

Subfamily Parapseudinae Gufu 1981 new rank

Genus Parapseudes Sars, 1882

Parapseudes blandowskii sp. nov.

Genus Saltipedis Gufu 1995(b)

Saltipedis nugoris Biazewicz-Paszkowycz & Bamber, 2007

Saltipedis floccus sp. nov.

Genus Remexudes Biazewicz-Paszkowycz & Bamber, 2007

Remexudes toompani Biazewicz-Paszkowycz & Bamber,

2007 [also Queensland 7 ]

Family Pagurapseudidae Lang 1970

Subfamily Hodometricinae Gufu 1981

Genus Indoapseudes Bacescu 1976(a)

Indoapseudes macabre Bamber, 2005 [also south-western

Australia 3 ]

Genus Similipedia Gufu 1989

Similipedia diarris Biazewicz-Paszkowycz & Bamber, 2007

Subfamily Pagurapseudinae Lang, 1970

Genus Pagurapseudes Whitelegge 1901

Pagurapseudes victoriae sp. nov.

Pagurapseudes kimbla sp. nov.

Genus Macrolabrum Bacescu, 1976(b)

Macrolabrum tangaroa sp. nov.

Macrolabrum sarda sp. nov.

Macrolabrum haikung sp. nov.

Suborder Tanaidomorpha Sieg, 1980(a)

Superfamily Paratanaoidea Lang, 1949

Family Paratanaidae Lang, 1949

Subfamily Paratanaidinae Lang, 1949

Genus Paratanais Dana, 1952

Paratanais malignus Larsen, 2001 [also New South Wales 4 ]

Paratanais tanyherpes sp. nov.

Paratanais vetinari Bamber, 2005 [also south-western

Australia 3 ]

Subfamily Bathytanaidinae Larsen & Heard, 2001

Genus Bathytanais Beddard, 1886

Bathytanais bathybrotes (Beddard, 1886(a)) [also south¬

western Australia 5 and Queensland 6 ]

Bathytanais fragilis Larsen & Heard, 2001

Bathytanais parageios sp. nov.

The Shallow-water Tanaidacea (Arthropoda: Malacostraca: Peracarida) of the Bass Strait, Victoria, Australia (other than the

Tanaidae)

Family Leptocheliidae Lang, 1973

Genus Leptochelia Dana, 1849

Leptochelia billambi sp. nov.

Genus Araleptochelia gen. nov.

Araleptochelia macrostonyx sp. nov.

Genus Pseudoleptochelia Lang, 1973

Pseudoleptochelia occiporta sp. nov.

Genus Bassoleptochelia gen. nov.

Bassoleptochelia verro sp. nov.

Family Tanaellidae Larsen & Wilson, 2002

Genus Araphura Bird & Holdich, 1984

Araphura pygmothymos sp. nov.

Araphura yarra sp. nov.

Araphura doutagalla sp. nov.

Genus Araphuroides Sieg, 1886

Araphuroides stabastris sp. nov.

Araphuroides batmania sp. nov.

Araphuroides sala sp. nov.

Genus Inconnivus gen. nov.

Inconnivus billibunteri sp. nov.

Family Mirandotanaidae Blazewicz-Paszkowycz & Bamber,

2009

Genus Pooreotanais Blazewicz-Paszkowycz & Bamber, 2009

Pooreotanais gari Blazewicz-Paszkowycz & Bamber, 2009

Family Typhlotanaidae Sieg, 1984

Genus Typhlotanais Sars, 1882 sensu lato

Typhlotanais herthio sp. nov.

Genus Antiplotanais Bamber, 2008

Antiplotanais actuarius sp. nov.

Genus Hamatipeda Blazewicz-Paszkowycz, 2007

Hamatipeda sima sp. nov.

Genus Paratyphlotanais Kudinova-Pastemak & Pasternak,

1978

Paratyphlotanais colouros sp. nov.

Genus Peraeospinosus Sieg, 1986

Peraeospinosus tanytrix sp. nov.

Genus Meromonakantha Sieg, 1986

Meromonakantha anarsios sp. nov.

Family Nototanaidae Sieg 1976

Subfamily Nototanainae Sieg 1976

Genus Tanaissus Norman & Scott, 1906

Tanaissus giraffa sp. nov.

Genus Protanaissus Sieg, 1983

Protanaissus huberti sp. nov.

Family Agathotanaidae Lang 1971

Genus Paragathotanais Lang, 1971

Paragathotanais wurundjeri sp. nov.

Genus Ozagathus gen. nov.

Ozagathus watharongus sp. nov.

Family Akanthophoreidae Sieg, 1986

Genus Gejavis gen. nov.

Gejavis corsotos sp. nov.

Family Tanaopsidae fam. nov.

Genus Tanaopsis Sars, 1896

Tanaopsis boonwurrungi sp. nov.

Tanaopsis oios sp. nov.

Family Colletteidae Larsen & Wilson, 2002

Genus Parafilitanais Kudinova-Pastemak, 1989

ParafUitanais vadosus sp. nov.

Genus Bascestus gen. nov.

Bascestus melmackenziae sp. nov.

Family incertae sedis

“Species 33”

1 Lang, 1970

2 Drumm & Heard, 2006

3 Bamber, 2005

4 Larsen, 2001

5 Lang, 1972

6 Bamber, 2008

7 Blazewicz-Paszkowycz & Bamber, 2007

Memoirs of Museum Victoria 69: 237-243 (2012)

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

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

Synidotea poorei , a new isopod from the fouling community in Singapore waters

(Valvifera, Idoteidae)

Yixiong Cai 1 ' 2 and Serena L. M. Teo 3

1 Tropical Marine Science Institute, National University of Singapore, 14 Kent Ridge Road, Singapore 119223, Republic of

Singapore- (caiyixiong@yahi»com)

2 Current address: National Biodiversity Centre, National Parks Board, 1 Cluny Road, Singapore 259569

2 Tropical Marine, Science Institute, National University of Singapore, 14 Kent Ridge Road, Singapore 119223, Republic of

Singapore (tmsteolm@nus. edu. sg)

Abstract Cai, Y. and Tee S.L.M. 2012. Synidotea poorei, a new isopod from the fouling community of Singapore (Valvifera,

Idoteidae). Memoirs of Museum Vicioria&. 237-243.

Synidotea poorei new species (Idoteidae) is described from fouling community on navigational buoys and jetties in

Singapore waters. It belongs to the S. hirtipes species group. The new species is characterized by a smooth uropodal

peduncle, sub-parallel pleotelson lateral margins and a moderately excavate pleotelson posterior margin. It represents a

new record for the genus in Southeast Asia.

Keywords Fouling, Synidotea , new species, marine isopod, Singapore

Introduction

Knowledge of the marine isopod fauna from Southeast Asia is

relatively poor, and that of Singapore is even less so. Ten species,

represent ten genera in eight families have been reported from

Singapore waters. Most of these consist of records within larger

works (e.g. Schioedte andMeinert, 1879; Barnard, 1925; Monod,

1926; Menzies and Barnard, 1951; Stock, 1960; Bruce 1986a). A

new species of wood-borer, Limnoria cristata CooksOn and

Cragg, 1991, was described from intertidal driftwood in a

mangrove forest, and two new species of branchial parasitic

isopods have been described from hermit crabs collected in

Singapore (Williams and Sehuelein, 2005). In 2002, during a

survey of the fouling community on the navigational buoys and

jetties in Singapore coastal waters, four species of marine

isopods, belonging to three families were found in the epifauna

and epiflora on the surface of the navigational buoys. Sphaeroma

walkeri Stubbing, 1905 (Sphaeromatidae) and Cirolana willeyi

Stebbing, 1904 (Cirolanidae) are reported here for the first time

from Singapore. Sphaeroma walkeri, originally described from

Sri Lanka, has been reported tobe a widely distributedintroduced

species found in various fouling communities (Miller, 1968;

Carlton and Iverson, 1981; Mak and Morton, 1985) but it may not

necessarily have been introduced to Singapore. Cirolana willeyi,

also described from Sri Lanka, has been subsequently reported

from estuarine and mangrove habitats from East Africa to

Australia (Bruce, 1981, 1986b). A third species, of the genus

Cilicaea (Sphaeromatidae) was also found. The fourth is a new

species of Synidotea (Idoteidae) described here. Specimens

examined have been deposited in Raffles Museum of Biodiversity

Research, National University of Singapore, Singapore (2fiCj

and Museum Victoria, Melbourne (NMV ). Terminology follows

that of Poore and Lew Ton (1993).

Idoteidae Samouelle, 1819

Synidotea Larger, 1878

Synidotea poorei sp. nov.

Figures 1 4

Material examined, Holotype: male, tl 11.4 mm, ZRC.2005.0118, North

Pandan Buoy, l°f5'48.12" K 103°45T0.81" E, Terumbu Pandan,

Singapore, coll. Y. Cai, S. L. M. Teo, K. S. Tan and T. M, Sin, 11 Apr.

2002. Paratypes: 25males, tl4.6-12.2 mm, I5females, tl 5,6-84 mm,

ZRC.2005.0119, data same as holotype: 2 females, 74 10,2 mm;

4 ovigerous females, tl 69-8.6 mm; 4 males, tl 10,2-12.9 mm, NVM

J62812, data same as holotype. Other specimens: 4 males, tl 6.0-9.1 mm,

2 females, tl 7.0-7.4 mm, ZRC.2005.pi20, North West Sudong Buoy,

l°13 l tf7.22'’ N, IQ3°42’59.I0" E, near Pulau Sudong, Singapore, 19 Jul.

2002; 1 female, tl 8.4 mm, ZRC.2005.0121, Perimbi. Buoy, 1°25'45.11" N,

103°S'14,75" E, East Johor Strait Singapore, coll, Y. Cai, 21 May 2002;

1 male, tl 12.0 mm, ZRG.2005.0122, Retan-D Buoy, 1°17'35.63" N,

103 <1 45'25.48" E, off Sungei Pandan, West Coast, Singapore, coll. Y. Cai,

K. S. Tan* S. C. Lim, 17Oct. 2003; 1 female, tl 5.9mm, ZRC.2Q0S.dl23,

Sirdhana Buoy; 1 < ’14!43.0Q 1 ' N, 103°52'55.02" E, off Marina Bay,

Singapore, coll. Y, Cai, K S. Tan, T. M, Sin & S. L. M Teo, 5 Jun. 2002;

6 males, tl 6.2-10.8 mm, 1 female, tl 7.0 mm, ZRC.2005.0124, Mooring

Buoys at Marina Bay; Singapore, 1°17'0626'' N, :lQ3°Sr20,09 ,r E, coll.

Memoirs of Museum Victoria 69:237-243 (2012)

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

http://museum

Fouling, Synidotea, new species, marine isopod, Singapore

Introduction

Knowledge of the marine isopod fauna from Southeast Asia is

relatively poor, and that of Singapore is even less so. Ten species,

represent ten genera in eight families have been reported from

Singapore waters. Most of these consist of records within larger

works (e.g. Schioedte and Meinert, 1879; Barnard, 1925; Monod,

1926; Menzies and Barnard, 1951; Stock, 1960; Bruce 1986a). A

new species of wood-borer, Limnoria cristata Cookson and

Cragg, 1991, was described from intertidal driftwood in a

mangrove forest, and two new species of branchial parasitic

isopods have been described from hermit crabs collected in

Singapore (Williams and Schuelein, 2005). In 2002, during a

survey of the fouling community on the navigational buoys and

jetties in Singapore coastal waters, four species of marine

isopods, belonging to three families were found in the epifauna

and epiflora on the surface of the navigational buoys. Sphaeroma

walkeri Stebbing, 1905 (Sphaeromatidae) and Cirolana willeyi

Stebbing, 1904 (Cirolanidae) are reported here for the first time

from Singapore. Sphaeroma walkeri, originally described from

Sri Lanka, has been reported to be a widely distributed introduced

species found in various fouling communities (Miller, 1968;

Carlton and Iverson, 1981; Mak and Morton, 1985) but it may not

necessarily have been introduced to Singapore. Cirolana willeyi,

also described from Sri Lanka, has been subsequently reported

from estuarine and mangrove habitats from East Africa to

Australia (Bruce, 1981, 1986b). A third species, of the genus

Cilicaea (Sphaeromatidae) was also found. The fourth is a new

species of Synidotea (Idoteidae) described here. Specimens

examined have been deposited in Raffles Museum of Biodiversity

Research, National University of Singapore, Singapore (ZRC)

and Museum Victoria, Melbourne (NMV). Terminology follows

that of Poore and Lew Ton (1993).

Idoteidae Samouelle, 1819

Synidotea Harger, 1878

Synidotea poorei sp. nov.

Figures 1-4

Material examined. Holotype: male, tl 11.4 mm, ZRC.2005.0118, North

Pandan Buoy, 1°15'48.12 M N, 103°45'10.81" E, Terumbu Pandan,

Singapore, coll. Y. Cai, S. L. M. Teo, K. S. Tan and T. M. Sin, 11 Apr.

2002. Paratypes: 25 males, tl 4.6-12.2 mm, 15 females, tl 5.6-8.1 mm,

ZRC.2005.0119, data same as holotype; 2 females, 7.4, 10.2 mm;

4 ovigerous females, tl 6.9-8.6 mm; 4 males, tl 10.2-12.9 mm, NVM

J62812, data same as holotype. Other specimens: 4 males, tl 6.0-9.1 mm,

2 females, tl 7.0-7.4 mm, ZRC.2005.0120, North West Sudong Buoy,

1°13’07.22" N, 103°42’59.10" E, near Pulau Sudong, Singapore, 19 Jul.

2002; 1 female, tl 8.4mm,ZRC.2005.0121,Perimbi Buoy, 1°25'45.11" N,

103°53’14.75" E, East Johor Strait, Singapore, coll. Y. Cai, 21 May 2002;

1 male, tl 12.0 mm, ZRC.2005.0122, Retan-D Buoy, 1°17’35.63" N,

103°45’25.48" E, off Sungei Pandan, West Coast, Singapore, coll. Y. Cai,

K.S.Tan&S.C.Lim, 17 Oct. 2003; 1 female, tl 5.9 mm, ZRC.2005.0123,

Sirdhana Buoy, 1°14'43.00" N, 103°52'55.02" E, off Marina Bay,

Singapore, coll. Y. Cai, K. S. Tan, T. M. Sin & S. L. M Teo, 5 Jun. 2002;

6 males, tl 6.2-10.8 mm, 1 female, tl 7.0 mm, ZRC.2005.0124, Mooring

Buoys at Marina Bay, Singapore, 1°17'06.26" N, 103°51'20.09" E, coll.

238

Y. Cai & S.L. M. Teo

K. S. Tan & S. C. Lim, 14 Jul. 2003; 84 males, tl 6.8-13.2 mm, 54

females, tl 7.3-10.0 mm, ZRC.2005.0125, CAAS2 Buoy, 1°23'38.65" N,

103°59'37.13" E, off Changi Beach, Singapore, coll. Y. Cai, S. L. M. Teo

& T. M. Sin, 23 Apr. 2002; 3 males, tl 4.8-10.3 mm, 1 female, tl 8.0 mm,

ZRC.2005.0126, Jetty of St John’s Island, 1°13'20.95" N, 103°50'56.33"E,

Singapore, coll. Y. Cai, 22 Mar. 2004.

Description of male. Maximum size: 11.4 x 4.0 mm (holotype).

Body 2.9-3.1 times as long as wide; depressed and smooth,

without tubercles or carina, lateral margin smooth. Colour

brownish with darker spots. Cephalon frontal margin almost

straight, with indistinct median excavation, dorsal surface with

anterior and posterior transverse grooves and longitudinal

lateral grooves. Eyes bulge outward, forming part of contour of

lateral margin of head. Ratio of post-orbital head width to

width of pereonite 3 (widest pereonite) 0.56.

Antenna 1 flagellum uniarticulate, with 10 pairs of jointed

aesthetascs. Antenna 2 0.6 body length; article 4 2.4 times as

long as wide; article 5 3.6 times as long as wide; flagellum with

16-22 articles, 1.2 length of peduncle.

Dorsum of each pereonite smooth, margin slightly arched;

dorsomarginal areas of pereonites 2-7 slightly enlarged,

distinctly depressed, lateral margins slightly upturned,

marginal areas becoming progressively less depressed, and

sloping gently on posterior pereonites; lunettes on pereonites

2- 4 with posterior margin subtriangular or rounded; distolateral

angle of pereonites 1-4 rounded, those of pereonites 5-7

subrectangular.

Pleotelson about 1.4 times as long as wide, dorsum smooth,

evenly convex, lateral margin sub- parallel over anterior two-

thirds, then tapering beyond curved margin to rounded

posterior margin, with a shallow medial excavation.

Mandible incisor with 4 strong, unequal cusps. Lacinia

mobilis stout, 4-cusped, with additional large serrate spine-like

process. Molar process large, truncate, surrounded by short

spines, bearing laterally 3 stiff setulose setae and many

denticles along distal end.

Mesial lobe of maxilla 1 with 2 stout distally serrated

robust apical setae with mesial setules; outer lobe with 9 robust

tooth-like serrated setae.

Maxilla 2 3-lobed, with plumose, simple and comb setae on

endopod as figured; mesial lobe of exopod lined with comb

setae, Outer lobe enlarged, recurved laterally, fringed with

extremely long plumose setae.

Endite of maxilliped with 1 recurved coupling hook, lined

with 7-10 apical moderately slender plumose setae. Palp

3- articulate, last article expanded and fringed with 6-10 long

setae. Epipod laminar, distal margin rounded, outer and distal

margin fringed with fine setae.

Pereopod 1 carpus triangular, flexor margin densely lined

with simple setae and denticles ; propodus 1.9 times as long as

greatest depth, tapering and curving distally, flexor margin

with long simple setae; dactylus elongated, with simple setae.

Pereopods 2-7 similar in form and size, slightly longer than

pereopod 1; carpus subrectangular; flexor margins of ischium

to propodus densely fringed with simple setae and pubescence;

extensor margins of carpus and merus armed with 1 or 2 simple

setae; dactylus more elongate and straighter than that of

pereopod 1. Pereopod 2 propodus as long as merus and carpus

together, 2.8 times as long as wide. Pereopod 4 propodus 2.4

times as long as wide. Pereopod 7 propodus 3.5 times as long

as wide.

Penes fused along entire length, 1.6 times as long as wide,

swollen distally, with notched lateral and distal margins.

Pleopods 1 and 2 with plumose marginal setae on endopods

and exopods, both rami without sutures. Pleopods 1-3 with

about 11, 5 and 3 coupling hooks on inner margin of peduncles

respectively. Pleopod 2 with appendix masculina elongated,

reaching beyond distal margin of endopod by one-sixth of its

length, mostly straight, distal quarter slightly curving

medisally, with numerous spinules distally. Pleopods 3-5 with

few and short simple marginal setae, incomplete transverse

suture present from near middle of the outer margin of exopod.

Uropod 2.9 times as long as distal peduncle width, with

short, simple setae, no oblique ridges on peduncle, distolateral

angle with 3 plumose setae; endopod 0.3 length of peduncle,

mesial length 0.7 proximal suture length, suture at 75° to long

axis, distal margin truncate, at 75° to long axis, lateral margin

gently convex between lateral and distal margins

Female. Maximum size, 7.6 x 3.1 mm (one of paratypes). Body

stouter than male, 2.3-2.6 times as long as wide; pleotelson 1.2

times as long as wide; pereonal margins more evenly curved

than in male. Antenna 2 with 13-15 articled flagellum. Maxilla

2 3-lobed, with plumose, simple and comb setae on endopod as

figured; both inner and outer lobes of exopod lined with comb

setae; no dense pubescence on pereopods 2-7. Oostegites

lamellar on pereonites 1-4.

Etymology. The new species is named after Gary C. B. Poore,

who has contributed significantly to our understanding of

marine isopods in the Indo-Pacific region.

Habitats. The new species was most commonly found in

association with macroalgae and hydroids in the fouling

community of Singapore waters.

Remarks. Currently, 14 species of the genus Synidotea belong

to the ‘5. hirtipes group’, which was defined as a group of

similar species characterized by a smooth body, entire or

slightly excavate front of the head, and excavated pleotelson

apex (Monod, 1931; Menzies and Miller, 1972; Poore, 1996).

The group contains: S. hirtipes (Milne Edwards, 1840), S.

laevidorsalis (Miers, 1881), S. laticauda Benedict, 1879, S.

harfordi Benedict, 1879, S. variegata, Collinge, 1917, S.

marplatensis Giambiagi, 1922, S. fluviatilis Pillai, 1954, S.

worlinensis Joshi and Bal, 1959, S. brunnea Pires and Moreira,

1975, S. hunumantharoei Kumari and Shyamasundari, 1984,

S. keablei Poore and Lew Ton, 1993, S. grisea Poore and Lew

Ton, 1993, S. oahu Moore, 2004, and S. fosteri Schotte and

Heard, 2004. Two more species, S. innatans and S. karumba

from Australia, were just described and added into the group

(Poore, 2012).

Synidotea poorei sp. nov. can be separated from S. hirtipes

easily by its smooth uropodal peduncle (vs. two ridges in S.

hirtipes ). The new species is superficially similar to S.

lavidorsalis, S. laticauda, S. grisea and S. keablei, but it can

be separated from these species by the sub-parallel lateral

margin of the pleotelson. It also differs from S. laevidorsalis

Synidotea poorei, a new isopod from the fouling community in Singapore waters (Valvifera, Idoteidae)

239

Figure 1. Photographs of Synidotea poorei , sp. nov. A. female, tl 7.4 mm, B. male, tl 12.9 mm, NMV. Scale bar = 1 mm.

by the shape of uropodal endopod (fig. 3G vs. fig. If in Poore,

1996) and the fused penial plates (fig. 3H vs. fig. Ik in Poore,

1996); from S. laticauda by the less excavated posterior end of

the pleotelson; and from S. keablei by sexual dimorphism of

the maxilla 2, and the elongated pleotelson. Synidotea poorei

sp. nov. can be distinguished from S. harfordi by the shape of

the lunette on the pereonites 2-4 (rounded vs. triangular) and

the overall body form, which is more slender in the latter.

Synidotea poorei is also similar to S. brunnea from which it

can be separated by the more elongated antennae 1 and 2, and

the shape of uropodal endopod (fig 3G vs. fig. 38 in Pires &

Moreira, 1975). Synidotea poorei , also resembles S. variegata

(cf. Collinge, 1917; Pillai, 1963) from which it differs by the

more stout peduncle of antenna 2 (fig 3 in Collinge, 1917 vs.

fig. 2C), stouter pleotelson (1.3 times as long as wide in female

and 1.5 times in male of S.poorei vs. 1.7 times in S. variegata)-,

and the shallower pleonal suture. With respect to the body

form, the cephalon and the pleotelson, S. poorei is very similar

to the Argentinean species S. marplatensis. It can be separated

by the much longer appendix masculina (Fig. 3C, D vs. Fig. 4

in Giambiagi, 1922); and the smooth uropodal peduncle (vs.

with an oblique ridge). Synidotea poorei can also be easily

separated from the two recently described species, S.fosteri

and S. oahu by its much longer antenna 2, and the smooth

uropodal endopod.

Acknowledgements

The authors would like to thank Tan Koh Siang, Sin Tsai Min

and Lim Swee Cheng (TMSI), for assisting in field collections,

to Gary Poore (NMV) for helping with photographs and

critically reading the manuscript, Niel Bruce (Museum of

Tropical Queensland, Queensland Museum, Townsville,

Australia), who has confirmed identity for some of fouling

isopod species from Singapore. Thanks are also due to MPA

(Maritime and Port Authority of Singapore), for granting the

permission and providing logistic support for the filed

samplings. The present study was financially supported by a

research grant from ASTAR (Agency for Science, Technology

and Research, Singapore).

240

Y. Cai & S.L. M. Teo

Figure 2. Synidotea poorei, sp. nov. A. dorsal view of male specimen, B. dorsal view of female specimen, C. antenna 2, D. right mandible, E. left

mandible, F. right maxilla, G, I. maxilla 2, male, H. inner and median lobes of left maxilla 2, male, close up. Scales: A, B=1.5 mm, C=0.5 mm,

D, E=0.2 mm, F, H=0.1 mm, G=2 mm.

Synidotea poorei, a new isopod from the fouling community in Singapore waters (Valvifera, Idoteidae)

241

Figure 3. Synidotea poorei, sp. nov. A. left maxilliped, B. pleopod 1, male, C. pleopod 2, male, D. appendix masculina of pleopod 2, E. base of

pleopod 1, F. pleopod 3, G. right uropod, H. penial papilla. Scales: A, D, E, G, H=0.2 mm, B, C, F=0.6 mm.

242

Y. Cai & S.L. M. Teo

Figure 4. Synidoteapoorei , sp. nov. A. left antenna 1, B. left pereopod 1, C. left pereopod 2, D. left pereopod 4, E. left pereopod 7. Scales: A-E=0.2 mm.

Synidotea poorei, a new isopod from the fouling community in Singapore waters (Valvifera, Idoteidae)

243

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Memoirs of Museum Victoria 69:245-258 (2012)

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

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

Two further species groups and new species among Australian Hydrobiosella

Tillyard: new species from south-eastern Australia (Trichoptera: Philopotamidae)

David I. Cartwright

13 Brolga Crescent, Wandana Heights, Victoria 3216, Australia (cartwright@hotkey.net.au)

Abstract Cartwright, D.1.2012. Two further species groups and new species among Australian Hydrobiosella Tillyard: new species

from south-eastern Australia (Trichoptera: Philopotamidae). Memoirs of Museum Victoria 69: 245-258.

Eleven philopotamid caddisfly species in the genus Hydrobiosella Tillyard are newly described from Australia,

based on small differences in features of the male genitalia. These species are placed in two new species groups, one based

on H. letti Korboot, the other on H. bandabanda sp. nov.

The five species of the Hydrobiosella bandabanda group, all newly described here: H. bandabanda, H.jibboor, H.

jirrima, H. tarrong and H. tiarka, share features of phallus with two or more embedded spines subapically and both preanal

appendages and parameres absent. Females of four of these species are also figured in part. All five species are endemic to

the eastern half of Victoria, mainland Australia.

The H. letti species group comprises seven species: Hydrobiosella bijurca, H. bos, H. excilatas, H. fibra, H.

incisura, H. letti Korboot and H. tenuitas, united in having features such as elongate, nearly straight parameres arising near

middle to base of phallus, no obvious spines in phallus and presence of preanal appendages. Females of two of these

species are also partly figured. These seven species are all are endemic to eastern coastal mainland Australia, from north¬

eastern Victoria to eastern Queensland.

Keywords Trichoptera, caddisflies, Philopotamidae, Hydrobiosella, Australia.

Introduction

The recent history of the genus Hydrobiosella Tillyard in

Australia is revised and summarized by Cartwright (2010,2012a,

2012b). In brief, the first Australian species in the genus were

described in 1953 by Mosely (in Mosely and Kimmins, 1953).

Further species were added by Korboot (1964); Jacquemart

(1965); (Neboiss, 1977,1982,2003) and Cartwright (2010,2012a,

2012b). A total of 53 species of Hydrobiosella are known

worldwide: from Australia (43 species — Cartwright 2012b),

New Zealand (4 species — Morse 1999) and New Caledonia

(6 species — Espeland and Johannson 2007).

Groups were first recognized among Australian

Hydrobiosella by Neboiss (1977) for Tasmanian species). He

diagnosed three groups based primarily on male genitalia and

these, the H. bispina, H. eminentia and H. waddama species

groups were detailed more recently and included in an updated

key to the Australian species and species group (Cartwright

2010, 2012a, 2012b). Two further species groups are diagnosed

here and include 11 newly described species which bring to 54

the total number of Australian species of Hydrobiosella.

Australian mainland species in the H. waddama group have

recently been reviewed (Cartwright 2012b).

In this study of the newly recognised Hydrobiosella

bandabanda group, 60 male and 10 female specimens were

examined, determined to be referable to five very similar species.

The more common species in the group were, H. tiarka

comprising about 41% of specimens, H. bandabanda 30%, H.

jirrama 17%, and H.tarrong with about 10% of specimens. One

species, H. jibboor, is known from only one specimen. No

differentiation of female genitalia has been established as yet. All

five species were collected from south-eastern Australia, within

the Bassian region, a distribution suggestive of a ‘southern’ origin.

This revision of the Australian ‘ Hydrobiosella letti ’ group is

based on some 37 male and 36 female specimens, referred to

seven species. The most common species, H. bifurca, contributed

about 79% of all H. letti species group specimens. All six other

species are known from fewer than five specimens. The seven

species of this group, including the six new species were

collected from eastern Australia, north-eastern Victoria,

central-eastern New South Wales and eastern Queensland.

Three of the seven species are from north-eastern Queensland,

so southern or northern Australia origins could be postulated

equally for the group.

Methods and abbreviations

Among species in this genus, size, and body and wing colour

can be useful distinguishing characters, but are often variable.

Wing and body colour can be most useful in freshly preserved

246

D.l. Cartwright

material but the colour often fades with time in alcohol. Most

of the material studied has been preserved in alcohol for many

years and was on loan from Museum Victoria, made available

by the late Dr Arturs Neboiss. Depositories for specimens are

abbreviated as follows: Museum Victoria, Melbourne (NMV),

Australian National Insect Collection, Canberra (ANIC), the

Natural History Museum, London (BMNH) and the

Queensland Museum, Brisbane (QM). All specimens,

including types, mentioned in the text are lodged in the NMV

unless stated otherwise.

Males of each species are most easily distinguished by

genitalic features, but usually require clearing of the abdomen

in potassium hydroxide. Females were paired with respective

males on the basis of similarities in size and coloration, and on

wing venation and locality.

Figured specimens are identified by the notebook numbers

of Dr Arturs Neboiss (NMV), prefix PT-; or the author; prefix

CT-. Terminology used generally follows that of Neboiss

(1977, 1982), Blahnik (1998, 2005) and Holzenthal et al.

(2007). Terminology used for genitalic parts are indicated on

selected figures for each genus and additionally where

necessary. Typically, setae or spines are illustrated only on the

right side of the figure (as viewed) to enable a better view of

the underlying structures. Length/width measurements

generally mean maximum length divided by maximum width.

Descriptions

Hydrobiosella Tillyard 1924

Hydrobiosella Tillyard 1924: 288; Mosely and Kimmins 1953:

387; Neboiss 1977: 45; Neboiss 2003: 55.

Type species. Hydrobiosella stenocerca Tillyard by monotypy.

Generic descriptions are given by Tillyard (1924: 288); Mosely

and Kimmins (1953: 387) and Neboiss (1977: 45).

Key to males of known Australian groups (or ungrouped

species) of Hydrobiosella Tillyard (after Cartwright 2010,

2012)

1. Phallus without pair of parameres (Figs 2, 3, 5, 6;

Cartwright 2012a, figs 2, 3; Neboiss 1986, figs pp. 99, H.

amblyopia ; 101, H. tasmanica ; 102, H. corinna ).2

Phallus with pair of parameres (Cartwright 2010, figs 2,3,

5, 6; Neboiss 1986, figs pp. 99, H. michaelseni, H.

waddama-, 101, H. letti; 102, H.bispina-, 103, H. arcuata)

2. Preanal appendages present, usually small

(Cartwright2012a, figs 2, 3; Neboiss 1977, figs 204, 205,

216, 217; Neboiss 1986, figs pp. 101, H. tasmanica-, 102, H.

corinna-, Neboiss 2003, figs 8a-h).3

Preanal appendages absent (Figs 2, 3, 18, 19; Neboiss

1986, figs pp. 99, H. amblyopia-, 101, H. tasmanica ) 4

3. Preanal appendages relatively slender, elongate and

‘unattached’ to segment IX (Cartwright 2012a, figs 2,3,5);

NE-Qld. Hydrobiosella eminentia group

Preanal appendages often short and bulbous or ‘attached’

to segment IX (Neboiss, 1977, figs 204-211; Neboiss 1986,

figs p. 102, H. corinna-, Neboiss, 2003, figs 8A-H); Tas

. Hydrobiosella corinna group

4. Phallus apically with downward projecting spine(s)

(Neboiss 1977, figs 216-221, 225, 226; Neboiss 1986, figs

p. 101, H. armata, H. tasmanica-, Neboiss 2003, figs 10A-

G, 11A-G, 12A-F); Tas. H. tasmanica group

Phallus apically without downward projecting spine(s)

(Figs 2, 3; Neboiss 1982, fig. 12; Neboiss 1986, figs p. 99

H. amblyopia ).5

5. Segment IX with a large disto-lateral projection (Neboiss

1982, fig. 12; Neboiss 1986, figs p. 99 H. amblyopia );

S-WA. H. amblyopia (ungrouped)

Segment IX without a large disto-lateral projection (Figs

2, 3, 5, 6); C and NE Vic. H. bandabanda group

6. Inferior appendages with harpago with dark row of setae

forming fringe along ventral margin (Cartwright 2010,

figs 3, 6; Neboiss 1986, figs pp. 102, H.bispina-, 103, H.

arcuata)-, E-Vic, E-NSW, E-Qld.

. Hydrobiosella bispina group

Inferior appendages with harpago without dark row of

setae forming fringe along ventral margin (Figs 18, 19;

Neboiss 1986, figs pp. 99, H. michaelseni, H. waddama)

7. Parameres elongate and sinusoidal, attached ventrally to

base of phallus (Cartwright 2012b, figs 2, 3, 5, 6; Neboiss

1977, fig. 233; Neboiss 1986, figs p. 99, H. waddama)-, Tas,

SE Aust.. Hydrobiosella waddama group

- Parameres not elongate and sinusoidal, not attached

ventrally to base of phallus (Figs 18, 19; Neboiss 1982,

figs 9, 10; Neboiss 1986, figs pp. 99, H. michaelseni-, 101,

H. letti) .8

8. Parameres curved strongly and crossed (Neboiss 1982,

figs 9, 10; Neboiss 1986, figs p. 99; H. michaelseni)-, S-WA

... Hydrobiosella michaelseni (Ulmer) (unplaced to group)

Parameres not curved strongly and crossed (Figs 18, 19;

Neboiss 1986, figs p. 101, H. letti)-, NE-Vic, CE-NSW,

E-Qld. Hydrobiosella letti group

Hydrobiosella bandabanda group

Diagnosis. Key characters of males in the group are phallus

without parameres but with two or more embedded spines

subapically and preanal appendages absent.

Description. Head and nota dorsally brown to dark brown

with pale setal warts and scutellum, abdomen brownish

dorsally and ventrally, paler laterally; wings light brown to

brown. Medium-sized adults. Forewing length about 2.8-3.1

times maximum width, males: 5.9-9.8 mm; females: 6.1-10.8

mm; wing venation (Fig. 1), similar to the type species H.

stenocerca (Mosely and Kimmins 1953, fig. 265a), H. bispina

(Cartwright, 2010, fig. 1) and H. waddama (Mosely and

Two further species groups and new species among Australian Hydrobiosella Tillyard: new species from south-eastern

Australia (Trichoptera: Philopotamidae)

247

Kimmins 1953, fig. 269a), R1 simple, forks 1, 2, 3, 4 and 5

present; forks 1 and 2 sessile; fork 2 with nygma present,

length about 1.3-1.4 times length fork 1; fork 3 shorter, length

0.7 times length fork 2, fork 3 length ranging from between

1.8-1.9 times length footstalk, cross-veins r-m and m

contiguous at fork 3; fork 4 similar length to fork 3, fork length

about 7 times length footstalk; fork 5 very long, length between

1.6-1.7 times length fork 4; discoidal cell closed, length

between 3.7-3.8 times maximum width. Hind wing length

about 2.4-2.5 times maximum width; forks 1, 2, 3 and 5

present; forks 1 and 2 sessile; fork 2 with nygma, length fork 2

between 1.4-1.5 times length fork 1; fork 3 shorter, length

about 0.6 times length fork 2, fork 3 slightly longer than

footstalk, length fork ranging between 1.3-1.4 times length

footstalk; fork 5 very long, length between 1.8-1.9 times

length fork 3; discoidal cell closed, length between 4.0-4.1

times maximum width; with three longer anal veins (Fig. 1).

Male. Segment IX with a shallow, wide V notch medially on

distal margin (Figs 4, 7, 10). Preanal appendages absent.

Segment X mainly sclerotised, dorso-ventrally compressed; in

dorsal view, with a ‘tongue-shaped’ process, tapered distally,

rows of fine hairs meso-laterally and usually a pair of small

knobs baso-laterally (Figs 2, 5). Phallus generally robust, tube¬

like, with a pair of spines sub-apically, sometimes with one or

two spines more basally (Figs 2, 3, 5, 6). Inferior appendages

with two segments, in lateral view, basal segment usually more

robust and longer than harpago. Harpago has a small field of

dark spines meso-distally (Figs 3, 6).

Female. Genitalia typical of genus with a small projection on

sternite IX mesodistally (Figs 38-41).

Larva. Confirmed larvae are known for H. bandabanda. These

larvae match Hydrobiosella sp. AV2 (Cartwright, 1997). The

diagnostic features are head wide and angular laterally, and

forecoxa with two sclerotised processes on anterior margin,

one longer than the other (Cartwright, 1997, fig. 1.3).

Hydrobiosella sp. AV2 larvae are recorded mainly from riffle

habitats of very small to medium-sized streams between 0.4-8

m wide at moderate to high altitudes between 800-1460 m

(Suter et al. 2006).

Key to males of species of the Hydrobiosella bandabanda

group from Australia

1. Segment X dorsally with a central ridge bearing pair of

acute lateral processes (Figs 2, 2a); EC-Vic.

. H. bandabanda

Segment X without a central dorsal ridge (Figs 5, 8).2

2. Segment X with sub-apical spine (Fig. 6); E-Vic.

. H. tiarka

Segment X without apical spine (Figs 9, 12).3

3. Segment X in lateral view with apical half slender, not

dilated apically; inferior appendages with terminal

segment dilated slightly in apical half (Fig. 9); NE-Vic

. H. tarrong

Segment X in lateral view with apical half robust, dilated

apically; inferior appendages with terminal segment not

dilated in apical half (Figs 12, 15).4

4. Segment X with a dorsal ‘bump’ subapically (Fig. 12);

NE-Vic. H.jibboor

Segment X without a dorsal ‘bump’ subapically (Fig. 15);

NE-Vic. H.jirrima

Hydrobiosella bandabanda sp. nov.

Figures 1, 2, 2a, 3, 3a, 4, 4a, 38, 38a

Holotype. Male. Victoria, Toorongo Falls, 7 km NE of Noojee,

(about 37°51'S, 146°00'E), 27 Nov 198F J. Morse & A. Neboiss (NMV,

T- 21392).

Paratypes. Victoria. 4 males (specimen CT-593 figured), 1 female

(specimen CT-594 figured), collected with holotype; 7 males

(specimen PT-581 figured), 2 females (specimen CT-612 figured),

Newlands Ck, Upper Thomson R. (about 37°37'S, 146°10'E), 26 Feb

1978, TR Survey (NMV).

Other material examined. Victoria. 2 males. Falls Ck Ski Village,

26 Mar 1957, A. Neboiss; 1 male, Cumberland Falls, SE of Marysville,

7 Jan 1971, A. Neboiss; 1 male, Baw Baw Ski Village, 4900 ft, 8 Jun

1974, J.C.; 1 male. Cement Ck, Mt Donna Buang Rd, 3 Mar 1980, J.

Dean?; 1 male, small ck 1 km N Rum Ck, 9 Nov 1983, D. Cartwright

(NMV).

Diagnosis. Males of Hydrobiosella bandabanda can be

distinguished from those of other species in the group by a

central dorsal ridge or groove with a pair of lateral acute

processes on segment X.

Description. Wings typical of the genus (Fig. 1), similar to

those of H. bispina (Cartwright, 2010, fig. 1). Length of

forewing; male 6.7-9.1 mm, female 1.2-93 mm.

Male. Segment IX with a shallow, wide V-shaped notch medially

on distal margin (Figs 4, 4a). Segment X in dorsal view, a

‘tongue-shaped’ process, with sides almost parallel, a pair of

small knobs baso-laterally, tapered distally and with a central

dorsal ridge or groove bearing a pair of lateral acute processes

(Figs 2, 2a), length about 3-3.5 times width; in lateral view

broad-based, slightly curved (Fig. 3), or straight (Fig. 3a) slender

in middle, usually slightly dilated distally with a sub-apical

‘knob’ (Fig. 3), sometimes without ‘knob’ (Fig. 3a). Phallus

generally tube-like, robust, with a pair of sub-apical spines and

two smaller spines basally (Figs 2,2a, 3,3a). Inferior appendages

in lateral view, with basal segment broadest near middle, length

about twice maximum width; harpago shorter, length about half

length basal segment, more slender, length about twice

maximum width, apex broadly rounded (Figs 3,3a).

Female. Genitalia typical of genus, with a small acute projection

on sternite IX meso-distally (Figs 38, 38a).

Etymology. Bandabanda - Australian aboriginal word for split

or broken open (dorsal groove on segment X).

Remarks. Hydrobiosella bandabanda shows slight variation

among the 21 specimens collected from 7 localities in eastern-

central and north-eastern Victoria (latitudinal range 36°52'-

37°51'S).

248

D.l. Cartwright

Figures 1-7. Hydrobiosella spp.; 1, Hydrobiosella bandabanda sp. nov., distal part of wings; 2-7, Hydrobiosella spp., male genitalia in dorsal,

lateral and part ventral views; 2-4, Hydrobiosella bandabanda sp. nov.; 2, dorsal; 3, lateral; 4, ventral, mesodistal margin of segment IX; 2a-4a,

Hydrobiosella bandabanda sp. nov. (variant); 2a, dorsal; 3a, lateral; 4a, ventral, mesodistal margin of segment IX; 5-7, Hydrobiosella tiarka sp.

nov.; 5, dorsal. 6, lateral; 7, ventral, mesodistal margin of segment IX.

Two further species groups and new species among Australian Hydrobiosella Tillyard: new species from south-eastern

Australia (Trichoptera: Philopotamidae)

249

Figures 8-16. Hydrobiosella spp.; male genitalia in dorsal, lateral and part ventral views; 8-10, Hydrobiosella tarrong sp. nov.; 8, dorsal, 9,

lateral, 10, ventral, mesodistal margin of segment IX; 11-13, Hydrobiosella jibbor sp. nov.; 11, dorsal; 12, lateral; 13, ventral, mesodistal margin

of segment IX; 14-16, Hydrobiosella jirrima sp. nov.; 14, dorsal; 15, lateral; 16, ventral, mesodistal margin of segment IX.

250

D.l. Cartwright

Figures 17-23. Hydrobiosella spp.; 17, Hydrobiosella letti Korboot., wings. 18-23, Hydrobiosella spp., male genitalia in dorsal, lateral and part

ventral views; 18-20, Hydrobiosella fibra sp. nov.; 18, dorsal; 19, lateral; 20, ventral, mesodistal margin of segment IX; 21-23, Hydrobiosella

bifurca sp. nov.; 21, dorsal; 22, lateral; 23, ventral, mesodistal margin of segment IX.

Two further species groups and new species among Australian Hydrobiosella Tillyard: new species from south-eastern

Australia (Trichoptera: Philopotamidae)

251

Figures 24-32. Hydrobiosella spp.; male genitalia in dorsal, lateral and part ventral views; 24-26, Hydrobiosella incisura sp. nov.; 24, dorsal;

25, lateral; 26, ventral, mesodistal margin of segment IX; 27-29, Hydrobiosella tenuitas sp. nov.; 27, dorsal; 28, lateral; 29, ventral, mesodistal

margin of segment IX; 30-32, Hydrobiosella exilatis sp. nov.; 30, dorsal; 31, lateral; 32, ventral, mesodistal margin of segment IX.

252

D.l. Cartwright

Figures 33-37. Hydrobiosella spp.; male genitalia; 33-35, Hydrobiosella bos sp. nov.; male genitalia in dorsal, lateral and part ventral views; 33,

dorsal; 34, lateral; 35, ventral, mesodistal margin of segment IX; 36-37, Hydrobiosella letti Korboot; male genitalia in dorsal and lateral views;

36, dorsal; 37, lateral.

Hydrobiosella tiarka sp. nov.

Figures 5-7, 39

Holotype. Male. Victoria. Cement Ck nr Warburton (about

37°43'S, 145°43'E), 8 Dec. 1970, A. Neboiss (NMV, T-21407).

Paratypes. Victoria. 7 males, collected with holotype; 4 males

(specimen PT-584 figured), same loc. and coll., 27 Mar 1972; 1 male,

same loc., 29 Sep 1990, B. Armitage; 1 male. Cement Ck, 5 Feb 1955,

A. Neboiss; 1 male, same loc. and coll., 19 Nov 1955; 1 female

(specimen CT-614 figured). Cement Ck, Mt Donna Buang, 8 Apr 1976,

Cartwright; 1 male, same loc. and coll., 1 Dec 1976 (NMV).

Other material examined. Victoria. 3 males. Cement Ck Mt

Donna Buang-Warburton Rd, 23 Feb 2012, D. Cartwright; 1 male.

Erica, 29 Jan 1960, A. Neboiss; 1 male, 1 female, Toorongo Falls, NE

Noojee, 17 Dec 1970, A. Neboiss; 1 male, Brittania Ck, 6km S of

Warburton, 27 Feb 1976, A. Neboiss; 3 males, 1 female, Ada R. on

Ada River Rd, 37°50.8'S 145°52'E, 19 Jan 1979, Latrobe C Survey; 1

male, same loc. and coll., 10 Feb 1980 (NMV).

Diagnosis. Males of Hydrobiosella tiarka can be separated

from those of other species in the group by the small apical

spine and slightly bulbous apex on segment X.

Description. Wings similar to those of H. bandabanda (Fig. 1);

length of forewing, male 6.7-9.8 mm; female 8.2-10.8 mm.

Male. Segment IX with a shallow, wide V-shaped notch medially

on distal margin (Fig. 7). Segment X in dorsal view, a ‘tongue¬

shaped’ process, with almost parallel sides in basal half, with a

pair of very small knobs meso-laterally, tapered strongly distally

to a small apical, slightly bulbous knob, length about 2.9 times

width, without a central dorsal ridge or groove (Fig. 5); in lateral

view broad-based, narrowed at basal third, slender with almost

parallel sides in distal two thirds, with short apical spine (Fig.

