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CONTENTS

Revision of Tertiary species of Anaskopora Wass (Bryozoa: Cribrimorpha)

P. E. Bock and P. L. Cook .

Revision of the multiphased genus Corbulipora MacGillivray (Bryozoa: Cribrimorpha)

P. E. Bock and P. L. Cook .

Notes on the genera Nordgaardia and Uschakovia (Bryozoa: Bugulidae)

P. L Cook .215

New species and a new record of Chimarra Stephens (Trichoptera: Philopotamidae) from

Bougainville Island, Papua New Guinea

D. L Cartwright .223

New species of Hydropsychidae (Insecta: Trichoptera) from northern Australia

J. C. Dean .231

Rcdcscriplion of Bimgona Barker with new synonyms in the Australian Baetidae

(Insecta: Ephemeroplera)

P. J. Suter and M. J. Pearson .247

Descriptions of new species of Biruhius (Amphipoda: Phoxocephalidac) from Australia and

Papua New Guinea with comments on the Biruhius-Kulgaphoxus-Tickalerus-Yan complex

J. Tavlor and G. C. B. Poore .255

Bathyal Jocropsididae (Isopoda: Asellota) from south-eastern Australia, with description

of two new genera

J.Just .297

Platy’protus phyllosoma, gen. nov and sp. nov., from Enderby Land, Antarctica, an unusual

munnopsidid without natatory pereopods (Crustacea: Isopoda: Asellota)

J. Just .235

Three new species ofCirolana Leach, 1818 (Cnistacea: Isopoda: Cirolanidac) from Australia

S. J. Keable .247

Redescriplion of the tropical Australian isopod, Lyidotea nodata Hale, 1929 (Crustacea: Idoteidae)

R. King and G. C. B. Poore .265

Plakarthriiim australiense, a third species of Plakarthriidae (Crustacea: Isopoda)

G. C. B. Poore and A. Brandi .273

A phylogeny of the Leptostraca (Crustacea) with keys to families and genera

G. K. Walker-Smith and G. C. B. Poore .383

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Memoirs of Museum Victoria 58(2): 179-189 (2001)

REVISION OF TERTIARY SPECIES OF ANASKOPORA WASS

(BRYOZOA: CRIBRIMORPHA)

P.E. Bock' and P.L. Cook^

'School of Ecology and Environment, Deakin University, 661 Blackburn Road, Clayton, Vic. 3168, Australia

(pbock(§}deakin.edu.au)

^Honorary Associate, Museum Victoria, GPO Box 666E, Melbourne, Vic. 3001, Australia

Abstract

Bock, P.E. and Cook, P.L., 2001. Revision of Tertiary species oi Anaskopora Wass (Bryozoa:

Cribrimorpha). Memoirs of Museum Victoria 58(2): 179-189.

The subgenus Anaskopora Wass, 1975 is raised to generic rank, separated from the genus

Corhiilipora, and redefined. The type species. Crihrilina elevata MacGillivray, 1895, is a Ter¬

tiary fossil from Victoria with small globular colonies formed principally by a special kind of

interzooidal frontal budding. Other Tertiary fossil species with a similar colony structure, here

assigned to Anaskopora, arc Chbrtlma comma MacGillivray, 1895 and Lepratia rotundaia

MacGillivray, 1895. Two further new Tertiary species, A. simplex and A. mesa, from Victoria

and South Australia have small encrusting colonies. A key to species is given.

Introduction

Revision of the cribrimorph genus CorbiiUpora

MacGillivray, 1895 means that its subgenus

Anaskopora Wass, 1975 also requires redefini¬

tion (Bock and Cook, 2001). Anaskopora is con¬

sidered here to be generically distinct Irom

Corhiilipora and to include Crihrilina conwta

MacGillivray, 1895, a species which Wass (1975)

referred to Corhnlipora.

Frontal budding is unusual in the Cribri¬

morpha. It occurs in Corhnlipora principally as a

result of expansions of cuticle and coelomic

tissue derived from the pclmatidia on the frontal

surface of the costae which form the calcified

frontal shield or pcricyst. Pelmatidia arc usually

absent in Anaskopora and if present arc infre¬

quent and restricted to the margins of the

horizontal median area of costal fusion of the

pericyst. Frontal buds arc however commonly

present and these arise in a series, often sur¬

rounding each autozooid and originate from a

special type of pore-chamber. These also oecur in

Corhnlipora and arc termed chambered pores

(Bock and Cook, 2001). Chambered pores consist

of a calcified chamber arising around one or

more septular pores in a vertical zooid wall. The

chamber is uncalcificd frontally and usually

expands to form an interzooidal kenozooid.

These pores somewhat resemble the pore-

chambers found in CeUeporeUa and Hippothoa

(Gordon and Hastings, 1979) but arc larger and

less regularly positioned. Avicularia occur in

series with these pores which arc also the origin

of rhizoids in Recent species (Arnold and Cook,

1997).

All the species described here arc fossils from

Victoria and South Australia and the specimens

are from the collections of Museum Victoria

(NMV). All species are illustrated by SEM, in

many cases for the first time.

Key to species Anaskopora

1. Autozooids large (Lz >1 mm). Costae curved, median area of costal fusion

narrow, lacunae rare. A. simplex sp. nov.

— Autozooids smaller (Lz <0.75mm). Costae with vertical marginal, and

horizontal median elements, median area of costal fusion wide.2

2. Vertical costal components distinct, well separated.3

— Vertical costal components very short, closely apposed....4

3, Colony usually a globular ectoproclolith. Costae <11, median area with 8-12

large, rounded lacunae. Pclmatidia absent. A. elevata

Colony enenisting minute substrata. Costae >14, marginally raised with a

single series of pclmatidia; medially horizontal with >15 scattered lacunae.

. A. mesa sp. nov

179

180

P. E. BOCK AND P. L. COOK

4. Median area of fusion with 12-24 regularly radiating costae, alternating with

rows of 6-8 lacunae. A. cornuta

— Median area of fusion with no obvious costae but with numerous scattered

minute lacunae with intervening tubercles. A. rotimdata

Anaskopora Wass

Corbulipora {Anaskopora)'H?iS?,, 1975: 170.

T\pe species. Crihrilina elevata MacGillivray,

1895.

Description. Colonics encrusting, usually on very

small substrata, with subsequent intcrzooidal

frontal budding and overgrowth forming globular

ectoproctoliths. Basal zooid walls calcified only

peripherally, autozooids with marginal gymno-

cyst and extensive, costatc pericyst. Peripheral,

vertical component of costae without lateral

fusions; central, horizontal component constitut¬

ing the median area of fusion. Lacunae present,

sometimes numerous; pelmalidia absent or rare,

marginal only. Secondary calcified orifice with or

without lateral condyles: peristome raised dis-

tally, forming a curv'cd plate which is often

marginally fimbriated. Oral spines paired, some¬

times wide and flattened, or inflexcd and fused

medially above the secondary calcified orifice.

Chambered pores large, forming kenozooids sur¬

rounding each autozooid; I -3 distally and 1-A lat¬

erally; calcification becoming extrazooidal later

in ontogeny. Avicularia interzooidal, budded in

series with kenozooids, usually distal or distolat-

eral and single, occasionally proximal and even

paired. Rostrum raised, palate rounded or sub-

triangular, orientated distally or laterally, with

paired condyles. Dimorphic brooding zooids and

ovicells unknown.

Remarks. The principal characters separating

Anaskopora from Corbulipora s.s. are the

regular presence of small interzooidal kenozooids

surrounding autozooids, the distal or distolatcral

interzooidal avicularia present beside each auto¬

zooid and the raised distal oral fimbriated plate.

The chambered pores resemble those of Corhuli-

pora but are more regularly placed surrounding

each autozooid and are presumably in communi¬

cation with both neighbouring autozooids. In the

globular colonies, nearly all budding after the ear¬

liest encrusting stages, is intcrzooidal and frontal.

Each new layer of autozooids is derived from

enlarged kenozooids of the previous layer. The

virtual absence of pelmatidia and the apparent

complete absence of multiphase growth also

distinguishes Anaskopora from Corbulipora.

In nearly all species of Anaskopora the median

area of fusion of each autozooid is delineated

sharply from the peripheral component of the

costae, making a distinct angle at the margin of

the shield where there is occasionally a single

pelmatidium. The vertical component is often

elongated and never bears any costal fusions

although the costae may be closely apposed in

some species. The species described here form a

series. In A. simplex sp. nov. the costae have no

distinct vertical clement and arc cur\^cd with only

terminal lateral fusions. At the other extreme, in

A. rotundata., costae arc hardly discemable and

the median area is wide, consisting of numerous

scattered lacunae and small tubercles. In contrast,

A. cornuta has regular rows of costae with numer¬

ous costal fusions whereas in A. elevata and

A. mesa sp. nov. the median area consists of scat¬

tered lacunae. The constant occurrence of a distal

avicularium and a distal plate, together with the

distinctive kenozooids arising from chambered

pores, suggests that although the frontal shield

shows a great diversity of structures these species

all form a fairly natural grouping. All species

appear to encrust minute substrata early in

aslogeny. A. elevata and A. rotimdata are known

only with globular frontally budded colonics;

A. simplex and A. mesa have few or no frontally

budded zooids. Colonies of A. cornuta. however,

occur in both states. All these species occur in the

Tertiary deposits of Victoria and South Australia.

Living species assignable to Anaskopora have

only recently been been recognised; they occur

from Queensland and New South Wales, and

were described by Arnold and Cook (1997) as

A. doliaris (Maplcstonc) and A. parkeri. These

Recent colonics arc small and globular and arc

anchored by well-developed rhizoids.

The complete absence of ovicells or of any

zooid with modified skeletal morphology which

might be intcr|:)reted as a brooding zooid raises

the suspicion that some kind of multiphase

growth may occur in Anaskopora as it does in

Corbulipora (Bock and Cook, 1994). However if

this were tnic any alternative growth phase or

phases which might have had a reproductive func¬

tion have not as yet been recognised in any fossil

or Recent assemblage. One species which was

referred to Anaskopora by Wass (1975;

A. ampulla, p. 170, pi. 8, figs 5, 6) remains

TERTIARY SPECIES OF THE BRYOZOAN ANASKOPORA

181

unrecognised in the material examined here.

Maplestone (1901: 207, pi. 37, llg. 9) originally

described Corbulipora ampulla from Momington

and the type slide (NMV PI0156) was examined

by Wass together with one of the specimens

described as C. sp. aff. ampulla by Brown (1958:

54). The type material consists of a few isolated

zooids only, one of which was presumably that

figured by Wass (1975). None of the zooids has a

distal avicularium which was, however, men¬

tioned by Brown in his specimens, and which arc

therefore assumed to belong to Anaskoponi. The

peripheral vertical component of the costae forms

an angle with the median area which has approx¬

imately 20 large rounded lacunae but no regularly

placed costal fusions. There are also about 20

costae, the vertical components being closely

apposed; according to Wass (1975) the distal pair

arc elongated to form a hood over the orifice.

Anaskopora ampulla has some characters similar

to those of/f mesa sp. nov. but differs in several

respects from that species.

Brown (1958: 55, fig. 33) also described as

Corbulipora pennata an encrusting Tertiary

species with costae raised and extended laterally.

A large distal avicularium was described but not

figured by Wass who redescribed and figured the

unique holotype. The encrusting habit and distal

avicularium suggest that C. pennata may be

referable to Anaskopora but as the specimen

is no longer present in the NMV collection its

assignment must remain uncertain.

Anaskopora elevata (MacGillivray)

Figures 1-2, Table 1

Crihrilhia elevata MacGillivray, 1895: 59, pi. 8, fig.

19.—Maplestone, 1904:20! (listed).

Corbulipora elevata. —Brown, 1958: 54.

Corbulipora (Anaskopora) elevata. —Wass, 1975:

170, pi. 8, figs 1.2,4.

Material examined. Lectotype. NMV P27641, Schnap-

per Point, Victoria, Miocene (selected by Wass, 1975).

Other material. Balcombe Bay, Bairnsdale, Fyans-

ford, Warrambine Creek, Muddy Creek.

Distribution. Additional records given by Maple-

stone (1904) include Sheiford, Griffins, Camp¬

bells Point, Momington and Mitchell River;

records in Brown (1958) include Localities XI

and XIV on the Glcnelg River at Werrikoo

(south-western Victoria), and at Mount Gambier,

South Australia. All records are Miocene

(Appendix).

Description. Anaskopora with globular colonies

formed by inlerzooidal frontal budding from an

early stage encrusting a very small substratum.

Autozooids with a distinct gymnocyst, frontal

shield with 8-10 costae, the vertical components

widely spaced, inflexed sharply to form a median

area of fusion. Costae bifurcate terminally, and

with some lateral fusions produce 8-12 relatively

large lacunae. Secondary calcified orifice not

greatly raised distally, with a pair of short oral

spines. Recessed junction of suboral costae form¬

ing a proximally sinuate secondary orifice. Distal

avicularium fairly large, length of rostrum equal

or exceeding that of autozooid orifice; rostmm

rounded and directed distally.

Remarks. The secondary orifice is distinctly sinu¬

ate. Many zooids communicate directly through

scptular pores and the chambered pores arc small,

forming kenozooids at the comers of the auto¬

zooids. The calcification becomes extrazooidal

late in ontogeny surrounding the autozooids.

Powell (1967: 223) suggested that C. elevata

might be ‘closely allied’ to the type species of

Crihralaria Silen, C. cunirostris Silen (1941:

122, figs 182-183) from the north-western

Pacific. Crihralaria curvirostris has autozooids

with numerous, rounded lacunae and no cham¬

bered pores. The ovicell is partially immersed in

the subrostra! chamber of the large distal avicu¬

larium which has a raised acute rostmm. The only

feature superficially similar to the avicularium of

A. elevata is its distal position (sec also Gordon,

1989: 15). Anaskopora elevata differs from the

other globular species in its small number of

widely spaced costae and scattered median

lacunae.

Table 1. Measurements in mm of species of Anaskopora. Lz, Iz, length and width of zooid; Lo, lo,

length and width of orifice; Lav, lav, length and width of avicularium; Lr, length of rostmm.

Lav

lav

A. elevata

0.46-0.54

0.37-0.50

0.14-0.15

0.13-0.17

0.17-0.23

0.10-0.12

0.09-0.15

A. conmta

0.66-0.78

0.45-0.66

0.15-0.18

0.12-0.16

0.12-0.18

0.08-0.10

A. rofundata

0.60-0.70

0.41-0.51

0.10-0.14

0.12-0.13

0.08-0.10

0.06-0.08

0.05-0.07

A. simplex sp. nov.

0.95-1.40

0.58-0.70

0.18-0.23

0.23-0.25

0.13-0.19

0.12-0.14

0.07-0.10

A. mesa sp. nov.

0.53-0.88

0.40-0.50

0.09-0.13

0.17-0.23

0.10-0.18

0.06-0.09

0.07-0.10

182

P. E. BOCK AND P. L. COOK

Anaskopora simplex sp. nov.

Figures 3—4, Table 1

Material examined. Holotype. NMV PI 40964,

Balcombe Bay, Victoria, Miocene.

Other material. Princelown, Cape Otway. Victoria.

Distribution. See above, Miocene to Oligoccnc

(Appendix).

Etymudogy. Simplex (Latin) — simple, referring

to the pericyst.

Description. Colonies encrusting minute sub¬

strata, with occasional isolated frontal buds aris¬

ing from small chambered pores at the comers of

the autozooids. Autozooids large, gymnocyst

marginal; pericyst fomied by 18-28 flattened

costae, widely spaced and curv'cd over the frontal

with no sharp angle of inflexion. Median area of

fusion narrow, formed by apposition and fusion

of terminal and subtcrminal costal bifurcations,

with very few lateral fusions. Secondary calcified

orifice raised to form fimbriated distal plate. Sub¬

oral bar raised and swollen; with 1 pair of oral

spines. Distal avicuiarium relatively small, the

rostrum less than half length of autozooid orifice,

subtriangular to rounded, with large paired

condyles. Chambered pores include more than 1

septular pore and tend to occur at comers of auto¬

zooids, forming small kenozooids and extra-

zooidal calcification late in ontogeny.

Remarks. The specimens from Princctown and

Cape Otway include only a few isolated groups of

zooids but the material from Balcombe Bay is

plentiful comprising more than 250 zooids most

of which encrust fragments of the bryozoans

Porina and Laminopora. The largest specimen

has more than 80 autozooids. The relative sim¬

plicity of the costal shield resembles those of

Figularia and Membraniporella and the apposed

ends of the costae arc raised in small tubercles

similar to those of of the species illustrated as A/.

bifurca Powell by Gordon (1984: pi. 19A).

Anaskopora cornuta (MacGillivray)

Figures 5-7, Table 1

Cribrilina cornuta MacGillivray, 1895: 58, pi. 8, figs

10-12.—Maplestone, 1904: 201 (listed).

Corbulipora cornuta.—Brown, 1958: 55.—Wass,

1975: 168. pi. 7 figs 3-6, pi. 8 fig. 3.

Material examined. Lectotype. NMV P27635,

Schnapper Point, Victoria. Miocene (selected by Wass,

1975).

Other material. Balcombe Bay, Warrambine Creek.

Muddy Creek, Cooriemungle, Princetowii, Victoria and

Mount Schanck, South Australia.

Distribution. Additional records given by

MacGillivray (1895) include Corio Bay and Gel-

librand; by Maplcstonc (1904) include Spring

Creek, Shelford, Griffins, Campbells Point,

Momington and Mitchell River: and by Brown

(1958) include localities XL Glenelg River at

Werrikoo, and XVI, Crawford River at Glenaulin,

all in Victoria. All records are Miocene, except

Spring Creek and Glenaulin which arc Oligocene

(Appendix).

Description. Colonics encrusting minute sub¬

strata, or cylindrical to globular, principally

formed by interzooidal frontal budding. Auto¬

zooids with a narrow gymnocyst and a pericyst

fonned by 12-24 costae; vertical components

short and closely apposed. Median area effusion

very regular, with radiating rows of small lacunae

between the costae, which overlap or abut ter¬

minally. Secondary calcified orifice small with a

slight but distinct distal plate, paired oral spines

and condyles. Recessed junction of the suboral

costae forming a proximal sinus. Avicuiarium

distal or dislolatcral, sometimes in pairs or

triads, with a short, rounded, raised rostrum about

two-thirds length of aulozooidal orifice. Cham¬

bered pores not obvious but small interzooidal

kenozooids arc numerous surrounding each auto¬

zooid and becoming extrazooidal in older

colonics.

Remarks. Like ,4. rotundata, the globular colonies

have randomly orientated zooids and are

almost completely composed of frontal buds.

Anaskopora cornuta differs from A. elevata in its

dimensions and much more numerous straight

rows of costae and lacunae. It differs from

A. rotundata in the regularity of the rows of

costae and lacunae and its slightly larger

avicularia. A significant number of the many

specimens examined encrust small shell and bry-

ozoan fragments like the colonics of A. simplex

and A. mesa.

Anaskopora rotundata (MacGillivray)

Figures 8, 12-16, Table I

Lagenipora rotundata MacGillivray, 1895: 78, pi.

10, fig. 18.

Lagenipora morningtonien.sis Maplcstone. 1902: 24,

fig. 15.— Maplestone. 1904: 216 (listed).

Material examined. Holotype. NMV P27688, Lake

Bullenmerri, Victoria. Miocene.

Holotype of L. morningtoniensis. NMV PI0205.

Momington. Victoria, Miocene.

Other material. Balcombe Bay, Baimsdale, Fyans-

ford, Cooriemungle, Victoria and Mount Schanck,

South Australia.

TERTIARY SPECIES OF THE BRYOZOAN ANASKOPORA

183

Distribution See above, Miocene (Appendix).

Description. Colonics globular, fomied princi¬

pally by interzooidal frontal budding and some

overgrowth, from zooids encrusting very small

substrata. Autozooids with distinct, narrow gym-

nocyst, pericyst fomicd by 20-24 closely apposed

costae with very little vertical component. Hori¬

zontal area of fusion formed by random and

repealed bifurcations and lateral fusions, produc¬

ing a finely punctate shield, with small tubercles

among lacunae fomicd where various costal ele¬

ments have met and fused. Tubercles may occur

in central longitudinal scries in some zooids. Sec¬

ondary calcified orifice has raised fimbriated

distal plate and paired oral spines which are often

fiatlencd and curved. Proximal sinus formed by

recessed costae of suboral bar together with

paired condyles. Distal aviculariuin is small,

rounded rostrum less than half length of auto-

zooid orifice. Autozooids become surrounded by

kenozooids late in ontogeny.

Remarks. MacGillivray’s (1895) account and

figure did not adequately describe this species

which he noted was represented by a ‘single

minute fragment'. The type specimen consists of

approximately 15 worn zooids encrusting and

almost completely enclosing a minute shell frag¬

ment on both sides. Most of the zooid orifices and

the avicularia arc so worn that few characters

remain. The minutely tubcrculate frontal shields

are, however, better preser\'cd and can be seen to

be identical with those of the lectotype of Lageni-

pora morningtoniensis. MacGillivray's (1895)

figure did not show the distal aviculariuin which

he had not recognised as being present but did

illustrate the frontal shield and the distal plate ol

a single zooid. Not suiprisingly, Mapicstone

(1902) did not consider that his material had been

described before. He referred L morningtoniensis

to the ascophoran genus Lagenipora because he

considered that the finely punctate and tubcrcu¬

late pericyst was a ‘granulated' frontal shield and

that the distal plate and fiattened curved oral

spines which tend to surround the secondary

orifice constituted an ‘irregularly elevated peris¬

tome.' This stmeture he thought was ‘very similar

to L simplex. MacG.' Lagenipora simplex

MacGillivray (1890: 109, pi. 5 fig. 8) is a Recent

species from Western Port, Victoria, referable to

Celleporina bideniiculata Busk, 1881 (Busk,

1881, 1884; Bock, personal observation).

Anaskopora rotiuidata occurs frequently

together with A. elevata and A. cornuta from a

wide range of Victorian and South Australian

localities. It has not been reported since its

description and neither has L. morningtoniensis,

probably because the colonies have not

been recognised as distinct from those of

A. cornuta which they greatly resemble. The auto¬

zooids and avicularia are very similar in size and

shape but the structure of the zooidal frontal

shields of A. cornuta and A. rotundata are totally

dissimilar.

Anaskopora mesa sp. nov.

Figures 9-11, Table 1

Material examined. Ilolotype. NMV P140965,

Balcombc Bay, Victoria , Miocene.

Other material. I^alconibe Bay, Bairnsdale, Victoria;

Mount Schanck, South Australia.

Distribution. See above, Miocene (Appendix).

Etymology'. Mesa (Spanish) — a plateau at the

summit of a mountain, referring to the shape of

the autozooid pericyst.

Description. Colonies encrusting minute sub¬

strata. Autozooids with distinct vertical gymno-

cyst and peripheral component of 16-24 costae.

Median area of fusion wide, with crenulatcd

peripheral rim fomicd by spinous processes bor¬

dering single marginal series of pclmatidia. Hori¬

zontal component of costae not obvious, shield

formed by calcification surrounding 25*35 large,

often irregular lacunae, some with raised rim.

Secondary calcified orifice rounded, distal plate

distinct, sometimes with paired .spinous pro¬

cesses. Oral spines large, paired, occasionally

with additional distal spine and often raised and

inflexed above orifice and fused medially. Distal

avicularium small, length of rostrum less than that

of orifice. Rostrum subtriangular to rounded, not

raised, with paired condyles. Chambered pores

and kenozooids present at comers of autozooids,

small pore-chambers and seplular pores also

present.

Remarks. The material from Mount Schanck is

not well preserved and none of the autozooids

shows the raised, inflexed and fused oral spines

present in the Victorian specimens. These spines

arc remarkably similar to those which sometimes

occur in Corbulipora ornata (encrusting and

kenozooidal phases only, Bock and Cook, 2001).

One specimen from Balcombc Bay differs from

all the others. It comprises only a few, rather worn

zooids (Fig. 11). Although the frontal shield

resembles those of the other colonies, the sec¬

ondary calcified orifice differs in having little or

no raised distal plate but a large square tooth pro¬

jecting proximally into the orifice from the distal

184

P. E. BOCK AND P. L. COOK

wall. In a few zooids the suboral bar is raised

above the orifice, forming a partial peristome

similar to that described by Wass (1975) in mate¬

rial he assigned to^. ampulla. Until this and addi¬

tional specimens become available for examina¬

tion it is not possible to decide whether this

specimen represents a species distinct from A.

mesa sensu slriclo or is identical with A. ampulla.

Acknowledgements

We should like to thank Dr David Holloway for

access to the collections of Museum Victoria; and

Mr David McDonald for his help in preparation of

the paper.

References

Arnold, P. and Cook, P.L., 1997. Some Recent species

of the genus Anasknpora Wass (Bryozoa: Cribri-

oniorpha) from Queensland. Memoirs of the

Queensland Museum 42: 1-11.

Bock, P.E. and Cook, P.L., 1994. Occurrence of three

phases of growth with taxonomically distinct zooid

morphologies. Pp. 33-36 in Hayward, P.J.,

Ryland, J.S., and Taylor, P.D. (eds). Biology and

palaeobiology of bryozoans. Olsen and Olsen:

Fredensborg.

Bock, P.E. and Cook. P.L., 2001. A revision of the

multiphased genus Corbulipora MacGillivray

(Bryozoa: Cribrimorpha). Memoirs of Museum

Victoria 58: 191-213.

Brown, D.A.. 1958. Fossil cheilo.stomatous Bryozoa

from south-west Victoria. Memoirs of the Geo-

logical Society'of Victoria 10: 1-90.

Busk, G., 1881. Descriptive catalogue of the species of

Cellepora collected on the ‘Challenger’ Expedi¬

tion. Journal of the Linnean Society Zoology* 15:

341-356.

Busk, G., 1884. Report on the Polyzoa collected by

H. M.S. Challenger during the years 1873-76. Part

I. The Chcilostomata. Report on the Scientific

Results of the Voyage of H.M.S. Challenger,

Zoology 10 (30): xxiv + 216.

Gordon. D.P., 1984. The marine fauna of New Zealand:

Bryozoa: Gymnolaemata from the Kcrmadec

Ridge. New Zealand Oceanographic Institute

Memoir 9\: 1-198.

Gordon, D.P., 1989. The marine fauna of New Zealand:

Bryozoa: Gymnolaemata (Chcilostomida Asco-

phorina) from the western South Island continental

shelf and slope. New Zealand Oceanographic

Institute Memoir 97: I -158

Gordon, D.P., and Hastings, A.B., 1979. The inter-

zooidal communications of Hippothoa sensu lato

(Bryozoa) and their value in classification. Journal

of Natural History 13 (5): 561-579.

MacGillivray, P.H., 1890. Descriptions of new or little

known Polyzoa. Part 13. Proceedings of the Royal

Society of Victoria n.s. 2: 106-110.

MacGillivray. P.ll., 1895. A Monograph of the Tertiary

Polyzoa of Victoria. Transactions of the Royal

Society of Victoria n.s. 4: 1-166.

Maplestone. C.M.. 1901. Further Descriptions of the

Tertiary Polyzoa of Victoria. Pan 6. Proceedings

of the Royal Society of Victoria n.s. 13: 204-213.

Maplestone. C.M., 1902. Further descriptions of the

Tertiary Polyzoa of Victoria. Part 8. Proceedings

of the Royal Society of Victoria n.s. 15; 17-27.

Maplestone, C.M., 1904. Tabulated list of the fossil

cheilostomatous f*olyzoa in the Victorian Tertiary

deposits. Proceedings of the Royal Society of

Victona 17: 182-219.

Powell. N.A., 1967. Polyzoa (Bryozoa) — Ascophora

— from north New Zealand. Discovery Reports 34:

199-393.

Silen. L„ 1941. Chcilostomata Anasca (Bryozoa) col¬

lected by Prof. Dr. Sixien Bock's expedition to

Japan and the Bonin Islands 1914. Arkiv Jor

Zoo/og/33A(12): 1-130.

Wass, R.E., 1975. A revision of the bryozoan genus

Corbulipora MacGillivray. Proceedings of the

Royal Society’ of Victoria 87: 167-184.

Appendix. Details of Tertiary localities

mentioned in the text

Baimsdale (Skinners): Mitchell River bank, about

12 km W of Baimsdale, Vic. 37°47.9'S,

147°29.5'E.

Balcombe Bay; Also known as Fossil Beach, Morning-

ton, Mount Martha and possibly ‘Schnapper Poin’

(MacGillivray); on coast of Port Phillip Bay, about

3 km S of Momington. Vic. 38°!4.5'S,

145°01.7'E. Fyansford Clay. Balcombian; Middle

Miocene, (Langhian).

Balesford Quarry: Upper levels of limestone Quarry,

7 km W of Geelong. Vic. 38°06.5'S, I44°17.3'E.

Fyansford Clay. Middle Miocene (Balcombian).

Bird Rock; Coastal section, about 3 km W of Torquay,

Vic. 38°21.4'S, 144®17.8'E. Jan Juc Fomialion.

Janjukian; Late Oligocene, (Chattian). Also known

as Spring Creek.

Cariipbclls Point: On Lake Connewarre, 12.5 km SE of

Geelong. Vic. 38°13.7'S, 144°26.7'E. Middle

Miocene.

Cape Otway: Also Point Flinders, Locality AWl.

Coastal section 2 km NW of Cape Otway, Vic.

38°51.rS, 143°29.5'E. Glen Aire Clay. Early

Oligocene (Rupelian/Latorfian).

Cooriemungle area: Road cuttings about 18 km N of

Princetown. 38“32.4'S, 143°08.1'E. Gellibrand

Marl. Balcombian.

Corio Bay; Coastal exposures between 2 and 6 km N of

Geelong. Vic. Middle to Late Miocene.

Fyansford: This locality was mentioned by Maplestone

(1904) and is almost certainly in the same area as

the Batesford Quarry (see above).

Gellibrand: This locality was mentioned by

MacGillivray (1895) and is in the same area as

Princetown.

TERTIARY SPECIES OF THE BRYOZOAN ANASKOPORA

185

Glenaulin: Locality XVI of Brown (1958). Limestone,

South landslip. Allotment 1. Parish of Glenaulin,

south-western Vic. Janjukian, Late Oligocene

(Chattian).

Gleneig Group: Locality XI of Brown (1958). Lime¬

stone Creek clifl'. Allotment 16, and Locality XIV,

near base of section at Caldwells Clilf, Allotment

68b, both Parish of Werrikoo, south-western Vic.

Myaring Beds, upper Gambier Limestone. Long-

fordian. Early Miocene (Burdigalian).

Griffins; This locality is listed in Maplestone’s check¬

list (1904). The locality was one of the limestone

quarries on the Moorabool River, about 8 km NW

of Gee. This is close to the Batesford Quarry

(above).

Lake Bullenmerri: One of the rare inland outcrops of

the uppermost Gellibrand Marl. 38°16.9 S,

143°06.2'E. Middle Miocene.

Mount Gambier: Abandoned quarry on road to Port

MacDonnell, about 7 km S of Mount Gambier, SA.

37°53.5'S. 140°43.2'E. Gambier Limestone.

LongfordianC?) Early Miocene.

Mount Schanck: Limestone quarry about 1 km W of

Mount Schanck, about 15 km S of Mount Gambier,

SA. 37°57'S, 140°43.2'E. Gambier Limestone.

Longfordian; Early Miocene.

Muddy Creek: Clifton Bank, Muddy Creek, 8 km W of

Hamilton, Vic. 37°44.6'S, I41°56.4'E, Muddy

Creek Marl (= Gellibrand Marl). Balcombian.

Princetown (Gigantocypraea locality): Coastal section,

about 2 km W of Princetown, Vic. 38°41.9'S,

143°08.3'E. Gellibrand Marl. Balcombian.

Shelford (Red BlulT): Road cutting 35 km WNW from

Geelong, Vic., 38°3.rs. I43°59.2'E. Middle

Miocene.

Warrambine Creek: Stream section beside the bridge on

the Inverleigh-Winchelsea Road, 31 km W of Gee¬

long, Vic. 38°7.0'S, 144°00.4'E. Middle to Late

Miocene.

