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Contents

151 > Deep-water hydroids (Hydrozoa: Leptolida) from Macquarie Island

Jeanette E. Watson

181 > A molecular and morphological review of the asterinid, Patiriella gunnii (Gray) (Echinodermata: Asteroidea)

R Mark O’Loughlin, Jonathan M. Waters and Michael S. Roy

197 > A new genus of Tasmanian millipedes (Diplopoda: Polydesmida: Dalodesmidae) with unusual spiracles and a mosaic distribution

Robert Mesibov

207 > The millipede genus Gasterogramma (Giplopoda: Polydesmida: Dalodesmidae) in Tasmania, Australia, with descriptions of seven

new species

Robert Mesibov

221 > New species of Goreopagurus (Decapoda: Anomura: Paguridae) from Tasmania and reevaluation of sexual tubes in hermit crab sys

tematics

Rafael Lemaitre and Patsy A. McLaughlin

229 > A new genus and species of hermit crab (Decapoda: Anomura: Paguridae) from seamounts off south-eastern Tasmania, Australia

Patsy A. McLaughlin

237 > Gnathophyllum taylori, a new species of caridean shrimp from south-eastern Australia (Crustacea: Decapoda: Gnathophyllidae)

Shane T. Ahyong

243 > Rediagnosis of Palaemon and differentiation of south-eastern Australian species (Crustacea: Decapoda: Palaemonidae)

Terry Walker and Gary C. B. Poore

257 > Linguimaera Pirlot, 1936 (Crustacea: Amphipoda: Melitidae), a valid genus

TraudI Krapp-Schickel

285 > Four new species of Ischnomesidae (Crustacea: Isopoda: Asellota) from off south-eastern Australia

Kelly L. Merrin and Gary C. B. Poore

309 > New genera and species of sphaeromatid isopod crustaceans from Australian marine coastal waters

Niel L Bruce

371 > Neastacilla Tattersall, 1921 redefined, with eight new species from Australia (Crustacea: Isopoda: Arcturidae)

Rachael King

Memoirs of Museum Victoria 60(2): 151-180 (2003)

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

http://www.museum.vic.gov.au/memoirs

Deep-water hydroids (Hydrozoa: Leptolida) from Macquarie Island

Jeanette W. Watson

Honorary Associate, Marine Biology Section, Museum Victoria, GPO Box 666E, Melbourne 3001, Victoria, Australia

Abstract Watson, J.E. 2003. Deep-water hydroids (Hydrozoa: Leptolida) from Macquarie Island. Memoirs of Museum Victoria

60(2); 151-180.

A cruise conducted by RV Southern Surveyor between latitudes 53°0"S and 56°17"S and longitudes 158°30"E and

159°25"E sampled 1 1 sites by dredging at depths ranging from 364 to 1422 m off subantarctic Macquarie Island. Twenty-

seven species of hydroids including six new, were identified from the collection. There was no discernible distri-

butional pattern of species with depth. The most diverse hydroid assemblage was recovered from 500-600 m depth and

the second richest at 1064 m.

Keywords Hydrozoa, Leptolida, Macquarie Island, hydroids, taxonomy

Introduction

In January 1999 the Australian Commonwealth Science and

Industry Research Organisation (CSIRO) undertook a survey of

subantarctic areas to the north, south and west of subantarctic

Macquarie Island. Cruise S SO 1/99 by RV Southern Surveyor

was conducted over a period of three days in the region

53°0^S-56°17'S, 158°30^E-159°25T. While primarily

designed to assess commercial fish stocks the cruise also

included benthic sampling to provide a scientific basis for

development of a Commonwealth of Australia Marine

Protected Area around Macquarie Island (Butler et ah, 2000).

Sampling of the epifauna was undertaken by benthic sled at 11

sites at depths ranging from 364 to 1422 m (Table 1). Video

imaging of the bottom revealed a barren rocky or debris-strewn

bed with a sparse epifauna (Butler et al., 2000).

Twenty-seven species of hydroids including six new, and

three probably new (but due to insufficient or poorly preserved

material unidentifiable) were recorded (Table 2). Calamphora

quadrispinosa sp. nov. and Tulpa diverticula Totton, 1930 were

recorded from five stations; Lafoea tenellula Allman, 1877 and

Acryptolaria patagonica El Beshbeeshy, 1991 at four stations;

Eudendrium ?cyathiferum Jaderholm, 1904, Tripoma arboreum

Hirohito, 1995 and Zygophylax sagamiensis Hirohito, 1983 at

three stations. The remaining species were recorded at only one

or two stations. No discernible distributional pattern of species

occurred with depth, the richest fauna (14 species) coming

from stn 44 at a depth of 500-600 m, the second richest haul

being from stn 120 at a depth of 1064 m.

The most abundant free-growing species were large

colonies of Eudendrium ?cyathiferum recorded from depths of

Table 1. CSIRO cmise SSOl/99 stations (All material collected by benthic dredge)

Stn

Date

Latitude/Longitude, Start-Einish

Depth (m)

19 Jan 1999

56°15.7'S, 158°30.2T-56°18'S, 158°28.7T

500-600

19 Jan 1999

56H5.7'S, 158‘’30.2T-56‘’18.7'S, 158‘’25.099'E

600-1300

22 Jan 1999

54°40.8'S, 158°38.999'E-54°41.5'S, 158°42.2T

444

26 Jan 1999

54°31.6° S, 158°59.3° E-54°35.5° S, 159°0.8° E

818

26 Jan 1999

53°55.8'S, 159°5.5^E-53°55.7'S, 159°4.7^E

453°

26 Jan 1999

53°54.7'S, 159°1.90LE -no finish data

1024

26 Jan 1999

53°55.9'S, 158°5.9T-53°54.9^S, 159°2.2T

364

119

30 Jan 1999

53°38.rS, 159°9 599T -53°36.4'S, 159°8.2T

1046

120

30 Jan 1999

53°38'S, 159°9.5T-53°36.8^S, 159H0.799T

1064

122

30 Jan 1999

53‘’37.2'S, 159°11.299T-53°34.4'S, 159°17.40LE

1158

130

31 Jan 1999

52 59.4 'S, 159°59'E-53°2'S, 159°58.2'E

1422

152

Jeanette E. Watson

1046-1158 m, Tripoma arboreum at 444-1158 m, Acryptolaria

patagonica at 453-1422 m and an arborescent species of

Halecium. Smaller species epizoic on these and other hydroids

included Filellum conopeum sp. nov. growing on haleciids and

acryptolariids; Tulpa diverticula abundant on a variety of

hydroid hosts, Calamphora quadrispinosa sp. nov. abundant on

Symplectoscyphus paulensis and Eudendrium, and Lafoea

tenellula on Tulpa diverticula and Eudendrium. The diverse

fauna from stn 44 consisted of a tangled mass of hydroids

growing on the stem of a dead primnoid gorgonian.

Type and voucher material is lodged in the Tasmanian

Museum, Hobart (TM) and Museum Victoria, Melbourne

(NMV).

Table 2. Species and their occurrence at stations

Species

SSOl/99 Stn No.

Hydractinia sp.

Eudendrium ?cyathiferum Jaderholm, 1904

119, 120, 122

Eudendrium deforme Hartlaub 1905

44, 46

Eudendrium macquariensis sp. nov.

Eudendrium sp.

Tripoma arboreum Hirohito, 1995

63, 97, 122

Lafoea tenellula AWman, 1877

44, 90 119, 120

Lafoea dumosa (Fleming, 1828)

Lafoea annulata sp. nov.

Eilellum conopeum sp. nov.

94, 119, 130

Zygophylax sagamiensis Hirohito, 1983

Acryptolaria conferta (Allman, 1877)

Acryptolaria patagonica El Beshbeeshy 1991

44, 90, 92, 130

Acryptolaria rninuta sp. nov.

133

Halecium ralphae Watson and Vervoort, 2000

119

Halecium tenellum Hincks, 1861

120

Halecium ovatum Totton, 1930

120

Halecium delicatulum Coughtrey, 1876

119, 120, 122

Halecium sp.

90, 122

Calamphora quadrispinosa sp. nov.

44, 46, 94, 120, 130

Staurotheca affinis (Jaderholm, 1904)

44, 122

Symplectoscyphus paulensis Stechow, 1923

94, 120

Symplectoscyphus tuba Totton, 1930

94, 120, 122

Thyroscyphoides sympodialis sp. nov.

Gymnangium japonicum Watson and Vervoort,

122

2001

Tulpa diverticula Totton, 1930

44, 90, 94, 119, 120

Campanularia hicksoni Totton, 1930

Anthoathecata

Hydractiniidae L. Agassiz, 1862

Hydractinia van Beneden, 1841

Hydractinia sp.

Figures lA-C

Material examined. Stn 44, TM K2768, a small infertile colony on axis

of dead primnoid gorgonian, spreading onto Eudendrium stem,

specimen alcohol preserved.

Description. Hydrorhiza consisting of anastomosing stolons

embedded in a fine trabeculate meshwork of perisarc to 1 .4 mm

thick; meshwork penetrated and in some places externally

Figures lA-C. Hydractinia sp., stn 44. A, part of colony. B, trabecul-

ate meshwork of basal perisarc. C, nematocysts, possibly euryteles,

undischarged. Scale bar; A, 1 mm; B, 0.1 nun; C, 10 pm.

covered by coenosarc. Mature gastrozooids columnar, stout,

emerging from perisarcal meshwork; hypostome dome-shaped,

surrounded by c. 12 stubby, slightly tapering tentacles (pre-

served material); tentacles often alternately long and short.

Tentaculozooids styloid, tapering, very extensible, arising from

a broad base in perisarc meshwork. Bases of gastrozooids and

tentaculozooids may be surrounded by a collar of coenosarc or

erupt from a shallow pit in perisarc. Many short zooids bearing

4 or 5 tentacles just emergent above basal meshwork. No solid

spines present but instead, hollow mounds of perisarc common,

circular in cross-section and invested by the perisarcal mesh-

work.

Nematocysts of two sizes present, probably euryteles, none

discharged: (i) bean-shaped, 11-12 x 4-5 pm, in body of

gastrozooid and in tentaculozooid; (ii) elongate ovoid, 8-10 x

4-5 pm, in tentacles of gastrozooid.

Gonozooids absent.

Colour. Gastrozooids and tentaculozooids white, basal

perisarcal meshwork shining golden-yellow.

Remarks. The hydrorhizal meshwork is composed of chitinous

strands surrounding subcircular holes which vertically pene-

trate the meshwork. No structures which could be identified as

spines occur, the scattered mounds and peaks being hollow

protrusions of the perisarcal meshwork

Apart from Millard (1975) and Schuchert (1996) few

authors have adequately described the cnidome of Hydractinia.

Such knowledge would undoubtedly lead to easier identifi-

cation of species. In the present formalin-preserved sample the

nematocysts are undischarged and cannot be further identified.

Small zooids with four or five tentacles among the larger

gastrozooids are probably developing gastrozooids; other

unstructured white rings flush with the surface of the

hydrorhizal meshwork may be introverted gastrozooids. The

finely structured hydrorhizal meshwork investing the hydroid

host bears a remarkable resemblance to the ectoderm of

Solanderia. Despite careful search and examination of sections

of perisarc, no gonozooids were found.

Stepanjants (1979) gave a key to the following Antarctic

species of Hydractinia: H. antarctica Studer, 1879, H. angusta

Hartlaub, 1904, H. clavata, Jaderholm, 1905, H. pannspina

Hartlaub, 1905, H. pacifica Hartlaub, 1905 and H. vallini

Jaderholm, 1926. The absence of basal spines in the perisarc

Deep-water hydroids from Macquarie Island

153

considerably narrows the possible matches to known species of

Hydractinia, including those of the wider concept of

Hydractinia of Bouillon et al. (1997) and Boero et al. (1998).

The present specimen most closely resembles H. angusta but in

the absence of reproductive structures it cannot be confidently

assigned any of these species nor described as new.

Eudendriidae L. Agassiz, 1862

Eudendrium Ehrenberg, 1834

Eudendrium ?cyathiferum Jaderholm

Figures 2A-D

ing a hydranth, clusters seated on short, corrugated pedicels;

gonophore encircled by a large spadix shed as gonophore

matures; a single mature gonophore ovoid, borne on a slender

peduncle with dish- shaped distal end, gonophore at this

stage 0.4 mm long and 0.25 mm wide, enclosed in a distinct

transparent pellicle, containing 6-8 developing ova.

No nematocysts seen on gonophores; a few undischarged

eury teles present in coenosarc of stems; capsule small, droplet

shaped, 8 x 4.5 pm.

Colour. Fascicled stems and branches shining brown, fading

to yellowish on monosiphonic branches; gonophores were

probably creamy-pink.

Eudendrium cyathiferurn JMerholm, 1904: 11. — Jaderholm, 1905:

10, pi. 4 figs 1-3. — Bedot, 1925: 184. — Stepanjants, 1979: 18, pi. 1

fig. 13. — Marques et al., 2000: 92, fig. 42.

Material examined. Stn 120, NMV F91307, colony 50 mm high,

attached to fragment of calcareous bryozoan, specimen alcohol pre-

served. Stn 122, TM K2769, four colonies, the largest 90 mm high and

50 mm wide, attached to a dead solitary coral, Caryophyllia sp.,

specimen alcohol preserved.

Description. Colonies arborescent with thick, heavily fascicled

main stems up to 3 mm wide at base; hydrorhiza a mass of

stolons entwining substrate, becoming erect as polysiphonic

tubes of stem. Branching twiggy, in many orders, almost in one

plane, polysiphonic tubes running almost to tips of branches

but ultimate branches and branchlets (pedicels) monosiphonic,

given off from within the mass of tubes. Pedicels short, arising

more or less alternately from branches, with up to 10 clear basal

corrugations, remainder of pedicel more or less rugose. No

hydranths present on pedicels.

Several clusters of female gonophores on colony from stn

122; 6-10 immature gonophores in cluster completely replac-

Figures 2A-D. Eudendrium ? cyathiferum. A, female colony, stn 122.

B, cluster of immature female gonophores on colony. C, mature female

gonophores with ova from same colony. D, undischarged eurytele

from coenosarc of branch. Scale bar: A, 50 mm; B, C, 0.5 mm;

D, 10 pm.

Distribution. Previously recorded from South Georgia

(Jaderholm, 1905; Stepanjants, 1979) but now known to have a

wider distribution in deep subantarctic and Patagonian waters

(A. Marques, pers. comm.).

Remarks. There are two differences between the present speci-

men and available published and unpublished descriptions of

E. cyathiferum. These are: (i) the length of the peduncle of the

nearly mature gonophore; and (ii) the presence of only one size

of eurytele in the cnidome.

Three species of Eudendrium with pedunculate (i.e. second-

ary pedicellate) female gonophores are known: E. cyathiferum,

E. vervoorti Marques and Migotto, 1998 and E. glomeratum

Picard, 1951 (see Marques et al., 2000). E. glomeratum was

rejected as it is a temperate to tropical species (Watson, 1999),

E. vervoorti is discounted because of the small size of the

colony and the fact that the developing female gonophore lacks

an encircling spadix. Although the “golf-tee” peduncle sup-

porting the nearly mature female gonophore of the present

specimen is typical of E. cyathiferum it is, however, somewhat

longer and slenderer than that shown in a photomicrograph of

the type specimen (Marques et al., 2000).

The cnidome of the holotype specimen of E. cyathiferum

contains two size classes of euryteles, the smaller associated

with the hydrocaulus and the larger with the gonophore (A.

Marques, pers. comm.). In view of the limited amount of pres-

ent material available for study it could not be ascertained if

large nematocysts are always absent from the gonophore. The

few nematocysts in the coenosarc are similar in size to those in

the hydrocaulus of the holotype.

For the above reasons, together with the fact that the

colonies are smaller than previously reported for E. cyathi-

ferum, the present specimens are doubtfully referred to that

species.

Eudendrium deforme Hartlaub

Figures 3A-D

Eudendrium deforme Hartlaub, 1905: 508, 514, 552, figs B', C. —

Bedot, 1925: 184.

Material examined. Stn 44, NMV F91308, many sparsely fertile

colonies on dead stem of a primnoid gorgonian, colonies intergrown

and entangled with Tulpa diverticula, sertularian and Eudendrium

colonies, specimen alcohol-preserved. Stn 46, TM K2770, many

colonies on dead stem of primnoid gorgonian, specimen alcohol-

preserved.

154

Jeanette E. Watson

Description. Colonies comprising many tangled stems up to 70

mm long and 0. 3-0.4 mm diameter arising from a hydrortiiza

entwining stem of host. Stems more or less straight to curved,

monophonic, with 5-10 strong basal annulations and groups of

annul ati on s at intervals along stems.

Stems sparsely, alternately, but irregularly branched, more

or less in one plane, occasionally secondary branching occurs;

branchlets (pedicels) more or less alternate, spaced well apart,

short to moderately long, beginning with up to 10 deep annul-

ations; stems sometimes annulated throughout; perisarc other-

wise smooth. A few hydranths in poor condition remain on

colonies; hydranth large, with c. 24 tentacles. Perisarc of

lower stems thick and smooth, thinning distally along smaller

branches and pedicels.

Nematocysts all undischarged euryteles of one size; small,

droplet-shaped, a few in coenosarc of stems and tentacles of

hydranth, capsule 10 x 6 pm.

Male gonophores present on several colonies; clustered on

short, smooth to corrugated pedicels near base of stems; up to

5 in a beaded chain, apical gonophore single, gonophores

below paired; tentacles of hydranth partially resorbed and

reduced in number, hypostome intact.

Colour. Lower stem shining in dark brown, fading distally to

pale yellowish; gonophores white.

Distribution. Calbuco, Chile; Mar del Plata, Argentina

(Hartlaub, 1905).

Remarks. Due to entanglement of the stems it is difficult to

decide if one or many colonies are present. The few hydranths

are in very poor condition, due either to damage during collec-

tion or senescence of the colony. The species is distinguished

by its strictly monosiphonic, shining brown stems and paired

chain of immature gonophores.

Although Hartlaub ’s description of the species is brief I

have no doubt that the present specimens are referrable to

Eudendrium deforme. Like much of the present material,

Hartlaub’s specimens apparently comprised bare stems lacking

hydranths or reproductive structures. The thin brown stems, as

described and figured by Hartlaub are characteristic. The

species has not been recorded since its first description.

Eudendrium macquariensis sp. nov.

Figures 3E-G

Material examined. Stn 44, NMV F91309, holotype, abundant infertile

colonies on stems and branches of other species of Eudendrium,

specimen alcohol-preserved.

Description (of holotype). Stolons creeping over stems and

branches of hydroid host, stems numerous, to 7 mm long; stems

(pedicels) and stolons morphologically identical and 0.1 mm

diameter. Stems monosiphonic, unbranched, straggling,

straight or flexed, perisarc of stolons and pedicels distinctly to

weakly rugose throughout. Hydranths very small, distal on

pedicel, hypostome prominent, dome-shaped, surrounded by

10-14 tentacles. Perisarc of stolon and stems firm and of same

thickness throughout.

Nematocysts of two kinds present, both undischarged: (i)

small droplet- shaped euryteles, capsule 9-10 x 4-5 pm, very

Figures 3A-G. A, Eudendrium deforme. A, part of colony on dead

branch of primnoid gorgonian, stn 46. B, single stem. C, cluster of

male gonophores with paired immature gonophores and partially

resorbed tentaclesof hydranth. D, undischarged nematocyst, probably

eurytele from coenosarc of stem. E-G, Eudendrium macquariensis sp.

nov., stn 44. E, stems from holotype colony on Eudendrium deforme.

F, undischarged nematocyst from tentacles. G, undischarged nema-

tocyst, site unknown. Scale bar: A, 25 mm; B, 10 mm; C, 0.5 mm;

E, 1 mm; F, G, 10 pm.

abundant in tentacles; (ii) bean-shaped capsule 9x4 pm, rare,

site unknown.

Colour. Stolons white, stems clear pale yellowish, hydranths

white.

Etymology. The specific name refers to the type locality.

Remarks. The colonies of Eudendrium macquariensis so

thickly invest the stems and tips of branches of the hydroid host

that when first examined they seemed to be part of the host

colony. Closer examination revealed white stolons of E. mac-

quariensis running along the brown perisarc of the host. The

thick aggregations on the tips of the host permits this tiny

species to occupy an exceedingly crowded microhabitat. As

both stolons and stems are morphologically identical it is

difficult to judge at what stage the stolon becomes an erect stem.

The species is unusual in that the straggling stolon-stems are

almost entirely rugose. The rugosity is usually most conspic-

uous in the proximal pedicel region, as usual in Eudendrium.

Although the hypostome is dome-shaped rather than annular,

this may be an artefact of preservation; it is, however, quite

constant throughout the material. If the dome-shape is diagnos-

tic it sets E. macquariensis somewhat apart from other known

species of Eudendrium.

Deep-water hydroids from Macquarie Island

155

Eudendrium sp.

Figures 4A-C

Material examined. Stn 44, NMV F91310, many sparsely fertile

colonies intergrown with Eudendrium deforme on a dead branch of

primnoid gorgonian, specimen alcohol-preserved.

Description. Colonies short, arborescent and shrubby, the

largest 20 mm high and 20 mm wide. Main stem and branches

strongly fascicled, branching irregular in many planes, poly-

siphonic tubes running up branches to distal region, ultimate

branchlets (pedicels) monosiphonic, 0.1 mm in diameter, base

of pedicel with up to 20 annulations, perisarc thereafter smooth

to faintly corrugated.

Few hydranths remain, all in poor condition; small, with

12-14 moderately long tentacles.

Nematocysts small euryteles in tentacles and coenosarc,

droplet- shaped, capsule 8.5 x 4.5 pm, undischarged.

Male gonophores borne on a short, basally annulated

pedicel; a small cluster in series of 3 or 4 surrounding a

completely resorbed hydranth.

Colour. Lower stems pale honey-brown, fading to almost

transparent on ultimate branches; hydranths white.

Remarks. While the small, shrubby colony with male

gonophores surrounding a completely atrophied hydranth

Figures 4A-C. Eudendrium spec., stn 44. A, colony on dead branch of

primnoid gorgonian, stn 44. B, cluster of male gonophores. C, undis-

charged nematocyst, probably eurytele; in tentacles and coernosarc.

Scale bar: A, 20 mm; B, 0.5 mm; C, 10 pm.

reduces the possible matches with known species, in the

absence of hydranths and female gonophores and details of the

cnidome, identification is not taken further.

Leptothecata

Campanulinidae Hincks, 1868

Tripoma Hirohito, 1995

Tripoma arboreum Hirohito

Figures 5A-C

Tripoma arboreum Hirohito, 1995: 98, figs 28a-e, pi. 6, fig. A. —

Watson and Vervoort, 2000: 249, figs lA-D, 2A-J. — Watson and

Vervoort, 2001: 156, fig. 3a-b.

Material examined. Stn 63, NMV F91311, several infertile colonies,

the tallest 40 mm high on a primnoid gorgonian, specimen alcohol pre-

served. Stn 97, TM K2771, specimens alcohol-preserved. NMV

F91332, Stn 97, malinol-mounted microslide.

Description. Colonies erect, branched, the tallest 75 mm high.

Hydrorhiza reptant on stem of host, tubular, undulating, some-

times contorted. Erect stems arising from hydrorhiza, some

hydrothecae arising directly from stolon. Most stems poly-

siphonic, several younger stems monosiphonic and

unbranched, older stems with increasingly polysiphonic. Stem

internodes variable in length, cylindrical, smooth, nodes

usually distinct, transverse to slightly oblique, marked by a

constriction in perisarc. Apophyses alternate, long, distal on

internode, directed upwards at c. 45° to internodal axis, often

shifted slightly around stem, distal node of apophysis trans-

verse, sometimes extended into two or three segments, each

with deep transverse node.

Hydrothecae inverted conical, long, slender, slightly asym-

metrical in section, arising from stem apophysis, usually a well

to poorly marked internode between apophysial segment and

hydrotheca; adcauline wall weakly convex, abcauline wall

straight to weakly concave, a ring of tiny punctae near base of

hydrotheca, a band of remnant tissue extending diagonally

upwards into hydrotheca from desmocyte ring. Margin with 2

long, sharp triangular cusps separated by deeply scalloped

embayments, demarcation between margin and hydrothecal

body faint, operculum splits into at least 4 thin flaps. Perisarc

of hydrotheca delicate, thinning markedly to margin.

Colour Colony pale honey-yellow, distal parts and hydrothecae

almost colourless.

Measurements (pm)

Hydrorhiza

diameter 80-160

Stem

length of intemode 960-1200

width at node 59-86

diameter at node 72-88

adcauline length of proximal apophysis 52-100

diameter at proximal apophysial node 64-80

length of apophysis including extensions 80-200

Hydro theca

length including pedicel 770-941

length, base to apex of cusps 784-848

width of margin across base of cusps 200-264

depth of opercular embayment 136-160

156

Jeanette E. Watson

Figures 5A-C. Tripoma arboreum, stn 97. A, part of stem.

B, hydro theca with closed operculum. C, hydrotheca with operculum

open. Scale bar: A, 1 mm; B, C, 0.5 mm.

Distribution. Japan, Tasman Sea and Bass Strait, Australia

(Watson and Vervoort, 2001).

Remarks. The material conforms to the descriptions of colonies

of Tripoma arboreum given by Watson and Vervoort (2000,

2001). A feature not noted in previous descriptions is a line of

very small punctae in some hydrothecae a short distance above

the distalmost apophysial node, passing transversely to

obliquely across the hydrotheca and then continuing upward as

a line of granules, gradually merging with the hydrothecal wall

below the margin. Tissue remnants attached to the granules in

some hydrothecae suggest they provide support for the

hydranth.

Lafoeidae A. Agassiz, 1865

Lafoea Lamouroux, 1821

Lafoea tenellula Allman

Figures 6A-C

Lafoea tenellula Allman, 1877: 12, pi. 8 figs 3, 4. — Ritchie, 1911:

820, pi. 88 fig. 5.— Stechow, 1913: 110.— Stechow, 1923a: 10.—

Stechow, 1923b: 143.— Stechow, 1925: 453, fig. 23.— Leloup, 1940:

14. — ^Fraser, 1943: 90. — Fraser, 1944: 227, pi. 47 fig. 211. — ^Fraser,

1948: 232.— Deevey, 1954: 646.— Riedl, 1959: 646.— Yamada, 1959:

49.— Vervoort, 1968: 101.— Millard, 1973: 28.— Hirohito, 1995: 128,

figs 36d-f

Material examined. Stn 44, NMV F91312, specimen alcohol-pre-

served, NMV F91339, malinol mounted microslide of coppinia; fertile

colonies on Eudendrium and on primnoid gorgonian. Stn 90, NMV

F91333, malinol-mounted microslide, fragment detached from

substrate. Stn 119, TM K2772, specimen alcohol-preserved, NMV

F91334, malinol-mounted microslide, colony on Eudendrium. Stn 120,

NMV F91313, sparingly fertile colony on Tulpa diverticula and

Eudendrium, specimen alcohol-preserved.

Description. Hydrorhiza reptant on host hydroids, stolons

crumpled, colony predominantly stolonal but sometimes a short

length of stolon becoming free as an erect monosiphonic stem.

Hydrothecae given off irregularly in all directions from

hydrorhiza; hydrothecae long, elongate conical, radially

symmetrical, straight to faintly sinuous, no true pedicel but a

tubular narrowing of hydrotheca. A faint transverse to slightly

oblique ring of scattered punctae sometimes present marking

distal junction of pseudopedicel with hydrotheca, sometimes a

faint constriction in perisarc of hydrotheca at desmocyte ring.

Margin transverse to hydrothecal axis, circular, rim weakly

everted with up to seven, usually widely spaced, regenerations.

Persiarc of hydrotheca moderately thick near base, thinning

distally to margin. Hydranths not well preserved, deeply

contracted into hydrothecae, but probably with eight to 10

tentacles.

Coppinia comprising a tightly packed circle of gonothecae

seated on a thin, dish- shaped basal plate adherent to stem of

hydroid host; protective nematophore tubules scattered in irreg-

ular groups around and throughout coppinia. Gonothecae small,

crowded, flask- shaped, bases rounded, widening from base to

shoulder, surmounted by a short tubular neck with transverse,

slightly everted circular orifice; perisarc of gonothecae rather

thin. Ovoid gonophores (or planulae) present in some gono-

thecae but too degenerate for description. Protective tubules

varying in length from short to very long, tubular in section,

single or bifid, some completely or partially conjoined proxi-

mally, straight or curved, narrowing from base to apex; ter-

minal orifice circular; perisarc thick, usually smooth; some

undulated; others showing scars from interruptions to growth.

Colour. Colonies and coppiniae transparent colourless;

gonophores pale creamy white.

Measurements (pm)

Hydrorhiza

diameter 68-88

Hydrotheca

length of unregenerated hydrotheca 360-500

Figures 6A-C. Lafoea tenellula, stn 97. A, stolonal colony on branch

of Eudendrium, hydrorhiza produced into free stolon. B, regenerated

hydrotheca. C, longitudinal section through coppinia. Scale bar:

A, 1 mm; B, C, 0.5 mm.

Deep-water hydroids from Macquarie Island

length of regenerated hydrotheca

1060-1440

width at puncta ring

76-112

diameter at margin

168-240

Coppinia

overall length of gonotheca

320-440

maximum width of gonotheca

80-112

diameter of gonothecal orifice

48-60

length of nematophorous tubules

500-1700

width of tubule at base

96-104

diameter of orifice of tubule

48-56

Distribution. Australian east coast (Ritchie, 1911), California

(Fraser, 1948), Carribean, West Indies (Vervoort, 1968), Japan

(Hirohito, 1995). The geographical and bathymetric distribu-

tion of L. tenellula ranges from moderately deep tropical and

temperate waters to deep subantarctic waters (this collection).

Remarks. The hydrothecae on younger parts of the colonies are

less crowded than those on older regions. The hydranths are

deeply contracted into the hydrothecae and the tentacles of

reasonably well preserved ones appear to be enclosed in a

sheath of tissue.

The present material generally matches the description,

dimensions and figure of Lafoea tenellula given by Stechow

(1925) for specimens from Madeira and the Canary Islands but

conforms less well with the figure of hydrothecae given by

Hirohito (1995) for the species from Japan.

Three coppiniae were found among colonies in the collec-

tion; one was attached to the stem of Eudendrium from Stn 119

and the others to a primnoid gorgonian stem (Stn 44). Because

of intergrowth of the material with hydrorhizae of several other

hydroid species the coppiniae are ascribed with some hesitation

to L. tenellula, the gonosome of which is not known. Vervoort

(1966) renamed L. tenellula recorded by Ritchie (1911) from

off the Australian temperate east coast Hebella ritchei. I have

examined Ritchie’s material and other Australian east coast

specimens held in the Australian Museum, Sydney; as they are

identical with the present specimen, Ritchie’s identification of

L. tenellula is correct.

Lafoea dumosa (Fleming)

Figures 7A-E

Sertularia dumosa Fleming, 1820: 83 (nomen nudum).

Campanularia dumosa . — Fleming, 1828; 548-549.

Lafoea dumosa. — Hutton, 1904; 319. — Fraser, 1911: 51. — Billard,

1912: 464. — Stechow, 1912: 357. — Stechow, 1913: 30. — ^Fraser, 1914:

86.— Fraser, 1918: 333, 354.— Stechow, 1919: 80, fig. AT- Fraser,

1921: 73.— Fraser, 1922; 5.— Stechow, 1923a: 10.— Billard, 1927:

331.— Fraser, 1927: 326.— Totton, 1930: 158, fig. 14.— Billard, 1931:

245.— Leloup, 1934; 8.— Fraser, 1935: 144.— Kramp, 1935: 123, figs.

52a, 53.— Fraser, 1937: 119-120, pi. 25 fig. 137.— Fraser, 1938:

110.— Fraser, 1939: 160.— Leloup, 1940: 14.— Vervoort, 1942; 285.—

Fraser, 1944: 221, pi. 45 fig. 205, pi. 46 fig. 205.— Fraser, 1948:

229.— Teissier, 1950: 17.— Deevey, 1954; 270.— Hamond, 1957: 295,

307.— Ralph, 1958; 310.— Riedl, 1959: 646.— Yamada, 1959; 50.—

Leloup, 1960: 221. — Naumov, 1960; 276, fig. 165. — Rees and

Thursfield, 1965: 79.— Teissier, 1965: 19.— Redier, 1967: 389.—

Vervoort, 1968; 100. — Calder, 1970; 1524, pi. 5 fig. 3. — Christiansen,

1972: 296.— Calder, 1975; 299, fig. 3D.— Cornelius, 1975: 385, fig.

4.— Millard, 1975: 185.— Millard, 1977a: 15.— Millard, 1978: 195.—

157

Figures 7A-E. Lafoea dumosa, stn 44. A, colony. B, part of branch.

C, hydrotheca with typically twisted pedicel and diaphragm.

D, hydrotheca with replicated margin. E, radial arrangement of

hydrothecae around stem. Scale bar: A, 10 mm; B, E, 1 mm;

C, D, 0.5 mm.

Garcia Corrales et al., 1979: 19, fig. 8.— Millard, 1980: 13 L—

Stepanjants, 1980: 116. — Hirohito, 1983: 6, 21. — Stepanjants, 1985:

127, 131. — Antsulevich, 1987; 49, fig. IIB. — Rees and Vervoort,

1987: 40, figs 7-8.— Cornelius, 1988: 76.— Llobetet al., 1988; 38, fig.

4E. — Gili, Murillo and Ros, 1989: 23. — Gili, Vervoort and Pages,

1989: 73, fig. 3B.— Cornelius and Ryland, 1990: 135, fig. 4.13.— El

Beshbeeshy, 1991: 84, fig. 20. — Pena Cantero, 1991: 70, pi. 5 figs

a-d.— Calder, 1992: 1080.— Cornelius, 1992: 254, 257.— Ramil and

Vervoort, 1992; 55. — Boero and Bouillon, 1993; 263. — Branch and

Williams, 1993: 10.— Cornelius, 1995: 261, fig. 60.— Hirohito, 1995:

126, figs 36a-c, pi. 8 fig. A. — Medel and Lopez-Gonzalez, 1996:

198. — Pena Cantero and Garcia Carrascosa, 1995; 23, figs 4A-D. —

Stepanjants et al, 1996: 7. — ^Pena Cantero and Garcia Carrascosa,

1999: 212.— Schuchert, 2000; 413.— Schuchert, 2001: 67, figs 54

A-D, 55, 56.

Lafoea ?dumosa. — Ralph, 1958; 310.

Campanularia fruticosa M. Sars, 1850; 131, 138.

Campanularia gracillima Alder, 1856: 361, pi. 14 figs 5, 6.

Lafoea gracillima. — ^Ralph, 1958: 310, figs ly, 2a-c.

Lafoea capillaris G.O. Sars, 1874: 115, pi. 4 figs 4, 5.

Lafoea elegantula Broch, 1903: 5-6, pi. 1, figs 5, 6, pi. 2, figs 7-9.

Material examined. Stn 44, TM K2773, NMV E91314, specimen

alcohol-preserved. NMV E91340, malinol-mounted microslide from

colony, abundant infertile colonies on primnoid gorgonian.

Description. Colonies arborescent, delicate and rather lax, the

largest 25 mm high and 15 mm wide across the branches, stem

0.8 mm wide above base; smallest colonies 3-5 mm high.

Hydrorhiza thread-like tubular stolons reptant on substrate;

stolons bunched at base of complex colonies. Smallest stems

monosiphonic, taller colonies with 2 or more polysiphonic

tubes running up stem and along branches. Colonies branched

158

Jeanette E. Watson

at acute upward angles but in no particular order around

stem; branches fascicled proximally, becoming monosiphonic

distally, perisarc smooth without nodes. In proximal stem

region of larger colonies tubes rather twisted, gradually

becoming more or less parallel.

Hydrothecae given off from stem and branches, on mono-

siphonic branches (hydrocladia) arranged in whorls of three at

an angle of c. 120° around hydrocladium, but occasionally an

opposite pair; each hydrotheca well separated from its neigh-

bour; in polysiphonic branches where hydrothecae given off

from outer tubes of stem or branch this verticil arrangement

obscured. Hydrothecal pedicel long, directed at an acute

upward angle to hydrocladium, tubular, expanding distally to

merge smoothly into hydrotheca, pedicel undulated with to 1-4

bends; a thin transverse line of punctae marking junction of

hydrotheca with pedicel, a small annular internal ledge in

hydrothecal wall at puncta line. Hydrothecae long, narrowly

conical, widening smoothly to margin; body typically asym-

metrically bent, adcauline side weakly convex in proximal

third, then becoming almost straight or faintly sinuous to

margin; abcauline side straight to weakly concave; both walls

expanding a little below margin. Margin of hydrotheca circular,

perceptibly everted but with no outrolling of rim; some

hydrothecae with two or three marginal replications, basal-

most replication typically at some distance below margin. Most

hydrotheca contain remnants of hydranths with c. 12 tentacles.

Perisarc thickest on lower stem and branches, thinning

distally along branches, hydrothecae very thin and smooth.

Colour. Colonies pale yellowish in lower regions, becoming

translucent distally. Hydrothecae transparent, shining.

Measurements (pm)

Hydrorhiza

width of stolon 88-120

Branch

diameter of monosiphonic part 112-136

length from axil to first hydrotheca 680-720

Hydrotheca

distance between successive hydrothecae on

hydrocladium 240-464

length of pedicel to puncta line 240-280

diameter of pedicel 80-88

diameter at puncta line 96-112

length, puncta line to margin (including replications) 600-664

diameter at margin 176-232

Distribution. Near-cosmopolitan, in Atlantic, Pacific and Indian

Oceans, Arctic and Antarctic, depth range from the sublittoral

zone to deep sea (Cornelius, 1995; Schuchert, 2001).

Remarks. The hydrothecal pedicels are bent rather than twisted

or smoothly annulated with up to 4 kinks; the perisarc is

smooth throughout. Most colonies are fascicled, only the very

youngest and shortest being monosiphonic. Single hydrothecae

are abundant on the hydrorhiza between complex stems. The

colonies are quite lax and unable to support their weight out of

fluid.

The present material conforms in most respects with earlier

descriptions of Lafoea dumosa. It does, however, differ some-

what from most descriptions in having an almost invariable

triseriate arrangement of the hydrothecae on monosiphonic

hydrocladia, this condition usually being obscured in poly-

siphonic stems and branches, rather than the incipiently

triseriate or multiseriate arrangement described by some authors.

Lafoea annulata sp. nov.

Figures 8A-E

Material examined. Stn 44, NMV F91315, holotype, specimen

alcohol-preserved, NMV F91341 malinol-mounted microslide from

holotype colony; infertile colony on a fragment of primnoid gorgonian,

hydrorhiza intergrown with those of other hydroids.

Description (of holotype). Colony 70 mm high and 40 mm

wide; hydrorhiza a broken tangled mass of tubular stolons coa-

lescing to form polysiphonic tubes of stem; colony branched

almost from base in disorderly flabellate fashion, base of stem

and lowest branch 1 mm thick, both heavily fascicled, poly-

siphonic tubes almost linear, sometimes a little twisted in lower

stem region then running up branches; distal region of some

branches with 2 or more tubes, a few branches monosiphonic.

Branches bent at various angles, becoming more or less straight

distally; walls of ultimate branches smooth without nodes.

Apical stolonisation occurring on several branches where a

tube becomes free of branch, walls of free tubes contorted and

undulated, sometimes rejoining branch.

Figures 8A-E. Lafoea annulata sp. nov., stn 44. A, colony.

B, distal stolonisation at end of branch. C, hydrotheca with twisted and

partially annulated pedicel. D, hydrotheca with almost smooth pedicel.

E, undischai'ged nematocyst from coenosarc of branch. Scale bar:

A, 25 mm; B, 1 mm; C, D, 0.5 mm; E, 20 pm.

Deep-water hydroids from Macquarie Island

159

Hydrothecae given off without apophyses from mono-

siphonic branches or from peripheral tubes of polysiphonic

branches; hydrothecae in opposite pairs or in groups of three,

one arising from each tube of branch, if three, the third arising

close to opposite pair, forming an incipient whorl of 3, groups

well separated at variable distances along branch. Hydrothecal

pedicels tubular, length variable but mostly long, straight to

bent, widening distally and merging smoothly into base of

hydrotheca below puncta line; pedicels usually deeply annul -

ated to almost smooth, frequently one side more deeply

annulated than other.

Hydrothecae rather fragile, long, tubular, slightly asymmet-

rically bent with one wall slightly convex in basal third,

opposite wall weakly concave to straight; diaphragm very thin,

transverse to slightly oblique, saucer-shaped with central

hydropore marked by an indefinite double ring of small scat-

tered punctae; a slight thickening of hydrothecal wall at puncta

line. Margin of hydrotheca circular, transverse to hydrothecal

axis, rim smooth, noticeably everted, many hydrothecae with

up to 5 everted marginal replications, usually well separated,

some well below rim.

Perisarc in polysiphonic stem region and branches quite

thick, thinning in monosiphonic parts; in branches with fewer

tubes, perisarc of youngest tube markedly thinner than others;

perisarc of hydrothecae thin, transparent and shining.

Hydranths too poorly preserved for description; remnants

emerging from many hydrothecae as a long strand of tissue.

Nematocysts large, bean-shaped, probably isorhizas, 21-22

X 8.5-9 pm, none discharged, but containing a long, probably

isometric, closely coiled tubule; abundant in coenosarc of

polysiphonic tubes of branches.

Colour. Colony translucent pale honey-yellow, mono-

siphonic branches and hydrothecae almost colourless.

Measurements (pm)

Branch

diameter of monosiphonic branch 56-88

distance between groups of pedicels 900-2000

length of pedicel to puncta line 1720-4000

proximal diameter of pedicel 80-144

Hydrotheca

diameter at puncta line 128-160

length, puncta line to margin (including regenerations) 520-704

diameter of margin (including eversion) 200-240

Remarks. An empty conical structure at the broken distal end of

a branch may be a damaged gonotheca; the structure is trans-

parent, adpressed to the branch with the wider end facing dis-

tally and is overgrown by several polysiphonic tubes; it could

be an empty corophiid tube. The diaphragm at the puncta line

of the hydrotheca may have a downwardly directed hydropore

but because of its delicacy the actual structure could not be

ascertained. The free stolons at the ends of several branches

seem to have developed from hydrothecal pedicels which have

reverted to polysiphonic tubes, growth then proceeding onward

without development of a hydrotheca. The thin strands of tissue

issuing from many hydrothecae may be collapsed protective

sheaths of hydranths. Several hydrothecae have remnants of

tissue adhering to the rim which could easily be mistaken for

opercular fragments.

Similar species considered were Lafoea fruticosa (M. Sars,

1850), L. gracillima 1856, L. capillaris G. O. Sars, 1874,

L. elegantula (Broch, 1903), L. dumosa Fleming, 1828 and L.

benthophila Ritchie, 1909. While L. annulata has a hydrotheca

with everted margin similar to L. benthophila that species has

an upwardly directed pedicel so that the hydrothecae are held

close to the branch. Similarly to L. annulata, hydrothecae of

L. dumosa from western Europe are arranged in groups of

two or three (Cornelius, 1995) but lack an everted margin.

Vervoort (1972a) described the range of variation of South

Atlantic L. fruticosa from latitudes 42° to 60° S. However,

it is much smaller in critical dimensions of hydrotheca and

pedicel and the hydrothecae have less everted margins than

those of L. annulata. A small colony from the Ross Sea recog-

nised as L. gracillima by Totton, 1930 approaches the dimen-

sions of the present specimen but his figures do not show the

annulated pedicels characteristic of L. annulata.

Filellum Hincks, 1868

Filellum conopeum sp. nov.

Figures 9A-C

Filellum sp. — ^Watson and Vervoort, 2001: 161, figs 6a, b.

Material examined. Stn 94, NMV F91342, holotype, malinol-

mounted microslide, sparse fertile colony on stem of Acryptolaria

patagonica.

Description (of holotype). Hydrothecae stolonal, arising from a

tunnel-shaped stolon with flattened base, wall rugose, perisarc

thin. Proximal quarter to one third of hydrotheca adnate to

Figures 9A-C. Filellum conopeum sp. nov., stn 94. A, hydrotheca with

ridged and frilled basal perisarc. B, lateral section through coppinia

showing gonothecae and protective tubes. C, transverse section

through coppinia. Scale bar; A, 0.2 mm; B, 0.5 mm.

160

Jeanette E. Watson

stolon, dorsal abcauline wall furrowed by many close,

sharp-edged ridges with minute ragged frill of perisarc; ridges

fading on adnate wall. Adnate wall becoming free at a sharp

upward bend, free part cylindrical or weakly expanding from

bend to margin, free part straight to broadly curved, walls

smooth, occasionally with several regenerations. Margin cir-

cular, transverse, with smooth, distinctly everted rim. Perisarc

of walls fairly thick, thinning distally. Hydranth with c. 12

tentacles and clavate hypostome.

Coppinia bud- shaped, c. 1 mm wide and 1 mm high, com-

prising many tightly packed gonothecae enclosed within a cone

of protective nematophorous tubules. Gonotheca flask-shaped

(lateral view), base rounded, body expanding a little from base

to shoulder then narrowing into a short straight or slightly

curved neck tapering to a circular aperture; in transverse view

gonothecae polygonal. Nematophorous tubules similar in

length, not forked, conjoined just above gonothecae then

becoming free, most narrowing distally and inwardly curved to

meet above gonotheca; terminal orifice circular. Perisarc of

gonothecae and tubes moderately thick; perisarc of tubes some-

what roughened. Planulae enclosed in gonothecae small,

spherical.

Colour. Colonies colourless, planulae creamy pink.

Measurements (pm)

Hydrorhiza

width

Hydrotheca

dorsal length of adnate part

180-240

depth of adnate part

92-116

length of free part

320-420

width of free part at bend

88-128

diameter at margin

140-168

Coppinia

diameter of nematophorous tubules

64-120

length of gonotheca

320-440

maximum width of gonotheca

128-160

diameter of orifice

64-80

diameter of planula

80-120

Etymology. Named for the cone of tubes protecting the

gonothecae.

Remarks. Epizoic colonies of Filellum conopeum occur

sparsely on the stem of Acryptolaria. The thin ragged peri-

sarcal frill surmounting the abcauline ridges of the hydrotheca,

together with the bud-shaped coppinia are characteristic.

The trophosome of F. conopeum resembles that of F. serra-

tum, however, the coppinia of that species as described and

figured by Pena Cantero et al. (1998) is quite different, the

gonothecae being protected by a canopy of apically divided

tubes, some of which originate from within the gonothecal

mass. Watson and Vervoort (2001) reported a species of

Filellum from the deep-water seamounts south-east of

Tasmania, Australia, but as their material was infertile and

dimensions somewhat greater than those of F. serratum

(Clarke, 1879) (the only other known species with a wrinkled

abcauline wall), they hesitated to identify it with that species.

While the hydrothecae of F. conopeum are somewhat smaller

than those of the Tasmanian specimens I consider the two are

conspecific.

Zygophylax Quelch, 1885

Zygophylax sagamiensis Hirohito

Figures lOA-F

Zygophylax sagamiensis Hirohito, 1983: 30, fig. 11. — Rees and

Vervoort, 1987: 85.— Hirohito, 1995: 144, figs 44a-e, pi. 9 fig. C.

Material examined. Stn 44,TM K2775, infertile colony fragment 10

mm high, specimen alcohol-preserved. Stn 63, TM K2774, specimen

alcohol-preserved, NMV F91335, malinol mounted microslide from

fertile colony fragment 10 mm high on primnoid gorgonian stem. Stn

122, NMV F91316, specimen alcohol-preserved, NMV F91336

mahnol-mounted microslide, complete branched colony 60 mm high

and 50 mm wide, detached from substrate.

Description. Hydrorhiza a mat of stolons running along axis of

gorgonian, bunching together at base of colony then passing

upwards to become fascicular tubes of stem. Polysiphonic

stems stiff, sparingly subalternately branched, primary

branches widely spaced, occasionally secondary subdicho-

tomous branching, one or two polysiphonic tubes running

halfway to two thirds along branch or hydrocladium; branch

P) ,

Figures lOA-F. Zygophylax sagamiensis, stn 63. A, colony.

B, distal part of branch. C, hydrotheca with multiple marginal replica-

tions. D, pedicellate nematotheca on hydrorhiza. E, coppinia.

F, hooded gonothecae from coppinia. Scale bar: A, 25 mm; B, 0.5 mm;

C, 0.2 mm; D, 0. 1 mm; E, E, 0.25 mm.

Deep-water hydroids from Macquarie Island

161

and hydrocladia thereafter monosiphonic; a hydrotheca in axil

of each branch. Hydrocladial intemodes long, slender, perisarc

moderately thick, smooth, nodes merely transverse constric-

tions, distinct when present, but often absent. Hydrothecae

alternate to subalternate, widely spaced, facing frontally, 1 or 2

on internode, if one, about halfway to two-thirds up internode,

if 2, one just below and the other just above node. Hydrocladial

apophyses short, distal node transverse, deep.

Hydrothecal pedicel cylindrical, slender, variable in length,

sometimes with one to several regenerations. Hydrothecae

long, slender, slightly asymmetrical, adcauline wall convex,

abcauline wall weakly concave, diaphragm distinct, clearly

demarcated from hydrothecal wall, transverse or oblique

(depending on angle of view) with central wide hydropore.

Hydrotheca margin circular, transverse, rim distinctly everted,

some hydrothecae with up to 8 marginal replications; perisarc

of hydrotheca smooth.

Nematothecae numerous on peripheral tubes of fascicled

stem, on hydrorhiza, on apophyses below hydrothecal pedicels,

on surface of coppinia and on coppinial tubes; nematothecae

small, vase-shaped, sometimes slightly asymmetric, pedicel

very short, margin transverse, circular, rim slightly everted.

Coppinia (Stn 122) scarcely visible, embedded in poly-

siphonic tubes of stem between primary branches, marked by a

faint swelling of the stem and numerous projecting

nematophorous tubules. Gonothecae small, cylindrical, tightly

packed, with low hooded semicircular orifice, a few with an

apical peak, walls thick. Tubules issuing from coppinial mass,

long, with one or two basal constrictions, some with one or two

nematothecae, some with a few nodes along length and some

incipiently branched.

Colour. Stems pale honey yellow, hydrocladia paler.

Measurements (pm)

Branch

length of internode

360-392

width at node

36-52

Hydrotheca

length of pedicel

108-120

length diaphragm to margin, including replications

392-600

diameter at diaphragm

56-64

diameter at margin

116-148

Nematotheca

length

76-108

diameter at margin

36-40

Coppinia

maximum length of tube

1050

diameter of tube

56-60

width of gonotheca

64-100

Distribution. Previously known from a depth of 300 m in

Sagami Bay, Japan (Hirohito, 1995).

Remarks. Branches are given off from just inside the peri-

pheral tubes of the stem. Hydrothecae in the axils of branches

usually have long, undulated or regenerated pedicels.

Hydrocladial nodes may be faint or altogether absent. As there

is only one coppinia in the sample and the gonothecae are

deeply embedded in the mass, sex could not be determined.

Nematothecae are so numerous on the hydrorhizal stolons and

peripheral tubes of the lower stem region that they impart a

rough, prickly appearance to these structures. Hirohito (1995)

mentioned the presence of nematothecae on the peripheral

tubes of the stem but not on the hydrorhiza.

In all but arrangement of the hydrothecae the present speci-

mens resemble Zygophylax tottoni Vervoort, 1987 in which the

branching is strictly planar and hydrothecae are not frontally

directed. As the present specimens generally conform with

descriptions and figures of Z. sagamiensis given by Hirohito

(1983, 1995) the material is referred to that species. Differences

with the present specimen are: (i) the fewer apical peaks on the

gonothecae; and (ii) the numerous coppinial nematothecae.

Acryptolaria Norman, 1875

Acryptolaria conferta (Allman)

Figures llA-D

Cryptolaria conferta Allman, 1877: 17, pi. 12 figs 6-10. — Stechow,

1913: 30.— Jarvis, 1922; 335.— Keller et al., 1975: 148.

Grammaria conferta. — Broch, 1913: 10. — Broch, 1918: 17-18.

Cryptolaria conferta var. australis Ritchie, 1911: 826, pi. 84 fig. 2,

pi. 87 fig. 1. — Jaderholm, 1919; 7, pi. 2 fig. 1. — Totton, 1930; 163, figs

19c-e. — Ralph, 1958: 315, figs 4a-g. — Yamada, 1959: 49. — Rees and

Thursfield, 1965: 82, 194.— Millard, 1967: 172.

Oswaldaria conferta var. australis. — Stechow, 1923a: 11.

Oswaldaria conferta. — Stechow, 1923b: 147. — ^Leloup 1940: 15. —

Picard, 1958: 193. — Marinopoulos, 1981; 176.

Acryptolaria conferta. — Totton, 1930; 164, figs 19a, b. — Kramp,

1932: 68, fig. 32.— Leloup, 1937: 4, 29, fig. 19.— Fraser, 1943: 90.—

Fraser, 1944: 210, pi. 40 fig. 189.— Ki-amp, 1947; 8.— Fraser, 1948:

Figures 11 A-D. Acryptolaria conferta, stn 44. A, colony with coppinia

on stems and branch. B, hydrothecae. C, distal end of branch.

D, lateral section through part of coppinia. Scale bar: A, 10 mm;

B, 1 mm; C, D, 0.5 mm.

162

Jeanette E. Watson

228. — Rossi, 1950; 201, fig. 4a. — Deevey, 1954: 270. — Ki'amp, 1963:

106.— Millard, 1964: 7, fig. lA-C, E.— Millard, 1968: 253, 260.—

Vervoort, 1968: 99. — ^Vervoort, 1972a; 41, fig. 12a. — Millard, 1973:

28, fig. 4c.— Millard, 1975: 169, fig. 56.— Millard, 1977b: 106.—

Millard, 1978: 188. — Stepanjants, 1979: 51, pi. 9 figs 4A-G. —

Millard, 1980: 131.— Hirohito, 1983; 6, 19.— Calder, 1991: 33, figs

19, 20.— El Beshbeeshy, 1991: 63, fig. 13.— Park, 1992: 287.— Boero

and Bouillon, 1993: 263. — Calder, 1993: 67. — Blanco et al., 1994: 8,

figs 4, 5. — Altuna Prados, 1995: 54. — Bouillon et al., 1995: 51. —

Hirohito, 1995: 104.— Calder. 1997: 87.— Schuchert, 2001: 62, figs

48A, B.

Acryptolaria conferta var. conferta. — Ralph, 1958: 317.

Acryptolaria conferta australis Millard, 1964: 9, figs ID,

E-G. — Vervoort 1966; 115, fig. 15. — Rees and Vervoort, 1987; 37,

fig. 6e.

Acryptolaria conferta conferta. — ^Vervoort, 1985: 282. — Ramil and

Vervoort 1992: 41, figs 7a, b.

Material examined. Stn 44, TM K2776, NMV E91317, many colonies

on pebbles and primnoid gorgonian fragment; one colony consisting of

2 large stems (one stem fertile) and several simple stems, other

colonies of several large infertile stems, specimens alcohol-preserved.

Description. Simple and branched colonies connected by

hydrorhiza replant on gorgonian. Larger branched colonies

flabellate, up to 50 mm high and 35 mm wide, stems to 1 mm

wide at base; colonies arising from a thick plug of stolons,

simple stems from single stolons. Stem and branches of flabel-

late colonies heavily fascicled, with roughly longitudinal poly-

siphonic tubes running upwards from stem and along branches;

ultimate parts of branches (hydrocladia) monosiphonic.

Branching irregular, of up to 3 orders, branches given off

behind a hydrotheca, best seen in distal parts of colony.

Monosiphonic branches weakly geniculate, given off from

behind a hydrotheca.

Hydrothecae biseriate, alternate, scarcely overlapping, often

frontally directed; on monosiphonic branches biseriate arrange-

ment of hydrothecae sometimes replaced by incipient whorls of

three around hydrocladium; on fascicled parts of stem and

branches hydrothecae often not strictly alternate, immersed in

polysiphonic tubes with only distal part or margin visible.

Hydrocladial hydrothecae long, tubular, bending outwards at an

angle of 40-50° to hydrocladial axis; adcauline wall convex,

often a slight change in convexity where wall becomes free;

free part about same length as adnate part, hydrotheca narrow-

ing proximally along adnate wall, sometimes fading into a

ragged septum. Abcauline wall curving smoothly into hydro-

cladium, usually a transverse ring of desmocytes just above

narrowest part of hydrotheca, marking site of attachment of

hydranth. Margin circular, rim transverse to hydrothecal axis,

slightly but distinctly everted; up to seven marginal replications

common. Perisarc of stem and branches firm, thinning a little

on hydrotheca. Hydranths too poorly preserved for description.

Coppinia situated about halfway up stem, spreading for 10

mm along a branch and 5 mm along adjoining stem; coppinia

cylindrical, diameter 2 mm, texture rather spongy. Gonothecae

radially arranged within coppinia around polysiphonic tubes of

stem and branch; flask-shaped, sides more or less straight,

conjoined to neighbours, expanding from narrow base to a

rounded shoulder surmounted by an erect tubular neck narrow-

ing into a circular non-e verted terminal orifice. No protective

tubules present. Small spherical planulae visible in some

gonothecae.

Colour. Colony translucent honey-gold; hydranths probably

of same colour; gonophores creamy pink.

Measurements (pm)

Branch

distance between hydrothecae on same side of branch 980-1160

width of hydrocladium at junction of free and adnate

adcauline wall 144-192

Hydrotheca

length of free adcauline wall including marginal

replications 712-860

length of adnate adcauline wall to desmocyte ring 200-A00

diameter at margin 184-200

Coppinia

overall length of gonotheca 520-528

width of gonotheca across shoulder 136-192

length of gonothecal terminal neck 96-160

diameter of gonothecal temiinal orifice 44-48

Distribution. Moderate to deeper waters of all oceans (Vervoort

and Watson, 2003).

Remarks. The verticil of three hydrothecae occasionally present

on monosiphonic hydrocladia approaches the generic definition

of Cryptolarella Stechow, 1913 in which the hydrothecae may

be triserially disposed around the stem. I cannot find in the

literature any mention of the more consistent feature in the

present specimens of the frontally directed hydrothecae, all

descriptions reporting hydrothecae either lying in the plane of

ramification or exceptionally, backwardly or forwardly

directed at the base of each new hydrocladium (Ramil and

Vervoort, 1992). However, as the colonies conform in all other

respects with Acryptolaria conferta, the material is presently

assigned to that species.

Acryptolaria patagonica El Beshbeeshy

Figures 12A-C

Acryptolaria patagonica El Beshbeeshy, 1991: 67, fig. 14.

Material examined. Stn 90, TM K2777, stem 50 mm high, NMV

E91318 stem fragments 5 mm high, all detached from sub-

strate, specimens alcohol-preserved. Stn 90, NMV E91337,

malinol-mounted microslide from stem fragment. Stn 94, NMV

E91319, colony 100 mm high, two stems 50 mm high joined at base

but detached from substrate, three stems to 5 mm high and stem frag-

ment; on primnoid gorgonian, specimens alcohol-preserved. Stn 130,

TM K2778, stem 50 mm high, detached from substrate, specimen

alcohol-pre served.

Description. All colonies infertile. Stolons replant on substrate,

tubular, narrow, coalescing into erect, almost straight stems;

sometimes single hydrothecae arising from stolons between

stems. Taller stems to 2 mm thick at base, basal 5-10 mm

unbranched; lower stem region strongly fascicled, polysi-

phonic tubes mostly linear, running up stem and along primary

branches. Complex colonies with up to four orders of branch-

ing; branches more or less alternate, usually in one plane,

primary and subsequent branches often originating just above

a hydrotheca, ultimate branches (hydrocladia) and some

branches in lower stem region monosiphonic.

Deep-water hydroids from Macquarie Island

163

Figures 12A-C. Acryptolaria patagonica, stn 44. A, colony.

B, hydrothecae. C, multiple replications of hydrothecal margin. Scale

bar: A, 12 mm; B, C, 1 mm.

Hydrothecae biseriate, tubular, subalternate, usually frontal

on branch, long, gracefully arched outwards, abcauline wall

concave, often a minor swelling in proximal wall; adcauline

wall smoothly convex, one third adnate to hydrocladium,

adnate part narrowing a little proximally and fading into hydro-

cladium; hydrocladium fairly narrow behind adnate hydro-

thecal wall. Margin circular, transverse to hydrothecal axis, rim

slightly everted, almost parallel to hydrocladial axis; margin

may have up to 14 close replications bending plane of rim away

from axis of hydrotheca. Cauline hydrothecae partially

immersed in polysiphonic tubes of stem and primary branches;

tubes then often contorted around hydrothecae.

Hydranths contracted and wrinkled; with c. 20 tentacles and

clavate hypostome; hydranth attached to base of adnate

hydrothecal wall by a ring of tissue.

Perisarc of stem and branches thick and smooth, perisarc of

hydrothecae thinner and shining.

Colour. Stems pale honey-yellow fading to almost colour-

less at tips of branches. Hydranths may have been deep golden-

brown.

Measurements (pm)

Branch

distance between hydrothecae on monosiphonic branch 900-1200

width of hydrocladium where adnate hydrothecal wall

becomes free 104-136

Hydrotheca

length (diagonal) of free adcauline wall including

marginal replications 1500-1600

length of adnate adcauline wall from base of hydranth 880-1040

width at base of hydranth 232-256

diameter at margin 416-448

Distribution. Patagonian shelf (El Beshbeeshy, 1991). Also

recorded from 415-1060 m at 39°-55°S, near Macquarie Island

(Vervoort and Watson, 2003).

Remarks. The larger colonies are rather flexuous and the ulti-

mate monosiphonic branches are quite lax out of fluid. The

long, curved hydrothecae are characteristic. The free part of the

hydrotheca is tubular but the adnate adcauline hydrothecal wall

narrows, becoming increasingly indefinite and rather fibrous in

appearance as it passes downward into the hydrocladium.

The frontally directed, long curvaceous hydrothecae resemble

Acryptolaria patagonica more than any other known species of

the genus. Although the habit of the present specimens differs

somewhat from descriptions of A. patagonica, this may be due

to immaturity of the colonies. It is unfortunate that the present

material is infertile, since a coppinia would confirm its

identity.

Acryptolaria minuta sp. nov.

Figures 13A-C

Material examined. Stn 130, NMV F 91338, holotype, small infer-

tile colony of five small stems, two branched, on dead primnoid

gorgonian, malinol-mounted microslide.

Description (of holotype). Tallest stem 9 mm high, broken off at

tip, 3.5 mm wide at base; stems branched once in one plane;

branched stems arising from a small matted plug of stolons,

simple stems from junction of stolonal tubes; taller stems with

up to four polysiphonic tubes intergrown and rather contorted

proximally, becoming linear distally; polysiphonic tubes run-

ning almost to top of stems; stems lightly fascicled; branches

given off beside a hydrotheca.

Hydrothecae biseriate, alternate, long, tubular, somewhat

frontally directed, scarcely overlapping, curving gracefully out-

wards at an angle of 50-60° from hydrocladial axis; single

hydrothecae on proximal region of larger stems partially

immersed in fascicular tubes. Adcauline hydrothecal wall

smoothly convex, free wall slightly more than half length of

adnate wall; adnate wall narrowing proximally downwards into

hydrocladium, base of wall ending in minute knot of perisarc;

abcauline wall smoothly concave, passing without interruption

into hydrocladium. Margin evenly circular, transverse to

hydrothecal axis, very weakly everted, sometimes with several

replications of rim. Perisarc of stem and branches firm, thin-

ning a little on hydrotheca. Hydranths deeply retracted into

hydrotheca; with c. 12 tentacles.

Colour Pale honey-yellow to colourless.

Measurements (pm)

Branch

distance between hydro thecae on same side 624-728

width at junction of adnate and free adcauline walls 152-168

Hydrotheca

length of free adcauline wall including marginal

replications 200-368

length of adnate adcauline wall 408-464

diameter at margin 104-128

Etymology. Describes the minute nature of the colonies.

Remarks. The habit of Acryptolaria minuta is similar to that of

A. patagonica described above. The species is, however, con-

siderably smaller in critical dimensions of the hydrocladium

and hydrotheca. The slightly everted margin of the hydrotheca

164

Jeanette E. Watson

Figures 13A-C. Acryptolaria minuta sp. nov., stn 133. A, whole

colony. B, part of branch. C, hydrotheca with replicated margin. Scale

bar: A, 3 nun; B, 0.5 mm; C, 0.2 mm.

resembles A. conferta minor Ramil and Vervoort, 1992 but it is

smaller and the hydrothecae are frontally directed in contrast to

those of A. conferta minor which, according to these authors,

lie in the plane of ramification of the branches.

Even if the colonies are immature specimens of a larger

species their smaller dimensions and sparsely branched habit

matches no other known species of Acryptolaria.

Haleciidae Hincks, 1868

Halecium Oken, 1815

Halecium ralphae Watson and Vervoort

Figures 14A-D

Halecium beanii. — Ralph, 1958; 332, fig. lOe, pro parte.

Halecium sessile. — Hirohito, 1995; 27, fig. 7g, pro parte.

Halecium ralphae Watson and Vervoort, 2001: 162, figs 7a-e.

Material examined. Stn 119, TM K2795, infertile colony on dead soli-

tary coral, specimen alcohol-preserved. Stn 44, NMV F91320, small

infertile stem detached from substrate, specimen alcohol-preserved.

Description. Colony 60 mm high and 2 mm thick at base;

smaller colony 30 mm high. Hydrorhizal filaments reptant on

coral; colony aborescently branched in up to three orders,

several branches broken off. Branching occurs from below a

hydrotheca, stem and branches heavily fascicled, polysiphonic

tubes almost linear to undulating, sometimes knotted around

origin of branch, tubes running along branches, ultimate

branches monosiphonic. Perisarc of stem and proximal

branches thick, thinning out on monosiphonic parts.

Monosiphonic branches (hydrocladia) arising from poly-

siphonic tubes on an apophysis of stem comprising one to three

subspherical segments with strong transverse nodes.

Hydrocladial internodes long, cylindrical, smooth, widening

distally, nodes slightly oblique, tilted away from hydrotheca,

marked by a deep constriction and tumescence in perisarc,

sometimes an additional short subspherical internode similar to

apophysis above node.

Hydrophores alternate, distal on intemode, almost entirely

adnate to intemode, adcauline wall marked by a seam in perisarc,

a semicircular thinning in perisarc below some hydrothecae;

abcauline wall contiguous with wall of intemode. Hydrotheca

free, closely adpressed but not adnate to intemode, very shallow,

tilted at slightly away from intemode, hydrothecal margin very

weakly everted, diaphragm distinct, a ring of desmocytes above.

Hydranths too poorly preserved for description.

Colour. Stem and fascicled branches honey brown;

monosiphonic parts pale brown.

Measurements (pm)

Branch

length monosiphonic branch internode 580-800

diameter at node 96-120

Hydrotheca

length adnate wall to diaphragm 100-128

diameter at diaphragm 128-144

depth margin to diaphragm 30-38

diameter of margin 152-154

Distribution. Previously recorded from 475-512 m off the

Chatham Islands (Ralph, 1958), shallow water in Japan

(Hirohito, 1995) and from 700-1122 m south of Tasmania

(Watson and Vervoort, 2001).

Figures 14A-D. Halecium ralphae, stn 119. A, colony. B, distal part of

branch. C, apophysis of stem and proximal part of branch.

D, hydrophore and hydrotheca. Scale bar: A, 5 nun; B, C, 0.5 mm;

D, 0.1 mm.

Deep-water hydroids from Macquarie Island

165

Remarks. Some large undischarged bean-shaped nematocysts

visible in the coenosarc of the branches could not be identified.

The marginal rims of some hydrothecae have one or two

obscure regenerations. Other than the noticeable thinning of the

perisarc of the hydrophore below some hydrothecae the

material conforms to the description and dimensions of

Halecium ralphae given by Watson and Vervoort (2001).

Halecium tenellum Hincks

Figures 15A-C

Halecium tenellum Hincks, 1861: 252, pi. 6 figs 1-4. — Hartlaub,

1904: 13, pi. 1 fig. 5. — Hartlaub, 1905: 609, fig. 63. — Jaderholm,

1905: 13, pi. 4 fig. 8. — Hickson and Gravely, 1907: 28. — Ritchie,

1907: 525, pi. 2 fig. 4.— Vanhoffen, 1910: 320, fig. 36.— Hilgendorf,

1911: 540.— Linko, 1911: 26, 240, fig. 5.— Ritchie, 1913: 10, 14.—

Broch, 1918: 46, fig. 20. — Jaderholm, 1919: 5, pi. 1 fig. 3. — Stechow,

1919: 41, figs J-K.— Stechow 1923a: 5.— Hargitt, 1927: 507.— Broch,

1927: 115.— Broch, 1928: 61.— Broch, 1933: 17.— Fraser, 1937: 110,

pi. 23, fig. 121. — ^Leloup, 1937: 4, 17, fig. 8. — Fraser 1938: 133. —

Fraser, 1939: 159. — Fraser, 1948: 225. — Dawydoff, 1952: 54. —

Hamond, 1957: 307, fig. 14.— Millard, 1957: 193, fig. 5.— Vervoort,

1959: 229, fig. 8.— Yamada, 1959: 31.— Leloup, 1960: 220, 230.—

Naumov, 1960: 454, fig. 344. — Mammen, 1965: 9, fig. 35. — Vasseur,

1965: 52, 70.— Millard, 1966: 471, figs IIC-R— Vervoort, 1966: 102,

fig. 2.— Millard, 1968: 253, 258.— Vervoort, 1968: 95.— Day et al.,

1970: 12.— Hirohito, 1974: 8, fig. 2.— Leloup, 1974: 1 1 .—Millard and

Bouillon, 1974: 5, fig. 22. — Rho and Chang, 1974: 136, pi. 1 figs

1-4.— Vasseur, 1974: 158.— Cornelius, 1975: 409, fig. 12.— Millard,

1975: 156, figs 5 OF-L.— Millard, 1977a: 11.— Millard, 1977b: 106.—

Rho, 1977: 252, 414, pi. 71 fig. 63.— Millard, 1978: 193.—

Stepanjants, 1979: 104, pi. 20 figs 5A-V. — Millard 1980: 130. —

Hirohito 1983: 5, fig. 11. — Stepanjants 1985: 137. — Antsulevich

1987: 106. - Gili, Vervoort, and Pages, 1989: 81, fig. lOA. — Cornelius

and Ryland, 1990: 140, fig. 4.— Calder, 1991: 22, fig. 14.— El

Beshbeeshy, 1991: 40, fig. 6. — Ramil and Vervoort, 1992: 90, figs 2 If,

g. — Medel and Vervoort, 2000: 23.-Schuchert, 2001: 85, figs 70A-E.

Halecium (?) tenellum. — Ralph, 1958: 340, figs Ilf, g.

Halecium geniculaturn Nutting, 1899: 744, pi. 63 figs la-d (not

Halecium geniculaturn Norman, 1867 (= Halecium halecinum

(Linnaeus, 1758)).

For full synonymy see Cornelius (1975).

Material examined. Stn 120, TM K2779, infertile colony of many stems

on stem of Eudendrium ?cyathiferum, specimen alcohol-preserved.

Description. Hydrorhiza tubular, undulating, reptant on hydroid

host. Stems to 5 mm high, straggling, given off irregularly from

hydrorhiza; stems monosiphonic, beginning with two or three

deep transverse annulations, branching thereafter mostly alter-

nate in one plane, straight to sympodial or irregularly dichoto-

mous; stem internodes long, thin, cylindrical, variable in

length, perisarc smooth, nodes oblique to transverse, marked by

one to three distinct constrictions in perisarc.

Primary hydrophore given off below node, cylindrical, base

contiguous with or inclined outwards from internode, variable

in length but usually fairly short, hydrotheca moderately deep,

diaphragm distinct, transverse, a slight thickening of hydro-

thecal wall around diaphragm, a faint ring of desmocytes

above; margin wide, strongly everted with recurved rim.

Hydrophores regenerated linearly up to 10 times, each arising

from diaphragm of preceding one, regenerations similar to

primaries but length highly variable; base contracted into

diaphragm of preceding hydrotheca. Perisarc of hydrorhiza and

proximal stem thinning distally, hydrothecal rim very thin.

Colour. White to colourless.

Measurements pm

Hydrorhiza

diameter

68-80

Stem internode

length of proximal internode

1200-1600

length of succeeding internodes

440-1320

diameter at node

60-80

Hydrophore

adcauline length of primary hydrophore

136-232

length of succeeding hydrophores

68-260

diameter of hydrotheca at diaphragm

78-80

depth, margin to diaphragm

24-36

diameter of marginal rim

130-150

Distribution. Near-cosmopolitan in Atlantic, Indian and Pacific

Oceans (if all previous identifications of the species are cor-

rect). Depth range, 0-550 m (Millard, 1975).

record (1046 m) is the deepest for the species.

The present

Figures 15A-E. A-C, Halecium tenellum, stn 120. A, single stem from

colony. B, linear series of hydrophores. C, hydrophores and hydro-

thecae with outrolled rims. D-E. Halecium ovatum, stn 120.

D, linear series of hydrophores. E, secondary hydrophore given off

below primary. Scale bar: A, 1 mm; B, 0.25 mm; C, 0.1 mm;

D, E, 0.5 mm.

166

Jeanette E. Watson

Remarks. The species is small and elegant. The diameter of the

widely flared and outrolled, thin hydrothecal rim is twice that

of the diaphragm. The desmocytes above the diaphragm are

very faint and visible only under high magnification. Some

stems have apical tendrils which revert to stolons.

Although the colonies are not strictly fascicled a few stems

have one or two extra tubes that become free above the base as

independent monosiphonic branches. Because of its small size

and epizoic habit H. tenellum is easily overlooked and can be

identified with certainty only when fertile. The present speci-

mens conform reasonably well with descriptions and dimen-

sions (where given by authors) of H. tenellum.

Halecium ovatum Totton

Fig. 15D, E

Halecium ovatum Totton, 1930: 143, fig. 3. — Vervooit, 1972b; 339,

fig. 1. — Stepanjants, 1979: 103, pi. 20 figs la-g. — Pena Cantero,

1991: 48, pi. 2 figs g, h. — Blanco, 1994: 156. — Pena Cantero and

Garcia Carrascosa, 1995: 12, figs 2G, H. — Pena Cantero and Garcia

Carrascosa, 1999: 212.

Material examined. Stn 120, TM K2780, NMV F91321, sparse infer-

tile colonies on Halecium ralphae, specimen alcohol-preserved.

Description. Colony minute, stolonal; stolons tubular, walls

crumpled, thin. Primary hydrophore seated on a short

apophysis of the stolon, a transverse to weakly oblique node

at base; hydrophore relatively long, cylindrical, smooth,

gradually expanding to hydrotheca.

Secondary and tertiary hydrophores arising in a series with-

out basal node from diaphragm of preceding hydrotheca; suc-

cessive hydrophores progressively shorter. Branching of

hydrophore at right angles from below a hydrotheca common,

secondary hydrophores shorter, with 2 or 3 partial basal con-

strictions. Hydrotheca shallow, expanding smoothly from well

marked diaphragm to wide margin with strongly recurved and

outrolled rim, a clear ring of desmocytes above diaphragm.

Perisarc of hydrorhiza very thin, that of hydrophores

thicker, rim of hydrotheca thin.

Colour. Colourless.

Measurements (pm)

Hydrorhiza

width of stolon 80-104

Hydrophore

length of primary, proximal node to diaphragm 120-496

width, primary proximal node 59-62

length of succeeding hydrophores 142-316

diameter at diaphragm 1 20- 136

depth, margin to diaphragm 32—44

diameter, marginal rim 176-208

Distribution. Antarctic (Stepanjants, 1979; Pena Cantero and

Garcia Carrascosa, 1999).

Remarks. The species is similar to Halecium tenellum for which

it was originally mistaken in samples from Stn. 120. In size and

habit of colony the present specimens most resemble Halecium

ovatum Totton, 1930 redescribed and figured by Vervooit

(1972b). Although the hydrotheca is narrower and shallower

than that of H. ovatum, in the absence of gonosome the

material is assigned to that species.

The stolons of H. tenellum and H. ovatum entwine on the same

substrate and although difficult to differentiate, the two species

can be distinguished by the strictly stolonal habit of H. ovatum,

its broader, undulating and thinner-walled hydrorhizal stolons,

branching of the subsidiary hydrophores from below the pri-

mary hydrophore, the greater overall cauline dimensions, the

less strongly flared and outrolled rim and the ratio of diameter

of hydrothecal rim to width of diaphragm (in H. ovatum 1.5:1,

in H. tenellum 2:1).

Halecium delicatulum Coughtrey

Figures 16A-C

Halecium delicatulum Coaghtiey, 1876b: 299. — Coughtrey, 1876a;

26, pi. 3 figs 4, 5.— Stechow, 1913; 144.— Stechow, 1913: 9, 79.—

Stechow, 1923a: 5.— Bale, 1924: 235.— Ralph, 1958; 334, figs lie,

h-n, 12 a-p. — Pennycuik, 1959; 173. — Yamada, 1959: 31. — Naumov

and Stepanjants, 1962: 94, figs 16, 17. — Rees and Thursfield, 1965;

106.— Millard, 1966; 464, fig. lOL.— Ralph, 1966; 158.— Millai'd,

1968: 253, 256. — Day et al., 1970: 12. — Blanco and Bellusci de

Miralles, 1972: 7, figs 3-5. — Naumov and Stepanjants, 1972: 34,

52. — Stepanjants, 1972: 72. — Vervoort, 1972a: 27, figs 4, 5. —

Vervoort, 1972b: 341, fig. 2a.— Watson, 1973: 166.— Leloup, 1974:

10.— Millard, 1975; 145, figs 47F-L.— Watson, 1975: 159.— Millard,

1977a: 7, figs 1C, D.— Millard, 1978: 193.— Stepanjants, 1979: 105,

pi. 20 figs 4A-V.— Watson, 1979: 234.— Hirohito, 1983: 5, 11.— Rho

and Park, 1983: 41, pi. 2 figs 1-3. — Aguirrezabalaga et al., 1984:

90. — Rees and Vervoort, 1987: 25, fig. 5. — Aguirrezabalaga et al.,

1988: 222. — Ramil et al., 1988; 72, fig. 2; Gili, Vervoort, and Pages,

1989: 78, fig. 7B. — ^Altuna and Garcfa Carrascosa, 1990: 54. —

Genzano, 1990: 38, figs 2-5. — ^E1 Beshbeeshy, 1991: 32, figs 4a, b.- —

Roca et al., 1991 : 70: 14. — Genzano and Zamponi, 1992: 40, fig. 17. —

Park, 1992: 286. — Ramil and Vervoort, 1992: 82, figs 20a-c. — Branch

and Williams, 1993; 11. — Genzano, 1994; 5. — ^Watson, 1994: 66. —

Altuna Prados, 1995; 54. — Bouillon et al., 1995: 45. — Hirohito 1995:

20, figs 5a-c, pi. 1, fig. C. — Park, 1995: lO.-Genzano, 1996: 290. —

Medel and Vervoort, 2000: 12.

Halecium flexile Allman, 1888: 11, pi. 5 figs 2, 2a.

Halecium gracile Bale, 1888: 759, pi. 14 figs 1-3

Halecium parvulum Bale, 1888: 760, pi. 14 figs 4, 5.

Material examined. Stn 122, NMV F91343, NMV F91344, malinol-

mounted microslides, colony of three damaged infertile stems, on

Eudendrium.

Description. Two stems simple, stolonal, the other, the tallest

16 mm high, lightly fascicled. Tubes of fascicled stem running

about two thirds distance up stem; stem thereafter mono-

siphonic with a few alternate branches standing out stiffly

almost at right angles to axis, smaller branchlets arising at inter-

vals along stem. Monosiphonic branch intemodes long, nodes

oblique to almost transverse, sloping almost parallel to primary

hydrophore, weakly to deeply constricted, deeper nodes with

tumescence above and below. Hydrophores alternate, distal on

internode, a smooth, outwardly directed continuation of

intemode.

Primary hydrophores variable in length, cylindrical, expand-

ing a little to below hydrotheca, perisarc smooth; some older

hydrophores on lower stem region deeply corrugated; a sec-

ondary hydrophore sometimes given off from primary; linear

series of up to eight hydrophores common, each hydrophore

Deep-water hydroids from Macquarie Island

167

Figures 16A-C. Halecium delicatulum, stn 122. A, distal part of

colony. B, monosiphonic branch. C, hydrophores, enlarged. Scale bar;

A, 5 mm; B, 0.5 mm; C, 0.3 mm.

arising from diaphragm of preceding one, usually a strong basal

constriction present, successive hydrophores usually becoming

progressively shorter. Hydrotheca moderately deep, expanding

smoothly to a wide, strongly everted margin with outrolled rim;

diaphragm very strong, transverse, some concave with central

hydropore, usually a row of desmocytes above.

Perisarc of polysiphonic tubes of stem quite thick, thinning

along monosiphonic branches and hydrophore, becoming thin

at hydrothecal margin.

Colour. Clear white (preserved material).

Measurements (pm)

Branch

length of intemode 549-765

width at node 109-152

Hydrophore

adcauline length of primary, to diaphragm 78-117

length of succeeding hydrophores, base todiaphragm 78-312

diameter at diaphragm 137-164

depth, margin to diaphragm 32-59

diameter, marginal rim 220-257

Distribution. Circumglobal in tropical, subtropical and boreal

waters (Vervoort and Watson, 2003).

Remarks. The rather stiff mode of branching in the largest

colony does not precisely accord with the usually rather lax

habit of H. delicatulum (pers. obs.) but may be an artefact of

preservation. Although the hydrothecae are a little shallower

than is normal for H. delicatulum I have no doubt the present

material is referrable to that species.

Halecium sp.

Figures 17A-E

Material examined. Stn 122, NMV F91328, fragmentary remaining

upper branches of a large fertile colony, specimen alcohol-preserved.

Stn 90, TM K2791, specimen alcohol-preserved, NMV F91345,

malinol-mounted microslide, from lower stem and denuded branches

of very large colony broken off from base.

Description. Stem (or lower branch) from Stn 122, 5 mm wide

at base and 150 mm high, rigid, irregularly branched; branches

heavily fascicled, tubes running almost to tips of branches.

Ultimate branches (hydrocladia) monosiphonic, irregularly

alternate, short, hydrocladium issuing from inside a hydrotheca

on peripheral tube of stem or polysiphonic tube of branch,

perisarc of stem and proximal branches thick, thinner on

hydrocladia. Proximal hydrocladial internode with 1 to 4 deep

transverse septa, internodes thereafter short, more or less

cylindrical, walls smooth to broadly undulated, nodes opposed,

tilted away from hydrophore, deeply incised, internode tumid

above and below node.

Hydrophores alternate, occupying distal half of internode,

abcauline wall sloping smoothly outwards from axis at c. 30°,

adcauline wall variable in length, adnate to below hydrotheca,

free wall short. Hydrotheca shallow, free of internode, opposite

to or just above node, tilted away from intemodal axis at

c. 110°, expanding slightly to a weakly everted rim; eversion

more pronounced on adcauline than abcauline side; in frontal

view hydrothecal margin slightly ovoid; diaphragm distinct,

transverse to saucer-shaped with central circular hydropore, a

circle of inward-facing thorn-shaped desmocytes above

diaphragm. Hydranth with c. 16 tentacles, none well preserved.

Gonothecae arising without pedicel on proximal part of

internode opposite a hydrophore; a few immature or broken

gonothecae present, minute to small, base subspherical,

perisarc very thin.

Colour. Colony from Stn 122 honey brown, fading to

white on monosiphonic branches. Colony from Stn 90 pale

yellowish-green.

168

Jeanette E. Watson

Figures 17A-E. Halecium spec., stn 90. A, colony. B, polysiphonic

stem with monosiphonic branches. C, monosiphonic branch with

hydrophores. D, young gonotheca erupting from stem. E, more

advanced gonotheca, distal end damaged. Scale bar: A, 50 mm;

B, 1 mm; C, 0.2 mm; D, E, 0.1 mm.

Measurements (pm)

Monosiphonic branch

length of proximal internode 120-440

length of succeeding internodes 600-820

width at node 160-280

Hydrophore

length of adnate adcauline wall 304-416

length of free adcauline wall 56-100

Hydrotheca

diameter at diaphragm 232-264

depth margin to diaphragm 48-60

diameter at margin 304-320

Gonotheca

width of immature gonotheca 1 20

Remarks. The heavily fascicled branches are woody and very

brittle. There is little tendency to secondary branching and there

are few regenerated hydrophores in the colonies. Although

most hydrophores are oppositely arranged on the hydro-

cladium, there is a tendency to frontal displacement on some

branches.

The few small gonothecae present were noted only during

detailed examination of the material. They may be female, but

the very thin perisarc is so collapsed and tom that the shape of

the mature gonotheca could not be reconstmcted or the sex

determined. Species with strongly fascicled colonies con-

sidered were the near-cosmopolitan Halecium beanii

(Johnston, 1838), H. luteum Watson, 1975 from Tasmania and

H. jaederholmi Vervoort, 1972b, known from Antarctic and

subantarctic waters. However, structure and dimensions of the

hydrotheca and shape of the internode of these species all dif-

fer from the present specimen. As the material is fragmentary it

is not to assigned to species.

Sertulariidae Lamouroux, 1812

Calamphora AWmm, 1888

Calamphora quadrispinosa sp. nov.

Figures 18A-E

Material examined. Stn 94, NMV E91325, holotype, specimen

alcohol-preserved, NMV E91346 malinol- mounted microslide from

holotype, large colony on Symplectoscyphus. Stn 44, TM K2783,

paratype, small colony on Eudendrium, specimen alcohol-preserved.

Stn 120, NMV E91324, paratype, colony on Symplectoscyphus paul-

ensis, specimen alcohol-preserved. Stn 130, TM K2784, paratype, on

dead coral fragment, specimen alcohol-preserved. Stn 46, TM K2785,

small colony on Eudendrium, specimen alcohol-preserved.

Eigures 18A-E. Calamphora quadrispinosa sp. nov., stn 94. A, three

stems, part of colony. B, stem, enlarged. C, hydrotheca with intact

flanged operculum. D, hydrotheca with ragged opercular valves

after eruption of the hydranth; ligaments connecting hydranth to

wall of hydrotheca. E, gonotheca. Scale bar; A, 10 mm; B, 1 mm;

C, D, E, 0.5 mm.

Deep-water hydroids from Macquarie Island

169

Description (ofholotype and paratypes). Stolonal colonies tan-

gled amongst host hydroids. Stolons simple, tubular, smooth,

perisarc moderately thick, smooth to undulated, firmly attached

to host, becoming free at intervals as monosiphonic stems,

sometimes sparingly branched.

Hydrothecae pedieellate, arising singly, more or less regul-

arly along one side of stolon or branch; hydrothecae upright or

inclined at various angles, barrel-shaped or slightly asym-

metrical (depending upon angle of view), body narrowing a

little below margin, walls smooth to weakly undulated, usually

in proximal third; body narrowing into a tubular pedicel;

diaphragm thick, transverse to slightly oblique, hydropore

central, circular, with short upturned collar. Margin transverse

to axis, quadrate, distinctly everted with four broad, sharply

pointed cusps separated by low embayments. Operculum of 4

thin triangular valves; in immature hydrothecae valves meet in

a low upturned flange decreasing in height from hydrothecal

margin to centre; in mature hydrotheca remnants of valves

meeting in a central, ragged, upturned tuft. Perisarc of

hydrotheca moderately thick proximally, thinning distally.

Hydranth (preserved material), columnar, with c. 16

tentacles and a wide, annular hypostome; no diverticulum or

annular fold but a long bifid ligament issuing from below

tentacle ring, joining hydrothecal wall in distal third.

Gonothecae arising from stolon, usually beside a hydro-

theca; pedicel short, thick; mature gonotheca barrel- shaped,

body with six to nine deep flanges, deepest in distal third, shal-

lower proximally; aperture a dome of tissue surrounded by four

long, equidistant, more or less inwardly curved spines. Sex of

gonophores could not be determined.

Colour. White (preserved material); may have been pale

yellow in life.

Measurements (pm)

Stolon-stem

diameter 192-216

distance between hydrothecal pedicels 1060-1040

Hydrotheca

length pedicel, adcauline side 64-224

length diaphragm to margin 1100-1280

maximum width 5 06-5 6 1

diameter at diaphragm 184-216

diameter at margin 440—480

height of marginal cusp 70-80

Gonotheca

length of pedicel 1 20-1 7 6

distal width of pedicel 160-200

maximum diameter 520- 640

depth of ridges 68-100

width across margin 208-240

height of marginal cusps 76-120

Etymology. Named for the four claw-like spines of the

gonotheca.

Remarks. There are no nodes in either the hydrorhizal or free

stolons, only infrequent constrictions marking probable sites of

breakage and repair. The free stolons (branches) are predomin-

antly monosiphonic with some tendency to polysiphony by

fusion of two stolons over short distances. The branches are

usually loosely curved, and this together with adherence of the

stolons to the substrate results in considerable tangling around

the stems and branches of the hydroid host. The hydrothecae

are usually single but occasionally two may be given off from

opposite sides of the branch. Although the hydropore is cen-

trally located, the base of the hydranth sometimes appears to be

attached beside, rather than through the hydropore; a ring of

large granules (seen in partially cleared specimens) marks the

site of attachment. The operculum is retained after eruption of

the hydranth and becomes torn into apical tufts presumably

from repeated movement of the hydranth. This difference

between the flanged operculum of immature hydrothecae and

the fragmented tufts of mature hydrothecae is striking, and if

the two forms were not present on the same colony the

hydrothecae could easily be mistaken for two different species.

The bifid ligaments supporting the extended hydranth are

visible only when the hydranth is retracted. Nematocysts present

in the tentacles and coenosarc of the stolons could not be iden-

tified. The colonies are abundantly fertile, the claw-like mar-

ginal spines of the gonotheca being characteristic. As no ova

were seen in the gonophores the sex is presumed to be male.

Genera considered were: Sertularella Gray, 1848,

Thyroscyphus Allman, 1877, Symmetroscyphus Calder, 1986

and Calamphora Allman, 1888. Sertularella was rejected as the

specimen has no clear abcauline diverticulum; as the material is

stolonal Thyroscyphus was rejected; Symmetroscyphus was also

rejected as the hydrotheca of that species is symmetrical.

Calamphora is said to possess a diverticulum situated on the

adhydrorhizal side of the hydranth (Millard, 1975) which in a

stolonal colony is assumed to be the adcauline side. No evi-

dence of such a structure was found in the present material; it

is possible however, that from some angles of view, strands of

ligament in inadequate material could be mistaken for divertic-

ula.

Although Vervoort (1968) considered Calamphora to be

inseparable from pedicellate Sertularella it nevertheless seems

useful to retain the genus for exclusively pedicellate sertulariid

species.

Calamphora quadrispinosa is the most abundant species in

the collection.

Staurotheca Allman, 1888

Staurotheca vanhoeffeni (Pena Cantero, Garcia Carrascosa

and Vervoort)

Figures 19A-E

Staurotheca vanhoeffeni Pena Cantero et. al., 1996: 1-10, figs

1-3. — Pena Cantero et al., 1997: 373, fig. 12.

Dictyocladiurn affine. — Vanhoffen, 1910: 331, figures 44a-e.

Thuiaria affinis. — Stepanjants, 1979; 95, pi. 18 figs lA, B, pi. 24

figs D-F.

Selaginopsis vanhoeffeni. — Pena Cantero and Garcia Carrascosa,

1994: 121, figs 3j-n.

Material examined. Stn 44, NMV F91322, several sparingly fertile

branch fragments, specimen alcohol-preseived. Stn 122, TM K2781,

specimen alcohol-preserved, NMV F91347, malinol mounted

microslide from i nf ertile colony detached from substrate

Description. Colony stiff and woody, stem 2 mm thick at base,

detached from hydrorhiza. Stem sparsely branched in one

170

Jeanette E. Watson

Figures 19A-E. Staurotheca affinis stn 44. A, part of colony. B, branch

internode. C, D, replicated hydrothecal margins. E, female gonotheca.

Scale bar: A, 50 mm; B, E, 1 mm; C, D, 0.5 mm.

plane, primary branches rather geniculate, pointing upwards,

straight or curved, a few second order branches present. Stem

and lower branches fascicled, tubes consisting of a sheath of

more or less concentric layers of perisarc surrounding stem;

sheath thick proximally, layers becoming fewer but lumpy

along branches, completely enclosing branches but branches

visible through transparent outer perisarc. Hydrocladia

(branches) long, straight, internodes variable in length, 1-10

groups of hydrothecae on internode, nodes oblique to

transverse, deeply constricted.

Hydrothecae flask-shaped, immersed in internode, walls not

in contact, typically 3 but sometimes 4 (on older hydrocladia)

arranged in a verticil around internode, base of each slightly

upwardly displaced with respect to others; a proximal athecate

section of internode below basal-most hydrotheca. Adcauline

hydrothecal wall straight to weakly convexly curved proxi-

mally, curvature more convex in distal third, abcauline wall

almost straight to faintly concave with outward flexure below

marginal rim; base of hydrotheca flat to weakly concave, a

thick knot of perisarc at base of adcauline wall. Margin of

hydrotheca circular, a tilted slightly upwards, edentulate, not

everted, protruding just clear of internode, rim often ragged and

produced into a short collar by numerous fine replications;

operculum a thin, low dome.

Female gonothecae given off branch above a hydrotheca;

pedicel short, wide, slightly bent, merging into gonotheca;

body of gonotheca heart-shaped, widening from base to

rounded shoulder, walls smooth to faintly undulated, distal end

a platform with central wide orifice suiTounded by a low collar

and a pair of wing-shaped lobes, side of collar slightly out-

wardly turned and facing adcaudally. Perisarc very thick. No

male gonothecae found.