6). Phallus generally tube-like, robust, with a pair of slightly

divergent sub-apical spines (Figs 5, 6). Inferior appendages in

lateral view, slender, with basal segment sub-rectangular, length

Two further species groups and new species among Australian Hydrobiosella Tillyard: new species from south-eastern

Australia (Trichoptera: Philopotamidae)

253

Figures 38-43. Hydrobiosella spp. Female genitalia (part segment VIII) in lateral view; 38, Hydrobiosella bandabanda sp. nov.; lateral; 38a,

Hydrobiosella bandabanda sp. nov. (variant) lateral; 39, Hydrobiosella tiarka sp. nov.; lateral; 40, Hydrobiosella tarrong sp. nov.; lateral; 41

Hydrobiosella jirrima sp. nov.; lateral; 42, Hydrobiosella bifurca sp. nov.; lateral; 43, Hydrobiosella bos sp. nov

about 2.8 times maximum width; harpago shorter, length about

half length basal segment, length about 2.1 times maximum

width, with broad rounded apex (Fig. 6).

Female. Genitalia typical of genus, with a small projection on

sternite IX meso-distally (Fig. 39).

Etymology. Tiarka - Australian Aboriginal word for toothpick

or sharp (apical spine on tergum X).

Remarks. Twenty-six male and three female specimens of

Hydrobiosella tiarka have been collected from six localities in

eastern-central Victoria (latitudinal range 37°43'-37°58' S).

Hydrobiosella tarrong sp. nov.

Figures 8-10, 40

Holotype. Male. Victoria, Cement Ck nr Warburton (about

37°43'S, 145°43'E), 26 Mar 1958, A. Neboiss (NMV, T-21425).

Paratypes. Victoria. 2 males (specimen CT-609 figured), 3 females

(specimen CT-610 figured), collected with holotype; 1 male, same loc.

and coll., 27 Mar 1972 (NMV).

Diagnosis. Males of Hydrobiosella tarrong can be separated

from those of other species in the group by the slender apical

half of segment X, in lateral view and the inferior appendages

with harpago dilated slightly in apical half.

Description. Wings similar to those of H. bandabanda (Fig. 1);

length of forewing: male 7.8-9.0 mm, female 8.5-9.0 mm.

Male. Segment IX with a shallow, wide notch medially on

distal margin (Fig. 10). Segment X in dorsal view, a ‘tongue-

shaped’ process, with sides almost parallel in basal half, with a

pair of very small knobs baso-laterally, tapered distally to a

slightly rounded apex, without a central dorsal ridge or groove

(Fig. 8); length about 1.7 times width; in lateral view broad-

based, narrowed strongly at midpoint, slender with almost

parallel sides in distal half (Fig. 9). Phallus generally tube-like,

robust, slightly truncate, with a pair of sub-apical spines and

two groups of chitinous spines basally (Figs 8, 9). Inferior

appendages in lateral view, robust, with basal segment sub-

rectangular, slightly angular meso-dorsally, length about 1.8-

1.9 times maximum width; harpago shorter, length about 0.8

times length basal segment, length about 1.9 times maximum

width, with slightly dilated, broadly rounded apex (Fig. 9).

Female. Genitalia typical of genus, with a small projection on

sternite IX meso-distally (Fig. 40).

Etymology. Tarrong - Australian aboriginal word for teeth

(pair of ‘chitinous spines 4 in middle of phallus).

Remarks. Four male and three female specimens of

Hydrobiosella tarrong have been collected from the type

locality in eastern-central Victoria (latitude 37°43'S).

Hydrobiosella jibboor sp. nov.

Figures 11-13

Holotype. Male (specimen CT-611 figured). Victoria, Mt Buller,

headwaters Chalet Ck, 1400 m (about 37°10'S, 146°25'E), 19 Mar.

1993,1. Campbell (NMV, T- 21437).

Diagnosis. Males of Hydrobiosella jibboor can be separated

from those of other species in the group by the dorsal ‘raised

ridge’ sub-apically on segment X and rounded apex and

segment IX with almost straight distal margin.

Description. Wings similar to those of H. bandabanda (Fig. 1),

length of forewing: male 6.9 mm.

254

D.l. Cartwright

Male. Segment IX with a weak notch in meso-ventral margin

(Fig. 13), and with an almost straight distal margin (Fig. 12).

Segment X in dorsal view, a ‘tongue-shaped’ process, with almost

parallel sides in basal third, with a pair of very small knobs baso-

laterally, tapered distally to a rounded apex, length about 2.3

times width, without a central dorsal ridge or groove (Fig. 11); in

lateral view tapered slightly in basal two thirds, slightly bulbous

sub-apically with a small dorsal ‘raised ridge’ (Fig. 12). Phallus

generally tube-like, robust, with a pair of narrowly separated sub-

apical spines, with two more spines basally (Figs 11, 12). Inferior

appendages in lateral view, robust, with basal segment sub-

rectangular, length about twice maximum width; harpago shorter,

length about half length basal segment, length about 1.6-1.7

times maximum width, with broad rounded apex (Fig. 12).

Female. Unknown.

Etymology. Jibboor - Victorian Aboriginal word for mountain

(type locality — Mt Buller).

Remarks. A single male specimen of Hydrobiosella jibboor has

been collected from the type locality in north-eastern Victoria

(latitude 37°10'S).

Hydrobiosella jirrima sp. nov.

Figures 14-16, 41

Holotype. Male. Victoria, McKay Ck, Sassafras Gap (about

36°37'S, 147°47'E), 2 Feb. 1974, A. Neboiss (NMV, T- 21438).

Paratypes. Victoria. 3 males (specimen CT-586 figured), collected

with holotype; 5 males, 1 female (specimen CT-613 figured). Sassafras

Gap, 13 Feb 1963, A. Neboiss; 1 male, Gibbo R., S of Donovan Ck jn,

3 Feb 1974, A. Neboiss; 1 male, roadside trickles, 1.2km N Sassafras

Gap, It tr., 11 Feb 2010, R. StClair and D. Cartwright (NMV).

Diagnosis. Males of Hydrobiosella jirrima can be distinguished

from those of other species in the group by the absence of a short

dorsal ‘raised ridge’ subapically and slender apex on segment X.

Description. Wings similar to those of H. bandabanda (Fig. 1);

length of forewing, male 6.6-7.8 mm, female 6.6 mm.

Male. Segment IX with a weak wide notch in meso-ventral

margin (Fig. 16), and an almost rounded distal margin (Fig.

15). Segment X in dorsal view, a ‘tongue-shaped’ process, with

almost parallel sides in basal quarter, with a pair of very small

acute knobs baso-laterally, tapered distally to a narrow apex,

without a central dorsal ridge or groove (Fig. 14); length about

1.9 times width; in lateral view weakly broad-based, tapered

slightly in basal two thirds, very slightly bulbous apically (Fig.

15). Phallus generally tube-like, robust, with a pair of narrowly

separated sub-apical spines (Figs 14, 15). Inferior appendages

in lateral view, robust, with basal segment sub-rectangular,

length about 2.3 times maximum width; harpago shorter,

length about 0.6 times length basal segment, length about twice

maximum width, with broad rounded apex (Fig. 15).

Female. Genitalia typical of genus, with a small projection on

sternite IX meso-distally (Fig. 41).

Etymology. Jirrima - Australian aboriginal word for a

mountain (type locality Sassafras Gap).

Remarks. Eleven males and one female of Hydrobiosella jirrima

have been collected from three sites near the type locality in

north-eastern Victoria (latitudinal range 36°37'-36 0 39'S).

Hydrobiosella letti group

Diagnosis. Key characters of males in the group are parameres

elongate and straight or slightly curved, arising near base or

middle of phallus, and preanal appendages absent.

Description. Head and body generally brown; wings light brown

to brown. Medium sized adults. Forewing length, males: 4.5-8.0

mm; females: 5.8-9.4 mm; forewing length about 2.9-3.0 times

maximum width, wing venation (Fig. 17), similar to H.

stenocerca (Mosely and Kimmins 1953, fig 265a), H. bispina

(Cartwright, 2010, fig. 1) and H. waddama (Mosely and Kimmins

1953, fig. 269a), R1 simple, forks 1, 2,3, 4 and 5 present; forks 1

and 2 sessile; fork 2 with nygma, about 1.3-1.4 times length fork

1; fork 3 shorter, length about two-thirds length fork 2, fork 3

length ranging from between 1.6-17 times length footstalk,

cross-veins r-m and m contiguous at fork 3; fork 4 similar length

to fork 3, fork length about 2.0-2.2 times length footstalk; fork 5

very long, length between 1.9-2.0 times length fork 4; discoidal

cell closed, length between 37-3.9 times maximum width. Hind

wing length about 2.6-27 times maximum width, with forks 1,2,

3 and 5 present; forks 1 and 2 sessile; fork 2 with nygma present,

fork 2 length between 1.6-17 times length fork 1; fork 3 shorter,

length about two-thirds length fork 2, fork 3 longer than footstalk,

length fork ranging between 1.9-2.1 times length footstalk; fork

5 very long, length between 1.6-1.7 times length fork 3; discoidal

cell closed, length between 4.1-4.3 times maximum width; with

three longer anal veins (Fig. 17).

Male. Segment IX in lateral view, length between 1.2-1.9 times

width, usually sub-rectangular (Figs 19, 37), occasionally

projecting and tapered basally (Fig. 25) with a shallow, wide V

notch (Figs 29,32) or wide, shallow concavity medially on distal

margin (Figs 23, 26). Segment X mainly sclerotised dorsally,

membranous ventrally, usually broadbased, tapered distally; in

dorsal view, with a ‘tongue-shaped’ process (Figs 18, 21, 24).

Phallus generally tube-like, parameres slender, elongate and

straight or slightly curved, arising near base of phallus (Figs 19,

22, 28). Inferior appendages with two segments, in lateral view,

basal segment usually more robust than harpago. Harpago has a

small field of dark spines meso-distally (Figs 19, 22, 31).

Female Genitalia typical of genus (Figs 42, 43).

Larva. No confirmed larvae are known for this group.

Remarks. The seven species in this group are known from NE-

Victoria, eastern New South Wales and eastern Queensland.

Females of only two species have been associated.

Key to males of species of the Hydrobiosella letti group

from Australia

1. Segment X with pair of baso-lateral lobes (Figs 18, 19);

SE-Qld. H.jibra

Segment X without pair of baso-lateral lobes (Figs 21, 22,

24, 25).2

Two further species groups and new species among Australian Hydrobiosella Tillyard: new species from south-eastern

Australia (Trichoptera: Philopotamidae)

255

2. Segment X in dorsal view, with pair of small apico-lateral

projections (Fig. 22); NE-Qld. H. bifurca

Segment X in dorsal view, without pair of small apico-

lateral projections (Figs 25, 28).3

3. Segment X laterally compressed apically, with distinctive

notch (Figs 24, 25); SE-Qld. H. incisura

Segment X not laterally compressed apically, without

distinctive notch (figs 27, 28).4

4. Inferior appendages in lateral view with harpago length >

3.5 times width (Figs 28, 31).5

Inferior appendages in lateral view with harpago length <

2.5 times width (Figs 34, 37).6

5. Segment X in lateral view, slightly dilated apically (Fig.

28); in dorsal view, slender; length about 2.8 times

maximum width (Fig. 27); NE-Qld. H. tenuitas

Segment X in lateral view, not dilated apically, instead

slender and upturned (Fig. 31); in dorsal view, not slender;

length about 1.4 times maximum width (Fig. 30); NE-Qld

. H. exilatis

6. Inferior appendages with basal segment angled dorso-

mesally (Fig. 34); NE-Vic. H. bos

Inferior appendages with basal segment not angled dorso-

mesally (Fig. 37); CE-NSW. H. letti

Hydrobiosella fibra sp. nov.

Figures 18-20

Holotype. Male. Queensland, Kenilworth rainforest (about

26°36'S, 152°44'E), 7 Apr. 1967, N. Dobrotworsky (NMV, T- 21450).

Paratypes. Queensland. 2 males, collected with holotype

(specimen PT-578 figured) (NMV).

Diagnosis. Males of Hydrobiosella fibra can be separated from

other species in the group by the presence of a pair of stout

baso-lateral lobes on tergum X.

Description. Wings similar to H. letti (Fig. 17); length of

forewing, male 63-6.6 mm.

Male. Segment IX in lateral view, sub-quadrate, length about

1.8-1.9 times width, projecting slightly basally (Fig. 6.3), with a

shallow rounded notch medially on distal margin (Fig. 20).

Segment X with a pair of large sub-ovate baso-lateral lobes

(Figs 18, 19); in dorsal view, ‘tongue-shaped (fig. 18); in lateral

view slender, slightly tapered distally (Fig. 19). Phallus

generally tube-like, with a pair of straight, slender lateral

‘parameres’ arising basally (Figs 18, 19). Inferior appendages

in lateral view, with basal segment sub-rectangular, length

about 1.7-1.8 times maximum width; harpago slightly shorter,

length about 1.6 times maximum width; bulbous dorso-basally,

tapered slightly distally, broadly rounded apically (Fig. 19).

Female. Unknown.

Etymology. Fibra - Latin for lobe (on tergum X).

Remarks. Three male specimens of Hydrobiosella fibra have

been collected from the type locality in south-eastern

Queensland (latitude 26°36'S).

Hydrobiosella bifurca sp. nov.

Figures 21-23, 42

Holotype. Male. Queensland, unnamed Ck, Paluma Dam Rd, Mt

Spec State Forest, 18°57'S, 146°10'E, 860m, 24 Feb 1994, A.L. Sheldon

(NMV, T- 21453).

Paratypes. Queensland. 2 males, 3 females, same site and collector,

17 Jan 1994; 2 females, same site and collector, 6 Jul 1994, A.L.S.; 2

males, 5 females, unnamed Ck, ‘cascade’ on Paluma Dam Rd, 840m, 6

July 1994, A.L.S.; 1 female (specimen CT-656 figured), same site and

collector, 31 Oct 1993; 1 male. Camp Ck trib., Mt Spec State Forest,

18°57'S, 146° 10'E, 760m, 22 Feb 1994, A.L.S.; 1 female, same site and

collector, 23 Apr 1994; 4 females, same site and collector, 6 Jul 1994; 1

male, 1 female. Birthday Ck, below falls, Mt Spec State Forest, 18°57'S,

146°10'E, 760m, 29 Mar 1994, A.L.S.; 1 male (specimen PT-1768

figured), Cairns, Lake Morris Rd, 16 Nov 1988, K. Walker (NMV).

Other material examined. Queensland. 1 male, Barron Falls,

Kuranda, 16 Jun 1971, E.F. Riek (ANIC); 1 female, U Freshwater Ck,

Whitfield Range, Cairns, 24 Aug 1974, M.S. Moulds; 1 male, 1 female,

Lock-Davies Ck Rd, Lamb Ra, Mareeba Dist. 10 Nov 1974, M.S.

Moulds; 2 males. Fishery Falls, 17°11S, 145°52E, 10-11 Nov 2007, A.

Cairns, A. Wells, W. Cairns (ANIC); 1 male, small waterfall on Kirrama

Range Rd, 9.2km from Nat. Pk sign, -18.20°, 145.83°, 9 May 2011, M.

Shackleton and J. Mynott; 1 male, 1 female, 1 km N of Tully Falls, 8 Jan

1976, A. Walford-Huggins; 1 male, Tully Falls, S of Ravenshoe, 11 Jan

1977, Moulds; 1 male, same locality and date, M.S. and B.J. Moulds; 1

male. Birthday Ck, 3.5 km WNN Paluma, 18°59'S, 146°10'E, 7 Apr

1990, R. StClair; 1 male, 1 female, same site and collector, 23 Dec 1989;

1 female, same site and collector, 19 Jan 1990; 1 female. Birthday Ck,

above weir, Mt Spec State Forest, 18°57’S, 146°10’E, 820m, 22 Oct 1993,

A.L.S.; 1 female, same site and collector, 6 Nov 1993; 1 female, same

site and collector, 23 Apr 1994; 1 female. Birthday Ck, below falls, Mt

Spec State Forest, 18°57'S, 146°10'E, 760m, 17 Mar 1994, A.L.S.; 1

female, same site and collector, 6 Jul 1994; 2 females. Birthday Ck, Iron

Cabin, Mt Spec State Forest, 18°57'S, 146°10’E, 790m, 23 Apr 1994,

A.L.S.; 1 female. Camp Ck trib.,Mt Spec State Forest, 18°57’S, 146°10'E,

760m, 15 May 1994, A.L.S.; 1 female, same site and collector, 15 Mar

1994; 1 female, unnamed Ck, Paluma Dam Rd, Mt Spec State Forest,

18°57'S, 146°10'E, 860m, 11 Jun 1994, A.L. Sheldon; 1 female, Goodard

Ck, Kirrama State For., 18°06'S, 145°41'E, Apr 1993, G. Theischinger; 8

males, 2 females, river on L. Morris Rd, 16.94°S, 145.72°E, M.

Shackleton and J. Mynott (NMV).

Diagnosis. Males of Hydrobiosella bifurca can be separated

from those of other species in the group by the pair of small

apico-lateral projections on segment X.

Description. Wings similar to H.letti (Fig. 17); length of

forewing, male 5.4-6.6mm, female 5.8-7.5 mm.

Male. Segment IX in lateral view, sub-quadrate, length about

1.5 times width, projecting slightly basally (Fig. 22), with a

very shallow, wide concavity medially on distal margin (Fig.

23). Segment X robust, in dorsal view, sub-quadrate, length

about 1.1-1.2 times width, with a pair of small apico-lateral

projections (Fig. 21); in lateral view slightly tapered distally,

with a slightly upturned apex (Fig. 22). Phallus generally tube¬

like; with a pair of straight, slender lateral ‘parameres’ arising

from the phallus near middle (Figs 21, 22). Inferior appendages

256

D.l. Cartwright

in lateral view, with basal segment sub-rectangular, length

about 1.3 times maximum width; harpago slightly shorter,

more slender, length about 1.5 times maximum width, tapered

slightly distally, broadly rounded apically (Fig. 22).

Female. Genitalia typical of genus (Fig. 42).

Etymology. Bifurca - Latin for having two prongs or forks

(apico-lateral projections on tergum X).

Remarks. Some 24 male and 34 female specimens of

Hydrobiosella bifurca have been collected from at least twelve

sites in north-eastern Queensland (latitudinal range 16°49'-

18°59'S). Hydrobiosella bifurca was cited in a checklist

(Walker et al., 1995) as Hydrobiosella sp. nov. PT-1768.

Hydrobiosella incisura sp. nov.

Figures 24-26

Holotype. Male (specimen CT-557 figured), Queensland, National

Park (about 28°14'S, 153°09'E), 5 Jun. 1955, collector? (NMV,

T-21478).

Diagnosis. Males of Hydrobiosella incisura can be separated

from those of other species in the group by the laterally

compressed apex on segment X with distinctive notch.

Description. Wings similar to H. letti (Fig. 17), length of

forewing: male 6.5 mm.

Male. Segment IX in lateral view, length about 1.3 times width,

projecting and tapered basally (Fig. 25), with a very shallow,

wide concavity medially on distal margin (Fig. 26). Segment X

robust in basal two-thirds, narrowed in apical third (Figs 24,

25), in dorsal view, slender in apical third (Fig. 24); in lateral

view, narrowed subapically, dilated apically with distinctive

notch (Fig. 25). Phallus generally tube-like; with a pair of

straight, slender lateral ‘parameres’ arising from the phallus

near base (Figs 24, 25). Inferior appendages in lateral view, with

basal segment sub-rectangular, length about 2.1 times maximum

width; harpago shorter, length about 0.7 times length basal

segment, more slender, length about 2.5 times maximum width,

tapered slightly distally, broadly rounded apically (Fig. 25).

Female. Unknown.

Etymology. Incisura - Latin for notch (apex of segment X).

Remarks. The single male specimen of Hydrobiosella incisura

was collected from the type locality in south-eastern

Queensland (latitude 28°14'S).

Hydrobiosella tenuitas sp. nov.

Figures 27-29

Holotype. Male (specimen PT-1038 figured), Queensland,

Bellenden Ker Range, summit TV stn, 1560m (about 17°16'S,

145°54'E), 1-7 Nov 1981, Earthwatch-Qld Mus. (NMV, T-21479).

Diagnosis. Males of Hydrobiosella tenuitas can be separated

from those of other species in the group by the slender segment

X, also slightly dilated apically in lateral view, and the long and

slender harpago of the inferior appendages.

Description. Wings similar to H. letti (Fig. 17), length of

forewing: male 7.6 mm.

Male. Segment IX in lateral view, length about 1.2 times

width, sub-rectangular, not projecting and tapered basally,

projecting slightly distally (Fig. 28), with a shallow, wide

V-shaped notch medially on distal margin (Fig. 29). Segment

X slender, almost parallel sided in basal two-thirds, length

about 2.8-2.9 times width, narrowed in apical third (Figs 27,

28), in dorsal view, narrow ‘tongue-shaped, tapered slightly in

distal third, narrowly rounded apex (Fig. 27); in lateral view,

narrowed at about apical third, dilated slightly distally with

bulbous apex (Fig. 28). Phallus generally tube-like; with a pair

of slightly curved, slender lateral ‘parameres’ arising from the

phallus near middle (Figs 27, 28). Inferior appendages in

lateral view, with basal segment sub-trapezoidal, length about

1.4 times maximum width; harpago longer, length about 1.6

times length basal segment, more slender, length about 3.8-3.9

times maximum width, tapered slightly distally, broadly

rounded apically (Fig. 28).

Female. Unknown.

Etymology. Tenuitas - Latin for slender (segment X and

harpago of inferior appendages).

Remarks. The single male specimen of Hydrobiosella tenuitas

was collected from north-eastern Queensland (latitude 17°16'S).

Hydrobiosella tenuitas was cited in a checklist (Walker et al.,

1995) as Hydrobiosella sp. nov. PT-1038.

Hydrobiosella exilatis sp. nov.

Figures 30-32

Holotype. Male: Queensland, Bellenden Ker Range, cable tower 3,

1054m (about 17°16'S, 145°54'E), 25-31 Oct 1981, Earthwatch-Qld

Mus. (NMV, T-21480).

Paratypes. 3 males (specimen PT-1037 figured), Mt Bartle Frere,

0.5 km N of South Peak, 1500m, 6-8 Nov 1981, Earthwatch-Qld Mus.

(NMV).

Diagnosis. In lateral view, males of Hydrobiosella exilatis can

be distinguished from those of other species in the group by the

slender and upturned distal half of segment X; in dorsal view,

very flat and appearing broad, length about 1.4 times maximum

width, and long and slender harpago of the inferior appendages.

Description. Wings similar to H. letti (Fig. 17), length of

forewing: male 6.4-7.4 mm.

Male. Segment IX in lateral view, length about 1.3-1.4 times

width, sub-rectangular, slightly projecting basally (Fig. 31),

with a shallow, wide V-shaped notch medially on distal margin

(Fig. 32). Segment X length about 1.3-1.4 times width, in dorsal

view, ‘tongue-shaped’, broad-based, tapered distally (Figs 30,

31), apex broadly rounded (Fig. 30); in lateral view, narrowed

near middle, dilated slightly distally with bulbous apex (Fig.

31). Phallus generally tube-like; with a pair of long, slender,

slightly curved, lateral ‘parameres’ arising from the phallus

near middle (Figs 30, 31). Inferior appendages in lateral view,

with basal segment sub-rectangular, length about 1.8-1.9 times

maximum width, harpago slightly longer, length about 1.1

Two further species groups and new species among Australian Hydrobiosella Tillyard: new species from south-eastern

Australia (Trichoptera: Philopotamidae)

257

times length basal segment, more slender, length about 3.5

times maximum width, ‘dumb bell- shaped’, narrowed near

middle, broadly rounded apically (Fig. 31).

Female. Unknown.

Etymology. Exilatis - Latin for slender (harpago of inferior

appendages).

Remarks. Four male specimens of Hydrobiosella exilatis have

been collected from two adjacent localities in north-eastern

Queensland (latitudinal range 17° 16'- 17°24'S). Hydrobiosella

excilatas was cited in a checklist (Walker et ah, 1995) as

Hydrobiosella sp. nov. PT-1037.

Hydrobiosella bos sp. nov.

Figures 33-35, 43

Holotype. Male. Victoria, Eurobin Falls, Mt Buffalo Rd (about

36°42'S, 146°50'E), 3 Dec. 1982, A. Neboiss (NMV, T- 21484).

Paratype. Victoria. 1 male (specimen CT-553 figured), 2 females

(specimen CT-554 figured), collected with holotype (NMV).

Diagnosis. Males of Hydrobiosella bos can be distinguished

from those of other species in the group by the dorso-mesally

angled basal segment on the inferior appendages.

Description. Wings similar to H. letti (Fig. 17), length of

forewing: male 7.6-8.0 mm, female 8.9-9.4 mm.

Male. Segment IX robust, in lateral view, length about 1.4 times

width, projecting and tapered basally (fig. 34), with a shallow,

wide concavity medially on distal margin (Fig. 35). Segment X

broad-based, tapered distally (Figs 33, 34), in dorsal view,

‘tongue-shaped’, length about 1.8-1.9 times width, broadly

rounded apex (Fig. 33); in lateral view, straight, with slightly

bulbous apex (Fig. 34). Phallus generally tube-like; with a pair

of long, slightly curved, ‘parameres’ arising from the phallus

near base (Figs 33, 34). Inferior appendages in lateral view,

with basal segment sub-pentangular, length about 1.4 times

maximum width, harpago shorter, length about 0.7 times length

basal segment, more slender, length about 2.6 times maximum

width, sub-ovate, broadly rounded apically (Fig. 34).

Female. Genitalia typical of genus with a small acute projection

on sternite IX meso-distally (Fig. 43).

Etymology. Bos - Latin for buffalo (type locality-Mt Buffalo).

Remarks. Two males and two female specimens of

Hydrobiosella bos have been collected from the type locality

in north-eastern Victoria (latitude 36°42'S).

Hydrobiosella letti Korboot

Figures 17, 36, 37

Hydrobiosella letti Korboot, 1964: 36, figs 40-57. - Neboiss,

1987: 132, figs 7, 8. - Neboiss, 1986: 101.

Holotype. Male (not seen): New South Wales, Lett R. near

Lithgow, 25 Sep 1962, K. Korboot, T-6182 (QM).

Other material examined. Queensland, 1 male (CT-655), Morans

Ck above Morans Falls, Lamington Nat. Pk, 28.23°S 153.13°E, 16 Nov

2011, J. Mynott & M. Shackleton.

Diagnosis. Hydrobiosella letti can be distinguished from other

species in the group, especially H. bos, by the basal segment of

the inferior appendages not angled dorso-mesally and harpago

sub-triangular in lateral view.

Description. (Revised after Korboot, 1964; Neboiss, 1987).

Wings (Fig. 17), length of forewing: male 4.5 mm.

Male. Segment IX robust, in lateral view, length about 1.4 times

width, projecting and tapered basally (Fig. 37). Segment X

broad-based, tapered distally (Figs 36, 37), in dorsal view,

‘tongue-shaped’, length about 1.8-1.9 times width, broadly

rounded apex (Fig. 36); in lateral view, straight, with slightly

bulbous apex (Fig. 37). Phallus generally tube-like; with a pair of

long, slightly curved, ‘parameres’ arising from the phallus near

base (Figs 36,37). Inferior appendages in lateral view, with basal

segment sub-pentangular, length about 1.4 times maximum

width, harpago shorter, length about 0.7 times length basal

segment, more slender, length about 2.6 times maximum width,

sub-triangular, tapered to broadly rounded apex (Fig. 37).

Female. Unknown.

Remarks. A single male specimen of Hydrobiosella letti is

known from the type locality in central-eastern New South

Wales (latitude 33°24'S). Korboot’s (1963, fig. 45) and Neboiss’s

(1987, figs 7, 8) figures have been redrawn to allow direct

comparisons and to accompany the description that is revised

in light of new interpretations of Hydrobiosella genitalic and

wing structures. Neboiss (1987) commented on the condition of

the type specimen and errors in Korboot’s original description

and that the vial containing the remaining body parts had two

locality labels: Lett River via Lithgow and Montville,

Queensland. The recently collected male specimen from

Lamington National Park in south-eastern Queensland is here

referred to H. letti, although there are several minute differences

in the genitalia (latitude 28°14'S).

Acknowledgements

I thank the then Department of the Environment and Water

Resources, in particular Australian Biological Resources

Study (ABRS) for providing a grant to undertake this work,

the late Dr Arturs Neboiss, and Dr Alice Wells for providing

access to the specimens and, together with John Dean, for

helpful advice on earlier drafts of this manuscript. The referee

is thanked for constructive comments on this manuscript. I am

indebted to John Dean and Ros St Clair for technical assistance

with scanning the figures and moral support during the project.

References

Blahnik, R.J. 1998. A Revision of the Neotropical Species of the

Genus Chimarra, Subgenus Chimarra (Trichoptera:

Philopotamidae). Memoirs of the American Entomological

Institute 59: 1-318.

Blahnik, R.J. 2005. Alterosa, a new caddisfly genus from Brazil

(Trichoptera: Philopotamidae). Zootaxa 991: 1-60.

Cartwright, D.I. 1997. Preliminary guide to the identification of late

instar larvae of Australian Ecnomidae, Philopotamidae and

Tasimiidae (Insecta: Trichoptera). Identification guide no. 10. Co¬

operative Research Centre for Freshwater Ecology, Albury. 33 pp.

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Cartwright, D.L 2010. Studies of Australian Hydrobiosella Tillyard: a

review of the Australian species of the Hydrobiosella bispina

Kimmins group (Trichoptera: Philopotamidae). Memoirs of

Museum Victoria 67: 1-13.

Cartwright, D.l. 2012a. Studies of Australian Hydrobiosella Tillyard:

two new Australian species from North Queensland (Trichoptera:

Philopotamidae). Australian Entomologist 39: 109-116.

Cartwright, D.l. 2012b. Studies of Australian Hydrobiosella Tillyard:

a review of the species of the Hydrobiosella waddama Mosely

group (Trichoptera: Philopotamidae). Proceedings of the Royal

Society of Victoria 124: 117-132.

Espeland, M. and Johanson, K. A. 2007. Revision of the New

Caledonian Hydrobiosella (Trichoptera: Philopotamidae) with

description of five new species. Pp 91-102, In: Bueno-Soria, J.,

Barba-Alvarez, R. and Armitage, B. (eds). Proceedings oftheXIIth

International Symposium on Trichoptera , The Caddis Press.

Holzenthal, R.W., Blahnik, R.J., Prather, A.L. and Kjer, K.M. 2007.

Order Trichoptera Kirby, 1813 (Insecta), Caddisflies. Zootaxa

1668: 639-698.

Korboot, K. 1964. Four new species of caddis-flies (Trichoptera) from

eastern Australia. Journal of the Entomological Society of

Queensland. 3: 32-41.

Jacquemart, S. 1965. Contribution a la connaisance de la faune

Trichopterologique de la Tasmanie et de la Nouvelle-Zealande.

Bulletin Institut royal des Sciences naturelles de belgique 41: 1-47.

Morse, J. C. (ed.). 1999. Trichoptera World Checklist. [Accessed 11

May 2011.] Available from URL: http://entweb.clemson.edu/

database/trichopt/index.htm, effective 27 March 1999.

Mosely, M.E. and Kimmins D.E.1953. The Trichoptera (Caddis-flies) of

Australia and New Zealand. London: British Museum (Natural

History). 550 pp.

Neboiss, A. 1977. A taxonomic and zoogeographic study of Tasmanian

caddis-flies (Insects: Trichoptera). Memoirs of the National

Museum of Victoria 38: 1-208.

Neboiss, A. 1982. The caddis-flies (Trichoptera) of south-western

Australia. Australian Journal of Zoology 30: 271-325.

Neboiss, A. 1986. Atlas of Trichoptera of the SW Pacific-Australian

Region. Dr W. Junk, Dordrecht. 286 pp.

Neboiss, A. 1987. Identity of species of Trichoptera described by K.

Korboot 1964-65 (Insecta). Memoirs of the Museum of Victoria 48:

131-140.

Neboiss, A. 2003. New genera and species, and new records, of

Tasmanian Trichoptera (Insecta). Papers and Proceedings of the

Royal Society of Tasmania 136: 43-82.

Suter, P., Dean J, Cartwright D, Sutcliffe K, Davies P, Pinder A and

Bryce, C. 2006. Habitat Profiles of Selected Australian Aquatic

Insects. Australian Biological Resources Study. Accessed 30 July

2010 at http://www.environrnent.gov.au/biodiversity/abrs/

publicationslelectronic-bookslaquatic-insects.html.

Tillyard, RJ. 1924. Studies of New Zealand Trichoptera or caddis flies

No. 2. Descriptions of new genera and species. Transactions of the

New Zealand Institute 55: 285-314.

Walker, K., Neboiss, A., Dean, J. and Cartwright, D. 1995. A preliminary

investigation of the Caddis-flies (Insecta: Trichoptera) of the

Queensland Wet Tropics. Australian Entomologist 22: 19-31.

Memoirs of Museum Victoria 69:259-267 (2012)

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

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

Two new species of Photonectes (Teleostei: Stomiidae) from the Indo-Pacific, and a

re-examination of P . achirus

Flynn, A. J. 1 - 2 - 3 and Klepadlo, C. 4

1 The University of Queensland, School of Biomedical Sciences/Queensland Brain Institute, St. Lucia 4071, Queensland,

Australia (adrian.flynn@uqconnect.edu.au)

2 CSIRO Marine and Atmospheric Research, PO Box 1538, Hobart 7001, Tasmania, Australia

3 Museum Victoria, Ichthyology, PO Box 666, Victoria 3001, Australia

4 Marine Vertebrate Collection, Scripps Institution of Oceanography, University of California at San Diego, 9500 Gilman

Drive, La Jolla, California, USA 92093-0208. (cklepadlo@ucsd.edu)

Abstract Flynn, A. J.and Klepadlo, C. 2012. Two new species of Photonectes (Teleostei: Stomiidae) from the Indo-Pacific, and a

re-examination of P. achirus. Memoirs of Museum Victoria 69: 259-267.

Two new species of the mesopelagic fish genus Photonectes (family Stomiidae) from the Indo-Pacific are described.

Both are referred to the subgenus Photonectes because they lack pectoral fins, have no fleshy tissue on their dorsal- and

anal-fin rays, have a pelvic-fin insertion closer to the caudal-fin base than the snout, and have IP photophores located on the

isthmus at a position midway along the mandible, posterior to the mandibular symphysis. Photonectes waitti sp. nov.,

known from the tropical western and central Pacific Ocean, northwestern Coral Sea and north Indian Ocean, is most similar

to P. coffea , differing from it by the number of PV photophores (24—25 vs. 29—31), gill filaments reduced (vs. greatly

elongate), jaw teeth long and short canines (vs. short canines only), and barbel length less than (vs. equal to or greater than)

head length. Photonectes paxtoni sp. nov., known from the tropical western Pacific Ocean, off Papua New Guinea, has a

short barbel, less than head length, ending in a very large bulb without any terminal filaments or appendages, and different

photophore numbers compared to other members of the subgenus: VAV 15—16 (vs. 10—15), AC 13 (vs. 9—12), and IV 29

(vs. 30—46). Photonectes achirus is re-examined with respect to previously unreported dark pigment on the head.

Keywords Photonectes, Stomiidae, new species, Indo-Pacific

Introduction

The Stomiidae is a very speciose family of midwater fishes,

currently comprising approximately 280 species in 27 genera

(Nelson, 2006; Eschmeyer, 2011; Froese and Pauly, 2011).

The genus Photonectes has 25 nominal species, 16 of which

are regarded as valid species (Eschmeyer, 2011; Froese and

Pauly, 2011).

In 2009, 21 specimens of an apparently undescribed

species of Photonectes were collected from the equatorial

western Pacific Ocean, in the vicinity of Howland and Baker

islands. In 2010, an additional specimen was collected in the

Coral Sea off Cairns, Australia. The examination of material

from museum collections revealed two more specimens,

extending the range of the species into Hawaiian waters and

the Gulf of Aden. The search for additional material resulted

in the discovery of another undescribed species collected in

1969 by the Australian Museum off Papua New Guinea. Both

species are members of the subgenus Photonectes based on the

lack of pectoral fins, their dorsal and anal fins not enveloped

with fleshy tissue, their pelvic-fin insertion located closer to

caudal-fin base than to the snout tip, and their IP photophores

located on the isthmus at a position midway along the

mandible, posterior to the mandibular symphysis (Regan and

Trewavas, 1930; Morrow and Gibbs, 1964; Klepadlo, 2011).

Descriptions of these two species are provided herein.

Besides photophores and barbels, other dermal features

that aid in identification of species of Photonectes include the

presence or absence of white luminous tissue patches on the

head and body and the presence or absence of ventro-lateral

blue luminous tissue markings on the body, along the isthmus

and inside the mouth.

Specimens of P. achirus were collected along with

specimens of the former undescribed species in the equatorial

western Pacific Ocean in the vicinity of Howland and Baker

islands. While examining museum specimens of P. achirus,

patches of dark pigment were observed on the head that were

previously unreported (e.g., in Regan and Trewavas, 1930;

Beebe and Crane, 1939; Morrow and Gibbs, 1964; Klepadlo,

2011). The patches of dark pigment were only observed on faded

museum specimens, usually rusty-brown in colour, that had

been preserved for a long period of time. The dark pigment was

not detectable in freshly-caught specimens that were usually

260

A. Flynn & C. Klepadlo

black or black-brown in color. Faded museum specimens of

other Photonectes species were examined, but these markings

were found only on specimens of P. achirus. A re-examination

of P. achirus is provided on the basis of this new finding.

Methods

Values for selected morphometric and meristic features in the

new species are given in Table 1. Measurements were made

with vernier calipers to the nearest 0.1 mm and include:

standard length (SL), head length (HL), barbel length, and

lengths from snout to pelvic fin (Sn-V), from pelvic fin to vent

(V-vent), and from vent to base of caudal fin (vent-C). Percent

of SL (% SL) was calculated for P. waitti and P.paxtoni and %

SL values for P. achirus were derived from Morrow and Gibbs

(1964) and Froese and Pauly (2011). Descriptions of teeth are

given and these were separated as premaxillary, maxillary,

mandibular, vomerine, palatine and basibranchial; maxillary

teeth are further divided into ’’erect” (anterior canines) and

’’oblique” (posterior series of very small inclined teeth).

Photophore terminology (Figure 1) follows Morrow and Gibbs

(1964), Harold (2003), and Klepadlo (2011). In addition to

diagnostic counts of the primary photophores, descriptions are

given of small secondary photophores that occur as clusters or

form lateral bars in the genus Photonectes. Further,

descriptions are made of two opercular (OP) photophores: the

pre-opercular photophore (PRO) at the posteriodorsal margin

of the operculum and the subopercular photophore (SO)

located in a cup-like depression ventral to the PRO (notation

follows Weitzman, 1974: Table 1).

Gill-filament length (Klepadlo, 2011: Fig. 2) on first

branchial arch is defined as ”l st reduced” (less than one-half

arch depth), ’’normal” (longer than arch depth but not extending

beyond opercular opening) or ’’very long” (much greater than

depth or arch and extending beyond opercular opening).

Most specimens were collected with a neuston net. Other

collection methods included Isaacs-Kidd midwater trawl

(IKMT), IKMT modified to collect plankton (IKPT)

(Williamson and McGowan, 2009), rectangular midwater trawl

(RMT) and a bottom trawl. Collection depths are given in meters

(m), fathoms (fm) or meters-wire-out (mwo). Time of collection

listed for P. waitti sp. nov. is given in UTC/GMT. Notes on the

color (under visible light) of fresh specimens of P. waitti are

based on observations made by the first author in the field.

Material examined is in the collections of the Australian

Museum (AMS), Museum of Victoria (NMV), Scripps Institution

of Oceanography (SIO) and Smithsonian Institution (USNM).

Photonectes waitti new species

Figures 2a,b, 5; Table 1

Holotype: NMV A 30913-001 (71.4mmSL),0°34.836'-35.329'N,

176°56.432’—49.956’W, neuston net, 0—0.5 m depth, R/V Seward

Johnson, Catalyst II expedition, 06 March 2009, 14:04—15:05.

Paratypes: AMS 1.49494-010 (48.7 mm SL), 16°34.25'S,

147°08.01'E, 11 December 2010, 19:43 -20:43, RMT8 trawl, 330 m

depth; NMV A 30908-001/-002 (2: 22.5-32.6 mm SL), 0°21.243'-

21.257'N, 176°50.117'-49.575'W, 23 February 2009, 12:17-13:17,

neuston, 0-0.5 m depth; NMV A 30915-001 (1: 32.1 mm SL) and SIO

11-301/-302 (2: both 48.5 mm SL), 0°27.568'-27.658'N, 176°51.848’-

50.574’W, 28 February 2009, 12:32—13:32, neuston, 0—0.5 m depth;

NMV A 30906-001/-002/-003/-004/-005 (5: 31.6-51.0 mm SL),

collected with holotype; NMV A 30908-001/-002 (2: 22.5-32.6 mm

SL), 0°21.243 —21.257'N, 176°50.117'-49.575 , W, 23 February 2009,

12:17-13:17, neuston, 0-0.5 m depth; NMV A 30907-001/-002/-

003/-004 (4: 25.2-30.7 mm SL), 0°54.289'N, 177°09.476'W, 02 May

2009, neuston net, 0—0.5 m depth; SIO 11-300 (37.6 mm SL),

0°13.518'—13.212’N, 176°46.428'-45.634'W, 19 February 2009,

07:43—08:43, neuston, 0—0.5 m depth; SIO 11-303 (2: 25.4—41.5 mm

SL), 0°53.000'—50.586’N, 176°57.332'-56.657'W, 20 March 2009,

12:59-13:57, neuston net, 0-0.5 m depth; USNM 402774 (32.3 mm

SL), 01°19.045'—18.335'N, 176°47.913'-47.673'W, 19 April 2009,

RMT trawl M205, 1200 m depth; USNM 300149 (23.4 mm SL),

12°40’N, 51°13'E, 1 m plankton net, captured at surface, R/V Anton

Bruun Cr. 5, sta. 287A, IIOE Expedition, 3 February 1964; USNM

300199 (26 mm SL), 21°20'-30'N, 158°20'-30'W, 26 September

1973, 3 m IKMT, 0-110 m depth, T. Clarke 73-9-33; USNM 300201

(24.8 mm SL), 21°20'-30'N, 158°20'-30'W, 30 August 1973, 03:18-

05:15,3 m IKMT 0-350 m depth, T. Clarke 73-8-31.

Diagnosis: Differs from other species in the subgenus

Photonectes in the following combination of characters:

dorsal-fin rays 11—14, anal-fin rays 12—16; IV photophores

38—39, PV photophores 24—25; absence of blue luminous

tissue; length of gill filaments on first branchial arch reduced,

less than depth of gill arch; and barbel with two large bulbs in

series. Few species of Photonectes have more than one large

bulb or luminous area on the barbel (secondary photophores

excluded). Above the main bulb in P. mirabilis are small bulbs

on the stem and a tiny terminal bulb. P. phyllopogon has a

single bulb and a terminal luminous appendage (Morrow and

Gibbs, 1964). P. barnetti has a large pale area along the stem

below the bulb (luminescent colour uncertain) (Klepadlo,

2011). P. coffea has two bulbs in series without any terminal

bulb (Klepadlo, 2011). For P. waitti there are two large bulbs

in series and a tiny bulb at the end of the filament. Photonectes

waitti differs from P. mirabilis and P. phyllopogon in having

11— 14 dorsal-fin rays (vs. 16—17 and 20—23, respectively),

12— 16 anal-fin rays (vs. 19—20 and 22—25, respectively), and

38—39 IV photophores (vs. 33—34 and 30—31, respectively).

It differs from P. coffea in having gill filaments on first gill

arch reduced (vs. very long, extending beyond gill opening),

and PV photophores 24—25 (vs. 29—31). It differs from P.

achirus, P. caerulescens, and P. mirabilis in lacking blue

luminous tissue.

Description: Body elongate, 20.9—71.4 mm SL; depth about

eight times into length. HL 3.2—8.7 mm (11.0—18.1% SL,

mean 15.3%); Sn-V 13.3—47.9 mm (58.5—67.4% SL, mean

62.1%); V-vent 3.4-10.5 mm (13.9-21.4% SL, mean 18.2%);

vent-C 4.3—13.0 mm (15.6—23.4% SL, mean 19.9%) (see Table

1). Eye 0.8—1.9 mm (17.2-29.3% HL, mean 23.5%). Opercle

concave dorsally and lobate posteriorly. Gill filaments on first

branchial arch reduced, length less than arch depth; tips of

filaments dark. Color of body in life dark brown to black. On

fresh specimens or undamaged preserved specimens, narrow

black lateral bands in line with PV, VAV and AC photophores.

On freshly caught specimens, these lateral bands could be seen

to harbor minute secondary photophores and luminous patches

(described further below).

Two new species of Photonectes (Teleostei: Stomiidae) from the Indo-Pacific, and a re-examination of P. achirus

261

Dorsal-fin rays 12 (11—14, rarely 15—16); anal-fin rays

14 (12—16, rarely 17—19); pelvic-fin rays 7 (rarely 6);

pectoral fins absent. Dorsal and anal fins not covered with

black fleshy skin. Pelvic fins inserted closer to caudal fin than

to snout (33.1—42.4% SL vs. 58.6—67.4% SL); longest ray

extending at least to anus (tips broken). Caudal fin forked and

elongate; ventral lobe longer than dorsal lobe. All fin-rays

covered with melanophores extending halfway along the

length; membranes clear.

Photophores: IV 38—39, rarely 35—37 (IP 12, rarely 11 or

13; PV 24-25, rarely 22, 23 or 26); VAV 13-14, rarely 12 or

15; AC 11-12, rarely 13; OA 38-39, rarely 35-37 (OV

24—26, rarely 23; VAL 12—13, rarely 14—15), last 1 to 2

photophores over anal-fin base; BR-7, rarely 6. IP series

beginning about halfway along isthmus length, approximately

opposite BR-7; space between IP-8 and IP-9 about twice space

between IP-1 and IP-2 making series 8 + 4. Anterior end of AC

series raised, approximately even with last VAL photophore,

tapering downward over anal fin base and ending along

ventro-lateral caudal peduncle. Secondary photophores on

head and body, mainly concentrated in vertical lines from

dorsum to each PV, VAV and AC photophore; also between IV

and OA series and along ventrum in rather horizontal lines; no

secondary photophores on any fin-ray. Postorbital organ ovoid,

0.7—0.9 mm (12.3—18.8% HL, mean = 14.6%; skin flap over

posterior end in NMV A.30907-003. One ovoid PRO

photophore; one smaller circular SO photophore. SO

photophore directed ventrally and placed in a cup-like

structure. PRO and SO photophores of fresh specimens reddish

pink under visible light. Blue luminous tissue and dark

markings absent.