186

P. E. BOCK AND P. L. COOK

Figures I^. Anaskopora elevata. Balcombc Bay, Vic. Fig. 1, globular colony, showing frontal buds (x4()). Fig. 2.

enlargement of zooids from the same colony. Note distal avicularia and large area of extrazooidal calcification

surrounding zooids (arrowed) (x80). Figures 3-4. Amtskopora simplex sp. nov. Balcombe Bay, Vic. Fig. 3, part

of a small encnisling colony (x23). Fig. 4. enlargement of zooids from the same colony, distal avicularium

arrowed (x50).

TERTIARY SPECIES OF THE BRYOZOAN ANASKOPORA

187

Figures 5-8. Anaskopora cornula. Balcombc Bay, Vic. Fig. 5, slightly worn zooids, showing regular rows of inter¬

costal lacunae (x52). Fig. 6, enlarged zooids (x70). Fig. 7, frontally budded zooids and avicularia. Interzooidal keno-

zooids and developing frontal buds arrowed (x60). Fig. 8. Anaskopora rotumiata. Balcombe Bay, Vic. Globular

colony with developing frontal bud (arrowed) {x65).

188

P. E. BOCK AND P. L. COOK

Figures 9-12. Anaskopora mesa sp. nov. Balcombe Bay, Vic. Fig. 9, encrusting colony; zooids with raised

medially fused oral spines (x37). Fig. 10, enlargement of zooids from the same colony, marginal pelmatidia

arrowed (xl28). Fig. 11, zooid of A. aff. mesa showing distal oral tooth (arrowed) (x72). Figure 12. Anaskopora

rotundata Balcombe Bay, Vic. Zooids from growing edge of encrusting colony, showing lateral septular pores and

developing chambered pore (arrowed) (xl25).

TERTIARY SPECIES OF THE BRYOZOAN ANASKOPORA

189

Figures 13-16. Anaskopora rotundata. Balcornbc Bay, Vic. Fig. 13, colony encrusting basal side of erect bryozoan

(x48). Fig. 14. Secondary calcified orifice, showing condyles and fimbriated distal plate (arrowed) (x360). Fig. 15,

zooids showing tuberculate frontal shield and chambered pores developing as kenozooids at ‘comers’ of zooids

(arrowed) (xl27). Fig. 16, orifice, showing distal plate and oral spines (x240).

Memoirs of Museum Victoria 58(2): 191-213 (2001)

REVISION OF THE MULTIPHASED GENUS CORBULIPORA MACGILLIVRAY

(BRYOZOA: CRIBRIMORPHA)

P.E. Bock' and P.L. Cook^

'School of Ecology and Environment, Deakin University, 661 Blackburn Road, Clayton, Vic. 3168, Australia

(pbock(^deakin.edu.au)

-Honorary Associate, Museum Victoria, GPO Box 666E, Melbourne, Vic. 3001, Australia

Abstract

Bock. P.E. and Cook. P.L., 2001. Revision of the multiphased genus Corbulipora

MacGillivTay (Bi^ozoa: Cribrimorpha). Memoirs of Museum Victoria 58(2): 191-213.

Corbulipora MacGillivray is redefined to include only species which occur in successive

growth phases. Tlie fossil type species, Corbulipora ornata MacGillivray, occurs in an

encrusting ancc.stailale phase, an erect quadriserial, ovicellatc phase, and a frontally-budded

partially kenozooidal phase. The enenisting anccstrulate phase of the Recent species, C. tubu-

lifera (Hincks), is the type species of the genus Acauthocella Canu and Bassler, which is a

junior synonym of Corbulipora. The succeeding, ovicellaie, fluslrinc phase, known as IVater-

sia miliutris (Waters), is the type species of IVatersia, another junior synonym of Corbulipora.

It produces a third bilaminar phase known as C oriparma, a synonym of C. tubuUfera. This

has rhizoids and develops further lliistrine phases. Fossil specimens assigned to Acanibocella

tubulifera in the past are here considered to be the primary encrusting phase of a bilaminar

phase, known as Corbulipora suggereus (Waters), from which it has become separated. A

thinly calcified intervening erect phase similar to the flustrinc phase of C. tubulifera is inferred

to have existed but not to have been preserved as a fossil. Some species previously referred to

IVaiersia are assigned to Klugeflustra Moyano which, like Neqflustra Lopez Gappa, has flus-

trine colonies with large, hyperstomial ovicells, unlike those of the family Flustridae sensu

stricto. A key to species of Corbulipora and their various phases is given.

Introduction

The superfamily Cribrilinoidea Hincks, as rede¬

fined and interpreted by Gordon (1984: 60),

includes not only all the genera usually assigned

to the family Cribrilinidac Hincks (for example,

Bassler, 1953: G184), but all those referred to the

Cribrimorpha by Lang (1916). Many of the

Cretaceous genera described by Lang were

included in the family Pelmatoporidae by

Larwood (1962: 51). Voigt (1993) also assigned

Cretaceous species to this family.

‘Cribrimorphs’ are characterized by the pres¬

ence of a frontal shield of hollow calcified spines

which overarch a membranous frontal wall with

contiguous, uncalcified primary orifice and oper¬

culum. Obviously, many ‘anascaif genera share

these characteristics and the resulting shifts in

systematic position of various species of CctUo-

pora and Memhraniporella (Ryland and Hay¬

ward, 1977), emphasise Levinsen’s (1909: 156)

opinion of the ‘unnaturalness’ of the family Crib-

rilinidae. Gordon (1984: 60-61) noted the simi¬

larities among some species of Calloporidae with

those of Memhraniporella and Figularia and

remarked ‘at present it is difficult to define

positively the boundaries between the two

families.’ Moyano (1984, 1991) suggested some

preliminary groupings of cribrimorphs based on

frontal shield structures. These include some with

umbonulomorph affinities but do not deal with

some intennediate forms. It remains difficult to

define any polythelic boundaries among super¬

family and family groups, which arc themselves

almost certainly non-homogcncous.

However difficult it may be to define cribri¬

morphs exclusively, many species with distinc¬

tive characteristics have been described. Detailed

examination of some apparently ubiquitous taxa

has recently yielded much more infonnation on

oviccll ontogeny and on kinds and degrees of

polymorphs, which have revealed many exclu¬

sive groups of character correlations, and have

greatly increased the number of described species

(Ristedt. 1985; Bishop and Househam, 1987;

Hamiclin and Aristegui, 1988).

Briefly, the frontal shield (pcricyst) is formed

by spinous outgrowths (costae) which are hollow

outfoldings of the calcifying marginal frontal

191

192

P. E. BOCK AND P. L. COOK

wall (gymnocyst). The costae extend over the

flexible part of the frontal wall (frontal mem¬

brane) and fuse centrally (median area of fusion,

Larwood, 1962). The central coelomic space of

each costa is in contact, at its origin, with the peri¬

gastric (or visceral) coelom of the zooid (Wass,

1975). Small uncalcified areas regularly spaced

along the lateral vertical faces of each costa have

the potential to expand their overlying cuticle

intussusceptively and to calcify and form fusions

with those of neighbouring costae leaving holes

between successive costae (lacunae). Other small

uncalcified areas, equally regularly distributed

along the upper (frontal) surface of each costa

(lumen pores or pelmatidia), allow similar

expansions of cuticle and underlying coelom

frontally. These may also calcify fomiing

spinous processes. Sometimes the most marginal

pelmatidia, which arc usually the largest, extend

as spinous processes which meet those of

neighbouring zooids, fonning inlerzooidal

fusions quite late in ontogeny. This may produce

secondary calcification and even tertiary inlcr-

zooidal shields or cxtrazooidal expanses (Wass,

1975). (n some species frontally budded zooids

arise from expansions originating from enlarged

pelmatidia, which then function very like the

frontal septular pores of many ascophorans

allowing transfer of coelom and nutrients to the

developing buds,

Interzooidal communication through both sep-

tular pores, pore-chambers and a special kind of

pore-chamber (here called a chambered pore) all

occur in species of Corbiilipora. The chambered

pores allow another form of frontal budding.

They originate as a fairly simple uniporous or

multiporous pore-plate at the base of a vertical

wall. This rapidly develops a large calcified

chamber with a window covered by cuticle on the

upper and outer side. These chambers may

encompass more than one septular pore. Some

zooids of the same astogenetic generation may

communicate directly through septular pores,

others through septular pores on both sides of a

chambered pore. This often expands frontally

between the vertical zooid walls, forming a keno-

zooid with an uncalcified window at its frontal

side. These kenozooids tend to occur at the

‘comers' between autozooids and arc very evi¬

dent at the growing edges of colonics; they arc

also the origin of rhizoids and ‘stalk’ kenozooids.

Somewhat similar kinds of pore-chambers have

been described in the family Hippothoidac by

Gordon and Hastings (1979) but these are much

more regular in size, shape and distribution than

the chambered pores. It is interesting, however,

that Gordon (1989) noted that an ability to

produce frontal buds from the windows of such

pore-chambers was a definitive character of his

Infraorder Hippothoomorpha and he considered

that there was a ‘kind of conceptual linkage, from

the Cribrilinoidea through Catcnicelloidea to

Hippolhooidea'.

The potential for considerable astogenetic and

ontogenetic change within a colony is thus pre¬

sent in cribrimorph morphology. Although

almost every possible resultant structure seems to

be produced in Corbidipora^ in general, this

potential does not seem to be realized among

other cribrimorph genera. Pelmatidia are not

always present, or are little-developed, and inter¬

zooidal frontal buds arc rare. Species may

thus be defined by such characters as the

proportion of the gymnocyst to the pcricyst, the

number and form of costae, and the extent and

nature of the median area of fusion. Patterns of

kenozooids and ovicclls, together with avicularia,

also seem to define many taxa in detail

and to have character consistency (Ristedt, 1985;

Bishop and Househam, 1987). The species of

Corbiilipora described below pose problems

precisely because they display a wide range of

ontogenetic changes, coupled with the occur¬

rence of multiple astogenetic phases in

which the contributing zooids are often com¬

pletely unlike one another; in one case being

totally ‘un-cribrimorph' in appearance (Bock

and Cook, 1994).

Colonies in which the zooids exhibit zones or

areas of totally differing morphologies are not

unknown in other Bryozoa. Frequently isolated

zones or areas have been assigned to completely

unrelated taxa and Voigt (1975) has illustrated

several examples drawn principally from the Late

Cretaceous of Northern Europe. Among these,

one cribrimorph, Castanopora bipunctata (Gold-

fuss), shows remarkable autozooidal helcro-

morphy. Colonies are bilaminar, unilaminar, or

occasionally encrusting. In all these colony fonns,

entire fragments or large areas within colonies

have membraniporinc zooids with no trace of a

calcified costal shield frontally. In other speci¬

mens, all or most of the zooids have a shield com¬

posed of 10-18 costae with pelmatidia, which

alternate with rows of small lacunae. Both types

ofautozooid bear identical ovicclls, and like some

phases of Corbiilipora tuhidifera. identical kinds

of large avicularia occur in both kinds of colony.

Castanopora bipunctata does not, however, seem

to occur in isolated subcolonies, each repre-

THE MULTIPHASED BRYOZOAN GENUS CORBULIPORA

193

senting a distinct phase, and thus differs from

Corhulipora.

Specimens from the collection of Dr Shizuo

Mawalari, and from the following institutions

have been examined; Australian Museum,

Sydney (AM), Natural History Museum, London

(BMNH), Museum of Victoria, Melbourne

(NMV) and the Zoological Museum, Copenhagen

(ZMC).

Corhulipora MacGillivray

Corhulipora MacGillivray, 1895: 60.—Wass 1975:

168.

Watersia Levinsen, 1909: 94.

Acaiuhocella Canu and Bassler, 1917: 35.

Type species. Corhulipora ornata MacGillivray,

1895.

Description. Complete colonies consisting of sub¬

colonies with encrusting, erect and/or semi-erect

phases of growth. Each phase with zooids of

consistent morphologies, which may differ

completely from those of a preceding or suc¬

ceeding phase. Each species including at least 1

cribrimorph phase, usually more. Growth

sequences always beginning with an encrusting

anccstrulate phase but other sequences may alter¬

nate or co-exist, although sometimes in physical

isolation. All phases are known, or assumed, to be

capable of a separate existence. Ancestrula tati-

fonn, with marginal spines. Zooids of all cribri¬

morph phases with gymnocyst and pericyst of

costae with pelmatidia and lacunae. Oral spines

1-3. Ovicells often confined to I phase only but

not necessarily the same phase in different

species. Ovicells are prominent, hyperstomial,

closed by operculum, with median suture and

paired entooecial frontal areas. Brooding zooids

of tlustrine phase with flattened spines or with

exposed frontal membrane and enlarged oral

spines only. Avicularia inlerzooidal, large, with

distally orientated rostrum, rounded or expanded

terminally.

Remarks. Multiphased growth is known or may

be reliably inferred to occur in each of the

three species here referred to Corhulipora. In

C ornata, all zooids of the three phases recog¬

nized have costate pericysts. The first phase is

encrusting, the second erect and quadriserial, and

this phase develops both avicularia and ovicells.

The third phase is frontally budded from the pel¬

matidia of the second erect phase zooids and is

often purely kcnozooidal. In C suggerens, two

cribrimorph phases have been recognized and

inferred to belong to one taxon. One phase is

encrusting, the other is bilaminar. Its colony and

zooid morphologies arc so akin to those of the

third, bilaminar phase of C. tuhulifera, that it

seems almost certain that it represents a third

phase, and that a missing, intervening, erect

second phase, with very thin calcification, and

perhaps flustrine or ccllularine growth, once

occurred, but was not preseiwed in the fossil

material. The bilaminar phase of C, suggerens

occasionally bears ovicells, unlike that of

C. tuhiilifera. This last species has been observed

with its three phases growing from one another.

The anceslnilatc encrusting phase gives rise to

several fiustrinc subcolonics which alone bear

ovicells. Large avicularia occur in this phase and

in the third phase, which develops from the

tips of the fiustrine fronds. These two phases may

alternate (Bock and Cook, 1994). Recently a fur¬

ther living species of Corhulipora, C. inopinata,

has been described from southwestern Australia

by Bock and Cook (1998). It closely resembles

C. tuhiilifera and also occurs in three sub¬

colony phases. The flustrine, second phase

differs in having zooids with frontal spines, some¬

what resembling those of the earlier ontogenetic

stages of the second, quadriserial phase of

C. ornata.

The genus Corhulipora was introduced for

C. ornata alone. Wass (1975) redefined the genus

and assigned some species to a new subgenus,

Anaskopora. These species, together with Crih-

rilina cornuta MacGillivray (1895), which Wass

(1975) referred to Corhulipora, are now regarded

as gencrically distinct and are discussed else¬

where (Bock and Cook, 2001).

Corhulipora collaris Canu and Bassler (1920:

308, pi. 43 fig. 14), trom the North American

Eocene was also redescribed by Wass (1975: 170,

pi. 8, figs 7-9). It has medially orientated

adventitious oral avicularia and an cndozooidal

ovicell. It is therefore excluded here from

Corhulipora which is considered to have a Ter¬

tiary to Recent, Australasian distribution only.

When Wass (1975) revised Corhulipora,

C. collaris was the only species assigned to the

genus in which an ovicell had been described.

This was probably the reason why he used the

temi ‘entozooidaP for the ovicell type, which he

then applied to the whole genus. All ovicells now

observed in species referred to Corhulipora are

hyperstomial, with paired, entooecial frontal

areas.

194

P. E. BOCK AND P. L. COOK

Key to species and phases of CorhuUpora

I. Subcolony tlustrine, aulozooids with exposed frontal membrane and 1 pair

of oral spines. Ovicells and large interzooidal avicularia present

.C tuhulifera phase 2

— Subcolony not flustrine, autozooids with costal frontal shield.2

2. Subcolony encrusting a substratum, mostly single layered.3

— Subcolony erect, suberect and bilaminar or formed by frontal budding.5

3. Autozooids large (Lz >0.75 mm), costae 16-24, with 3-4 concentric series

of long, sometimes branched, spinous processes arising from pelmatidia

.C. sug}^erefus phase 1

— Autozooids smaller, costae 10-18, with 1-2 concentric series of blunt tuber¬

cles arising from pelmatidia.4

4. Autozooids very small (Lz <0.50 mm), orifice small (lo 0.10 mm). Lateral

oral spines raised, inflexed and fused above the orifice.C oniata phase 1

— Autozooids larger (Lz >0.60 mm), orifice larger (lo >0.11 mm). (3ral spines

not raised or fused.C. tubulifera phase 1

5. Subcolony erect, quadriserial; autozooids with long gymnocysl and small

costal shield. Avicularia rare, ovicells present.C. ornata phase 2

— Subcolony composed of autozooids and kenozooids, budded frontally from

phase 2-zooids, orientation random. Gymnocyst and orifice not generally

visible; avicularia and ovicells absent.C. ornata phase 3

— Subcolony bilaminar, small: autozooids orientated distally, interzooidal

avicularia present.6

6. Subcolony triangular, arising from 2-4 zooids with partially calcified frontal

shields, orifices not occluded by oral spines. Avicularia not raised distally,

with a bar. Ovicells present.C. su^gerens phase 3

— Subcolony rectangular, arising from 6-8 zooids with partially calcified

frontal shields, orifices often occluded by growth and fusion of oral spines.

Avicularia raised distally, with paired condyles. Ovicells absent, rhizoids

present.C. tubulifera phase 3

CorhuUpora ornata MacGillivray

Figures 1-9, 27, Tables 1, 2

CorhuUpora ornata MacGillivray, 1895: 60, pi. 8

figs 20, 21.—Maplestone, 1904: 201 (listed).—Brown,

1958: 54.—Wass, 1975: 168. pi. 7 figs 1, 2.

Material examined. Lcctotype (chosen by Was.s, 1975):

NMV P27642, Schnapper Point (= Balcombe Bay).

Victoria. Middle Miocene. Balcombian.

Other material. Balcombe Bay; Cooriemunglc, near

Princelown and Princetown, Victoria (Middle Miocene,

Balcombian); Bird Rock, near Torquay. Victoria

(Upper Oligoccne. Janjukian); Mount Schanck, S of

Mount Gambler. South Australia (Lower Miocene.

Longfordian).

Distribution. Additional localities given by

MacGillivray (1895) and Maplestone (1904)

include: Cape Otway, Spring Creek, Campbells

Point, Momington, Mitchell R. and Baimsdale;

localities given by Brown (1958) include;

Localities XVI (Glenaulin Clay) and XXVI

(Watacpoolan Limestone) near Crawford R.: and

XXII and XXIII on Glenelg R., Nangeela, south¬

western Victoria. Miocene to Oligoccne (see

Appendix).

Description. CorhuUpora with subcolonies of 3

growth phases; all zooids with costate pericysts.

Subcolonies of encrusting phase very small, com¬

prising at most 40 zooids; basal walls calcified

only at margins. Anccstnila has 14-16 marginal

spines and buds a triad of distal primary zooids.

As in C. suggerens. appearance of some zooids

differs from those budded later in astogeny. Auto¬

zooids have small, distinct gymnocyst, and peri-

cyst with 10-16 costae, each with 2 or 3 interven¬

ing lacunae and 2 pelmatidia, raised as blunt

tubercles. Secondary calcified orifice small, with

paired condyles and 3 oral spines, lateral pair

often raised vertically, inflexed and fused above

orifice (Figs 1, 2). This condition also occurs in

some zooids of third phase, which may resemble

those of first phase closely. Zooids communicate

through small pore-chambers at base of vertical

walls.

Second phase erect and quadriserial, zooids

THE MULTIPHASED BRYOZOAN GENUS CORBULIPORA

195

Table 1. Comparative measurements in mm among phases in species of Corbtdipora, Lz, Iz, length and

width of zooid; lo width of orifice.

C. ornata

C suggerens

C tubulifera

Phase 1

Lz 0.45-0.48

Lz 0.58-0.87

Lz 0.58-0.74

lz 0.37-0.40

lz 0.37-0.54

lz 0.38-0.50

lo 0.08-0.09

lo 0.11-0.13

lo 0.12-0.13

Phase 2

Lz 0.70-1.00

—

Lz 0.78-1.32

lz 0.28-0.36

—

lz 0.25-0.33

lo 0.13-0.14

—

Phase 3

Lz 0.33-0.37

Lz 0.58-0.87

Lz 0.54-0.87

lz 0.33-0.37

lz 0.37-0.54

lz 0.32-0.54

lo 0.07-0.10

lo 0.06-0.12

lo 0.10-0.14

Table 2. Comparative measurements in mm among phases of Corhulipora and species of Klugeflustra.

Lz, lz, length of zooid; Lav, lav, length and width of avicularium; Lp, length of palate; Lov, lov, length

and width of ovicell.

Lz lz Lav lav Lp Lov lov

C omi/a phase 2 0.70-1.00 0.28-0.36 0.55-0.66 0.17-0.21 0.43-0.55 0.24-0.25 0.23-0.25

C .si/ggermv phase 3 0.58-0.87 0.37-0.54 0.28-0.43 0.12-0.16 0.12-0.20 0.12-0.16 0.16-0.21

C./nWi/tra phase 2 0.78-1.32 0.25-0.33 0.65-0.96 0.22-0.26 0.31-0.35 0.23-0.27 0.28-0.33

K. vanhoejfeni 1.23-1.56 0.40-0.50 1.03-1.25 0.25-0.33 0.40-0.50 0.50-0.60 0.44-0.48

K. kishakaensis 1.32-1.43 0.32-0.36 0.25-0.37 0.32-0.31 0.08-0.10 0.33-0.43 0.40-0.47

K. simplex _0.44-0.50 0.15-0.19 —_—_— 0.21-0.28 0.28-0.31

communicating through simple septular pores.

Branches bifurcate occasionally. Intennediate

developmental stages linking encrusting with

quadriscrial phase unknown, and in isolation,

component zooids of the 2 phases totally unlike.

Erect-phase autozooids have elongated, smooth

gymnocyst surrounding costal shield which may

be subrcctangular or small and oval. Zooid length

nearly twice that of those from other phases,

12-22 costae, distal pair forming suboral bar,

prominent and sloping forward on distal edge

(Fig. 4). Secondary calcified orifice large, almost

perpendicular to frontal shield, with 1 pair of

small lateral spines. Costae expanded and fused

terminally; variable median area of fusion which

changes its aspect with ontogeny. Costae bearing

2 or 3 small pcimalidia which alternate with 2-4

lacunae. Ovicclls generally rare, narrow, with

paired, crescentic, entooecial frontal areas. Oral

spines of brooding zooids greatly enlarged, raised

and fused above orifice, like those of phase 1

zooids; .spines also bear some pelrnatidia (Fig.

27). Avicularia also rare; large, with distinct gym¬

nocyst, in series with autozooids. Rostrum

rounded or expanded terminally, and orientated

distally; small paired condyles (Fig. 3).

Subcolonies of third phase arise by frontal bud¬

ding directly from autozooids of second phase.

Zooid buds develop from greatly expanded pcl-

matidia; kenozooids and autozooids produced;

ovicells and avicularia absent. Frontally budded

zooids have random orientations; frontal shields

have 10-16 costae, each with 2 or 3 pelrnatidia

raised into long spinous processes. These inter-

digitate and sometimes fuse interzooidally, espe¬

cially at margins of zooids, forming continuous

secondary shield in some specimens. Secondaiy,

calcified orifice very small, with 3 oral spines.

These may be long and prominent, occasionally

raised, infiexed and fused above orifice, exactly

196

P. E. BOCK AND P. L. COOK

as in encrusting zooids of first phase, not occlud¬

ing orifices (compare C tuhuUfera), but many

zooids have no obvious orifice. Some may have

been occluded by general interzooidal fusion of

spinous processes but others seem to have been

budded with no orifice, and are therefore true

kenozooids.

Remarks. The taxonomic identity of the first and

second phases of C ornata can only be inferred,

since zooids of intermediate morphologies have

not been found linking them in situ. The ontoge¬

netic changes in second-phase zooids leading to

the third phase can, however, be traced in many

specimens from Balcombe Bay and Mount

Schanck. Early in ontogeny, the pcricysts of the

erect second-phase zooids form a small, oval area

distal to an elongated gymnocyst (Fig. 3). The

costae are long and virtually without lateral

fusions, so that the intervening, slit-like lacunae

extend to a narrow median area of fusion. Onto¬

genetic changes begin with a widening of this

area by the development of lateral co.stal fusions,

and a thickening of the shield by expansion of the

pelmatidia and growth of spinous processes

frontally (Fig. 4. 5). Some pelmatidia continue to

increase in size, particularly those on the thick¬

ened suboral bar, and they become blister-like,

while the more lateral spinous processes fuse

interzooidally. The third-phase buds arise directly

from expanded pelmatidia and some appear to

develop as chambered pores or small interzooidal

kenozooids (Figs 6. 7). The larger kenozooids and

some autozooids have costate pcricysts with

numerous, long spinous processes arising from

their pelmatidia. The third phase forms small,

globular masses, budding from and overgrowing

the second-phase zooids especially at the bifur¬

cations of erect branches. In some specimens sec¬

ondary calcification results in the third phase

appearing as a globular mass of spinous or ridged

costae with no orifices and no distinct zooidal

structure remaining. The zooids of the second

phase remain visible protruding at the proximal

and distal ends of the mass (Figs 8, 9).

Corhidipora suggerens (Waters)

Figures 10-16, 28-29, Tables 1, 2

Crihrilina suggerens Waters 1881: 327, pi. 17 fig.

75.—MacGillivray, 1895: 57, pi. 8 fig. 8.—Maple-

stone, 1904: 201 (listed),

Cribhlimi tuhulifera Waters, 1883: 436 (not Hincks,

1881).

Acanthocella tubuUfera. —Brown, 1958: 54 (fossil

specimens only).

Cribrilina jonesi Brown 1958: 52, fig. 28.

Corbulipora sp.—Wass and Yoo, 1975: 811, pi. 2

fig. 4.

Corbulipora suggerens. —Bock and Cook, 1994: 34,

fig. 6.

Material examined. NMV P73147, Holotype of

C. jonesi, locality IX, K bank of Cilcnelg R., Myaring,

south-western Victoria (Middle Miocene, Balcombian).

Other material. Balcombe Bay (ovicells); Coorie-

mungle: Princetown (ovicells); Grices Creek; Baims>

dale (all Middle Miocene, Balcombian); Narrawaturk

Bore 2, near Peterborough, Victoria and Mount

Schanck, South Australia (both Lower Miocene,

Longfordiaii).

Distribution. Additional localities given by

MacGillivray (1895) and Maplcstonc (1904)

include: Campbells Point, Curdics Creek, Muddy

Creek, Mitchell R. and Schnapper Point; and by

Brown (1958) include Locality XIV, section at

West end of Caldwell’s Cliff, Glcnelg R.

(Miocene, Myaring beds) (.sec Appendix).

Description. Corbulipora with subcolonics of 2

known growth phases. Ancestrula tatifoim, with

10-12 marginal spines. Autozooids of first,

encrusting phase large, oval, with distinct rim of

smooth gymnocyst suirounding costal shield with

16-24 costae (Fig. 10). Secondary, calcified ori¬

fice smaller than that of C. tubulifera (compare

figs 16, 17), with 3 oral spines and paired

condyles. Costae with 3, occasionally 4 pelma¬

tidia and 3 or 4 lacunae. Median area of fusion

increases in width with asiogcny, and pelmatidia

become raised and tubcrculate, with long, some¬

times branched spinous processes, which inter-

digitate and occasionally fuse laterally. Zooids

with typical pore-chambers, with 3 distal and 1 or

2 lateral windows within gymnocyst boundary

(Fig. 10). Avicularia and ovicells absent.

Bilaminar phase fomiing small, subtriangular

subcolonies, each arising from 2-4 zooids which

have frontal shields uncalcificd or calcified only

at distal end (Fig. 12). Subcolonics expand by

bifurcation of zooid rows to 4 6 within two asto-

genciic generations. Basal lamina between the 2

zooid faces with few septular pores; autozooids

with 2 or 3 large chambered pores or kenozooids

laterally and distally (Figs 13 15). Autozooids of

first 2 generations of completely calcified zooids

often have elongated costal shields with little or

no gymnocyst visible, and 16-22 costae. Distal

pair of costae fonn prominent suboral bar sloping

forw'ard on distal edge, obscuring large, wide,

vertical secondary calcified orifice. Costae well

separated marginally with 2 or 3 inter\'cning

lacunae and 3 concentric rows of tubcrculate pcl-

matidia. Autozooids of next 2 or 3 generations

with increasingly short, rounded costal shields.

THE MULTIPHASED BRYOZOAN GENUS CORBULIPORA

197

with some marginal gymnocyst visible; iO-16

costae, distal pair fonning suboral bar defining a

minute, semicircular orifice, with paired condyles

and 3 oral spines, which is visible frontally (Fig.

16). Pclmalidia arranged in 3 raised concentric

scries, with spinous processes which may inter-

digitate and fuse laterally. Oral spines often long

but do not fuse above or occlude orifice. At grow¬

ing margin of subcolonies, large, blister-likc

chambered pores or kenozooids occur in adjacent

series between autozooids (Figs 13-15). Avi-

cularia only among zooids of proximal, earlier

astogcnetic generations of subcolonies; large and

interzooidal, with distally orientated, subspathu-

late rostrum and complete bar (Fig. 12). Oviccils

rare, only on those zooids on proximal margin of

phase 3 subcolonies, where zooids lack any calci¬

fied, costal frontal shield. Ovicells slightly flat¬

tened frontally, with paired entooecial areas,

which in a few cases show signs of costatc struc¬

ture (Figs 28, 29). Unlike brooding zooids of

C. ornata and C. tubiilifera, paired oral spines not

enlarged.

Remarks. The encrusting and bilaminar phases of

C. stiggcrefis resemble the equivalent first and

third phases of C. tuhulifera closely. The encrust¬

ing zooids are usually larger and in early astogeny

exhibit a dramatic change in appearance between

the generations like those of C. ornata at the same

asiogenelic stage. The vertical, marginal parts of

the costae arc well separated and the median area

of fusion is narrow. There are only two concentric

scries of pelmatidia with short, but distinct

spinous processes. In the next generations, zooids

have wider median areas of fusion and three or

more scries of spinous processes. In some speci¬

mens from Princctown, a large distal pore-

chamber fonns an irregular distal kenozooid with

a single pore in its frontal surface. The pore-

chambers do not nomially protrude beyond the

base of the gymnocyst, unlike those of

C. liibidifera. The zooids of the bilaminar phases

of the two species resemble one another closely,

but the size and shape of the subcolonics differ.

The narrow origin of those of C suggerens sug¬

gest that they may have been derived from the tips

of a quadri.scrial, or very narrow, thinly calcified

phase which has not been preserved. This in con¬

trast to the much wider colonies of C tiibulifera,

which are known to be derived from the broader,

flustrine, IVatersia-phasc fronds. The orifices of

the zooids of C suggerens arc not occluded by

oral spine development, and the avicularia also

differ from those of C. tubulifera in having a flat

rostrum and a complete bar. Finally, as mentioned

above, the bilaminar phase of C suggerens occa¬

sionally bears ovicells proximally, unlike that of

C. tubulifera.

Waters’s (1883) record of C. tubulifera from

Muddy Creek seems to have been based on spec¬

imens of the encrusting phase of C. suggerens,

which he described as growing on shell (see

above). He noted that the autozooids had larger

orifices than those of his specimens of bilaminar

C. suggerens where they arc extremely small.

Brown (1958) introduced Cribrilina jonesi for

another bilaminar species from the Victorian

Tertiary. The zooids had 16 costae and a thick

suboral bar, and he noted their resemblance to

Acanthocella tuhulifera, which he regarded as a

senior synonym of C. suggerens. Brown

remarked on the small size of the orifice. The

holotype of C. jonesi has been examined and

proves to be identical with the bilaminar phase of

C. suggerens.

Corhulipora tuhulifera (Hincks)

Figures 17-26, 30-32, Tables 1, 2

Cribrilina tubulifera Hincks 1881: 8, pi. 1 fig. 7.—

Water.s, 1887b: 187, pi. 5 figs 2, 6.

Acanthocella tuhulifera. —Brown, 1958: 52 (lecto-

type only, see C. suggerens).