Colour. Colony uniformly dark brown

Measurements (pm)

Branch (hydrocladium)

length of internode 1360-6000

width at node 320-400

length of infrathecal internode 360-840

width across hydrothecal pair, margin to margin 840-1100

Hydrotheca

length (diagonal) across adcauline wall

900-1000

length of abcauline wall

780-1000

maximum width

312-384

width across floor

344-400

diameter at margin

280-336

Gonotheca

length, base (excluding pedicel) to shoulder

1360-1460

width across shoulder

1360-1420

length of pedicel

140-200

width of pedicel at base

200-300

height of apertural collar

360-480

diameter of collar

700-900

Distribution. Circumantarctic (Pena Cantero et al., 1997).

Remarks. The stiff woody colony of the most intact specimen

matches previous descriptions of Staurotheca vanhoejfeni.

Dimensions of the two undamaged female gonothecae from stn

44 fit those given by Vanhoffen (1910). Stepanjants (1979) and

Pena Cantero et al., (1997) described and figured male

gonothecae: none are present in the Macquarie Island material.

None of these authors mention the distinct knot of perisarc at

the base of the adcauline hydrothecal wall. The fascicular tubes

envelop the proximal part of the stem in contorted, more or less

concentric layers of thickened, tough perisarc.

Symplectoscyphus paulensis Stechow

Figures 20A-D

Symplectoscyphus paulensis Stechow, 1923a: 8. — Stechow, 1923b:

172.— Stechow, 1925: 467, fig. 28.— Millard, 1967: 183, figs 4G, H.—

Vervoort, 1972b: 180, figs 60b, 61.— Millard, 1975: 317, figs

102A-C.— Millard, 1977b: 107.— Millard, 1978: 199.— Stepan-jants,

1979: 71, pi. 17 fig. 2.— Hirohito, 1983: 51, fig. 24.— Veivoort, 1993:

263, figs 63a-d, 65a. — Blanco, 1994: 154. — Bouillon et al., 1995: 74.

Material examined. Stn 120, TM K2790, NMV F91329, specimens

alcohol-preseived; NMV F91348, malinol-mounted microslide, thi'ee

complete infertile colonies, the tallest 100 mm high and fragments of

others, possibly branches shed from these colonies. Colonies heavily

overgrown by Calamphora quadrispinosa.

Description. Colonies irregularly and profusely branched more

or less in one plane. Hydrorhiza a tuft of stolons detached from

substrate. Stem of tallest colony 3 mm wide above hydrorhiza,

heavily fascicled, stolons growing upwards to become poly-

siphonic tubes of stem. Branches (hydrocladia) to 30 mm long,

slender, polysiphonic tubes often extending a short distance

along branch. Branches thereafter monosiphonic (hydrocladia),

branched subdichotomously several times at c. 120°, a

hydrotheca in each dichotomy.

Hydrocladial internodes long, cylindrical, widening a little

below hydrotheca, perisarc smooth, sometimes undulated,

internode widening distally; nodes distinct, oblique to almost

transverse, marked by a narrowing of perisarc, frequently a

tumescence above or below node; a node on each side of

hydrotheca in dichotomy.

Hydrothecae alternate, distant, given off almost in one plane

just below node at an angle of c. 80° to internodal axis, almost

cylindrical, narrowing very slightly to margin, walls smooth,

adcauline wall gently convex, adnate adcauline wall short.

Deep-water hydroids from Macquarie Island

171

Figures 20A-D. Symplectoscyphus paulensis, stn 120. A, colony.

B, distal monosiphonic branches. C, part of distal branch. D, rim of

hydrotheca showing obtuse cusps. Scale bar: A, 50 mm; B, 10 mm;

C, 1 mm; D, 0.5 nun.

becoming free opposite or just below node, free wall convex to

almost straight, but less curved than adnate part, at least twice

length of adnate part; abcauline wall slightly concave, curving

smoothly outward, contiguous with internode. Hydrotheca

widest at junction of adnate and free adcauline wall, narrowing

a little to margin, floor narrow, flat, a minor thickening of

perisarc at base of adcauline wall, a triangular foramen and

thinning of perisarc in intemode beneath floor.

Margin with 3 low, equidistant cusps separated by broad,

rather shallow embay ments; margin often with numerous fine

replications, operculum of 3 triangular valves. Perisarc of

hydrotheca thin, slightly thicker at marginal replications,

operculum thin.

Hydranth with c. 24 tentacles, a strand of tissue attaching hyd-

ranth to hydrotheca about one-third distance up adcauhne wall.

Colour. Pale yellow-brown.

Measurements (pm)

Hydrocladium

length of internode

1000-1440

diameter at node

120-176

Mrotheca

length of free adcauline wall

792-840

length of adnate adcauline wall

328-392

length of abcauline wall

880-960

width at floor

184-280

width at margin

400-480

Distribution. A moderately deep-water species from 680 m in

the southern Indian Ocean (Stechow, 1923b), 440 m in the

south-west Indian Ocean, 347 m off Mozambique (Millard,

1967), 424-428 m on Verna Seamount (Vervoort, 1972a) and

399-500 m in Antarctica (Stepanjants, 1979).

Remarks. Although flexuous, the apical branches are rather

brittle and easily broken. The hydrothecae are mostly arranged

in one plane, but on some hydrocladia there is a tendency to

face frontally. The small, thin marginal replications are slightly

everted and are probably remnants of opercular attachments.

The hydrothecae in the branch dichotomies do not differ in size

or shape from those on the internodes.

The colonies are so heavily overgrown by Calamphora

quadrispinosa that it is difficult to distinguish the stolons of

that species from the polysiphonic tubes of S. paulensis. The

syntype of S. paulensis is strongly polysiphonic (Vervoort,

1993) while monosiphonic hydrocladia of S. paulensis are sim-

ilar to S. bathyalis Vervoort, 1972. S. bathyalis was rejected

because of its weakly expanding hydrothecae.

The present specimens are the largest colonies of S. paulen-

sis ever recorded. Previous records are of small polysiphonic

colonies or monosiphonic fragments.

Symplectoscyphus tuba Totton

Figures 21A-D

Symplectoscyphus tuba Totton, 1930: 186, figs 37a, b. — Ralph,

1961: 816, figs 18f, g. — ^Leloup, 1974: 42, fig. 41. — Vervoort, 1993:

272, figs 67a-^.

Sertularella tuba. — Stepanjants, 1979: 76, pi. 1 fig. 4.

Material examined. Stn 94, NMV F91323, specimen alcohol- pre-

served, NMV F 91349 malinol mounted microslide, fragments of

colony detached from substrate. Stn 119, TM K2793, small fertile

colony on primnoid gorgonian stem, specimen alcohol-preserved. Stn

120, TM K2794 many broken stems, longest intact stem 40 mm long,

specimen alcohol-preserved. Stn 122, TM K2782, small infertile

colony on Eudendrium, specimen alcohol-preserved.

Description. Hydrorhiza composed of sparse stolonal

tubes. Stems monosiphonic, of same diameter as stolons.

Figures 21A-D. Symplectoscyphus tuba stn 120. A, single stem

from colony. B, branch with axillar hydrotheca. C, hydrotheca.

D, gonotheca. Scale bar: A, 10 mm; B, 1 mm; C, D, 0.5 mm.

172

Jeanette E. Watson

lower stem rather lax, almost straight proximally, narrowing a

little distally. Stems branched alternately, branches widely

spaced, usually simple but sometimes rebranched once or

twice.

Primary branches up to 12 mm long, directed upward at an

acute angle to stem; secondary branching (hydrocladia) if pres-

ent, pseudodichotomous, forking from below a hydrotheca.

Lower branch internodes sub-sympodial, this structure becom-

ing more pronounced distally along hydrocladia. Some hydro-

cladia terminating in tendrils that rejoin other hydrocladia to

form a loose meshwork. Intemodes variable in length, slender,

widening distally to accommodate hydrotheca, perisarc

smooth, node a weak oblique constriction in perisarc sloping

away from hydrotheca.

Hydrothecae alternate, distal on internode, short, con-

tracting from base to margin, but sometimes almost tubular;

directed upwards at an angle of 35-45° to internodal axis,

abcauline wall straight to weakly concave, smoothly contig-

uous with internode, adcauline wall convex, adnate adcauline

wall slightly longer than free wall, almost straight, wall becom-

ing free at node, curvature greater than adnate wall, imparting

a distinct angularity to wall. Floor of hydrotheca narrow, flat, a

small subcircular foramen in perisarc below. Margin with 3

cusps, adcauline the longest, slightly everted, with 2 abcauline

laterals, these less pronounced, rounded; some hydrothecal

margins slightly thickened, many replicated. Hydranths with c.

12-16 short tentacles, not well preserved.

Gonothecae abundant, arising on a very short, bent pedicel

inserted close beneath hydrothecae on stem and hydrocladia;

gonotheca adpressed to hydrocladium, facing distally, ovoid

with 8-10 deep, upturned flanges, proximal 2 or 3 shallowest,

distalmost flange a collar surrounding orifice; gonothecal

wall between flanges deeply concave and strongly vertically

striated. Orifice central, wide, trumpet- shaped, upright.

Remains of gonophores present in many gonotheca but

sex indeterminate.

Colour. Colony colourless, transparent; gonophores may

have been pink in life.

Measurements (pm)

Stem

diameter

Internode length

780-1260

diameter at node

88-120

Hydrotheca

length of abcauline wall

416-440

length of adnate adcauline wall

296-360

length of free adcauline wall including replications

248-312

width across margin (lateral view)

192-216

width across floor

96-120

Gonotheca

total length including pedicel

1120-1200

maximum diameter including flanges

760-800

diameter of orifice

240-280

length of tube

160-192

length of pedicel

80-112

Distribution. Northern New Zealand, 183 m (Totton, 1930);

Antarctica, 145^10 m (Stepanjants, 1979); Chile, 30-60 m

(Leloup, 1974); New Caledonia, 650-680 m

(Vervoort,

1993).

Remarks. Although the width of the hydrothecal margin and

diameter of the gonotheca are somewhat greater than measure-

ments given by Totton (1930), Ralph (1961a) and Vervoort

(1993) for Symplectoscyphus tuba I have no doubt that the

present material is referrable to that species. It is a rather deli-

cate species, of which only small colonies or fragments of

colonies have been previously reported. Its long, slender inter-

nodes, widely spaced hydrothecae and ovoid, deeply ridged

gonothecae are characteristic.

Thyroscyphoides Naumov, 1955

Thyroscyphoides sympodialis sp. nov.

Figures 22A-D

Material examined. Stn 44, NMV F91330, holotype, specimen alco-

hol-preserved, NMV F91350, malinol-mounted irdcroslide from holo-

type colony; 20 infertile stems on hydrorhiza of Eudendrium.

Description (of holotype). Hydrorhiza tubular, rather contorted,

reptant on substrate. Stems simple, to 12 mm high, proximal

part of stem of same diameter as stolon; basal-most part of stem

with 2 to 4 broad, indefinite annulations, stem thereafter

sympodial, sometimes a side branch given off from lower

stem. Internodes long, smooth, tubular, becoming shorter along

stem, widening distally to below hydrotheca; nodes usually

with 1 or 2 deep, oblique constrictions almost parallel with

adcauline wall of hydrotheca, a slight tumescence in perisarc

above node, sometimes 1 or 2 additional constrictions along

intemode.

Hydrothecae alternate, arising just below node, body tubu-

lar, wide, most sessile but some pedicellate; in sessile

hydrothecae abcauline wall smoothly contiguous with outward

bend of internode, free adcauline wall smooth, weakly convex

to almost straight, a short length of wall adnate to intemode;

floor of hydrotheca flat, saucer- shaped or funnel-shaped

(depending upon angle of view), hydropore slightly offset, fair-

ly wide, a row of desmocytes above marking place of attach-

ment of hydranth. Pedicellate hydrothecae arising from floor of

a broken hydrotheca; pedicels of variable length, smooth to

cormgated with some internal thickenings; floor of hydrothecae

rounded to sinusoidal. In both forms, hydrothecal margin

almost circular (anterior view) with 4 sharply pointed triangu-

lar cusps separated by deep embay ments; operculum of 4 thin

pyramidal valves. Hydranth without diverticulum, but with a

distinct annular fold; hydranth with c. 20 tentacles.

Perisarc of lower stems comparatively thick, thinning

apically along internodes, hydrothecal margin and operculum

extremely thin and fragile.

Colour. Transparent, colourless

Measurements (pm)

Stem internode

length

diameter at node

Hydrotheca

length of adnate adcauline wall

length of free adcauline wall

length of abcauline wall (to marginal embayment) 336-520

width at margin 248-296

height of marginal cusps 120-184

width of floor 120-136

1000-1640

80-96

168-200

480-584

Deep-water hydroids from Macquarie Island

173

Figures 22A-D. Thyroscyphoides sympodialis sp. nov., stn 44.

A, single stem from colony. B, sessile hydrotheca, C, pedicellate

hydrotheca. D, anterior view of hydrothecal operculum. Scale bar:

A, 1 mm; B, C, D, 0.5 mm.

Distribution. This is the second record of the genus. T. biformis

is known from deep water in the northern Pacific Kurile Islands

(Naumov, 1955).

Etymology. Refers to the strongly sympodial habit.

Remarks. The colonies are almost flaccid out of fluid and the

thin, almost transparent perisarc of the hydrothecal margin and

operculae are so crushed that few remain intact.

Sessile and pedicellate hydrothecae on the same stem and a

hydranth lacking a diverticulum places the species in

Thyroscyphoides Naumov, 1955. The only other known species

of the genus, T. biformis Naumov, 1955, differs from T. sympo-

dialis in its alternate hydrothecae and its strongly sympodial

habit.

Aglaopheniidae L. Agassiz, 1862

Gymnangium Hincks, 1874

Gymnangium japonicum Watson and Vervoort

Figures 23 A, B

Gymnangium japonicum Watson and Vervoort, 2001: 182, figs

15a-g.

Halicetta sp. — Hirohito, 1995: 293, figs 103e-g, pi. 13 fig. D.

Material examined. Stn 122, TM K2789, specimen alcohol-preserved,

NMV F91351, malinol-mounted microslide from same colony; broken

infertile stem 50 mm long, detached from substrate.

Distribution. Originally recorded (as Halicetta sp.) from a

depth of 250 m from Japan (Hirohito, 1995), from 750-900 m

south-east of Tasmania (Watson and Vervoort, 2001).

Remarks. The material conforms exactly with the description

and dimensions of Gymnangium japonicum Watson and

Vervoort, 2001 reported from deep water seamounts south-east

of Tasmania. The stem is golden brown, hy drocladia yellowish-

green.

Figures 23A-C. Gymnangium japonicum, stn 122. A, colony.

B, hydrocladial hydrothecae. C, cauline internodes with tubular nema-

tocysts (after Watson and Vervoort 2001). A, 20 mm; B, C, 0.5 mm.

Campanulariidae Johnston, 1837

Tulpa Stechow, 1921

Tulpa diverticula Totton

Figures 24A-D

Tulpa diverticula Totton, 1930: 145, fig. 5.— Ralph, 1957: 844, fig.

7.— Millard, 1977a: 20, figs 5G-H.— Stepanjants, 1979: 35, pi. 6 fig

2. — Gravier-Bonnet, 1979: 33. — Bouillon et al., 1995: 86.

Campanularia diverticula Naumov and Stepanjants, 1962: 72. —

Dawson, 1992: 13.

Material examined. Stn 44, TM K2786, specimen alcohol-preserved,

infertile colony on Eudendrium and other hydroids. Stn 90, TM

174

Jeanette E. Watson

Figures 24A-G. A-D, Tulpa diverticula, stn 94. A, stem from colony.

B, hydrotheca, C, rim of hydrotheca, enlarged, D, base of hydrotheca,

enlarged. E-G, Campanularia hicksoni, stn 44. E, hydrotheca.

F, base of hydrotheca showing thickening and diaphragm. G, rounded

marginal cusps. Scale bar: A, 10 mm; B, 1 mm; C, D, 0.5 mm;

E, F, G, 0.5 mm.

K2788, specimen alcohol-preserved, sparse infertile colony on

another hydroid. Stn 94, NMV F91326, specimen alcohol-preserved,

NMV F91352 malinol-mounted microslide from same colony; infertile

colony detached from substrate. Stn 119, TM K2787, specimen

alcohol-preserved, sparse infertile colony on another hydroid. Stn 120,

NMV F91327, sparse infertile colony, specimen alcohol-preserved.

Description. Colony stolonal, hydrorhiza tubular, perisarc

smooth and thick with close, fine, vertical internal striae.

Hydrothecal pedicels tubular, of variable length, given off more

or less regularly, usually from same side of stolon; pedicel end-

ing in a weak constriction below hydrotheca, sometimes 1 or 2

regeneration joints along length; perisarc thinner than on

hydrorhiza.

Hydrotheca large, deeply cylindrical to weakly bell-shaped,

base rounded with transverse diaphragm with central

hydropore, wall thickened below diaphragm, thinning above.

Hydrotheca gracefully everted below circular margin, rim with

12-14 shallow crenulations sometimes finely replicated,

usually six to eight faint pleats extending partially or com-

pletely down into hydrotheca from embayments between

crenulations. Hydranth large, with 20-24 tentacles.

Colour. Preserved specimens colourless.

Measurements (pm)

Hydrorhiza

diameter

280-380

Pedicel

length

1900-4900

diameter

160-300

Hydrotheca

diameter at diaphragm

220-340

length, diaphragm to margin

2320-2820

diameter of margin

820-1000

Distribution. North and South Islands of New Zealand (Ralph,

1957) and Antarctic (Stepanjants,1979).

Remarks. The straggling hydrorhiza readily detaches from the

substrate. The thin hydrotheca is easily crushed along the pleat-

lines during mounting. Many pedicels are segmented with up to

five regenerations, apparently repair after breakage. The

species is a very common epizooite on other hydroids in the

collection.

Campanularia hicksoni Totton

Figures 24E-G

Campanularia hicksoni Totton, 1930: 148, figs 7a-e. — Briggs,

1938: 15. — Rees and Thursfield, 1965: 90, 195. — Blanco and Bellusci

de Miralles, 1972, 145: 10, figs 6-19. — ^Naumov and Stepanjants,

1972: 34, 37. — Stepanjants, 1972: 67, fig. 12. — Stepanjants, 1979: 29,

pi. 5 fig. 1.— Blanco, 1984: 18, pi. 13 figs 29-31.— El Beshbeeshy,

1991: 94, fig. 22a.— Blanco, 1994: 159.

Campaularia laevis Hickson and Gravely, 1907: 25, pi. 4 fig. 26. —

Ritchie, 1913: 19, fig. 5.— Vanhbffen, 1910: 298, fig. 18 (not

Campaularia laevis Hartlaub, 1905: 565, fig. PI).

Material examined. Stn 44, TM K2792, NMV F91331, infertile

colonies on Tulpa diverticula and Eudendrium, specimens alcohol-

preserved.

Description. Colonies comprising many pedicellate hydrothe-

cae arising from hydrorhiza overrunning stems and hydrorhizae

of other hydroids. Stolons tubular, sometimes contorted, peri-

sarc thick. Pedicels simple, unbranched, variable in length,

long, tubular, of same diameter as stolon; most pedicels with 3

or 4 obscure proximal annulations, pedicels thereafter smooth,

sometimes with a few groups of undulations or joints marking

site of regeneration. Pedicel ending in a slightly expanded

distal shoulder, a flattened spherule between stem and basal

chamber of hydrotheca.

Hydrotheca deeply campanulate, walls expanding smoothly

from base to margin, basal chamber narrow, enclosed by outer

walls, diaphragm marked by an annular thickening of wall;

margin circular, crenulate, with 8-10 apically flattened cusps,

embayments between U-shaped, an almost exact reverse image

of cusps. Hydrothecal margins often with 2 or 3 widely

separated regenerations. Hydranth with c. 16-18 tentacles.

Perisarc of stems fairly thick, thinning distally along

hydrothecal body to margin.

Colour Colonies transparent and colourless to white.

Measurements (pm)

Pedicel

length 1200-3800

diameter 64-72

diameter of spherule 52-64

Deep-water hydroids from Macquarie Island

175

Hydro thee a

depth of basal chamber 40-64

length, diaphragm to margin 640-664

diameter at margin 320-440

height of marginal cusp 40

width of marginal embayment 48-64

Distribution. Antarctic (Totton, 1930; Briggs, 1938;

Stepanjants, 1979).

Remarks. Many hydrothecal margins are damaged and most of

the undamaged ones fade in stained mounts The few faint

pleats extending downwards from the margin in many

hydrothecae may result from collapse in mountant.

Dimensions of the present specimens agree fairly well with

those of the “short race” of Campanularia hicksoni recorded

from a depth of 92 m from McMurdo Sound (Totton, 1930) but

with the following minor differences: (i) the hydrothecae of the

present specimens are a little more campanulate than shown in

Totton’s figure; (ii) overall length of the hydrotheca of the

present specimens is at the lower range of Totton’s specimens;

and (iii) the marginal diameter of the present specimens falls

between Totton’s “short race” and those he considered normal

for specimens from Cape Adare. Totton’s specimens had a vari-

able number of marginal crenulations (9-20); the number of

crenulations on the present specimens are at the lower end of

this range.

Acknowledgments

I thank the CSIRO for making the collection available for

examination and the CSIRO Science and Industry Endowment

Fund for financial assistance with the study. Thanks are due to

Dr Antonio Marques of Universidade de Sao Paulo for advice

concerning Antarctic species of Eudendrium, and to the

Australian Museum, Sydney, and the Museum fiir Naturkunde,

Humbold Universitat zu Berlin, for loan of comparison and

type material. I particularly thank Dr W. Vervoort of the

National Museum of Natural History, Leiden, The Netherlands

for helpful advice.

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Memoirs of Museum Victoria 60(2): 181-195 (2003)

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

http://www.museum.vic.gov.au/memoirs

A molecular and morphological review of the asterinid, Patiriella gunnii (Gray)

(Echinodermata: Asteroidea)

P. Mark O’Loughlin*, Jonathan M. Waters^ and Michael S. Roy^

' Honorary Associate, Marine Biology Section, Museum Victoria, GPO Box 666E, Melbourne, Victoria 3001, Australia

(pmo@bigpond.net.au)

^ Department of Zoology, University of Otago, PO Box 56, Dunedin, New Zealand

(Jonathan, waters @ stonebow.otago.ac.nz)

Abstract O’Loughlin, P.M., Waters, J.M., and Roy, M.S. 2003. A moleeular and morphologieal review of the asterinid, Patiriella

gunnii (Gray) (Eehinodermata: Asteroidea). Memoirs of Museum Victoria 60(2): 181-195.

The six-rayed species of the asterinid genus Patiriella Verrill from the rocky shallows of southern Australia are

reviewed. Based on moleeular and morphological evidence, it is judged that Patiriella brevispina H.L. Clark is a junior

synonym of Patiriella gunnii (Gray). Three new species are described: Patiriella medius sp. nov., Patiriella occidens sp.

nov. and Patiriella oriens sp. nov. A key to the four species of Patiriella is provided.

Keywords Echinodermata, Asteroidea, Patiriella, new species, taxonomy, Australia

Introduction

Typically, six-rayed asterinid seastars that occur on the rocky

coast of southern Australia are currently referred to either the

variably-coloured Patiriella gunnii (Gray, 1840) or dark-crim-

son Patiriella brevispina H.L. Clark, 1938. Molecular evidence

indicated to Hart et al. (1997) and Byrne et al. (1999) that there

were genetically divergent eastern and western forms of what is

currently referred to as P gunnii in southern Australia, and that

a taxonomic revision was necessary.

Gray (1840) described Avtenna gunnii from material sent to the

British Museum (Natural History) (BMNH, now The Natural

History Museum, London) from “Van Diemen’s Land”

(Tasmania, Australia) by Ronald Gunn. One lot (four speci-

mens, BMNH 40.3.9.10-13) was collected from Sandy Bay,

Hobart, in south-eastern Tasmania, and has a note “presumably

syntypes, judging from source and Gray’s label.” A second lot

(numerous specimens, BMNH 49.11.19) was from George

Town, northern Tasmania, and was considered by A.M. Clark

(1966) to be “presumably the types.” Dartnall (1970) estab-

lished one of the Hobart specimens as a lectotype (BMNH

40.3.9.10), and three of the George Town specimens as para-

lectotypes (BMNH 49.11.19-10, 14, 33). The type locality is

thus the Derwent River estuary at Hobart. Verrill (1913)

referred Axfer/na gunnii Gray, 1840 to his new genus Patiriella.

The lectotype and only one of the paralectotypes (BMNH

49.11.19.33) were present when the BMNH collection was seen

by O’Loughlin in April 2002. Two paralectotypes (BMNH

49.11.19.10 and 14) were absent and the remaining individuals

from George Town comprised a mixture of Patiriella species.

H.L. Clark (1938) correctly distinguished as discrete species

variably-coloured and crimson forms of P. gunnii. Without ref-

erence to the types of P. gunnii, he referred the variably-

coloured form to P. gunnii, and described the crimson form as

a new species, Patiriella brevispina H.L. Clark, 1938. In this

study the taxonomic status of six-rayed species of Patiriella is

re-assessed after further collecting across southern Australia,

and reference to molecular evidence and morphological exam-

ination of specimens in the Australian Museum, Sydney (AM),

Museum Victoria, Melbourne (NMV), South Australian

Museum, Adelaide (SAM), Tasmanian Museum, Hobart (TM),

and Western Australian Museum, Perth (WAM). We set out to

show by morphological comparison of the lectotype of P. gun-

nii (type locality, Hobart) with the paratypes of P. brevispina

(type locality, Bunbury in southern Western Australia) that the

crimson P. brevispina is conspecific with P. gunnii, and is thus

a junior synonym. The typically six-rayed asterinid seastars

which occur in southern Australia are thus the dark crimson

P. gunnii and three new variably-coloured species described

below.

In a few cases it has been possible to assign mentions of

''Patiriella gunnii” in the literature to one of the species referred

to below, and this has been done in the synonymies, but in most

cases this has not been possible. Authors who have referred to

"Patiriella gunnii” are: Lamarck (1816, as Asterias calcar

variety b; synonymy by Perrier (1876)); Muller and Troschel

(1842, as Asteriscus australis (part) and Asteriscus diesingi;

synonymies by Perrier (1876)); Dujardin and Hupe (1862, as

Asteriscus calcar, synonymy by Perrier (1876)); Perrier (1869,

182

R Mark O’Loughlin, Jonathan M Waters and Michael S. Roy

as Asteriscus exiguus; synonymy by Perrier (1876)); Penier

(1876, asAsterina gunnii); Sladen (1889, as A. gunnii)\ McCoy

(1890, as A. gunnii)-, Farquhar (1895 and 1898, as A. gunnii)-,

Verrill (1913, as Patiriella gunnii)-, Fisher (1919); H.L. Clark

(1928, 1938, 1946); Cotton and Godfrey (1942); A.M. Clark

(1966); Shepherd (1968); Dartnall (1969, 1971, 1980); Rowe

and Vail (1982); Zeidler and Shepherd (1982); A.M. Clark

(1983); O’Loughlin (1984); A.M. Clark and Downey (1992);

Marsh and Pawson (1993); A.M. Clark (1993); Rowe and Gates

(1995); Campbell and Rowe (1997); Edgar (1997). The pres-

ence of six-rayed specimens of Patiriella regularis Verrill, 1867

(normally five-rayed) in New Zealand led to reports by Muller

and Troschel (1842 (part), as Asteriscus australis), Perrier

(1876) and Mortensen (1925) that Asterina gunnii occurred in

New Zealand. Farquhar (1898) considered the six-rayed

material to be P. regularis.

Genetics

Methods. Sixty ethanol-preserved specimens provisionally identi-

fied as P. 'gunnii’ (21 locations) and 18 provisionally identified as

P. ‘brevispina’ (9 locations), sampled from a broad geograpliic range,

were included in genetic analyses. Genomic DNA was extracted from

tube foot tissue using a 5% chelex solution (Walsh et al., 1991) or a

CTAB -proteinase K extraction buffer (Saghai-Maroof et al., 1984). A

1580 base pair (bp) portion of the mitochondrial cytochrome oxidase 1

gene (COl) and the adjacent tRNA-Pro was amplified and sequenced

using universal primers F210-CO1 (5’ GGTAATGCCAATTAT-

GATTGG 3’) and COII 14098-14078 (5’ CCTARTTGGGTTCAR

TTTGCC 3’) (Hart et al., 1997). Subsequently, an internal 830 bp frag-

ment was amplified and sequenced for all specimens using specific

primers GUN COl-L (5’ TCCCAAAGCTATCATTCT 3’) and GUN

COl-R (5’ AGAGATCATTCCAAATCC 3’).

PCR (polymerase chain reaction) amplifications (25 pi) contained

50 mM KCl, 10 mM Tris-HCl pH 8.3, 0.1% Triton X-100, 0.005%

gelatine, 1.5 mM MgC12, 800 pM dNTPs, 0.5 pM of each primer, 0.75

units of Taq DNA polymerase, and 1 pi of extracted DNA. All ampli-

fications were performed in a PTC- 100 cycler (MJ Research,

Watertown, MA) with 40 cycles of 94°C 1 min, 48°C 30 sec, 72°C 30

sec. PCR products were purified with a High-Pure PCR Product

Purification Kit (Roche, Mannlieim, Germany) and sequenced using an

ABI Prism Big-Dye kit. Completed reactions were purified by ethanol

precipitation prior to electrophoresis on an automated DNA sequencer

(Perkin Elmer, Foster City, CA).

Phylogenetic analysis of aligned DNA sequence data was per-

formed under maximum parsimony (MP) using PAUP 4.0b 10

(Swofford, 1998). Phylogenetic confidence was estimated by boot-

strapping (Felsenstein, 1985) with 500 replicate data sets analysed with

the “full heuristic” option. Published COl sequences from P ‘gunnii’

(U50047-48), P ‘brevispina’ (U50049-50) and P. calcar (U50046)

(Hart et al., 1997) were also included in analyses. Genetic divergences

were calculated using the Kimura (1980) 2-parameter model of

sequence evolution.

Results. Phylogenetic analysis of DNA sequence data yielded

over 1000 equally parsimonious trees (863 steps; Fig. 1).

However, bootstrap analysis revealed strong phylogenetic

structure among haplotypes, with four well- supported clades

(100% bootstrap support). Three of the clades corresponded to

the so-called P. ‘gunnii ’ samples, and the fourth to the so-called

P. ‘brevispina’ (Fig. 1). All four clades were deeply divergent

(7.5-14.1%), and all were supported by strict consensus

analysis of 1000 MP trees. By contrast, haplotypes within each

clade exhibited small divergences (typically less than 1.0%)

and showed little phylogenetic structure.

Strong phylogeographic structure was detected within

P. ‘gunnii’: all eight Western Australian samples were placed in

a western clade, whereas all seven sequences from New South

Wales were placed in an eastern clade. The eastern clade was

also detected in Victoria and Tasmania, and the western clade

was also represented in South Australia. Although no geo-

graphic overlap was detected between eastern and western

clades, a third central clade exhibited an intermediate distri-

bution, encompassing Tasmania, Victoria, and South Australia

(Fig. 1). There was moderate bootstrap support (73%) for a

sister relationship between central and western clades.

Morphology

Methods. Specimens for which molecular sequences were obtained,

and consequent clades established, were used for the selection of

morphological diagnostic characters (MOL codes in lists of material,

refers to the code of the tissue sample used to obtain sequence data for

molecular phylogeny). These morphological characters were then used

to identify specimens in Australian museums. Descriptions of species

are based on a combination of observations of specimens in wet and

dried and cleared condition. Some specimens were cleared of surface

spines and spinelets and thin body wall (skin) using commercial

bleach, in order to more clearly observe skeletal plate form and size,

and the numbers of secondaiy plates and papulae in papular spaces.

Diagnostic characters are most readily observed on dried and cleared

specimens. Most diagnostic characters were found to vary and in many

cases to overlap amongst the four species. An attempt was made to

relate species and size of specimen to numbers of carinal plates on a

ray, width of proximal carinal plates, numbers of spinelets on proximal

carinal plates, and length of actinal spines, but these characters were

found to be so variable that they were unreliable for diagnostic

purposes. Moi-phological characters finally selected for diagnostic pur-

poses all vary within a species and with size. For determination, more

than one diagnostic character is most reliable.

Terminology follows that defined in the glossary and illustrated in

Clark and Downey (1992, figs 2, 3), except that “papular space” is

used for “papular area” (“restricted area with papular pores”) and

“papulate areas” is used to refer to the parts of the abactinal surface

where papulae occur. “Adradial” refers to the series of actinal interra-

dial plates adjacent to the ambulacral plates.

Results. The morphological analysis confirmed the existence of

four six-rayed species of Patiriella, corresponding to the four

clades established in the molecular study: Patiriella gunnii

(Gray, 1840) to the so-called P. ‘brevispina’ clade; Patiriella

medius sp. nov. to the central clade; Patiriella occidens sp. nov.

to the western clade; and Patiriella oriens sp. nov. to the

eastern clade (Fig. 1).

Asterinidae Gray, 1840

Patiriella Verrill, 1913

Synonymy and remarks. O’Loughlin et al. (2002) reviewed the

status of Patiriella Verrill, 1913, and noted that recent molecu-

lar phylogenetic analyses of species of Asterinidae (Hart et al.,

1997; Byme et al., 1999; J. M. Waters and M. S. Roy, unpubl.

Review of the sea-star, Patiriella gunnii

183

data) were beginning to provide a basis for a reassessment of

asterinid taxonomy. Pending publication of a review of the

assignment of species to genera within Asterinidae, the four

species treated in this work are retained provisionally in

Patiriella, most recently diagnosed by Campbell and Rowe

(1997).

Key to six-rayed species of Patiriella

1. Subambulacral spines predominantly 1 per plate, up to

twice length of adradial actinal spines; actinal interradial

spines very short, frequently bulbous; abactinal surface

uneven; abactinal spinelets prominently spinous, fre-

quently low to subcapitate; proximal papular spaces large,

frequently with more than 10 secondary plates and more

than 10 papulae per space when R = 30 mm; abactinal

colour of adults consistently uniform crimson to brownish

red, tube feet orange Patiriella gunnii (Gray, 1840)

— Subambulacral spines predominantly 2-3 per plate, not up

to twice the length of adradial actinal spines; actinal inter-

radial spines not very short or bulbous; abactinal surface

even; abactinal spinelets moderately to minutely spinous,

frequently columnar; proximal papular spaces not large,

fewer than 10 secondary plates and 10 papulae per space

when R = 30 mm; abactinal colour variable, not uniform

crimson to brownish red, with orange tube feet 2

2. At least a few suboral spines frequently present; adradial

actinal spines up to about two-thirds length of subambu-

lacral spines; actinal interradial spines short and fine;

carinal plates normally doubly papulate for less than two-

thirds ray length, frequently less than half ray length; prox-

imal papular spaces small, frequently up to 3 secondary

plates and 3 papulae per space when R = 30 mm; abactinal

spinelets relatively small and fine, predominantly narrow-

ing distally, minutely spinous; abactinal colour variable,

frequently overall maroon red (not reported with grey or

blue, or with black disc); actinal colour off-white,

frequently with prominent flecking

Patiriella medius sp. nov.

— Suboral spines rarely present; adradial actinal spines fre-

quently more than two-thirds length of subambulacral

spines; actinal interradial spines digitate; carinal plates

normally doubly papulate for more than half ray length,

frequently more than two-thirds ray length; proximal

papular spaces fairly open, frequently with about 6

secondary plates and 6 papulae per space when R = 30

mm; abactinal spinelets relatively coarse, predominantly

columnar, slightly narrowing to slightly widening distally,

moderately spinous; abactinal colour variable, not overall

maroon red 3

3. Form variable, commonly distinctive short rays with inter-

radial margin deeply indented; subambulacral spines fre-

quently projecting fairly prominently; abactinal spinelets

coarse, columnar, moderately spinous, frequently widened

distally; up to about 8 spinelets on inferomarginal plates

when R = 20 mm, up to about 11 when R = 30 mm;

abactinal colour frequently dark, with grey or brown or

blue, infrequently with red . . Patiriella occidens sp. nov.

— Form variable, commonly subhexagonal with interradial

margin slightly incurved; subambulacral spines not pro-

jecting significantly; abactinal spinelets fairly coarse,

columnar, moderately spinous, frequently narrowing dis-

tally, up to about 10 spinelets on inferomarginal plates

when R = 20 mm, up to about 15 when R = 30 mm; abac-

tinal colour frequently pale, with white or pink or mauve or

orange or bright red, disc frequently black (not reported

with grey (except in NSW), or blue)

Patiriella oriens sp. nov.

Patiriella gunnii (Gray)

Figures 1 (as P. ‘brevispina’), 2a-f, 3a-f, 7b

Asterina gunnii Gray, 1840: 289-290. — Gray, 1866: 16. — McCoy,

1890: 372, pi. 200 fig. 2 (part).

Patiriella gunnii. — Venill, 1913: 484. — Dartnall, 1970: 74-76, pi.

Patiriella brevispina H.L. Clark, 1938: 166-167, pi. 22 figs 2-3. —

Cotton and Godfrey, 1942: 202.— H.L. Clai'k, 1946: 134-135.— A.M.

Clark, 1966: 320.— Shepherd, 1968: 745, 747.— Dartnall, 1969: 55.—

Dartnall, 1970: 75-76.— Dartnall, 1971: 47, fig. 1.— Dartnall, 1980:

34, 65.— Rowe and Vail, 1982: 222.— Zeidler and Shepherd, 1982:

402, 412; figs 10.7c, d. — O’Loughlin, 1984: 136. — Bennett, 1987:

346-347, fig. — Rowe and Gates, 1995: 39. — Campbell and Rowe,

1997: 130.— Edgar, 1997: 346, fig.— Hart etal., 1997: 1848-1861, figs

1-4, tabs 1, 2. — Byrne et al., 1999: 188-191, figs 1, 3C, 6 (new

synonymy).

Material examined. Asterina gunnii Gray, 1840. Australia, Tasmania,

Hobart, Sandy Bay, 2 m, R. Gunn, BMNH 40.3.9.10 (Lectotype: dry;

partly eleared; designation by Dartnall, 1970); BMNH 40.3.9.-11, 12,

13 (3 specimens).

Patiriella brevispina H.L. Clark, 1938. Western Australia,

Koombana Bay, Bunbury, 9-14 m, E.W. Bennett and H.L. Clark, 26

Oct 1949, AM J6181 (2 paratypes, dry).

Other material (selected for moleeular confirmation, distribution

and depth data). Vic. East Gippsland, off Ninety Mile Beach, 38°42’ S,

147°53’ E, 22 m, NMV E73255 (1); Western Port, Honeysuckle Point,

TM H723 (4); Elinders, ocean platforms, NMV E71744 (3) (MOL 54);

NMV E93430 (1) (MOL 180); NMV E93429 (1) (MOL 10); NMV

E93435 (4) (MOL 130-131); Port Phillip Bay, Altona, NMV E72130

(1) (MOL 53); Geelong, Mackey St jetty, 2 m, NMV E93441 (1) (MOL

51, 56); “Mullet Holes”, 10 km NE Apollo Bay, NMV E93433 (3)

(MOL 97-99); Port Eairy, Griffith L, roeky shallows, NMV E93432 (1)

(MOL 81); Portland, below lighthouse, 2-3 m, NMV E93440 (1). Tas.

Eaglehawk Neck, rocky shallows, NMV E71873 (1); Bass Strait, Cape

Portland, TM H596 (1); Jacobs Boat Harbour, TM H2941 (1). SA. Gulf

Saint Vincent, Normanville, NMV E74628 (1) (MOL 60); NMV

E74629 (1) (MOL 59); NMV E93434 (1) (MOL 111); NMV E93436

(1) (MOL 102); Eyre Peninsula, Point Westall, near Streaky Bay, NMV

E93437 (1) (MOL 55); Nuyts Arehipelago, Goat L, 29 m, NMV

E93446 (1). WA. Esperance, Sandy Hook L, WAM Z9471 (1);

Busselton, jetty piles, 4 m, WAM Z8948 (3) (MOL 146); Cockburn

Sound, TM HI 116 (4); Trigg L, WAM Z9539 (2); Yanchep, lagoon,

NMV E93443 (2).

Description (dry and cleared specimens). Up to R = 56 mm;

5-9 rays, predominantly 6 (102 of 116 AM specimens with

6 rays, 10 with 7 rays, 2 with 5 rays, 1 with 8 rays, 1

with 9 rays); form variable from 6 short rounded to pointed

rays with interradial margin incurved, to hexagonal; body

thick, flat orally, flattened dome aborally, acute angle at

184

margin; madreporite conspicuous; lacking pedicellariae;

gonopores abactinal.

Abactinal surface uneven; papulate areas more extensive

than non-papulate areas; secondary plates abundant, very irreg-

ular in size and form; proximal radial and interradial plates

openly imbricate; proximal papular spaces large, frequently up

to 16-20 secondary plates and 16-20 papulae in proximal

papular spaces outside disc when R = 30 mm (5-6 secondary

plates and 5-6 papulae when R = 20 mm); abactinal plates

thick, raised, prominent, crescentic in papulate areas, carinally

with double notch and proximal lobe; carinal series variably

regular from close to disc to end or near end of rays, frequent-

ly doubly papulate to near end of ray when R = 20 mm and

larger; distal interradial non-papulate plates closely imbricate,

domed, rounded proximally; disc variably distinct, bordering

plates variably regular crescentic radial plates and smaller

interradial plates, disc frequently obscured by irregular large

plates within and distal to disc; abactinal plates granular,

covered by glassy convexities, lacking spine-bearing ridge;

abactinal spinelets with variable forin, frequently widened

distally, some capitate or truncate or columnar or narrowing

distally or with swollen base, long spines distally, typically

about 0.40 mm long when R = 30 mm (0.32 mm when R = 20

mm), distributed over projecting surface of plates; supero-

marginal plates aligned longitudinally with inferomarginal

plates; lacking internal superambulacral plates between

ambulacrals and actinals; distal abactinal and actinal inter-

radial plates with internal tapered vertical contiguous

projections.

Projecting inferomarginal plates form margin, up to about

11 spinelets per plate when R = 20-30 mm; actinal plates in

regular series, curving acutely from furrow to margin, some

proximal actinal areas not calcified; actinal interradial spines

generally thick, very short, columnar to bulbous, distally

minutely spinous; actinal interradial proximal plates with 1-2

spines, distally 2-3 (4 rare); adradial row of actinal interradial

proximal plates with predominantly 1 short, thick, bulbous to

tapered spine, typically up to only half the length of sub-

ambulacral spines; adambulacral proximal plates with pre-

dominantly 1 (2 rare) thick subambulacral spines, frequently

bulbous with waist, form variable from columnar to sub-

spatulate to spatulate, minutely spinous distally; furrow spines

2-3 (4 rare) proximally, fairly thick, form variable from tapered

to slightly widened distally; suboral spines rare (7 suboral

spines on 1 of 116 AM specimens); oral spines 4-6,

predominantly 5.

Live colour. Abactinally uniform dark crimson or reddish-

brown (a few AM labels refer to “purple” and “indigo blue”),

paler actinally; orange tube feet.

Distribution. Eastern Vic. (off Ninety Mile Beach) continuous

to Abrolhos Islands off WA (Loisette Marsh, pers. comm.);

Bass Strait; Tas.; under rocks; 0-29 m (molecular confirmation

for Flinders (Vic.) to Busselton (WA)).

Remarks. The lectotype of Asterina gunnii and the two

paratypes of Patiriella brevispina seen in this study do not

exhibit any significant morphological differences and in

R Mark O’Loughlin, Jonathan M Waters and Michael S. Roy

particular have single stout subambulacral spines per plate and

very short actinal interradial spines. These two diagnostic char-

acters were used by Clark (1938) to distinguish his new species.

Clark (1938) also considered the “consistent deep purple to

brownish-crimson colour and orange tube feet” of R brevispina

to be diagnostically reliable. This observation is confirmed and

specimens with these characteristic colours consistently

exhibit the morphological diagnostic characters of A. gunnii.

Dartnall (1970) followed H.L. Clark (1938) when designating

and describing the lectotype of A. gunnii, and noted that

paired subambulacral spines distinguished P. gunnii from

P. brevispina. Dartnall (1970) then considered the lectotype of

A. gunnii to be exceptional in having single subambulacral

spines. The lectotype of A. gunnii (R = 24 mm) and paratypes

of P. brevispina (up to R = 20 mm) are similar in size, and are

small specimens of A. gunnii (up to R = 56 mm). On the mor-

phological evidence P. brevispina is considered here to be a

junior synonym of A. gunnii.

Hart et al. (1997) and Byrne et al. (1999) found from molec-

ular evidence that P. ‘brevispina’ specimens from eastern

Australia (Mornington Peninsula, Vic.) and Western Australia

(Perth) were closely related. Eastern and Western Australian

specimens of P. ‘brevispina’ were found to be conspecific in

this study. Byrne (1995, 1996), Byrne and Cerra (1996), and

Long and Byrne (2001) have reported on the reproductive and

developmental biology of P. gunnii (as P. ‘brevispina ’).

The combination of characters which distinguishes P. gunnii

from other six-rayed species of Patiriella is: consistent uniform

dark crimson to reddish brown colour; orange tube feet; larger

maximum size; prominent papular spaces with numerous

papulae and secondary plates; noticeably spinous abactinal

spinelets; predominantly single thick subambulacral spines per

plate; and very short thick actinal interradial spines. No evi-

dence was found to confirm the occurrence of P. gunnii in

NSW, or the depth of 39 m reported by Rowe and Gates (1995,

as P. ‘brevispina’).

Patiriella medius sp. nov.

Figures 1 (as ‘central’), 4a-f, 7c

Material examined. Holotype. Victoria, Port Fairy, Griffith I., rocky

shallows, 0-2 m, M. O’Loughlin and M. Mackenzie, 29 Dec 2001,

NMV F92986 (MOL 72).

Paratypes. Type locality and date, NMV F92987 (3) (MOL 73, 74,

79); NMV F92988 (2 dry, 2 cleared) (MOL 69, 70, 75, 76).

Other material (selected for molecular confirmation, distribution

and depth data). Vic. W of Wilsons Promontory, Walkerville South,

Bear Gully, NMV F71869 (1); Western Port, McHaffie Point, NMV

F93174 (4); Flinders, ocean platform, NMV F93171 (2); Port Phillip

Bay, Popes Eye, 5-12 m, NMV F93730 (1); Point Addis, 8-13 m,

NMV F93165 (3) (MOL 170, 172); “Mullet Holes”, 10 km NE Apollo

Bay, 0-2 m, NMV E92990 (3) (MOL 94-96); Port Eairy, Griffith L,

rocky shallows, NMV E83593 (2) (MOL 50); NMV E93164 (1) (MOL

176); NMV E87171 (1) (MOL 177); Portland, Nelson Bay, 24 m,

NMV E73193 (2). Tas. Port Aithur, below low tide, TM H809 (1);

Bass Strait, Waterhouse Passage, rocky shallows, NMV E71872 (1)

(MOL 46); Tamar River, Greens Beach, TM H1107 (21); Somerset,

near Burnie, under rocks at low tide, WAM Z9394 (5); Rocky Cape, 5

m, NMV E92989 (2) (MOL 211). SA. Cape Jaffa, 14-20 m, SAM

K1938 (2); Kangaroo L, Western River, 8-10 m, SAM K1917 (1); Gulf

Review of the sea-star, Patiriella gunnii

185

100

U 50049 V

MOL 99 V

AM J22ei9T

MOL 10 V

MOL 51 V

MOL 97 V

MOL 53 V

MOL 59 S

MOL 54 V

MOL 131 V

MOL 130 V

MOL 1B0 V

MOL 55 S

MOL 111 3

h MOL 60 S

MOL9S V

MOL 102 S

^ MOL 31 3

MOL 146W

L U50050 W

100

P. calcar

P. ‘brevispina’

US0046 N

5 changes

U50047 N

MOL 134 N

MOL 215 N

MOL 219 N

[r MOL 117 V

^ MOL 78 V

, MOL 1 75 V

■1 MOL 5 V

' MOL 2 V

- MOL 71 V

- MOL 65 V

- MOL 77 V

MOL 192 T

AM J14016 N

MOL 201 T

MOL 193 T

MOL 205 T

MOL 202 T

MOL 191 T

MOL 220 N

MOL 213 N

MOL 116 V

MOL SO V

MOL 203 T

MOL 1 70 V

MOL 1 72 V

MOL 96 V

MOL 177 V

MOL 75 V

MOL 47 S

MOL 164 S

MOL 72 V

MOL 163 S

MOL 165 S

MOL 103 S

- MOL 105 3

L MOL 95 V

MOL 73 V

MOL 94 V

MOL 1 76 V

- MOL 50 V

j MOL 69 V

■* MOL 76 V

- MOL 70 V

MOL 74 V

MOL 79 V

I, MOL 46 T

n MOL 211 T

MOL 106 S

I- MOL 107 S

MOL 104 3

eastern’

100

‘central’

I MOL 1 S2 S

MOL 148 W

MOL 155 W

U50048 W

MOL 149 W

MOL 67 S

MOL 1B1 S

MOL 154 W

MOL 150W

MOL 156W

MOL 151 W

100

ri ^

‘w/estern’

Figure 1. One of 1000+ optimal MP trees (863 steps) derived from analysis of COl sequence data. Numbers at nodes are bootstrap percentages

based on 500 replicate analyses. MOL codes refer to specimens obtained for the current study. AM codes refer to Australian Museum specimens.

Previously published sequences (Hart et al., 1997) are indicated in bold. Geographic origin of samples is indicated: Tasmania (T), Victoria (V),

New South Wales (N), South Australia (S), Western Australia (W).

186

Saint Vincent, Fleurieu Peninsula, Rapid Bay jetty, 8-12 m, NMV

F93166 (4) (MOL 163-165); Normanville, rocky shallows, NMV

F93163 (4) (MOL 104-107); F93167 (1) (MOL 103); Port Noarlunga,

under stones, 1 m, WAM Z9390 (1); Yorke Peninsula, Edithburg,

Troubridge Light, 18 m, TM H1359 (1); Sir Joseph Banks Group, 3-14

m, SAM K1923 (3); Investigator Group, 6-8 m, SAM K1901 (1); Eyre

Peninsula, near Streaky Bay, Point Westall, rocky shallows, NMV

E71856 (1 cleared) (MOL 47, 57); Nuyts Archipelago, 34 m, SAM

K1933 (1). WA. Hopetoun, east of jetty, rock platform, NMV E73201

(3); Cheyne Bay, intertidal reef, WAM Z9472 (1); Yallingup, under

boulders with P. ‘brevispina’ (= P. gunnii here) and P ‘gunnii’ (= P

occidens here), 0-1 m, WAM Z9477 (5); Cape Naturaliste, 9 m, WAM

Z9404 (1); Dunsborough, Eagle Bay, under rock, 10 m, NMV E93172

(1); Bunbury, 4 km N, WAM Z9411 (1); Eremantle, Halls Bank, under

rock, 8 m, NMV E73 178 (1).

Description (dry and cleared specimens). Up to R = 38 mm;

5-7 rays, predominantly 6 (134 of 145 SAM, TM and WAM

specimens with 6 rays, 9 with 7 rays, 2 with 5 rays); form vari-

able from 6 short predominantly pointed rays with interradial

margin incurved, to hexagonal; body flat orally, flattened dome

aborally, acute angle at margin; madreporite conspicuous;

lacking pedicellariae; gonopores abactinal.

Abactinal surface even; papulate areas slightly more

P. Mark O’Loughlin, Jonathan M Waters and Michael S. Roy

extensive than non-papulate areas; secondary plates few, very

irregular in size and form; proximal radial and interradial plates

fairly closely imbricate; proximal papular spaces not large, fre-

quently 1-3 secondary plates and 2-4 papulae in proximal

papular spaces outside disc when R = 20-30 mm; abactinal

plates crescentic in papulate areas, carinally with double notch

and proximal lobe; carinal series variably regular from close to

disc to near end of ray, rarely doubly papulate beyond half ray

length; distal interradial non-papulate plates closely imbricate,

domed, rounded proximally; disc variably distinct, bordering

plates variably regular crescentic radial and smaller interradial

plates, frequently obscured by irregular large plates within and

distal to disc; abactinal plates granular, covered by glassy con-

vexities, lacking spine-bearing ridge; abactinal spinelets with

variable form, subcolumnar to narrowing and rounded distally

with swollen base, minutely spinous distally, typically about

0.36 mm long when R = 30 mm (0.30 mm when R = 20 mm),

distributed over projecting surface of plates; superomarginal

plates aligned distally with inferomarginal plates; lacking inter-

nal superambulacral plates between ambulacrals and actinals;

distal abactinal and actinal interradial plates with internal

tapered vertical contiguous projections.

Eigure 2. a-c, Asterina gunnii Gray, 1840, lectotype, R = 24 mm (BMNH 40.3.9.10, dry), a, abactinal view (extensively cleared); b, abactinal ray

(extensively cleared); c, actinal ambulacra and interradius, with single stout subambulacral spines per plate (arrow) and very short interradial

spines.

d-f, Patiriella brevispina H.L. Clark, 1938, paratype, R = 20 mm (AM J6181, dry), d, abactinal view; e, abactinal ray; f, actinal ambulacra and

interradius, with single stout subambulacral spines per plate (arrow) and very short interradial spines.

Review of the sea-star, Patiriella gunnii

187

Projecting inferomarginal plates form margin, widely

rounded distally, frequently 10-14 spinelets per plate when

R = 20-30 mm; actinal plates in regular series, curving acutely

from furrow to margin; some proximal actinal areas not cal-

cified; actinal interradial spines generally fairly slender,

tapered, short; actinal interradial proximal plates with 1-4

spines (predominantly 2-3), distally 2-4 very short tapered

spines (frequently 3), minutely spinous distally; adradial row of

actinal interradial proximal plates with predominantly 2 tapered

spines, minutely spinous distally, significantly shorter than sub-

ambulacral spines (about half to two-thirds length); adam-

bulacral proximal plates with frequently 2-3 thick tapered sub-

ambulacral spines, minutely spinous distally, frequently

unequal, form variable from digitate to spatulate to distally

bulbous, commonly slightly longer than furrow spines; furrow

spines slender, webbed, tapering, predominantly 3-4 (some-

times 5) per plate proximally when R = 20 mm and larger,

minutely spinous distally; suboral spines frequently present (at

least 1 suboral spine on 68 of 115 AM, SAM and TM speci-

mens examined; 10-12 suboral spines on 42 of 115 specimens);

oral spines 5-7, predominantly 6.

Live colour. Abactinally very variable; dark-coloured disc not

reported; frequently overall maroon red or red or reddish brown

appearance, sometimes fairly uniform pale brown or orange or

pink, sometimes with red or mauve or orange or brown or

cream or white flecks; a few with margin or rays and interradii

coloured differently, or with colour patches; some mottled with

dark red, red, maroon, mauve, brown, orange, white; actinally

typically off-white with prominent maroon flecking.

Distribution. Walkerville South (W of Wilsons Promontory,

Vic.), Bass Strait, Tas., continuous to Fremantle (WA); under

rocks; 0-34 m (molecular confirmation for Point Addis (Vic.)

to western Eyre Peninsula (SA)).

Etymology. From medius (Latin, as a noun in apposition) mean-

ing “between the two”, and referring to a distribution across

southern Australia between the most easterly distribution

of Patiriella oriens sp. nov. (below) and the most westerley

distribution of Patiriella occidens sp. nov. (below).

Remarks. The combination of characters which distinguishes

P. medius from other six-rayed species of Patiriella is: rarely

doubly papulate carinally for more than half ray length;

Figure 3. a-f, Patiriella gunnii (Gray), a, abactinal view, R = 40 mm (NMV F73273); b, abactinal ray, carinally doubly papulate for most of ray,

R = 40 mm (NMV F73273); c, abactinal interradius with coarse spination and fringe of stout inferomarginal spinelets, R = 22 mm (WAM Z8948);

d, cleared abactinal view, with numerous papulae and secondary plates, R = 35 mm (WAM Z8952); e, actinal ambulacra and interradius, lacking

suboral spines (arrow), R = 40 mm (NMV F73273); f (above), common forms of abactinal spinelets; f (below), common forms of actinal spines

proximal to ambulacrum, with very short interradial spines (arrow).

188

abactinal spinelets small and fine; frequent presence of some

sub-oral spines; adradial row of actinal interradial spines up to

about two-thirds the length of subambulacral spines; actinal

interradial spines short, thin. The limited live colour data avail-

able for confirmed determinations indicates that maroon red is

a common abactinal overall colour, with no reports of grey or

blue or a black disc.

Patiriella occidens sp. nov.

Figures 1 (as ‘western’), 5a-f, Id

Patiriella gunniiW. — Hartetal., 1997; 1848-1861, figs 1-4, tabs 1,

2. — Byrne et al., 1999: 188-194, figs 1, 3D, 6 (non Patiriella gunnii

(Gray, 1840)).

Material examined. Holotype. Western Australia, Perth, Cottesloe, on

reef amongst algae, 1 m, L. Marsh, 29 Dec 2001, WAM Z8951 (MOL

151).

Paratypes. Type locality and date, NMV F92971 (1, cleared) (MOL

150); Albany, Cape Vancouver, Quaranup, amongst boulders, 1 m, L.

Marsh, 13 Dec 2001, WAM Z8949 (1) (MOL 148); under boulder, 1 m,

WAM Z8950 (1) (MOL 149); Cockbum Sound, Woodman Point, under

rocks, 1 m, L. Marsh, 1 Jan 2002, WAM Z8953 (3) (MOL 154-156).

P. Mark O’Loughlin, Jonathan M Waters and Michael S. Roy

Other material (selected for molecular confirmation, distri-

bution and depth data). Vic. Port Fairy, causeway beach,

NMV F73149 (1). SA. Victor Harbour, The Bluff wharf, NMV

F92975 (1) (MOL 67); Cape Jervis, rocky shallows, NMV

F74638 (1) (MOL 182); Kangaroo L, Eastern Cove, rocky shal-

lows, NMV F71862 (2) (MOL 181); Gulf Saint Vincent,

Glenelg, SAM K1932 (3); Yorke Peninsula, Edithburg, 0-4 m,

SAM K1904 (3); Sir Joseph Banks Group, 0-1 m, SAM K1899

(2); Eyre Peninsula, Point Labatt, 0-1 m, SAM K1907 (1);

Nuyts Archipelago, 14 m, SAM K1898 (2). WA. E of

Hopetoun, Mason Bay, East Mason Point, granite/dolerite with

algae and seagrass, 0-3 m, WAM Z9470 (1); Cheyne Bay,

under stones, WAM Z9479 (6); Two Peoples Bay, WAM Z9576

(2); Albany, Middleton Beach, under rocks, WAM Z9478 (2);

Torbay, Mutton Bird L, under boulders, intertidal, WAM Z9474

(2); Kilkamup, Cape Mentelle, WAM Z9466 (1); Cowaramup

Bay, under boulders, 0-1 m, WAM Z9400 (1); Yallingup, lime-

stone reef, under boulders with P. ‘brevispina ’ (= P gunnii here),

WAM Z9396 (4); Cape Naturaliste, under intertidal granite

boulders, WAM Z9405 (2); Geographe Bay, Dunsborough,

WAM Z9402 (1); Rockingham, Point Peron, H.L. Clark, Oct

Figure 4. a-f, Patiriella medius sp. nov. a, actinal view of holotype, R = 25 mm (NMV F92986); b, abactinal ray of paratype, carinally doubly

papulate for about half ray length (ending at arrow), R = 23 mm (NMV F92988); c, abactinal interradius of paratype, with fine spination and fringe

of fine inferomarginal spinelets, R = 24 mm (NMV F92988); d, cleared abactinal view of paratype, with few papulae and secondary plates, and

carinal plates doubly papulate for about half the ray length, R = 22 mm (NMV F92988); e, actinal ambulacra and interradii of paratype, with

suboral spines (arrows), and atypically 7 ambulacra, R = 23 mm (NMV F92988); f (above), common forms of abactinal spinelets; f (below),

common forms of actinal spines proximal to ambulacrum, with adradial actinal spines (arrow) much shorter than subambulacral spines.

Review of the sea-star, Patiriella gunnii

189

1929, WAM Z9440-2 (3); SAM K712 (3); Rottnest L, reef,

WAM Z9530 (9); Trigg L, reef platform, WAM Z9454 (1);

Yanchep, reef, WAM Z9433 (1); Port Gregory, N of Geraldton,

top of reef in pool, NMV F73179 (1); Kalbarri (Murchison

River ), reef top, WAM Z9413 (2).

Description (dry and cleared specimens). Up to R = 38 mm;

4-9 rays, predominantly 6 (292 of 331 AM, SAM, TM and

WAM specimens with 6 rays, 25 with 7 rays, 9 with 8 rays, 3

with 5 rays, 1 with 4 rays, 1 with 9 rays); form variable from 6

distinct pointed rays with interradial margin deeply incurved

(common for larger specimens) to subhexagonal (rare except

for smaller specimens); body flat orally, flattened dome abor-

ally, acute angle at margin; madreporite conspicuous; lacking

pedicellariae; gonopores abactinal.

Abactinal surface slightly uneven; papulate areas more

extensive than non-papulate areas; secondary plates numerous,

very irregular in size and form; proximal radial and interradial

plates fairly openly imbricate; proximal papular spaces fairly

large, frequently 2-6 secondary plates and 6-7 papulae in prox-

imal papular spaces outside disc when R = 30 mm (2-4

secondary plates and 4-6 papulae when R = 20 mm); abactinal

plates crescentic in papulate areas, carinally with double notch

and proximal lobe; carinal series frequently regular from close

to disc to end or near end of rays, doubly papulate for at least

three quarters ray length when R = 20 mm and larger; distal

interradial non-papulate plates closely imbricate, domed,

rounded proximally; disc variably distinct, bordering plates

variably regular crescentic radial plates and smaller interradial

plates, disc frequently obscured by irregular large plates within

and distal to disc; abactinal plates granular, covered by glassy

convexities, lacking spine-bearing ridge; abactinal spine-

lets frequently twice as long as wide, variable form, most fre-

quently columnar, sometimes slightly widened or narrowing

distally, sometimes with slight waist, truncate and prominently

spinous distally, typically up to 0.48 mm long when R = 30 mm

(0.36 mm when R = 20 mm), distributed over projecting sur-

face of plates; superomarginal plates aligned longitudinally

with inferomarginal plates; lacking internal superambul-

acral plates between ambulacrals and actinals; distal abactinal

and actinal interradial plates with internal tapered vertical

contiguous projections.

Figure 5. a-f, Patiriella occidens sp. nov. a, actinal view of holotype, with distinctively elongate rays, R = 35 mm (WAM Z8951); b, abactinal

ray, carinally doubly papulate for most of ray (ending at arrow), and coarse cover of abactinal spinelets, R = 25 mm (WAM Z9491); c, abactinal

interradius with coarse spination and fringe of stout inferomarginal spinelets, up to 10 per plate, R = 30 mm (WAM Z9483); d, cleared abactinal

view of paratype, with carinal plates doubly papulate for all of ray shown, R = 31 mm (NMV F92971); e, actinal ambulacra and interradii, with

oral plates lacking suboral spines, R = 25 mm (WAM Z9491); f (above), common forms of abactinal spinelets; f (below), common forms of

actinal spines proximal to ambulacmm, with projecting subambulacral spines (arrow).

190

Projecting inferomarginal plates form margin, frequently

8-11 spinelets per plate when R = 20-30 mm; actinal plates in

regular series, curving acutely from furrow to margin, some

proximal actinal areas not calcified; actinal interradial spines

generally thick, long, digitate, distally spinous; actinal inter-

radial proximal plates with 1-2 spines, distally 2 (rarely 3)

shorter, digitate to slightly bulbous, distally spinous spines;

adradial row of actinal interradial proximal plates with pre-

dominantly 1 thick digitate spine, rarely 2, typically about four

fifths length of subambulacral spines, some slightly bulbous;

adambulacral proximal plates with 1-3, predominantly 2, thick

subambulacral spines, frequently unequal, form variable from

digitate to slightly bulbous to subcapitate to spatulate to

widending distally, minutely spinous distally; furrow spines

slender, tapering, webbed, 2-4 per plate proximally, pre-

dominantly 3, minutely spinous distally, subequal in length

with subambulacral spines; suboral spines rare (at least 1

suboral spine on 23 of 307 AM, SAM, TM and WAM speci-

mens examined; more than 10 suborals on 2 of 307); oral spines

4-6, predominantly 5.

P. Mark O’Loughlin, Jonathan M Waters and Michael S. Roy

Live colour. Very variable abactinally; commonly dark

coloured disc, sometimes red; frequently grey appearance;

sometimes fairly uniform grey or red or blue-green or grey-blue

or blue or brown or orange, sometimes with black or white

flecks; some with rays, interradii and margin coloured differ-

ently, or with colour patches; frequently mottled with grey,

white, green, red, brown, blue, orange, mauve, black.

Distribution. Port Fairy (Vic.) continuous to Kalbarri (WA);

predominantly on reef flat, sometimes with seagrass; 0-14 m

(molecular confirmation for Victor Harbour (SA) to Perth

(WA)).

Etymology. From occidens (Latin, as a noun in apposition)

meaning “west” and referring to the westerly distribution in

southern Australia.

Remarks. Hart et al. (1997) and Byrne et al. (1999) found from

molecular evidence that specimens of Patiriella ‘gunnii’ from

eastern and western Australia had divergent lineages. The west-

ern material was collected from Margaret River in WA

(L. Marsh, pers. comm.). Western Australian specimens are

Figure 6. a-f, Patiriella oriens sp. nov. a, actinal view of holotype, R = 28 mm (NMV F92983); b, abactinal ray, carinally doubly papulate for

about two-thirds of ray length (ending at arrow), R = 25 mm (NMV F73152); c, abactinal interradius with moderately coarse spination and fringe

of fairly fine inferomarginal spinelets, up to 15 per plate, R = 25 mm (NMV F73152); d, cleared abactinal view of paratype, with carinal series

doubly papulate for most of ray length, R = 25 mm (NMV F92984); e, actinal ambulacra and interradii, R = 30 mm (NMV F73155); f (above),

common forms of abactinal spinelets; f (below), common forms of actinal spines proximal to ambulacmm, with relatively long furrow spines

(arrow).

Review of the sea-star, Patiriella gunnii

191

confirmed by molecular and morphological evidence and

described here as Patiriella occidens sp. nov.

Grice and Lethbridge (1988) reported on the reproductive

and developmental biology of Patiriella ‘gunnii’. Since the

research was based on material collected from the region of

Perth, it is assumed here that the species was not P gunnii

but probably the readily found and collected new species

P. occidens (used hereafter for P ‘gunnii’ from western

Australia). It is improbable that the collections included the

cryptic P. medius, which is sympatric with P occidens in the

Perth region. Grice and Lethbridge (1988) found that spawning

by P occidens occurred in late summer and early autumn.

Spawning is thus later than P. oriens (see below), a factor which

may be significant in the maintenance of genetic identity in

these similar species. P occidens is found most frequently on

intertidal reef platform, a habitat frequently occupied by

P. calcar. This potential competitive factor may have resulted

in the absence of P. calcar in most of the distribution range of

P. occidens.

The combination of morphological characters which

distinguishes P. occidens from other six-rayed Patiriella

species is: frequently distinct long rays; carinal series of plates

frequently doubly papulate for at least three-quarters ray

length; abactinal spinelets frequently columnar, distally

slightly swollen and spinous, and creating a very coarsely

spinous surface appearance; normal absence of suboral spines;

subambulacral spines projecting significantly above furrow and

actinal interradial spines; actinal interradial spines digitate; up

to about 1 1 spinelets per inferomarginal plate. The limited live

colour data available for confirmed determinations indicates

that grey or brown or blue are frequently evident abactinally,

and red infrequently.

Patiriella oriens sp. nov.

Figures 1 (as ‘eastern’), 6a-f, 7e

Patiriella gunnii E. — Hart et al., 1997: 1848-1861, figs 1-4, tabs 1,

2. — Byrne et al., 1999: 188-194, figs 1, 3D, 6 (non Patiriella gunnii

(Gray, 1840)).

Material examined. Holotype. Tasmania, Recherche Bay, Black Reef,

8 m, N. Barrett, 6 Jun 2002, NMV F92983 (MOL 193).

Paratypes. Type locality and date, NMV F92984 (1 dry, 1 cleared)

Figure 7. a, the temperate region of southern Australia.

b-e, geographic distributions based on morphological determinations, with type localities for each of the species indicated by crossed symbols;

b, P. gunnii sensu lato; c, P. medius sp. nov.; d, P. occidens sp. nov.; e, P. oriens sp. nov.

192

(MOL 191, 192); Bicheno, rocky coast, 5 m, N. Barrett, 1 Apr 2002,

NMV F92982 (6) (MOL 201-203, 205).

Other material (selected for molecular confirmation, distribution

and depth data). Lord Howe L, AM J 193 39 (1). Qld. Rockhampton,

Satellite L, 6 m, NMV F73165 (1). NSW. Byron Bay, Julian Rocks, 12

m, AM J14869 (1); Minnie Water, AM J12953 (26); Wooli, AM J15570

(6); Woolgoola, North Solitary L, 30 m, AM J14910 (4); Coffs

Harbour, Solitary L, 9 m, AM J14942 (13); South Solitary L, 27 m, AM

J14899 (1); Broughton L, near Port Stephens, 25 m, AM J12963 (3);

Wyong, Nora Head, WAM Z9382 (1); Manly, Long Reef, SAM K1909

(18); Port Jackson, Camp Cove, AM J20059 (1); Swansea Channel,

near Heads, 3 m, AM J21893 (4); Clovelly, NMV F93180 (9) (MOL

215, 218, 219, 220); Little Bay, AM J4794 (5); Shellhai'bour, rock

pools, AM J4419 (15); Jervis Bay, under rocks, 25 m, AM J15610 (1);

Ulladulla, 24 m, AM J14179 (1); Batemans Bay, 12 km S, Pretty Point

Bay, rocky shallows, NMV F71858 (1, cleared) (MOL 184); Montague

L, 27-30 m, AM J13972 (1); Merimbula, rocky, 9 m, AM J14651 (2);

Eden, Twofold Bay, sublittoral platform, AM J19853 (2). Vic.

Walkerville South, NMV F93448 (2); Western Port, McHaffie Point,

NMV F93175 (1); Flinders, ocean platforms, 0-2 m, NMVF92979 (1)

(MOL 175); NMV F92985 (2) (MOL 2); NMV F92980 (1) (MOL 5);

NMV F92981 (2) (MOL 116, 117); Port Phillip Bay, Portsea jetty;

under rubble, 4-5 m, NMV F73200 (1); Torquay, Point Danger, rocky

shallows, NMV F92976 (1, cleared) (MOL 65); Port Fairy, Griffith L,

rocky shallows, NMV F92978 (3) (MOL 77, 78, 80); NMV F92977 (1,

cleared) (MOL 71); Portland, Nelson Bay, 24 m, NMV F93462 (4).

Tas. Cape Tourville, rocky shallows, NMV F71870 (3); Maria L, 10 m,

TM H1792 (1); Forestier Peninsula, 4-9 m, SAM K1911 (4);

Eaglehawk Neck, TM HI 109 (2); Hobart, Tinderbox, under rocks, 2 m,

NMV E73993 (2); Port Davey, Sarah L, 3 m, TM H1789 (4); Bass

Strait, Ringarooma Bay, TM HI 114 (5); Tamar River, Low Head, TM

H1352 (1); Circular Head, TM H1765 (34); King L, Narracoopa, NMV

E93447 (1). SA. Encounter Bay, 2-4 m, SAM K1919 (1); Kangaroo L,

Western River, 10-12 m, SAM K1915 (1); Spencer Gulf, Gambler Is,

Wedge L, under stones, 1-1.5 m, AM J23763 (2); Nuyts Archipelago,

6 m, SAM K1937 (1). WA. Perth, Cottesloe, beach after stoma, TM

H2945 (1).

Description (dry and cleared specimens). Up to R = 39 mm;

4-8 rays, predominantly 6 (373 of 396 AM, SAM and TM

specimens with 6 rays, 18 with 7 rays, 3 with 5 rays, 1 with 4

rays, 1 with 8 rays); form variable, from 6 short pointed to

rounded rays with interradial margin incurved, to sub-

hexagonal; body flat orally, flattened dome aborally, acute

angle at margin; madreporite conspicuous; lacking

pedicellariae; gonopores abactinal.

Abactinal surface even; papulate areas more extensive than

non-papulate areas; secondary plates numerous, very irregular

in size and form; proximal radial and interradial plates fairly

openly imbricate; proximal papular spaces fairly large, fre-

quently 3-5 secondary plates and 4-6 papulae in proximal

papular spaces outside disc when R = 30 mm (2-4 secondary

plates and 4-5 papulae when R = 20 mm); abactinal plates cres-

centic in papulate areas, carinally with double notch and prox-

imal lobe; carinal series frequently regular from close to disc to

end or near end of rays, doubly papulate for more than half up

to three quarters ray length when R = 20 mm and larger; distal

interradial non-papulate plates closely imbricate, domed,

rounded proximally; disc variably distinct, bordering plates

variably regular crescentic radial and smaller interradial plates,

disc frequently obscured by irregular large plates within and

P. Mark O’Loughlin, Jonathan M Waters and Michael S. Roy

distal to disc; abactinal plates granular, covered by glassy

convexities, lacking spine-bearing ridge; abactinal spinelets

slightly less than twice as long as wide, variable form, not

widened distally, most frequently with swollen base and nar-

rowing distally, sometimes columnar, sometimes with slight

waist, rounded with small spines distally, typically up to 0.44

mm long when R = 30 mm (0.34 mm long when R = 20 mm),

distributed evenly over projecting surface of plates; supero-

marginal plates aligned distally with inferomarginal plates;

lacking internal superambulacral plates between ambulacrals

and actinals; distal abactinal and actinal interradial plates with

internal tapered vertical contiguous projections.

Projecting inferomarginal plates form margin, frequently

10-15 spinelets per plate when R = 20-30 mm; actinal plates in

regular series, curving acutely from furrow to margin; some

proximal actinal areas not calcified; actinal interradial spines

generally fairly thick, digitate, moderately tall; actinal inter-

radial proximal plates with 1-3 slightly tapered spines, pre-

dominantly 1, distally 3-4 short thick spines, columnar to

slightly tapered, spinous distally; adradial row of actinal inter-

radial proximal plates with predominantly 1-2 thick spines,

minutely spinous distally, slightly shorter than subambulacral

spines; adambulacral proximal plates with 1-3, predominantly

2, thick subambulacral spines, frequently unequal, form vari-

able from digitate to spatulate to widening distally, minutely

spinous distally, frequently slightly shorter than furrow spines;

furrow spines slender, tapering, webbed, minutely spinous dis-

tally, 2-4 per plate proximally, frequently slightly longer than

subambulacral spines; suboral spines very rare (9 of 273 AM

speciems with at least 1 suboral spines, 1 with 7 spines, 1 with

9 spines, 1 with 11 spines); oral spines 4-6, predominantly 5.

Live colour. Abactinally very variable; frequently pale or light

coloured; commonly dark coloured disc; some fairly uniform

white or pink or mauve or orange or bright red, sometimes with

dark red or brown or white flecks; some with rays, interradii or

margin coloured differently; some finely mottled with brown,

red, mauve, pink, orange, yellow, green, white (NSW speci-

mens sometimes with grey); actinally typically off-white with

rare to sparse colour flecks.

Distribution. Lord Howe L; Rockhampton (Qld) continuous to

Nuyts Archipelago (western SA); Cottesloe, Perth (WA); Bass

Strait; Tas.; under rocks; 0-30 m (molecular confirmation for

Batemans Bay (NSW) to Port Fairy (Vic.)).

Etymology. From oriens (Latin, as a noun in apposition) mean-

ing “east”, and referring to the uniquely easterly distribution in

Australia.

Remarks. Hart et al. (1997) and Byrne et al. (1999) found from

molecular evidence that specimens of Patiriella ‘gunnii ’ from

eastern and western Australia had divergent lineages. The east-

ern material was collected from Clovelly, Sydney (M. Byrne,

pers. comm.). Eastern Australian specimens are confirmed by

molecular and morphological evidence and described here as

Patiriella oriens sp. nov. Byrne (1991, 1992, 1995, 1996),

Byrne and Anderson (1994), Byrne and Cerra (1996), Cerra

and Byrne (2001), and Long and Byrne (2001) reported

Review of the sea-star, Patiriella gunnii

193

extensively on the reproductive and developmental biology of

Patiriella ‘gunnii’. Since this research was based on material

collected from Clovelly (M. Byrne, pers. comm.), the species

was not P. gunnii but the new species P oriens (used hereafter

for P. ‘gunnii’ from Clovelly).

Byrne (1992) reported broadcast spawning during spring

and summer and some habitat overlap for Clovelly populations

of P oriens (typically under subtidal boulders) and P calcar

(typically intertidal reef). Byrne and Anderson (1994) sub-

sequently reported viable laboratory hybrids (high frequency of

seven rays) of P oriens (typically six rays) and P. calcar (typic-

ally eight rays). Field and museum specimens of P oriens

observed by Byrne and Anderson (1994), and material seen in

this Study, showed a low frequency of seven rays, suggesting

variation of arm number (within the species) or hybridization or

both. Relevant to this consideration is the fact that about 10%

of specimens of P occidens (above) have more than six arms

across a distribution range where P. calcar is mostly absent.

The spawning of P oriens is earlier (spring and summer) than

P. occidens (late summer, discussed above), a factor which may

be significant in the maintenance of genetic identity in these

similar species.

In this study a single specimen (TM H2945) which was col-

lected on a beach at Cottesloe (Perth) after a storm was identi-

fied as P. oriens. It is the only specimen in Australian museums,

determined as P. oriens, which has been collected west of

Nuyts Archipelago.

The combination of morphological characters which distin-

guishes P. oriens from other species of six-rayed Patiriella is:

frequently subhexagonal form; carinal series of plates frequent-

ly doubly papulate for about two-thirds ray length; abactinal

spinelets frequently columnar and moderately spinous distally,

creating a fairly coarsely spinous surface appearance; normal

absence of suboral spines; furrow spines frequently slightly

longer than subambulacral spines; actinal spines continuous in

declining height with subambulacral spines; actinal interradial

spines digitate; up to about 15 spinelets per inferomarginal

plate. The limited live colour data available for confirmed

determinations indicates that white or pink or mauve or

orange or bright red, with a dark centre, are frequently evident

abactinally.

Discussion

Byrne et al. (1999) concluded that “P. gunnii from eastern and

western Australia are different species, while P. brevispina

from eastern and western Australia are the same species”.

These conclusions were premature as they were based on rela-

tive levels of mtDNA divergence rather than any direct assess-

ment of reproductive isolation (biological species concept,

Dobzhansky, 1937; Mayr, 1942), or monophyly (phylogenetic

species concept, Cracraft, 1983). Nevertheless, Byrne et al.

(1999) were correct that multiple species are present: inde-

pendent morphological and molecular data reveal consistent

evidence for three new taxa. The newly recognised taxa are

deeply divergent for mtDNA, but exhibit only subtle morpho-

logical differences. This discrepancy might reflect the lack of

morphological novelty typically associated with the echino-

derm body plan (Janies, 2001), but could also stem from rapid

coalescence of mitochondrial genes relative to nuclear genes

(Palumbi et al., 2001).

Although the overlapping distributions of these taxa (Fig. 7)

provide compelling evidence for some form of reproductive

isolation, we cannot rule out the possibility of a small degree of

hybridization among some or all of the species in zones of

sympatry. Future ecological and genetic studies should help

elucidate the strength and nature of reproductive barriers.

Parallel analyses of molecular and moiphological characters

represent a powerful technique for asteroid systematics

(Flowers and Foltz, 2001; O’Loughlin et al., 2002).

Acknowledgements

We are grateful to: Penny Berents (AM), Clare Valentine and

Miranda Lowe (BMNH), Thierry Laperousaz (SAM), Liz

Turner (TM), and Jane Fromont, Loisette Marsh and Mark

Salotti (WAM) for assistance in accessing museum collections;

Neville Barrett, Maria Byrne, Georgina Cunningham, Clarrie

Handreck and the Marine Research Group (Victoria), Rebecca

McIntosh, Melanie Mackenzie, Tim O’Hara, Janine Ortenburg

and the numerous other fieldworkers who did the collecting of

many of the specimens on which this work has been based;

Leon Altoff, Audrey Falconer and Loisette Marsh for providing

data and the loan of colour slides; Cynthia Ahearn (United

States National Museum) for assistance with literature; Chris

Rowley (NMV) for technical assistance; Mark Darragh for

photography; Ben Boonen for the presentation of the photo-

figures; and referees Tim O’Hara and Frank Rowe for their

much appreciated critique of the manuscript.

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Memoirs of Museum Victoria 60(2): 197-206 (2003)

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

http://www.museum.vic.gov.au/memoirs

A new genus of Tasmanian millipedes (Diplopoda: Polydesmida: Dalodesmidae)

with unusual spiracles and a mosaic distribution

Robert Mesibov

Queen Victoria Museum and Art Gallery, Launceston, Tasmania 7250, Australia (mesibov@southcom.com.au)

Abstract Mesibov, R. 2003. A new genus of Tasmanian millipedes (Diplopoda: Polydesmida: Dalodesmidae) with unusual spira-

cles and a mosaic distribution. Memoirs of Museum Victoria 62(2): 197-206.

Dasystigma gen. nov. is erected for Lissodesmus margaretae Jeekel, 1984 (type species), D. bonhami sp. nov.,

D. huonense sp. nov. and D. tyleri sp. nov. A dense cluster of hair-like structures of unknown function emerges from each

spiracle in all Dasystigma species, and the four geographic distributions form a closely fitted mosaic.

Keywords Diplopoda, Polydesmida, Dalodesmidae, millipede, taxonomy, Australia, Tasmania

Introduction

R.M. Shelley suggested that distribution mosaics may be com-

mon in the Diplopoda. Mosaics of closely related species, here

called lineage mosaics, are “typically characterized by large

generic distributions, limited component ranges, tightly fitted

parapatry patterns with limited instances of sympatry, cluster-

ing of components, and slight or minor differences between

some species” (Shelley, 1990a: 23). A lineage mosaic of five

parapatric and two allopatric species has been documented in

the Tasmanian polydesmidan genus Gasterogramma (Mesibov,

2003). In the Gasterogramma mosaic, the components are read-

ily distinguished by cursory examination of the male gonopods,

i.e. the species differences are not “minor”.

Here, I describe a group of four similar Tasmanian

dalodesmids forming a closely fitted lineage mosaic. The group

is also unusual in having hair-like structures of unknown func-

tion emerging from all spiracles. A new genus Dasystigma is

erected for the group, based on Lissodesmus margaretae Jeekel,

1984.

As is also the case for components in North American mil-

lipede mosaics (Shelley and Whitehead, 1986; Shelley, 1990b),

it is not yet known whether the four Dasystigma taxa

recognised here are fully reproductively isolated. Although D.

margaretae and D. tyleri sp. nov. have been found in sympatry

in south-western Tasmania (with D. huonense sp. nov. known

from a locality only 5 km distant), the overlap zone appears to

be narrow and may be a tension zone (Key, 1982) maintained

by hybridisation, i.e. isolation is incomplete. I regard the four

Dasystigma as evolutionary species sensu Wiley (1978).

Specimens were killed and preserved in 75-80% ethanol.

Preliminary drawings on graph paper were made using material

cleared in 60% lactic acid and viewed at 100 or 200x magnification

through an eyepiece graticule. A Philips Electroscan ESEM 2020 oper-

ated in high-vacuum mode was used to examine preserved material

which had been air-dried before sputter-coating with gold. SEM

images were acquired digitally.

To save space in the printed version, full details of localities, dates,

collectors, specimens and registration numbers for the 202 samples

examined are provided separately on the Memoirs of Museum Victoria

website, www.museum.vic.gov.au/memoirs/. The specimen data table

is also available from the author and a copy has been deposited at the

QVM.

Collecting sites for all but a few of the specimens examined were

estimated in the field to be within particular 100 m Universal

Transverse Mercator (UTM) grid squares on 1:25000 scale maps pub-

lished by the State of Tasmania. Grid squares are recorded below in

2-letter, 6-digit form, e.g. ‘EN700712.’ The maximum horizontal error

in these estimates is likely to be less than 100 m. Latitude/longitude

equivalents were calculated using GeoCalc 4.20 (GeoComp Systems,

Blackburn, Victoria) and are not as precise as the UTM grid references.

LGRSS transect locations (see separate specimen data table) were

derived from 1:2000 survey charts made available to the QVM by the

Hydro-Electric Commission, Tasmania, in 1994.

Abbreviations and codes are as follows: AM, Australian Museum,

Sydney, NSW; ANIC, Australian National Insect Collection, Canberra,

ACT; LTV Latrobe University, Bundoora; LGRSS, Lower Gordon R.

Scientific Survey, 1976-1978; NMV, Museum Victoria, Melbourne,

Vic.; NRCP, National Rainforest Conservation Program invertebrate

survey, 1989-1990; PCS, posterior corner seta (a long, prominent seta

arising dorsally near each posterior corner of the collum and all ter-

gites); QVM, Queen Victoria Museum and Art Gallery, Launceston,

Tas. Male and female refer to stadium 8 adults.

198

Robert Mesibov

Order Poly desmida Leach, 1815

Suborder Dalodesmidea Hoffman, 1977

Dalodesmidae Cook, 1896

Dasystigma gen. nov.

Type species. Lissodesmus margaretae Jeekel, 1984, by present

designation.

Diagnosis. Differs from Lissodesmus and other known

Australian dalodesmids in (a) the dense brush of hair-like

structures emerging from each spiracle and (b) the unusually

wide separation between the bases of the solenomerite and

tibiotarsus on the gonopod telopodite.

Description of males. Adult length 18-22 mm when contracted

in alcohol, diameter of midbody metazonite c. 1.8 mm. Overall

colour pale yellow-brown to deep chestnut brown. Head with

labrum weakly emarginate in center; clypeus very weakly con-

vex in lateral view, moderately setose; vertex bare, strongly

convex in lateral view, vertigial sulcus extending forward to a

point about one antennal socket width from an imaginary line

joining socket centres. Antennal sockets separated by about

twice the diameter of a socket, weakly impressed; anten-

nomeres (Fig. 2) setose, more densely and finely so on 5-8,

antennomere lengths decreasing in the order 2, (3, 6), (5, 4),

antennomere 6 the widest. Collum slightly wider than head in

front, widening posteriorly, anterior margin broadly convex,

lateral margin with typically 3 small, seta-bearing teeth, pos-

terior corner rounded and not projecting, posterior margin

squarely transverse laterally but with the central third slightly

emarginate; several transverse rows of sparse, long setae anter-

Figure 1. Whole-animal view of Dasystigma margaretae (Jeekel,

1984) comb. nov. A, Rocka Rivulet male, QVM 23:24949; B, Tiger

Creek male, QVM 23:24950. Scale bar = 10 mm.

Figure 2. Dasystigma bonhami sp. nov., Sandspit R. male, QVM

23:15219. A, Antenna; B, leg 10; C, cross-section of somite 8. Setation

not shown.

iorly on collum; a long seta extending posteriolaterally from

point near posterior corner of collum (= posterior corner seta,

PCS). Paranota inflated (Fig. 2), maximum width at about one-

third the ring diameter from the dorsum in midbody segments.

Somites 2-4 from above about equal in width and slightly

wider than collum; somites 5-17 about equal in width and

slightly wider than 2-A\ somite 18 narrower than 17. Tergites

unsculptured, bare apart from PCS near posterior corners (Fig.

3). Paranota on most somites with 4 or 5 (3-6) small marginal

teeth, each bearing a seta (Fig. 3); paranotal margin a straight

line in lateral view, rising posteriorly; margin in dorsal view

either nearly straight (parallel to long body axis) or slightly

convex (see also “Derwent form” under D. margaretae (Jeekel,

1984) comb, nov., below); posterior corner variably projected

(Fig. 3), with minute terminal seta. Ozopores on somites 5, 7,

9, 10, 12, 13, 15-19; pore opening dorsally on paranotum, just

mesal to marginal thickening and typically about one-fourth of

lateral margin length from tip of posterior corner. Spiracles

(Fig. 5) variably enlarged, all with hair-like structures variably

emergent (Fig. lA) (at low magnification, the swollen, “hairy”

spiracles in D. bonhami and D. margaretae resemble ectopara-

sitic mites). Legs (Fig. 2) incrassate, much more so anteriorly

beginning with leg-pair 3, prefemur and femur dorsally

swollen, tibia on anterior legs in D. bonhami, D. margaretae

and D. tyleri with slight ventrodistal swelling; tarsus about as

long as or slightly longer than femur; dense pubescence ven-

trally on coxa, prefemur, femur and postfemur; numerous

sphaerotrichomes ventrally on postfemur, tibia and tarsus; long,

prominent seta at ventrodistal end of coxa and prefemur and at

dorsodistal end of tibia. Genital opening inconspicuous on

Dasystigma, a new genus of Tasmania millipede

199

Figure 3. Dorsal views of right margin of tergite 14 of Dasystigma females. A, D. bonhami sp. nov., Sand R., QVM 23:40811; B, D. huonense sp.

nov., Edwards Rd, QVM 23:15269; C, D. margaretae (Jeekel, 1984) comb, nov., Fingal Tier, QVM 23:15230; D, D. margaretae (Jeekel, 1984)

comb, nov., Tarraleah, QVM 23:15252; E, D. tyleri sp. nov., Trackham Creek, QVM 23:15260. PCS = posterior corner seta.

slight distal swelling of leg 2 coxa. Preanal ring with numerous

long setae, densest dorsahy; epiproct in dorsal view a trun-

cated triangle with weakly concave sides; hypoproct broadly

paraboloid in ventral view.

Gonopod aperture one-third to one-half ring 7 prozonite

diameter in width, about 1.5 times as wide as long; in ventral

view with straight anterior and lateral margins, posterior mar-

gin slightly curved, concave anteriorly; in lateral view anterior

aperture margin not raised, lateral margin not raised or slightly

convex upwards, and higher than slightly raised posterior

margin.

Gonopods (Fig. 4) retracted reaching as far as leg-pair 5

bases on ring 5, solenomerites and tibiotarsi of opposing

gonopods interlaced. Telopodite in posterior view more or less

straight, posterior and mesal faces sparsely setose from base to

about level of solenomerite origin. Solenomerite arising at

about half telopodite length on anteriomesal face, just proximal

to origin of prefemoral process, directed first distad and mesad,

then curving caudad and distad, tapering to a sharp point from

about two-thirds its length and terminating at about three-

quarters length of telopodite. A thin, curved ridge of cuticle on

anterior surface of telopodite appearing to extend the line of the

solenomerite proximad and strengthening it at its base; pro-

static groove running along anterior surface of telopodite Just

lateral of this ridge. Tibiotarsus arising on posterior face of

telopodite at about level where prefemoral process arises,

smoothly curving mesad and distad, tapering near its apex to a

blunt point on the telopodite just proximal to tip of

solenomerite. Prefemoral process arising about midlength on

telopodite, curved (concave posteriorly) and flattened anterio-

posteriorly, bearing a large uncus on posterior surface at about

half its length, tip of uncus pointed caudad and mesad. Femoral

process arising from lateral surface of prefemoral process

proximal to uncus, variably shaped, not extending further

distad than prefemoral process.

Females longer and heavier-bodied than males. Legs not

swollen apart from slight dorsal swelling on prefemur and

femur on anterior leg-pairs; no ventrodistal swelling of tibia;

no sphaerotrichomes or ventral pubescence. Cyphopods not

examined.

Juveniles considerably smaller than adults, midbody meta-

zonite diameters c. 1 mm in stadium VII and c. 0.8 mm in

stadium VI. Paranotal teeth much more prominent than in

adults, spiracles generally placed as in adults (see

species descriptions, below) in stadium VII, but typically well-

separated in stadium VI and younger.

Figure 4. Mesal view of right gonopod of male holotype of

Dasystigma margaretae (Jeekel, 1984) comb, nov., redrawn from

Jeekel (1984). f = femoral process, pf = prefemoral process, s =

solenomerite, tt = tibiotarsus, u = uncus. Dashed line marks path of

prostatic groove.

200

Robert Mesibov

Figure 5. Spiracles on midbody segment of A, Dasystigma bonhami sp. nov., Sandspit R. male, QVM 23:15219; B, D. huonense sp. nov.,

Huon R. (Arve Rd) male paratype, QVM 23:40805; C, D. margaretae (Jeekel, 1984) comb, nov., Tooms Lake male, QVM 23:15214; D,

D. margaretae (Jeekel, 1984), Dromedary Creek male, QVM 23:15212; E, D. tyleri sp. nov.. White Spur male, QVM 23:15190. Scale bar in all

cases = 0.25 mm; anterior to right for A-D, anterior to left for E.

Dasystigma, a new genus of Tasmania millipede

201

Figure 6. Dasystigma bonhami sp. nov. Gonopods in situ. Scale bar =

0.5 mm. Flash Tier male, QVM 23:15222.

Figure 8. Dasystigma huonense sp. nov. Gonopods in situ. Scale bar =

0.5 mm. Picton R. male, QVM 23:40801.

Figure 7. Dasystigma bonhami sp. nov. Approximately lateral (left)

and mesal (right) views of left gonopod telopodite of Ravens Hill

male, QVM 23:15223. Setation not shown.

Figure 9. Dasystigma huonense sp. nov. Approximately lateral view

(left) of left gonopod telopodite and mesal view (right) of right gono-

pod telopodite of Edwards Rd male paratype, QVM 23:15195.

Setation not shown.

202

Robert Mesibov

Figure 10. Dasystigma margaretae (Jeekel, 1984) comb. nov. Gonopods in situ, scale bar = 0.5 mm in all cases. A, Tooms Lake male, QVM

23:15214; B, Lake Augusta male paratype, (DPI-NT) 19A17; C, Coles Creek male (“Derwent form”), QVM 23:24932.