Teeth caniniform, long and short; premaxillary teeth

longest, needle-like. Premaxillary teeth 5—8; 4—8 maxillary

teeth erect and 4—10 oblique; mandibular teeth 11—25;

vomerine teeth in 2 to 3 pairs, lateral teeth longest; palatine

teeth in 2 pairs, about equal length; basibranchial teeth in 2 to 4

pairs (anterior and posterior pairs separated by gap, 1 to 2 pairs

followed by 1 to 2 pairs; posterior tooth in each group longest).

Barbel shorter than head, 2.3-S.5 mm (59.0—85.4% HL,

mean 74.0%), with two main bulbs and a fine terminal filament.

One specimen (SIO 11-304) with very tiny bulb at distal end of

terminal filament. Stem pigmented, no secondary photophores;

first segment (-45% of barbel length) tapering into first bulb

with pigment cup-like around bulb base. First bulb ovoid

(-18% of barbel length), width about three times stem width,

tapering into short segment of stem (-9% of barbel length)

followed by second bulb. Section of stem between bulbs with

melanophores increasing in concentration distally. Second

bulb elongate (-18% of barbel length), narrow, about twice

stem width, and with melanophore-line from stem extending

length of bulb onto short terminal filament (-9% of barbel

length). In freshly-caught specimens bulbs lavender to pink

under white light.

Distribution: Known from tropical western and central Pacific

Ocean, northwestern Coral Sea and Indian Ocean (Gulf of

Aden) at depths ofO—1200m.

Etymology: Named in recognition of Theodore (Ted) Waitt, the

founder of the Waitt Family Foundation and the Waitt Institute.

The Waitt Institute sponsored and directed the expedition of the

R/V Seward Johnson to the equatorial western Pacific Ocean

(Catalyst II expedition) during which this species was collected.

Figure 1. (A) Photophore notation: post - postorbital organ; VAV - ventrally, from pelvic fin base to anal fin origin; AC - from anal fin origin to

caudal fin base; OV - laterally, from pectoral fin origin to pelvic fin origin; VAL - laterally, from pelvic fin origin to anal fin origin or slightly

beyond; (B) and (C) IP - along isthmus to pectoral fin base commencing at mandibular symphysis or posteriorly halfway along isthmus length,

respectively.

262

A. Flynn & C. Klepadlo

Figure 2 . Photonectes waitti new species a) NMV A.30906-005,51.0 mm, paratype; b) lateral view of head and barbel, NMV A.30913-001, 71.4

mm, holotype. Photograph by NMV.

Two new species of Photonectes (Teleostei: Stomiidae) from the Indo-Pacific, and a re-examination of P. achirus

263

Figure 3. Photonectespaxtoni new species, holotype, AMS 1.1972043, 23.4 mm SL. Photograph by NMW.

Figure 4. Photonectes achirus, AMS 1.19739-018, 32.3 mm, lateral view of head of showing dark marking at upper operculum (Photograph by

C. Klepadlo).

264

A. Flynn & C. Klepadlo

130° 160° W 140" W 120" W 100" W 30° W 60° W 40" W

—I-1-

j Area of Inset below right

Area of Inset below left

ro’N-

o°o-

■A

▲

Howland Island

Baker Island

-1-r

177°0'W 176

0'W

23°O f

22 6 0 f

21 °0 f

2Q"ET

19"0 T M-

13"0 T N-

1 - h

^ - f

'w 1 isro-w

i r

15S°0'W

■30° N

-10° N

10° S

■30° S

■50° S

■70° S

60"0’E

30°0'E

100°0’E

120"0'E 14D°0'E

30"0'N

10 6 O f N

10"Q'S

30°0'S

Figure 5. Distribution of Photonectes waitti (black square), P. paxtoni (black circle), P. achirus (black triangle; white triangle = literature

records). Symbols may represent more than one record

Two new species of Photonectes (Teleostei: Stomiidae) from the Indo-Pacific, and a re-examination of P. achirus

265

Table L Selected meristics and morphometries of Photonectes waitti , P. paxtoni , and P. achirus. Values for holotypes followed by range for

paratypes in parentheses where they vary from the holotype. Values for P. achirus from Morrow and Gibbs (1964) and Froese and Pauly (2011).

Character

Photonectes waitti

(n=24)

Photonectes paxtoni

(n=2)

Photonectes achirus

Standard Length (mm)

71.4 (20.9-74.0)

23.4 (22.4)

20.0-87.0

Fin Rays:

Dorsal

12 (11-16)

16 (18)

14-16

Anal

14(11-19)

18 (18)

14-19

Pelvic

7 (6-7)

Photophores:

12(11-13)

8-11

23 (22-26)

22-28

VAV

14 (12-15)

16 (15)

11-13

11 (10-13)

10-11

26 (21-26)

19-22

VAL

13 (12-15)

11-12

7 (6-8)

6-8

%SL:

Head

12.9 (12.1-18.9)

17.1 (15.6)

15.4-18.7

Sn-V

66.9 (58.5-67.4)

61.1 (61.6)

58.2-73.2

V-vent

14.7 (14.9-21.3)

15.4 (15.2)

8.7-21.1

Vent-C

18.2 (15.6-23.4)

23.5 (23.2)

15.6-22.8

Remarks: Photonectes waitti is a widespread tropical open-

ocean species. Twenty-one specimens were collected in 2009

from the equatorial western Pacific Ocean. One specimen was

collected in 2010 from Australian waters in the Coral Sea.

While examining older (40+ years) museum specimens labeled

Photonectes sp., three additional specimens were found that

extended the distribution of P. waitti into the Central Pacific, in

the vicinity of Hawaii, and the northwestern Indian Ocean

(Gulf of Aden).

Photonectes paxtoni new species

Figures 3, 5; Table 1

Holotype: AMS 1.1970-243 (23.4 mm SL), 05°05'S, 145°56'E, off

Madang, Papua New Guinea, 27 October 1969, 2 m IKMWT, 135 m

depth.

Paratype: AMS 1.19727-023 (22.4 mm SL), 05°05'S, 145°54'E, off

Madang, Papua New Guinea, 27 October 1969, 2 m IKMWT, 130 m

depth.

Diagnosis: Photonectes paxtoni differs from other members of

the subgenus Photonectes in the following combination of

characters: barbel short, with a very enlarged ovoid terminal

bulb lacking terminal appendages; absence of blue luminous

tissue; length of gill filaments on first branchial arch reduced,

less than arch depth; and VAV photophores 15—16 (v.v. 10—15),

AC photophores 13 (v,v. 9—12) and IV photophores 29 (vs. SO¬

TO). The bulb is remarkably large and plain (without filaments).

The enlarged bulb is similar to P. ovibarba (synonym of P.

braueri). However, P. paxtoni has no pectoral fins (vs. present,

with 2 rays each), IV photophores 29 (vs. 32—33), AC photophores

13 (vs. 10—12), and the bulb lacks any terminal appendage (vs.

bearing a small ovoid appendage). Another species with a large

bulb, P. fimbria (synonym of P. parvimanus), can also be

eliminated based on the following: IV photophores 43—49 (vs.

P. paxtoni with 29), IP photophores insert near mandibular

symphysis or no gap (vs. insert halfway along the isthmus, or

with gap), thick skin over dorsal- and anal-fin rays (vs. thick skin

absent), and bulb with terminal flap increasing with growth (vs.

no flap or terminal appendage; growth change unknown).

Description: Body elongate, 22.4—23.4 mm SL; depth about

seven times into length. HL 3.5—4.0 mm (15.6—17.1% SL);

Sn-V 13.8-14.3 mm (61.1-61.6% SL); V-vent 3.4-3.6 mm

(15.2-15.4% SL); vent-C 5.2-5.5 mm (23.2-23.5% SL) (see

Table 1). Eye 1.0—1.1 mm (25.0—27.5% HL). Opercle lobate,

slightly concave dorsally. Gill filaments on first branchial arch

reduced, length less than arch depth; tips of gill filaments un-

pigmented. Color of body rusty-brown in preservative, assumed

black in life.

Dorsal-fin rays 16—18; anal-fin rays 18; pelvic-fin rays 7;

pectoral fins absent. Dorsal and anal fins not covered with

black fleshy skin; fin-rays covered with minute white luminous

spots; membranes clear. Pelvic fins inserted closer to caudal

fin than to snout tip (38.4—38.9% SL vs. 61.1—61.6% SL);

longest ray extending to anal-fin origin. Caudal-fin rays

broken; fin assumed forked.

Photophores: IV 29 (IP 8; PV 21); VAV 15-16 (last 2 to 3

over anal-fin base); AC 13; OA 34 (OV 20; VAL 14, last two

photophores over anal-fin base); BR 8. IP series beginning

posteriorly about halfway along isthmus length, opposite BR-8;

photophores evenly spaced. Anterior end of AC series beginning

on same level as last VAV. Secondary photophores scattered

over head and body, in clusters along dorsum, tapering ventrally

to between each OA photophore, continuing ventrally between

each IV photophore, and in clusters along ventral surface; none

266

A. Flynn & C. Klepadlo

on any fin rays. Postorbital organ ovoid, elongate, about equal to

eye diameter. One PRO and one SO photophore on operculum,

and one postorbital photophore. Specimen AMS 1.1927-023

with a pair of white luminous spots on snout between nostrils.

Blue luminous tissue and dark markings absent.

Teeth caniniform, long and short; premaxillary teeth

longest, needle-like. Premaxillary teeth 4; 5—7 maxillary teeth

erect and 8 oblique; mandibular teeth 12—15; vomerine teeth

one pair; palatine teeth absent; basibranchial teeth 4 (one pair

anteriorly and two single teeth midlength). Vomerine teeth long,

length equal to longest premaxillary tooth.

Barbel short, 1.5—1.8 mm (42.9—45.0% HL), with very

large ovoid bulb. Stem short, ~1.0 mm (~25% HL), pigment

tapering anteriorly onto base of bulb in a small V-shape; no

secondary photophores on stem. Bulb large and simple, with no

appendages or terminal filaments; width ~1.0 mm. Color of bulb

in life unknown.

Distribution: Known only from type locality off Madang,

Papua New Guinea; depth 130—135 m.

Etymology: The name recognises Dr. John Paxton for his many

contributions to the study of mesopelagic fishes and for his

encouragement to the authors.

Remarks: Photonectes paxtoni is currently known from two

specimens from shallow collections (depth 130—135 m). They

were located among 46-year-old museum specimens labeled

Photonectes sp.

Photonectes achirus Regan and Trewavas, 1930

Figures 4, 5; Table 1

Material examined: AMS 1.19739-018 (32.3 mm SL), 07°09.0'S,

148°52.0'E (western Solomon Sea), 110 m depth, 7 November 1969;

AMS 1.19753-037 (2: 25.3-30.9 mm SL), 05°51.0'S, 147°20.0'E

(Papua New Guinea, Vitiaz Straits), IKMT, 0—110 m depth, 4

November 1969; AMS 1.24859-002 (78.0 mm SL), 33°43.0'-40.0'S,

152°03.0’—05.0’E (Australia, off Sydney, NSW), bottom trawl,

0-1135 m depth, 16 October 1984; NMV A 30906-006 (46.4 mm

SL), 0°34.836’—35.329'N, 176°56.432'-49.956'W, neuston net,

0-0.5 m depth, 6 March 2009; NMV A 30917-001 (46.7 mm SL),

0° 14.213'—14.354'N, 176°45.843'-45.64rW, neuston net, 0-0.5m

depth, 19 February 2009; SIO 11-305 (72.2 mm SL), 01°11.503'-

10.113'N, 176°45.152'—46.050'W, RMT, 600 m depth, 26 April 2009;

SIO 11-306 (83.5 mm SL), 0°20.264'-18.657'N, 177°02.297'-

00.516'W, RMT, 600 m depth, 9 May 2009; SIO 70-333 (33.5 mm

SL), 19°11.0'—04.8’N, 125°12.7'45.0'E, 10 ft IKMT, 0-2000 m

depth, 13 September 1970; SIO 88-194 (21.4 mm SL), 24°40.5’N,

76.16'W (Exuma Sound), IKMT, 200 m depth, 14 July 1986; SIO 88-

197 (23.0 mm SL), 25°28.0'N, 78°07.3’W (Tongue-of-the-Ocean),

IKMT, 200 m depth, 8 January 1987; USNM 300205 (36.7 mm SL),

21°10’N, 158°10’W, IKMT, 0-775 m depth, 8 November 1974,

12:35-17:12, T. Clarke 74-11-4.

Diagnosis: Photonectes achirus differs from other species in

the subgenus Photonectes in the following combination of

characters: presence of blue luminous tissue in a band

extending from pectoral region to pelvic fins, with short

transverse streaks between OV photophores, and in patches on

sides of isthmus, under lower jaw and above end of maxillary;

length of gill filaments very long, extend beyond opercular

opening; and barbel shorter than head length, with small bulb

ending in a terminal appendage with a tiny bulb at tip.

Re-Examination: Body elongate, largest recorded specimen

87.0 mm SL. Morphometric and meristic values for four

specimens collected from waters around Howland and Baker

islands were in agreement with Morrow and Gibbs (1964):

body depth 11.8—16.7% SL; head 15.4—18.7% SL (mean

14.8%); Sn-V 58.2-73.0% SL (mean 65.6%); V-vent 8.7-

21.1% SL (mean 14.4%); vent-C 15.6-22.8% SL (mean 20.1%).

Eye 7.7—28.8% HL (mean 20.7%). Opercle slightly concave

dorsally. Gill filament very long, feathery; extending slightly

beyond gill cover; tips of filaments uncolored. Color of body

black, fading to rusty brown in preservative.

Dorsal-fin rays 14—16; anal-fin rays 14—19; pelvic-fin

rays 7; pectoral fins absent. Dorsal and anal fins not covered

with black fleshy skin. Pelvic fins inserted closer to caudal

fin than to snout tip; longest ray extending posteriorly to vent.

Caudal fin forked; ventral lobe longer than dorsal lobe. All

fin rays covered with melanophores along full length;

membranes clear.

Photophores: IV 31-36 (IP 8-11 + PV 22-28); VAV

11-13; AC 10-11; OA 30-35 (OV 19-22; [rarely 23] + VAL

11—12 [rarely 9—10 or 13—15], last two photophores over

anal-fin base); BR 6—8. IP series begins about halfway along

isthmus length, approximately opposite posterior most BR; IP

photophores noticeably smaller than body photophores.

Anterior end of AC series raised, about one photophore

diameter above last VAV, gently tapering to ventral profile.

Secondary photophores very small, scattered over head and

body; most densely clustered along dorsum; no secondary

photophores on any fin-ray. Postorbital organ ovoid, with dark

cap of melanophores along dorsal margin or with cluster of

melanophores at anterior margin; length up to 30% HL.

Clusters of small white luminous spots on opercle. Blue

luminous patch under OV-1 to OV-2; two patches along

isthmus from symphysis to BR-1; pair of spots between eyes

and small spots dorsal to occipito-vertebral articulation

(“nape”); one pair anteriorly inside mouth; midventral stripe

from below pectoral fins to pelvic fins, with short transverse

streaks alternating with serial photophores.

Teeth caniniform, long and short; premaxillary teeth

longest, needle-like. Premaxillary teeth 7—12; 6—8 maxillary

teeth erect and 7—10 oblique; mandibular teeth 17—22;

vomerine teeth 1 or 2 pairs; palatine teeth 1 pair; basibranchial

teeth 4 pairs (anterior and posterior pairs separated by gap: 1

or 2 pairs followed by 2 or 3 pairs).

Barbel shorter than head length (70.0—80.0% HL). Stem

covered with melanophores, terminating in a small bulb,

slightly wider than stem width, with a long slender terminal

appendage ending in a small bulb at tip.

Comment on Pigmentation: In older, faded museum specimens

small to large dark semi-circular patches on snout and upper

opercle (Fig. 4; see Discussion). The dark patches were observed

on older faded specimens of P. achirus (AMS 1.19739-018, SIO

70-333, SIO 88-194, SIO 88-197) on the snout between the

nostrils and on the upper opercle (see Fig. 4). They are circular

to slightly ovoid in shape, contain darker spots and are bordered

Two new species of Photonectes (Teleostei: Stomiidae) from the Indo-Pacific, and a re-examination of P. achirus

267

by heavy pigment. Examination of more recently collected

(unfaded) specimens of P. achirus (SIO 11-305, SIO 11-306) did

not reveal the colouration observed in older (faded) specimens.

Several specimens did have clusters of secondary photophores

at the upper opercular margin that are probably associated with

the dark spots. However, other areas on the body with secondary

photophores were not surrounded by any dark patches. One

specimen of P. achirus (USNM 300205) had noticeable clusters

of white luminous tissue on the nape and along the opercular

margin with no dark patches surrounding the clusters; snout

tissue was damaged. The skin on the head of P. cf. gracilis

(USNM 300157) is in very good condition with both secondary

photophores and a large cluster of white luminous tissue at the

upper opercular margin; there is no indication of dark patches.

Other faded Photonectes were examined for dark patches: P.

albipennis (AMS 1.221809-030, SIO 73-149), P. braueri (AMS

1.20305-013), P. caerulescens (SIO 76-6), P. margarita (SIO 69-

354), P. mirabilis (AMS 1.19739-018), P. parvimanus (SIO 10-

177), P. paxtoni (AMS 1.19727-023, AMS I. 19702-043), P.

waiiti (USNM 300149, USNM 300201); none showed any trace.

The significance of the tissue is unknown; it is not noticeable in

fresh specimens nor is it reflective or photogenic. In reviews of

dragonfishes ascribed to the family Melanostomiatidae by

Beebe and Crane (1939) and Morrow and Gibbs (1964), neither

work mentions these dark patches. Whether it occurs only on P.

achirus is unknown at this time.

Distribution: Known from western Atlantic and Caribbean, north

Pacific near Hawaii and Tasman Sea at depths of 75—1400 m

(Clarke, 1974; Froese and Paul, 2011; Morrow and Gibbs, 1964;

Sutton and Hopkins, 1996) and now recorded from the

equatorial western Pacific Ocean in the vicinity of Howland

and Baker islands.

Comparative material examined

Photonectes albipennis (AMS 1.22809-030, SIO 73-149).

Photonectes braueri (AMS 1.20305013).

Photonectes caerulescens (SIO 76-6, USNM 256901).

Photonectes cf. gracilis (AMS 1.20941-010, USNM 300157).

Photonectes margarita (SIO 69-354).

Photonectes mirabilis (AMS 1.19739-018) [re-identified as P.

achirus].

Photonectes parvimanus (SIO 10-177).

Acknowledgements

Curatorial assistance was provided by H. J. Walker, Jr. (SIO),

M. McGrouther (AMS), and S. Jewett (USNM). The first

author’s research was supported by The University of

Queensland Research Scholarship and a Graduate School

Research Travel Grant from The University of Queensland.

We thank Ted Waitt and the Waitt Institute for the invitation to

take part in a voyage to the equatorial western Pacific Ocean

aboard R/V Seward Johnson (Catalyst II expedition) and this

expedition was supported by The University of Queensland

Deep Ocean Australia Project (ARC-linkage grant #

LP0775179 to Prof. Justin Marshall). Alan Goldizen and David

Wheeldon (The University of Queensland) made collections

of P. waitti and P. achirus on the second leg of the Catalyst II

expedition. We thank M. Gomon and D. Bray (NMV) for their

support throughout this project and for use of NMV facilities.

M. Gomon provided valuable comments on a draft of this

manuscript. We are grateful to J. Paxton (AMS) for bringing

the authors together and encouraging our research; R.

Rosenblatt, P. Hastings and H. J. Walker, Jr. (SIO) for support

and encouragement, and for use of laboratory facilities.

References

Beebe, W. and Crane, J. 1939. Deep-sea fishes of the Bermuda

Oceanographic Expeditions. Family Melanostomiatidae.

Zoologica, N.Y., 24(6): 65-238.

Clarke, T.A. 1974. Some aspects of the ecology of stomiatoid fishes in

the Pacific Ocean near Hawaii. Fisheries Bulletin 72(2): 337-351.

Eschmeyer, W.N. (ed.). 2011. Catalog of Fishes. World Wide Web

electronic publication http://research.calacademy.org/ichthyology/

catalog/fishcatmain.asp accessed on 29 March 2011.

Froese, R. and Paul, D. (eds.). 2011. FishBase. World Wide Web

electronic publication (version 02/2011) http://www.fishbase.org/

search.php accessed on 29 March 2011.

Harold, A.S. 2003. Melanostomiidae: scaleless dragonfishes, Pp. 907-

912. In: Carpenter, K.E. and Walker, R.W. (eds.), FAO Species

Identification Guide for Fishery Purposes. The living marine

resources of the western central Atlantic. Vol. 2: Bony fishes part

1 (Acipenseridae to Grammatidae). FAO, Rome.

Klepadlo, C. 2011. Three new species of the genus Photonectes

(Teleostei: Stomiiformes: Stomiidae: Melanostomiinae) from the

Pacific Ocean. Copeia 2011(2): 201-210.

Morrow, J.E., and Gibbs, R.H Jr. 1964. Family Melanostomiatidae, Pp.

351-510 In: Bigelow, H.B., Breder, C.M., Cohen, D.M., Mead,

G.W., Merriman, D., Olsen, Y.H., Schroeder, W.C., Schultz, F.P.

and Tee-Van, J. (eds.). Fishes of the Western North Atlantic. Vol.

1. Part 4. Yale University, New Haven.

Nelson, J.S. 2006. Fishes of the World. John Wiley and Sons, New

York.

Regan, C.T., and Trewavas, E. 1930. The fishes of the families

Stomiatidae and Malacosteidae. Danish Dana Expedition 1920-

22 6: 1-143.

Sutton, T.T., and Hopkins, T.F. 1996. Species composition, abundance,

and vertical distribution of the stomiid (Pisces: Stomiiformes) fish

assemblage of the Gulf of Mexico. Bulletin of Marine Science 59:

530-542.

Weitzman, S.H. 1974. Osteology and evolutionary relationships of the

Sternoptychiae, with a new classification of stomiatoid families.

Bulletin of the American Museum of Natural History 153: 327-

478.

Williamson, M., and McGowan, J.A. 2009. The copepod communities

of the north and south Pacific central gyres and the form of

species-abundance distributions. Journal of Plankton Research

32: 273-183.

Memoirs of Museum Victoria 69:269-308 (2012)

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

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

The phyllophorid sea cucumbers of southern Australia (Echinodermata:

Holothuroidea: Dendrochirotida: Phyllophoridae)

P. Mark O’Loughlin 1 , Shari Barmos 2 and Didier VandenSpiegel 3

1 Marine Biology Section, Museum Victoria, GPO Box 666, Melbourne, Victoria 3001, Australia (pmoloughlin@

edmundrice.org)

2 Marine Biology Section, Museum Victoria, GPO Box 666, Melbourne, Victoria3001, Australia (shari_barmos@hotmail.

com)

3 Musee royal de PAfrique centrale. Section invertebres non-insectes, B-3080, Tervuren, Belgium (dvdspiegel@

africamuseum.be)

Abstract O’Loughlin, P. M., Barmos S. and VandenSpiegel D. 2012. The phyllophorid sea cucumbers of southern Australia

(Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae). Memoirs of Museum Victoria 69: 269-308.

A new monotypic Phyllophoridae (Phyllophorinae) genus Phyllostauros , with author O’Loughlin, is erected for

Thyone vercoi Joshua and Creed. We raise Phyllophorella Heding and Panning (subgenus of Phyllophorus Grube) to

generic rank. The holotype and three paratypes of Phyllophorus ventripes Joshua and Creed are conspecific with Thyone

vercoi Joshua and Creed. We make Phyllophorus ventripes Joshua and Creed a subjective junior synonym of Thyone

vercoi Joshua and Creed. One paratype of Phyllophorus ventripes Joshua and Creed is conspecific with Phyllophorus

thyonoides H. L. Clark. We reject the synonymy of Thyone okeni Bell with Thyone venusta Selenka. Twelve new species

of Phyllophoridae are described for southern Australia, with author O’Loughlin: Lipotrapeza eichleri, Lipotrapeza litusi,

Massinium melanieae, Massinium vimsi, Massinium watsonae, Phyllophorella notialis, Thyone flindersi, Thyone joshuai,

Thyone kerkosa, Thyone spenceri, Thyone tourvillei, Thyonidiella kungi. Phyllophoridae species reported previously for

southern Australia are reviewed: Lipotrapeza ventripes (Joshua and Creed); Lipotrapeza vestiens (Joshua); Neothyonidium

dearmatum (Dendy and Hindle); Phyrella thyonoides (H. L. Clark); Thyone nigra Joshua and Creed; Thyone okeni Bell;

Thyone vercoi Joshua and Creed. Species Phyrella thyonoides (H. L. Clark) is re-assigned to genus Phyllophorella Heding

and Panning. A key is provided for the southern Australian species of Phyllophoridae. We acknowledge the rejection for

nomenclatorial purposes by the ICZN (Opinion 417) of the publication by Oken 1815, and hence the current invalid status

of genus Thyone Oken. A petition has been sent to the ICZN for re-validation of Thyone Oken and we retain the use of

Thyone Oken provisionally. The petition includes a similar request for re-validation of Psolus Oken.

Keywords Sea cucumber, Dendrochirotida, Phyllophoridae, Lipotrapeza, Massinium, Neothyonidium, Phyllophorella, Phyrella,

Psolus, Thyone, Thyonidiella, new genus, new species, synonymies, Australia, key, ICZN.

Introduction

The first phyllophorid species described for southern Australia

(Victoria) was Phyllophorus vestiens Joshua, 1914. This species

subsequently became the type species for Lipotrapeza H. L.

Clark, 1938. The following year species Phyllophorus ventripes

Joshua and Creed, 1915 and Thyone nigra Joshua and Creed,

1915 were erected for South Australia. Phyrella thyonoides H. L.

Clark, 1938 was erected for southwest Australian specimens. A

second Thyone Oken, 1815 species occurring in the rocky

shallows of southern Australia has been mistakenly identified as

the New South Wales (Port Jackson) species Thyone okeni Bell,

1884. And Joshua (1914) and Hickman (1978) mistakenly

identified specimens from southeast Australia as the New

Zealand species Neothyonidium dearmatum (Dendy and Hindle,

1907). Thus at the time of writing six phyllophorid species have

been reported for southern Australian waters, two with mistaken

identities. We review the systematic status of these species.

During 1981 the former Victorian Institute of Marine

Sciences conducted a survey of the benthic fauna of eastern Bass

Strait using the Taiwanese FRV Hai Rung and the then New

Zealand Oceanographic Institute RV Tangaroa. Collecting and

sorting yielded specimens as small as a few millimeters in

length. Many small phyllophorid specimens were collected,

representing four new phyllophorid species. We describe the new

species in this work. The Museum Victoria south-east Australia

continental slope survey on RV Franklin yielded a small

phyllophorid from off the Freychinet Peninsula in eastern

Tasmania that is described here. During 1986 and 1987 the South

Australian Department of Fisheries collected many small

phyllophorid specimens from the upper Spencer Gulf that are

lodged in the South Australia Museum and were available for

270

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

this study. We describe two new species from this collection.

And we examined relevant collections of southern Australian

phyllophorid specimens from the Australian Museum, South

Australian Museum, Western Australian Museum and

Tasmanian Museum and Art Gallery.

Recently the new phyllophorid genus Massinium Samyn and

Thandar, 2003 was erected for some former Neothyonidium

Deichmann, 1938 species. We have found three new species of

Massinium in southern Australian waters. There has been

considerable uncertainty about the systematic status of the two

species Phyllophorus ventripes Joshua and Creed , 1915 and

Thyone vercoi Joshua and Creed, 1915 and we have attempted to

resolve the issues.

We are attempting a comprehensive review of southern

Australian phyllophorid species and have therefore described

new species based in some cases on limited and damaged

specimens that nonetheless have distinguishing morphological

characters. We have recognized and noted some cases of what we

consider to be inappropriate species assignments to higher taxa

such as species to genera and genera to sub-families, but we have

generally refrained from altering current assignments until more

comprehensive revisions are possible with the aid of emerging

genetic data. One such current revision that we are aware of is a

review by Fran£ois Michonneau and Gustav Paulay (University

of Florida) of genus Phyrella Heding and Panning, 1954.

We have noticed throughout this work that for a particular

species the number of polian vesicles and the form of the

calcareous ring may vary considerably for individual

specimens. These realities are significant for attempts to

distinguish species systematically.

We acknowledge the rejection for nomenclatorial purposes

by the ICZN (Opinion 417, 42 pp., 1956) of the publication by

Oken 1815, and hence the current invalid status of genus Thyone

Oken, 1815. Pawson and Miller 1981 consequently referred

Thyone to author Jaeger 1833 with type species Holothuria jusus

Muller, 1776 (following Jaeger 1833). This maintained the type

species of Oken for Thyone. But the first author after Oken to use

genus Thyone validly was Lesson 1830, not Jaeger 1833. Lesson

1830 used Thyone as a sub-genus in Holothuria for one new

Aspidochirotida species ( Holothuria (Thyone , Oken) edulis) and

for a new Echiura (spoon-worm) species ( Holothuria ( Thyone ,

Oken) eaouari ). Neither species would be satisfactory as a type

species for Thyone. Rowe (in Rowe and Gates 1995) retained

Oken 1815 as author of Thyone as a nomen conservandum,

following H. L. Clark 1946. This retention has not been validated

by the ICZN. On the grounds of anticipating considerable

uncertainty and instability in holothuroid literature if Thyone

Oken remains invalid and has to be replaced, a petition has been

sent by Gustav Paulay and Mark O’Loughlin (July 2012) to the

ICZN for validation of Thyone Oken, 1815. Reasons have been

given in this petition for the re-validation also of Psolus Oken,

1815. We retain the use of Thyone Oken provisionally.

Methods

Scanning electron microscope (SEM) images were taken by

Didier VandenSpiegel after clearing the ossicles of associated

soft tissue in commercial bleach, air-drying, mounting on

aluminum stubs, and coating with gold. Observations were

made using a JEOL JSM-6480LV SEM. Measurements were

made with Smile view software. Photos of the preserved

specimens were taken by Shari Barmos with a Nikon 300s

DSLR camera, using a Nikkor 105 mm lens and 2x adaptor /

teleconverter. The photo of a live juvenile specimen of

Lipotrapeza vestiens was taken by Leon Altoff and Audrey

Falconer using a Pentax K10D with bellows mounted

Olympus 38mm lens and dual flashes. The photo of a live

specimen of Thyone nigra was taken by John Eichler using a

Pentax W30.

Abbreviations

AM Australian Museum (registration number prefix J).

ICZN Appropriately the International Commission on

Zoological Nomenclature, or the International Code

of Zoological Nomenclature.

MRG Marine Research Group of the Field Naturalists Club

of Victoria.

NMV Museum Victoria (registration number prefix F).

NIWA New Zealand National Institute of Water and

Atmospheric Research (est. 1992).

PPS Port Phillip Survey (1957-1963).

SAM South Australian Museum (registration number

prefix K, photo index number prefix PK).

TMAG Tasmanian Museum and Art Gallery (registration

number prefix H).

VIMS Victorian Institute of Marine Sciences (1974-1998).

WAM Western Australian Museum (registration number

prefix Z).

Numbers in brackets after registrations refer to numbers of

specimens in lots.

Order Dendrochirotida Grube, 1840

Family Phyllophoridae Oestergren, 1907 (sensu Pawson and

Fell 1965)

Diagnosis (after Pawson and Fell 1965). Dendrochirotida with

calcareous ring composite with posterior extensions to the

plates, the extensions composed of small pieces.

Remarks. Pawson and Fell 1965 abandoned tentacle number as

the principal diagnostic criterion for family Phyllophoridae,

and based their new diagnosis on the presence of a composite

calcareous ring as described in the diagnosis above.

Key to southern Australian species of Phyllophoridae

(sensu Pawson and Fell 1965)

1. Tentacles 20 or 15.2

— Tentacles 10, 8 large, 2 small ventral.

genus Thyone Oken.11

2. Tentacles 15, 5 outer pairs large, 5 single small inner ones

. Thyonidiella kungi

O’Loughlin sp. nov. (Bass Strait offshore)

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

271

— Tentacles 20.genera Lipotrapeza H. L. Clark,

Massinium Samyn and Thandar, Pliyllophorella Heding

and Panning, Phyllostauros O’Loughlin gen. nov.. 3

3. Outer tentacles 15, variable sizes, 5 single small inner

ones.genus Phyllophorella

Heding and Panning.4

— Tentacles 5 pairs large, 5 pairs small. genera Lipotrapeza

H.L. Clark , Massinium Samyn and Thandar, Phyllostauros

O’Loughlin gen. nov..5

4. Table ossicle discs regular, never incomplete; tube foot

support ossicles sometimes tables with curved discs, never

dumbbell-shaped rods. Phyllophorella notialis

O’Loughlin sp. nov. (south New South Wales)

— Table ossicle discs frequently irregular, frequently

incomplete; tube foot support ossicles frequently

dumbbell-shaped rods. Phyllophorella

thyonoides (H. L. Clark) (South Australia and south

Western Australia)

5. Calcareous ring composite, long, tubular, plates joined

posteriorly by continuous calcareous band of small plates

creating posterior inter-radial oblong non-calcareous

spaces. Massinium Samyn and Thandar.6

— Calcareous ring with posterior prolongations not joined

by continuous band of small plates; not long, not tubular;

posterior composite extensions short, free.

.genera Lipotrapeza H. L. Clark,

Phyllostauros O’Loughlin gen. nov..8

6. Large and small tube feet uniformly distributed; 4 polian

vesicles; anal body wall with table ossicles.

. Massinium melanieae

O’Loughlin sp. nov. (Great Australian Bight)

— Tube feet not uniformly distributed; fewer than 4 polian

vesicles; anal body wall lacking table ossicles. 7

7. Body wall soft; tube feet scattered dorsally, concentrated

ventrally; peri-oral table discs with up to 20 perforations;

tentacles with rods and some rosettes; endplate support

ossicles perforate rods. Massinium vimsi

O’Loughlin sp. nov. (Bass Strait offshore)

— Body wall thin, firm; tube feet more concentrated along

longitudinal muscles; peri-oral table discs with up to 40

perforations; tentacles with rare fine rods, lacking rosettes;

endplate support ossicles rare. Massinium watsonae

O’Loughlin sp. nov. (SE Tasmania)

8. Adult mid-body wall ossicles lacking .. genus Lipotrapeza

H.L. Clark.9

— Mid-body wall ossicles crosses Phyllostauros vercoi

(Joshua and Creed) (South Australia)

9. Tube feet ossicles include abundant rosettes intergrading

with few small plates. Lipotrapeza litusi

O’Loughlin sp. nov. (SW Australia)

10. Body reddish-brown; tube feet usually concentrated

ventrally, scattered dorsally; tube foot endplate support

ossicles and tentacle ossicles predominantly dumbbell¬

shaped . Lipotrapeza eichleri

O’Loughlin sp. nov. (SE Australia)

— Body pale brown / flesh-pink; tube feet usually evenly

distributed around body; tentacle ossicles predominantly

rods with small distal widening; tube foot endplate

support ossicles irregular narrow perforated plates.

. Lipotrapeza vestiens (Joshua)

(across southern Australia)

11. Adult body wall with table ossicles.12

— Adult mid-body wall lacking table ossicles.16

12. Table ossicles with spires comprising 2 pillars; anterior

third at least of calcareous ring tubular.13

— Table ossicles with predominantly single pillar spires;

none of calcareous ring tubular. Thyone tourvillei

O’Loughlin sp. nov. (eastern Tasmania, Bass Strait)

13. No adult body wall table discs with more than 30

perforations.14

— Many adult body table discs with more than 30 perforations

.15

14. Tentacles with rod and rosettes ossicles; adult body wall

table discs with predominantly 4 and often 8 perforations,

discs typically about 56 pm long. Thyone joshuai

O’Loughlin sp. nov. (South Australia, SE Australia)

— Tentacles with rosette ossicles only; adult body wall table

discs rarely with 4 perforations, discs frequently about 88

pm long. Thyone nigra

Joshua and Creed (SE to SW southern Australia)

15. Adult body wall table discs small to large, large discs

sometimes narrow with solid distal extensions, spires long

with long splayed distal ends. Thyone kerkosa

O’Loughlin sp. nov. (Recherche Archipelago)

— Adult body wall table discs round to oval, uniform size,

spires short. Thyone spenceri

O’Loughlin sp. nov. (Upper Spencer Gulf)

16. Peri-oral table ossicles with irregular discs, up to 72 pm

long, fewer than 40 perforations, spires with 2 pillars;

tentacles branches with thin rods, distally with fine

branches or few perforations, some rods with sigmoid

form, some straight, rods up to 88 pm long.

. Thyone flindersi

O’Loughlin sp. nov. (southern Australia)

— Peri-oral table ossicles with regular discs, up to 192 pm

diameter/length, up to more than 60 small perforations,

table spires thick short columnar mesh; tentacle branches

with stout rods, some dumbbell-shaped, rods up to 320

pm long. Thyone okeni Bell (New South Wales)

Tube feet never with rosette ossicles

272

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Subfamily Phyllophorinae Heding and Panning, 1954

Diagnosis. Radial plates of the calcareous ring with posterior

prolongations composed of a few large pieces, prolongations

sometimes arising jointly from radial and inter-radial plates;

inter-radial plates frequently lacking any posterior prolongation;

ring not tubular.

Southern Australia genera. Lipotrapeza H. L. Clark, 1938;

Phyllophorella Heding and Panning, 1954; Phyllostauros

O’Loughlin gen. nov. (below); Thyonidiella Heding and

Panning, 1954.

Remarks. We raise subgenus Phyllophorella Heding and Panning,

1854 (subgenus of Phyllophorus Grube, 1840) to generic rank.

Lipotrapeza H. L. Clark, 1938

Lipotrapeza H. L. Clark, 1938: 494-495.-H. L. Clark 1946:

411.-Heding and Panning, 1954:173-175.

Diagnosis. Dendrochirote holothuroids with 20 tentacles, 5 pairs

large, 5 pairs small; tube feet around body, not radial, sometimes

concentrated ventrally and scattered dorsally; calcareous ring

stout with paired short tapered posterior composite prolongations

arising from radial (early development) or jointly from radial and

inter-radial plates (later development); mid-body wall of adults

lacking ossicles; mid-body wall of juveniles with tables, spires

with 4 pillars; tube feet with endplates and endplate supporting

ossicles, sometimes with rosettes and small plates.

Type species. Phyllophorus vestiens Joshua, 1914 (original

designation).

Other species. Lipotrapeza ambigua Cherbonnier, 1988

(Madagascar); L. capilla Cherbonnier, 1958 (Sierra-Leone); L.

eichleri O’Loughlin sp. nov. (SE Australia); L. incurva

Cherbonnier, 1988 (Madagascar); L. japonica Heding and

Panning, 1954 (Japan); L. litusi O’Loughlin sp. nov. (SW

Australia).

Remarks. We make Lipotrapeza ventripes (Joshua and Creed,

1915) a subjective junior synonym of Phyllostauros vercoi

(Joshua and Creed, 1915) (see below). Cherbonnier 1988 referred

five small specimens (up to 20 mm long) from Madagascar to

Lipotrapeza ventripes (Joshua and Creed, 1915). This referral is

discussed under Lipotrapeza vestiens (below).

Lipotrapeza ambigua Cherbonnier, 1988, erected for 12

specimens from Madagascar, is described as having 10 large

outer tentacles and two inner circles of five small ones, a

calcareous ring with paired composite radial posterior

prolongations and composite inter-radial posterior prolongations,

and some tables in the adult anal body wall. We judge that the

species does not fit the diagnostic characters of Lipotrapeza H.

L. Clark, 1938.

Lipotrapeza capilla Cherbonnier, 1958, erected for two

specimens from Sierra-Leone, is described as having a tentacle

arrangement of 15 outer tentacles comprising five pairs of large

with a very small tentacle between each pair, and five inner

tentacles. There are rosette and plate ossicles in the mid-body

wall, and long paired composite posterior radial elongation of

the calcareous ring. We judge that the species does not fit the

diagnostic characters of Lipotrapeza H. L. Clark, 1938.

Lipotrapeza incurva Cherbonnier, 1988, erected for eight

specimens from Madagascar, is described as having 10 large

outer tentacles and 10 very small ones in two inner circles. On

this character we judge that the species does not fit the

diagnostic characters of Lipotrapeza H. L. Clark, 1938,

although for most characters it does.

We also query the inclusion of the northern Lipotrapeza

japonica Heding and Panning, 1954, specifically for the

presence of three series of tentacles (10 + 5 + 5) and the form

of the calcareous ring with discrete paired posterior radial

prolongations (as illustrated by Heding and Panning 1954).

The current referrals of species to Lipotrapeza H. L. Clark,

1938 suggest to us that from a morphological viewpoint the

genus is currently polyphyletic and requiring review.

Lipotrapeza eichleri O’Loughlin sp. nov.

Figures la, b, 2

Material examined. Holotype. Victoria, Phillip I., Kitty Miller Bay,

rocky shallows, M. O’Loughlin and J. Monagle, 20 Apr 1987, NMV

F174906.

Paratypes. Type locality and date, NMV F161502 (4); Westernport

Bay, Phillip I., McHaffie Reef, sea-grass beds, J. Eichler and V. Stajsic,

1 Mar 2008, NMV F157403 (1).

Other material. Cape Paterson, 4 m, 13 Mar 2001, NMV F97432

(2); Westernport Bay, Shoreham, 16 Feb 1972, NMV F45265 (1);

Honeysuckle reef, 21 Mar 1976, NMV F161493 (1); Port Phillip Bay,

Point Lonsdale, J. Kershaw, Jan 1902, NMV F76558 (3); Portland,

Anderson Point, 23 Feb 2007, NMV FI25359 (1 juvenile).

Diagnosis. Up to 75 mm long, up to 25 mm diameter (preserved,

tentacles withdrawn), cylindrical body, narrow rounded oral and

anal ends, thin firm body wall minutely papillate; external anal

scales not detected; 20 tentacles, 5 large pairs, alternating with

5 small pairs; complete cover of tube feet, crowded ventrally,

scattered dorsally, diameter about 0.5 mm; calcareous ring with

10 posterior composite tapering projections arising from both

radial and inter-radial plates, posterior ends free thin tails, ring

not tubular, length of plates with anterior projections sub-equal

with posterior composite prolongations, form of the plates and

composite components variable; short stone canal, madreporite

close to vascular ring; 1-3 tubular polian vesicles, variable

lengths; longitudinal muscles flat, lacking longitudinal

indentation centrally; gonad tubules arise in series along

gonoduct on each side of dorsal mesentery, tubules not branched.

Adult mid-body wall lacking ossicles; tube foot ossicles

endplates and support rods only; endplates with denticulate

margin, irregular perforations similar size, diameters up to

480 pm\ tube foot support ossicles predominantly dumbbell¬

shaped, enlarged distally, perforate, denticulate margin, up to

192 pm long; peri-oral region with abundant rosettes about 40

pm long; tentacles with rods and rosettes, larger rods dumbbell¬

shaped with small perforations and denticulate margin distally,

up to 208 pm long, few small rosettes up to 40 pm long; anal

ossicles tube foot endplates and support rods, tentacle-like

rods, rosettes, some large up to 64 pm long.

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

273

Figure 1. Photos of preserved type specimens, a, Lipotrapeza eichleri O’Loughlin sp. nov. (holotype NMV F174906); b, Lipotrapeza eichleri

calcareous ring (paratype NMV F161502); c, Lipotrapeza litusi O’Loughlin sp. nov. (holotype WAM Z13475); d, Thyonidiella kungi O’Loughlin

sp. nov. (holotype NMV F76637) (insert: form of the calcareous ring plates); e, Phyllophorella notialis O’Loughlin sp. nov. (holotype NMV

F132691) (insert: form of the calcareous ring plates); f, Phyllophorella notialis calcareous ring (holotype).

274

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 2. SEM images of ossicles from specimens of Lipotrapeza eichleri O’Loughlin sp. nov. Top, rods and rosettes from tentacles (holotype

NMV F174906); middle, tables from juvenile (NMV F161502,5 mm long); bottom, endplate and dumbbell-shaped endplate support ossicles from

same juvenile (NMV F161502).

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

275

Juvenile (5 mm long) with mid-body wall tables and tube

foot endplates and support rods; tables with irregular discs up

to 60 pm long, short irregular 4-pillared spires; endplates up to

180 pm diameter, dumbbell-shaped endplate support rods up

to 240 pm long.

Colour (preserved). Body dark brown to grey-brown with

variable red colouration, dark grey-brown oral end, white anal

end; many tube feet red; tentacles pale brown.

Distribution. South-east Australia, Victoria, Cape Paterson to

Portland, rocky shallows and sea-grass, 0-4 m.

Etymology. Named for John Eichler, with appreciation of his

contribution to Museum Victoria through the MRG, and his

discovery and field photograph of a paratype of this new species.

Remarks. We note that unlike the other two species of

Lipotrapeza H. L. Clark, 1938 the large and small pairs of

tentacles in Lipotrapeza eichleri O’Loughlin sp. nov. alternate

in one series, not inner and outer series. Lipotrapeza eichleri is

distinguished from other Lipotrapeza species in the key (above).

Lipotrapeza litusi O’Loughlin sp. nov.

Figures lc, 3

Material examined. Holotype. Western Australia, Cottesloe Beach,

32°00'S 115°45'E, beach washed, L. M. Marsh, Aug 1990, WAM Z13475.

Diagnosis. Up to 70 mm long, 22 mm diameter (preserved,

tentacles withdrawn); form cylindrical, upturned oral and anal

rounded tapers; 20 tentacles, 5 outer pairs large, 5 inner pairs

small; tube feet all around body, scattered dorsally, close

ventrally, across introvert, diameters up to 0.7 mm; calcareous

ring stout, irregular, with some short posterior composite

projections arising jointly from radial and inter-radial plates,

not tubular, lacking thin radial composite posterior elongations;

short stone canal, madreporite close to vascular ring; single

polian vesicles; thin branched gonad tubules arise in series

along gonoduct on each side of dorsal mesentery.

Mid-body wall lacking ossicles; tube feet with endplates,

support rods, abundant rosettes, few small plates; endplate

diameters up to 536 pm\ support rods stout, distally enlarged

and perforate, up to 144 pm long; rosettes oval, up to 48 pm

long, intergrade with small plates; small plates irregular,

perforate, denticulate margin, up to 104 pm long; tentacles with

rods and rosettes, rods stout, distally enlarged and perforate, up

to 200 pm long, abundant rosettes up to 64 pm long.

Colour (preserved). Body off-white, body and tube feet with

some residual red; tentacle trunks white, branches pale brown.