Flustra miliiaris Waters 1887a: 93, pi. 14 fig. 2.

IVatersia militaris. —Leviasen, 1909: 94, 95 (not

Livingstone, 1929: 53; not Gordon, 1986: 28).

Corhulipora sp.—Wass, 1973; 361, pi. 1 figs 1^.

Corhulipora oriparma Wass 1975: 169, pi. 9 figs

1-7, pi. 10 figs 1-6, 9-11.—Cook, 1979; 271, fig.

ID.—Wass and Yoo. 1983; 323.

Corhulipora tubulifera.—Bock and Cook, 1994: 33,

figs 1-5.

Material Examined. BMNH 1899.5.1.726, photograph

of lectotype of C tubulifera, Bass Strait. BMNH

1887.10.21.1, ?part of type material of W. militaris.

Port Jackson, Waters coH. BMNH 1926.6.1.2, Port

Jack.son, 9-15 m. label in Waters’s hand ‘sandy mud

bottom near the Heads'. BMNH 1897.5.1.447, Port

Jackson. AM U5006, ‘Endeavour’ Exped.,

T.Mortensen, S of Lakes Entrance. Victoria,

38"15'-25'S, 148°2()' 43'E, 128-220 m, 8-9 Sep

1914, sand. NMV Bass Strait Sur\'cy: Kimhla stn 99

40'^06'S, 143®I6'E, II Oct 1980, 139 m; stn 100, as

above, 158 m; stn 101, as above, 187 ni. Sarda sin 112,

40^22'S, 145°127'E, 3 Nov 1981,40 m. Hai Kung stn

135,40°50 S, 146^31.3 E,4 Feb 1981,68 m. Tangaroa

stn 155, 38°34'S. 144°54.3'E, 12 Nov 1981, 70 m; as

above, 75 m; stn 164, 40°40.7'S, 148^36.9'E, 14 Nov

1981, 67 m. SLOPE stn 22, 37°0.6'S, I50°20'E 21 Jul

1986,363 m; stn45,42‘'2.2'S, 148M8.7'E, 27 Jul 1986,

800 m; stn 46, as above, 720 m; stn 47, 41‘^59'S,

148°38.8'E, 27 Jul 1986, 500 m. ZMC, Port Jackson,

NSW (label in Waters’ hand).

198

P. E. BOCK AND P. L. COOK

Description. Corbulipora with subcolonics of 3

growth phases, 2 of which may be sequential as

well as alternating. Observed sequences include

encrusting (phase 1) to tlustrine (phase 2);

flustrinc to bilaminar (phase 3): and bilaminar to

flustrine (Fig. 18).

Phase 1. Subcolony encrusting small fragments

of shell, dead bryozoans etc. Ancestrula tatiform

with 10-14 fine marginal spines (La 0.45 mm)

(Figs 23). Subcolony forming small patch of

50-100 zooids, basal walls calcified only

marginally. Autozooids with marginal gymnocysl

and pericy.st of 10-16 stout costae. Vertical

marginal part of costae well separated, inllexcd

and abutting centrally to form wide median area

of fusion, alternating with short rows of 2 or 3

lacunae. Pelmatidia in 2 concentric circles, raised,

forming blunt tubercles, occasionally spinous.

Suboral bar swollen, with 3 or 4 pelmatidia.

Secondary, calcified orifice semicircular with pair

of small condyles and 3 oral spines. Operculum

golden brown, closely apposed to the calcified

orifice. I distal and 2 or 3 large lateral chambered

pores, each enclosing several small septular

pores. Each marginal chamber capable of giving

rise to 1 rhizoid or stalk kenozooid. Avicularia

and ovicells absent.

Phase 2. Subcolony arising from long stalk of

thinly calcified kenozooids derived from cham¬

bered pores at margins of phase I or phase 3 sub¬

colonies. Stalk kenozooids in apposed pairs or

triads, and bud elongated autozooids by the

second to fifth astogcnetic generation, followed

by bifurcation of zooid rows to form bilaminar

flustrine frond (Fig. 18b). Fronds 6-8 rows wide,

increasing to 10-12 before simple bifurcation.

Largest subcolonics extend for >90 astogcnetic

generations, bifurcate 4 times, and include over

2000 zooids. Marginal zooids elongated, but not

kenozooids. Vertical walls of opposing laminae

do not correspond, and simple septular pores

communicate through basal walls. Zooids also

have 1 distal, and 1 or 2 lateral multiporous sep¬

tular pores. Autozooids very thinly calcified, with

no gym nocyst or cryptocyst; frontal membrane

continuous with simple, terminal crescentic oper¬

culum. Oral spines paired, tcmiinal, arising from

lateral walls, wide at base; short and frontally

directed in autozooids; massive, swollen and

curved over operculum in brooding zooids. Ovi-

cclls may be developed on all zooids except

marginal zooids; large, prominent, hyperstomial

and closed by operculum. Frontally ovicells

slightly flattened, with median suture and paired,

crescentic cntooecial areas (Figs 30, 31 ). Number

of tentacles 12-14 (Lt 0.75 mm, Lt sheath 0.20

mm). Avicularia scattered, usually proximal to, or

at a bifurcation of zooid rows. Gymnocyst thinly

calcified, elongated, shorter in those avicularia at

bifurcations; opesia uncalcified, rostrum raised

and thickly calcified dislally. Mandible sub-

spathulate, hinged on paired condyles, directed

distally (Fig. 3). Rhizoids arising from multi-

porous septular pores in walls of stalk keno¬

zooids, or from autozooids at proximal end of

fronds.

Phase 3. Subcolony bilaminar, fonning sub-

rectangular masses of 100-150 zooids, anchored

by numerous rhizoids originating from enlarged

marginal chambered pores. Subcolonies at tips of

flustrinc fronds, and proximal ends fonned by 2

apposed series of 6-8 partially calcified zooids,

alternating with next generation of zooids, which

have frontal shield calcified at di.stal end only

(Figs 25, 26). Following generation of 8-10

zooids elongated, with 16-18, often irregular

costae, and prominent suboral bar. Subsequent 4

or 5 astogcnetic generations have zooids with

increasingly shorter and rounder costal shields,

less prominent suboral bar. and smaller, semi¬

circular calcified orifice. Pelmatidia in 2 concen¬

tric series, rai.sed as blunt tubercles. 3 oral spines,

becoming very large, swollen and fused, occlud¬

ing orifice completely and developing large,

blister-like pelmatidia on frontal surface. Large

chambered pores at margins give ri.sc to stalk

kenozooids or rhizoids (Fig. 27, 28). Avicularia

infrequent, occurring in earlier astogcnetic gener¬

ations, morphology same as those occurring in

flustrinc fronds. Ovicells absent.

Remarks. The nomenclalural consequences of

the identity of Crihrilina tiibulifera. Watersia

militaris and Corbulipora oriparma are consider¬

able, and arc summarized in the synonymy given

above. Although Acanthocella was introduced

by Canu and Bassler (1917) with the Recent,

Australian, Crihrilina (ubulifera Flincks as type

species, it was apparently their intention to erect a

genus to receive the Eocene, Gulf of Mexico

species, A. erinacea (Canu and Bassler, 1917: 36,

pi. 4 fig. 1; 1920: 309, pL 43 figs 101-103).

Acanthocella erinacea has a frontal shield like

that of A. tubiilifera but has elongated ovicells

without cntooecial areas. It resembles the Recent,

amphiAtlantic Crihrilina Jigitlaris floridana

Smitt (1873), which was referred to Reginclla by

Chectham and Sandberg (19(>4), Cook (1985:

125, pi. 9F) and Winston (1982: 134, fig. 59), and

which does not develop ovicells. One further

species of Acanthocella., A. clypeata Canu and

Bassler (1928: 39, pi. 4 figs 9, 10, Fig. 5) from the

THE MULTIPHASED BRYOZOAN GENUS CORBULIPORA

199

Gulf of Mexico, has complex oral spines which

incorporate the suboral bar (Rislcdl, 1979).

No ovicells have been described. AcanthoceUa

ehnacea, R. floriclana and A. clypeata are not

known to occur in more than one phase and arc

excluded from Corhulipora here. AcanthoccUa

erinacea has been made the type species of

Castanoporina. a new subgenus of the Cretac¬

eous genus Castanopora by Voigt (1993). He

described a new species, together with one

originally introduced by Larwood (I9()2) as

Castanopora voigti. Castanoporina voigti was

illustrated by Voigt (1993: 148, pis 7, 8,9) having

both colonies and zooids with a strong resem¬

blance to those of erect, phase 2 Corhulipora

ornata. However, the zooids each have paired,

adventitious oral avicularia and the ovicells have

no cniooecial frontal areas.

Watersia Levinsen (1909: 99) was infonnally

introduced in a key to genera of Bicellariidac. In

addition. Watersia militaris was mentioned twice

(pp. 94, 95), although with no indication of author

or dale. However, Levinsen seems to have thus

indicated Flustra militaris Waters as type species,

as no other taxon was assigned to the genus

Watersia, which was not mentioned again.

Flustra militaris was originally described by

Waters (1887a) from Port Jackson, New South

Wales. 1 lis figure shows three apparently verticil-

late rows of brooding zooids. with membranous

frontals, stout, ‘club-shaped’ oral spines, and ovi-

celis with a median suture and paired frontal

areas. The colony was described as ‘bilaminate'

and Waters noted the similarities in appearance

of the ovicells with those of 'Flustra episcopalis'

Euthyroides).

Some specimens (ZMC, BMNH), four of

which have labels in Waters' hand, or are from his

collection, and ail of which are from Port Jackson,

have been examined. All the subcolonies were

dry and very shmnkcn laterally, none possessed

the earliest astogenetic stages of the flustrine

phase. None of the subcolonies had any avicu¬

laria, which are generally rare in flustrine sub¬

colonies with ovicells. Waters (1887a) did not

mention avicularia in his original description. The

verticillate appearance of the ovicclled specimens

illustrated by Waters (1887a) is explained by the

considerable lateral shrinkage which occurs when

ilustrine colonics are dried. The ovicells are

prominent, but the very thinly calcified proximal

parts of the zooids almost disappear from view.

The length of the zooids of the subcoionies from

New South Wales is consistently greater than I.O

mm; longer than the zooids of this phase in the

specimens from Bass Strait.

The structure of the ovicells resembles that of

species of Figidaria, as well as Euthyroides, and

the similarities have been discussed and illus¬

trated by Gordon (1989, pis 2, 30). It is interest¬

ing that the species illustrated as Figularia

mernae Gordon (1989: 15, pi. 2C, D) from New

Zealand, occurs with an encrusting and a flustrine

phase.

Corhulipora oriparma was described in detail

by Wass (1973, 1975). He particularly noted the

astogenetic and ontogenetic changes occurring in

the small, bilaminar subcolonies. He described

the increasingly early occlusion of the zooidal

orifices with aslogeny, and speculated on the

apparent impossibility of any tentacle protrusion

in these zooids. He also described the changes in

zooid shape with astogeny and spinous processes

arising from the pelmatidia.

Among the 94 specimens examined, 27 include

the transitional stages between phases. The com¬

monest transition is that from the bilaminar to the

flustrine phase (15 examples), with the transition

from the encrusting to the flustrine phase being

less frequent (nine examples). The transition from

the flustrine to the bilaminar phase occurs in only

three examples (Fig. 24). The total number of

phase 1 subcolonies is 27, the majority (18)

occurring from Tangaroa stn 155. The small sub¬

colonies all encrust minute shell fragments, and

some have the ancestrula visible. Some have a

few marginal rhizoids, and one has a flustrine

phase growing from stalk kenozooids. Stalk keno-

zooids developing from enlarged pores also occur

in subcolonics from Kimhla stns 99 and 100, and

Tangaroa stn 164. The kenozooids originate from

adjacent chambered pores and are closely

apposed in pairs or triads; their walls become

attached to each other by the first astogenetic gen¬

eration, and communicate with one another

through septular pores. Zooids of the following

generations arc just as elongated, but have traces

of an operculum present, and may have been

capable of feeding early in their ontogeny. Some

of these zooids develop thin calcification over

their frontal walls later in ontogeny, and these

develop multiporous septular pores, which are the

origins of rhizoids 10-20 mm long. The total

number of Ilustrine subcolonics is 40 and the

largest occur from SLOPE stn 22 and Tangaroa

stn 164, comprising 90 astogenetic generations

and bifurcating four times. Ovicells are frequent,

and when they arc present, avicularia arc rare.

The flustrine subcolonics from Kimbla stn 100

and the ‘Endeavour' sample, which have bila¬

minar phase 3 subcolonies developing at the tips

of their fronds, have numerous avicularia but few

200

P. E. BOCK AND P. L. COOK

or no ovicells. The 27 bilaminar phase 3 sub¬

colonies have the largest number of rhizoids

(Cook, 1979). The number of flustrine sub¬

colonies arising from the bilaminar phases seems

to be limited only by the number of marginal

kenozooidal chambers which have not already

produced rhizoids. One specimen from the

Endeavour sample has approximately 120 zooids,

spanning five astogenetic generations, with par¬

tially calcified proximal zooids and nearly 60

marginal rhizoids, 10 mm long. Only one stalk

kenozooid is present. On the other hand, one sub¬

colony from SLOPE stn 22 has 11 stalks and 20

rhizoids emanating from a group of 70 zooids. II

all these stalks developed into large, flustrine sub¬

colonies, they would have the potential to pro¬

duce more than 20,000 brooding zooids with

ovicells. Some of the zooids from bilaminar sub¬

colonics from Kimbla sins 99 and 100 have their

interiors filled w'ilh orange tissue. Their orifices

are completely occluded by the development and

fusion of the oral spines. The tissue is reminiscent

of that which fills the 'closed' zooids of some

species of the genus Conopeum (Cook, 1985: 87),

which can apparently function as hibcniacula. It

is inferred that if the closure of orifices is corre¬

lated with a non-feeding role for bilaminar sub¬

colonies, the tissue-filled, closed zooids of Cor-

hidipora may function as similar storage areas.

Notes on niultiphased growth

Cloning by accidental fragmentation and sub¬

sequent isolation has been discussed for several

colonial marine animals including Bryozoa by

Jackson (1983). Cloning by specific scries of

astogenetic changes leading to the production of

isolated subcolonies is knowm in the free-living,

lunulilifotm species of Cupuladriidae, particu¬

larly in the genus Discoporella (Cook, 1985;

Winston, 1988). These species are components of

sand fauna assemblages, and similar environ¬

ments are associated w'ith records of Corhulipora

tubidifera, particularly those from the Endeavour

.sample (see above). Flustrine and bilaminar sub¬

colonies were attached by their rhizoids to very

large colonies of Caienicellidae and fiustrinc

colonies of Spiralaria. These in turn were

anchored to coelentcrate colonies, and the entire

mass was covered and intertwined with worm-

tubes, sponges and smaller encrusting and

creeping bryozoan colonies. Quantities of shell

fragments and foraminiferans, together with sand

grains were also present. This sample was from a

locality and depth close to that from which

another sample from Gabo Island originated,

which was analysed by Cook (1979). This com¬

prised a similar assemblage of animals and colony

forms, including subcolonies of bilaminar

C tubidifera. Both assemblages probably formed

a mat on the surface of the bottom sediments. The

minute shell fragments encaistcd by the ancestru-

lale phase of C tubidifera suggests that these sub¬

colonies may have lived both on the surface and

within the the upper layers of the sediments. Inter¬

stitial colonies living on shell fragments were

described by Winston and Hdkansson (1986), and

included several cribrimorph species. Although

the records of C tubidifera arc from much deeper,

and quieter waters, most from more than 50 m.

and many from more than 100 m depth, an inter¬

stitial life w'ould still be possible. The rhizoids

which sometimes emanate from the growing

margins of encrusting subcolonics would also

help to anchor the large, erect, flustrine sub¬

colonies as they grew, and before their own

proximal rhizoids were developed later in

ontogeny.

Bock and Cook (1994) have already speculated

briefly on the roles of the different phases of

C. tubidifera in establishing, extending and main¬

taining the colony asexually and sexually (Fig.

32). It is possible to infer similar roles among dif¬

ferent phases of the two fossil species, C. ornafa

and C. suggerens. The encrusting subcolonics of

both species are small; they inhabit a similar size

of shell fragment substratum as lho.se inhabited by

C. tubidifera. and may w'ell have lived in similar

interstitial and semi-interstitial conditions. The

erect, ovicellatc phase of C. ornata is much more

heavily calcified than the equivalent, fiustrinc

phase of C. tubidifera, but would extend the

colony and aid dispersal of lar\'ae in the same

w^ay. Although there are numerous specimens of

the small, bilaminar subcolonics of C suggerens

in the material examined, very few possess

ovicells, and these occur only singly, near the

proximal margins of the subcolonies on zooids

with no costal frontal shield (Figs 28, 29). It

seems reasonable to infer that, if the ‘missing’

cellularinc or flustrine phase of C suggerens

existed, it too would have borne the majority of

the ovicells. This inference is strengthened by the

close similarity between the bilaminar phases of

C. tubidifera and C. suggerens. Subcolonics of

both species develop large chambered pores and

kenozooids at the margins w'hich arc known to

produce both stalk kenozooids and rhizoids in the

Recent species. It has been inferred that the bil¬

aminar phase is anchored by its rhizoids. After

fracture of the delicate flustrine subcolonics it is

isolated from the encrusting phase but stabilizes

THE MULTIPHASED BRYOZOAN GENUS CORBULIPORA

201

and maintains the extended colony, also produc¬

ing more flustrine phases and increasing the area

occupied by the clone even further. To a certain

extent this stabilizing role may be enhanced by

the non-feeding nature of many zooids of the bil-

aminar phase and it is interesting that, although

quite diflcrent in origin and subcolony structure,

the kenozooidal third phase of C ornata may

have had a similar function. The kenozooidal

masses also have large chambered pores etc., at

the surface which may have given rise to rhizoids.

Although much more robust than the erect phase

of C. tubitlifera, the quadriscrial subcolonies of

C ornata may have also been susceptible to frac¬

ture, especially if they had originated from stalk

kenozooids.

Notes on sonic other records of JVatersia

Watersia rnilitaris has not been fully described

since its introduction and the genus Watersia has

rarely been mentioned. Mawatari (1956) rclerred

to "Watersia kishakaensis\ The synonymy

included two previously described taxa, Fliistra

simplex kishakaensis Okada, 1918 and Euthy-

roides simplex Okada, 1921. These species are

quite distinct from one another, although both

have flustrine colonies and hyperstomial ovicclls.

Watersia kishakaensis has large ovicells with

paired frontal entooecial areas, and small, inter-

zooidal avicularia with rounded mandibles, orien¬

tated distally. Euthyroides simplex has much

smaller ovicells and no avicularia. Dr Shunsuke

Mawatari has kindly lent us some preparations of

W. kishakaensis, together with some manuscript

notes on species, made by his father. Dr Shizuo

Mawatari. These confirm the separate identity of

the two species, and the details of Mawatari’s

(1956: 121, fig. 5a) description. The ancestrula

and early astogeny of neither W. kishakaensis nor

W. simplex have been obscrv'cd and described.

Both species bear several resemblances to some

Antarctic forms assigned to KlngeJJustra by

Moyano (1972), who introduced the genus for

two flustrine species which had large, hyper¬

stomial ovicclls with paired, entooecial frontal

areas. Both the type species, Flustra vanhoeffeni

Kluge (1914), and Flustra drygalskii Kluge

(1914), have colonics with broad bilaminar

fronds, possess intcrzooidal avicularia, develop

numerous rhizoids, and have recently been

described from the Antarctic by Rosso (1994) and

by Hayward (1995). Gordon (1985) regarded

Klugeflustra as a junior synonym of Watersia,

and although much more robust, both the Antarc¬

tic species resemble the Watersia-phaso of C.

tiihulifera in their colony form, thinly calcified

zooids, spathulate avicularia and large ovicells.

However, Ristedt (1991: 378, pi. 2 fig. 1) has

described the early astogeny of K. vanhoeffeni in

a specimen from the Weddell Sea (76°55.2'S,

40°58.9'W, 216-264 m). The ancestrula is very

large (Lan 1.0 mm), encrusting and membrani-

porine. with no marginal or oral spines, gymno-

cyst or cryptocyst. The encrusting base of the

colony consists of approximately 50 similar auto-

zooids. A specimen of K. vanhoeffeni from the

South Shetland Islands {Discovety stn 1872, near

Clarence and Elephant Islands, 63°29.6'S,

54°()3.rW, 247 m, 12 Nov 1936), has been

examined. The robust fronds arc 45 mm in length,

the zooids large, very thinly

calcified, with no cryplocyst and two to three

large, multiporous lateral septular pores. The

avicularia arc large, spathulate, and occur proxi-

mally to each bifurcation of zooid rows. The avic¬

ularia are raised distally and the descending part

of the subrostral chamber is more heavily calci¬

fied than the rest, and appears white in contrast to

the brown cuticle of surrounding walls. The prox¬

imal opcsia is large and the mandible is hinged on

a bar. Ovicells occur in identical bands on both

sides of the fronds, five to seven astogcnctic

generations deep, and at intervals of about five

generations apart. Ovicclls arc very large, arc

closed by the partially open operculum, and

have a pair of long, irregular frontal entooecial

areas Hanking a central suture. These may be

subdivided into four foramina in some cases.

In spile of the close similarities between

K. vanhoeffeni and the rF^7/env/t/-phasc of C tubii-

lifera, the complete lack of a cribrimorph phase in

early astogeny, suggests that the genus Kluge¬

flustra should be maintained for the Japanese and

Antarctic species described above.

One further Antarctic Hustrinc genus, Neo-

flustra Lopez Gappa (1982), is also known to

have hyperstomial ovicclls, but these do not have

frontal entooecial areas. The avicularia are

numerous but adventitious and occur on a distinct

proximal zooidal gymnocysl. It is interesting that

although the autozooids of the type species, N.

dimorpha, have no spines, the brooding zooids

bear several pairs of Hattencd. overarching,

marginal spines, the distal pair being enlarged

like those on the brooding zooids of the Watersia-

phasc of C. tubulifera. Klugeflustra kishakaensis,

K. simplex, K. vanhoeffeni and K. drygalskii,

together with Neoflustra, form a distinctive sub¬

group, at present classified within the Flustridae.

Their ovicell stnicturc is so unlike that found in

all other members of the family, however, that it

202

P. E. BOCK AND P. L. COOK

seems almost certain that a new family grouping

may be eventually required to accommodate

them. However, little more can be inferred with¬

out much more information about the early asto-

genetic stages of colonies. The relationship of

these genera with Corbulipora, and the Crib-

rimorpha in general, is even more obscure and

cannot be evaluated until more is known about the

occurrence of multiphase growth within the

Bryozoa.

The records of Watersia sp. (Gordon 19S5) and

of Watersia militaris from New Zealand given by

Livingstone (1929) and by Gordon (1986), are not

referable to the genera and species discussed here,

but belong to the anascan family Calloporidac

(Cook and Bock, 2000).

Acknowledgements

We are very grateful to several colleagues for

notes on, and the loan of specimens from the col¬

lections in their care. These include Drs Peter

Arnold (Museum of Tropical Queensland,

Townsville), Penny Berents (Australian Museum,

Sydney), P.J. Chimonidcs and Mary Spencer

Jones (Natural History Museum, London),

Dennis Gordon (NIWA Wellington), Peter

Hayward (University of Wales, Swansea),

Shunsuke Mawatari (Hokkaido University,

Sapporo), David Holloway (Museum Victoria),

and Claus Nielsen (Zoological Museum. Copen¬

hagen). We also thank Mr David McDonald for

his help in preparation of this paper.

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from the southern Australian continental shelf.

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Research. 34; 303-354.

Waters, A.W. 1881. On fossil chilo.slomatous Bryozoa

from south-west Victoria. Australia. Quarterly

Journal of the Geological Society 37: 309-347.

Waters, A.W. 1883. Fossil chilostomatous Bryozoa

from Muddy Creek, Victoria etc. Quarterly Jour¬

nal of the Geological Society 39: 423-443.

Waters. A.W. 1887a. Bryozoa from New South Wales,

north Australia, etc. Part I. Annals and Magazine

of Natural History’ (5) 20: 81-95.

Waters, A.W. 1887b. Bryozoa from New South Wales,

north Australia, etc. Part 2. Annals and Magazine

of Natural History' (5) 20: 181 -203.

Winston. J.E.. 1982. Marine Bryozoans (Ectoprocta) of

the Indian River area (Florida). Bulletin of the

American Museum of Natural Histoty 173 (2):

99-176.

Winston, J.E., 1988. Life histories of free-living

bryozoans. National Geographic Research 4 (4):

528-539.

Winston, J.E. and IL^kansson, E. 1986. The interstitial

bryozoan fauna from Capron Shoal, Florida.

American Museum Noviiates, 2865: 1-50.

204

P. E. BOCK AND P. L. COOK

Appendix. Details of Tertiary localities in south¬

eastern Australia

Baimsdale (Skinner’s): Mitchell R. bank, about 12 km

W of Baimsdale, Vic. 37°47.9’S, 147°29.5’E.

Miocene.

Balcombe Bay: also known as Fossil Beach, Moming-

lon. Mount Martha and possibly ‘Schnapper Point’

(MacGillivray); on coast of Port Phillip Bay, about

3 km S of Momington. Vic. 38®14.5’S,

145°0l.7’E. Fyansford Clay. Balconibian; Middle

Miocene, (Langhian).

Bird Rock: Coastal section, about 3 km W of Torquay,

Vic. 38°2I.4’S, 144“17.8’E. Jan Juc Formation.

Janjukian; Late Oligocene, (Chattian). Also known

as Spring Creek.

Campbells Point: On Lake Connewarre, 12.5 km SE of

Geelong. Vic. 38°I3.7' S, 144°26.7’ E. Middle

Miocene.

Cape Otway: also Point Flinders. Locality AW I.

Coastal section 2 km NW of Cape Otway, Vic.

38°51.rS, 143°29.5’E. Glen Aire Clay. Early

Oligocene (Rupelian/Latdorfian).

Cooriemungle area: road cuttings about 18 km N of

Princetown. 38°32.4’S, 143°08.rE. Gcllibrand

Marl. Balcombian.

Curdies Creek: this is the locality name applied to the

bryozoans described by Waters (1881). The

original paper mentions ‘Yarra Yarra’, which is the

river on which Melbourne is located, and from

which no bryozoan fossil localities have been

recorded. Later publications (e.g. MacGillivray,

1895) have used the locality Curdies Creek for this

collection. The coastal section at the mouth of the

Curdies R. exposes Late Miocene limestone with a

sparse bryozoan fauna. Inland sections are gener¬

ally in limestone but some of the muddy sediments

of the Gcllibrand Marl arc exposed near Timboon.

However, it is believed that the sample is more

likely to have come from coastal sections, which

were much more accessible at that time. If the

sample was obtained from the coastal exposure, it

almost certainly was close to the locality listed as

‘Princetown’ below.

Grices Creek; also known as Gunyong Creek: on coast

of Port Phillip Bay, about 8 km N of Momington,

Vic. 38^n.9’S. 145“03.9'E. Fyansford Clay. Bal¬

combian (some material may be Baimsdalian);

Middle Miocene, (Langhian).

Mount Schanck: limestone quarry about 1 km W of

Mount Schanck, about 15 km S of Mount Gambicr,

SA. 37°57'S. 140'^43.2’E. Gambicr Limestone.

Longfordian; Early Miocene.

Muddy Creek: Clifton Bank, Muddy Creek, 8 km W of

Hamilton. Vic. 37'^44.6’S. 141°56.4'E, Muddy

Creek Marl ( = Gcllibrand Marl). Balcombian.

Narrawaturk Bore 2: at Peterborough, Vic. 38°36.3'S,

142°52.3'E. Core 9, 522.1-526.4 m. Gcllibrand

Marl. Early Miocene (Longfordian).

Princetown {Gigantocypraea locality): coastal section,

about 2 km W of Princetown, Vic. 38°41.9’S,

I43°08.3’E, Gcllibrand Marl. Balcombian.

THE MULTIPHASED BRYOZOAN GENUS CORBULIPORA

205

Figures 1-5. Corhulipora ornata. Figs 1-2, Mount Schanck, SA, encrusting, phase I subcolonies. Fig. 1, ancestrula

(a) and primary triad of zooids. Note that the first distal zooid (zl) has costae with no lateral fusions: the second

and third zooids have costae with lateral fusions and raised marginal spinous processes. Note pore chambers (p),

potential frontal bud arising from raised chambered pore (c), and raised, medially fu.sed oral spines on tlic third

zooid (arrowed) (x70). Fig. 2, encrusting, globular subcolony, showing raised, medially fused oral spines (arrowed)

(x62). Figs 3-5, Balcombe Bay, Vic., erect, pha.se 2 subcolonies. Fig. 3, zooids early in ontogeny with long gym-

nocyst and small, simple coslalc shield (incomplete or worn), and avicularium with distally expanded rostrum and

paired condyles (x8()). Fig. 4, zooids later in ontogeny with median area of frontal shield with pelmalidia (x50). Fig.

5, zooids late in ontogeny with numerous, raised pelmatidia and prominent suboral bar (x42).

206

P. E. BOCK AND P. L. COOK

Figures 6-9. Corbulipotu oniata. Balcombc Bay, Vic., subcolonics of erect, phase 2 and kenozooidal, phase 3

zooids. Fig. 6, note central column of phase 2 zooids surrounded by frontally budded costalc zooids and kenozooid.'

(arrowed) x45). Fig. 7, enlargement of zooids, note marginal chambered pores (small arrows), and raised walls of

partially calcified kenozooids (large arrows), all originating above the pelniatidia of underlying, phase 2 zooid>

(x7()). Figs 8-9. later astogenetic stages. Fig. 8, subcolony showing mass of spinous frontal shields of aulozooids

and kenozooids, the outlines of which arc no longer visible. The central core of the original erect, phase 2 subcolony

zooids is visible distally and proximally (x30). Fig. 9, enlarged portion of the same subcolony (xlOO).

THE MULTIPHASED BRYOZOAN GENUS CORBVLIPORA

207

Figures 10-13. Corhulipora sitggerens. Fig. 10, Princetown. Vic., encnisting. phase I subcolony, zooids showing

spinous, costate frontal shields: pore chambers (arrowed) (x50). Fig. 11, Balcombc Bay, Vic., bilaminar, phase 3

subcolony arising from paired, proximal zooids (x70). Fig. 12, Mount Schanck, SA, slightly wider, bilaminar, phase

3 subcolony, with paired proximal avicularia and zooid with uncalcified frontal (arrowed) (x64). Fig, 13. Balcombc

Bay, Vic., enlargement of zooid from distal part of subcolony in Fig. II, showing large, frontally uncalcified,

distal kenozooid arising from chambered pore (arrowed) (xl60).

208

P. E. BOCK AND P. L. COOK

Figures 14-17. Corbulipora siiggerens. Figs 14-15, Balcombe Bay. Vic., zooids from distal margin of subcolony

in Fig. 11 , showing large kenozooids arising from distal chambered pores; note relatively small size of the calcified

orifice. Fig. 14 (xl20). Fig. 15 (xl50). Compare both figures with the origin of stalk kenozooids of Cjiihuliferii

illustrated in l*ig.20. Fig. 16, Princclown, Vic., sing'c zooid from encrusting, phase 1 subcolony showing frontal

shield and oral spines. Note large chambered pore surrounding distal oral spine, probably derived from a pore cham¬

ber, and relatively small size of calcified orifice. Compare with C. tuhulifera. Fig. 17 (x210). Figure 17. CorhuH-

pora tuhuUfera. Tangaroa sin 155, Bass Strait, single zooid showing frontal shield and oral spines. Note relatively

large size of calcified orifice (x200).

THE MULTIPHASED BRYOZOAN GENUS CORBULIPORA

209

Figure 18. Corbulipora tuhulifera, characteristics of phases, a. encrusting, phase 1 zooids with stalk kenozooids (k),

arising from marginal chambered pores, developing rhizoids (r) and budding autozooids (z) at the base of an erect,

flustrinc phase 2 subcolony, b. tip of a frond of a tlustrine, phase 2 subcolony, with an avicularium (av), and devel¬

oping distal transitional zooids and bilaminar, phase 3 zooids with costate frontal shields (see also Fig. 24). c. zooids

of flustrine, phase 2 subcolony with ovicells (ov) and large oral spines. Scale ^ 0.60 mm.