Remarks. The four species of Dasystigma recognised here are

very similar in overall appearance (Fig. 1), distinguished partly

on differences in the size, placement and “hairiness” of spira-

cles on diplosegments, but primarily on details of gonopod

structure. The various processes on the gonopod are named

here in accordance with the terminology used by Jeekel (1984)

for Lissodesmus margaretae. Interspecific differences in

gonopod structure are consistent in Dasystigma but subtle, and

I have therefore provided for each species three different views

of the gonopod, emphasising the prefemoral and femoral

processes.

The dense spiracular “brush” of hairs is present in the type

specimens of D. margaretae but appears to have been over-

looked by Jeekel (1984). In 1972, P.M. Johns collected speci-

mens of D. bonhami near Triabunna, in south-eastern Tasmania

and later noted “spiracles greatly swollen, densely setose, the

setae fine and short” (Johns, in litt., 15 Oct 1991). The function

of the hair-like structures is unknown. Throughout their ranges,

Dasystigma species co-occur in forest litter with dalodesmids

of similar size and habits but with non-hairy spiracles.

Distribution and microhabitat. Tasmania south of 4TS; in and

under rotting wood, in leaf litter and in humus.

Etymology. Greek dasys (“hairy”) -i- stigma (in entomology,

“opening to tracheal system”), neuter.

Dasystigma bonhami sp. nov.

Figures 2, 3A, 5A, 6, 7, 14 (map)

Material examined. Holotype. Male, Australia, Tasmania. Sandspit R.,

EN700712 (42°4230'^S 147°51T7'T), 230 m, 31 Jul 1991, R.

Mesibov, QVM 23:41726.

Paratypes. 2 males, details as for holotype, QVM 23:15219; 1 female,

details as for holotype, QVM 23:15261; 2 males, Sandspit R., EN688712

(42°4230^'S UT5t)'2A"E), 200 m, 26 Jun 1988, R. Mesibov, NMV

K-8803, K-8804 (formerly QVM 23: 15220); 2 males, Nugent, EN559711

(42°4237^'S 147°40'57^T), 400 m, 9 Aug 1998, K. Bonham and R.

Crookshanks, AM KS85095 (formerly QVM 23:40807).

Other material. 30 males, 25 females and 45 juveniles from 27

unique localities including Baldy Creek, Bellettes Creek, Bishop and

Clerk (Maria L), Black Hill, Blind Creek (Maria L), Blue Gum Spur,

Carlton R., Chauncy Vale, Douglas Creek, Elash Tier, Macgregor Peak,

Maclaines Creek, Mother Browns Bonnet, Mt Walter, Ravens Hill,

Sand R., Sandspit R., Sheepdip Creek and Three Thumbs.

Diagnosis. Differs from other Dasystigma in its slender, upright

femoral process with a small mesal spike, from D. huonense

and D. tyleri in having posterior spiracle on diplosegments

located above anterior leg, and from D. tyleri in having large

spiracles (unusually large for dalodesmids) with much greater

spiracular “hairiness.”

Description. As for the genus except in the following details.

Both spiracles on diplosegments (Fig. 5A) positioned over

anterior leg, hair-like structures emergent from spiracles and

apparent at low magnification, spiracles unusually large for a

dalodesmid. Paranotal margin very slightly convex; posterior

comer projected caudad (Fig. 3A). Gonopod telopodite (Figs 6,

7) with prefemoral process narrowing slightly distad, apex

curving caudad, with 2 teeth on lateral edge near apex and

single tooth on mesal edge more proximad. Femoral process

projecting parallel to prefemoral process and terminating just

proximal to flexed apex of latter. Femoral process a narrow,

somewhat flattened rod with a few, minute terminal teeth, a

small, slender spike arising at about three-quarters of process

length and projecting mesad.

Distribution and macrohabitat. Common in dry and wet

eucalypt forest over c. 2000 km^ in south-eastern Tasmania,

from Campania east to Maria I. and from the Forestier

Peninsula north to the Little Swanport River valley (Fig. 14); c.

100-600 m elevation.

Etymology. In honour of the Tasmanian malacologist Kevin

J. Bonham, a very talented collector whose “bycatch” of

millipedes nearly always contains specimens of interest.

Remarks. D. bonhami varies little in size and form across its

range. However, even syntopic adults differ considerably in the

depth of dorsal body coloration, with some pale and others

honey- or chestnut-coloured.

Dasystigma, a new genus of Tasmania millipede

203

Figure 11. Dasystigma margaretae (Jeekel, 1984) comb. nov.

Approximately mesal views of left gonopod telopodite of (left) Lake

Augusta male paratype (DPI-NT) 19A17 and (right) Tarraleah male

(“Derwent form”), QVM 23:15203. Setation not shown.

Dasystigma huonense sp. nov.

Figures 3B, 5B, 8, 9, 14 (map)

Material examined. Holotype. Male, Australia, Tasmania. Edwards Rd,

DN797310 (43°04^23^'S 146°45^02'T), 110-130 m, 15 Mar - 6 Apr

1988, R. Mesibov, QVM 23:41728.

Paratypes. 3 males, details as for holotype, QVM 23:15195; 1 male,

Huon R. (Arve Rd), DN788280 (43°06^00'^S 146°44^22^T), 150 m,

22 May 1997, R. Mesibov, plot 3M5, QVM 23:40805; 1 female, Huon

R. (Manuka Rd), DN769286 (43°05AL^S 146°42^58^T), 100 m, 29

Apr 1997, R. Mesibov, plot 1R4, QVM 23:40800; 2 males. Coopers

Creek, DN507635 (42°46A5^^S 146°23^50^T), 460 m, 23 Feb 1994,

R. Mesibov, AM KS 85096 (formerly QVM 23:15199); 1 male, Picton

Valley, DN737146 (43°13H4^^S 146°4034^T), 300 m, 15 Jul 1994,

K. Michaels, pitfall collection, NMV K-8806 (formerly QVM

23:21294).

Other material. 13 males, 18 females and 18 juveniles from 22

unique localities including Collins Cap, Coopers Creek, the

Huon/Picton Rivers junction, the Huon R. crossing on the Port Davey

track. Lake Sydney, Mt Mangana and Mt Tobin (Bruny L), Mystery

Creek Cave environs. Palmers Lookout, Picton R., South Cape Bay,

Waterfall Bay and Arve, Edwards and Manuka Rds in the vicinity of

Tahune Bridge on the Huon R.

Diagnosis. Differs from other Dasystigma in the sinuous curve

of lateral edge of prefemoral process and corresponding curva-

ture of apposed femoral process; from D. bonhami and D. mar-

garetae in having posterior spiracle on diplosegments located

between anterior and posterior legs; from D. tyleri in lacking a

toothed anterior margin on femoral process and in closer

Figure 12. Dasystigma tyleri sp. nov. Gonopods in situ. Scale bar = 0.5

mm. Wedge Inlet male, QVM 23:24953.

spacing of spiracles on diplosegments, spiracles being larger

than those in D. tyleri (or in other dalodesmids).

Description. As for the genus except in the following details.

Posterior spiracle on diplosegments (Fig. 5B) between anterior

and posterior leg, hair-like structures emergent from spiracles;

spiracle ‘hairiness’ and separation of spiracles both apparent at

low magnification, spiracles unusually large for a dalodesmid.

Paranotal margin slightly convex; posterior corner projected

caudad and slightly laterad (Fig. 3B). Gonopod telopodite (Figs

8, 9) with prefemoral process narrowing slightly distad and

strongly flattened anterioposteriorly, the apex broadly rounded

with a small terminal notch, mesal edge of prefemoral process

straight, lateral edge sinusoidal, convex near apex with a small

tooth projecting caudad. Femoral process curving mesad in par-

allel with concavity on lateral edge of prefemoral process, and

terminating about halfway between thickened uncus and apex

of prefemoral process. Femoral process flattened with a few,

minute terminal teeth, a short thick spike arising at about three-

quarters of process length and projecting mesad and distad.

Distribution and macrohabitat. In wet eucalypt forest and rain-

forest over c. 6000 km^ in southern Tasmania including South

Bruny L, from Tasman Peninsula south to South Cape and west

to the vicinity of Lake Pedder (Fig. 14); from near sea level to

c. 700 m elevation.

Etymology. After the Huon River in southern Tasmania.

Remarks. Dasystigma huonense varies very little in size and

form across its range. Adult colour varies considerably, from

very pale yellow-brown to deep chestnut brown.

Dasystigma margaretae (Jeekel, 1984) comb. nov.

Figures 1, 3C, 3D, 4, 5C, 5D, 10, 11, 14 (map)

Lissodesmus margaretae Jeekel, 1984: 99.

Material examined. Holotype and paratypes. Australia, Tasmania.

“Lake Augusta, 25. IV. 1979 [on cushion plant] 25 April 1979,

204

Robert Mesibov

Tasmanian Department of Agriculture 19A17, holotype, 6,1$

(fragm.), 2 juv. $ (19 somites), 1 juv. $ (18 somites) paratypes” (Jeekel,

1984: 99). When I inspected the type-containing vial in 2001, 1 found

two cotton-plugged genitalia tubes and (under a cotton pad) a number

of body fragments. The genitalia tubes contained the male holotype

and the fragmented mature female paratype, respectively, both in good

condition. After removing a paratype male segment 7 for SEM exami-

nation, I placed the remaining body fragments in a small cotton-

plugged glass tube. The three tubes and all accompanying labels were

then sealed in an alcohol-filled McCartney vial for continuing storage

at the New Town Laboratories of the Tasmanian Department of

Primaiy Industries, Water and Environment (formerly the Department

of Agriculture).

Other material. 97 males, 66 females and 72 juveniles from 52

unique localities including Alberts Marsh, Anglers Creek, Blackman

R., Blue Tier Creek, Boyer, Brumbys Creek, Butlers Gorge, Coal

Marsh, Coles Creek, Dromedary Creek, Eingal Tier, Elexmore Creek,

Elorentine R., Gulf Creek, Halls Creek, Lake Dobson, Liawenee, Little

Elorentine R., Lookout Hill, Lost Ealls, Meehan Range, Mossy Marsh

Creek, Mt Mismanagement, Native Tier, Old Mans Head, Pinnacles

Creek, R. Dee, Rocka Rivulet, Sassafras Hill, St Pauls Dome,

Tarraleah, Tiger Creek, Tooms Lake, Tooms White Gum Reserve and

Yangena Hill.

Diagnosis. Differs from D. bonhami in having a broader, more

flexed and more prominently toothed femoral process; from

D. huonense and D. tyleri in having the posterior spiracle on

diplosegments located above the anterior leg; from D. tyleri in

having much “hairier” spiracles, spiracles also being larger than

those in D. tyleri (or in other dalodesmids).

Description. As for the genus except in the following details.

Both spiracles on diplosegments (Fig. 5C) positioned over

anterior leg, hair-like structures emergent from spiracles and

apparent at low magnification, spiracles unusually large for a

dalodesmid. Paranotal margin slightly convex; posterior comer

variably projected (Figs 3C, D; see Remarks). Gonopod

telopodite (Figs 10, 11) with prefemoral process bent mesad,

sharply narrowing distad, apex curving caudad, apical edge

slightly serrulate, a single tooth on mesal edge of process at

about three-quarters of process length. Femoral process curving

caudad and mesad and terminating just proximal to level of

tooth on mesal edge of prefemoral process. Femoral process

massive, divided into a broadly cuneate anterior portion and a

posterior spike; distal and posterior margin of cuneate portion

with numerous heavy teeth, spike crossing below cuneate

portion in manner of a thumb bent slightly towards palm.

Distribution and macrohabitat. Common in dry and wet euca-

lypt forest and in subalpine woodland over c. 12 000 km^ in

eastern and central Tasmania (Fig. 14), from the south side of

the Fingal Valley south to the north side of the Little Swanport

R. valley, and from near the east coast west to the Little

Florentine R.; so far known at altitudes c. 100-1100 m ele-

vation. Locally abundant in places in the Eastern Tiers, on the

eastern fringe of the Central Plateau and in wet forests in the

Derwent valley.

Remarks. Jeekel (1984) gave a complete description of this

species from the type locality. Lake Augusta on Tasmania’s

Central Plateau; for the sake of consistency I have included my

own summary. This taxon is the most variable within

Figure 13. Dasy stigma tyleri sp. nov. Approximately lateral (left) and

mesal (right) views of left gonopod telopodite of Franklin R. male,

QVM 23:15189. Setation not shown.

Dasystigma. The prefemoral process is straighter and narrower

in the eastern portion of the range (Fig. lOA) than in the west

(Fig. lOB) and the femoral process less bent mesad. A distinc-

tive variant, here called the “Derwent form,” is restricted to the

valley of the Derwent R. (Fig. 14), where it replaces more

typical D. margaretae. In the “Derwent form” the spiracles are

somewhat less “hairy” (Fig. 5D) and the prefemoral process

somewhat broader (Figs IOC, 11). The most striking difference,

however, is in the form of the paranota: the margins have

moved ventrad relative to those in typical Dasystigma and the

posterior corners are very strongly projected caudad and

laterad (Fig. 3D). In future, genetic data may justify the erec-

tion of a new species for this geographically and morpho-

logically distinctive variant. I am reluctant at this time to name

the “Derwent form” formally, as it differs less from more

typical D. margaretae in gonopod details than do the three

forms recognised here as new species.

Dasystigma tyleri sp. nov.

Figs 3E, 5E, 12, 13, 14 (map)

Material examined. Holotype. Male, Australia, Tasmania.

Donaghys Hill, approx. DP120270 (42°12'S 145°560, 480 m, 29 Apr

1987, N. Platnick, R. Raven and T. Churchill, QVM 23:41729.

Paratypes. 1 male. Little Florentine R., DN525683 (42°44T0""S

146°25T0''E) 440 m, 2 Dec 1986, R. Bashford, QVM 23:40796; 1

Dasystigma, a new genus of Tasmania millipede

205

male, Trackham Creek, CQ822092 (41°2736^'S 145°35'22''E), 630

m, 28 Oct 1991, R. Mesibov, QVM 23:15180; 1 female. Wedge Inlet,

DN379569 (42°50'16''S 146°14^24'T), 350 m, pitfall emptied 16

Nov 2001, D. Driscoll, sample EY4-75, QVM 23:24954.

Other material. 16 males, 4 females and 9 juveniles from 23 unique

localities including Acheron Cave environs, Algonkian Mountain,

Dismal Creek, Dohertys Range, Eranklin R., Goderich Rd, Gordon R.,

Hardwood R., Hermit Hill, Laughing Jack Lagoon, Little Elorentine

R., Loddon R., Mt Rufus, Olga R., The Clump, Trackham Creek,

Wakefield Creek, Wedge Inlet and White Spur.

Diagnosis. Differs from other Dasystigma in having a

heavily toothed femoral process with teeth on anterior margin

and small spiracles with no “hairiness” visible at low

magnification, with posterior spiracle located just anterior to

posterior leg on diplosegments.

Description. As for genus except in following details. Posterior

spiracle on diplosegments (Fig. 5E) positioned just anterior to

posterior leg and very clearly separated from anterior spiracle;

hair-like structures only just emergent from spiracles, not

apparent at low magnification; spiracles of size typical for

dalodesmids. Paranotal margin slightly convex; posterior cor-

ner projected caudad and slightly mesad (Fig. 3E). Gonopod

telopodite (Figs 12, 13) with prefemoral process narrowing

sharply distad, truncated apex curving caudad and shallowly

notched; on posterior surface a small tooth near apex; uncus

apparently bifid, with second, caudally projected tip arising

from its lateral edge. Femoral process arising in small depres-

sion on lateral surface of telopodite, projecting distad and

slightly mesad. Femoral process massive, divided into broadly

lanceolate anterior portion and robust posterior spike; entire

margin of lanceolate portion with numerous heavy teeth, spike

extending just past most distal teeth and nearly reaching as far

distad as apex of prefemoral process.

Distribution and macrohabitat. An uncommon species in rain-

forest and wet eucalypt forest over c. 11 000 km“ in western

Tasmania, from Lake Pedder north to Balfour in a band 70-90

km wide extending inland from the west coast (Fig. 14); c.

50-1000 m elevation.

Etymology. In honour of the Australian limnologist Peter A.

Tyler, whose investigations in Tasmania led to the recognition

of the biogeographic divide known as Tyler’s Line.

Remarks. There is little morphological variation over the

D. tyleri range, but mature specimens tend to be somewhat

larger and more heavily pigmented in northwest Tasmania than

in the Southwest.

Dasystigma sp.

A number of female and juvenile specimens cannot yet be

assigned with confidence to any of the named species; this

unidentified material is shown as ''Dasystigma sp.” in the spec-

imen data table and the distribution map (Fig. 14). Males from

Little Quoin (Yarlington Tier) seem closest to D. huonense, yet

this locality, an isolated forest fragment, lies between the

ranges of D. bonhami and D. margaretae. The Little Quoin and

Boyd R. material includes DNA vouchers in absolute ethanol,

and genetic analysis can be used in future to clarify taxonomic

placement.

Biogeography and conservation

Where species of Dasystigma are not locally abundant, they

can be hard to find and it has so far not been possible to map

range boundaries on as fine as scale as has been done for other

Tasmanian dalodesmids (Mesibov, 1997, 1999). It seems

likely, however, that the apparently narrow parapatric boundary

between D. tyleri and D. margaretae in western Tasmania (Fig.

14) is congruent with Tyler’s Line (Mesibov, 1994), a major

biogeographic divide in Tasmania which is also commonly a

species boundary for millipedes. An uncertain divide in eastern

Tasmania is the one between D. bonhami on Forestier

Peninsula and D. huonense on Tasman Peninsula (Fig. 14); it is

not yet known whether the two species meet in parapatry on

one or the other of the peninsulas, or are separated by the nar-

row strip of interpeninsular land (Eaglehawk Neck). Also

uncertain is the gap between D. bonhami and D margaretae in

the valley of the Little Swanport River. The upstream portion of

the valley now carries agricultural grassland and is unsuitable

Dasystigma habitat, and access difficulties have so far limited

sampling in the woodlands on the lower portion of the river.

The nearest currently known localities for D. bonhami and

Figure 14. Known localities (to Jun 2002). Dasystigma bonhami sp.

nov., A. D. huonense sp. nov., #. D. margaretae (Jeekel, 1984) comb,

nov., ■. D.margaretae (Jeekel, 1984) comb. nov. “Derwent form”,

Q D. tyleri sp. nov., +. Unidentified Da.vy.vrig/nfl species, •. Polygons

have been drawn through outermost known localities of each of the

four named species to more clearly delineate the species ranges. Arrow

marks Little Quoin (see text).

206

Robert Mesibov

D margaretae are 14 km apart. About 12 km currently separates

the nearest localities for D. huonense and D. margaretae

(“Derwent form”) across the Derwent River valley just down-

stream from New Norfolk. Forest in this portion of the Derwent

valley has been largely cleared or degraded, and a search for

evidence of D. huonense/D. margaretae overlap should begin

much further up the Derwent system; the two species have been

collected only 5 km apart just west of Maydena. Another map-

ping exercise for the future is documentation of the apparent

parapatry (Fig. 14) between the typical and “Derwent” forms of

D. margaretae,.

Dasystigma bonhami, D. margaretae (all known variants)

and D. huonense are often locally abundant and are found in

formal State reserves. Much of the range of the less common

D. tyleri is formally reserved, notably in national parks. The

three eastern species have all been found in logged and regen-

erated native forest, including older regrowth (20-1- years) from

clearfall-and-bum operations. For evolutionary studies and for

clarification of taxonomic boundaries it would be worthwhile

to seek special, conservative management for public land forest

patches in which different forms meet in narrow parapatry, and

the author hopes to identify suitable patches in the near future.

Acknowledgements

I am grateful to the Plomley Foundation for financial assis-

tance; to David Steele (University of Tasmania) for acquiring

the SEM images; to Graeme Anderson and Owen Seeman

(Department of Primary Industries, Water and Environment,

Tasmania) for access to the L. margaretae types; to Graham

Compton (CSIRO Entomology) and Dennis Black (La Trobe

University) for access to ANIC material; and to Kevin Bonham

and my wife Trina Moule for assistance in collecting

Dasystigma specimens.

References

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genus Lissodesmus Chamberlin, with the description of four new

species from Tasmania (Diplopoda, Polydesmida, Dalodesmidae).

Papers and Proceedings of the Royal Society of Tasmania 118;

85-102.

Key, K.H.L. 1982. Species, parapatry, and the morabine grasshoppers.

Systematic Zoology 30: 425^58.

Mesibov, R. 1994. Faunal breaks in Tasmania and their significance for

invertebrate conservation. Memoirs of the Queensland Museum 36:

133-136.

Mesibov, R. 1997. A zoogeographical singularity at Weavers Creek,

Tasmania. Memoirs of the Museum of Victoria 56: 563-573.

Mesibov, R. 1999. The Mersey Break: an unexplained faunal boundary

on the north coast of Tasmania. Pp. 246-252 in Ponder, W., and

Lunney, D. (eds.). The Other 99%. The Conservation and

Biodiversity of Invertebrates. Transactions of the Royal Zoological

Society of New South Wales. Royal Zoological Society of New

South Wales; Mosman (NSW).

Mesibov, R. 2003. The millipede genus Gasterogramma (Diplopoda;

Polydesmida: Dalodesmidae) in Tasmania, Australia, with descrip-

tions of seven new species. Memoirs of Museum Victoria 60:

000 - 000 .

Shelley, R.M. 1990a. Are allopatric/parapatric mosaic complexes

widespread in the Diplopoda? (Abstract) P. 23 in Minelli, A. (ed.),

Proceedings of the 7th International Congress of Myriapodology.

E.J. Brill: Leiden.

Shelley, R. M. 1990b. Revision of the milliped family

Eurymerodesmidae (Polydesmida: Chelodesmidea). Memoirs of the

American Entomological Society 37: 1-112.

Shelley, R.M. and Whitehead, D.R. 1986. A reconsideration of the mil-

liped genus Sigmoria, with a revision of Deltotaria and an analysis

of the genera in the tribe Apheloriini. Memoirs of the American

Entomological Society 35: 1-223.

Wiley, E.O. 1978. The evolutionary species concept reconsidered.

Systematic Zoology 21 \ 17-26.

Memoirs of Museum Victoria 60(2): 207-219 (2003)

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

http://www.museum.vic.gov.au/memoirs

The millipede genus Gasterogramma (Diplopoda: Polydesmida: Dalodesmidae) in

Tasmania, Australia, with descriptions of seven new species

Robert Mesibov

Queen Victoria Museum and Art Gallery, Launceston, Tasmania, Australia 7250 (mesibov@southcom.com.au)

Abstract Mesibov, R. 2003. The millipede genus Gasterogramma (Diplopoda: Polydesmida: Dalodesmidae) in Tasmania,

Australia, with descriptions of seven new species. Memoirs of Museum Victoria 60(2): 207-219.

Eight species of Gasterogramma (Diplopoda: Polydesmida: Dalodesmidae) occur in Tasmania: G. austrinum sp. nov.,

G. extremum sp. nov., G. imber sp. now, G. plomleyi sp. now, G. psi Jeekel, 1982, G. rusticum sp. now, G. tarkinense sp.

nov. and G. wynyardense sp. nov. The genus is largely confined to areas with an annual rainfall greater than 1000 mm.

G. psi is widespread and abundant in the northern and western third of Tasmania, while G. plomleyi is restricted to the

area of endemism known as Plomleys Island in Tasmania’s north-east. G. plomleyi and the six other new species form

an allopatric/parapatric distribution mosaic in northern, western and southern Tasmania. All eight species are burrowers

in soil and deep litter.

Introduction

C.A.W. Jeekel (1982) established the genus Gasterogramma

for Gasterogramma psi Jeekel, 1982, a peculiar Tasmanian

dalodesmid millipede in which the distal portion of the gono-

pod telopodite appeared to Jeekel to have rotated nearly 180° in

development, giving the prostatic groove a spiral path to the

solenomerite. Further collections in Tasmania have yielded

seven more species of Gasterogramma, described here. The

new species strongly resemble G. psi in general appearance and

habits and all have, like G. psi, a helical solenomerite, but other

features of gonopod structure differ markedly from those in

G. psi. The generic description is amended accordingly.

Specimens were killed and preserved in 75-80% ethanol.

Preliminary drawings on graph paper were made using material

cleared in 60% lactic acid and viewed at 100 or 200x magnification

through an eyepiece graticule. A Philips Electroscan ESEM 2020 oper-

ated in high-vacuum mode was used to examine preserved material

which had been air-dried before sputter-coating with gold. SEM

images were acquired digitally.

More than 600 Gasterogramma samples were examined. To save

space in the printed version, full details of localities, dates, collectors,

specimens and registration numbers are provided separately on the

Memoirs of Museum Victoria website, www.museum.vic.gov.au/

memoir s/index. html. The specimen data table is also available from the

author and a copy has been deposited at the QVM.

Universal Transverse Mercator (UTM) grid references are the spa-

tial locators used by most field workers to define collecting localities

in Tasmania. Collecting sites for all but a few of the specimens exam-

ined were estimated in the field to be within particular 100 m UTM

grid squares on 1:25000 scale maps published by the State of

Tasmania. Grid squares are recorded below in 2-letter, 6-digit form.

e.g. ‘EN700712.’ The maximum horizontal error in these estimates is

likely to be less than 100 m. Latitude/longitude equivalents were cal-

culated using GeoCalc 4.20 (GeoComp Systems, Blackburn, Victoria)

and are not as precise as the UTM grid references. LGRSS transect

locations (see separate specimen data table) were derived from 1:2000

survey charts made available to the QVM by the Hydro-Electric

Commission, Tasmania, in 1994.

Abbreviations are: AM, Australian Museum, Sydney; ANZSES,

Australia and New Zealand Scientific Exploration Society; QVM,

Queen Victoria Museum and Art Gallery, Launceston. Unless other-

wise indicated, male and female refer to stadium 8 adults.

Order Polydesmida Leach, 1815

Suborder Dalodesmidea Hoffman, 1977

Dalodesmidae Cook, 1896

Gasterogramma Jeekel, 1982

Gasterogramma Jeekel, 1982: 10. — Shelley et ah, 2000: 102.

Type species. Gasterogramma psi Jeekel, 1982, by original

designation.

Included species. G. austrinum sp. nov., G. imber sp. nov.,

G. extremum sp. nov., G. plomleyi sp. nov., G. psi Jeekel, 1982,

G. rusticum sp. nov., G. tarkinense sp. nov. and G. wynyardense

sp. nov.

Remarks. Jeekel (1982: 10) gave the following diagnosis for

somatic features of Gasterogramma: “Medium- sized

Dalodesmidae with 20 somites and a normal poreformula.

Head without particulars; antennae of moderate length, clavate,

the 6th antennomere longer than the 5th. Collum without

208

Robert Mesibov

paranota; the sides almost perpendicular. Somites very weakly

constricted, almost cylindrical; the waist broad. Metatergites

without sculpture. Paranota weakly developed, ridgelike, only

dorsally demarcated by a furrow in which the pores are situ-

ated. Sternites much longer than wide, without particular mod-

ifications. Legs of moderate length, incrassate in the male and

with dense setation of short stiff bristles on ventral side of

podomeres 2 to 4 and dense granulation on ventral side of

podomeres 5 and 6. Anal somite without particulars.” This

general description also applies to the seven new species

described here but the following generic features should be

included: integument well calcified; narrow but clearly

distinguishable paranota on second segment, lower on body

than the lateral collum margins and the paranotal traces on

third segment; gonopod aperture ovoid with long axis trans-

verse, rim not raised in front and variably raised at rear;

epiproct blunt, prominent, extending caudad past the anal

valves.

Jeekel’s diagnosis of the Gastewgramma gonopod (1982;

10) is here amended: telopodites elongate, loosely joined

medially; solenomerite arising at one-half to two-thirds the

length of the telopodite, each solenomerite a small, distally

tapering rod twisted in the following sense; a right-handed

helix (clockwise turning away from the observer) on the right

gonopod, a left-handed helix on the left gonopod.

In an attempt to identify (i.e., homologise) the elements of

the G. psi gonopod, Jeekel (1982: 11) hypothesised that “the

course of the spermal channel [here called “prostatic groove”]

is homologous in all polydesmoid millipedes, and that the

location of the base of each process in relation to the spermal

channel defines its morphological identity.” He named the

process arising caudad of the prostatic groove the tibiotarsus,

and the structure arising cephalad the femoral process. If the

distal portion of the G. psi gonopod is untwisted, in imagin-

ation, then the lateral process with a tip shaped like a bird’s

head (fig. 6) is identifiable as the femoral process; the anterior,

laterally fringed process bearing a rod-like structure is a pro-

longation of the prefemur; and the rod-like structure is the

tibiotarsus. In the notch between the femoral and prefemoral

processes arises the solenomerite.

An alternative interpretation is that the distal end of the

telopodite is not torted, and that the ‘femoral process’ is a

growth from the caudomedial side of the gonopod which has

displaced the prostatic groove laterally. It is important to note

that the course of the prostatic groove in G. psi is actually two

opposed spirals. From the base of the telopodite of the right

gonopod, the groove first runs distally in a left-handed spiral to

encircle the “femoral process”, then in a right-handed spiral

upon entering the solenomerite. The cheirality of the spirals on

the left gonopod mirrors that on the right. In the three species

here assigned to Gastewgramma which lack a femoral process

(G. austrinum, G. rusticum and G. wynyardense), the basal

spiral is missing but the solenomerite spiral persists.

Jeekel’s hypothesis sought to make each gonopod process in

polydesmoids a homologue of either the prefemur, femur or

tibia-i-tarsus of the millipede leg, and uses a “fixed”

solenomerite as a morphological landmark. If the “no torsion”

interpretation of the Gasterogramma gonopod is correct, then

either the simple notion of homologisation is inadequate, or the

solenomerite is not “fixed.” Unfortunately, polydesmoid

gonopods develop suddenly from unsegmented primordia in

the ultimate moult, and there are no visible intermediate stages

which would allow us to unambiguously identify individual

gonopod processes with prototypical leg segments, in

Gasterogramma or any other polydesmoid. A conect inter-

pretation of gonopod structure may be possible in future with

advances in the developmental genetics of arthropods. For the

present, Jeekel’s labelling of the gonopod processes in

Gasterogramma is a useful hypothesis, and is adopted for

descriptive purposes in this paper.

Distribution. Western and north-eastern Tasmania (fig. 13); not

recorded from King I. or the Fumeaux Group in Bass Strait.

Habits. The eight species of Gasterogramma are burrowing

millipedes typically found in moist, organic-rich soil, in deep

moist leaf litter and in and under wet rotting logs. As noted by

Jeekel (1982: 9) for G. psi, Gasterogramma are “sluggish” and

at first sight they resemble “a worm or a dipteran larva rather

than a millipede.” They are often locally abundant in loose,

mixed-age aggregations. All Gasterogramma species release a

pungent defensive secretion which presumably includes hydro-

cyanic acid, although this is not the most prominent odour com-

ponent. The pinkish-purple Gasterogramma species have been

called “stinky pinkies” by local collectors impressed by the

pungency of the secretion. G. plomleyi has the strongest smell

in the genus, and can be detected by treading heavily through

its wet forest habitat and sniffing attentively. Unfortunately, no

somatic characters have been so far noted which can reliably be

used to distinguish the seven western species (i.e.,

Gasterogramma spp. other than G. plomleyi), and at many

western sites two Gasterogramma species are present. I have

not, therefore, assigned non-plomleyi females and juveniles to

species; these individuals are recorded in the separate specimen

data table as Gasterogramma sp.” Males and females have

been found in copula in all months of the year, but mating

is most frequent in the austral spring; the only two pairs so

far preserved in copula (G. psi, QVM 23:8086 and QVM

23:40560) were collected in late November. Females are

often found curled around an egg mass; it is unclear

whether such females are in the process of constructing an

egg chamber (as expected for Polydesmida) or are brooding

their eggs.

Relationships. Jeekel (1982: 12) noted that Gasterogramma

and the Chilean Semnosoma Silvestri, 1903 were the only

dalodesmid genera in which the prostatic groove had been

reported to be helical, and “It seems likely, therefore, that the

new genus is closely related to Semnosoma. Eventually, the two

may form the nucleus of a tribe for which the family-group

name Semnosomatidae Brolemann, 1916 is available.” The

illustrations of Semnosoma gonopods in Demange and Silva

(1976a, b) show the prostatic groove running first caudally,

then laterally, then anteriorly as in G. psi. Further, species in the

Semnosoma group of Chilean dalodesmids (i.e. Semnosoma,

Anaulacodesmus Attems, 1898, Chiloriis Chamberlin, 1957

New Tasmania species of the millipede Gasterogramma

209

Figure 1. (left) G. austrinum sp. nov., Tahune Bridge, southern Tasmania, QVM 23:16195; SEM of gonopods in situ. Scale-bar = 0.5

mm. (right) G. austrinum sp. nov.. Gold Creek, southern Tasmania, QVM 23:14021; outline sketches of right gonopod showing

course of prostatic groove (dashed line): lateral view, left; posterior view, right.

Figure 2. (left) G. extremum sp. nov.. Lake Sydney, southern Tasmania, QVM 23:14019; SEM of

gonopods in situ. Scale-bar = 0.5 mm. (right) G. extremum sp. nov., Hastings Caves, southern

Tasmania, QVM 23:8059; outline sketch of right gonopod, lateral view, showing course of prostatic

groove (dashed line).

210

Robert Mesibov

Figure 3. (left) G. imber sp. nov., Tarraleah, central Tasmania, QVM 23:8047; SEM of gonopods in

situ. Scale-bar = 0.5 mm. Prefemoral processes are parallel in life and in alcohol-preserved material;

the divergence seen in this SEM is an artefact of specimen preparation, (right) G. imber sp. nov.,

Argent R., western Tasmania, QVM 23:8051; outline sketch of right gonopod, lateral view, showing

course of prostatic groove (dashed line).

Figure 4. (left) G. plomleyi sp. nov.. Rattler Hill, north-east Tasmania, QVM 23:8068; SEM of gonopods in

situ. Scale-bar = 0.5 mm. (right) G. plomleyi sp. nov.. Crystal Hill, north-east Tasmania, QVM 23:8069; out-

line sketch of right gonopod, lateral view, showing course of prostatic groove (dashed line).

New Tasmania species of the millipede Gasterogramma

211

and Tsagonus Chamberlin, 1957) all have, like Gastero-

gramma, greatly reduced paranoia (Chamberlin, 1957: 19). The

argument for a Gondwanan dalodesmid lineage including the

Chilean and Tasmanian genera thus appears reasonable.

Relationships within Gasterogramma are unclear. Grouping

phenetically on gonopod form, we might have (G. extremum +

G. imber + G. plornleyi + G. psi + G. tarkinense) with a femoral

process and (G. austrinum + G. rusticum + G. wynyardense)

without such a process, but relationships within these groups

are far from obvious. Both groups are widely dispersed across

Tasmania, suggesting that the two might have differentiated

independently in the landscape. However (see below) the two

groups are not independently distributed: the seven non-psi

species currently form an allopatric/parapatric mosaic.

Gasterogramma austrinum sp. nov.

Figures 1,5, 12 (map)

Material examined. Holotype. Male. Australia, Tasmania. Huon R.

(Arve Rd), DN784273 [43°06'23'' 146°44'04'd, 120 m, 2 May 1997,

R. Mesibov, plot 3M1, oldgrowth wet eucalypt forest with rainforest

understorey, QVM 23:40628.

Paratypes. 3 males, Huon R. (Edwards Rd), DN790284 [43°05'48'"

146°443rd, 90 m, 23 May 1997, R. Mesibov, plot 3R6, regrowth wet

eucalypt forest following clearfelling and burning, QVM 23:40630; 2

males, Edgar Bay area, DN483404 [42°59T3^' 146°2r57^1, 350 m,

pitfall emptied 16 Nov 2001, D. Driscoll, QVM 23:24802; 4 males,

Huon R. (Arve Rd), DN798284 [43°05'48'' 146°45'06'1, 200 m, 16

May 1997, R. Mesibov, regrowth wet eucalypt forest following clear-

felling and burning, AM KS85097 (formerly QVM 23:40629).

Other material. 19 males from 8 unique localities in southern

Tasmania, including Gold Creek, Huon R., Kallista Creek, Picton R.,

Reuben Palls and Tahune Bridge.

Diagnosis. Gonopod with no femoral process or tibiotarsus;

telopodite in posterior view ending in broad, mesodistally

directed point.

Description. Males c. 17-20 mm long and c. 1. 8-2.0 mm wide

at midbody. Body in both sexes darkly mottled with pink-pur-

ple pigmentation. Gonopod telopodites slender in posterior

view, lightly joined for about half thek length, a few short,

coarse setae at their bases, setate area extending just outside

aperture (fig. 1). Aperture more or less ovoid, long axis trans-

verse, posterior margin not noticeably raised. Telopodite with a

posteromesal concavity near base, somewhat sinuous in lateral

view, curving anteriorly, then distally, then anteriorly and

slightly tapering, finally tapering mesally to end in blunt,

mesodistally pointed tip. Femoral process and tibiotarsus not

evident. Solenomerite short, helical, tapering process arising at

about four-fifths length of telopodite on its posterior face.

Prostatic groove running distally and posteriorly along mesal

side of telopodite, then curving laterally before running straight

distally to enter solenomerite on its mesal side.

Distribution. In rainforest and wet eucalypt forest over at least

750 km- in southern Tasmania (fig. 12), at altitudes c. 50-600

Etymology. From Latin austrinus, southern, adjective. This

species is restricted to southern Tasmania.

Gasterogramma extremum sp. nov.

Figures 2, 12 (map)

Material examined. Holotype. Male. Australia, Tasmania. Deadmans

Bay, DM592802 [43°3L47^' 146°29'42'1, <40 m, 26 Jan 1994, R.

Mesibov, wet eucalypt forest and rainforest, QVM 23:14018.

Paratypes. Male (dissected), near Hastings Caves, approx.

DM870960 [43°23T8^' 146°50^22^d, 5 Dec 1952, VV Hickman,

QVM 23:8059; male (dissected for SEM), Lake Sydney, DN686070

[43°17'20'' 146°36'46"], 690 m, 2 Eeb 1994, R, Mesibov, rainforest,

QVM 23:14019.

Diagnosis. Gonopod without tibiotarsus; femoral process

unbranched; prefemoral process curved laterally, then poster-

iorly, with short, tooth-like projections near tip; solenomerite

arising near base of prefemoral process.

Description. Males c. 15-17 mm long and c. 1.6-1. 8 mm wide

at midbody. Body in both sexes ivory-coloured with faint pink-

purple mottling. Gonopod telopodites massive at base (fig. 2),

lightly joined mesally for most of their length, with a few short,

coarse basal setae, setate area extending just outside aperture.

Aperture more or less ovoid, long axis transverse, posterior

margin not noticeably raised. Telopodite axis more or less

straight. Telopodite base ending abruptly at about one-half

length of telopodite, with a relatively slender, anterioposterior-

ly flattened prefemoral process arising from anteromesal corner

of top of base. Prefemoral process tapering gradually to a fine

point, bending first laterally, then posteriorly, and bearing set of

10-20 short, simple or bifid teeth on posterolateral surface just

proximal to tip. Tibiotarsus not evident. Femoral process

massive, tapering gradually to a fine point, arising from pos-

teromesal corner of top of telopodite base and curving later-

ally, then anteriorly, then distally and slightly laterally.

Solenomerite a short, helical, somewhat flattened process aris-

ing at base of prefemoral process on its posterior face. Prostatic

groove running first posteriorly and distally across mesal face

of telopodite base to its posterior surface, then curving later-

ally, distally and mesally around base of femoral process, then

extending distally to enter solenomerite base on its mesal side.

Distribution. Rainforest and wet eucalypt forest at 3 locations

in far southern Tasmania (fig. 12) from near sea level to 690 m,

with maximum linear range of c. 30 km. Likely to be more

widespread in this little- sampled part of Tasmania.

Etymology. From Latin extremus, farthest, adjective. This

species extends to the southern-most part of mainland

Tasmania.

Gasterogramma imber sp. nov.

Figure 3, 12 (map)

Material examined. Holotype. Male. Australia, Tasmania. Little

Florentine R., DN525683 [A2°AA'W' 146°25H0''], 440 m, pitfall

emptied 10 Apr 1986, R. Bashford, QVM 23:24958.

Paratypes. Male, details as for holotype, QVM 23:40586; male,

Tan-aleah, DP473196 [42°16'25'' 146°2L39^1, 750 m, 4 May 1992,

R. Mesibov, oldgrowth wet eucalypt forest with rainforest understorey,

QVM 23:8048; male. Needles Picnic Ground, DN5 12656 [A2°A531"

\A6°2A'\2"}, 470 m, 23 Feb 1994, R. Mesibov, oldgrowth wet euca-

lypt forest with rainforest understorey, QVM 23:13840; 5 males.

212

Robert Mesibov

Tarraleah, DP473197 [42°16^22'' 146°2r39"], 780 m, 5 May 1992,

R. Mesibov, oldgrowth wet eucalypt forest with rainforest understorey,

AM KS85098 (formerly QVM 23:8045).

Other material. 49 males from 26 unique localities in central, south-

western and western Tasmania, including Argent R., Boyd R., Denison

R., Franklin R., Frenchmans Cap, Gordon R., Humboldt Divide, Huon

R., Kallista Creek, Little Florentine R., Mossy Marsh Creek,

Queenstown, Scotts Peak Dam, Taffys Creek, Tarraleah, Trappes Inlet,

Wedge Inlet and Wedge R..

Diagnosis. Gonopod without tibiotarus; femoral process

unbranched; prefemoral process slender, straight, without

tooth-like projections, flexed sharply at tip; solenomerite rising

near base of prefemoral process.

Description. Males c. 17-20 mm long and c. 1. 8-2.0 mm wide

at midbody. Body in both sexes darkly mottled with pink-

purple pigmentation, Gonopod telopodites massive at base (fig.

3), lightly joined mesally for about half their length, a few

short, coarse basal setae, setate area extending just outside

aperture. Aperture more or less ovoid, long axis transverse,

posterior margin not noticeably raised. Telopodite narrowing

near its base, bending slightly in anterior direction at about half

telopodite length, narrowing greatly and abruptly at bend and

extending from anteromesal corner of top of base as a spear-

like, sharply pointed prefemoral process flexed sharply poster-

iolaterally at about three-quarters of its length. (In life and in

alcohol preservative, axis of prefemoral process parallels that

of telopodite; prefemoral processes in SEM image in fig. 3

bend laterally at their bases as a result of drying-out during

specimen preservation) Tibiotarsus not evident. Femoral

process massive, arising from posterior face of telopodite base

and directed laterally and distally, bending sharply anteriorly

and distally at about two-thirds its length and tapering to a blunt

point. Solenomerite a short, helical, tapering, somewhat flat-

tened process arising between bases of femoral and prefemoral

processes. Prostatic groove running first posteriorly and dis-

tally across mesal face of telopodite base to its posterior sur-

face, then curving laterally, distally and mesally around base of

femoral process, then extending distally to enter solenomerite

base on its mesal side.

Distribution. In rainforest and wet eucalypt forest over at least

6000 km- in western Tasmania (fig. 12), at altitudes c. 50-800

m. Syntopic with G. psi at various locations through its range.

Etymology. Latin imber, shower, noun in apposition. This

species occurs in the highest rainfall zone in Tasmania.

Gasterogramma plomleyi sp. nov.

Figures 4, 5, 12 (map)

Material examined. Holotype. Male. Australia, Tasmania. Rattler Hill,

EQ744353 [41°13A8'' 147°53T5'1, 650 m, 29 Aug 1990, R.

Mesibov, oldgrowth rainforest, QVM 23:24960.

Paratypes. 2 males (1 dissected for SEM), details as for holotype,

QVM 23:8068; 2 females, details as for holotype, QVM 23:8307; 3

males, Eorest Lodge, EQ786296 [41°16'5r' 147°56T8'3, 410 m, 13

Jan 1993, R. Mesibov, wet eucalypt forest, QVM 23:8066; 1 male.

Pecks Hill, EQ282251 [41°19'29^' 147“20T3^1, 450 m, 14 Jan 1993,

R. Mesibov, wet eucalypt forest, QVM 23:8061; 3 males, Williams

Hill, EQ558548 [41°03^21'^ 147°39'50^3, 300 m, 10 Mar 1993, R.

Mesibov, wet eucalypt forest, QVM 23:16477; 3 males, Mathinna

Plains, EQ616221 [4F2r00'' 147°44T0'1, 800 m, 12 Jan 1993, R.

Mesibov, AM KS85099 (formerly QVM 23:8063).

Other material. 26 males, 40 females and 52 juveniles from 33

unique localities in north-east Tasmania, including Carters Creek,

Chinaman Corner, Crystal Hill, Cuckoo, Eorest Lodge, Golconda,

Hogarth Rivulet, Joseph Creek, Lisle, Mathinna Plains, Milly Brook,

Mt Arthur, Mt Michael, Mt Victoria, Northallerton Valley, Patersonia

Rivulet, Pecks Hill, Peddles Hill, Rattler Hill, Rayners Hill, Sideling

Range, South Springfield, Tombstone Creek, Weldborough and

Williams Hill.

Diagnosis. Epiproct unusually broad, with bumpy appearance

(fig. 5B). Gonopod without tibiotarsus; femoral process

unbranched; prefemoral process short, without tooth-like pro-

jections, bent anteriorly, then distally and mesally;

solenomerite seemingly fused with prefemoral process for half

length of latter.

Description. Males c. 20-23 mm long and c. 2.0-2.4 mm wide

at midbody, females somewhat larger. Body in both sexes fair-

ly uniformly ivory-coloured. Epiproct (fig. 5B) enlarged rela-

tive to other Gasterogramma (fig. 5A), lateral and terminal

setae on prominent bumps (fig. 5C). Gonopod telopodites (fig.

4) fairly straight and slender, lightly joined mesally for most of

their length, a few long setae at their bases just inside aperture.

Aperture more or less ovoid, long axis transverse, posterior

margin very slightly raised and broadly, medially notched.

Femoral process massive, arising on posteromesal face of

telopodite at about two-thirds its length, directed laterally and

slightly distally before turning abruptly anteriorly at about half

length of process and tapering to a blunt point. Telopodite

extending from femoral process base as a short, slender

prefemoral process which bends anteriorly, then curves distally

and mesally to terminate in a fine point. Tibiotarsus not evident.

Solenomerite about as long as prefemoral process, arising near

base of prefemoral process on its posterior face and apparently

fused with it to point where prefemoral process bends anterior-

ly, from this point helical and slightly flattened. Prostatic

groove running distally and posteriorly across mesal surface of

telopodite, then curving laterally, distally and mesodistally

around femoral process before entering solenomerite on its

mesal side. Cyphopods not examined.

Distribution. In rainforest and wet eucalypt forest over c. 1700

km in high-rainfall parts of north-east Tasmania (fig. 12), at

altitudes 150-850 m.

Etymology. In honour of Brian Plomley (1912-1994),

Tasmanian scientist and scholar who encouraged biological

research in north-east Tasmania.

Gasterogramma psi Jeekel, 1982

Figures 6,1, 11 (map)

Gasterogramma psi Jeekel, 1982: 12. — Shelley et al., 2000: 102.

Holotype. Male. Australia, Tasmania. “Sta. 100, Hellyer Gorge, 32 km

SSW Somerset, 25.XI.1980 (temperate rain-forest (Nothofagus,

Eucalyptus, Dicksonia) along the Hellyer R., under logs)” (Jeekel,

1982: 12). Holotype probably in Zoological Museum, Amsterdam;

not examined.

New Tasmania species of the millipede Gasterogramma

213

Figure 5. Sketches of epiproct form. A, Typical form in Gasterogramma; G. austrinum paratype, QVM

23:40629; lateral view. B, G. plomleyi, male paratype, QVM 23:8068, lateral view. C, same specimen as

B, ventral view.

Figure 6. A-D, G. psi gonopod variations, seen in situ; see fig. 11 for locations. A, Washpond Forest, north-west Tasmania,

QVM 23:8085. B, Notley Gorge, north-central Tasmania, QVM 23:8139. C, Coles Creek, central Tasmania, QVM

23:24797. D, Henty R., western Tasmania, QVM 23:8143. Scale-bar in all cases = 0.5 mm.

214

Robert Mesibov

Figure 7. A-C, body pattern variations in G. psi. A, almost uniformly pale; Cam R., north-west Tasmania. B, dark-spotted; Dial Range, north-west

Tasmania. C, dark with faint mottling; Florentine R., south-west Tasmania.

Figure 8. (left) G. rusticum sp. nov., Don Reserve, north-central Tasmania, QVM 23:16199; SEM

of gonopods in situ. Scale-bar = 0.5 mm. (right) G. rusticum sp. nov., Gog Range, north-central

Tasmania, QVM 23:8053; outline sketches of right gonopod showing course of prostatic groove

(dashed line): lateral view, left; posterior view, right.

New Tasmania species of the millipede Gasterogramma

215

Figure 9. (left) G. tarkinense sp. nov., The Clump, north-west Tasmania, QVM 23:8056; SEM of

gonopods in situ. Scale-bar = 0.5 mm. (right) G. tarkinense sp. nov.. Wombat Hill, north-west

Tasmania, QVM 23:8057; outline sketch of right gonopod showing course of prostatic groove

(dashed line).

Figure 10. (left) G. wynyardense sp. nov., Inglis R., north-west Tasmania, QVM 23:41909; SEM of gonopods in situ.

Scale-bar = 0.5 mm. (right) G. wynyardense sp. nov., Seabrook Creek, north-west Tasmania, QVM 23:40632; outline

sketches of right gonopod showing course of prostatic groove (dashed line): lateral view, left; posterior view, right.

216

Robert Mesibov

Material examined. Paratypes. 10 males, 12 females, 3 stadium VII

males, 3 stadium VI males, collection details as for holotype; locations

unknown.

Other material. 393 males from 190 unique localities in northern

and western Tasmania, including Allen Creek, Animal Creek, Arthur

R., Balfour, Bellana Creek, Big Creek, Black Bog Creek, Black R.,

Blackfish Creek, Bond Tier, Bonneys Tier, Borradale Creek, Branchs

Creek, Broadsword cave (Gunns Plains), Brooks Creek, Bubs Hill,

Burnie Park, Burning Down the House cave (Junee-Florentine), Cam

R., Cann Creek, Central Castra, Chester Creek, Christmas Hill,

Christmas Hills, Coles Creek, Companion Hill, Companion Rd,

Crayfish Creek, Dalgarth Forest Reserve, Dark Creek, Dawson R.,

Dead End Den cave (Mt Cripps), Dee Lagoon, Deep Creek Bay,

Denison R., Dip R., Don R., Duck Creek, Dundas R., East Ridgley,

Emu R., Fisher R., Florentine R., Flowerdale, Gawler R., Gibson

Creek, Goderich Rd, Gordon R., Guildford Rd, Hardwood R.,

Heemskirk Rd, Henty R., Hermit Hill, Hoi well Gorge, Inglis R.,

Interview R., Jessie Rd, Jones Creek, Julius R., Kelcey Tier, Kenzies

Hill, Lake Chisholm, Lake Lea, Lake Mackenzie, Laughing Jack

Lagoon, Lawson Plains, Library Creek, Little Claytons Rivulet,

Loongana cave L9, Maggs Mountain, Mahoneys Creek, Marine Creek,

Maxwell R., Mersey R., Meunna, Meunna Hills, Milkshakes, Montagu

Swamp, Mossy Marsh Creek, Mostyn Hardy cave (Loongana), Mt

Oakleigh, Mt Sprent, Newall Creek, Nietta Creek, No Mans Creek,

Nook, Notley Gorge, Olga R., Orange R., Ordnance Point, Parrawe

Creek, Pelion Valley, Pencil Pine Creek, Philrod cave (Mt Cripps),

Poppys Lagoon, Punchs Terror, Richardsons Flats, Roger R., Roger R.

West, Rosebery, Salmon R., Sassafras Creek, Savage R., Saxons

Creek, Scopus, Seabrook Creek, Serpentine Dam, Shadow Lake,

Sisters Creek, Smithton, Somerset, Sterling R., Stony Rises, Sundown

Point, Table Cape, Tarraleah, Three Hummock L, Trowutta Caves,

Tunnel, Upper Natone, Wakefield Creek, Wandle R., Washpond Forest,

Weaning Paddock Creek, Welcome Heath, West Montagu, Whisky

Creek, Wild Wave R., Williamsford, Wilsons Creek and Wombat Hill.

Diagnosis. Gonopod with femoral process terminating in a

mesally directed tip with shape of bird’s head; prefemoral

process fringed with tooth-like projections; tibiotarsus promi-

nent, distally or distally and mesally directed; solenomerite

arising near base of prefemoral process.

Descriptive notes. For a very detailed description of this

species, see Jeekel (1982: 12-14). Variations in gonopod struc-

ture and body colour pattern (see below) occur across the range

of G. psi, but I am reluctant to divide G. psi into subspecies,

Gonopod: The form described by Jeekel (1982) from north-

west Tasmania (site “TL” in fig. 11) extends with little variation

to central Tasmania. In the latter area (“C” in fig. 11), the tip of

the prefemoral process is shortened and bent slightly poster-

iorly (fig. 6C). In the central west (“D” in fig. 11), the tip of the

prefemoral process is further flexed and lies lateral to the tibio-

tarsus, which is directed posteriomesally (fig. 6D) rather than

standing more or less “upright” as in north-west forms. In the

far north-west (“A” in fig. 11), the tibiotarsus extends well

distal of the prefemoral process, and a second small, lateral

tooth is present towards the base of the femoral process (fig.

6A). In the north-east of the G. psi range (“B” in fig. 11), the

tibiotarsus is enlarged and directed slightly mesally, and the tips

of the femoral process turn slightly posteriorly (fig. 6B).

Colour: Over much of the western and north-western

portion of its range, G. psi is pale with very faint, brownish

gray markings (fig. 7A). In the north of the range almost all

specimens have a striking pattern of dark lateral spots on the

segments with ozopores (5, 7, 9, 10, 12, 13, 15-18; fig. 7B),

while specimens from the south and central portions of the

G. psi range are dark with faint mottling (fig. 7C).

Distribution. From sea level to c. 1250 m over c. 25 000 km” in

northern and western Tasmania, including Three Hummock

Island in Bass Strait (fig. 11). In coastal, subalpine and riparian

scrubs; in swamp forest, wet eucalypt forest and rainforest; in

exotic tree plantations; and occasionally in caves. Syntopic

with G. imber, G. rusticum, G. tarkinense, and G. wynyardense

over parts of its range.

Remarks. Gasterogramma psi is the most ecologically tolerant

species of the genus in north-west Tasmania, and is generally

more abundant than co-occurring G. rusticum, G. tarkinense or

G. wynyardense.

Gasterogramma rusticum sp. nov.

Figures 8, 12 (map)

Material e.xamined. Holotype. Male. Australia, Tasmania. Christmas

Hill, DQ697095, 340 m, 10 Nov 1993, R. Mesibov, wet eucalypt

forest, QVM 23:16476.

Paratypes. 2 males. Little Claytons Rivulet, DQ321385 [41°12"06"'

146°lL24^"b 80 m, 31 Oct 1996, R. Mesibov, wet eucalypt forest,

QVM 23:40594; 2 males, Elizabeth Town, DQ633091 [4L28T7''

146°33"37'd» 210 m, 17 Sep 1997, R. Mesibov, wet eucalypt forest,

QVM 23:40600; 5 males, Gawler R., DQ277347 [41°14'08''

146°08T4'd» 150 m, 29 Apr 1999, K. Bonham, site 6b, blackwood

forest, AM KS 85100 (formerly QVM 23:41901).

Other material. 31 males from 21 unique localities in northern

Tasmania, including Caroline Creek, Christmas Hill, Dalgarth Forest

Reserve, Dasher R., Don Reserve, Dulverton, Dysodile Hills, East

Gawler R., Gog Range, Kelcey Tier, Latrobe, Lobster Rivulet, Long

Hill, Marine Creek, Mersey R., Nook, Staggs Hill, Stella Glen,

Warners Sugarloaf and Winter Brook.

Diagnosis. Gonopod with no femoral process or tibiotarsus;

telopodite in posterior view with broadly notched tip, i.e. with

2 well- separated ter mi nal teeth.

Description. Males c. 17-20 mm long and c. 1.8-2.0 mm wide

at midbody. Body in both sexes lightly mottled with brownish-

pink pigmentation. Gonopod telopodites slender, more or less

straight, lightly joined for about middle third of their length, a

few short, coarse setae near bases just inside aperture (fig. 8).

Aperture more or less ovoid, long axis transverse, posterior

margin not noticeably raised. Telopodite slightly sinuous in

lateral view, curving first anteriorly, then distally, then anter-

iorly, anterioposteriorly flattened towards its end and bending

slightly laterally, broadly notched at its terminus to form two

subequal teeth. No evidence of femoral process or tibiotarsus.

Solenomerite short, helical, tapering process arising at about

three-quarters of length of telopodite on its posterior face.

Prostatic groove running distally and posteriorly on mesal side

of telopodite, then bending first laterally, then mesally on pos-

terior telopodite surface before running distally to enter

solenomerite on its mesal side.

Distribution. In rainforest and wet eucalypt forest over c. 2300

km^ in north-central Tasmania (fig. 12), from sea level to

New Tasmania species of the millipede Gasterogramma

217

c. 700 m. Syntopic with G. psi at various locations through its

range.

Etymology. From Latin rusticus, rural, adjective. Much of the

range of this species is now an intensively farmed landscape.

Gasterogramma tarkinense sp. nov.

Figures 9, 12 (map)

Material examined. Holotype. Male. Australia, Tasmania. The Clump,

CQ213361 [41°12'23^' I44°52m'1, 190 m, 6 Feb 1992, R. Mesibov,

wet eucalypt forest, QVM 23:8058.

Paratypes. 3 males (1 dissected for SEM), details as for holotype but

17 Sep 1989, QVM 23:8056; 1 male. Little Donaldson R. area,

CQ552256 [41°18'28'' U5°W\T'}, 500 m, 7 Jan 1997, ANZSES

personnel, rainforest, AM KS85101 (formerly QVM 23:40584).

Other material. One male each from Newdegate Creek and Waratah

in north-west Tasmania.

Diagnosis. Gonopod with branched femoral process; tibio-

tarsus prominent, directed mesally and posteriorly from

prefemoral process fringed with tooth-like projections;

solenomerite arising near base of prefemoral process.

Description. Males c. 17-20 mm long and c. 1. 8-2.0 mm wide

at midbody. Body in both sexes ivory-coloured with traces of

pink-purple pigmentation middorsally and anteriorly on each

segment. Gonopod telopodites slender, narrowing somewhat

near base, lightly joined mesally for about two-thirds their

length, at which point they bend slightly anteriorly (fig. 9). A

projection like a shark’s dorsal fin, pointed distally, on anterior

surface of telopodite at about one-third its length. A few short,

coarse setae at telopodite bases, setate area extending just out-

side aperture. Aperture more or less ovoid, long axis transverse,

posterior margin not noticeably raised. Telopodite flattened

anterioposteriorly past bend and tapering laterally, with a

lateral and terminal fringe of c. 20 short teeth, simple and bifid,

directed posteriorly on tapered portion. Tibiotarsus about as

long as tapered portion of prefemoral process, a straight, point-

ed rod arising on mesal side of prefemoral process at about

halfway along tapered portion and directed mesally and poste-

riorly. Femoral process beginning as a bulge on posterior sur-

face of telopodite just proximal to bend, extending laterally in

two diverging, rod-like branches. Distal branch straight, direct-

ed distally and laterally and terminating abruptly in a finely

pointed, posteriorly directed hook. Proximal branch about one-

third length of distal branch, directed proximally and laterally

and curving posteriorly at its blunt tip. Solenomerite a short,

helical, tapering process arising near base of prefemoral

process on its posterior face. Prostatic groove running distally

and posteriorly on mesal side of telopodite, then curving later-

ally, then distally, then mesally around femoral process before

running distally to enter solenomerite on its mesal side.

Distribution. Rainforest and wet eucalypt forest at four sites in

north-west Tasmania (fig. 12), at altitudes c. 200-900 m, with

an estimated minimum range of c. 2300 km^. Syntopic with G.

psi on Wombat Hill, near Waratah.

Etymology. Adjectival form of Tarkine, a popular name for the

area between the Arthur and Pieman Rivers where the range of

this species is centred.

Gasterogramma wynyardense sp. nov.

Figures 10, 12 (map)

Material examined. Holotype. Male. Australia, Tasmania. Inglis R.,

CQ850382 [41°ir57'^ 145°37A2'1, 390 m, 27 May 1999, K.

Bonham, site 35b, wet eucalypt forest, QVM 23:41910.

Paratypes. 2 males, details as for holotype, QVM 23:41910; 1 male,

Meryanna, CQ388491 [4L0535'^ 145°04'50'1, 190 m, 10 Apr 1999,

R. Mesibov, rainforest, QVM 23:41044

Other material. 9 males from 8 unique localities in north-west

Tasmania: Arthur R., Blackfish Creek, Cam R., Inglis R., Julius R.,

Oonah, Seabrook Creek and Sumac Rivulet.

Diagnosis. Gonopod with no femoral process or tibiotarsus;

telopodite in posterior view ending in a narrow, distally

directed point

Description. Males c. 15-17 mm long and c. 1.6-1. 8 mm wide

at midbody. Body in both sexes lightly mottled with brownish-

pink pigmentation. Gonopod telopodites slender (fig. 10), aris-

ing from syncoxite well-separated, but lightly joined from

about one-third their length, a few short, coarse setae at

telopodite bases within aperture. Aperture more or less ovoid,

long axis transverse, posterior margin slightly raised and nar-

rowly, medially notched. Each telopodite bends anteriorly at

about half its length, bears a narrow, smoothly curved ridge on

its anterior surface at about three-quarters of its length, and

tapers mesally to blunt point. Solenomerite short, helical

process arising from flattened area on posterior side of

telopodite just distal to its bend, solenomerite cradled at its base

in a shallow depression in telopodite. Prostatic groove running

distally on mesal side of telopodite to vicinity of bend, then

curving to posterior side of telopodite, entering base of

solenomerite on a slightly sinuous, distal course. No indication

of a femoral process or a tibiotarsus.

Distribution. Rainforest and closed wet eucalypt forest (and

exotic tree plantations) over at least 900 km^ in north-west

Tasmania (fig. 12), from near sea level to c. 500 m. Syntopic

with G. psi at various locations through its range.

Etymology. Adjectival form of Wynyard, a Tasmanian town

close to which this species is abundant.

Gasterogramma sp.

Material. 2 males. Australia, Tasmania. Cam R. area, DQ019512

[41°05'03^' 145°49^55'1, 140 m, 30 Jul 1997, R. Mesibov and R. van

Riet, QVM 23:40634.

Remarks. These specimens were found at the extreme eastern

end of the G. wynyardense range, close to a locality for that

species. However, they seem closest to G. rusticum: the

prefemoral prolongation has a wide terminal notch, the pro-

static groove has an “S-bend” as it crosses the posterior face of

the telopodite, and the body is larger and more deeply pig-

mented than it is in G. wynyardense. They differ from

G. rusticum in having a mesodistally directed, spike-like

process arising on the mesoposterior face of the telopodite at

about the level of the solenomerite. One interpretation is that

the specimens represent a disjunct G. rusticum population (the

nearest known G. rusticum locality is 30 km to the east) which

218

Robert Mesibov

has long been separated from the main population and is some-

what differentiated. It is curious that the specimens were col-

lected at the edge of the wynyardense/rusticum distribution gap,

in which only G. psi has so far been found. Genetic studies of

Gasterogramma populations in the area may help to clarify the

taxonomic situation.

Biogeography and conservation

Gasterogramma is largely restricted to areas with an annual

rainfall greater than 1000 mm (fig. 13), i.e. to areas which at

least potentially carry wet forest habitat. The distributions of

individual species, however, are not as simply explained.

Gasterogramma psi occupies the north-western third of

Tasmania (fig. 11) from sea level to nearly 1300 m, and is sym-

patric with G. imber, G. rusticum, G. tarkinense and G. wyn-

yardense. There is no obvious ecological reason why G. psi

could not extend into the ranges of G. austrinum and G.

extremum in the south, or into the G. plomleyi range in the

north-east (the closest known approach to the latter is only 12

km through formerly forested country).

The seven newly described species in the genus form an

allopatric/parapatric spatial mosaic (fig. 12). In north-east

Tasmania, the G. plomleyi range is closely congruent with that

of the land snail Anoglypta launcestonensis (Reeve, 1853).

These two distributions partly define the area of endemism

known as Plomleys Island (Mesibov, 1994), which is also char-

acterised by the presence of a number of other narrow-range

endemic invertebrates and by the absence of some wide-

ranging species. On the north coast, the 30-km-wide gap

between the ranges of G. wynyardense and G. rusticum is now

largely farmland. A search of forest remnants in the gap has so

far yielded only G. psi, so it is not known whether G. wyn-

yardense met G. rusticum in parapatry before forests in the gap

were cleared for agriculture. Parapatric boundaries may yet be

found between G. wynyardense and G. tarkinense and between

G. tarkinense and G. imber in the relatively inaccessible forests

of the Arthur-Pieman area in north-west Tasmania, and between

G. austrinum and G. extremum in the wilderness forests of the

upper Picton R. catchment. The Gordon River Rd in south-west

Tasmania crosses the G. imber/G. austrinum parapatric zone

and the easy access afforded by this road has facilitated pre-

liminary fine- scale mapping of the two species. As with para-

patric millipedes in north-east Tasmania (Mesibov, 1997), the

two ranges overlap in a zone less than 1 km wide, and a site has

been found along the road where G. austrinum and G. imber

co-occur in litter at the base of a single large tree (see separate

specimen table). Possible historical explanations for the

mosaic distribution pattern seen in Gasterogramma will be

discussed elsewhere.

Where rotting logs and deep, well-drained forest soils are

available as refuges, the four northern Gasterogramma species

(G. plomleyi, G. psi, G. rusticum and G. wynyardense) and the

southern G. austrinum appear to tolerate burning, logging and

partial clearing of their wet forest habitat. G. psi and G. wyn-

yardense are also known to tolerate replacement of native

forest by exotic tree plantation. A survey of litter invertebrates

in north-west Tasmania (Bonham et ah, 2002) found these two

species to be equally abundant in forest plantations (mainly

Pinus radiata) and nearby native forest. Remarkably, G. psi has

been collected in tiny (<0.25 ha) degraded remnants of riparian

eucalypt forest on farms, having evidently persisted in the litter

under a few surviving eucalypt trees for many years.

Although G. rusticum and G. wynyardense have lost sub-

stantial portions of their ranges to agriculture on the more

fertile soils of northern Tasmania, all eight Gasterogramma

species can be regarded as well conserved. All are known to

occur (or are likely to occur) in formal, forested reserves with-

in their respective ranges, and all seem likely to persist indefi-

nitely outside reserves under land management regimes which

perpetuate closed forest cover, i.e. logging and regeneration of

native forest, and plantation forestry.

Acknowledgments

I am grateful to the Plomley Foundation for financial assis-

tance, to David Steele (University of Tasmania) for acquiring

the SEM images.

References

Bonham, K.J., Mesibov, R., and Bashford, R. 2002. Diversity and

abundance of some ground-dwelling invertebrates in plantation vs.

native forests in Tasmania, Australia. Forest Ecology and

Management 158; 237-247.

Chamberlin, R.V. 1957. Reports of the Lund University Chile

Expedition 1948-49. 30. The Diplopoda of the Lund University and

California Academy of Sciences Expeditions. Lunds Universitets

Arsskrift (N.E. Avd. 2) 53(8): 1-44.

Demange, J.M., and Silva, E. 1976a. Etudes de quelques genres

chiliens de la famille des Sphaerotrichopidae (Myriapodes,

Diplopodes, Polydesmoidea). Bolletino del Laboratorio di

Entomologia Agraria “Filippo Silvestri” di Portici 33: 15-33.

Demange, J.M., and Silva, E. 1976b. Contribution a la connaissance

des especes originaires du Chili decrites par E. Silvestri en 1902.

Bolletino del Laboratorio di Entomologia Agraria “Filippo

Silvestri” di Portici 33: 34-43.

Jeekel, C,A.W. 1982. Millipedes from Australia, 4; a new genus and

species of the family Dalodesmidae from Tasmania (Diplopoda,

Polydesmida). Bulletin Zoologisch Museum, Universiteit van

Amsterdam 9(2): 9-15.

Mesibov, R. 1994. Eaunal breaks in Tasmania and their significance for

invertebrate conservation. Memoirs of the Queensland Museum

36(1): 133-136.

Mesibov, R. 1997. A zoogeographical singularity at Weavers Creek,

Tasmania. Memoirs of the Museum of Victoria 56: 563-573.

Shelley, R.M., Sierwald, P, Kiser, S.B., and Golovatch, S.I. 2000.

Nornenclator generum et familiar urn Diplopodorum II. A list of the

genus and family-group names in the Class Diplopoda from 1958

through 1999. Pensoft; Sofia.

New Tasmania species of the millipede Gasterogramma

219

Figure 11. Localities (squares) for G. psi males. Circled and labelled

localities correspond to the four sites noted in fig. 6: A, Washpond

Forest, B, Notley Gorge, C, Coles Creek, D, Henty R. “TL” points to

the type locality, Hellyer Gorge.

Figure 12. Localities for males of G. austrinum sp. nov. (■ south), G.

extremum sp. nov. (+, far south), G. imber sp. nov. (A, west and south-

west), G. rusticum sp. nov. (□, north-central), G. tarkinense sp. nov.

(*, north-west) and G. wynyardense sp. nov. (•, north-west), and for all

specimens of G. plomleyi sp. nov. (half-filled squares, north-east).

Figure 13. All known localities for Gasterogramma spp. (■). Shaded

areas have annual rainfall of at least 1000 mm.

Memoirs of Museum Victoria 60(2): 221-227 (2003)

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

http://www.museum.vic.gov.au/memoirs

New species of Goreopagurus (Decapoda: Anomura: Paguridae) from Tasmania

and reevaluation of sexual tubes in hermit crab systematics

Rafael Lemaitre' and Patsy A. McLaughlin^

'Department of Systematic Biology, National Museum of Natural History, Smithsonian Institution, Washington, D.C.

20013-7012, USA (lemaitre.rafael@nmnh.si.edu)

-Shannon Point Marine Center, Western Washington University, 1900 Shannon Point Road, Anacortes, Washington

98221-908 IB, USA (hermit@fidalgo.net)

Abstract Lemaitre, R. and McLaughlin, P.A. 2003. New species of Goreopagurus (Decapoda: Anomura: Paguridae) from

Tasmania and reevaluation of sexual tubes in hermit crab systematics. Memoirs of Museum Victoria 60(2): 221-227.

Anew, deep-water species is assigned to the hermit crab genus Goreopagurus McLaughlin, 1988, previously known

only from western Atlantic and eastern Pacific waters. The assignment is made because of the unusual and sexually

dimorphic expansion of the right cheliped characteristic of species of this genus. However, in contrast to its congeners,

Goreopagurus poorei sp. nov. has distally quadriserial gills and exhibits no male sexual tube development. The latter

condition has made it necessary to reevaluate the emphasis placed on sexual tubes in paguroid systematics.

Goreopagurus has been emended to accommodate this species.

Keywords Cmstacea, Anomura, Paguroidea, Paguridae, Goreopagurus, taxonomy

Introduction

McLaughlin (1988) proposed the genus Goreopagurus for a

small hermit crab, Pagurus piercei Wass, 1963. Wass’ (1963)

original description of P. piercei was based on a single male

from off Port Aransas, Texas, USA., and while the species was

found in substantial numbers in the Middle Atlantic Bight

(Wenner and Boesch, 1979), it was not until it was collected off

the east Florida continental shelf that a critical morphological

examination was conducted. McLaughlin (1988) found that

adult females of P. piercei were provided with paired and mod-

ified first pleopods; mature males were found to have a short

sexual tube produced from the gonopore on the right fifth coxa,

and frequently a very short tube or papilla was produced from

the left gonopore. At that time, only species of Nematopagurus

A. Milne-Edwards and Bouvier, 1892, and “Pagurodes” limat-

ulus Henderson, 1888 (subsequently reassigned to

Michelopagurus McLaughlin, 1997) were known to have sexu-

al modifications in both sexes. McLaughlin (1988) reasoned

that Pagurus piercei could not be assigned to Nematopagurus

because of the structure of the male sexual tube, which was

long and distally filiform, rather than short and “stubby”.

Although Pagurus piercei and Pagurodes limatulus agreed in

the form of the sexual tube and the female paired first pleopods,

McLaughlin (1988) considered them not to be congeneric

because the gills of Pagurus piercei were biserial, the chelipeds

were grossly unequal and the ambulatory dactyls relatively

short. In contrast, the gills of Pagurodes limatulus were

quadriserial, the chelipeds were subequal and the ambulatory

dactyls relatively long.

In addition to the chelipeds being grossly unequal in

Goreopagurus piercei, the right cheliped was distinctively sex-

ually dimorphic. The carpus in small males and females was

somewhat broader than the elongate chela, but with increased

size, the male carpus developed a marked flare of the dorso-

mesial margin, the dorsal surface doubled or tripled in breadth,

the lateral face became appreciably produced ventrally and the

ventral surface became noticeably concave. The chela became

correspondingly more elongate, while surface and marginal

spination was reduced.

McLaughlin and Haig (1995) described a second species

from the eastern Pacific. Although paired first pleopods were

present in females, the male right sexual tube was distinctly

shorter in G. garthi McLaughlin and Haig, 1995, and the left

tube, if developed at all, was even shorter. Despite this differ-

ence in sexual tube development, McLaughlin and Haig had no

difficulty in assigning their new species to Goreopagurus,

because the dimorphic right cheliped, while not identical to that

of G. piercei, was morphologically very similar and showed

comparable changes in the carpus and chela in large males.

Mature males of the Tasmanian species, Goreopagurus poorei

222

Rafael Lemaitre and Patsy A. McLaughlin

sp. nov. exhibit the same development and dimorphism of the

right cheliped, but have no trace of sexual tube development.

Additionally, the gills in the new species are distally quad-

riserial (Fig. la), whereas the gills of both G. piercei and

G. garthi are biserial. Nevertheless, in all other morphological

attributes, G. poorei agrees well with the generic diagnosis. The

genus is herein emended to accommodate this new species.

Shield length is measured from the tip of the rostrum to the

midpoint of the posterior margin of the shield. The ratio of

corneal diameter to ocular peduncle length was obtained by

measuring the length of the left ultimate peduncular segment,

including the cornea along the lateral surface; corneal diameter

was the maximum measured width of the left cornea. Sexual

tube lengths are based on the criterion proposed by McLaughlin

(2003). The holotype and most paratypes are deposited in

Museum Victoria, Melbourne, Australia (NMV); three

paratypes have been deposited in the National Museum of

Natural History, Smithsonian Institution, Washington, DC,

USA (USNM).

Goreopagurus McLaughlin, 1988

Emended diagnosis. Eleven pairs of biserial or distally quad-

riserial phyllobranchiate gills. Ocular acicles triangular, with

prominent submarginal spine, separated basally by basal width

or more of 1 acicle. Antennal peduncle with supernumerary

segmentation. Maxillule with internal lobe moderately well

developed and provided with 1 or 2 stiff distal bristles; external

lobe produced, not recurved. Third maxilliped with well devel-

oped crista dentata and prominent accessory tooth; merus with

or without dorsodistal and ventral marginal spines. Sternite of

third maxillipeds with or without small spine on either side of

midline.

Right cheliped with elongate, slender chela; propodal-carpal

articulation generally perpendicular. Carpus tending to be

strongly produced ventrally, at least in large males, dorsomesial

margin slightly to prominently expanded. Left cheliped appre-

ciably shorter than right; chela slender, triangular in cross-

section; propodal-carpal articulation perpendicular. Sternite of

third pereopods with anterior lobe variable in shape. Fourth

pereopods with propodal rasp consisting of single row of cor-

neous scales; dactyl with or without preungual process. Fifth

pereopods chelate. Sternite of fifth pereopods consisting of 2

ovate or subcircular lobes.

Mature males with coxae of fifth pereopods generally sym-

metrical; gonopore of right without or with vas deferens pro-

duced as short sexual tube; gonopore of left without or with vas

deferens produced as very short tube or papilla; 3 unpaired uni-

ramous or weakly biramous pleopods developed on left side of

abdominal somites 3-5, very rarely vestigial pleopod on somite

2. Females with paked gonopores; well developed, paired and

modified first pleopods; 4 unpaired pleopods, second to fourth

unequally biramous, fifth with endopod rudimentary or absent.

Abdomen flexed. Uropods asymmetrical. Telson with trans-

verse indentations; posterior lobes symmetrical or slightly

asymmetrical; separated by small median cleft; terminal

margins oblique or straight, each armed with few to several

small spines; lateral margins frequently delimited by narrow

chitinous plate.

Goreopagurus poorei sp. nov.

Figures 1-3

Michelopagurus n. sp. — Koslow and Gowlett-Holmes, 1998: 32.

Michelopagurus sp. — ^Poore et al., 1998: 71.

Material examined. Holotype. Australia, Tasmania, 84 km SSE of SE

Cape, “Jl” seamount, 44°16.2'S, 147°19.8^, 1300 m, 27 Jan 1997

(CSIRO stn SSOl/97 37), NMV J44757 (male, 5.2 mm).

Paratypes. Australia, Tasmania, off Ereycinet Peninsula, 4r58.6'S,

148°38.9'E, 500 m, 27 Jul 1986 (stn SLOPE 47), NMV J17433 (2

males, 3.6, 4.2 mm). 85.4 km SSE of SE Cape, “Main Pedra”

seamount, 44°15.6'S, 147°06.0'E, 741 m, 21 Jan 1997 (CSIRO stn

SSOl/97 03), NMV J44807 (5 males, 2.9-3.6 mm); NMV J44767 (2

ovigerous females, 2.4, 2.7 mm). 83.8 km SSE of SE Cape, “Jl”

seamount, 44°16.2'S, 147°19.8'E, 987 m, 27 Jan 1997 (stn SSOl/97

36) , NMV J448051 (male, 2.4 mm; ovigerous female, 2.6 nun);

USNM 1007889 (female, 3.7 mm). 84.0 km SSE of SE Cape “Jl”

seamount, 44°16.2'S, 147°19.8'E, 1300 m, 27 Jan 1997 (stn SSOl/97

37) , NMV J52355 (3 males, 3.3^.9 mm; ovigerous female, 3.2 mm).

85.8 km SSE of Cape, “Bl” seamount, 44° 18.6 'S, 147°16.2U, 1150 m,

28 Jan 1997 (stn SSOl/97 43), NMV J448021 (male, 3.0 mm). 69.7 km

SSE of SE Cape, “Mackas” seamount, 44°12.6"S, 147°02.4'E, 640 m,

29 Jan 1997 (stn SSOl/97 50), NMV J44808 (male, 3.0 mm). 65.5 km

SSE of Cape, SE “Andys” seamount, 44° 10.8 'S, 147°00.0'E, 800 m,

29 Jan 1997 (stn SSOl/97 56), NMV J44804 (5 males, 2.9-4.7 mm ),

USNM 1007890 (2 males, 2.6, 3.9 mm). 65.1 km SSE of Cape, SE

“Andys” seamount 44°10.8'S, 146°59.4T, 900 m, 29 Jan 1997 (stn

SSOl/97 57), NMV J44809 (male, 3.0 mm).

Other material. Australia, Tasmania. 66.5 km SSE of Cape, SE

“Andys” seamount, 44° 11. 4 'S, 148°57.0'E, 620 m, 29 Jan 1997 (stn

SSOl/97 55), NMV J44806 (2 males, 2.9, 3.4 mm). 65.6 km SSE of

Cape, SE “Andys” seamount, 44°10.8'S, 147°00.0"E, 800 m, 29 Jan

1997 (stn SSOl/97 56), NMV J44764 (ovigerous female, 2.6 mm).

Exact locality not recorded (CSIRO cmise 1/97), NMV J44810 (male,

2.4 mm; 3 ovigerous females, 2.4-3. 3 mm).

Description. Gill lamellae distally quadriserial (Fig. la). Shield

(Fig. lb) as broad or broader than long; anterior margins

between rostrum and lateral projections weakly concave;

anterolateral margins sloping or slightly terraced; posterior

margin roundly truncate, frequently with slight median con-

cavity; dorsal surface with few sparse tufts of setae. Rostrum

(Fig. Ib-d) acutely or obtusely triangular, with or without small

terminal spine or spinule, sometimes also with 1 small spine or

spinule laterally on one side. Lateral projections obtusely

triangular, each with small submarginal spine.

Ocular peduncles 0.4-0.6 length of shield, moderately stout

and with distinct medial constriction; dorsomesial surface with

2 or 3 short, transverse rows of stiff setae in distal half, 1 sparse

tuft of setae on dorsal surface; corneal diameter 0.4-0. 5 of

peduncular length. Ocular acicles narrowly and acutely trian-

gular, with strong submarginal spine; separated basally by at

least basal length of 1 acicle.

Antennular peduncles overreaching distal margins of

corneas by at least entire length of ultimate segments. Ultimate

segment with row of long, stiff setae at dorsodistal margin and

additional few scattered, short setae. Penultimate segment with

few scattered setae. Basal segment with slender spine on

lateral margin of statocyst lobe.

Antennal peduncles overreaching distal comeal margins by

New Tasmanian hermit crab and reevaluation of sexual tubes

223

Figure 1. Goreopagurus poorei sp. nov.: paratypes: a-c, f, male (3.9 mm), USNM 1007889; d, male (si =2.6 mm), USNM 1007890; e, female

(3.7 mm), USNM 1007889. a, gill lamella; b, shield and cephalic appendages, dorsal view; c, d, rostrum, dorsal view; e, female coxae, eight tho-

racic sternite, and first abdominal somite with paired pleopods, ventral view; f, telson, dorsal view.

0.7 to entire length of ultimate segments. Fifth segment with

several sparse tufts of setae. Fourth segment with few scattered

setae. Third segment with prominent, acute spine at ventro-

distal margin. Second segment with dorsolateral distal angle

produced, terminating in acute spine, mesial margin with 3-7

small spines or spinules (occasionally only weakly apparent)

and usually with 1 prominent spine on lateral margin in distal

half; dorsomesial distal angle with well developed spine. First

segment usually with small spine on dorsolateral distal margin

and 1 spine on ventrolateral margin distally. Antennal acicle

long, reaching beyond midlength of ultimate peduncular seg-

ment, arcuate, terminating in small spine; mesial surface with

row of tufts of stiff setae. Antennal flagella long, but usually not

overreaching tip of outstretched right cheliped; articles each

usually with 2 or 3 short (< length of 1 article) setae.

Third maxilliped with 2 or 3 spines on basis; ischium with

accessory tooth on well developed crista dentata; merus with 1

spine on ventral margin and 1 spine at dorsodistal margin.

Sternite of third maxillipeds with tiny spinule on either side of

midline.

Right cheliped (Fig. 2a-e) of large males very elongate,

somewhat shorter in females and small males; palm, fixed

finger and dactyl slender, dorsoventrally compressed. Dactyl

0.7-0.9 length of palm; cutting edge with 1 moderately promi-

nent calcareous tooth at midlength, 3 or 4 smaller calcareous

teeth proximally and row of very small corneous teeth in distal

0.3, terminating in small corneous claw, but often worn;

dorsomesial margin with row of very small spines, spinules or

spinulose tubercles, dorsal surface somewhat elevated in mid-

line, occasionally unarmed but usually with single or double

row of very small spinules, tubercles or protuberances. Palm

slightly shorter than carpus; dorsomesial margin not distinctly

delimited, rounded mesial face with irregular rows of very

small tubercles, granules or spinules; dorsolateral margin with

row of very small spinules, dorsal surface with covering of very

small spinules or granules and very short setae, most prominent

in females and small males, occasionally additional short row

of slightly larger spinules or granules adjacent to dorsolateral

margin proximally, dorsal midline with 1 or short, longitudinal

row of 2-4 small spines proximally; cutting edge of fixed fin-

ger with entire calcareous margin in proximal half, usually few

individual calcareous teeth distally, terminating in corneous

claw, often worn. Carpus (Fig. 2b, c, e) longer than merus, pro-

duced ventrally, particularly in large males; dorsomesial margin

weakly to notably expanded, armed with row of prominent,

blunt or acute spines, dorsodistal margin with 1 or 2 spines

mesially, dorsal surface usually with numerous very small

tubercles, granules or low protuberances and scattered setae,

dorsolateral margin usually delimited at least distally by row of

small granules or spinules; mesial face usually sloping, often

224

Rafael Lemaitre and Patsy A. McLaughlin

Figure 2. Goreopagurus poorei sp. nov.: a-c, f, paratype male (3.9 mm), USNM 1007890; d, e, holotype male (5.2 mm), J44757. a, mems, car-

pus and chela of right cheliped, dorsal view; b, carpus of same, lateral view; c, same, mesial view; d, carpus and chela of right cheliped, dorsal

view; e, carpus of same, ventromesial view; f, mems, carpus and chela of left cheliped, dorsal view.

Figure 3. Goreopagurus poorei sp. nov.: paratype male (3.9 mm), USNM 1007890. a, second right pereopod, lateral view; b, dactyl of same,

mesial view; c, third right pereopod, lateral view; d, dactyl of same, mesial view; e, anterior lobe of sternite of third pereopods, ventral view;

f, propodus and dactyl of left fourth pereopod, lateral view.

New Tasmanian hermit crab and reevaluation of sexual tubes

225

appearing somewhat concave, ventromesial margin unarmed;

ventrolateral margin with row of granules, tubercles or spines.

Merus subtriangular; dorsodistal margin sometimes with 1

spine, dorsal surface with few low protuberances and setae;

ventromesial and ventrolateral margins each with row of

spines, ventral surface usually with few scattered spines and

sparse, moderately long setae. Ischium with row of small

spinules on ventromesial margin, at least proximally, and scat-

tered moderately long setae.

Left cheliped (Fig. 2f) long and slender, but not reaching

much beyond proximal margin of palm of right in large males;

dactyl and fixed finger weakly arched ventrally. Dactyl 1. 5-2.0

length of palm; cutting edge with row of tiny corneous teeth,

terminating in small corneous claw; dorsal surface elevated in

midline and with sparse row of setae; dorsomesial margin with

irregular row of small spinules or tubercles, at least in proximal

half. Palm 0.4-0.6 length of carpus; dorsomesial margin with

irregular row of small spines or tubercles, dorsal surface

usually with covering of very short setae, midline prominently

elevated and armed with row of small spines, extending onto

fixed finger but usually not to tip, dorsolateral margin with row

of small spinules or granules, adjacent dorsal surface granular;

ventral surface with scattered tufts of long setae; cutting edge

of fixed finger with row of very small calcareous teeth, inter-

spersed distally with corneous teeth. Carpus slightly shorter to

slightly longer than merus; dorsal surface somewhat flattened,

dorsomesial and dorsolateral margins each with row of spines,

strongest distally; mesial face with numerous protuberances

and setae, distomesial margin with few spinules ventrally,

sometimes extending onto ventromesial margin; ventral surface

with few spinules or low protuberances and scattered, moder-

ately long setae; lateral face unarmed, ventrolateral margin with

row of spines. Merus subtriangular; dorsal surface with few

low protuberances and sparse setae; ventromesial and ventro-

lateral margins each with row of spines and moderate to

long setae, ventral surface frequently with moderately dense

covering of short setae. Ischium with row of small spines or

spinules on ventromesial margin and scattered moderately

long setae.

Ambulatory legs (Fig. 3a-d) similar from left to right.

Dactyls 1.2-1. 4 length of propodi; dorsal margins each with

sparse row of short stiff setae; ventral margins each with row of

10-13 corneous spines; lateral faces each with weak longitud-

inal sulcus; mesial faces each with row of corneous spinules

dorsally. Propodi 1.3-1. 7 length of carpi; each with 1 or 2 cor-

neous spinules at ventrodistal angle and row of widely-spaced,

small corneous spinules on ventral margin, dorsal surfaces each

with low protuberances and sparse stiff setae or bristles. Carpi

each with small spine at dorsodistal margin, few stiff setae aris-

ing from low protuberances on dorsal surface, 1 or 2 stiff setae

on ventral margin distally and occasionally also medially. Meri

with low protuberances and sparse tufts of setae on dorsal sur-

faces, tufts of stiff setae and occasionally 1 to few, often spinu-

lose, protuberances or small spinules on ventral surfaces, distal

angles of second pereopods occasionally each with acute spine

(not present in holotype). Ischia unarmed but with few stiff

setae or bristles. Anterior lobe of sternite of third pereopods

(Fig. 3e) subsemicircular with stiff bristles on anterior margin.

Fourth pereopods (Fig. 3f) with small preungual process at base

of claw.

Males without sexual tube on either coxa of fifth pereopods;

unpaired left pleopods 3-5 with endopods rudimentary, very

rarely with vestigial second pleopod present. Females with

paired and modified first pleopods (Fig. le); second to fourth

unpaired left pleopods with endopods stout, egg-bearing,

exopods long, slender and usually non egg-bearing; fifth pleo-

pod as in males, Uropods markedly asymmetrical. Telson (Fig.

If) with transverse indentation; posterior lobes separated by

moderately shallow median cleft, terminal margins slightly

oblique to nearly horizontal, each with 3-5 small spines,

lateral margins sometimes delimited by narrow chitinous band.

Colour (in preservative). Cephalothorax, carpi and meri of

chelipeds and entire ambulatory legs with tint of reddish-

orange; chelas dirty-white. Ocular peduncles dull red.

Distribution. Tasmania, off Freycinet Peninsula and seamounts

SSE of Southeast Cape: “Andys”, “Bl”, “Jl”, “Main Pedra”,

and “Mackas” seamounts; 500-1300 m depth.

Etymology. For Gary C. B. Poore, Curator of Crustacea at

Museum Victoria, Melbourne, Australia, in recognition of his

major contributions to crustacean systematics.

Remarks. On the basis of a few individuals, this species was

tentatively but incorrectly assigned to Michelopagurus by P.A.

McLaughlin in correspondence to Koslow and Gowlett-

Holmes (1998).

Individuals of Gorepagurus poorei are significantly larger

than those of G. piercei and G. garthi. Specimens range in

shield length from 2.4 to 5.2 mm, whereas the specimens of G.

piercei and G. garthi, all sexually mature and including some

ovigerous females recorded by McLaughlin (1988) and

McLaughlin and Haig (1995) range from 0.5 to 1.8 mm, and

1.3 to 2.3 mm, respectively. McLaughliin (1988) and

McLaughlin and Haig (1995) attributed variation of the right

cheliped, particularly the carpus, of G. piercei and to some

extent also G. garthi, to sexual dimorphism. Although speci-

mens of G. poorei as small as those of G. piercei and G. garthi

are not known, it appears that in G. poorei variations of the

carpus are related to allometric growth rather than sexual

dimorphism. The dorsomesial margin of the carpus in large

specimens of G. poorei develop a more marked flare (Fig.

2a-d) than in smaller specimens; the strength and sharpness of

the spines on the dorsomesial and ventrolateral margins of the

carpus diminish with increased size.

Not only is G. poorei immediately distinguishable from

R piercei and P. garthi by the absence of male sexual tubes, the

gill lamellae are distally quadriserial rather than biserial.

Although uniformity in gill structure is usual in most genera,

Goreopagurus is not the only genus in which both biserial and

distally quadriserial gills are found. A similar condition exists

in the parapagurid genus Sympagurus (Lemaitre, in press; fig.

1) and the pagurid Xylopagurus A. Milne-Edwards, 1880 (pers.

obs.). McLaughlin and de Saint Laurent (1998) reported that

gill lamellae vary from deeply quadriserial to only distally so

within a single species or even within a single individual, De

Saint Laurent-Dechance (1966) considered quadriserial gills

226

Rafael Lemaitre and Patsy A. McLaughlin

more primitive than biserial gills, thus one might envision the

evolution of the gill lamellae from completely divided to dis-

tally divided to entire (biserial). In the case of the three species

of Goreopagurus, the most primitive species then would be

G. poorei. Support for this hypothesis is also apparent in the

transition from no sexual tubes in G. poorei to very short tubes

in G. garthi to short tubes in G. piercei.

Sexual tube development and its role in systematics of

Paguroidea

H. Milne Edwards (1837) described specialised tubular stmc-

tures found on the coxae of the fifth pereopods in a species of

the semiterrestrial hermit crab genus Coenobita Latreille, 1829.

De Haan (1849) described an apparently similar structure in the

pagurid, Pagurus spiriger De Haan, 1849, a species subse-

quently transferred to the genus Spiropagurus Sdmpson, 1858.

Although these tubular structures on the coxae of male pagurids

and coenobitids were recognised by early carcinologists (e.g.,

Stimpson, 1858; Miers, 1881; Henderson, 1888; A. Milne-

Edwards and Bouvier, 1892, 1893; Borradaile, 1903; Alcock,

1905), it was de Saint Laurent-Dechance (1966) and de Saint

Laurent (1968, 1970a, 1970b) in particular who discussed their

role in the systematics of Paguridae. Importantly, she pointed

out that early classifications had placed species in genera on the

basis of tube position, without consideration of other charac-

ters, and as a result, species with disparate morphologies often

were assigned to the same genus. At that time, only eight

genera had been described with male sexual tubes. There are

now many more (McLaughlin, 2003) and the number has

increased dramatically in the past 35 years.

Two types of sexual tubes can be differentiated, although

most descriptions make no distinction. Those in several, but not

all, species of Coenobita represent prolongations of the coxae

that are heavily calcified, and are diagnostic at the specific level

(Nakasone, 1988). In contrast, sexual tubes in Paguridae are

diverse and may be coxal prolongations or external prolonga-

tions of the vas deferens that may be calcified, chitinous or

membranous. Their development and form have had a major,

although not always informed, impact on classification at the

generic level. McLaughlin’s (2003) key to genera emphasises

the length and direction of sexual tubes but there are many

instances where intrageneric variation requires alternative

pathways. Her figures 7h-t illustrate the varibility.