Distribution. Southwest Australia, Cottesloe beach, off-shore

sediments.

Etymology. Named from the Latin litus (beach) with reference

to the beach-washed source of the type specimen.

Remarks. Lipotrapeza litusi O’Loughlin sp. nov. is distinguished

from other Lipotrapeza species in the key (above). The single

specimen that this species is based on was found on a beach

and the species presumably lives in off-shore sediments.

Lipotrapeza vestiens (Joshua, 1914)

Figures 4, 5a, 6

Phyllophorus vestiens Joshua, 1914: 5, pi. 1 fig. 2a-f.

Lipotrapeza vestiens.— H. L. Clark, 1938: 496-497.—H. L. Clark,

1946: 412.—Heding and Panning, 1954: 176, fig. 84 (from Joshua

1914).—Rowe, 1982: 462, pi. 31.2.-Rowe and Gates, 1995: 311.—

Edgar, 1997: 369,-Gowlett-Holmes, 2008: 263.

Material examined. Holotype. Victoria, Torquay, NMV F45144.

Paratypes. Torquay, NMV F45145 (1); slide of body wall, NMV

F45146 (mounted body wall, 33 mm long, 20 mm wide, not from either

type specimen, endplates and support rods only).

Other material (selection). Victoria, Cape Conran, 2 Oct 1988,

NMV F73832 (1); Walkerville south. Bear Gully, 7 Mar 1982, NMV

F73838 (1); Cape Paterson Survey (CPA), 1982, Petrel Rock, NMV

F73823 (1); Shack Bay, NMV F73824 (1); Cape Paterson, NMV F73825

(1); Harmers Haven, NMV F73822 (1); Westernport Bay Survey by

MRG, 1969-71, Merricks, NMV F45239 (3); 29 Oct 1980, NMV F75976

(many); 28 Jan 1983, NMV F73831 (2); Shoreham, NMV F73854 (1);

Flinders, 26 Feb 1977, NMV F74235 (5); 10 Mar 1980, NMV F73830

(many); Port Phillip Bay, Beaumaris, Ricketts Point, 25 Apr 2008, NMV

FI57402 (1); Cheltenham beach after storm, 20 Jul 1891, NMV F73856

(6); Mordialloc beach. May 1897, NMV F73857 (4); Sandringham, Jul

25 1891, NMV F73858 (4; no. 60642-5; det. by Joshua 1914; det. as

Phyllophorus ventripes by F. W. E. Rowe 1976); Kennett River, 29 Dec

1982, NMV F73827 (2); Marengo, 26 Mar 1977, NMV F73850 (2);

Portland, MRG, 23 Feb 2007, NMV F125356 (1).

Tasmania. Bass Strait, Lulworth, 22 Nov 1982, NMV F174904 (13);

North Head, mouth of Tamar River, 28 Aug 1978, NMV F97070 (1).

South Australia. Gulf St Vincent, Willunga Reef, 23 Nov 1976,

SAM K2583 (1); Cape Jervis, 3-5 m, 22 Apr 2005, SAM K2590 (1,

PK0296); S side of Kangaroo I., Hanson Bay, rocky shallows, 6 Mar

1978, AM J12559 (3); Whittlebee Point, 1 Mar 1975, SAM K2591 (4);

Baird Bay, 27 Feb 1975, SAM K2596 (3).

Western Australia, Two Peoples Bay, 5 Dec 1968, WAM Z31964

(1); Cape Naturaliste, 31 Dec 1971, WAM Z31971 (1); Yanchep,

31°33'S 115°4rE, reef flat, 1959, WAM Z8994 (1).

Diagnosis. Up to 175 mm long, up to 35 mm diameter

(preserved, tentacles withdrawn, largest WAM Z8994),

cylindrical body with slight anal taper, thick soft body wall;

external anal scales not detected; 20 tentacles, 5 outer pairs

large, 5 inner pairs small; complete cover of close tube feet, not

contiguous, sometimes more concentrated ventrally than

dorsally, diameter about 0.7 mm, radial to scattered tube feet

cross introvert; calcareous ring with 10 posterior composite

tapering projections arising predominantly from radial plates

but sometimes jointly from both radial and inter-radial plates,

posterior ends free thin tails comprising small elongate plates,

ring not tubular, posterior projections shorter than combined

height of plates and anterior projections, form of plates and

composite components variable; short stone canal, madreporite

multi-lobed, close to vascular ring; up to 5 polian vesicles,

variable sizes, frequently one; longitudinal muscles flat,

longitudinal indentation along centre of muscle; gonad tubules

arise in series along gonoduct on each side of dorsal mesentery,

tubules not branched.

Mid-body wall lacking ossicles; tube foot ossicles endplates

and support rods and narrow plates only; endplates with

denticulate or smooth margin, irregular perforations similar

size, endplate diameters up to 480 pm\ endplate support rods

276

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 3. SEM images of ossicles from holotype of Lipotrapeza litusi O’Loughlin sp. nov. (WAM Z13475). Tube foot endplate (top right),

rosettes, endplate support rods, rare perforated plate (bottom right).

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

277

Figure 4. Photos of preserved specimens of Lipotrapeza vestiens (Joshua, 1914). a, adult specimen with tentacles withdrawn, close cover of tube

feet, paired series of tube feet along interior longitudinal muscle attachment (NMV F73830, 80 mm long); b, calcareous ring with posterior

prolongations arising from both radial and inter-radial plates (NMV F73830); c, oral view of tentacle crown with inner and outer rings of 5 paired

small and 5 paired large tentacles (NMV F75976); d, lateral view of bare introvert and tentacle crown showing outer paired large and inner paired

small tentacles (NMV F73830); e, juvenile with radial paired series of tube feet (NMV F75976); f, juvenile cucumariid-like calcareous ring

lacking posterior prolongations (NMV F75976).

278

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 5. Photos of live specimens, a juvenile of Lipotrapeza vestiens (Joshua, 1914) from Eagles Nest, coast of Victoria, with body wall tables

and ventral tube feet only (3 mm long; photo by Leon Altoff); b, specimen of Thyone nigra Joshua and Creed, 1915 from McHaffie Reef,

Westernport Bay, with calcareous ring eviscerated (NMV F151855, photo by John Eichler).

and narrow plates straight to curved, larger ossicles perforate

and denticulate, up to 200 pm long; tentacles with rods and

rosettes, larger rods branched and denticulate distally,

sometimes with small perforations, up to 160 pm long, rosettes

oval to irregular, up to 80 pm long; peri-oral region with

tentacle-like rods and rosettes; introvert with rosettes, fine

rods up to 80 pm long; anal ossicles tube foot endplates and

support rods, tentacle-like rods, rosettes, 5 scales with base up

to 320 pm wide and digitiform column up to 360 pm long,

base and column comprising massed short branched rods.

Juveniles. For specimen 20 mm long tentacles 20; for

specimen 10 mm long tube feet radial only; for specimen 15 mm

long calcareous ring cucumariid-like, lacking posterior

projections, not composite; for specimens 7 mm long and smaller,

tables present in mid-body wall, discs regular and irregular with

frequently 8 perforations, lobed margin, discs typically 72 pm

long, spires with 4 pillars, spires up to 64 pm long.

Colour (preserved). Body pink-brown, dark brown orally and

anally; tube feet off-white; large tentacle branches dark brown,

trunks off-white, small tentacles off-white.

Distribution. Southern Australia, from E Victoria, Cape

Conran, west to S Western Australia (Yanchep); rocky shallows,

0-5 m.

Remarks. Cherbonnier 1988 referred five small specimens (up

to 20 mm long) from Madagascar to Lipotrapeza ventripes

(Joshua and Creed, 1915). As noted above and discussed below

we make Lipotrapeza ventripes (Joshua and Creed, 1915) a

subjective junior synonym of Phyllostauros vercoi (Joshua and

Creed, 1915). In his observations Cherbonnier 1988 based his

referral of these specimens to Lipotrapeza ventripes on Clark

1938. We judge from their descriptive remarks that both authors

are referring to Lipotrapeza vestiens (Joshua, 1914). The

Madagascar specimens have the diagnostic characters of the

presence of tentacle rods, the form of the calcareous ring,

presence of tube foot endplates and support rods, and absence

of ossicles in the mid-body wall that characterize Lipotrapeza

vestiens. But Lipotrapeza vestiens specimens attain a much

larger size (up to 175 mm long), have five distinct pairs of inner

small tentacles (not two circles of five), do not normally have

fewer dorsal than ventral tube feet, and lack the “pseudo¬

plates” referred to by Cherbonnier 1988. We judge that the

Madagascar specimens are not Lipotrapeza vestiens.

Lipotrapeza vestiens (Joshua, 1914) is distinguished from the

other species of Lipotrapeza in the key (above).

Phyllophorella Heding and Panning, 1954

Diagnosis (after Heding and Panning 1954). Dendrochirote

holothuroids with 20 tentacles in 2 series of 15 variably large

outer and 5 small inner; body covered with tube feet; calcareous

ring with 5 paired short composite tapering posterior

extensions, arising principally from the radial plates but

sometimes jointly from the radial and inter-radial plates; body

wall ossicles tables with 4-pillared spires.

Type species. Phyllophorus (Phyllophorella) robusta Heding

and Panning, 1954 (original designation; locality Thailand).

Other species. P. contractura Cherbonnier, 1988; P. drachi

Cherbonnier and Guille, 1968; P. dubius Cherbonnier, 1961; P.

kohkutiensis Heding and Panning, 1954; P. liuwutiensis (Yang,

1937); P. longipeda (Semper, 1867); P. notialis O’Loughlin sp.

nov.; P. perforata (H. L. Clark, 1932); P. purpureopunctata

(Sluiter, 1901); P. rosetta Thandar, 1994; P. roseus Cherbonnier

and Feral, 1981; P. spiculata (Chang, 1935); P. thyonoides (H.

L. Clark, 1938).

Remarks. We re-assign species Phyllophorus thyonoides H. L.

Clark, 1938 from Phyrella Heding and Panning, 1954 to

Phyllophorella Heding and Panning, 1954 on the basis of

having 15 variably large outer tentacles and 5 small inner ones,

and frequent cases of the posterior prolongations of the

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

279

Figure 6. SEM images of ossicles from juvenile of Lipotrapeza vestiens (Joshua, 1914). Mid-body wall tables with 4-pillared spires, irregular

discs with about 12 perforations, endplate with support rods (5 mm long, NMV F125356).

280

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

calcareous ring arising from both radial and inter-radial plates

(as illustrated by Heding and Panning 1954 for most

Phyllophorella species). We note that Rowe (in Rowe and Gates

1995) made Phyllophorus parvipedes H. L. Clark, 1938 (type

locality Broome in NW Australia) a subjective junior synonym

of Phyllophorus spiculata Chang 1935 (type locality China).

Phyllophorella notialis O’Loughlin sp. nov.

Figures le, f, 7

Material examined. Holotype. SE Australia, New South Wales,

Disaster Bay, RV Southern Surveyor SS0404 stn 102, 37°17.22'S

150°4.15'E, 79 m, 28 April 2004, NMV F132691.

Description. Body 20 mm long, diameter up to 7 mm (preserved,

tentacles slightly extended, part of posterior body missing); 20

tentacles in 2 series, 15 outer variable sizes from large to medium,

5 inner very small; complete cover of tube feet, diameter about

0.3 mm; calcareous ring not tubular, radial plates with paired

composite tapering tails; inter-radial plates not composite,

lacking posterior extensions, truncate, less than half total length

of radial plates; single polian vesicle; lacking gonad tubules.

Mid-body ossicles tables only, table discs variably round to

oval, margins smooth and slightly lobed, disc lengths variable

from 64 pm to 192 pm, frequently 96 pm, table spires short,

about 24 pm long, predominantly 4 pillars, rarely 5, blunt

apical spines; endplates up to 144 pm diameter, endplate

support ossicles tables with curved discs, discs up to 136 pm

long; tentacle ossicles abundant small, fine rods, intergrading

with stout rods with swollen multi-perforate and bluntly

denticulate ends, each end with a large and up to 10 small

marginal perforations, rods up to 176 pm long.

Colour. Body grey-brown, introvert white, tentacles pale brown

with some dark brown markings.

Distribution. Australia, south coast of New South Wales,

Disaster Bay, 79 m.

Etymology. Named from the Latin notialis (southern), for the

research vessel Southern Surveyor.

Remarks. Phyllophorella notialis O’Loughlin sp. nov. is

distinguished from other southern Australian species of

Phyllophoridae in the key (above).

Phyllophorella thyonoides (H. L. Clark, 1938)

Figures 9, 10, 13f

Phyllophorus thyonoides H. L. Clark, 1938: 492-494, fig. 48.—H.

L. Clark, 1946: 409.

Phyllophorus ventripes Joshua and Creed, 1915: 19, pi. 2 fig. 1, pi.

3 fig. 5. (part).

Lipotrapeza ventripes— H. L. Clark, 1938: 495-496— H. L.

Clark, 1946: 412.—Heding and Panning, 1954: 175, fig. 83 (from

Joshua and Creed 1915).—Smith, 1970: 94.—Rowe, 1982: 464.—Rowe

and Gates, 1995: 311. (part).

Phyrella thyonoides— Heding and Panning, 1954: 183-184, fig.

88 (from H. L. Clark 1938).-Rowe and Gates, 1995: 313-314.

Material examined. Phyllophorus ventripes Joshua and Creed, 1915.

Paratype. South Australia, J. C. Verco, SAM K1375 (1; earliest label

“type material”; recent label “type”; judged here to be a paratype;

determined as Phyrella thyonoides by F. W. E. Rowe 1977; confirmed

here).

Other material. South Australia, J. C. Verco, SAM K2597 (1;

determined as Phyllophorus (Phyllophorella) thyonoides by F. W. E.

Rowe 1977; confirmed here).

Western Australia, Garden I., Careening Bay, intertidal sand flats,

26 Nov 1961, WAM Z31837 (1); sand flats, 20 Dec 1969, WAM Z31962

(1); Cockburn Sound, Woodman’s Point, seagrass bank, 1 Dec 1957,

WAM Z31834 (2); 1 m, 18 Mar 1972, WAM Z31975 (1); 100 km S of

Geraldton, Cliff Head, seagrass rhizomes, 2 m, 2 Sep 1985, WAM

Z21214 (2); 3 Jun 1986, AM J21716 (4).

Diagnosis. Up to 60 mm long, up to 25 mm diameter (preserved,

tentacles withdrawn), curved cylindrical body tapered to rounded

upturned ends orally and anally, variably with ventral belly, thick

firm body wall; external anal scales not detected; 20 tentacles, 15

variably large outer, 5 inner small; complete spaced cover of

large and small tube feet, larger and more numerous ventrally,

disc ends up to 0.7 mm diameter, smaller dorsally, orally, anally,

radial to scattered tube feet cross introvert, spaced paired

posterior radial series; calcareous ring with 10 posterior

composite tapering projections arising from radial plates only

(smaller specimens) or jointly from both radial and inter-radial

plates (larger specimens), ring not tubular, posterior ends free

tapering tails generally sub-equal with anterior plates and

projections, spaces between posterior prolongations extending

anteriorly to similar extents radially and inter-radially, form of

plates and composite components variable; short stone canal,

multi-lobed madreporite close to vascular ring; 1-3 polian

vesicles; gonad tubules arise in series along gonoduct on each

side of dorsal mesentery, tubules not branched.

Mid-body wall with scattered tables, discs regular to

irregular in form, some with appearance of being incomplete,

rounded quadrangular to oval, commonly 96 pm long,

frequently up to 112 pm long, rarely up to 144 pm long, margin

lobed, lobes smooth or denticulate, commonly central and

about 8 outer perforations, sometimes many perforations

anally, spires frequently with 4 pillars up to 48 pm high, rarely

3, 5, 6 pillars; tube foot ossicles endplates and endplate support

ossicles only; endplates with denticulate margin, irregular

perforations similar size, diameters up to 600 pm\ endplate

support ossicles up to 184 pm long, dumbbell-shaped rods with

rounded ends with large and small perforations and denticulate

margin, and narrow multi-perforate plates with denticulate

margin; tentacles with rods and rosettes, larger rods dumbbell¬

shaped, widened and perforate distally with denticulate margin,

up to 320 pm long, rosettes up to 104 pm long; introvert with

some rosettes up to 48 pm long; peri-oral region with tentacle¬

like rods and rosettes; anal ossicles tube foot endplates and

support rods, tentacle-like rods, rosettes, 5 scales comprising

base up to 320 pm wide with digitiform column up to 360 pm

long, base and column comprising massed short branched rods.

Specimen 4 mm long with radial tube feet; cucumariid

ring lacking posterior prolongations; mid-body ossicles typical

solid 4-pillared tables, some margins closely denticulate, rare

larger irregular tables with discs up to 120 pm wide.

Colour (preserved). Body off-white with close grey-brown/

black flecking creating dark grey/black appearance, body

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281

Figure 7. SEM images of ossicles from holotype of Phyllophorella notialis O’Loughlin sp. nov. (NMV F132691). Tentacle rods (top right) and

4-pillared tables from the body wall.

282

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 8. SEM images of ossicles from specimens of Thyonidiella kungi O’Loughlin sp. nov. Tentacle rosettes and rods (left; holotype NMV

F76637); endplate, dumbbell-shaped endplate support ossicles, and 4-pillared tables, from body wall of a juvenile (3 mm long; NMV F76639).

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283

Figure 9. Photos of preserved specimens of Phyllophorella thyonoides (H. L. Clark, 1938). a, specimen from Western Australia (AM J21716); b,

calcareous ring with posterior prolongations from radial plate only (AM J21716); c, specimen from South Australia (SAM K2597); d, calcareous

ring with posterior prolongations jointly from radial and inter-radial plates (paratype of Phyllophorus ventripes Joshua and Creed, 1915, SAM

K1375).

sometime red-brown to orange, not darker orally and anally;

tube feet off-white, sometimes red; tentacle branches dark

brown, white lumps (rosette ossicle clusters) on trunks.

Distribution. St. Vincent Gulf (South Australia) to Cliff Head

(100 km S of Geraldton) (Western Australia), intertidal and

offshore sediments, seagrass beds, 0-2 m.

Remarks. Specimen SAM K1374 is judged here (and so reported

by Rowe and Gates 1995) to be the holotype for Phyllophorus

ventripes Joshua and Creed, 1915. Thus specimens of

Phyllophorus ventripes SAM K1375 (labelled “type material”)

and NMV F45143 (3 specimens labelled “syntypes”, and so

reported by Smith 1970) are judged here to be paratypes. Rowe

and Gates 1995 listed four paratype specimens of Phyllophorus

ventripes in lot SAM K1375. We found only one specimen in

this lot. The three missing specimens may be those registered as

syntypes (NMV F45143). We examined these SAM and NMV

type specimens. The holotype of Phyllophorus ventripes Joshua

and Creed, 1915 and the three NMV paratypes are conspecific

with Thyone vercoi Joshua and Creed, 1915, described in the

same paper and based on a single specimen (see below under

Phyllostauros vercoi ). The remaining SAM K1375 paratype of

Phyllophorus ventripes is conspecific with Phyllophorus

thyonoides H. L. Clark, 1938.

There are no labels or other records of where and when Dr.

J. C. Verco collected the specimens for which Thyone vercoi

and Phyllophorus ventripes were erected as new species.

However H. L. Clark 1938 records Le Fevre Peninsula (sandy

shallows 15 km NW of Adelaide in Gulf St. Vincent) as the

precise locality for the collection of two loan specimens of

Thyone vercoi (information presumably pers. comm). Apart

from the original specimens collected by Dr. Verco there are

no other SAM specimens of either of these two species. WAM

specimens of P. thyonoides collected from sea grass beds and

284

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 10. SEM images of ossicles from specimen of Phyllophorella thyonoides (H. L. Clark, 1938) (AM J21716). Tentacle rods, some dumbbell¬

shaped, and rosettes (top); mid-body wall tables and tube foot endplate and support plate ossicle (bottom).

• I

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285

Figure 11. Photos of preserved specimens of Phyllostauros vercoi (Joshua and Creed, 1915). a, lateral view showing tube feet close ventrally and

ventro-laterally, and sparse dorsally (paratype of Phyllophorus ventripes Joshua and Creed, 1915; NMV F45143) (insert: form of the calcareous

ring plates); b, calcareous ring (NMV F45143); c, ventro-lateral view (holotype of Phyllophorus ventripes Joshua and Creed, 1915, dry, oral end

detached, SAM K1374); d, calcareous ring (SAM K1374).

the sandy Le Fevre Peninsula provide evidence of the eco-

niche for both P. thyonoides and P. vercoi.

The drawing of the calcareous ring for Phyllophorus

ventripes Joshua and Creed, 1915 ( P. thyonoides and P. vercoi

here) is somewhat misleading as it shows posterior

prolongations arising from the radial plates only, whereas in

many specimens they arise jointly from radial and inter-radial

plates. H. L. Clark 1938 noted for Phyllophorus thyonoides

“obviously the lines of division between radial and inter-radial

pieces of the calcareous ring are indefinite and arbitrary”. We

agree. We noted that in one specimen of Phyllophorella

thyonoides (WAM Z31975, illustrated in fig. 13f) the posterior

prolongations of the calcareous ring are joined as in genus

Massinium Samyn and Thandar, 2003 species (see below).

The dorsal tube feet are deeply buried in pits for

Phyllophorus ventripes paratype SAM K1375 ( P. thyonoides

here), and this presumably led the original authors to describe

the species as lacking tube feet dorsally.

Phyllostauros O’Loughlin gen. nov.

Diagnosis. Dendrochirote holothuroids with 20 tentacles, 5

large pairs alternating and 5 small pairs; body covered with

tube feet; calcareous ring with 5 paired composite posterior

tapering prolongations, arising jointly from the radial and

inter-radial plates; body wall ossicles crosses with bifurcate

ends, 1 or 2 surface spines near the ends.

Type species. Thy one vercoi Joshua and Creed, 1915 (monotypic).

Distribution. Central southern Australia.

Etymology. From the first part Phyllo of the relevant family

name Phyllophoridae, with stauros (Greek for cross) referring

to the body wall ossicle form.

Remarks. Thyone vercoi Joshua and Creed, 1915 was erected

for one specimen that is now dry and in poor condition

(holotype, SAM K1374). The authors made three permanent

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P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 12. SEM images of ossicles of specimen of Phyllostauros vercoi (Joshua and Creed, 1915) (from paratype of Phyllophorus ventripes

Joshua and Creed, 1915; NMV F45143). Surface-knobbed crosses from the body wall.

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

287

Figure 13. Photos of preserved specimens, a, Massinium melanieae O’Loughlin sp. nov. (holotype NMV F174897); b, Massinium vimsi

O’Loughlin sp. nov. (holotype, NMV F76629) (insert: form of the calcareous ring plates); c, Massinium watsonae O’Loughlin sp. nov. (holotype,

NMV F97435); d, calcareous ring, stone canal (along alimentary canal) and madreporite, and 2 polian vesicles (top left corner) of Massinium

watsonae (TMAG H1988); e, colour morph of Massinium watsonae (NMV F174899) (insert: form of the calcareous ring plates); f, exceptional

calcareous ring from specimen of Phyllophorella thyonoides (H. L. Clark, 1938) with posterior prolongations joined as in Massinium species

(WAM Z31975) (insert: form of the calcareous ring plates).

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P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

microscope slides from parts of the dorsal body wall, pharynx

and a tentacle. These slides are in good condition, and held by

SAM. H. L. Clark 1946 refers to additional slides held in MCZ.

The tentacle crown is damaged, but there are more than the 10

tentacles reported originally, and close to 20. The calcareous

ring is composite with posterior prolongations, but is not

tubular as in southern Australian Thyone Oken, 1815 species.

Based on the cross ossicles in the body wall H. L. Clark

1946 referred Thyone vercoi to Staurothyone H. L. Clark, 1938,

but with reservations. He noted that Deichmann thought that the

species was identical with Lipotrapeza vestiens (Joshua, 1914).

Genus Staurothyone is in family Cucumariidae. Thyone vercoi

has close to 20 tentacles, composite posterior extensions to the

plates of the calcareous ring, and tube feet all over the body

(close ventrally, scattered and sparse dorsally). Thyone vercoi

belongs in family Phyllophoridae, not Cucumariidae, and

cannot be retained in Staurothyone. Thyone vercoi is unique

amongst Phyllophoridae species for the presence of body wall

cross ossicles and absence of table ossicles. Rowe 1982, and

Rowe in Rowe and Gates 1995, anticipated the need for a new

genus. Phyllostauros O’Loughlin gen. nov. is erected here as a

monotypic genus, and is referred to family Phyllophoridae and

sub-family Phyllophorinae with reservations because of the

unique cross ossicles in the body wall.

Phyllostauros vercoi (Joshua and Creed, 1915)

Figures 11, 12

Thyone vercoi Joshua and Creed, 1915: 19-20, pi. 2 figs 2-4, pi. 3

fig. 1, pi. 4.-H. L. Clark, 1938: 463-464.

Phyllophorus ventripes Joshua and Creed, 1915: 19, pi. 2 fig. 1, pi.

3 fig. 5. (part).

Lipotrapeza ventripes.— H. L. Clark, 1938: 495-496.—H. L.

Clark, 1946: 412.—Heding and Panning, 1954: 175, fig. 83 (from

Joshua and Creed 1915).—Rowe, 1982: 464.—Rowe and Gates, 1995:

311. (part).

Staurothyone vercoi.— H. L. Clark, 1946: 397-398.—A. M. Clark,

1966: 345.—Rowe and Gates, 1995: 281.

‘ Staurothyone ’ vercoi.— Rowe, 1982: 464, fig, 10.29b.

Material examined. Holotype. South Australia, SAM K517 (with

microscope slides of “skin”, tentacle, and pharynx; specimen strongly

contracted, dry).

Phyllophorus ventripes Joshua and Creed, 1915. Holotype. South

Australia coast, SAM K1374 (specimen dry; oldest labels “type

M.S.S.”, “ Phyllophorus sp. n. type”, “holotype”; determined as

“ Sturothyone ” vercoi by F. W. E. Rowe 1977). Paratypes. South

Australia coast, 18 Sep 1913, NMV F45143 (3; no. 60676-8; labelled

syntypes; determined by Joshua and Creed in 1914; from SAM “in

exchange” 19 Sep 1919; AM label u Staurothyone , ’ > vercoi ; judged here

to be paratypes).

Diagnosis. Up to 50 mm long, up to 15 mm diameter (preserved,

tentacles withdrawn), cylindrical body tapered to rounded

slightly upturned ends orally and anally, distinct ventral

surface, thick firm body wall; external anal scales not detected;

20 tentacles, 5 large pairs alternating with 5 small pairs;

complete cover of tube feet, larger and concentrated ventrally,

disc ends about 0.7 mm diameter, smaller and scattered

dorsally, orally, anally, disc ends about 0.4 mm diameter; tube

feet cross introvert; calcareous ring with 10 posterior composite

digitiform projections arising jointly from both radial and

inter-radial plates, projections each comprise about 6 end-to-

end plates, projections close radially creating long narrow gap

extending higher anteriorly into the radial plates than the wide

inter-radial gap, ring not tubular, posterior prolongations sub-

equal in length with plates and anterior projections, form of

plates and composite components variable; short stone canal,

multi-lobed madreporite close to vascular ring; 1 tubular polian

vesicle; longitudinal muscles with mid-muscle groove; gonad

tubules in series along gonoduct on each side of dorsal

mesentery, not branched.

Mid-body wall with abundant crosses, typically 64 pm

long, some with 1 or 2 blunt surface spines near bifurcate ends,

rare large crosses 88 pm long with bifurcations joined to

create large perforations; tube foot ossicles endplates and

support rods only, endplates with denticulate margin, irregular

perforations similar size, diameters up to 320 pm, endplate

support ossicles predominantly dumbbell-shaped, up to 104

pm long, rounded ends with large and small perforations and

denticulate margin; tentacles with rods and rosettes, larger

rods dumbbell-shaped, widened and perforate distally with

denticulate margin, up to 240 pm long, rosettes up to 40 pm

long; pharynx with abundant rosettes, typically 32 pm long.

Colour. Body red-brown, rusty, not darker orally and anally;

tube feet disc ends off-white; tentacles with white lumps

(rosettes), trunks white, branches pale brown.

Distribution. South Australia, Gulf St. Vincent, intertidal and

offshore sediments.

Remarks. The new generic referral and unique diagnostic

characters of Phyllostauros vercoi (Joshua and Creed, 1915)

are discussed above under the new genus Phyllostauros

O’Loughlin.

As the basis of their erection of this new species Joshua

and Creed 1915 refer specifically to a single specimen collected

by Dr. J. C. Verco, “considerably contracted” and 36 mm long

(SAM K517). H. L. Clark 1938 refers to a loan of two specimens

of Thyone vercoi, 50 and 14 mm long. He further notes that

both came from Le Fevre Peninsula (15 km NW of Adelaide in

Gulf St. Vincent), and thus from sandy shallows. Curiously H.

L. Clark 1946 makes no mention of these two specimens in

discussing Staurothyone vercoi.

The two oldest labels with the holotype specimen of

Phyllophorus ventripes (SAM K1374) record “type M.S.S”

and “type, SA coast, Dr. Verco”. Other recent labels indicate

“holotype”. Rowe and Gates 1995 record “holotype”. We

accept that this specimen is the holotype. A determination by

F. W. E. Rowe in 1977 indicates ‘ Staurothyone ’ vercoi. We

agree with this species determination and synonymy. Although

the specimen is dry it is still possible to extract cross ossicles

from the body wall.

Furthermore we have found that the three paratypes of

Phyllophorus ventripes Joshua and Creed 1915 (NMV F45143)

are conspecific with Thyone vercoi (type status of paratypes

discussed above under Lipotrapeza thyonoides ).

The holotype specimen of Thyone vercoi is dry and in

poor condition but there is an excellent set of slides from the

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

289

type available. The species itself has unique characters within

Phyllophoridae, has never been uncertain in terms of type

status, has not been erroneously reported in the literature, and

has received considerable attention in the literature (see above

under Phyllostauros). On the other hand, for Pliyllophorus

ventripes, considerable confusion has surrounded the history

of the status and identity and description of the type specimens,

and subsequent determination of other material. Thus as “first

revisers” (see ICZN Article 24.2.2) we give name precedence

to Thyone vercoi over Pliyllophorus ventripes in Joshua and

Creed 1915. Pliyllophorus ventripes is a subjective junior

synonym of Thyone vercoi.

Thyonidiella Heding and Panning, 1954

Diagnosis (after Heding and Panning 1954). Dendrochirote

holothuroids with 15 tentacles in 2 series, 10 large outer and 5

small inner; body covered with tube feet; calcareous ring with

composite tapering posterior extensions comprising discrete

single series of plates, arising principally from radial plates but

sometimes also joined with inter-radial plate posterior

extensions; body wall ossicles tables with 4-pillared spires,

sometimes only present in juveniles; endplates with support

plates, not support tables.

Type species. Thyonidiella tenera (Ludwig, 1875) (senior

synonym of original type species Thyonidiella oceana Heding

and Panning, 1954).

Other species. T. cherbonnieri Rowe and Richmond, 2004; T.

drozdovi (Levin and Stepanov, 1999); T. exigua Cherbonnier,

1988; T. kungi O’Loughlin sp. nov.

Remarks. Rowe and Richmond (2004) commented in detail on

genus Thyonidiella Heding and Panning, 1954. They noted that

Thyonidiella oceana Heding and Panning, 1954 is a junior

synonym of Thyonidiella tenera (Ludwig, 1875), and that

Semperiella Heding and Panning, 1954 is a junior synonym of

Thyonidiella Heding and Panning, 1954. They further judged

that Thyonidiella exigua was probably based on a juvenile

specimen of Selenkiella paradoxa Cherbonnier, 1970.

Thyonidiella kungi O’Loughlin sp. nov.

Figures Id, 8

Material examined. Holotype. Australia, Bass Strait, VIMS 81-T-l,

NZOIRV Tangaroa stn 176,38°54.3'S, 147°13.4'E, 58 m, coarse shell,

18 Nov 1981, NMV F76637.

Paratypes. Type locality and date, NMV F174910 (1); VIMS 81-

T-l stn 171, 38°53.7'S, 147°55.2'E, 71 m, shelly sand, 17 Nov 1981,

NMV F76635 (3); VIMS 81-T-l stn 177, 38°53.7'S, 147°06.5'E, 58 m,

coarse shell, 18 Nov 1981, NMV F76638 (5).

Other material. VIMS 81-T-l stn 161, 39°47.3’S, 147°19.3'E, 60

m, muddy shell, 14 Nov 1981, NMV F76636 (1); VIMS 81-T-l stn 174,

39°14.8'S, 147°31.5'E, 57 m, muddy shell, 18 Nov 1981, NMV F76639

(5, very small).

Diagnosis. Up to 16 mm long, up to 5 mm diameter (preserved,

tentacles withdrawn), cylindrical body, rounded end orally,

tapered to rounded end anally, thick soft body wall; external

anal scales not detected; 15 tentacles in 2 series, 5 outer pairs

large, 5 single inner small; complete spaced cover of tube feet,

diameter up to about 0.2 mm; calcareous ring with 10 posterior

composite tapering projections arising in pairs from radial

plates only, posterior projections free thin tails each comprising

about 5 elongate plates, ring not tubular, anterior radial plate

narrowing slightly with deep central split, lacking lateral small

notches, interradial plates not composite; single polian vesicle;

longitudinal muscles flat, not divided.

Mid-body dorsal wall lacking ossicles; tube foot ossicles

endplates and support ossicles only, endplates with denticulate

margin, irregular sub-equal perforations, diameters up to 184

pm, endplate support ossicles dumbbell-shaped, distally

perforate with denticulate margin, up to 120 pm long; tentacles

with rods and rosettes, larger rods dumbbell-shaped, widened

and perforate distally with denticulate margin, up to 304 pm

long, rosettes up to 88 pm long; anal ossicles tube foot

endplates and support rods, tentacle-like rods, rosettes, 5

scales comprising base up to 320 pm wide with digitiform

column up to 240 pm long, base and column comprising

massed short branched rods.

Specimen 3 mm long with radial tube feet; cucumariid

ring lacking posterior prolongations; mid-body ossicles solid

4-pillared tables, irregular multi-perforate discs up to 184 pm

long, high spires with 4 cross connections, up to 136 pm high;

mid-body tube feet with endplates up to 152 pm diameter,

perforations small centrally, large peripherally, margin

denticulate, endplate support rods dumbbell-shaped, up tol84

pm long.

Colour (preserved). Body pale brown.

Etymology. Named for the Taiwanese FRV Hai Kung that

conducted part of the survey of the benthic marine fauna of

eastern Bass Strait for the former Victorian Institute of Marine

Sciences.

Distribution. Eastern Bass Strait, 38-40°S, 147-148°E, 58-71

m, coarse shell, mud and sand.

Remarks. Thyonidiella kungi O’Loughlin sp. nov. is

distinguished from other southern Australia Phyllophoridae

species in the key (above).

Subfamily Semperiellinae Heding and Panning, 1954

Diagnosis. Calcareous ring tubular with long posterior

prolongations, both ring and projections made up of a mosaic

of small pieces; tentacles 20; body wall ossicles tables, each

with a spire of 2 or 4 pillars.

Genera. Cladolella Heding and Panning, 1954; Massinium

Samyn and Thandar, 2003; Neopentadactyla Deichmann,

1944; Neothyonidium Deichmann 1938; Pentadactyla Hutton,

1879; Phyrella Heding and Panning, 1954.

Remarks. We note that Rowe and Richmond (2004) judged that

Semperiella Heding and Panning, 1954 (family Semperiellinae

Heding and Panning, 1954) is a junior synonym of Thyonidiella

Heding and Panning, 1954 (family Phyllophorinae). This

refines the diagnosis for genera of subfamily Semperiellinae,

removing the inclusion of genera with 15 tentacles. We query

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P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

the inclusion in Semperiellinae of genus Phyrella Heding and

Panning, 1954 with its calcareous ring that is not tubular and is

similar to that in genera of Phyllophorinae. We note that

Cladolella Heding and Panning, 1954 has a Massinium-\ ike

calcareous ring (see illustration in Heding and Panning 1954),

20 tentacles, and tables with 2-pillar spires, and might be

considered to be a senior synomym of Massinium Samyn and

Thandar, 2003 with its variety of body wall ossicles, although

the tube feet are radial in Cladolella.

Massinium Samyn and Thandar, 2003

Massinium Samyn and Thandar, 2003: 136—Samyn et al., 2010: 2.

Material examined. Massinium granulosum Samyn and Thandar,

2010. Holotype and paratype. Queensland, Moreton Bay, Stradbroke

I., intertidal seagrass, 14 Nov 1977, AM J13578 (2); Queensland, 32

km NNE of Double Island, AM J2093 (1).

Massinium magnum (Ludwig, 1882). Queensland, Heron I., 10 m,

sandy rubble, 21 Nov 1974, AM J9490 (1).

Diagnosis (Samyn et al. 2010). Dendrochirotid holothuroids

with 20 tentacles arranged in two circles of 10 + 10; anus

usually encircled by calcareous teeth; calcareous ring elongate,

tubular, with both radial and interradial plates fragmented into

a mosaic of small pieces and prolonged posteriorly, with the

posterior processes distally linked to processes of neighbouring

plates forming a loop beneath the water vascular ring; polian

vesicles elongate, usually 4; ossicles of mid-body wall comprise

granuliform rods, and/or rosette-shaped deposits; tables present

in juveniles or occasionally in adults as scarce reduced deposits.

Type species. Massinium maculosum Samyn and Thandar,

2003 (original designation).

Other species. Massinium albicans Samyn et al. 2010; M.

arthroprocessum (Thandar, 1889); M. dissimilis (Cherbonnier,

1988); M. granulosum Samyn et al., 2010; M. magnum (Ludwig,

1882); M. melanieae O’Loughlin sp. nov.; M. vimsi O’Loughlin

sp. nov.; M. watsonae O’Loughlin sp. nov.

Remarks. We have not further emended the generic diagnosis in

Samyn et al. 2010 but note the possibly polyphyletic nature of

Massinium Samyn and Thandar, 2003, with the variety of body

wall ossicle forms in the assigned species that include tables

with 2-pillar spires, pseudobuttons, and thick granuliform rods.

Massinium melanieae O’Loughlin sp. nov.

Figures 13a, 14

Material examined. Holotype. Southern Australia, Great Australian

Bight, Southern Surveyor cruise SS01/00 stn 378, 31°50'S 130°46'E,

55 m, 13/14 May 2000, NMV F174897.

Diagnosis. Massinium species up to 42 mm long, up to 24 mm

diameter (preserved, tentacles withdrawn), slight oral to anal

taper, thin firm body wall; exterior anal scales not detected; 20

tentacles, 5 pairs of large, 5 pairs of small; complete uniform

cover of small and large tube feet, diameter up to 0.4 mm;

white tubular composite calcareous ring 25 mm long; radials

up to 3 mm wide, anterior taper with apical notch; inter-radials

up to 2 mm wide, taper to anterior point, distal indentation

closed by narrow calcareous posterior bridge creating oblong

non-calcareous section; 4 polian vesicles, shorter than

calcareous ring; stone canal extending from end of calcareous

ring with madreporite free in coelom (dissected condition);

cylindrical longitudinal muscles, not divided; gonad tubules

thin, branched.

Lacking mid-body wall ossicles; tube feet with endplates

only, diameter up to 320 pm, irregular perforations, small

central, larger peripheral; tube feet lacking support ossicles;

peri-oral region with abundant regular tables, discs oval to

rounded rectangular, up to 96 pm long, 4 large central and

numerous small peripheral perforations, spires up to 24 pm

long, 2 pillars each with up to 5 blunt apical spines, some bifid;

tentacle branches with thin rods, knobbed or finely perforate

distally, up to 104 pm long; tentacle trunks lacking ossicles;

inner wall of tubular calcareous ring with rods up to 104 pm

long, widened and rounded distally, each end with up to 8

perforations; anal ossicles scales, endplates, tables; scales

single-layered, thick, irregular, perforated; endplates about

160 pm diameter; tables irregular, frequently about 16 disc

perforations, 3 or 4 large central, small peripheral, 2 pillars,

each with blunt apical spines, most discs up to 72 pm long,

pillars about 40 pm high.

Colour (preserved). Body off-white with grey-brown irregular

patches; tentacle trunks grey, branches grey-brown.

Type locality. Southern Australia, Great Australian Bight, 55 m.

Etymology. Named for Melanie Mackenzie, collection curator

in the Department of Marine Biology in Museum Victoria, and

valued colleague in echinoderm systematic research.

Remarks. Massinium melanieae O’Loughlin sp. nov. is erected

for a single specimen in good condition. The type locality is

unlikely to be sampled again in the foreseeable future and we

judge that it is important to erect this species. The distinguishing

combination of diagnostic characters for Massinium melanieae

O’Loughlin is given in the key (above).

Massinium vimsi O’Loughlin sp. nov.

Figures 13b, 15

Material examined. Holotype. Australia, eastern Bass Strait, VIMS,

NZOI RV Tangaroa 81-T-l stn 174, 39°16.8'S 147°32.2'E, 57 m, mud

/ shell, 18 Nov 1981, NMV F76629.

Diagnosis. Massinium species 12 mm long, 5 mm diameter

(preserved, tentacles withdrawn), form cylindrical with

rounded ends, soft body wall; exterior anal scales not detected;

20 tentacles, 5 pairs of large, 5 pairs of very small; complete

cover of tube feet, more prominent ventrally, up to 0.2 mm

diameter; tubular composite calcareous ring 9 mm long, plates

up to 1 mm wide, posterior radial notch, posterior deep inter¬

radial indentation closed posteriorly by continuous band of

small plates to create elongate non-calcareous space; radials

tapered anteriorly with anterior notch; inter-radials tapered to

anterior point; single tubular polian vesicle; stone canal and

madreporite lying anteriorly on calcareous ring; cylindrical

longitudinal muscles, not divided; lacking gonad tubules.

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

291

Figure 14. SEM images of ossicles from holotype of Massinium melanieae O’Loughlin sp. nov. (NMV F174897). Regular tables from peri-oral

body wall (top left); mid-body tube foot endplate (top right); large irregular table from anal region (central right); thin rods from tentacle branches

(bottom right).

292

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 15. SEM images of ossicles from holotype of Massinium vimsi O’Loughlin sp. nov. (NMV F76629). Tube foot endplate (left); tentacle rods

(right); (insert: drawings of table disc, developing disc, and table spire from peri-oral body wall table ossicles).

Lacking mid-body wall ossicles; tube feet endplates up to 184

pm diameter with irregular small perforations and slightly

undulating to denticulate margin, rod-like support plates around

margin of endplates, narrow, curved, perforations distally,

sometimes along rod, large central, small distal, rods up to 96 jim

long; peri-oral region with tables, discs up to 72 jim long, lobed

margin, up to about 18 perforations, spires about 32 jim long, 2

pillars each with 2-3 sometimes bifid spines distally, single cross¬

bridge; introvert ossicles endplates, endplate support rods and

rosettes, endplates up to 136 jim diameter with irregular small

perforations, rods up to 80 jim long, straight to slightly bracket¬

shaped, slightly swollen distally with 1-2 terminal perforations,

rosettes rare; tentacles ossicles rods and rare rosettes, rods up to

96 jim long, curved, ends of rods swollen, smooth or denticulate,

1-2 distal perforations; anal ossicles endplates and endplate

support rods, anal endplates up to 144 jim diameter, undulating

margin, some with slightly larger peripheral perforations, support

rods perforated as in tube feet, up to 80 jim long.

Colour (preserved). Body pale brown; tentacles dark brown.

Type locality and distribution. Southeast Australia, eastern

Bass Strait, 57 m.

Etymology. Named for the former Victorian Institute of Marine

Sciences (VIMS), with appreciation of the thoroughness of the

1981 benthic survey of eastern Bass Strait with a significant

contribution of specimens to this study.

Remarks. Massinium vimsi O’Loughlin sp. nov. is erected for a

single specimen in good condition. The small specimen size

and absence of gonad tubules suggest that it is probably a

juvenile. The type locality is unlikely to be sampled again in

the foreseeable future and we judge that it is important to erect

this species. The distinguishing combination of diagnostic

characters for Massinium vimsi O’Loughlin is given in the key

(above).

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

293

Massinium watsonae O’Loughlin sp. nov.

Figures 13c, d, e, 16

Lipotrapeza dearmatus.—H. L. Clark, 1938: 494.—H. L. Clark,

1946: 411-412. (part; Australian material non Pliyllophorus

dearmatus Dendy and Hindle, 1907).

Neothyonidium dearmatum.— Heding and Panning, 1954: 191—

192 (part; Australian material non Phyllophorus dearmatus Dendy

and Hindle).—Hickman, 1978: 32, pi. 1, figs 15-24 (non Phyllophorus

dearmatus Dendy and Hindle).

Neothyonidium sp.—Rowe, 1982: 466, pis 31.5, 31.6.

Neothyonidium ? dearmatum— Edgar, 1997: 369 (with photo of

tentacles).

Material examined. Holotype. Southeast Tasmania, Bruny Island,

Simpsons Bay, 43°17'S 147°18'E, 11 m, 15 Feb 1972, J. E. Watson,

NMV F97435.

Paratypes. Type locality and date, NMV F174898 (1); NMV

F174899 (1); off Bruny Island, D’Entrecasteaux Channel, 12 m, Feb

1972, J. E. Watson, NMV F97420 (2).

Other material. D’Entrecasteaux Channel, off Green Island, 56 m,

D. F. Turner, 21-25 July 1948, TMAG H169 (1; det. F. W. E. Rowe

1974 as Neothyonidium dearmatum)-, Derwent River, Blackmans Bay,

on beach after storm, E. Turner, 12 Jun 1988, TMAG H1988 (1,

tentacle crown and ring only); Kettering, 43°08’S 147°15'E, 10 m, A. J.

Dartnall and T. Sward, 29 Mar 1977, AM J12385 (1; as Lipotrapeza

dearmatus-, ring damaged posteriorly prohibiting type status); slides

for AM J12385 specimen prepared by V. V. Hickman, tentacles and

body wall TMAG H2089, introvert TMAG H2090, dorsal body wall

TMAG H2091, mid-body wall TMAG H2092, anal TMAG H2093,

posterior TMAG H2094, anal papilla TMAG H2095, cloaca TMAG

H2096; Eastern Tasmania, Great Oyster Bay, no other data, AM J1538

(1). “New South Wales”, no other data, AM J18132 (1; presumed

locality error, SE Tasmania distribution).