210

P. E. BOCK AND P. L. COOK

Figures 19-22. Corbulipora tiihulifera. Bass Strait. Fig. 19. Tangaroa sin 155, encrusting, phase 1 autozooids with

distal stalk kenozooid (s), arising from chambered pores; note aulozooid frontal membranes and opcrcula {x80)

Figs 20-22, Kimhla stn 100. Fig. 20, small, bilaminar. phase 3 .subcolony, with rhizoids (r) and a pair of large stalk

kenozooids (s), sec al.so Fig. 21 (x35). Fig.21, enlargement of zooids from Fig. 20, showing origin of stalk keno-

zooid.s from distal chambered pores; compare with Figs 14 and 15 (x80). Fig. 22. bilaminar. phase 3 subcolony with

distal rhizoids and a stalk kenozooid (s); proximal autozooids with partially calcified frontal shields (arrowed); sec

also Figs 25 and 26. Note progressive, distally directed closure of zooid orifices by fused oral spines (x28).

THE MULTIPHASED BRYOZOAN GENUS CORBULIPORA

211

Figures 23-26. CorhuHpora tuhulifeni. Bass Strait. Fig. 23, Tangaroa stn 155, early astogeny of an encrusting,

phase 1 subcolony, on an aclconid bryozoan; anccstrula (a) (x46). Figs 24-26, Kimbla stn 100. Fig. 24, part of a tlus-

trinc, phase 2 frond, developing transitional distal zooids, and phase 3, calcified zooids at the tip (x31): compare

with Fig. 18b. Figs 25 and 26, zooids with uncalcificd or partially calcified frontal shields from proximal zone of

transition from phase 2 to phase 3 subcolony shown in Fig. 22. (x90).

212

P. E. BOCK AND P. L. COOK

Figures 27-31. Ovicclls in Corhulipora. Fig.27. CorbuUpora onuita^ Balcombe Bay, Vic. Brooding zooids of erect,

phase 2 subcolony, showing ovicclls. Crescentic cntooecial area (arrowed) and raised, medially fused oral .spines

(arrowed) (x75). Figures 28 and 29. Corhulipora suggerens. Fig. 28, Balcombe Bay, Vic., brooding zooid with

uncalcificd frontal shield and ovicell with crescentic cntooecial areas (xl30). Fig. 29, Princctown, Vic., brooding

zooid with uncalcificd frontal .shield and ovicell with paired pores (x200). Figures 30 and 31, Corhulipora tuhiilif-

era^ Bass Strait, Tangaroa stn 164. Fig. 30, brooding zooids from ilustrine, phase 2 subcolony, showing uncalcifieil

frontal membranes, enlarged oral spines and ovicclls with crescentic entooccial areas (x80). Fig. 31, part of a

flustrine, phase 2 subcolony, showing uncalcificd frontal membranes, one ovicell and two avicularia (xlOO).

THE MULTIPHASED BRYOZOAN GENUS CORBUUPORA

213

Figure 32. Diagram of inferred roles of different phases in Corhulipora tubulifera. 1. encrusting phase (e) estab¬

lishes subcolony on shell fragment which becomes anchored by rhizoids in and on the surface layers of sediment.

2. erect ilustrine phase arises from stalk kenozooids (s), budded from marginal chambered pores of phase I zooids.

Flustrinc phase produces brooding zooids with ovicells (o), which brood embryos of motile larvae (1), which dis¬

perse and settle on shell fragments, metamorphosing and producing anccstrulae, which bud new, phase I sub¬

colonies (c). 3. bilaminar phase originates from the tips of fronds of the flustrinc phase, and becomes anchored on

surface of sediment by rhizoids (r). These subcolonies maintain the area occupied by the Oustrine phase 2 sub¬

colonies, even if they become isolated. Phase 3 subcolonics then bud further phase 2, flustrinc subcolonies from

marginal stalk kenozooids (s), and repeat the sexually reproductive phase. Not to scale.

Memoirs of Museum Victoria 58(1): 215-222 (2001)

NOTES ON THE GENERA NORDGAARDIA AND USCHAKOVIA

(BRYOZOA: BUGULIDAE)

Patricia L. Cook

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

Abstract

Cook, P.L., 2001. Notes on the genera Nordgacmiia and Uschakovia (Bryozoa: Bugulidae).

Memoirs of Museum Victoria 58(2): 215-222.

The rare deep-water bryozoan genera Nordgaardia and Uschakovia share important simi¬

larities in budding paltenis and zooid structure together with a type ofavicularium which has

an extended subrostral chamber. Ovicells are known only in Nordgaardia and are here

described in Australian specimens of N. cornucopioides for the first time. The remarkable

spines of Uschakovia gorhimovi arc described from a colony from the Faroe Islands; they are

extensions of the frontal body wall, and each has a modibed parietal muscle at its base.

Introduction

Two species of the genus Nordgaardia Kluge,

1962 have been described in some detail:

N. pusilla (Nordgaard, 1907), type species, by

Hayward (1978), and N. cornucopioides by

d’Hondt (1983). The type and only species of

Uschakovia Kluge. 1946, U. gorhimovi Kluge,

1946, has been described solely from Russian

Arctic specimens by Kluge (1946, 1962). Deep¬

water bryozoans often have delicate colonics with

thinly calcified zooids and the oppominity to

examine wcll-prcser\'ed specimens is rare. A

single almost complete colony of U. gorbunovi,

together with 18 colonies of A. cornucopioides,

seven of which possessed both ancestrulac and

ovicells, arc here compared directly from mate¬

rial in the collections of Museum Victoria. The

relationships of the two genera with each other

and with some other genera of the superfamily

Buguloidea are assessed.

The specimen of V. gorhimovi is part of a col¬

lection including several colonics of this species,

collected as part of the Undcrsogclsor af den

marine bunddyrfauna omkring Faeroesne (BIO-

FAR, Investigations of the marine benthic fauna

of the Faroe Islands). The other specimens are

stored at the Marine Station. Vidskridvbakka,

FR-180, Kaldbak, Foroyar. The figured speci¬

men, was presented to Museum Victoria by Dr

P.J. Hayward (Marine Research Group, Univer¬

sity of Swansea, UK), who investigated the

Bryozoa from this collection (Hayward, 1994),

The specimens of N. cornucopioides were among

material sorted by P.E. Bock from Museum

Victoria’s SLOPE stations (taken on the south¬

eastern Australian slope from RV Franklin using

a WHOI cpibcnthic sled) and arc stored in

Museum Victoria (NMV).

Nordgaardia Kluge

Nordgaardia Kluge, 1962: 437.

Tvpe species. Svnnotuni pusillum Nordgaard,

f9()7.

Diagnosis. Colony erect, arising from an ances-

Irula anchored by numerous rhizoids, followed by

a short chain of uniserial autozooids. Autozooids

thinly calcified, elongated, tubular; proximal

gymnocyst expanding to surround an extensive

opesia with an almost terminal operculum. Distal

part of zooids free, projecting from branch sur¬

face. Autozooids arranged in alternating pairs or

in triads. Avicularia arising proximal to opesia,

subrostral chamber greatly extended, expanded

terminally into an acute, beaked rostrum. Ovicell

with partially membranous cctooecium, sur¬

rounding a thinly calciUcd capsule of entooe-

cium. attached to distolatcral end of the brooding

zooid, with a laterally directed aperture which

partially obscures the brooding zooid operculum.

Remarks. Nordgaardia pusilla, originally intro¬

duced for material from 1000 m in the northern

Atlantic, was redescribed in detail by Hayward

(1978: 215, Figs 3d-h), who had 12 samples from

the western European continental slope from a

depth range of 610-1400 m. This material

included colonies with ancestrulac and ovicells

which he described for the first time. D’Hondt

(1983) introduced a second species, Nordgaardia

215

216

PATRICIA L. COOK

cormicopioides from Brazil which differs from N.

pusilla in having a greater proportion of each

zooid freely projecting from the branch surface

and in having more robust avicularia. D'Hondt's

specimens possessed neither ancestrulae nor ovi-

cells. The colonies described below from the con¬

tinental slope of eastern Tasmania and Victoria

appear to be referable to N. conmeopioides

and allow description of both these features, and

comparison with N. pusilla.

Camptoplites marchemarcluidi Redier and

d’Hondl (1976) was assigned to Nordgaardia by

d’Hondt (1983: 82). Although d'Hondt (1983)

distinguished the two genera from each other on

the basis of their avicularian types, that of N.

marchemarchadi having an elongated subrostral

chamber, not a flexible ‘stalk’ as is present in

Camptoplites, he mentioned that avicularia

occurred only on the secondary branches. The

development of these secondary branches is a

characteristic found only in Camptoplites, and

was fully described by Hastings (1943). The

generic position of C. marchemarchadi is there¬

fore equivocal. The type specimen was from West

Africa, and it is possible that the fragmentary

specimen from Senegal assigned to Kinetoskias

sp. by Cook (1968: 63), which had similar

avicularia, may have belonged to the same

species.

Nordgaardia cornucopioides d’Hondt

Nordgaardia cormicopioides d’Hondt. 1983: 80,

pl.l, figs 3^ (off Brazil, 770-805 in).—Harmelin and

d'Hondt, 1992: 31 (Gulf of Cadiz, 281 m).—d’Hondt

and Gordon, 1996: 85, fig. IID (New Caledonia,

1175-1950 m).

Material examined. Stn SLOPE-32, 38°21.90’S,

149°20'E, 23 Jul 1986, S of Point Hicks, Vic., 1000 m.

stn SLOPE-33, 38°19.60’S, I49‘=24.30'E, 23 Jul 1986,

S of Point Hicks, Vic., 930 m. stn SLOPE-34,

38°16.40’S, 149°24.30’E, 23 Jul 1986, S of Point

Hicks, Vic., 800 m. stn SLOPE-45, 42°2.20’S

148°38.70’E. 27 Jul 1986, off Freycinet Peninsula'

Tas., 800 m. Ail NMV F74868.

Description. Nordgaardia with at least half the

distal part of each zooid free and projecting front

the branch surface. Avicularia expanded and trun^

cate distally, with a hooked rostrum. Ovicelh

very large.

Remarks. The four colonies from Tasmania (stp

SLOPE-45) and three of those from Victoria

(stn SLOPE-32) have ancestrulae and rhizoidsi

present, and branch three to four times. Ovicells

are present in all these colonies, which include, on

average, more than 100 zooids each. The ances^

trula is tubular, attached by 10-12 proximal

rhizoids, and has a narrow opesia extending

nearly the whole of its length. It gives rise to onQ

distal zooid which bears 15-20 rhizoids along itsi

entire length. The first pair of autozooids follows.

Subsequent autozooids are arranged in alternating

pairs or in triads with frontals inclined to one side

of the branch so that the long tubular proximal

parts of the zooids tend to be visible only on the

basal side. Hayward (1978) described the slightly

different budding and branching patterns of

N. pusilla in detail and also noted the difficulty in

understanding the morphology of the ovicelU

because their delicate calcification was distorted

by collection and preservation. The ovicclls of

N. cornucopioides are also very thinly calcified

and are somewhat distorted. The ovicclls arc

larger than those of N. pusilla (Table 1) and hav'c

an almost completely membranous ectooeciuni

which surrounds a distinctly smaller calcified

capsule ofcntooecium. The two layers merge at a

small attachment area on the basal side of the dis-

tolateral wall of the brooding zooid which has a

wider border of gymnocyst on the outer side. The

large aperture of the oviccll opens laterally above

the operculum of the brooding zooid which is

Table 1. Measurements in mm of Nordgaardia pusilla, N. cornucopioides, and Uschakovia gorbunovi

from specimens and literature. ‘ .

N. pusilla

N. cornucopioides

U. gorbunovi

Length of ancestrula

1.30-1.50

1.70-1.85

1.00-1.70

Length of autozooids

1.40-1.80

0.90-1.20

0.76-0.83

Length of opesia

0.76-1.00

0.65-0.75

0.50-0.54

Length of subrostral chamber

0.54-0.85

0.67-0.74

0.45-0.66

Length of rostrum

0.12-0.19

0.19-0.30

0.12-0.27

Length of oviccll

0.36

0.45-0.50

Width of ovicell

0.28

0.39-0.45

Length of tentacles

—

0.40-0.45

0.25-0.29

NORDGAARDIA AND USCHAKOVIA (BRYOZOA, BUGULIDAE)

217

Figure 1. Nordgaardia conmcopioides d’Hondt, Tasmania, a, primary aslogenetic zone: ancestrula {an), rhizoids

(r). b, later astogcnctic zone: avicularia ovicell (ov). Scale = 1 mm.

therefore partially obscured (Fig. lb). Ovicells

with an uncalcified ectooecium were figured in

Bugula longissima Busk by Hastings (1943: Fig.

39D) and ovicells with a laterally facing aperture

obscuring the brooding zooid operculum were

figured in Kinetoskias mitsukurii by Yanagi and

Okada (1918: 425, Fig. 7a) and in K. ehngata by

1 farmer (1926: 469, pi. 34, fig. II). The closely

similar ovicelled zooids of Enoplo 2 oum cirratwn

(Busk) were illustrated by d’Ffondt and Gordon

(1996: 86, fig. 9D).

A few autozooids have partially protruded ten¬

tacle crowns. The tentacles are closely opposed

aiid contracted in most cases but in a few zooids

they are slightly expanded with an estimated total

of 14-16 tentacles in the crown. The avicularia do

not reach the largest rostral dimension given by

dM Iondt (1983, i.e. length of rostrum 0.30 mm)

but are within the range he figured. They are more

robust than those of N. pusilla and their rostra are

hooked terminally. The extensive muscle systems

occupy nearly all the distal, expanded part of the

elongated subrostral chamber.

D'Hondt’s specific name refers to the similar¬

ity of the avicularia with those occurring in some

species of the genus Cormteopina Levinsen.

hfamicr (1926) described several species

including C. moluccensis (Busk) which has

large avicularia almost exactly like those of

N. conmcopioides. The extended proximal parts

of the autozooids and the complex budding

patterns of the two genera are also somewhat sim¬

ilar but Cornucopina differs completely in the

expanded shape of the distal part of its autozooids

and in the occurrence of numerous marginal

spines.

The locality of the Tasmanian specimens of

N. conmcopioides is remote from most other

records of Nordgaardia which arc all from the

Atlantic. However, recently, specimens from very

deep water from New Caledonia have been

recorded by d’Hondt and Gordon (1996). It is just

218

PATRICIA L. COOK

Figure 2. Uschakovia gorhunovi Kluge, Faroe Islands, a, primary astogenetic zone: founding zooids (fz), rhizoids

(/•), primary linking zooid {h /). b, later astogenetic zone: secondary linking zooid {Iz 2), spines (.v), inward-facing

zooids (/), avicularia (av). Many spines not shown completely. Scale ^ I mm.

possible that a single rather variable taxon is con¬

cerned. Harmelin and d'Hondl (1992) noted that

their colonies from the north-eastern Atlantic (off

Cadiz, 35“43.5\ 6°I8.2’W, 281 m) had some

avicularia which approached the more slender

type found in N. pitsiiia. The avicularia of the

Tasmanian population of N. corniicopioides are

generally less robust than those originally

described by d'Hondt (1983) from Brazil, and

the very delicate, slender avicularia described

for N. marchemarchadi. from Senegal,

suggest that a direct comparison of all the

available material might reveal a continuum of

character states.

NORDGAARDIA AND USCHAKOVIA (BRYOZOA, BUGULIDAE)

219

Figure 3. Uschakovia gorhunovi Kluge, Faroe islands. Two autozooids and avicularia, showing modified parietal

muscles (w). Spines not shown completely. Scale = 0.5 mm.

Uschakovia Kluge

Uschakovia Kluge, 1946; 196.

Type species. Uschakovia gorbunovi KXxxgQ, 1946

(monotypic).

Diagnosis. As for U. gorbimovi.

Uschakovia gorbunovi Kluge

Uschakovia gorbunovi Kluge, 1946: 196 , pi. 1, figs

I^.—Kluge, 1962: 438, Figs 228A-D (Arctic, Kara

and East Siberian Seas, 64-648 m).— Hayward, 1994:

183 (Faroe Islands, 610 1400 m).

Materia! examined. BIOFAR stn 095, ()60°41.5rN,

05°I8.63'W, south-east Faroe Islands, 23 Jul 1987, 803

m, NMV F74869.

Description. Colony erect, arising from an ances-

trula anchored by rhizoids which gives rise to a

series of 'founding’ zooids which also develop

rhizoids. Erect branched part of colony originat¬

ing from one zooid. or a short uniserial chain of

'linking’ zooids which arc extremely elongated.

Branches of autozooids in alternating pairs and

triads, becoming quadriscrial before a bifurcation.

Primary branch bifurcating several times to form

a cluster of 4~6 branches. Autozooids with very

long tubular proximal gymnocyst expanding dis-

tally to surround an elongated opesia. Outward¬

facing zooids with bipartite opesia, distal part

covered by frontal membrane which surrounds

operculum, the proximal part formed by the

swollen bases of pair of long partially cuticular

spines. Inward-Hieing zooids without spines,

opesia elongated. Avicularia arising from proxi¬

mal gymnocyst of both kinds of zooids; the sub-

rostral chamber considerably elongated and

expanded distally to form a tenninally hooked

acute rostrum. Ovicells unknown.

Remarks. The colony examined here does not

differ materially from those described from the

high Arctic by Kluge (1962). Colonics attain a

height of 10-15 mm and a width of 5-8 mm, and

arc composed of two groups of distinctly different

zooids, each group connected by a single elon¬

gated linking zooid (length 1.00-1.14 mm). The

colony from the Faroes has no ancestrula; those

ligured by Kluge (1962) have anccstmlae which

are more elongated than, but which otherwise

resemble, the succeeding founding zooids. The

groups of founding zooids (length 0^.50-0.55 mm)

alternate with one or tw'o linking zooids and this

succession may be repeated. The colony from the

Faroes has a few founding zooids followed by a

linking zooid which has no calcification. A fur¬

ther group of approximately 20 triserially budded

founding zooids, the more proximal with rhizoids,

220

PATRICIA L. COOK

is then followed by a second, single calcified link¬

ing zooid. The remainder of the colony is formed

by bifurcation of a biserial-to-quadriscrial branch

which is budded distally to the linking zooid

(Fig. 2). The resulting colony has six branches.

All zooids arc very thinly calcified early in

astogeny at least. The tubular proximal gymno-

cysts are closely apposed and each expands dis¬

tally to surround the opesia: the tcnninal third of

this upper part of the zooid is free and projects

from the branch surface. The founding zooids

appear to have passed through ontogenetic

changes including resorption of calcification and

they have completely cuticular body walls. A line

marking the presumed foniier position of the edge

of the gymnocyst is present in some zooids and all

are swollen, irregularly orientated, and have open

opcrcula. The underlying orifices are however

scaled by cuticle. The more distal zooids have

paired cuticular tubercles near the distal ends of

the former opesia; the more proximal zooids have

long uncalcified, spinous processes, or paired rhi-

zoids in the same position. Other rhizoids arise

from the proximal part of these zooids. The more

proximal of the two linking zooids is more

slender and longer than the neighbouring found¬

ing zooids but also has cuticular walls. The more

distal linking zooid is separated from the nearest

proximal founding zooid by two cuticular nodes

with an intcr\^cning small calcified intemode

(Fig. 2b). The gymnocyst is thinly calcified and

very elongated and tubular, expanding distally to

surround a bipartite opesia which is divided by

lateral constrictions formed by the swollen bases

of paired spinous processes. The opesia proximal

to the constriction is short and both parts of the

frontal membrane form an angle at the constric¬

tion. The distal part is almost circular and the

small almost semi-circular operculum has a

marginal sclcritc. The autozooids of the next three

to four astogenetic generations resemble the link¬

ing zooid but have a shorter, tubular gymnocyst

and longer narrower proximal opesia. The oper¬

culum and the circular distal part of the frontal

membrane are sunken with a cowl-like raised rim

of semitransparent gymnocyst with a crenulated

border.

After four to five astogenetic generations the

first bifurcation occurs and branches become tri-

to quadriserial; they arc close to each other and

the autozooids facing inw'ard dilTcr from those

facing outward which are in the majority. The

inward-facing zooids are flattened and have a

wide elongated opesia without any trace of spines

or lateral constrictions; a distal pair of parietal

muscles is visible in many zooids. The outward¬

facing zooids all have paired spines and laterally

constricted bipartite opcsiac. The spines are

1.55-3.50 mm long and arc direct expansions of

the body wall. On the proximal face they are

derived from the cuticle of the frontal membrane;

on the distal face they are thinly calcified and

continuous with the lateral gymnocyst which

forms a curved shoulder round the base of each

spine. The spines are curved at their origin but

extend distally and are numerous and long enough

to obscure the underlying zooids completely.

Although flattened and partially calcified at first,

they become tubular and completely cuticular. At

the base of each spine the frontal cuticle is

expanded to form a tubercle and muscles extend

through this from the cuticular frontal face to the

interior of the shoulder and the lateral part of the

gymnocyst (Fig. 3). These muscles appear to be

modified and enlarged parietal muscles. All the

zooids are so thinly calcified that they are some¬

what shrunken by preservation in alcohol. In life

the frontal membranes w'ould be convex espe¬

cially when the tentacle crowns were retracted.

The base of the spines would therefore not con¬

strict the opesia as much as they do in the pre¬

served state. However, contraction of the parietal

and other muscles during tentacle protrusion

would presumably cause the proximal part of the

frontal membrane to become concave. This would

alter the position and direction of the elongated

frontal spines. The avicularia which arise from

the proximal gymnocyst are similar to but much

less robust than those of N. conuicopioUIes; the

elongated proximal part of the subrostral chamber

is narrower and the ‘head' more elongated.

Discussion

In colony fonn and budding patterns, and in the

possession of erect rhizoid-bearing ancestrulae

and elongated avicularia, Nordgaardiu and

Uschakovia resemble each other closely. Kluge

(1962) suggested that the founding zooids of

Uschakovia might function in nutrient storage as

well as in attachment of colonics. The swollen,

cuticular zooids resemble the ba.sal vesicular

kenozooids of Caidibugida (I lastings, 1939)

which may also function in both of these ways-

The greatly extended cuticular peduncle of KitW'

toskias (see Kluge. 1962) which has rhizoids at its

base is analogous to the founding zooid of

Uschakovia but is an extrazooidal structure in

direct contact with all autozooids. It presumably

expands continuously w'ilh growth in the same

manner as the peduncle of the ascophoran genus

Parmularia (Cook and Chimonides, 1985). The

NORDGAARDIA AND USCHAKOVIA (BRYOZOA. BUGULIDAE)

221

apparent resorption of calcification in Uschakovia

seems to be an ontogenetic change which pro¬

ceeds distally from the anccstrular region. The

earliest founding zooids and the more proximal

linking zooid bear only rhizoids, the more distal

founding zooids bear long uncalcified spines and

show traces of cuticle which may mark the foniier

position of a gymnocyst. The distal linking zooid

and first pair of autozooids have completely cal¬

cified gymnocysts but like all the founding zooids

their open opercula have an underlying orifice

sealed by cuticle. This gradient of ontogenetic

change suggests that the resorption of calcifica¬

tion is a successive process which, in the larger

colonies figured by Kluge (1962), may have pro¬

ceeded further than in the specimens from the

Faroes.

Nordgaardia and Uschakovia exhibit an inter¬

esting mosaic of characteristics of other genera

within the Bugulidae. Both genera have similar

avicularia which also closely resemble those of

several species oi' Cornucopina (Hanncr, 1926)

as noted above. However, Cornucopina differs in

autozooid morphology although its budding

pattern does resemble that oiN. pusilla. The avic¬

ularia of some species of Kinetoskias although

less elongated are similar to those of Nordgaardia

in structure and position (Yanagi and Okada,

1918; Manner, 1926). As noted above, the

ovicells of Nordgaardia resemble tho.se of

Kinetoskias which has however a very different

colony form (Kluge, 1962). The modified parietal

muscles at the base of the spines in U. gorhunovi,

are, like the spines themselves, apparently unique.

However, the 'flexor zooecii’ muscles of both

Kinetoskias and Euoplozown (Busk, 1881;

Harmer, 1926) arc analogous in some ways

although certainly not homologous.

An elongated ancestrula attached by long rhi¬

zoids is found in several genera of Bugulidae.

Hastings (1943) illustrated the ancestnilac of

several species of Camptoplifes in detail. The

zooid shape and to a certain extent the budding

patterns and the avicularia of all the deep-water

genera, including Nordgaardia, Uschakovia,

Kinetoskias and some fonns of Camptoplifes,

show a large number of similarities with one

another which cannot all be ascribed to the com¬

mon features of their environments. In all

respects, the extensive continuum of character

correlations found in the family Bugulidae

(Gordon, 1984) seems to accommodate all four

genera. Hayward (1978) considered that the bud¬

ding and bifurcation patterns of Nordgaardia

were so close to those of the genera Epistomia

and Synnotwn that both Nordgaardia and

Uschakovia could be retained within the family

Epistomiidae (Hastings, 1943; Ryland and

Hayward, 1977). The members of this family do

not however possess elongated avicularia or

hyperstomial ovicells. Kluge (1962) introduced

the family Sadkoidae to include Nordgaardia and

Uschakovia. Nordgaardia has been included in

this family by dMIondt (1983) and by Harmclin

and d'Hondt (1992). However, as noted by

Ryland (1982: 760), the family name is invalid

as no genus Sadkoa, from which the family

name should have been derived, seems ever to

have been described. In any case, the family is

unnecessary as both genera are assignable to the

Bugulidae.

Acknowledgements

1 thank Dr Peter J. Hayward (Marine Research

Group, Swansea University), for presenting the

colony of Uschakovia to Museum Victoria for

examination, and to Philip E. Bock for all his help

and encouragement.

References

Busk, G., 1881. Notes on a peculiar form of Polyzoa

closely allied to Bugula (Kinetoskias, Kor. and

Dan.). Quarterlv Journal of Microscopical Science

21; 1-14.

Cook, P.L., 1968. Bryozoa (Polyzoa) from the coasts of

tropical West Africa. Atlantide Report 10:

115-262.

Cook, P.L. and Chimonides, P.J., 1985. Larval settle¬

ment and early astogeny of Panmilaria

(Chcilostomata). Pp. 71-78 in Nielsen, C. and

Larwood, G.P. (eds). Rryvzoa: Ordovician to

Recent. Olsen and Olsen: Fredensborg.

Gordon. D.P., 1984. The marine fauna of New

Zealand: Bryozoa: Gymnolaemata from the Ker-

madcc Ridge. Memoirs of the New Zealand

Oceanographic Institute 91; 5-198.

Harmclin, J.G. and d'Hondt, J.L., 1992. Bryozoaires

des parages de Gibraltar (campagne

oceanographique BALGIM. 1984) I —

Cheilostomes. Bulletin du Mushnn National

d'Histoire Naturelle (Serie 4) I4A: 23-67.

Harmer, S.F., 1926. The Polyzoa of the Siboga

Expedition. Part 2. Chcilostomata, Anasca.

Sihoga-Espedilie Monograph 28b: 181-501.

Hastings, A. B., 1939. Notes on some cellularine Poly¬

zoa (Biy'ozoa). Novitates Zoologicae 4\ : 321-344.

Hastings, A. B., 1943. Polyzoa (Bryozoa), 1.

Scrupoccllariidae, Epistomiidae, Farciminariidae,

Bicellariellidae, Aeteidae, Senipariidae. Discovery

Reports 22\ 301-510.

Hayward, P.J., 1978. Bryozoa from the West European

continental Journal of Zoology’, London 184:

207-224.

222

PATRICIA L. COOK

Hayward. P.J., 1994. New species and new records of

cheiloslomatous bryozoa from the Faroe Islands,

collected by BIOFAR. Sarsia 79: 181-206.

d’Hondt, J.-L., 1983. Nouvclle contribution a Fetude

des Bryozaires Eurystomes bathyaux et abyssaux

de 1’ocean Allantique. Bulletin dii Museum

National d'Histoire Naturelle (Serie 4) 5A:

73-99.

d’Hondl, J.-L. and Gordon, D.P., 1996. Bryozoa:

Ctenostomes ct Cheilostomes (Cellularines. Scru-

pariines et Malacosteges) des campagnes

MUSORSTOM autour de la Nouvelle-Caledonie

(Rcsultats des campagnes MUSORSTOM 15).

Menwires du Museum National d Hisloire

Naturelle 168; 55-123.

Kluge, G.A., 1946. [Novye i maloizvestnye mshanki iz

sevemogo Ledovitogo okeana]. New and little

known Bryozoa from the Northern Ice Ocean.

Trudi dreifmmschei ekspeditslva Glavsevmorputi

na l/s G. Sedov, 1937-I940; III: 194-223 [In

Russian].

Kluge, G.A.. 1962. [Mshanki Sevemykh Morei SSR].

Bryozoa of the northern seas of the USSR.

Opredelitii po Faune SSSR. Izdavaemye ZoO'

logicheskim Muzeem Akademii Natik 76: 1-584 [In

Russian]. (1975, English translation, Smithsonian

Institution and the National Science Foundation,

New Delhi).

Nordgaard, O., 1907. Bryozoen dem norwcgischen

Fischereidampfer ‘Michael Sars’ in den Jahren

1900-1904 gesammell. Bergens Museum Aarhok

2: 3-20.

Redier, L. and d’Hondt. J.-L. 1976. Contribution a Pe-

tude des Bryozaires de POucsl africain (rccoltes de

M.l. Marche-Marchad au large du Senegal el de la

Mauritanic). Bulletin de /' Institut Fondamental

d'Afrique Noire A38 (4): 851 -858.

Ryland, J.S., 1982. Bryozoa. Pp. 743-769 in Parker,

S.P. (ed.) Synopsis and classification of living

organisnLs. McGraw Hill: New York.

Ryland. J.S. and Hayward, P.J., 1977. British anascan

Bryozoans. Synopses of the British Fauna 10.

Linnean Society and Academic Press: London.

Yanagi, N. and Okada, Y., 1918. On a collection of

Japanese Cheiloslomatous Bryozoa 1. Annota-

tiones Zoologicae Japonenses 9 (4): 407-429.

Memoirs of Museum Victoria 58(2): 223-230 (2001)

FOUR NEW SPECIES AND A NEW RECORD OF CHIMARRA STEPHENS

(TRICHOPTERA: PHILOPOTAMIDAE) FROM BOUGAINVILLE ISLAND,

PAPUA NEW GUINEA

David 1. Cartwright

13 Brolga Crescent, Wandana Heights. Victoria 3216, Australia

Abstract

Cartwright, D.I., 2001. New species and a new record of Chimarra Stephens (Trichoptera:

Philopotamidae) from Bougainville Island, Papua New Guinea. Memoirs of Museum Vietoha

58(2): 223-230

Descriptions and keys are provided for males of five species of caddisllies of the widespread

genus Chimarra from Bougainville Island, Papua New Guinea, including four new species.

Females of four species are also keyed and described.

Introduction

The cosmopolitan caddisHy genus Chimarra

Stephens, 1829 has not been recorded from

Bougainville Island, although 21 species have

been described from mainland New Guinea

(Papua New Guinea and Irian Jaya, Indonesia)

(Neboiss, 1986; Malicky, 1994). Two have been

described from the nearby Solomon Islands,

Guadalcanal Island {9°32'S, I6()°12'E) (Kim-

mins, 1957a). Specimens of the genus were col¬

lected during 1987-1990 by Ms Cathy Yule near

Arawa (6°I2'S, 155°32'E) and deposited in

Museum Victoria, Melbourne. A total of 21

males and 18 females were sorted to five species.

Females were paired with respective males on the

basis of similarities in coloration particularly on

the head, and on wing venation and locality.