De Saint Laurent (1968, 1970a, 1970b) was of the opinion

that just the presence or position of tubes was not indicative of

close phylogenetic relationships. McLaughlin and Lemaitre

(2001), and Lemaitre and McLaughlin (2003) emended the

diagnoses of Pylopagurus A. Milne-Ed wards and Bouvier,

1891, Enallopaguropsis McLaughlin, 1981, and Enallopagurus

McLaughlin, 1981, to indicate minor sexual tube development

in some species. Their rationale was that members of each

genus shared suites of other morphological characters. In con-

trast, Komai (1998, 1999) transferred four species from

Pagurus to Parapagurodes McLaughlin and Haig, 1973 simply

because males were found to have very short sexual tubes. As

a result, the presence of a right sexual tube is the only com-

monly shared character among these species currently assigned

to Parapagurodes. On the basis of cheliped morphology

and other attributes, P gracilipes (Stimpson, 1858) and

P nipponensis Yokoya, 1933 appear most closely related to

McLaughlin’s (1974) ''bernhardus'' group of Pagurus.

Similarly Parapagurodes imalli (Yokoya, 1939) and P. con-

stans (Stimpson, 1858), while not immediately identifiable

with specific groups within Pagurus, are not phylogenetically

related to the species for which the genus Parapagurodes was

proposed. This view is also supported by the larval data of

Hong and Kim (2002) for P. constans.

Correspondingly, Asakura (2001; 827) transferred

Catapagurus doederleini Doflein, 1902 to Parapagurodes,

stating that Doflein’s (1902) species agreed in all diagnostic

characters with McLaughlin and Haig’s (1973) genus. Yet

Asakura (2001: 888) pointed out how the structure of the

ambulatory legs of P. doederleini differed from all species

presently included in Parapagurodes. Additionally, there are

marked differences in telson structure of P doederleini and all

of the other assigned species. The only shared character of any

potentially phylogenetic significance is the presence in P doed-

erleini, as in the other species, of a very short right sexual tube

and a slight protrusion of the vas deferens from the left gono-

pore. We concur with Asakura’s conclusion that Doflein’s

(1902) species is not assignable to Catapagurus or

Hemipagurus as redefined; however, its inclusion in

Parapagurodes is equally inappropriate.

Our only precise information on the structure and function

of hermit crab sexual tubes is the recent study by Tudge and

Lemaitre (2003) of Micropagurus acantholepis (Stimpson,

1858), a species with a moderate to long (> 2 coxal lengths) left

sexual tube. These authors demonstrated that a well developed

sexual tube is used in the transport of spermatophores and

described the ultrastructure. Sexual tube structure is just one of

a number of morphological characters that should be consid-

ered when making generic evaluations.

Acknowledgements

We are indebted to Dr Gary C. B. Poore, Museum Victoria, for

making the material available for study. This is, in part,

a scientific contribution from the Shannon Point Marine Center,

Western Washington University.

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Museum Victoria 60: 111-144.

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Memoirs of Museum Victoria 60(2): 229-236 (2003)

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

http://www.museum.vic.gov.au/memoirs

A new genus and species of hermit crab (Decapoda: Anomura: Paguridae) from

seamounts off south-eastern Tasmania, Australia

Patsy A. McLaughlin

Shannon Point Marine Center, Western Washington University, 1900 Shannon Point Road, Anacortes, WA 98221-9081B,

USA (hermit@fidalgo.net)

Abstract McLaughlin, PA. 2003. A new genus and species of hermit crab (Decapoda: Anomura: Paguridae) from seamounts off

south-eastern Tasmania, Australia. Memoirs of Museum Victoria 60(2): 229-236.

A new and highly variable species, representing a new genus of hermit crabs, is described and illustrated. In addition

to the morphological changes in shape and armature of the chelipeds associated with growth in Bythiopagurus macro-

colus, gen. nov., sp. nov., this species exhibits a singular development of the left chela that may be an adaptation to its

specialised habitat among colonies of the stony coral Solenosmilia variabilis Duncan. The similarities seen among the

genera Bythiopagurus, gen. nov., Michelopagurus McLaughlin and Icelopagurus McLaughlin appear the be superficial

and convergent; Bythiopagurus appears phylogenetically related to the Pylopaguropsis group of genera characterised by

13 pairs of bi- or quadriserial gills.

Key words Cmstacea, Anomura, Paguridae, new genus, new species, Tasmania, seamounts, Bythiopagurus

Introduction

The seamount region of south-eastern Tasmania is a distinctive

deep-water environment extensively surveyed for the first time

between 20 January and 1 February 1997. Among the approxi-

mately 242 species of invertebrates cited by Koslow and

Gowlett-Holmes (1998) in their report on the survey, only three

were hermit crabs. Two of the three were new to science.

Propagurus deprof undis (Stebbing, 1924), described initially

from South Africa, was reported by McLaughlin and de Saint

Laurent (1998) to range from South Africa to Hawaii, and had

been collected previously in Tasmanian waters (McLaughlin,

unpub.). Of the latter two, the species of Goreopagurus

described by Lemaitre and McLaughlin (2003) is the first truly

deep-water record (620-1300 m) for this genus, and the first

report of the genus outside the continental waters of Atlantic

and Pacific coasts of the United States. The unusual species

reported herein represents both a new species and a new genus

that is noteworthy not only for its characteristic growth-related

variability, but for its unusual habitat. It was found only in asso-

ciation with colonies of the stony coral, Solenosmilia variabilis

Duncan, 1873.

Measurements include shield length, measured from the tip

of the rostrum to the midpoint of the posterior margin of the

shield. However, in this genus, as in a few others, e.g.

Goreopagurus McLaughlin, 1988, Alainopaguroides

McLaughlin, 1997, the shield breadth: length ratio increases

appreciably with increased animal size, thus making the

measurement of shield length less informative than for most

pagurid genera. The ratio was calculated of corneal diameter

(maximum measured width of the left cornea) to length of the

ocular peduncle (length of the left ultimate peduncular seg-

ment, including the cornea, along the lateral surface). Male

sexual tube development varies from a simple papilla-like pro-

trusion to very short (less than coxal length) tube(s), but

occasionally the vas deferens may not be extruded at all. The

holotype and most paratypes are deposited in Museum Victoria,

Melbourne, Australia (NMV); and three paratypes in the

National Museum of Natural History, Smithsonian Institution,

Washington, DC, USA (USNM).

Terminology. For the most part, the terminology utilised in

the species description follows that of McLaughlin (1997) but

the interpretation of gill structure is based on the definition of

Lemaitre (in press). The varying extent to which the paguroid

cephalothorax is delineated and/or calcified has resulted in a

certain amount of confusion regarding the correct terminology

to be applied to various portions (e.g. Boas, 1880, 1926;

Bouvier, 1895; Borradaile, 1906; Jackson, 1913; Pilgrim, 1973;

McLaughlin, 1974; Forest, 1987). Morgan and Forest (1991)

corrected the misinterpretation by McLaughlin (1974) regard-

ing the sulcus cardiobranchalis and assigned the name cardiac

sulci to the previously unnamed sulci bordering the postero-

median plate. Lemaitre (1995) added another term “accessory

portion” for the calcified portion of the carapace presumably

delineated anteriorly by the cervical groove and posteriorly by

230

Figure 1. Bythiopagurus macrocolus, gen. nov., sp. nov.: a,Jparatype

(6.4 mm), USNM 1007810; b-h, ? paratype (6.0 mm), J44760; a,

cephalothorax; b, gill lamella; c, left mandible (internal view, palp

removed); d, right mandible (internal view); e, left maxillule (external

view, setal details omitted); f, left maxilla (external view, setal details

omitted); g, left maxilliped 1 (external view, setal details omitted); h,

coxa, basis and ischium of left maxilliped 3 (external view, setae omit-

ted). Abbreviations: be: branchiostegal areas of calcification; ebs: sul-

cus cardiobranchialis; eg: cervical groove; cll: carapace lateral lobe;

cs: cardiac sulcus; la, linea anomurica; Ip: lateral projection; It: linea

transversalis; peme: posterior carapace median element; pip: postero-

lateral plate; pmp; posteromedian plate; r: rostmm; s: shield. Scales

equal 1 mm (b-h) and 5 mm (a).

the linea transversalis. These calcified areas have now become

incorporated into species descriptions as “accessory portion of

the shield” (e.g. Asaknra, 2001; McLaughlin and Lemaitre,

2001). Personal examination of the internal structure of the

cephalothorax of Propagurus deprofundis, Porcellanopagurus

tridentatiis Whitelegge, 1900, Solitariopagurus triprobolus

Poupin and McLaughlin, 1996 and the new species have con-

firmed the structure and position of the linea transversalis as

defined by Boas (1926) and Pilgrim (1973). The linea transver-

salis is a chitinous hinge separating the posterior portion of the

shield from the median anterior portion of the posterior cara-

pace. It does not curve anteriorly to delineate areas of calcifi-

cation as illustrated by McLaughlin (1974: fig. 2), Lemaitre

(1995: fig. 1), or McLaughlin and Lemaitre (2001: fig. 2); nor

is it calcified as reported by McLaughlin (1997) and Asakura

(2001). Although its position can be recognised externally, the

linea transversalis itself often is not visible on the surface of the

carapace and may be covered by a calcified plate.

In Bythiopagurus macrocolus gen. nov., sp. nov., the shield

is completely delineated from the surrounding well calcified

integument by the cervical groove (Fig. la), much as it is in the

genera Porcellanopagurus Filhol, 1885 and Solitariopagurus

P. A. McLaughlin

Figure 2. Bythiopagurus macrocolus, gen. nov., sp. nov.:

a-d, 6 paratype (6.4 mm), USNM 1007810; e, d holotype (6.0 mm),

J44765. a, shield and cephalic appendages (dorsal view, aesthetascs

omitted); b, anterior lobe of sternite of pereopods 3; c, dactyl

and propodus of left pereopod 4 (lateral view); d, coxae and sternite

of pereopods 5; e, telson. Scales equal 3 mm (a), 2 mm (b-d) and

1 mm (e).

Tiirkay, 1986 (cf. McLaughlin, 2000: fig. 1). The linea trans-

versalis in B. macrocolus is contiguous with the cervical groove

centrally and delimits the calcified lateral elements posteriorly

on either side of the posterior margin of the shield. In this

respect, the linea transversalis of B. macrocolus is comparable

to that of Porcellanopagurus, which directly abuts the poster-

ior carapace lobes. However, the carapace is not broadened in

B. macrocolus as it is in Porcellanopagurus and

Solitariopagurus, thus the lateral calcified elements do not

appear posterior as they tend to in the latter genera, particul-

arly in Solitariopagurus. Nevertheless, these calcified elements

cannot correctly be considered accessory portions of the shield,

because by definition, the shield is delimited by the cervical

groove (Forest, 1987: 18). Despite the positional homologies in

the three genera, the anterolateral position of these calcified

areas in Bythiopagurus necessitate that they be referred to as

carapace lateral lobes (Fig. la) rather than posterior carapace

lateral lobes. It would appear that these carapace lateral lobes

are partially delimited anterolaterally by the linea anomurica,

but these lineae are difficult to follow posteriorly in the

membranous regions of the branchiostegites. Other terminolo-

gy applied to the elements of the posterior portions of the

A new hermit crab from seamounts off Tasmania

231

carapace follow those used by McLaughlin (2000) for

Porcellanopaguriis and Solitariopagurus.

Bythiopaguriis gen. nov.

Type species. Bythiopaguriis macrocohis sp. nov.

Diagnosis. Shield and carapace lateral lobes well calcified;

posterior carapace with posterior carapace element contiguous

or fused with lateral carapace lobes, posteromedian and pos-

terolateral plates also well calcified at least in anterior halves

(Fig. la). Sulci cardiobranchialis extending approximately 0.50

length of posterior carapace; cardiac sulci extending nearly to

posterior margin. Cervical groove distinct; linea transversalis

usually partially discernable externally; linea anomurica not

readily apparent in posterior portion of carapace. Frequently

small areas of branchiostegal regions weakly calcified .

13 pairs of distally quadriserial (Fig. lb) phyllobranchiate

gills.

Ocular peduncles short and stout; ocular acicles acutely tri-

angular. Antennal peduncles with supernumerary segmentation.

Mandibles sometimes weakly asymmetrical (Fig. Ic, d);

strongly calcified; palp well developed. Maxillule (Fig. le)

with anterior lobe of endopod weakly developed, posterior lobe

moderately well developed, not recurved. Maxilla (Fig. If)

with slender scaphognathite. First maxilliped (Fig. Ig) with

slender two-segmented exopod; no epipod. Third maxilliped

with basis and ischium not fused; crista dentata of ischium well

developed (Fig. Ih), 1 or 2 accessory teeth.

Chelipeds subequal; left sometimes longer, but not stronger;

dactyls and fixed fingers opening in horizontal plane. Fourth

pereopod very weakly semichelate; propodal rasp consisting of

single row of corneous scales (Fig. 2c). Fifth pereopod chelate.

Male usually with papilla or very short sexual tube extruded

from both right and left gonopores (Fig. 2d). Female with

paired gonopores.

Abdomen somewhat reduced, dextrally twisted. Male with

left unpaired pleopods on somites 3-5; exopods moderately

long and very slender, endopods reduced. Female with paired

and modified pleopod 1, unpaired left pleopods 2-4 with both

rami of approximately equal length, endopods much thicker;

pleopod 5 as in male.

Uropods markedly asymmetrical. Telson with transverse

indentations; posterior lobes separated by median cleft; termi-

nal margins each with few small spines.

Etymology. From Greek bythios meaning from the deep, and

pagurous meaning crab, and referring to the archibenthic and

archiabyssal depths from which the type species was collected.

Bythiopagurus macrocolus sp. nov.

Figures 1-3

Paguridae n. gen. n. sp. — Koslow and Gowlett-Hohnes, 1998: 30

(in part, see remarks).

Pagurid sp. — Poore, et al., 1998; 71 (in part, see remarks).

Material examined. Holotype. (6.0 mm), NMV J44765, CSIRO

SSOl/97 stn 41, 44“19.2^S, 147“07.2'E, 1083 m; 82.8 km SSE of SE

Cape “U” seamount, Tasmania, Australia.

Paiatypes. All from Tasmanian seamounts, collected on cruise

CSIRO SSOl/97. Stn 15, 82.9 km SSE of SE Cape “Sister 1”,

44°16.2/S, 147° 17.4 'E, 1100 m, 23 Jan 1997, 6 (2.6-5.9 mm), 9 ?

(3.2-6.4 mm), 5 ovigerous ? (4.2-6.2 mm), NMVd44760. Stn 28, 89.5

km SSE of SE Cape “Kl“, 44°17.4'S, 147°24.6'E, 1225 m, 25 Jan

1997. 4 (4.3-6. 8 mm), 1 ovigerous ? (3.6 mm), NMV J44762. Stn 34,

85.4 knr^SSE of SE Cape “U”, 44°19.8'S, 147°10.2'E, 1083 m, 27 Jan

1997, 3 ? (4.2-5 .3 mm), NMV J44759. Stn 37, 84.0 km SSE of SE

Cape “Jl”, 44°16.2'S, 1477°19.8T, 1300 m, 27 Jan 1997, 1 (4.9

mm), 1 ? (4.2 mm), 1 ovigerous ? (6.2 mm), NMV J44803. Sin 40,

82.6 km SSE of SE Cape “Jl”, 44°14.4'S, 147°21.8T, 1200 m, 27 Jan

1997, 2 (4.4, 5.1 mm), 5 ? (5.1-6.4 mm), 5 ovigerous ? (si = 5. 1-6.3

mm), NMV J44758. Stn 41, 82.8 km SSE of SE Cape “U”, 44°19.2^S,

147°07.2'E, 1083 m, 27 Jan 1997, 1 (4.2 mm), 1 ? (4.3 mm), 3

ovigerous ? (5.0-5.5 mm), NMV J44765; 2 (3.8, 6.4 mm), 1 oviger-

ous ? (5.3 nun), USNM 1007810. Stn 58, ^1.2 km SSE of SE Cape

“38”, 44°13.2'S, 147°22.8'E, 1140 m, 30 Jan 1997, 1 (5.6 mm),

NMV J44768. Stn 59, 81.6 km SSE of SE Cape “38cJ, 44°13.8'S,

147°22.8'E, 1200 m, 30 Jan 1997, 1 (4.5 mm), NMV J44763. Stn 62,

87.8 km SSE ofSE Cape “Al”, 44°lt$.8^E, 147°16.2^, 1200 m, 30 Jan

1997, 1 ovigerous ? (5.4 mm), NMV J44761.

Eigure 3. Bythiopagurus macrocolus, gen. nov., sp. nov.:

a, c, d paratype (2.6 mm), J44760; b, d-f,d paratype (6.4 mm), USNM

1007810. a, b, chela and caipus of right cheliped (dorsal view);

c, d, chela and carpus of left cheliped (dorsal view); e, dactyl and fixed

finger of left chela (ventral view); f, left pereopod 2 (lateral view).

Scales equal 5 mm (b, d-f) and 1 mm (a, c).

232

P. A. McLaughlin

Other material examined. CSIRO SS91/97, no other data, 1 (5.6

mm), NMV J44801.

Description. Shield (Figs, la, 2a) broader than long; anterior

margin between rostrum and lateral projections concave;

anterolateral margins sloping; posterior margin truncate; dorsal

surface with numerous sparse tufts of simple setae. Rostrum tri-

angular, reaching to or slightly beyond bases of ocular acicles;

with or without terminal spinule. Lateral projections reaching

level of rostrum or slightly beyond, each with prominent

marginal or submarginal s|>ine.

Ocular peduncles very short and stout, 0.40-0.50 length of

shield; dorsal, mesial and lateral surfaces calcified, median

longitudinal region of^yentral surface membranous; dorsal sur-

faces of peduncles each with 3 or 4 short transverse rows of

simple setae; corneas reduced dorsally by projection of pedun-

cular integument, corneal diameter 0.40-0.50 of peduncular

length. Ocular acicles each with slender triangular projection,

terminating acutely and with small submarginal spine or

spinule not usually visible dorsally; separated basally by

approximately 0.50 basal width of 1 acicle.

Antennular peduncles overreaching distal margins of

corneas by length of ultimate segments to nearly entire lengths

of penultimate segments. Ultimate segments nearly twice

length of penultimate segments, dorsal surfaces each with 3-5

short transverse rows of simple setae. Penultimate segments

with few sparse tufts of setae. Basal segments each with

slender spine on dorsolateral margin of statocyst lobe.

Antennal peduncles overreaching distal margins of corneas

by entire lengths of fifth and fourth segments, but reaching only

to approximately midlength of ultimate segments of antennular

peduncles. Fifth and fourth segments with few sparse tufts of

short setae. Third segment with spine at ventrodistal margin.

Second segment with dorsolateral distal angle prominently pro-

duced, terminating in small spine and with 1 or 2 small spines

on mesial margin; dorsomesial distal margin with well devel-

oped spine. First segment sometimes with spine on dorsolater-

al distal margin, ventrolateral margin with 1 spine. Antennal

acicles reaching to or nearly to distal margin of fifth pedun-

cular segment, terminating in small spine; mesial margin with

row of sparse tufts of stiff setae.

Mandibles with upper incisor edge of left (Fig. Ic) with 2 to

several calcareous denticles, cutting edge with corneous-tipped

tooth at lower angle, sometimes also at upper angle, stout inter-

nal ridge presumably representing molar process with usually

corneous-tipped tooth at lower angle; right mandible with

upper edge of incisor process (Fig. Id) unarmed or with 1 or 2

calcareous denticles, upper and lower angles of cutting edge

each usually with corneous-tipped tooth, presumed molar

process with usually corneous-tipped tooth at lower angle; palp

three- segmented, distal segment with row of short marginal

setae. Maxillule (Fig. le) with internal lobe of endopod not

produced, but provided with 1 stiff bristle, external lobe some-

what produced, not recurved, with 3 terminal setae. First

maxilliped (Fig. Ig) with setae of external margin of

exopod only on proximal half. Third maxilliped (Fig. Ih)

with prominent tooth at ventrodistal angle of coxa; basis with

2 or 3 spine-like teeth on ventral margin; ischium with well

developed crista dentata, usually 1 or occasionally 2 widely

separated accessory teeth; merus and carpus each with

dorsodistal spine.

Right cheliped (Figs 3a, b) long and slender. Dactyl 0.50 to

nearly equal to length of palm; dorsomesial margin not delim-

ited, rounded mesial face unarmed or with numerous minute

spinules in proximal half; dorsal surface often with scattered

small pits and individual or sparse tufts of short to moderately

long setae; ventral surface with scattered tufts of longer setae;

cutting edge usually with 2 or 3 large calcareous teeth, ter-

minating in small corneous claw and slightly overlapped by

fixed finger, occasionally crossed with fixed finger in larger

specimens. Palm 0.70-0.95 length of carpus; armature of dor-

somesial margin varying from ill-defined and irregular single

or double row of small spinules or tubercles over entire length

in small specimens to row of prominent spines in proximal

0.80-90 of margin and small spines or spinules in distal

0.10-0.20 in large individuals; dorsal surface with scattered

very small tubercles or spinules, particularly in mesiodistal

0.35, dorsal midline with longitudinal row of 6-13 tiny to

prominent spines, not extending to junction of dactyl and fixed

finger; dorsolateral margin not delimited except faintly in some

small specimens, but armed with numerous very small and tiny

spinules, tubercles or granules, extending to distal 0-25-0.50

of fixed finger; mesial, lateral and ventral surfaces with tiny

tubercles or spinules, sometimes forming short, transverse

rows; surfaces all with scattered sparse tufts of short to moder-

ately long setae, most numerous ventrally; cutting edge of fixed

finger with 2 or 3 prominent calcareous teeth, calcareous ridge

or sometimes few to several small calcareous teeth distally.

Carpus slightly shorter to slightly longer than merus; dorsome-

sial margin with row of small to moderately prominent spines,

dorsodistal margin usually with 1 small to large spine mesially

and several very small spines laterally, occasionally only single

small spine in small specimens; dorsal midline with row of

small spines or spinules becoming row of prominent spines in

large individuals, dorsolateral surface sloping, dorsolateral

margin armed with weakly defined row of tiny spinules or

tubercles in small specimens, increasing in size and number

with increased body size; mesial and lateral faces each with

scattered to moderately dense covering of very small tubercles

or spinules, most numerous in small individuals, distal margins

each sometimes with several spines or spinulose tubercles; ven-

tromesial margin often tuberculate, ventral surface with scat-

tered tubercles; ventrolateral margin usually not delimited, but

with 1 or 2 spines at ventrolateral distal angle. Merus subtri-

angular; dorsal surface with row of short transverse ridges and

sparse tufts of setae; mesial face with few sparse tufts of setae,

ventromesial margin with row of conical, subacute spines, ven-

tral surface often with scattered spines; ventrolateral margin not

distinctly delimited but with short transverse rows of small

tuberculate spines extending onto ventral surface. Ischium with

row of small spines or spinules on ventromesial margin, ventral

surface with few low protuberances or small spines and sparse

setae. Coxa with spine on distal margin ventromesially and

additional spine ventrolaterally.

Left cheliped (Figs 3c-e) often as long as or slightly longer

than right; dactyl approximately 0.50 longer than palm; dorsal

A new hermit crab from seamounts off Tasmania

233

surface slightly elevated proximally, dorsomesial margin not

delimited, but with transverse tuberculate or minutely spinulose

ridges and sparse setae, few spinules on dorsal surface in prox-

imal half, dorsal surface also with scattered stiff setae; cutting

edge with row of small corneous spines, rounded tip of dactyl

slightly overlapped by corneous claw of fixed finger (Fig. 2c)

in small specimens (e.g. 2.6 mm), just begihhing to show

overlap by fixed finger (e.g. ovigerous ? 3.6 mm), or com-

pletely overlapped by hooked end of fixed finger (Figs 2d, e) in

large specimens (e.g. 3.5 mm, ? 5.0 mm); inner surface of

dactyl concave, small corneous claw laterally positioned at tip

of dactyl. Palm 0.65-0.80 length of carpus; dorsomesial margin

variable: not delimited in small specimens, but rounded surface

frequently armed with numerous tiny spinulose tubercles or

small spines often increasing in size proximally and tufts of

setae; large specimens with well delimited margin armed with

row of small spines not quite extending to distal margin, 1 or 2

prominent spines at proximal angle; dorsal surface elevated in

midline and armed with row of 3-11 minute to moderately

prominent tuberculate spines, not reaching to articulation of

dactyl, but often continued as irregularly-set very small spin-

ules in large individuals; dorsomesial surface unarmed in small

specimens, but with increasing number of small or very small

spinules with increased size, primarily in distal half; dorsolat-

eral surface sloping and armed with numerous minute to small

spinules and/or tubercles, continued onto proximal half of fixed

finger, dorsolateral margin varying from well delimited by row

of small spines to not delimited, but rounded surface with

numerous tiny to small spinules or spinulose tubercles also con-

tinued onto fixed finger but not extending to tip, dorsolateral

proximal angle with blunt tubercle or spine; ventral surface

with small spinules or spinulose tubercles laterally, remainder

of ventral surface with tufts of sparse setae; distal portion of

fixed finger curved and slightly overlapping dactyl in small

specimens, but becoming drawn out into hook-like tip with

increased size, terminating in small corneous claw; inner sur-

face of fixed finger oblique in small individuals to prominently

concave, particularly distally in larger specimens. Carpus

slightly shorter to approximately equal to length of merus; dor-

sodistal margin with 1 to several small spines; dorsomesial

margin with irregular row of small to moderately large spines,

dorsal surface often with few small spines, at least distally and

tufts of sparse setae, irregular row of small to moderately large

spines beginning proximally at dorsolateral angle but becoming

almost median distally in larger specimens because of sloping

dorsolateral surface; in smaller specimens lateral face tending

to be more perpendicular, surface armed with numerous tiny to

small spinules or tubercles, ventrolateral margin delimited by

row of small spines or simply rounded with surface armature

continued onto ventral surface laterally; mesial face with scat-

tered tufts of sparse setae, distomesial margin and mesial sur-

face ventrally spinulose or tuberculate; remainder of ventral

surface with few low protuberances and tufts of sparse setae.

Merus subtriangular; dorsal surface with short transverse rows

of sparse setae; ventromesial margin with row of irregularly-

sized spines; ventrolateral margin with row of somewhat small-

er spines, lateral face ventrally with short transverse rows of

very small tubercles or granules accompanied by tufts of setae;

ventral surface with few small spines, particularly laterally, and

scattered tufts of sparse setae. Ischium with row of small spines

on ventromesial margin, 1 small spinule on ventrolateral mar-

gin distally, ventral surface occasionally with few scattered

small spines. Coxa with spine on ventromesial distal margin

and additional spine on distal margin ventrolaterally.

Ambulatory legs (Fig. 3f) similar from left to right, usually

only slightly overreaching tips of chelipeds, if at all. Dactyls

0.10-0.30 longer than propodi; dorsal surfaces each with row

of tufts of moderately short sparse setae, interspersed with cor-

neous spinules in distal 0.25-0.45; lateral faces generally flat-

tened, each with faint longitudinal sulcus flanked dorsally and

ventrally by row of sparse setae; mesial faces each with weak

longitudinal sulcus and row of corneous spinules dorsally; ven-

tral margins each with row of 15-23 corneous spines; terminat-

ing in sharp corneous claw. Propodi 0.50-0.70 longer than

carpi; dorsal surfaces each with low protuberances and numer-

ous tufts of sparse setae; lateral faces each usually with row of

tufts of sparse setae dorsally; ventral surfaces each with few

tufts of sparse setae, 1 or 2 corneous spines on each ventro-

distal margin, at least on second. Carpi each with small dor-

sodistal spine; dorsal surfaces with few low, occasionally

weakly spinulose, protuberances and tufts of sparse setae;

lateral faces each with row of sparse setae dorsally; ventral sur-

faces each with 2 or 3 tufts of setae. Meri each with transverse

ridges and sparse setae dorsally; ventral margins of second

pereopods each with irregular row of small spines and tufts of

sparse setae, lateral faces often with few to numerous spinules

or granules in lower half; third with ventral margins usually

unarmed, rarely with few minute spinules or granules, but with

scattered tufts of setae. Ischia each with dorsal and ventral tufts

of setae. Anterior lobe of sternite of third pereopods (Fig. 2b)

subrectangular, anterior margin with 1 or 2 pairs of small sub-

acute or acute spines.

Telson (Fig. 2e) with posterior lobes symmetrical or only

weakly asymmetrical; terminal margins somewhat oblique,

each armed with 1 or 2 to several spines.

Colour (in preservative). Cephalothorax and appendages all

generally orange to reddish-orange.

Reproduction. Slightly more than half of the females col-

lected were ovigerous, but few if any of the eggs were near to

hatching (eyed). The eggs were relatively small (diameter

0.75-0.98 mm) and quite numerous, although precise counts

were not possible because of obvious loss during preservation

and transport.

Variation. Variation in shape and armature of the chelipeds is

appreciable in B. macrocolus as is indicated in the description,

and this appears to be a function of size rather than sexual

dimorphism. Small specimens of both sexes do not have the

prominent spines on the dorsomesial margin of the right chelae

(Fig. 3a) or on the dorsomesial margins of the right and left

carpi seen in large specimens. Similarly, the median row of

spines on the right chela becomes increasingly more

pronounced with increasing animal size. In contrast, the row of

spines often delimiting the dorsolateral margin of the chela of

the right in small individuals becomes indistinguishable in

larger specimens. The very unusual development of the dactyl

234

P. A. McLaughlin

and fixed finger of the left cheliped may be an adaptation to its

distinctive environment; however, it is unquestionably correlat-

ed with animal growth. In the smallest specimen (62.6 mm) the

corneous claw of the fixed finger overlaps the rounded tip of

the dactyl (Fig. 3c) but the terminal portion of the fixed finger

has not yet developed into a hook. The ventral surfaces of the

dactyl and fixed finger, while sloping inward in this small male,

do not exhibit the prominent concavities seen in larger individ-

uals (Fig. 3e). Although these morphological variations are

seen in both sexes, there appears to be a developmental “lag” in

females. In males and females of similar size, spination on the

dorsomesial margins of chela and carpi tends to be more

prominent in males, whereas the loss of marginal distinction

dorsolaterally on the chela occurs more rapidly.

Distribution and habitat. Seamounts U, JI, Kl, Al, 38 and

Sister 1 off south-eastern coast of Tasmania; 1083-1300 m;

typically occupying small gastropod shells; found in associa-

tion with primarily dead colonies (Gowlett-Holmes, 1998: 47)

of stony coral, Solenosmilia variabilis.

Etymology. From Greek makros, meaning long, and kolon

meaning leg, and indicative of the long-leggedness of this

hermit crab.

Remarks. Because size-related diagnostic characters are not

apparent in small specimens, individuals of shield lengths <2.8

mm, particularly females, Goreopagurus poorei Lemaitre and

McLaughlin, 2003 look superficially like small B. macrocolus.

Consequently, specimens reported by Koslow and Gowlett-

Holmes (1998: 21) from “Andys” and “Main Pedra” seamounts

and Poore et al. (1998: 77) from stations 03, 06, and 56 were

incorrectly referred to B. macrocolus. Two lots, one from

“Andys” (stn 56) and one from “Main Pedra” (stn 03), are

G. poorei. The single female (4.2 mm) from “Main Pedra”

seamount (stn 06) belongs to Pagurodes inarmatus Henderson,

1888, which shares with the aforementioned species distinctly

shortened ocular peduncles.

The majority of specimens of B. macrocolus had been

removed from their shells shortly after capture but a few exam-

ined still occupied shells. These shells were worn and frequent-

ly damaged but the most notable observation was that the shells

rarely even completely covered the abdomens of the crabs.

Correspondingly, there appeared to be reduction in overall

abdomen length with increased animal size. The need for only

minimal abdominal protection is undoubtedly correlated with

the appreciable calcification of the cephalothorax of B. macro-

colus as it is in species of Labidochirus Benedict, 1892,

Porcellanopagurus, and Solitariopagurus. Whether a similar

correlation exists between abdomen length and shell size

cannot be adequately determined from the limited material.

Discussion

Certain morphological characters shared by Bythiopagurus and

Icelopagurus McLaughlin, 1997 might suggest a relationship

between the two. Both monotypic genera are defined as having

broad, well calcified anterior carapaces, very short ocular

peduncles, long antennular and antennal peduncles and elon-

gate chelipeds and ambulatory legs. However, that the similar-

ities are superficial and not phylogenetic is clearly apparent

when gill number and structure, mandibular and maxillulary

development, male and female secondary sexual characters and

telsonal structure are considered. Bythiopagurus has 13 pairs of

distally quadriserial gills, whereas Icelopagurus has 11 biserial

pau's. The mandibles, or at least the left, has a denticulate upper

margin in Bythiopagurus, while this margin is smooth in

Icelopagurus. The external lobe of the endopod of the new

genus is produced but it is rudimentary or vestigial in

Icelopagurus. Papillae or very short male sexual tubes usually

are produced from both gonopores in Bythiopagurus but the

right sexual tube in Icelopagurus is of moderate length (> 2

coxal lengths) and directed toward the exterior. Females of the

new genus have pleopod 1 paired and modified; no first pleo-

pod development occurs in females of Icelopagurus. The very

distinctive telson, which is armed on the terminal margin of

each posterior lobe with several long corneous spines sets

Icelopagurus apart from all other genera.

Papillae or paired, very short sexual tubes, paired and mod-

ified female pleopod 1, quadriserial gill structure, development

of the external lobe of the endopod of the maxillule, short stout

ocular peduncles, and subequal chelipeds are characters that

Bythiopagurus shares With Michelopagurus McLaughlin, 1997,

and these characters certainly might suggest more than a super-

ficial relationship. That the gills are only distally divided in the

new genus and deeply divided in the latter genus is of little sig-

nificance. McLaughlin and de Saint Laurent (1998) found that

gill lamellae in their genus, Propagurus, varied from biserial to

distally quadriserial within a single gill of P. deprofundis.

Lemaitre (2003b) has shown that in species of Sympagurus

Smith, 1883 considerable variation occurs in the extent of

lamellar division, both within and among species. More impor-

tant is the fact that there are 13 pairs of gills in Bythiopagurus,

but only 11 in Michelopagurus . Other characters also indicate

that the observed similarities most probably are convergent.

Perhaps most significant are the major differences in

cephalothoracic calcification among species of the two genera,

the dentition on the mandible(s) in Bythiopagurus, and the lack

of fusion between the basis and ischium of the third maxilliped

in the latter genus.

De Saint Laurent-Dechance (1966) considered those few

genera of Paguridae that have 13 pairs of gills to be the most

primitive genera. At the time of her report, only five genera,

Munidopagurus A. Milne-Edwards and Bouvier, 1893,

Xylopagurus A. Milne-Edwards, 1880, Pylopaguropsis Alcock,

1905, Tomopaguropsis Alcock, 1905, and Tomopaguroides

Balss, 1912, were included in her Pylopaguropsis-gvonp

although she was aware of additional undescribed genera. In

addition to gill number, this group of genera was characterised

by the presence of an accessory tooth on the crista dentata of

maxilliped 3, by paired first and/or second pleopods in males or

paired pleopod 1 in females, but with males lacking sexual

tubes.

Since de Saint Laurent-Dechance’s (1966) discussion of

relationships among genera of Paguridae, an additional 41 gen-

era have been described, of which only five can be included in

her Pylopaguropsis-gyoup-. Lithopagurus Provenzano, 1968,

Bathypaguropsis McLaughlin, 1994, Propagurus, Chano-

A new hermit crab from seamounts off Tasmania

235

pagurus Lemaitre, 2003a, and Bythiopagurus. All have 13 pairs

of gills but pleurobranchs of the fifth and sixth thoracic somites

(cephalothoracic somites XI and XII, McLaughlin and

Lemaitre, 2001) are reduced or rudimentary in Chanopagurus

and Propagurus. De Saint Laurent-Dechance (1966; 259) was

of the impression that all genera in the Pylopaguropsis-group

had quadriserial gills but Asakura (2000) has demonstrated that

the gills in species of Pylopaguropsis are really biserial. The

gills in Lithopagurus and Munidopaguriis are also reported to

be biserial (McLaughlin, 2003). Lemaitre (1995) had described

the gills of Xylopagurus as biserial whereas McLaughlin (2003)

indicated that both bi- and quadriserial gills occurred in species

of that genus. Lemaitre (pers. comm.) has now found that gills

in species of Xylopagurus vary from very weakly divided

distally (practically biserial) to distally divided (quadriserial).

With the exception of Tomopaguroides where the crista

dentata has not been described, all have an accessory tooth;

however, there may be one or two teeth in Bythiopagurus. The

mandible has been described as having a denticulate upper

margin only in Bythiopagurus and Pylopaguropsis but in the

former genus, the incisor process usually is provided with small

corneous-tipped teeth. A truly corneous-toothed incisor process

has been described only in the Pylojacquesidae McLaughlin

and Lemaitre, 2001 but even calcified individual teeth are rare

in Paguroidea.

Males of Xylopagurus have both pleopods 1 and 2 paired,

whereas only pleopod 2 is paired in Lithopagurus and

Tomopaguroides. Males of Chanopagurus are unknown, but in

none of the remaining six genera are either pleopods 1 or 2

paired, although unpaired pleopods may or may not be present.

Contrary to de Saint Laurent-Dechance ’s (1966) diagnosis of

the Pylopaguropsis-group, males of Bythiopagurus do have

extruded papillae or very short sexual tubes. As noted by

Lemaitre and McLaughlin (2003), whether the presence of very

short sexual tubes or simple papillae reflects an early stage in

evolutionary sexual tube development or is a function of

sexual activity in species lacking sexual tube development is

not known. Paired female gonopores are characteristic of the

Pylopaguropsis-group but females of Chanopagurus have a

single left gonopore. Paired pleopod 1 occurs in females of

several genera, but not in Bathypaguropsis, Lithopagurus,

Propagurus, or Tomopaguropsis. Females of Tomopaguroides

are unknown.

Clearly, the presence of 13 pairs of gills is the single unify-

ing character of the Pylopaguropsis-group, as in virtually all

other morphological attributes there is extreme diversity.

However, for the four genera of the group in which some larval

information is available, Provenzano (1971) commented that

the several unusual features found in the zoeas of

Pylopaguropsis atlantica Wass, 1963, Lithopagurus yucatani-

cus Provenzano, 1968, Munidopagurus macrocheles (A. Milne-

Edwards, 1880), and Xylopagurus rectus A. Milne-Edwards,

1880 could not be without phylogenetic significance.

McLaughlin and de Saint Laurent (1998) noted that species

of Propagurus exhibited morphological variations that sug-

gested this genus was undergoing evolutionary transformations

from those of the typical Pylopaguropsis-group genera to those

seen in Pagurus-liko genera. In addition to the overall develop-

ment of gill lamellae that varied from deeply quadriserial to

only distally divided, these authors pointed to reduction in the

pleurobranch of the fifth thoracic somite and to the develop-

ment of the external lobe of the maxillulary endopod, which

varied from well developed to rudimentary. If the other mor-

phological variations demonstrated among genera of the

Pylopaguropsis-group are considered in a similar context, it

might be possible to develop a model of evolutionary change

using this small, and presumably monophyletic, potentially

primitive group of pagurids that would be applicable to the

larger conglomerate of taxa with 11 or fewer paks of gills.

Acknowledgements

The author is particularly indebted to Dr G. C. B. Poore,

Museum Victoria, for making the specimens available for

study, and to Karen Gowlett-Holmes, CSIRO, for providing a

copy of the Koslow and Gowlett-Holmes final report on the

seamount fauna. This is a scientific contribution from the

Shannon Point Marine Center, Western Washington University.

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Memoirs of Museum Victoria 60(2): 237-242 (2003)

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

http://www.museum.vic.gov.au/memoirs

Gnathophyllum taylori^ a new species of caridean shrimp from south-eastern

Australia (Crustacea: Decapoda: Gnathophyllidae)

Shane T. Ahyong

Department of Marine Invertebrates, Australian Museum, 6 College St, Sydney, NSW 2010, Australia (shanea®

austmus.gov.au)

Abstract Ahyong, S. T. 2003. Gnathophyllum taylori, a new species of caridean shrimp from south-eastern Australia (Crustacea:

Decapoda: Gnathophyllidae). Memoirs of Museum Victoria 60(2): Til-lAl.

Anew species of caridean shrimp, Gnathophyllum taylori sp.nov., is described from south-eastern Australia. The new

species most closely resembles two Atlantic species, G. modestum Hay, 1917, and G. elegans Risso, 1816, but is

readily distinguished by its banded rather than spotted body colour pattern. Gnathophyllum taylori is the third Indo-West

Pacific species to be recognised. It differs from G. americanum, with which it can be sympatric, by having fewer and

narrower dark bands on the body as well as by the rostral dentition, morphology of the orbit and dactyli of the walking

legs. The identity of G. fasciolatum Stimpson, 1860, a junior synonym of G. americanum, is fixed by neotype selection.

Keywords Cmstacea, Decapoda, Caridea, Gnathophyllum taylori, new species

Introduction

The most recent treatment of the caridean genus

Gnathophyllum Latreille, 1819, recognised eight species

worldwide with two in the Indo-West Pacific - G. americanum

Guerin-Meneville, 1855, and G. precipuum Titgen, 1989 (see

Chace and Bruce, 1993). Gnathophyllum precipuum is known

only from Hawai’i. Gnathophyllum americanum is presently

recognised as having a circumtropical distribution and in the

Indo-Pacific is known from the Red Sea and South Africa to

Australia, Japan and French Polynesia. Off eastern Australia,

G. americanum is known as far south as Port Jackson, Sydney,

New South Wales. To date, specimens of Gnathophyllum from

south-eastern Australia have not been studied in any detail but

a popular account recognised two colour forms of G. ameri-

canum from eastern Australia (Healy and Yaldwyn, 1970). One

form bears numerous narrow pale bands on a dark background,

and the other form bears relatively few, narrow dark bands on

a pale background. The first form is G. americanum sensu

stricto, whereas the second form represents a new species,

described below.

Measurements given for each specimen refer to orbital cara-

pace length followed by total carapace length, given in mil-

limetres (mm). Specimens are deposited in the collections of

the Australian Museum, Sydney (AM).

Gnathophyllum taylori sp. nov.

Figures 1-2

Gnathophyllum americanum. — Healy and Yaldwyn, 1970: 46, fig.

21. — Debelius and Baensch, 1994: 506 [not G. americanum Guerin-

Meneville, 1855]

Gnathophyllum sp. — Debelius, 1999: 196 [colour figure].

Material examined. Holotype: AM P64829, ovigerous female (3 .9/5. 6

mm), S side of Long Bay, Malabar, New South Wales, 33°58'S,

151°15"E, 4 m, under Centrostephanus rogersi, R. Taylor, 3 Apr 2001.

Paratypes: AM P306, 2 ovigerous females (5. 0/7. 0-5. 7/7.7 mm).

Port Jackson; AM P1420, 3 females (4.2/5. 9 mm, 5.4/7. 5 mm, oviger-

ous 6.9Z9.3 mm). Port Stephens, 32°42'S, 152°06T; AM P12423, 1

ovigerous female (4.0/5.7 mm). Long Reef, Collaroy, 33°44'S,

151°19"E, west end of reef among weeds and boulders, low tide, E.

Pope, 15 Eeb 1953; AM P38007, 1 ovigerous female (3. 8/5.5 mm), SW

Elizabeth Reef, Tasman Sea, 29°57.7"S, 159°02.8"E, outer slope,

among coral heads and overhangs, A. Gill and S. Reader, 11 Dec 1987.

Comparative material of Gnathophyllum americanum Guerin-

Meneville, 1855. AM P62995, ovigerous female (4. 1/5.8 mm). Rose

Bay, Port Jackson, NSW, seine net, D. Hoese 19 Jan 1976 (neotype of

G. fasciolatum Stimpson); AM P20302, 1 male (4.0/5. 4 mm), 1 female

(4. 2/5. 7 mm). Gun Island, Houtman Abrolhos Group, Western

Australia, 1.8 m, under rocks near reef edge, N. Coleman, 9 May 1972;

AM P18558, 1 male (4.2/5.4 mm), 1 ovigerous female (4.3/5.4 nun).

Heron Island, Queensland, Aug 1960.

238

Shane T. Ahyong

Figure 1. Gnathophyllum taylori sp. nov., holotype female cl. 5.6 mm. A, right lateral. B, anterior carapace, right lateral. C, eye, right lateral. D,

antennule, right dorsal. E, basal segment of antennule, right ventral. F, antennal peduncle, right dorsal. G, antennal peduncle, right ventral. H, pos-

terior margin of third abdominal somite, dorsal. I, tailfan, dorsal. Scale A = 2 mm, B-I = 1 mm.

Diagnosis. Rostrum with 5 or 6 dorsal teeth and minute sub- reaching base of second segment. Pereopods 3-5 with dactylus

distal ventral denticle; proximal 1 or 2 teeth behind level of biunguiculate, ventral process shorter than dorsal, broad, sub-

posterior orbital margin. Orbit without narrow posterodorsal trapezoid, not evenly tapering. Body pale dull yellow with

sinus. Telson posterior lateral spines arising well anterior to narrow, widely spaced dark bands; pereopods without banding

outer pair of posterior spines; telson posterior margin with but with white speckling; sixth abdominal tergite and tail-fan

blunt median tubercle. Cornea of eye with narrow, rounded transparent,

point. Antennular peduncle with apex of stylocerite not

A new shrimp of the genus Gnathophyllum

239

Figure 2. Gnathophyllum taylori sp. nov., holotype female cl. 5.6 mm. A, pereopod 1, right lateral. B, pereopod 2, left lateral. C, pereopod 3, right

lateral. D, dactylus of pereopod 3, right. E, mandible, right posterior. F, mandible, distal posterior. G, mandible, distal anterior. H, maxilla 1, right

posterior. I, maxilla 2, right posterior. J, maxilliped 1, left anterior. K, maxilliped 2, right posterior. L, maxilliped 3, right posterior. Scale A-C,

E-L = 1 mm, D = 0.5 nun.

Description. Rostrum not overreaching basal-antennular-

segment, with 5 or 6 dorsal teeth and minute subdistal ventral

denticle; proximal 1 or 2 teeth behind level of posterior orbital

margin. Orbit without narrow posterodorsal sinus. Antennal

spine well developed, arising slightly submarginally. Inferior

orbital angle with blunt anterior projection at level of antennal

spine, extending anteriorly to or slightly beyond apex of

antennal spine. Pterygostomian angle of carapace rounded and

produced anteriorly beyond antennal spine.

Abdomen smooth, pleura 1-4 rounded; pleuron 5 postero-

laterally angular; medial margin of somite 3 rounded, slightly

overhanging somite 2. Sixth abdominal somite three-quarters

length of telson; height about three-quarters length. Telson

twice as long as wide; with 2 pairs of lateral spines and 3 pairs

240

Shane T. Ahyong

of spines on posterior margin; anterior lateral spines arising

well behind midlength of telson; posterior lateral spines arising

well anterior to outer pair of posterior spines; intermediate

posterior spines more than twice length of lateral and mesial

posterior spines; mesial posterior spines somewhat soft; telson

posterior margin with blunt median tubercle.

Cornea of eye pigmented; with blunt apical point.

Antennular peduncle with broad basal segment, apex of

stylocerite not reaching base of second segment, with stout dis-

tolateral spine overreaching second segment, ventromesial

margin with spine at midlength; dorsolateral flagellum totalling

19-28 articles, fused basal portion of 7-13 short articles.

Antennal scale overreaching antennular peduncle; about

twice as long as wide; lateral margin straight, distal spine about

as long as wide, not reaching to apex of blade.

Mandible small, unequal, without incisor process. Maxillule

with upper endite expanded with double row of robust setae

medially, setose laterally; lower endite small, blunt, distally

setose. Maxilla without endites; scaphognathite well devel-

oped, marginally setose; palp elongate, distally tapered, non-

setose.

First maxilliped with well developed caridean lobe, exopod

slender, distally setose; endite large, setose. Second maxilliped

exopod slender, distally setose; endopod with broad basal

articles and elongate, curved distal segment, setose on outer

margin. Third maxilliped well-developed; endopod with oper-

culiform ischiomerus, about twice as long as wide, evenly

setose medially, sparsely setose laterally; exopod long, slender,

setose distally and proximolaterally; ischiomerus completely

fused with basis.

Pereopods symmetrical left to right. Pereopod 1 distinctly

oven'eaching antennal scale by length of chela and three-

quarters length of carpus; occlusal margins of dactylus and

pollex smooth; dactylus almost as long as palm. Pereopod 2

oven'eaching antennal scale by length of chela and carpus;

occlusal margins of dactylus and pollex smooth on distal three-

quarters sinuous on proximal third; dactylus more than Vh.

times palm length. Pereopods 3-5 similar, merus 6-7 times as

long as wide. Distal half of propodus with 7-10 movable spines

along lower margin. Dactylus biunguiculate, dorsal process

slender, evenly tapering to sharp point; ventral process shorter

than dorsal, broad, trianguloid to subtrapezoid, not evenly

tapering.

Uropodal protopod unarmed dorsally or ventrally; exopod

with stout laterodistal spine flanked by smaller movable spine

mesially; endopod ovate, unarmed.

Colour in life. Body pale dull translucent yellow with

narrow, widely spaced dark purplish bands; 4-5 bands on cara-

pace, 6-8 bands on abdomen; margins of dark bands slightly

brighter than base colour of body. Ground colour of

cephalothorax sometimes appearing deeper yellow than that of

abdomen owing to colour of viscera. Pereopods without band-

ing but with white or yellowish speckling. Sixth abdominal

somite and tail-fan transparent. Th ir d maxilliped with dark pur-

ple “O” on ischiomerus.

Etymology. Named for Richard Taylor, Leigh Marine

Laboratory, New Zealand (formerly of University of New

South Wales, Australia), who collected the holotype.

Distribution. NSW, Elizabeth Reef south to Long Bay.

Remarks. Gnathophyllum taylori sp.nov. is the third species of

the genus to be recognised from the Indo-West Pacific.

Gnathophyllum precipuurn, from Hawai’i is readily distin-

guished from G. taylori by having two instead of five or six

dorsal rostral teeth, and by lacking the dark banding on the

body. Instead, G. precipuurn has a basic colour pattern of

“irregular reddish-brown spots on light yellow background”

(Titgen, 1989: 206). The only other species of Gnathophyllum

known from the Indo-West Pacific is G. americanum. As in

G. taylori, G. americanum bears dark banding on the body, but

of a different pattern. Morphologically, however, G. taylori

most closely approaches the Atlantic species, G. elegans

(Risso, 1816) and G. modestum Hay, 1917, in having the

posterior-most of the dorsal rostral spines situated behind the

posterior orbital margin, in having the posterior pair of lateral

telson spines distinctly anterior to the posterior pair of spines,

and in having the apex of the stylocerite not reaching the base

of the second antennular segment. The new species is readily

distinguished from G. modestum and G. elegans by the consid-

erably shorter laterodistal spine on the antennal scale in which

it is about as long as wide instead of about twice as long as

wide. Gnathophyllum taylori also differs from G. modestum

and G. elegans by the banded rather than uniform or spotted

colour pattern of the body (Manning, 1963).

Both G. taylori and G. americanum bear transverse banding

on the body, but the nature of the banding is the most obvious

distinguishing feature. In G. americanum the dark bands are

broader and about twice as numerous as in G. taylori. The

broader and more numerous dark bands in G. americanum give

the body a dark overall appearance whereas in G. taylori, the

background colour is dull pale yellow with narrow, widely

spaced dark purplish bands giving the body a light overall

appearance. In addition, the pereopods are banded in G. amer-

icanum, but not banded in G. taylori. The ground colour of the

body of G. taylori may vary somewhat in the density of pale

yellow pigmentation. Where the base pigmentation is less

dense, the colour of the internal organs is visible through the

carapace giving the appearance of a deeper yellow cast to the

cephalothorax as figured by Debelius and Baensch (1994) and

Debelius (1999). Conversely, the base yellow pigmentation in

the holotype was denser, giving a more uniformly yellow cast

to the whole body.

Aside from differences in colour pattern, G. taylori differs

morphologically from G. americanum in having the proximal

tooth of the dorsal rostral series on the carapace arising behind

instead of anterior to the posterior orbital margin, in having a

higher range of dorsal rostral teeth (5 or 6 instead of 3-6), in

lacking a short narrow sinus in the posterodorsal margin of the

orbit, and in having a broad rather than slender ventral tooth on

the dactyli of pereopods 3-5.

A character that might prove useful in distinguishing species

of Gnathophyllum is the fusion of the ischiomerus with the

basis of the third maxilliped. In G. taylori and apparently in

G. ascensione Manning and Chace, 1991 (see their fig. 5P), the

ischiomerus and basis of the third maxilliped are fused.

Conversely, the ischiomerus and basis appear to be clearly

demarcated in G. americanum from the Canary Islands and

A new shrimp of the genus Gnathophyllum

241

Micronesia, and G. precipuum as figured by Kubo (1940: fig.

8F), Holthuis (1949b: fig. 6H) and Titgen (1989: fig. 21)

respectively. The accuracy of these figures requires verifi-

cation, but if accurate, indicate that the degree of fusion of the

ischiomerus and basis of the third maxilliped has diagnostic

value for species of Gnathophyllum. It is noteworthy then, that

all Australian specimens of G. americanum examined resemble

G. taylori in the fusion of the ischiomerus and basis of the third

maxilliped, unlike those figured by Holthuis (1949b) and Kubo

(1940). The possible heterogeneity in G. americanum also indi-

cates the potential validity of one of its synonyms,

Gnathophyllum fasciolatum Stimpson, 1860.

Gnathophyllum fasciolatum Stimpson, 1860, was described

from Port Jackson, Sydney, which is well within the range of

G. taylori. Stimpson’s (1860) species has long been established

as a junior synonym of G. americanum (e.g. Rathbun, 1901;

Armstrong, 1940; Manning, 1963; Chace and Bruce, 1993).

Gnathophyllum americanum, first described from Cuba, is

presently regarded as a near-cosmopolitan species but could

prove to be a species complex (this study, Davie, 2002). The

holotype of G. fasciolatum, however, is lost and Stimpson’s

(1860) brief account could apply to either species of

Gnathophyllum from the Sydney region (let alone almost any

Other species of the genus were colour pattern not considered).

Gnathophyllum fasciolatum, an older name, threatens the

nomenclatural stability of G. taylori. Therefore, a neotype des-

ignation for G. fasciolatum is justified. A specimen of G. amer-

icanum from Rose Bay, Port Jackson, is herein designated as

the neotype of G. fasciolatum to fix the identity of the species

(Fig. 3). The neotype is an ovigerous female, cl. 5.8 mm, regis-

tered as AM P62995. Revision of G. americanum is beyond the

scope of this study but should G. fasciolatum be removed from

synonymy, its identity is now stabilised through the present

neotype selection. Other synonyms of G. americanum are not

Figure 3. Gnathophyllum americanum, (neotype of G. fasciolatum),

female cl. 5.8 mm. A, anterior carapace, right lateral. B, antennule,

right dorsal. C, dactylus of pereopod 3, right. D, dactylus of pereopod

5, right. Scale A, B = 1 mm, C, D = 0.5 mm.

identifiable with G. taylori. In both G. zebra Richters, 1880

(described from Mauritius) and G. miniscularium Armstrong,

1940 (described from Bermuda), the posterior-most dorsal ros-

tral tooth is placed anterior to the orbit instead of behind as in

G. taylori. Gnathophyllum tridens (type locality Rikitea,

French Polynesia; redescribed by Holthuis, 1949a) bears three

dorsal rostral teeth and is within the range of G. americanum

(3-6) but not of G. taylori (5-6). Gnathophyllum pallidum

Ortmann, 1890, also described from French Polynesia, was neither

illustrated nor described in detail, but is well within the known

range of G. americanum, and well outside that of G. taylori.

The holotype of G. taylori was collected from beneath the

echinoid, Centrostephanus rogersi, but other specimens were

collected from under boulders or amongst marine algae.

Gnathophyllum taylori is temperate water species, in contrast

to the primarily tropical G. americanum. All known specimens

of G. taylori are from New South Wales, Australia.

Gnathophyllum americanum is known as far south as Port

Jackson on the east coast and as far south as the Abrolhos

Islands on the west coast. The distribution of G. taylori thus

overlaps that of G. americanum in south-eastern Australia.

Gnathophyllum americanum was figured in colour from near

Grafton and Lord Howe Island, New South Wales by Healy and

Yaldwyn (1970: pi. 20), and Coleman (2002: 50) respectively.

Debelius and Baensch (1994) and Debelius (1999: 196) figured

G. taylori in colour, photographed by R. Kuiter, at Seal Rocks,

New South Wales at 2 m depth.

Acknowledgements

Thanks to Richard Taylor and Peter Schiipp, both formerly

University of New South Wales, for collecting and photograph-

ing the holotype respectively, and thanks to Rudie Kuiter for

information regarding his photograph of G. taylori from Seal

Rocks. Comments from two anonymous reviewers are grate-

fully acknowledged. This study was supported by an Australian

Postdoctoral Fellowship from the Australian Research Council

to STA.

References

Armstrong, J.C. 1940. New species of Caridea from the Bermudas.

American Museum Novitatcs 1906: 1-10.

Chace, F.A., and Bruce, A.J. 1993. The caridean shrimps (Crustacea:

Decapoda) of the Albatross Philippine Expedition 1907-1910, pait

6: Superfamily Palaemonoidea. Smithsonian Contributions to

Zoology 543: 1-152.

Coleman, N. 2002. Lord Howe Island Marine Park: Sea Shore to Sea

Floor. Neville Coleman’s Underwater Geographic: Springwood. 96

pp.

Davie, P.J.F. 2002. Crustacea: Malacostraca: Phyllocarida,

Hoplocarida, Eucarida (Part 1). In; Wells, A. and Houston, W. W. K.

(eds) Zoological Catalogue of Australia. Vol. 19.3 A. CSIRO

Publishing: Melbourne, xii + 551 pp.

Debelius, H. 1999. Crustacea Guide of the World. IKAN -

Unterwasserarchiv, Erankfurt. 321 pp.

Debelius, H., and Baensch, H.A. 1994. Marine Atlas.Mergus: Melle.

1215 pp.

Guerin-Meneville, EE. 1855-1856. Crustaceos. In: Ramon de la

Sagra, Historia fisica politica y natural de la isla de Cuba, Historia

Natural 7: v-xxxii, 8, pis 1-3.

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Hay, W.P. 1917. Preliminary descriptions of five new species of crus-

taceans from the coast of North Carolina. Proceedings of the

Biological Society of Washington 30; 71-73.

Healy, A., and Yaldwyn, J. 1970. Australian crustaceans in colour.

Reed: Sydney. 112 pp.

Holthuis, L.B. 1949a. Note sur le type du Gnathophyllum tridens

Nobili, 1906 (Crustaces, Decapodes). Bulletin du Museum

d’Histoire Naturelle, Paris (2) 21(2); 210-214.

Holthuis, L.B. 1949b. The Caridean Crustacea of the Canary Islands.

Zoologische Mededelingen, Leiden 30; 227-255.

Kubo, T. 1940. On some littoral shrimps collected from Micronesia.

Journal of the Imperial Fisheries Institute, Tokyo 34(1); 77-99.

Latreille, PA. 1819. Salicoques, Carides, Latr. Nouveau Dictionnaire

d’Histoire Naturelle 30: 68-73.

Manning, R.B. 1963. The east American species of Gnathophyllum

(Decapoda, Caridea), with the description of a new species.

Crustaceana 5(1): 47-63.

Nobili, G. 1906. Diagnoses preliminaires de Crustaces, Decapodes et

Isopodes nouveaux recueillis par M. le Dr. G. Seurat aux iles

Touamotou. Bulletin du Museum d’Histoire Naturelle, Paris 12(5):

256-270.

Ortmann, A. 1890. Die Unterordnung Natantia Boas: Die Decapoden-

Krebse des Strassburger Museums, mit besonderer Beriicksichtung

der von Herrn Dr. Dbderlein bei Japan und bei den Liu-Kiu-Inseln

gesammelten und z. Z. im Strassburger Museum aufbewahrten

Formen, I. Zoologische Jahrbiicher. Abtheilung fUr Systematik,

Geographie und Biologie der Tiere 5: 693-750

Rathbun, M.J. 1901. The Brachyura and Macrura of Porto Rico.

Bulletin of the United States Fish Commission 20(2): 3-126.

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Meersfauna der Insel Mauritius und der Seychellen. Berlin.

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Librairie Grecque-Latine-Allemande: Paris 175 pp, 173 pis.

Stimpson, W. 1860. Prodromus Descriptionis Animalium

Evertabratorum, quae in expeditione ad Oceanum Pacificum

septentrionalem a Republica Federata missa, Cadwaladaro

Ringgold et Johanne Rogers ducibus, observavit descripsit. Pars

VIII. Crustacea Macmra. Proceedings of the Academy of Natural

Sciences of Philadelphia 1860: 22-47.

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Memoirs of Museum Victoria 60(2): 243-256 (2003)

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

http://www.museum.vic.gov.au/memoirs

Rediagnosis of Palaemon and differentiation of southern Australian species

(Crustacea: Decapoda: Palaemonidae)

Terry M. Walker' and Gary C. B. Poore^

'WATER ECOscience, PO Box 34, New Town, Tas. 7008, Australia, or 41 South St, Bellerive, Tas. 7018, Australia

Eormerly: Zoology Department, University of Tasmania, GPO Box 252-55, Hobart, Tas. 7001, Australia

(twalker@wes.com.au)

"Museum Victoria, PO Box 666E, Melbourne, Vic. 3001, Australia (gpoore@museum.vic.gov.au)

Abstract Walker, T.M., and Poore, G.C.B. 2003. Rediagnosis of Palaemon and differentiation of southern Australian species

(Crustacea: Decapoda: Palaemonidae). Memoirs of Museum Victoria 60(2): 243-256.

Examination of the morphology of palaemonine shrimps from Tasmania and south-eastern Australia, and molecular

analyses by other authors, necessitated an examination of the validity of some characters used for generic differentiation

within the Palaemoninae and a redefinition of the genus Palaemon. A new species, Palaemon dolospina sp. nov., is

described from Margate Beach in south-eastern Tasmania. Palaemon intermedins (Stimpson, 1 860) is redescribed. Both

species vary between localities across southern Australia. Keys are presented for the small estuarine and marine shrimps

of Palaemoninae in southern and south-eastern Australia belonging to genera, Palaemon and Palaemonetes. One species

of Palaemon has been introduced to Australia from overseas.

Key words Crustacea, Caridea, Palaemonidae, Palaemon, new species, Australia

Introduction

An investigation of Tasmanian palaemonids shrimps usually

known as Macrobrachium intermedium from the North West

Bay River Estuary and D’Entrecasteaux Channel in south-

eastern Tasmania revealed not only the presence of two forms

in these environments but also the presence of a similar new

species in the estuary. Both species are common in seagrass

and, in eastern and south-eastern Tasmania, occur sympatric-

ally in the seagrasses Zostera muelleri and Heterozostera

tasmanica. Eurther study showed that the first is widespread in

marine and estuarine waters from Western Australia to

Queensland. The new species has not been located outside

South Australia, Tasmania and Victoria. It is generally found in

estuarine waters or, in Tasmania, in shallow marine waters

where the other is absent.

Neither species complies well morphologically with the

generic diagnoses of Macrobrachium or the alternative,

Palaemon. Analyses using allozymes and molecular infor-

mation from 16S rRNA strongly suggested that M. inter-

medium was more closely related to Palaemon serenus (Heller,

1 862) (Murphy and Austin, 2002) or to Palaemonetes australis

Dakin, 1976 (Boulton and Knott, 1984; Murphy and Austin,

2003) than to other species of Macrobrachium. This paper

re-examines morphological characters used in generic classifi-

cation within the subfamily Palaemoninae and redefines

Palaemon. Both species are described and assigned to

Palaemon, henceforth P. intermedins (Stimpson, 1860) and

P. dolospina sp. nov.

We follow Bruce (1992) in treating the gills of maxilliped 3

as two arthrobranchs (one rudimentary) rather than one pleuro-

branch, said by Holthuis (1993) and earlier workers to define

the subfamily Palaemoninae.

Palaemon is separated from Palaemonetes, Macrobrachium

and other palaemonine genera principally on the basis of the

presence and form of the branchiostegal groove, the presence

or absence of branchiostegal and hepatic spines, the form of the

second pereopods and the presence or absence of a mandibular

palp. Problems with the use of these characters were noted

previously by Eujino and Miyake (1968), Chace (1972),

Bray (1976) and Boulton and Knott (1984). We reviewed these

characters in eight southern Australian species (Table 1).

Palaemon is said to be separated from Macrobrachium by

the presence of a branchiostegal spine and smooth second

pereopods while Macrobrachium has an hepatic spine and

spinulose second pereopods in males of many species. In

Palaemon, adult females are larger than males while males are

usually larger in Macrobrachium.

Most species of Macrobrachium have an hepatic spine set

well back from the carapace edge on or just below the ridge of

the antennal spine (Pig. 1). The branchiostegal groove is short.

Table 1 . Characters of some Australian species of Palaemon, Palaemonetes and Macrobrachium

244

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tiX) P C/I

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Terry M. Walker and Gary C. B. Poore

Palaemon and its southern Australian species

245

Figure 1 . SEMs of carapace spines, a, Macrobrachium msenhergir, b, Palaemon seremis', c, Palaemonetes australis; d, Palaemon intermedins;

e, Palaemon dolospina.

246

Terry M. Walker and Gary C. B. Poore

and runs from the carapace edge to the front of the base of the

spine and stops there. The back of the spine is closed with a

suture line sun'ounding the spine where it makes contact with

the carapace and it is usually flattened dorsoventrally. The

hepatic spine is well removed from the anterior edge of the

carapace (Holthuis, 1950) by a distance greater than its length.

In most species of Palaemon (and Palciemonetes) there is a

branchiostegal groove running back from the anterior edge

of the carapace in a shallow arc near the junction of the

branchiostegite and the thorax (Fig. 1). Its length generally

varies from a quarter to a half of the carapace length. A branch-

iostegal spine is found below this groove, usually on the cara-

pace edge close to the start of the groove. However, the spine

may be found removed ventrally from the groove somewhat

and set back a little from the carapace edge. In Australian

material, the spine is usually rounded rather than flattened and

there is no distinct suture line where it is in contact with the

carapace.

Within Australia, there is a continuum of spine/groove

patterns between those typically found in Palaemon and

Macrobrachium. Palaemon serenus, Palaemon litoreus

(McCulloch, 1909), Palaemon macrodactlyus (Rathbun, 1902)

and Palaemonetes atrinubes Bray, 1976 have a typical branch-

iostegal groove and the spine on the edge of the carapace (Fig.

lb). Palaemonetes australis has the branchiostegal spine set

well back from and not overlapping the carapace edge, with the

groove of the usual form passing over it (Fig. Ic). Palaemon

dolospina has a unique arrangement where a circular spine is

set back from the carapace edge, but with the tip Just over-

lapping it in some individuals (Fig. le). The spine is displaced

dorsally to an almost hepatic position. The branchiostegal

groove runs back into the spine but is then displaced dorsally,

loops over the spine and runs past it in a ventroposterior

direction for a distance equal to about twice the length of the

spine. As the spine is below the branchiostegal groove, it can

considered to be branchiostegal.

Palaemon intermedins possesses what appears to be an

hepatic spine, but it is not set as far back from the carapace

edge as is usual in that genus, and is lower on the carapace,

being nearer the junction of the branchiostegite and thorax

(Fig. Id). The groove runs into the front of the spine but the

latter is open posteriorly, having no suture line where it meets

the carapace at that point. The groove or suture line dorsal to

the spine extends further posteriorly than that ventral to it and

may represent a short extension of the branchiostegal groove

over the spine. The spine is flattened dorsoventrally.

Macrobrachium rosenbergii (De Man, 1879) (Fig. la),

M. lar (Fabricius, 1798) and other Australian species of this

genus have a typical hepatic spine set high up on the carapace

with the branchiostegal groove running up into it and stopping.

The spine is surrounded by a suture line where it joins the cara-

pace, and its tip is well removed from the anterior carapace

edge.

The intermediate nature of the two species can be appreci-

ated by examination of post-larvae and juveniles. Both species

have no branchiostegal groove and a branchiostegal spine on

the edge of the carapace (Fig. 2). Over a series of moults, the

spine migrates away from the carapace edge to its final adult

position as the branchiostegal groove forms. This was also

found to occur in Macrobrachium australe (Guerin-Meneville,

1838), M. latimanus (Von Martens, 1868) and M. lar by

Holthuis (1950). Other species are similar: M. nipponense (De

Haan, 1849) (Kwon and Uno, 1969), M. rosenbergii (see Uno

and Kwon, 1969), M. australiense Holthuis, 1950 (Fielder,

1970), M. niloticum (Roux, 1833) (Williamson, 1972),

M. novaehollandiae (De Man, 1938) (Greenwood et al., 1976),

M. equidens (Dana, 1852) (Ngoc-Ho, 1976) and M. lar (see

Atkinson, 1977).

Thus it appears that the “ hepatic” spine of Macrobrachium

and “branchiostegal” spine of Palaemon and related genera are

homologous. An “hepatic” spine appears to be a branchiostegal

spine that has migrated during post-larval and juvenile devel-

opment. The groove appears to be displaced upwards and

remains developed only between the carapace edge and the

spine. One exception is found in M. palaemonoides Holthuis,

1950, where the spine is in an hepatic position but the groove

passes below the spine (Holthuis, 1950). A consequence of this

realisation is that while adults of Macrobrachium and

Palaemon can be assigned to to their respective genera on this

character, juveniles cannot and other characters must be applied

to distinguish juveniles.

This ontological process appears to be arrested in Palaemon

intermedius and P. dolospina but other characters such as

features of pereopod 2 place them clearly in Palaemon.

Palaemonetes differs from Palaemon solely by the absence

of a mandibular palp. Incidentally, the same character differen-

tiates Pseudopalaemon Sollaud, 1911 from Macrobrachium.

Most specimens of the two Tasmanian species have a three-

articled palp. Boulton and Knott (1984) noted difficulties

identifying the joint between the second and third article in

Palaemon intermedius from Swan River, WA. The palp is

absent in the first post-larval stage and present as a small bump

in the second post-larval stage, developing into its normal form

over the next few moults (Fig. 4m). This developmental change

is widespread in Palaemon, Macrobrachium and Leptocarpus

(e.g. Ngoc-Ho, 1976).

The mandibular palp varies in adults of some species or in

some populations of some species. Chace (1972) reported

specimens of Palaemon debilis Dana, 1852 from Hawaii with

between one and three articles or no mandibular palp

at all. Fujino and Miyake (1968) found the same for five

species of Palaemon from Japan, and the same discrepancy

between two sides of a single individual. Among 20

specimens of Palaemon cf. debilis from Coila Lake, NSW, the

number of palp articles ranged from none to two (65%).

Bray (1976) found Palaemonetes australis Dakin, 1915 from

WA with a mandibular palp of one or two articles or without

a palp.

The doubtful status of Palaemonetes has been noted

previously on grounds of morphology (Chace, 1972; Bray,

1976), morphology and allozymes (Boulton and Knott, 1984)

and mitochondrial genes (Murphy and Austin, 2003). Placing it

in synonymy with Palaemon may not be appropriate since the

form of the mandibular palp is consistent once adult state is

achieved in at least some Australian species, Palaemonetes

atrinubes and the two described below. Nevertheless, it is

Palaemon and its southern Australian species

247

variable in Palaemonetes australis and Palaemon cf. debilis. A

final decision on the synonymy would require a more thorough

examination of species beyond Australia.

The second pereopods of most, though not all, species of

Macrobrachium (and Chryphiops Dana, 1852) are larger and

often spinulose in adult males.This is a useful character for the

generic identification of sexually mature males, but it is of no

use in juveniles or females which resemble members of other

genera such as Palaemon and Palaemonetes.

Pending a major revision of the Palaemoninae, Australian

species may be divided into three groups. The first includes all

species of Macrobrachium sensu stricto, which occupy fresh-

water or estuarine habitats, have markedly sexually dimoiphic

second pereopods, larger males than females and the hepatic

spine/groove arrangement typical of that genus. The second

group is Palaemon serenus, P. litoreus, P macrodactylus (intro-

duced), P. dolospina and P intermedius. These occupy marine

and estuarine habitats, are robust, females are larger than

males, the second pereopods are similar in both sexes, a prox-

imal tuft of setae is present dorsally on the telson and they

always possess a mandibular palp of three articles. The third

group contains Palaemonetes atrinubes, Palaemonetes

australis, Palaemon debilis and possibly another species close

to P debilis. These occupy marine and estuarine habitats, are

less robust than those of the second group, have more slender

pereopods, have larger females than males, the second pere-

opods are similar in both sexes, there is no proximal tuft of

setae on the telson, and the number of articles in the mandib-

ular palp appears to be variable with a tendency to be reduced

or absent.

In spite of the shortcomings of the characters discussed

above, it is possible to redefine Palaemon around the second

and third groups of species and include the following two

species.

Palaemon Weber, 1795

Palaemon Weber, 1795: 94. — Holthuis, 1950: 42—44 (extended

diagnosis). — Holthuis, 1993: 112-114 (synonymy).

Diagnosis. Rostrum well developed, toothed dorsally and ven-

trally, without an elevated basal crest; upper margin bearing

single row of setae between dorsal teeth. Carapace smooth,

bearing distinct branchiostegal spine and groove; branchio-

stegal spine usually situated on carapace edge but may be set

back; groove usually running straight back from carapace edge

in a shallow arc but may rise upwards and over spine before

continuing posteriorly or, in P intermedius, into and over but

not past its posterior edge. Antennal spine generally strong, on

anterior margin of carapace some distance below rounded

lower orbital angle. Mandibular palp usually of 2 or 3 articles,

but may be absent. Eyes distinctly pigmented, cornea well

developed. Anterior margin of antennular peduncle rounded,

anterolateral spine small. First pleopod of male without or with

rudimentary appendix interna on endopod. Propodus of fifth

pereopod with several transverse rows of setae on distal part of

the posterior margin. Telson with 2 slender median setae on

posterior margin. Pleura of fifth abdominal somite usually

ending in small sharp point.

Remarks. The revised diagnosis allows for the variable position

of the branchiostegal spine and accommodates the hepatic

position of this spine seen in some Australian species.

Palaemon intermedius (Stimpson)

Figures Id, 2a-e, 3a, 3b, 4, 5

Leander intermedius Stimpson, 1860: 41. — Haswell, 1882: 195. —

Whitelegge, 1889: 224 (record in Port Jackson). — Thomson, 1893: 51

(record in Tas.). — Stead, 1898: 210. — Sayce, 1902: 155 (record in Port

Phillip Bay).— McCulloch, 1909: 309, pi. 89 figs 13, 14.— Hale,

1927a: 58-59, fig. 53 (record in SA).— Hale, 1927b: 309.— Anderson,

1938: 351 (record in Spencer Gulf). — Mack, 1941: 108 (diet of cor-

morants, Gippsland Lakes). — Thomson, 1946: 59, 67 (record in

WA).— Holthuis, 1950: 20.

Palaemon (Leander) intermedius. — Miers, 1884: 295.

Macrobrachium intermedium. — Holthuis, 1952: 207-209

(redescription). — Poore et al., 1975: 35, 71 (distribution in Port Phillip

Bay). — ^Walker, 1979 (redescription and ecology in Tas.). — Boulton

and Knott, 1984: 769-784 (ecology in WA). — Davie, 2002: 296.

Material examined for redescription. 15 males, 12.7-27.3 mm body

length , 20 females including 7 ovigerous, 12.7-34.5 mm body length,

Margate Beach, mouth of North West Bay River Estuary, Tasmania, T.

Walker, 1975. 4 adult females, D’Enttecasteaux Channel, Middleton,

14 m, Tasmania, T. Walker, Khin Khin U and T. Sward, 11 Apr 1975.

4 ovigerous females, 2 mature males. Western Port, Victoria, Eisheries

and Wildlife Department, Victoria, 29 Nov 1973. 3 ovigerous females,

1 mature male, Spalding Cove, 4-12 m. South Australia, 5 Nov 1969.

Additional material. Numerous sites in Tasmania, Victoria, South

Australia, Western Australia, New South Wales, Queensland; including

many lots registered in Museum Victoria.

Diagnosis. Carapace smooth; branchiostegal spine set back

from anterior edge of carapace and open posteriorly; branch-

iostegal groove running from carapace edge upwards and pos-

teriorly to dorsoposterior edge of the spine where it stops.

Rostrum upper border with 7-10 teeth (usually 8 or fewer,

commonly 9); 2-3 of these (usually 2) postorbital; dorsal teeth

evenly spaced although proximal 3 or 4 often closer together

than remainder and incompletely articulated; ventral rostral

border with 4-6 teeth (usually 5) evenly spaced over distal two-

thirds;

Description, {based on 15 males, 20 females from Margate

Beach, Tasmania) Maximum body length (orbit to telson tip)

c. 30 mm in males, c. 40 mm in females.

Carapace smooth; antennal spine strong and marginal;

branchiostegal spine set back from anterior edge of carapace

and open posteriorly; branchiostegal margin rounded, branch-

iostegal groove running from carapace edge upwards and

posteriorly to dorsoposterior edge of the spine where it stops.

Eyes well developed, with ocellus; interocular tooth well

developed.

Rostral tip level with or exceeding terminal setae of scapho-

cerite; length:depth ratio 6.9-10.0 in males, 6.2-8. 8 in females;

upper border straight to concave; body length;rostral length

ratio 2.5-3. 1 in males, 2.8 - 3.6 in females; lateral carina

gently and concavely curved, degree of curvature variable;

third to half rostrum above lateral carina; upper border with

7-10 teeth (usually 8 or less commonly 9); 2-3 of these

(usually 2) postorbital; dorsal teeth evenly spaced although

248

Terry M. Walker and Gary C. B. Poore

Figure 2. Migration of the carapace spine during growth to final posi-

tion. a-e, Palaemon intermedius', f-j, Palaemon dolospina. Body

lengths of shrimps (mm): a, 6.0; b, 6.2; c, 6.5; d, 6.6; e, 6.6; f, 6.3; g,

6.2; h, 6.3; i, 6.8; j, 8.4.

proximal 3 or 4 are often closer together than remainder and

incompletely articulated; rostral tip usually bifid but may be tri-

fid or a single point; single row of plumose setae along upper

border between but not on teeth; ventral rostral border with 4-6

teeth (usually 5) evenly spaced over distal two-thirds; 2 rows of

plumose setae on ventral border between but not on teeth.

Antennule well developed, article 1 of peduncle about 2.5

times length of articles 2 and 3; article 2 slightly longer than 3,

bearing elongate, blunt projection on medial edge; basal article

elongate and rectangular, about 2.3 times as long as wide,

expanded and flattened on its inner edge, bearing several trans-

verse and longitudinal rows of plumose setae, including

subapical row extending across dorsal surface; setae longer

over inner half of this row; basolateral spine slender, sharply

pointed, reaching over half article 1 ; outer flagellum of anten-

nular exopod much longer than inner flagellum; inner flagellum

fused with outer usually over basal third of its length, about 13

articles in adult, fused up to half its length in juvenile; each free

article of inner flagellum with medial and terminal transverse

row of sensory setae, medial row bearing 3-4 setae and trans-

verse row 4 setae in males, both rows bearing 3 setae in

females.

Antenna with well developed peduncle with strong disto-

lateral tooth, well developed flagellum and scaphocerite; prox-

imal article of flagellum with blunt anteroventral projection;

flagellum naked. Scaphocerite elongate, rectangular, extending

well beyond peduncle, widest just anterior to its base, about one

quarter of body length, 3. 3-4.3 times as long as wide; distal and

inner lateral edges bearing row of long plumose setae; outer

edge naked, slightly concave, ending distally in well developed

tooth that just fails to overlap anterior edge of lamella; inside

edge straight or slightly concave.

Mandible- incisor of one side with 3 teeth (either side),

incisor of the other usually with 4; palp with 3 articles; article

3 longest, with 3 terminal setae longer than article.

Maxillule endopod with bifid tip; distal lobe acute, naked;

proximal lobe bearing 1 plumose seta; coxal endite with long

setae and terminally 4-5 stout setae; basal endite with 3 rows

of stout setae along terminal, inner edge extending down

proximal margin as longer setae.

Maxilla basal endites elongate, nearly rectangular, both

bearing long, stout setae terminally, distal endite with row of 5

setae along distal margin near its base; endopod with 3 short

setae; scaphognathite proximal lobe wider and shorter than

distal; coxa with small expansion near base (?epipod).

Maxilliped 1 with rectangular coxa with endite and bilobed

epipod; basis with endite; endopod with 1 long, subterminal

seta; well developed exopod with prominent lateral lobe;

endites with stout setae along mesial edges, and distal on basal

endite; both with single row of similar setae on posterior face.

Maxilliped 2 with well developed exopod; coxa with poorly

developed endite (on its outer edge), epipod and podobranch;

ischiobasis about twice as long as wide, with scattered setae;

endopod merus half length of ischiobasis, carpus about half

length of merus, propodus expanded, about as long as merus

but twice as wide, dactylus short and wide; anterior and mesial

margin of propodus bearing rows of setae, some on anterior

margin strong.

Maxilliped 3 epipod, medial margin setose; exopod extend-

ing beyond article 1 of endopod. Endopod ischio-merus 1.4-2

times as long as propodo-dactylus; carpus 1.2-1. 6 times as long

as latter; ischio-merus with 2 rows of long setae, one on each of

posterior and anteromesial margins; carpus with similar rows

and additional row on lateral surface; propodo-dactylus with

stout terminal seta and numerous plumose and serrate setae,

parallel rows of shorter, serrate setae across anteromesial sur-

face and longer plumose setae elsewhere. Two arthrobranchs,

one rudimentary and obscured by the larger; maxilliped endo-

pod relatively shorter in males than females. Body length:

maxilliped 3 length ratio 3.7- 4.5 in males, 3. 5-4. 3 in females.

Pereopod 1 articles smooth; reaching approximately to level

of scaphocerite spine; tufts of serrate setae on posteroproximal

region of palm and distomesial surface of carpus; outer surface

of propodal finger with tufts of serrate setae, simple setae on

both fingers; posteromesial ridge of ischium and merus with

single row of long setae; fingers with simple cutting edge,

0.8-1. 2 times as long as palm; carpus 1. 5-2.5 times as long as

chela and 1. 1-1.4 times as long as merus; merus 1.1-1. 9 times

as long as ischium. Body length: pereopod 1 ratio 2.5-3. 1 in

males, 2.0-2. 6 in females.

Palaemon and its southern Australian species

249

Figure 3. Habitus with colour pattern, a, Palaemon dolospina\ b, telson of P. dolospina', c, Palaemon intermedins .

Pereopods 2 equal, similar in males and females, articles

smooth, usually exceeding scaphocerite by all of palm; fingers

slender, curved, with small tooth at base, cutting edges serrate,

sometimes one or both smooth, particularly in smaller individ-

uals; palm swollen, laterally flattened, 2. 8-3. 4 times as long as

wide; palm 0.9-1. 2 as long as fingers, usually longer; carpus

long and slender, narrow proximally, 6.0-8. 5 times as long as

wide at that point, distally 1. 7-2.4 times as wide as at base,

length 6.0-8.5 times distal width; chela 0.9-1. 4 times as long

as carpus and 1.2-1. 5 times merus; merus 1.0-1. 3 times as long

as ischium. Body length: pereopod 2 ratio 1.6-1. 9 in males,

1.3-1. 8 in females.

Pereopods 3-5 progressively longer, mainly owing to

increase in lengths of propodi.

Pereopod 3 simple, slightly less than half body length;

dactylus with small cutting edge on posterior border; propodus

with 2 rows of long setae anteriorly, 2 variable rows of short

plumose setae, outer row with 5-9 setae; propodus 2.0-2. 5

times as long as dactylus and 1.5-1. 9 times as long as carpus in

males, 1. 6-2.4 in females; merus 1. 1-1.3 times as long as

propodus and 1. 1-1.5 times as wide; merus 1. 9-2.4 times as

long as ischium; 1.9-2. 1 times as long as carpus in males,

1. 9-2.4 in females.

Pereopod 4 similar to 3, but slightly longer; propodus length

about 2.5 times as long as dactylus, about twice as long as

carpus; merus about same length as propodus and twice as long

as ischium.

Pereopod 5 similar to 3 and 4, slightly longer than 4; row of

5-7 short setae on lateral posterior margin of propodus with

5-7 parallel rows of serrate setae distally; mesial posterior edge

with row of 7-10 short setae. Propodus 2.6-3. 1 times as long as

dactylus in male, 2.0-2. 8 in female.

Branchial formula (r = rudimentary).

Maxillipeds

Pereopods

Total

gills

1 2

1 2 3 4 5

Pleurobranchs

- -

11111

Arthrobranchs

- -

1+lr

_____

1 + Ir

Podobranchs

- 1

_____

Exopods

1 1

_____

Epipods

1 1

_____

Pleopod 1 with well developed exopod and smaller endo-

pod; appendix interna absent. Endopod of male little more than

half as long as exopod, with convex outer edge, concave inner

edge, about 4 times as long as wide. Endopod of female

smaller, about one third as long as exopod, about 3 times as

long as wide.

Pleopods 2-5 with equally developed exopod and endopod;

endopod with appendix interna. Endopod of male pleopod 2

with appendix masculina, longer than appendix interna, usual-

ly with 22 setae along its length, 6 of which are apical or sub-

apical; each seta up to half as long as appendix masculina.

250

Terry M. Walker and Gary C. B. Poore

Figure 4. Palaemon intertnedius. a, rostrum (female, 33.2 mm); b, rostmm (male, 20.8 mm); e, earapace spine (female, 30.2 mm); d, e, antennule

and scaphocerite (female, 30.2 mm); f, scaphocerite (male, 28.0 mm); g, mandible; h, maxillule; i, maxilla; j-1, maxillipeds 1-3; m, mandible of

post-larva (all mouthparts, female, 30.2 mm).

Pleopods 1-2 and to a lesser extent 3-5 with keel-shaped,

flattened extension on lateral edge, smaller in male than female.

First abdominal pleuron about 2-3 times as long as wide;

second abdominal pleuron about 1.5 times as long as

wide; apex of fifth pleuron acute, with short terminal

spine; sixth abdominal article 1.4- 1.8 times as long as fifth.

Ventral edges of abdominal pleura with row of short, plumose

setae.

Telson 2.0-2. 5 times wider at base than apex, length 2.0-4.2

times basal width; 1. 1-1.4 times as long as sixth abdominal

somite; with 2 pairs of dorsal stout setae and larger inner and

smaller outer terminal pair; apex produced into acute spine

flanked by a 1 simple seta on each side and occasionally a

shorter second pair; apical spine much shorter than the terminal

setae; tuft of long plumose setae dorsomedially near base of

telson.

Uropods endopod elongate, fringed with long plumose

setae; exopod rectangular; transverse suture on exopod about

two-thirds of distance from base; outer edge of exopod entire,

ending at suture line in an immovable spine; second movable

seta often present inside spine; remainder of margin fringed by

row of long plumose setae; posterior to immovable spine, dor-

sally and ventrally, a row of about 12 long, non-plumose setae

set back from margin, extending almost to apex of exopod; ven-

trally, outer margin from base of the exopod to the immovable

spine with row of stout, simple setae just inside the margin.

Colour pattern. Carapace chromatophore lines usually

red; abdominal markings generally olive green, black and red;

accessory flagellum distinctly marked with alternate red and

white bands.

Sexual dimorphism in adults. Males are smaller than females;

with larger sternal process on the eighth thoracomere; smaller

keel-like expansion on outer edge of peduncles of pleopods;

appendix masculina on pleopod 2; significantly larger endopod

on pleopod 1 ; 4 setae in distal sensory row on each article of

antennular inner flagellum (3 in adult female); rostrum more

slender and longer; longer carpus in pereopod 2; maxilliped 3,

pereopods 1 and 3 shorter relative to body length; the ratio of

length of propodus and dactylus of pereopod 5 significantly

greater.

Females ovigerous from 19.5 mm body length; bearing

approximately 100-700 ovoid eggs, larger females bearing

more eggs; egg size 0.55-1.0 mm depending on developmental

stage, 0.9-1. 0 mm when ready to hatch.

Ontogenetic changes. Some characters alter as the size of males

and females increases. In females, the scaphocerite becomes

stouter and shorter relative to body length; the number of

articles over which the antennular flagella are fused increases;

the carpus and merus of pereopod 1 become longer relative to

other articles; the number of parallel rows of setae on the

propodus of pereopod 5 increases, and pereopod 2 becomes

longer. In males, the number of articles over which the anten-

nular flagella are fused increases; length ratios between articles

of pereopods 1-5 differ; pereopod 1 becomes slightly shorter

relative to body length.

In early post-larval stages, the mandibular palp and the

branchiostegal groove are absent and a branchiostegal spine is

present on the edge of the carapace. Over a series of moults, a

palp of three articles and the branchiostegal groove form and

Palaemon and its southern Australian species

251

Figure 5. Palaemon intermedius . a-f, pereopods 1-5; b details of chela of pereopod 1 (male, 28.0 mm), g, h, pleopods 1, 2 (male); i, j, pleopods

1, 2 (female), k, telson. 1, appendix masculina.

the branchiostegal spine migrates posteriorly and upwards to its

final position in adults.

Variation. The type or types of Leander intermedius Stimpson,

1860 are lost. They were collected from Port Jackson, NSW.

Examples of the species from nearby Port Hacking most

resemble the typical Tasmanian form from Margate Beach but

examples from other locations including Tasmania differ.

Deep-water form (5 adult females, 29.3-36.6 mm,

D’Entrecasteaux Channel). More slender and longer legged

than shallow water form. Scaphocerite more slender, rostrum

with 2-3 but usually 3 postorbital teeth rather than 2;

Pereopods 1, 3, 5 longer, body length 2.2-1.3 times as long as

pereopod 1, 1.7-1. 9 times as long as pereopod 2, 1.5-1. 6 times

as long as pereopod 5; carpus of pereopod 2 longer, carpus

length 2.0-2. 3, 3.7^.7, 1.2-1. 4 and 1.9-2. 1 times as long as

chela, fingers, merus and ischium respectively; palm more

swollen, length 2.6-2. 8 times its maximum width; apex of car-

pus more expanded; pereopods 3 and 5 with longer propodus;

telson stouter, at least at its base, 2. 2-3. 8 times as long as basal

width.

Intermediate form (4 ovigerous females, 21.2-33.0 mm, 2

adult males 26.2 mm, Western Port, Vic.). More similar to the

deep-water form particularly in number of postorbital rostral

teeth, length of pereopods 2, 3, 5, and the degree the palm is

swollen on pereopod 2.

South Australian form (3 females (2 ovigerous) 21.9-35.9

mm, 1 adult male 20.8 mm, Spalding Cove, Port Lincoln. SA).

Resembles closely Tasmanian material except for 3 postorbital

rostral teeth and a more swollen palm on pereopod 2.

Remarks. Palaemon intermedius bears what appears to be an

hepatic spine but it is set lower down and not as far back from

the carapace edge as is usual in Macrobrachium, the genus to

which it was assigned by Holthuis (1952). The branchiostegal

groove appears to run back to the spine and over it, but ceasing

at the back of the spine. The spine is open posteriorly, having

no suture line between it and the carapace in that region. The

migration of the branchiostegal spine from the carapace edge

during ontogeny is arrested partly completed and the value of

this character for generic assignment is therefore limited.

Generic assignment is based principally on the other characters.

Palaemon dolospina sp. nov.

Eigures le, 2f-j, 3c, 6, 7

Palaemon sp. — Walker, 1979.

Material examined. Holotype. Margate Beach, North West Bay, Tas.

(lat long) in Zostera muelleri and Heterozostera tasmanica on sand,

0.2-1 m, T. Walker, 12 Dec 1973, NMV J52659 (male, 6.5 mm pos-

torbital carapace length, 42 mm total length).

Allotype. Collected with holotype, NMV J52660 (ovigerous female,

8.9 mm postorbital carapace length, 45 mm total length)

Paratypes. Collected with holotype, NMV J52661 (92 specimens,

4.7-9.0 nun postorbital carapace length)

Material described. 26 males, 12-35 mm body length, 28 females,

including 6 ovigerous, 12.6-40 mm body length, Margate Beach and

in brackish water from North West Bay River Estuary, Margate,

Tasmania, in seagrasses Zostera muelleri and Heterozostera tasmani-

ca, T. M. Walker, 1975. 1 ovigerous female, 32.1 mm body length.

Western Port, Victoria, intertidally on mud. Fisheries and Wildlife

252

Terry M. Walker and Gary C. B. Poore

Figure 6. Palaemon dolospina. a, rostrum (female, 31.8 mm); b, rostrum (male, 32.8 mm); c, carapace spine (female, 31.8 mm); d, e, antennule

and scaphocerite (female, 31.8 mm); f, scaphocerite (male, 31.8 mm); g, mandible; h, maxillule; i, maxilla; j-1, maxillipeds 1-3 (all mouthparts,

female, 31.8 mm).

Department, Victoria, 8 Jan 1974. 8 females, 2 ovigerous 13.9-34.6

mm, 6 males, 15.3-23.8 mm, ICI Saltfields, Port Gawler, South

Australia, ICI staff ,13 Jan 1973. 6 ovigerous females, 37.7-50.7 mm,

Newport Power Station, Victoria. H.A. Morrison, 21 July 1949.

Additional material from 50 sites in Tasmania (including Flinders

and King Islands), 2 in Victoria, 3 in South Australia.

Diagnosis. Carapace smooth; branchiostegal spine set back

from and its tip barely reaching anterior carapace edge, set high

up on brachiostegite; branchiostegal groove runs up to, loops

over the spine and sweeps down again posteriorly. Rostrum

upper border with 6-9 teeth; 1-2 of these (usually 2) postor-

bital; dorsal teeth unevenly spaced, first set back from rest on

carapace, next 3 or 4 evenly spaced, incompletely articulated,

increasingly longer gap between next 3 until penultimate tooth

close to distal tooth; ventral rostral border with 3-6 teeth (usu-

ally 4), evenly spaced over distal two-thirds.