Neothyonidium dearmatum (Dendy and Hindle, 1907). New

Zealand, near Cook Strait, 40°S 174°E, 67 m, NIWA 4187 (2); 55 m,

NIWA 73992 (1); off Christchurch, 43°S 173°E, 59 m, NIWA 28443

(1); 44°S 173°E, 54 m, NIWA 76793 (1); 64 m, NIWA 73987 (4); off

Stewart I., 47°S 168°E, 33 m, NIWA 45522 (2).

Diagnosis. Massinium species up to 110 mm long, up to 40 mm

diameter (preserved, tentacles withdrawn), wide orally, tapered

to rounded end anally, thin firm body wall; exterior anal scales

not detected; 20 tentacles, 5 outer pairs large, 5 inner pairs

small; complete cover of tube feet, single series of close tube

feet on each side of longitudinal muscle interior attachment,

scattered inter-radially, across introvert in irregular paired

radial series, diameters up to 0.4 mm; white leathery tubular

composite calcareous ring up to 50 mm long; radials with

anterior taper and notch, posterior notch; inter-radials tapered to

anterior point, posterior inter-radial indentation closed by

continuous band of small plates to create oblong non-calcareous

space; 1-2 tubular polian vesicles, as long and longer than

calcareous ring; long thin stone canal with globular madreporite

attached by mesentery to pharynx; cylindrical longitudinal

muscles, not divided; gonad tubules long thin.

Ossicles disappear with age / size; per-oral tables only in

110 mm long holotype; eroded ossicle remnants only in 70 mm

long paratype (NMV F97420); in specimen 65 mm long (AM

J12385), mid-body wall lacking ossicles; tube feet with few

endplates up to 216 pm diameter, irregular small perforations,

larger peripherally, rare curved support rods up to 80 pm long

around endplate margin; peri-oral region with tables, sometimes

rosettes, table discs oval to rounded, irregular, up to 144 pm

long, up to about 40 perforations, spires with 2 pillars, distal

cross-bar sometimes perforated, pillars with few blunt spines

apically; introvert ossicles tables with oval to rounded

rectangular discs, sometimes rosettes, table discs up to 96 pm

long, up to 24 perforations, 4 large central perforations, many

smaller surrounding ones, spires up to 48 pm long, 2 (rarely 1)

pillars with 2-3 blunt spines apically; larger specimens

tentacles lacking ossicles in trunks and branches, 50 mm long

specimen with rare rods, up to 80 pm long, few distal

perforations; posterior/anal body with tube foot endplates up to

192 pm diameter, rare curved endplate support rods up to 80

pm long; anal papilla with anal scale fragments, tube foot

endplates, rods and rosettes; anal scales comprising multi¬

layered base with digitiform calcareous papilla about 800 pm

long comprising mass of thick linear, X, Y, H shaped rods; anal

tube foot endplates up to 152 pm diameter; numerous short

irregular anal rods intergrading with rosettes, up to 48 pm long.

Colour (preserved). Variable, body pale pinkish-brown with

dark brown patches to uniform pale brown, tentacle trunks

brown to dark brown, branches dark brown to grey-brown.

Type locality. Tasmania, Bruny Island, Simpsons Bay, 11m.

Distribution. Southeast Tasmania, D’Entrecasteaux Channel to

Derwent Estuary; 11-56 m.

Etymology. Named for Jan E. Watson, Honorary Associate of

Museum Victoria, hydroid systematist, significant contributor

to the marine invertebrate collections of Museum Victoria, and

collector of the types of this species.

Remarks. The diagnostic description of ossicles is based

principally on the comprehensive set of V. V. Hickman slides

(see Material examined) of a smaller Kettering specimen (65

mm long, up to 15 mm diameter) registered to the Australian

Museum (AM J12385). The presence and form of tables in the

introvert and absence of ossicles in the tentacles are consistent

with the Blackmans Bay part-specimen, except that there are

also rosettes in the introvert. The presence or absence of

rosettes is judged to be variable in specimens of some

phyllophorid species.

H. L. Clark 1938 referred Phyllophorus dearmatus Dendy

and Hindle, 1907 to his new genus Lipotrapeza with considerable

reservation, based on the entirely different calcareous ring. In

the same year Deichmann erected the genus Neothyonidium to

which Heding and Panning 1954 subsequently referred

Phyllophorus dearmatus. No specimen of Neothyonidum

dearmatum (Dendy and Hindle, 1907) has been found in

Australian waters. Material determined by Joshua (1914) to be

Phyllophorus dearmatus has been examined and found to

represent our new species Thyone flindersi O’Loughlin sp. nov.

(below). Deichmann (in H. L. Clark 1946) correctly judged that

the Joshua specimens should be referred to Thyone.

Hickman (1978) reported a specimen of Neothyonidium

dearmatum from Kettering on the D’Entrecasteaux Channel in

southeast Tasmania, but noted some differences from the

descriptions of this New Zealand species. The specimen is not

294

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 16. SEM images of ossicles from specimen of Massinium watsonae O’Loughlin sp. nov. (TMAG H1988). Tables and rosette from introvert

(top); tables and rosettes from peri-oral body wall (bottom).

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

295

held in the TMAG but ossicle slides prepared by Hickman are

available and were examined by us. The specimen is now held by

the AM and has been examined here. Based on the description

by Hickman (1978) and the ossicle slides, the Kettering specimen

is conspecific with specimens referred here to our new species

Massinium watsonae O’Loughlin sp. nov. (above). Rowe (1982)

referred to the Kettering specimen as Neothyonidium sp.

Based on the descriptions by Dendy and Hindle (1907),

Mortensen (1925), and Pawson (1970) and our observations the

diagnostic characters that distinguish Neothyonidium

dearmatum (Dendy and Hindle) from Massinium watsonae

O’Loughlin sp. nov. (characters in bracket) are: size up to 65 mm

long (up to 110 mm); tentacle rod ossicles present (lacking

tentacle ossicles); inter-radial plates of calcareous ring divided

for most of their length by non-calcareous space (short distal

closed oblong non-calcareous space); distal end of each inter-

radial plate with two narrowing tails that may be contiguous but

not joined by small plates (distal end of calcareous ring a

continuous collar of small plates, the generic diagnostic character

for Massinium Samyn and Thandar, 2003); single polian vesicle

(2 polian vesicles); if present tables in anterior and posterior body

wall (in introvert only); table disc lengths average 75 pm (average

about 90 pm); perforations in table discs usually 8 (up to 24). The

combination of unique diagnostic characters of Massinium

watsonae O’Loughlin sp. nov. is given in the key (above).

Subfamily Thyoninae Panning, 1949

Thy one Oken, 1815

Diagnosis (emended from Pawson and Miller 1981). Tentacles

10; tube feet scattered on body wall, never restricted to radii;

calcareous ring tubular with long posterior prolongations

comprising a mosaic of small pieces; body wall ossicles tables

with a spire of one or two pillars.

Type species. Holothuria fusus Muller, 1776 (monotypy).

Southern Australian species. Thyone flindersi O’Loughlin sp.

nov.; Thyone joshuai O’Loughlin, sp. nov.; Thyone kerkosa

O’Loughlin sp. nov.; Thyone nigra Joshua and Creed, 1915;

Thyone okeni Bell, 1884; Thyone spenceri O’Loughlin sp. nov.;

Thyone tourvillei O’Loughlin sp. nov.

Remarks. We acknowledge the current invalid status of Thyone

Oken, 1815 and our provisional retention is discussed in the

Introduction. We emend the diagnosis of Thyone Oken to

include tables with one pillar in the spire to accommodate a

new species described in this work. Pawson and Miller 1981

remarked on the need for a revision of the “supergenus” Thyone.

This need continues for what will be a major undertaking. The

revision should await further supportive evidence from

emerging genetic data.

Thyone flindersi O’Loughlin sp. nov.

Figures 17a, 18

Phyllophorus dearmatus.— Joshua, 1914: 4-5 (non Phyllophorus

dearmatus Dendy and Hindle, 1907).

Thyone sp. —Joshua and Creed, 1915: 20.

Thyone okeni.— Rowe, 1982: 462 (“provisional referral”; Port

Phillip and Westernport Bays specimens non Thyone okeni Bell,

1884).—Rowe and Gates, 1995: 316 (southern coast specimens non

Thyone okeni Bell, 1884).—Gowlett-Holmes, 2008: 263 (southern

coast specimens non Thyone okeni Bell, 1884).

Material examined. Holotype. Victoria, Flinders ocean platform,

intertidal rocky shallows, 17 Feb 2008, M. O’Loughlin and E.

Whitfield, NMV F151847.

Paratypes. Westernport Bay, McHaffie Reef, 38°28'S 145°10'E,

MRG, 16 Feb 2008, NMV F151848 (1); Shoreham, Honeysuckle Point,

sieved from root of Amphibolus, A. Falconer (MRG), 12 May 2012,

NMV F174912 (1); MRG, 12 Apr 2012, NMV F174909 (1); Flinders,

Mushroom Reef, 17 Feb 1990, NMV F73805 (1).

Other material (selection). Westernport Bay, Merricks, rocky

shallows, 27 May 1989, NMV F73804 (1); Westernport Bay Survey,

stn 57/02/69, Merricks, 22 Feb 1969, NMV F45261 (1; determined by

D. Pawson 1974 as Thyone okeni Bell, 1884); Shoreham, rocky

shallows, Jun 1979, NMV F73806 (1); San Remo, 28 Jan 1909, NMV

F73819 (1) (no. 60641; determined in Joshua 1914 as Phyllophorus

dearmatus Dendy and Hindle, 1907).

Tasmania, Bass Strait, Waterhouse Passage, 40°49’S 147°38'E,

rocky shallows, 23 Nov 1982, NMV F97430 (1).

South Australia, J. C. Verco, Jan 1914, NMV F97433 (1; as Thyone

sp. by Joshua and Creed 1915); Gulf St Vincent, off Adelaide, between

North Haven and Largs Bay, seagrass, 1 Dec 1980, SAM K2585 (2);

Willunga Reef, 23 Nov 1976, SAM K2582 (1); Aldinga Reef, 22 Nov

1979, SAM K2584 (3); Yorke Peninsula, Port Giles jetty, 15 m, Jan

1983, SAM K2595 (1); Wool Bay, 4 m, sand, AM J16647 (2);

Edithburgh Jetty, 3-4 m, in sand, 14 Mar 1994, SAM K2600 (1), SAM

PK0072; Kangaroo Island, Kingscote Jetty, 3-4 m, in sand, 1 May

1999, SAM K2598 (1), SAM PK0191; 3 May 1999, SAM K2599 (2),

SAM PK0183; Sir Joseph Banks Group, Smith Rocks, 28 Jan 1986,

SAM K2607 (3); Waterloo Bay, seagrass, 5 m, 1980, SAM K2586 (2);

Great Australian Bight, Point Westall, near Streaky Bay, 15 Jan 1991,

NMV F97427 (1); near Ceduna, Cape Vivonne, 32°12'S 133°41'E,

rocky shallows, 16 Jan 1991, NMV F97428 (1).

Western Australia, Albany, 35°03'S 117°92'E, 2 m, Jan 1988, AM

J24966 (2); Bunbury, 13 Apr 1963, WAM Z31974 (1); Cockburn

Sound, rubble and muddy sand, 29 Apr 1989, WAM Z31977 (1);

Jervoise Groyne, 1 Jan 1958,3 m, WAM Z31969 (1); Garden I., 29 Apr

1989, WAM Z31984 (1); Mangles Bay, 31 Jul 1988, WAM Z31961 (2);

Woodman’s Point, WAM Z31982 (2).

Diagnosis. Thyone species up to 100 mm long, up to 15 mm

diameter (relaxed, preserved, excluding tentacles, introvert;

WAM Z31984), wide mid-body, tapered to upturned rounded

oral and anal ends, thick firm body wall; external anal scales

not detected; 10 tentacles, 8 large, 2 much smaller ventrally;

complete cover of spaced tube feet, contiguous paired series on

each side of longitudinal muscle interior attachments, more

scattered inter-radially, more abundant ventrally than dorsally,

diameter about 0.2-0.3 mm; tubular composite calcareous

ring, posterior ends of composite radial plates not joined;

radials tapered anterior with terminal split, divided posterior

into paired narrowing composite tails, tails sometimes split;

inter-radials tapered to anterior point, posterior ends truncate at

a point anterior to the division of each radial; short stone canal

and globular split-pea-like madreporite lie on the calcareous

ring; single elongate thin polian vesicle; longitudinal muscles

cylindrical to flat, undivided; gonad tubules unbranched.

Mid-body wall lacking ossicles; tube feet with endplates only,

endplates with irregular perforations, distinctly larger peripherally,

296

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 17. Photos of preserved specimens, a, holotype of Thy one flindersi O’Loughlin sp. nov. (NMV FI 51847) (insert: form of the calcareous

ring plates); b, specimen of Thyone okeni Bell, 1884 (AM J10868); c, holotype of Thyone joshuai O’Loughlin sp. nov. (SAM K2566) (insert: form

of the calcareous ring plates); d, holotype of Thyone kerkosa O’Loughlin sp. nov. (WAM Z31838); e, holotype of Thyone spenceri O’Loughlin

sp. nov. (SAM K2562) (insert: form of the calcareous ring plates); f, holotype of Thyone tourvillei O’Loughlin sp. nov. (NMV F174902) (insert:

form of the calcareous ring plates).

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

297

Figure 18. SEM images of ossicles from paratype of Tliyone flindersi O’Loughlin sp. nov. (NMV F174909). Ossicles from peri-oral body wall

(top); rods from tentacles (bottom).

298

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

diameters up to 160 pm: lacking endplate support rods or plates

or tables; per-oral body wall with tables and sometimes rosettes,

table discs irregular, rounded rectangular to oval to rounded

triangular, up to 35 perforations, discs up to 72 p m long, spires

with 2 pillars, single cross connection, blunt spines distally, spires

up to 32 pm high, rosettes predominantly oval up to 40 pm long;

anal ossicles scales with spires, tube foot endplates, rosettes, rods,

sometimes tables; scales thick, single-layered with lace-like

thickening, fragment 440 pm long, multilayered spire 424 pm

long, anal endplate diameters up to 136 pm, rosettes up to 48 pm

long, rods with finely perforated ends, some with short branching

distally, up to 96 pim long, table discs up to 56 pim long, up to > 8

perforations, spires with 2 pillars; tentacles branches with rods

predominantly bracket-shaped, distally with lumps or fine

branches or few perforations, some rods with sigmoid form, some

straight, rods up to 88 pim long, tentacle trunks sometimes with

rosettes, round to oval to rod-like, up to 56 pim long.

Colour (live and preserved). Body brown to dark reddish

brown, body frequently dark grey orally and anally, introvert

brown; tube feet red to dark brownish red, off-white orally and

anally; tentacle trunks brown, lacking white spots (rosette

clusters), tentacle branches dark grey.

Distribution. Southern Australia, east to San Remo

(Westernport Bay), northern Tasmania (Bass Strait), west to

Cockburn Sound (Western Australia near Perth), rocky

shallows, seagrass; 0-15 m.

Etymology. Named for the type locality, Flinders, in Victoria,

in turn named for the English explorer Matthew Flinders who

first charted the coastline of Victoria.

Remarks. The specimen reported in Joshua and Creed 1915 as

Thyone sp. is registered to Museum Victoria and determined as

Thyone flindersi O’Loughlin sp. nov. A specimen from San

Remo collected in 1909 and reported in Joshua 1914 as

Phyllophorus dearmatus Dendy and Hindle, 1907 was examined

and determined as Thyone flindersi. Rowe 1982 “provisionally”

referred Port Phillip and Westernport Bay specimens to Thyone

okeni Bell. Thyone flindersi is distinguished from other southern

Australia Thyone species in the key (above).

Thyone joshuai O’Loughlin sp. nov.

Figures 17c, 19

Material examined. Holotype. South Australia, Upper Spencer Gulf,

Whyalla, 33°02'S 137°40'E, 12 m, E. Oks, Sep 1987, SAM K2566.

Paratypes. South Australia, Upper Spencer Gulf, SA Department

of Fisheries, Backy Point W, 18 m, Sep 1987, SAM K2560 (1); Feb

1986, SAM K2563 (1); East Shoal NE, 8 m, Feb 1986, SAM K2569 (3);

Lowly Point W, 24 m, Sep 1987, SAM K2571 (4); Feb 1987, SAM

K2572 (2); False Bay, 10 m, Sep 1987, SAM K2573 (1); Cockle Spit W,

7 m, Aug 1976, SAM K2574 (1); Ward Spit S, 9 m, Feb 1987, SAM

K2576 (1); Douglas Bank NE, 11 m, Feb 1987, SAM K2578 (1); Fairway

Bank W, 13 m, Sep 1987, SAM K2580 (1); Whyalla, 12 m, Sep 1987,

SAM K2581 (1); Port Bonython, 15 m, Sep 1987, SAM K2622 (1).

Other material. Victoria, Eastern Bass Strait, 38-40°S 143-149°E;

VIMS, FRV Hai-Kung 81-HK-l, stn 119, 92 m, 31 Jan 1981, NMV

F76632 (2); stn 120, 84 m, sand, 31 Jan 1981, NMV F76630 (4); VIMS,

NZOI RV Tangaroa 81-T-l stn 162, 51 m, shell, 14 Nov 1981, NMV

F76631 (1); 81-T-l stn 167,124 m, mud/fine sand, 14 Nov 1981, NMV

F76633 (1; 2.5 mm long); stn 170, 140 m, muddy sand, 15 Nov 1981,

NMV F76624 (3; up to 5 mm long); 81-T-l stn 174, 57 m, mud/shell,

18 Nov 1981, NMV F76628 (1); stn 174, 57 m, mud/shell, 18 Nov 1981,

NMV F76634 (1; 3 mm long); stn 180, 65 m, mud/sand, 18 Nov 1981,

NMV F76626 (1; 1.5 mm long); stn 200, 48 m, sand/shell, 22 Nov

1981, NMV F76625 (1; 2 mm long).

New South Wales, Disaster Bay, RV Southern Surveyor SS0404

stn 102, 37°17.22'S 150°04.15'E, 79 m, 28 Apr 2004, NMV F132693

(1); Port Hacking, Jibbon Head, 34.0708°S 151.1306°E, 23 m, Sep

1976, AM J22366 (1).

Diagnosis. Thyone species up to 21 mm long, up to 8 mm

diameter (preserved, tentacle withdrawn), body fusiform, tapered

to rounded end orally and anally, short tail anally, soft thin body

wall; external anal scales not detected; 10 tentacles, 8 large, 2

ventral small; tube feet in close single paired series on radii,

scattered inter-radially, diameter about 0.15 mm; composite

calcareous ring, anterior third tubular, posterior ends of

composite radial plates not joined, radials tapered anteriorly with

terminal split, deep posterior division more than half composite

plate length, paired composite ends thin, narrow, free; inter-

radials tapered to anterior point, truncate posteriorly about where

the radial plates split, inter-radial composite plates about one

third the length of the radial plates; gonad tubules unbranched;

long stone canal, globular madreporite; single polian vesicle,

elongate, tubular; longitudinal muscles sub-cylindrical.

Mid-body wall ossicles scattered tables, discs regular, oval,

margin slightly lobed, perforations typically 4 large central,

frequently 4 small corner, discs 56-64 pm long, spires with 2

pillars, distally spinous, single distal cross connection, spires

up to 32 pm long; tube feet with endplates, variably with and

lacking tube foot support tables, endplates with distinct outer

ring of large perforations, small central perforations, endplate

diameters up to 160 pm, if present tube foot support tables with

curved narrow discs up to 112 pm long; tentacles ossicles stout

and fine rods and rosettes, rods irregular, up to 200 pm long,

frequently with 1 or 2 side branches bluntly spinous distally,

widened perforated ends, rosettes up to 64 pm long, some inter¬

grade with rods; anal ossicles tables and scales, tables as in

mid-body wall, scales single and multi-layered.

Small specimens (10 mm long): abundant mid-body tables,

discs with typically 4 perforations, regular, lobed margin, 40

pm long; tube feet with endplates, ring of outer large perforations,

88 pm diameter, tube foot support tables with narrow curved

discs, 72 pm long; anal tables as in larger specimen mid-body.

Smallest specimens (1.5-3 mm long): mid-body tables

regular with 8 perforations and irregular with many more than

8 perforations, discs up to 128 pm long, spires up to 48 pm

long; tube feet table discs curved and narrow with more than 8

perforations, discs up to 112 pm long, large mesh endplates,

112 pm diameter.

Colour(preserved). Body off-white, sometimes fine brown

flecking; tentacles brown to pale brown.

Distribution. Off-shore sediments, Victoria, eastern Bass

Strait, 39-40°S 143-149°E, 48-140 m; New South Wales,

Disaster Bay, Port Hacking, 34-37°S 150-151°E, 23-79 m;

South Australia, Upper Spencer Gulf, 32-33°S 137°E, 7-24 m.

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

299

Figure 19. SEM images of ossicles from specimens of Thyone joshuai O’Loughlin sp. nov. Tables from mid-body wall (right), tube foot endplate

(top), and tube foot support tables (central and left bottom) (from holotype, SAM K2566); rods from tentacles (top left) (specimen NMV F76631).

300

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Etymology. Named for E. C. Joshua, who described the first

phyllophorid species on the coast of southern Australia.

Remarks. Significant changes with age/size in body wall and

tube foot ossicle forms are recorded in the diagnosis (above).

Notable changes are the increase in size and number of

perforations in tables with decreasing size of specimen, the

mesh-like form of endplates with decrease in specimen size,

and the loss of tube foot support tables with increasing

specimen size. The morphological characters that distinguish

Thyone joshuai O’Loughlin sp. nov. from other southern

Australian Thyone species are given in the key (above).

Thyone kerkosa O’Loughlin sp. nov.

Figures 17d, 20

Material examined. Holotype. Southern Australia, Recherche

Archipelago, Sandy Hook Island, 34°02'S 122°00'E, Scallop Trawl

Survey 1986, L.F.B.E. Triumph , D. Richards et al., 26 July 1986, WAM

Z31838 (body with tentacles and ring eviscerated and lost).

Diagnosis. Body 45 mm long (preserved), up to 15 mm

diameter, cylindrical, tapered to rounded oral and anal ends,

body wall soft, thick; external anal scales not detected; tentacles

lost; complete close cover of small tube feet, diameter 0.3 mm;

calcareous ring lost.

Body wall with table ossicles only, tables range greatly in

form and inter-grade; numerous small tables with discs rounded

rectangular with smooth lobed margins, smallest with typically

8 perforations, discs 48-56 pm long, spires with 2 pillars, short

blunt pointed spines apically, spires 16 pm long; medium-sized

tables with elongate oval discs, sometimes bi-lobed, up to about

40 small perforations, no perforations between disc margin and

2 large central perforations adjacent to spire, discs up to 144

pm long, spires with 2 pillars, up to 3 cross-connections, spires

up to 64 pm long; large tables with irregular narrow tapered

discs, 2 large central perforations, up to about 30 small

perforations, ends of disc sometimes extended into solid non-

perforate ‘tail’, sometimes solid digitiform side branches off

discs, discs and tails up to 124 pm long, spires with 2 pillars,

solid, up to 3 small perforations, 2 long laterally extended

apical points together as wide as disc length, pillars 48 pm

long; tube feet with endplates and table support ossicles;

endplates up to 120 pm diameter; tables with curved discs; anal

ossicles tables, single and multi-layered scales.

Colour (preserved ). Body pale brown.

Distribution. SW Australia, Recherche Archipelago.

Etymology. Named from the Greek kerkos (tail), with reference

to the elongate tail-like extensions on some narrow table discs

and some table spires apically.

Remarks. We are conscious of the limited morphological data

available for the single damaged specimen on which we are

erecting Thyone kerkosa O’Loughlin sp. nov. But we judge that

it is important to establish this distinctive species in a work that

is attempting a comprehensive treatment of southern Australian

phyllophorid species. We are confident that the species belongs

to genus Thyone Oken because of the distribution and size of

tube feet, and similarity of the tables to those of another new

Thyone species from the Upper Spencer Gulf that has a

characteristic Thyone ring and tentacles (see below). Thyone

kerkosa is distinguished from other southern Australian

Thyone species in the key (above).

Thyone nigra Joshua and Creed, 1915

Figures 5b, 21

Thyone nigra Joshua and Creed, 1915: 20-21, pi. 3 figs 3a-e, 4.—

H. L. Clark, 1946: 401 .-A. M. Clark, 1966: 347-348, figs 10b, c.-

Rowe, 1982: 462, fig. 10.29a.—Rowe and Gates, 1995: 316.

Material examined. Holotype. South Australia, Dr. Verco, SAM

K1376 (specimen and slide).

Other material. Victoria, Westernport Bay, McHaffie Reef, 16 Feb

2008, NMV F151855 (1; photo live); Port Phillip Bay, Corio Bay, PPS,

Area 26 stn 300-301, 6 m, 16 May 1963, NMV F73807 (2); PPS, Corio

Bay, Area 27 stn 41, 3 m, 16 Feb 1958, NMV F73808 (1).

Tasmania, Bass Strait, Waterhouse Passage, 40°49'S 147°38'E,

rocky shallows, 23 Nov 1982, NMV F97425 (2); 26 Feb 1991, NMV

F174901 (1).

South Australia, Upper Spencer Gulf, Backy Point, 18 m, Sep

1987, SAM K2561 (1); Ward Spit, 9 m, Feb 1986, SAM K1799 (1);

Eyre Peninsula, Elliston, 33°39'S 134°53'E, rocky shallows, 18 Jan

1991, NMV F97426 (1); Point Westall (near Streaky Bay), 32°55'S

134°04'E, 0 m, 15 Jan 1991, NMV F97427 (1); Great Australian Bight,

Point Sinclair, 32°06'S 132°59'E, 10 May 1973, NMV F97424 (3).

Western Australia, Busselton, 1 Mar 1962, WAM Z31985 (1);

Cockburn Sound, Woodman’s Point, 13 Feb 1972, WAM Z31983 (4); 1

m, 25 Aug 1971, WAM Z31963 (2); Jervoise Groyne, 2 m, 1 Nov 1959,

WAM Z31835 (1); Perth, Ocean Reef boat harbour, 10-14 m. May

1990, WAM Z31981 (1).

Diagnosis. Thyone species up to 35 mm long, up to 11 mm

diameter (preserved; NMV F151855 specimen 42 mm long

live, 17 mm long preserved), body fusiform, tapered to rounded

end orally and anally, sometimes oral and anal ends upturned,

thick firm body wall; external anal scales not detected; 10

tentacles, 8 large, 2 ventral small; complete cover of spaced

tube feet, paired to multiple series of contiguous tube feet

radially, scattered inter-radially, diameter about 0.2 mm;

tubular composite calcareous ring, posterior ends of composite

plates not joined; radials tapered anteriorly with terminal split,

deep posterior division about half composite plate length,

paired composite ends narrow, free; inter-radials tapered to

anterior point, truncate posteriorly, inter-radial composite plate

extending about half the length of the radial plate; stone canal

attached to side of calcareous ring, split pea madreporite form;

single polian vesicle, elongate, tubular; longitudinal muscles

cylindrical; gonad tubules unbranched.

Mid-body wall ossicles tables, discs irregularly oval to

rounded rectangular with 4 central large perforations, 4 and up

to about 26 outer smaller perforations, small table discs

typically 80 pm long, large table discs up to 120 pm long,

tables inter-grade with endplate support tables with narrow

curved discs up to 136 pm long, table spires with 2 pillars,

single cross-bar, spinous distally, spire height up to 32 pm\

tube feet with endplates, endplate support tables (above);

endplates with irregular perforations, small centrally, large

marginally, up to about 160 pm diameter; peri-oral body wall

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

301

Figure 20. SEM images of ossicles from holotype of Thyone kerkosa O’Loughlin sp. nov. (WAM Z31838). Mid-body wall tables small and large

tables with attenuated discs and distal spires, and tube foot endplate (bottom left).

302

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 21. SEM images of ossicles from specimen of Thyone nigra Joshua and Creed, 1915 (NMV F151855). Mid-body wall tables and tube foot

support tables with narrow curved discs.

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

303

ossicles tables, rosettes, tables irregularly oval to elongate, 2

pillars, up to 30 disc perforations, discs up to 90 pm long,

rosettes oval to bi-lobed, up to 50 pm long; tentacles rarely

with rods, perforated ends, predominantly straight and curved

more than bracket-shaped, up to 72 p m long, rods inter-grade

with rosettes, rosettes abundant, many linear.

Colour (preserved). Body white with dark brown flecking/

blotching, darker to black orally and anally, tube feet white

with rare dark brown flecks; tentacle trunks brown with white

spots (clusters of rosettes), tentacle branches dark grey.

Type locality. South Australia (precise collecting locality not

recorded by collector Dr J. C. Verco).

Distribution. Southern Australia, east to Westernport Bay,

north Tasmania (Bass Strait), west to Perth (Western Australia),

rocky shallows and offshore sediments, 0-18 m.

Remarks. Thyone nigra Joshua and Creed, 1915 is distinguished

from other southern Australia Thyone species in the key (above).

Thyone okeni Bell, 1884

Figures 17b, 22

Thyone okeni Bell, 1884: 149-150, pi. 9D.-H. L. Clark, 1921:

167.—H. L. Clark, 1946: 402—Rowe, 1982: 462 (non specimens from

Port Phillip and Westernport in Victoria).—Rowe and Gates, 1995:

316 (non specimens from southern Australia).

Material examined. Australia, New South Wales, Port Jackson, AM

J16905 (1); Vaucluse, Bottle and Glass Rock, under boulders, 21 Nov

1968, AM J7731 (1); 5 m, 29 Aug 1977, AM J10868 (2); mid-tide rock

pools, 23 Oct 1968, NMV F174900 (1); off Nielsen Park, dredged, 21

Jan 1951, AM J16869 (1); Newcastle, Swansea Channel, 5 m, 21 May

1987, AM J20267 (1).

Diagnosis. Thyone species up to 75 mm long, up to 20 mm

diameter (preserved, tentacle withdrawn; 85 mm long by Bell

1884); body cylindrical, oral and anal ends tapered, upturned;

typically 10 tentacles, 8 large, 2 small ventral; body closely

covered with tube feet, about 0.3-0.4 mm diameter, not more

concentrated radially; no external evidence of anal scales or

papillae; calcareous ring composite, tubular, no evidence of

discrete plates for most of length, posterior composite tails not

joined distally; radial plates with split anterior point, posterior

end divided into two thin composite tails posterior to the end of

the inter-radial plates; inter-radial plates tapered to anterior

point, truncate posteriorly, about two thirds the length of radial

plates; single long tubular polian vesicle; stone canal and

madreporite lie anteriorly from attachment along calcareous

ring; longitudinal muscles cylindrical, distinctly divided by

deep groove mid-body.

Mid-body wall lacking ossicles; tube feet with endplates

only, up to 240 pm diameter lacking any support rods or tables

or plates, endplate with irregular perforations, large marginally

small centrally, slightly denticulate margin; peri-oral body

wall with numerous tables, discs round to slightly oval, up to

192 pm diameter/length, up to > 100 small perforations, spires

thick short columnar mesh; tentacles branches with stout

dumbbell-shaped ossicles, few to about 12 distal perforations,

rods up to 320 pm long, tentacle trunks lacking ossicles, no

rosettes seen; introvert ossicles rosettes; anal ossicles rods as

in tentacles, rosettes as in introvert, single and multi-layered

scale fragments, tube foot endplate diameters up to 200 pm.

Colour (preserved). Body brown to dark brown to grey-brown;

tube feet off-white to grey to speckled to red; tentacle trunks

off-white with brown patches, branches dark brown.

Type locality. New South Wales, Port Jackson.

Distribution. New South Wales, Port Jackson to Newcastle;

0-5 m.

Remarks. The distribution extensions of Thyone okeni Bell,

1884 to Thursday Island off Cape York in northern Queensland

(H. L. Clark 1921) and to Guam in the eastern Pacific (Rowe and

Doty 1977) are not confirmed. Thandar 1990 followed the

comment by Rowe and Doty 1977 that Thyone okeni Bell was

“probably distributed throughout the western Pacific area”, and

without examining specimens was “inclined to think that T.

venusta Selenka, 1868 and Thyone okeni were conspecific” and

then formally relegated Thyone okeni Bell, 1884 to junior

synonymy with Thyone venusta Selenka, 1868 (type locality the

Red Sea). In the key in his paper Thandar (1990) described the

body of Thyone venusta Selenka, 1868 as being violet at both

ends, and the tube feet as lacking ossicles. Neither character is

true of Thyone okeni Bell, 1884. There are distinctive dumbbell¬

shaped endplate support rods in the tube feet of Thyone okeni

Bell. We reject this synonymy on the basis of inadequate

systematic evidence for these two species with widely separated

type localities. Thyone okeni Bell, 1884 is distinguished from

other southern Australia Thyone species in the key (above).

Thyone spenceri O’Loughlin sp. nov.

Figures 17e, 23

Material examined. Holotype. South Australia, Upper Spencer Gulf,

Backy Point - West, 32°50'S 137°50.52'E, 18 m, SA Fisheries

Department, Sep 1987, SAM K2562 (ring, tentacle crown, anterior

part of body wall).

Paratype. Upper Spencer Gulf, Port Bonython, 33°01'S 137°45'E,

15 m, SA Fisheries Department, Sep 1987, SAM K2565 (1, ring,

tentacle crown, anterior part of body wall).

Diagnosis. Anterior body and calcareous ring 13 mm long

(preserved), body wall soft; 10 tentacles, 8 large 2 ventral

small; complete close cover of small tube feet; calcareous ring

composite, tubular, posterior ends of composite plates not

joined; radial plates tapered anterior with terminal split, long

paired composite posterior prolongations separated for most of

length, narrowing and further divided distally for about a

quarter of total plate length, not joined; inter-radial plates

composite, tapered to anterior point, truncate posteriorly; stone

canal attached to alimentary canal mesentery, ‘split-pea’

madreporite form; single polian vesicle.

Anterior body wall with abundant tables, tables regular,

discs round to oval to triangular, up to 96 pm long, 2 large

central perforations, smaller perforations nearer margin,

predominantly 40-50 small perforations, spires with 2 pillars,

24 pm long, 2 tapered ends distally; some table discs rosette¬

like; some table discs curved for tube foot support; tentacles

304

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 22. SEM images of ossicles from specimen of Thyone okeni Bell, 1884 (AM J7731). Peri-oral body wall rosette and tables with multi-

branched spires (top); dumbbell-shaped ossicles from tentacles (bottom).

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

305

Figure 23. SEM images of ossicles from paratype of Thyone spenceri O'Loughlin sp. nov. (SAM K2565). Tables from anterior body wall (top);

plates and rosettes from tentacles (bottom).

306

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

with abundant perforated plates, rosettes; plates variable in

shape, some plates narrow, elongate with large perforations and

distal blunt teeth, sometimes small branches along margin of

plate, plates up to 280 pm long, some plates cross-shaped with

perforate arms, some plates oval with four large central

perforations, smaller surrounding ones and distal blunt teeth,

plates up to 160 pm long, some plates curved with denticulate

margin and numerous small perforations, plates up to 120 pm

long; rosettes oval, up to 88 pm long; peri-oral body wall with

tables and rods; tables with 2 spires, discs with 2 central large

perforations, small perforations between central large ones and

disc margin, each disc with up to more than 30 small perforations,

discs about 104 p m long; peri-oral ossicles irregular wide

perforated rods, up to 272 p m long, sometimes crosses.

Colour (preserved). Body off-white; tentacles brown with dark

brown to black markings.

Distribution. South Australia, Upper Spencer Gulf, 15-18 m.

Etymology. Named with reference to Spencer Gulf in South

Australia where the type specimens were collected.

Remarks. Both the holotype and paratype of Thyone spenceri

O’Loughlin sp. nov. are damaged and most of the body wall has

been lost. But there are good morphological systematic

characters for the erection of this new Thyone Oken species.

Thyone spenceri is distinguished from other southern

Australian Thyone species in the key (above).

Thyone tourvillei O’Loughlin sp. nov.

Figures 17f, 24

Material examined. Holotype. Eastern Tasmania, 37 km NE of Cape

Tourville, RV Franklin stn SLOPE 85, 41°56'S 148°35'E, upper

continental slope, 124 m, G. C. B. Poore et al., 30 Oct 1988, NMV

F174902 (1, small, damaged, 15 mm long).

Paratype. Eastern Bass Strait, VIMS, RV Tangaroa, 81-T-l stn

170, 38°52.6’S 148°25.2’E, 140 m, mud / sand, 15 Nov 1981, NMV

F76627 (1, small, damaged, 6 mm long).

Diagnosis. Thyone species 15 mm long (preserved body and

partly extended tentacles), up to 3 mm diameter, body

cylindrical, rounded ends, body wall soft; external anal scales

not detected; 11 tentacles (some lost in holotype), large pairs,

small singly; complete close cover of tube feet, single contiguous

series on each side of longitudinal muscle interior attachments,

more scattered inter-radially, diameter about 0.2 mm; composite

calcareous ring; radial plates tapered anteriorly with terminal

split, long paired composite posterior prolongations, narrowing

distally; inter-radial plates tapered to anterior point, not

composite, lacking posterior extensions, contiguous postero-

laterally with composite radial posterior extensions; stone canal

and madreporite lie on calcareous ring; single polian vesicle.

Mid-body wall with tables, regular, discs oval, up to 64 pm

long, 4 narrow perforations in cross formation centrally, 4

larger corner perforations, single pillar spire with apical spines,

spire 24 pm long; tube feet with endplates and endplate support

tables and plates; endplates with small central perforations,

transversely elongate perforations marginally, marginal

denticulations, diameters up to 144 pm\ tube foot support tables

with curved narrow disc, frequently 4 central perforations, 2

distally, discs up to 88 pm long, single pillar spire 24 pm long;

tube foot support plates, slightly curved, sub-rectangular, larger

perforations centrally, smaller distally, digitiform projections

on one edge, up to 160 pm wide; tentacles with rods, tables,

rosettes; rods up to 280 pm long, up to 4 distal perforations, up

to 2 spines along shaft; rosettes up to 80 pm long.

Colour (preserved). Body and tube feet off-white, tentacles

pale brown.

Type locality. Eastern Tasmania, off Cape Tourville, 124 m.

Distribution. Eastern Bass Strait and Tasmania, 124-140 m.

Etymology. Named for the type locality, Cape Tourville, in

eastern Tasmania; in turn named for the French naval

commander Comte de Tourville by the explorer Nicolas Baudin.

Remarks. Both the holotype and paratype of Thyone tourvillei

O’Loughlin sp. nov. are damaged, but calcareous components

are well-preserved and the specimens sufficiently intact to

provide distinctive diagnostic characters for the new species.

The diagnostic characters of this new species are not those of

any current phyllophorid genus.

Thandar 1989 erected subfamily Sclerothyoninae Thandar,

1989 with two monotypic genera Sclerothyone Thandar, 1989

and Temparena Thandar, 1989 to accommodate two previously

erected species. In both genera the calcareous ring is not

tubular and is similar to that in the new species Thyone

tourvillei. But in both Thandar genera the tube foot distribution

is radial only, there are eight large and two small tentacles, and

the tables have spires with two pillars. Neither genus is suitable

for referral of the new species.

We refer this new species to Thyone Oken on the basis of

tube feet and ossicle characters, but with considerable reservation

because of the single pillar spire tables, Sclerothyoninae-like

calcareous ring, and uncertain tentacle state. We are reluctant to

erect a new genus on damaged and very small specimens, with

uncertainty about tentacle characters, and with a major revision

needed of genus Thyone. Thyone tourvillei is distinguished from

other southern Australian Thyone species in the key (above).

Acknowledgments

We are most grateful for the invaluable contribution to our

work of the following: Stephen Keable (AM), Kareen Schnabel,

Niki Davey and Sadie Mills (NIWA), Thierry Laperousaz

(SAM), Liz Turner and Ruth Mollison (TMAG), Mark Salotti

and Jane Fromont (WAM) for their gracious assistance with

loan materials; Leon Altoff, Audrey Falconer and John Eichler

(MRG) for providing live specimen photographs; Melanie

Mackenzie and Chris Rowley for their technical assistance in

the Marine Invertebrate Department (NMV); Ben Boonen for

the preparation of the figures; Gustav Paulay and Francois

Michonneau (University of Florida), David Pawson

(Smithsonian Institution), Frank Rowe (Australian Museum),

and Ahmed Thandar (University of KwaZulu-Natal) for their

helpful and valued communications on systematic issues. We

are especially grateful for the review advice offered by Dr. F.

W. E. Rowe (Australian Museum).

The phyllophorid sea cucumbers of southern Australia (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae)

307

Figure 24. SEM images of ossicles from holotype of Thyone tourvillei O'Loughlin sp. nov. (NMV F174902). Mid-body wall tables with single

pillar spires (top); rods and rosettes from tentacles (bottom).

308

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

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species in the phyllophorid genus Massinium (Echinodermata:

Holothuroidea) with redescription of Massinium magnum.

Zootaxa 2399: 1-19.

Selenka, E. 1868. Nachtrag zu den Beitragen zur Anatomie und

Systematik der Holothurien. Zeitschrift fiir Wissenschaftliche

Zoologie 18: 109-119 pi. 8.

Smith, B. J. 1970. Catalogue of echinoderm types in the National

Museum of Victoria, Australia. Memoirs of the National Museum

Victoria 31: 91-96.

Thandar, A. S. 1989. The sclerodactylid holothurians of southern

Africa, with the erection of one new subfamily and two new

genera (Echinodermata: Holothuroidea). South Africa Journal of

Zoology 24(4): 290-304.

Thandar, A. S. 1990. The phyllophorid holothurians of southern

Africa with the erection of a new genus. South Africa Journal of

Zoology 25(4): 207-223.

Memoirs of Museum Victoria 69: 309-325 (2012)

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

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

A review of Australian fossil penguins (Aves: Sphenisciformes)

Travis Park 1 and Erich M.G. Fitzgerald 2

1 School of Life and Environmental Sciences, Deakin University, Vic. 3125, Australia and Geosciences, Museum Victoria,

GPO Box 666, Melbourne, Vic. 3001, Australia (tpark@museum.vic.gov.au)

2 Geosciences, Museum Victoria, GPO Box 666, Melbourne, Victoria 3001, Australia (efitzgerald@museum.vic.gov.au)

Abstract Park, T. and Fitzgerald, E.M.G. 2012. A review of Australian fossil penguins (Aves: Sphenisciformes). Memoirs of Museum

Victoria 69: 309-325.

Australian fossil penguins (Sphenisciformes) are reviewed as a basis for future primary research. The five named

species are based on type specimens of Eocene, Miocene—Pliocene and Holocene age collected from South Australia,

Victoria and Tasmania. The phylogenetic affinities of these taxa remain unresolved. Only one type specimen is represented

by clearly associated elements of a skeleton; the rest are single bones (isolated partial humeri and a pelvis). Further

research is required to establish the taxonomic status of Pachydyptes simpsoni, Anthropodyptes gilli, Pseudaptenodyes

macraei, ?Pseudaptenodytes minor and Tasidyptes hunteri. Additional described specimens include isolated postcranial

elements from the Late Oligocene of South Australia and Late Miocene—Early Pliocene of Victoria. Other Miocene and

Pliocene specimens are housed in Museum Victoria. These specimens have the potential to shed fight on the Neogene

palaeobiogeography and diversification of crown group penguins.

Keywords South Australia, Tasmania, Victoria, Cenozoic, evolution, bird

Introduction

The published fossil record of penguins (Sphenisciformes) in

Australia, although limited compared to that of Antarctica,

New Zealand and South America, spans some 40 million

years from the late Eocene to Recent (Ksepka and Ando,

2011). The majority of previous work has been produced by

one author (Simpson, 1957, 1959, 1965, 1970) with the last

primary research conducted by Van Tets and O’Connor (1983).

Since then, several summaries have been published (Jenkins,

1985; Fordyce & Jones, 1990; Vickers-Rich, 1991). Current

work underway by the authors indicates that many undescribed

diagnostic specimens reside in museum collections. The aim

of this work is to introduce the Sphenisciformes, summarise

current knowledge of Australian fossil penguins, discuss

implications for penguin evolution, and outline both gaps in

knowledge and opportunities for future research.

Definitions and Terminology

This review follows the traditional classification of penguins

where Sphenisciformes is the Order to which the single family,

Spheniscidae, belongs. Spheniscidae contains all known

species of penguin, fossil and extant. The terms

Sphenisciformes and Spheniscidae are therefore used

interchangeably throughout this paper. Osteological

terminology and terms of orientation follows that of Baumel

and Witmer (1993).

Institutional Abbreviations

ANWC, Australian National Wildlife Collection, CSIRO

Division of Wildlife and Rangelands Research, Canberra,

Australia; NM V P, Museum Victoria Palaeontology Collection,

Melbourne, Australia; SAM P, South Australian Museum

Palaeontology Collection, Adelaide, Australia.

The Sphenisciformes

Sphenisciformes (penguins) are a group of flightless marine birds

confined to the southern hemisphere. Contrary to popular

stereotype, not all species reside in Antarctica, with the highest

species diversity found in New Zealand (Ksepka et al., 2012) and

one species (the Galapagos penguin, Spheniscus mendiculus

Sundevall, 1871 actually living at the equator (Vargas et al., 2005;

Jadwiszczak, 2009). Fossil species are found in the same regions

as extant species (Simpson, 1975), with Antarctica, Australia,

New Zealand, South Africa, and South America all possessing

both fossil and extant assemblages. Present regional species

diversities roughly correspond to past levels, with areas such as

New Zealand and Antarctica well represented by numerous fossil

and living species, despite the fossil record not being continuous

throughout the Cenozoic (Ksepka & Ando, 2011; Ksepka et al.,

2012)(Fig. 1). One of the most specialised avian groups (Kaiser,

2007), the morphology of living penguins is well known (Pycraft,

1898; Lowe, 1933; Marples, 1952) and they have evolved a range

of adaptations to an aquatic lifestyle including: small and scale-

310

T. Park & E.M.G. Fitzgerald

like feathers; increased underwater visual acuity (Sivak &

Millodot, 1977; Bowmaker & Martin, 1985); several retia

mirabilia systems for efficient thermoregulation (Frost et al.,

1975; Thomas & Fordyce, 2007, 2012); stiffening of wing joints

(Raikow et al., 1998); relative shortening of the wing;

hydrodynamic flattening of wing elements; and shortening of the

tarsometatarsus. The Sphenisciformes differ from most avian

groups in that total diversity was greater in the past than present,

with 19 extant species (sensu Ksepka & Ando, 2011), and 53

recognised fossil species (Fig. 2). This is testament to their aquatic

lifestyle and the fact that particular penguin bones (e.g. humerus,

tarsometatarsus) are more likely to fossilise than their equivalents

in other avian groups due to their pachyostotic histology (Meister,

1962). Extant species feed on small fish, cephalopods, crustaceans

and plankton, and show little interspecific postcranial

morphological variation (Olson, 1985). Interspecific differences

in cranial morphology are minimal; the differences that do occur

probably reflect disparate feeding ecology (Zusi, 1975).