The Bougainville Island Chimarra species dis¬

play the following character states: in the

forewing: the curved or sinuous Rs vein basal to

the discoida! cell is associated with thickened

veins in four of the species; abdominal segments

VIII and IX arc elongate in females of three of the

four known species; lateral lobe processes arc

present on tergum X in males of all species; and

in one species, the dense brush of small spines is

present on the phallus. The forewing with vein Rs

curved has been cited as a key diagnostic feature

for the subgenus Chimarra {Chimarra) separat¬

ing it the from other subgcnera of Chimarra in

the New World (Blahnik, 1998). This feature is

also present in many species from Australia

(Cartwright, in prep.), several species from Asia

(Kimmins, 1957b, 1964) and Africa (Kimmins,

1963). Kimmins (1957b, 1962) reported that

both C. crepidata Kimmins from India and

C. papuami Kimmins from New Guinea have the

phallus containing numerous spines at the apex, a

characteristic also of C. pinga sp. nov.

Both males and females arc most readily distin¬

guished by gcnitalic features, often requiring the

clearing of the abdomen in potassium hydroxide.

Figured specimens arc identified by the note¬

book numbers of Dr Arturs Neboiss (NMV), pre¬

fix PT-; or the author; prefix CT-. Abbreviations

for genitalic parts are indicated on Figs 1-3 and

21 .

Depositories are abbreviated as follows:

BMNH, the Natural History Museum, London;

NMV, Museum Victoria, Melbourne.

Chimarra Stephens

Chimarra Stephens, 1829; 318.—Mosely and

Kimmins, 1953: 398.

Type species. Phryganea marginata. Linnaeus,

1767 by monotypy.

Remarks. A diagnosis of the genus Chimarra and

subgenus Chimarra was provided by Blahnik

(1998).

Key to males and females of species of Chimarra from Bougainville Island

1. Males.2

— Females.6

2. Inferior appendages in lateral view, long and slender, length > 3.5 times

width (Figs 1,4).3

— Inferior appendages in lateral view, not long and slender, length < twice

width (Figs 7, 10).4

223

224

D. I. CARTWRIGHT

3. Inferior appendages, very long and slender over the whole length, length > 5

times width (Figs 1-3). Chimarra longpela

— Inferior appendages, long and slender over the apical half, broadened in

basal third, length <4 times width (Figs 4-6). Chimarrapanguna

4. Inferior appendages with large process present on mcsal margin (Fig. 8);

phallus with dense brush of pale spines apically (Figs 7-9).. Chimarra pinga

— Inferior appendages without process present on mesal margin; phallus with¬

out dense brush of pale spines apically (Figs 11.14).5

5. Inferior appendages in lateral view widely bifid (Fig. 10)

. Chimarra hiramosa

— Inferior appendages in lateral view not bifid (Fig. 13). Chimarrayuleae

6. Abdominal segments VUI and IX short and robust (Fig. 21)

. Chimarra hiramosa

— Abdominal segments VIII and IX long and slender (Figs 22, 23). 7

7. Abdominal segments VII! and IX in ventral view, of similar width, and not

tapered apically (Fig. 22)... Chimarra pinga

— Abdominal segments VIII and IX in ventral view, not of similar width,

tapered apically (Figs 23, 24).8

8. Abdominal segment IX extended apically to form an elongated point Figs

23, 23a). Chimarra lon^ela

— Abdominal segment IX not extended apically to form an elongated point

(Fig. 24). Chimarra yuleae

Chimarra longpela sp. nov.

Figures 1-3, 17, 23, 23a

Type material. Holotype male, Papua New Guinea,

Bougainville I., Panguna, light trap, 7 Dec 1989, C.

Yule (NMVT-17489).

Paratypes, all same locality and collector as holotype:

1 male, 13 Apr 1988 (genitalia prep. PT-1795. figured);

1 male. Mar 1989; 1 male, 19 Apr 1988; 3 males, 17

Dec 1988; 2 males, 2 females (genitalia prep. CT-284.

figured), 29 Jan 1989; 1 female, Feb 1989; I female,

Apr 1989, I female, 1 Oct 1988 (all NMV).

Other material examined. Papua New Guinea,

Bougainville I.. Konaiano Ck, f*anguna, 24 Sep 1988,

C. Yule, I female (NMV).

Description. General colour brownish, including

head, mesolhorax and wings; although triangular

area on head between ocelli dark brownish. Wing

venation: forewing veins Rs and M curved and

thickened basal to discoidal and median cells

(Fig. 17); forewing with forks 1, 2, 3 and 5

present; hindwing with forks 1,2, 3 and 5 present.

Male. Genitalia dark brown. Ventral process on

segment IX absent; inferior appendages in lateral

view, very long and slender, length about 6 times

width (Fig. 1), in ventral view, straight and

slender, length about 5 times width, tapered

slightly apically (Fig. 2); lobes of tergum X

simple, divided dorsomedially for most of length

(Fig. 3); phallus tubular with a conspicuous dark

spine embedded subapically (Figs I, 3).

Female. Abdominal segments VIII and IX

elongate and slightly laterally compressed,

segment IX extended apically to form an

elongated point; cerci short (Figs 23, 23a).

Length of forewing: male 5.0-5.9 mm, female

4.9-5.8 mm.

Etymology^ Longpela — New Guinea pidgin

word for long, referring to the inferior

appendages of the male.

Distribution. Papua New Guinea, Bougainville I.

(males known only from the type locality).

Remarks. In the male, the extremely elongate and

slender inferior appendages are distinctive. The

female genitalia are also extremely elongate and

similar to C. yuleae sp. nov. and C. pinga sp. nov,

Chimarra longpela females can be distinguished

by the elongated point on abdominal segment IX.

Chimarra panguna sp. nov.

Figures 4-6, 18

Type material. Holotype male, Papua New Guinea,

Bougainville I.. Panguna. 29 Aug 1988, C. Yule (NMV

T-I7503).

Paratype male, same data as holotype: genitalia prep.

CT-282, figured (NMV).

Description. Male. General body colour brown,

head and mesolhorax dark brown contrasting with

pale warts; wings brownish. Wing venation:

forewing vein Rs curved anteriorly basal to dis-

coida! cell (Fig. 18); forewing with forks I, 2, 3

and 5 present; hindwing with forks 1, 2, 3 and 5

present.

NEW CADDIS-FLIES FROM BOUGAINVILLE ISLAND

225

Genitalia brown. Ventral process on segment

IX absent; inferior appendages broadest basally,

narrowed at about half length, apical half slender,

length about 3 times width (Figs 4, 5); lateral

lobes of tergum X simple, divided dorsomcdially

for most of length (Fig. 6), shorter than inferior

appendages, slightly downtumed apically; phallus

tubular, inserted between lateral lobes, with 2

dark spines embedded subapically and basally

(Figs 4, 6).

Female unknown.

Length of forewing: male 4.1-5.0 mm.

Etymology’. Panguna — named after the type

locality (Panguna), noun in apposition.

Distribution. Papua New Guinea, Bougainville I.

(known only from type locality).

Remarks. The males of C. panguna can be distin¬

guished from other Bougainville species by the

slender apical half and robust basal half of the

inferior appendages.

Chimarra pinga sp. nov.

Figures 7-9, 19, 22

Type material. Holotype male, Papua New Guinea,

Bougainville I., Panguna, light trap, 17 Dec 1988, C.

Yule (NMV T-17505).

Paratypes, all same locality and collector as holotype:

1 male, 20 Jan 1988 (genitalia prep. PT-1793, figured);

I male, Feb 1989; 1 female, 19 Dec 1989 (genitalia

prep. CT-286. figured) (all NMV).

Description. General body colour pale, including

head and mesothorax, although triangular area on

head between ocelli dark brownish; wings brown¬

ish. Wing venation: forewing discoidal cell veins

R4-5 and R2-3 thickened basally, Rs curved ante¬

riorly basal to thickening (Fig. 19); forewing with

forks 1,2, 3 and 5 pre.sent; hindwing with forks 1,

2, 3 and 5 present.

Male. Genitalia dark brown. Ventral process on

segment IX absent; inferior appendages in lateral

view subtriangular, length about 1.5 times width

(Fig. 7) with a mesal digitifomi projection sub¬

apically (Figs 8, 9); lateral lobes of tergum X sim¬

ple, slightly truncated apically, divided dorsome-

sally for most of length (Fig. 8); phallus tubular,

dilated apically with a field of small, pale spines,

and with conspicuous dark spine embedded sub¬

apically (Figs 7-9).

Female. Abdominal segments VIII and IX in

ventral view, long and slender, of similar width,

and not tapered apically; cerci short (Fig. 22).

Length of forewing: male 5.1-5.2 mm, female

5.7 mm.

Etymology. Pinga — New Guinea pidgin word

for finger, referring to finger-like mesal projec¬

tion on the inferior appendages of the male.

Distribution. Papua New Guinea, Bougainville I.

(known only from type locality).

Remarks. Chimarra pinga is a distinctive species

in that the male has digitiform projections on the

inferior appendages and a phallus with an apical

"brush' of spines, separating it from all known

Australasian species. The female differs from

other Bougainville species on the basis of the

abdominal segments VIII and IX, having the

combination long and slender, of similar width,

and not tapered apically.

Chimarra biramosa Kimmins

Figures 10-12, 16, 21

Chimarra biramosa Kimmins, 1957a; 292, figs 4a,

5.—Neboiss, 1986: 108.

Type material. Holotype male, Solomon Islands,

Guadalcanal I., Tapenanje, 10-15 Dec 1953, J.D.

Bradley (BMNH). Type not seen.

Paratypes, 1 male, 6 females, all same data as holo¬

type (BMNH). Paratypes not seen.

Material e.xamined. 1 male, Papua New Guinea,

Bougainville I., Arawa 10/39, 21 Dec 1989,

C. Yule (NMV); 1 male, same locality and col¬

lector, 13 Jun 1988 (genitalia prep. PT-1797, fig¬

ured); 1 female, same loc. and collector, 12 Mar

1988; 1 female, same loc. and collector, 14 Aug

1988 (genitalia prep. CT-283, figured); 1 female,

same loc. and collector, 25 Dec 1989; 1 female,

Panguna, light trap, 29 Jan 1989, Q. Yule; 2

females, Panguna, 3 Dec 1987, C. Yule (all

NMV).

Description (revised after Kimmins, 1957a).

General body colour brown, head and mesothorax

dark brown contrasting with pale warts; wings

brownish. Wing venation: forewing vein Rs

slightly curved basal to discoidal cell, forewing

with forks 1, 2, 3 and 5 present (Fig. 16); hind¬

wing with forks 1, 2, 3 and 5 present (Kimmins

1957a: fig. 4a).

Male. Genitalia dark brown. Ventral process on

segment IX absent; inferior appendages in lateral

view short with dorsal extension basally (i.c. bifid

or biramous) (Fig. 10), in ventral view, broad,

tapering apically (Fig. 11); lateral lobes of tergum

X simple, length almost as long as inferior

appendages (Fig. 12); phallus tubular, with con¬

spicuous asymmetrical dark spine subapically

(Figs 10-12; Kimmins 1957a: fig. 5a-c).

Female. Abdominal segments VI11 and IX

226

D. I. CARTWRIGHT

short; segment VIII dark brownish; segment IX

with pair of dark brown sub-triangular ventral

plates; cerci short (Fig. 22; Kimmins 1957a: fig.

5d).

Length of forewing: male 4.1-4.5 mm, female

5.0-5.2 mm.

Distribution. Solomon Islands, Guadalcanal I.

and Papua New Guinea, Bougainville I.

Remarks. Chimarra biramosa males differ from

all other Australasian species in the widely bifid

inferior appendages of the male (Kimmins,

1957a). The female differs from the other

Bougainville species in having short abdominal

segments VIII and IX. Kimmins' (1957a) figures

have been redrawn to allow direct comparisons

and to accompany the description that is revised

in light of new interpretations of Chimarra

genitalic structures.

Chimarra yuleae sp. nov.

Figures 13-15, 20, 24

Type material. Holotype male, Papua New Guinea,

Bougainville I., Panguna, light trap. 17 Dec 1988,

C. YulefNMV T-17509).

Paralypcs. all same locality and collector as holotype:

I male, 3 Dec 1987 (genitalia prep. PT-I792. figured);

1 male, 29 .Uil 1988 (genitalia prep. PT-1796); 1 male,

Feb 1989; 1 female, 17 Dec 1988 (genitalia prep. CT-

285, figured); 1 female. Apr 1989 (all NMV).

Description. General body colour pale, including

head, mesothorax and wings. Wing venation:

forewing with a heart-shaped raised thickening at

the junction of veins Ml-2 and M3 (Fig. 20);

forewing with forks 1, 2, 3 and 5 present; hind¬

wing with forks 1, 2, 3 and 5 present.

Male, Ventral process on segment IX absent;

inferior appendages in lateral view triangular,

narrowed basally, broadened apically, with

apicodorsal projection, length about same as

width, with mcsal flange and tapering to acute

apex (Fig. 13), in ventral view, basically spb-

rectangular (Fig. 14); lateral lobes of tergum X

elongate, extending further than inferior

appendages, slightly upturned apically; phallus

tubular, with a conspicuous ventral dark spine

embedded subapically (Figs 13-15).

Female. Abdominal segments VIII and IX in

ventral view, not of similar width, tapered

apically; cerci short (Fig. 24).

Length of forewing: male 5.1-5.3 mm, female

6.0-6.1 mm.

Etymology. The species is named for Cathy Yule

(collector).

Distribution. Papua New Guinea, Bougainville 1,

(known only from type locality).

Remarks. In the male, the distinctive triangular

inferior appendages allow separation from other

Bougainville species. The female genitalia is very

similar to C longpela but lacks the elongate point

on segment IX.

Acknowledgements

I thank Ms Cathy Yule for collecting all the mate¬

rial studied. Dr Arturs Neboiss (NMV) for pro¬

viding access to the specimens, and him, John

Dean and two anonymous referees for comments

on drafts of this manuscript.

References

Blahnik, RJ., 1997. Systematics oi' Cliimarrita, a new

subgenus of Chimarra (Trichoptcra: Philopotami-

dae). Systematic Entomology' 22: 199-243.

Blahnik. R.J., 1998. A revision of the neotropical

species of the genus Chimarra., subgenus Chi*

marra (Trichoptcra; Philopolamidae). Memoirs of-

the American Entomological Institute 59: i-vi

1-318.

Kimmins, D.E., 1957a. Neuroptera and Trichoplera col¬

lected by Mr. J.D. Bradley on Guadalcanal Island.

1953-4. Bulletin of the British Museum (Natural

Histoty) Entomology 5: 289-308.

Kimmins, D.E.. 1957b. Entomological results from the

Swedish c.xpcdition 1934 to Bumia and British

India. Trichoptcra. The genus Chimarra Stephens

(Fam. Philopotamidae). Archiv fur Zoolosi 11*

53-75.

Kimmins, D.E., 1962. Miss L.E. Checseman's expedi¬

tions to New Guinea. Trichoplera. Bulletin of the

British Museum (Natural History) Entomoloiiy 11 ■

99-187.

Kimmins, D.E., 1963. On the Trichoplera of Ethiopia.

Bulletin of the British Museum (Natural History)

£'/i/omo/oj^v 13: 119-170.

Kimmins. D.E.. 1964. On the Trichoptcra of Nepal.

Bulletin of the British Museum (Natural Histoty)

Entomology 15: 35-55.

Malicky, IL, 1994. Neue Trichopleren aus Nepal, Viet¬

nam. China, von den Philipen und vom Bismarck-

Archipcl (Trichoptcra). Entomologische Berichte

Luzern M : 163-172.

Mosely, M.E. and Kimmins D.E., 1953. The Tri*

choptera (Caddis-flies) of Australia and New

Zealand. British Museum (Natural History): Lon¬

don. 550 pp.

Neboiss, A.. 1986. Atlas of Trichoptcra of the Sii'

Pacific-Australian Region. Dr W. Junk: Dordrecht.

286 pp.

Stephens. J.F., 1829. A systematic catalogue of British

insects. Part /. Baldwin: London. 416 pp.

NEW CADDIS-FLIES FROM BOUGAINVILLE ISLAND

227

Figures 1-6. Chimarra spp. Male genitalia in lateral, ventral and dorsal views. 1-3: Chimatra longpela sp. nov. 4-6

Chimarra panguna sp. nov.

Abbreviations: i.a., inferior appendages; X, tergum X; pha, phallus. All scale lines 0.1 mm.

228

D. I. CARTWRIGHT

Figures 7-12. Chimarra spp. Male genitalia in lateral, ventral and dorsal views. 7-9; Chimarra pinga sp. nov.

10-12: C. biramosa Kimmins.

All scale lines 0.1 mm.

NEW CADDIS-FLIES FROM BOUGAINVILLE ISLAND

229

Figures 13-20, Cfiimarra spp. Male. 13-15: Genitalia in lateral, ventral and dorsal views. Chimarrayuleae sp. nov.

16: Chimarra biramosa Kimmins, forewing. 17-20: section of forewing. 17: Chimarra longpela sp. nov. 18:

Chimarra panguna sp. nov. 19: Chimarra phtga sp. nov. 20: Chimarra yuleae sp. nov.

Abbreviation: dc, discoidal cell. Scale lines. Figs 13-15: 0.1 mm; Figs 16-20: 0.5 mm.

230

D. I. CARTWRIGHT

Figures 21-24, Chimarra spp. Female genitalia. 21: Chimarm hiramosa Kimmins, ventral view; 22: Chimana

pinga sp. nov., ventral view. 23, 23a: Chimarra longpela sp. nov. 23: ventral view; 23a: lateral view. 24: Chimarra

yuleae sp. nov. ventral view.

Abbreviations: IX, abdominal segment nine; VIII, abdominal segment eight. All scale lines 0.1 mm.

Memoirs of Museum Victoria 58(2): 231-246 (2001)

NEW SPECIES OF HYDROPSYCHIDAE (INSECTA: TRICHOPTERA)

FROM NORTHERN AUSTRALIA

John C.Dhan

Environment Protection Authority, Freshwater Sciences, GPO Box 4395 QQ, Melbourne,

Victoria 3001. Australia

(john.dcan@cpa.vic.gov.au)

Abstract

Dean. J.C., 2001. New species of Hydropsychidae (Insecta; Trichoptera) from northern Aus¬

tralia. Memoirs of Museum Victoria 58(2):231-246.

Five new species of hydropsychid caddisilies of northern Australia are described from

males, females and larvae: Cheumatopsyche kakaduensis sp. nov., C. suteri sp. nov., C well-

sae sp. nov., C. dostinei sp. nov. and Asmicridca capricornica sp. nov. Cheumatopsyche

kakaduensis sp. nov. has thus far only been recorded from Kakadu National Park, while the

remaining four species are widely distributed across northern Australia. Distribution maps are

presented with keys for identification of males, females and larvae.

Introduction

Caddisflics of the family Hydropsychidae of

northern Australia are poorly known, in recent

years a large amount of material from the

Alligator Rivers region. Northern Territory, has

been accumulated and a smaller amount from

north-western Australia. Four new species of

Cheumatopsyche Wallengrcn, 1891 and one of

^.w 7 ;/cr/V/ei/ Moscly, 1953 arc described here

from this material. Distribution records for north¬

ern Australia (including North Queensland) arc

presented. Additional undescribed species are

known from Cape York Peninsula and the

Queensland wet tropics but are outside the scope

of the present study.

Although only one species of Cheumatopsyche

has been described from Australia, recent exami¬

nation of larval material has revealed the exis¬

tence of at least 18 species (Dean, 1999).

Cheumatopsyche modica (McLachlan. 1871) was

described from material collected in Victoria.

While the name has subsequently been applied to

all adult Australian Cheumatopsyche material,

probably in part a consequence of the conserva¬

tive male and female genitalia, the identity of the

described species has yet to be determined. Of six

larval species currently known from Victoria, any

one of three or four could be C. modica. Since

none of these species occurs in northern Aus¬

tralia, description of northern species can proceed

prior to the identity of C. modica being estab¬

lished. A similar situation exists in Asmicridea.

Although lar\ac of the two described species

have yet to be delennined, type localities for both

species are in southern Australia. The northern

Australia larva described below is clearly

different from all larvae known from southern

Australia (Dean, 1999).

Associations of adults and larvae have been

achieved by collection of pupal chambers con¬

taining pharate male pupae and associated larval

scleritcs. Observations on colour arc based on

material preserved in 70% ethanol. Adult geni¬

talia have been prepared for examination by

clearing in cold potassium hydroxide for 24^8

hours. Terminology for adults follows Nimmo

(1987), while that for larvae follows Wiggins

(1977) and Shefter and Wiggins (1986). Type

material has been lodged in Museum Victoria,

Melbourne (NMV) or the Australian National

Insect Collection, Canberra (ANIC). Material

without a stated repository is lodged in Museum

Victoria. Not all of the material examined is listed

below and a full listing is available from the

author. Abbreviations for material examined are;

M, adult male; F, adult female; MP, male pupa;

FP, female pupa; P, unsexed pupa: L, larva.

Abbreviations for collectors are: AW (A. Wells),

DC (D. Cartwright), JB (J. Blyth), JD (J. Dean),

JEB (J.E. Bishop), MBM (M.S. and B.J.Moulds),

PD (P. Dostinc), PS (P. Suter).

Cheumatopsyche kakaduensis sp. nov.

Figures 1-11

Type material. Holotype: adult male, Gulungul Creek,

Radon Springs, Northern Territory (I2°45'S

I32°55'E), 13-14 Apr 1989. P.Siitcr and A.Wells

(NMV T-I7415). Paratypes collected with holotype: 6

males, 6 females (NMV); 5 males, 5 females (ANIC).

231

232

J. C. DEAN

Figures 1-5 Cheumatopsyche kakadiiensis sp. nov. Adult; 1, wings; 2, male genitalia, dorsal; 3, male genitalia, lat¬

eral; 4, female genitalia, lateral; 5, female genitalia, dorsal.

Other material examined. Northern Territory.

Numerous M, F, P, and L, collected with holotype; 1M,

Graveside Creek (13°18'S 132°32'E), 18Jul 1988, PD;

2P. IL, Little Nourlangie Rock Creek {12°52'S

I32°48'E), 22 Apr 1989, PS and AW; 2L. erdek 5 km

W of Gimbat OSS field station (13°33'S 132“34'E). 19

Apr 1989, AW and PS; 2P, 47L, Baroalba Springs

{12'’49'S 132°52'E), various dates, AW and PS, DC;

2L, Magela Creek, d/s Magela Falls (I2°46’S 133°6'E),

various dates, AW and PS, DC; 2L. Carbo Creek

(\3°\rS 132°51'E), 29 May 1988, AW and PS; IL,

Barramundie Creek, d/s Falls (13°22'S 132°28'E), 26

May 1988, AW and PS; 3L, Lone Spring (12°i7'S

132°36'E),2I Aug 1999, JD.

Description. Adult. General colour dark brown,

almost black. Vertex of head dark brown; anten¬

nae straw-coloured; thorax medium brown.

Wings (Fig. 1): forewdng dark brown, irrorate;

hindwing uniformly paler brown. Length of

forewing 4.9-6.1 mm (male), 5.0-6.3 mm

(female). Male genitalia (Figs 2, 3): distal lobes

of tergum X rounded, not turned up apically.

NEW HYDROPSYCHID CADDISFLIES FROM NORTHERN AUSTRALIA

233

Figures 6-11 Cheumatopsyche kakaduensis sp. nov. Pupa: 6, mandibles; 7, abdominal hook plates (3-8: abdominal

segment number, a: anterior hook plate, p: posterior hook plate); 8, apical process of abdomen. Larva: 9, fronto-

clypeus; 10, pronotum, anterolateral margin; 11, prostemites.

234

J. C. DEAN

separated by a distance approximately 3 times

their width; prc-anal appendages squat, located

well forward of base of distal lobe; terminal seg¬

ment of claspcr about quarter length of basal seg¬

ment; dorsum of abdominal segment IX broad in

lateral view. Female genitalia (Figs 4, 5): clasper

receptacle small, shallow; inner aperture aligned

with outer aperture, clearly visible in lateral view;

separation of receptacles in dorsal view 2-3 times

width of each receptacle.

Pupa. Right mandible with 3 subapical teeth,

left mandible with 4 (Fig. 6). Paired hook plates

on abdominal segments 3-8; anterior plates on

segments 3-8 and posterior plates on segments 3

and 4 only (Fig. 7); some variation in number of

teeth on all plates. Apical processes of abdomen

(Fig. 8) with apicolateral angles acute, separated

by shallow concave surface covered by small pro-

tmbcnces; outer margin of each process fringed

with long black setae, ventral surface with scries

of pale spine-like setae which protrude beyond

apex.

Mature lana. Head and thoracic sclerites pre¬

dominantly medium-dark brown; unpigmenled

area surrounding each eye; small pale region in

centre of frontoclypeus. Frontoclypeus length:

width ratio 1.54-1.59; anterior margin finely

crenulate with about 25 lobes (Fig. 9), Head and

pronotum densely covered with conspicuous

spine-like setae; those near posterior angle of

frontoclypeus bru.sh-like (Fig. 9); primary seta 17

on head short, length much less than distance

from setal base to apex of frontoclypeus; primary

seta 22 near anterolateral margin of pronotum

short, robust, not much longer than adjacent

secondary setae (Fig. 10). Posterior prostemites

reduced to small flecks, one adjacent to each

posterolateral angle of anterior prostemite (Fig.

11). Abdominal gills present on segments 1-7.

Etymology. The name reflects the apparent

restriction of the species to Kakadu National

Park.

Comments. C. kakaditensis appears to have a lim¬

ited distribution having been recorded only from

a few small creeks close to the escarpment and

from one lowland spring (Fig. 53). The lar\^a has

previously been designated Cheumatopsyche sp.

10 (Wells, 1991) and Cheumatopsyche sp. AVIO

(Dean, 1999).

Cheumatopsyche suteri sp. nov.

Figures 12-21

Type material. Ilolotype: adult male, South Alligator

River, Gimbat OSS Field Station, Northern Territory

(I3°35'S i32®36*E), 24 May 1988. A.Wells and

P.Sutcr (NMV T-17428). Paratypes collected with

hololype; 7 males. 4 females (NMV); 6 males, 2

females (ANIC).

Other material examined. Western .\ustralia. IM, 2F,

Drysdale River, Kalumbuni Crossing, 28 Sep 1979, JB;

20M. 35F. Camp Creek, Mitchell Plateau, various

dates. JEB, PS; 4M. Barnett River Gorge, 1 Oct 1979,

JB; 15L, Manning Gorge, near Gibb River Rd, Kimber¬

ley. 28 Jul 1994. DC: lOL, King Edward River,

Mitchell River Rd. 25 Sep 1995, L.Metzcling; IL. trib¬

utary of Bell Creek, Kimberley, 27 Jul 1994, DC.

Northern Territory. 6L, South Alligator River,

Kakadu Hwy crossing (13"17 S 132°19'E), 26 May

1988. PS and AW; 2M. 2F. Jim Jim Ca-ck, 3 km d/s

falls, 1 Sep 1979. JB: IM, IMP. IL, Katnbolgie Creek

(13°32'S 132‘^23'E). various dates, PS and AW, DC;

15L, Magcia Creek, d/s MagcIa Falls (I2M6'S

133°06'E), various dales. AW and PS, DC; 3M, IF,

16L, Graveside Gorge (I3°18'S 132°32'H). various

dates. AW and PS. PD; 3L, Baroalba Springs, Gubara

(12°49‘S 132^52'E). 28 Apr 1990, DC; 4 L. Koolpin

Creek (13°29'S I32'»35*E), 25 May 1988, PS and AW;

IL. Nourlangic Creek (12°28 S I32°44'E|. 26 May

1988, PS and AW; 2L. Lone Spring. Kakadu National

Park (12°17*S 132^36'E), 21 Aug 1999. .ID; IM, 2F.

Howard Springs, near Darwin. 23 Jiin 1969. Lc Souef;

IM. Devil Devil Creek. 70 km SW of Daly River Mis¬

sion. 23 Aug 1979; JB; 4M, IF. Katherine Gorge

National Park. 13 Aug 1979, JB; 2M. Roper River,

Mataranka Homestead. 25 Jan 1977, MBM: 6L. Mann

River, Amhemland (12^22'S 134°08*E), 26 Aug 1999,

JD; 18L, Liverpool River, Arnhemland (12°2rs

134°07‘E), 26 Aug 1999, JD: IL. Florence Falls. Litch¬

field National Park (13'^06 S 13°047 E). 31 Aug 1999.

JD. Queensland. lOM, 21F. 36L. Gunshot Creek, Tele¬

graph Crossing (1 ]*’44'S 142“29*E), Feb 1992. DC and

AW; 4M. 3F, 15L, Cockatoo Creek, Telegraph Cross¬

ing (1^395 I42°27 C), Feb 1992, DCandAW;2MP.

4L, Canal Creek, u^s Eliot Ck junction (11°23'S

142°25*E). 6 Feb 1992. DC and AW; IL. Peaches

Creek, 3 km NE of Coen (13'*42'S 143°I5'E), 4 Nov

1988, K.Walker; I5M. 8F, Upper Jardinc River, various

sites, Oct 1979. MBM: IM. Gordon Creek, Iron Range,

19 Apr 1975, MBM.

Description. Adult. General colour dark brown,

almosl-black. Venex of head dark brown; anten¬

nae stravv coloured, basal segments usually with

oblique darker bands; thorax medium brown.

Wings (Fig. 12): forewing medium-dark brown,

weakly irrorate; hindwing uniformly paler brown.

Length of forewing 5.9-1.2 mm (male), 5.7-7.1

mm (female). Male genitalia (Figs 13, 14): dor¬

sum of icrgum X mesally elevated; distal lobes of

tergum X rounded, not turned up apically, sepa¬

rated by approximately 4 times their width;

pre-annal appendages raised, elongate in lateral

view, situated close to base of distal lobes; ter¬

minal segmeni of claspcr almost a third length of

NEW HYDROPSYCHID CADDISFLIES FROM NORTHERN AUSTRALIA

235

Figures 12-21 Cheumatopsyche suteri sp. nov. Adult: 12, wings; 13, male genitalia, dorsal; 14, male genitalia, lat¬

eral; 15, female genitalia, lateral; 16, female genitalia, dorsal. Larva: 17, frontoclypeus; 18, secondary setae near

apex of frontoclypeus; 19, pronotum, anterolateral margin; 20, prosteniites; 21, foretrochantin.

236

J. C. DEAN

basai segment, narrow, apically turned inwards;

dorsum of segment JX narrow in lateral view.

Female genitalia (Figs 15, 16): clasper receptacle

large, broadly tubular; inner aperture clearly dor¬

sal to outer aperture, outer aperture wide, semi¬

circular; in dorsal view separation of receptacles

usually less than width of each receptacle.

Pupa. Abdominal hook plates similar to

C. kakaduensis.

Mature larva. Head and thoracic sclerites pre¬

dominantly pale-medium brown; unpigmented

area surrounding each eye. Dorsum of head some¬

what tlattcncd. Frontoclypeus lengthrwidth ratio

1.34-1.43; anterior margin finely crcnulalc with

approximately 27 lobes (Fig. 17). Secondary setae

numerous on posterior half of frontoclypeus,

sparser on anterior half; some setae dark, but

mostly pale and inconspicuous under dissecting

microscope; setae of frontoclypeus hair-like with

4 or 5 apical filaments (Fig. 18). Primary seta 17

on head moderately long, about as long as dis¬

tance from base of seta to apex of frontoclypeus.

Pronotum with dark secondary setae restricted to

anterior margin and region of middorsal line,

setae towards lateral margins paler; primary seta

22 near anterolateral margin of pronotum long,

tapered, about half length of pronotal scleritc

(Fig. 19). Posterior prostcmilcs reduced to small

flecks, one adjacent to each postero-latcral angle

of the anterior prosternite (Fig. 20). Abdominal

gills present on segments 1 7.

Etymologyi The species is named for Phil Suter

who collected much of the material on which the

present description is based.

Comments. The species is widely distributed

across northern Australia with many records from

the Northern Territory, north-western Australia

and Cape York Peninsula in North Queensland

(Fig. 54). The larva has previously been referred

to as Cheumatopsyche sp. 12 (Wells, 1991) and

Cheumatopsyefw sp. AVI2 (Dean, 1999).

Cheumatopsyche wellsae sp. nov.

Figures 22-30

Type material, llolotypc: adult male. East Alligator

River, Cahills Crossing, Northeni Territory (12°26'S

132°58 E), 27 May 1988, A.Wells and P.Suter (NMV

T-17440). Paralypes collected with holotype: 11 males,

7 females (NMV); 4 males, 4 females (ANiC).