Description, (based on 26 males, 28 females from Margate

Beach, Tasmania. Characters resembling P. intermedins not

described) Maximum body length (orbit to telson tip) c. 42 mm

in males, c. 5 1 mm in females.

Carapace smooth; antennal spine strong and marginal;

branchiostegal spine set back from and its tip barely reaching

anterior carapace edge, set high up on brachiostegite;

branchiostegal groove runs up to, loops over the spine and

sweeps down again posteriorly.

Eyes well developed, with ocellus; interocular tooth well

developed.

Rostral tip exceeding lamella of scaphocerite; length:depth

ratio 6.5-9. 3 in males, 5. 3-7.6 in females; upper border straight

to concave; body length:rostral length ratio 2.4-3. 6 in males,

2.5-3. 5 in females; lateral carina gently concave and directed

upwards at tip; third to half rostrum above lateral carina; upper

border with 6-9 teeth; 1-2 of these (usually 2) postorbital;

dorsal teeth unevenly spaced, first set back from rest on cara-

pace, next 3 or 4 evenly spaced, incompletely articulated,

increasingly longer gap between next 3 until penultimate tooth

close to distal tooth; rostral tip appears bifid; single row of

plumose setae along upper border between but not on teeth;

ventral rostral border with 3-6 teeth (usually 4), evenly spaced

over distal two-thirds; 2 rows of plumose setae on ventral

border between but not on teeth.

Antennule well developed; basal article bearing subapical

transverse row of 10 plumose setae, not extending onto stylo-

cerite; basilateral spine moderately slender, sharply pointed,

reaching over half basal article of peduncle; inner flagellum

fused with outer usually over basal third of its length, about

11-12 articles in male and 14-15 in female, fused up to half its

length in juveniles; each free article of inner flagellum with

mesial and terminal transverse row of sensory setae, mesial row

of 3-4 and distal of 4 setae in female, mesial row of 4 and dis-

tal of 5 (occasionally 4) setae in male; sensory setae noticeably

shorter than those in P. intermedins.

Antennal scaphocerite elongate, rectangular, widest near

base, 3.2-4. 1 times as long as wide; outer edge slightly convex

to slightly concave, with terminal spine that just fails reach

anterior edge of lamella; inside edge straight or concave over

distal two thirds, convex proximally.

Mandible incisor of one side with 3 teeth (either side),

incisor of the other usually with 4; palp with 3 articles; article

Palaemon and its southern Australian species

253

Figure 7. Palaemon dolospina. a-f, pereopods 1-5; b details of chela of pereopod 1 (male, 32.8 mm), g, h, pleopods 1, 2 (male); i, j, pleopods 1,

2 (female), k, telson. 1, appendix masculina.

2 with 10 or more setae; article 3 slightly swollen, setose

including 3 terminal and 2 subterminal setae; article 3 length

about twice article 2, article 1 about 1.5 times article 2.

Maxillule endopod with bifid tip, naked, distal lobe acute.

Maxilla endopod with 2 short setae on inner edge and about 10

on proximal half of outer edge. Maxilliped 1 endopod without

seta.

Maxilliped 3 epipod, mesial margin setose; exopod extend-

ing to end of article 1 of endopod. Endopod ischio-merus 1 .3-2

times as long as propodo-dactylus; carpus 1.2-1. 6 times as long

as latter; ischio-merus with 2 rows of long setae, one on each of

posterior and anteromesial margins; propodo-dactylus with

stout terminal seta Body lengthimaxilliped 3 length ratio

3. 8-4.5 in both sexes.

Pereopod 1 articles smooth; reaching approximately to tip of

scaphocerite lamella; tufts of serrate setae on posteroproximal

region of palm and distomesial surface of carpus; outer surface

of propodal finger with tufts of serrate setae, simple setae on

both fingers; posteromesial ridge of ischium and merus with

single row of long setae; fingers with simple cutting edge,

0.8-1. 4 times as long as palm; carpus 1.3-2. 2 times as long as

chela and 1.0-1. 4 times as long as merus; merus 1.3-1. 8 times

as long as ischium. Body length:pereopod 1 ratio 2.4-2. 9 in

both sexes.

Pereopods 2 equal, similar in males and females, articles

smooth, usually exceeding scaphocerite by at least fingers;

fingers slender, curved, with small tooth at base, cutting edges

serrate, sometimes one or both smooth, particularly in smaller

animals; palm slightly swollen, laterally flattened, 2. 6^.0

times as long as wide; palm 1.0-1. 4 as long as fingers; carpus

long and slender, narrow proximally, thickens distally to be

1. 5-2.5 times wider than at base, length:apical width ratio

6.4- 8. 8 in males, 5. 2-7.7 in females; chela 0.8-1. 2 times as

long as carpus and 1.0-1. 4 times merus; merus 1.0-1. 5 times as

long as ischium. Body length: pereopod 2 ratio 1.6-2. 2 in both

sexes.

Pereopods 3-5 progressively longer, mainly due to increase

in lengths of propodi. Pereopod 3 reaching about end of

scaphocerite; dactylus with small cutting edge on posterior bor-

der; propodus with 2 rows of long setae anteriorly, 2 variable

rows of short flagellate setae, outer row usually with 4-5 setae

in male, 5-6 in female, inner row usually with 3-5 in male, 5

in female; propodus 1.5-2. 1 times as long as dactylus and

1.4- 1. 8 times as long as carpus in males, 1. 6-2.0 in females;

merus 1.0-1. 4 times as long as propodus and 0.9-1. 5 times as

wide; merus 1. 8-2.4 times as long as ischium; 1.6-2. 2 times

as long as carpus in male, 1. 8-2.4 in female.

Pereopod 4 similar to 3, but slightly longer. Pereopod 5 sim-

ilar to 3 and 4, slightly longer than 4.

Branchial formula as for P. intermedius.

Pleopod 1 with well developed exopod and smaller endo-

pod; appendix interna absent. Endopod of male little more than

half as long as exopod, with convex outer edge, concave inner

edge, about 4 times as long as wide. Endopod of female

smaller, about one third as long as exopod, about 3 times as

long as wide.

Pleopods 2-5 with equally developed exopod and endopod;

endopod with appendix interna. Endopod of male pleopod

2 with appendix masculina, longer than appendix interna,

usually with 24 setae along its length, 6 of which are apical or

subapical; each seta less than third as long as appendix

masculina. Pleopods 1-2 and to a lesser extent 3-5 with keel-

shaped, flattened extension on lateral edge, smaller in male

than female.

254

Terry M. Walker and Gary C. B. Poore

First abdominal pleuron about 2-3 times as long as wide;

second abdominal pleuron about 1.5 times as long as wide;

apex of fifth pleuron acute, with short terminal spine; sixth

abdominal article 1.3-1. 9 times as long as fifth. Ventral edges

of abdominal pleura with row of short, plumose setae.

Telson 2.0-2. 8 times wider at base than apex, length 3.0-4.3

times basal width; 1.0-1. 3 times as long as sixth abdominal

somite; with 2 pairs of dorsal stout setae and larger inner and

smaller outer terminal pair; apex produced into acute spine

flanked by 1 simple seta on each side and occasionally a

shorter second pair; apical spine much shorter than terminal

setae; tuft of long plumose setae dorsomedially near base

of telson.

Uropods endopod elongate, fringed with long plumose

setae; exopod rectangular; transverse suture on exopod incom-

plete laterally, about two-thirds of distance from base; outer

edge of exopod entire, ending at suture line in an immov-

able spine; second movable seta often present inside spine;

remainder of margin fringed by row of long plumose setae;

posterior to immovable spine, dorsally and ventrally, a row of

about 9 long, non-plumose setae set back from margin, extend-

ing almost to apex of exopod; ventrally, outer margin from base

of the exopod three quarters of the way to the immovable spine

with row of stout, simple setae just inside the margin.

Colour pattern. Carapace chromatophore lines red; distinct

transverse abdominal bars generally red but may be olive

green; abdominal bars present on first post-larva as single lines

of orange or yellow chromatophores; lateral carina of rostrum

often with row of large, white chromatophores; hexagonal

matrix of abdominal segments most distinct in large females,

made up of small, olive green and some scattered white chrom-

atophores; viewed dorsally, tail fan with white, transverse

bands or patches.

Sexual dimorphism in adults. Males are smaller and more

slender than females; with larger sternal process on the eighth

thoracomere (almost absent in female); smaller keel-like

expansion on outer edge of peduncles of pleopods; appendix

masculina on pleopod 2; significantly larger endopod on pleo-

pod 1 ; 5 setae in distal sensory row on each article of antennu-

lar inner flagellum (3-4 in adult female); rostrum more slender

and longer; broader carpus in pereopod 2; longer carpus in

pereopod 3; ratio of length of propodus and dactylus of

pereopod 5 greater.

Females ovigerous from 25 mm body length; bearing

approximately 140-840 ovoid eggs, larger females bearing

more eggs; egg size 0.65-1.15 mm depending on develop-

mental stage, 0.9-1.15 mm when ready to hatch.

Ontogenetic changes. Some characters alter as the size of

recognisably male and female animals increases. In females,

the rostrum and scaphocerite becomes stouter and shorter

relative to body length; the number of articles over which the

antennular flagella are fused increases; pereopods 2, 3, 5

become longer relative to the body; the fingers of pereopod 1

become shorter relative to other articles; the palm of pereopod

2 becomes less swollen, the carpus more expanded and fingers

shorter; the number of parallel rows of setae on the propodus of

pereopod 5 increases; and the telson becomes stouter. In males.

the number of articles over which the antennular flagella

are fused increases; length ratios between articles of pereopods

1-5 differ; caipus of pereopod 1 becomes relatively longer;

pereopod 2 palm becomes less swollen and fingers shorter.

In early post-larval stages, the mandibular palp and

branchiostegal groove are absent and a branchiostegal spine is

present on the edge of the carapace. Over a series of moults, a

palp of 3 articles and branchiostegal groove form and the

branchiostegal spine migrates posteriorly and upwards to its

final position in adults.

Etymology. From Latin, dolus meaning deceit and spina, a spine,

alluding to the deceptive position of the branchiostegal spine.

Remarks. Palaemon dolospina bears a branchiostegal groove

looping over a spine set back from the carapace edge albeit

displaced dorsally somewhat. Consequently, although the spine

is situated unusually high on the carapace between the usual

branchiostegal and hepatic position, it can be defined as

branchiostegal. Females are larger than males and the second

pereopods of males are not spinulose or markedly sexually

dimorphic. The mandibular palp is of three articles. In spite its

unusual branchiostegal spine/groove arrangement, this species

clearly belongs to Palaemon.

Keys to species of Palaemon and Palaemonetes from

southern Australia

The shallow marine and estuarine Australian palaemonid fauna

includes eight, possibly nine, species distributed as follows:

Palaemon debilis Dana, 1852 - Widespread in Indo-West

Pacific, Qld, redescribed by Holthuis (1950) and Chace (1972).

Palaemon cf. debilis Dana, 1852 - NSW, eastern Vic., also

similar to Palaemonetes atrinubes but its status remains

unresolved.

Palaemon dolospina sp. nov. - Vic., Tas., SA.

Palaemon intermedins (Stimpson, 1860) - Qld (N to

Moreton Bay), NSW, Vic., Tas., SA, WA.

Palaemon litoreus (McCulloch, 1909) - NSW, Vic., SA,

WA. Palaemon serenus and P. litoreus have in the past, been

separated on the basis of the carpus exceeding the palm of pere-

opod 2 in P. serenus but being shorter in P. litoreus (McCulloch,

1909; Hale, 1927b; Bray, 1976). Examination of material of

both species from Western Australia has shown that, the carpus

exceeds the palm in both species.

Palaemon macrodactylus Rathbun, 1902 - NSW, SA [native

to Japan, Korea, northern China, introduced to San Francisco

Bay and two Australian localities (Buckworth, 1979; Walker,

1979). Identifications of Australian material confirmed by TW

and L. Holthuis, 1976].

Palaemon serenus (Heller, 1862) - southern Qld, NSW,

Vic., Tas., SAWA.

Palaemonetes australis Dakin, 1915 - WA.

Palaemonetes atrinubes Bray, 1976 - NT, Qld, Vic, WA

1 . Carapace spine set back from carapace edge by less than its

length and situated between branchiostegite and ridge of

antennal spine; open posteriorly; branchiostegal groove

runs up to dorsoposterior edge of spine but not past it (Figs

Palaemon and its southern Australian species

255

Id, 4c); pereopod 2 smooth and equal; rostrum with 7-10

dorsal teeth, usually 8, 2-3 postorbital, proximal 4 incom-

pletely articulated; 4-6, usually 5, ventral teeth

Palaemon intermedius

— Carapace spine set below branchiostegal groove which

may deviate up and over the spine; spine may be on or set

back from edge of carapace; pereopod 2 and rostrum not as

above 2

2. Branchiostegal spine set back from the carapace edge,

overlapping it at most only with tip 3

— Branchiostegal spine at or very near carapace edge just

under start of branchiostegal groove; groove running back

from carapace edge in a shallow arc, not upwards sharply

and down again 4

3. Branchiostegal spine almost or just reaching carapace edge

with tip only; set above level of starting point of bran-

chiostegal groove which runs upwards to front of spine,

loops sharply over it and sweeps down again posteriorly

(Figs le, 6c); pereopod 2 carpus 1-1.5 times as long as

merus; mandibular palp with 3 articles

Palaemon dolospina

— Branchiostegal spine set well back from the carapace edge

which it never overlaps; branchiostegal groove deviating

only slightly as it passes over spine (Fig Ic); pereopod 2

carpus 1.3-1. 5 times as long as merus; mandibular palp

absent Palaemonetes australis

4. Antennule inner flagellum fused for greater than half

of its length, usually over about two thirds, 16-20 articles

fused in adults; mandibular palp absent

Palaemonetes atrinubes

— Antennule inner flagellum fused for half or usually less of

its length, less than 15 articles fused in adults 5

5. Pereopods slender (carpus of pereopod 2 1. 5-2.0 times as

long as chela, 1.4-1. 7 times as long as merus) 6

— Pereopods not unusually slender (carpus of pereopod 2

shorter than chela, 0.9-1. 1 times as long as merus .... 7

6. Rostrum exceeding scaphocerite by at least quarter of its

length; proximal 2 dorsal teeth fully articulated; 5-7

ventral teeth Palaemon debilis

— Rostrum exceeding scaphocerite by less than quarter of its

length; no fully articulated dorsal teeth; 3-4 ventral teeth

Palaemon cf. debilis

7. Pereopod 2 stout but short, 0.4-0. 5 times body length

Palaemon litoreus

— Pereopod 2 stout but long, 0.7-0. 9 times body length . .8

8. Rostrum with 9-15 dorsal teeth (usually 10-12), all

incompletely articulated; teeth directed upwards; only one

sixth of the antennule inner flagellum fused (4-6 articles)

Palaemon macrodactlyus

— Rostrum with 6-9 dorsal teeth, proximal 3 incompletely

articulated; about one third of antennule inner flagellum

fused (about 12 articles in adults) .... Palaemon serenus

In fresh material, pigmentation pattern can help separate

species in this alternative key. The colour description for

Palaemon litoreus was taken from a description by McNeill in

Holthuis (1952) and those for Palaemonetes australis and

P atrinubes from Bray (1976).

1 . Without dorsal hump on abdominal somite 3 2

— Distinct dorsal hump on abdominal somite 3 4

2. Long, slender rostrum, substantially exceeding scapho-

cerite Palaemon debilis

— Rostrum at most just exceeding scaphocerite 3

3. Distinct black spot at posterolateral end of sixth abdominal

somite; 3 distinct red and blue lines on carapace; 1 trans-

verse line across posterior edge of third abdominal somite

Palaemonetes atrinubes

— Without black spot at base of sixth abdominal somite . . .

Palaemon cf. debilis

4. Broad, red band across most of palm of pereopod 2; fingers

white; abdomen with longitudinal rows of few large spots

of red, blue and black chromatophores; distinct longitudi-

nal stripes on carapace Palaemon serenus

— Without wide red band or only a narrow one across

palm of pereopod 2; if narrow red band present,

abdomen with saddle stripes of red on dorsal and lateral

surfaces 5

5. Grey or olive-green; distinct wide, grey band across palm

of pereopod 2; diffuse longitudinal and oblique rows on

carapace; diffuse transverse lines on posterior edges of

abdominal articles Palaemon macrodactylus

— Without wide grey or olive-green band on palm of

pereopod 2 6

6. Saddle stripes of red across back and sides of whole body;

less distinct on carapace; palm of pereopod 2 with narrow

red band; fingers with similar band half way along

Palaemon litoreus

— Without longitudinal stripes on abdomen 7

7. Distinct red lines either obliquely on carapace or trans-

versely on abdomen 8

— Overall olive-green or brown due to red, white, yellow

and blue chromatophores; carapace with diffuse dorsal

and anterior bands and indistinct mottling posteriorly;

antennule inner flagellum pale red

Palaemonetes australis

8. Distinct transverse red and/or occasionally olive stripes

across all abdominal somites; less distinct longitudinal

and oblique red stripes on carapace (Fig. 2a, b); anten-

nule inner flagellum pink with few white flecks; often

row of white chromatophores along lateral carina of the

rostrum Palaemon dolospina

— Distinct oblique and transverse red lines on carapace; few

scattered red, olive and black chromatophores forming

indistinct transverse lines on abdominal segments (Fig.

2c); antennule inner flagellum with definite red and

white bars, persisting as purple bars in ethanol-preserved

specimens Palaemon intermedius

Acknowledgements

We thank RS. Lake (Monash University) and David Ritz

(University of Tasmania) who supervised the PhD research by

the first author that is the most part of this contribution, and

J.L. Hickman for taxonomic advice. Sherri Lehmann prepared

the plates from original drawings by the first author.

256

Terry M. Walker and Gary C. B. Poore

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Memoirs of Museum Victoria 60(2): 257-283 (2003)

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

http://www.museum.vic.gov.au/memoirs

Linguimaera Pirlot, 1936 (Crustacea, Amphipoda, Melitidae), a valid genus

Traudl Krapp-Schickel

Forschungsinstitut und Museum A. Koenig, Adenauerallee 160, D-53113 Bonn, Germany (traudl.krapp@uni-bonn.de)

Abstract Krapp-Schickel, T. 2003. Linguimaera Pirlot, 1936 (Cmstacea, Amphipoda, Melitidae), a valid genus. Memoirs of

Museum Victoria 62(2): 257-283.

The genus Maera sensu lato is among the largest genera in the Melitidae, used as a catch-all for species lacking the

defining characters of other genera. Recently, several authors have started splitting this species flock into better-defined

genera and the present paper continues this task. It revives Linguimaera erected in 1936 by Pirlot, but soon after syn-

onymised again with Maera. The most shaking characters are a short third article of the mandibular palp, asymmetrical

gnathopods in both sexes, epimeral plate 2 with a sinus, epimeral plate 3 serrate on its posterior margin and telsonic lobes

asymmetrically incised. Eight species can be attributed to Linguimaera, of which seven are new: L. pirloti (type species),

L. bogombogo, L. caesaris, L. garitima, L. kellissa, L. leo, L. mannarenis and L. tias.

Key words Taxonomy, Amphipoda, Linguimaera, Indo-Pacific

Introduction

The suspicion that the amphipod family Melitidae may be para-

phyletic is not new: Bousfield (1973) conceived it as compris-

ing marine members of Barnard’s Gammaridae (Barnard,

1969); neither he nor following researchers could find a single

synapomorphy. Thus, genera are grouped on the basis of shared

character states which may be homoplasic. Recent results of

chromosome research confirmed big differences in chromo-

some numbers and karyotype-morphology between genera

(Libertini and Krapp-Schickel, 2000).

Besides this, within Maera Leach, 1814 sensu lato, charac-

ters vary significantly. Barnard and Barnard (1983: 623) listed

59 species in their overview of the genus and diagnosed it:

“Like Elasmopus but article 3 of mandibular palp not falciform.

Like Ceradocus but inner plate of maxilla 2 lacking facial row

of setae; maxillae generally poorly setose medially. Species of

Maera probably polyphyletic, sources from Ceradocus,

Elasmopus, Mallacoota etc.” In recent years, several papers by

me and others have started revision of this taxon by defining

Maera sensu stricto (Krapp-Schickel, 2000; Krapp-Schickel

and Jarrett, 2000), and by splitting off clades into new genera

(Quadrimaera Ruffo and Krapp-Schickel, 2000, Zygomaera

Krapp-Schickel, 2000 and Othomaera Krapp-Schickel, 2000).

Here, the process continues and the present paper deals with the

validation of Linguimaera Pirlot, 1936.

Walker (1904) described a new species of Maera found in

Ceylon. He offered only small sketches and stressed eight dif-

ferences from Maera othonis Milne Edwards, 1830; he named

his species Maera othonides. The type material is no longer

extant and the description is too poor to give an exact idea

about most of the crucial character states. Pirlot (1936) found

amphipods during the Siboga expedition to Indonesia which he

attributed to Walker’s species. As he noticed an enlarged sagit-

tal lobe of the labium, he erected a new genus Linguimaera,

choosing not his material but ''Maera othonides Walker 1904”

as type species. Shortly after, Schellenberg (1938: 49) denied

the taxonomic value of the shape of the upper lip at generic

level (it occurs also in other species of Maera sensu lato), and

synonymised Linguimaera with Maera.

Barnard (1972a: 224) suspected that there might be a group

of species closely related to M. othonides Walker sensu Pirlot,

but opined that the name Linguimaera Pirlot unfortunately was

not available. In my opinion, Pirlot, who probably never saw

Walker’s type material of M. othonides (which I consider to be

a species dubia), wrongly identified his Indonesian material as

that species. In reality he based his description of Linguimaera

on his Indonesian material which in the present paper is

described as a new species, L. pirloti. According to ICZN (4th

edition, 1999) Article 70.3.1, this may now be selected as the

type species, replacing the dubious Maera othonides Walker

chosen by Pirlot.

While studying the rich collections of Museum Victoria,

Melbourne, it became obvious to me that Maera othonides

Walker sensu Pirlot, 1936 shows a series of character states

shared with M. mannarensis Sivaprakasam, 1968 and with

other undescribed species. These peculiarities seem always to

occur together, are probably not convergent but are syna-

pomorphies of a clade of related species as Barnard had already

surmised.

258

TraudI Krapp-Schickel

Abbreviations are as follows: AM, Australian Museum, Sydney;

MCNCr, Museo Civico di Storia Naturale Verona; NMV, Museum

Victoria, Melbourne; USNM, Naturaly History Museum, Smithsonian

Institution, Washington; ZMA, Zoological Museum, Amsterdam.

Symbols used in the figures are as follows: Epl-3, epimeral plates

1-3; Gnl, 2, gnathopods 1, 2 (1 =large, s=small); Hd, head; LL, lower

lip; Md, mandible; Mdp, mandibular palp; Mxp, maxilliped; Mxl, 2,

maxilla 1, 2; ov., ovigerous; P3-7, pereopods 3-7; T, telson; Ul-3,

uropods 1-3; UL, upper lip; Us, urosome.

Maera othonides Walker, 1904 species dubia

Maera othonides Walker, 1904: 273, fig. 29

Remarks. The original description of this species from Ceylon

is slim. One is required to examine illustrations of M. othonis

(Milne Edwards) from Europe to make assumptions on char-

acters Walker did not describe other than by “similar to

M. othonis."' He reported a length of 8 mm, article 3 mandibu-

lar palp “considerably shorter” than article 2, antenna 1 acces-

sory flagellum with 3 articles, the third pleon segment with no

teeth on its lower and 1-3 teeth on the posterior margin, third

uropods truncate, telsonic lobes distally incised having one

short robust seta sitting there, and a second notch on the inside

of the lobe. This description would fit many species of the old

Maera flock. Another species within this geographical distri-

bution is Linguimaera mannarensis but we cannot check the

suspicion that the two are the same.

Indian material cited under the same name by Chilton

(1921: 535, fig. 5; no body length) could well be the same as

that of K.H. Barnard (1935: 285, fig. 5; adults 11 mm) or of

Nayar (1959: 26, pi. 8 figs 1-18; up to 12 mm) and is certainly

not the species from Ceylon. In this, the antenna 1 accessory

flagellum is up to 5 articles, the gnathopod 2 of the female has

the palm slightly convex and of the male regularly excavated;

the epimeral plate 3 is serrate on the posterior (and inferior)

margins; uropod 3 and the posterior portion of the pleon are

densely beset with delicate woolly setae; the telson is cleft

halfway and its lobes distally pointed, with one long setule and

two smaller ones, and no notch on inner margin. The characters

of the mandibular palp (article 3 short, although much shorter

still than in all Linguimaera species here described), the serrate

posterior epimeral plate 3, and the lanceolate uropod 3 would

match the definition of Linguimaera. But all three authors

reported a “pubescent” pleon, densely setose uropod 3 and

(only Nayar) pleon segments serrate along the posterodorsal

margins of the segments. As already surmised by Krapp-

Schickel (2000; 432), the citations of Chilton, K.H. Barnard

and Nayar are synonyms of Ceradomaera plumosa Ledoyer,

1973 and not Maera othonides Walker. That species, according

to Walker (1904) was “very common” in Sri Lanka. Nor is their

material referable to Pirlot’s species, Linguimaera pirloti sp.

nov. Eor the time being, Maera othonides Walker must remain

species dubia.

Distribution. Sri Lanka.

Linguimaera Pirlot, 1936

Eigure 1

Linguimaera Pirlot, 1936: 309. — Schellenberg, 1938: 49 (syn-

onymy of Linguimaera with Maera)

Type species. Linguimaera pirloti sp. nov. (=Maera othonides

Walker sensu Pirlot, 1936), not Maera othonides Walker, 1904;

herein selected, see ICZN (4th edition, 1999) Article 70.3.1

Diagnosis. Body smooth. Eyes reniform, more than twice as

long as wide. Upper lip thickened, in side-view linguiform,

lengthened, reaching between peduncles of antenna 2 (Eig. 1);

mandibular palp article 1 rounded or subquadrate, distally not

produced; article 3 considerably shorter than article 2, maxilla

1 inner plate with 3 robust setae, maxilla 2 not marginally

setose. Antenna 1 longer than antenna 2; antenna 1 accessory

flagellum of 2-6 articles; antenna 2 peduncle article 2 gland

cone not longer than article 3; Ceradocus-Ytke, cephalic cheek

having notch or slit. Coxa 1 anteriorly lengthened, pointed or

rounded, posterodistal corner notched. Gnathopod 1 carpus

CeradocusAYke, swollen, with distoinferior margin usually

lengthened to short, acute tooth, often hardly visible under

dense robust setae; gnathopod 2 sexually dimorphic and asym-

metrical in both sexes, palm ornate and excavate. Pereopod 3

equal to or longer than pereopod 4, merus often somewhat

swollen; pereopods 5-7 slender, basis clearly longer than wide,

often rectolinear with right-angled posterodistal corner, weakly

to strongly serrated posteriorly; dactyli simple, with stiff robust

seta on inner side functioning like a pincer. Pleon dorsally

smooth. Epimeral plates 1, 2 posterodistal comer upwards

curved, acute, followed by shallow sinus defined by another

acute tooth; epimeral plate 3 posterior margin densely serrate,

up to 9-11 teeth; uropod 3 rami much longer than peduncle,

richly spinose, robust setae always much shorter than length of

rami; telson deeply cleft, lobes characteristically asymmetric-

ally incised, the outer end being longer; robust setae clearly

shorter than telson length.

Included species. Linguimaera bogombogo sp. nov., L. caesaris

sp. nov., L. garitima sp. nov., L. kellissa sp. nov., L. leo sp. nov.,

L. mannarensis Sivaprakasam, 1968, L. pirloti sp. nov., L. tias

sp. nov.

Discussion. Lowry et al. (2001) redescribed Megamoera mas-

tersii Haswell, 1879 from type material and figured round, not

sexually dimorphic, eyes and symmetrical not sexually

dimorphic gnathopods. They also redescribed Moera hamigera

Haswell, 1879 on the basis of abundant recent collections. The

latter is obviously a common species on Australian coasts

which nevertheless totally seems to have escaped collecting in

the last 130 years. It has different second epimeral plates, third

uropods and telsonic lobes. Lowry et al. (2001) also

redescribed Megamoera boeckii Haswell, 1879 with an emar-

ginate telson shorter than broad. All these species do not seem

to be closely related to the species flock presented here.

The genus Anamaera Thomas and Barnard, 1985 (from

Elorida, Thomas and Barnard, 1985) appears to be similar to

Linguimaera also having asymmetrical gnathopod 2 and the

same epimeral shape. But the mandibular palp article 3 is not

shorter than article 2 and the eyes are rounded. Maera

williamsi Bynum and Eox, 1977 was given erroneously as a

synonym of Anamaera hixoni Thomas and Barnard, 1985 by

Krapp-Schickel and Jarrett (2000) but has since been

recognised as distinct. Another species remains to be described.

The amphipod genus Linguimaera

259

Figure 1: Linguimaera sp. male (Adelaide, South Australia) SEM pictures. Above: head with thickened upper lip laterally; mandible with palp,

maxillae; left half of upper lip ventrally. Second line: head with antennae, upper lip, mandible with palp, maxillae and maxilliped ventrolaterally.

Below: head with first coxae and mouthparts from lateral and ventral; gnathopod 2 male.

260

Traudl Krapp-Schickel

Ceradomaera Ledoyer, 1973 also has asymmetrical second

gnathopods and differs mainly in the emarginate telson and

dorsal teeth.

Serrations on epimeral plate 3, both below and above the

posteroventral tooth, also occur in other species of Maera sensu

lato (e.g. Othomaera othonis (Milne-Edwards, 1830),

Quadrimaera serrata (Schellenberg, 1938), Maera tepuni

Barnard, 1972), but extra teeth defining an excavation on

epimeral plate 1 and (more clearly visible in) epimeral plate 2,

appear to be confined in the Indo-Pacific to Linguimaera, and

in the Atlantic to Anamaera and the Maera williamsi-c\di6&.

Barnard (1972a) cited also Maera othonopsis Schellenberg,

1938 in connection with the present species flock. It was

described with few figures on the basis of only two incomplete

ovigerous females from the Gilbert Is (Tropical West-Pacific)

as having subequal mandibular palp articles 2 and 3, quite dif-

ferent third uropods and telson (cf. Quadrimaera Ruffo and

Krapp-Schickel, 2000 or Mallacoota Barnard, 1972), and was

never found again. It certainly does not belong to Linguimaera.

The Indo-Pacific genus Linguimaera is well differentiated

from other genera of this region (Quadrimaera, Ceradocus

Costa, 1853, Elasmopus Costa, 1853, Maeracoota Myers,

1997, Mallacoota) by asymmetrical second gnathopods in both

sexes, a sinus on the posterodistal corner of the first and second

epimeral plates and a serrate posterior margin on the third

epimeral plate. It shares the asymmetry of the gnathopods with

the Atlantic Anamaera and the Indopacific Zygomaera Krapp-

Schickel, 2000, but differs mainly by characters of the

mandible (palp articles 2 and 3 subequal) and telson (in

Anamaera lobes cuspidate, in Zygomaera lobes partly

coalesced). The Maera sensu lato flock of Barnard and Barnard

(1983), containing 59 species, is now mostly unravelled, but

there are still a dozen species remaining in Maera sensu lato,

thereby stressing that they do not belong to the well-defined

Maera sensu stricto.

Key to genera similar to Linguimaera

1 . Gnathopod 2 asymmetrical 2

— Gnathopod 2 symmetrical 5

2. Telson fused halfways or more, distally only emarginate 3

— Telson deeply cleft 4

3. Metasome and urosome minutely toothed dorsally, beset

with plumose setae; mandibular palp article 3 as short as

article 1 Ceradomaera

— Metasome and urosome smooth; mandibular palp article 3

clearly longer than article 1 Zygomaera

4. Eyes round, mandibular palp article 2 equal to article 3,

distal robust setae of telson longer than telson length . . .

Anamaera

— Eyes reniform, mandibular palp article 3 shorter than arti-

cle 2, distal robust setae of telson shorter than telson length

Linguimaera

5. Dorsally carinate or toothed 6

— Dorsally smooth 8

6. Gnathopod 2 palmar corner rectangular, urosomites 1, 2

with dorsal teeth Maeracoota

— Gnathopod 2 palmar corner absent or less than 90° ... 7

7. Epimeron 3 smooth or carinate, urosome carinate

Mallacoota

— Epimeron 3 with serrations, urosome smooth

Parelasmopus

8. Mandibular palp article 3 falcate Elasmopus

— Mandibular palp article 3 linear 9

9. Mandibular palp article 1 distally tooth-shaped, lengthened

— Mandibular palp article 1 rounded 11

10. Gnathopod 2 dactylus outer margin densely setose;

maxillae not fully setose Maera

— Gnathopod 2 dactylus outer margin with 1 seta; maxilla 1

inner plate fully setose, maxilla 2 with oblique facial row

of setae Ceradocus

11. Pereopod 5 basis not longer than coxa 5; uropod 3 very

short, rami scarcely longer than peduncle .... Lupimaera

— Pereopod 5 basis longer than coxa 5; uropod 3 rami

clearly longer than peduncle 12

12. Gnathopods 2 without palmar corner, dactyli smooth;

uropod 3 lanceolate, medially widened, distally pointed .

Othomaera

— Gnathopod 2 distally widened, palmar corner well defined

13. Palmar corner subrectangular, pereopod 5-7 dactylus

simple, uropod 3 slightly unaequiramous .... Maeropsis

— Palmar corner rectangular, pereopods 5-7 with bifid

dactylus, uropod 3 aequiramous Quadrimaera

Key to species of Linguimaera

1. Telson distal robust setae longer than half telson length;

gnathopod 2 male palm transverse

L. garitima (300 m depth, Australia)

— Telson distal robust setae equal or shorter than half telson

length; gnathopod 2 male palm oblique 2

2. Uropod 3 ratio peduncle : rami greater or equal to 2.5 .3

— Uropod 3 ratio peduncle : rami less than 2.5 5

3. Longest distal robust seta on telson equal to half telson

length; male gnathopod 2 palmar comer thumb- shaped . .

L. leo (shallow water, Australia)

— Longest distal robust seta on telson shorter than half telson

length; male gnathopod 2 without thumb 4

4. Male gnathopod 2 palm J-shaped excavated, palmar corner

sharp L. kellissa (infralittoral, Australia)

— Male gnathopod 2 palm oblique, neither excavated nor

convex, without palmar corner

L. pirloti (infralittoral, Indonesia)

5. Male gnathopod 2 palm with deep U-shaped excavation,

dactylus strongly inwards bent

L. rnannarensis (no depth reported, India)

— Male gnathopod 2 palm with excavation, not U-shaped 6

6. Male gnathopod 2 propodus palm with V-shaped incision

near palmar corner; pereopods 5-7 very strong, pos-

terodistal corner lengthened and broadened, pereopod 7

propodus widened . . L. bogombogo (littoral, Micronesia)

— Male gnathopod 2 propodus palm with shallow semi-

circular excavation; pereopods 5-7 slender 7

The amphipod genus Linguimaera

261

7. Male gnathopod 1 propodus more than twice as long as

wide; pereopod 7 basis posterior margin straight, pos-

terodistal corner with right angle

L. tias (infralittoral, Australia and New Zealand)

— Male gnathopod 1 propodus twice as long as wide; pereo-

pod 7 basis posterior margin and posterodistal corner

rounded

. . .L. caesaris (littoral, eastern Mediterranean, Red Sea)

Linguimaera pirloti sp. nov.

Figure 2

Linguimaera othonides. — Pirlot, 1936: 309-311, fig. 132.

Material examined. Holotype. Sulawesi, Sailus ketjil, lies Paternoster,

27 m depth, corals and sand, 30, 31 Mar 1899 (Siboga Expedition stn

37), ZMA Amph. 204584 (1 male, 7.1mm on 2 slides).

Pai'atypes. Detroit de Molo, 54-90 m, sand, 19 Apr 1899 (Siboga

Expedition stn 51) ZMA Amph. 204585 (4 females, all on slides; 11

inadults in alcohol). 4°20'S, 122°58"E, sand and shells, 20 Sep 1899,

75-94 m (Siboga Expedition stn 204), ZMA Amph. 204586 (2 males,

1 adult female, partly on slides).

Diagnosis. Female gnathopod 1 propodus ratio of length :

width = 3, in male = 2; palm not defined, posterior margin

regular; palmar corner proximally followed by shallow excava-

tion. Gnathopod 2 in both sexes similar in shape, but asymmet-

rical; in female palmar excavations deeper. Pereopod 7 basis

ratio of length : width = 1.25. Telson with 2 or 3 distal robust

setae, maximum lengths half length of telson.

Description. Adult male and female 6-7.1 mm.

Head: lateral cephalic sinus antero ventral comer blunt,

nearly right-angled. Eyes with upper half narrower. Coxae 1-4

with small notch posterodistally.

Antenna 1 peduncle with 1 robust seta on article 1 distally;

peduncle article 1 subequal article 2; article 3 about one third

of article 1; accessory flagellum of 3-4 articles; antenna 2 slen-

der, gland cone short, peduncle reaching end of antenna 1

peduncle; article 4 longer than article 5, flagellum as long as

article 5, of about 12-14 articles.

Mandibular palp article 1 longer than wide; ratio of article 2

: article 1 = 3.6; article 2 : article 3 = 1.3; article 2 with 4 long

setae but no groups of setae along margin; article 3 with 8 long

setae. Mandibular incisor, lacinia mobilis and molar with

medium sized blunt teeth. Molar medium Labium with

rounded inner lobes, outer ones densely setose. Maxilla 1

unknown. Maxilla 2 outer plate wider than inner plate, outer

plate 8 robust setae only distally, no setae marginally.

Maxilliped unknown.

Gnathopod 1 weakly sexually dimorphic. Coxa 1 anteriorly

acutely produced. Basis ratio length : width = 3, anterior mar-

gin with 3 long setae, posteriorly more and longer ones; merus

posteroventrally with acute tooth; carpus with nearly parallel

margins, length to width about 2.5, with stiff marginal and sub-

marginal robust setae; propodus in female slim, narrower than

carpus, ratio length : widths 2.5-3.0, in male wider, twice as

long as wide, in both palm not defined.

Gnathopod 2 oi female slightly asymmetrical in size, simi-

lar in shape. Coxa 2 longer than wide, basis with few short

setae on anterior margin, many very long ones on posterior

margin; merus posterodistally with acute tooth; carpus pos-

teroventral corner rounded; length ratio carpus : propodus =

0.8, about same width; propodus slender, ratio length : width =

2. 5-3.0, palm concave, weakly defined by comer; 1 subdistal

prominent robust seta on the inner surface next to the palmar

corner, 2 smaller ones along the palm. Gnathopod 2 of male

dimorphic both in size and shape: the smaller is as described for

the female, the other has a longer, less excavated palm.

Dactylus not much curved.

Pereopods spinose, propodus longer than merus, carpus

shorter, length of basis : propodus = 1.3; basis posterior margin

serrate, posterodistal corner somewhat lengthened. Dactyli

forming a “chela” with their nail and the stiff, towards the

dactylus bent robust seta on inner margin.

Epimeral plates 1, 2 postero ventral corners acute, followed

by a shallow short sinus ending with a blunt tooth. Epimeral

plate 3 with up to 9 small teeth on posterior margin, in adult

males many irregular teeth, in juvenile specimens less; ventral

margin smooth.

Uropod 2 the shortest, uropod 3 extending much more than

uropod 1 ; uropod 1 peduncle longer than rami, 1 sub-basofacial

strong curved robust seta in about one third of length on outer

margin, 1 very long robust seta subdistally; inner ramus longer

than outer one. Uropod 2 distally with 2 longer and some

shorter robust setae. Uropod 3 ratio peduncle : rami less than

0.5, rami subequal, outer somewhat narrower than inner one; 1

article; both distally tapering, scarcely truncate, with 2-3 distal

robust setae, marginal ones on outer ramus in 3-4 groups, on

inner one many short single ones.

Telson, inner side shorter and outer corner acutely

prominent; in the excavation 2-3 strong robust setae (0.2-0. 5

of telson length).

Etymology. Dedicated to J.M. Pirlot.

Distribution. Sulawesi, Indonesia; coral rubble, shells, sand,

27-94 m.

Remarks. The “disproportionate” insertion between propodus

and carpus, which Barnard (1972a: 126) stressed for this

species flock, and which leads to a deeper “gap” dorsally in

other species, is here very scarcely visible.

Linguimaera bogombogo sp. nov.

Figures 3-4

Maera hamigera. — Barnard, 1965: 507-510, fig. 16.

IMaera species A. — Barnard, 1970: 158, fig. 98, 99

(not Maera hamigera Haswell, 1879a: 333, pi. 21 fig. 1)

Material examined. Holotype. Eniwetok Atoll, Bogombogo Island,

Micronesia, USNM 108926 (male, 4.3 mm).

Paratype. Eniwetok Atoll, Bogombogo Island, Micronesia, USNM

108926 (ovigerous female, 5.2 mm).

Other material. Eniwetok Atoll, Igurin I., lagoon side, dXgdiBryopsis

sp., attached to rocks, 27 Sep 1956, USNM 108928 (male 5 mm, drawn

by Barnard, 1965; 2 ovigerous females 5 mm, 4.3 mm, immature and

juvenile, incomplete). Igurin I., lagoon side, preserved rocks, 27 Sep

1956, USNM 108949 (1 juvenile). Igurin I., lagoon side, sand wash-

ings from under rocks, 27 Sep 1956, USNM 108927 (1 male? 3.5 mm.

The amphipod genus Linguimaera

265

1 specimen 3 mm, 3 juveniles). Boden L, ocean side, 12 Oct 1956,

USNM 108929 (1 specimen 3.3 mm)..

Diagnosis. Gnathopod 1 propodus rectangular, twice as long as

wide, palm oblique, straight. Gnathopod 2 male palm well

defined as acute tooth, followed by V-shaped incision; palm

with small shallow excavations. P7 rectangular, widened and

posterodistally lengthened, ratio length : width = 1.5. Telson

with 1 distal strong robust seta of about one third of telson

length.

Description. Ovigerous female 5-5.5 mm, male 4-5 mm.

Head about same length as first 2 body segments. Lateral

cephalic lobes rounded, with notch, anteroventral corner

rounded. Eyes narrowed in middle.

Antenna 1 about 0.6 body length, peduncle = flagellum,

peduncle article 1 shorter than article 2; flagellum of up to 25

articles, accessory flagellum of 3^ articles; antenna 2 slender,

gland cone reaching about half of peduncle article 3, article 4

longer than article 5, flagellum longer than peduncle article 5,

of 8 articles.

Mandibular palp article 1 clearly longer than wide; ratio

article 2 : article 3 = 1.4, article 2 with 4 long setae, article 3

distally 4 setae, laterally 2-3.

Gnathopod 1 not sexually dimorphic. Coxa 1 anterodistally

lengthened and pointed. Basis ratio length : width = 3, poster-

iorly 5-6 long setae. Merus posteroventrally rounded, charac-

teristical pointed tooth lacking, ratio length : breadth less than

2; carpus triangular, ratio length : breadth^ 2.3; propodus rec-

tangular, less broad than carpus, ratio length : breadth^ 2.3,

palm defined, oblique.

Gnathopod 2 oi female slightly dimorphic in size and shape.

Coxa 2 quadrangular, basis with few long setae on posterior

margin, merus posterodistally with tooth; carpus triangular,

ratio length : breadth^ 1.45, carpus: propodus = 0.55, both

about same width; propodus slender, similar shape to male, but

narrower; anterior and posterior margin parallel, palm defined

by prominent tooth and beset with strong robust setae, anterior

: posterior margin = 4:3. Smaller gnathopod 2 lacking promi-

nent defining tooth, longer and narrower. Gnathopod 2 of male

strongly dimorphic, carpus in larger gnathopod shorter, length

subequal to width. Both propodi similar to female, but larger

one more broadened and defining tooth as well as robust seta

sitting next to it more developed.

Pereopods 3, 4 similar in shape and size. Pereopods 5-7

robust, basis rounded, very small serrations on posterior

margins. Pereopod 6 the longest, in pereopod 7 the propodus

shortened and thickened. Dactylus with uneven outer and inner

margins, distally on inner one 1 stiff seta and one short and thin

next to it.

Epimeral plate 3 with small dense serration.

Uropod 1 peduncle inferior margin on proximal third with 1

strong robust seta, subequal rami shorter than peduncle; uropod

2 subequal rami shorter than peduncle; uropod 3 rami sub-

equal distally rounded, beset with many robust setae being

maximally of a quarter of ramuslength.

Telson longer than wide, lobes outer end longer than inner

one; in excavation 1 strong robust seta inserted, about one third

of telson length. Outer margin about halfways another, some-

what shorter robust seta, distally 1-2 fine setae, proximally a

stiff robust seta.

Etymology. Erom the island where the type specimens were

found (noun in apposition).

Distribution. Eniwetok Atoll (Micronesia); green algae

{Bryopsis, Caulerpa, Halimeda) and surrounding sand of rocky

intertidal; wash of old coral heads in about one-third metre of

water, together with amphipods Cymadusa filosa, Elasmopus

pseudaffinis, Gammaropsis digitatus, G. pacificus, Lembos

aequimanus, L. bryopsis, L. cf. intermedius, Quadrimaera ser-

rata, Mallacoota cf. insignis, Paragrubia vorax.

Discussion. This species clearly belongs to this clade, having

bean-shaped eyes, a (not very pronounced, but present) notched

cephalic lobe, mandibular palp article 3 much shorter than

article2, coxa 1 anteriorly acutely lengthened, gnathopod 1

carpus thickened, gnathopod 2 asymmetrical, epimeral plates

1-3 with characteristic excavations or serrations, uropod 3 rami

long and with many short robust setae, telsonic lobes distally

incised.

Linguimaera caesaris sp. nov.

Maera hamigera. — Walker, 1909: 335, pi. 43, fig. 5. — Karaman

and Ruffo, 1971; 152-158, figs 21-23.— Lyons and Myers, 1993: 587,

fig. 10.

(not Maera hamigera Haswell 1879a; 333, pi. 21, fig. 1)

Material examined. Holotype. Cesarea, Mediterranean coast of Israel,

MCNCr 425 (ovigerous female 8.5 mm).

Paratype. Same locality, MCNCr 1209-1212 (male 6 mm).

Diagnosis. Gnathopod 1 propodus rectangular, less broad than

carpus, 3 times as long as large. Gnathopod 2 male, female

propodus oval, palm one third of posterior margin, slightly

excavated and defined by blunt to rectangular corner. Pereopod

7 basis oval, posterior margin with fine serration. Telson with 1

bigger and 2 tiny robust setae distally, maximum length about

one third of telsonic length.

Description. Adult female 8.5 mm, male (immature?) 6 mm.

Head: lateral cephalic lobes rounded, anteroventral corner

rounded. Eyes more than twice as long as large, upper part

narrowed.

Antenna 1 peduncle scarcely longer than flagellum,

peduncle article 1 shorter than article 2; flagellum of up to 28

articles, accessory flagellum of 5 articles; antenna 2 slender,

gland cone short, article 4 longer than article 5, flagellum

subequal to peduncle article 5, of 9 articles.

Mandibular palp article 1 longer than wide; article 2 : article

3 = 2, both with long setae, article 3 also distally.

Coxa 1 anterodistally a bit upturned, bluntly pointed, pos-

terodistal comer with small notch. Basis ratio length : breadth

= 3, posteriorly 5 long setae. Merus posteroventrally rounded,

(sharp tooth lacking here), less than twice as long as wide;

carpus triangular, swollen; propodus rectangular, less broad

than carpus, about 3 times as long as wide, palm defined,

oblique.

Gnathopod 2 of female slightly dimorphic in size and shape.

Coxa 2 rectangular, basis with 9 long setae on posterior margin.

266

Traudl Krapp-Schickel

merus posterodistally with sharp tooth; carpus triangular, ratio

length : breadth = 1.5, carpus: propodus = 3:5, both about same

width. Propodus slender, similar shape to male, but shorter;

anterior : posterior margin = 5:3; palm defined by blunt cor-

ner beset with groups of robust setae, proximal part shallow

excavate, in distal third defined by strong robust seta sitting ele-

vated. The other gnathopod 2 lacks the palmar excavation as

well as the defining robust seta near dactylus insertion and is

narrower. Gnathopod 2 of male strongly dimorphic, carpus in

larger gnathopod shorter, length subequal to width. Both pro-

podi similar to female, but larger one broader, palmar defining

tooth well developed, hump defining the palmar excavation

more developed.

Pereopods 3, 4 very similar in shape and also size.

Pereopods 5-7 robust, basis rectangular, small serrations on

posterior margins; pereopod 5 small; pereopods 6, 7 subequal.

Dactylus distally on inner margin with 2 stiff setae bent to tip

of nail.

Epimeral plate 3 with serration of 4 or 5 teeth.

Uropod 1 peduncle inferior margin subproximally with 1

strong robust seta, subequal rami shorter than peduncle; uropod

2 with subequal rami as long as peduncle; uropod 3 peduncle

less than half length of rami, rami subequal, beset with many

short robust setae of maximally one seventh ramus length.

Telson longer or subequal to width, lobes outer end longer

than inner one; in excavation 1 strong robust seta inserted with

small additional ones, robust seta about one third of telson

length. On outer margin of first and second third, another short-

er robust seta.

Etymology. There are two reasons for the choice of this name:

at first sight, because the Mediterranean material (see Karaman

and Ruffo, 1971) comes from Cesarea (Israel), but more impor-

tantly it should remind of Sandro Ruffo, the grand old man and

“emperor” (= caesar) of amphipodologists.

Distribution. Suez Channel (Walker, 1909); Mediterranean

coast of Israel; Red Sea: Gulf of Aqaba (Karaman and Ruffo,

1971: 158; Lyons and Myers, 1993: 587-590); 4-5 m, coral

rubble (Lyons and Myers, 1993).

Discussion. Figures and description of Ledoyer (1982:

523-527) match perfectly with the ones given by Karaman and

Ruffo (1971), except the fact that the telson of the Madagascar

material shows on the inner side of the lobes some short robust

setae, which lack in the figures of Karaman and Ruffo (1971)

and the ones by Lyons and Myers (1993).

Linguimaera garitima sp. nov.

Figures 5-7

Material examined. Holotype. Australia. Tasmania, eastern Bass Strait,

82 km ENE of North Point, Elinders I. (39“27.7^S, 148°41.4T), 293 m,

coarse sand, naturalist’s dredge, G.C.B. Poore on HMAS Kimbla, 28

Mai- 1979 (stn BSS 36), NMV J52321 (1 male 7 mm).

Paratype. Collected with holotype, NMV J52322 (1 female 7 mm).

Other material. Collected with holotype, NMV J20371 (8 males, 16

females 17 juveniles).

Diagnosis. Similar to L. leo sp. nov. but: body smaller, articles

narrower, coxa 1 anterodistally more lengthened and anteriorly

excavated, gnathopod 1, 2 propodus rectangularly narrow, setae

long, gnathopod 2 male the smaller ones similar to gnathopod

2 female in totally lacking palmar tooth; pereopod 7 basis about

twice as long as wide. Telson about as long as wide, distal

strong robust seta between half and two thirds of telsonlength.

Description. Adult male, female 5.5-8 mm.

Head: lateral cephalic lobes rounded, antero ventral corner

rectangular. Eyes more than twice as long as large, upper part

narrowed.

Antenna 1 peduncle scarcely longer than flagellum, pedun-

cle article 1 shorter than article 2; flagellum of up to 34 articles,

accessory flagellum of 6 articles; antenna 2 gland cone short,

article 4 longer than article 5, flagellum subequal to peduncle

article 5, of 9 articles.

Mandibular palp article 1 longer than wide; article 2 : article

3 = 1.6, both with long setae, article 3 also distally.

Coxa 1 anterodistally acute and anteriorly excavated;

gnathopod 1 propodus narrow, more than twice as long as wide.

Gnathopod 2 of female with narrow and long propodus

lacking a defining tooth on the palmar comer, with palm

oblique, crenulate. Gnathopod 2 of male propodus palm almost

transverse, blunt robust defining tooth on (about rectangular)

palmar corner; ratio of propodi gnathopod 1 : gnathopod 2 =

0.5.

Pereopods 3, 4 merus not widened; female pereopod basis

narrower.

Uropod 1 peduncle as long as rami.. Uropod 3 rami with

thinner robust setae.

Telsonic lobes with long robust seta which is clearly longer

than half telsonlength.

Etymology. During my stay at Museum Victoria, Melbourne,

Gary Poore and Tim O’Hara were extremely helpful on many

occasions, and furthermore were strongly involved in the col-

lection of the material. The specific epithet is a combination of

their names and expresses my gratitude; used as an adjective.

Distribution. Bass Strait, coarse shell, 293 m.

Discussion. This clade seems strikingly conservative and

though checking very thoroughly, there are not many easy char-

acters separating this species from the much shallower L. leo (if

not dealing with mature males), except: much greater depth,

smaller body size, mandibular palp ratio aiticle 2 to article 3

longer (1.65 vs 1.8 in L. leo), palmar corner of male gnathopod

2 with about right angle (vs oblique) and (most reliable charac-

ter) a long robust seta on telsonic lobes (much shorter in L. leo).

Linguimaera kellissa sp. nov.

Figures 8, 9

Material examined. Holotype. Australia. Victoria, eastern Bass Strait,

8 km S of South East Point, Wilsons Promontory (39°12.9'S,

146°27.3'E), 65 m, medium sand, R.S. Wilson on RV Tangaroa, 18

Nov 1981 (stn BSS 180), NMV J20370 (1 male 5 mm).

Paratype. Vic., eastern Bass Strait, 11.2 km E of eastern edge of

Lake Tyers (37°51.4US, 148°13.16'E), 32 m, sand-shell, Sniith-

Mclntyre grab. Marine Science Laboratories, 25 Sep 1990 (stn MSL-

EG 27), NMV J25482 (1 male 7 mm).

Other material. Vic., eastern Bass Strait, 8 km S of South East Point,

The amphipod genus Linguimaera

267

Figure 5. Linguimaera garitima sp. nov. (Tasmania). Habitus in scale = 1 mm; Gnl, Gn2 male large and Ep3 in scale = 0.4 mm; Gn2" male large

in scale = 0.2 mm.

The amphipod genus Linguimaera

271

Wilsons Promontory (39°12.9'S, 146°27.3T), 65 m, NMV J52344 (5

males, 8 females, 2 juv.); NMV J52343 (1 male). 13.3 km E of eastern

edge of Lake Tyers (37°51.74'S, 148N4.77T), 37 m, NMV J25491 (1

female). 15.5 km SW of Pt Ricardo (37°53.14^S, 148°28.94^E), 45 m,

NMV J25478 (4), NMV J25486 (1 male, 1 juv.), NMV J25485 (1 male,

1 female).

Diagnosis. Gnathopod 1 propodus and carpus relatively robust,

propodus length in male twice of width; gnathopod 2 male

palm distally next to palmar corner regularly J-shaped excav-

ated, in female distally narrowing, with serrated palm.

Pereopods 3-5 length subequal. Pereopod 7 basis ratio length :

breadth^ 1.5, posterior margin rounded. Uropod 3 very long,

slim, rami about twice length of peduncle, always having a

“knee” between peduncle and rami. Telson with short distal

robust setae, distal one about one third to half of telson length.

Description. Adult male, female 7-9 mm.

Body slim, slender, fragile. Eyes reniform, medially

narrowed, lower part longer and wider.

Antenna 1 half length of body, article 1 slightly shorter than

article 2, accessory flagellum of 5 articles; antenna 2 gland

cone long; peduncle reaches end of antenna peduncle article 2;

flagellum of 11 articles.

Mandible incisor and lacinia mobilis with strong blunt teeth,

accessory robust setae smooth; molar huge and oblong; palp

article 2: article 1= 3.75, article 2 : article 3 = 1.8, article 2 with

9 single long setae (some plumose) without groups.

Gnathopod 1 coxa 1 anteriorly lengthened, rounded, basis

ratio length : breadth^ 2.8; merus posterodistally acutely point-

ed; carpus rounded, ratio length : breadth^ 2; propodus palm

oblique, scarcely defined.

Gnathopod 2 of female propodus ratio length : breadth = 2.6,

palm nearly straight, with many shallow excavations about one

272

Traudl Krapp-Schickel

third of total length, corner not defined. Gnathopod 2 of male

propodus rectangular, about twice as long as wide, palm in the

shape of a question mark or J, well defined by a prominent

acute tooth, next to it palm distally smoothly excavated, no ser-

rations, proximally ending in a rounded hump, beset with many

short robust setae.

Pereopods basis in female slender, in male robust, in

pereopod 7 basis ratio length : breadth = 1.5, posterior margin

rounded, only very small serrations.

Epimeral plate 1 shorter than epimeral plate 2; epimeral

plates 1, 2 posterodistal comer with scarcely visible very shal-

low excavation and second tooth. Epimeral plate 3 with few but

strong teeth distally.

Uropod 3 very characteristic, always having a “knee”

between peduncle and rami; peduncle ratio length : breadth =

2.2, outer ramus ratio length : breadth = 6.75, with many robust

setae marginally and distally in 8-9 groups, apically long setae,

that are easily lost.

Telson with 1 strong short robust seta distally, less than half

telson length, accompanied by 1 other, of half length, on each

side; 1 plumose seta on the outer margin distally, 2 robust setae

and 1 seta marginally.

Etymology. Dedicated to Kelly Merrin and Melissa Storey with

whom I shared the lab at Museum Victoria and who at any time

were helpful and friendly “daughters” to their guest!

Distribution. Bass Strait, Australia, muddy, medium to coarse

sand, sand-shell, 33-65 m.

Linguimaera leo sp. nov.

Eigures 10-12

Maera rnastersi. — Barnard, 1972a: 226-227, fig. 132.

{not Megamoera mastersii Haswell, 1879b: 265, pi. 11 fig. 1)

Material examined. Holotype. Australia. Victoria, Port Phillip Bay,

Prince George Light (38°6.3"S, 144°44.25T), 9.6 m, silty sand with

broken rock, SCUBA, Fisheries and Wildlife Dept and Museum, (stn

PPS 10), NMV J35851 (1 male 12 mm).

Paratypes. Collected with holotype NMV J52309 (1 male, 2 juve-

niles, 11.8, 6-7 mm). Vic., Portland Bay, reef below lighthouse

(38°22'S, 14r36.2'E), 3 m, sand and rubble, SCUBA airlift, R.S.

Wilson, 26 Feb 1992 (stn CRUST 141); NMV J24121 (1 female, 15

mm). Western Port, off Crib Point (38°20.83'S, 145°13.49'E), 13 m,

sandy gravel, Smith-Mcintyre gi'ab, A.J. Gilmour on EV Melita, 23

Mar 1965 (stn CPBS-N 32); NMV J48856 (more than 20 males,

females).

Other material. Numerous specimens in 61 NMV collections from

Vic. (Western Port, Port Phillip Bay, Cheviot Beach, Point Nepean,

Bass Strait) and SA (Cape Northumberland, Wallaroo), 0-26 m, algal

and sedimentary substrates. Port Phillip Bay (stn PPS 47 Area 40),

USNM 275759 (1 male 12 mm, 1 female 10.5 mm, 2 male ?juvenile

10 mm); (stn PPS 83 Ai'ea 69), USNM 275759 (6 males 8-10.5 mm, 3

females, 7.5-8 mm).

Diagnosis. Gnathopod 1 propodus rectangular, ratio length :

width = 2. Gnathopod 2 of male adult with prominent stout

hump on palmar-comer, distally followed by a small incision;

palm convex, with 3 incisions; dactylus near insertion not fit-

ting totally to palm, leaving a hole-shaped gap. Pereopod 7

basis ratio length : width = 1.45. Telson with 1 long distal

robust seta, between half and total telson length, and 3 short

ones. (Pereopods 5-7 of ov. female strikingly twisted in

articulation between merus and ischium.)

Description. Adult female 8-15 mm, male 7-12 mm.

Head nearly as long as first 2 segments, anteroventral comer

acute. Eyes medially narrowed.

Antenna 1 about three fifths of body, peduncle article 1 sub-

equal to article 2; flagellum of up to 30 articles, accessory

flagellum of 3 or 4 articles; antenna 2 gland cone reaching half

of article 3; flagellum of about 12 articles.

Mandible incisor, lacinia mobilis and molar medium; ratio

palp article 2 : article 1 = 3; article 2: article 3= 1.8; palp

article 2 densely setose with 5-6 groups. Maxilla 1 inner plate

narrower than outer plate, oval, with 3 plumose robust setae;

outer plate 6 simple to pectinate robust setae, about twice as

long as large, palp article 1 quadrangular, article 2 twice as long

as large, 8 robust setae only apically. Maxilla 2 outer and inner

plates equal, robust setae only dis-tally, no fine hairs marginal-

ly. Maxilliped inner plate reaching one third of palp article 2,

apically truncate with dense distal and a few lateral robust

setae; outer plate large, oval, reaching two thirds of article 2 of

palp, with curved robust setae, gradually increasing in length

from inner to outer side; palp article 1 shorter than one third of

article 2, article 3 half article 2, oval.

Gnathopod 1 not sexually dimorphic; coxa 1 anteriorly

acutely lengthened; basis anterior margin with 4 or 5 medial

robust setae, posteriorly longer robust setae; mems postero-

ventrally with short tooth; carpus regularly rounded on both

margins, with stiff marginal and submarginal robust setae,

about twice as long as wide; propodus rectangular, less broad-

ened than carpus, ratio length ; breadth = 2, palm oblique, well

defined by blunt corner.

Gnathopod 2 of female strongly dimorphic in size, similar

in shape; merus posterodistally with blunt tooth; carpus pos-

teroventral corner with sharp tooth; carpus : propodus = 2:3,

about same width; propodus rectangular, twice as long as

wide, palm scarcely concave, scarcely defined by corner,

no posterodistal “thumb”; 1 subdistal prominent robust

seta on inner surface next to palmar corner, 6 smaller ones

along palm. Gnathopod 2 of male strongly dimorphic in size

and shape: the smaller is as described for smaller female

gnathopod 2, the other has a palmar corner defined by a blunt

and prominent “thumb”, distally followed by a short exca-

vation, palm with rounded hump having 3 short excavations;

dactylus strongly curved, leaving a hole- shaped gap near

insertion.

Pereopods 3, 4 shape very similar, pereopod 3 reaching dis-

tally about half of gnathopod 2 propodus. Pereopods 5-7 simi-

lar, very spinose, on both margins serrate, propodus equal to

merus, carpus shorter, basis : propodus =1.7; pereopod 5 basis

posterior margin straight to concave and weakly serrate,

posterodistal corner broadened and lengthened; pereopod 6

subequal or somewhat longer pereopod 7, posteroventral comer

lengthened.

rn2 1 large male,

le in scale y = 1

276

Traudl Krapp-Schickel

Uropods 1, 2 ending at similar level; uropod 2 inner margin

with 2 strong robust setae; uropod 3 peduncle half length of

rami, outer ramus nan'ower than inner one; distally clearly

truncate, with marginal robust setae on outer ramus in 7 or 8

groups.

Epimeral plate 3 seiTation with 7 or 8 teeth.

Telson quadrangular, 1 or 2 setae and no or 1 strong robust

seta marginally; each lobe distally excavated as U-shape, with

outer corner acutely prominent; in excavation 1 long robust seta

(half or more telson-length) and 3 additional ones, not much

surpassing length of the sinus.

Etymology. Dedicated to little Leo, the “most beautiful grand-

son in the world”! (noun in apposition).

Distribution. Victoria (Port Phillip Bay, Western Port, Portland

Bay), South Australia (Wallaroo), Western Australia (Bunbury,

Favourite Is, Point Peron) (J.L. Barnard, 1972a); littoral,

gravel pools, under rocks and holdfasts on reef, sand and

rubble, sandy gravel; occun'ed in 56 samples with depth

average of 12 m; most robust adults in intertidal.

Remarks. Already Barnard (1972a: 226) noticed the somewhat

“disproportionate” insertion between propodus and carpus of

gnathopod 1, which leads to a deeper “gap” dorsally.

Discussion. This species is found sometimes together with

L. tias sp. nov. and has for more than a century been mixed up

with other similar species. The above cited bibliography there-

fore contains only the proven citation by Barnard (1972a).

There are several records of “M. mastersi (Haswell)”, mainly

by Chevreux (1908: 481 - French Polynesia); Stebbing (1910:

457 - South Africa); Chilton (1916: 367 - New Zealand);

Chilton (1925: 317 - Chatham Islands, New Zealand). But

there is not enough information to make a conclusion.

Linguimaera aff. leo

Material examined: Numerous specimens in 1 8 NMV collections from

eastern and central Bass Strait, 13-60 m depth, and Western Port,

Victoria; sandy sediments.

Remarks. There are robust specimens of 10-15 mm in the first

10 metres of Port Phillip Bay, with oblique to nearly transverse

gnathopod 2 palm in male, and robust ovigerous females with

characteristically upwards twisted pereopods. Below 10 m

down to about 45 m or even 60 m, in coarse sands or sandy

shells of southern and eastern Bass Strait, there is a population

of smaller and more delicate specimens, adult males with well

developed penis papillae never reaching more than 8 mm, and

ovigerous females of 6-7 mm. Their legs (especially gnathopod

1 male , pereopods 5-7 basis) are more slender, the setation on

gnathopods 1, 2 basis or uropod 3 is richer and some distal

robust setae are very long. But pereopods are mostly missing,

robust setae are easily broken or setae lost, and there is no

obvious and clearly reliable morphological difference to offer

at the moment, to allow defining it as a separate species, and all

differences found may also occur in less adult specimens of

Linguimaera leo. But I mistrust that Linguimaera leo could

have such a wide depth range, and also the ecology is quite

different.

Linguimaera mannarensis (Sivaprakasam)

Ceradocoides chiltoni. — Sivaprakasam, 1968a: 109-111, fig. 11

(not Ceradocoides chiltoni Nicholls, 1938).

Maera mannarensis Sivaprakasam, 1968b: 274-278, figs 1-2.

Maera mastersi. — Sivaprakasam, 1970; 36, fig. 1 a-g.

{not Megarnoera mastersii Haswell, 1879a; 265, pi. 11, fig. 1).

Type locality. Gulf of Mannar, India.

Diagnosis. Gnathopod 1 propodus medially widened, 2.4 times

longer than wide, palm oblique, straight; gnathopod 2 male

strongly asymmetrical, larger propodus pyriforai, with rectan-

gular hump near dactylus insertion and deep U-shaped incision,

defined by a sharp tooth; carpus triangular, shorter than broad;

pereopod 7 basis rounded, posterior margin serrated, propodus

posterior margin with 3 groups of long robust setae and a fourth

posterodistally; uropod 3 rami twice as long as peduncle, api-

cally truncate, richly beset with robust setae. Telson with long

apical robust seta, length twice the depth of incision of lobes

and more than one third of telson length, with 2 strong robust

setae mediolaterally.

Description. Adults 8-9 mm.

Lateral cephalic lobes rounded, with notch, anteroventral

comer rounded. Eyes inferior part a bit widened, medially not

narrowed.

Antenna 1 about 0.6 of body length; peduncle as long as

flagellum, peduncle article 1 shorter than article 2; flagellum of

26-29 articles, accessory flagellum of 4-5 articles; antenna 2

slender, gland cone short, peduncle article 4 shorter than article

5, flagellum longer than article 5, of 11 articles.

Mandibular palp article 1 longer than wide; ratio article 2 :

article 3 = 1.2 (thus article 3 relatively long compared to other

species); both with long setae, especially many dense on article

3 distally.

Gnathopod 1 sexual dimorphism not found. Coxa 1

anterodistally a bit upturned, bluntly pointed; basis ratio length

: breadth = 3, posteriorly 5 long setae and some shorter ones;

merus posteroventrally rounded (sharp tooth lacking here),

twice as long as wide; carpus triangular, swollen; propodus

slender, less broad than carpus, 3 times as long as wide, palmar

comer scarcely defined.

Gnathopod 2 male strongly dimorphic, carpus in larger

gnathopod short, shorter than broad; palmar corner well devel-

oped, with upturned point, followed by U-shaped incision

defined by a straight blunt distal elevation of palm; no espec-

ially prominent robust seta except some submarginal along

palm; dactylus strongly curved, the bend being stronger than

the outline of propodus, thus inwards folded.

Pereopods 3, 4 very similar in shape and also size; pereo-

pods 5, 6 robust, basis rectangular, small serrations on poster-

ior margins, posterodistal corner slightly lengthened and not

widened; pereopod 7 basis much larger, posterior margin

rounded, serrated; robust setae on posterodistal comer of

carpus reaching or surpassing half length of propodus.

Epimeral plate 3 with semation of 3 or 4 teeth.

Uropod 1 peduncle inferior margin subproximally with 1

strong robust seta, rami shorter than peduncle, outer a bit

shorter; uropod 2 outer ramus as long as peduncle, inner a bit

The amphipod genus Linguimaera

277

longer; uropod 3 much longer than uropods 1 and 2, rami

subequal, truncate, beset with many short robust setae of

maximally one fifth ramus length.

Telson longer than wide, lobes outer end longer than inner

one; in excavation 1 strong robust seta inserted (one third of

telson length) without small additional ones. Outer margin

medially and on proximal third, 1 other, shorter robust seta.

Distribution. Gulf of Mannar, India; seaweeds.

Discussion. I have not seen this species, thus the description

relies on Sivaprakasam's papers. According to them, this

species differs from all other species by the the inwardly-bent

dactylus and cup-shaped short carpus on the larger male

gnathopod 2, the widened and serrate basis of pereopod 7 with

long, rich setation on other articles and a relatively long

mandibular palp article 3.

Linguimaera tias sp. nov.

Figures 13-15

Maera mastersi. — Barnard, 1972b: 108-10, figs 55-56. — Sheard,

1936: 177-178 fig. 3.— Sheard, 1937: 24.

Maera mastersii. — Hale, 1929: 215, fig. 213. — Chilton, 1916:

367. — Chilton, 1925: 317. — Hurley, 1954: 603. — ^Lowry and Fenwick,

1983: 236.

?Moera mastersi. — Chilton, 1911: 564. — Chilton, 1921: 72-73.

(not Megarnoera mastersii Haswell, 1879b: 265, pi. 1 fig. 1).

Material examined. Holotype. New Zealand, Otago Harbour, Shelly

Beach, gravel pools, USNM 149478 (male 11.2 mm).

Paratype. Locality like above, USNM 149478 (ovigerous female

9.9 mm).

Other material. Numerous specimens in 39 NMV collections from

Vic. (Western Port, Port Phillip Bay, Portland Bay), SA (Cape

Northumberland), and eastern and central Bass Strait, 0-40 m depth,

sedimentary and algal substrates.

Diagnosis. Gnathopod 1 propodus ratio length ; breadth =

2. 1-2.7, changing with age. Gnathopod 2 male, female propo-

dus palm excavated, palmar corner in male without “thumb”-

shaped prolongation; gnathopod 2 female similar in shape and

not much different in size, slender. Pereopod 7 basis ratio

length : breadth = 1.75. Telson with apical robust setae between

half and total telson length.

Description. Adult male 10-17 mm, female 10-13 mm.

Eyes reniform, medially narrowed.

Antenna 1 0.8 of body length, peduncle article 1 shorter than

article 2; flagellum of up to 46 articles, accessory flagellum of

6-7 articles. Antenna 2 gland cone nearly reaching end of

article 3; peduncle reaches half of antenna 1 peduncle article 2;

flagellum of 16-17 articles.

Mandible incisor and lacinia mobilis with strong blunt teeth,

accessory robust setae serrate; molar huge and oblong; palp

article 2: article 1 = 2.25, article 2 : article 3 = 1.3, article 2 with

12-13 long setae in 4-5 groups; maxilla 1 inner plate width

subequal to outer plate; outer plate with 7-8 simple to pectinate

robust setae; maxilla 2 setae only distal, but many fine hairs

also marginally.

Gnathopod 1 weakly sexually dimorphic. Coxa 1 anteriorly

bluntly lengthened; basis ratio length : breadth = 2.5; merus

posteroventally bluntly lengthened; carpus regularly rounded

on posterior margin, length about twice to 2.3 width; propodus

palm oblique, scarcely defined.

Gnathopod 2 weakly sexually dimorphic, different in size,

not in shape, subchelate.

Female slightly dimorphic in size, similar in shape. Coxa 2

quadrangular, merus posterodistally with sharp tooth; propodus

palm concave with blunt hump medially, defined by a pos-

terodistal tooth, a straight part distally and shallow exca-

vation proximally; 1 subdistal prominent robust seta on

inner surface next to palmar corner, 6 smaller ones along

the palm. Male dimorphic in size and shape, but in hyper-

adults both gnathopods again similar in size and shape; when

dimorphic, one is as described for female, the other has a

stronger defined palmar corner, distally followed by

deeper semicircular excavation, while straight distal half of

female has 1 or more blunt humps medially; dactylus

stout, curved.

Pereopod 3 not much shorter than gnathopod 2 in male;

pereopods 3, 4 basis and merus strong, other articles slim.

Pereopods 5, 6 basis : propodus = 1.5; pereopod 7 male ratio

length : width =1.7.

Uropod 3 marginal setae on outer ramus in 4 or 5 groups.

Telson distomarginally with 2 small additional setae, no

robust seta; in the excavation of lobes, 1 robust seta of about

half telson length and another of one third telson length.

Etymology. Dedicated to our newest family member and son-

in-law Matthias, shortened to Tias (noun in apposition).

Discussion. This species is very similar to Linguimaera

mannarensis (Sivaprakasam, 1968). Differences are: eyes

oval, width medially narrowing (vs width not narrowing);

lateral cephalic lobe anterodistal corner pointed and curved

(vs very little developed, rounded); mandibular palp article 3

distally oblique (vs regularly rounded); gnathopod 1 propodus

twice as long as wide (vs longer and narrower); gnathopod

2 carpus longer than wide (vs wider than long); pereopod

7 basis posterior margin only very weakly rounded (vs

evenly excavate); telson subquadrate (vs longer than wide);

marginally on first third no robust seta (vs. one stout robust

seta).

Remarks. Thomson (1882; 235, fig. 4a) illustrated a New

Zealand amphipod as Moera quadrimana with characters sim-

ilar to the present species, although his fig. 4b probably deals

with the true Quadrimaera quadrimana (Dana).

Distribution. New Zealand: Otago Harbour, Shelly Beach

(Barnard, 1972b). Australia: Victoria: Port Phillip Bay, Western

Port, Portland Bay, Cape Northumberland, Bass Strait. South

Australia: Sellicks Beach (Sheard, 1936). Gravel pools, sand,

silty clay; 3.5^0 m depth.

Linguimaera sp.

Figure 16

Materia] examined. South-western Bass Strait (39°32.8'S, 144° 1 6 U),

18 m, 1 Nov 1980, fine sand, epibenthic sled, G.C.B. Poore on FV

Sarda (stn BSS 107), NMV J 2505 (2 males 10 mm).

282

Traudl Krapp-Schickel

Diagnosis. Length 10 mm. Coxa 1 anteriorly acutely produced.

Gnathopod 1 propodus slender, carpus much longer than

propodus, widened. Gnathopod 2 male palm slightly excavated

and crenulated. Pereopods 5, 6 basis rectangular, pereopod 7

posterodistal corner rounded, lengthened. Uropod 3 long, slim.

Telson with short distal robust setae that are scarcely exceeding

tip; 4 robust setae along inner margin of telsonic lobes.

Distribution. Bass strait.

Remarks. This material is very similar to L. leo sp. nov. and

L. tias sp. nov. but the robust setae on the inner margin of tel-

sonic lobes (naked in all other species) seem to be a good

character to distinguish this species within the group. Not a

large animal, in many respects these specimens seem more

slender than L. leo : antenna 2 peduncle article 4, gnathopod 1

propodus and carpus, and gnathopod 2 dactylus, propodus are

all narrower than in L. leo. Linguimaera sp. shares the narrow

articles of appendages with L. tias, and also the rich setation on

the posterior margin of the basis of gnathopods and the

relatively long accessory flagellum, but again the spination of

the telson is much different. Most probably these species have

different ecological niches. The present material is too poor for

defining a new species.

Megamoera thomsoni Miers, species dubia

Megamoera thomsoni Miers, 1884: 318, pi. 34, fig. B.

Remarks. Miers’ description of his Australian Megamoera

thomsoni could apply to a species of Linguimaera (especially

the slim gnathopod 1, propodus without palmar comer, carpus

very long would fit L. pirloti). But coxa 1 is definitely rounded

anteriorly in Miers’ species (vs very acute), the serrated exca-

vation on gnathopod 2 palm could fit some of the described

species, while the telson is figured very differently as densely

beset with robust setae on the inner margin of the lobes, and

apically without any incision. Thus, as the type material is

apparently lost and the description short, Megamoera thomsoni

Miers seems to be related to Linguimaera and may be even to

L. pirloti, kellissa, young L. leo or the unnamed species, but

should be considered as species dubia.

Maera aequimana Ledoyer, 1979, species dubia

Maera aequimana Ledoyer, 1979: 77-78, fig. 43.

Remarks. The figures of Ledoyer (1979) match well with

species of Linguimaera. However, it is stressed that the second

male gnathopods are not asymmetrical (therefore the name - it

may be an immature specimen?) and we don’t know the shape

of the third uropods. In the slide of the holotype the telson is

broken in pieces, so it is not clear how far it is cleft, while the

remaining material in alcohol (1 male, 1 female, 1 immature) is

not available. For the time being this species must remain

dubious.

Conclusions

The genus Linguimaera has an Indo-Pacific distribution. It

shows close relationship to Zygomaera, but the latter has an

uncleft and more or less emarginate telson, while that of

Linguimaera has a constant and quite characteristic structure in

being deeply cleft with the tips of the lobes asymmetrically

incised. The two genera share many other characters, such as

the produced anterodistal corner of coxa 1, the thickened car-

pus of gnathopod 1, dimorphic gnathopods 2 in males (at least

known in two species of Zygomaera), a shallow excavation on

the posterodistal corner of epimeral plates 1 and 2, a serrate

posterior margin of epimeral plate 3, and uropod 3 with a short

peduncle and long rami with many short robust setae that are

never longer than the rami. The differences with Zygomaera

seem mainly to reside in the shape of the eyes (in Zygomaera

rounded, often scarcely visible) and of the telson, but also in the

always truncated tip of uropod 3 rami, which show in some

species a minute second article. The last article of the mandibu-

lar palp is in Zygomaera (where known) only a little shorter or

subequal to the second article (always clearly shorter in

Linguimaera) and the falcate interramal robust seta distally on

the peduncle uropod 1 is in Zygomaera strikingly strong and

even on a special peduncle (vs less striking). While members of

Zygomaera are not all described and known with all their cru-

cial character states (and it might be that the emarginate telson

is homoplastic), members of the new genus Linguimaera seem

to form a natural group.

Acknowledgements

I am grateful to my friends Wim Vader, Joergen Berge (Troms0,

Norway) and Sandro Minelli (Padova, Italy), the latter also

member of the committee of ICZN (International Commission

for Zoological Nomenclature), for helping me out in nomen-

clatory questions. Many thanks to Gary Poore and his helpful

crew for offering me a desk, equipment and time for discus-

sions at then' museum in Melbourne, and to John Moverley

who for the second time took me into his house and family. Jim

Lowry (Australian Museum, Sydney) most kindly sent me the

draft of his Australian melitid paper before finishing my manu-

script. I am obliged to the Deutsche Forschungsgemeinschaft

who provided partial support for travel expenses for my last

two stays in Australia. Dirk Platvoet (Museum Amsterdam,

Netherlands) kindly provided the SEM pictures. Like always,

Wim Vader revised the manuscript and replied with patience to

my reference-queries. Careful critical reading of the submitted

manuscript was finally done by Jim Lowry, Alan Myers and last

but not least Gary Poore.

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Memoirs of Museum Victoria 60(2): 285-307 (2003)

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

http://www.museum.vic.gov.au/memoirs

Four new species of Ischnomesidae (Crustacea: Isopoda: Asellota) from off

south-eastern Australia

Kelly L. Merrin'’^ and Gary C. B. Poore'

' Museum Victoria, GPO Box 666E, Melbourne, Vic. 3001, Australia

^ Department of Zoology, The University of Melbourne, Vic. 3010, Australia; present address: Marine Biodiversity and

Systematics, National Institute of Water and Atmospheric Research, PO Box 14-901, Kilbirnie, Wellington, New Zealand

(k.merrin@niwa.co.nz; gpoore@museum.vic.gov.au)

Abstract Merrin, K.L. and Poore, G.C.B. 2003. Four new species of Ischnomesidae (Cmstacea: Isopoda: Asellota) from off south-

eastern Australia. Memoirs of Museum Victoria 62(2): 285-307.

Four new species of Ischnomeidae Hansen, 1916 are described from the continental slope of south-eastern Australia:

Haplomesus franklinae, Ischnomesus tasmanensis, I. justi and Stylomesus sarsi. The genera Haplomesus and Stylomesus

are recorded from Australain waters for the first time. The family and three genera are rediagnosed to accommodate the

new species.

Keywords Cmstacea, Isopoda, Ischnomesidae, Haplomesus, Ischnomesus, Stylomesus, deep-sea, Australia

Introduction

The asellote isopod family Ischnomesidae is distributed

throughout the world’s deep oceans and continental slopes with

28% of the 94 described species known from the Southern

Hemisphere. Little is known of its diversity in Australia and

previous to this study only two species had been described from

these waters. These, Ischnomesus anacanthus Wolff, 1962 and

Mixomesus pellucides Wolff, 1962, were collected from the

Tasman Sea during the voyage of the Danish research vessel

Galathea during the early 1950s (Wolff, 1962). More recently,

Poore et al. (1994) reported on the distribution patterns of a rich

fauna of continental slope isopods from south-eastern

Australia, western Tasman Sea and Bass Strait. These collec-

tions, made between 1979 and 1988, are part of Museum

Victoria’s SLOPE program. Among this abundant material

were about 13 undescribed ischnomesid species. This paper

describes four of these, now lodged at Museum Victoria,

Melbourne (NMV) and supplementary specimens from the

Australian Museum, Sydney (AM).

Due to their brittle nature, few specimens are intact and sev-

eral individuals were illustrated to complete the sets of figures.

The left side was dissected and drawn in preference, except

when an appendage was damaged or missing. If both sexes

were available, the most complete specimen was designated

holotype and pleopods from the opposite sex were included in

the description. Pereopod 1 is drawn figured for male and

female as it is sexually dimorphic, being larger and more robust

in females. Due to the rarity of material representing both sexes

in ischnomesid samples, there is no consistency concerning the

sex of the holotype in the literature.

Ischnomesidae Hansen

Ischnomesini Hansen, 1916: 54. — ^Wolff, 1956: 86.

Ischnomesidae. — Gurjanova, 1932: 40. — Menzies, 1962: 111. —

Wolff, 1962: 71-73.— Birstein, 1971: 198-199.— Menzies and

George, 1972: 971.— Chardy, 1974: 1537.— Kussakin, 1988: 418.

Diagnosis. Body elongate, subcylindrical and narrow.

Pereonites 4-5 elongate, pereonite 5 longest, at least twice as

long as wide, pereonite 4 widest anteriorly, 5 widest posterior-

ly. Head fused to and embedded in pereonite 1. Pereonites 1

(posterior margin) to 4 free and articulating. Pleon with maxi-

mum of 2 free pleonites plus pleotelson. Anus separated from

branchial chamber. Eyes absent. Antenna 1 terminating with

simple setae, article 1 squat and globular, article 2 elongate, at

least twice as long as article 1. Antenna 2 length more than half

body length, without squama. Maxilla 2 inferior margin with 2

medial pectinate setae. Pereopod 1 robust, strongly subchelate

and haptorial; pereopods 2-7 ambulatory; dactylus with single

unguis and 2 simple setae. Pleopod 3 endopod with 3 distal

plumose setae, exopod tapering, shorter than endopod, with

long distal plumose seta. Pleopod 4 unadorned, simple lobe,

exopod absent; pleopod 5 absent. Uropod uniramous, terminal.

In females, operculum wider at midpoint than proximally.

Remarks. This family is unique amongst the asellotes in having

the fifth pereonite at least twice as long as wide and the head

completely fused to the first pereonite, with only a slight

286

K. L. Merrin and G. C. B. Poore

indentation between the somites, and medial pectinate setae on

the inferior margin of maxilla 2. This paper acknowledges the

often overlooked generic synonymies of Kussakin (1988).

Haplomesus Richardson

Haplomesus Richardson, 1908: 81. — Hansen, 1916: 59. —

Gurjanova, 1932: 42. — Birstein, 1960: 6. — Menzies, 1962: 117. —

Wolff, 1962: 86. — Birstein, 1963: 59. — Birstein, 1971: 209. — Menzies

and George, 1972: 973. — Kussakin, 1988: 445.

Type species. Ischnosoma quadrispinosum Sars, 1879 (by

monotypy).

Diagnosis. Pereonites 5-7 fused with pleonites; pleotelson

fully fused to pereonite 7. Antenna 1 of 6 articles. Mandible

palp absent. Maxilliped palp narrower than basal endite,

articles 2 and 3 expanded. Pereopod 1 carpus of subequal width

throughout length. Male pleopod 2 stylet does not extend

beyond sympod. Uropod moderately short and of 1 article

(except Haplomesus franklinae sp. nov. — with 2 articles).

Remarks. In his description of Haplomesus, Hansen (1916)

stated that the uropods are of only one article. This is true of all

except the new species described below which has two uro-

podal articles. In Haplomesus franklinae, the presence of

biarticulate uropods is treated as a specific autapomorphic char-

acter. The fusion of pereonites 5-7 with the pleonites and pleo-

telson, and the stylet of male pleopod 2 not extending beyond

the sympod, are two of the characters which preliminary phy-

logenetic analysis have shown to be key synapomorphies for

this genus (unpubl. data). This analysis also shows that if

Haplomesus franklinae were placed in another genus it would

render Haplomesus paraphyletic.

Haplomesus franklinae sp. nov.

Figures 1-6

Material examined. Holotype. Australia, Vic., S of Point Hicks

(38°25'S, 149°00"E), 1500 m, compacted clay, WHOI epibenthic sled,

G.C.B. Poore et al, RV Franklin, 22 Jul 1986 (stn SLOPE 27), NMV

J20300 (preparatoiy female, 6 mm).

Paratypes. Australia, Vic., 76 km S of Point Hicks (38°29.33'S,

149°19.98'E), 1840-1750 m, sandy mud, fine shell, WHOI epibenthic

sled, G.C.B. Poore et al., RV Franklin, 26 Oct 1988 (stn SLOPE 69),

NMV J20303 (1 male, 6 mm); same data as holotype, NMV J40686 (1

female, 6 mm).

Other material. NSW, 54 km ESE of Nowra (34°52.72'S,

15ri5.04'E), 996-990 m, mud, fine sand, fine shell (stn SLOPE 53),

NMV J20301 (1 female). Off Nowra (34°58.40'S, 15r23.20T),

1750-1650 m, NMV J20305 (1 fragment). Off Nowra (34°51.28'S,

15r21.3LE), 1725-1701 m, (stn SLOPE 13), NMV J20304 (1 frag-

ment). 40° 45 'S 149° 09.3 T; 3000-2500 m (stn ER1086-4) Vic., 67

km S of Point Hicks (38°23.95'S, 149° 17.02^), 1277-1119 m, fine

mud (stn SLOPE 67), NMV J20302 (2 females, 5 fragments). 76 km S

of Point Hicks (38°29.33'S, 149°19.98'E), 1840-1750 m, sandy mud

(stn SLOPE 69), NMV J40688 (2 males, 15 females, 41 fragments). S

of Point Hicks (38°25'S, 149°0'E), 1500 m, compacted clay (stn

SLOPE 27), NMV 140687 (2 females, 2 fragments).

Diagnosis. Body granulate, about 6 times as long as wide.

Pereonite 1 with pair of long frontal-facing spines on antero-

lateral margin and pair of stout dorsal spines on posterior

margin. Pereonites 2-A each bearing pair of anterolateral

spines, posterolateral margins of pereonites 5-7 rounded.

Pereonites 4 and 5 about 2 and 7 times as long as pereonite 2

respectively. Pleotelson longer than broad, with wide dorsal

keel. Antenna 1 of 6 articles, article 2 13 times as long as wide,

0.6 times as long as entire antenna 1, bearing 3 long flagellate

setae and 1 aesthetasc. Antenna 2 article 3 more than twice as

long as article 4. Mandible covered with microtrichs, molar

elongate, with many small circular pits. Maxilliped palp articles

2 and 3 expanded, narrower than basal endite. Pereopod 1 car-

pus inferior margin with 1 simple seta, 2 long, robust flagellate

setae and 1 long robust seta; propodus inferior margin with 2

short flagellate robust setae. At least pereopods 2-4 carpi with

finely serrate distal margin. Pleopod 3 exopod with microtrichs,

lateral margin with fringe of fine setae. Uropod of 2 articles,

article 1 0.5 times length of article 2; article 2 with 1 brush seta.

Description of female holotype. Body length 6.3 times maximum width

of pereonite 3 (excluding spines); cuticle calcified, brittle and granul-

ated, all pereonites bearing numerous short simple setae. Head+pere-

onite 1 1.1 times as wide as long, with 2 anterolateral spines each as

long as head+pereonite 1 and 2 short dorsal spines extending from

medial ridge. Ratio of lengths of pereonite 2: pereonite 3 : pereonite 4

: pereonites 5+6+7+pleon (together), 1.0:1.1:1.9:12.1. Pereonite 2

slightly wider than pereonite 1, with transverse ridge. Pereonites 2-7

progressively decreasing in width posteriorly. Pereonites 2-4 each with

pair of anterolateral spines. Anterolateral spines on pereonites 2 and 3

0. 4 times as long as spines on pereonite 1; spines on pereonite 4 0.2

times as long as those on pereonite 1. Posterolareral margins on pere-

onites 5-7 rounded. Pereonites 5-7 and pleon with faint suture lines

present at joins of these pereonites on dorsal surface, ventral surface

fused completely. Pleotelson broad, dorsally with wide longitudinal

ridge. Posterior margin rounded, swollen, with 2 medial sub ventral

lobes.

Antenna 1 article 1 with 3 simple setae and 1 brush seta; article 2

4.0 times as long as and 0.2 times as wide as maximal width of article

1, with 8 simple setae of different lengths, 3 long flagellate setae, 1

biarsh seta and 1 distal aesthetasc; article 3 0.7 times as long as article

1, with 1 simple seta; article 4 0.6 times as long as article 1, with 3

simple setae and 1 brarsh seta; article 5 0.5 times as long as article 1,

without setation; article 6 0.4 times length of article 1, with 5 distal

simple setae. Antenna 2 article 1 quadrangular, without setae; article 2

approximately 0.6 times as long as longest margin of article 1, with no

setae; article 3 elongate, about 4.3 times as long as article 1, with at

least 11 simple setae; artiele 4 about 1.7 times as long as article 1, with

3 simple setae and few spinules on distal margin.

Mandible surface covered with microtrichs; incisor process with 5

cusps; lacinia mobilis with 2 cusps; spine row of 6 spines; molar long,

reetangular, with numerous, small round pits, 9 fine simple setae on

inferior margin and 5 fine simple setae on proximal margin. Maxilla 1

with both lobes bearing numerous microtrichs and fine simple setae on

both margins; mesial lobe 0.9 times as wide as lateral lobe, with 1 den-

tate and 1 distal simple seta; lateral lobe with fine simple setae and

microtrichs on surface and 12 distal, robust, dentate setae. Maxilla 2

lateral and middle lobes of equal width, lateral lobe with clusters of

microtrichs on surface and 5 simple setae on inferior margin; lateral

and middle lobes both with 4 distal pectinate setae of different lengths;

mesial lobe 2.8 times as wide as lateral lobe, with many microtrichs

and scale-setae covering surface, 2 long pectinate setae set medially on

inferior mai'gin, and distally, 4 dentate, 2 pectinate and 6 simple setae.

Maxilliped coxa small, narrow, oblong; basis 1.8 times as long as

widest point, with fine simple setae and 1 simple seta each on lateral

Four new Australian species of Ischnomesidae (Isopoda)

287

Figure I . Haplomesus franklinae sp. nov., female holotype, NMV J20300: A, dorsal view; B, lateral view; C, antenna 1 (right); D, antenna 2. Scale

bar = 1 mm, dorsal and lateral views only.

margin and surface, endite with 3 coupling hooks and distally 3 fan, 5

simple and many fine simple setae. Palp tapering, 2.6 times as long as

basal endite; palp article 1 shortest, subrectangular, with few fine

simple setae and 1 simple seta; article 2 0.6 times as wide as endite at

maximum width, 2.1 times as long and 1.3 times as wide as article 1,

with micro trichs on both margins, 2 simple setae and 1 small robust

seta; article 3 width and length equal to that of article 2 surface and

superior margin with microtrichs and 2 short setae and inferior margin

with fringe of fine simple and 3 longer simple setae; article 4 much

narrower than article 3, 1.6 times as long as article 1, with few

microtrichs and 2 simple setae on distal margin; article 5 length equal

to article 4 but narrower, bearing 5 simple setae and 1 thick distal seta.

Epipod elongate, semicircular, length 3.5 times width and 1.1 times

basis length.

Pereopod 1 basis 0.1 body length; ratio of lengths of articles, basis

to dactylus, 1.0;0.4;0.2:0.4;0.3:0.3; basis with 12 scattered simple

setae; ischium with 3 long and 2 short simple setae; merus with 1 long

and 1 short seta on distal superior margin and 1 simple, 1 flagellate

robust and 1 long robust seta on distal inferior margin; carpus width

subequal, superior margin with 3 long, simple slender setae, inferior

margin with 1 simple and 2 long flagellate robust setae on proximal

half, and more distally with 1 robust seta; propodus width subequal

throughout, superior margin with 3 long simple setae and 4 smaller

simple setae, 1 simple seta set medially and inferior margin with 1 dis-

tal simple seta and 2 robust flagellate setae; dactylus with 4 simple

setae on distal superior margin. Pereopods 2, 3 and 5-7 missing.