Origin of Sphenisciformes

The fossil record of penguins is one of the longest and relatively

complete of any of the neornithine groups, potentially allowing

scientists to test hypotheses regarding the physical drivers of

vertebrate evolution e.g. climate change, palaeoceanography

(Baker et al., 2006; Ksepka & Thomas, 2012), biogeography

(Clarke et al., 2007), secondary adaptation to water (Thomas &

Fordyce, 2007), and stratigraphic calibration of molecular

divergence estimates (Slack et al., 2006). Thus the penguin fossil

record informs broader issues in macroevolution.

The oldest known penguin, Waimanu manneringi Jones,

Ando and Fordyce, 2006, is from the early Paleocene (60.5 -

61.6 Ma) of New Zealand (Slack et al., 2006). Although archaic,

it is clearly a penguin and already flightless. Simpson (1946)

summarised previous theories of penguin evolution (e.g. Lowe,

1933) and concluded that penguins evolved directly from a volant

ancestor, with no intermediate terrestrial stage. Molecular data

estimate the divergence of Sphenisciformes from their sister

taxon, Procellariiformes, about 71 Ma during the Cretaceous

(Baker et al., 2006; Brown et al., 2008). Slack et al., (2006) wrote

that the origins of Sphenisciformes took place 90-100 Ma as

part of the Late Cretaceous neornithine radiation. It has been

proposed that once the loss of aerial flight had occurred the

adaptation of penguins to an aquatic lifestyle occurred rapidly

due to the opening of ecological niches left by the extinction of

most marine reptiles at the end of the Cretaceous and the

intensive selection pressures of entering a new “adaptive zone”

(Fordyce & Jones, 1990).

Some fossil species reached giant sizes (e.g. Anthropornis

nordenskjoeldi Wiman, 1905, Pachydyptes ponderosus Oliver,

1930) of 1.5 - 1.6 m in standing height (Jadwiszczak, 2001), far

exceeding that of today’s largest species Aptenodytes forsteri

Gray, 1844 (emperor penguin), which rarely exceeds 1.0 m

(Friedmann, 1945; Stonehouse, 1975; Ksepka et al., 2012).

Nonetheless, estimated heights of giant taxa may be overestimates

following the first discovery of body proportions in a nearly

complete stem penguin (Ksepka et al., 2012). These giant species

(and potentially all stem species) are thought to have fed on fish,

using their slender bills to spear large prey (Olson, 1985; Myrcha

et al., 1990,2002; Ksepka et al., 2008), This contrasts with extant

species, which tend to have shorter beaks {Aptenodytes is an

exception) and feed on smaller fish (Zusi, 1975). This trophic

specialisation is thought to have occurred relatively late in

penguin evolution (Ksepka & Bertelli, 2006), with elongate,

narrow beaks representing the ancestral condition (Clarke et al.,

2007). Fossil feathers are known from Inkayacu paracasensis

Clarke et al., 2010, a species from the Eocene of Peru. These

show not only that the key features of penguin wing feathers had

evolved early in penguin evolution, but that this particular

species was reddish-brown and grey, considerably different from

the iconic black and white colouration of extant penguins (Clarke

et al., 2010).

Considerable effort over the last two decades has been aimed

at resolving penguin phylogeny including extinct taxa (Sibley &

Ahlquist, 1990; Baker et al., 2006; Gianni & Bertelli, 2004;

Bertelli & Gianni, 2005; Bertelli et al., 2006; Ksepka et al.,

2006; Slack et al., 2006; Walsh & Suarez, 2006; Clarke et al.,

2007; Acosta Hospitaleche et al., 2007, 2008). There is a general

consensus between morphological and molecular data (Fig. 2),

apart from the issues of where the phylogenies are rooted

(Livezey, 1989), and the timing of the divergence of the crown

Spheniscidae (Clarke et al., 2007). Basal penguins form a

paraphyletic group, with higher morphological disparity

compared to crown Spheniscidae (Davis and Renner, 2003). This

is most likely due to the relatively recent common ancestry and

broadly similar feeding ecology of the crown Spheniscidae

(Zusi, 1975). The timing of the crown clade’s divergence from

stem Sphenisciformes is still unresolved, as molecular and

morphological data give different estimates of ca. 41 Ma (Middle

Eocene) and 11-13 Ma (Middle-Late Miocene), respectively

(Baker et al., 2006; Gohlich, 2007). All pre-Miocene taxa are

stem Sphenisciformes (Ksepka & Clarke, 2010), rendering the

fossil record incongruent with the ancient divergence estimated

from molecular data.

Australian fossil Sphenisciformes

The record of Sphenisciformes in Australia is less extensive than

that of New Zealand, South America and Antarctica, with a

chronologically scattered distribution and the majority of fossils

being fragmentary. This limited record probably reflects a lack

of systematic field exploration, collecting and research, rather

than real rarity of fossils. Until now, the majority of fossil

penguin discoveries have been fortuitous in nature. Despite this

relatively meagre record, penguins are known from ten localities

limited to southeast Australia in every geologic epoch from the

Eocene onwards (Figs. 1 and 3; Table 1).

Eocene. Six specimens in total have been described from this

epoch (Finlayson, 1938; Glaessner, 1955; Simpson, 1957; Jenkins,

1974), including the partial skeleton (SAM P14157) of an

indeterminate form resembling the Antarctic genus Anthropornis

(Jenkins, 1974; Jenkins, 1985) (Fig. 4; Table 2). Found at Blanche

Point, South Australia and originally named as Pachydyptes

simpsoni, it is the most complete fossil penguin yet discovered in

Australia. Other specimens referred to this Anthropornis-like

form include a partial right humerus (SAM P14158a) (Fig. 5;

A review of Australian fossil penguins (Aves: Sphenisciformes)

311

Table 1. Summary of Australian fossil penguin localities. Abbreviations: E, east; NNW, north northwest; S, south; SSW, south southwest;

Locality

Geographic Location

Coordinates

Age (Ma)

Stratigraphic

position

Reference(s)

Blanche Point

37 km SSW of Adelaide,

South Australia.

35°14'S,

138°27'E

Late Eocene

(36.5-38.0)

Blanche Point

Formation

Alley et al., 1995; Jenkins

et al., 1982; James and

Bone, 2000

Witton Bluff

South end of Christie’s

Beach, about 26 km SSW of

Adelaide, South Australia.

35°09'S,

138°28'E

Late Eocene

(36.5-38.0)

Mount Gambier

Pritchard Brothers’ building

stone quarry, about 11 km

west of Mount Gambier,

South Australia.

37°49'S,

140°38'E

Early-Late

Oligocene

(23.0-30.0)

Gambier

Limestone

Alley et al., 1995; Li et

al., 2000; Fitzgerald, 2004

Devil’s Den

On east bank of Glenelg

River, about 17 km NNW of

Dartmoor, Victoria. Site is

marked “Bw” on the map

published by Singleton

(1941, p. 46).

37°46'S,

141°14'E

Early Miocene

(17.6-21.0)

Gellibrand

Marl

Gill, 1959a; Jenkins,

1974; Abele etal., 1988;

Dickinson et al., 2002

Batesford Quarry

Australian Cement Company

quarry south of Batesford, on

the western bank of the

Moorabool River, west of

Geelong.

38°06'S,

144°17'E

Early-Middle

Miocene

(15.9-17.6)

Batesford

Limestone

Abele et al., 1988;

Gourley and Gallagher,

2004

Middle Miocene

(13.7-15.9)

Fyansford

Formation

Abele et al., 1988;

Gourley and Gallagher,

2004

Portland

Beach on western side of

Portland Bay, at Portland,

Victoria.

38°20'S,

141°36'E

Late Miocene

(6.0-9.8)

Port Campbell

Limestone

Mallett, 1977; Abele et

al., 1988; Dickinson et al.,

2002

Spring Creek

Northeast of Minhamite, 41

km southeast of Hamilton,

Victoria.

37°58'S,

142°23'E

Late Miocene-

Early Pliocene

(5.0-6.0)

Goodwood

Formation

Gill, 1964; Simpson,

1970; Abele etal., 1988;

Holdgate and Gallagher,

2003

Beaumaris

East of Rickett’s Point on

west shore of Beaumaris

Bay, on northeast shore of

Port Phillip Bay.

37°59'S,

145°03'E

Late Miocene-

Early Pliocene

(5.0-6.2)

Black Rock

Sandstone

Abele et al., 1988;

Dickinson and Wallace,

2009

Red Bluff

About 3.5 km southwest of

Lake Tyers, east of Lakes

Entrance, East Gippsland,

Victoria.

37°52'S,

148°03'E

Late Pliocene

(2.5-3.5)

Jemmy’s Point

Formation

Abele etal., 1988;

Wallace et al., 2005

Amphitheatre

Cave

Approximately 6 km north of

Nelson, south-western

Victoria.

38°03'S,

141°0rE

Holocene

Cave pitfall

assemblage

Baird, 1992

Hunter Island

Stockyard Site, Hunter

Island, 5 km north of

Tasmania.

40°32'S,

144°45'E

Holocene

Aboriginal

midden

Van Tets and O’Connor,

1983

312

T. Park & E.M.G. Fitzgerald

Figure 1. Chronostratigraphy and correlation of Australian fossil penguin-bearing units, compared with the fossil record of Sphenisciformes on

other continents. Note that the vertical ranges of stratigraphic units represent estimates of geologic age maxima and minima for penguin-bearing

horizons, not necessarily discrete time-spans of deposition. Geologic timescale is after Gradstein et al. (2004) with updates from Walker and

Geissman (2009). See Table 1 for references to ages of Australian units. Ages of African units: Hendey (1981); Roberts et al. (2011); Ksepka and

Thomas (2012). Ages of Antarctic units: Dingle and Lavelle (2000); Jadwiszczak (2006); Marenssi et al. (2012). Ages of New Zealand units:

White and Waterhouse (1993); Cooper (2004); Slack et al. (2006); Ksepka et al. (2012). Ages of South American units: Devries (1998); Scasso et

al. (2001); Celma and Cantalamessa (2007); Achurra et al. (2009); Malumian and Nanez (2011); Uhen et al. (2011).

Table 2), a partial right radius (SAM P14158b) (Fig. 5; Table 2),

and a partial rib (SAM P17913; Table 2). A second form originally

thought to be closely related to Palaeeudyptes was described by

Finlayson (1938), who reported a left humerus (SAM P7158)

(Fig. 6; Table 2) from Witton Bluff, South Australia. A right

tibiotarsus (SAM P10862) (Fig. 7; Table 2), also from Witton

Bluff was noted by Glaessner (1955), described by Simpson

(1957), and also referred to this form. Simpson (1971)

subsequently reassigned this material to Sphenisciformes,

indeterminate. A third form, intermediate in size between the

Palaeeudyptes and Anthropornis forms is known from two

bones, one allegedly found at Blanche Point and the other from

late Eocene rocks near Browns Creek, Otway Peninsula, Victoria

(Jenkins, 1985; Vickers-Rich, 1991). These two specimens have

not been described, and recent efforts by one of us (EMGF) to

locate them in the SAM and MV collections have failed.

A review of Australian fossil penguins (Aves: Sphenisciformes)

313

OLIGOCENE

TPUlPl

0 AGE (Ma)

MIOCENE

EPOCH

Delphinornfs ,

Marambiomis exilis

Mesetaornis poiaris

Perudyptes devriesi

Anthropomis i

Pataeeudyptes

inkayacu paracasensis

Icadyptes satasi

Pachydyptes ponderosus

Katruku

m Archaeospheniscus

parvus

'j (- Paraptenodytes antarcticus

M r-^Platydyptes

i Palaeospheniscps

1 Eretiscus tonnii

> Dege hendeyi

iMarplesornis novaezealandiae

r—^Aptenodytes forsteri

' — Aptenodytes patagonicus

,i— 'fy<

"i—t Py

' "— iPy<

Pygoscalis adeliae

ygoscelis papua

‘ anrsj

|-sp"

Pygoscelis antarctic a

Pygoscetis grandis

lEudyptuta minor

Spheniscus megarhampus

Spheniscus urbinat

HEIongate hegks

Deeper beaks h

Spheniscus mui2oni_ Sphenlscl]s mage/te „ teus

" heniscus demersus

eniscus mendiculus

eniscus humboldti

Meaadyptes antipodes

i Madrynpmis mirandus

J Eudyptes filholi

iEudyptes chrysocome

1 Eudyptes moseieyi

1 Eudyptes schiegti

- Eudyptes chrysolvphus

■Eudyptes set atari

1 Eudyptes pachyrhynchus

^Eudyptes robustus

' j— i i

Giafit body

siza^-

Morphological

trends

First occurrence

Figure 2. Temporally-calibrated phylogeny of Sphenisciformes modified from Ksepka and Ando (2011) and Ksepka et al. (2012), showing major

trends in Sphenisciform evolution and the first occurrence of Sphenisciformes in each region with extant species. Stem sphenisciform branches

are shown in black and crown clade Spheniscidae branches are shown in blue. Thick horizontal bars on branches indicate stratigraphic range

maxima and minima. Abbreviations: AF, Africa; AN, Antarctica; AU, Australia; Ma, million years ago; SA, South America.

Oligocene. Only two fossil penguin specimens are known from

the Oligocene of Australia. Both of them were derived from the

Camelback Member of the Gambier Limestone, which has been

correlated to the P21/22 planktonic foram zone (Lower-Upper

Oligocene) (Li et al., 2000). Both specimens, a partial right

humerus (SAM P10863) (Fig. 8; Table 2) and a partial left femur

(SAM P10870) (Fig. 9; Table 2), were first noted by Glaessner

(1955) and later described by Simpson (1957). Neither of the

specimens has been assigned to a genus or species, but they are

considered to be separate taxa (Simpson, 1957).

Early-Middle Miocene. Fossil penguin specimens have been

described from the Early-Late Miocene of Australia. A single

specimen was found as float on the banks of the Glenelg River at

Devil’s Den, Victoria (Gill, 1959a; Simpson, 1959). The large

right humerus (NMV P17167) was named as Anthropodyptes

gilli by Simpson (1959) (Fig. 10; Table 2). From the Early

Miocene (Gill, 1959a; Jenkins, 1974; Dickinson et al., 2002), it is

the latest surviving giant stem Spheniscid known, although some

extinct crown Spheniscid taxa (e.g. Spheniscus megaramphus )

would have been larger than Aptenodytes forsteri (Ksepka &

Clarke, 2010; Stucchi, 2003). As yet undescribed material has

also been collected from the Lower-Middle Miocene Batesford

Limestone at Batesford, near Geelong, Victoria including: a

partial right femur (NMV P222904), a partial left femur (NMV

P201867), and a partial left coracoid (NMV P231933). A partial

left tibiotarsus (NMV P231836) and a partial right femur (NMV

P201856) have also been collected from the Middle Miocene

Fyansford Clay at the same locality.

Late Miocene-Pliocene. Mio-Pliocene penguins have been

described from two localities: Spring Creek, near Minhamite,

and Beaumaris, Victoria. Simpson (1965,1970), described a total

of 21 penguin specimens from these localities. From Spring

Creek an incomplete left humerus (NMV P26668, holotype of

Pseudaptenodytes macraei Simpson, 1970) (Fig. 11; Table 2),

314

T. Park & E.M.G. Fitzgerald

Figure 3. Fossil sphenisciform localities in Australia. 1 = Devil’s Den; 2 = Portland; 3 = Spring Creek; 4 = Batesford Quarry; 5 = Beaumaris; 6

= Red Bluff; 7 = Hunter Island; 8 = Witton Bluff; 9 = Blanche Point; 10 = Mount Gambier

and from Beaumaris: a partial left coracoid (NMV P24065)

(Simpson, 1965) (Fig. 12; Table 2); three partial left humeri

(NMV P26671, NMV P26676, NMV P27059) (Fig. 13; Table 2);

four partial right humeri (NMV P26669, NMV P26677, NMV

P27057, NMV P26670) including the holotype of

IPseudaptenodytes minor Simpson, 1970 (NMV P26669) (Figs.

13 and 14; Table 2); four partial carpometacarpi (NMV P27055,

NMV P27056, NMV P27058, NMV P26903) (Figs. 13 and 15;

Table 2); and eight isolated fragments (registration numbers

unspecified by Simpson, 1970). The P. macraei humerus is the

larger of the two species, being similar in size to the humerus of

a king penguin ( Aptenodytes patagonicus Miller, 1778), whereas

the IP. minor humerus approaches that of the gentoo penguin

(Pygoscelis papua Forster, 1781) in size. IPseudaptenodytes

minor was referred to Sphenisciformes indet. by Ksepka and

Ando (2011: 159). The Upper Miocene Port Campbell Limestone

at Portland, Victoria, has yielded a nearly complete left humerus

(NMV P221273) and a partial left humerus (NMV P232062).

One definitively Pliocene specimen is known from Australia, an

undescribed partial right femur (NMV P41738) from the

Pliocene Jemmys Point Formation at Red Bluff, west of Lake

Tyers, Victoria.

Holocene. Van Tets and O’Connor (1983) described penguin

remains from a ca. 760 year-old aboriginal midden on Hunter

Island, Tasmania and described these as a new genus and species,

Tasidyptes hunteri. Other workers (Fordyce and Jones, 1990;

Ksepka and Clarke, 2010) have doubted this identification due to

the fragmentary nature of the fossils: the coracoid (ANWC

BS2669) (Table 2) and tarsometatarsus (ANWC BS2668) (Table

2) are indistinguishable from Eudyptes\ and the four specimens

comprising the hypodigm (ANWC BS2667, ANWC BS2668,

ANWC BS2669, ANWC BS2670) (Table 2) come from three

different horizons within the midden (Van Tets and O’Connor,

1983; Fordyce and Jones 1990; Ksepka and Clarke, 2010). Baird

(1992) reported little penguin ( Eudyptula minor Forster 1781)

material from a pitfall assemblage in Amphitheatre Cave,

Victoria which has been dated to ca. 4670 ybp (Table 2). However,

this Eudyptula material is unlikely to be from the original pitfall

assemblage due its differential preservation (Baird, 1992: 31-32).

A review of Australian fossil penguins (Aves: Sphenisciformes)

315

Table 2. Described fossil penguin specimens from Australia. Abbreviations: ANWC, Australian National Wildlife Collection; indet.,

indeterminate; Ma, million years ago; NMV P, Museum Victoria Palaeontology Collection; SA, South Australia; SAM P, South Australian

Museum Palaeontology Collection; TAS, Tasmania; VIC, Victoria.

Specimen

Taxon (previous

assignment)

Material

Locality (Formation)

Age (Ma)

Reference(s)

SAM P7158

Sphenisciformes indet. (cf.

Palaeeudyptes)

left humerus

Witton Bluff, S A (Blanche Point

Formation, Tuketja Member)

Late Eocene

(36.5-38.0)

Finlayson, 1938;

Simpson, 1957

SAM P10862

right tibiotarsus

Witton Bluff, SA (Blanche Point

Formation, Gull Rock Member)

Glaessner, 1955;

Simpson, 1957

SAM P14157

(a-g)

Sphenisciformes indet.

(Pachydyptes simpsoni )

partial skeleton

Blanche Point, SA (Blanche Point

Formation, Gull Rock Member)

Jenkins, 1974

SAM P14158a

partial right humerus

Blanche Point, SA (Blanche Point

Formation, Tuketja Member)

SAM P14158b

“

partial right radius

“

SAM P17913

“

partial rib

“

SAM P10863

Sphenisciformes indet.

partial right humerus

Mt. Gambier, SA (Gambier

Limestone)

Early-Late

Oligocene

(-23.0-30.0)

Glaessner, 1955;

Simpson, 1957

SAM P10870

partial left femur

Mt. Gambier, SA (Gambier

Limestone)

Early-Late

Oligocene

(-23.0-30.0)

Simpson, 1957

NMV P17167

Anthropodyptes gilli

right humerus

Glenelg River, Devil’s Den, VIC

(Gellibrand Marl)

Early Miocene

(17.6-21.0)

Simpson, 1959;

Gill, 1959a

NMV P24065

Sphenisciformes indet.

partial left coracoid

Beaumaris, VIC (Black Rock

Sandstone)

Late Miocene-Early

Pliocene (5.0-6.0)

Simpson, 1965

NMV P26668

Pseudaptenodytes macraei

partial left humerus

Spring Creek, Minhamite, VIC

(Goodwood Formation)

Late Miocene-Early

Pliocene (5.0-6.0)

Gill, 1964;

Simpson, 1970

NMV P27055

cf. Pseudaptenodytes

macraei

partial right

carpometacarpus

Beaumaris, VIC (Black Rock

Sandstone)

Late Miocene-Early

Pliocene (5.0-6.0)

Simpson, 1970

NMV P27056

cf. Pseudaptenodytes

macraei

partial left

carpometacarpus

NMV P26669

Sphenisciformes indet.

(?Pseudaptenodytes minor)

partial right humerus

NMV P26677

“

NMV P26670

“

NMV P27057

“

NMV P26671

“

partial left humerus

“

NMV P26676

“

NMV P27058

partial right

carpometacarpus

NMV P26903

right

carpometacarpus

NMV P27059

Sphenisciformes indet.

partial left humerus

“

N/A

“

isolated fragments (8)

“

ANWC BS2667

Sphenisciformes indet.

(Tasidyptes hunteri )

juvenile synsacmm

Hunter Island, TAS (Aboriginal

midden)

Holocene (ca. 760

ybp)

Van Tets and

O’Connor, 1983

ANWC BS2668

Eudyptes sp. ( Tasidyptes

hunteri)

left tarsometatarsus

ANWC BS2669

“

left coracoid

“

ANWC BS2670

Sphenisciformes indet.

('Tasidyptes hunteri)

pelvis

NMV P178677

Eudyptula minor

proximal pelvis

Amphitheatre Cave, VIC (pitfall

assemblage)

Holocene (ca. 760

ybp)

Baird, 1992

NMV P178678

Eudyptula minor

right femur

“

NMV P178679

Eudyptula minor

left femur

“

NMV P178680

Eudyptula minor

partial left tibiotarsus

“

NMV P178681

Eudyptula minor

partial left

tarsometatarsus

316

T. Park & E.M.G. Fitzgerald

Figure 4. Pachydyptes simpsoni holotype, SAM P14157: A, right radius in ventral view, left carpometacarpus and left phalanx II-l in dorsal view;

B, head of right humerus in dorsal view; C, left coracoid in dorsal view; D, ?twelfth cervical vertebra in ventral view.

Figure 5. Pachydyptes simpsoni paratype, SAM P14158: proximal end of right radius in (A) ventral view and (C) dorsal view; partial right

humerus in (B) ventral and (D) dorsal views.

A review of Australian fossil penguins (Aves: Sphenisciformes)

Figure 6. Sphenisciformes indet. left humerus, SAM P7158: A, dorsal

view; B, ventral view.

Systematic Palaeontology

A total of five species of fossil penguin have been named from

Australia. Only one taxon is based on a type specimen

consisting of associated remains. The remaining four species

are established on isolated elements. Only two of these five

species are currently considered taxonomically distinct

(Anthropodyptes gilli and Pseudaptenodytes macraei ).

Aves Linnaeus, 1758

Sphenisciformes Sharpe, 1891

Spheniscidae Bonaparte, 1831

Pachydyptes Oliver, 1930

Pachydyptes simpsoni Jenkins, 1974

Holotype. Partial skeleton (SAM P14157) consisting of: a

partial left coracoid; partial right humerus; partial left humerus;

a right radius; a partial left carpometacarpus; a left phalanx II-

1; and a partial vertebra. (Fig. 4; Table 2).

Type locality. Blanche Point, 37 km SSW of Adelaide, South

Australia (35°14'S, 138°27'E).

Horizon and age. Occurs in the Gull Rock Member and the

Tuketja Member of the Blanche Point Formation. The Gull

317

Figure 7. Sphenisciformes indet. partial right tibiotarsus, SAM

PI0862: A, cranial view; B, caudal view.

Rock Member consists of green and grey, glauconitic and

fossiliferous calcareous mudstone with a few limestone lenses;

the Tuketja Member consists of alternating bands of tough,

dark grey chert and friable clays, silts and calcareous clays

(Jenkins et al., 1982). Both members are in the P15 foraminiferal

zone; Late Eocene (Bartonian-Priabonian), 36.5-38.0 Ma

(James and Bone, 2000).

Referred material. A partial right humerus (SAM P14158a)

(Fig. 5), a partial right radius (SAM P14158b) (Fig. 5) and a rib

fragment (SAM P17913). (Table 2)

Diagnosis. Following Marples (1952), the generic diagnosis of

Pachydyptes is as follows : humerus relatively wide; m.

deltoideus minor insertion (referred to as the ‘external

tuberosity’ by Marples, 1952) projects distally; articular surface

flattened; fossa pneumotricipitalis undivided; m.

supracoracoideus insertion slightly oblique, almost parallel to

long axis of shaft and widely separated from the m. latissimus

dorsi insertion; shaft has slight sigmoid curve and slight

angulation of the cranial border; angle between long axis of

shaft and tangent of condylus dorsalis and condylus ventralis is

acute; shelf adjacent to condylus ventralis approximately the

same width as condylus ventralis; and coracoid convex at base.

Following Jenkins (1974), Pachydyptes simpsoni differs from

Pachydyptes ponderosus Oliver, 1930 by having: more concave

medial margin of the coracoid; more pronounced angulation of

the cranial margin of the humerus (referred to as the “preaxial

318

I ,1 1

F\ V* 1 I ; 1

I j 1 *1

I y V 1

/ 110 mm

Figure 8. Sphenisciformes indet. partial right humerus, SAM P10863:

A, dorsal view; B, ventral view.

tuberosity on the shaft at the proximal limit of attachment of

brachialis internus” by Jenkins, 1974); more widely separated

insertions of the musculi supracoracoideus and the musculi

coracobrachialis caudalis (referred to as the pectoralis secundus

and pectoralis tertius respectively by Jenkins, 1974); metacarpal

III extends further distally than metacarpal II; and the bones

are generally less robust.

Remarks. The referred humerus (SAM P14158a) is similar to

Pachydyptes ponderosus with its large head, expanded muscle

attachments and wide shaft. The skeleton however is overall

less robust than P. ponderosus and the overall morphology of

the coracoid, radius and carpometacarpus shows similarities to

Anthropornis and Palaeeudyptes (Jenkins, 1974). The coracoid

has a broadly flared base and an oval foramen nervi

supracoracoidei. On the radius, the insertion site of the m.

brachialis is hollowed, forming a distinct notch similar to that

of Paraptenodytes robustus Ameghino, 1905, although the

bone itself resembles that of Palaeeudyptes and Anthropornis

(Jenkins, 1974). Systematic revisions of Pachydyptes simpsoni

have seen it first synonymised with Anthropornis nordenskjoeldi

(Jenkins, 1985), and most recently considered as

Sphenisciformes indet. (Ksepka and Clarke, 2010). The latter

authors concluded that it occupied a more crownward position

than Antarctic A. nordenskjoeldi specimens. We therefore

consider the systematics of P. simpsoni to be unresolved.

T. Park & E.M.G. Fitzgerald

Figure 9. Sphenisciformes indet. partial left femur, SAM P10870: A,

dorsal view; B, ventral view.

Anthropodyptes Simpson, 1959

Anthropodyptes gilli Simpson, 1959

Holotype. Right humerus (NMV P17167). (Fig. 10; Table 2).

Type locality. Specimen was found as float on top of Miocene

marl on east bank of Glenelg River at Devil’s Den, about 17 km

NNW of Dartmoor, Victoria. Site is marked “Bw” on the map

published by Singleton (1941: 46) (37°46'S, 141°14'E).

Horizon and age. Gill (1959a) determined that NMV P17167

was derived from the Gellibrand Marl, which at this locality

represents planktonic foram zones N5-N6, Early Miocene

(Aquitanian-Burdigalian), 17.6-21.0 Ma (Jenkins, 1974: 292;

Abele et al., 1988:285; Dickinson et al., 2002).

Diagnosis. Simpson (1957: 118) notes that Anthropodyptes does

not share any diagnostic characters with any previously named

genus. Generic characteristics as follows: humerus slender and

elongate; shaft slightly sigmoid, with moderate angulation of the

cranial margin; the proximal part of the shaft is narrower than

the distal part; fossa pneumotricipitalis undivided and large

proximo-distally; m. supracoracoideus insertion wide and

slightly oblique, almost parallel to long axis of the shaft; angle

between long axis of shaft and tangent of condylus dorsalis and

condylus ventralis is about 42°; the condylus ventralis is only

slightly ventral to the condylus dorsalis; shelf adjacent to

condylus ventralis smaller than condylus ventralis.

A review of Australian fossil penguins (Aves: Sphenisciformes)

A B

Figure 10. Anthropodyptes gilli holotype right humerus, NMV P17167:

A, dorsal view; B, ventral view.

Remarks. This species is apparently most similar to

Archaeospheniscus (Gill, 1959b; Simpson, 1959: 118), a Late

Oligocene New Zealand form. Based on synapomorphies,

Anthropodyptes gilli has a most exclusive placement of clade 8

in the phylogenetic analysis of Ksepka and Clarke (2010: Fig.

21), giving it a more crown-ward position than earlier ‘giant’

forms from the late Eocene, but a similar phylogenetic position

to late Oligocene giant forms such as Kairuku and

Archaeospheniscus. Anthropodyptes gilli bears the distinction

of being the latest surviving giant stem penguin, all other

‘giant’ stem taxa having a Palaeogene age (Ksepka and Clarke,

2010: 45). Comparisons of body proportions with other giant

taxa are not possible until more complete material is found.

Pseudaptenodytes Simpson, 1970

Pseudaptenodytes macraei Simpson, 1970

Holotype. Partial left humerus (NMV P26668). (Fig. 11; Table

2 ).

Type locality. Spring Creek near Minhamite, 41 km southeast

of Hamilton, Victoria (37°58'S, 142°23'E).

Horizon and age. The holotype is derived from the Goodwood

Formation, a green-grey marly fine sand with abundant

pebbles (Gill, 1964:332). The macroinvertebrate assemblage is

similar to that of the Upper Miocene-Lower Pliocene Black

319

Figure 11. Pseudaptenodytes macraei holotype left humerus, NMV

P26668: A, dorsal view; B, dorso-caudal view; C, ventral view.

Rock Sandstone (Gill, 1964; Simpson, 1970), and the

Goodwood Formation is possibly laterally equivalent to the

Upper Miocene-Lower Pliocene Grange Burn Formation

(Fitzgerald, 2004).

Referred material. A partial right carpometacarpus (NMV

P27055) and a partial left carpometacarpus (NMV P27056)

were tentatively referred to this species (Simpson, 1970) (Fig.

15; Table 2). Both specimens were derived from the Upper

Miocene-Lower Pliocene Black Rock Sandstone at Beaumaris.

Diagnosis. Simpson (1970) noted the very close similarity of

Pseudaptenodytes macraei to Aptenodytes patagonicus, both

in terms of size and the features of the proximal end of the

humerus. Nevertheless, P. macraei differs from Aptenodytes

by having a humerus with: a more sigmoid shaft; a smaller

volume of the fossa pneumotricipitalis; a distinctly oval

opening of the internal division of the fossa pneumotricipitalis;

and a rounded cranial margin lacking a distinct ‘preaxial

angle’ (Acosta Hospitaleche et ah, 2008: Fig. 5, char. 11). It

further differs from A. forsteri by lacking the pit for ligament

insertion on the proximal surface adjacent to the head (Ksepka

et al., 2006: Fig. 8). In A. patagonicus this feature is variable

(Ksepka et al., 2006).

Remarks. Despite the similarities of the type specimen to

Aptenodytes patagonicus, Pseudaptenodytes macraei is not

ancestral to it or any of the modern species (Simpson, 1970:

320

T. Park & E.M.G. Fitzgerald

Figure 12. Sphenisciformes indet. partial left coracoid, NMV P24065:

A, dorsal view; B, ventral view.

20). Although similar, the autapomorphies of NMV P26668

preclude this specimen from referral to Aptenodytes or any

extant genera. We consider P. macraei to be a distinct taxon

established on the basis of a diagnostic type specimen.

?Pseudaptenodytes minor Simpson, 1970

Holotype. Partial right humerus (NMV P26669). (Fig. 14;

Table 2).

Type locality. East of Rickett’s Point on the western shore of

Beaumaris Bay, northeast shore of Port Phillip Bay, Victoria

(37°59'S, 145° 03'E).

Horizon and age. Black Rock Sandstone, which consists of a

basal layer of phosphatic and ferruginous intraclasts overlain

by calcareous sandstone (Dickinson and Wallace, 2009).

Planktonic foram and molluscan biostratigraphy indicate a

Late Miocene-Early Pliocene age range, which is corroborated

by Sr dates of 5.0-6.2 Ma (Dickinson and Wallace, 2009).

Referred material. Distal end of right humerus (NMV P26677),

proximal end of left humerus, (NMV P26671), proximal end of

left humerus (NMV P26676), right humerus (NMV P26670),

right humerus (NMV P27057), partial right carpometacarpus

(NMV P27058), right carpometacarpus (NMV P26903) (Fig.

13; Table 2). All referred material was collected from the Upper

Miocene-Lower Pliocene Black Rock Sandstone at Beaumaris.

Diagnosis. Differs from Pseudaptenodytes macraei by having:

a more slender and less sigmoid shaft; and a less expanded

distal section (Simpson, 1970). It also differs by having a

distinct angle on the cranial margin, although this angulation is

still less than that of any modern penguin (Simpson, 1970).

Remarks. Simpson (1970) notes the similarities between

?Pseudaptenodytes minor and Paraptenodytes robustus yet the

holotype of IP. minor is too incomplete to permit meaningful

comparisons. The lack of diagnostic morphology in the type

specimen of IP. minor has resulted in Ksepka and Clarke (2010)

referring this taxon to Sphenisciformes indet. More completely

preserved material is required to confirm or reject the placement

of this species in Pseudaptenodytes. The additional material

referred by Simpson (1970) to IP. minor displays little overlap in

morphology with the holotype (NMV P26669). Furthermore, it

is only on the basis of the referred material that the holotype

was designated a species of Pseudaptenodytes. We therefore

recommend restricting the concept of IP. minor to the holotype.

All referred material should be considered Sphenisciformes

indet. pending further study.

Tasidyptes Van Tets and O’Connor, 1983

Tasidyptes hunteri Van Tets and O’Connor, 1983

Holotype. Pelvis in three parts (ANWC BS2670) (Table 2).

Type locality. Stockyard Site, Hunter Island, 5 km north of

Tasmania (40°32'S, 144°45'E).

Horizon and age. Material found in an aboriginal midden.

Carbon dating resulted in an age of 760 ± 70 ybp (Holocene).

Referred material. The paratype specimen is a left

tarsometatarsus (ANWC BS2668). Also referred to the species

are a juvenile synsacrum (ANWC BS2667) and a left coracoid

(ANWC BS2669). (Table 2)

Diagnosis. Differs from Eudyptula and Megadyptes by having:

a caudal part of the synsacrum with relatively broader fused

vertebrae and long slender lateral processes; and the lateral

foramen vasculare proximale situated more distal than the

medial foramen vasculare proximale on the plantar surface of

the tarsometatarsus. However, this character is not clear from

the figure in Van Tets and O’Connor (1983: Fig. 4).

Remarks. This taxon is no longer considered valid due to the

fragmentary nature of the fossils, the fact that the coracoid and

tarsometatarsus are indistinguishable from Eudyptes and the

fact that the four specimens come from three different

stratigraphic layers of the midden (Van Tets and O’Connor,

1983; Fordyce and Jones, 1990; Ksepka and Clarke, 2010).

However, Ksepka and Clarke (2010) note that due to the young

age of the specimens, DNA testing to confirm their identity

may well be possible. Ksepka and Ando (2011: 178) also draw

attention to the synsacrum stating that the long slender lateral

processes may be “a possible diagnostic character, certainly in

need of quantitative evaluation”.

A review of Australian fossil penguins (Aves: Sphenisciformes)

321

Figure 13. Sphenisciformes from the Upper Miocene-Lower Pliocene Black Rock Sandstone, Victoria. A-J and M-P, specimens referred to

IPseudaptenodytes minor : partial left humerus, NMV P 26671, in (A) dorsal and (B) ventral views; partial left humerus, NMV P26676, in (C)

dorsal and (D) ventral views; partial right humerus, NMV P26677, in (E) dorsal and (F) ventral views; partial right humerus, NMV P26670, in

(G) dorsal and (H) ventral views; partial right humerus, NMV P27057, in (I) dorsal and (J) ventral views; partial right carpometacarpus, NMV

P27058, in (M) dorsal and (N) ventral views; partial right carpometacarpus, NMV P26903, in (O) dorsal and (P) ventral views. Sphenisciformes

indet. partial left humerus, NMV P27059: K, dorsal view; L, ventral view.

322

T. Park & E.M.G. Fitzgerald

Figure 15. Specimens referred to Pseudaptenodytes macraei : partial right carpometacarpus, NMV P27055, in (A) dorsal and (B) ventral views;

partial left carpometacarpus, NMV P27056, in (C) ventral and (D) dorsal views.

A review of Australian fossil penguins (Aves: Sphenisciformes)

Discussion

The Australian penguin record has thus far played a minor role

in the interpretation of sphenisciform evolutionary history.

Despite there being general summaries (Jenkins, 1985;

Fordyce and Jones, 1990; Vickers-Rich, 1991), no primary

systematic research has been conducted since 1983.

With this limited quantity of described material, what

patterns may be deduced from the fossil record of penguins in

Australia? As shown above, the record has a scattered

chronologic distribution and is based on fragmentary fossils.

Nonetheless, the vast majority of all fossil penguin specimens

ever found are isolated and often incomplete elements (see

Acosta Hospitaleche et al., 2007; Ksepka et al., 2008, 2012 for

exceptions). Second, there is a disparity between past and

present taxonomic diversity. During the Eocene, Oligocene

and Miocene there were more than one species of penguin

inhabiting Australia. All these species were larger than the

sole extant species, Eudyptula minor Forster, 1781 (little

penguin), although body sizes have yet to be estimated using

established regression equations (e.g. Simpson, 1946;

Jadwiszczak, 2001). This higher taxonomic diversity and

morphological disparity relative to the present remains

unexplained. A possible causal factor promoting higher

diversity is the former increased availability of suitable

breeding grounds due to higher sea levels forming numerous

small offshore islands. An example of this is known from the

Pliocene of Africa (Ksepka and Thomas, 2012).

The Australian record includes penguin material from the

Early through late Miocene. This interval is inadequately

sampled worldwide (Ksepka and Ando, 2011: 157-163), and

thus Australian fossils may provide insights into this pivotal

period in penguin history, including the diversification of

crown Spheniscidae.

Acknowledgements

We thank: B. Crichton, E. Eidelson, T. Flannery, R. Hamson, J.

Long, J. Rowe and S. Wright for donating specimens to Museum

Victoria; D. Pickering, W. Longmore and K. Roberts for

providing access to collections at Museum Victoria; M.-A.

Binnie, N. Pledge and T. Worthy for providing access to

collections and research facilities at the South Australian

Museum; the Harold Mitchell Foundation and Museum Victoria

for supporting this research; and the South Australian Museum

for facilitating a visit by EMGF to study their collections.

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Memoirs of Museum Victoria 69:327-340 (2012)

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

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

Four new valviferan isopods from diverse tropical Australian habitats (Crustacea:

Isopoda: Holognathidae and Idoteidae)

Gary C. B. Poore

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

Abstract Poore, G.C.B. 2012. Four new valviferan isopods from diverse tropical Australian habitats (Crustacea: Isopoda:

Holognathidae and Idoteidae). Memoirs of Museum Victoria 69: 327-340.

Two new isopods of the family Holognathidae are described from tropical Australia: Cleantioides Carpentaria sp.

nov. from shallow seagrass sediments in the Gulf of Carpentaria and an eastern Queensland port; and Zenobianopsis

cidaris sp. nov. from 500 m depth in the Coral Sea. The latter is the third in its genus. Two new species of Synidotea

(Idoteidae), S. innatans sp. nov. and S. karumba sp. nov. are described. Both are distinguished from others of the S.

hirtipes- group of species, on the basis of subtle differences in colour and proportions of the body and limbs. The material

of S. innatans was taken from flotsam in the Timor Sea and is potentially an obligate rafting species. Synidotea karumba

is from shallow sedimentary environments in the Gulf of Carpentaria.

Keywords Crustacea, Isopoda, Valvifera, Holognathidae, Idoteidae, Cleantioides, Zenobianopsis, Synidotea, new species, Australia

Introduction

Two valviferan families of isopods are characterised by having

more or less flattened body shape and slight differentiation of

body segments, Holognathidae Thomson, 1904 and Idoteidae

Samouelle, 1819. The Holognathidae comprise 21 species in

five genera, four reviewed by Poore and Lew Ton (1990) and a

fifth added by Liu et al. (2010). Three genera inhabit temperate

shallow sediments in both hemispheres and two occur only in

the deep South Pacific and Southern oceans. Three species

occur in shallow temperate southern Australia (Poore and

Lew Ton, 1993). The Idoteidae on the other hand are found

circum-globally in shallow algal and seagrass habitats, mainly

at temperate latitudes. The family comprises 23 species in

Australia confined, with one exception, to temperate coasts

(Poore and Lew Ton, 1993). No new species have been reported

since the date of their review.

The only tropical species in Australia is the doubtful

Idotea brevicorna Milne Edwards, 1840. Poore and Lew Ton

(1993) debated the possible synonymy of this species, and its

probable synonym I. duplicata Nierstrasz, 1941 from

Indonesia, with the European species I. balthica (Pallas, 1772).

Idotea balthica is a facultative drifter on a variety of floating

material in the North Atlantic (Thiel and Gutow, 2005; Thiel

and Haye, 2006) and the probability of this species occurring

naturally in tropical Australian waters seems low. Neither I.

brevicorna nor I. duplicata has been reported since from the

region and the possibility remains that the records, based on

collections in the 19th century, are of translocated specimens

from ships' hulls.

In this contribution, new species are described from

tropical northern Australia, one holognathid from a shallow

environment and another from bathyal depths, plus two new

species of Idoteidae, one from shallow sedimentary habitat

and another from flotsam.

Abbreviations in figures are: al(f), antenna 1 (flagellum);

a2(p), antenna 2 (peduncle); md, right mandible; mp,

maxilliped; mx2, maxilla 2; p, penial plate; pl-p7, pereopods

1-7; pl2, pleopod 2; u, uropod. Material is lodged in Museum

Victoria, Melbourne (NMV), Queensland Museum, Brisbane

(QM) and the Museum of Tropical Queensland, Townsville

(MTQ).

Holognathidae Thomson, 1904

Cleantioides Kensley and Kaufman, 1978

Cleantioides Kensley and Kaufman, 1978: 658. - Poore and Lew

Ton, 1990: 59.

Remarks. The genus contains 12 species, four in Central

America (Kensley and Kaufman, 1978; Brusca and Wallerstein,

1979; Kensley, 1987), five in the north-western Pacific

(Richardson, 1912; Kussakin, 1982; Kwon and Kim, 1992), one

in South Africa (Barnard, 1925), and two in southern Australia

(Poore and Lew Ton, 1990). Poore and Lew Ton (1990)

diagnosed the genus and discussed its synonymy. Cleantis

annadalei Tattersall, 1921, previously included in this genus,

has been removed to its own genus, Chongxidotea Liu, Poore

and Lu, 2010.

328

G.C.B. Poore

Cleantioides Carpentaria sp. nov.

Figures 1-2

Material examined. Holotype. Queensland, Karumba, Norman River

mouth (17°28'S, 140°49'E), sandy sediment with seagrasses Halodule

uninervis and H.pinifolia, 1 m, K. Neil, 2001, NMV J62815 (ovigerous

female, length 13.5 mm).

Other material. Queensland, Lucinda, Sugar Loading Jetty Outer

(18°31.3'S, 146°19.9'E), sediments, 5 m, K. Neil, CRC Reef Research,

01 Jun 1999, QM 462915 (juvenile, 7.6 mm).

Description. Ovigerous female. Body approximately 6 times as

long as wide. Dorsal surface largely smooth, without setae.

Head 1.4 times as wide as long, excavate and depressed

anteriorly at midpoint, with small medial pseudorostrum;

posterior margin convex; transverse maxillipedal segmental

groove obvious. Pleotelson 0.32 whole length, twice as long as

wide; pleonite 1 free and articulating, barely visible under

pereonite 7; pleonites 2 and 3 well defined but not articulating,

pleonite 3 fused medially, lateral margin hidden under pleonite

2; remaining pleotelson parallel-sided and with semicircular

apex; dorsum of distal half with oblique (c. 30° from horizontal)

circular plane occupying 0.4 of pleotelson length, well-defined

by a sharp ridge extending three-quarters around, with obscurely

rugose surface.

Antenna 1 reaching to end of article 2 of antenna 2 peduncle;

peduncle article 1 as long as wide; flagellum about third length

of last article of peduncle, with 4 apical aesthetascs. Antenna 2

0.3 length of body, article 2 with narrow ventromesial projection

with bilobed apex; flagellum of 1 article, 0.25 total antenna

length, with fine setae all over and a dense clump at apex.

Maxillipedal endite with 2 plumose setae mesially and 8 on

transverse apex, palp width 0.45 length, 5 articles visible but

suture between 2 and 3 not articulating; articles 2-5 mesially

setose; article 3 mesiodistally lobed; article 5 wider than long,

0.2 length of article 4; epipod tapering and obliquely truncate.

Pereopod 1 with dense robust setation on margins of

merus, carpus and propodus; propodus 1.8 times as long as

wide, with 13 robust setae on mesial face. Pereopods 2 and 3

similar, anterodistal setae on merus and carpus, and margins

of propodus; propodus 2.4 times as long as wide. Pereopod 4

0.4 length of pereopod 3, ischium without setae; merus, carpus

and propodus with posterodistal U-shaped rows of 11, 18 and

12 robust setae respectively; dactylus reduced to a compact

unguis only. Pereopod 5 longer than 4, posteriorly with few

spines; dactylus slightly hooked. Pereopods 6 and 7 more

elongate than 5, propodus of pereopod 7 4.6 times as long as

wide. Oostegites on pereopods 1-5.