Other material e.xamined. Western Australia. IM, IF,

Drysdale River. Kalumburu Crossing, 28 Sep 1979, JB:

13M, 26F, Camp Creek and vicinity, Mitchell Plateau.

variou.s dates, JEB. PS; IM, Barnett River Gorge,

Kimberley. 1 Oct 1979, JB; I6M, 2IF, Crossing pool.

Millstream, Pilbara, 21 Oct 1979, JB; 19M, 29F, Fortes-

cue Falls. Hamersley Range National Park. 27 Oct

1979. JB; 39M. 34F, Ord River, 9 km N ofKununurra,

19 Sep 1979. JB; numerous M and F, Spillway Creek.

Ord River Dam, 2 Feb 1978, JEB. Northern Territory,

Numerous L, Fa.si Alligator River, E of Jim Jim Falls

(13=*I2'S 133°20'E), 29 May 1988. AW and PS; 2F.

3MP, numerous L, South Alligator River, Gimbat OSS

Field Station (13®35'S 132'^36'E), various date.s, AW

and PS, PD; IM, 4F, Jim Jim Creek. 3 km d/s falls. 1

Sep 1979, JB; IIL, Coobanrbora Spring. Kakadu

National Park (I2“24'S 132°40*E), 21 Aug 1999, .ID;

2M, 30L. Kambolgie Creek (I3®32'S 132°23’E), vari¬

ous dates. PS and AW, DC; 1 MP, 8L. Magella Creek,

u/s Magela Falls (I2°47'S 133'’06'E). various dates,

AW and PS; IM. 2F. 4L. Graveside Gorge (13°18'S

I32®32 E), various dales, PD. AW and PS; IL.

Nourlangie Creek (12°28'S I32'’44’E). 26 May 1988.

PS and AW; numerous M. F, Katherine Gorge National

Park, 13 Aug 1979. JB; 4M, Roper Bar, 15 Jul 1969. Lc

Souef; 4F, Groote Eyiandt. Amagule Pool, 6 Feb 1984,

M.Davies; I5L. Mann River, Amhemland (I2°22'S

I34°08'E), 26 Aug 1999. JD; I3L, Liverpool River.

Amhemland (12°2rS I34''07'E). 26 Aug 1999. JD;

2L, Florence Falls, Litchfield National Park (13°06'S

I30°47'E),31 Aug 1999. JD. Queensland. IM, IF, IP,

I4L. Bertie Creek. Telegraph Crossing (I1°50'S

142^30'E), Feb 1992. DC and AW; 2MP. 36L, Cocka¬

too Creek, Telegraph Crossing (1P39'S 142‘^27'E),

Feb 1992, DC and AW; 7L. Pascoe River. 60 km W of

Lockhart (12°53S 143°0rE), lONov 1988. K.Walker;

5M, 4F, Archer River Crossing, Capo York Peninsula,

9 Sep 1974, MBM; IM, 4F, Curmnda Creek, trib. of

Freshwater Creek. Cairns district, 30 April 1979, AW.

Description. Adult. General colour pale-medium

brown. Wings (Fig. 22): forewing golden-pale

brown, without irroration; hind wing paler brown.

Length of forewing 5.2-6.3 mm (male), 4.9-6.1

mm (female). Vertex of head pale-medium

brown; antennae straw coloured, basal segments

without darker bands. Thorax golden. Male geni¬

talia (Figs 23-25): distal lobes of tergum X

strongly upturned, apices pointed both in lateral

and dorsal views; width of lobes somewhat vari¬

able. Terminal segment of clasper short, triangu¬

lar, about fifth length of basal segment; inner

basal angle acute. Female genitalia (Figs 26, 27):

clasper receptacle small; in lateral view consisting

of short, narrow chimney with circular aperture at

upper end, delicate sclerotisation extended

antcroventral from base of chimney; separation of

receptacles in dorsal view greater than 8 limes

width of each receptacle.

Pupa. Abdominal hook plates similar to C.

kakaduensis.

Mature laira. Dorsum of head medium brown,

lateral margins predominantly pale yellow, venter

of head extensively dark brown-black. Fronto¬

clypeus length;width ratio 1.54-1.62, anterior

margin coarsely crenulatc with about 15 lobes

NEW HYDROPSYCHID CADDISFLIES FROM NORTHERN AUSTRALIA

237

Figures 22-30 Cheumatopsyche wellsae sp. nov. Adult: 22, wings; 23, male genitalia, dorsal; 24, distal lobes of

male genitalia, variant; 25, male genitalia, lateral; 26, female genitalia, lateral; 27, female genitalia, dorsal. Larva:

28, frontoclypeus; 29, secondary setae near apex of frontoclypeus; 30, prosternites.

238

J. C. DEAN

(Fig. 28); secondary setae dear, appressed, incon¬

spicuous under dissecting microscope, those near

posterior angle of frontoclypcus predominantly

simple, the apex thin and whip-like (Fig. 29).

Pronolum pale-medium brown; secondary setae

clear, appressed and inconspicuous; primary seta

22 near anterolateral margin of pronotum long,

tapered, about half Icngtii of pronotal sclcritc.

Posterior prostemites consisting of pair of small

lateral scleritcs and pair of large mesa! scleriles.

(Fig. 30). Mesonolal and metanotal scleritcs

densely clothed with appressed fine, forward-

directed dark setae, forming short fringe along

anterior margin of both segments. Abdominal

gills present on segments 1-7.

Etvnw!og\\ The species is named for Alice Wells

who collected much of the material on which the

present description is based.

Comments. The species is widely distributed

acro.ss northern Australia, and has been collected

from the Pilbara and Kimberley regions of West¬

ern Australia, the Northern Territory, Cape York

Peninsula in North Queensland and further south

in eastern Queensland (Fig. 55). The larva has

previously been designated Chewnatopsyche sp.

11 (Wells, 1991) and Chenmatopsyche sp. AVI 1

(Dean, 1999).

Chenmatopsyche dostinei sp. nov.

Figures 31-38

Tx'pe material. Holotype: adult male. Adelaide River.

l '5 km E of Stuart 1 lighway. Northem Territory. 15 Aug

1979. J.Blyth (NMV T-17459). Paratypes collected

with holotype: 5 males, 1 1 females (NMV); 3 males, 3

females (ANIC).

Other material examined. Western Australia. IIM.

I2F, 15 km S of Windjana Gorge, 4 Aug 1989. McCub-

bin: 9M, 5F. Ord River. 9 km N of Kununurra. 19 Sep

1979, JB: IM. Geiki Gorge National Park. 9 Oct 1979,

JB: IM. Dunham River. 100 km S of Wyndham, 7 Feb

1977, MBM; 3M. I OF, Filzroy River Crossing, Derby-

Broome Road, 3 Nov 1978, MBM. Northern Terri¬

tory. 2M, 8F. 4L. East Alligator River, CahiUs Cross-

ing*(I2°26'S 132°58'E). 27 May 1988, AW and PS;

2MP, South Alligator River, Kakadu Hwy crossing

(13°I7 S I32‘'I9'H), 26 May 1988. PS and AW; 22L.

Wildman River. Arnhem Highway (I2°50 S I32°02 E),

22 Apr 1989, PS and AW; IM, IF. 14L, Kambolgie

Creek (I3'’32 S 132“23'E), various dates, DC. AW and

PS; IM. Jim Jim Creek, Kakadu llwy (12‘^57'S

132°33’E). 28 May 1988. PS and AW; 2MP. 9L.

Nourlangie Creek (12'^28'S I32"44 H), 26 May 1988,

PS and AW. Queensland. 9M, 6F. Upper Ro.ss River,

SW of Townsville, S May 1979, AW; IM. 3F, Cairns,

Lake Morris Rd (16’^55‘S I45°46 E), 16 Nov 1988,

K.Walker; 2M, 20F, Forty Mile Scrub, 65 km SW of Mt

Garnet, 19 Dec !974, MBM; IM, 2F, Archer River

Crossing. Cape York Peninsula. 9 Sep 1974. MBM.

Description. Adult. General colour palc-mcdiuni

brown. Wings (Fig. 31): forewing pale-mcdiuni

brown, without irroralion; hind wing very pale

fawn, almost white. Length of forewing 4.2-5.^

mm (male), 5.0-5.5 mm (female). Vertex of head

medium-dark brown; antennae with basal seg¬

ments yellow, without oblique darker bands.

Thorax medium brown. Male genitalia (Figs 32,

33): distal lobes of tergum X broad, somewhat

truncate in dorsal view, separated by a distance

about 3 times the width of each lobe: in lateral

view each lobe upturned. Pro-anal appendages

located well anterior of posterior margin of

tergum X. Ciasper with apical segment about

quarter length of basal segment. Female genitalia

(Figs 34, 35): ciasper receptacle moderately large,

subtriangular in lateral view; outer margin

oblique, orientated anteroventral to posterodorsal;

inner aperture small and dorsal: separation of

receptacles in dorsal view about 2 times the width

of each receptacle.

Pupa. Abdominal hook plates similar to

C kakadiiensis.

Mature lan^a. Dorsum of head pale-mediunt

brown, lateral margins predominantly pale

yellow, venter of head pale-medium brown. Froii'

toclypeus lcngth:widlh ratio 1.5-1.6, anterior

margin coarsely crcnulatc with about 15 lobes

(Fig. 36); secondary setae clear, appressed. incon¬

spicuous under dissecting microscope, those near

posterior angle of frontoclypcus predominantly

bi- or multi furcate (Fig. 37). Pronotum pale-

medium brown; secondary setae clear, appressed

and inconspicuous; primary seta 22 near antero¬

lateral margin of pronotum long, tapered, about

half length of pronotal sclerile. Posterior proster-

nites consisting of pair of small lateral scleritcs

and pair of large mesal scleritcs (Fig. 38).

Mesonolal and metanotal scleriles densely

clothed with appressed fine, forward-directed

dark setae, fonning short fringe along anterior

margin of both segments. Abdominal gills present

on segments 1-7.

Etymology. The species is named for Peter

Dostinc in recognition of his contribution to

knowledge of the aquatic insects of northem

Australia.

Comments. The larva is very similar to C

wellsae. The venter of the head capsule is

medium brown in C. dostinei as opposed to dark

brown, almost black in C. wellsae, and the

primary seta on the dorsum of the pronotum is

longer in C dostinei than in C. wellsae^ but these

NEW HYDROPSYCHID CADDISFLIES FROM NORTHERN AUSTRALIA

239

Figures 31-38 Chenmatopsyche dostinei sp. nov. Adult: 31, wings; 32, male genitalia, dorsal; 33, male genitalia,

lateral; 34, female genitalia, lateral; 35, female genitalia, dorsal. Larva: 36, frontoclypeus; 37, secondary setae near

apex of frontoclypeus; 38, prostemites.

240

J. C. DEAN

characters are not completely reliable. To sepa¬

rate the two species, setae near the posterior angle

of the frontoclypeus must be examined under high

magnification, C dostinei is widely distributed

across northern Australia (Fig.56). The Iar\'a has

previously been referred to as CIwumatopsyche

sp. 13 (Wells, 1991) and Cheximatopsyche sp.

AV13 (Dean, 1999).

Asmicridea capricornica sp. nov.

Figures 39-52

Type material. Hololypc: adult male. Graveside Creek,

Northern Territory (13° 18*S I32°32 E), 18Jul 1988, P.

Destine (NMV T-I7476). Paratypes collected with

holotype: 6 males. 6 females (NMV); 6 males, 3

females (ANIC).

Other material examined. Western Australia. IL.

King Edward River, Mitchell Plateau, 25 Sep 1995,

L.Metzeling; 3P. 14L. Manning Gorge, nr Gibb River

Rd. Kimberley. 28 Jul 1994, DC; 2P. 12L. Bell Creek

Gorge. Mt Hart Station, Kimberley, 26 Jul 1994. DC:

2L, Trib. Mitchell River, Mitchell Plateau. 19 Feb 1979,

JEB. Northern Territory. 4M, 41F. 149L, Gulungul

Creek, Radon Springs t'l2°45'S 132°55'E), various

dates, PS and AW, JD; 7P. 143L. Magella Creek, u/s

Magcla Falls (12°47'S 133°06'E), various dales. AW

and PS; 5P, 33L, East Alligator River, E of Magela

Falls (12°47'S 133°22'E), various dates, PS and AW;

IL, South Alligator River. C}imbal OSS station

(13°35 S i32°36'E), 20 Apr 1988. PS and AW; 3P,

64L, Baroalba Creek, Kubarra Pools. 12°49'S

132°52'E. various dates, AW and PS. DC. JD; IP, 3L,

Barramundie Creek, d/s Falls (13°22'S 132°28'E), 26

May 1988. AW and PS: IL, Liverpool River. Am-

hemland (I2°2rs 134°07'E), 26 Aug 1999, JD.

Queensland. 4L, Babinda Creek, The Boulders, 24

Nov 1979, DC; IL, Little Mulgrave River. 10 km SW

ofGordonvale, 16Nov 1988, K.Walker; 3L. Millstream

Creek, 1 Aug 1980, S.Bunn and Gray; 1L. Annan River.

30 km S of Cooktown. 20 Jun 1971, E.F.Riek.

Description. Adult. General colour pale. Wings

(Fig. 39): forewing whitish, with golden tinge

(female more intense than male), no obvious

colour pattern except for small area of medium

brown near base of wing around humeral.

crossvein; hindwing white, almost hyaline.

Length of forewing 5.8-6.9 mm (male), 6.1-7.3

mm (female). Vertex of head and thorax pale

golden colour; antennae straw coloured. Abdomi¬

nal slemile 5 with lateral process about half

length of segment. Male abdominal segments 6

and 7 with internal membranous sacs (Fig. 40).

Male genitalia (Figs 41^5): tergite X elongate,

deeply cleft, apices upturned; phallus elongate,

terminating in a pair of rounded cndothecal pro¬

cesses, and between these a pair of elongate phal-

lotrcmal scicrilcs, each sclerite with a strongly

acute apex. Female genitalia (Fig. 46): simple,

without obvious clasper receptors or pockets.

Pupa. Mandibles slender (Fig. 47); right

mandible with 3 subapical teeth, left mandible

with 4. Abdominal segment 3 with paired anterior

and posterior hook plates, segments 5-7 with

anterior hook plates only (Fig. 48), segment 4

without hook plates; some variation in numbers of

teeth on all plates. Apical processes of abdomen

with long dorsal projection at outer apical angle,

inner apical angle with 2 short projections (Fig,

49); numerous long black setae apically and also

along outer lateral margin.

Mature lamt. Head with distinctive pattern of

yellow and pale brown, dorsal surface covered

with numerous small pale dimples, particularly

conspicuous on darker pigmented areas (Fig. 50).

Head capsule broadest at or a little posterior to

midlength; lateral margins rounded; posterior

ventral apotome about third length of eedysial

line linking it with anterior ventral apotome.

Frontoclypeus length about two-lhirds head cap¬

sule length; anterior margin with deep notch on

left side, mesa! lobes projecting well forward of

base of notch. Foretrochantin not forked (Fig. 51).

Mesostemum and metastemum each with 2 gill

tufts. Abdominal segments 1 and 2 with trans¬

verse double row of stout appressed setae on dor¬

sal surface (Fig. 52).

Etymolog}’. The name recognises the broad distri¬

bution of this species across northern Australia in

latitudes north of the Tropic of Capricorn.

Comments. The structure of male genitalia within

the Australian genera Asmicridea and Smicrophy-

la.x is conservative and the gcnilalic stmeture of

A. capricornica is similar to species of both gen¬

era. A. capricornica can be distinguished from the

two described species of Asmicridea by the pos¬

session of paired membranous sacks within

abdominal segments 6 and 7. This character state

has previously been used to distinguish adult

males of Asmicridea and Sniicrophyla.x (Neboiss,

1977) and separation of adults becomes tenuous.

Fortunately, larval characters are of greater

diagnostic value and enable separation of the

genera and species of Asmicridea. A. capricor¬

nica is widely distributed across northern

Australia (Fig. 57), and while there is some over¬

lap the species tends to occur further upstream in

river systems than do species of the genus

Cheumatopsyche. The larva of A. capricornica

has previously been reported as Asmicridea sp.

AV 3 (Dean. 1999).

NEW HYDROPSYCHID CADDISFLIES FROM NORTHERN AUSTRALIA

241

Figures 39-46 Asmicridea capricornica sp. nov. Adult: 39, wings; 40, abdominal segments 5-10, lateral; 41, male

genitalia, dorsal; 42, male genitalia, lateral; 43-45, apex of phallus, dorsal, ventral and apical; 46, female genitalia,

lateral.

242

J. C. DEAN

Figures 47-52 Asmicridea capricornica sp. nov. Pupa: 47, mandibles; 48, abdominal hook plates (3, 5-7: abdomi¬

nal segment number, a: anterior hook plate, p: posterior hook plate); 49, apical process of abdomen. Larva: 50, head

capsule, dorsal; 51, foretrochantin; 52, dorsum of abdominal segment 1.

NEW HYDROPSYCHID CADDISFLIES FROM NORTHERN AUSTRALIA

243

Keys to species of Hydropsychidae of north-western Australia and the

Northern Territory

Adults

Abdominal stemite 5 with lateral processes (Fig. 40); posterior wing with Sc

and RI separate right to wing margin (Fig. 39). Asmicridea capricornica

Abdominal stemite 5 without lateral processes; posterior wing with Sc and

R1 fused prior to wing margin (Figs 1, 12,22,3!). Cheumatopsyche...!

Males.3

Females.6

Colour pale—medium brown; distal lobes of tergum X distinctly upturned in

lateral view (Figs 25, 33).4

Colour dark brown—black; distal lobes of tergum X rounded in both dorsal

and lateral views, not upturned (Figs 3, 14).5

Distal lobes of tergum X acute in both dorsal and lateral views (Figs 23-25)

,. Cheumatopsyche wellsae

Distal lobes of tergum X rounded in both dorsal and lateral views (Figs 32,

33) . Cheumatopsyche dostmei

Pre-anal appendages squat, located well anterior of base of distal lobes;

dorsum of segment IX broad in lateral view (Figs 2, 3)

. Cheumatopsyche kakadueusis

Pre-anal appendages elongate, located close to base of distal lobes; dorsum

of segment IX narrow in lateral view (Figs 13, 14). Cheumatopsyche suteri

Clasper receptacle in lateral view small, consisting of very short chimney

(Fig. 26); separation of receptacles in dorsal view about 8 times width of

each receptacle (Fig. 27). Cheumatopsyche wellsae

Clasper receptacle moderate to large (Figs 4, 15, 34); separation of

receptacles in dorsal view less than 3 times width of each receptacle (Figs 5,

16,35).7

Inner aperture of clasper receptacle aligned with outer aperture (Fig. 4)

. Cheumatopsyche kakaduensis

Inner aperture of clasper receptacle clearly dorsal to outer aperture (Figs 15,

34) .8

Outer margin of clasper receptacle semicircular; inner aperture large;

separation of receptacles in dorsal view less than or equal to width of each

receptacle (Figs 15, 16). Cheumatopsyche suteri

Outer margin of clasper receptacle oblique, orientated anteroventral to pos-

terodorsal; inner aperture small; separation of receptacles in dorsal view

about twice width of each receptacle (Figs 34, 35)..Cheumatopsyche dostinei

Mature larvae

Forctrochantin simple (Fig. 51); terga of abdominal segments 1 and 2 each

with 2 transverse rows of stout appressed setae (Fig. 52).

. Asmicridea capricornica

Forctrochantin forked (Fig. 21); terga of abdominal segments 1 and 2 with¬

out transverse row of setae... Cheumatopsyche....!

Anterior margin of frontoclypcus coarsely crcnulate, fewer than 20 lobes

(Figs 28, 36).3

Anterior margin of frontoclypeus finely crcnulate, more than 20 lobes (Figs

9, 17).4

Setae near apex of frontoclypeus predominantly simple (Fig. 29).

. Cheumatopsyche wellsae

Setae near apex of frontoclypeus predominantly divided (Fig. 37).

. Cheumatopsyche dostinei

Pronolum with primary seta 22 short, less than twice length of adjacent

secondary setae (Fig. 10). Cheumatopsyche kakaduensis

Pronotum with primary seta 22 long, about half length of pronotum (Fig. 19)

. Cheumatopsyche suteri

244

J. C. DEAN

Acknowledgments

The present study would not have been possible

without the extensive collection and rearing of

material from Kakadu National Park by Alice

Wells and Phil Suter. They are thanked for mak¬

ing the material available. Chris Humphrey, Peter

Destine, John Hawking and ERJSS provided sup¬

port and assistance during my visit to the region.

Ken Walker is thanked for providing access to

additional material held in Museum Victoria and

for preparing the distribution maps.

References

Dean, J.C., 1999. Preliminary keys for the identification

of Australian Trichoptera larvae of the Family

Hydropsychidac. Cooperative Research Centre for

Freshwater Ecology, Australia, Identification

Guide 22: 1-40.

McLachlan. R., 1871. On new forms, &c., of extra-

European Trichopterous insects. Journal of the

Linnean Society of London, Zoology 11: 98-141.

Neboiss, A., 1977. A taxonomic and zoogeographies

study of Tasmanian caddis-flies (Insecta: Tri-v

choptera). Memoirs of the National Museum of

Victoria 38: 1-208.

Nimmo, A.P., 1987. The adult Arctopsychidae an4

Hydropsychidac (Trichoptera) of Canada and adja^

cent United States. Quaestiones Entomological

23: 1-189.

Shefter, P.W. and Wiggins, G.B., 1986 A systematio

study of the nearctic lar\’ae of the Hydropsyche

morosa group (Trichoptera: Hydropsychidae),

Royal Ontario Mu.seum: Toronto. 94 pp. (Life Sci^

ences Miscellaneous Publications)

Wells, A., 1991. A guide to the caddisflics (Tri-v

choptera) of the Alligator Rivers region. Northern

Territory. Super\’ising Scientist for the Alligator

Rivers Region. Open File Record 84.

Wiggins, G.B., 1977. Larvae of the North Americart

caddisfiy genera (Trichoptera). University of

Toronto Press: Toronto, xi + 401 pp.

NEW HYDROPSYCHID CADDISFLIES FROM NORTHERN AUSTRALIA

245

Figures 53-55. Distribution of Hydropsychidae in northern Australia. 53, Cheumatopsyche kakaduensis sp. nov.

54, Cheumatopsyche suteri sp. nov.; 55, Cheumatopsyche wellsae sp. nov.

246

J. C. DEAN

Figures 56-57. Distribution of Hydropsychidae in Australia. 56, Cheumatopsyche dostinei sp. nov.; 57, AsmicricieC

capricornica sp. nov.

Memoirs of Museum Victoria 58(2): 247-254 (2001)

REDESCRIPTION OF BUNGONA MARKER WITH NEW SYNONYMS IN

THE AUSTRALIAN BAETIDAE (INSECTA: EPHEMEROPTERA)

P. J. SuTER AND M. J. Pearson

CRC for Freshwater Ecology, Department of Environmental Management and Ecology,

La Trobc University Albury-Wodonga Campus, PO Box 821,

Wodonga. Victoria 3689, Australia

p.suter(gaw.Iatrobe.edu.au

Abstract

Suter, P.J. and Pearson, M.J., 2001. Redcscription of Bungona Marker with new synonyms

in the Australian Baetidae (Insecta: Ephemeroptera). Memoirs of Museum Victoria 58(2):

247-254.

The monospecific genus Bungona Marker is redefined and descriptions of the imago and

nymphs of Bungona narilla Marker are provided. Two species described by Lugo-Ortiz and

McCafferty (1998) as Cloeodes fiistipalpus and C. illiesi are recognised as being conspecific

and junior synonyms of Bungona narilla. Bungona narilla is distributed along the east coast

of Australia from North Queensland to Tasmania and nymphs occur in shallow, slow Rowing,

cobble streams.

Introduction

Marker (1957) erected the genus Bungona and

Bungona narilla, type species, on the basis of a

nymph and adult from Coal and Candle Creek,

Sydney. The adult was distinguished from the

genus Pseudocloeon by the shape and number of

segments of the forceps and the nymphs were dis¬

tinguished by the shape of the labial palpi and the

maxillary palpi being 3-segmented. As the Aus¬

tralian baetid fauna became belter known through

the Monitoring River Health Initiative it was

clear that the original description of the nymph

was inadequate to distinguish Bungona from

other baetid genera. The British Museum of

Natural History has no record of type material

having been deposited in spite of assertions that

types were lodged (J. Marker, pers. comm). Dean

and Suter (1996) and Suter (1997) redefined the

genus in terms of the nymph, and noted that a

number of distinguishing characters were not

recorded by Marker (1957).

Direct association of an adult and nymph from

the Rose River in Victoria and adults and nymphs

from the New England area in New South Wales

has enabled verification that the nymphs

described by Dean and Suter (1996) and Suter

(1997) are conspecific with Bungona narilla

Marker.

Lugo-Ortiz and McCafTcrty (1998) recorded

the genus Cloeodes Traver, 1938 from Australia.

They described two new species C. fustipalpus

and C. illiesi, without citing any Australian

papers. The two species clearly belong in Bung¬

ona as redefined by Dean and Suter (1996) and

subsequently verified. Bungona has numerous

nymphal and imaginal characteristics which dis¬

tinguish it from Cloeodes as defined by Traver

(1938) and redefined by Waltz and McCafferty

(1987a, b). Character states of C. fustipalpus and

C illiesi arc within the variation of Bungona

narilla and both are treated as junior synonyms of

BimgofUi narilla.

Nymphs were collected using a hand-held 250

pm mesh, dip net held downstream of disturbed

substrate, or by hand picking of nymphs clinging

to the under-surface of rocks. Mature nymphs

were kept in IL plastic rearing containers which

had mesh lids and which contained river water

and a cobble. These were maintained in the

stream or in the laboratory until the subimago

emerged. The subimago was then removed and

placed in a dry rearing container to complete the

final moult. All specimens were preserved in

75% ethanol.

Nymphs were dissected and mounted on slides

in polyvinyl lacto-phenol mounting medium.

Illustrations were prepared with the aid of a

camera lucida, Mouthparts were viewed vcntrally

(except labnim) and the labium is illustrated with

the ventral surface shown on the right hand side

of the illustration and the dorsal surface on the

left. Comparative measurements of segments of

labial palpi, maxillary palpi and legs are

expressed as ratios compared with the proximal

247

248

P. J. SUTER AND M. PEARSON

segment length, which is given in parentheses.

Ranges of segment lengths are also presented in

parentheses. At least 20 specimens were used for

nymphal morphometric parameters. All measure¬

ments are given in millimetres.

The specimens examined were collected by

numerous people identified by initials, as follows;

AB (Andrew Boulton), BC (Bnicc Chessman)

DNR (Department of Natural Resources, Queens¬

land), DO (David Oldmeadows), JD (John Dean),

MN (Mark Nelson). MP (Melanie Pearson), PM

(Phil Mitchell), PS (Phil Sutcr), SB (Stuart Bunn)

and TC (Tim Cumii).

The ncotype is placed in the Australian

National Insect Collection (ANIC), CSIRO,

Canberra, and all other material is held in the

senior author’s collection.

Bnngona Hanker

Bungona Marker, 1957: 73.—Campbell, 1988: 9.—

Dean and Suter, 1996: 22.— Sutcr, 1997; 2.

Cloeodes.-Ux^o-OxWz and McCafferty 1998:

122-128. (not Cloeodes Traver, 1938).

Type species. Bungona narilla Marker, 1957

(original designation).

Diagnosis. Male Imago (Figs 1-5). Forewings

with paired marginal intcrcalaries from radial to

cubital sectors (Fig. 3); pterosligmatal veinlets

entire; base of vein MA2 attached to crossvein

between MAI and MPI. Hind wings absent.

Tarsal claws dissimilar. Posterior margin of

metanotum not deeply emarginate (Fig. 1); metas-

cutellar hump lacking a dorsoposterior projection

(Fig. 2). Forceps 4-segmentcd, segment 3 elon¬

gate, segment 4 longer than wide; segment 1 lack¬

ing long basal bristles; segment 2 with very short

bristles (Figs 4 and 5).

Mature Nymph (Figs 6-22): Head hypo-

gnalhous. Labrum slightly broader than long, with

a shallow median notch (Fig. 13). Left mandible

with incisors apically separate, proslheca robust

and digitate, margin between incisors and molars

lacking tuft of setae, thumb of molar area triangu¬

late and elevated above plane of incisor base (Figs

16 and 17). Right mandible with incisors separate

apically, prostheca bifid, margin between incisors

and molars with tuft of setae present (Figs 14 and

15). Maxillae palpi with 3 segments, palpi sub¬

equal to length of galeolacinia, galeolacinia with

4-5 apical teeth (Figs 18-20). Labium with 3-

segmented palpi, terminal segment short,

bulbous, medially broader than basal width, seg¬

ment 2 without inner apical lobe; glossae and

paraglossae equal in length (Fig. 22).

Thorax lacking hindwing pads. Femora lack a

femoral patch or villopore, with long robust blunt

setae on dorsal margin, apically with a pair of

contiguous setae (Fig. 6). Tibiae with subproxi-

mal arc of long fine setae, with a longitudinal row

of long fine setae on dorsal margin, adjoining

proximal arc (Fig. 6). Tarsi with a longitudinal

row of long fine setae on dorsal margin (Fig. 6).

Tarsal claw short, edentate (Fig. 7).

Abdominal terga with triangular, serrated,

scales which lack fine setae (Fig. 10), posterior

margins with triangular teeth (Fig. 1 1). Abdomi¬

nal stemites with scales and a row of short, fine

setae on segments 4-6; posterior margins with

long triangular teeth. Abdominal colour pattern

with tergites 9 and 10 dark. Gills asymmetrical,

ovate to pointed apically, with serrated and cili¬

ated inner margins (Figs 8 and 9). 3 caudal fila¬

ments; terminal filament shorter than cerci,

fringed with setae on lateral margins; cerci

fringed on inner margin.

Remarks. While both adults and nymphs of the

type species were described. Marker (1957) did

not record the edentate claws, tarsal and tibial

setal fringes, stcnial setae and other nymphal

characters which distinguish this genus from

other Australian bactids. The problem posed by

the absence of type material of the type species

has been overcome by successful rearing of

adult material. Comparison of adults with

Marker’s descriptions has confinned the identity

of Bungona narilla.

Waltz and McCafferty (1987a, b) revised

Cloeodes and closely related genera (Waltz and

McCafferty, 1987b). Lugo-Ortiz and McCafferty

(1998; 122) commented that Cloeodes had “eden¬

tate tarsal claws, a conspicuous arc of long, fine,

simple setae on the tibiae and setal tufts on stema

2-6.” They considered the two species they

described from Australia to belong in Cloeodes.

However, the Australian species possesses a row

of setae only on sterna 4-6, and can be clearly dis¬

tinguished from Cloeodes and other closely

related genera by the following combination of

adult and nymphal characters;

Male imago with pterostigmatic veinlets entire,

base of vein MA 2 attached to crossvein between

MA| and MP|. Mind wings absent. Tarsal claws

dissimilar. Posterior margin of metanotum not

deeply emarginate, mctascutcllar hump lacking a

dorsoposterior projection. Forceps four-

segmented, segment 3 elongate, segment 4 longer

than wide, segment 1 lacking long basal bristles,

segment 2 with very short bristles. Mature nymph

with incisors of left mandible apically separate.

THE EPHEMEROPTERAN GENUS BUNGONA

249

Right mandible with incisors apically separated,

prostheca billd, margin between incisors and

molars with tuft of setae present. Labial palpi

with tenninal segment short, bulbous, medially

broader than basal width, segment 2 without inner

apical lobe. Maxillae with 3 segmented palpi

which is subcqual to length of galeolacinia. Legs

with tibiae possessing proximal arc of long fine

setae adjoined to longitudinal row of long fine

setae and tarsi with a row of long ime setae.

Scales on abdominal tergites and stemites lacking

fine setae, row of fine setae on stemites 4-6. Gills

with serrated and ciliated margins.