Pereopod 4 basis 0.2 body length; ratio of lengths of articles, basis

to dactylus, 1.0:0.6:0.2:0.8: 0.5:0.2; basis with 22 scattered simple

setae; ischium with 10 scattered simple setae; merus with 5 simple

setae; carpus with 6 simple setae and 1 distal brush seta on superior

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K. L. Merrin and G. C. B. Poore

Figure 2. Haplomesus franklinae sp. nov., female holotype, NMV J20300: A, mandible; B, maxilla 1; C, maxilla 2; D, maxilliped (right).

margin, 2 simple setae set medially, 3 robust flagellate setae and dis-

tally, 1 long robust seta on inferior margin, distal margin finely serrate;

propodus superior margin with 7 simple setae and 1 brush seta, inferi-

or margin with at least 5 flagellate robust setae, 1 long robust seta and

1 simple seta; dactylus with 4 simple setae on superior margin.

Additional pereopods from female paratype. Pereopod 1 articles,

length ratios same as in holotype; basis with 5 simple setae scattered;

ischium with 3 simple setae; merus same as that of holotype, carpus

width generally subequal throughout distally, widens slightly proxi-

mally, with 2 long and 1 short simple setae on superior margin, infer-

ior margin setation same as holotype; propodus width subequal

throughout, with 3 long simple setae and at least 1 smaller simple seta

on superior margin, medially 2 simple setae and inferior margin with 2

robust flagellate setae; dactylus same as in holotype.

Pereopod 2 ratio of lengths of articles, basis to dactylus,

1.0:0.4:0.2:0.6:0.3:0.3; basis with 3 robust flagellate setae and 1 bmsh

seta proximally, and 7 simple scattered setae; ischium with 4 simple

setae; merus with 4 simple setae; carpus with 1 simple seta on super-

ior margin and 1 on inferior margin, 2 robust flagellate setae and 1 long

robust setae, distal margin finely serrate; propodus bearing 1 simple

setae proximally, 1 short robust flagellate and 1 long robust seta on

inferior margin and distally 3 simple setae, 1 brush seta and 1 robust

flagellate setae and 1 long robust seta; dactylus with 4 short setae on

superior margin.

Pereopod 3 ratio of lengths of articles, basis to dactylus,

1.0;0.6:0.1:0.8:0.5:0.3; basis bearing 3 proximal robust flagellate

setae, 1 brush seta and 6 scattered simple setae; ischium with 4 simple

setae; mems with 3 simple setae; carpus with 4 simple setae and 1

brush seta on superior margin, and inferior margin with 2 flagellate

robust and 1 long distal robust setae, distal margin finely serrate;

propodus with 1 simple seta on superior margin, inferior margin with

2 robust flagellate setae and 1 long robust setae with few fringed scales

in between and distally with 1 robust flagellate, 1 brush and 2 simple

setae; dactylus bearing 4 short setae on superior margin.

Four new Australian species of Ischnomesidae (Isopoda)

289

Figure 3. Haplomesus franklinae sp. nov., female holotype, NMV J20300: A, pereopod 1; female paratype, NMV 40686: B, pereopod 1 (right).

Pereopod 4 ratio of article lengths similar to that of holotype; basis

bearing 13 simple and 2 brush setae; ischium with 7 simple setae;

mems with 5 simple setae; carpus with 5 simple setae, 2 robust flagel-

late setae and 1 distal brush seta, distal margin finely serrate; propodus

with 2 medial simple setae and on inferior margin 2 robust flagellate,

1 long and 1 short robust seta and 4 simple and 1 robust setae distally;

dactylus same as in holotype.

Pereopods 5-7 missing.

Operculum length 0.9 times maximum width; numerous simple

setae on both lateral margins and scattered on ventral surface, 9 long,

plumose setae on posterior margin. Pleopod 3, exopod 0.8 times length

of endopod, with few surface microtrichs and fringe of fine simple

setae on superior margin. Pleopod 4 length 1 .5 times width.

Uropod straight, 0.5 times length of pleotelson; article 1 short, with

at least 5 simple setae; article 2 twice as long as article 1, tapering, with

3 simple setae and 1 brush seta.

Male. Pleopod 1 proximal end damaged; lateral margins indented,

distal portion covered with microtrichs; medial keel extends distally

with 4 simple setae on each side of suture line; distal margin with 5

simple setae on both left and right sides; prominent distolateral horns.

Pleopod 2 sympod length 2.4 times maximum width; lateral margin

curved with 7 simple setae and 6 distolateral plumose setae; exopod a

small lobe, 0.2 times length of sympod; stylet 0.3 length of sympod,

narrowing to a point; sperm duct 0.6 length of stylet. Pleopods 3 and 4

similar to female.

Distribution. South-eastern Australia, from Nowra, New South

Wales, to Point Hicks, Victoria; 990-1840 m depth.

Etymology. For the CSIRO Research Vessel Franklin, from

which the material was collected.

Remarks. Haplomesus franklinae is the fifth species of this

genus to be described from the Southern Hemisphere, and the

first to be described from Australian waters. In body form it

most closely resembles Haplomesus modestatenuis Menzies

and George, 1972, with both species having anterolateral spines

on pereonites 1^ and the posterior margin of the pleotelson

with two medial sub ventral lobes. However, H. franklinae dif-

fers from H. modestatenuis in having stouter spines on pereo-

nites 4 and 5, uropods with two articles that extend well past the

pleotelson, lack of dorsal hooks on pereonite 7, and no robust

setae on the posterolateral margins of the pleotelson (Menzies

and George, 1972).

Ischnomesus Richardson

Ischnosoma Sars, 1868; 34 (preoccupied).

Ischnomesus Richardson, 1908: 81. — Hansen, 1916: 56. —

Gurjanova, 1932: 40.— Wolff, 1956: 88.— Menzies, 1962: 111.—

Wolff, 1962: 73. — Menzies and George, 1972: 971. — Kussakin, 1988;

419 (replacement name).

Rhabdomesus. — Richardson, 1908; 81.

Bactromesus. — Wolff, 1962: 83.

Type species. Ischnosoma bispinosum Sars, 1868 (by monotypy).

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K. L. Merrin and G. C. B. Poore

Figure 4. Haplomesus franklinae sp. nov., female holotype, NMV J20300: A, pereopod 4 (right); female paratype, NMV J40686: B, pereopod 2;

C, pereopod 3 (twisted at mems); D, pereopod 4.

Diagnosis. Pereonites 5-7, pleonite 1 and pleotelson free and

articulating. Remaining pleonites fused with pleotelson.

Antenna 1 of at least 6 articles. Mandible palp usually absent.

Maxilliped palp width at least equal with or wider than basal

endite; articles 2 and 3 expanded. Carpus of pereopod 1 infer-

ior margin usually expanded proximally. Stylet of male pleopod

2 either longer or shorter than sympod, if longer, can be whip

like. Uropod long and of 2 articles (except Ischnomesus justi

sp. nov. — with 1 article).

Remarks. The presence of a mandibular palp, the ancestral state

in isopods, is confined to five species of Ischnomesus (namely

/. justi sp. nov., I. roseus Wolff, 1962, 1, simplissimus Menzies,

1962, 1, calcificus Menzies and George, 1972 and 1. carolinae

Chardy, 1974), being absent in all other species in the family.

The presence of a mandibular palp has never been considered

as a character to remove these species from the family or even

the genus Ischnomesus. A preliminary phylogenetic analysis

placed those species with a palp within the 'Ischnomesus clade’

(unpubl. data).

In describing Ischnomesus, Hansen (1916) stated that the

uropods are of two articles. In Ischnomesus justi the uropod is

of one article. This character state has never been reported in

this genus and, while such uropod characters are usually con-

sidered to be of generic significance (for example see Wolff,

1962), 7. justi has free and articulating pereonites 5-7 and

pleonite 1. This is the principal diagnostic of this genus. For

further discussion, see Remarks under 7. justi below.

Four new Australian species of Ischnomesidae (Isopoda)

291

Figure 5. Haplomesus franklinae sp. nov., female holotype, NMV J20300: A, operculum; B, pleopod 3; C, pleopod 4; D, uropod.

Ischnomesus tasmanensis sp. nov.

Figures 7-11

Material examined. Holotype. Australia, Vic., 76 km S of Point Hicks

(38°29.33'S, 149°19.98T), 1840-1750 m, sandy mud, fine shell,

WHOI epibenthic sled, G.C.B. Poore et ak, RV Franklin, 26 Oct 1988

(stn SLOPE 69), NMV J20280 (immature male, 7 mm).

Paratypes. Australia, type locality, NMV J40684 (1 male, 7 mm; 1

female, 8 mm).

Other material. Australia, Vic., 67 km S of Point Hicks (38°23.95"S,

149H7.02T), 1277-1119 m, fine mud, WHOI epibenthic sled, G.C.B.

Poore et ak, RV Franklin, 25 Oct 1988 (stn SLOPE 67), NMV J20279

(2 fragments). Type locality, NMV J40685 (4 fragments).

Diagnosis. Body covered in short setae, about 6 times as long

as wide. Pereonite 1 with pair of 2 frontal-facing, long antero-

lateral spines and 2 stout dorsal spines. Pereonites 2-4 each

with pair of stout dorsal spines; pereonites 2-4 each with pair

of anterolateral spines; pereonites 5-7 each with pair of pos-

terolateral spines. Pereonites 4 and 5, about 4 and 6 times the

length of pereonite 2 respectively, pereonite 5 with 7 pairs of

protrusions extending from each lateral margin, most terminat-

ing with long, thick simple setae. Pleotelson longer than broad,

lateral margins with 3 large spines. Antenna 1 of 6 articles, with

long simple setae present on articles 2 and 3. Antenna 2 article

1 fused to head, with 3 spines; articles 2-4 with numerous

simple setae. Mandible without palp. Maxilliped palp article 2

widest; epipod with 3 simple setae. Pereopod 1 carpus with 2

long robust setae and several robust flagellate setae on inferior

margin. Pleopod 3 exopod over half epipod length, superior

margin with numerous fine simple setae and 9 simple setae.

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K. L. Merrin and G. C. B. Poore

Figure 6. Haplomesus franklinae sp. nov., male paratype, NMV J20303: A, pleopod 1; B, pleopod 2 (right); C, pleopod 3 (right); D, pleopod 4

(right).

Uropod elongate, with article 2 1.4 length of article 1, both

articles with many simple setae of different lengths,

terminating with 3 long simple setae.

Description of immature male holotype. Body length 5.8 times maxi-

mum width of pereonite 3 (excluding spines). Body cuticle calcified

and brittle, with numerous scattered simple long setae. Head-i-pereonite

1 length 0.8 times maximum width (excluding spines), with 2 long

anterolateral spines that do not reach beyond front of head and 2 stout

dorsal spines set posteriorly on pereonite. Although fused, head differ-

entiated from pereonite 1, maximum width 0.7 times width of pere-

onite 1 (excluding spines), with pair of dorsal tubercles. Ratio of

lengths of pereonites 2:3;4:5:6:7: pleonite 1 : pleotelson,

1.0:1.0:4.2:5.5:1.3:1.0:0.3:3.2. Pereonite 1 widest, pereonites 2-4

width subequal at widest point, pereonites 5-7 decreasing in width

posteriorly Pereonites 2-4 with 2 anterolateral and 2 stout dorsal

spines. Pereonites 5-7 with 2 posterolateral spines. Pereonite 5 with 7

pairs of long, thick, simple setae extending from small projections of

cuticle. Pleotelson length 1.2 times width (excluding spines), elliptical,

with 3 pairs of posterior-facing lateral spines extending from margin;

posterior margin dorsal surface with small projection each side of

median large projection; smaller projections are insertion point for uro-

pod; ventral surface with triangular medial extension, not extending

past margin.

Antenna 1 not complete, article 1 short, rectangular, distal end

rounded, with 3 simple setae; article 2 elongate, 5.3 times as long

as article 1, bearing 16 simple setae of different lengths and distally,

1 brush seta; article 3 2.9 times as long as article 1, with 12 simple

setae of different lengths; article 4 1.0 times as long as article 1,

with 1 simple seta and distally, 1 bmsh seta. Antenna 2 not complete,

article 1 fused to head with 3 robust setae and 1 simple seta; article

2 with 11 simple setae; article 3 0.5 times as long as article 2, with

2 simple setae; article 4, damaged, portion present with 40 simple

setae.

Four new Australian species of Ischnomesidae (Isopoda)

293

Figure 7. Ischnomesus tasmanensis sp. nov., male holotype, NMV J20280; A, lateral view; B, dorsal view; C, left antenna base; D, right antenna

base. Scale bar = 1 nun, dorsal and lateral views only.

Mandible incisor process and lacinia mobilis each with 4 cusps;

spine row of 7 spines; molar wide, rectangular, with row of 6 medial

simple setae. Maxilla 1, both lobes with numerous fine simple setae on

both margins; mesial lobe 0.7 times width of lateral lobe; with 1 short

dentate and 7 simple distal setae; lateral lobe with 8 simple setae on

distolateral margin, 12 distal, strong, dentate setae and few proximal

microtrichs. Maxilla 2 lateral lobe 1.2 times width of middle, with

many rows of microtrichs on superior margin and 6 simple setae on

inferior margin; middle lobe with few microtrichs; both lateral and

middle lobes with 4 long distal pectinate setae of different lengths;

mesial lobe 2.4 times width of lateral lobe, with numerous fine simple

setae and microtrichs on surface, 5 simple setae and 2 long pectinate

setae medially set on inferior margin, and distally, 3 dentate setae, 2

long robust simple setae, 2 small pectinate and 1 1 simple setae of var-

ious lengths. Maxilliped coxa small, rectangular; basis length 2.1 as

long as wide at widest point, endite with 3 coupling hooks, distally

with 6 simple setae, 6 pectinate setae and 3 fan setae, flap with margin

of fine simple setae. Palp narrowing, 3.2 times length of basal endite;

article 1 squat and rectangular, with 1 simple setae on superior margin;

article 2 1 .4 times as wide as basal endite at maximum width, 1 .6 times

wider and 2.3 times as long as article 1, with 1 simple setae on super-

ior and 4 simple setae inferior margin; article 3 width equal with arti-

cle 2 at widest point, 2.0 times as long as article 1, with 1 simple seta

on superior and 6 simple setae on inferior margin; article 4 1.7 times as

long as article 1, much narrower than article 3, with 2 long simple setae

on distal inferior margin; article 5 length equal to 4, with 2 simple setae

and terminating with 2 robust simple setae; posterior face of palp, arti-

cle 1 with 1 additional simple setae, article 2 with 2, article 3 with 2,

article 4 with 2 and no additional setae on article 5. Epipod elongate,

length 2.8 times width, 1.3 times basis length, with 3 long simple setae

and microtrichs on superior margin.

Pereopod 1 basis 0.1 body length; ratio of lengths of articles, basis to

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K. L. Merrin and G. C. B. Poore

Figure 8. Ischnomesus tasmanensis sp. nov., male holotype, NMV J20280; A, antenna 1 (right); B antenna 2 (right); C, pereopod 1 (right); D,

pereopod 2 (right); E, pereopod 3 (right).

dactylus, 1.0;0.7:0.3;0.8:0.7;0.4; basis with 7 simple setae, all except 1 on

inferior margin; ischium with 8 simple setae of various lengths; merus

with 2 long simple and 1 short robust setae on distal superior margin

and 2 simple and 2 robust setae of different lengths on inferior margin;

carpus wider proximally, superior margin with 2 long and 1 short sim-

ple setae and inferior margin with 1 short simple seta, 1 short and 2

long robust setae and 7 flagellate robust setae; propodus width sube-

qual throughout, with 9 long simple setae on superior margin and infer-

ior margin with 6 simple setae of different lengths interspersed

between 7 flagellate robust setae; dactylus with 4 simple setae in

inferior margin and clump of 4 simple setae on distal superior margin

Pereopod 2 basis 0.2 body length; ratio of lengths of articles, basis

to mems (carpus-dactylus missing) same as in pereopod 1; basis with

7 simple setae and 1 medial brush seta; ischium with 1 1 simple setae;

merus with 6 simple setae and 1 distal flagellate robust seta; part of

carpus intact, with 1 simple seta and 5 robust flagellate setae.

Pereopod 3 basis 0.2 body length; ratio of lengths of articles, basis

to dactylus, 1.0;0.8:0.4;1.0:0.5:0.3; basis with 17 simple setae, and

superior margin also with 2 fork and 1 bmsh setae; ischium with 13

simple setae; merus with 6 simple setae; carpus with at least 8 simple

setae and 13 flagellate robust setae, 3 of these on distal margin; pro-

podus with at least 11 simple setae and inferior margin with 6 long and

5 short alternating flagellate robust setae on inferior margin; dactylus

with 4 small simple setae on distal superior margin.

Pereopods 4-7 missing.

Pleopod 1 length 3.5 times proximal width; lateral margins each

with 4 simple setae; ridge overlapping dorsal lobes at 0.9 length, with

6 simple setae on left and 7 on right; small distolateral horns extend

from dorsal lobes. Penes triangular, 0.2 length of pleopod 1. Pleopod 2

sympod elongate, length 2.6 times maximum width, with 12 long, dis-

tal, simple setae; exopod short, roughly oval, length 0.2 times sympod

length; stylet 0.8 length of sympod undeveloped. Pleopod 3 exopod 0.9

Four new Australian species of Ischnomesidae (Isopoda)

295

Figure 9. Ischnomesus tasmanensis sp. nov., male holotype, NMV J20280: A, mandible; B, maxilla 1; C, maxilla 2; D, maxilliped; E, basal endite

of maxilliped, distal margin.

times endopod length with fine simple setae on both margins and 9

simple setae on superior margin. Pleopod 4 slightly elongate, length

1.8 times width.

Uropod (from male paratype) of 2 articles, elongate and slender;

article 1 with at least 1 1 long simple setae; article 2 length 1 .4 times

article 1, narrower, with 21 simple setae and 3 long simple setae

distally.

Female. Pereopod 1 thicker and more setose than in male; ratio of

lengths of articles, basis to dactylus, 1. 0:0.5 :0.7:0.6:0.6:0.3; basis with

15 simple setae; ischium with 10 simple setae of different lengths;

merus with 7 simple setae, 2 short robust setae and 2 long robust setae;

carpus similar in shape but more robust than in male, with at least 6

simple setae on superior margin, inferior margin with 9 robust flagel-

late setae, 2 short robust setae, 2 long robust setae and at least 5

simple setae; propodus with 21 simple setae of different lengths on

superior margin and 3 simple setae on surface, inferior margin with 1 1

robust flagellate setae and 4 long simple setae; dactylus similar to that

in male although with 5 simple setae on inferior margin and clump of

3 on distal superior margin.

Distribution. South-eastern Australia, eastern Bass Strait;

1119-1840 m depth.

Etymology. For the Tasman Sea.

Remarks. This is the second species of Ischnomesus reported

from the Tasman Sea off south-eastern Australia. The first, 7.

anacanthas Wolff, 1962, was described from only the anterior

half of one specimen. The first four pereonites of 7. anacanthus

are totally devoid of both lateral and dorsal spines. Of other

species, 7. tasmanensis most resembles 7. birsteini Wolff, 1962

(the anterior half of the type specimen was too damaged for an

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K. L. Merrin and G. C. B. Poore

Figure 10. Ischnomesus tasmanensis sp. nov., male holotype, NMV J20280; A, pleopod 1; B, pleopod 2; C, pleopod 3; D, pleopod 4. Male

paratype, NMV J40684: E, uropod.

accurate description) and I. spaercki Wolff, 1956, both from the

Kermadec Trench. All three species have long simple setae on

lateral projections of the cuticle on pereonite 5, the pleotelson

with posterior-facing lateral spines extending from margin and

the second uropod article terminated with long simple setae.

However, Ischnomesus tasmanensis can be distinguished from

those two species by its pleotelson, which has three lateral

spines on each margin and lacks spines on the dorsal surface. It

can be further distinguished from /. spaercki by a pair of short

dorsal spines on each of the first four pereonites as opposed to

/. spaercki having a pair of dorsal spines on pereonite 1, 4 dor-

sal spines on pereonites 2 and 3, and short dorsal spines on

pereonites 6 and 7.

Ischnomesus justi sp. nov.

Figures 12-14

Material examined. Holotype. Australia, NSW, 54 km ESE of Nowra

(34°52.72'S, 151° 15.04 T), 996-990 m, mud, fine sand, fine shell,

WHOI epibenthic sled, G.C.B. Poore et al., RV Franklin, 22 Oct 1988

(stn SLOPE 53), NMV J20306 (female, 5 mm).

Paratype. Tasmania, eastern Bass Strait, 87 km ENE of North Point,

Elinders I. (39°28.2"S, 148°52.4"E), 841 m, muddy sand, naturalists’

dredge, G.C.B. Poore, HMAS Kimbla, 29 March 1979 (stn BSS 37),

NMV J20309 (1 immature male, 5 mm).

Other material. Tasmania, 48 km ENE of Cape Tourville (42°00.25"S,

148°43.55"E), 1264-1130 m, gravel with lumps of sandy mud aggre-

gate, WHOI epibenthic sled, G.C.B. Poore et al., RV Franklin, 30 Oct

Four new Australian species of Ischnomesidae (Isopoda)

297

Figure 11. Ischnomesus tasmanensis sp. nov., male holotype, NMV J20280: A, pleotelson, ventral view. Female paratype, NMV J40684:

B, pereopod 1.

1988 (stn SLOPE 81), NMV J20307 (1 male, 1 female). Eastern Bass

Strait, 87 km ENE of North Point, Elinders I. (39°28.2'S, 148°52.4'E),

841 m, muddy sand, naturalists’ dredge, G.C.B. Poore, HMAS Kimbla,

29 Mar 1979 (stn BSS 37), NMV J20308 (1 female). Paratype locali-

ty, NMV J40690 (1 fragment). Type locality, NMV J40689 (2 females,

1 fragment).

Diagnosis: Body smooth, about 6 times as long as wide.

Pereonite 1 bearing pair of stout anterolateral spines. Pereonites

2^ anterolateral margins and pereonites 5-7 posterior margin

rounded, lacking spines. Pereonites 4 and 5 elongate, about 2

and 3 times length of pereonite 2 respectively. Pleotelson pad-

dle-like, proximally with wide proximal keel. Antenna 1 with 6

articles, article 2 with several long flagellate setae. Antenna 2,

article 1 not fused to head; articles 2 and 3 with 1 long robust

seta on distolateral margin. Mandible with palp. Maxilliped

palp article 2 widest; epipod broad, rounded, with no setae.

Pereopod 1 carpus inferior margin with only long pectinate and

simple setae. Pleopod 3 exopod small, length less than half of

endopod. Uropod of single article, inserted at about 90° in

pleotelson with 4 simple setae.

Description of female holotype. Body length 5.8 times maximum

width of pereonite 3. Body cuticle smooth, calcified and brittle, with

few short simple setae scattered over pereonites. Head+pereonite 1

curved around in lateral view, with 2 short anterolateral spines. Ratio

of lengths of pereonites 2;3;4;5;6;7: pleonite 1 : pleotelson,

1.0; 1.3:2. 2:2. 9:0. 9:0. 8:0. 3; 1.6. Widths of pereonites 1 (including

spines) to 3 equal; pereonites 4-7 widths decreasing (at widest part)

posteriorly. Pleotelson long, paddle-like in dorsal view, length 1.3

times widest point; apex broadly rounded, with angular posterolateral

corners.

Antenna 1 article 1 conical, bearing 1 short simple seta and 1 distal

bmsh seta; article 2 elongate, 4.4 times as long as article 1, with 6 sim-

ple setae, 4 long flagellate setae and 1 brush seta; article 3 1.8 times

length of article 1, with 2 distal simple setae; article 4 0.6 times length

of article 1, with 3 simple setae; article 5 0.8 times length of article 1,

with 1 simple seta; article 6 length equal to article 5, with 2 short sim-

ple setae and 4 simple setae of different lengths distally. Antenna 2 not

complete; article 1 broad and short, with 1 simple seta; article 2 with

longer lateral margin, 1.4 times length of article 1, with 1 simple seta

and 1 distal lateral robust simple seta; article 3 elongate, 2.1 times

length of article 1, with 2 simple setae and 1 distal lateral long robust

seta; article 4 quadrangular, 1.5 times length of article 1, with 3 simple

setae.

Mandible body with 1 simple seta near palp; incisor process with 5

cusps; lacinia mobilis with 4 cusps; spine row of 5 spines; molar rec-

tangular, angled tightly towards spine row, with microtrichs and cusp

on proximal margin; palp article 1 with 1 simple seta; article 2 1.7

times as long as article 1 with many microtrichs and 2 pectinate setae

on inferior distal margin; article 3 0.5 times as long as article 1 with

microtrichs on the inferior margin and terminating with 1 pectinate

seta. Maxilla 1 with mesial lobe 0.6 times width of lateral lobe, with

numerous fine simple setae on superior margin, and distally 2 simple

setae, 1 pectinate seta and 1 dentate seta; lateral lobe with many fine

simple setae on face and superior margin, distally 2 simple and 12

strong, dentate setae. Maxilla 2 lateral lobe wider than middle, with a

few fine simple setae, 19 simple setae and distally, 1 long pectinate and

3 long simple setae; middle lobe 0.8 times width of lateral lobe, with 2

simple setae on inferior margin and distally, 1 long pectinate and 2 long

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K. L. Merrin and G. C. B. Poore

Figure 12. Ischnomesus justi sp. nov., female holotype, NMV J20306; A, lateral view; B dorsal view; C, antenna 1; D, antenna 2; E, uropod

(right). Scale bar = 1 nun, dorsal and lateral views only.

simple setae; mesial lobe 2.0 times as wide as lateral lobe with many

fine simple setae covering face, inferior margin with 7 simple setae,

medially 2 long pectinate setae, and distally 3 small pectinate, 1 den-

tate and 13 simple setae. Maxilliped coxa large, rectangular; basis 1.7

times as long as widest point, curved over on inferior margin with

fringe of fine simple setae, 2 simple setae and endite with 3 coupling

hooks, and distally 3 fan and 7 simple setae. Palp tapering, 2.6 times

length of basal endite; palp article 1 shortest, wide, rectangular, bear-

ing 4 simple setae; article 2 wide, triangular, 0.9 times maximum width

of basal endite, 3.0 times length of article 1, with 3 simple and 1 robust

setae; article 3 same width as widest point of article 2, 2.1 times length

of article 1, with 1 robust and 11 simple setae; article 4 narrower than

article 3, 1.4 times length of article 1 with 6 simple setae; article 5

twice as long as wide, 1.8 times length of article 1, with 4 simple setae

and 1 thick distal seta. Epipod with broad rounded tip, length 2.9 times

width and 1 .2 times basis length.

Pereopod 1 basis 0.2 body length; ratio of lengths of articles, basis

to dactylus, 1.0:0.4:0.2;0.4:0.4;0.2; basis with 5 simple setae; ischium

with 7 simple setae of various lengths, all distal; mems with 3 simple

setae on distal superior margin and medially, at least 4 simple and 2

long pectinate setae; carpus width subequal throughout, with 3 long

and 2 short simple setae on distal superior margin and inferior margin

with at least 7 simple and 13 pectinate setae of different lengths, with

small fringed scale-setae between; propodus width subequal through-

out, superior margin with 1 1 long simple setae and inferior margin with

4 robust flagellate setae, at least 20 simple setae of various lengths and

with small fringed scale-setae between on margin; dactylus with 8

short simple setae.

Pereopod 2 basis 0.2 body length; ratio of lengths of articles, basis

to dactylus, 1.0:0.8:0.4:0.7:0.3:0.2; basis with 7 simple and 1 distal

short flagellate seta; ischium with 5 simple setae and at least 5 short

flagellate setae; merus with at least 9 simple setae of different lengths;

Four new Australian species of Ischnomesidae (Isopoda)

299

Figure 13. Ischnomesus justi sp. nov., female holotype, NMV J20306: A, mandible; B, maxilla 1; C, maxilla 2; D, maxilliped (right).

carpus with at least 16 simple setae of various lengths, at least 3 robust

flagellate setae, 1 of these on distal inferior margin and 1 brush seta on

distal superior margin; propodus with 20 simple setae and 5 short

robust flagellate setae on inferior margin; dactylus with 4 simple setae.

Operculum length 1.3 times maximum width; heart-shaped, margin

with small simple setae, 4 slightly longer simple setae on posterior

margin. Pleopod 3 exopod small and slender, 0.3 times endopod

length; endopod elongate, rounded distal margin. Pleopod 4 elongate,

length 2.7 times maximum width.

Uropod of 1 article, straight, 0.4 times length of pleotelson; sitting

up in socket almost 90° degrees to body, with 4 simple setae.

Immature male. Pereopod 1 more slender than in female; ratio of

lengths of articles, basis-dactylus, 1.0:0.3;0.2:0.5:0.4:0.2; basis with 6

simple setae; ischium with 3 long simple setae; merus with 2 long

simple setae on distal superior margin and medially 2 long simple setae

and 2 long pectinate setae; carpus width subequal throughout, with 2

long simple setae on distal superior margin, inferior margin with at

least 20 pectinate setae of various lengths and small fringed scale-setae

between; propodus with 17 simple setae of different lengths and

inferior margin with 4 fringed scale- setae and 3 short, robust flagellate

setae; dactylus with 5 simple setae.

Pleopod 1 length 2.6 times maximum width; completely fused, with

no ornamentation, margin slightly compressed, widens towards distal

end and curves inwards, indenting in the middle. Pleopod 2 sympod

length 2.6 times maximum width, with 1 small simple seta; exopod

rounded distally, 0. 1 times as long as sympod; stylet, undifferentiated,

fused with endopod, rounded distally. Pleopod 3 exopod short, length

0.4 times that of endopod; endopod distal margin coming to a wide

point. Pleopod 4 elongate, length 2.0 times width.

Distribution. South-eastern Australia, from Nowra, New South

Wales, to Cape Tourville, Tasmania; 841-1264 m depth.

Etymology. For Jean Just, distinguished isopod taxonomist,

who sorted and made preliminary identifications of the materi-

al on which this study is based.

300

K. L. Merrin and G. C. B. Poore

Figure 14. Ischnomesus justi sp. nov., female holotype, NMV J20306; A, pereopod 1; B, pereopod 2 Male paratype, NMV J J20309:

C, pereopod 1 .

Remarks. Although the uropods of Ischnomesus justi have only

one article, the species displays all the other diagnostic

characters that characterise Ischnomesus. Ischnomesus justi

is similar to /. calcificus Menzies and George, 1972 and

I. simplex Menzies and George 1972 (both species from the

Peru-Chile Trench) and /. roseus Wolff, 1962 (from the

Gulf of Panama and Eastern Pacific Ocean, off Costa Rica).

There are many characters that link these four: pereonites

2-4 rounded, with pereonite 1 having either small spines or

tubercles; 7. calcificus, I. justi and 7. roseus all have a three-

articled mandibular palp (not illustrated in 7. simplex) with two

pectinate setae on the distal inferior margin; and pereopod 1 in

both 7. calcificus and 7. justi has scale-setae on the inferior

margin of the carpus and propodus. The uropods are

undescribed for 7. calcificus and 7. roseus, and are of two

articles in 7. simplex. I. justi is distinguished from the others by

the uropod having a single article, pereonites 1-4 being

broadly rounded, and the pleotelson with only slight lateral

bulges and an anterior keel.

Stylomesus Wolff

Stylomesus Wolff, 1956: 97. — Birstein, 1960: 15. — Menzies, 1962:

123.— Wolff, 1962: 83.— Birstein, 1971: 204.— Kussakin, 1988: 437.

Gomphomesus. — Wolff, 1962: 84.

Helomesus. — ^Wolff, 1962: 84.

Type species. Rhabdomesus inermis Vanhoffen, 1914 (by

monotypy).

Four new Australian species of Ischnomesidae (Isopoda)

301

Figure 15. Ischnomesus justi sp. nov., female holotype, NMV J20306; A, Operculum; B, pleopod 3; C, pleopod 4. Male paratype, NMV J20309:

D, pleopod 1; E, pleopod 2; F, pleopod 3 (right); G, pleopod 4 (right).

Diagnosis. At least pereonite 7 fused with pleon. Antenna 1

with 6 articles. Antenna 2 supported by anterolateral

projections extending from head. Mandibular palp absent.

Maxilliped basis endite large, wider than palp, palp articles

2 and 3 not expanded. Pereopod 1 carpus width usually sub-

equal throughout length. Stylet of male pleopod 2 not

extending past distal margin of sympod. Uropod long and of 2

articles.

Remarks. The fusion of at least pereonite 7 to the pleotelson

and the autapomorphy of the presence of anterolateral projec-

tions extending from the head and supporting the antennae is a

modification of Wolff’s original description but agrees with

that given by Kussakin (1988).

Stylomesus sarsi sp. nov.

Figures 16-20

Material examined. Holotype. Tasmania., continental slope (40°45"S,

149° 09.3 T - 40° 46.54'S 149°00.27T), 3000-2400 m, WHOI

epibenthic sled, P. Hutchings et al., RV Franklin, 10 Dec 1986 (stn

FR1086-4), AM P63900 (male, 5 mm).

Paratypes. Collected with holotype, AM P63901 (1 female, 5 mm;

1 male, 5 mm).

Other material. Australia, Tasmania., 48 km ENE of Cape Tourville

(42°00.25'S, 148°43.55T), 1264-1130 m, gravel with lumps of sandy

mud aggregate (stn SLOPE 81), NMV J20299 (2 females, 1 male, 1

fragment). Tasmania., continental slope, 40°45"S, 149°09.3"E

_40°46.54'S, 149°00.27T, 3000-2400 m, (stn ER1086-4), AM

302

K. L. Merrin and G. C. B. Poore

Figure 16. Stylomesus sarsi sp. nov., male holotype, AM P63900: A, dorsal view; B, lateral view; C, antenna 1; D, antenna 2. Scale bar = 1 mm,

dorsal and lateral views only.

P63902 (17 females, 8 males, 22 fragments). NSW, 67 km ENE of

Nowra (34°41.97'S, 15r22.44'E), 1896-1642 m (stn SLOPE 59),

NMV J20294 (8 females, 2 fragments). Off Nowra (34°58.40'S,

15r23.20T), 1750-1650 m, (stn SLOPE 15), NMV J20291 (1 frag-

ment). Vic., 67 km S of Point Hicks (38”23.95'S, 149H7.02T),

1277-1119 m, fine mud (stn SLOPE 67), NMV J20296 (7 females, 1

fragment). 76 km S of Point Hicks (38°29.33'S, 149H9.98T),

1840-1750 m, sandy mud, fine shell (stn SLOPE 69), NMV J20297

(12 females, 2 males, 7 fragments), NMV J20298 (4 females, 2 males,

1 fragment). 96 km S of Point Hicks (38°40.29'S, 149° 1 8.06 T),

2900-2900 m, compacted clay (stn SLOPE 66), NMV J20295 (8

females, 8 fragments). S of Point Hicks (38°25"S, 149°0"E), 1500 m,

compacted clay (stn SLOPE 27), NMV J20293 (12 females, 2 males,

7 fragments). S of Point Hicks (38°25.90"S, 148°58.60"E), 1850 m,

muddy, sandstone (stn SLOPE 25), NMV J20292 (1 male).

Diagnosis. Body granulate, about 7 times as long as wide.

Head-i-pereonite 1 bearing 2 anterolateral extensions supporting

antennae. Pereonites 1 and 2 with pair of stout anterolateral

spines. Pereonites 2-4 each with pair of elevated dorsal bosses.

Pereonites 4 and 5, about 1.4 and 4.5 times length of pereonite

2 respectively. Pleotelson longer than wide, with wide medial

ridge. Antenna 2 articles 2 and 3 length subequal, article 3

about quarter length of article 4, flagellum of 26 articles, each

setose. Mandible molar quadrangular, with broad face.

Maxilliped with fringe of fine simple setae on basis, palp arti-

cle 1 widest, with each article narrower than the previous.

Pereopod 1 merus with 2 robust setae on distal inferior margin,

carpus width generally subequal throughout length, 0.4 length

of basis, bearing 1 long robust seta and 3 smaller, robust flag-

Four new Australian species of Ischnomesidae (Isopoda)

303

ellate setae. Pereopods 2-7 slim, carpus with finely serrate dis-

tal margin. Uropod with 2 articles, article 2 0.8 times length of

article 1.

Description of male holotype. Body length 6.5 times maximum width

of pereonite 3. Body cuticle highly granulated, calcified and brittle,

with all pereonites bearing short simple setae. Head+pereonite 1 1.7

times as wide as long, with 2 anterolateral projections of head sup-

porting antennae. Ratio of lengths of pereonites 2 ; 3 : 4 : 5 : 6 :

7+pleon, 1.0:1.0:1.4;4.5:0.9;2.8. Pereonites 1 and 2 with pair of stout

anterolateral spines. Pereonite 2 slightly narrower than pereonite 1,

pereonites 2-4 widths subequal (excluding anterolateral spines), 5-7

decreasing posteriorly Pereonites 2-4 each with pair of raised dorsal

bosses. Pleotelson oval in dorsal view, length 1.1 times width, with

wide, medial longitudinal ridge; posterior margin with 2 subventral

lobes.

Antenna 1 article 1 broad, roughly oval, bearing 1 simple seta;

article 2 length 2.2 as long and 0.25 times as wide of article 1, bearing

3 long flagellate setae and 5 shorter simple setae. Antenna 2 of 31 arti-

cles; article 1 small, lacking setae, with a lateral protrusion; article 2

granulated, 5.5 times as long as article 1, with 6 simple setae; article 3

short, 1.7 times as long as article 1 with 1 simple setae; article 4 5.6

times length of article 1, with 7 simple setae; article 5 longest, 8.0

times as long as article 1, with many simple and 2 distal bmsh setae;

flagellum of 26 articles decreasing in length, each setose.

Mouthparts (of male paratype). Mandible incisor process with 5

cusps; lacinia mobilis with 4 cusps; spine row of 4 spines; molar wide,

rectangular, with broad, distal grinding surface with 3 fine simple setae

on face and proximally 6 fine simple setae. Maxilla 1 with mesial lobe

0.6 times width of lateral lobe; both lobes with numerous fine simple

setae on inferior and superior margins; mesial lobe distal margin with

1 short dentate and 1 simple setae; lateral lobe with 12 distal, robust,

dentate setae. Maxilla 2 lateral lobe with several rows of microtrichs

on superior margin; middle lobe 0.9 times width of lateral lobe, no

304

K. L. Merrin and G. C. B. Poore

Figure 18. Stylomesus sarsi sp. nov., male holotype, AM P63900; A, pereopod 1; B, pereopod 2. Female paratype, AM P63901: C, pereopod 1;

D, operculum; E, pleopod 3 (right); F, pleopod 4.

ornamentation, both lateral and middle lobes with 4 distal pectinate

setae of different lengths; mesial lobe with many micro trichs and fine

simple setae covering surface, 2 long, pectinate setae set medially on

inferior margin, and distally, 3 dentate, 2 small pectinate and 6 simple

setae. Maxilliped coxa small, rectangular; basis 1.8 times as long as

widest point, with fringe of fine simple setae on superior margin and in

its surface, endite with 3 coupling hooks and distally 3 fan and 3 sim-

ple setae. Palp tapering, 2.4 times length of basal endite; palp article 1

shortest, almost rectangular bearing 2 simple setae; article 2 0.5 times

as wide as maximum width of basal endite and width equal to and 1.4

times length of article 1, with 2 simple setae and few fine simple setae;

article 3 as wide as and 1.8 times as long as article 1, with 1 simple

setae and 1 stronger distal seta on inferior margin and numerous fine

simple setae; article 4 1.6 times length of article 1, narrower than arti-

cle 3, with 2 distal simple setae and numerous fine simple setae; arti-

cle 5 narrower than article 4, 1.3 times the length of article 1, with 1

thick distal seta and 4 slightly thinner simple setae. Epipod length 2.8

times width, 1.1 times basis length, with proximal end square, distal tip

curved inward and a few microtrichs.

Pereopod 1 basis 0.1 body length; ratio of lengths of articles,

basis-dactylus, 1.0;0.4:0.2;0.4;0.3:0.2; basis with 8 simple setae;

ischium with 3 simple setae; merus with 2 simple and on inferior dis-

tal margin, 1 simple and 2 long, robust setae; carpus width subequal

throughout, inferior margin with 3 flagellate robust setae and 1 long

robust seta; propodus width subequal throughout, with 4 simple setae

on superior margin, 1 simple seta on surface and inferior margin with

2 flagellate, robust and 1 long simple seta; dactylus with 4 short

simple setae on distal superior margin.

Pereopods 2-7, basis 0.2 body length; ratio of lengths of articles,

basis to dactylus, 1.0:0.5:0.3:0.6:0.4:0.2.

Pereopod 2 basis with 12 simple setae; ischium with 8 simple setae;

mems with 4 simple setae; carpus with 5 simple setae on superior

Four new Australian species of Ischnomesidae (Isopoda)

305

Figure 19. Stylomesus sarsi sp. nov., male holotype, AM P63900: A, pereopod 3 (right); B, pereopod 4; C, pereopod 5; D, pereopod 6;

E, pereopod 7 (right).

margin and on inferior margin 5 flagellate robust and 1 long, robust

seta, distal margin finely serrate; propodus with 3 simple setae on supe-

rior margin, 6 flagellate robust setae with small fringed scale-setae

between each on inferior margin; dactylus with 2 short distal setae.

Pereopod 3 basis with 10 simple setae; ischium with 10 simple

setae; merus with 6 simple setae; carpus with 7 simple setae on super-

ior margin and at least 5 flagellate robust setae on inferior margin, dis-

tal margin finely serrate; propodus with at least 2 simple setae, infer-

ior margin with 7 flagellate robust setae with fringed scale-setae

between each and 1 brush seta on distal superior margin; dactylus with

4 small distal simple setae.

Pereopod 4 basis with 15 simple setae; ischium with 13 simple

setae; merus with 8 simple setae; carpus with at least 8 simple setae

and inferior margin with 8 flagellate robust setae, distal margin finely

serrate; half of propodus intact with superior margin with 3 simple

setae and inferior margin with 4 flagellate robust setae with fringed

scale- setae between them.

Pereopod 5 basis with 13 simple setae; ischium with 11 simple

setae; merus with 8 simple setae; carpus with at least 8 simple setae

and at least 4 flagellate robust setae on inferior margin; propodus with

6 simple setae, 10 flagellate robust setae with fringed scale-setae

between them and 1 bmsh seta on distal superior margin; dactylus

superior margin with 2 small simple setae.

Pereopod 6 basis with 13 simple setae; ischium with 9 simple setae;

merus with 9 simple setae; carpus superior margin with 6 simple setae

and inferior margin with 1 simple and 4 flagellate robust setae, distal

margin finely serrate; propodus with 4 simple setae on superior mar-

gin, 6 flagellate robust setae with fringed scale- setae between each on

inferior margin and 1 brush seta on distal superior margin; dactylus

with 3 small simple setae.

306

K. L. Merrin and G. C. B. Poore

Figure 20. Stylomesus sarsi sp. nov., male holotype, AM P63900: A, pleopod 1; B, pleopod 2; C, pleopod 3; D, pleopod 4; E, uropod (right).

Pereopod 7 basis with at least 8 simple setae; ischium with 3 simple

setae; mems with 5 simple setae; carpus with 3 simple setae on super-

ior margin, 2 flagellate setae on inferior margin and distal margin fine-

ly serrate; propodus with 3 simple setae on superior margin and on the

inferior margin, 4 flagellate robust setae with some fringed scale- setae

between; dactylus with 4 small distal simple setae.

Pleopod 1 length 2.6 times maximum width; margins indent at 0.6

of length from proximal end; 5 simple setae near a small transverse

ridge near distal margin; 6 simple setae on right and 5 simple setae on

left of distal margin; prominent distolateral horns. Pleopod 2 sympod

length 2.2 times maximum width; lateral margins curved, with few

simple setae and 6 distolateral, plumose setae; exopod short, 0.1 times

sympod length, rounded distally; stylet 0.4 length of sympod tapering

to a point; sperm duct 0.5 length of stylet. Pleopod 3 exopod length 0.7

times endopod length with a fringe of fine simple setae on surface and

extended from superior margin. Pleopod 4 length 1.8 times width.

Uropod straight, 0.6 times length of pleotelson; article 1 with 2

simple setae; article 2 tapering, 0.8 times length of article 1, with 3

simple setae.

Female. Pereopod 1 shorter and more robust than in male; ratio of

lengths of articles, basis-dactylus, 1.0:0.4:0.2:0.5:0.4;0.2; basis with 8

simple setae; ischium with 3 simple setae; mems with 2 simple setae

on distal superior margin and 2 long robust setae on distal inferior mar-

gin; carpus slightly broader at proximal end, 1 long robust seta on

superior margin, inferior margin with 1 simple seta, 5 robust flagel-

late setae and 1 long robust seta; propodus with 5 simple setae and 2

flagellate, robust setae on inferior margin; dactylus narrower than

propodus, with 4 short simple distal setae.

Operculum slightly wider than long, length 0.9 times maximum

width; curves to slight keel at apex, with few simple setae on lateral

margins, plumose setae and microtrichs on posterior margin. Pleopod

3 similar to male, exopod length 0.8 times endopod. Pleopod 4 length

1.6 times width, with 3 slight dents on lateral margin.

Distribution. South-eastern Australia, from Nowra, New South

Wales, to Cape Tourville, Tasmania; 1119-3000 m depth.

Etymology. In honour of the Norwegian carcinologist, Georg

Four new Australian species of Ischnomesidae (Isopoda)

307

Ossian Sars (1837-1927), the first person to describe taxa of

Ischnomesidae.

Remarks. Stylomesus sarsi is the first species of Stylomesus

recorded from Australian waters. It resembles Stylomesus

natalensis Kensley, 1984 from south-western Indian Ocean off

South Africa. Both species have granulate integuments, short

anterolateral spines on pereonites 1 and 2 and in both the

mandible molars have a broad distal surface. However, S. sarsi

can be distinguished from S. natalensis by the lack of tubercles

set medially on pereonites 1 and 2, less elongate pleotelson, and

longer flagellum of antenna 2. The maxilliped palp of S. sarsi

has setae present on all articles, while on S. natalensis, setae are

present only on the last three articles.

Acknowledgements

This contribution results from material collected as part of a

wide-ranging exploration of the continental slope of south-

eastern Australia, commenced in 1984, and supported by grants

from the Australian Research Grants scheme and by the then

Victorian Institute of Marine Sciences. We acknowledge the

considerable effort devoted to the project by Jean Just

(Museum Victoria, now of Museum of Tropical Queensland,

Townsville) who extracted most of the animals from the

samples and sorted the isopods to putative species. We are

grateful to the RV Franklin Steering Committee and to CSIRO

Marine Laboratories, Hobart, for the provision of ship-time and

to the master and crew of the vessel for help aboard. We are

also grateful to Niel Bruce (NIWA, Wellington) and the anony-

mous reviewers for reviewing the manuscript and providing

feedback.

References

Birstein, J.A. 1960. The family Ischnomesidae (Cmstacea, Isopoda,

Asellota) in the north-western part of the Pacific and the problem of

amphiboreal and bipolar distribution of the deep-sea fauna.

Zoologicheskii Zhurnal, Moscow 34: 3-28. In Russian.

Birstein, J.A. 1963. Deep water isopods (Crustacea. Isopoda) of the

north-western part of the Pacific Ocean. Akademia Nauk, SSSR:

Moscow. 213 pp. English translation by the Indian National

Scientific Documentation Centre, New Dehli, 1973.

Birstein, J.A. 1971. Fauna of the Kurile-Kamchatka Trench. Additions

to the fauna of isopods (Crustacea, Isopoda) of the Kurile-

Kamchatka Trench. Part II. Asellota 2. Trudy Instituta

Okeaonogiya, Akademiya Nauk SSSR, Moscow 92: 162-238. In

Russian.

Chardy, P. 1974. Complements a I'etude systematique des

Ischnomesidae (Isopodes Asellotes) de I'Atlantique. Description de

quatre especes nouvelles. Bulletin Mensuel de la Societe Linneenne

de Lyon (3) 179: 1537-1552.

Gurjanova, E. 1932. Tableaux analytiques de lafaune de TURSS, pub-

lies par Tlnstitut Zoologique de TAcademie des Sciences. Les

isopodes des mers arctiques. Moscow. 181 pp. In Russian.

Hansen, H.J. 1916. Cmstacea Malacostraca III: V. The Order Isopoda.

Danish IngolfExpedition 3: 1- 262 pis 1-16.

Kussakin, O.G. 1988. Marine and brackish-water Crustacea (Isopoda)

of cold and temperate waters of the Northern Hemisphere. 3.

Suborder Asellota 1. Janiridae, Santiidae, Dendrotionidae,

Munnidae, Haplomunnidae, Mesosignidae, Haploniscidae,

Mictosomatidae, Ischnomesidae. Opredeliteli po Faune SSR,

Akademiya Nauk, SSSR 152: 1-501 [in Russian].

Menzies, R.J. 1962. The isopods of abyssal depths in the Atlantic

Ocean. Vema Research Series 1: 79-206.

Menzies, R.J., and George, R.Y. 1972. Isopod Cmstacea of the

Pem-Chile Trench. Anton Bruun Report 9: 1-124.

Poore, G.C.B., Just, J. and Cohen, B.F. 1994. Composition and diver-

sity of Crustacea Isopoda of the southeastern Australian continental

slope. Deep-Sea Research 41: 677-693.

Richardson, H. 1908. Some new Isopoda of the superfamily Aselloidea

from the Atlantic coast of North America. Proceedings of the

United States National Museum 35: 71-86.

Sars, G.O. 1868. Beretning om en i Sommeren 1865 foretagan zoolo-

gisk Reise ved Kysterne af Christianias og Christiansands Stifter.

Forhandlinger i Videnskaps-Selskapet in Kristiania 1868: 1^7.

Vanhoffen, E. 1914. Die Isopoden der Deutschen Siidpolar-Expedition

1901-1903. Deutsche Siidpolar-Expedition, 1901-03 15: 449-598.

Wolff, T. 1956. Isopoda from depths exceeding 6000 m. Galathea

Reports 2: 85-157.

Wolff, T. 1962. The systematics and biology of bathyal and abyssal

Isopoda Asellota. Galathea Reports 6: 1-320

Memoirs of Museum Victoria 60(2): 309-369 (2003)

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

http://www.museum.vic.gov.au/memoirs

New genera and species of sphaeromatid isopod crustaceans from Australian

marine coastal waters

Niel L. Bruce

Marine Biodiversity and Biosecurity, National Institute of Water and Atmospheric Research, Private Bag; 14-901,

Kilbimie, Wellington, New Zealand (n.bruce@niwa.co.nz)

Abstract Brace, N.L. 2003. New genera and species of sphaeromatid isopod crustaceans from Australian marine coastal waters.

Memoirs of Museum Victoria 60(2): 309-369.

Australia has a great diversity of sphaeromatid isopods, both in terms of species and genera, with 187 recorded species

and subspecies in 58 genera. This contribution records and describes a further four genera: Austrasphaera gen. nov.,

Cassidias Richardson, 1906, Koremosphaera gen. nov. and Pedinura gen. nov. and ten new species: Austrasphaera

berentsae sp. nov., A. springthorpei sp. nov., Cassidias australiensis sp. nov., Exosphaeroma agmokara sp. nov.,

Exosphaeroma alveola sp. nov., Koremosphaera colonus sp. nov., Margueritta sandyi sp. nov., Moruloidea perionasus

sp. nov., Pedinura flindersia sp. nov. and Pedinura mokari sp. nov. Several appear to be common and widely distributed,

with both species of Austrasphaera and Pedinura flindersia ranging from Vic. to WA; Pedinura mokari is recorded from

Vic. and SA while Cassidias australiensis is recorded from the NT to the North-West Shelf, WA. The remaining species

are restricted to or near to their type locations. A brief diagnosis is given for Exosphaeroma and an annotated list of all

accepted species is provided, including incertae sedis species and some recent exclusions from this genus.

Key words Taxonomy, Isopoda Sphaeromatidae, Austrasphaera, Cassidias, Koremosphaera, Pedinura, Exosphaeroma, Australia

Contents

Introduction 309

Austrasphaera gen. nov 310

Austrasphaera berentsae sp. nov 311

Austrasphaera springthorpei sp. nov 315

Richardson, 1906 319

Cassidias australiensis sp. nov 320

Exosphaeroma Stebbing, 1900 327

Exosphaeroma agmokara sp. nov 328

Exosphaeroma alveola sp. nov 333

Koremasphaera gen. nov 339

Koremasphaera colonus sp. nov. 340

Margueritta Bruce, 1993 345

Margueritta sandyi sp. nov 346

Moruloidea Baker, 1908 348

Moruloidea perionasus sp. nov 350

Pedinura gen. nov 355

Pedinura flindersia sp. nov 356

Pedinura mokari sp. nov 363

Acknowledgements 365

References 365

Appendix. Species currently placed in Exosphaeroma

Stebbing, 1900 368

Introduction

Over the last two decades of the twentieth century Museum

Victoria and the Australian Museum made extensive shallow-

water collections from southern Australian coastal regions from

New South Wales to southern Western Australia. The collectors

used fine-scale collecting techniques such as epibenthic sleds

and in shallow coastal waters, SCUBA, which resulted in a

wealth of undescribed crustacean material. These collections

include new species many of which, though not collected pre-

viously, proved to be widely distributed and common. This

contribution draws principally from these collections, but is

supplemented by material from the South Australian Museum,

Adelaide and some collected by myself.

Australia has a rich sphaeromatid fauna (Poore et al., 2002)

that displays extraordinary morphological diversity. The docu-

mentation of this fauna results from three discrete periods of

research - by Baker (1908-1929), Harrison and Holdich

(1980-1984 in either combination) and Bruce (1982 to the

present). Poore et al. (2002) listed 187 species and subspecies

of Sphaeromatidae which, together with the species described

herein, are nearly 28% of all sphaeromatid species known

world- wide, and approximately 17% (the figures are continu-

ally changing) of all isopod species known from Australia. It is

310

N. L. Bruce

estimated that this may represent less than 50% of the potential

total.

Methods follow those outlined in Bruce (1994a, 1995,

1997). Abbreviations are: AM, Australian Museum, Sydney;

NTM, Museum and Art Gallery of the Northern Territory,

Darwin; NMV, Museum, Victoria, Melbourne; QM,

Queensland Museum, Brisbane; SAM, South Australian

Museum, Adelaide; WAM, Western Australian Museum, Perth;

CP, circumplumose; PMS, plumose marginal setae; RS, robust

seta/setae; SEM, scanning electron microscope.

Austrasphaera gen. nov.

Type species. Austrasphaera berentsae sp. nov., here

designated.

Diagnosis. Anterior margin of head overriding antennule and

antennae bases. Pleon with all segments entirely fused, with no

visible sutures, entirely fused to pleotelson. Antennule pedun-

cle articles 1 and 2 subequal in length, each about twice as long

as article 3; article 3 posteriorly offset. Pereopods basis and

ischium subequal in length, short, about two-thirds as long as

propodus. Pleopod 1 operculate, rami collinear, endopod with

mesial margin indurate. Male appendix masculina massive,

0. 8-1.0 as wide as endopod, 1.2-1. 5 times as long as endopod,

distally widest. Uropods subterminal.

Description of female. Body elongate, 3^ times as long as

greatest width, strongly vaulted; dorsal surface smooth; lateral

margins subparallel to weakly ovate; unable to conglobate.

Head weakly to strongly anteriorly produced, weakly immersed

in pereonite 1, anteriorly overriding antennule and antennal

bases; rostral process minute. Eyes small, lateral. Pereon seg-

ments without raised posterior margins. Coxal plates distinct,

overlapping anterior over posterior. Membrana cingula absent.

Pleotelson without foramen or excision, but with shallow

ventral exit channel. Pleonal sternite present.

Antennule and antenna in ventral position on head.

Antennule peduncles narrowly separated by epistome;

peduncle articles 1 and 2 robust, anteriorly flattened, expanded;

article 2 inferodistal angle produced forming a lobe, plane of

articles 1 and 2 of projecting ventrally; peduncle article 3 short,

about 0.17 as long as articles 1 and 2, posteriorly offset at mid-

point of article 2; flagellum of 3 articles. Antenna slender, held

beneath body, peduncle articles 1-3 short, together about 1.3

times as long as article 4, article 5 longest; flagellum as long as

or slightly shorter than peduncle.

Epistome narrow, unornamented, anteromedian portion

weakly produced. Labrum unornamented. Mandible incisor

unicuspid or with weakly defined cusps; left mandible with or

without prominent lacinia mobilis; molar process smooth,

without serrations or ridges; palp article 1 longest, 3 shortest.

Maxillule with lateral lobe with 13 RS on gnathal surface,

mesial lobe with 2 long CP slender setae and 1 short slender

seta. Maxilla with all articles well developed; lateral and

middle lobes with flat finely serrate RS, mesial lobe with blunt

and acute long RS, some of which are basally CP. Maxilliped

endite distally with cactus and club setae, laterally with 1 long

curved CP seta; palp articles not mesially produced, mesial

margins with numerous setae, lateral margins without setae.

Pereopods all robust, inferior margins flattened; dactylus

with prominent recurved trifid secondary unguis; propodus

longest, about 1.0-1. 6 times as long as basis. Pereopods

lacking serrate and trifid RS.

Brood pouch composed of an anterior and posterior pocket.

Pleopods with all rami elongate, pleopods 1-3 with PMS, 4

and 5 without PMS. Pleopod 1 exopod 1.3- 1.6 as long as endo-

pod, endopod about half as wide as exopod, apex distally nar-

rowed. Pleopods 3 and 4 without transverse suture. Pleopods 4

and 5 exopod and endopods with transverse thickened ridges;

pleopod 5 endopod with 2 scale patches and transverse suture.

Uropods endopod and exopod lamellar, exopod smaller than

endopod.

Male. Similar to female except for sexual characters.

Pleopod 2 appendix masculina basally or subbasally attached.

Penes short, adjacent, basally not widely separated. Pereopod

1 lateral margin with 2 proximally positioned submarginal

serrate RS.

Composition. Austrasphaera berentsae sp. nov.; A. spring-

thorpei sp. nov.

Etymology. Prom Latin australis (meaning south, in reference

to the ‘South Land’, i.e., Australia), with the ending -sphaera

indicating the family affinity (feminine).

Remarks. This distinctive genus is characterised by the follow-

ing characters: elongate and strongly vaulted body; pleonite

and pleotelson entirely fused; posterior margin of the pleo-

telson entire; antennules ventrally positioned on head, with

articles 1 and 2 expanded, plane of articles ventrally directed;

antennule peduncle articles 1 and 2 subequal in size, article 2

with an anterior lobe and article 3 posteriorly offset on article

2; pereopods robust with a very short basis and a long pro-

podus; operculate pleopods, pleopod 2 exopod about half as

long as endopod, in the male endopod with a massive appendix

masculina; pereopods 5-7 without the usual serrate and trifid

setae; and uropods with the exopod smaller than the endopod

(but not minute).

Pleopods 4 and 5 have thickened ridges on both rami plac-

ing this genus within the broad subfamily concept of

Dynameninae Bowman, 1981. Bruce (1995) defined the

Ischyromene-gmup, an informal grouping of genera within the

Dynameninae. Austrasphaera shows all the characters of that

group of genera: proportions of the antennule peduncle articles,

pleopod 1 with the endopod mesial margin being indurate, and

pleopod 2 with the exopod distinctly shorter than the endopod.

Austrasphaera is similar to four southern Australian genera

(Juletta Bruce, 1993, Maricoccus Poore, 1994 and Margueritta

Bruce, 1993) which share: anterior margin of the head pro-

duced, overriding the antennule and antennal peduncles, anten-

nule peduncle article 3 being posteriorly offset (also present in

Diclidocella Bruce, 1995, and some species of Cymodocella

Pfeffer, 1887, see Bruce, 1995), pleopod 1 being operculate,

pereopods 5-7 with none (or very few) trifid or serrate setae

and the pleotelson largely fused to the pleon. The body shape

of the four genera varies from scale-like in Maricoccus,

New Australian sphaeromatrid isopod crustaceans

311

flattened in Juletta, vaulted in Margueritta, to elongate and

semicylindrical in Austrasphaera.

Two South African species, incorrectly placed in

Dynamenella, are similar to Austrasphaera and to the

Ischyromene-group. These are “D.” tauriis Barnard, 1940 and

“D.” navicula Barnard, 1940. “D.” tauriis has a similar anten-

nule with peduncular article 2 being large and anterodistally

produced, and the anterior margin of the head produced with

ventral antennules and antennae (rather than anterior).

Dynamenella ” navicula has a similar head and body shape to

Austrasphaera, but those appendages that were figured by

Barnard otherwise differ. Both species have a pleon with four

segments and collinear antennule peduncle articles; pereopod

and pleopod morphology is unknown and further comparison is

not possible without the redescription of the South African

species.

Austrasphaera berentsae sp. nov.

Figures 1-4

Material examined. Holotype. Female (ovigerous 3.6 mm), Vic., Bay

of Islands, 38°35.0'S, 142°49.5'E, 28 Apr 1988, 2.5 m, red algae, R.T.

Springthorpe and P.B. Berents (AM P51061).

Paratypes. Vic. 10 males (2.4-2.5 dissected, 2.6 mandibles dis-

sected, 2.8 mm), female (ovigerous. 3.5 nnn, dissected), same data as

holotype (AM P41342). 8 females (ovigerous 3.5-3. 8 SEM, 3.9 SEM,

3.9 mm), 1 km E of Harmers Haven, 38°34'S, 145°40'E, 6 Mar 1982,

6 m, 300 m offshore, rocky, R.S. Wilson and C. Larsen (NMV J26391).

Eemale (3.7 mm), same data as previous (NMV J26394). Male (2.8

mm), same data as previous, but 11m (NMV J26229). Male (2.9 mm),

female (3.4 mm), 50 m E of Petrel Rock, Venus Bay, 38°39'S,

145°42'E, 5 Mar 1982, 8 m, rocky, M. MacDonald, M. Gomon and

G. Baiker (NMV J26393). Male (2.8 mm), female (3.5 mm), 75 m SW

of Eagles Nest, Venus Bay, 38°40'S, 145°40'E, 5 Mar 1982, 8 m,

rocky, R.S. Wilson and G. Barker (NMV J26390). Male (2.5 mm), 50

m offshore, east side of S point, Twin Reefs, 38°41'S, 145°39T, 11m,

C. Larsen, G. Barber and R.S. Wilson (NMV J26395).

Non-type material. Vic. Laurence Rocks, Portland, 38°34.0'S,

141°40.5'E, 23 m, Herdmania momus with encmsting sponge and red

algae (AM P41379). Whalers Point Lighthouse, Portland, 38°20.5'S,

141°37.5T, 10 m, brown algae from boulder bottom (AM P41399).

SA, “The Hotspot” reef, Blinders L, 33M0.8'S, 134°22.5T, 21 m,

large red algae (NMV J39716); 33°40.5'S, 134°22.05^E, 12 m, assort-

ed algae (NMV J39724); 33°40.5'S, 134°22.0'E, 17 m, algae, large

forms (NMV J39696). NE end of West L, 35°37.0'S, 138°36.0T, 12

m, red algae (NMV J39697). Western River Cove, Kangaroo L,

35°43'S, 137°35'E, 6 m, coralline algae (AM P51064). WA.

Mississippi Bay, E of Esperance, 34°0.0'S, 122°170'E, ?2 m, algae

(AM P41125). Point Peron, S of Perth, 32°16'S, 115°4LE, 4 m,

Posidonia and algae (AM P51063). Green L, Rottnest L, 32°0LS,

115°30'E, mixed algal turf (AM P41113). Dongara-Port Denison

Beach, 29°16.0'S, 114°55.0'E, 5 m, red algae, mainly Laurencia

(NMV J26152); 3 m, drift red algae on sand (NMV J26168). Seven

Mile Beach, Dongara, 29°12.0"S, 114°53.0'E, 1 m, epiphytes on

Amphibolus (NMV J39700); D. Edgar (NMV J39698). Red Bluff,

Kalbarri, 27°42'S, 114°0.9T, 10 Jan 1984, mixed corallines (AM

P51062).

Description of female. Body about 3.5 times as long as greatest

width, ovate, widest at pereonites 3 and 4; dorsal surfaces

smooth, without scattered setae. Cephalon anterior margin

without transverse ridge, ventral rostral process weakly

developed. Head and pereonites 1 subequal in length in dorsal

view, pereonite 1 about twice as long as pereonite 2, with 1 or

2 low indistinct sublateral bosses; pereonites 2-7 of approxi-

mately equal length; pereonite 7 with low indistinct median

boss; coxae with sutures, ventrally directed. Pleon with weak

median boss, posterior margin indicated by short lateral

sutures. Pleotelson posterior margin smoothly rounded; ventral

margin with shallow exit channel not extending to posterior of

pleotelson.

Antennule peduncle article 2 2.6 times as long as wide,

about 1.3 times as long as article 2; article 2 flattened, inferior

margin convex; inferior margins of both articles 1 and 2 with

mass of flattened scales and fine tubular setae; article 3 about

one-quarter as long as article 2, inserted midpoint of posterior

margin; flagellum 3-articled, extending to pereonite 1, about

1.7 times as long as article 3. Antenna peduncle articles 1-3

short, combined lengths about equal to that of article 5; article

4 about 0.8 as long as article 5, both articles 4 and 5 with

inferior margins convex; flagellum about equal in length to

peduncle, extending to anterior margin of pereonite 2, with 8

articles.

Epistome smooth, narrow, laterally encompassing labrum,

anteriorly truncate. Left mandible incisor without distinct

cusps, lacinia mobilis without cusps, spine row of 3 serrate

curved spines; right mandible with 3 indistinct cusps, spine row

of 1 broad-based multidigitate spine and 3 serrate blunt spines;

molar process irregular, largely smooth, without serrate mar-

gins or ridged surfaces; palp article 1 longest, without setae;

article 2 with 4 biserrate setae, article 3 with 6 biserrate setae,

terminal seta being largest. Maxillule mesial lobe with 2 long,

weakly pectinate setae and 1 shorter simple seta; lateral lobe

with 10 terminal and peripheral RS on gnathal surface, eleventh

seta set between these; lateral most RS are weakly serrate on

distal part of seta. Maxilla lateral lobe and middle lobe each

with 3 and 4 curved RS respectively, mesial lobe with 6 RS,

variously serrate, mesial-most seta only being acute, remainder

terminally rounded. Maxilliped endite extending about half

way along palp article 4, distal margin with 1 conical RS, 3

rounded RS, 2 cactus RS and 2 slender CP RS; palp articles 2-5

with about 10, 15, 14 and 11 setae respectively.

Pereopod articles generally with abundant widely-spaced

microtrichs. Pereopod 1 basis about 1.7 times as long as great-

est width, approximately half as long as propodus; inferodistal

angle with 1 simple setae; ischium as long as basis, twice as

long as greatest width; merus about 0.4 as long as ischium, 0.8

times as long as greatest width, caipus (inferior margin) 0.6 as

long as merus, 0.5 as long as wide; propodus twice as long as

ischium, 3 times as long as greatest width; dactylus about half

as long as propodus, unguis strongly recurved, inferior margin

with prominent serrate cuticular scales, secondary unguis

recurved with 2 basal cusps. Pereopods 2 and 3 similar to pere-

opod 1; pereopod 2 with 1, 1 and 4 setae on inferior margins of

merus, carpus and propodus respectively; pereopod 3 with 2, 2

and 3 setae on inferior margins of merus, carpus and propodus

respectively. Pereopods 5 and 6 similar, shorter than pereopods

1 and 2. Pereopod 6 propodus inferior margin without setae.

Pereopod 7 short, slightly longer than pereopod 6, articles

robust and laterally flattened, merus slightly shorter than wide.

312

N. L. Bruce

Figure \ . Austrasphaera berentsae sp. nov. A, B, D, female holotype, G-I, female paratype, (AM P41342), remainder as indicated. A, lateral view;

B, dorsal view; C, frons and anterior of head in ventral view; D, pleon and pleotelson, ventral view; E, head and anterior pereonites, female

3.5 nun Flinders I. (NMV J39716); F, head and anterior pereonites, male 2.8 mm paratype (AM P41342); G, antennule; H, antennule, detail of

marginal setae; I, antenna.

appearing rounded in lateral view, inferior margin with setulose

fringe and CP setae; propodus 2.8 times as long as proximal

width, tapering distally, inferior margin with fringe of CP setae,

superior margin with numerous scale-setae; dactylus 0.7 times

as long as propodus.

Pleopod 1 exopod and endopod with 13 and 8 PMS respec-

tively; endopod 0.8 times as long as exopod, slender, 4 times as

long as greatest width, proximal lateral margin weakly con-

cave. Pleopod 2 exopod and endopod with 16 and 10 PMS

respectively, those of the proximal two-thirds of the exopod lat-

eral margin only feebly plumose; endopod 1.8 times as long as

exopod. Pleopod 3 exopod and endopod with c. 17 and 8 PMS

respectively. Pleopod 4 both rami with prominent ridges, exo-

pod lateral Omargin with 4 fine setae. Pleopod 5 both rami with

New Australian sphaeromatrid isopod crustaceans

315

process and pleonal sternite; D, pleopod 2.

prominent ridges; exopod with 3 scale patches and transverse

suture. Uropod rami flattened, margins provided with continu-

ous flattened scales and fine setae; exopod 0.4 times as long as

fused endopod, 1.8 times as long as greatest width, distal mar-

gin broadly rounded; endopod broadly rounded, dorsal with 3

clusters of sensory setae.

Male. Anterior margin of head produced slightly dorsally (in

contrast to female in which is produced anteriorly). Males lack

indistinct nodules on pereonite 1, pleon and pleotelson.

Pereopod 1 propodus proximomesial surface with 2 strongly

pectinate robust submarginal setae. Penial processes separate,

adjacent, about 2.5 times as long as basal width, tapering to

rounded apex. Pleopod 2 endopod 1.7 times as long as exopod;

appendix masculina lateral margins diverging slightly, surface

with abundant microtrichs, about 5.3 times as long as wide, and

1.2 times as long as endopod, apex obliquely truncate; widest

at approximately three-quarters of its length.

Size. Males 2. 1-2.8 mm, ovigerous females 3. 5-3. 8 mm,

non-ovigerous females 3. 2-3. 6 mm.

Etymology. For Dr Penny B. Berents (Australian Museum),

amphipod taxonomist and collector of much valuable peracarid

material.

Distribution. Vic., SA, WA, north to Kalbarri; 2.5-23 m, with

red and brown algae, including the red alga Laurencia and the

seagrass Posidonia.

Remarks. Austrasphaera berentsae is readily distinguished by

the head being strongly produced anteriorly, an ovate body

shape, the uropods being posterolateral in position and not

meeting posteriorly and by the short uropod exopod. The

uropods of the sympatric A. springthorpei fold across the pos-

terior margin of the pleotelson, and the penial processes are rel-

atively shorter and the appendix masculina less massive than

that of A. berentsae.

Austrasphaera springthorpei sp. nov.

Figures 5-7

Material examined. Holotype. Female (ovigerous 3.2 mm), WA, Red

Bluff, Kalbarri, 27°42"S, 114°0.9"E, 10 Jan 1984, mixed corallines,

R.T. Springthorpe (AM P51057).

Paratypes. 10 males (2. 2-2. 5 mm), 17 females (with visible ova

2.5-3. 5 dissected, 3.5, non-ovigerous 2.4-3. 3 mm), c. 64 unmeasured

male, female, and mancas, same data as holotype (AM P41124). Male

(2.4 nun), female (ovigerous 3.0 mm), same data as holotype except

mixed algae and sediment (AM P41128). 2 males (2.4, 2.5 nun),

female (2.5 mm). Point Peron, 48 km S of Perth, 32°16'S, 115°4rE,

27 Jan 1972, 4 m, Posidonia and algae, W.E and J.M. Ponder (AM

P51060). 2 females (ovigerous 2.7, 2.9 mm), Mississippi Bay, 48 km E

of Esperance, 34°0.0'S, 122°170'E, 71972, 2 m, mixed algae, W.E. and

J.M. Ponder (AM P51058).

Non-type material. Vic. Henty Reef, Apollo Bay, Mounts Bay,

38°47.0^S, 143°40.5T, 18 m, red algae on boulder (AM P51059). SA.

W of “The Hotspot” reef. Blinders I., 33°40.8^S, 134°22.5'E, 21 m,

316

N. L. Bruce

Figure 5 . Austrasphaera springthorpei sp. nov. A, B, holotype, remainder female paratype (AM P41124). A, dorsal view; B, lateral view; C, anten-

nule; D, antenna; E, maxilliped; F, left mandible.

large red algae (NMV J39699); 17 m, assorted algae (NMV J39726);

21 m, large red algae (NMV J39718). North of Tiparra Light, Tiparra

reef, 34°4'S, 137°23T, 10 m, red algae (NMV J26209). WA. Dongara

(NMV J39725).

Description of female. Body about 3.8 times as long as greatest

width, ovate, widest at pereonites 5 and 6; dorsal surfaces

smooth, without setae or microtrichs. Cephalon anterior margin

without transverse ridge, ventral rostral process weakly devel-

oped. Head and pereonites 1 subequal in length in dorsal view,

pereonite 1 about 1.2 times as long as pereonite 2;

pereonites 2-7 of approximately equal length, 5 and 6 slightly

longer than 7 and 4; coxae with sutures, ventrally directed,

inferior margins with setulose fringe. Pleon mesial portion

indistinctly raised, posterior margin indicated short lateral

suture. Pleotelson posterior margin depressed, terminally with

minute indentation, forming open exit-channel with uropods;

ventral margin with shallow exit channel not extending to

posterior of pleotelson.

Antennule peduncle article 2 1.6 times as long as wide,

about 0.9 times as long as article 2; article 2 flattened, with

large anterodistal lobe; anterior margins of both articles 1 and 2

with gel-like layer and fine tubular setae; article 3 about one-

third as long as article 2, inserted at midpoint of posterior

New Australian sphaeromatrid isopod crustaceans

317

Figure 6. Austrasphaera springthorpei sp. nov. Female paratype (AM P41124). A-C, pereopods 1, 2, and 7.

New Australian sphaeromatrid isopod crustaceans

319

margin; flagellum 3-articled, extending to pereonite 1, about

1.7 times as long article 3. Antenna peduncle articles 1-3 short,

combined lengths about equal to that of article 5; articles 4 and

5 subequal in length; flagellum about 0.7 as long as peduncle,

extending to anterior margin of pereonite 2, with 7 articles.

Epistome and mouthparts conforming to the generic charac-

ters and generally similar to that of A. berentsae. Maxilliped

palp articles 2-5 mesial margins with 7, 7, 6 and 8 setae respec-

tively.

Pereopods similar to A. berentsae.

Pleopod 1 exopod and endopod with 18 and 10 PMS respec-

tively; endopod 0.9 times as long as exopod, slender, 3.3 times

as long as greatest width, proximal lateral margin distinctly

concave. Pleopod 2 exopod and endopod with 18 and 9 PMS

respectively; endopod 1.4 times as long as exopod. Pleopod 3

exopod and endopod with 10 and 8 PMS respectively. Pleopod

4 and 5 with both rami with prominent ridges, both with incom-

plete transverse suture; pleopod 5 exopod with 3 scale patches.

Uropod rami flattened, margins provided with gel-like layer

and fine setae; exopod 0.6 times as long as fused endopod

ovate, twice as long as greatest width, distal margin broadly

rounded; endopod broadly rounded, mesial margin oblique to

lateral margin, dorsally with 2 clusters of sensory setae.

Male. Penial processes about 1.3 times as long as basal

width, tapering to rounded apex. Pleopod 2 endopod 1.3 times

as long as exopod; appendix masculina weakly sinuate, mesial

margin curving laterally slightly, surface with scattered

microtrichs, about 8.6 times as long as wide, 1.4 times as long

and 0.6 times as wide as endopod, apex narrowly rounded.

Size. Males 2. 2-2. 6 mm, ovigerous females 2. 5-3. 5 mm,

non-ovigerous females 2.4-3. 3 mm.

Etymology. For Roger T. Springthorpe (Australian Museum),

whose field-collecting has over the years contributed many new

discoveries.

Distribution. Western Vic., SA and WA to Kalbarri; 2-18 m,

from coralline and red algae and Posidonia.

Remarks. See Remarks for A. berentsae for the differences that

separate the two species of Austrasphaera. As the appendages

of these two species are so similar an abbreviated description is

given for this species.

Cassidias Richardson

Cassidias Richardson, 1906: 20. — Barnaid, 1920: 374. — Kensley,

1978: 87.— Harrison, 1984a: 373.— Hanison and Ellis, 1991: 934.

Type species. Cassidias argentinea Richardson, 1906, by

monotypy. Types held at USNM, Smithsonian Institution,

Washington, DC, USA.

Diagnosis. Pleotelson with prominent median process; poster-

ior margin with simple median notch and distinct ventral exit

channel. Antennule peduncle article 1 more than twice as long

as article 2, articles 1 and 2 robust; article 3 slender, all articles

collinear. Maxilliped palp articles 2-4 with distomesial angle

moderately produced, mesial margins with numerous setae.

Pleopods 1-3 with both rami longitudinally oblique. Pleopod 2

with appendix masculina distally attached. Male uropods

attached subdistally on pleon, endopod reduced to short stub,

exopod round in section with apical spike, entire ramus

extending beyond posterior margin of pleotelson.

Description of male. Body stout, about twice as long as great-

est width, strongly vaulted; dorsal surface finely granular, with

scattered setae; unable to conglobate. Head weakly immersed

in pereonite 1. Eyes large, lateral, facets distinct, prominent

posterior lobe. Pereon segments without raised posterior mar-

gins. Coxae not distinctly demarcated, overlapping anterior

over posterior, ventrally directed. Membrana cingula absent.

Pleon of 4 segments, segment 1 largely concealed by pereonite

7, segments 2-4 indicated by 2 distinct suture lines running to

lateral margins of pleon. Pleotelson with prominent median

process; posterior margin with simple median notch and

distinct ventral exit channel. Pleonal sternite present.

Antennule and antenna anteriorly positioned on head.

Antennule peduncle articles 1 and 2 robust, article 1 more than

twice as long as article 2; article 3 slender, all articles collinear;

flagellum shorter than peduncle, extending to anterior of

pereonite 1. Antenna peduncle article 1 and 2 short, subequal,

shorter than 3-5, which become progressively longer; flagel-

lum about as long as peduncle, extending to posterior of

pereonite 1.

Epistome anteriorly acute, with weak mesial constriction;

with indistinct ventral tubercle Labrum unornamented.

Mandible incisor multicuspid; molar process prominent, crush-

ing, provided with marginal scale teeth; left mandible with

prominent lacinia mobilis both mandibles with spine row of 5

or 6 spines; palp article 1 longest, 3 shortest. Maxillule lateral

lobe with about 13 RS on gnathal surface, most of which are

serrate; mesial lobe with 5 long RS, 3 of which are prominent-

ly serrate. Maxilla with all articles well developed; lateral and

middle lobes with flat strongly curved and finely serrate RS,

mesial lobe with several and acute long RS, some of which are

basally CR Maxilliped endite distal margin numerous long

acute CP setae and 3 long CP RS on distomesial margin; palp

articles 2^ each with distomesial angle produced, mesial mar-

gins with numerous setae, lateral margins of articles 2 and 3

without setae, article 4 with 1 seta.

Pereopods all ambulatory, pereopods 1-3 subsimilar, more

robust than 4-7 ; inferior margins of merus, carpus and propo-

dus with serrate and CP RS; setulose fringe weakly developed;

dactylus with prominent simple secondary unguis and 1 flat-

tened seta arising at lateral margin, 2 flat setae at distolateral

margin. Pereopods 6 and 7 inferior and distal margins of merus,

carpus and propodus with numerous serrate and biserrate RS.

Penes paired, separated by about basal width of penial

process; short, not reaching pleopod peduncles.

Pleopods 1-3 both rami with PMS, longitudinal axis of both

rami oblique Pleopod 1 exopod extending beyond endopod dis-

tal margin subtruncate, proximolateral angle with single short

acute RS; endopod distinctly triangular in shape. Pleopod 2

with appendix masculina distally attached on mesial margin.

Pleopod 3 exopod with complete suture, suture not distinct on

pleopod 4 and incomplete on pleopod 5. Pleopods 4 and 5 exo-

pod and endopods with well-developed transverse thickened

ridges; pleopod 5 endopod with 3 lobate scale patches and

320

N. L. Bruce

transverse suture. Uropod exopod 3 times as long as wide, sub-

distally attached on pleon, endopod reduced to short stub, exo-

pod round in section with apical spike, entire ramus extending

beyond posterior margin of pleotelson.

Female. Pleotelsonic process considerably smaller than in

the male; uropods with both rami lamellar, positioned midlater-

ally on pleon, endopod extending only slightly beyond posteri-

or margin of pleotelson; both rami with numerous CP setae.

Mouthparts metamorphosed. Brood pouch of the type species

made up of 4 pairs of oostegites on pereonites 1-4 (Harrison

1984, but see remarks).

Composition. Cassidias argentinea Richardson, 1906, C.

australiensis sp. nov., C. africana Barnard, 1920. Cassidias

tritube rculata Thielemann, 1910 has been synonymised with

Holotelson tuberculatus Richardson, 1909 (Kwon, 1990).

Remarks. Harrison (1984a), on the basis of differences in

brood-pouch morphology, excluded Cassidias africana

Barnard, 1920 from the genus. The type species is known only

from females (Richardson, 1906), and until such time as males

are known it is impossible to assess the generic characters for

this genus. Although brood-pouch morphology is generally

consistent within sphaeromatid genera, there is insufficient

evidence to consider such differences to be axiomatically of

generic merit, and this character is known to vary in Sphaeroma

(Harrison, 1984a) and now also in Margueritta (present study).

At present it seems appropriate to retain the genus, diagnosed

on the basis of the known males, with recognition that its status

is uncertain. A new genus cannot adequately be differentiated

from Cassidias on the basis of type material.

Males of this genus, as here defined can be readily recog-

nised by subterminal uropods which have the endopod reduced

in size, by the presence of a broad, blunt, posteriorly directed

process on the pleotelson and by the terminal appendix mas-

culina. In addition, the posterior margin of the pleotelson has a

distinct simple median notch. Females have the uropods in a

more midlateral position and both rami are flat. The process on

the pleotelson is reduced compared with that of the male, but

does allow identification of females.

The most similar genus is Neonaesa Harrison and Holdich,

1982b which is readily differentiated by the males having the

appendix masculina mesially attached, the pleotelson posterior

margin with three small notches and the pleotelson dorsal surface

with a small median boss flanked by two low submedian bosses.

Females of Neonaesa are separated by having a cylindrical

uropodal exopod and a reduced endopod in contrast to female

Cassidias which have uropodal rami both present and flat.

Cassidias australiensis sp. nov.

Figures 8-13

Material examined. Holotype. Male (4.0 mm), NW end of McCluer I.,

NT, 11°02"S, 132°58"E, 16 Oct 1982, hydroids etc. from bommies,

G.C.B. Poore (NMV J39703).

Paratypes. NT. Males (3.4-4.0 dissected, 4 each 3. 5-4.0 [all

crushedl mm), females (ovigerous 4.3 mm, dissected, non-ovigerous

2.6— 4.2 dissected), immature (2.3-2. 7 mm), 54 umneasured immature,

males and females, same data as holotype (NMV J26384, J39709).

Female (non-ovigerous 3.2 mm), Table Head, Coburg Peninsula, 11

May 1983, mixed substrata, rocks and sponges, N.L. Bmce (NTM

Cr012315). Male, 25 females and immature, McCluer L, S end,

ir06'S, 133°00'E 17 Oct 1982, 8.0 m, hydroids etc., G.C.B. Poore

(NMV J26388). 4 males, c. 40 females and immature, McCluer I., NW

end, 11°02'S, 132°58^E 16 Oct 1982, 8.0 m, yellow hydroids, J.K.

Lowry (NMV J26389). 13 males, c. 100 females and immature, Fannie

Bay, Dai-win, H°24'S, 130°48'E , 26 Oct 1982, 8 m, hydroids etc, J.K.

Lowry (NMV J26386). WA. Females (ovigerous 3.0, non-ovigerous

3.1 mm), North-West Shelf, 19°29.7^S, 118°52.2^, 24 Oct 1983, 37

m, sled, CSIRO, RV Soela (WAM C28895).

Description of male. Body about twice as long as greatest

width, strongly vaulted, ovate, widest atpereonite 5; dorsal sur-

faces anteriorly smooth, becoming minutely granular towards

posterior, scattered setae present on posterior of pereonites and

on pleotelsonic process. Cephalon anterior margin with 2 trans-

verse ridges, ventral rostral process weakly developed. Head

and pereonite 1 subequal in length in dorsal view, pereonite 1

about 1.4 times as long as pereonite 2, unomamented; pere-

onites 2>3<4<5>6>7. Pleon about twice as long as pereonite 7,

with evident sutures and sublateral ‘keys’. Pleotelson process

proximal half with convex lateral margins distal portion with

lateral margins straight, posteriorly subtruncate with weak

median indentation; distal part with 2 distinct ranks of setae

merging to one across posterior margin; in lateral view appear-

ing distally narrowly rounded; ventral margin with deep exit

channel, ventral margin wide and flattened with sublateral

depression mesial to uropods.

Antennule peduncle article 1 1.9 times as long as wide,

about 4.7 times as long as article 2, distal one-third of anterior

margin with obscurely indented blade; both articles 1 and 2

finely pilose, posterior margin with sensory setae; article 3

about half as long as article 1 4 times as long as wide, twice as

long as article 2; flagellum 7-articled, extending to posterior of

pereonite 1, about 2.5 times as long article 3. Antenna peduncle

articles 1 and 2 short, article 1 anterior margin with mass of

setae, combined lengths about equal to that of article 5; article

3 about 0.8 times as long as article 4; article 4 about 0.7 as long

as article 5, articles 3-5 collinear; flagellum about equal in

length to peduncle, extending to anterior margin of pereonite 2,

with 12 articles.

Epistome anteriorly acute, minutely granular, with weak lat-

eral constriction; with distinct anteromedial nodule. Left

mandible incisor with 4 cusps, lacinia mobilis with 3 cusps,

spine row of 5 serrate curved spines; right mandible with 3

indistinct cusps, spine row of 2 broad-based multidigitate

spines, 5 serrate spines; molar process round; palp articles 1

and 2 subequal in length, article 2 distolateral margin with 3

biserrate setae; article 3 with 11 biserrate setae, terminal seta

being largest. Maxillule mesial lobe with 3 long, weakly pecti-

nate RS and 2 shorter simple RS, lateral lobe with 11 peri-

pheral RS on gnathal surface, twelfth seta set between these;

most RS are weakly and bluntly serrate on distal part, proximal

RS strongly serrate. Maxilla lateral lobe and middle lobe each

with 7 and 6 curved finely serrate RS respectively, mesial lobe

with 6 serrate and biserrate RS. Maxilliped endite lateral mar-

gin strongly convex, distal margin with 1 simple RS at sublat-

eral angle, 4 curved CP RS, 3 sinuate CP RS; distomesial

margin with 3 large stout CP RS, increasing in size proximally;

324

N. L. Bruce

palp slender, article 2 4 times as long as basal width; articles 3

and 4 with finger-like distomesial lobes; articles 2-5 with about

12, 13, 14 and 16 setae respectively.

Pereopod 1 basis about 2.9 times as long as greatest width,

approximately twice as long as propodus; superior margin with

few widely-spaced short simple setae; ischium about as long as

propodus, twice as long as greatest width, proximal superior

margin with 2 acute short simple RS, inferior margin with short

setulose fringe; merus about 0.7 as long as ischium, 1.2 times as

long as greatest width, superior distal angle with 2 acute biser-

rate RS, inferior margin with distally plumose setae set among

setulose fringe; carpus approximately as long as wide, inferior

margin 0.6 times as long as merus, with 2 distally plumose

setae; propodus 2.5 times as long as greatest width, inferior

margin with 3 stout serrate and 2 slender distally plumose

setae; dactylus 0.6 times as long as propodus, unguis inferior

margin with prominent serrate cuticular scales, secondary

unguis recurved simple. Pereopods 2-7 subsimilar. Pereopods

2 and 3 similar to pereopod 1 . Pereopod 2 basis 3 times as long

as greatest width, inferodistal angle with single simple seta,

superior margin with widely spaced small setae; ischium 0.7

times as long as basis, 3 times as long as greatest width.

326

N. L. Bruce

Figure 13. Cassidias australiensis sp. nov. SEMs. Male, McLuer Island, NT, NMV J26384. A, cuticle on pleon; B, left mandible; C, pereopod 1,

dactylus; D, maxilla; E, mandible palp article 3; F, pleopod 5, exopod dorsal.

superior distal margin with 2 simple setae, inferior margin with

short setulose fringe and 4 widely-spaced short simple setae;

merus about half as long as ischium, superior distal angle with

1 long simple seta, inferior margin setulose with 2 long dis-

tally plumose setae, distal seta being twice as long as proximal

seta; carpus anterodistal angle with single small simple seta,

inferior margin setulose, with 4 setae, longest of which is

simple, remainder distally plumose; propodus about as long as

ischium, superior distal angle with 3 setae one of which is pap-

pose, inferior margin finely setulose, with 3 RS; dactylus 0.6 as

long as propodus. Pereopods 5-7 similar, longer and with more

RS than pereopods 1 and 2. Pereopod 7 basis 4 times as long as

greatest width, inferodistal angle with single simple seta, super-

ior margin with 2 widely-spaced small setae, distally with 1

palmate seta; ischium 0.8 times as long as basis, 3.8 times as

long as greatest width, proximal superior margin with 1 short

acute RS, superior distal margin with 1 simple setae, inferior

margin with 2 widely- spaced short simple setae; merus one-

third as ischium, superior distal margin with 2 simple setae,

inferior margin setulose with 3 simple RS; carpus 1.3 times as

long as merus, anterodistal angle with 2 acute RS, 1 simple and

1 palmate seta, simple seta, inferior margin setulose, with 1

New Australian sphaeromatrid isopod crustaceans

327

simple and one serrate RS, inferodistal angle with 4 serrate RS;

propodus 1.7 times as long as ischium, superior distal angle

with 1 simple and 1 palmate seta, inferior margin finely

setulose, with 4 simple RS; dactylus 0.4 as long as propodus.

Penes twice as long as basal width, distally subtruncate;

separated by 1.1 times basal width of penial process.

Pleopod 1 exopod and endopod with 26 and 12 PMS respec-

tively; endopod 0.4 as long as exopod, 4 times as long as great-

est width, proximal lateral margin weakly concave. Pleopod 2

exopod and endopod with 28 and 10 PMS respectively; appen-

dix masculina 5 times as long as wide, bluntly rounded, mesial

margin straight, lateral margin weakly convex. Pleopod 3 exo-

pod and endopod with 28 and 13 PMS respectively. Pleopod 4

both rami with prominent thick ridges, exopod lateral margin

with c. 10 fine setae, small proximal submarginal lobe. Pleopod

5 both rami with prominent thick ridges; exopod with 3 scale

patches distal to suture, lateral margin with numerous short

simple setae, distal margins with longer scale-setae; endopod

with scale-setae on distal margin only. Uropod 3 times as long

as wide.

Female. Body shape similar to that of male, but pleotel-

son process prominent, less developed; with fewer setae on

dorsal surfaces. Uropod endopod 4 times as long as wide

(including peduncular portion), distal margin subtruncate;

exopod about 3 times as long as wide, distal margin concave.

Brood pouch of 3 pairs of overlapping oostegites arising from

pereonites 2-4.

Size. Adult males 3.4-4.0 mm, ovigerous females 3.0-4. 3

mm, non-ovigerous females 2.6-4.2 mm.

Etymology. All species of Cassidias have been named after the

country from which they were collected (coincidentally all

starting with the letter ‘a’), and I continue with that practice.

Distribution. NT, North-West Shelf, WA; 8-37 m. The species

has been recorded from hydroids, while field observations sug-

gest that it is a commensal of gorgonian ‘corals’ . A colour pho-

tograph shows a similar isopod from Indonesia, identified as

Cassidias sp., on a gorgonian (Bruce, 1999).

Remarks. Neither Cassidias argentinea nor Cassidias africana

have been described in detail. The male of Cassidias argentinea

remains unknown, but the female differs from C. australiensis

in being larger, with distally rounded uropods (in C. aus-

traliensis the female uropodal endopod is truncate, the exopod

distally indented). Males of Cassidias africana differ in having

the pleotelsonic process subacute rather than truncate and have

a far longer uropodal endopod which is distally swollen giving

a somewhat club-shaped appearance.

Exosphaeroma Stebbing

Exosphaeroma Stebbing, 1900: 553; 1902: 54 (part). — Stebbing,

1910a: 220.— Stebbing, 1910b: 428.— Hansen, 1905: 103, 118.—

Richardson, 1905: 287. — Barnard, 1914: 374. — Monod, 1933:

9-20. — Menzies, 1962: 132. — Menzies and Frankenberg, 1966: 45. —

Menzies and Glynn, 1968: 65. — Schultz. 1969: 131. — Hurley and

Jansen, 1977: 55. — Kussakin, 1979: 398. — Hairison, 1984a: 381. —

Brusca and Iverson, 1985: 26. — Jacobs, 1987: 67. — Kensley and

Schotte, 1989: 229. — Hairison and Ellis, 1991: 939 (key).

Type species: Sphaeroma gigas Leach 1818, by original desig-

nation.