Uropodal endopod as wide as long, distally truncate at

right-angles to mesial margin over about 0.4 width.

Colour. Holotype unpigmented. Juvenile brown, darkest across

front of head, antenna 1 peduncle, articles 1, 2 and 5 of antenna

2, laterally on body segments and coxa.

Etymology. From the type locality, Gulf of Carpentaria (noun

in apposition).

Distribution. Australia, Queensland, east and west coasts of

Cape York at c. 18°S; to 5 m depth.

Remarks. Cleantioides Carpentaria is similar to the southern

Australian species C. albaniensis Poore and Lew Ton, 1990.

It differs in being narrower (6 times vs 5 times as long as

wide) and having slightly narrower limbs. It differs from all

other species in the obscurely rugose surface of the

pleotelsonic plane. The other Australian species, C. striata

Poore and Lew Ton, 1990, from NSW, is longitudinally

striped, as are several species in this genus, and has a steeper

pleotelsonic plane. The Asian species differ as follows: C.

emarginata Kwon and Kim, 1992 has an emarginate telsonic

apex, C. poorei Kwon and Kim, 1992 is more compact, C.

japonica Richardson, 1912 has a median tubercle on the

pleotelsonic plane, and C. rotundata Kussakin, 1982 has a

more acute pleotelson. Cleantioides natalensis (Barnard,

1925) is more elongate (Kensley, 1978).

Zenobianopsis Hale, 1946

Zenobianopsis Hale, 1946: 164-165.-Poore and Lew Ton, 1990: 74.

Remarks. Poore and Lew Ton (1990) rediagnosed the genus and

redescribed the type species, Z. caeca Hale, 1946. This species

and the second, Z. rotundicauda Kussakin, 1967 are both from

the Southern Ocean. Here, a third is described from deep water

at a more tropical latitude.

The head of this species and of Z. caeca (confirmed on

NMV material) possesses an obvious horizontal groove at the

base of antenna 2 reaching back about one third of the head

length. The groove is not seen in species of any of the other

genera, Cleantis, Cleantioides or Holognathus.

Zenobianopsis cidaris sp. nov.

Figures 3, 4

Material examined. Holotype. Coral Sea, 17°34.58'S, 146°53.21'E,

458-500 m, M. Pichon et al., 15 May 1986, sledge (CIDARIS I stn

43.2), MTQ W34049 (ovigerous female, 14.2 mm).

Description. Ovigerous female. Body 5.6 times as long as

greatest width at pereonite 3, pereonites 5-7 noticeably

narrower than 1-4, dorsal surface smooth, with fur of fine setae

on pleotelson. Head 1.1 times as wide as long, front with broad

obtuse pseudorostrum as long as lateral margins; lateral margin

with horizontal groove at base of antenna 2. Pleotelson 0.25

total length, 1.8 times as long as greatest width; pleonites 1 and

2 freely articulating; pleonite 3 indicated by lateral suture

barely visibly in dorsal view; pleonite 4 very short, with short

suture; remaining pleotelson parallel-sided, with semicircular

apex, dorsally evenly domed curving posteriorly to oblique

profile in lateral view, posterodistal margin not elevated.

Antenna 1 reaching to distal margin of second article of

antenna 2; flagellum of major article plus 2 minute articles.

Antenna 2 0.25 body length; flagellum broken.

Mandible (right) molar truncate, with acute toothed

accessory blade, with 20 molar setae; spine row of 9 spines,

lacinia mobilis bifid; incisor with 3 blunt teeth.

Coxa 2 subrectangular; coxa 3 tapering posteriorly, 0.9

pereonite dorsal length; coxa 4 smaller than 3; coxae 5-7

overlapping, ventrally concave, projecting acutely

Four new valviferan isopods from diverse tropical Australian habitats

329

Figure 1. Cleantioides Carpentaria sp. nov. Holotype.

posteroventrally; coxa 7 reaching back to mid-pleonite 2.

Pereopod 1 ischium with prominent proximal facial robust

seta; merus with 2 robust setae on flexor margin, complex seta

distally on extensor margin; propodus 2.5 times as long as

greatest depth, palm with 5 strong setae and comb of fine short

setae, mesial face with 10 pectinate setae; dactylus almost

linear except for curved unguis, reaching back to mid-carpus.

Pereopod 2 1.3 times as long as pereopod 1; propodus 4.6

times as long as wide; dactylus as long as propodus. Pereopod

3 1.1 times as long as pereopod 2; propodus 6.4 times as long

as wide; dactylus almost as long as propodus. Pereopods 4, 5

missing. Pereopod 6 0.4 times length of pereopod 3; propodus

extensor margin projected beyond articulation with merus;

merus-dactylus 1.5 times length of propodus; dactylus

twisted, compact. Pereopod 7 as long as pereopod 6; propodus

extensor margin scarcely projected beyond articulation with

merus; merus-dactylus 3.7 times length of propodus; dactylus

twisted, damaged.

Uropod 2.5 times as long as greatest width; endopod about

0.4 length of total uropod, triangular with rounded apex,

suture at 75° to long axis; with 1 seta.

Etymology. Cidaris, from the name of the cruise during which

the specimen was taken; noun in apposition.

Distribution. Australia, Coral Sea, eastern continental slope of

Queensland, c. 17°S; 458-500 m depth.

Remarks. The single specimen has been partially dissected by

a previous investigator and most mouthparts, all pereopods of

the left side and pereopods 4 and 5 of the right are missing.

Nevertheless, the species is undoubtedly of this genus. The new

species differs from the two others in the absence of a median

carina on the anterior part of the pleotelson, absence of

upcurved lateral pleotelson margins and more elongate anterior

pereopods. Zenobianopsis cidaris has a more rounded uropodal

endopod than Z. rotundicauda. In Z. caeca , the pleotelson is

decidedly truncate.

Idoteidae Samouelle, 1819

Synidotea Harger, 1878

Synidotea Harger, 1878: 374. - Poore and Lew Ton, 1993: 261—

262.

Remarks. The valviferan idoteid genus Synidotea occurs world¬

wide and is represented by 59 species, most highly endemic

(Schotte et al., 2008 onwards). Chapman and Carlton (1991,

1994) suggested that some described species are in fact a single

one, the Japanese Synidotea laevidorsalis Miers, 1881

translocated elsewhere, especially to ports. This assertion was

disputed by Poore (1996) who showed that morphological and

ecological differences could be shown between several

examples. Nevertheless, one other species does appear to be

translocated. Synidotea laticauda Benedict, 1897, first

described from the estuarine parts of San Francisco Bay, is

now known, though wrongly identified, from the Gironde

Estuary, France (Mees and Fockedey, 1993), the Guadalquivir

River estuary, Spain (Cuesta et al., 1996) and Delaware Bay,

USA (Buschek and Boyd, 2006).

Most idoteid genera are restricted to temperate shores

but Synidotea is not (Schotte et al., 1995 onwards). Northern

Australia is within the broad Indo-West Pacific biogeographic

region and species recorded there could potentially occur in

the Australian tropics. Four species have been described

from India: S. variegata Collinge, 1917, S. fluviatilis Pillai,

1954, S. worliensis Joshi and Bal, 1959 and S.

hanumantharaoi Kumari and Shyamasundari, 1984. Two, S.

fecunda Javed and Yasmeen, 1994 and S. indica Javed and

Yasmeen, 1994 occur in Pakistan and S.poorei Cai and Teo,

2012 in Singapore. Further afield, S. oahu Moore, 2004 was

described from Hawaii and S. pacifica Nobili, 1906 from

Tuamoto. All of these belong to the so-called S. hirtipes

Milne Edwards, 1840 group of species; S. hirtipes is from

southern Africa. Both new species differ from the Indian

Ocean species as follows: from S. hirtipes in the absence of

330

G.C.B. Poore

Figure 2. Cleantioides Carpentaria sp. nov. Holotype. Left antennae 1 and 2 detail in ventral view, x = ventral view of left side of pereonite 7 and

anterior pleotelson showing pleonal epimera 1-3. Scale bar = 1 mm (habitus only) and 0.2 mm (pereopods).

Four new valviferan isopods from diverse tropical Australian habitats

331

Figure 3. Zenobianopsis cidaris sp. nov. Right limbs from holotype. Scale bars = 1 mm (habitus only) and 0.2 mm (pereopods, uropod).

332

G.C.B. Poore

Figure 4. Zenobianopsis cidaris sp. nov. Right limbs from holotype; detail of p6 untwisted.

ridges on the uropod (Kensley, 1978), from S.fecunda in the

absence of bosses on the head, from S. indica in the less

angled pereonite margins, from S. variegata in the more

robust antenna 2 and less obviously triangular uropodal

endopod, and from S. hanumantharaoi and S. worliensis in

the more elongate antenna 2. The new species differ from S.

poorei in absence of or less dense persistent colour and more

elongate antenna 2.

The four Australian species already described are

temperate: S. grisea Poore and Lew Ton, 1993 and S. keablei

Poore and Lew Ton, 1993 from southeastern states, S. watsonae

Poore and Lew Ton, 1993 from southern WA and Vic., and an

undescribed species from WA. Of these, the new species differ

from S. grisea and S. keablei most obviously in the broader

pleotelsonic apex, and from S. watsonae in the absence of

sculpture. The two species differ from each other most

obviously in colour, proportions of the articles of antenna 2,

pereopods and uropodal endopod.

Synidotea innatans sp. nov.

Figures 5-7

Material examined. Holotype, Australia, Timor Sea, (11°42.4'S,

125°06'E), discarded fragment of fishing net floating on sea surface,

G.C.B. Poore et al. (with Consulting Environmental Engineers), 26

Oct 2001, NMV J62816 (male, 11.3 mm). Paratypes, same locality

NMV J62817 (ovigerous female, 7.1 mm), NMV J62818 (12 males,

6.6-10.8 mm, 17 ovigerous females, 6.6-7.3 mm, 15 juveniles, 4.4-7.5

mm ). All specimens fixed in 70% alcohol.

Description. Male. Body 3.0 times as long as greatest width at

pereonite 3, dorsal surface smooth, pale, ornamented with

numerous, small distinct chromatophores, arranged in a dense

median stripe, densely on head, pleotelson and laterally, in

oblique wavy bands midlaterally on pereonites, and with even

single rows along posterior margins of pereonites; without dorsal

sculpture. Head 2.0 times as wide as long, 0.7 times width of

pereonite 3, front straight, strongly tapering in front of eyes, with

shallow transverse depression; eye bulging, 0.40 times as long as

Four new valviferan isopods from diverse tropical Australian habitats

333

Figure 5. Synidotea innatans sp. nov. Holotype male top, paratype female below.

median head length. Pereonite 1 0.80 width of pereonite 3,

margin with rounded obtuse angle one-quarter from anterior

suture, posterior three-quarters with parallel margins. Pereonite

2 with rounded anterolateral margins, parallel-sided over

posterior three-quarters. Pereonite 3 lateral margin broadly

convex. Pereonite 4 lateral margin broadly convex. Pleotelson

1.33 times as long as greatest width; tapering beyond pleonite 1

suture to 0.85 of greatest width to an obtuse angle at 0.7 length,

then more steeply to narrow, barely-excavate posterior margin.

Antenna 1 flagellum 0.9 length of peduncle, with 9 pairs of

aesthetascs. Antenna 2 0.5 body length; article 4 2.5 times as

long as wide; article 5 4.3 times as long as wide; flagellum

with 12 articles, 0.8 length of peduncle.

Maxilla 2 outer lobe strongly produced laterally, with

marginal row of 24 long plumose setae. Maxillipedal basal

endite longer than wide, with 1 coupling hook, apex setose.

Maxillipedal palp 1.6 times as long as greatest width; article 3

1.4 times as long as wide. Epipod 1.2 times as long as wide,

with broad transverse apex.

Pereopods with dense mat of setation on flexor margins of

merus-propodus. Pereopod 1 propodus 1.5 times as long as

greatest depth, palm excavate, mesial face with about 90

plumose setae; dactylus almost linear except for curved

unguis, reaching back to base of carpus. Pereopod 2 propodus

1.2 times as long as merus and carpus together, 3.0 times as

wide as long. Pereopod 4 propodus 2.9 times as wide as long.

Pereopod 7 propodus 3.2 times as wide as long.

Penial plate 1.6 times as long as wide, double-waisted,

with excavate distal margin.

Pleopod 2 with appendix masculina 1.27 times as long as

endopod, with rounded apex, distally with numerous

superficial spines. Uropod 3.8 times as long as distal peduncle

width; endopod about 0.25 length of peduncle, mesial length

0.7 proximal suture length, suture at 80° to long axis, distal

margin convex-truncate, at 80° to long axis, lateral margin

curved into distal margins; with 3 setae.

Ovigerous female. Body 2.5 times as long as greatest

width at pereonite 3. Head 2.4 times as wide as long, 0.64

times width of pereonite 3; eye bulging, 0.55 times as long as

median head length. Pereonite 1 0.80 width of pereonite 3,

margin barely convex, oblique. Pereonite 2 margin barely

convex, oblique. Pereonite 3 margin barely convex, parallel¬

sided. Pereonite 4 margin barely convex. Pleotelson 1.3 times

as long as greatest width; tapering beyond pleonite 1 suture to

0.75 of greatest width to an obtuse angle at 0.7 length, then

more steeply to narrow, barely-excavate posterior margin.

1 mm

334

G.C.B. Poore

Figure 6. Synidotea innatans sp. nov. Holotype male left, paratype female right. Scale bar = 1 mm (habitus only).

Four new valviferan isopods from diverse tropical Australian habitats

335

Figure 7. Synidotea innatans sp. nov. Limbs from holotype male.

Antenna 2 0.5 body length; article 4 2.2 times as long as

wide; article 4 3.6 times as long as wide; flagellum with 13

articles, 1.3 times length of peduncle. Maxilla 2 outer lobe not

expanded. Pereopods with mat of setae on flexor margins, less

dense on pereopods 4 and 5 and sparse on 6 and 7.

Etymology. From innatare, Latin, to float, alluding to the

discovery of this species on flotsam.

Distribution. Australia, Timor Sea, c. 11°S; on flotsam.

Remarks. The colour pattern of small chromatophores arranged

in regular rows persists and is characteristic of Synidotea

innatans. The new species differs in several respects from all

other species in the region including S. poorei collected from

buoys near Singapore (Cai and Teo, 2012) and from others as

detailed above.

The discovery of numerous specimens of this species on

flotsam (an abandoned fragment of a fishing net), in the

company of numerous shrimps, crabs and fishes, 170 km from

land (the island of Timor) raises questions about the

distribution of this species. This is only the second record of

a species of Synidotea from this habitat. Hobday (2000)

reported the normally benthic Synidotea harfordi Benedict,

1897 from the kelp Macrocystis pyrifera off the Californian

coast. The new species is less elongate than S. harfordi and

differs from the only two others known from benthic habitats

near to where it was taken, that from Singapore and S.

karumba, as detailed above. Thiel and Gutow (2005) reviewed

the distribution of 13 rafting species of Idotea worldwide.

Only one, I. metallica (Bose, 1802), would appear to be an

obligate rafter and it is cosmopolitan in the world’s oceans

(Poore and Lew Ton, 1993 for numerous citations; Abello and

Frankland, 1997). There is no suggestion that the new species

is similarly widespread. The ability of these species to raft

can not be used to suggest that this is how S. laticauda became

distributed between estuaries in San Francisco Bay, Delaware,

France and Spain (Mees and Fockedey, 1993; Cuesta et al.,

1996; Buschek and Boyd, 2006). Transportation by shipping

seems more probable.

336

G.C.B. Poore

Synidotea karumba sp. nov.

Figures 8-10

Material examined. Holotype. Queensland, Karumba, Norman River

mouth (17°28'S, 140°49'E), sandy sediment with seagrasses Halodule

uninervis and H.pinifolia, 1 m, K. Neil, 2001 NMV J53202 (male, 9.1

mm). Paratypes, same locality, 5 m, NMV J53203 (2 females, 6.3,

7.1 mm).

Description. Male. Body 3.1 times as long as greatest width at

pereonite 3, dorsal surface smooth, pale, ornamented with

numerous, evenly spaced, small distinct chromatophores;

without dorsal sculpture. Head 1.6 times as wide as long, 0.57

times width of pereonite 3, middorsum evenly domed, without

ornamentation, with shallow transverse depression near

anterior margin, front barely excavate, strongly tapering in

front of eyes; eye bulging, 0.37 times as long as median head

length. Pereonite 1 0.75 width of pereonite 3, lateral margin

with definite rounded obtuse angle one-third from anterior

suture, posterior two-thirds with parallel margins. Pereonite 2

with rounded anterolateral margins, narrower at posterior

suture. Pereonite 3 lateral margin evenly convex. Pereonite 4

lateral margin broadly convex. Pleotelson 1.2 times as long as

greatest width; tapering beyond pleonite 1 suture to 0.8 of

greatest width to an obtuse angle at 0.75 length, then more

steeply to excavate posterior margin.

Antenna 1 flagellum 0.9 length of peduncle, with 12 pairs

of aesthetascs. Antenna 2 0.7 body length; article 4 3.6 times

as long as wide; article 5 6.2 times as long as wide; flagellum

with 21 articles, as long as peduncle.

Maxilla 2 outer lobe expanded, with 17 long plumose

setae. Maxillipedal basal endite longer than wide, with 1

coupling hook, apex setose. Maxillipedal palp 2.0 times as

long as greatest width; article 3 1.3 times as long as wide.

Epipod 1.5 times as long as wide, with broad oblique apex.

Pereopods with dense mat of setation on flexor margins of

merus-propodus. Pereopod 1 propodus 1.7 times as long as

greatest depth, palm excavate, mesial face setose; dactylus almost

linear except for curved unguis, reaching back to base of carpus.

Pereopod 2 propodus as long as merus and carpus together, 2.9

times as wide as long. Pereopod 4 propodus 3.3 times as wide as

long. Pereopod 7 propodus 3.8 times as wide as long.

Penial plate 1.6 times as long as wide, slightly waisted,

with rounded apex.

Pleopod 2 with appendix masculina 1.25 times as long as

endopod, with rounded apex, with few superficial spines. Uropod

3.9 times as long as distal peduncle width; endopod 0.28 length

of peduncle, mesial length 0.8 proximal suture length, suture

almost at right-angles to long axis, distal margin truncate, at 75°

to long axis, lateral margin straight and with curve between

lateral and distal margins; with 1 seta.

Female. Body 2.6 times as long as greatest width at

pereonite 3. Head twice as wide as long. Pereonite 1 0.8 width

of pereonite 3, lateral margin more evenly curved than in

male. Pereonite 2 with rounded anterolateral margins, parallel¬

sided over posterior two-thirds. Pleotelson 1.1 times as long as

greatest width; tapering to 07 of greatest width beyond

pleonite 1 suture to an obtuse angle at 0.75 length, then more

steeply to excavate posterior margin.

Antenna 2 0.7 body length; article 4 3.2 times as long as

wide; article 4 6.9 times as long as wide; flagellum with 16

articles, 1.2 times length of peduncle. Maxilla 2 outer lobe not

expanded. Pereopods without mat of setae on flexor margins.

Etymology. Karumba, from the town where the specimens

were found, noun in apposition.

Distribution. Australia, Queensland, west coast of Cape York

at c. 17°S; 1 m depth.

Remarks. Synidotea karumba was compared with others above.

Acknowledgements

This paper was supported in part by Australian Biological

Resources Study National Taxonomy Research Grant Program

contract CN211-13. Collecting expeditions in southern

Australia were also supported by ABRS grants. I thank Scott

Chidgey, Consulting Environmental Engineers, for the

opportunity to join him in fieldwork in the Timor Sea and all

those whose names appear below. Isopods from the CIDARIS

I cruise undertaken by M. Pichon at the Australia Institute of

Marine Sciences were lent by the late Peter Arnold and

returned to Niel Bruce, Museum of Tropical Queensland,

Townsville. Further material came from surveys undertaken

for the Introduced Marine Pests Group, Queensland Fisheries

Service, Cairns, by Kerry Neil, CRC Reef Research. Peter

Davie, Queensland Museum, Brisbane, chased and lent

missing specimens. Initial illustrations of two of these species

were prepared by Rainbo Dixon and Anna McCallum in my

lab. Martin Thiel, Coquimbo, provided leads on the biology of

rafting animals and Cai Yixiong shared his description of the

new species from Singapore.

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Memoirs of Museum Victoria 69:341-354 (2012)

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

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

A review of giant roughies of the genus Hoplostethus (Beryciformes,

Trachichthyidae), with descriptions of two new Australasian species.

Clive D. Roberts 1 and Martin F. Gomon 2

1 Museum of New Zealand Te Papa Tongarewa, PO Box 467, Wellington, New Zealand (cliver@tepapa.govt.nz)

2 Ichthyology, Sciences Department, Museum Victoria, GPO Box 666, Melbourne, Victoria, 3001, Australia (mgomon@

museum .vie .gov.au)

Abstract Roberts, C.D. & Gomon, M.F. 2012. A review of giant roughies of the genus Hoplostethus (Beryciformes, Trachichthyidae),

with descriptions of two new Australasian species. Memoirs of Museum Victoria 69: 341-354.

Hoplostethus gigas McCulloch, 1914 and two previously unnamed species of the genus that reach a similarly

‘giant’ size are described. The redescription of H. gigas, which is confined in distribution to the southern coast of Australia,

is based on specimens identified as comprising at least part of McCulloch’s type series (one herein designated lectotype),

together with subsequently collected material. The very similar H. melanopeza sp. nov., occurring in northern New

Zealand and southeastern Australian waters, as well as seamounts in the intervening Tasman Sea, is distinguished by the

distinctly black outer margins of fins in large adults and count of predorsal scales. The New Caledonian H. grandperrini

sp. nov., known only from the two types, differs from previously described species in having the combination of 17 or 18

pectoral-fin rays, 13 or 14 abdominal scutes, 19 or 20 total gill rakers and a buccal cavity that is mostly pale, the only black

pigmentation occurring posterior to the gill arches.

Keywords Giant roughy, Trachichthyidae, Hoplostethus, H. gigas, lectotype, species nov., Australasia

Introduction

In synthetic treatments of the Trachichthyidae and the

Beryciformes, Kotlyar (1986, 1996) listed 23 species in the

roughy genus Hoplostethus Cuvier (in Cuvier & Valenciennes,

1829). Thirteen of these he referred to the subgenus

Hoplostethus (Hoplostethus ), which he diagnosed as having

pale colouration, scales on the ventral midline of the abdomen

modified into enlarged, thickened scutes, 25-27 total

vertebrae, dorsal fin with 4-8 (usually 6) spines and 14-18

(usually 15 or 16) soft rays, 14-19 (usually 15 or 16) pectoral-

fin rays, 14-25 (usually 16-21) predorsal scales and 15-50

simple pyloric caeca. The resurrection of H. latus McCulloch,

1914 by Gomon (in Gomon et ah, 1994) and descriptions of H.

vniro by Kotlyar (1995), H. ravurictus by Gomon (2008) and

H. robustispinus by Moore and Dodd (2010) brought the

number referrable to the subgenus, according to Kotlyar’s

criteria, to 17. Moore and Dodd (2010) listed H. intermedius

(Hector, 1875) as another species that would be referrable to

this assemblage, but genetic evidence supports its synonymy

with H. mediterraneus Cuvier (in Cuvier & Valenciennes,

1829) (Smith and Roberts, 2004). Although the validity of

Kotlyar’s four subgenera remains to be tested, the greatest

diversity of species in this large complex is clearly in the

Indo-Pacific. However, the conservative nature of the

morphology of species in the group has caused great confusion

about their numbers and identities (e.g. Kotlyar, 2011),

especially in the geographical areas of greatest diversity.

Of the 17 described species of Hoplostethus ( Hoplostethus ),

only four have been reliably recorded in the literature as

reaching what might be regarded as a truly large size, well in

excess of 250 mm SL, although we question one of these. The

southern Australian H. gigas McCulloch is certainly the

largest of the subgenus with one of the type specimens

measuring 525 mm SL (McCulloch, 1914). Hoplostethus

robustispinus Moore and Dodd, 2010, follows based on the

sole type from the Philippines reported to be 340 mm SL and

a 354 mm SL specimen from southern Japan (HUMZ 148072).

Kotlyar (1996) recorded H. mediterraneus as reaching 30 cm,

but only provided specimen information for specimens to 240

mm SL, the collection locality of these specimens reported as

the North Atlantic. No reliable, published account of the

species exceeding 200 mm SL is known from the Australasian

region. The fourth species reaching a size in the 250 mm SL

range is H. crassispinus from the Emperor Seamounts, which

Kotlyar (2011) stated was confined to that seamount group and

the Hawaiian Island chain. Although only reported by Kotlyar

(1996) to a size of 123 mm SL, specimens identified by Kawai

(in Inada and Wudianto, 2006) as H. crassispinus Kotlyar,

1980 (HUMZ 193945: 253 mm SL) and H. sp. (HUMZ 191163:

289 mm SL) are specimens of H. confinis Kotlyar, 1980 that

are considerably larger than any of Kotlyar’s specimens.

342

C.D. Roberts & M.F. Gomon

In 1989, Paulin et al. reported Hoplostethus gigas in New

Zealand waters, based on captures of surprisingly large

roughies from knolls and ridges in the Bay of Plenty on the

northeastern coast of New Zealand’s North Island. Subsequent

work on these specimens motivated Roberts (1995, 1996,

2012) to implore New Zealand fishers to bring in additional

examples of what he determined not to be H. gigas, but instead

a yet undescribed species. The largest of the specimens now in

hand exceeds a standard length of 500 mm, putting the species

on par with H. gigas with respect to size. The species is very

similar in appearance to H. gigas but is separable by colour

pattern and minor though consistent morphological

differences. In addition, a recent comparison of mitochondrial

DNA sequences clearly separate the two (Te Papa/ NIWA/

CSIRO/NMV unpublished data).

At about the same time as the discovery of the New

Zealand species, French surveys in New Caledonian waters

turned up examples of a roughy that attains a size of similar

magnitude, and which is clearly separable from H. gigas and

the New Zealand species on the basis of meristic features.

The purpose of this publication is to provide a detailed

redescription of H. gigas, together with descriptions and

names for these two new large congeners. The three are clearly

separable from each other and from other nominal species of

the genus on the basis of a variety of features.

Methods and Materials

Terminology and methodology is that of Kotlyar (1996). The

number and size range in standard length (SL) for each lot of

specimens examined is presented as a parenthetical expression

after the respective registration number; if a lot comprises a

single specimen only the standard length is presented.

Institutional abbreviations are listed in Leviton et al. (1985).

The descriptions are based on the lectotype or holotypes with

variations observed in paratypes following in parentheses.

Pectoral fins of the two new species were counted on both

sides. Gill rakers reported are those on the lateral face of the

first gill arch of the right side. Scale terminology is that of

Roberts (1993). The considerable variation in transverse scale

counts is attributable to the apparent irregular distribution of

scales on the sides of Hoplostethus species with lateral line

scales considerably larger than those elsewhere; although most

fishes have scales arranged in discrete oblique rows, a

homologous arrangement is not apparent in species of this

genus; in addition, the profusion of spines on scales of some

species makes the distinction between individual scales

extremely difficult. Modified scales on the ventral midline of

the abdomen are regarded as scutes when they have acquired a

laterally compressed keel-like form with a pointed apex or

apices; numbers of scutes in species having them appear to

increase slightly with growth.

Comparative material examined:

Hoplostethus confinis HUMZ 191395 (282), Indian Ocean,

off Sumatra, Indonesia, 03°29.03' N, 94°57.59' E - 03°29.23'

N, 94°57.22' E, 760-790 m, 7 October 2004, coll, by K. Odani;

HUMZ 194238 (270) Indian Ocean, off Java, Indonesia,

08°19.07' S, 109°53.09' E- 08°19.00' S, 109°52.08' E, 864-950

m, 7 May 2005, coll, by T. Kawai; H. robustispinus: HUMZ

148071 (195) and HUMZ 148072 (354) southern Japan,

27°53.20' N, 128°28.72' E -27°51.70' N, 128°29.05' E, 603-586

m, 18 July 1994.

Hoplostethus gigas McCulloch, 1914

Giant sawbelly

Figures 1 & 2; Tables 1 & 2

Hoplostethus gigas McCulloch, 1914: 101, plate xix, original

description. Great Australian Bight.

Hoplostethus gigas. McCulloch, 1929: 132, listed (“holotype on

deposit in Austr. Mus.”), Great Australian Bight. -Munro, 1958; 79,

fig. 548, description ex McCulloch, Great Australian Bight. -Whitley,

1964: 40, listed, Australia. -Scott, 1962: 108, fig., description ex

McCulloch, Great Australian Bight. -Woods and Sonoda, 1973: 306,

listed, Australia (“possibly a very large japonicus ”). -Kotlyar, 1980:

197, fig. 10, family revision, description ex McCulloch, Australia. -

Maxwell, 1980: 66, plate 155, description ex McCulloch, temperate

Australia. -Kotlyar, 1986: 126, generic revision, key, description ex

McCulloch, Australia. -Paxton and Hanley, 1989: 366, synonymy and

Australian distribution. -Gomon, in Gomon et al., 1994 (in part): 403,

Fig. 361, description. Great Australian Bight. -Paxton et al., 2006:

769, taxonomy, central south coast of Australia. -Gomon, in Gomon et

al., 2008: 424, fig., description, central south coast of Australia. -

Moore and Dodd, 2010: 138-141, morphological characters.

Hoplostethus latus (not McCulloch, 1914: 97, fig. 5). May and

Maxwell, 1986: 219, fig., description ex McCulloch, Great Australian

Bight.

Material examined. Lectotype. AMS 1.12766 (307) (herein

designated). Great Australian Bight, FIS Endeavour, one of six

registered, 27 March 1913.

Paralectotypes (5, 292-426 mm SL). AMS 1.15710-001 (ca. 375;

skeleton, S.1285) same data as AMS 1.12766; NMV A.20541 (385;

formerly E.4298) same data as AMS 1.12766; QMB 1.1423 (426;

formerly E.3238) Great Australian Bight, 33°18' S, 126°47' E, 238-311

m (130-170 fms), February 1913, FIS Endeavour.

Non-types (7, 293-378 mm SL). CSIRO H.4874-01 (366),

CSIRO H.4874-02 (368), CSIRO H.4874-03 (349), CSIRO

H.4874-04 (356), CSIRO H.4874-05 (373), CSIRO H.4874-06

(378) Great Australian Bight, 33°19’ S, 128°25’ E, 180-350 m,

collected at end of tow over rough ground near a drop-off to canyon,

FV Noble Pearl, demersal trawl, 21 September 1998; NMV A.21541

(293) Victoria, south-west of Portland, 38°48' S, 141°44' E, 432-522

m, 19 June 2000, FV Zeehaan, demersal trawl, coll, by K. Graham.

Diagnosis. Pectoral-fin rays 15, rarely 14; total gill rakers on

outer side of first arch 18; predorsal scales 9-15; abdominal

scutes 9 or 10, few scutes in large individuals with multiple

apical points; isthmus lacking scales; body scales adherent;

lateral-line scales with tuberculate medial ridge, but no spine;

scales on predorsal midline forming low raised ridge; body

ovoid and deep, depth 2.0-2.2 in SL; nape distinctly curved,

forehead almost straight to above upper lip; dorsal- and anal-

fin spines of moderate thickness; body of adults grey,

superimposed with orange-red to red in life, outer margin of

soft portions of dorsal, anal and caudal fins greyish to almost

blackish; buccal cavity and opercular recess black, vomer,

underside of tongue and upper surface of lower jaw stark white.

Reaches 525 mm SL.

Australasian giant Hoplosthethus

343

Table 1. Selected meristic, standard length and morphometric values for types and other specimens examined of three species of Hoplostethus.

Morphometric values are expressed as percent SL.

lectotype

.gigds

all specimens

H. melan

holotype

opeza sp.nov.

paratypes

H. grandpe

holotype

rrini sp.nov.

paratype

(n=9)

(n=27)

Dorsal fin

VI, 13

VI, 13-14

VI, 13

VI-VII, 12-13

VI, 13

Anal fin

III, 9

III, 9-10

III, 9

Caudal fin

6+2+17+2+6

6-7+2+17+2+6-7

6+2+17+2+6

6-7+2+17+2+6

6+2+17+2+6

Pectoral fin

14-15

14-16

17-18

Pelvic fin

1,6

Lateral line scales

26+2

26-28+1-2

27+1

26-29+1-2

28+1

Transverse scales

13/1/23

9-13/1/22-24

12/1/25

9-15/1/20-28

12/1/26

11/1/35

Predorsal scales

9-15

16-22

Abdominal scutes

9-10

9-12

Gill rakers

5+13=18

6+14=20

5-6+12-15=18-21

6+14=20

6+13=19

Pseudobranch

~46*

16-21

Vertebrae

11+15

Standard length

307

292-378

286

72.6-515

455

131

Body depth

47.6

46.2-50.7

52.0

47.9-56.4

55.6

53.2

Head length

37.1

34.7-39.1

40.6

32.6-42.6

41.4

42.9

Forehead height

39.8-40.1 1

41.2

34.1-46.4

41.0

44.2

Eye diameter

10.9

9.1-10.9

10.9

9.4-14.7

9.6

12.5

Postorbital length

19.1-20.0 1

20.3

17.8-21.8

22.4

22.0

Interorbital width

11.5

11.5-12.9

10.9

10.2-12.8

14.1

13.4

Maxillary length

26.0

24.4-27.5

26.5

23.3-29.2

26.3

30.4

Lower jaw length

24.7-28.4 1

28.2

23.6-29.6

28.6

31.1

Snout length

9.9

6.5-10.4

9.8

7.09-11.3

10.5

9.5

Caudal peduncle depth

11.3

9.9-12.5

12.0

10.4-15.3

12.5

13.0

Caudal peduncle length

25.2

21.9-27.1

23.0

20.2-26.1

22.6

22.9

Predorsal length

45.3

40.4-47.9

47.6

45.0-52.2

38.8

49.1

Preanal length

71.9-72.9 1

70.6

64.4-73.5

76.7

73.2

Prepectoral length

37.7-38.2 1

40.7

36.5-41.2

40.8

40.4

Prepelvic length

42.8-44.9 1

44.3

39.1-45.3

42.6

43.8

Pectoral Pelvic length

11.6-12.3 1

15.8

12.0-16.6

18.3

15.8

Pelvic Anal length

34.0-38.2 1

32.1

25.3-36.8

42.4

35.5

Dorsal base length

38.4-39.3 1

38.9

36.0-41.3

38.3

36.0

Anal base length

17.1-18.7 1

19.0

15.3-21.8

17.2

17.0

Pectoral fin length

23.8

23.8-27.3

23.8

21.1-34.9

25.2

34.0

Pelvic fin length

19.5

19.1-21.0

19.5

17.0-26.4

18.4

23.5

1st dorsal spine length

2.5-3 .T

3.9

1.8-7.5

2.9

4.7

2nd dorsal spine length

4.3-6.8 4

8.1

3.1-12.1

4.1

8.9

Last dorsal spine length

14.9

13.7-15.0 2

14.6

12.0-23.2

12.1

17.6

1st anal spine length

1.4-2.1 2

2.1

1.2-2.8

1.6

2.1

Last anal spine length

11.0

10.8-12.3 4

11.1

8.8-16.4

7.7

13.3

* 1 specimen only, 1 2 of 9 specimens, 2 6 of 9 specimens, 3 7 of 9 specimens, 4 8 of 9 specimens

344

C.D. Roberts & M.F. Gomon

Table 2. Frequency of abdominal scute numbers in specimens examined of Hoplostethus gigas, H. melanopeza sp. nov. and H. grandperrini sp. nov.

Holotypes or lectotype indicated by *.

No. of scutes

H. gigas

H. melanopeza sp.nov.

7 *

H. grandperrini sp.nov.

Figure 1. Hoplostethus gigas McCulloch, 1914. CSIRO H4874.02, 368 mm SL, photo by Thor Carter.

Description. (See Table 1 for frequencies of values for selected

meristic characters.) Dorsal-fin rays VI, 13 (VI, 14 in 1 of 9);

anal-fin rays III, 9; caudal-fin rays 6 + 2 + 9 + 8 + 2 + 6(6or

7 + 2 + 9 + 8 + 2 + 6or7); pectoral-fin rays 15 (14 in 1 of 9);

pelvic-fin rays I, 6; gill rakers 5+1 + 12; lateral-line scales 26

+ 2 (26 to 28 + 1 or 2 = 28 or 29); transverse scales 13/1/23

(9-13/1/22-24); predorsal scales 9 (9-15); scutes 10 (9 or 10;

Table 2); vertebrae 11 + 15; pseudobranch about 46;

branchiostegal rays 8.

(See Table 1 for comparative ranges of selected

morphometric characters.) Body ovoid and deep, depth 1.98-

2.16 in SL. Head large, its height slightly greater than its

length, 110-115% HL; upper profile in front of dorsal fin

distinctly curved to above rear of eye, then mostly straight to

upper jaw, slightly concave above rear half of eye in some;

anterodorsal profile well separated from upper orbital rim;

space between eyes wide, interorbital width 28.8-35.2% HL;

eye of moderate size, orbital diameter 24.0-32.8% HL; crests

of head bones strong, fine spinules on apices at skin surface;

depressions between crests moderately deep, hidden by thick

skin in adults; infraorbital bones becoming progressively

broader with growth; nostrils immediately preceding orbits,

posterior nostril two to four times area of anterior nostril;

mouth reaching just beyond vertical through hind margin of

eye; large, fine denticulate teeth covering oral margins and

exposed lateral surfaces of premaxilla and dentary, palatine

with posteriorly tapering band of similar teeth, vomer

apparently lacking teeth, at least in adults; tip of dentary with

ossified knob at symphysis. Preopercular spine short, reaching

about 1/4 way from preopercular margin to pelvic-fin base,

broad basally in large specimens. Humeral and preopercular

spines of similar size. Longest gill raker about 2/3 eye

diameter; gill filaments at angle of first gill arch very short,

about 1/8 eye diameter and about 1/3 length of longest

filaments of pseudobranch.

Body covered with adherent scales, with densely covered,

finely spinoid scales above lateral line, posteriorly and low on

side, scales above and covered laterally by pectoral fin cycloid

(cycloid scales distributed more ventrally in smaller individuals);

head naked except for patch of scales on cheek posterior to rear

tip of maxilla in about four vertical rows, posteriormost row with

about 17 scales; isthmus scale-less; each lateral-line scale with

tuberculate ridge but lacking distinct spine; deep serrated

abdominal keel formed from greatly enlarged scales (scutes)

with slender spine-like apices, most without sculpturing or

multiple tips, some striated; scales on dorsal midline in front of

dorsal fin slightly but noticeably raised, their spinules complex

but not greatly enlarged. First dorsal-fin spine short, subsequent

spines distinctly longer but only increasing progressively in

length slightly; spines of moderate thickness, those posteriorly

progressively thicker with distinct lengthwise striations; first soft

ray distinctly longer than last spine, third ray longest, subsequent

Australasian giant Hoplosthethus

345

Figure 2. Collection localities for specimens examined of Hoplostethus gigas (squares), H. melanopeza sp. nov. (circles) and H. grandperrini sp.

nov. (diamonds). Red symbols indicate holotypes or lectotype.

rays progressively decreasing in length, outer margin of soft

dorsal fin curved anteriorly, straight posteriorly. First anal-fin

spine short, second short or of intermediate length, and third

long; spines of similar robustness to those in dorsal fin. Caudal

fin distinctly forked, lobes of moderate breadth and rounded;

middle rays about 40% length of longest rays. Pectoral fin

moderately short, reaching to beyond anus in large specimens.

Pelvic fin reaching to seventh or eighth scute in large specimens.

A large species, largest specimen examined 426 mm SL;

reported to 525 mm SL (McCulloch, 1914).

Colour in life. Head and body grey, obscured by deep reddish

orange to red; opercle black suffused with red; buccal cavity

and opercular recess black, vomer, underside of tongue and

upper surface of lower jaw stark white; fins deep red;

membranes between dorsal fin spines grey, marginal strip on

soft portions of dorsal, anal and caudal fins dark grey to almost

blackish in large individuals (fig. 1).

Pigmentation in alcohol. Slightly dusky above, pale below;

opercle dark; buccal and branchial chambers dark, including

gill arches and rakers; vomer, palatines, underside of tongue

and lower jaw uniformly pale; medial fins and pelvic fins dusky

near outer edges.

Distribution. Confined to the southern coast of Australia,

documented from the western part of the Great Australian

Bight (126°47' E) to southwest of Portland, Victoria (141 °44' E)

with collection depths recorded between about 188 and 522 m

(fig. 2). It is reported to “hang-out at canyon edges” and in areas

with “rough ground” (pers. comm. T. Parsons, skipper of ‘FV

Noble Pearl').

Comments. The type locality given by McCulloch (1914) in his

original description of Hoplostethus gigas does not accurately

match registration records for specimens collected by the FIS

Endeavour we have been able to locate. In his account,

McCulloch presented it as Great Australian Bight, 33° 18' S,

126°42' E, 130-170 fms (= 238-311 m). The closest locality we

have been able to find for specimens identifiable as this species

is that of three specimens registered in the South Australian

Museum (SAM F137, formerly AMS E.3236), Western

Australian Museum (WAM P.63-001, formerly E.3237) and

Queensland Museum (QMB 1.1423, formerly E.3238). So far,

the specimens at the South Australian Museum and Western

Australian Museum have not been found, although specimens

of Hoplostethus collected by the Endeavour were received and

registered by the institutions as indicated, and in both cases,

like the Australian Museum material, they were recorded as H.

intermedius. As no further Endeavour material identifiable as

H. gigas appears to be in the Australian Museum collection, we

assume the other six specimens of the type series were

distributed to museums elsewhere in Australia, along with an

assortment of other “E series” specimens currently in their

collections. For instance, in addition to the assumed type of H.

346

C.D. Roberts & M.F. Gomon

gigas in the Museum Victoria collection listed above among

Material examined, a specimen of H. latus was received as part

of the same gift. That specimen, NMV R5962 (formerly AMS

E.2350) is most likely one of the nine specimens on which

Hoplostethus mediterraneus var. latus was based in the same

McCulloch publication and is consequently considered a

syntype of what is now regarded as the valid species, H. latus.

The ledger of original Endeavour numbers, which is in the

care of the Australian Museum, records E.3236, E.3237 and

E.3238 as having been collected at 33°18'S, 126°47'E, 130-170

fms and having been exchanged to the Adelaide, Perth and

Queensland Museums, respectively. Although identified in the

ledger as Hopostethus intermedius these are the specimens of

H. gigas listed above. We suspect the discrepancy in minutes

longitude is a misinterpretation of the handwritten record or a

transcription error. Despite his lone locality for the species,

McCulloch indicated the account is based on eleven specimens.

We consider it unlikely that all were collected at the one

station as specimens of this species are uncommonly rare in

collections and the few that do exist were not taken in large

numbers. As McCulloch was zoologist at the Australian

Museum at the time of publication, we assume he retained at

least one type and AMS 1.12766 is one of two specimens

currently registered in that collection as H. gigas, even though

it was registered as Hoplostethus intermedius without locality

information on 27 April 1913. No subsequent annotations were

made to that ledger entry. The other specimen so identified in

the AMS collection is a skeleton (AMS 1.15710-001) prepared

about the time of registration. AMS Skeleton Register records

in the hand of McCulloch “S.1285, 28 Mar 1913, Hoplostethus

intermedius” with intermedius crossed out and “gigas”

inserted, in the same hand.

Based on our perceptions of human nature, we assume that

McCulloch chose to retain the specimen he regarded as the

most representative of the species (now regarded as a holotype).

We adjudge AMS 1.12766 to be that specimen and here

designate it lectotype of Hoplostethus gigas McCulloch, 1914.

Other specimens listed above as paralectotypes are considered

to also be from the original series.

The redescription presented here is the first since the original

description of the species nearly 100 years ago, and includes

additional specimens and data ranges. This is the largest of

currently described species and is easily separated from other

nominal species by its maximum size attained and the combined

morphological features comprising 9-15 predorsal scales, 9-10

enlarged abdominal scutes, 15 (rarely 14) pectoral-fin rays, and

18 total gill rakers. It is separable from the two species described

below as discussed in the commentary following each treatment.

Hoplostethus melanopeza sp. nov.

New Zealand giant sawbelly

Figures 2-5; Tables 1 & 2

Hoplostethus gigas (not McCulloch, 1914). Paulin & Stewart,

1985: 31, listed. Bay of Plenty, 100-300 m, first record for New

Zealand. -Paulin et al., 1989: 153, 257, colour plate (opposite page

163), key. -Gomon, in Gomon et al., 1994 (in part): 403, Fig. 361,

description, off Sydney, NSW, and Bay of Plenty, New Zealand.

Hoplostethus Igigas (not McCulloch, 1914). Roberts, 1995: 106,

colour figs, description. Bay of Plenty. -Roberts, 1996: 40, colour fig.,

description. Bay of Plenty.

Hoplostethus cf. gigas (not McCulloch, 1914). Roberts et al.,

2009: 532 (listed). -Roberts, 2012: 38, colour fig., description, off

Mayor Island, Bay of Plenty.

Material examined. Holotype. NMNZ P.053205 (286) New Zealand,

North Island, Bay of Plenty, Mayor Knolls, 12 km east of Mayor

Island, 37°19.07' S, 176°25.35' E, 320 m, gill net, 3 hr soak, 5 March

2012, FV Ruben Jack, A. Oliver & C. Molloy.