Bimgona and Cloeodes arc closely related

genera, but there are at least three adult and nine

nymphal characters which clearly distinguish the

Australian material from the revised characterisa¬

tion Cloeodes given by Waltz and McCatTerty

(1987a, b). Lugo-Ortiz and McCafTerty (1998)

did not mention the distinctive characters of the

Australian material which here have been used to

support the maintenance of Bungona. The

"^Cloeodes group" is widely distributed in South

America (Waltz and McCafferty (1987a, b),

Africa (Waltz and McCalTerty , 1994), Madagas¬

car (Lugo-Ortiz and McCafferty, 1999), Sri

Lanka, China (Waltz and McCafferty, 1987a, b)

and Australia (Lugo-Ortiz and McCafferty, 1998;

this paper) and all share a number of charac¬

teristics but re-examination of this material

is now warranted to establish the phylogenetic

relationships.

Btingotui narilla Marker

Bungona narilla llarkcr, 1957: 73, figs 48-57.

Cloeodes fuslipalpus Lugo-Ortiz and McCafferty,

1998: 123-124, figs 1-9. (syn. nov.)

Cloeodes illiesi Lugo-Ortiz and McCafferty, 1998:

124^127, figs 10-18. (syn. nov.)

Material examined. Neotype herein selected. Adult

male from Gara R. at Thalgarrah Field Study Centre,

NSW, 30°26'S I5I°29'E, 25 Nov 1998, PS and JD,

ANIC.

Adults. Vic. I male reared from Rose R. RWC Gauge

Station, 35°52'S 146°03'E, 18 Jan 1997, PS and MP; 1

male, same locality, 4 Feb 1997. PS and MP. NSW. I

male reared from Gara R. at Thalgarrah Field Study

Centre, 30°26'S 15r29'E, 25 Nov 1998. PSandJD: 10

males. Commissioners Waters on Andersons Rd E of

Armidalc.3()°34S 151M7'E, 24 Nov 1998, PSandJD;

4 males Gara R. E of Amiidale on Amiidale to Coffs

Harbour Rd, 30°32'S I5I°48'E, 24 Nov 1998, PS and

JD.

Nymphs. Qld. 4 nymphs, Booloumba Ck, Conondalc

Ranges, 26“42'S ]52“38'E, 4 May 1993, SB; 2

nymphs, Sunday Ck site 5, 17*^55'S I45°09‘E, 30 May

1992, SB; 2 nymphs, Bundaroo Ck, Conondale Ranges,

26°42*S 152'’37'E, 20 May 1993, SB: 3 nymphs, Kool-

moon Ck near Tully, 17°45'S I45°38'E, 31 Jul 1990,

29 Nov 1990, SB; 6 nymphs. Stony Ck, Conondale

Ranges, 26°50'S 152°46'E, 15 Mar 1993, 15 Nov 1993,

SB; 6 nymphs, un-named Ck, Upper Conondale

Ranges, 26°52'S 152°44'E, 20 May 1993. SB; 5

nymphs, Mt Barney Ck at Mt Maroon, 28°I4'S

i52®44'E, 2 Nov 1998, DNR; I nymph. Little Yabba

Ck at Sunday Ck Rd, 26'’36'S 152°37'E, 18 May 1999.

DNR. NSW. 1 nymph, Kangaroo R., Upper Kangaroo

Valley, 34“42 S 150^35'E, 23 Sep 1972, JD; 1 nymph.

Wollondilly R. at Murphys Crossing, 33°43'S

150°30'E, 16 Nov 1990, BC; 4 nymphs, Bellinger R. at

Cool Ck, 30°27'S 152°37E. 23 Sep 1994, AB; 1

nymph, Imlay Ck/Wallaugh R. junction, 37°I4'S

149°42'E. 27 Oct 1995. AB; 2 nymphs, Collombatta Ck

(MACL02), 30^54'S 152®44'E, 18 Sep 1994, AB; 3

nymphs. Chandler R. at Carten, 30°44'S 152'^02'E, 5

Nov 1995, AB. 6 nymphs, Goorudee Rivulet. North of

Adaminaby, 35°59'E I48‘’46'E, 11 Mar 2000, PS and

TC; I nymph. Commissioners Waters on Andersons Rd

E of Annidale. 30“34'S 151°47'E, 24 Nov 1998, PS

and JD; 2 nymphs, Gara R.. E of Armidalc on Annidale

to Coffs Harbour Rd.30°32'S 151^48'E, 24 Nov 1998.

PS and JD; 10 nymphs, Woolomombi R. near Kilcoy

Cemetery, 30*^26'S 15P49'E,25 Nov 1998, PSandJD:

10 nymphs, Gara R. at Thalgarrah Field Study Centre,

30°26'S 15r29'E, 25 Nov 1998, PS and JD; 24

nymphs, Tilbuster Ck (Commissioners Waters),

30”29'S l51°42'E,25Nov 1998. PS and JD; 5 nymphs,

Dumaresq Ck on Weir Rd u/s Amiidale. 30°29'S

151°37'E, 25 Nov 1998, PS and JD; 28 nymphs, Gara

R. on Guyra-Dorrigo Rd., 30'’12'S 15r4'E, 25 Nov

1998, PS and JD; 9 nymphs, Marowan Ck near Glen¬

coe, 29'’56'S 151^43'E, 26 Nov 1998, PS and JD; 5

nymphs, Maybolc Ck, SW of Glen Innes. 29°53'S

l5t°38'E. 26 Nov 1998, PS and JD; 3 nymphs. Chan¬

dler R. on LynockRd, 30°18'S 152^05'E. 26 Nov 1998.

PS and JD; 7 nvmphs, Guy Fawkes R. at Ebor, 30°24'S

!52°20'E.27Nov 1998. PS and JD; 1 nymph, Bellinger

R., 23.7 km u/s of Thora, 30°28'S 152°35'E, 29 Nov

1998, PS and JD; 4 nymphs, Moncarlowe R. 1.8 km d/s

Monga Setllcment, 35°33'S 149°55'E, 1 Dec 1998, PS

and JD; 1 nymph, Trib. Mongarlowe R. at Monga

turnoff to Monga, 35®32'S 149°56'E. 1 Dec 1998, PS

and JD: 12 nymphs, Mongarlowe R. 100 m W of

turnoff to Monga, 35°32'S 149°56'E, 1 Dec 1998, PS

and JD; 3 nymphs, Shoalhaven R. at Bombay Bridge,

7.7 km W of Braidwood, 35®26'S I49°43'E, 1 Dec

1998, PS and JD. Vic. 2 nvmphs. King R. upstream of

Lake William Hovell, 36o56'S 146o27'E, 7 Jun 1990,

12 Nov 1991, PM; 10 nymph.s. Rose R. at “Bennies",

36°58'S MO'^Jl 'E, 6 Dec 1996, 14 Jan 1997, PS and

MP; 8 nymphs. Rose R. RWC Gauge Station, 35°52'S

146^03'E, 18 Jan 1997, PS and MP. Tas. 1 nymph,

Wilmot R. on Spellmans Rd. 4l°3rS 146°10'E. 13 Oct

1994, DO and MN; 3 nymphs. Rubicon R., Smiths Rd,

41°19'S 146°34'E, 6 Oct 1994, DO and MN.

Description. Male Imago. Body length: 4.6 mm.

Cerci length: 8.4 mm. Forewing length: 4.2 mm.

General colour brown with cream abdominal

250

P. J. SUTER AND M. PEARSON

markings, abdominal segments 1-2 with central

cream marking, 3 dark brown. 4 cream, 5-6 dark

brown, 7-10 light brown. Head: turbinate eyes on

high stalks, separate, reddish-brown dorsally

(Fig. I). Thorax brown; posterior margin of

metathorax with shallow cmarginalion (Fig. 1).

mela-sciitcllar hump lacking a posterior projec¬

tion (Fig. 2); legs cream, ratio of fore leg seg¬

ments 1.00: 1.05: O.OI: 0.58: 0.34: 0.20: 0.13 (I.O

mm); middle and hind leg measurements similar,

ratio of segment lengths 1.00: 0.74: 0.09: 0.18:

0.10: 0.08: 0.15 (0.72 mm). Wings hyaline (Fig.

3), plcrosligma with 4-5 cross veins, intercalaries

paired from R 7 - Cu vein, MA2 extending to and

beyond MA|~MP| crossvein. Forceps long, 4-

segmented; segment 1 and 2 subcqual. third seg¬

ment elongate 1.3 times as long as segment 2.

with slight constriction in basal half; segment 4

longer than wide, approx half segment 3 length

(Fig. 4); segment I lacking tufts of basal bristles,

segment 2 with few short basal bristles (Fig. 5).

Female Imago. F3ody length; 4.5 mm. Ccrci

length: 6,7 mm. Forewing length: 4.7 mm.

General colour brown with cream central mark¬

ings on abdominal segments. Head: eyes not

turbinate, smaller, placed laterally. Legs all

similar in length. Wings hyaline, similar to male.

Mature nymph, (all measurements based on 21

mature specimens). Body length: 3.9 mm (3.0-6.2

mm). Ccrci length: 1.7 mm (1.1-2.1 mm). Ter¬

minal filament; 1.5 mm (1.0-1.9 mm). Antennal

length: 0.9 mm (0.8-1.1 mm). Females generally

larger than males. The description given by Lugo-

Ortiz and McCafferty (1998) for the nymph is

adequate and generally well illustrated. Additions

only arc given below.

Thorax: Fore legs (Fig. 6 ) with upper margins

of femora with 4-9 blunt simple setae, apically

with a pair of contiguous setae, lower margin with

numerous short, fringed setae; fore tibiae with

proximal arc of long fine setae and a longitudinal

row of long fine setae, inner margin with 2-14

short fine setae, apically with pair of short fringed

setae; tarsi with longitudinal row of long fine

setae, inner margin with 4-10 short fringed setae

with 2 apical setae; tarsal claw short lacking teeth

(Fig. 7). Middle and hind legs similar, with outer

margins of femora lined with 5 10 blunt setae,

apically with pair of contiguous setae; tibiae with

proximal arc of long fine setae and longitudinal

row of long fine setae, inner margin with 4-10

short fine setae, apically with pair of short fringed

setae; tarsi with longitudinal row of long fine

setae, inner margin with 5-10 short fringed

setae with 2 apical setae; tarsal claw short lacking

teeth.

Ratios of leg segments: fore leg. 1.00: 0.62:

0.67 (0.72 mm); middle leg, 1.00: 0.60: 0.55 (0.72

mm); hind leg, l.OO: 0.56: 0.48 (0.75 mm).

Femur length to width ratios: fore leg, 4.83

(3.55-6.62); middle leg, 4.92 (3.79-5.92); hind

leg, 4.62 (3.65-6.54).

Abdominal segments 1-7 with plate-Hke gills

with serrated and ciliated inner margins (Figs 8 b,

9b), first, fifth-seventh gills narrow apically

pointed (Fig. 8 a), second-fourth gills ovate (Fig.

9a), rounded apically (shape of gills variable

within populations from narrow-pointed to

broadly ovate). Abdominal colour pattern vari¬

able but with ninelh and tenth segments brown.

Scales on tergites and stemites triangular and ser¬

rated (Fig. 10), posterior margins of tergites with

triangular spines (Fig. 11 ). Paraprocls with 10-18

well developed sharf> marginal spines (Fig. 12),

number varies with instar.

Mouthparts. Labrum (Fig. 13) almost square

length 0.8 width, anterior margin with a medial

concavity, anterior margin fringed with short

bifid setae, submarginal row of spine setae. Right

mandible (Fig. 14) outer incisor with 3-4 teeth

and small subapical tooth, inner incisor with 3

teeth, prostheca bifid (Fig. 15) with lateral section

strongly serrated (damaged in specimen described

and illustrated as Fig. 3 by Lugo-Ortiz and

McCafferty, 1998), margin between incisors and

molars smooth or serrated with tuft of setae near

molars. Left mandible (Fig. 16) outer incisor with

2-4 teeth, inner incisor with 3 large teeth, pros-

ihcca robust (Fig. 17), margin between incisors

and molars smooth or serrated, lacking tuft of

setae. Maxilla rectangular (Fig. 18) with 4 apical

teeth (Fig. 19), maxillary palpi 3-scgmentcd,

palpi length subcqual to galeolacinia (Fig. 20),

apical segment longer than basal segment: seg¬

ment ratios l.OO: 0.66; 1.14 (0.07 mm. 0.04-0.09

mm). Hypopharynx as in Fig. 21. Labium (Fig.

22 ) with glossae and paraglossae long, narrow

and pointed, labial palpi with 3 segments, apical

segment ovoid, basal segment 1.73 times as long

as broad, ratio of segments 1.00 : 0.87: 0.56 (0.15

mm. 0.12-0.18 mm).

RemarLs.Uo new material has been found from

the type locality but nymphs consistent with

Harker's description have been recorded from

streams of the Great Dividing Range in Queens¬

land, New South Wales, Victoria and one speci¬

men from Tasmania. Adults reared from the Rose

R. in Victoria confirmed its identity, Lugo-Ortiz

and McCafferty (1998) described Clacodes fusti-

palpus from nymphs collected from the Chandler

and Bellinger Rivers near Armidalc in New South

THE EPHEMEROPTERAN GENUS BUNGONA

251

Wales. This species cannot be differentiated from

B. narilla, and is here synonymised.

Lugo-Ortiz and McCafferty (1998) differenti¬

ated a second species (Cloeodes illiesi) from a

single nymph from Cascade Falls near Cairns on

the basis of abdominal colour pattern, bifid right

prosthcca and a reduced maxillary palp. They

stated that the colour pattern of C.fustipalpus ..

varies somewhat among specimens ...” . The

pattern illustrated for C. illiesi is within the range

of variation oi' B. narilla. The bifid prosthcca is

also consistent with B. narilla. The specimen they

illustrated and described as C. fustipalpus had a

broken prosthcca (subsequently confirmed by

McCafferty, pers. comm.). The maxillary palpi in

C. illiesi may also be an aberration, as at least one

specimen examined in this study had a short.

2 -segmcnled maxillary palp on one side of the

body, and a long, 3-scgmentcd palp on the other.

Such an aberration has also been observed in

specimens of Cloeon in northern Australia.

Although not specifically mentioned as distin¬

guishing features in their paper, Lugo-Ortiz and

McCafferty (1998) noted that the margin between

the incisors and molars of the right mandible of

C illiesi differed from C. fustipalpus in that it was

smooth, not serrated, the gills were narrow and

less tracheated and the paraprocts had few scale

bases. In the extensive material available for this

study this combination of characters was not con¬

sistent even within a single population. For

example, specimens with a smooth mandibular

margin had broad, strongly tracheated gills and

numerous paraproct scale bases, and specimens

with narrow, poorly tracheated gills had numer¬

ous paraproct scales and serrated madibular mar¬

gins. Early instar nymphs of {\\q fustipalpus type

had more pointed gills than older nymphs in the

same population suggesting this character

may be influenced by age and environment.

Cloeodes illiesi is considered a junior synonym of

B. narilla.

Distribution and ecology’. North Queensland to

Tasmania. Nymphs have been collected from

streams of low gradient and low turbidity, mainly

in foothill and lowland reaches. Large numbers

have been recorded from shallow water (less than

20 cm) at the edge of cobble streams where veloc¬

ities were very low. In the Conondale Ranges,

southern Queensland, specimens were collected

from bedrock in shallow, still, edge habitat. They

were observed on the upper surface of the rock

during the day and when disturbed only swam a

few centimetres and then settled. Adults were not

collected with UV light traps, and few were

collected by sweep netting the riparian vegeta¬

tion. A single specimen was collected from a

swarm ofbaetid males at a height of 8-10m above

the Rose River.

Acknowledgements

The financial support from the Land and Water

Resources Research and Development Corpora¬

tion under the National Healthy Rivers Program

and the Australian Biological Resources Study is

gratefully acknowledged. Thanks to all who have

provided us with material particularly Dr Stuart

Bunn, John Dean, Phil Mitchell, David Oldmcad-

ows, Mark Nelson, Dr Andrew Boulton, Dianne

Conrick, Dr Satish Choy and Dr Richard

Marchant. Special thanks to John Dean for com¬

ments on an earlier draft.

References

Campbell, I.C., 1988. Ephemcroplera. Zoological

Catalogue of Australia 6; 1-22. Australian

Govemmeni Publishing Service: Canberra

Dean, J. C. and Suter, P.J., 19%. Mayily nymphs of

Australia; A guide to genera. CRC for Freshwater

Ecology Identification Guide 7: 1 82,

Harker, J. E., 1957. Some new Australian

Ephemeroptera. Proceedings of the Royal Ento¬

mological Society of London (B) 26: 63-78.

Lugo-Ortiz. C. R. and McCafferty, W. P., 1998. First

report and new species of the genus Cloeodes

(Ephemeroptera: Baetidae) from Australia.

Entomological News 109(2): 122-128.

Lugo-Ortiz. C. R. and McCaffeity, W. P., 1999. The

small minnow mayfly genus Cloeodes Traver

(Ephemeroptera: Baetidae) in Madagascar.

Proceedings of the Entomological Society of

IVashington 101 (I): 208-211.

Suter. P. J., 1997. Preliminary guide to the identifica¬

tion of nymphs of Australian baetid Mayflies

(Insecta : Ephemeroptera) found in Rowing waters.

CRC for Freshwater Ecology Identiflcation Guide

14: 1-36.

Traver, J. R.. 1938. Mayflies of Puerto Rico. Journal of

Agriculture of the University of Puerto Rico 22:

5 ^ 2 .

Waltz, R. D. and McCafferty, W. P., 1987a. Revision of

the genus Cloeodes Traver (Ephemeroptera:

Baetidae). Annals of the Entomological Society of

America 191-207.

Waltz, R. D. and McCafferty. W. P., 1987b. Generic

revision of Cloeodes and description of two new

genera (Ephemeroptera; Baetidae). Proceedings of

the Entomological Society of Washington 89(1):

177-184.

Waltz. R.D. and McCafferty, W.P., 1994.. Cloeodes

(Ephemeroptera: Baetidae) in Africa. Aquatic

Insects 16(3): 165 169.

Figures 1-5. Bungona narilla, male imago. 1, dorsal view of head and thorax; 2, lateral view of head and thorax; 3.

forewing; 4, ventral view of male forceps; 5, enlarged view of right forcep. Scale lines: 1 mm (Figs 1-3); 0.1 mm

(Figs 4-5).

THE EPHEMEROPTERAN GENUS BUNGONA

253

Figures 6-12. Btmgona narilla, nymph. 6, foreleg: 7, tarsal claw; 8, sixth abdominal gill, (8a, whole gill; 8b, lateral

margin of gill); 9, third abdominal gill, ( 9a, whole gill; 9b, lateral margin of gill); 10, scales on tergites; 11, poste¬

rior margin of fourth abdominal tergite; 12, paraproct. Scale lines: 0.1 mm.

254

P. J. SUTER AND M. PEARSON

Figures 13-22. Bungona narilla, nymph. 13, labrum; 14, right mandible: 15, incisors and prostheca of right

mandible; 16, left mandible; 17, incisors and prostheca of left mandible; 18, maxilla; 19, apex of maxilla; 20, max¬

illary palp; 21, hypopharynx; 22, labium, dorsal (left) and ventral (right) aspects. Scale lines: 0.1 mm.

Memoirs of Museum Victoria 58(2): 255-295 (2001)

DESCRIPTIONS OF NEW SPECIES OF BIRUBIVS (AMPHIPODA:

PHOXOCEPHALIDAE) FROM AUSTRALIA AND PAPUA NEW GUINEA WITH

COMMENTS ON THE BIRUBIUS-KULGAPHOXUS-TICKALERUS-YANCOUVLEX

Joanne Taylor’’^ and Gary C. B. Poore*

'Museum Victoria, GPO Box 666E, Melbourne, Victoria 3001, Australia

•Department of Zoology, University of Melbourne, Parkville, Victoria 3010, Australia

(jtaylor@museum.vic.gov.au and gpoorc@museum.vic.gov.au)

Abstract

Taylor, J. and Poore, G.C.B., 2001. Descriptions of new species Binihius (Amphipoda:

Phoxocephalidae) from Australia and Papua New Guinea with comments on the Binihius-Kul-

gaphoxtis-Tickalerus-Yan complex. Memoirs of Museum Victoria 58(2): 255-295.

Five new species of Binthius Barnard and Drummond, 1976 (Crustacea: Amphipoda: Phox-

occphalidac) are reported: B. drummomiae sp. nov and B. heislersi sp. nov from Victoria, Aus¬

tralia: B. w'ullisae sp. nov. from Queensland, Australia; and B. lowryi sp. nov., and B. wilsoni

sp. nov. from Papua New Guinea. The present records extend the range of the genus previously

reported from Australia and Indonesia. The genus Birubius is discussed and compared with the

Australian genera Tickalcrus Barnard and Drummond, 1978 (monotypic), Kuigaphoxus

Barnard and Drummond, 1978 (two species) and Van Barnard and Dmmmond, 1978 (two

species) in light of the new species exhibiting a combination of characters from all genera.

Their synonymy is foreshadowed.

Introduction

Barnard and Drummond (1978) discussed the

relationships between all Australian species of

the Phoxocephalidae. The subfamily Birubiinac

was established to include the Australian type

genus Birubius Barnard and Drummond, 1976,

Australian genera Tickalerus Barnard and

Drummond, 1978, Kuigaphoxus Barnard and

Dnimmond. 1978 and Van Barnard and

Drummond, 1978 and the North and South

American genera Microphoxus J.L. Barnard,

1960 and Metharpinia Schel Icnberg, 1931.

Barnard and Karaman (1991) expanded the sub¬

family to include the North and South American

genera Foxiphalus J.L. Barnard, 1979 and

Grandiphoxus J.L. Barnard, 1979. The genus

Linca Alonso de Pina, 1993 was erected based on

a single specimen from the Argentine continental

shelf and although showing some convergence

with the Brolginae, its similarity to Birubius best

placed it in the Birubiinac. Linca dilTers from

Birubius by the presence of ventral setae on

uropod I peduncle, an aulapomorphy of this

monotypic genus. Jarrett and Bousficid (1994)

reassessed the North and South American genera

and removed them from the Birubiinac. They

erected the new subfamily Mclharpiniinae to

include Beringiaphoxus Jarrett and Bousficid.

1994, Foxiphalus J.L. Barnard, 1979, Grandi¬

phoxus J.L. Barnard, 1979, Majoxiphalus Jarrett

and Bousficid, 1994, Metharpinia Schellcnberg,

1931, Microphoxus J.L. Barnard, 1960 and Rhep~

oxynius J.L. Barnard, 1979. The genus Linca was

not included in the new subfamily and so remains

a member of the Birubiinac.

Our preliminary cladistic analysis (work in

progress) of most species of Birubiinac, Brol¬

ginae, Leongathinac, Metharpiniinae, Parhar-

piniinac and Tipimeginae has failed to support

the monophyly of any of the subfamilies or

genera. We arc unable to identify a synapo-

morphy for the subfamily Biaibiinae, even in the

restricted sense, nor for its type genus Birubius.

The Birubiinac shares a broad form of the basis of

pereopod 5 with all subfamilies except Harpini-

inae which exhibit a narrow basis unique to that

subfiimily. It shares a biarticulatc palp of maxilla

1 with all subfamilies except Phoxocephalinae

and those members of the Harpiniinac that exhibit

a uniarticulate palp, it differs from Brolginae,

Harpiniinac, Phoxocephalinae and Ponthar-

piniinac by the medium to elongate (rather than

short) length of peduncular article 2 of antenna 1,

a state that it also shares with Tipimeginae,

Parharpiniinae and some members of the

Joubincllinac.

Examination of unidentified phoxocephalid

amphipods from the Australian Museum,

Museum Victoria and Queensland Museum col¬

lections revealed five new species belonging to

255

256

J. TAYLOR AND C. B. POORE

the nominal subfamily Birubiinae. The generic

placement of some of the species was uncertain

using Barnard and Karaman's (1991) generic

diagnoses. Although closely fitting the descrip¬

tion for Binihius four species exhibited a large

dorsal hook on urosomite 3, a character restricted

within the Birubiinae to members of Tickalents

and Kulgaphoxus.

Barnard and Drummond (1978) defined

Kulgaphoxus, Tickalerus and Van only on the basis

of differences from Birubius, the largest genus.

Tickalerus differs from Bintbius in the presence

of a dorsal hook on urosomite 3, shortened outer

ramus of uropod 3 and rectangular coxa 4. Kul-

gaphoxus differs from Biruhiiis in the presence of

a dorsal hook on urosomite 3, shortened outer

ramus of uropod 3, proximal placement of setae

on peduncular article 2 of antenna 1 and the ves¬

tigial dactyl of pereopod 7. Yan ditTers from Bint¬

bius in the proximal placement of setae on pedun¬

cular article 2 of antenna 1 and the vestigial dactyl

of pereopod 7. The new species share some but

not all of the diagnostic features of Kulgaphoxus,

Tickalents and Yan and could not be placed in any

of the genera as presently diagnosed.

The discovery of four species exhibiting a

dorsal hook on urosomite 3 is significant. Pre¬

viously it was a trait obscr\'ed in only five phoxo-

cephalid species belonging to Kulgaphoxus^

Microphoxus and Tickalerus and was partly used

to split these species from Bintbius. The trait was

fonnerly believed to be sexually dimorphic and

restricted to females with males having a reduced

hump at best. Males of the new species B. drum-

momiae sp. nov. and B. wallisae sp, nov. however

exhibit a w'eil developed dorsal hook as in

females. It appears that sexual dimoqihism is

variable but the possibility that males with¬

out hooks belong to other species is a remote

possibility.

Barnard and Drummond did not use cladistic

methodology to define genera. Rather, small

genera were picked off from larger clusters on the

basis of few differences that may or may not be

unique synapomorphies. The inevitable conse¬

quence of this is that the large genus, Bintbius in

this case, is paraphylctic because its numerous

species lack a synapomorphy. We are forced to

conclude that either (a) the small genera.

Kitlgaphoxus^ Linca, Tickalents and Yan, as

presently constituted arc gradal offshoots of

Birubiinae which cannot be supported in a classi¬

fication based on cladistic principles; or (b) the

type species of the four genera represent much

larger clades which may be redefined using very

different character suites (synapomorphies).

Until the cladistic analysis is completed we are

reluctant to complicate the taxonomy further by

erecting new small genera simply because they do

not comply with existing diagnoses. It is unclear

whether the minor genera in question will come to

encompass larger clades but it is certain that they

cannot be justified as currently defined. Therefore

in this contribution we describe the new laxa as

members of Bintbius in spite of their similar¬

ities to some members of Kulgaphoxits.

Tickalents and Yan. The synonymy of tlicse gen¬

era with Bintbius is foreshadowed. Our revised

diagnosis of Bintbius is written to include all

species included in Bintbius, Kulgaphoxus,

Tickalerus and Yan.

Abbreviations arc; A, antenna; H, head; rLM,

right lacinia mobilis; MD, mandible; MX,

maxilla; MP, maxilliped; GN. gnathopod; P, pere¬

opod; EP, epimeron; U, uropod; PL, pleopod; T.

telson; r, right; m, male; tl., total length; MAFRI.

Marine and Freshwater Resources Institute,

Quccnscliff; NMV, Museum Victoria, Mel¬

bourne; AM, Australian Museum, Sydney, QM,

Queensland Museum, Brisbane. All dissections

and illustrations follow the methods of Barnard

and Drummond (1978) whereby the Icfi side of

the animal is illustrated unless olhenvise stated.

Descriptions of the new species closely follow

that of other species of the genus described in

Barnard and Drummond (1978).

Biruhius Barnard and Drummond

Biruhiu.*{ Barnard and Drummond, 1976: 543.—

Barnard and Drummond, 1978: 191.—Barnard and

Karaman, 1991: 635.

Type species. Bintbius pananiunus Bamard and

Drummond. 1976 (by original designation).

Diagnosis. Rostrum variably constricted. Eyes

present. Antenna I peduncular article 2 length

variable, ventral setae not confined apically.

Antenna 2 peduncular article 1 not or scarcely

ensiform. article 3 with 2 facial setules, facial

robust setae on article 4 in 2+ rows, all robust

setae thick, article 5 ordinary. Right mandibular

incisor with 3-4+ teeth, right lacinia mobilis bifid

or simple, often fiabellate or absent, molar not

triturativc, with 4+ splayed robust setae; palpar

hump small to medium, apex of palp article

3 oblique. Maxilla I inner plate with 3-4 setae,

palp 2-arliculate. Maxillipedal plates small to

ordinary, apex of palp article 3 not strongly

protuberant, dactyl elongate, apical nail distinct.

Gnathopods small, similar, gnathopods 1-2

carpus length medium to elongate, not cryptic

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

257

(posterior margin not concealed by the abutment

of propodus and merus), palms oblique, gnat-

hopods 1-2 propodus ordinary to natrow, ovate to

rectangular, poorly setiferous anteriorly. Pere-

opods 3-4 carpus with (rarely without) postero-

proximal robust setae, propodus with robust setae.

Pereopod 5 basis of broad form (basis equal to or

greater than twice width of ischium), pereopods

5-6 menis-carpus broad to narrow: pereopod 7

unreduced, article 3 not enlarged, dactyl well

developed, vestigial or absent.

Epimera I -2 with or without long facial

brushes of setae, without posterior setae,

epimeron 3 bearing long setae. Urosomite 3 with

or without dorsal hook in females, sometimes in

male only if in female. Uropod 1 peduncle with¬

out intcrramal robust setae, without major dis¬

placed robust seta (seta that is shifted onto the

apical margin disjunclly from the true inner mar¬

gin), uropods 1-2 rami occasionally continuously

setose to apex (thus with minute apical robust

setae or nails), uropod 1 inner ramus with I row

of marginal robust setae, Uropod 2 inner ramus

ordinary to shortened. Uropod 3 variable, cither

unreduced (outer ramus longer than peduncle), or

reduced (outer ramus shorter than or subcqual to

peduncle), bearing a second article on outer

ramus, with 2 long apical setae, Telson ordinary

to elongate.

Species. Biribius batei (Haswell, 1879); B. ros-

fratus (Dana, 1853) = B. barnardi Pirlot, 1932.

Species described by Barnard and Drummond,

1978: B. apari; B. hahaneekus; B. booleiis:

B. cartoo; B. chintoo; B. eake: B. eleebanus:

B. gallcmgus; B. gambodeni: B. gelarus; B. jir~

randits; B. kahbulimts: B. kareus: B. karobrani:

B. kinkiis: B. kokoriis; B. kyeemits; B. lorus;

B. knvamnis: B. maamus; B. maldits; B. rnaya-

mayi: B. nnddarpus: B. mimggai: B. myaflus:

B. ' nammiddits; B. narus: B. panamumts:

B. qiiearus: B. teddeus: B. tfudmus; B. idaritiis:

B. wirakiis; B. wvlgam; B. yandus; B. yorhmus.

Species added after 1978; B. hali Ortiz and Lalana,

1999; B. drummondae sp. nov.; B. heislersi sp.

nov.; B. lowty'i sp. nov.; B. murariui Ortiz and

Lalana, 1997; B. walUsae sp. nov.; B. wilsoni

sp. nov.

Habitat and distribution. Marine 0-70 m. Aus¬

tralia; Indonesia; Papua New Guinea.

Remarks. Barnard and Karaman's (1991)

generic diagnosis has been altered to accommo¬

date the new species and to rellect the fore¬

shadowed synonymy of Tickalenis, Kidgaphoxus

and Yan.

Birubius drummondae sp. nov.

Figures 1-6

Material examined. Holotype. Australia, Victoria,

Western Port (38°22'S, 145‘’32'E) no further data,

NMV J47227 (1 female, tl. 3.8 mm).

Allotype. Same locality as holotype, NMV J47228 (1

male, tl. 5.25 mm).

Paralypes. Same locality as holotype, NMV J47226

(27 females, tl. 3.0-5.7 mm).

Diagnosis. Rostrum constricted. Antenna 2, arti¬

cle 4 without well developed dorsal setalion.