Diagnosis. Pereonite 7 posterior margin even or forming a

mesial point, without processes; pleon and pleotelson without

process. Pleonite 1 dorsal posterior margin with pair of flat and

flush submedian lobes. Pleotelson posterior margin entire, ven-

trally excavate, without distinct exit channel. Maxilliped palp

articles 2-4 mesial margin with distinct lobes. Pereopods 1-3

with inferior margins of mems-propodus densely setulose;

ischium superior margin usually with cluster of long simple

setae at midpoint. Penial process basally set apart, slender usu-

ally between 3-5 times as long as basal width. Pleopod 2

appendix masculina slender, basal in position, extending well

beyond distal margin of ramus, apex may be glandular in

appearance; pleopods 3-5 exopods with complete transverse

suture; pleopods 4 and 5 exopods with or without thickened

ridges or folds. Uropods with both rami prominent in dorsal

view, lamellar, subequal in length, margins not serrated.

Mouthparts not metamorphosed. Brood pouch (Haiiison,1984)

formed of four pairs of oostegites that do not overlap at the

midline.

Species included and distribution. See Appendix.

Exosphaeroma appears to be distributed world-wide, though

apparently absent from the North Atlantic with those species

currently known from the western Atlantic and Caribbean being

incorrectly placed. Its presence in the eastern South Atlantic is

uncertain as there are insufficient data from the African coast to

be sure of its absence. Loyola e Silva (1979) reviewed the dis-

tribution of the genus as then composed.

Remarks. Characters which best serve to identify

Exosphaeroma include the lamellar uropodal rami with the exo-

pod being about as large as the endopod, with both rami lack-

ing serrate margins, the entire posterior margin to the pleo-

telson which lacks a ventral exit channel, and the superior mar-

gin of the ischium of pereopods 1-3 provided with long setae.

Pleopod 2 usually has the appendix masculina longer than the

endopod, and the distal portion is often folded back on itself.

Pleonite 1 has two flat submedian lobes forming part of the

pleonite outline, a character shared with at least Isocladus

Miers, 1976. That genus is readily separated by males having a

prominent backwardly-directed process on pereonite 7.

Exosphaeroma is in critical need of revision, the most recent

diagnosis being that of Kensley and Schotte (1989). A minimal

diagnosis is offered here, based on those species for which the

characters mentioned are described. Currently 32 species are

included (Appendix), although many more species have been

incorporated, and some removed, notably to

Gnorimosphaeroma Menzies, 1954, Harrieta Kensley, 1987,

Ptyosphaera Holdich and Harrison, 1983, Thermosphaeroma

Cole and Bane, 1978, Tholozodium Eleftheriou, Holdich and

Harrison, 1980 and also Clianella Boone, 1923, Cymodoce

Leach, 1814, Paracerceis Hansen, 1905, Pseudosphaeroma

Chilton, 1909, Sphaeroma Bose, 1892 and Sphaeromopsis

Holdich and Jones, 1973.

The relationship of Exosphaeroma to the similar Isocladus

Miers, 1976 and Zuzara Leach, 1818 is unresolved (Bruce and

328

N. L. Bruce

Holdich, 2002). Isocladus and Zuzara are not precisely defined

and include numerous ill-defined and poorly described species.

Many of the 39 species, including those regarded as incertae

sedis (see Appendix), have minimal descriptions. All of the

Southern Hemisphere species need to be redescribed for there

to be sufficient data to assess species differences, relationships

or the biogeography of this genus.

Exosphaeroma agmokara sp. nov.

Figures 14-17

Material examined. Holotype. Male (7.7 mm), Broken Head, NSW,

c. 28°42'S, 153°37'E, 30 Mar 1980, intertidal, on rocks at sand-rock

interface, N.L. Bruce (QM W26727).

Paratypes. 6 males (7. 0-7.4, immature 5.4, 5.6 mm), 5 females

(ovigerous 5.8, 5.9, non-ovigerous 5.7-6.6 mm), 3 mancas (3.9-5.0

mm), same data as holotype (QM W8573).

Description of male. Body 1.8 times as long as greatest width,

ovate, widest at pereonite 5; dorsal surfaces smooth, anteriorly

with fine ridges. Cephalon anterior margin without transverse

ridges, ventral rostral process weakly developed, not visible in

dorsal view. Head about one-third as long as pereonite 1, p

ereonite 1 about 1.5 times as long as pereonite 2; pereo-

nite 2>3=4>5<6<7; pereonite 7 laterally shorter than 6, longer

than pereonite 6 at median point. Pleon laterally about twice as

long as pereonite 7, with sublateral ‘keys’. Pleotelson strongly

vaulted, posterior margin produced to acute apex; ventral

margin anteriorly excavate.

Antennule peduncle article 1 1.4 times as long as wide,

about 2.4 times as long as article 2, anterior and posterior mar-

gins convex; article 3 about two-thirds as long as article 1, 2.5

times as long as wide, 1.6 times as long as article 2; flagellum

16-articled, extending to posterior of pereonite 1, about 2.8

times as long article 3. Antenna relatively robust, peduncle

articles 1 and 2 short, article 1 anterior margin with mass of

setae, combined lengths about equal to that of article 5; article

3 about 1.2 times as long as article 4; article 4 about 0.6 as long

as article 5, articles 3-5 collinear; flagellum about 1.4 times as

long as peduncle, extending to middle of margin of pereonite 3,

with 17 articles.

Epistome anteriorly truncate, anterior lateral margins

straight, diverging to mid point, then narrowing to medial con-

striction. Left mandible incisor with 3 cusps, lacinia mobilis

with 3 cusps, spine row of 7-8 curved serrate spines; right

mandible with 3 indistinct cusps, spine row of 2 broad-based

multidigitate spines, 11 serrate spines; molar process round,

crushing surface strongly ridged; palp articles 1 and 2 subequal

in length, article 2 distolateral margin with 16 finely biserrate

setae; article 3 with 18 biserrate setae, terminal 2 setae being

longest. Maxillule mesial lobe with 4 long, strongly CP RS,

lateral lobe with 10 RS on gnathal surface, twelfth seta set

between these; 4 lateral-most and mesial-most RS simple, others

strongly serrate. Maxilla lateral lobe and middle lobe each with

9 curved finely serrate RS respectively, mesial lobe with about

20 serrate and biserrate RS, many of which are weakly

spatulate, proximal seta longest. Maxilliped endite lateral

margin strongly convex, distal margin with 1 simple RS at sub-

lateral angle, 5 curved CP RS, 8 mesially bent CP RS; distomesial

margin with 5 large stout CP RS, increasing in size proximally;

palp articles 2-5 with about 26, 30, 30 and 24 setae respectively.

Pereopod 1 with inferior margin of distal half of ischium and

entire margin of merus to propodus with dense setulose fringe;

basis about 2.8 times as long as greatest width, approximately

twice as long as propodus; superior margin with 3 close-set

short simple setae at midpoint; ischium 0.8 times as long as

basis, 2.3 times as long as greatest width, superior margin

weakly setulose with 4 prominent acute simple setae; merus

short, about 0.3 times as long as ischium, about 0.7 times as

long as greatest width, superior distal angle with 5 acute long

simple setae, inferior margin with 5 setae distal-most only

being long and extending beyond setulose fringe; carpus 0.8

times as long as wide, inferior margin with 1 long seta and dis-

tally with 1 RS; propodus 2.0 times as long as greatest width,

0.7 times as long as ischium, inferior margin with 2 setae and

distally with 1 RS; dactylus 0.8 times as long as propodus,

unguis inferior margin with prominent serrate cuticular scales,

secondary unguis recurved simple. Pereopods 2-7 subsimilar,

pereopod 7 slightly shorter than pereopod 6. Pereopod 2 basis

2.4 times as long as greatest width, inferior proximal margin

and submarginal surface with setulose fringe; inferodistal angle

with single simple seta, superior margin with widely spaced

small setae; ischium 0.8 times as long as basis, 2.3 times as

long as greatest width, superior distal margin with 4 prominent

simple setae, inferior margin with setulose fringe at distal angle

only; merus about half as long as ischium, superior distal angle

with 2 long simple seta, inferior margin with dense setulose

fringe with 4 short and 1 long simple seta; carpus slightly (1.1)

times longer than merus, 1.6 times as long as greatest width,

anterodistal angle with 2 small simple seta, inferior margin with

dense setulose fringe, with 5 long simple setae; propodus 0.9

times as long as ischium, superior distal angle with 3 setae one

of which is palmate, inferior margin with dense setulose fringe,

with 8 simple setae none of which greatly exceed length of

setulose fringe; dactylus 0.4 as long as propodus. Pereopods

5-7 similar, longer and with more RS than pereopods 1 and 2,

pereopod 7 noticeably more slender that pereopod 6. Pereopod

7 basis 4.2 times as long as greatest width, inferodistal angle

with single simple seta, superior margin with 4 widely spaced

small setae, distally with group of 4 palmate seta, proximal

inferior margin with setulose patch; ischium 0.6 times as long

as basis, 2.9 times as long as greatest width, proximal superior

margin with 4 prominent long simple setae, inferior distal angle

with 1 short simple setae; merus half as long as ischium, super-

ior distal angle truncate, with 5 long simple setae, inferior dis-

tal margin with setulose fringe with 2 simple setae; carpus 1.3

times as long as merus, anterodistal margin with 8 acute biser-

rate and 3 simple RS, inferior margin with setulose fringe with

4 simple setae, inferior distal angle with 2 stout biserrate RS;

propodus about as long as ischium, 4.8 times as long as wide,

inferior margin setulose but less dense than merus and carpus,

distally with 2 short simple RS, superior distal angle with 1

simple and 1 palmate seta; dactylus 0.4 as long as propodus.

Penes slender, 4.5 times as long as basal width; separated by

about twice basal width of penial process, angled mesially,

weakly curved.

Pleopod 1 exopod and endopod with c. 35 and 27 PMS

332

N. L. Bruce

Figure 17. Exosphaeroma agmokara sp. nov. A-E, pleopods 1-5; F, uropod.

respectively, exopod proximolateral RS prominent; endopod

0.7 as long as exopod, 1.5 times as long as greatest width.

Pleopod 2 exopod and endopod with c. 42 and 30 PMS respec-

tively; appendix masculina 12 times as long as wide, distally

highly folded and glandular in appearance, apically rounded,

Pleopod 3 exopod and endopod with c. 45 and 19 PMS respec-

tively; exopod transverse suture entire. Pleopod 4 endopod with

prominent thick ridges, lateral, exopod transverse suture entire.

distal margin with 3 short simple setae, lateral margin with con-

tinuous evenly spaced fine simple setae. Pleopod 5 both endo-

pod with feeble thickenings, without raised ridges, lateral mar-

gin with evenly spaced fine simple setae; exopod with 4 scale

patches, 3 distal to suture, lateral margin with numerous even-

ly spaced simple setae, with obscure proximal lobe. Uropod

exopod extending slightly beyond endopod, about 3 times as

long as wide; endopod about twice as long as wide; both

New Australian sphaeromatrid isopod crustaceans

333

rami with lateral margins evenly convex distal margin evenly

rounded.

Female. Similar to male.

Size. Males 1 .Q-1 .1 mm, ovigerous females 5.8 5.9 mm,

non-ovigerous females, 5. 7-6.6 mm, mancas 3. 9-5.0 mm.

Etymology. From Greek agmo (break) and kara (head), alluding

the type locality; noun in apposition.

Distribution. Known only from the type locality. Broken Head,

northern NSW.

Remarks. Many species of Exosphaeroma (Appendix) are

similar. Exosphaeroma agmokara sp. nov. is best identified by

the posterior margin of pereonite 7 being weakly produced to

form a median point which overrides pleonite 1, the evenly

rounded uropodal rami which extend to about the end of the

pleotelson (i.e. not extending noticeably beyond nor falling

short of the pleotelson apex), the apex of the pleotelson being

weakly produced and acute, the narrowly truncate anterior

margin of the epistome and, in the male, by the distal margin of

the appendix masculina being bent with the subdistal part being

heavily folded (concertina-like).

No Australian species of Exosphaeroma has pereonite 7 pro-

duced to form a median point, while E. serventii Baker, 1928

has the uropodal exopod distally acute; E. bicolor Baker, 1926

has the uropodal exopod longer than the endopod and distally

acute, and an anteriorly rounded epistome; E. laevis (Baker,

1910) has an anteriorly acute epistome; E. aliae Baker, 1926

has a broadly subtruncate pleotelson and anteriorly acute epis-

tome; and E. varicolor Barnard, 1914 has proportionally wider

uropods, the pleotelson bearing two short anteriorly-positioned

submedian ridges.

Similar species are the larger £. gigas (Leach, 1818) (Brandt

and Wagele, 1989) from the Southern Ocean, E. obtusum

(Dana, 1853, sensu Hurley and Jansen, 1978) from New

Zealand and E. pallidum Barnard, 1940 from South Africa. Of

these only E. obtusa has pereonite 7 forming a median point but

can be distinguished by the widely truncate epistome, more

bluntly rounded uropodal rami and the subdistal margins of the

pleotelson being convex rather than straight. The shape of

pereonite 7 is not mentioned in the poorly-known E. pallidum,

which can be separated from E. agmokara by having lanceolate

uropodal rami.

Exosphaeroma alveola sp. nov.

Figures 18-22

Material examined. Holotype. Male (6.5 mm), near mouth of Moona

Moona Creek, Jervis Bay, NSW, 34°04.5'S, 150°41.0'E, 23 Jan 1982,

4.5 m, on test of ascidian Herdmania, P.B. Berents (AM P51055).

Paratypes. NSW. 3 males (6.0 mm), 2 females (non-ovigerous 3.5,

4.2 mm), E of Eaiiy Bower, Manly, 33°48'S, 151°17'E, 22 Nov 1984,

6 m, sand between rocks, J. Just (AM P41189). 2 females (non-oviger-

ous 4.3, 4.4 mm), Edwards Beach, Middle Harbour, Sydney,

33°49.4'S, 151°15.rE, 17 Mar 1985, under rocks at low tide, N.L.

Bruce (AM P41389). Male (7.1 mm), same as previous, 23 Mar 1985

(AM P41382). 5 males (4.6-[6.7 dissected] mm), 11 females (non-

ovigerous 3.0-4. 6 mm), same data as holotype (AM P51056). Male (5

mm, rolled), Murrumbulga Point, Twofold Bay, 37°05'S, 149°54T, 17

Sep 1985, intertidal rock platform, PA. Hutchings and S.J. Keable

(AM P41201). 2 males (5.5, 6.6 mm). Quarantine Bay, Murrumbulga

Point, Twofold Bay, 37°05'S, 149°54T, 17 Sep 1985, subtidal break-

water, PA. Hutchings and S.J. Keable (AM P41202). 2 females (5.5,

3.0 mm), Murmmbulga Point, Twofold Bay, 37°05'S, 149°54'E, 9 Oct

1985, subtidal rock, PA. Hutchings and S.J. Keable (AM P35955).

Tas. 4 females (4.0-5. 5 mm), 3 females (?3.8, 4.0, 4.3 mm). Governor

I., Bicheno, 41°52'S, 148°19'E, 29 May 1984, 30 m, on bryozoan,

(NMV J26419).

Description of male. Body heavily calcified, 1.7 times as long

as greatest width, dorsum noticeably flattened, lateral margins

distinctly thickened, approximately straight in dorsal view,

diverging, widest at pereonite 6; dorsal surfaces minutely punc-

tate. Head about 1.3 times as long as pereonite 1, dorsally

deeply corragated; pereonite 1> 2>3=4>5<6<7; pereonite 7

laterally narrower than 6, not forming part of body outline,

laterally wholly overlapped by coxae of pereonite 7. Pleon

laterally about as long as pereonite 7 in lateral view (flexure

makes this difficult to measure accurately), with median

weakly elevated region, with evident sublateral ‘keys’.

Pleotelson strongly vaulted, posterior margin produced to

bluntly rounded apex; ventral margin anteriorly excavate; dor-

sal margin mesially raised, with 2 obscure punctate submedian

ridges uniting at about midpoint, expanding at posterior

margin.

Antennule peduncle article 1 1.4 times as long as wide,

about twice as long as article 2, anterior margins convex, distal

one-third somewhat produced; article 3 about two-thirds as

long as article 1, 3.7 times as long as wide, 1.3 times as long as

article 2; flagellum 7-articled, extending to posterior of pereo-

nite 1, about 1.5 times as long article 3; antennule peduncles

prominently visible in dorsal view. Antenna relatively robust,

peduncle articles 1 and 2 short, article 1 anterior margin with

mass of setae, combined lengths of articles 1 and 2 about equal

to that of article 5; article 3 about 0.8 times as long as article 4;

article 4 about 0.6 as long as article 5, articles 3-5 collinear;

flagellum about 0.8 times as long as peduncle, extending to

posterior margin of pereonite 1, with 12 articles.

Epistome prominently visible in dorsal view, anteriorly

truncate, anterior lateral margins straight, diverging to mid-

point, then narrowing to medial constriction; ventral surface

pitted, centrally depressed. Left mandible incisor with 4 cusps,

lacinia mobilis with 3 cusps, spine row of 8 curved serrate

spines; right mandible with 3 indistinct cusps, spine row of 2

broad-based multidigitate spines, 8 serrate spines; molar

process round, crushing surface strongly ridged; palp article 2

longest, article 2 distolateral margin with 9 stout and promi-

nently biserrate setae; article 3 with 12 prominently biserrate

setae, 2 terminal setae being longest. Maxillule mesial lobe

with 4 long, strongly CP RS and 1 simple slender seta, lateral

lobe with 11 RS on gnathal surface, twelfth seta set between

these; gnathal RS variously serrate, submesial RS being most

strongly seiTate. Maxilla lateral lobe and middle lobe each with

9 curved finely serrate RS respectively, mesial lobe with about

between 12-18 serrate and biserrate RS (number could not be

clearly observed), proximal seta longest. Maxilliped endite

lateral margin strongly convex, distal margin with 1 simple RS

at sublateral angle, 5 curved CP RS, 2 short straight CP RS;

distomesial margin with 3 large stout CP RS, increasing in size

338

N. L. Bruce

Figure 22. Exosphaeroma alveola sp. nov. SEMs. Female, AM P51056. A, head; B, pleon, dorsal view; C, right mandible, spine row; D, right

mandible, molar; E, maxillule, lateral lobe; F, maxillule, mesial lobe; G, pereopod 1, dactylus; H, pleopod 5, exopod.

New Australian sphaeromatrid isopod crustaceans

339

proximally; palp articles 2-5 with about 11, 14, 16 and 14 setae

respectively.

Pereopod 1 basis about 2.7 times as long as greatest width,

1.7 times as long as propodus; margins without setae, infero-

distal angle with single seta; ischium 0.6 times as long as

basis, about as long as propodus, twice as long as greatest

width, superior margin with scattered scale-setae, with 2 promi-

nent mesial acute simple setae, one proximal and 1 at midpoint,

inferior margin without setae; merus short, about half as long as

ischium, about 0.8 times as long as greatest width, superior dis-

tal angle with 2 short RS, inferior margin with dense setulose

fringe, with 1 apically bifid RS at distal angle; carpus 0.8 as

long as wide, inferior margin with dense setulose fringe, dis-

tally with 1 apically bifid RS; propodus 2.0 times as long as

greatest width, about as long as ischium, inferior margin setu-

lose, with 3 apically bifid setae; dactylus 0.6 times as long as

propodus, unguis inferior margin with prominent serrate cuti-

cular scales, secondary unguis recurved, blunt, simple.

Pereopods 2-7 subsimilar, slender. Pereopod 2 basis 3.3 times

as long as greatest width, inferior margin with 2 minute widely

spaces setae; inferodistal angle with single simple seta, super-

ior margin proximally with 2 palmate setae, with widely spaced

small setae; ischium 0.9 times as long as basis, 3.3 times as

long as greatest width, superior margin with 3 prominent acute

RS, one set proximally, 2 set at midpoint, slender seta distally,

inferior margin setulose at distal angle only; merus about half

as long as ischium, superior distal angle with 2 apically serrate

RS, inferior margin with dense setulose fringe with 4 short

setae and 1 long simple seta; carpus slightly (1.1) times longer

than merus, 1.9 times as long as greatest width, anterodistal

angle with 1 short RS, distal half of inferior margin densely

setulose; propodus 1.4 times as long as ischium, superior distal

angle with 1 palmate seta, inferior margin with dense setulose

fringe, with 6 simple setae none of which greatly exceed length

of setulose fringe; dactylus about half as long as propodus.

Pereopods 5-7 similar, longer and with more RS than pereo-

pods 1 and 2. Pereopod 7 basis 4.8 times as long as greatest

width, inferior margin without setae, superior margin with

numerous widely spaced small setae, distally with 1 palmate

seta; ischium 0.8 times as long as basis, 3.6 times as long as

greatest width, proximal superior margin setulose with 4 promi-

nent acute RS, inferior distal angle with 1 short simple setae;

merus about half as long as ischium, superior distal angle with

2 short acute distally serrate RS, inferior margin with dense

setulose fringe with 1 distally bifid RS; carpus 1.3 times as long

as merus, anterodistal margin with 6 acute biserrate and 1

simple RS, inferior margin with dense setulose fringe, inferior

distal angle with 1 simple and 2 stout biserrate RS; propodus

about as long as ischium, 4 times as long as wide, inferior

margin setulose but less dense than merus and carpus, distally

with 2 short simple RS, superior distal angle with 1 simple and

1 palmate seta; dactylus 0.4 as long as propodus.

Penes 3.4 times as long as basal width; separated by about

half basal width of penial process, straight with subacute apex.

Pleopod 1 exopod and endopod with c. 33 and 17 PMS

respectively, exopod proximolateral RS prominent; endopod

about as long as exopod, 1.7 times as long as greatest

width. Pleopod 2 exopod and endopod with c. 34 and 22 PMS

respectively; appendix masculina 18 times as long as wide,

straight, distally glandular in appearance, apically acute,

extending beyond endopod by 0.4 of its length. Pleopod 3 exo-

pod and endopod with c. 37 and 14 PMS respectively; exopod

transverse suture entire. Pleopod 4 rami without folds, distally

with 1 PMS; exopod transverse suture entire, distal margin with

2 short PMS, lateral margin proximal to suture with continuous

evenly spaced fine simple setae. Pleopod 5 with both rami lack-

ing folds, endopod lateral margin with evenly spaced fine sim-

ple setae; exopod transverse suture entire, with 3 scale patches,

2 distal to suture, lateral margin with numerous evenly spaced

simple setae. Uropod rami subequal in length, rami not extend-

ing beyond posterior margin of pleon, exopod with lateral

margin convex with apex laterally falcate, endopod distally

subtruncate with distolateral angle produced.

Female. Slightly smaller than males; body shape generally

similar to that of male, dorsal surface markedly more orna-

mented.

Size. Males 4.6-6. 6 mm, females 3. 0-4. 6 mm.

Etymology. From Greek alveus (cavity, pit), alluding the pitted

surfaces of this species.

Distribution. Southern NSW, Tas.; from ascidians, bryozoans,

under rocks and in sand from intertidal to 6 m in NSW, 30 m in

Tas.

Remarks. Although the large uropods and coarsely pitted dorsal

surfaces of this species immediately separates it from all other

species, Exosphaeroma alveola is strikingly similar to the

South African E. planum Barnard, 1914 in the shape of the

pleotelson and uropods, pleotelson ornamentation and the

somewhat flattened body shape. In contrast to E. planum the

posterior margin of the pleotelson extends well beyond the

uropods.

Four Southern Hemisphere species of Exosphaeroma have

similar pleotelson ornamentation, with the anterior dorsal sur-

face with two submedian ridges and the posterior part being

somewhat produced, with a median ridge. These species are

E. antikraussi Barnard, 1940, E. kraussi Tattersall, 1913,

E. varicolor Barnard, 1914 (also recorded from Australia by

Hale, 1929) all from South Africa, and E. montis (Hurley and

Jansen, 1978) comb. nov. (Appendix) from New Zealand.

Most species of Exosphaeroma have a group of long, simple

setae at the midpoint of the superior margin of the ischium and

at the distal margin of the merus of the pereopods. This is

shown by the type species (Brandt and Wagele, 1989),

E. agmokara sp. nov., E. bruscai (Espinosa-Perez and

Hendrickx, 2002) and E. amplicauda (Stimpson, 1857)

(Kussakin, 1979), but has rarely been illustrated for other

species. In the present species these setae are absent.

Koremasphaera gen. nov.

Type species. Koremasphaera colonus sp. nov., here designated.

Diagnosis. Pleotelson posterior margin entire, without ventral

exit channel. Dorsal surfaces of pereonites 2-7, pleon and pleo-

telson densely setose. Pleon with 4 segments, sutures running

to lateral margin. Antennule peduncle article 1 more than twice

as long as article 2, articles 1 and 2 robust; article 3 slender, all

340

N. L. Bruce

articles collinear. Maxilliped palp articles 2-4 each with dis-

tomesial angle strongly produced, those of 3-4 finger-like; arti-

cle 5 elongate and finger-like; mesial margins with numerous

long simple setae. Pleopods 4 and 5 without thickened folds or

ridges. Uropods attached subdistally on pleon, both rami

semicylindrical in section, apically acute, exopod apex with

cuticular spike; pleotelson posterior margin entire.

Description of male. Body stout, about twice as long as

greatest width, strongly vaulted; dorsal surface granular, with

abundant setae. Head weakly immersed in pereonite 1. Eyes

small, facets distinct. Pereon segments with raised posterior

margins. Coxae distinctly demarcated, overlapping anterior

over posterior, ventrally directed. Membrana cingula absent.

Pleon of 4 segments, segment 1 largely concealed by pereonite

7, segments 2-A indicated by 2 distinct suture lines running to

lateral margins of pleon. Pleotelson posterior margin entire

without distinct ventral exit channel. Pleonal sternite absent.

Antennule and antenna anteriorly positioned on head.

Antennule peduncle articles 1 and 2 robust, article 1 more than

twice as long as article 2; article 3 slender, all articles collinear;

flagellum about as long as peduncle, extending to middle of

pereonite 1. Antenna peduncle articles 1-2 short, subequal,

shorter than 4-5, which become progressively longer;

flagellum shorter than peduncle, extending to posterior of

pereonite 1.

Epistome anteriorly narrowly rounded, apex overlapped by

rostrum, medial constriction not present. Labrum unorna-

mented. Mandible incisor multicuspid; molar process promi-

nent, crushing, provided with marginal scale teeth; left

mandible with prominent lacinia mobilis both mandibles with

spine row of 5 or 6 spines; palp article 1 longer than articles 2

and 3. Maxillule lateral lobe with about 13 RS on gnathal sur-

face, most of which are serrate; mesial lobe with 4 long RS, 3

of which are prominently serrate, and 2 short acute simple RS.

Maxilla with all articles well developed; lateral and middle

lobes with flat strongly curved and finely serrate RS, mesial

lobe with several acute long RS, some of which are basally CR

Maxilliped endite distal margin numerous long acute CP setae

and 3 long CP RS on distomesial margin; palp articles 2-4 with

distomesial angle strongly produced, that of articles 3-4 finger-

like, article 5 elongate and finger-like; mesial margins with

numerous long setae, lateral margins of articles 2 and 3 without

setae, article 4 with 1 distal seta.

Pereopods all ambulatory, robust; pereopods 1-3 subsimilar,

more robust than 4-7; inferior margins of merus, carpus and

propodus with serrate and CPRS; setulose fringe weakly devel-

oped; dactylus with prominent simple secondary unguis and 2

flattened setae arising at lateral margin, 2 flat setae at disto-

lateral margin. Pereopods 6 and 7 inferior and distal margins

of merus, carpus and propodus with numerous serrate and

biserrate RS.

Penes paired, adjacent; short, about twice as long as basal

width; not reaching pleopod peduncles.

Pleopods 1-3 both rami with PMS. Pleopod 1 exopod distal

margin subtruncate, proximolateral angle with single short

acute RS; endopod distinctly triangular in shape. Pleopod 2

with appendix masculina basally attached on mesial margin.

Pleopod 3-5 exopods with complete suture. Pleopods 4 and 5

exopod and endopods without transverse thickened ridges;

pleopod 5 endopod with 2 scale patches. Uropod attached in

ventromesial position; both rami subcylindrical in section,

subequal in length; both rami narrowing evenly to an acute

apex, not extending significantly beyond posterior margin of

pleotelson, exopod distally with hardened spike.

Female. Similar to male; brood-pouch unknown.

Etymology. Prom Greek korema (brush), coupled with the end-

ing -sphaera to indicate family affinity; alluded to the densely

setose dorsal surfaces (feminine).

Remarks. Koremasphaera is another monotypic genus difficult

to characterise and define but its species defies placement in

any existing genus. The most similar genera are Cymodoce

Leach, 1814 and Oxinasphaera Bruce, 1997, both large genera.

Oxinasphaera is unambiguously defined, the principle diag-

nostic apomorphic characters being the antennule peduncle

spikes, epistome and pereon with cuticular spikes, and the short

uropod exopod with a deeply bifid apex, all of which are absent

in Koremasphaera. In addition Oxinasphaera usually has an exca-

vate pleotelson posterior margin. Points of similarity include

the very long finger-like prolongation of the mesial lobes of the

mandible palp articles 3, 4 and 5, the penial morphology and

the presence of weakly developed spikes on the pleon and pleo-

telson. Cymodoce is less similar, and differs in the pleotelson

having an excavate posterior margin, in having both uropod

rami lamellar (European species) or the exopod only lamellar

(Indo-Pacific species; Bruce, 1997), slender and elongate

penial processes, and in having the posterior of the pleotelson

with a prominent dorsal hardened hemispherical medial dome.

The characters which serve to identify Koremasphaera are

the densely setose dorsal body surfaces; the posterior margin of

the pleotelson entire; uropod rami thickened and terminally

acute; penial processes short, wide and adjacent; maxilliped

palp articles 3-5 strongly produced and provided with long

setae; and pleopods 4 and 5 without thickened folds or ridges.

Uropod morphology in Sphaeromatidae is consistent within

genera, at least when monophyly seems assured (e.g.

Sphaeroma and Oxinasphaera) and the uropods of

Koremasphaera, with both rami semicylindrical and apically

acute and subequal in size, are unique and a putatives synapo-

morphy.

Koremasphaera colonus sp. nov.

Pigures 23-27

Material examined. Holotype. Male (7.5 mm), “The Whaleback” bom-

mie, 0.5 km S of Point Hicks, Vic., 37°48.5^S, 149°16.8^E, 8 Apr 1989,

sponge/yellow zoantliid community in roof of cave, 13 m, G.C.B.

Poore and R.S. Wilson (NMV J39723).

Paratypes. Female (non-ovigerous 11.5 mm), 5 immature (4. 5-5. 5

mm), 36 mancas (2.3-2.6 mm), same data as holotype (NMV J26403).

Description of male. Body twice as long as greatest width, lat-

eral margins subparallel, widest at pereonite 5; dorsal surfaces

of pereon, pleon and pleotelson densely setose, pereonites 6

and 7, pleon and pleotelson with small cuticular spikes. Head

anterior margin without transverse ridges, rostral process

visible in dorsal view, overlapping epistome ventrally; head

New Australian sphaeronnatrid isopod crustaceans

343

Figure 25. Koremosphaera colonus sp. nov. A, pereopods 1; B, pereopod 1, dactylus; C, pereopod 2; D, pereopod 7; E, pereopod 6, ischium-dacty-

lus; F, penial processes.

about half as long as pereonite 1 in dorsal view; pereonite 1

dorsally smooth, pereonites 2-7 posteriorly with raised setose

ridge, postero-ventral angles of coxae 5-7 produced to acute

point, pereonite 1 about 1.5 times as long as pereonite 2; pereo-

nite 2>3>4<5>6>7. Pleon laterally about twice as long as

pereonite 7, with evident sutures, without pleonal sublateral

‘keys’. Pleotelson strongly vaulted, posterior margin weakly

produced and flat; dorsal surface with 4 indistinct subparallel

longitudinal ridges, lateral ridges being shortest.

Antennule peduncle article 1 1.5 times as long as wide,

about 1.9 times as long as article 2, anterior margin convex,

with submarginal ridge, posterior straight, angled obliquely

New Australian sphaeromatrid isopod crustaceans

345

distally; article 3 about as long as article 1, about 4.1 times as

long as wide, 1.7 times as long as article 2; flagellum 8-articled,

extending to posterior of pereonite 1, about twice as long

article 3. Antenna relatively robust, peduncle articles 1 and 2

short, article 1 quadrate, combined lengths of articles 1 and 2

about equal to that of article 5; article 3 about 0.6 times as long

as article 4; articles 4 and 5 subequal in length, articles 3-5

collinear; flagellum stout, about 0.7 times as long as peduncle,

extending to middle of margin of pereonite 3, with 6 densely

setose articles.

Epistome anteriorly narrowly rounded, posterior surface

with transverse row of 4 nodules. Left mandible incisor with 4

cusps, lacinia mobilis with 3 cusps, spine row of 5 curved

serrate spines; right mandible with 2 indistinct cusps, spine row

of 1 broad-based multidigitate spine and 6 distally serrate

spines; molar process round, crushing surface strongly ridged;

palp article 1 longest, articles 2 and 3 subequal in length, arti-

cle 2 distolateral margin with 6 finely biserrate setae; article 3

with 17 biserrate setae, terminal 2 setae being longest.

Maxillule lateral lobe with 12 terminally acute RS on gnathal

surface, twelfth seta set between these. Maxilla lateral and mid-

dle lobes with 11 and 9 curved finely serrate RS respectively,

mesial lobe with about 8 serrate and biserrate RS. Maxilliped

endite lateral margin strongly convex, distal margin with 3 CP

RS at sublateral angle, 2 cactus setae, 6 curved CP RS;

distomesial margin with 35 large stout CP RS, increasing in

size proximally.

Pereopod 1 basis about twice as long as greatest width,

approximately twice as long as propodus; superior margin with

few widely-spaced short simple setae, inferior lateral surface

with numerous scale-setae; ischium 1.6 times as long as propo-

dus, twice as long as greatest width, superior margin with 1

proximal and 1 distal acute short simple RS; merus about 0.3

times as long as ischium, 0.8 times as long as greatest width,

superior distal angle with 3 acute simple RS inferior distal

margin with 4 acute RS and 1 simple slender seta; carpus 1.2

times as long as wide, inferior margin 1.2 times as long as

merus, distally with 1 biserrate, 4 acute RS and 1 slender seta;

propodus 1.9 times as long as greatest width, inferior lateral

margin with 5 short acute RS, inferior margin with 2 basally

biserrate RS; dactylus 0.7 times as long as propodus, inferior

margin with prominent serrate cuticular scales, secondary

unguis simple, acute. Pereopods 2 and 3 similar to pereopod 1,

less robust. Pereopod 2 basis 2.4 times as long as greatest

width, lateral surfaces dense with scales, 1 proximolateral

palmate seta; ischium 0.9 times as long as basis, 3 times as long

as greatest width, superior margin with 1 proximal and 1 dias-

tral acute short simple RS, inferior margin with 3 widely spaced

short simple setae; merus about one-third as long as ischium,

superior distal angle with 4 acute RS, inferior margin with 1

short stout acute RS and 1 long simple seta; carpus about as

long as merus, anterodistal angle with 5 RS, 2 of which are

biserrate, inferodistal margin with 5 acute RS, one of which is

biserrate; propodus about half as long as ischium, 1.8 times as

long as carpus, superior distal angle with 1 simple and 1

palmate setae, inferior margin with 3 short stout acute RS,

distal-most being longest; dactylus 0.5 times as long as pro-

podus. Pereopods 5-7 similar, ischium notably longer than for

pereopods 1-3, distal margins of carpus with more and longer

RS. Pereopod 7 basis 2.6 times as long as greatest width, infer-

odistal angle with single simple seta, superior margin with 2

proximal and 2 distal palmate setae, numerous widely spaced

small scale-setae; ischium 1.1 times as long as basis, 4.5 times

as long as greatest width, superior distal angle with 1 acute RS,

merus one-third as long as ischium, superior distal margin with

3 acute RS, inferior margin 3 minute setae and 1 minute RS at

distal angle; carpus about as long as merus, anterodistal angle

with 5 long acute simple and serrate RS, inferior margin with 2

stout short acute RS, distal angle 3 biserrate RS; propodus 1.4

times as long as carpus, 0.4 times as long as ischium, superior

distal angle with 2 palmate seta, inferior margin with 3 short

stout acute RS, 1 at mid point, 2 at base of dactylus; dactylus

0.4 as long as propodus.

Pleopod 1 exopod and endopod with c. 32 and 11 PMS

respectively, both rami densely setulose; endopod triangular,

0.6 times as long as exopod, 1.3 times as long as greatest width;

exopod lateral and distal margins quadrate. Pleopod 2 exopod

and endopod with c. 33 and 14 PMS respectively; appendix

masculina 10 times as long as wide, slightly wider proximally,

distally weakly bent laterally, apex bluntly rounded. Pleopod 3

exopod and endopod with c. 32 and 12 PMS respectively.

Pleopod 4 exopod lateral margin with 7 fine simple setae,

distal margin with continuous fine setae. Pleopod 5 exopod

with distal scale patch large, forming mediodistal lobe, lateral

margin with numerous simple setae, distal margins with long

scale-setae; endopod with scale-setae on distal margin only.

Uropod peduncle and rami densely covered with large distally

bifid simple setae, palmate setae and scale-setae.

Female. Body shape generally similar to that of male, but

dorsal surfaces lacking large setae, with smaller tubercles than

in male, but with a densely pilose appearance from the

abundant scale-setae.

Etymology. From Latin colo (dwell, inhabit), in the sense of a

colony.

Distribution. Known only from the type locality, off Point

Hicks, Vic.; possibly a commensal of sponges.

Remarks. The setose dorsal body surface in combination with

the pleotelson posterior margin being entire and subcylindrical

terminally acute uropod rami of about equal length all serve to

identify the genus and species.

Margueritta Bruce

Margueritta Bruce, 1993: 164.

Type species. Margueritta sylviae Bruce, 1993, by original

designation.

Species included and distribution. Margueritta sylviae Bruce,

1993; Margueritta sandyi sp. nov.; southern WA.

Remarks. The new species differs from the type species in the

brood-pouch morphology. The type species was re-examined to

ensure that the original diagnosis was correct. Harrison (1984a)

considered brood-pouch morphology to show important gener-

ic characters, and the differences between the two species could

be considered to be of generic merit. However some genera.

346

N. L. Bruce

such as Sphaewma (Harrison, 1984a) are known to be variable,

and the two species Margueritta otherwise agree.

Margueritta sandyi sp. nov.

Figures 28-30

Material examined. Holotype. Female (3.2 mm, ovigerous), western

side of Carnac 1., off Fremantle, WA, 18 Dec 1971, 4-7 m, on algae,

W.F. Ponder (AM P50939).

Paratypes. Female (2.9 mm, non-ovigerous), manca (1.5 mm).

North Lumps, 2 km off Mullaloo, WA, 31°47.30'S, 115°42.80'E, 2

May 1986, 8 m, red algal turf adjacent to reef, G.C.B. Poore and H.M.

Lew Ton (NMV J26053).

Holotype of Margueritta sylviae Bruce, 1993 (AM P41021).

Description of female. Body about 1.7 times as long as greatest

width, ovate, widest at pereonites 2 and 3; dorsal surfaces

smooth, with irregular series of low bosses these provided with

scattered setae. Cephalon anterior margin anteriorly projecting

over frons, medially indented; without transverse ridge, ventral

rostral process weakly developed. Pereonites 1 about 1.6 times

as long as head in length in lateral view, about 1.8 as long as

pereonite 2, with 2 clusters of low sublateral bosses; pereonites

2-7 of approximately equal length; pereonites 2, 3 and 5 with

submarginal irregular transverse row of low bosses; pereonite 4

with one large median boss and single boss at each lateral mar-

gin; pereonites 6 and 7 each with two submedian bosses, pere-

onite 6 with 2 additional low submarginal bosses; coxae with

sutures, ventrally directed, each with dorsal boss. Pleonite 1

entire; pleon otherwise without evident sutures, posterior mar-

gin indicated short lateral suture. Pleotelson with prominent

anteriorly positioned median boss; posterior margin with

distinctly produced ventrally open tube extending beyond

posterior of uropodal rami.

Antennule peduncle article 1 about 3.3 times as long as

distal width, anterior margin with 3 stout simples setae and 1

palmate seta; peduncle article 2 about half as long as article 1,

1.7 times as long as wide, anterodistal margin with 1 long and

1 short simple setae and 3 palmate sensory setae; article 3 about

0.7 as long as article 2, weakly offset on posterior margin of

article 2; flagellum 4-articled, extending to pereonite 1, about

2.8 times as long article 3. Antenna peduncle articles 1-3 short,

combined lengths 1.5 times as long as article 5; article 4 about

0.8 as long as article 5, both articles 4 and 5 with inferior

margins eonvex; flagellum about equal in length to peduncle,

extending to anterior margin of pereonite 2, with 8 articles.

Epistome smooth, narrow, laterally encompassing labrum,

not anteriorly produced. Mandible as for the genus. Maxillule

mesial lobe with 2 long, weakly pectinate setae and 2 shorter

simple seta, lateral lobe with 10 peripheral RS on gnathal sur-

face. Maxilla lateral lobe and middle lobe each with 2 and 3

curved RS respectively, mesial lobe with 6 setae, variously cir-

cumplumose, mesial-most seta only being acute, remainder ter-

minally rounded. Maxilliped endite extending to palp article 5,

distal margin with 1 conical RS, 3 rounded RS, 2 cactus RS and

3 slender CP RS; palp articles 2-5 with about 6, 12, 10 and 10

setae respectively.

Pereopod 1 basis about 2.4 times as long as greatest width,

1.6 times as long as propodus; inferodistal angle with 1 long

simple setae; ischium 0.9 times as long as basis, 2.4 times as

long as greatest width, margins with scale-setae more abundant

on inferior margin; merus about 0.2 as long as ischium, 1.8

times as long as greatest width, inferior margin with sparse

scale- setae and single long simple seta; carpus (inferior margin)

0.6 as long as merus, 0.4 as long as wide, with single simple

seta; propodus 0.7 times as long as ischium, twice as long as

greatest width inferior margin with single distal simple seta;

dactylus about o.8 times as long as propodus, inferior margin

with prominent serrate cuticular scales; unguis strongly

recurved, secondary unguis recurved with 2 basal cusps.

Pereopods 2 and 3 similar to pereopod 1, but with more and

longer setae. Pereopods 5 and 6 similar, shorter than pereopods

1 and 2. Pereopod 7 slightly longer than pereopods 2-6, other-

wise generally similar.

Pleopod 1 exopod and endopod with 8 and 9 PMS respec-

Figure 27. Koremosphaera colonus sp. nov. Female. A. lateral view; B, pleon and pleotelson, dorsal view.

348

N. L. Bruce

Figure 29. Margueritta sandyi sp. nov. Female paratype. A, maxilla; B, maxillule; C, pereopod 1; D, pereopod 2; E, pereopod 7.

lively, both rami distally narrowly rounded; endopod 0.6 as

long as exopod, 2.5 times as long as greatest width, proximo-

lateral margin weakly concave. Pleopod 2 exopod and endopod

with c. 18 and 15 PMS respectively, those of distal margin of

endopod submarginal; endopod twice as long as exopod.

Pleopod 3 exopod and endopod with c. 17 and 14 PMS respec-

tively. Pleopods 4 and 5 damaged, examined in situ, similar to

that of the type species. Uropods not dissected and not exam-

ined in detail; rami flat, distally rounded, not extending to dis-

tal margin of pleotelson.

Male. Unknown.

Etymology. For Dr A.J. (Sandy) Bruce in recognition of his con-

tribution to knowledge of the Crustacea, and to Caridea of the

tropical Indo-Pacific and Australia in particular.

Distribution. Carnac I. and off Mullaloo, southern WA; on

algae; 4-8 m.

Remarks. The numerous small dorsal bosses and the prominent

median bosses on pereonite 4 and the pleotelson separate this

species from its congener, Margueritta sylviae. Additional con-

spicuous points of differences are that in M. sandyi the body

shape is narrower, the body itself is more strongly vaulted and

the pleotelson extends posterior to the uropodal rami.

Moruloidea Baker

Moruloidea Baker, 1908: 150.— Baker, 1926; 276.— Hale, 1929;

292.— Harrison, 1984a: 383.— Harrison, 1984b: 268.

Vallentinia Stebbing, 1914a: 351 (name preoccupied).

New Australian sphaeromatrid isopod crustaceans

349

Figure 30. Margueritta sandyi sp. nov. Female paratype. A, pleopod 1; B, pleopod 2; C, pleopod 3.

Euvallentinia Stebbing, 1914b: 944 (replacement name). — Barnard,

1920: 374. — Nierstrasz, 1931: 218. — Loyola e Silva, 1974: 3.

Type species. Moruloidea lacertosa Baker, 1908, by monotypy.

Species included and distribution. Moruloidea lacertosa Baker,

1908; M. darwinii (Cunningham, 1871); M. tasmaniae (Baker,

1926); M. tumida (Harrison 1984b); M. perionasus sp. nov.;

Gondwanan, southern coasts of Australia (WA, SA and Tas.),

with one species from Atlantic coast of South America and

Falkland Is (Harrison, 1984b).

Diagnosis of male. Body stout, about twice as long as greatest

width, strongly vaulted; dorsal surfaces nodular. Pleotelson

with or without median process; posterior margin with simple

median notch and shallow exit channel. Coxae of pereonite 5

overlapping those of both pereonite 4 and 6. Pleon of 4 seg-

ments, segment 1 largely concealed by pereonite 7, segments

2-A indicated by 2 distinct suture lines running to lateral

margins of pleon. Antennule peduncle article 1 more than twice

as long as article 2, articles 1 and 2 robust; article 3 slender, all

articles collinear. Antenna articles 3 and 5 proportionally large,

article 5 strongly reflexed. Mandible incisor unicuspid, or cusps

indistinct; molar process prominent, crushing, provided with

marginal scale teeth. Maxilliped palp articles 2-4 with

distomesial angle moderately produced, mesial margins

with numerous setae. Pereopods all ambulatory, pereopod 1

massive, robust, propodus inferior margin with or without lobe-

like extension; pereopods 2-7 subsimilar, slender. Penes paired,

close set; short, not reaching pleopod peduncles. Pleopods 1-3

both rami with PMS, both rami of subequal length; pleopod 1

with longitudinal axis of both rami weakly oblique, remaining

pleopods with longitudinal axis of rami straight. Pleopod 2 with

appendix masculina basally attached. Pleopods 3-5 exopods

with complete suture. Pleopods 4 and 5 exopod and endopods

usually with well-developed transverse thickened ridges; pleo-

pod 5 endopod with 3 lobate scale patches. Uropods attached

anterolaterally on pleon, exopod moderate to minute in size;

both rami flat not extending beyond posterior margin of

pleotelson.

Female. Sexual dimorphism weak; mouthparts not meta-

morphosed. Brood pouch of the type species of 3 pairs of oost-

egites on pereonites 2-A\ brood housed in 4 pairs of internal

pouches (Harrison, 1984). Antenna peduncle and pereopod 1

not as robust as in the male.

Remarks. The genus was revised by Harrison (1984b) who

placed Vallentinia Stebbing, 1914a and Euvallentinia Stebbing,

1914b into synonymy.

The coxae of pereonite 5, antenna, antennule, pereopod and

pleotelson of Moruloidea are similar to those of

Caecocassidias Kussakin, 1967 (Brandt, 1998), Cymodopsis

Baker, 1926, CeratocephalusWoo&waid, 1877 (Bruce, 1994b),

350

N. L. Bruce

Kranosphaera Bruce, 1992 and Waiteolana Baker, 1926

(Harrison 1984b). All have coxal plates 5 overlapping both

anteriorly and posteriorly, robust pereopod 1 and the posterior

margin with a simple shallow exit channel. Most (Moruloidea,

Caecocassidias, Ceratocephalus and Kranosphaera) have the

antenna with peduncle article 5 strongly reflexed and flat

uropodal rami with the exopod varying from moderate to

absent Kranosphaera. All species of Cymodopsis are inade-

quately described and the genus is poorly understood. Until

Cymodopsis is revised the relationships of these genera to each

other and to others will remain unclear.

The presence of a prominent pleotelson process in

Moruloidea perionasus sp. nov. and additional data on M. dar-

winii (Brandt, 1998) necessitates modification of the diagnosis

of Holdich (1984b). M. darwinii was described by Brandt

(1998) as having lamellar rami on pleopods 4 and 5.

Moruloidea perionasus sp. nov.

Figures 31-34

Material examined. Holotype. Male (7.0 mm, immature). Thistle Cove,

WA, 34°0'S, 122°12'E, 11 Apr 1984, 5.0 m, G.C.B. Poore and H.M.

Lew Ton (NMV J39710).

Paratypes. SA. 2 males (6.2 immature, 5.5 adult [crushed] mm),

north side of West 1., Encounter Bay, 35°37'S, 138°36'E, 21 Mar 1985,

5 m, sediment at base of Heterozostera, G.C.B. Poore and H.M.

Lew Ton (NMV J26202). Male (4.6 mm, immature), West L,

Encounter Bay, 28 Jan 1990, under boulder fauna, S.A. Shepherd

(SAM C5744).

Description of male. Body 1.8 times as long as greatest width

(including anterior and posterior processes), widest at pereo-

nites 1 and 5; dorsal surfaces of pereon, pleon and pleotelson

granular. Head anterior margin strongly anteriorly produced to

form anteriorly medially indented and bifid process, rostral

process and frons distinctly ventral in position; head about 1.4

times as long as pereonite 1 in lateral view; pereonite 1 dor-

sally with ill-defined transverse band of tubercles, laterally

with distinct boss and oblique thick longitudinal ridge; pere-

onites 2-4 narrower that pereonites 1 and 5, each with trans-

verse row of small low tubercles; pereonite 5 wider than 4 and

6, coxae with prominent boss; pereonites 5-7 without distinct

tubercles, pereonite 6 narrower than 7, pereonite 7 narrower

than 6. Pleon posteriorly rounded in dorsal view, posterodorsal

margin with low tubercles; sublateral pleonal ‘keys’ present.

Pleotelson strongly vaulted, posterior margin with strongly

produced dorsally arched process dorsal surface of which is

provided with irregularly shaped nodules.

Antennule peduncle article 1 2.4 times as long as wide,

about 8 times as long as article 2, anterior margin convex, pos-

terior margin straight, angled obliquely distally; article 3 about

as 0.5 times as long as article 1, about 3.6 times as long as wide,

4.0 times as long as article 2; flagellum 13-articled, extending

to posterior of pereonite 1, about twice as long article 3.

Antenna peduncle article 1 short, setose; articles 2 and 3 rela-

tively elongate, article 2 2.5 times as long as wide, article 3 0.6

times as long as article 4, 1.7 times as long as wide; article 3,

single long simple seta at superior distal angle; articles 2-A

collinear; article 4 0.8 times as long as article 5, 2.5 times as

long as wide, superior margin with scale- setae; article 5 2.6

times as long as wide; flagellum stout, about 0.9 times as long

as peduncle, extending to anterior of pereonite 2, with 11

articles.

Epistome anteriorly acute, with median constriction, surface

hregular. Mandibles with both incisors unicuspid; left mandible

with lacinia mobilis distally narrow, with 3 small cusps, spine

row of 4 curved serrate spines; right mandible spine row of 1

broad-based bifid, multidigitate spine and 5 distally serrate

spines; molar process round, crushing surface strongly ridged,

marginally serrate; with basal group of 3 long plumose setae;

palp not observed. Maxillule lateral lobe with 11 terminally

acute serrate RS on gnathal surface, twelfth seta set between

these. Maxilla lateral and middle lobes each with 6 curved

finely serrate RS, mesial lobe with about 12 serrate and biser-

rate RS. Maxilliped endite lateral margin strongly convex,

distal margin with 7 CP RS, 2 cactus setae, distomesial angle

with 1 simple RS; distomesial margin with 3 large stout CP RS,

increasing in size proximally; mesial margin of palp articles

2-5 with 9, 10, 12 and 8 long simple setae respectively; palp

lateral margins without long setae, with 1 short simple seta at

distal angle of article 3 and 4.

Pereopod 1 basis about twice as long as greatest width,

approximately twice as long as propodus; ischium 1.2 times as

long as propodus, 1.9 times as long as greatest width, superior

margin with 1 proximal and 1 distal acute short simple RS;

merus about 0.4 as long as ischium, 0.6 times as long as great-

est width, superior distal angle with 2 acute simple RS inferior

distal margin with 2 short bifid and 1 long acute simple; carpus

inferior margin 1.2 times as long as merus, distally with 2 short

bifid; propodus 1.5 times as long as greatest width, inferior

lateral margin with 2 short biserrate RS, inferior margin with 3

stout bifid RS; dactylus 0.7 times as long as propodus, inferior

margin with prominent serrate cuticular scales, secondary

unguis simple, with distal point. Pereopod 2 basis 3.0 times as

long as greatest width, margins with scattered scale-setae,

superior margin with weak distal flange; ischium 0.75 times as

long as basis, 2.8 times as long as greatest width, superior mar-

gin with 1 proximal and 1 distal acute short simple RS, merus

about 0.7 times as long as ischium, superior distal angle with 2

short acute RS, inferior margin with 2 short stout acute setae

and 1 long simple seta; carpus about as long as merus, superior

distal angle with 1 simple seta, inferior margin with 2 acute

simple setae, distal angle with 1 RS; propodus about as long as

ischium, 1.6 times as long as carpus, superior distal angle with

1 simple and 1 palmate setae, inferior margin with 3 short stout

acute RS; dactylus 0.5 times as long as propodus. Pereopods

5-7 similar, basis and ischium relatively longer than for pereo-

pod 2, distal margins of carpus with more and longer RS.

Pereopod 7 basis 4.4 times as long as greatest width, inferodis-

tal angle with single simple seta, superior margin with widely-

spaced small scale-setae; ischium 0.9 times as long as basis, 5.7

times as long as greatest width, superior margin with 1 proxi-

mal short acute RS, merus 0.4 times as long as ischium, super-

ior distal margin with 1 acute RS, inferior distal angle with 1

stout acute seta; carpus about as long as merus, anterodistal

angle with 3 long acute finely serrate and 1 simple RS, inferior

margin with 2 stout short acute RS, inferior distal angle 3

New Australian sphaeronnatrid isopod crustaceans

353

finely biserrate RS; propodus 1.5 times as long as carpus, 0.7

times as long as ischium, superior distal angle with 2 palmate

seta, inferior margin with 3 short stout acute RS; dactylus 0.4

times as long as propodus.

Penes mutually adjacent, twice as long as basal width;

mesial margin straight, lateral margin angled mesially, apex

bluntly rounded.

Pleopod 1 exopod and endopod with c. 17 and 16 PMS

respectively, endopod mesial margin setulose; endopod sub-

triangular, 0.9 times as long as exopod, 1.5 times as long as

greatest width; exopod lateral and mesial margins subparallel,

distal margin rounded. Pleopod 2 exopod and endopod with c.

18 and 27 PMS respectively; appendix masculina 11 times as

long as wide, slightly wider proximally, weakly sinuate, apex

bluntly rounded. Pleopod 3 exopod and endopod with c. 50 and

18 PMS respectively. Pleopod 4 exopod lateral margin with 3

fine simple setae, distal part triangular, both margins with fine

setae; endopod without setae. Pleopod 5 exopod with 2 distal

bi-lobed scale patches, lateral margin with scattered minute

simple setae; endopod with fine setae on distal margin only.

Uropod peduncle dorsal surface densely covered with small

nodules and minute hemispherical structures; endopod mesial

margin straight, lateral margin with distal half angled mesially,

distolateral margin subapically excavate; lateral margin

entirely fringed with expanded cuticular scales; exopod small,

0.2 as long as endopod and peduncle, setin to anterolateral

angle.

Female. Similar to male; ovigerous females not known.

New Australian sphaeromatrid isopod crustaceans

355

Etymology. From Greek periosus (immense) and nasus (nose),

alluding the hugely projecting anterior margin of the head;

noun in apposition.

Distribution. Thistle Cove, Great Australian Bight, WA, to

Encounter Bay, SA; intertidal to 5 m.

Remarks. This remarkable spindle-shaped isopod is easily rec-

ognized by the prominent anterior cephalic and pleotelson pro-

jections. No other species in the genus has such ornamentation,

and in addition the uropodal exopod is reduced to an incon-

spicuous small flat stub. These characters distinguish

Moruloidea perionasus from all other sphaeromatids except

perhaps Bregmotypta Bruce, 1994c, the only genus with similar

cephalic and pleotelsonic projections. There are many differ-

ences at generic level, but in Bregmotypta the cephalic

process is single and the pleotelson has two prominent bosses,

while in M. perionasus the anterior process is doubled, and the

posterior process is single and elongate.

Character states that support inclusion in Moruloidea are:

antenna with expanded and reflexed articles 4 and 5; close-set,

temiinally rounded short, flat penial processes; appendix mas-

culina arising basally, not distally narrowed and longer that

ramus; pereopod 1 much more robust than pereopods 2-7, the

propodus of which has a proximal extension; coxae 5 over-

lapping anteriorly and posteriorly; uropodal rami lamellar with

exopod smaller than endopod. Other characters such as mouth-

parts and pleopods present a consistent appearance with other

species of the genus.

The immature specimen was selected as holotype as it was

the most intact specimen. The propodus of pereopod 1 in this

juvenile lacks the prominent ‘heeT of the adult male although

that part of the inferior margin is weakly produced; the

dactylus lacks the scales on the inferior margin.

Pedinura gen. nov.

Type species. Pedinura flindersia sp. nov., here designated.

Diagnosis. Pleotelson posterior margin entire, posteriorly pro-

duced, without exit channel, without ventral depression or

groove. Antennule and antenna anteriorly positioned, peduncle

articles 1 and 2 flattened and expanded; antennule peduncle

articles 2 and 3 short, less than half as long as article 1 ; article

3 and flagellum collinear. Pereopods 1, 4-7 robust, pereopods

2 and 3 slender, pereopods 2 and 3 dactylus with prominent

comb seta set against unguis. Pleopods 1-3 with rami of about

equal length; pleopod 1 with axis of rami straight, not oblique.

Pleopod 2 appendix masculina medially inserted, extending

beyond distal margin of endopod. Pleopods 4 and 5 endopod

and exopod with thickened ridges. Uropods ventrolateral in

position, not visible in dorsal view; exopod minute, inserted

into lateral margin, stub-like.

Description of male. Body elongate, 3 times as long as greatest

width, moderately vaulted; dorsal surface smooth; lateral mar-

gins subparallel; unable to conglobate. Head weakly immersed

in pereonite 1; rostral process minute or absent. Eyes small, lat-

eral. Pereon segments without raised posterior margins. Coxal

plates fused, without discernable suture, overlapping anterior

over posterior. Membrana cingula absent. Pleon with 4

segments, pleonite 1 entire, 2 lateral sutures running to posteri-

or margin of pleon. Pleotelson posterior margin produced as

plate-like extension posterior to pleopod chamber; without

foramen, without ventral exit channel or groove. Pleonal

sternite present.

Antennule and antenna anteriorly positioned. Antennule

peduncles not separated by epistome; peduncle articles 1 and 2

expanded, anteriorly flattened; plane of articles 1 and 2 of pro-

jecting ventrally; peduncle articles 2 and 3 short, article 3

shorter than article 2, together about 0.5-0.8 as long as article

1; flagellum of 4 articles, slightly shorter than combined

lengths of articles 1 and 2. Antenna slender, peduncle articles 1

and 2 short, appearing fused, peduncle articles 4 longer than

articles 3 and 5; flagellum as long as or slightly longer than

peduncle.

Epistome wide, unornamented. Labrum unornamented.

Mandible incisor 3- or 4-cuspid; left mandible with lacinia

mobilis or without; molar process with or without marginal ser-

rations; palp article 1 longest, 3 shortest. Maxillule with lateral

lobe with 1 1 RS on gnathal surface, mesial lobe with 2 long and

1 short CP slender setae and 2 short simple seta (type species).

Maxilla with all articles well developed; lateral and middle

lobes with flat RS, mesial lobe with blunt and acute long RS,

some of which are basally CP. Maxilliped endite distally with

cactus and club setae, laterally with 1 long curved CP seta; palp

articles not mesially produced, mesial margins with numerous

setae, lateral margins without setae.

Pereopods 1, 5-7 robust, 2 and 3 slender; pereopods 2 and 3

dactylus with prominent pectinate secondary unguis opposing

unguis, all other pereopods with prominent recurved trifid

secondary ungul.

Penes short, not extending to pleopod peduncles; mutually

adjacent or slightly set apart.

Pleopods 1 with axis of both rami straight, not oblique,

about equal in length. Pleopod 1 not operculate, not indurate.

Pleopod 2 appendix masculina medially attached. Pleopods 3

exopod with transverse suture, pleopods 4 with or without

suture, pleopod 5 without. Pleopods 4 and 5 exopod and

endopods with transverse thickened ridges; pleopod 5 endopod

with 2 or 3 distal scale patches. Uropods endopod lamellar,

exopod minute, stub-like, set into anterolateral margin of

endopod.

Female. Antennule peduncle of type species articles 1 and 2

greatly expanded anteriorly. Brood pouch of overlapping oost-

egites arising on pereonites 2, 3 and 4. Mouthparts not meta-

morphosed.

Composition and distribution. Pedinura flindersia sp. nov.;

Pedinura rnokari sp. nov.; subtropical WA to Vic., Australia.

Etymology: A combination of Greek pedinos (flat, even), and

oura (tail), alluding to the flattened pleotelson of the two

species (feminine).

Remarks. The characters that best distinguish Pedinura are: the

plate-like extension to the posterior margin of the pleotelson

entirely lacking any exit channel, groove or depression; ventral

uropods (not visible in dorsal view) with a minute exopod;

356

N. L. Bruce

expanded articles to antennule peduncle articles 1 and 2; and

the appendix masculina being mesially inserted and extending

beyond the distal margin of the exopod.

Pedinura resembles Amphowidea Milne Edwards, 1840, A.

angustata Baker, 1908 being the most similar. Although

Amphowidea and its species have not been fully described, the

two genera can immediately be separated by Amphowidea

having anterolateral uropods, prominent in dorsal view and

extending well beyond the posteriorly narrowed pleotelson.

Further points of distinction include the appendix masculina

being basal (mesial in Pedinura), pleopod 1 endopod distinctly

triangular with an indurate mesial margin, and the uropod with

both rami large and lamellar.

Cassidinopsis Milne Edwards, 1840 is similar (Brandt,

1998) but in that genus the antennule peduncular articles 1 and

2 are not expanded, pereopod 2 lacks the pectinate robust seta

opposite the dactylus, and most notably the uropods are lateral

(not ventral) and extend well to the posterior of the pleotelson

which is not posteriorly produced.

Pedinura flindersia sp. nov.

Figures 35-39

Material examined. Holotype. Male (3.8 mm), “The Hotspot” reef, 5 n.

miles W of north end of Flinders L, SA, 33°40.50'S, 134°22'E, 19 Apr

1985, 17 m, assorted red algae, S. Shepherd (NMV J39728).

Paratypes. Vic. 3 females (immature 2.0, 2.2, 2.3 mm), 2 mancas

(1.1, 1.2 mm). Whalers Point Lighthouse, Portland, 38°20.5'S,

141°37.5'E, 1 May 1988, 10 m, brown algae from boulder bottom,

R.T. Springthorpe and P.B. Berents (AM P50944). Male (5.0 mm),

38°40'S, 145°35T, 6 Mai- 1982, 0 m, rocky, G.C.B. Poore (NMV

J26380). Female (4.2 mm), 2 mancas (1.5, 2.5 mm), 500 m offshore, 1

km E of Harmers Haven, 38°34'S, 145°40'E, 6 Mai- 1982, 11 m, rocky,

C. Larsen and G. Barber (NMV J26377). Male (4.1 mm), female (3.0

mm), 300 m offshore, E of Harmers Haven, 38°34'S, 145°40'E, 6 Mar

1982, 6 m, rocky, R.S. Wilson and C. Larsen (NMV J26381). Male

(5.5. mm), female (3.5 mm), 50 m offshore, E side of South Point,

Twin Reefs, 38°4LS, 145°39'E, 4 Mar 1982, 11 m, rocky, C. Larsen,

G. Barber and R.S. Wilson (NMV J26383). Male (4.2 mm). The Oaks,

Bunurong Coast, 38°40'S, 145°38^E, 5 Mar 1982, rock, G.C.B. Poore

(NMV J26375). Male (3.1 mi-n). Eagles Nest, Venus Bay, 38°40'S,

145°40'E, 5 Mar 1982, rock, G.C.B. Poore (NMV J26378). Eemale

(5.2 mm), NW side of Henty Reef, Mounts Bay, 38°47.0'S,

143°40.5'E, 3 May 1988, 18 m, red algae on boulder, R.T.

Springthorpe and PB. Berents (AM P50946). SA. 2 males (3.3 dis-

sected, 3.4 mm), 6 females (ovigerous 3.7, 4.4, 5.0 dissected, non

ovigerous 2.9, 3.0, 3.1 mm), immature (2.4, 2.6, 2.8, 3.0, 3.8 mm),

mancas (1.7, 1.7, 1.8 mm), same data as holotype (NMV J39721). 2

males (3.0, 3.2 mm), 2 females (3.4, 3.6 nrai), Snapper Point,

Beachport, 37°29.3'S, 139°59.6'E, 14 May 1990, 6.0 m, brown algae,

on limestone reef, G.C.B. Poore (NMV J26231). WA. Eemale (3.0

mm). Seven Mile Beach, N of Dongara, 29°12'S, 114°53'E, 24 Apr

1986, 1 m, Amphibolus epiphytes, G.C.B. Poore and H.M. Lew Ton

(NMV J26173). Female (4.1 mm). Cliff Head, S of Dongara, 29°32'S,

114°59'E, 22 Apr 1986, 1 m, red algae on limestone, G.C.B. Poore and

H. M. Lew Ton (NMV J26158). Female (3.9 mm), off jetty. Green I.,

Rottnest I., 32°0ES, 115°30'E, 21 Dec 1983, 1 m, mixed algal turf on

rock, R.T Springthorpe (AM P50943).

Other material, unmeasured. Vic. Bay of Islands, 38°35.0'S,

142°49.5'E, 2.5 m, red algae (AM P50945). Portland, 38°24^S,

141°40.5'E, 23m, Herdmania niomus with encmsting sponge and red

algae (AM P50947). SA. Topgallant L, Investigator Group, 33°43'S,

133°36.60'E, 20 m, on Cystophora and Plocamium (NMV J39730).

“The Hotspot” reef, N end of Flinders I., 33°40.50'S, 134°22'E, 12 m,

assorted algae (NMV J39706); 21 m, red algae (NMV J39722). 7, 2.3

n. miles W of Tiparra Light, Tiparra Reef, Tiparra Bay, 15 Mar 1985,

10 m, red algae (NMV J39727). WA. Port Denison Beach, Dongara, 5

m, red algae, mainly Laurencia (NMV J39704).

Description of male. Body about 3.2 times as long as greatest

width, lateral margins subparallel, pereonites of subequal

width, pereonite 4 marginally widest; dorsal surfaces smooth,

unomamented. Head anterior margin without transverse ridge,

rostral process not visible in dorsal view. Head and pereonites

1 subequal in length in dorsal view, pereonites

1>2=3<4=5>6>7. Pleotelson posterior margin smoothly

rounded.

Antennule peduncle article 1 1.4 times as long as proximal

width, about 2.6 times as long as article 2; article 2 flattened,

about as long as wide, anterior margin with single small seta,

posterodistal angle with 2 small setae; article 3 about half as

long as article 2; flagellum 4-articled, extending to anterior of

pereonite 1, about 0.4 times as long article 3. Antenna peduncle

articles 1 and 2 about as long as combined lengths of article 3

and 4; article 4 about as long as article 5, about 1.3 times as

long as wide; flagellum about equal in length to peduncle,

extending to anterior margin of pereonite 2, with 6 articles.

Epistome smooth, narrow, laterally encompassing labrum,

anteriorly forming distinct point. Mandible incisor with 3

cusps; lacinia mobilis absent; spine row of 3 serrate curved

spines, on right mandible with additional broad-based multi-

digitate spine; molar process appearing largely smooth, without

serrate margins or ridged surfaces; palp article 1 longest, with-

out setae; article 2 with 3 biserrate setae, article 3 with 4 biser-

rate setae, terminal seta being largest. Maxillule mesial lobe

with 2 long and 1 shorter weakly CP and 1 short simple seta;

lateral lobe with 9 peripheral simple RS on gnathal surface,

eleventh seta set between these. Maxilla lateral lobe and mid-

dle lobe each with 4 curved flat simple RS respectively, mesial

lobe with about (not clearly observed) 6 RS, variously serrate.

Maxilliped endite extending about half way along palp article

3, distal margin with 1 conical RS, 3 rounded RS, 1 cactus RS

and laterally with 2-3 slender CP RS; palp articles 2-5 with

about 4, 9, 8 and 11 setae respectively.

Pereopod 1 basis about 1.6 times as long as greatest width,

about 1.1 times long as propodus; ischium 0.6 times as long as

basis, 1.8 times as long as greatest width, inferior surfaces with

abundant microtrichs; merus about 0.7 times as long as isch-

ium, 0.7 times as long as greatest width; carpus (inferior mar-

gin) 0.7 as long as merus, 0.8 times as long as wide; propodus

1.5 times as long as ischium, 1.9 times as long as greatest

width, inferior margin with 2 biserrate RS, inferodistal margin

with 2 biserrate RS and 2 short simple setae adjacent to base of

dactylus; dactylus 0.4 times as long as propodus, unguis

strongly recurved, inferior margin with few small cuticular

scales, secondary unguis recurved with 2 basal cusps. Pereopod

2 with all articles slender; basis about 3.1 times as long as

greatest width, about as long as propodus; ischium 0.8 times as

long as basis, 2.5 times as long as greatest width, inferodistal

angel with single seta; merus about 0.7 times as long as

ischium, 1.7 times as long as greatest width, superior distal

360

N. L. Bruce

margin strongly convex; carpus 1.7 as long as merus, 4 times as

long as wide; propodus 1.3 times as long as ischium, 5.3 times

as long as greatest width; dactylus 0.4 times as long as pro-

podus, unguis slender apically falcate, secondary unguis flat

and strongly pectinate extending alongside ungul. Pereopod 3

similar to 2, but a little more robust. Pereopods 4-7 similar to

pereopod 1, but carpus quadrate rather than subtriangular,

propodus inferior margin without biserrate RS; all pereopods

without trifid and biserrate setae. Pereopod 7 basis with

proximal flange; carpus distal margin with single simple

acute RS.

Penial processes separate, adjacent, about 1.7 times as long

as basal width, quadrate in shape, distal margin weakly oblique,

feebly indented.

Pleopod 1 exopod and endopod with c. 14 and 12 PMS

respectively; rami of similar size, endopod 0.9 as long as

exopod, 1.8 times as basal width, lateral margin converging

slightly to rounded distal margin; exopod 2.0 time as long as

greatest width, margins subparallel, distal margin broadly

rounded. Pleopod 2 exopod and endopod with c. 18 and 12

PMS respectively, pleopod 2 endopod 1.2 times as long as

exopod, 2.3 times as long as greatest width, lateral margin

strongly recessed at insertion point of appendix masculina;

appendix masculina attached about half way along mesial mar-

gin, lateral surface with abundant microtrichs, about 5.3 times

as long as wide, 0.8 times as long as endopod, apex bluntly

rounded. Pleopod 3 exopod and endopod with c. 24 and 11

PMS respectively; both rami elongate, endopod very slightly

(1.06) longer than exopod, slender, 2.9 times as long as great-

est width, proximal lateral margin convex. Pleopod 4 both rami

with prominent ridges, exopod lateral margin with fine setae,

exopod suture incomplete. Pleopod 5 both rami with prominent

New Australian sphaeromatrid isopod crustaceans

361

Figure 39. Pedinura flindersia sp. nov. SEMs. Female, ‘The Hotspot’ reef, Flinders I., NMV J39722. A, anterior, lateral view; B, mandible; C,

maxilliped, distal margin; D, pereopod 1 dactylus; E, pereopod 2, pectinate robust seta on propodus; F, uropod lateral margin; G, sensory setae,

uropods.

New Australian sphaeronnatrid isopod crustaceans

363

ridges; exopod with 2 distal scale patches, and transverse

absent. Uropod rami flattened, margins provided with mebrana

cingula; exopod inserted into lateral margin at about one-third

from posterior margin, 02 times as long as fused endopod,

anterior margin convex, posterior weakly concave, distal

margin narrowly rounded; endopod anterolateral margin

straight, posterolateral part subtruncate, mesial margin weakly

convex; broadly rounded, proximomesial cluster of 3 sensory

setae.

Female. Anterior body outline includes conspicuously

expanded antennule peduncular articles 1 ; rostral point evident

in dorsal view; eyes smaller than in males. Antennule peduncle

article 1 0.8 times as long as mesial width, about 2.6 times as

long as article 2, anteriorly produced, plate-like; article 2 flat-

tened, about as long as wide, anterior margin produced,

anteromesial angle at right angles, forming continuous margin

with peduncular article 1 ; article 3 anterior margin with small

distinct anterior boss. Antenna as for the male.

Size. Males 3. 1-5.5 mm, ovigerous females 3. 7-5.0 mm,

non-ovigerous females 2.9-5. 2 mm; juvenile (sex indeter-

minate) 2.4-3. 8 mm; mancas 1. 1-2.5 mm.

Etymology. From the type locality; noun in apposition.

Distribution. Vic. (Bunerong), SA, WA (Rottnest L, Seven Mile

Beach, Dongara); intertidal to 21 m; mainly on red and brown

algae, specifically on Cystophora, Laurencia and Plocamium.

Remarks. This species is distinguished by the posterior margin

of the pleotelson being widely subtruncate and without a

median indentation. Further distinguishing characters include

the strongly expanded antennule peduncle articles 1 and 2, the

anteriorly acute epistome not visible in dorsal view, robust

pereopods and the uropod exopod being set distally on the

endopod.

Pedinura mokari sp. nov.

Figures 40-42

Material examined. Holotype. Male (3.2 mm). Snapper Point,

Beachport, SA, 37°29.3'S, 139°59.6T, 14 May 1990, 6.0 m, brown

algae ete., G.C.B. Poore and R.S. Wilson (NMV J39714).

Paratypes. Vic. Male (3.1 mm), wharf at Port Campbell, 38°37.5"S,

142°59.5"E, 8 Apr 1988, 5 m, wood and encrusting algae from wharf

pile, R.T. Springthorpe and PB. Berents (AM P50942). Male (3.2.

mm), ?female (2.6 mm), juv. (1.8, 2.0 mm), 50 m offshore, E side of

South Point, Twin Reefs, 38°41"S, 145°39"E, 4 Mar 1982, 11m, rocky,

C. Larsen, G. Barber and R.S. Wilson (NMV J26228). ?female (3.0

Eigure 41. Pedinura mokari sp. nov. Male paratype NMV J39721. A-C, pereopods 1, 2 and 7.

364

N. L. Bruce

Figure 42. Pedinura mokari sp. nov. Male paratype NMV J39721. A-E,

mm), SW of Eagles Nest, Venus Bay, 38°40'S, 145°40'E, 5 Mar 1982,

8 m, rocky, R.S. Wilson and G. Barber (NMV J26226). 6 females

(ovigerous 3. 0-3. 5 mm, non-ovigerous 3. 1-3.2 mm), E side of Cape

Paterson, 38°4rS, 145°36'E, 5 Mar 1982, 6 m, rocky, R.S. Wilson,

G. Barber et al. (NMV J26334).

Description of male. Body about 3.0 times as long as greatest

width, lateral margins subparallel, pereonites of subequal

width; dorsal surfaces smooth, unornamented. Head anterior

margin without transverse ridge, rostral process present. Head

1.5 times as long as pereonite 1, pereonites 1>2=3=4=5>6>7.

Pleotelson lateral margins straight, converging towards

posterior, posterior margin flat with median excavation.

Antennule peduncle article 1 1.3 times as long as proximal

width, about 2.0 times as long as article 2; article 2 flattened,

0.9 times as long as wide; article 3 about 0.9 times as long as

article 2; flagellum 5-articled, extending to anterior of

pereonite 1, about as length of articles 2 and 3 combined.

Antenna similar to type species, flagellum with 9 articles.

Epistome smooth, narrow, laterally encompassing labrum,

widest anteriorly with distinct and convex anterior margin,

lateral margins shallowly concave. Mandible incisor with 4

cusps; lacinia mobilis present; spine row of 4-5 serrate curved

spines; molar process appearing large, with serrate margins or

ridged surfaces; palp article 1 longest, without setae; article 2

without setae, article 3 with 7 biserrate setae, distal 2 setae

being largest. Maxillule mesial lobe with 4 long and 2 shorter

CP; lateral lobe with 10 peripheral simple RS on gnathal

surface. Maxilla lateral lobe and middle lobe each with 4

curved, flat, finely serrate RS respectively, mesial lobe with

about (not clearly observed) 8 RS, variously serrate. Maxilliped

endite extending about half way along palp article 3, distal mar-

gin with 1 conical RS, 2 rounded RS, 1 cactus RS and 2-3

New Australian sphaeromatrid isopod crustaceans

365

slender CP RS; palp articles 2-5 with about 5, 9, 10 and 9 setae

respectively.

Pereopod 1 basis about 2.3 times as long as greatest width,

about 1.4 times long as propodus, widest proximally; ischium

0.6 times as long as basis, 2.58 times as long as greatest width,

inferior margin with 2 acute simple setae, one set at distal

angle; merus about 0.4 times as long as ischium, 0.8 times as

long as greatest width, inferior margin with 2 acute RS; carpus

(inferior margin) 0.7 as long as merus, 0.7 times as long as

wide, inferodistal angle with 2 simple setae; propodus 12 times

as long as ischium, 32 times as long as greatest width, inferior

margin with single proximal seta, inferomesial margin with 2

biseiTate RS, inferodistal margin with 2 simple RS adjacent to

base of dactylus; dactylus 0.5 times as long as propodus, unguis

strongly recurved, inferior margin with few small cuticular

scales, secondary unguis recurved with 2 basal cusps. Pereopod

2 with all articles slender; basis about 6.7 times as long as

greatest width, about 1.3 times as long as propodus; ischium

0.8 times as long as basis, 4.7 times as long as greatest width,

inferodistal angel with single seta; merus about 0.7 times as

long as ischium, 3.1 times as long as greatest width, superior

distal margin convex, inferodistal angle with single slender

seta; carpus 1.1 as long as merus, 4.7 times as long as wide,

inferodistal angle with 2 slender setae; propodus about as long

as ischium, 5.7 times as long as greatest width; dactylus 0.6

times as long as propodus, unguis slender, apically falcate,

secondary unguis flat and strongly pectinate, extending along-

side ungul. Pereopod 3 similar to 2, but a little more robust.

Pereopods 4-7 similar to pereopod 1, but more slender, carpus

quadrate rather than subtriangular, propodus inferior margin

without biserrate RS; all pereopods without trifid and biserrate

setae. Pereopod 7 basis with proximal flange; carpus distal

margin with 3 biserrate acute RS.

Penial processes separate, about 1.3-1. 6 times as long as

basal width, tapering distally to rounded apex.

Pleopod 1 exopod and endopod with c. 14 and 12 PMS

respectively; rami of similar size, endopod 0.9 as long as

exopod, 1.6 times as long as basal width, lateral margin

converging slightly to rounded distal margin; exopod 1.5 times

as long as greatest width, margins convex, distal margin

broadly rounded. Pleopod 2 exopod and endopod with c. 20 and

11 PMS respectively, pleopod 2 endopod about as long as

exopod, 2.0 times as long as greatest width, lateral margin

strongly recessed at insertion point of appendix masculina;

appendix masculina attached about one-quarter of way along

mesial margin, about 4.4 times as long as wide, 1.1 times as

long as endopod, apex bluntly rounded. Pleopod 3 exopod and

endopod with c. 22 and 12 PMS respectively; both rami elon-

gate, endopod very slightly (1.04) longer than exopod, 2.3

times as long as greatest width, proximal lateral margin convex.

Pleopod 4 both rami with obscure ridges, exopod lateral margin

with fine setae, exopod suture complete. Pleopod 5 both rami

with indistinct ridges; exopod with 3 distal scale patches, trans-

verse suture absent. Uropod rami flattened, margins provided

with mebrana cingula; exopod inserted into lateral margin at

about one-quarter from proximal margin, 02 times as long as

fused endopod, triangular, apex subacute; endopod antero-

lateral margin weakly convex, posterolateral part rounded.

mesial margin straight; mesially with 3 cluster of 2 sensory

setae.

Female. Sexes similar, the dimorphism of the type species

apparently not present in this species

Size. Males 3. 1-3.2 mm, ovigerous females 3.0-3. 5 mm,

non-ovigerous females 3. 1-3.2 mm.

Etymology. Mokari is an Aboriginal word meaning new; noun

in apposition.

Distribution. Vic. and SA, where it is sympatric with the more

widely distributed P. flindersia\ 5-11 m, rocky habitats.

Remarks. Pedinura mokari is separated from the only congener,

P. flindersia by the posteriorly indented pleotelson, the epis-

tome being widest anteriorly and visible in dorsal view and the

weakly expanded antennular peduncle articles 1 and 2.

Acknowledgements

I thank Drs Gary Poore, Robin Wilson and Elycia Wallis

(Museum Victoria, Melbourne); Penny Berents (Australian

Museum); and Wolfgang Zeidler (South Australian Museum,

Adelaide) — for loans of large amounts of unidentified

sphaeromatid material and for waiting patiently while I worked

on these valuable collections. I also thank Kim Larsen for ink-

ing the drawings and Geert Brovad (both Zoologisk Museum,

Copenhagen) for producing prints of the SEM photograph. I

thank Janet Bradford-Grieve (NIWA, Wellington) for her

comments on the manuscript.

This is the concluding contribution from Australian

Biological Resources Study Grant ABRS 89/1844, and was

completed in part at the Zoologisk Museum, University of

Copenhagen. This publication acknowledges National Science

Eoundation award DEB 997 8 193.

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Appendix. Species currently placed in Exosphaeroma

Stebbing, 1900

Many species of Exosphaeroma, including all those described

by Keppel H. Barnard from southern Africa and all those

described by W.H. Baker from southern Australia during the

first half of the twentieth century (13 of the 32 included here),

need redescription, but only those that are particularly

inadequately described or are of uncertain generic status are

mentioned as such. Australian species are shown in bold.

Exosphaeroma agmokara sp. nov. Type locality — Broken

Head, Northern NSW Australia; intertidal, under rocks on

sand.

Exosphaeroma aliae Baker, 1926. Type locality — Victor

Harbour, South Australia.

Exosphaeroma alveola sp. nov. Type locality — ^Jervis Bay,

New South Wales, Australia; intertidal.

Exosphaeroma amplicauda (Stimpson, 1857). Type locality —

Monterey Bay, California; in need of redescription

[original combination Sphaeroma].

Exosphaeroma antikraussi Barnard, 1940. Type locality —

Barnard cited several locations in South Africa.

Exosphaeroma bicolor Baker, 1926. Type locality — Kangaroo

Island, South Australia.

Exosphaeroma brevitelson Barnard, 1914. Type locality — Sea

Point, near Cape Town, South Africa; intertidal.

Exosphaeroma bruscai Espinoza-Perez and Hendrickx, 2001.

Type locality — Los Arcos, Jalisco, Pacific Mexico;

among inter- tidal algae and rocks to a depth of 3 metres.

Exosphaeroma diminutum Menzies and Frankenberg, 1966.

Type locality — Sapelo Island, Georgia, USA;

Chesapeake Bay to Florida and Venezuela (Kensley and

Schotte 1989); this species does entirely conform to the

genus.

Exosphaeroma echinensis Hurley and Jansen, 1977. Type

locality — Kaikoura, New Zealand; intertidal and subtidal.

Exosphaeroma estuarium Barnard, 1951. Type locality —

Umbagaga Estuary, near Umkomaas, Natal coast. South

Africa; among Zostera seagrass.

Exosphaeroma falcatum Tattersall, 1921. Type locality —

Spirits Bay, North Cape, New Zealand; subtidal to 20 m.

Exosphaeroma gigas (Leach, 1818). Type species. Type local-

ity — unknown {'Son pays inconnu.’)', this species has a

wide reported distribution in the Southern Hemisphere,

including Macquarie Island and Tasmania, but from the

time that the genus was established (Stebbing 1900; more

recently Poore et al. 2002) there has been at least a

subjective doubt that all material under this name is

correctly identified [original combination Sphaeroma]

Exosphaeroma hylecoetes Barnard, 1940. Type locality —

Barnard cites several locations in South Africa.

Exosphaeroma inornata Dow, 1958. Type locality — Palos

Verdes, Los Angeles County, California, under kelp hold-

fasts.

Exosphaeroma kraussi Tattersall, 1913. Type locality —

Saldanha Bay, Cape Town, South Africa; the uropodal

exopod is clearly figured as serrate, and the species is in

need of redescription.

Exosphaeroma laevis (Baker, 1910). Type locality — Gulf of

Saint Vincent, South Australia [original combination

Zuzara].

Exosphaeroma laevisculum (Heller, 1868). Type locality —

South Africa [original combination Sphaeroma.

Exosphaeroma media George and Stromberg, 1968. Type

locality — San Juan Island, Washington, USA.

Exosphaeroma montis (Hurley and Jansen, 1978), comb. nov.

Type locality — Mount Maunganui, New Zealand; inter-

tidal; conforms well with the diagnosis for Exosphaeroma

[original combination Cymodopsis] and is here trans-

ferred to that genus; see ‘Remarks’ for Exosphaeroma

alveola sp. nov. for comments.

Exosphaeroma obtusum (Dana, 1853). Type locality — Bay of

Islands, New Zealand; widely recorded around New

Zealand; also Stewart Island, Auckland Island, the Snares

Islands, Campbell Island and Chatham Island (Hurley

and Jansen 1977); intertidal [original combination

Sphaeroma].

Exosphaeroma octoncum (Richardson, 1899). Type locality —

Monterey Bay, California [original combination

Sphaeroma] ; in need of redescription.

Exosphaeroma pallidum Barnard, 1940. Type locality —

Woodstock Beach, Table Bay, South Africa; also

recorded from southern Australia.

Exosphaeroma parva Chilton, 1924. Type locality — ChiUca

Lake, India; in need of redescription.

Exosphaeroma planum Barnard, 1914. Type locality — Sea

Point, near Cape Town, South Africa; intertidal.

Exosphaeroma planulum Hurley and Jansen, 1977. Type

locality — Heathcote-Avon Estuary, Christchurch, New

Zealand; intertidal, estuarine; a replacement name for

homonym Exosphaeroma planum Hurley and Jansen,

1970.

New Australian sphaeromatrid isopod crustaceans

369

Exosphaeroma porrectum Barnard, 1914. Type locality — South

Africa; intertidal to 5 metres.

Exosphaeroma rhomburum (Richardson, 1899). Type

locality — California [original combination Sphaeroma].

Exosphaeroma serventii Baker, 1928. Type locality — Pallinup

Estuary, on the Great Australia Bight coast of Western

Australia.

Exosphaeroma studeri Vanhoffen, 1914. Type locality — Punta

Arenas, Chile.

Exosphaeroma truncatitelson Barnard, 1940. Type locality —

Kleinmond, Cape Province, South Africa.

Exosphaeroma varicolor Barnard, 1914. Type locality —

Woodstock Beach and Sea Point, near Cape Town, South

Africa.

Incertae sedis

Exosphaeroma alba Menzies and Glynn, 1968. Puerto Rico.

This species and its named variety E. alba chromata

Menzies and Glynn, 1968 were provisionally placed in

Exosphaeroma (as 1 Exosphaeroma) by Menzies and

Glynn (1968). The morphology or the antennule pedun-

cle, penial processes, shape and orientation of pleopod 1

rami, appendix masculina, and pleonal sutures (which run

to the posterior margin of the pleon) and the pleotelson

with a distal incision (or exit channel) all preclude its

inclusion in Exosphaeroma, and suggest a closer affinity

with Dynamenella. Remarkably, Menzies and Glynn

(1968) considered the most similar species to be a species

of Dynamenella, yet placed their new species in

Exosphaeroma.

Exosphaeroma antillense Richardson, 1912. Type locality —

Montego Bay, Jamaica (Kensely and Schotte, 1989); the

bidomed pleotelson together with the short uropodal exo-

pod and overlapping oostegites suggest that this is not a

species of Exosphaeroma (as suggested by Kensley and

Schotte 1989).

Exosphaeroma antarctica Richardson, 1908. Type locality —

Port Madryn, Patagonia; the description and figures are of

insufficient detail to distinguish this species from similar

species such as E. gigas and E. obtusum.

Exosphaeroma aphrodita Boone, 1923. Type locality — La

Jolla, California. This species has never been illustrated,

and from the description given by Boone it is not possi-

ble to assess its status. Boone’s description describes the

dorsal surfaces of the body as being highly nodular and

ornamented.

Exosphaeroma productatelson Menzies and Glynn, 1968.

Puerto Rico. This species is very similar to E. alba, and

the comments given for that species also apply entirely to

this species, except the posterior margin of the pleotelson

is entire.

Exosphaeroma yucatanum (Richardson, 1901). Included in

Exosphaeroma by Kensley and Schotte (1989). The trilo-

bate pleotelson and tri-domed pleotelson dorsum would

indicate that this is not a species of Exosphaeroma.

Exosphaeroma coatsii Tattersall, 1913. Port Stanley, Falkland

Islands. This species is based on females and has promi-

nent nodules across the pereonites, pleon and pleotelson.

The uropods have smooth margins, and the apex is of the

pleotelson is acute. I am unaware that this species has

been placed into synonymy, but as it is based on females

there is the possibility that it is a species of Isocladus. Not

included in Nierstrasz’s (1931) listing of species.

Exclusions

Exosphaeroma crenulatum Richardson, 1902. Synonym of

Dynamenella perforata (Moore, 1902) (see Glynn, 1974).

Exosphaeroma pulchellum Colosi, 1921. Synonym of

Sphaeroma hookeri (Leach, 1818) (see Jacobs, 1987).

Exosphaeroma calcareum (Dana, 1853). At times placed in

Exosphaeroma (e.g. Nierstrasz, 1931), more recently

retained in Isocladus (e.g. Menzies, 1962).

Memoirs of Museum Victoria 60(2); 371-416 (2003)

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

http://www.museum.vic.gov.au/memoirs

Neastacilla Tattersall, 1921 redefined, with eight new species from Australia

(Crustacea: Isopoda: Arcturidae)

Rachael A. King

Museum Victoria, PO Box 666E, Melbourne, Vic. 3001, Australia, and Department of Zoology, University of Melbourne,

Parkville, Vic. 3052, Australia

Present address: Southeastern Regional Taxonomic Center, Marine Resources Research Institute, PO Box 12599,

Charleston, SC 29422-2599, USA (rachaelking@crustacea.net)

Abstract King, R.A. 2003. Neastacilla Tattersall, 1921 redefined, with eight new species from Australia (Cmstacea: Isopoda:

Arcturidae). Memoirs of Museum Victoria 60(2): 371-416.

The arcturid isopod genus Neastacilla is rediagnosed and eight new species from Australia {Neastacilla coonabooloo

sp. nov., Neastacilla kanowna sp. nov., Neastacilla lawadi sp. nov., Neastacilla marrimarri sp. nov., Neastacilla soelae

sp. nov., Neastacilla tarni sp. nov., Neastacilla tharnardi sp. nov. and Neastacilla yuriel sp. nov.) are described. Five

species from Australia and New Zealand are rediagnosed. A key to differentiate Australian and New Zealand species is

presented.

Keywords Cmstacea, Isopoda, Arcturidae, Neastacilla, taxonomy, Australia

Table of Contents

Introduction 371

Neastacilla 372

Key to Australian and New Zealand species of Neastacilla 374

Neastacilla algensis 375

Neastacilla antipodea 375

Neastacilla attenuata 376

Neastacilla coonabooloo sp. nov 376

Neastacilla deducta 380

Neastacilla fusiformis 380

Neastacilla inaequispinosa 384

Neastacilla kanowna sp. nov 384

Neastacilla lawadi sp. nov 390

Neastacilla levis 390

Neastacilla macilenta 395

Neastacilla marrimarri sp. nov 395

Neastacilla monoseta 396

Neastacilla sheardi 401

Neastacilla soelae sp. nov 401

Neastacilla tarni sp. nov 401

Neastacilla tattersalli 405

Neastacilla tharnardi sp. nov 405

Neastacilla tuberculata 409

Neastacilla yuriel sp. nov 409

Acknowledgments 414

References 414

Introduction

The valviferan isopod family Arcturidae Dana, 1849 is repre-

sented in Australia by four genera. Parastacilla Hale, 1924 is an

endemic Australian genus, its four species reviewed by King

(2000). Amesopous Stebbing, 1905 is monotypic and its

species, A. richardsonae Stebbing, 1905, was recorded from

tropical Australia by Poore et al. (2002). Astacilla Cordiner,

1793 is represented by one species (King, 2003). The

Australian fauna is dominated by species of Neastacilla

Tattersall, 1921, here reviewed.

The systematics of Neastacilla has always been problemat-

ical. The genus was established by Tattersall (1921) for New

Zealand specimens of 'Astacilla falclandica Ohlin’, a species

originally described from the Falkland Islands. TattersalTs

record was shown to be a misidentification and TattersalTs

species is now Neastacilla tattersalli (Lew Ton and Poore,

1986a). Tattersall (1921) differentiated Neastacilla from

Astacilla Cordiner, 1793 primarily on the absence of an antero-

lateral expansion of the head and pereonite 1 over the mouth-

parts (apparent in Astacilla) and by the completely fused pleon

(as opposed to visible pleonite segmentation in Astacilla).

These characters were later shown to be variable as more

species were described (Nordenstam, 1933; Hale, 1946) and the

newer genus did not become widely accepted.

Nordenstam (1933) was the first to rediagnose Neastacilla.