Paratypes. (27, 72.6-515 mm SL). AMS 1.27085-001 (515)

Australia, New South Wales, Taupo Seamount, 32° S, 155° E, March

1982; AMS 1.30415-001 (2, 72.6-123) Australia, New South Wales,

Moruya, 36°03' S, 150°27' E, 383 m, 21 November 1979, K. Graham;

AMS 1.40390-004 (108) Australia, New South Wales, Bermagui, 36°47'

S, 150°21' E, 585 m, 23 July 2000, K. Graham; CSIRO H5321-04

(107), Australia, Victoria, Cape Everard, 38°14' S, 149°36’ E, 486-602

m, 23 July 2000, K. Graham; CSIRO H7387-01 (380) formerly NMNZ

P.053725, New Zealand, Bay of Plenty, east of Mayor Island, west of

Rangitira Knoll, 37°15.52’ S, 176°44.4P E, 500 m, set net, April 2012,

FV Ruben Jack, OPC Fish & Lobster Ltd, A Oliver & C Molloy; MNHN

2012-0268 (273) formerly NMNZ P.053206, same data as holotype;

NMNZ P.014162 (465) New Zealand, North Island, Bay of Plenty,

Rangitira Knoll, 37° 15.5’ S, 176°5P E, 140 m, June 1983, G. Schroeder;

NMNZ P.015181 (2, 440-450) New Zealand, Tokokemoke Knoll, 12

miles west of White Island, 37°28’ S, 176°54’ E, 256 m, longline, FV

Arapawa I, C. Walker; NMNZ P.015854 (3, 397-467) New Zealand,

North Island, Bay of Plenty, Rangitira Knoll, 37°15’ S, 176°5P E, 366 m,

February 1984, gill net, G. Schroeder; NMNZ P.031100 (3, 449-482)

New Zealand, North Island, Bay of Plenty, southeast of Rangitira Knoll,

37° 17.2' S, 176°53.6’ E, 240-500 m, April 1994, bottom trawl, FV

Margaret Philippa, J. & J. McGrath; NMNZ P.038312 (99) New

Zealand, southern Kermadec Ridge, outer Bay of Plenty, Rumble 3

submarine volcano, 35°44.51' S, 178°29.62' E, 270^-26 m, epibenthic

sled, 11:31-11:51 hrs, 19May 2001, GRV Tangaroa ; stn. TAN 0107/004;

NMNZ P.038325 (395) Gascoyne Seamount, 36°42.00' S, 155°54.00' E,

925 m, 8 August 2001, dropline, D. Smith; NMNZ P.038548 (178) same

data as NMNZ P.015854; NMNZ P.047799 (132) Tasman Sea, Lord

Howe Rise, Central Plateau 34° 11.95’ S, 162°38.90’ E, 712-760 m, 7

September 2010, trawl, FV Voyager, OBS 3177/058, J. Houston; NMNZ

P.053308 (486) New Zealand, North Island, Bay of Plenty, east of Mayor

Island, 37° 19.75' S, 176°29.57' E, 540 m, set net, March 2012, OPC Fish

& Lobster Ltd., coll. A Oliver & C Molloy; NMNZ P.053877 (468) New

Zealand, Bay of Plenty, Mayor Knolls, east of Mayor Island, 37° 18.6’ S,

176°31.32’ E, 347 m, set net, April 2012, FV Ruben Jack, A. Oliver & C.

Molloy; NMV A22070 (128) Tasman Sea, Lord Howe Rise, 33°38’ S to

33°40' S, 162°21' E to 162°28' E, demersal trawl, 300-750 m, 22 March-2

April 2001, coll. K. Smith; NMV A30942-001 (282) formerly NMNZ

P.053307, New Zealand, Bay of Plenty, east of Mayor Island, 37°19.75'

S,176°29.57' E, set net, 540 m, March 2012, FV Ruben Jack, OPC Fish

& Lobster Ltd., coll. A. Oliver & C. Molloy; NMV A30943-001 (393)

formerly NMNZ P.053726, New Zealand, Bay of Plenty, E of Mayor

Island, Rangitira Knoll, 37°15.522' S,176°44.408' E, set net, 500 m, April

2012, FV Ruben Jack, OPC Fish & Lobster Ltd., coll. A. Oliver & C.

Molloy; NMV A30944-001 (420) formerly NMNZ P.054048, New

Zealand, Bay of Plenty, E of Mayor Island, Maungaiti Knoll, 37°17.25’

S,176°51.63’ E, set net, 450 m, June 2012, FV Ruben Jack, OPC Fish &

Lobster Ltd., coll. A. Oliver; USNM 406863 (262) formerly NMNZ

P.053529, New Zealand, North Island, Bay of Plenty, Mayor Knolls, 12

km east of Mayor Island, 37°19.07' S, 176°25.35' E, 320 m, 5 March

2012, FV Ruben Jack, coll. A. Oliver & C. Molloy.

Diagnosis. Pectoral-fin rays 15, rarely 13, 14 or 16; total gill

rakers on outer side of first arch 18-21; abdominal scutes 9-12,

Australasian giant Hoplosthethus

347

Figure 3. Hoplostethus melanopeza sp. nov. A, NMNZ P.053205, holotype, 286 mm SL, photo by C. Struthers; B, NMNZ P.0533 08, paratype,

486 mm SL, photo by C. Struthers; C, NMV A22070, paratype, 128 mm SL, photo by M. Gomon.

348

C.D. Roberts & M.F. Gomon

Figure 4. Buccal colouration of Hoplostethus melanopeza sp. nov. NMNZ P.053206, paratype, 273 mm SL, photo by C. Struthers.

some scutes in large individuals with multiple apical points;

predorsal scales, 16-22; isthmus lacking scales; body scales

adherent; lateral-line scales with strong medial ridge, but no

spine in small and medium sized adults, with strong spine

centrally on posterior margin in large adults; scales on predorsal

midline forming low raised ridge; body ovoid and deep, depth

1.9-2.1 in SL; nape gently curved, forehead almost straight to

above upper lip; dorsal- and anal-fin spines of moderate

thickness; body of adults grey, superimposed with deep red in

life, outer margin of all fins with narrow black edge in adults;

buccal cavity and opercular recess black, vomer, margins of

mouth roof lateral to palatines, underside of tongue and upper

surface of lower jaw stark white. Reaches at least 515 mm SL.

Description. (See Table 1 for frequencies of values for selected

meristic characters.) Dorsal-fin rays VI, 13 (VI, 12 in 2 and

VII, 12 in 1 of 27); anal-fin rays III, 9 (III, 10 in 1 of 27);

caudal-fin rays 6 + 2 + 9 + 8 + 2 + 6(7 + 2 + 9 + 8 + 2 + 6in2

of 21); pectoral-fin rays 15 (13 in 1, 14 in 2 and 16 in 4 of 54);

pelvic-fin rays I, 6; gill rakers 6 + 1 + 13 (5 or 6 + 1 + 12-14 =

18-21, usually 6 + 1 + 13 = 20, mean total 19.7); lateral-line

scales 27+ 1 (26-29 + 1, rarely 2, mean 27.7 + 1.1 ); transverse

scales 12/1/25 (9-15/1/20-28; mean 11.1/1/23.4); predorsal

scales 18 (16-22, mean 18.5); scutes 10 (9-12; Table 2);

vertebrae 11 + 15; pseudobranch 19 (16-21); branchiostegal

rays 8.

(See Table 1 for comparative ranges of selected

morphometric characters.) Body ovoid and deep, depth 1.92-

2.08 in SL. Head large, its height slightly greater than its

length, 94.1-114% HL; upper profile in front of dorsal fin

gently curved to above rear of eye, then mostly straight to

upper jaw, slightly concave above rear half of eye in some;

anterodorsal profile only slightly separated from upper orbital

rim; space between eyes wide, interorbital width 28.8-35.2%

HL; eye of moderate size, orbital diameter 24.0-32.8% HL;

crests of head bones strong, fine spinules on apices at skin

surface; depressions between crests moderately deep, hidden

by thick skin in adults; infraorbital bones becoming

progressively broader with growth; nostrils immediately

preceding orbits, posterior nostril two to four times area of

anterior nostril; mouth reaching just beyond vertical through

hind margin of eye; large, fine denticulate teeth covering oral

margins and exposed lateral surfaces of premaxilla and

dentary, palatine with narrow band of similar teeth, vomer

apparently lacking teeth (with about 3 tiny teeth in small

specimens); tip of dentary with ossified knob at symphysis.

Preopercular spine short, reaching about 1/4 to 1/2 way from

preopercular margin to pelvic-fin base, broad basally in large

specimens. Humeral and preopercular spines of similar size.

Longest gill raker about 2/3 eye diameter; gill filaments at

angle of first gill arch very short, about 1/8 eye diameter and

about 1/3 length of longest filaments of pseudobranch.

Australasian giant Hoplosthethus

349

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350

C.D. Roberts & M.F. Gomon

Body covered with adherent scales, scales densely covered

with low, knob-like spinules above lateral line, posteriorly and

low on side, scales above and covered laterally by pectoral fin,

cycloid (cycloid scales distributed more ventrally in small

individuals); head naked except for patch of scales on cheek

posterior to rear tip of maxilla in about four vertical rows,

posteriormost row with 8-12 scales; isthmus scaleless; lateral

line with fine spinulation at free margin, each lateral-line

scale with strong ridge but lacking distinct spine (no spine in

small and medium sized adults, with strong spine centrally on

posterior margin in large adults), considerably larger than

other body scales, although adjacent scales intermediate in

size between lateral-line scales and others away from lateral

line; deep serrated abdominal keel formed from greatly

enlarged scales (scutes) with spine-like apices, most without

sculpturing or multiple tips, though with striations in some

especially posteriorly; scales on dorsal midline in front of

dorsal fin slightly but noticeably raised, their spinules not

greatly enlarged. First dorsal-fin spine short, subsequent

spines distinctly longer but only increasing slightly

progressively in length; spines of moderate thickness, those

posteriorly progressively wider with distinct lengthwise

striations; first soft ray longer than last spine, subsequent rays

progressively decreasing in length, outer margin of soft dorsal

fin slightly curved. First anal-fin spine short, second only

slightly or considerably longer, third long; spines of similar

robustness to those of dorsal fin. Caudal fin distinctly forked,

lobes of moderate breadth and rounded. Pectoral fin of

moderate length, reaching to or beyond last few scutes

(relatively longer in small individuals, to base of the third

anal-fin ray in 99.3 mm SL paratype). Pelvic fin reaching just

beyond middle scutes (to anal-fin origin in smallest paratype).

A large species, largest specimen examined 515 mm SL.

Colour in life. Head and body dark grey, obscured by deep red;

opercle black suffused with red; dark areas of buccal and

branchial cavities black; fins deep red with narrow black distal

edges (figs 3 & 4).

Pigmentation in alcohol. Freshly preserved individuals dark

grey (juveniles pale below a line between humeral spine and

dorsal side of caudal peduncle near termination of dorsal fin

base, dusky above); opercle dark; buccal and branchial

chambers dark, including gill arches and rakers; vomer, roof of

mouth lateral to palatines, underside of tongue and lower jaw

uniformly pale; fins pale with narrow dark margins (juveniles

with pale fins, except for slightly dusky outer portion of

membrane between dorsal-fin spines).

Etymology. The name melanopeza is from the Greek melano

for ‘black’ and peza ‘edge’, in reference to the characteristic

black edge on all fins in large individuals of this species. As a

noun in apposition, the spelling of melanopeza is not influenced

by masculine gender of genus Hoplostethus.

Distribution. Confined to sub-tropical and temperate latitudes

of the Tasman and South Fiji Basins in the south-western

Pacific, documented from localities between about 33° and 37°

S from south-eastern Australian slopes on the west to the outer

Bay of Plenty and southern Kermadec Ridge at the north end of

the North Island in New Zealand in the east (fig. 2). Occurs on

continental slopes, seamounts and submarine rises with

collection depths recorded between about 140 and 760 m, but

most often 250-400 m.

Comments. Hosplostethus melanopeza is very similar to H.

gigas, a species with which it was confused in the early 1980’s

when initial specimens were collected. The absence of detailed

descriptive information beyond McCulloch’s initial description

of H. gigas nearly 100 years ago no doubt contributed to this

confusion. McCulloch evidently received the specimens on

which his description is based well after they were preserved,

saying about the species’ original colouration only that “when

first received all the fins had traces of deep rose pink.” Although

both species have a similar overall red or reddish-orange

colouration, H. melanopeza has distinct black margins to all

fins in large individuals, while the edges of only the medial fins

appear to be no more than dark grey to blackish in large

individuals of H. gigas. The recognition of the two species is

supported by Cytochrome Oxidase subunit one sequences (Te

Papa/ NIWA/CSIRO/NMV unpublished data).

Morphologically H. gigas and H. melanopeza have nearly

identical ranges for meristic values, but the latter has a greater

number of predorsal scales (16-22 versus. 9-15) and a higher

mean value for total gill rakers of 19.7 versus 18.0, based on

material examined. Proportional measurements for the two

are also extremely similar. The greater size range of specimens

for the new species makes a full comparison of the two

impossible, although an examination of the distribution of

values in a number of other species shows morphometric

ranges converging at smaller sizes and the greatest disparities

evident as individuals approach their maximum size. At

comparable sizes H. melanopeza seems to have a slightly

deeper body and shorter pelvic to anal-fin length than H.

gigas (fig. 5).

Morphological comparisons with other nominal species

referred to the subgenus H. ( Hoplostethus ) are difficult because

of the relatively small sample sizes that literature descriptions

on which most are based and the considerably greater variation

detected when greater numbers are examined. In comparison

with H. robustispinus, which reaches a comparably large size,

H. melanopeza has 15, rarely 16, versus 17, rarely 16 pectoral-

fin rays, 9-12 versus 11-13 abdominal scutes, 26 versus 27

total vertebrae, and medial fin spines of a moderate breadth

versus broad in large individuals. The Indian Ocean H.

confinis, as mentioned in the Introduction, also reaches a

reasonably large size and has meristic values that are more

similar to those of H. melanopeza and H. gigas, apparently

differing from the former in having the same lower mean

value of total gill rakers as the latter. It is further separable

from H. melanopeza in lacking the black margins to its fins at

a large size. Two other species occurring in the Indo-Pacific

region have pectoral-fin ray numbers that appear to be

comparable with H. melanopeza, the Red Sea H. marisrubri

Kotlyar, 1986 and western Indian Ocean H. mikhailini Kotlyar,

1986. Neither has been reported to have dark margins to their

fins, but individuals of H. melanopeza do not appear to develop

dark margins until they reach a standard length well in excess

Australasian giant Hoplosthethus

351

of 150 mm SL, a size that is greater than the largest specimen

reported for either of these two species. Hoplostethus

melanopeza differs from H. marisrubri in having 26 versus 27

total vertebrae and from H. mikliailini in having 18-21 versus

23-27 total gill rakers.

Although H. melanopeza, together with most other species

of the genus reaching a significant size (apart from the

commercially important orange roughy Hoplostethus

atlanticus Collett, 1889), have long been regarded as very rare

in deepwater environments. The type series took over 30 years

to collect despite periodic requests to networks and the

commercial and recreational fishing sectors (e.g. Roberts,

1995, 1996, 2012). The use of fishing gear such as deep

droplines and set nets that are effective in capturing species

living in rough bottom environments, have shown them to

occur locally in greater numbers than originally thought. Even

though life history details are yet to be identified for this and

other giant roughy species, it is reasonable to believe their age

and growth rate approach species like the orange roughy for

which information is reasonably well known. Consequently,

we urge restraint in the development of targeted commercial

exploitation of what is potentially another long-lived, late

maturing species, for which we currently have inadequate

information on its abundance and biology.

Hoplostethus grandperrini sp. nov.

Grandperrin’s giant sawbelly

Figures 2, 5 & 6; Tables 1 & 2

Hoplostethus cf. g igas\ Grandperrin & Lehodey, 1992: 7, 26 and

35, listed, seamount “B”, Norfolk Ridge, New Caledonia.

Holotype. NMNZ P.027462 (455) New Caledonia, Norfolk Ridge,

Seamount “B”, 24°55.15'S, 168°20.95'E, 600-675 m, BERYX 2, stn 3,

N.O. Alis, chalut a poissons (= fish bottom trawl), 24 October 1991, R.

Grandperrin and C. Roberts. (Lodged in NMNZ collection at the

request of MNHN).

Paratype. MNHN 2012-0269 (131) New Caledonia, SE slope of

Grande Terre, 22°13.00' S, 167°14.00' E, 500-510 m, MUSORSTOM

4, stn 238, N.O. Vauban, chalut a perche (= beam trawl), 2 October

1985.

Diagnosis. Pectoral-fin rays 17 or 18; total gill rakers on outer

side of first arch 19 or 20; abdominal scutes 13 or 14, scutes

rectangular and rugose in large individuals; predorsal scales,

21-24; isthmus lacking scales; body scales adherent; lateral

line scales with strong medial ridge and spine posteriorly and

with numerous fine rather long spines in small individuals,

scales rugose, lacking a posterior spine in large adults; scales

on predorsal midline forming very low ridge in small

individuals, no ridge apparent in very large individuals; body

ovoid and deep, depth 1.8-1.9 in SL; dorsal profile of head

gently curved; dorsal- and anal-fin spines of moderate

thickness; pectoral fin of moderate length, reaching base of

second anal-fin spine in small individuals, to tenth or eleventh

abdominal scute in largest; adults orange-red; buccal cavity

mostly white, black only on roof of mouth posteriorly,

opercular recess black anteriorly, much paler near periphery.

Reaches 455 mm SL.

Description. (See Table 1 for frequencies of values for selected

meristic characters.) Dorsal-fin rays VI, 13; anal-fin rays III, 9;

caudal-fin rays 6 + 2 + 9 + 8 + 2 + 6; pectoral-fin rays 17 (18 in

1 of 4); pelvic-fin rays I, 6; gill rakers 6+1 +13 (6+1 + 12);

lateral-line scales 28 + 1; transverse scales 12/1/26 (11/1/35);

predorsal scales 21 (24); scutes 14 (13; Table 2); vertebrae 11 +

15; pseudobranch 15 (only holotype examined); branchiostegal

rays 8.

(See Table 1 for comparative ranges of selected morphometric

characters.) Body ovoid and deep, depth 1.8-1.9 in SL. Head

large, its height equal to or slightly greater than its length,

99.1- 103% HL; upper profile in front of dorsal fin gently curved

to upper jaw; anterodorsal profile moderately separated from

upper orbital rim; space between eyes wide, interorbital width

31.1- 34.0% HL; eye of moderate size, orbital diameter 23.2-

29.18% HL; crests of head bones strong, fine spinules on apices

at skin surface; depressions between crests moderately deep,

hidden by thick skin in adults; infraorbital bones becoming

progressively broader with growth; nostrils immediately

preceding orbits, posterior nostril two to four times area of

anterior nostril; mouth reaching just beyond vertical through

hind margin of eye; large, fine denticulate teeth covering oral

margins and exposed lateral surfaces of premaxilla and dentary,

palatine with narrow band of similar teeth, vomer apparently

lacking teeth (with three small teeth in 131 mm paratype); tip of

dentary with ossified knob at symphysis. Preopercular spine

short and rounded (slender and pointed, reaching to pelvic-fin

base in small individual). Humeral spines short and rounded

(short and pointed in smallest individual). Longest gill raker

about 2/3 eye diameter; gill filaments at angle of first gill arch

very short, about 1/10 eye diameter and just less than 1/2 length

of longest filaments of pseudobranch.

Body covered with adherent scales, those above lateral

line, posteriorly and low on side densely covered by rather

long fine spines, scales above and covered laterally by pectoral

fin, cycloid (cycloid scales distributed more ventrally in

smaller individual); head naked except for patch of scales on

cheek posterior to rear tip of maxilla in about three or four

vertical rows, posteriormost row with 13 or 14 scales; isthmus

scaleless; lateral-line scales rugose, without a strong spine

(smaller individual having each lateral-line scale with slender

ridge and small pointed spine posteriorly, peripheral row of

spinules noticeably longer than others); low serrated abdominal

keel formed from slightly rugose, enlarged scales (scutes) with

pointed apices (smaller individual with smooth scutes more

typical of other species), scales on midline preceding scutes

somewhat enlarged but not counted as scutes above; scales on

dorsal midline in front of dorsal fin not forming low ridge

(smaller specimen with posterior scales slightly raised

medially), their spinules not enlarged. First dorsal-fin spine

short, second and third spines progressively longer, last three

increasing in length only slightly; spines of moderate thickness,

those posteriorly progressively thicker with distinct lengthwise

striations; first soft ray longer than last spine, subsequent rays

progressively decreasing in length, outer margin of soft dorsal

fin slightly curved. First anal-fin spine short, second short or

of intermediate length, and third long; spines of similar

robustness to those in dorsal fin. Caudal fin distinctly forked,

352

C.D. Roberts & M.F. Gomon

Figure 6. Hoplostethus grandperrini sp. nov., NMNZ P.027462, 455 mm SL, holotype. A, fresh, photo by C. Roberts. B, preserved, photo by C.

Struthers.

lobes broad and rounded. Pectoral fin of moderate length

reaching tenth or eleventh abdominal scute (reaching base of

second anal-fin spine in smaller individual). Pelvic fin

reaching to middle scutes (to eighth scute in smaller individual).

A large species, largest specimen examined 455 mm SL.

Colour in life. Head orange-red; body orange-tan dorsally,

extending ventrally to pelvic-fin base anteriorly and to anal-fin

base posteriorly; opercle black suffused with red; dark areas of

buccal and branchial cavities black, pale areas stark white; fins

orange to orange-tan, without darker margins (fig. 6).

Pigmentation in alcohol. Freshly preserved individuals grey-tan;

opercle dark; buccal chamber pale, roof dark posteriorly; fins

pale, outer edge of membranes between dorsal-fin spines dusky.

Etymology. The specific name grandperrini recognises Dr Rene

Grandperrin, retired chief scientist of ORSTOM Noumea, ardent

fish researcher and leader of deepwater fish explorations off New

Caledonia (Chef de Mission of research cruises BERYX 2 and

HALIPR02), in recognition of his strong supportfor collaborative

fieldwork between French and New Zealand scientists.

Distribution. Known only from the types collected at Seamount

“B” in the northern portion of the Norfolk Ridge, south of New

Caledonia and on the south-eastern slope of New Caledonia’s

Grande Terre (fig. 2), from depths of 500-675 m.

Comments. The great similarity of many Hoplostethus species

to one another has led to confusion about the identity of this

species with two other apparently undescribed species living on

Australasian giant Hoplosthethus

353

seamounts in waters off southern New Caledonia. At least one of

the latter reaches a considerable size, the largest known specimen

measuring 355 mm SL, and has a distribution that includes

Australian territorial waters. The three are consistently separable

by a combination of characters, including numbers of pectoral-

fin rays and abdominal scutes, and the pigmentation of the buccal

cavity. The identities of the other two species will be the subject

of a forthcoming publication by Gomon and Roberts (ms).

With regard to the three New Caledonian species we

recognise, G. grandperrini has 17 or 18 pectoral-fin rays (versus

16 or 17, rarely 15 or 18, and 15, rarely 16 in the other two), 13 or

14 abdominal scutes (versus 10-15 and 11-13 scutes) and has a

buccal cavity that is mostly pale, any black pigmentation confined

to the rear of the mouth in the vicinity of the throat (versus buccal

cavity entirely black behind the oral valves, including the

underside of the tongue and top of the lower jaw, and buccal

cavity black with vomer, underside of the tongue and the top of

the lower jaw white). These same character combinations, plus

high count of predorsal scales (21-24 versus 9-15, 16-22),

separate H. grandperrini from the other two giant species treated

above. Based on the limited series of types, H. grandperrini also

appears to have a greater body depth, postorbital length,

interorbital width, and pelvic fin to anal fin distance (fig. 5).

Of the remaining nominal species referred to the subgenus

H. (Hoplostethus ) apparently only two regularly have 17 or 18

pectoral-fin rays, H. druzliinini Kotlyar, 1986 in the Arabian Sea

and H. vniro in the southeastern Atlantic. Both of these species

differ from H. grandperrini in having on average more numerous

gill rakers, 21-25, versus 19 or 20. In addition, H. duzhinini has

only 8-12 abdominal scutes (versus 13-14) and a mostly black

buccal cavity (versus mostly pale).

Acknowledgements

We thank all observers, skippers, crews and fishing companies

together with scientists, skippers and crews of research vessels

who collected the rare giant sawbelly specimens used in this

study. Specimens examined and collection data were provided

with the assistance of: M. McGrouther (AMS), A. Graham

(CSIRO), A. Stewart and J. Baker (NMNZ), D. Bray (NMV),

J. Rivaton (ORSTOM), J. Johnson (QMB), G. Moore (WAM),

J. Williams (USNM), and R Pruvost and R. Causse (MNHN).

Radiographs were prepared by S. Reader (AMS), J. Baker and

R. McPhee (NMNZ). Images used in the figures were prepared

and edited by C. Struthers (NMNZ). Helpful comments on the

manuscript were received from A. Stewart and C. Struthers.

This study was funded in part by the New Zealand Ministry of

Science and Innovation through Te Papa subcontract within

NIWA’s Coasts and Oceans Centre. Participation by the senior

author in research voyage BERYX 2 on N.O. Alis in 2001

during which the holotype of H. grandperrini was collected

was assisted by ORSTOM (now IRD) Noumea.

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Ichthyology. Proceedings of the Linnean Society of New South

Wales 89 (1): 11-127.

Woods, L.P. and Sonoda, P.M. 1973. Fishes of the western North

Atlantic. Order Berycomorphi (Beryciformes). Memoirs of the

Sears Foundation of Marine Research, 1 (6): 263-396, 66 figs, 10

tabs.

Memoirs of Museum Victoria 69:355-363 (2012)

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

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

Some hydroids (Hydrozoa: Hydroidolina) from Dampier, Western Australia:

annotated list with description of two new species.

Jeanette E. Watson

Honorary Research Associate, Marine Biology, Museum Victoria, PO Box 666, Melbourne, Victoria Australia 3001.

(hydroid@bigpond.com)

Abstract Jeanette E. Watson, 2012. Some hydroids (Hydrozoa: Hydroidolina) from Dampier, Western Australia: annotated list with

description of two new species. Memoirs of Museum Victoria 69: 355-363.

Eleven species of hydroids including two new ( Halecium corpulatum and Plumularia fragilia ) from a depth of 50

m, 50 km north of Dampier, Western Australia are reported. The tropical hydroid fauna of Western Australia is poorly

known; species recorded here show strong affinity with the Indonesian and Indo-Pacific region.

Keywords Hydroids, tropical species, Dampier, Western Australia

Introduction

A collection of hydroids provided by the Western Australian

Museum is described. The collection comprises 11 species

including two new. Material was collected 50 km north of

Dampier, Western Australia, from the gas production platform

Ocean Legend (019° 42' 18.04" S, 118° 42' 26.44" E). The

collection was made from a depth of 50 m by commercial

divers on 4 th August, 2011.

The species in the collection show a strong affinity with

the northern Indian Ocean, Indonesian and Indo-Pacific

regions. The tropical hydroid fauna of the western Australian

coast from Geraldton to Darwin is poorly known. Stechow

(1925) reported on some hydroids collected in Shark Bay and

Watson (1997, 2000) described collections from the Abrolhos

Islands and Darwin. Other reports relevant to the present study

are those from the Indonesian region by Billard (1913),

Vervoort (1941) and Schuchert (2003) and from the Aru Sea

(Stechow and Muller 1923). Other than those collected by

scuba diving from the Abrolhos and Darwin, all other material

is from dredging hence many new species are still being found

in habitats only collectable by hand sampling.

Type and voucher material is deposited in the Western

Australian Museum (WAM Z). In the following text only

synonymies relevant to the Indonesian, Australian and Indo-

Pacific and regions are given.

Family Eudendriidae L. Agassiz, 1862

Eudendrium racemosum (Cavolini, 1785)

Figure 1A

Sertolaria racemosa Cavolini, 1785: 160, pi. 6, figs 1-7, 14-15

Sertularia racemosa. - Gmelin, 1791: 3854

Eudendrium racemosum- Ehrenberg, 1834: 296- von

Lendenfeld, 1885: 351,353.-Millard and Bouillon, 1973: 33.-Watson,

1985: 204, figs 63-67

Material examined. WAM Z31857, material ethanol preserved. Four

infertile colonies, the tallest 40 mm long, on purple sponge.

Description. Hydrorhiza a tangled mass of stolonal tubes.

Stems fascicled, stolons becoming stems in a loose untidy mass

of tubes with much adventitious matter embedded between the

tubes. Largest colony comprised of several heavily fascicled

main branches, branching and rebranching roughly alternate,

ultimate branches monosiphonic. Hydranth pedicels more or

less alternate on ultimate branches, cylindrical, smooth, length

variable, with two to four obscure annulations at base. Hydranth

large with approximately 20 stubby tentacles.

Cnidome: all nematocysts undischarged,

- small euryteles, abundant in tentacles,

- large isorhizas, rare on hydranth body.

Measurements of Eudendrium racemosum, (/<m) preserved

material

Branch, distance between pedicels

1000-1800

Hydranth pedicel

length

760-1700

diameter

136-184

Hydranth, width below tentacles

320-448

356

J.E. Watson

Figure 1A-G. A. Eudendrium racemosum from photograph of whole colony. B. Filellum serratum, regenerated hydrotheca. C, D. Lafoeina

amirantensis. C, small hydrotheca and nematotheca on hydrorhiza. D, large hydrotheca. E-G. Halecium Itenellum. E, unbranched stem.

F, basally annulated hydrophore. G, hydrotheca with strongly everted rim and desmocytes. Scale bar, mm: A, 20. B, C, E, F, G, 0.2. D. 0.1.

Remarks. No supplementary cnidophores are present on the

hydranths (see Watson 1985). The colonies of E. racemosum are

substrate for most of the smaller hydroid species in the

collection.

Distribution. Mediterranean Sea, Seychelles, Vietnam, South

China Sea, Japan; previously recorded in Australia from the

Great Barrier Reef (Pennycuik 1959).

Family Lafoeidae A. Agassiz, 1865

Filellum serratum (Clarke, 1879)

Figure IB

Lafoea serrata Clarke, 1879: 242.

Reticularia serrata- Ralph, 1958: 312.

Filellum serratum - Millard, 1975: 178 - Gravier-Bonnet, 1979:

22 - Hirohito, 1995: 110.-Watson, 2000: 5, fig. 2C.

Material examined. WAM Z31858, microslide, malinol mount. One

small infertile colony creeping on stem of Eudendrium racemosum.

Description. Hydrothecae long, tubiform, base adnate to

substrate, distal two thirds of body erect, diameter narrower on

adnate section, abcauline surface closely transversely striated

above adnate adcauline wall. Margin circular, rim slightly

everted, some with some marginal replications.

Measurements of Filellum serratum , (/tm)

Hydrotheca

length of free part

240-640

diameter of free part

84-92

diameter at margin

104-116

Remarks. Although the material is in poor condition the

abcauline striations are a good indicator of identity as Filellum

serratum. The dimensions conform with F. serratum from

Darwin (Watson 2000).

Some hydroids (Hydrozoa: Hydroidolina) from Dampier, Western Australia: annotated list with description of two new

species.

357

Distribution. Cosmopolitan, previously recorded from Darwin

and temperate Western Australia.

Lafoeina amirantensis Millard and Bouillon, (1973)

Figure 1C, D

Egmundella amirantensis Millard and Bouillon, 1973: 40-

Millard, 1975: 133- Gibbons and Ryland, 1989: 389 - Ramil and

Vervoort, 1992: 22.

Lafoeina amirantensis .- Calder, 1991: 10 - Watson, 1994: 147-

Calder and Vervoort, 1998: 15 - Watson, 2000: 5, fig. 2A, B.

Material examined. WAM Z31850. Infertile colonies on Eudendrium

racemosum\ one microslide, malinol mount.

Description. Colonies stolonal, creeping on branches of host,

stolons flattened, roughened and coated with fine sediment.

Hydrothecae minute, arising at intervals along hydrorhiza,

subconical to cylindrical, very variable in size, sometimes

asymmetrically curved, base expanding from a short, wide

pedicel, operculum of numerous segments overlapping at apex,

no demarcation between opercular segments and body of

hydrotheca. Nematothecae sparse, on hydrorhiza between

hydrothecae, minute, clavate.

Measurements of Lafoeina amirantensis, (/tm)

Hydrorhiza, width

28-48

Hydrotheca

length including operculum

152-200

maximum width

52-80

Nematotheca, length

24-40

Remarks. The specimen conforms to previous descriptions of

Lafoeina amirantensis.

Distribution. Cosmopolitan, previously recorded in Australia

from Bass Strait and Darwin (Watson 1994a, 2000).

Family Haleciidae Hincks, 1868

Halecium tenellum Hincks, (1861)

Figure IE- G

Halecium tenellum Hincks, 1861a: 252, pi. 6, figs 1-4 - Vervoort,

1959: 229, fig. 8.-Vervoort, 1966: 102, fig. 2.-Millard, 1975: 156, fig.

50F-L - Vervoort and Watson, 2003: 98, fig. 19A, B.

Material examined. WAM Z31849. Abundant infertile colonies on

Eudendrium racemosum\ one microslide, malinol mount.

Description. Hydrorhiza creeping on host, stolons tubular,

undulating. Hydrocaulus monosiphonic, cylindrical, variable in

length, mostly unbranched but some branched once or twice.

Hydrophore smooth, cylindrical, a few transverse proximal

septa marking site of branching from below or within a

hydrotheca; hydrophore expanding slightly to below hydrotheca.

Hydrotheca shallow dish-shaped, expanding to margin, rim

strongly outrolled, diaphragm transverse, a row of desmocytes

above, no marginal replications. Perisarc smooth, thin.

Measurements of Halecium Itenellum, (pm)

Hydrocaulus, basal length

400-900

Hydrophore

length below hydrotheca

80-520

width

48-60

Hydrotheca

width at diaphragm

60-80

depth to diaphragm

20-22

width across margin

92-124

Remarks. Identification of Halecium to species level in closely

similar species groups is difficult with infertile material. The

present material is doubtfully referred to H. tenellum.

Distribution. Atlantic, Indian and Pacific Oceans, probably

New Zealand. Not previously recorded from Australia.

Halecium corpulatum sp. nov.

Figure 2A-F

Material examined. Holotype WAM Z31865, one microslide, malinol

mount and remaining preserved material from holotype colony. Three

sparsely fertile stems, probably fragmented colony.

Description. Hydrorhiza comprising tangled stolons becoming

fascicular tubes of erect stem. Stems fascicled, thick, arborescent,

almost planar, branching irregular from opposite to alternate,

polysiphonic tubes reaching to base of ultimate branches.

Hydrophores arising from just below a hydrotheca, usually

one but sometimes two opposite, one to six successive

secondary hydrophores arising linearly from diaphragm of

preceding hydrotheca. Hydrophores moderately long,

cylindrical, increasing slightly in diameter to below hydrotheca,

one to three proximal abcauline partial septa fading out on

adcauline wall. One or two thickenings of perisarc at the base

of a hydrotheca above junction with hydrophore.

Hydrotheca shallow dish-shaped, adcauline wall of

primary hydrotheca closely adpressed to primary hydrophore,

succeeding hydrothecae expanding smoothly from a transverse

diaphragm to margin, a row of desmocytes just above

diaphragm, sometimes a concave pseudo-diaphragm below.

Margin everted and strongly outrolled, not replicated, ratio of

depth of hydrotheca to diameter of margin 1:4—1:5.

Hydranth (preserved material) with a long conical or

cylindrical peduncle with about 20 moderately long tentacles.

Gonotheca inserted on a very short pedicel within a

hydrotheca, lenticular, wall smooth, perisarc thin, apex

slightly pointed, no evidence of aperture. Gonophore probably

male.

358

J.E. Watson

Figure 2A-F. Halecium corpulatum sp. nov. A, holotype colony. B, linear series of hydrophores. C, branching of stem. D, dichotomous branching

of hydrophores from below a hydrotheca showing basal thickening below hydrotheca. E, male gonotheca. F, hyranth (preserved material). Scale

bar, mm: A, 25. B, D, 0.2.C, E, F, 0.5.

Measurements of Halecium corpulatum , (/<m)

Hydrophore, primary

length

800-940

width

152-184

Hydrotheca

width at diaphragm

140-180

diameter of margin

216-248

depth, diaphragm to margin

40-56

Gonotheca

height

860

width

660

Remarks. The colony may have originally been much taller as

it is broken off at the top. In many respects including the

thickening at the base of many primary hydrophores Halecium

corpulatum resembles Hydrodendron sibogae (Billard, 1929)

but the absence of nematothecae places it in Halecium. The

few gonothecae available for study do not resemble those of

any large arborescent species such as Halecium muricatum

(Ellis & Solander, 1786), H. dichotomum Allman, 1888 and

H. lankesteri (Bourne, 1890). The gonothecae however,

resemble Gibbons and Ryland’s (1989) description of

Halecium sibogae from Fiji; their specimen may well have

been H. corpulatum.

While the unusually long skirt-like shape of the hydranth

is probably an artefact of preservation the living hydranth

nevertheless must have been extraordinarily large.

Etymology. Describes the large, corpulent hydranth.

Some hydroids (Hydrozoa: Hydroidolina) from Dampier, Western Australia: annotated list with description of two new

species.

359

Family Hebellidae Fraser, 1812

Hebella costata (Bale, 1884)

Figure 3A

Campanularia costata Bale, 1884: 56 - Stechow and Muller,

1923: 463.

Scandia corrugata Millard and Bouillon, 1973: 60.

Hebella muscensis Millard and Bouillon, 1975: 10.- Boero et al.,

1997: 22.

Hebellopsis costata- Watson, 2000: 6, fig. 3A.

Hebella costata- Watson and Vervoort, 2003: 64.

Material examined. WAM Z31866. Infertile colony on Eudendrium

racemosum\ microslide, malinol mounted.

Description. Hydrorhiza creeping on stem and branches of

host. Hydrothecal pedicels variable in length, almost smooth

and straight to gnarled, widening to base of hydrotheca.

Hydrotheca tubular, straight to slightly bent, body with five to

seven undulations becoming more pronounced distally, no

diaphragm visible. Margin circular, transverse or slightly

oblique to hydrothecal axis, rim weakly everted, not replicated.

Measurements of Hebella costata , (/<m)

Pedicel

length

176-440

maximum width

96-128

Hydrotheca

length

800-1080

diameter at margin

460-600

Remarks. The material falls within the dimensional range of

Hebella costata reported from Darwin (Watson 2000).

Distribution. Indian Ocean, Indonesia, tropical Australia.

Family Sertulariidae Lamouroux, 1812

Diphasia digitalis Busk, (1852)

Figure 3B, C

Sertularia digitalis Busk, 1852: 387, 393.

Diphasia digitalis (Busk).- von Lendenfeld, 1885a: 415, 633-

Bale, 1884: 101.-Bale, 1915: 265.-Jaderholm, 1920: 4.-Billard, 1931:

249.- Vervoort, 1972: 99.- Pennycuik, 1959: 191.- Millard, 1975:

257.- Watson, 1996: 78.- Watson 2000: 14, fig. 10A, B.- Schuchert,

2003: 166, fig. 25.

Material examined. WAM Z31867. Two broken infertile stems, the

longer 22 mm; microslide, malinol mount.

Description. Hydrorhiza creeping, stolons tubular. Stems

sparingly branched, proximal athecate stem segment long,

tubular, with a strong distal hinge joint. Hydrothecae paired,

one pair per internode, nodes transverse, indistinct to absent,

marked only by a narrowing of internode. Hydrotheca long,

tubular, expanding from base to margin, free adcauline wall

convex, abcauline wall concave. Margin quadrangular, with a

low abcauline cusp and an indistinct longitudinal pleat

extending downwards from margin, fading out near base of

hydrotheca. Remains of operculae visible inside many

hydrothecae. Perisarc smooth.

Measurements of Diphasia digitalis , (pm)

Internode

distance between hydrothecae

1020-1300

width across hydrothecal base pair

256-384

Hydrotheca

adcauline wall length adnate

800-840

adcauline wall length free

240-300

width across margin

264-296

width at floor

92-100

Remarks. The hydrothecae contain much adventitious matter

presumably acquired during collection. The margins of most

hydrothecae are damaged.

The specimens conform to previous descriptions of

Diphasia digitalis from Australia (Watson 2000). The

longitudinal hydrothecal pleat is inconspicuous and may be

due to the young age of the colony.

Distribution. Circumglobal in tropical and subtropical waters.

Australian distribution - Torres Strait (Busk 1852), Queensland

(Pennycuik 1959), north-western Australia (Watson 1996,2000).

Family Halopterididae Millard, (1962)

Halopteris glutinosa Lamouroux, (1816)

Figure 3D

Aglaophenia Glutinosa Lamouroux, 1816:171.

Plumularia glutinosa- Billard, 1909: 327-Billard, 1910: 36, fig. 16.

Plumularia buskii Bale, 1884: 125, pi. 10, fig. 3, pi. 19, figs 34-35-

Bale, 1887: 22.

Plumularia buski- Billard, 1913: 21, fig. 11.

Halopteris buski - Watson, 1973: 184 - Schuchert, 1997: 58, figs

18, 19 - Vervoort and Watson, 2003: 353.

Halopteris glutinosa- Watson, 2005: 537, fig. 37A, B.

Material examined. WAM Z31868. One stem fragment 5 mm long,

detached from substrate; microslide, malinol mounted.

Remarks. Billard (1913) recorded H. glutinosa (as Plumularia

buski) from nine sites on sand, shell sand and Lithothamnion

from depths of 13-522 m in Indonesia. A specimen from Japan

(author’s collection) is also H. glutinosa.

Distribution. Southern and south-western Australia, New

Zealand, Indonesia and Japan.

360

J.E. Watson

Figure 3A-D. A. Hebella costata. Hydrotheca with proximal corrugations. B, C. Diphasia digitalis. B, branched stem. C, stem internode with

opposite hydrothecae. D. Halopteris glutinosa. Hydrocladial athecate and thecate internodes. Scale bar, mm: A, 0.5. B, 1.0. C, D, 0.25.

Family Plumulariidae McCrady, (1859)

Plumularia fragilia sp. nov.

Figure 4A-D

Material examined. Holotype, WAM Z31846, One infertile stem on

Lytocarpia delicatula (Busk, 1852); microslide, malinol mounted.

Description. Hydrorhiza a tubular creeping stolon. Stems

pinnate, slender, to 13 mm long, monosiphonic, proximal

hydrocauline segment long, straight, athecate, following

internodes long, cylindrical, nodes transverse, distinct, a slight

tumescence below each node.

Hydrocladia distal on cauline internode, alternate, position

not quite planar. Apophysis smooth, directed upward at an

angle of about 45° to hydrocaulus, abcauline apophysial wall

contiguous with cauline internode, adcauline wall concave

with three deep septa, distal node transverse.

Hydrocladia with two or three hydrothecae, cylindrical,

narrow, beginning with a proximal athecate internode with

transverse proximal and oblique distal node, usually with

indistinct internal supplementary proximal and distal septa.

First athecate internode followed by alternate hydrothecate

and athecate internodes; athecate internodes variable in

length, sometimes much longer than hydrothecate internode;

hydrothecate internode long, straight with almost transverse

distal node; nodes often with one or two internal septa.

Hydrotheca seated about halfway along internode, small,

deep bowl-shaped, abcauline wall weakly concave to straight,

adcauline wall weakly convex, completely adnate to internode,

floor of hydrotheca transverse to hydrocladial axis, margin

circular, oblique to hydrocladial axis, rim everted.

Nematothecae all of same shape and similar in size,

bithalamic, moveable, slender conical, base long, cup wider

than deep, margin circular, not excavated. One median

nematotheca about halfway along athecate internode, one

median inferior on hydrothecate internode well below base of

hydrotheca, bases of twin laterals inserted just below

hydrothecal margin; one or two nematothecae on cauline

internode, standing out from internode, one about one third

distance up internode from apophysis, the other two thirds

distance up internode and on same side as hydrocladium, two

nematothecae in axil, a bun-shaped hydrostatic pore at base of

axillar nematothecae.

Colony colourless (preserved material), perisarc thin.

Some hydroids (Hydrozoa: Hydroidolina) from Dampier, Western Australia: annotated list with description of two new

species.

361

_ F

Figure 4A-E. A. Plumularia fragilia sp. nov., holotype colony. B, stem internodes and alternate hydrocladia. C, proximal part of hydrocladium

and apophysis with axial nematothecae. D, hydrocladium with athecate and thecate internodes. E. Plumularia bedoti. Distal part of hydrocladium

with developing anastomose. F. Lytocarpia delicatula. Two hydrocladial hydrothecae. Scale bar, mm: A 10. B, 1.0. C-F, 0.2

Measurements of Plumularia fragilia, (/<m)

Hydrorhiza, width

40-56

Stem

internode length

264-320

width at node

56-64

Hydrocladium

apophysis, adcauline length

36-48

first athecate internode length

84-120

succeeding athecate internodes, length

220-260

thecate internode, length

248-284

width at node

24-36

Hydrotheca

abcauline wall, length

40-46

width at margin

72-80

Twin lateral nematothecae

length of base

42-52

depth of cup

10-14

width of cup

Remarks. Plumularia fragilia is a very small species. It closely

resembles but is somewhat smaller than Plumularia setacea

recorded from Darwin by Watson (2000). Stems are flaccid out

of fluid. Cauline nematothecae may be absent leaving no scars

on the internode

362

J.E. Watson

Descriptions of Plumularia mossambicae Millard, 1975,

P. Antonbruuni Millard, 1967, P. strobilophora Billard 1913

and P. orientalis Billard, 1911 were compared with Plumularia

fragilia. P. mossambicae is larger in dimensions, lacks

cauline nematothecae, the abcauline hydrothecal wall is

straight, and the margin is not everted. Millard’s figure of P.

antonbruni shows differences in structure and is also larger

in all dimensions than P.fragilia. P. strobilophora is closer in

overall dimensions, but the hydrothecae are more bowl¬

shaped and have no marginal eversion. While P. orientalis

also has an outrolled margin, its dimensions are larger than P.

fragilia.

Etymology. The species is named to describe the fragility of

the colony.

Plumularia bedoti Billard, (1911)

Figure 4E

Plumularia bedoti Billard, 1911: 64- Billard 1913: 27-Watson,

2000: 54, fig. 42A-D.

Material examined. WAM Z31847. Infertile stem fragment 13 mm

long. Microslide, malinol mount.

Remarks. The material conforms with the description of

Plumularia bedoti from Darwin (Watson 2000). The stem is

lightly fascicled and there are distal anastomoses on the

hydrocladia.

Distribution. Indonesia, tropical Australia (Darwin).

Family Aglaopheniidae Marktanner-Turneretscher, (1890)

Lytocarpia delicatula Busk, (1852)

Figure 4F

Plumularia delicatula Busk, 1852: 396.

Aglaophenia delicatula - Bale, 1884: 167- Kirkpatrick, 1890:

604.-Billard, 1913: 106.-Pennycuik, 1959: 185.-Watson, 2000: 57,

fig. 46A-E.

Thecocarpus delicatulus Millard and Bouillon, 1973: 94.

Lytocarpia delicatula - Schuchert, 2003: 235, fig.76.

Material examined. WAM Z31848. Several pinnate stems to 35 mm

long, one stem fertile; microslide, malinol mounted.

Remarks. The material conforms with the description of

Lytocarpia delicatula from Darwin by Watson (2000) and

from Indonesia by Schuchert (2003). Male and female

gonophores are borne on the same corbula.

Distribution. Maldives, Mosambique, Indonesia, tropical

northern and eastern Australia.

Acknowledgements

I thank the Western Australian Museum for providing the

collection for examination.

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