Modifying TattersalTs diagnosis, Nordenstam argued that up to

372

Rachael A. King

three pleonite segments could be detected “indicated by shal-

low grooves,” the first pereopod lacked an unguis, and the

endopod of the uropod bore one seta. He also suggested (erro-

neously it turns out) that Astacilla amblyura Stebbing belonged

in Neastacilla. Nordenstam acknowledged the lack of descrip-

tive information concerning key Neastacilla characters and

despite his diagnosis was not fully convinced of the validity of

Neastacilla. He believed that Neastacilla would be made

redundant when more detailed information was known or when

species intermediate between Astacilla and Neastacilla were

found.

Later, Hale (1946) documented arcturid species from

Australia, New Zealand and the Southern Ocean. He described

five new species, which he placed in Astacilla, and transferred

his two previously described species of Neastacilla (N. algen-

sis, N. deducta) to Astacilla. While acknowledging the mor-

phological similarities of his new species to Neastacilla, as

defined by Nordenstam (1933), Hale argued that Neastacilla

was ambiguously defined; all his specimens possessed two

uropodal endopod setae and so he placed them in Astacilla.

Despite reservations, neither Hale (1946) nor Guiler (1949)

who later described Tasmanian species, attempted to syn-

onymise Neastacilla. Birstein (1963) was the first to argue that

Neastacilla should be synonymised with Astacilla on the basis

of its variable characters. Later, Monod (1970) synonymised

Neastacilla with Astacilla in his partial revision of Astacilla.

Following extensive examination of material from the

north-west Pacific, Kussakin (1972) re-established the genus,

placing more importance on the morphology of the anterior

pereopods. He defined Neastacilla as including those taxa pos-

sessing a dactylus and an unguis on pereopod 1 and possessing

shortened dactyli, without ungui, on pereopods 2-4. This was

compared to Astacilla, which he defined as including those

species lacking a dactylus on pereopod 1 and lacking dactyli on

pereopods 2-4. He subsequently re-assigned all Pacific Ocean

species of Astacilla to Neastacilla.

Kussakin’ s views were supported by H. M. Lew Ton

(unpublished B. Sc. (Hons) thesis. Monash University, 1980)

who studied Australian species of Neastacilla. She concluded

that the characters Kussakin used to separate the two genera

were valid but should be modified slightly, as some Australian

species of Neastacilla lack dactyli on pereopods 2-4 (some

species with dactyli on pereopod 4 only and others without

dactyli on pereopods 2-4). The loss of the dactyli on pereopods

2-4 occurs elsewhere in Arcturidae (in Parastacilla) and in

another valviferan family, Chaetiliidae (in Chaetilia Dana) and

is considered a synapomorphy for each genus. Lew Ton found

biogeographical support for the distinction of Astacilla from

Neastacilla. Most species of Neastacilla are found in the

Pacific and those of Astacilla in the Atlantic Ocean,

Mediterranean and Indian Ocean.

Without the evidence of Lew Ton’s comprehensive unpub-

lished work, other authors questioned Kussakin’s (1972) con-

clusions. Schultz (1981) attempted to synonymise Neastacilla,

placing all the known species into Astacilla, Arcturus Latreille

or Arcturella Sars as he regarded the presence of an unguis on

pereopod 1 as a variable character. Menzies and Kruczynski

(1983) argued that the elongation of pereonite 4 was a more

useful character than whether or not an unguis was present on

pereopod 1 and also treated Neastacilla and Astacilla as

synonyms.

While characters used in diagnosing the genus remain

debatable, Kussakin’s observations provide support for the

recognition of Neastacilla as a genus. There have been no

revisions of the genus in the past 20 years. As the type of

Astacilla falclandica has been lost. Lew Ton and Poore (1986a)

redescribed N. falclandica (Ohlin, 1901), proposing that it

should be kept as the type species for the genus despite

Tattersall’s mistake (Lew Ton and Poore, 1986b; ICZN, 1987).

Whether or not Neastacilla is a monophyletic group is yet to

be determined, but the genus remains valid. In this contribution

a new diagnosis of Neastacilla is given, eight new Australian

species are described, and five Australian and New Zealand

species are rediagnosed. Poore et al. (2002) listed Arcturus

brevicornis Haswell, 1881 as a possible Australian species of

Neastacilla but in the absence of a good description, and of

type material, the name cannot be applied to any known

species. A key is provided for all species from these two

countries.

Isopods came from collections in Australia of Museum Victoria,

Melbourne (NMV), the South Australian Museum, Adelaide (SAM),

the Australian Museum, Sydney (AM) and the Tasmanian Museum and

Art Gallery, Hobart (TM). Material of some species was unavailable

for study and short diagnoses derived from published literature are pre-

sented instead of full descriptions (see Remarks sections). Illustrations

were made using microscopes with a camera lucida attachment.

Australian Aboriginal names used as species epithets are to be treated

as nouns in apposition. States and territories of Australia are abbrevi-

ated as follows; NSW (New South Wales), Vic. (Victoria), WA

(Western Australia), SA (South Australia), Tas. (Tasmania), and NT

(Northern Territory). Dimensions are total body length. In figures

limbs are abbreviated: Al, antenna 1; A2, antenna 2; MXl, maxilla 1;

MX2, maxilla 2; MD, mandible; MXP, maxilliped; P1-P5, pereopods

1-5; PL1-PL2, pleopods 1, 2; U, uropod or its rami; Pe, penial plate.

Arcturidae Dana, 1849

Neastacilla Tattersall, 1921

Neastacilla Tattersall, 1921a: 243-244. — Nordenstam, 1933:

118-119.— Kussakin, 1972: 178-189-.— Kensley, 1978: 31.— Wagele,

1991: 91.

Type species. Astacilla falclandica Ohlin, 1901, by plenary

powers (ICZN, 1987).

Diagnosis. Body cylindrical, slender and strongly geniculate

between pereonites 4 and 5. Pereonite 1 fused to head, fusion

indicated by groove, occasionally by lateral incision. Pereonite

4 elongate, 3-10 times length of all other pereonites. Pleon

about same length as combined lengths of pereonites 5-7.

Antenna 2 slender, flagellum of 2 or 3 articles with claw.

Pereopod 1 included within margin of head, dactylus as long as

wide; carpus and propodus of subequal length; dactylus with-

out unguis, with distal setae. Pereopods 2-4 slender; with long,

closely spaced setae; dactylus reduced and claw-like [lost

entirely in a few Australian species]; flexion between carpus

and propodus absent. Pereopods 5-7 progressively shorter pos-

teriorly, dactylus with unguis and secondary unguis. Oostegites

present on pereopods 1^; oostegite 4 forming the major part of

Isopods of the genus Neastacilla from Australia

373

marsupium, thickened. Male pleopod 1 with lateral notch and

setae on posterior face. Male pleopod 2 appendix masculina

curved, with a ridge on posterior face, not extending much past

endopod. Penial plate simple, tapering to apex, straight.

Composition. 38 species, excluding those inquirenda (see

Table 1).

Remarks. Characters such as the morphology of pleopods 1 and

2 (including the short, curved appendix masculina), simple

fused penial plate and the morphology of oostegite 4 in females

are new characters used here to redefine Neastacilla.

Importance has also been placed on the absence of flexion

between the carpus and propodus in pereopods 2^ (apparent in

Astacilla). Many existing description do not include these

characters and so some species (especially those from the

North-west Pacific) remain uncertainly placed within

Neastacilla.

According to Kussakin’s criteria N. tzvetkowae belongs to

Astacilla or possibly Arcturus because it possesses an antenna

2 with 3 or more flagellar articles and probably has an unguis

Table 1. Distributional information for Neastacilla species

Species of Neastacilla

Distribution

Depth range

N. algensis Hale, 1924

Australia, SA

11-12 m

N. antipodea Poore, 1981

Subantarctic New Zealand

intertidal to 15

N. attenuata (Hale, 1946)

Australia, NSW

60-83 m

N. bacillus (Barnard, 1920)

South Africa

5-400 m

N. calif arnica (Boone, 1918)

California, USA

18-100 m

N. coonabooloo sp. nov.

south-eastern Australia (NSW, Vic., Tas.)

subtidal to 9 m

N. deducta Hale, 1924 (synonym: Astacilla vicaria Hale, 1946)

south-eastern Australia (NSW, Vic., Tas., SA)

9 m

N. diomedeae (Benedict, 1898)

Straits of Magellan

subtidal to 34 m

N. estadoensis (Schultz, 1981)

Argentina

intertidal

N. exilis Kussakin, 1971

North-west Pacific

400 m

N. falclandica (Ohlin, 1901)

Falkland Islands

subtidal

N.fusiformis (Hale, 1946)

New Zealand, Hauraki Gulf

? (tow net)

N. inaequispinosa (Guiler, 1949) (synonyms; Astacilla derwenti

south-eastern Australia, Vic., Tas.

18 m

Guiler, 1949; A. oculata Guiler, 1949; A. unicornis Guiler, 1949)

N. kanowna sp. nov.

South-eastern Australia, Vic., Tas., SA

subtidal to 29 m

N. kurilensis Kussakin, 1974

Kurile Islands

intertidal to 60 m

N. lawadi sp. nov.

Australia, Vic., Tas., SA, WA, NT

subtidal to 82 m

N. leucophthalma Kussakin, 1971

North-west Pacific

400 m

N. levis (Thomson and Anderton, 1921)

New Zealand, Cook Strait

31 m

N. littoralis Kussakin, 1974

Kurile Islands

0-45 m

N. macilenta (Hale, 1946)

Australia, NSW

2.5 m

N. magellanica (Ohlin, 1901)

Straits of Magellan

12-208 m

N. marionensis (Beddard, 1886) (synonym: Astacilla kerguelenensis

Marion Islands, Kerguelen Islands

45 to 340 m

Vanhbffen, 1914)

N. marrimarri sp. nov.

Australia, WA

subtidal to 25 m

N. monoseta (Guiler, 1949)

South-eastern Australia, Vic., Tas.

subtidal to 84 m

N. nodulosa Kussakin, 1982

North-west Pacific

460 m

N. ochroleuca Kussakin and Vasina, 1990

Kurile Islands

880 m

N. pallidocula Kussakin and Vasina, 1990

Kurile Islands

910-920 m

N. polita (Gurjanova, 1936)

Sea of Japan

25-60 m

N. richardsonae Kussakin, 1982 (replacement name fox Astacilla

North-west Pacific

128 m

dilatata Richardson, 1909)

N. sheardi (Hale, 1946)

South-eastern Australia, NSW, Vic., SA

subtidal

N. soelae sp. nov.

Australia, WA

subtidal to 52 m

N. tarni sp. nov.

Australia, SA

subtidal to 15 m

N. tattersalli Lew Ton and Poore, 1986a

New Zealand

20-129 m

N. tharnardi sp. nov.

south-eastern Australia, Vic., SA

subtidal to 20 m

N. tristanica Sivertsen and Holthuis, 1980

South Atlantic, Nightingale Island

intertidal to 10 m

N. tuberculata (Thomson, 1879)

New Zealand

8-10 m

N. y Uriel sp. nov.

New Zealand

subtidal to 201 m

N. vitjazi Kussakin, 1971

North-west Pacific

820-1050 m

Species inquirenda

Neastacilla sirenkoi Kussakin and Vasina, 1990

Kurile Islands

880 m

Neastacilla tritaeniata Kussakin, 1982

Bering Sea

5-77 m

Neastacilla tzvetkowae Kussakin, 1974

Kurile Islands

0-43 m

374

Rachael A. King

on pereopod 1. N. tritaeniata was not thoroughly described but

the flagellum of antenna 2 is composed of five articles and

pereonite 4 is not elongate, suggesting a species of Arcturus.

Similarly, N. sirenkoi has four antenna 2 flagellar articles and

an unguis on the dactylus of pereopod 1 and should be placed

in Arcturus. Further, these three species are all from the North-

west Pacific where other species of Arcturus occur.

It is possible that Astacilla cymodocea Menzies and Glynn,

1968 from the Caribbean may yet be included in Neastacilla as

it is described as lacking an unguis on pereopod 1 and lacking

dactyli on pereopods 2-4. With no information available on the

male sexual appendages (no specimens could be examined), it

is not included here.

The morphology of arcturid oostegites is considered here to

be taxonomically important, yet these characters have not often

been included in species descriptions or illustrations. The mor-

phology of oostegite 4, for example, was discovered to vary

within Neastacilla. In some species of Neastacilla (N. coona-

booloo, N. deducta, N. monoseta and N. soelae) there is a

midlength suture in oostegite 4 that is not found in any other

genus (Figs 3A, 4E, 18C, 20B). This suture is thought to be

homologous to that separating the posterior lobe found on

oostegite 4 in other species of Neastacilla (Fig. 7C, 9E, 14E)

and in other genera. This condition is found in females where

the pereonite 4 is extremely narrow and elongate. The smaller

sutured posterior lobe may help aerate the marsupium while the

female is brooding and it is possible that the suture found in

elongate females performs a similar function.

Sexual characters are also argued here to be taxonomically

important. Male pleopodal structure and female oostegites of

the Australian and New Zealand species of Neastacilla differ

from those in N. richardsonae, a north-western Pacific species.

While it is the only species examined from this region, others

from the north-western Pacific are figured similarly.

Neastacilla richardsonae possesses a straight appendix mas-

culina and a developed functional oostegite on pereopod 5.

Both these character states are shared with Arcturus and not

with Australian and New Zealand Neastacilla species. No

information is available on the structure of the appendix mas-

culina in other North-western Pacific species but the curved,

ridged appendix masculina has so far only been found in

Australian and New Zealand species of Neastacilla (and in

Parastacilla, endemic to Australia (King, 2000)). The North-

western Pacific species remain in Neastacilla for the time being

but they may belong to Arcturus, a new genus, or both.

Biogeographical evidence supports the view that the north-

western Pacific species may belong to another genus. The

majority of the species from around Australia and New Zealand

are found in the Pacific Ocean, with four of the new Australian

species described here (Neastacilla lawadi, N. marrimarri,

N. soelae and N. yuriel) reported from the Indian Ocean (coast

of Western Australia). The centre of diversity of Neastacilla is

evidently Australia and New Zealand with undisputed species

also reported from subantarctic islands. Clearly, the north-west

Pacific species are geographically separated from this radiation

and thus are only conditionally included within the genus until

further work can make clear their position. Species from south-

ern South America, South Africa and California, USA were

unavailable for study and are similarly included in the genus

until further clarification.

Key to Australian and New Zealand species of Neastacilla

* Species known only from this sex.

1. Head with distinct dorsal tuberculation(s) or elevations(s)

— Head dorsally smooth or with small rounded elevation 14

2. Fusion of head and pereonite 1 indicated by a suture . . 3

— Fusion of head and pereonite 1 without a suture 4

3. Anterolateral margin of pereonite 1 extended laterally,

head and pereonites 1-4 with large prominent branch-like

elevations, pereopods 2-A flattened Neastacilla tharnardi

— Anterolateral margin of pereonite 1 not extended laterally,

head and pereonites 1-4 with very small anteriorly

produced elevations, pereopods 2-4 cylindrical (male

extremely elongate and elevations reduced to blunt

tubercles) Neastacilla attenuata

4. Pereonite 4 with large, prominent dorsal elevation(s) . . 5

— Pereonite 4 smooth or with small dorsal elevation(s) . . 7

5. Pleotelson equal to or longer than pereonites 5-7; pereo-

nite 4 with proximal dorsal elevations (female pereonite 4

widened at midlength) 6

— Pleotelson shorter than pereonites 5-7; pereonite 4 with

dorsal elevations at midlength (female pereonite 4 not

laterally widened at midlength)

Neastacilla marrimarri (female)*

6. Pereonite 4 with large proximal dorsal elevation with 3

apices and 2 large lateral elevations; pleotelson with 2

pairs of lateral wings and widening distally to taper to a

pointed apex (male pereonite 4 with single dorsal elevation

proximally) Neastacilla tuberculata

— Pereonite 4 with large dorsal midlength elevation with 2

apices with 2 large lateral elevations; pleotelson with 1 pair

of lateral wings, not widened and sharply tapered to a

bluntly rounded apex Neastacilla tarni (female)

7. Pereonite 4 constricted (in dorsal view) for first third

length 8

— Pereonite 4 not constricted (in dorsal view) for first third

length 10

8. Pereonite 4 extremely elongate (10 times as long as

pereonite 3 length) . . Neastacilla inaequispinosa (male)

— Pereonite 4 not extremely elongate (less than 10 times

pereonite 3 length) 9

9. Pereonite 4 with dorsal and lateral elevations at midlength;

pereonites 5-7 with a pair of dorsal elevations plus lateral

elevations; pleotelson wider than pereonite 7, with 2 pairs

of acute lateral wings . . . .Neastacilla fusiformis (male)*

— Pereonite 4 with a pair of small anterodorsal round tuber-

cles; pereonites 5-7 smooth; pleotelson not wider than

pereonite 7, with pair of blunt lateral wings

Neastacilla tarni (male)

10. Head and pereonite 4 each with a pair of distinct spine-like

elevations (male with elevations reduced to blunt

tubercles) Neastacilla lawadi

— Head and pereonite 4 smooth or with weak/ blunt

elevations 11

Isopods of the genus Neastacilla from Australia

375

11. Pereonite 4 without elevations; pleotelson longer than

pereonites 5-7 12

— Peronite 4 with dorsal elevations; pleotelson not longer

than pereonites 5-7 13

12. Pleotelson apex acute; antenna 2 peduncular article 5 as

long as article 4 Neastacilla soelae (female)*

— Pleotelson apex truncate; antenna 2 peduncular article 5

slightly shorter than article 4

Neastacilla algensis (female)*

13. Pereonite 4 with dorsal rounded tubercles extending later-

ally at midlength; pleotelson as long as pereonites 5-7,

with 2 distinct blunt lateral wings

Neastacilla sheardi (female)*

— Pereonite 4 with a weak dorsal elevation at midlength;

pleotelson longer than pereonites 5-7, without defined

lateral wings Neastacilla inaequispinosa (female)

14. Fusion of head and pereonite 1 defined with a suture ....

— Fusion of head and pereonite 1 not defined with a suture . .

15. Eye large and triangular shaped; pleotelson with an apical

blunt notch Neastacilla levis

— Eye small, round or oval in shape; pleotelson without

apical blunt notch 16

16. Pleotelson with large, acute lateral wings with an acute

apex Neastacilla monoseta

— Pleotelson without defined lateral wings, apex blunt . 17

17. Pereonite 4 not more than 6 times as long as pereonite 3,

almost square in dorsal view (male without strong tuber-

culation and pereonite 4 not square in dorsal view)

Neastacilla antipodea

— Pereonite 4 extremely elongate (greater than 8 times as

long as pereonite 3), not square in dorsal view 18

18. Pereonite 4 with a pair of distinct dorsal elevations proxi-

mally (female with pereonite 4 laterally expanded and

upturned at midlength, male pereonite 4 narrow and con-

stricted (in dorsal view) for first third length

Neastacilla coonabooloo

— Pereonite 4 dorsally smooth (female with slight lateral

expansion of pereonite 4 at midlength)

Neastacilla deducta

19. Adult size 3-4 mm; pereonite 4 with a dorsal elevation at

midlength (males with pereonite 4 constricted (in dorsal

view) proximally but with dorsal elevation at first third

length) Neastacilla yuriel

— Adult size >7 mm; pereonite 4 without dorsal elevations .

20. Head with horizontal unsutured groove below eye; pleo-

telson with defined lateral wings (female pereonite 4 with

anterolateral extensions) Neastacilla macilenta

— Head without horizontal unsutured grove; pleotelson with-

out defined lateral wings 21

21. Pereonite 4 elongate (greater than 8 times as long as

pereonite 3); pleotelson with a truncated apex

Neastacilla kanowna

— Pereonite 4 around 6 times as long as pereonite 3;

pleotelson bluntly rounded

Neastacilla tatters alii (female)*

Neastacilla algensis Hale

Neastacilla algensis Hale, 1924; 213, fig. 3. — Hale, 1929: 313, fig.

314.— Poore et al., 2002: 258.

Astacilla algensis. — Hale, 1946: 174, fig. 7A. — Monod, 1970:

1139.

Diagnosis of female. Eyes small and triangular. Head with

rounded dorsal elevation slightly posterior to eyes; lateral mar-

gin of head and pereonite 1 not incised. Pereonites 1-7 dor-

sally smooth, without distinct lateral extensions. Pereonite 4

extremely elongate (more than 10 times as long as pereonite 3).

Pleon longer than pereonites 5-7 combined, lateral wings

absent, apex truncate. 12.3 mm.

Male. Undescribed.

Distribution. Australia: South Australia; subtidal.

Remarks. Neastacilla algensis is diagnosed here with reference

to the illustrations of Hale (1924, 1929), who described a sin-

gle female specimen. The elongate, truncated pleotelson and

extremely elongate pereonite 4 separates this species from all

others in Neastacilla. It should be noted that the specimen

drawn by Hale (1924, 1929) was probably an immature female

as he described the oostegites as not fully developed.

Neastacilla antipodea Poore

Neastacilla antipodea Poore, 1981: 333, figs 2-3.

Material examined. New Zealand: NMV J679 (1 male, 1 female).

Description of female. Head without dorsal elevation, antero-

lateral lobes rounded, rostral point absent; lateral margin of

head and pereonite 1 incised. Pereonite 1 without lateral exten-

sion, with dorsal elevation. Pereonites 2 and 3 progressively

wider; with distinct keel-like dorsal elevations, bilobed lateral

extensions visible on dorsal view. Pereonite 4 about 5 times as

long as pereonite 3, with single keel-like anterior dorsal eleva-

tion, anterolateral and posterolateral margins extended, forming

a square shape in dorsal view. Peronites 5-7 progressively

shorter posteriorly; without dorsal elevations. Pleon length

subequal to the combined lengths of pereonites 5-7, without

dorsal elevations, without lateral wings, apex bluntly rounded.

Eyes small and oval. Antenna 1 reaching to second pedun-

cular article of antenna 2; aesthetascs present distally on flagel-

lum. Antenna 2 robust, more than half as long as body; flagel-

lum of 3 articles, ending with claw, row of scales along full

length.

Maxilla 1 mesial lobe with 3 plumose terminal setae; lateral

lobe with 11 distal robust setae. Maxilla 2 mesial lobe with

about 20 plumose setae; middle lobe with 6 setae; lateral lobe

with 3 setae. Maxillipedal endite with numerous mesial setae, 2

coupling hooks; palp article 2 with mesial setal rows; articles 3

and 4 with mesial and lateral setal rows; article 5 with mesial

and distal setae.

Pereopod 1 propodus slightly shorter than carpus; dactlyus

as long as wide, without unguis. Pereopods 2-4 with dactylus.

Pereopods 5-7 dactylus denticulate, with unguis and secondary

unguis; secondary unguis three-quarters length of primary

unguis. Uropodal exopod with 2 setae of subequal length.

Oostegite 4 with sutured small posterior lobe.

376

Rachael A. King

Adult: 7.0-8.0 mm, juveniles 3.0-7.0 mm, mancas 2.0-3.0

Male. Pereonite 1 without lateral extensions, without dorsal

elevation. Pereonites 2 and 3 progressively wider; without dis-

tinct dorsal elevations, without lateral extensions. Pereonite 4

about 5 times as long as pereonite 3, without dorsal elevation,

anterolateral and posterolateral margins not extended. Peronites

5-7 progressively shorter posteriorly, without dorsal ele-

vations. Pleon length subequal to the combined lengths of

pereonites 5-7, without dorsal elevations, without lateral

wings, apex bluntly rounded.

Antennae, mouthparts, pereopods and uropods as for

female.

Pleopod 1 exopod with lateral notch and 2 plumose lateral

setae. Pleopod 2 appendix masculina with ridge on posterior

face, curved, extending twice as long as the endopod, apex

simple. Penial plate widened proximally, distally bulbous, apex

acute.

Adult: 7. 3-8.5 mm, juveniles 3. 0-7.0 mm, mancas 2.0-3.0

mm.

Distribution. New Zealand: The Snares; intertidal to 15 m

depth.

Remarks. Neastacilla antipodea is one of five species of

Neastacilla (N. fusiformis, N. tuberculata, N. levis, N. tatter-

salli) known from New Zealand. The ornamentation of

N. antipodea distinguishes it from all the other species immed-

iately: specifically, the bluntly rounded pleon and the square

shape of pereonite 4 of the female in dorsal view.

Neastacilla attenuata (Hale)

Astacilla attenuata Hale, 1946: 177-179, figs 9-10. — Monod,

1970: 1137-1139, figs 45-55.— Poore et al., 2002: 258.

Diagnosis of female. Eyes small and oval. Head with dorsal

pair of small anteriorly directed spine-like elevations slightly

posterior to eyes; lateral margin of head and pereonite 1

incised. Pereonites 2 and 3 with single dorsal, anteriorly direct-

ed spine-like elevations. Pereonite 4 extremely elongate (more

than 10 times as long as pereonite 3); anterolateral margins

expanded, with pair of small blunt anterodorsal elevations.

Pleon longer than pereonites 5-7 combined, 2 pairs of small lat-

eral wings present, apex acute. 7.5 mm.

Male. Similar to female except dorsal spines are reduced to

blunt elevations on head and pereonites. 9.8 mm.

Distribution. Australia: New South Wales; 60-80 m.

Remarks. This species is diagnosed from the description of

Hale (1946). This species is distinguished from other species by

the presence of anteriorly directed spine-like dorsal elevations

in the female, the elongate pereonite 4 (extremely elongate in

males), and long pleon.

Neastacilla coonabooloo sp. nov.

Figures 1-3

Material examined. Holotype. Australia: Vic., Crib Point, Western

Port, 38°21.15'S, 145°13.36'E, 9 m, 24 Mar 1965, NMV J1036 (1

female, 6.6 mm).

Paratypes. Australia: NSW. Moes Rock, S of Jervis Bay, AM

P32681 (1 female), AM P32680 (1 female). Vic. Crib Point, Western

Port, 38°21.63'S, 145°15.08'E, 9 m, 23 Eeb 1965, NMV J978 (1

female), NMV J979 (1 female). Western Port, 38°22.0'S, 145°32.0'E,

NMV J980 (1 immature female, 1 immature male). Mallacoota,

37°34.03’S, 149°46.02'E, 5 m, 6 Apr 1989, NMV J47324 (1 immature

female). Tas. Breaksea Island, Bathurst Harbour, 4 m, 16 Eeb 1989,

NMV J47323 (1 male, 5.5 mm).

Description of female. Head with small dorsal elevation poste-

riorly, anterolateral lobes angular, small rostral point present;

lateral margin of head and pereonite 1 incised. Pereonite 1

extended anterolaterally. Pereonites 2 and 3 without dorsal ele-

vations, progressively wider, with small lateral extensions.

Pereonite 4 about 8 times as long as pereonite 3, wider than pre-

vious pereonites, widest at midlength; with 2 anterior horn-like

dorsal elevations, lateral margins extended and curved dorsally

at midlength. Pereonites 5-7 progressively shorter posteriorly;

without dorsal elevations. Pleon length greater than combined

lengths of pereonites 5-7, without dorsal elevations, with small

anterior lateral wings, apex rounded.

Eyes small and oval. Antenna 1 reaching to third pedunc-

ular article of antenna 2; aesthetascs present distally and later-

ally on flagellum. Antenna 2 slender, more than half as long as

body; flagellum of 2 articles ending with claw, row of scales

along full length.

Maxilla 1 mesial lobe with 3 plumose terminal setae; lateral

lobe with 9 distal robust setae. Maxilla 2 mesial lobe with 22

plumose setae; middle lobe with 3 setae; lateral lobe with 3

setae. Maxillipedal endite with 13 mesial setae, 1 coupling

hook; palp article 2 with mesial setal rows; articles 3 and 4 with

mesial and lateral setal rows; article 5 with mesial and distal

setae.

Pereopod 1 propodus as long as carpus; dactylus twice as

long as wide, without unguis. Pereopods 2-3 dactylus present.

Pereopod 4 dactylus absent. Pereopods 5-7 dactylus somewhat

denticulate, with unguis and secondary unguis; secondary

unguis half-length of primary unguis. Uropodal exopod with 2

setae of subequal length. Oostegite 4 with suture at midlength.

5.0-6.6 mm.

Male. Head with a small dorsal elevation posteriorly, antero-

lateral lobes angular, small rostral point present; lateral margin

of head and pereonite 1 incised. Pereonite 1 extended antero-

laterally. Pereonites 2 and 3 of similar width; without dorsal

elevations, small lateral extensions present. Pereonite 4 about

nine times as long as pereonite 3, constricted for first third of

length; with 2 anterior horn-like dorsal elevations, antero-

lateral margins not extended. Pereonites 5-7 progressively

shorter posteriorly; without dorsal elevations. Pleon length

greater than combined lengths of pereonites 5-7 ; without dor-

sal elevations, with small anterior lateral wings, apex rounded.

Antenna 1 reaching half way along third peduncular article

of antenna 2; aesthetascs present distally and laterally on

flagellum. Antenna 2 slender, more than half as long as body;

flagellum of 3 articles ending with a claw, a row of scales full

length.

Mouthparts, pereopods and uropods as for female.

Pleopod 1 exopod with a lateral notch and 2 plumose lat-

eral setae of unequal length. Pleopod 2 appendix masculina

Isopods of the genus Neastacilla from Australia

379

Figure 3. Neastacilla coonabooloo sp. nov., female holotype (NMV J1036): PI, P3, P4, P5, U; A, ventral view. Male (NMV J47323): Pe, PLl,

PL2. Scales: a (PI, P3, P4) = 0.5 mm; b = (A) 1.0 mm; c (P5) = 0.5 mm; d (U, Pe, PLl, PL2) = 0.2 mm

380

Rachael A. King

with ridge on posterior face, curved, extending third length past

the endopod, apex simple. Penial plate widened proximally,

distally tapered, apex simple.

4. 5-5. 6 mm

Distribution. Australia: New South Wales, Victoria, Tasmania;

subtidal to 9 m depth.

Etymology. “Coonabooloo” is an Aboriginal word meaning two

shoulders and refers to the extended lateral margins of pere-

onite 4 in the female of this species.

Remarks. This species possesses an incised dorsolateral groove

at the fusion of the head and pereonite 1 as does Neastacilla

deduct a and N. monos eta. Females of N. coonabooloo are dis-

tinguished from these two species by the presence of extended

lateral margins on pereonite 4 and two anterior horn-like ele-

vations on pereonite 4. Males of N. coonabooloo possess

smaller horn-like elevations. N. coonabooloo can be further dif-

ferentiated from N. monoseta by possessing a pleon with small

rounded lateral wings, as opposed to the large acute wings in

N. monoseta. In the material examined here, the heights of the

dorsal elevations on pereonite 4 vary from small bumps to

spine like elevations.

Neastacilla deducta Hale

Figures 4-6

Neastacilla deducta Hale, 1925: 33, fig. 16.— Hale, 1929: 313, fig.

315.— Monod, 1970: 1139.— Poore et al, 2002: 259.

Astacilla deducta.— Hale, 1946; 174-175, fig. 7.

Astacilla vicariaHale, 1946: 175-176, fig. 8. — Monod, 1970: 1139.

Material examined. Holotype. Australia: SA, Port Adelaide (Gulf St

Vincent) (1 male) SAM C 273. Paratypes. Austtalia: SA, Port Adelaide

(Gulf St Vincent) SAM C 274.

Syntypes of Astacilla vicaria Hale, 1946. NSW, off Yarra Bay,

Botany Bay, AM P8967 (8 specimens).

Other material. Australia: NSW. Jervis Bay, 35°08.0'S, 150°43.0'E,

2 m, 23 Apr 1985, NMV J11199 (2 females, 4 males, 1 immature

female). Port Kembla, 34°29.0’S, 150°55.0'E, Mar 1978, NMV J16516

(1 female). Vic. Oberon Bay, 39°04.2'S, 146°19.4'E, 21 m, 05 Eeb

1982, NMV 111 200 (11 females, 7 males, 5 immature males, 2 manca

2). Tas. 43°17.0’S, 147°15.'E, 3 m, 20 Mar 1988, NMV J48664 (1

female); 39°32.8'S, 144°16.0'E, 18 m, 1 Nov 1980, NMV J8840. SA.

Venus Bay, 33°13.2'S, 134°40.1'E, 2 m, 23 Apr 1985, NMV J40673 (1

female, 11.5 mm), NMV J 40674 (1 male, 11.3 mm), NMV J16569 (12

individuals); Venus Bay, 33°13.8'S, 134°40.1'E, 3 m, 21 Apr 1985,

NMV J 16570 (2 females, 1 male, 2 immature females, 1 immature

male, 3 juveniles, 2 manca-2). NE side of Topgallant I., 33°43.0'S,

134“36.6'E, 20 m, 21 Apr 1985, NMV J16568 (3 females).

Description of female. Head without dorsal elevation, antero-

lateral lobes angular, small rostral point present; lateral margin

of head and pereonite 1 incised. Pereonite 1 with an anterolat-

eral expansion. Pereonites 2 and 3 similar width, without dor-

sal elevations, without lateral extensions. Pereonite 4 about 10

times as long as pereonite 3, with some small anterior dorsal

elevations, anterior anterolateral margins extended, not greatly

wider then previous pereonites. Pereonites 5-7 progressively

shorter posteriorly, without dorsal elevations. Pleon length

greater than combined lengths of pereonites 5-7, without

dorsal elevations, with small anterior lateral wings, apex blunt.

Eyes small and subtriangular. Antenna 1 reaching to the end

of second peduncular article of antenna 2; aesthetascs present

laterally and distally on flagellum. Antenna 2 slender, more

than half as long as body; flagellum of 2 articles, ending with

claw, with row of scales along full length.

Maxilla 1 mesial lobe with 3 terminal setae; lateral lobe with

9 distal robust setae. Maxilla 2 mesial lobe with 26 plumose

setae, middle lobe with 3 setae, lateral lobe with 3 setae.

Maxillipedal endite with 11 mesial setae; 1 coupling hook; palp

article 2 with mesial setal rows, article 3 with mesial and later-

al setal rows, article 4 with mesial setae, article 5 with distal

setae.

Pereopod 1 propodus as long as carpus; dactylus slightly

longer than wide, without unguis. Pereopods 2-3 dactylus pres-

ent. Pereopod 4 dactylus absent. Pereopods 5-7 dactylus not

denticulate, with unguis and secondary unguis; secondary

unguis two-thirds length of primary unguis. Uropodal exopod

with 2 setae of subequal length. Oostegite 4 with suture at

midlength.

8.0-10.5 mm.

Male. Head and pereonites 1-3 as for female. Pereonite 4 about

10 times as long as pereonite 3; without dorsal elevations, con-

stricted for the first quarter length. Pereonites 5-7 progres-

sively shorter posteriorly, without dorsal elevations. Pleon

length greater than combined lengths of pereonites 5-7, with-

out dorsal elevations, with small lateral wings, apex blunt.

Antenna 1 extending past the end of the second peduncular

article of antenna 2; aesthetascs present laterally and distally on

flagellum. Antenna 2 slender, more than half as long as body;

flagellum of 3 articles and claw; with a row of scales along full

length.

Mouthparts, pereopods and uropods as for female.

Pleopod 1 exopod with lateral notch and 2 plumose lateral

setae of unequal length. Pleopod 2 appendix masculina with

ridge on posterior face, curved, extending quarter length past

the endopod, apex simple. Penial plate widened proximally,

tapering distally, apex notched.

7-10 mm.

Distribution. Australia: New South Wales, Victoria, Tasmania,

South Australia; subtidal to 21 m depth.

Remarks. Neastacilla deducta was described from South

Australia from a single male by Hale (1925). He later described

a female and ‘subadult’ male of Astacilla vicaria from eight

syntypes from New South Wales (Hale, 1946). The distinction

between the two species has never been clear. Hale (1946)

argued that the structure of the second antenna, including the

number of flagellar articles; eye size and expansion of pere-

onite 1 were differences. Examination of type material of N.

vicaria and A. deducta and of other material at Museum

Victoria has shown that the two are synonymous, a view shared

by Poore et al. (2002). The type males of A. vicaria males were

all immature, some up to 1 or 2 moults from maturity as shown

by the underdeveloped appendix masculina. Juveniles and

females always possess two antenna 2 flagellar articles; only

mature males have a flagellum of 3 articles. This condition is

also seen in N. coonabooloo sp. nov.

Isopods of the genus Neastacilla from Australia

383

Figure 6. Neastacilla deducta Hale, 1924, male (NMV J40674): PLl, PL2, Pe. Female (NMV J40673); P5, U. Scales a (P5) = 0.5 mm; b (PLl,

PL2, Pe, U) = 0.5 mm.

Neastacilla deducta looks most similar to N. monoseta and

N. kanowna sp. nov. It is distinguished from N. monoseta by

the blunt pleotelson without lateral expansions and from

N. kanowna by the presence of a shallow dorsolateral groove

between the head and pereonite 1, which is incised laterally.

Neastacilla fusiformis (Hale)

Astacilla fusiformis Hale, 1946:185-186, fig. 14. — Hurley, 1961:

264.— Monod, 1970: 1139.

Diagnosis of male. Eyes small and subtriangular. Head with

small dorsal elevation anterior to eyes and pair of larger dorsal

elevations slightly posterior to eyes, lateral margin of head and

pereonite 1 not incised. Pereonites 1-3 with single dorsal ele-

vation and pair of lateral elevations. Pereonite 4 7 times as long

as pereonite 3, anteriorly restricted in dorsal view, with pair of

dorsal elevations and 2 pairs of dorsolateral elevations at

midlength, with single dorsal elevation and pair of dorsolateral

elevations posteriorly. Pereonites 5-7 with pair of dorsal evel-

ations and pair of lateral elevations. Pleon slightly longer than

pereonites 5-7 combined, with 3 pairs of dorsal elevations and

lateral elevations, 2 pairs of lateral wings present, apex acute. 5

mm.

Female. Undescribed.

Distribution. New Zealand, North Island, Huaraki Gulf;

subtidal.

Remarks. Neastacilla fusiformis was described by Hale (1946)

from a single male specimen taken in a tow net from the

Hauraki Gulf, New Zealand. This specimen is similar to the

384

Rachael A. King

male of N. tuberculata and N. sheardi, however neither of these

species possesses a pereonite 4 that is anteriorly constricted in

dorsal view or sharp dorsal and dorsolateral elevations.

Neastacilla inaequispinosa (Guiler)

Figures 7-8

Astacilla inaequispinosa. Guiler, 1949: 49-53, figs 3, 4. — Guiler,

1952: 24.— Monod, 1970: 1139.

Astacilla unicornis Guiler, 1949: 53-55, fig. 5. — Guiler, 1952:

24.— Monod, 1970: 1139-1140.

Astacilla clerwenti Guiler, 1949: 56-57, fig. 6. — Guiler, 1952: 24. —

Monod, 1970: 1139-1140.

Astacilla oculata Guiler, 1949: 59-61, fig. 7. — Guiler, 1952: 24. —

Monod, 1970: 1139-1140.

Neastacilla inaequispinosa. — Poore et al., 2002: 259.

Material examined. Syntypes of Astacilla inaequispinosa Guiler, 1949.

Tas., N end of D'Entrecasteaux Channel, TMG76a (male), TMG76b (1

specimen).

Holotype of Astacilla derwenti Guiler, 1949. Tas., N end of

D'Entrecasteaux Channel, TM (not registered).

Holotype of Astacilla oculata Guiler, 1949. Tas., N end of

D'Entrecasteaux Channel, TM (not registered).

Holotype of Astacilla unicornis Guiler, 1949. Tas., N end of

D'Entrecasteaux Channel, TM (not registered).

Australia: Vic. Port Phillip Bay, 38°17.6’S, 144“42.3’E, 17 m, 4 Mai-

1991, NMV J39217 (1 female, 6.4 mm), NMV J40691(male, 8 mm)

NMV J40695 (1 male, 6 mm). Western Bass Strait, 39°26.3’S,

143“06.8’E, 115 m, 21 Nov 1981, NMV J8843 (1 female, 7.0 mm; 2

males, 7 mm); 39°21.0’S, 143°06.0’E, 101 m, 10 Oct 1980, NMV

J8844 (1 female, 6.5 mm); 39°06.0’S, 143°21.0’E, 59 m, 8 Oct 1980,

NMV J8837 (1 immature male, 5.5 mm); 39°20.0’S, 143°34.0’E, 95 m,

10 Oct 1980, NMV J8842 (1 male, 10 mm). Western Port, 38°26.48’S,

145°13.03’E, 23 m, 25 Nov 1973, NMV J1014 (1 female, 10.5 mm; 1

immature male, 8 mm); 38°21.39’S, 145°14.03’E, 16 m, 25 Mar 1965,

NMV JlOll (1 male, 8.5 mm; 1 immature male, 7 mm). 9.5 km SW of

Port Albert, 38°44.0’S, 146°37.0’E, 9 m, 22 Nov 1983, NMV J12580

(1 female, 10.5 mm). Tas. Central Bass Strait, 40°31.rS, 145°04.0’E,

29 m, 3 Nov 1980, NMV J8841 (2 females, 10-11 mm; 1 immature

male, 8 mm).

Description of female. Head with a dorsal elevation between

the eyes; anterolateral margins angular; rostral point present;

lateral margin of head and pereonite 1 not incised. Pereonite 1

with small dorsal elevation. Pereonites 2 and 3 progressively

wider, with small dorsal elevations, with small anterolateral

extensions present. Pereonite 4 about 6 times as long as pereo-

nite 3; with a dorsal elevation at third length and an elevation

on the posterior dorsal margin, anterior anterolateral margins

rounded and extended, widest anteriorly. Pereonites 5-7 pro-

gressively shorter posteriorly, without dorsal elevations. Pleon

length greater than combined lengths of pereonites 5-7; with-

out dorsal elevations, with small lateral wings, tapered to a

narrow rounded apex.

Eyes large and round. Antenna 1 reaching to the end of the

second peduncular article of antenna 2; aesthetascs present dis-

tally and laterally on flagellum. Antenna 2 slender, more than

half as long as body; flagellum of 3 articles ending with claw,

with 2 rows of scales along full length.

Maxilla 1 mesial lobe with 3 terminal setae; lateral lobe with

12 distal robust setae. Maxilla 2 mesial lobe with 20 plumose

setae, middle lobe with 4 setae, lateral lobe with 3 setae.

Maxillipedal endite with 8 mesial setae; 1 coupling hook; palp

article 2 with mesial setal rows, articles 3 and 4 with mesial and

lateral setal rows, article 5 with mesial and distal setae.

Pereopod 1 propodus as long as carpus; dactylus slightly

longer than wide, without unguis. Pereopods 2-4 with small

dactylus. Pereopods 5-7 dactylus not denticulate, with unguis

and secondary unguis; secondary unguis half length of primary

unguis. Uropodal exopod with 2 setae of unequal length.

Oostegite 4 with sutured small posterior lobe.

6.5-11 mm.

Male. Head and pereonite 1 similar to female. Pereonites 2

and 3 without dorsal elevations. Pereonite 4 about 11 times as

long as pereonite 3, constricted for the first quarter length, with

a posterodorsal curved elevation. Pereonites 5-7 progressively

shorter posteriorly, without dorsal elevations. Pleon length

greater than combined lengths of pereonite 5-7, with a pair of

small dorsal elevations, with lateral wings, tapering to a narrow

rounded apex.

Antennae, mouthparts, pereopods and uropods as for

female.

Pleopod 1 exopod with lateral notch and 2 plumose lateral

setae of equal length. Pleopod 2 appendix masculina with ridge

on posterior face, with tufts of fine setae at three-quarters

length, curved, extending quarter length past endopod, apex

simple. Penial plate proximally widened, straight, apex simple.

6.0-10.5 mm.

Distribution. Australia: Victoria, Tasmania; subtidal to 115 m

depth.

Remarks. This species was briefly described by Guiler (1949)

along with three others from the same locality that are synony-

mous. The similarity between Guiler’s specimens, two of

A. inaequispinosa and one of each of the others, was first cor-

rectly noted by Lew Ton (unpublished) who attributed the

alleged specific differences to uneven mounting and illus-

trations of the material, and incorrect differentiation of males,

females and juveniles. The large dorsal elevation between the

eyes and the lack of an incision in the groove between the head

and pereonite 1 most easily distinguishes N. inaequispinosa

from other species of Neastacilla. Also, the shape of the pleo-

telson, the dorsal elevations on pereonites 1-4 of the female

and the posterior dorsal elevation on pereonite 4 of both males

and females are key characteristics.

Neastacilla kanowna sp. nov.

Figures 9-1 1

Material examined. Holotype. Australia: Vic., Oberon Bay,

39‘’04.2’S, 146°19.4’E, 21 m, 5 Feb 1982, NMV J3418 (1 female, 14

mm).

Paratypes. Australia: Vic. type locality, NMV J3213 (2 females,

15-16 mm; 2 males, 13.5-15 mm), NMV J3214 (1 male, 14 mm),

NMV J40679 (1 male, 10.5 mm), NMV J40680 (6 females, 13-15 mm;

7 males, 11-15.5 mm; 2 immature females, 9.0-9.5 mm; 1 immature

male, 11 mm). Swan Bay, 38‘’14.0’S, 144°39.0’E, 4 m, 26 Feb 1991,

NMV J20908 (1 female, 13 mm). Eastern Bass Strait, 38°15.0’S,

147°22.5’E, 16 m, 31 Jul 1983, NMV J8820 (1 female, 13.5 mm; 1

Isopods of the genus Neastacilla from Australia

387

Figure 9. Neastacilla kanowna sp. nov., female holotype (NMV J3418): A, lateral view; B, dorsal view; E, ventral view with oostegites. Male

(NMV J40679): C, lateral view; D, dorsal view. Scales = a (A-D) =1.0 mm; b (E) = 1.0 mm.

male, 11 mm; 1 immature female, 10 mm; 1 immature male stage 1,

7.5 mm). Tas. Central Bass Strait, off Three Hummock I., 40°31.1’S,

145°04.0’E, 29 m, 3 Nov 1980, NMV J8819 (1 male, 15.5 mm). SA.

Elinders L, 33°40.50’S, 134°22.0’E, 20 m, 19 Apr 1985, NMV J16578

(1 male, 15 mm).

Description of female. Head without dorsal elevation, antero-

lateral lobes rounded, small rostral point present; lateral margin

of the head and pereonite 1 not incised. Pereonite 1 without

anterolateral expansion. Pereonites 2 and 3 similar width, with-

out dorsal elevations, without lateral extensions. Pereonite 4

about 9 times as long as pereonite 3, without dorsal elevations,

small anterolateral extensions present, slightly wider than

pereonites 2 and 3. Pereonites 5-7 progressively shorter

posteriorly, without dorsal elevations. Pleon length greater than

combined lengths of pereonites 5-7, without dorsal elevations,

with small proximal lateral wings, apex truncated.

388

Rachael A. King

Figure 10. Neastacilla kanowna sp. nov., female holotype (NMV J3418): left MXP, MXl, MX2, MD, Al, A2, PI, P3. Scales: a (MXl, MX2, MD)

= 0.5 mm; b (MXP) = 0.5 mm; c (Al) = 0.5 mm; d (A2) = 0.5 mm; e (PI, P3) = 0.5 mm)

Eyes small and subtriangular. Antenna 1 reaching to the

distal edge of the second peduncular article of antenna 2;

aesthetascs present distally and laterally on flagellum. Antenna

2 slender, more than half as long as body; flagellum of 3

articles plus claw, with 2 rows of scales along full length.

Maxilla 1 mesial lobe with 3 terminal setae; lateral lobe with

11 robust setae. Maxilla 2 mesial lobe with 22 setae; middle

lobe with 7 setae; lateral lobe with 3 setae. Maxillipedal endite

with 14 mesial setae; 1 coupling hook; palp article 2 with

mesial setal rows; article 3 with mesial setal and lateral setal

Isopods of the genus Neastacilla from Australia

389

Figure n. Neastacilla kanowna sp. nov., female holotype (NMV J3418): U. Male (NMV J40679); PLl, PL2, Pe. Scales: a (P5) = 0.5 mm; b (PLl,

PL2, Pe) = 0.5 mm; c (U) = 0.25 mm.

rows; articles 4 and 5 with mesial and lateral setal rows; article

5 with mesial and distal setae.

Pereopod 1 propodus shorter than carpus; dactylus almost

twice as long as wide, without unguis. Pereopods 2-4 with

dactylus. Pereopods 5-7 dactylus not denticulate, with unguis

and secondary unguis; secondary unguis greater than half

length of primary unguis. Uropod exopod with 5 setae.

Oostegite 4 with sutured small posterior lobe.

10-16 mm .

Male. Smaller than female but similar morphologically.

Pereonites, mouthparts, pereopods all as for female.

Pleopod 1 exopod with lateral notch and 2 plumose lateral

setae of equal lengths. Pleopod 2 appendix masculina with

ridge on posterior face, with fine setae, curved, and extending

quarter length past endopod, apex simple. Penial plate widened

proximally, distally bulbous with fine setae, apex with shallow

notch.

7.5-15 mm.

Distribution. Australia: Victoria, Tasmania, South Australia;

subtidal.

Etymology. “Kanowna” is an Australian Aboriginal name for

the sea.

Remarks. This species is most easily distinguished from all

other species of Neastacilla by its large adult size (around 15

mm), unornamented body and truncate pleotelson. The pres-

ence of five setae on the uropodal exopod distinguishes this

species from all other Australian species of Neastacilla, which

generally have two or three apical setae.

390

Rachael A. King

Neastacilla lawadi sp. nov.

Figures 12-13

Material examined. Holotype. Australia: WA, between Dampier and

Port Hedland, 19M8.8'S, 117°52.2'E, 52 m, 2 Sep 1983, NMV J16933

(1 female, 8 mm).

Paratypes Australia: NT. Oxley Island, (W side), IFOO.O’S,

132°49.0'E, 14 m, 21 Oct 1982, NMV J16580 (2 females, 5 mm). NT,

station unknown, Oct 1982, NMV J16581 (1 female, 9 mm; 1 male, 6

mm). SA. upper Spencer Gulf, NMV J16614 (1 female, 7.5 mm); E of

Lowly Point, 33mO'S, 137°49.5'E, 18 m, Eeb 1986, NMV J16610 (1

female, 7.5 mm); N of Pairway Bank, 33°02.4'S, 137°45.0'E, 18 m, Eeb

1986, J 166 11 (1 female, 8 mm). WA. Between Dampier and Port

Hedland, 19°37.00'S, 118‘’53.00’E, 30 m, 3 Jun 1983, NMV J16634 (1

male, 5.5 mm); 19°05.82S, 118‘’56.7'E, 82 m, 14 Eeb 1983, NMV

J16669 (3 females, 7.5-8. 5 mm; damaged juvenile 5.5 mm; manca 1

3.5 mm); 19°48.8'S, 117°52.2'E, 52 m, 2 Sep 1983, NMV J16660 (1

female 7.5 mm); 19°29.0'S, 118°53.2’E, 40 m, 12 Eeb 1983, NMV

16931 (1 female 7.5 mm); 19°37.0’S, 118°53.0'E, 30 m, 3 Jun 1983,

NMV 16634 (3 mature females, 5-6.5 mm; 2 immature females, 5 mm;

1 male, 5 mm).

Description of female. Head with 2 large dorsal elevations and

2 small elevations posteriorly; anterolateral margins angular;

rostral point present; lateral margin of head and pereonite 1 not

incised. Head and pereonite 1 extended anterolaterally.

Pereonite 1 with small paired tuberculate dorsal elevations.

Pereonites 2 and 3 with small paired tuberculate dorsal eleva-

tions and lateral tuberculate elevations, small lateral extensions

present. Pereonite 4 about 9 times as long as pereonite 3, with

dorsal and dorsolateral tuberculate elevations anteriorly, large

paired dorsal elevations midlength, anterolateral margins

rounded and extended. Pereonites 5-7 progressively smaller

posteriorly, with some dorsal tuberculation. Pleon length

greater than combined lengths of pereonites 5-7, with 2 sets of

lateral wings, apex subacute.

Eyes small and suboval. Antenna 1 extending to midlength

of third peduncular article of antenna 2; aesthetascs present dis-

tally on flagellum. Antenna 2 slender, more than half as long as

body; flagellum of 2 articles ending with a claw, with row of

scales along full length.

Maxilla 1 with 3 terminal setae; lateral lobe with 10 distal

robust setae. Maxilla 2 mesial lobe with 22 plumose setae, mid-

dle lobe with 3 setae, lateral lobe with 3 setae. Maxillipedal

endite with 12 mesial setae; 1 coupling hook; palp article 2 with

mesial setal rows, articles 3 and 4 with mesial and lateral setal

rows, article 5 with distal setae.

Pereopod 1 propodus slightly shorter than carpus; dactylus

longer than wide, without unguis. Pereopods 2-4 with small

dactylus. Pereopods 5-7 dactylus not denticulate, with unguis

and secondary unguis; secondary unguis setiform and slender

and only slightly smaller than primary unguis. Uropodal exo-

pod with 2 setae of equal length. Oostegite 4 with a sutured

small posterior lobe.

5-9 mm.

Male. Ornamentation not as pronounced as for female. Head

with slight elevation and pereonites 1 to 3 with some tuber-

culation. Pereonite 4 about 10 times as long as pereonite 3;

without dorsal elevations except for posterodorsal curved ele-

vation. Pereonites 5-7 progressively smaller; pereonite 5 with

a dorsal curved elevation; pereonites 6 and 7 without dorsal

elevations. Pleon longer than combined lengths of pereonites

5-7, with 2 sets of lateral wings, apex subacute.

Antennae, mouthparts, pereopods and uropods as for

female.

Pleopod 1 exopod with lateral notch and 2 plumose setae of

unequal length. Pleopod 2 appendix masculina with ridge on

posterior face, curved, extending third length past endopod,

apex simple. Penial plate widened proximally, distally tapered,

apex simple.

5-7 mm.

Distribution. Australia: Western Australia, Northern Territory,

South Australia; subtidal to 82 m depth.

Etymology. “Lawadi” is an Australian Aboriginal word in the

Gooniyandi dialect from north-western Australia where this

species was first collected. It means “shoulder” and refers to the

rounded anterolateral extensions on pereonite 4 of the female.

Remarks. This species resembles Neastacilla attenuata Hale,

1946 from New South Wales and the eastern coasts of

Australia. However, the shape of the head is the most obvious

difference between the two. N. lawadi sp. nov. does not have a

lateral incision between the head and pereonite 1 and the head

is more ventrally expanded and more sculptured. The third

peduncular article of antenna 2 in N. lawadi sp. nov. is not

thickened as in N. attenuata. The pleotelson of N. lawadi sp.

nov. has a posterior dorsal elevation and more pronounced

lateral wings. Neastacilla attenuata has a slender pleotelson

with no elevation and is slightly turned up at the end.

Neastacilla levis (Thomson and Anderton)

Figures 14-15

Astacilla levis Thomson and Anderton, 1921: 114-115, text-fig. —

Hurley, 1957: 13-15, figs 50-66.— Hurley, 1961: 264, 281.

Material examined. New Zealand, off the W coast of South Island,

42°25.0’S, 171°06.0’E, 35 m, 5 Mar 1982, NMV J4736 (1 female, 9.5),

NMV J40692 (1 female, 10 mm), NMV J40675 (1 male, 9.5 mm).

Description of female. Head without dorsal elevations, antero-

lateral lobes rounded, rostral point present; lateral margin of

head and pereonite 1 incised. Pereonite 1 not extended antero-

laterally, with small dorsolateral tuberculate elevations.

Pereonite 2 and 3 progressively wider, without dorsal ele-

vations, small lateral extensions present. Pereonite 4 about 7

times as long as pereonite 3; dorsally elevated at first third

length, anterolateral margins extended and rounded. Pereonites

5-7 progressively smaller, without dorsal elevations. Pleon

length greater than combined lengths of pereonites 5-7, with-

out dorsal elevations, with small lateral wings, apex truncated

and notched.

Eyes large and triangular. Antenna 1 extending to the distal

edge of second peduncular article of antenna 2; aesthetascs

present distally and laterally on flagellum. Antenna 2 slender,

more than half as long as body; flagellum of 3 articles ending

in a claw, 2 rows of scales along full length.

Maxilla 1 mesial lobe with 3 terminal setae; lateral lobe with

11 distal robust setae. Maxilla 2 mesial lobe with 14 plumose

Isopods of the genus Neastacilla from Australia

395

setae, middle lobe with 5 setae, lateral lobe with 4 setae.

Maxillipedal endite with three coupling hooks present; palp

article 2 with mesial setal rows, articles 3 and 4 with mesial and

lateral setal rows, article 5 with mesial and distal setae.

Pereopod 1 propodus and carpus similar length; dactylus

longer than wide, without unguis. Pereopods 2-4 with dactylus.

Pereopods 5-7 dactylus with 3 denticulations (see Remarks)

and setae, with unguis and secondary unguis; secondary unguis

about third length of primary unguis. Uropodal exopod with 3

setae of equal length. Oostegite 4 with sutured small posterior

lobe.

10-12 mm.

Male. Head ornamentation similar to female. Pereonite 4

about 8 times as long as pereonite 3, without dorsal elevations,

anterolateral margins not extended. Pereonites 5-7, Pleon

similar to female.

Antennae, mouthparts, pereopods and uropods as for female.

Pleopod 1 exopod with lateral notch and 2 plumose lateral

setae of unequal lengths. Pleopod 2 appendix masculina with

ridge on posterior face, curved, extending third length past the

endopod, apex simple. Penial plate simple and straight.

10-13 mm.

Distribution. New Zealand; subtidal.

Remarks. First described by Thomson and Anderton (1921) as

Astacilla levis, only a lateral view of the single female speci-

men taken off Otago Heads was drawn. The description was

brief and relatively uninformative. Hurley (1957) provided fur-

ther description the species from two males and one female

taken from Cook Strait, figuring only the male and not describ-

ing the male pleopod 1, female oostegites, maxilla 1, maxilla 2

or mandible. The large triangular eye, shape of the head and

pereonite 1 and the truncate, notched pleotelson immediately

distinguish this species. Hurley (1957) described the denti-

culation of pereopods 5-7 as “4-5 small corrugations;” three

denticulations were found in all specimens examined for this

study so this character must be variable.

Neastacilla macilenta (Hale)

Astacilla macilenta Hale, 1946: 179-182, figs 11-12. — Monod,

1970: 1139.

Neastacilla macilenta . — Poore et al., 2002: 259.

Diagnosis of female. Eyes small and round. Head without dor-

sal elevations; lateral margins of head and pereonite 1 not

incised, extended anterolaterally; unsutured lateral groove

below eye present. Pereonites 2-3 without dorsal elevations.

Pereonite 4 7 times as long as pereonite 3, without dorsal ele-

vations, anterolateral margins extended and angular. Pleon

length longer than pereonites 5-7 combined, 2 pairs of lateral

wings, apex bluntly rounded. 8 mm.

Male. Similar to female except for pereonite 4, which is 15

times as long as pereonite 3.

Pleopod 1 exopod with lateral notch and 2 plumose lateral

setae of unequal lengths. Pleopod 2 appendix masculina with

ridge on posterior face, curved, not extending past the endopod,

apex simple. Penial plate undescribed. 9 mm.

Distribution. Australia: New South Wales.

Remarks. This species is diagnosed using the description of

Hale (1946). This species is superficially similar to Neastacilla.

soelae sp. nov., N. monoseta and N. deducta. However,

N. soelae possesses a distinctive long pointed pleotelson and

N. monoseta possesses a large, angular lateral wings on the

pleotelson. The pleotelson of N. macilenta is rounded and

blunt. N. macilenta can be further distinguished from N. mono-

seta and N. deducta by the absence of an incision in the suture

line between the head and pereonite 1 .

Hale (1946) illustrated three lateral plumose setae on the

exopod of pleopod 1 in males. In all specimens examined here,

there were two lateral plumose setae.

Neastacilla marrimarri sp. nov.

Figures 16-17

Material examined. Holotype. Australia: WA, King George Sound,

35°00.7'S, 118°10.1'E, 25 m, 15 Apr 1984, NMV J16641 (1 female, 6.4

mm).

Description of female. Head with 2 dorsal elevations between

eyes, anterolateral margins angular, rostral point absent; lateral

margin of head and pereonite 1 not incised. Pereonite 1 with

large dorsal elevation and small dorsolateral elevations.

Pereonites 2 and 3 with dorsal and dorsolateral elevations, with

lateral margins extended. Pereonite 4 about 5 times as long as

pereonite 3; with 2 dorsal elevations midlength each with 2

apices, small dorsolateral elevations at midlength, anterior dor-

solateral elevations, 2 posterodorsal elevations and 2 posterior

dorsolateral elevations, anterolateral margins rounded and

extended. Pereonites 5-7 progressively shorter posteriorly, with

dorsal and dorsolateral elevations. Pleon similar length to the

combined lengths of pereonites 5-7, with 2 sets of lateral

wings, apex subacute.

Eyes small and round. Antenna 1 reaching past the second

peduncular article of antenna 2; aesthetascs present distally and

laterally on flagellum. Antenna 2 slender, more than half as

long as body; flagellum of 3 articles ending with claw; with 2

rows of scales along full length.

Maxilla 1 mesial lobe with 3 terminal setae; lateral lobe with

9 distal robust setae. Maxilla 2 mesial lobe with 15 plumose

setae, middle lobe with 3 setae, lateral lobe with 3 setae.

Maxillipedal endite with 8 mesial setae; 1 coupling hook; palp

article 2 with mesial setal rows, articles 3 and 4 with mesial and

lateral setal rows, article 5 with distal setae.

Pereopod 1 propodus and carpus of a similar length; dacty-

lus longer than wide; without unguis. Pereopods 2-4 with small

dactylus. Pereopods 5-7 dactylus denticulate, with unguis and

secondary unguis; secondary unguis half size of primary

unguis. Uropodal exopod with 2 setae of equal length.

Oostegite 4 with a sutured small posterior lobe.

6.4 mm.

Male. Unknown

Distribution. Australia: Western Australia; subtidal.

Etymology. "Marrimarri" is an Australian Aboriginal word in

the Nyungar dialect from south-western Australia where this

specimen was found. It means “crustacean” or “crab”.

396

Rachael A. King

Figure 16. Neastacilla marrimarri sp. nov., female holotype (NMV J16641): A, lateral view; B, dorsal view; C, ventral view with oostegites. Scale

= 1.0 mm.

Remarks. Even though this species is known only from a single

female specimen, it is sufficiently distinct to warrant a new

species. Superficially this species resembles Neastacilla tuber-

culata from New Zealand and N. sheardi from southern

Australia. However, in N. marrimarri sp. nov. the dorsal eleva-

tion of pereonite 4 is not as pronounced and the lateral margins

not expanded to as great an extent as they are in N. tuber-

culata. The ornamentation of pereonite 4 is also arranged dif-

ferently. Neastacilla sheardi has less ornamentation than

N. marrimarri. Further, the pleotelson of N. marrimarri is more

narrow, than in N. tuberculata and more angular than that of

N. sheardi.

Neastacilla monoseta (Guiler)

Figures 18-20

Astacilla monoseta Guiler, 1949: 47-49, figs 1, 2. — Guiler, 1952;

24.— Monod, 1970: 1139-1140.

Neastacilla monoseta. — Poore et al., 2002: 259.

Material examined. Australia: Vic. Western Port, 38°21.48’S,

145°13.85’E, 15 m, 30 Mar 1965, NMV J1022 (1 female, 10 mm);

38°22.0’S, 145°32.0’E, NMV J1027 (1 female, 15.5 mm; 7 manca 1

individuals), NMV J1028 (1 female, 15 mm); 38°20.67’S,

145°14.74’E, 9 m, 4 Mar 1965, NMV J1018 (immature male, 8 mm);

38°21.0’S, 145°14.0’E, 8 m, 12 Oct 1964, NMV J1017 (1 male, 15

mm; 1 immature male 9 mm); 38°21.39’S, 145°14.03’E, 16 m, 25 Mar

1965, NMV J1023 (1 male, 15.5 mm); 38°21.17’S, 145°14.0’E, 18 m,

29 Mar 1965, NMV J1024 (immature female, 11 mm). Portsea,

38°19.0’S, 144°43.0’E, NMV J1029 (1 female, 17 mm). Bass Strait,

39°01.0’S, 143°22.1’E, 84 m, 31 Jan 1981, NMV J40641 (1 male, 8.5

mm). Eastern Bass Strait,38°18.0’S, 147°37.0’E, 55 m, 31 Jul 1983,

NMV J8818 (immature male, 8 mm). Bennison Channel, 38°49.0’S,

146°23.0’E, 6 m, 23 Nov 1983, NMV J12579 (manca 2, 6.5 mm). Tas.

Central Bass Strait, 39°32.8’S, 144°16.0’E, 18 m, 1 Nov 1980, NMV

J8830 (1 female, 13 mm).

Description of female. Head without dorsal elevations, antero-

lateral lobes rounded, rostral point present; lateral margin of

head and pereonite 1 incised. Pereonite 1 anterolaterally

extended. Pereonites 2 and 3 without dorsal elevations.

Pereonite 4 about 11 times length of pereonite 3, wider than

pereonites 2 and 3, anterolateral margins extended and angular.

Pereonites 5-7 progressively shorter posteriorly, lateral mar-

gins not expanded. Pleon length greater than combined lengths

of pereonites 5-7, with 2 sets of lateral wings, apex acute.

Eyes small and round. Antenna 1 reaching just past the

distal edge of the second peduncular article of antenna 2; aes-

thetascs present distally and laterally on flagellum. Antenna 2

slender, more than half length of the body; flagellum of 2

articles ending with claw, row of scales along full length.

Maxilla 1 mesial lobe with 3 terminal setae; outer lobe with

10 robust setae. Maxilla 2 mesial lobe with 20 setae; middle

lobe with 3 setae; lateral lobe with 3 setae. Maxillipedal endite

with 12 mesial setae; 1 coupling hook; palp article 2 with

mesial setal rows; article 3 with mesial and lateral setal rows;

article 4 with mesial and lateral setal rows; article 5 with mesial

and distal setae.

Pereopod 1 propodus and carpus similar length; dactylus

almost twice as long as wide, without unguis. Pereopods 2-4

slender, with dactylus. Pereopods 5-7 dactylus not denticulate.

Isopods of the genus Neastacilla from Australia

401

posteriorly; unguis and secondary unguis present; secondary

unguis less than half length of primary unguis. Uropod exopod

with 2 setae distally. Oostegite 4 with suture at midlength.

11-17 mm.

Male. Pereonites, mouthparts, pereopods as for female.

Pleopod 1 exopod with lateral notch and 2 plumose setae of

unequal lengths. Pleopod 2 with appendix masculina with ridge

on posterior face, apex simple, curved, not extending past the

endopod. Penial plate widened proximally, apex simple.

8-15.5 mm

Distribution. Australia: Victoria, Tasmania; subtidal to 84 m

depth.

Discussion. Guiler (1949) called the species 'mono seta'

because he found only one seta on the uropod exopod. This has

subsequently been found to have been erroneous and every

specimen examined for this redescription had two setae on the

uropodal exopod. The best distinguishing characteristic for this

species is its possession of an elongate distally acute pleotelson

that flares out laterally into two acute wings.

Neastacilla sheardi (Hale)

Astacilla sheardi Hale, 1946: 183-184, fig. 13.

Neastacilla sheardi. — Poore et al., 2002: 259.

Diagnosis of female. Eyes small and subtriangular. Head with

dorsal elevation (with 2 apices) slightly posterior to eyes, and

small single dorsal elevation posteriorly; lateral margin of head

and pereonite 1 not incised. Pereonite 1 with single small

dorsal elevation. Pereonites 2-3 dorsally smooth, with small

lateral expansions present. Pereonite about six times as long as

pereonite 3, with anterolateral expansions, with pair of blunt

dorsal elevations at midlength and blunt dorsal elevation

posteriorly. Pereonites 5-7 dorsally smooth. Pleon as long as

pereonites 5-7 combined, with 2 pairs of lateral wings, apex

subacute. 5 mm.

Male. Unknown.

Distribution. Australia: South Australia (Spencer Gulf):

subtidal.

Remarks. This species is diagnosed using the description of

Hale (1946). believe that Hale’s single specimen was an imma-

ture female, as the marsupium does not look to be fully formed.

The specimen drawn by Hale (1946) is similar to Neastacilla

inaequispinosa but can be easily distinguished from it by the

much broader pleon, the shape and ornamentation of the head

and slightly more robust antenna 2. If N. sheardi is found to be

an immature specimen, on further examination, it may more

closely resemble N. marrimarri or N. tuberculata.

Neastacilla soelae sp. nov.

Figures 20-21

Material examined. Holotype. Australia: WA, between Dampier and

Port Hedland, 19°59.3’S, 117°03.6'E, 52 m, 22 Feb 1983, NMV J16652

(1 female, 7 mm).

Paratypes. Australia: WA. Between Dampier and Port Hedland, type

locality, NMV J16930 (1 female, 8 mm); 19°27.2'S, 118°58.6'E, 36-46

m, 8 Dec 1982, NMV J40681 (1 female, 9 mm); 19°29.7'S, 118°52.2'E,

39 m, 24 Oct 1983, NMV J40682 (1 female, 7.5 mm); 19°56.7'S,

117°53.8'E, 43 m, 26 Aug 1983, NMV J40683 (1 female, 8.5 mm).

Description of female. Head with 2 dorsal elevations between

eyes and smaller dorsal elevation anterior to eyes, with setae on

elevations, anterolateral lobes angular, rostral point present; lat-

eral margin of head and pereonite 1 not incised, with small

flared anterolateral extension. Pereonite 2 with 1 dorsal tuber-

culate elevation with setae. Pereonite 3 with 2 dorsal elevations

with setae. Pereonite 4 about 10 times as long as pereonite 3;

with small tuberculate dorsal elevations with dorsal setae along

length, anterolateral margins extended and rounded. Pereonites

5-7 with tuberculate elevations with setae. Pleon longer than

combined lengths of pereonites 5-7; with small tuberculate

dorsal elevations with dorsal setae, with lateral wings, long and

tapering, apex acute.

Eyes small and round. Antenna 1 reaching past second

peduncular article of antenna 2; aesthetascs present distally and

laterally on flagellum. Antenna 2 slender, more than half as

long as body; flagellum of 2 articles ending with claw, row of

scales undetected.

Maxilla 1 mesial lobe with 3 terminal setae; lateral lobe with

10 distal robust setae. Maxilla 2 mesial lobe with 16 plumose

setae, middle lobe with 3 setae, lateral lobe with 3 setae.

Maxillipedal endite with 9 mesial setae; 1 coupling hook; palp

article 2 with mesial setal rows, articles 3 and 4 with mesial and

lateral setal rows, article 5 with mesial and distal setae.

Pereopod 1 propodus slightly shorter than carpus; dactylus

longer than wide; without unguis. Pereopods 2-4 without

dactylus. Pereopods 5-7 dactylus not denticulate, with unguis

and secondary unguis; secondary unguis setiform and slender,

only slightly smaller than primary unguis. Uropodal exopod

with 2 setae of equal length. Oostegite 4 with a posterior suture.

7-9 mm.

Male. Unknown.

Distribution. Australia: Western Australia; subtidal to 52 m

depth.

Etymology. This species is named after the research vessel from

which the specimen was collected, RV Soela.

Remarks. This species is from a region (north-western

Australia) where little is known about the arcturid fauna. The

long tapered pleotelson, loss of dactyli on pereopods 2-4 (a

state possessed by N. yuriel sp. nov.) and the setose secondary

dactlyus on pereopods 5-7 (also found in N. lawadi sp. nov.)

characterise this species.

Neastacilla tarni sp. nov.

Figures 22-24

Material examined. Holotype. Australia: SA, Topgallant I.,

Investigator Group, 33°43.0’S, 134°36.6’E, 15 m, , K. Brandon and

G.C.B. Poore, 21 Apr 1985, NMV J16579 (1 female, 6.0 mm).

Paratypes. Australia: SA. type locality, NMV J40676 (1 male, 7.0

mm); NMV J40677 (6 females, 4.5-5.0 mm; 7 males, 5.0-6.5 mm; 3

manca 2, 3.0-3. 5 mm; 2 manca 1, 2.5 mm).

Description of female. Head with dorsal elevation (with 2

apices) posterior to eyes, anterolateral lobes angular, rostral

point absent; lateral margin of head and pereonite 1 not incised.

402

Rachael A. King

Figure 21. Neastacilla soelae sp. nov., female holotype (NMV J 16652): PI, P3, P5, U. Scale: a (PI, P3) = 0.5 mm; b (U) = 0.5 mm

Pereonite 1 with posterior dorsal elevation. Pereonites 2 and 3

with dorsal and dorsolateral elevations, lateral margins

extended. Pereonite 4 about 6 times as long as pereonite 3; with

2 dorsal elevations at midlength, 2 dorsolateral elevations pos-

terior to them, a posterior dorsal elevation with 2 apices and 2

small dorsal elevations posteriorly; anterolateral margins

extended and rounded. Pereonites 5-7 progressively shorter

posteriorly, with small posterior dorsolateral and lateral ele-

vations. Pleon longer than combined lengths of pereonites 5-7,

with lateral wings, apex blunt.

Eyes small and subtriangular. Antenna 1 reaching to the end

of the second peduncular article of antenna 2; aesthetascs pres-

ent laterally on flagellum. Antenna 2 slender, more than half as

long as body; flagellum of 3 articles ending in claw, 2 rows of

scales along full length.

Maxilla 1 mesial lobe with 3 terminal setae; lateral lobe with

9 distal robust setae. Maxilla 2 mesial lobe with 13 plumose

setae, middle lobe with 4 setae, lateral lobe with 3 setae.

Maxillipedal endite with 10 mesial setae; 1 coupling hook; palp

article 2 with mesial setal rows, article 3 with mesial and

lateral setal rows, article 4 with mesial and lateral setae, article

5 with distal setae.

Pereopod 1 propodus as long as carpus; dactylus slightly

longer than wide, without unguis. Pereopods 2-4 with small

dactylus. Pereopods 5-7 dactylus denticulate, with unguis and

secondary unguis; secondary unguis half length of primary

unguis. Uropodal exopod with 2 setae of subequal length.

Oostegite 4 with sutured small posterior lobe and thickened dis-

tolateral edges. A pair of fleshy pads are visible on the ventral

surface of pereonite 5 that may be vestigial fifth oostegites.

4.5-6 mm.

Male. With less ornamentation than female. Head with 1

dorsal elevation between eyes. Pereonite 1 without dorsal ele-

vations. Pereonites 2 and 3 without dorsal elevations. Pereonite

4 about 8 times as long as pereonite 3, constricted for the first

quarter length, with 2 small elevations at quarter length and a

404

Rachael A. King

Figure 23. Neastacilla tarni sp. nov., female holotype (NMV J16579): Al, A2, left MXP, MXl, MX2, MD; PI, P3. Scales; a (Al) = 0.25 mm; b

(A2) = 0.5 mm; c (MXP, MXl, MX2, MD) = 0.25 mm; d (PI, P3) = 0.5 mm.

posterior elevation. Pereonites 5-7 progressively shorter pos-

teriorly, without dorsal elevations. Pleon longer than combined

lengths of pereonites 5-7 , with lateral wings, apex blunt.

Antennae, mouthparts, pereopods and uropods as for

female.

Pleopod 1 exopod with lateral notch and 2 plumose lateral

setae of equal length. Pleopod 2 appendix masculina with ridge

on posterior face; curved; extending quarter length past the dis-

tal edge of the endopod; apex simple. Penial plate widened

proximally, apex simple.

5-7 mm.

Distribution. Australia: South Australia; subtidal.

Etymology. “Tarni” is an Australian Aboriginal name from the

Kauma language group in South Australia. It means “the sea”.

Remarks. The female of this species resembles Neastacilla

tuberculata (Thomson, 1879) from New Zealand. However the

arrangement of the dorsal tubercles on pereonite 4 and the

shape of the pleotelson distinguish the two species; in N. tarni

the dorsal tubercles are medially and posteriorly placed on

pereonite 4 and the pleotelson is narrow and sculpted with a

blunt apex, whereas in N. tuberculata the tubercles are in the

Isopods of the genus Neastacilla from Australia

405

Figure 24. Neastacilla tarni sp. nov., female holotype (NMV J16579): P5, U. Male (NMV J40676): PLl, PL2, Pe. Scales: a (PLl, P12, Pe) = 0.2

mm; b (P5) = 0.5 mm.

anterior first half of pereonite 4 and the pleotelson is wide and

tapered to an acute apex.

Neastacilla tattersalli Lew Ton and Poore

Neastacilla falclandica. — Tattersall, 1921; 244, pi. 10, fig. 1 (not

Ohlin, 1901: 266, pi. 20).

Neastacilla tattersalli Lew Ton and Poore, 1986a: 193-195, fig.

3.— Lew Ton and Poore, 1986b: 99.— ICZN, 1987: 214.

Diagnosis of female. Eyes large and oval. Head without dorsal

elevations, anterolateral lobes angular; lateral margin of head

and pereonite 1 not incised. Pereonites 1-7 without dorsal ele-

vations. Pereonites 2-4 with only small lateral expansions.

Pereonite 4 9 times as long as pereonite 3. Pleon slightly longer

than pereonites 5-7 combined, no distinct lateral wings present,

apex bluntly rounded. Size unrecorded.

Male. Unknown.

Distribution. New Zealand, North Island.

Remarks. This species is diagnosed from the description of Lew

Ton and Poore (1986) and most closely resembles Neastacilla

falclandica. N. tattersalli can be distinguished from the former

species by the long tapered pleon without lateral wings, angu-

lar anterolateral lobes of the head and the presence of two setae

on the uropodal exopod.

Neastacilla tharnardi sp. nov.

Figures 25-27

Material examined. Holotype. Australia: Vic., Portland, Saxon Reef,

38°18.5’S, 14U38.5’E, 11 m, R. Wilson, 5 Mar 1992, NMV J24200 (1

female, 5 mm).

Paratypes. Australia: SA. Tiparra Bay, Tiparra Reef, 34°04.0’S,

137°23.0’E, 10 m, G.C.B. Poore and H.M. Lew Ton, 15 Mar 1985,

NMV J16575 (1 male, 5 mm). Investigator Group, 33°43.0’S,

134°36.6’E, 20 m, K. Brandon and G.C.B. Poore, 21 Apr 1985, NMV

J47326 (manca-2, 3 mm).

Description of female. Head with 2 dorsal elevations (each with

3 apices) between the eyes, anterolateral lobes angular with

small tubercles, small rostral point present; lateral margin of

head and pereonite 1 incised. Pereonite 1 with dorsal elevation

Isopods of the genus Neastacilla from Australia

407

Figure 26. Neastacilla tharnardi sp. nov., female holotype (NMV J24200): Al, A2, left MXP, MXl, MX2, MD; PI, P3, P4. Male (NMV J16575):

A16. Scales; a (Al, Al6, A2) = 0.5 mm; b (MXP, MXl, MX2, MD) = 0.2 mm; c (PI, P3, P4) = 0.5 mm.

with 3 apices; with large, laterally curved anterolateral expan-

sion. Pereonite 2 with dorsal elevation with single apex, with

narrow lateral extensions. Pereonite 3 with dorsal elevation

with 3 apices, with narrow lateral extensions. Pereonite 4 about

4 times as long as pereonite 3, with 4 anterior dorsal elevations,

2 dorsal elevations with many apices at midlength and 2 small

dorsal elevations posteriorly, with anterolateral margins

extended. Pereonites 5-7 progressively smaller, with small pos-

terior dorsolateral elevations. Pleon longer than combined

lengths of pereonites 5-7, with lateral wings, apex acute.

Eyes small and round. Antenna 1 reaching to the end of the

third peduncular article of antenna 2; aesthetascs present later-

ally on flagellum. Antenna 2 slender, more than half as long as

body; flagellum of two articles and claw, second article very

small, with a row of scales along full length.

Maxilla 1 mesial lobe with 3 terminal setae; lateral lobe with

10 distal robust setae. Maxilla 2 mesial lobe with 14 plumose

setae, middle lobe with 4 setae, lateral lobe with 3 setae.

Maxillipedal endite with 14 mesial setae; 1 coupling hook; palp

article 2 with mesial setal rows, article 3 with mesial and

lateral setal rows, article 4 with mesial and lateral setae, article

5 with mesial and distal setae.

Pereopod 1 propodus as long as carpus; dactylus slightly

longer than wide, without unguis. Pereopods 2 and 3 with small

dactylus. Pereopod 4 without dactylus. Pereopods 2-4 slightly

dorsoventrally flattened, with tuberculate lateral elevations at

proximal edges of the merus and carpus. Pereopods 5-7 dacty-

lus not denticulate, with primary and secondary unguis;

secondary unguis half length of primary unguis. Uropodal exo-

pod with 2 setae of subequal length. Oostegite 4 without a

408

Rachael A. King

Figure 27. Neastacilla thamardi sp. nov., male (NMV J16575): PLl, PL2, Pe. Female holotype (NMV J24200): U; A, ventral view with ooste-

gites. Scales: a (U) = 0.5 mm; b (P5) = 0.5 mm; c (A) = 1.0 mm; d (PLl, PL2, Pe) = 0.5 mm

suture, with ventrally projecting elevation posterior to insertion

of pereopod 4.

5 mm.

Male. With less ornamentation than female. Head with 2

dorsal elevations (each with 2 apices) between eyes, anterolat-

eral margins angular with small tubercles. Pereonite 1 with

dorsal elevation with a single apex. Pereonite 2 without dorsal

elevations, with lateral extensions. Pereonite 3 with dorsal

tubercle with single apex, with lateral extensions. Pereonite 4

about 7 times as long as pereonite 3, constricted for the first

quarter length; with 2 anterior dorsal elevations at midlength.

Pereonites 5-7 progressively smaller, with small posterior dor-

solateral elevations. Pleon longer than combined lengths of

pereonites 5-7, with lateral wings, apex acute.

Antennae, mouthparts, pereopods and uropods as for

female.

Pleopod 1 exopod with lateral notch and 2 plumose lateral

setae of unequal length. Pleopod 2 appendix masculina with

ridge on posterior face; curved; extending quarter length past

the endopod; apex simple. Penial plate proximally widened,

distally tapered and apex simple.

5 mm.

Distribution. Australia: Victoria, South Australia; subtidal to 20

m depth.

Etymology. “Tharnardi” is an Australian Aboriginal word in the

Yindjibarndi language from north-western Australia. It means

“the sea”.

Remarks. This species of Neastacilla superficially most resem-

bles species of Parastacilla Hale, 1946 (King, 2000). The lat-

eral extension of the head and pereonite 1 as well as the mor-

phology of the anterior pereonites are very similar and no other

Neastacilla species. However, in N. tharnardi pereopods 2 and

3 possess dactyli with claws, antenna 2 is elongate and there are

no dorsolateral wings on the pleotelson.

Isopods of the genus Neastacilla from Australia

409

Neastacilla tuberculata (Thomson)

Figures 28-29

Arcturus tuberculatus Thomson, 1879: 416-417, pi. 19 figs 1-4. —

Thomson, 1881: 206, pi. 7 fig. 2. — Thomson and Chilton, 1886:

156. — Filhol, 1885, 437. (not Arctums tuberculatus Latreille, 1829,

junior synonym of Arctums baffini Sabine).

Astacilla tuberculata. — Hurley, 1961: 264.

Neastacilla tuberculata. — Poore, 1981: 333.

Material examined. New Zealand: Lyttelton Harbour, 5 m, Aug 1997,

NMV J40643 (1 male, 4.5 mm). Otago Harbour, 12.1 m, 4 Oct 1965,

NMV J 16559 (1 female, 4.5 mm), NMV J 16560 (1 male, 4 mm), NMV

J 1655 8 (1 female, 5 mm) NMV J 16562 (1 male, 5 mm). Otago

Harbour, 6 m, 18 Jun 1965, NMV J 16560 (3 females, 4-4.5 mm; 5

males, 3.5-5 mm; 2 juveniles, 3 mm). Otago Harbour, 13 m, 30 Apr

1965, NMV J16561 (1 male, 5 mm). Otago Harbour, 13 m, 4 Oct 1965,

NMV J16556 (1 female, 5 mm). Otago Harbour, 4.1 m, 13 Jun 1965,

NMV J12927 (1 female, 4 mm). Otago Harbour, 2.9 m, 7 Jan 1967,

NMV J40644 (1 female, 4.5 mm). The Snares, 146 m, 26 Nov 1974,

NMV J 16557 (1 male, 4.5 mm).

Description of female. Head with 2 dorsal elevations between

eyes, small dorsal elevation in front of eyes and an elevation

posterior to eyes on dorsal midline, anterolateral margins

angular, rostral point absent. Lateral margin of head and pereo-

nite 1 not incised. Pereonite 1 with 2 elevations along dorsal

line and pair of dorsolateral elevations. Pereonites 2 and 3 pro-

gressively wider; with dorsal and dorsolateral elevations,

anterolateral margins extended. Pereonite 4 about 7 times as

long as pereonite 3, with a large dorsal elevation (with 3 apices)

covering the first two-thirds length, small anterior dorsolateral

elevations,dorsolateral elevations at midlength, a posterodorsal

elevation on the dorsal line and 2 small dorsolateral elevations

posterior to that, with anterolateral margins rounded and

extended. Pereonites 5-7 progressively smaller, with small dor-

sal and dorsolateral elevations, lateral margin extended. Pleon

longer than combined lengths of pereonites 5-7 , with 2 sets of

lateral wings, with small dorsal and dorsolateral elevations,

flared posteriorly and tapering to acute apex.

Eyes small and round. Antenna 1 reaching to the end of third

peduncular article of antenna 2; aesthetascs attached laterally

and distally on flagellum. Antenna 2 slender, more than half as

long as body; flagellum of 3 articles and claw, with row of

scales along full length.

Maxilla 1 inner lobe with 3 terminal setae; outer lobe with

10 distal robust setae. Maxilla 2 inner lobe with 18 plumose

setae, middle lobe with 3 setae, outer lobe with 3 setae.

Maxillipedal endite with 14 mesial setae; 1 coupling hook; palp

article 2 with mesial setal rows, article 3 with mesial and later-

al setal rows, article 4 with mesial setae, article 5 with mesial

and distal setae.

Pereopod 1 propodus as long as carpus; dactylus slightly

longer than wide, without unguis. Pereopods 2-4 with small

dactylus. Pereopods 5-7 progressively smaller; dactylus den-

ticulate, with unguis and secondary unguis; secondary unguis

half length of primary unguis. Uropodal exopod with 2 setae of

subequal length. Oostegite 4 with sutured small posterior lobe.

4-5 mm.

Male. With less ornamentation than female. Head with 2

dorsal elevations (sometimes fused) between eyes; antero-

lateral margins angular; rostral point absent; lateral margin of

head and pereonite 1 not incised. Pereonite 1 with small dorsal

and dorsolateral elevations. Pereonite 2 and 3 with dorsal and

dorsolateral elevations. Pereonite 4 about 6 times as long as

pereo-nite 3; with an anterior dorsal elevation at third length, a

small posterior dorsal elevation and 2 small posterior dorsolat-

eral elevations. Pereonites 5-7 progressively smaller without

distinct dorsal elevations except for pereonite 5 which has a

small posterior dorsal elevation. Pleon longer than combined

lengths of pereonites 5-7, with 2 sets of lateral wings, with

small dorsal elevations, tapering to an acute apex.

Antennae, mouthparts, pereopods and uropods as for

female.

Pleopod 1 exopod with lateral notch and 2 plumose lateral

setae of equal length. Pleopod 2 appendix masculina with ridge

on posterior face; curved; extending more than third length past

the endopod; apex tapered. Penial plate simple and straight.

3. 5-5.0 mm.

Distribution. New Zealand; subtidal to 201 m depth.

Remarks. This species was briefly described by Thomson

(1879) who illustrated only the lateral view of a female. The

highly ornamented body, particularly the dorsally elevated

pereonite 4, makes it distinguishable from other New Zealand

arcturids.

Neastacilla yuriel sp. nov.

Figures 30-32

Material examined. Holotype. Australia: SA, Flinders I., Hotspot Reef,

33M0.5'S, 134°22.0'E, 17 m, G.C.B. Poore, SCUBA, 19 Apr 1985,

NMV J16553 (1 female, 3.5 mm).

Paratypes. Australia: SA, collected with holotype, NMV J40678 (1

female, 3 mm), NMV J40693 (1 male, 3.5 mm), NMV J40694 (1 male,

4 mm), NMV J16554 (4 females, 3.5-4 mm), NMV J16549 (1 male,

3.5 mm). Flinders I., Hotspot Reef, 33°40.8’S, 134°22.5'E, 21 m, 20

Apr 1985, NMV J 16547 (manca 2, 3 mm). Topgallant I., 33°43.0'S,

134°36.6'E, 25 m, 21 Apr 1985, NMV J16546 (1 male, 3 mm);

33°43.0'S, 134°36.6'E, 12 m, 21 Apr 1985, NMV J16548 (2 females,

3-3.5 mm; 2 males, 4 mm; manca 2, 2.5 mm). Vic. Central Bass Strait,

38°33.4'S, 144°54.9'E, 55 m, 12 Nov 1981, NMV J8853 (1 male, 4

mm). Tas. Spiky Bridge coastal reserve, 42°08.0'S, 148°08.0E, 4 m, 21

Mar 1988, NMV J40648 (1 female, 3.5 mm; 2 manca 2, 2.5 nmi).

Bicheno, 41°53.0'S, 147°18.0'E, 7 m, 23 Mar 1988, NMV J40645 (1

female, 4 mm). WA. Breaksea I., 35°03.9'S, 118°02.9'E, 15 m, 7 Apr

1984, NMV J 16632 (2 females, 3 mm; 1 male, 3.5 mm). King George

Sound, 35°00.7'S, 118°10.1’E, 27 m, 15 Apr 1984, NMV J16630 (1

female, 3 nmi).

Description of female. Head with dorsal elevations slightly pos-

terior to the eyes, anterolateral margins angular, small rostral

point present; lateral margin of head and pereonite 1 not

incised. Pereonite 1 with 2 indistinct dorsolateral elevations.

Pereonites 2 and 3 with indistinct dorsal elevations, with

lateral margins extended. Pereonite 4 about 7 times as long as

pereonite 3, with large dorsal elevation at midlength, antero-

lateral margins rounded and extended. Pereonites 5-7 progres-

sively smaller, without distinct elevations. Pleon longer than

combined lengths of pereonites 5-7, with 2 sets of lateral

wings, without dorsal elevations, tapering to subacute apex.

Isopods of the genus Neastacilla from Australia

411

Figure 29. Neastacilla tuberculata (Thomson, 1879), female (NMV J16559); PI, P3, P5, U. Male (NMV J16560): PLl, PL2, Pe Scales: a (PI,

P3) = 0.5 mm; b (P5) = 0.5 mm; c (PLl, PL2, Pe) = 0.5 mm.

Antenna 1 reaching to end of third peduncular article of

antenna 2; aesthetascs present laterally and distally on flag-

ellum. Antenna 2 slender, more than half as long as body;

flagellum of 3 articles and claw; with row of scales along full

length.

Maxilla 1 inner lobe with 3 terminal setae; outer lobe with

10 distal robust setae. Maxilla 2 inner lobe with 18 plumose

setae, middle lobe with three setae, outer lobe with three setae.

Maxillipedal endite with 14 mesial setae; one coupling hook;

palp article 2 with mesial setal rows, article 3 with mesial and

lateral setal rows, article 4 with mesial setae, article 5 with

mesial and distal setae.

Pereopod 1 propodus as long as carpus; dactylus slightly

longer than wide, without unguis. Pereopods 2-4 with small

dactylus. Pereopods 5-7 progressively smaller, dactylus dentic-

ulate; with primary and secondary unguis; secondary unguis

half length of primary unguis. Uropodal exopod with 2 setae of

subequal length. Oostegite 4 with sutured small posterior lobe.

3^ mm.

Male. With less ornamentation than female. Head with

indistinct dorsal elevation between eyes, anterolateral margins

angular, very small rostral point present; lateral margin of head

and pereonite 1 not incised. Pereonite 1 with indistinct dorsal

and dorsolateral elevations. Pereonite 2 and 3 with indistinct

dorsal elevations, lateral margins not extended. Pereonite 4

about 6 times as long as pereonite 3; constricted in dorsal view

for first fifth of length, with 2 small posterior dorsal elevations

at midlength. Pereonites 5-7 progressively smaller, without

dorsal elevations. Pereonites 6 and 7 without dorsal elevations.

Pleon longer than combined lengths of pereonites 5-7, with 2

sets of lateral wings, with small dorsal elevations, tapering to a

subacute apex.

Antennae, mouthparts, pereopods and uropods as for

female.

Pleopod 1 exopod with lateral notch and 2 plumose setae of

equal length. Pleopod 2 appendix masculina with ridge on

414

Rachael A. King

Figure 32. Neastacilla yuriel sp. nov., male (NMV J40693); PLl, PL2, pleopod 2. Female holotype (NMV J16553): U; A, ventral view with oost-

egites; P5. Scales: a (A) =1.0 mm; b (P5) = 0.5 mm; c (PLl, PL2, Pe, U) = 0.2 mm.

posterior face; curved; extending third length past endopod;

apex simple. Penial plate unknown.

Distribution. Australia: Victoria, Tasmania, South Australia,

Western Australia; subtidal to 55 m depth.

Etymology. "Yuriel" is an Australian Aboriginal word for

"coastal bay" in reference to the area specimens were first taken

from.

Remarks. This species is the smallest of the Australian arcturids

(2.5-4 mm) making the species instantly recognisable. The

ornamentation of females has been seen to differ slightly with-

in populations. In a few specimens the dorsal elevation on

pereonite 4 possessed two apices and the small elevations on

the head and pereonites 1^ were more enlarged.

Acknowledgements

This work was conducted as part of a postgraduate degree at the

University of Melbourne and Museum Victoria with funding

from an Australian Postgraduate Award. I am extremely grate-

ful to Dr Gary Poore for encouragement and advice and also to

the staff and students at Museum Victoria for support. I thank

Helen Lew Ton, whose unpublished honours thesis provided a

basis for this study of Neastacilla and Professor Alan Myers for

helpful suggestions on the manuscript.

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