Right lacinia mobilis bifid, distal branch denticu¬

late. Pereopods 3-4 carpus with 2-3 proximopos-

terior robust setae. Pereopod 5 dactyl fully

formed. Pereopod 7 basis without long ventral

setae. Coxa 1 not expanded distally. Coxa 4 lack¬

ing long ventral setae. Epimeron 3 without large

tooth; without ventral setae; without long poste¬

rior seta; with oblique row of facial setae. Uro-

somile 3 with large dorsal hook. Uropod I with¬

out basofacial setae. Uropods 1-2 inner rami

lacking accessoiy apical nails. Uropod 3 unre¬

duced, outer ramus longer than peduncle.

Description of female. Head about 18% of total

body length, greatest width about 100% of length;

rostrum constricted, exceeding apex of pedunc¬

ular article 1 on antenna I. Eyes medium, clear of

pigment. Antenna 1 peduncular article 1 about 1.3

times as long as wide, about 1.8 limes as wide as

peduncular article 2, ventral margin with 4

setulcs, produced dorsal apex with 1 setule;

peduncular article 2 about 0.8 times as long as

peduncular article I, with 5 ventral setae; primary

flagellum with 10 articles, about 0.9 times as long

as peduncle, bearing long aesthetascs; accessory

flagellum with 8 articles. Antenna 2, peduncular

article 4 robust setae formula = I-3-4-3, dorsal

margin with notch bearing 3 setae, ventral margin

with 4-5 groups of 1-2 long to short setae, 1 long

ventrodistal robust seta; peduncular article 5

about 0.8 times as long as peduncular article 4,

facial robust seta fomiula = 1-2, dorsal margin

naked, ventral margin with 3 sets of 1-2 long to

short setae, without ventrodistal robust setae;

flagellum 1.51 limes as long as peduncular arti¬

cles 4-5 combined, with 11 articles. Mandibles

with medium palpar hump; right incisor with 4

teeth and notch; leff incisor with 2 humps in 2

branches; right lacinia mobilis bifid, distal branch

much shorter than proximal branch, denticulate,

proximal branch simple, pointed, with facial

hump; left lacinia mobilis sub-bifld; right raker 7;

left rakers 7 plus 1 rudimentary; molar in form of

bulbous hump, right molar with 4 long robust

setae, plus 1 short robust seta strongly disjunct,

258

J. TAYLOR AND C. B. POORE

Figure I. Birubius dnunmondae sp. nov., holotype female, tl. 3.80 mm (m = male allotype, 5.25 mm).

left molar with 5 long robust setae, plus ! short

robust seta strongly disjunct; palp article 1

slightly elongate, article 2 with 1 medium inner

apical seta and 2 other shorter inner setae, article

3 about equal in length to article 2, apex oblique

with 6 robust to slender setae, with 3 basofacial

setae. Maxilla I inner plate narrow, bearing 1

long apical seta, 1 similar apicomedial seta, 2 api-

colateral much shorter seta; palp article 2 with I

apicomedial marginal robust seta, 3 apicomedial

setae and 3 submarginal setae. Maxilla 2 inner

and outer plates extending equally, outer not

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

259

broader than inner. Maxilliped inner plates with 2

large thick apical robust seta, 3 apicofaciai setae,

5 medial setae; outer plate with 7 medial and api¬

cal robust setae, 1 apicolateral seta; palp articles

1-2 with 1 apicolateral seta, article 3 weakly pro¬

tuberant, with 3 facial setae, 1 lateral seta, nail of

article 4 medium length, with 1 accessory sctulc.

Coxa 1 not expanded distally; main ventral setae

of coxae 1-4 = 5-5-5-0, posteriormost seta of

coxae 1-3 shortened; anterior and posterior mar¬

gins of coxa 4 parallel, posterior margin straight,

posterodorsal comer sharp, posterodorsal margin

medium, width-length ratio of coxa 4 almost =

36:53. Long posterior setae on basis of

gnathopods 1-2 and pereopods 3-4 = 4-1-5-7,

short posteriors = 1 -0-1 -0, long antcriors = 4-8-0-

0 , short anteriors = 2-2-0-0.

Gnathopods, width ratios of carpus-propodus

on gnathopods 1-2 = 20:27 and 20:27, length

ratios = 21:26 and 1:1; palmar humps ordinary,

palms oblique; gnalhopod 1 carpus of medium

length gnathopod 2 carpus slightly elongate. Pere¬

opods 3-4 similar, facial setae on merus = 3 and

3, on carpus = 3 and 4; main spine of carpus

260

J. TAYLOR AND C. B. POORE

extending to M. 77 on propodus, carpus with 2

and 3 proximoposterior robust setae; robust setae

formula of propodus = 4 + 5 and 4 + 5; acclivity

on inner margin of dactyls of pcreopods 3-4

weak, midfacial seta ordinary. Coxae 5-7 pos-

teroventral seta formula = 2-2-1; merus-carpus of

pcreopods 5-6 broad, facial robust setae rows

dense, facial ridge formula on basis of pcreopods

5-7 = 0-2-2, anterior ridge of pereopod 7 long;

width ratios of basis, merus, carpus, propodus of

pereopod 5 = 11:12:11:5, of pereopod 6 =

18:13:10:5, of pereopod 7 = 107:27:23:11, length

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

261

Figure 4. Biruhius drummondae sp. nov., holotype female, tl. 3.80 mm (m = male allotype, 5.25 mm).

262

J. TAYLOR AND C. B. POORE

ratios of pereopod 5 = 45:21:24:22, of pereopod 6

= 58:32:24:29, of pereopod 7 = 63:15:13:13; basis

of pereopod 7 exceeding apex of merus, naked

venirally. Pleopods 1-3 with 2 coupling hooks;

rear facial setae on peduncle = 2-2-3; articles on

outer rami = 11-10-11, inner rami = 7-6-6.

Epimeron 1 posteroventra! comer rounded,

anteroventral margin with 3 setae, posteroventral

face with 2 medium setae, set vertically;

epimeron 2 posteroventral comer rounded, with 5

facial setae, posterioimost pair set almost verti¬

cally; epimeron 3 posteroventral comer barely

protuberant, with setule sinus, posterior margin

almost straight, with 2 setule notches, ventral

margin naked, midfacc with oblique row of 4

setae near posterior margin. Urosomite I naked,

articulation line almost complete; urosomite 3

with large hook dorsally. Uropods 1-2 rami with

articulate enlarged apical nails, uropod 1 outer

ramus with 3 dorsal robust setae, inner with 1

dorsomedial and I subapical robust setae, uropod

2 outer ramus with 2 dorsal robust setae, inner

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

263

Figure 6. Birubius drummondae sp. nov., holotype female, tl. 3.80 mm (m = male allotype, 5.25 mm).

with 1 broad dorsomcdial robust seta; iiropod I

peduncle w ith 3 apicolateral robust setae, without

basofacial slender setae, wath apical enlarged

robust seta; uropod 2 peduncle with 5 dorsal

robust setae; apicolateral corners of peduncles on

uropods 1-2 without comb. Uropod 3 unreduced,

outer ramus longer than peduncle. Uropod 3

peduncle with 6 ventral robust setae, dorsally

with 1 lateral robust seta; rami masculine, inner

extending to M. 100+ on article I of outer ramus,

apex with 3 setae, medial and lateral margins

naked, article 2 of outer ramus elongate, 0.38,

bearing 2 long setae, apicomedial margin of

article I naked, lateral margin with I acclivity,

robust setal formula = 2-2, without slender setae

formula. Telson long, length-width ratio = 34:29,

not fully cleft, each apex wide, rounded, lateral

acclivity broad, shallow, bearing ordinary lateral

setule, robust setae next medial little longer than

setule, midlatcral selules diverse.

Description of male. Similar to female but eyes

larger. Antenna I like female but with dense

medial setation on peduncular article 1; primary

flagellum bearing calceoli. Antenna 2 elongate,

peduncular articles 3-4 with dense dorsal seta-

264

J. TAYLOR AND C. B. POORE

tion, peduncular article 5 about as long as article

4, dorsal margin bearing 2 calceoli and 3 groups

of male setae, flagellum 28-articulate bearing cal-

ccoli. Maxiliiped and maxillae 1-2 similar to

female. Right lacinia mobilis bifid, distal branch

much shorter than proximal branch, flabcllate,

proximal branch simple, pointed, with facial

humps; left lacinia mobilis with 4 teeth; right

raker 8; left rakers 9; right and left molars with 5

long robust setae; palp similar to female, article 3

with 6 basofacial setae. Main ventral setae of

coxae 1-4 = 7-6-7-0. Gnathopods 1-2 similar to

female. Urosomite 3 with large hook dorsally.

Uropod I outer ramus with 5 dorsal robust setae,

inner with 1 dorsomedial and 1 subapical robust

setae, uropod 2 outer ramus with 3 dorsal robust

setae, inner with 1 broad dorsomedial robust seta;

uropod 1 peduncle with 2 apicolatcral robust

setae, with 1 basofacial slender seta; uropod 2

peduncle with 12 dorsal robust setae. Uropod 3

with inner ramus elongate, exceeding apex of

article 1 on outer ramus. Telson elongate, length-

width ratio = 7:6.

Etvmology. For Margaret Drummond who identi¬

fied this species as new from Museum Victoria

collections and contributed so much to knowledge

of Australian amphipods.

Remarks. The following variations from the holo-

type were observed in the paralypes. The main

ventral setae of coxae 1-4 = (4-5)-(4-6H4-6)-0.

Uropod 1 outer ramus with 3^ dorsal robust

setae, inner ramus with 1 dorsomedial and 1 sub-

apical robust setae. Uropod 2 outer ramus with

1-2 dorsal robust setae, inner ramus with 1 dorsal

robust setae.

Binihius drummondae shares the dorsal hook

of urosomite 3 with Tickalerus hirubi, both

species of Kulgaphoxus and three other new

species of Biruhius described herein. This species

can not be placed in the genus Tickalerus as it

lacks both the well developed dorsal setation on

article 4 of female antenna 1 and the shortened

outer ramus of uropod 3, characters diagnostic of

the type species, T, hirubi. It remains distinct

from both species Kidgaphoxiis in its lack of

accessory apical nails on the inner rami of both

uropods 1-2. the unreduced rostrum and the per¬

fectly rectangular coxa 4, a character it shares

with T. hirubi. Biruhius drummondae differs from

previously described species of Biruhius by the

presence of the dorsal hook on urosomite 3, and

from the other new species described herein by

the combination of characters listed in the diag¬

noses. The species is number MoV3679 in

Museum Victoria's TAXA database.

Biruhius heislersi sp. nov.

Figures 7-11

Material examined. Holotypc. Australia, Victoria

Ninety Mile Beach (38°30*S. 147°25.8'E), 40 ni'

Smith-Mcintyre grab, 8 May 1998 (MAFRI stn 37C)'

NMV J47320 (1 female, tl. 6.8 mm).

Paratypes. Australia. Victoria, Apollo Bay, Skenes

Creek (38°23.4*S. I44®15.6'E), 40 m, Smith-McIntyre

grab, 3 May 1998 (MAFRI stn ISC). NMV J47321 (2

females, tl. 6.5 9.75 mm).

Diagnosis. Rostrum constricted. Antenna 2, arti¬

cle 4 without well developed dorsal setation.

Right lacinia mobilis bifid, distal branch den¬

ticulate. Pcrcopods 3^ carpus with 3^-4

proximoposterior robust setae. Pereopod 5 dactyl

fully formed. Pereopod 7 basis with 1 medium

ventral setae, without long ventral setae. Coxa I

expanded disially. Coxa 4 lacking long ventral

setae. Epimeron 3 with small tooth; without ven¬

tral setae; without long posterior seta; with

oblique row of facial setae. Urosomite 3 without

dorsal hook. Uropod I with basofacial setae.

Uropods 1-2 inner rami lacking accessory apical

nails. Uropod 3 reduced, outer ramus shortened,

subcqual to peduncle.

Description of female. Mead about 18% of total

body length, greatest width about 78% of length;

ro-strum constricted, exceeding peduncular article

I on antenna 1. Eyes medium, clear of pigment.

Antenna 1 peduncular article 1 about 1.3 times as

long as wide, about 1.6 times as wide as pedun¬

cular article 2, ventral margin with 10 setulcs,

weakly produced dorsal apex with 3 setules;

peduncular article 2 about 0.8 times as long as

peduncular article 1, with 8 ventral setae; primary

tlagellum with 15 articles, about 0.95 limes as

long as peduncle, bearing aesthctascs; accessory

llagcllum with 13 articles. Antenna 2, peduncular

article 4 robast setae fonnula ^ 1-3-5-6, dorsal

margin with notch bearing 3 setae, ventral margin

with 6 groups of 1 -2 long to medium setae. I ven-

trodistal robust seta; peduncular article 5 about

0.7 times as long as peduncular article 4. facial

robust seta fonnula = 0-3, dorsal margin naked,

ventral margin with 3 sets of 1 -2 long to short

setae, 3 vcntrodistal long to medium robust setae;

llagelliim 1.6 times as long as peduncular articles

4-5 combined, with 16 articles. Mandibles with

medium palpar hump; right incisor with 3 teeth;

left incisor with 3 teeth in 2 branches; right lacinia

mobilis billd, distal branch shorter than proximal

branch, broad, denticulate, proximal branch

simple, pointed, with marginal denticles; left

lacinia mobilis with 4 teeth; right raker 10; left

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

265

rakers 7; molar in form of short protmsion dcmar- seta strongly disjunct; palp article 1 slightly elon-

cated mainly by robust setae, right and left molar gate, article 2 with 1 medium inner apical seta and

with 5-6 long robust setae, plus 1 short robust 2 other shorter inner setae, article 3 about 0.8

266

J. TAYLOR AND C. B. POORE

Figure 8. Birubius heislersi sp. nov., holotype female, tl. 6.80 mm.

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

267

Figure 9. Birubius heislersi sp. nov., holotypc female, tl. 6.80 mm.

268

J. TAYLOR AND C. B. POORE

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

Figure 11. Biruhius heislersi sp. nov., holotype female, tl. 6.80 mm.

270

J. TAYLOR AND C. B. POORE

times long as article 2, apex oblique with 8 robust

setae, with 6 basofacial setae. Maxilla I inner

plate large, bearing 1 long apical seta, 1 similar

apicomedia! seta, I similar apicolateral seta; palp

article 2 with 4 apicomedial marginal robust setae

and 6 submarginal setae. Maxilla 2 inner and

outer plates extending subequally. outer not

broader than inner. Maxillipcd inner plates with 2

large thick apical robust seta, 3 apicofacial setae,

4 medial setae; outer plate with 6 medial and

apical robust setae, 2 apicolateral setae; palp

article 1 with I apicolateral setae, article 2 with

3 apicolateral setae, article 3 protuberant, with

5 proximal facial setae, with 2 lateral setae, nail of

article 4 medium length, with 1 accessory setulcs.

Coxa 1 expanded distally, anterior margin weakly

concave; main ventral setae ot coxae 1-4 = 8-8-9-

0, postcrioimost seta of coxae 1-3 shortened;

anterior and posterior margins of coxa 4 diver¬

gent, posterior margin oblique, almost straight,

postcrodorsal comer sharp, postcrodorsal margin

short, width-length ratio of coxa 4 = 59:50. Long

posterior setae on basis of gnathopods 1-2 and

pcrcopods 3^ = 3-8-12-12, short to medium pos¬

teriors =2-1 -1-0 , long antcriors = 3-7-2-1, short

antcriors = 1-6-4-6.

Gnathopod propodus narrow; gnathopods 1-2

width ratios of carpus-propodus = 5:7 and 11:15,

length ratios = 1:1 and 25:26; palmar humps

ordinary, palms oblique; gnathopods 1-2 carpus

elongate. Pcrcopods 3-4 similar, facial setae on

mems = 5 and 6, on carpus = 4 and 5; main spine

of carpus extending to M. 75 on propodus, carpus

with 3-4 proximoposterior robust setae; robust

setae fomiula of propodus = 4 + 5; acclivity on

inner margin of dactyls of pcrcopods 3-4 weak,

midfacial seta ordinary. Coxae 5-7 postcrovcntral

setLile fomiula = 7-9-7; merus-carpus of pereo-

pods 5-6 broad, facial robust setae rows dense,

facial ridge fomiula on basis of pereopods 5-7 =

0-2-2, anterior ridge of pereopod 7 very short;

width ratios of basis, mcrus, carpus, propodus of

pereopod 5 = 29:30:26:12, of pereopod 6 =

50:33:25:13, of pereopod 7 = 65:15:14:6, length

ratios of pereopod 5 = 55:23:26:29, of pereopod 6

= 62:39:31:33, of pereopod 7 = 69:19:17:17;

pereopod 7 basis reaching or exceeding middle of

carpus, with 1 medium ventral setae. PIcopods

1-3 with 2 coupling hooks; pleopod 1 with 3 mid

and 3 proximal facial setae, pleopod 2 with

2 proximal, 3 mid and 2 distal facial setae, plco-

pod 3 with 5 proximal and 5 distal facial setae;

articles on outer rami = 15-14-16, inner rami =

10-9-10.

Epimeron 1 postcrovcntral comer rounded,

anteroventral margin with 8 short to medium

setae, postcrovcntral face with 2 long setae, pos¬

terior margin with 3-5 setulcs in sinuses;

epimeron 2 posteroventral comer rounded, with 5

facia! setae, posteriomiost pair set vertically, pos¬

terior margin with 5-6 setulcs in sinuses;

epimeron 3 posteroventral comer weakly pro¬

tuberant, with small tooth, posterior margin

straight, with selule sinuses, ventral margin

naked, face w'ith horizontal row' of 9 setae.

Urosomitc 1 naked, articulation almost complete;

urosomite 3 weakly protuberant dorsally, without

hook. Uropods 1-2 rami with articulate enlarged

apical nails, uropod 1 outer ramus with 6 dorsal

robust setae, inner with 1, uropod 2 outer ramus

W'ith 4 dorsal robust setae, inner with 1 dorso-

mcdial robust seta; uropod 1 peduncle with 7 api¬

colateral robust setae and 2 basofacial slender

setae, apical ly with 2 marginal robust setae, api-

calmost enlarged, medially with 3 slender setae;

uropod 2 peduncle with 7 dorsal robust setae; api¬

colateral comers of peduncles on uropods 1 2

without comb. Uropod 3 reduced, outer ramus

shortened, subcqual to peduncle. Uropod 3

peduncle with 5 ventral robust setae, dorsally

with 1 lateral robust seta; rami masculine, inner

extending to M. 100+ on article 1 of outer ramus,

apex with 2 setae, tnedial margin with 1 seta, lat¬

eral margin with 4 setae, article 2 of outer ramus

short, 0.21, bearing 2 long .setae, apicomedial

margin of article 1 with a single seta, lateral

margin with 2 acclivities, robust setal formula =

1-1-2, slender setal fomiula = 2-1-0. Telson

length-width ratio = 1:3, almost fully cleft, each

apex wide, rounded, lateral acclivity broad, shal¬

low, bearing ordinary lateral setule. robust setae

next medial longer than setule, midlatcral setulcs

diverse.

Male. Unknown.

Etymolog}\ For Simon lleislcrs. Museum Victo¬

ria, who identified this species as new from

Museum Victoria collections.

Remarks. The following variations from the holo-

type were obscr\'ed in the paratypes. The main

ventral setae of coxae 1-4 = (5-8)-(5-8)-(5-9)-0.

Uropod I outer ramus with 6-8 dorsal robust

setae, inner ramus with 1 dorsal robust seta. Uro¬

pod 2 outer ramus with 3^ dorsal robust setae,

inner ramus with I dorsal robust seta.

Birubhis heislersi conforms well to I3arnard

and Drummond's (1978) diagnosis of Biruhius

except that it exhibits shortened rami of uropod 3

as seen in Tiekalerus and Kttlgapltoxus. It differs

from these genera in the lack of a dorsal hook on

urosomite 3. Attempts to identify this species

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

271

using Barnard and Drummond’s (1978) key

failed. Biruhius heislersi appears most similar to

B. lowannus (Barnard and Drummond, 1978) but

differs on many accounts including the denticu¬

late vs simple distal branch of the right lacinia

mobilis, more ventral setae on coxae 1-3, the

presence of 3-4 vs 1 proximoposterior robust seta

on carpus of pcrcopods 3^ and the naked ventral

margin of uropod 3. The species is number

MoV367I in Museum Victoria’s TAXA

database.

Biruhius lowryi sp. nov.

Figures 12-17

Material examined. Ilololype. Papua New Guinea, NW

comer of Pig I. (05°9.98'S, 145^50.45’E), 21 m, J. D.

Thomas, 4 Feb 1990 (stn PNG 33K), AM P60004 (1

female, tl. 4.75 mm).

Allotype. Papua New Guinea. Barracuda Point, E of

Pig 1. (05*^10.26*5, 145°50.6rE). 30 m, J. D. Thomas,

8 Feb 1990 (stn PNG 37K). AM P56151 (1 male, tl.

3.75 mm).

Paratypes. Same data as allotype. AM P60005 (4

females, tl 3.0-3.75 mm). Papua New' Guinea, Horse¬

shoe Reef. Bootless Inlet (09°30.05’S, 147°15.50’E),

30 m. 28 Oct 1980, AM P60006 (2 females, ll. 3.75-3.9

mm).

Diagnosis. Rostrum constricted. Antenna 2, arti¬

cle 4 without well developed dorsal setation.

Right lacinia mobilis bifid, distal branch simple.

Pcrcopods 3-4 carpus with 1 proximoposterior

robust seta. Pereopod 5 dactyl fully formed.

Percopod 7 basis with long ventral setae. Coxa

1 strongly expanded dislally. Coxa 4 lacking long

ventral setae. Epimeron 3 with small tooth; with

ventral setae; without long posterior seta; without

facial setae. Urosomitc 3 with large dorsal hook.

Uropod 1 with basofacial setae. Uropods 1-2

inner rami lacking acce.ssory apical nails. Uropod

3 unreduced, outer ramus longer than peduncle.

Description of female. Head about 16% of total

body length, greatest width about 84% of length;

rostrum constricted, narrow, elongate, reaching

middle of peduncular article 2 on antenna 1. Eyes

large, clear of pigment. Antenna I peduncular

article I about 1.5 times as long as wide, about 2.0

times as wide as article 2, ventral margin w'ith

8 setulcs, unproduced dorsal apex without sctulc;

peduncular article 2 about 0.6 times as long as

peduncular article 1. with 5 ventral setae; primary

flagellum with 10 articles, about 0.8 times as long

as peduncle, lacking aeslhetascs; accessory tlag-

cllum with 8 articles. Antenna 2, peduncular

article 4 robu.st setae fonnula = 1-3-4-4, dorsal

margin with notch bearing 2 setae, ventral margin

with 6-7 groups of 1-2 long to short setae, with¬

out vcntrodistal robust seta; peduncular article 5

about 0.76 times as long as peduncular article 4,

facial robust seta formula = 1-2, dorsal margin

naked, ventral margin with 4 sets of 1-2 long to

short setae, 2 ventrodistal long to medium robust

setae; flagellum 1.07 times as long as articles 4-5

of peduncle combined, with 11 articles.

Mandibles with medium to large palpar hump;

right incisor with 3 teeth; left incisor with 2

humps in 2 branches; right lacinia mobilis bifid,

distal branch shorter than proximal branch,

simple, pointed, proximal branch simple, pointed;

left lacinia mobilis with 5 teeth; right raker 8: left

rakers 7; molar in fonn of short protrusion demar¬

cated mainly by robust setae, right molar with 6

long robust setae, left molar with 6 long robust

setae, no seta disjunct; palp article 1 slightly elon¬

gate, article 2 with 2 long-medium inner apical

.setae and 2 other medium inner setae, article 3

about 0.86 times long as article 2. apex oblique

with 6 robust to slender setae, without basofacial

setae. Maxilla 1 inner plate narrow, bearing 1

long apical seta plus I shorter apicomedial seta;

palp article 2 with 1 apicomedial marginal robust

seta, 3 apicomedial setae and 3 submarginal setae.

Maxilla 2 inner and outer plates extended equally.

Maxillipcd inner plates with I large thick apical

robust seta, 3 apicofacial setae, 1 medial seta;

outer plate with 5 medial and apical robust setae;

palp article I with 1 apicolaleral seta, article 2

with 2 apicolateral setae and 1 other lateral seta,

article 3 unprotuberant, with 2 facial setae, nail of

article 4 long, with 2 accessory setules. Coxa 1

strongly expanded distally; posterior setae of coxa

1-3 = 3-4-3, main ventral setae of coxae 1-4 =

6-6-8-(), posteriomiost seta oT coxae 1-3 elon¬

gate; anterior and posterior margins of coxa 4

strongly divergent, posterior margin oblique, pos-

tcrodorsal comer rounded, posterodorsal margin

medium, width-length ratio of coxa 4 almost =

1:1. Long posterior setae on basis of gnathopods

1-2 and pcrcopods 3-4 = 2-8-9-10, short posteri¬

ors = 5-6-6-7, long anteriors = 0-10-0-0, short

anteriors = 6-6-12-12.

Gnathopods, width ratios of carpus-propodus

on gnathopods 1-2 = 9:11 and 10:13, length ratios

= 23:26 and 25:19; palmar humps ordinary, palms

oblique; gnathopods 1-2 carpus of medium

length. Pcrcopods 3-4 similar, facial setae on

mems = 4 and 3, on carpus = 4 and 4; main spine

of carpus extending to M. 91 on propodus, carpus

with 1 proximoposterior robust seta; robust setae

fonnula of propodus =2 + 4; acclivity on inner

margin of dactyls of pcrcopods 3-4 weak, mid¬

facial seta short. Coxae 5-7 posteroventral seta

272

J. TAYLOR AND C. B. POORE

Figure 12. Birubius lowryi sp. nov., holotype female, tl. 4.75 mm (m = male allotype, 3.75 mm).

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

Figure 13. Biruhius lowryi sp. nov., holotype female, tl. 4.75 mm.

274

J. TAYLOR AND C. B. POORE

Figure 14. Birubius lowryi sp. nov., holotvpe female, tl. 4.75 mm (m = male allotype, 3.75 mm).

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

275

Figure 15. Binihitis lowryi sp. nov., holotype female, tl. 4.75 mm (m = male allotype, 3.75 mm).

276

J. TAYLOR AND C. B. POORE

Figure 16. Birubius lowryi sp. nov., holotype female, tl. 4.75 mm (m = male allotype, 3.75 mm).

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

111

Figure 17. Biruhius lowryi sp. nov., holotype female, tl. 4.75 mm.

formula = 6-1-3; articles 4-5 of pereopods 5-6

medium to narrow, facial robust setae rows

poorly developed, facial ridge formula on basis of

pereopods 5-7 = 0-2-2, anterior ridge of pcreopod

7 long; width ratios of basis, mcrus, carpus,

propodus of percopod 5 = 24:27:20:9, of pcrco-

pod 6 = 37:23:15:8, of pereopod 7 = 25:7:6:3,

length ratios of pcreopod 5 = 41:17:26:24, of

pereopod 6 = 49:32:26:36, of pereopod 7 =

53:14:13:15; Pereopod 7 basis of reaching apex of

merus, moderately setose vcntrally. Plcopods 1-3

with 2 coupling hooks; pleopod 1 with 5 distal

facial setae, pleopod 2 with 2 distal facial setae,

pleopod 3 with 2 mid and 5 distal facial setae;

articles on outer rami = 14-13-15, inner rami =

10 - 8 - 10 .

Epimeron 1 posteroventral comer rounded,

anterovcntral margin with 5 setae, posterovcntral

face with 3 medium setae; epimeron 2 pos-

tcroventral comer rounded, with 7 facial setae;

epimeron 3 posterovcntral comer with small to

medium tooth, posterior margin almost straight

with 3 medium setae, ventral margin with 6

medium setae. Urosomite 1 naked, articulation

line almost complete; urosomite 3 with large hook

dorsally. Uropods 1-2 rami with articulate

enlarged apical nails, uropod 1 outer ramus with 5

dorsal robust setae, inner with 1 dorsomedial

robust seta, uropod 2 outer ramus with 3 dorsal

robust setae, inner with 2 small robust seta; uro¬

pod 1 peduncle with 4 apicolatcral robust setae,

and 5 basofacial slender setae, medially with

many marginal setae plus apical enlarged robust

seta; peduncle of uropod 2 with 9 dorsal robust

setae; apicolatcral comers of peduncles on

uropods 1-2 without comb. Uropod 3 unreduced,

outer ramus longer than peduncle. Uropod 3

peduncle with 5 ventral robust setae, dorsally

with 1 lateral seta; rami feminine, inner extending

to M. 41 on article 1 of outer ramus, apex with 2

setae, medial and lateral margins naked, article 2

of outer ramus elongate, 0.26, bearing 2 long

setae, apicomedial margin of article 1 with 2

setae, lateral margin with 3 acclivities, robust

278

J. TAYLOR AND C. B. POORE

seta! formula = 1 -1 -1 -0, slender sctal formula = 1 -

1-1-1. Telson, length-width ratio = 27:26, not

fully cleft, each apex wide, rounded, faintly

setose, lateral acclivity broad, shallow, bearing

ordinary lateral selule, robust setae next medial

shorter than selule, midlatcral setulcs diverse.

Description of male. Similar to female but eyes

larger. Antenna 1 like female but with dense

medial sclation on peduncular article 1: primary

nagellum bearing calceoli. Antenna 2 elongate,

peduncular articles 3-4 with dense dorsal seta-

tion, peduncular article 5 about as long as article

4, dorsal margin lacking calccoii bearing 2 groups

of male setae, flagellum 28-articulate bearing cal-

ccoli. Maxilliped and maxillae 1-2 similar to

female. Right mandible damaged; left lacinia

mobilis with 6 spines; left rakers 9, left molar

with 7 long robust setae, palp similar to female,

article 3 with 1 basofacial seta. Main ventral setae

of coxae 1-4 = 5-7-6-0, Gnathopods 1-2 similar

to female. Urosomitc 3 without large hook dor-

sally. Uropod 1 outer ramus with 3 dorsal robust

setae, inner with 1 dorsomedial robust seta, uro¬

pod 2 outer ramus with 3 dorsal robust setae,

inner with I dorsal robust seta; uropod 1 peduncle

with 3 apicolateral robust setae, with 3 basofacial

slender setae; uropod 2 peduncle with 8 dorsal

robust setae. Uropod 3 with inner ramus falling

short of article 1 on outer ramus. TeLson elongate,

length-width ratio = 14:13.

Etymology'. For Dr Jim Lowry, in gratitude for his

assistance and advice during the first author’s

visit to examine Australia Museum collections.

Remarks. The following variations from the holo-

type were observed in the paratypes and material

examined. The main ventral setae of coxae 1-4 =

( 5 . 7 ).( 5 - 7 ).( 5 - 7 ). 0 . Uropod 1 outer ramus with

3-4 dorsal robust setae, inner ramus with 1 dorsal

robust seta. Uropod 2 outer ramus with 2-3 dor¬

sal robust setae, inner ramus with 1 dorsal robust

seta.

Biruhius Unviyi confonns well to Barnard and

Drummond’s (1978) diagnosis Biriibiiis except

that it exhibits a dorsal hook on urosomitc 3 as

seen in Tickalerus and Kulgaphoxits. It varies

from these genera in the lack of a shortened outer

ramus of uropod 3. It differs from the other new

species described herein by the combination of

characters listed in the diagnoses. It can be distin¬

guished from B. w'ilsoni, the only other species

described from Papua New Guinea, by the

absence of posterior setae on coxae 1-3, long ven¬

tral setae on coxa 4 and the presence of proximo-

posterior setae on the carpus of pereopods 3-4.

The species is number MoV3667 in Museum Vic^

toria’s TAXA database.

Biruhius nmlUsae sp. nov.

Figures 18-24

Materia!examined. Ilolotypc. Australia. Queensland, r\]

entrance to Moreton Bay (27''()2.85'S, I53°20.11'E)^

]9 m, Smith-Mclnlyre grab, G.C.B. l*oore, 16

1998 (stn ASB/6/24). NMV J47236 (I female, tl. 5.t)

mm).

Allotype. Australia, Queensland. Middle Banks

Moreton Bay (27°15‘S, 153°15'E). 18 November 197fs

(stn 6), QM VV25241 (1 male, ll 4.8 mm).

Paratypes. Australia. Queensland. N entrance t^^

Moreton Bay, all Smith-Mclntyre grab, G.C.B. Poore

16 Mar 1998; 27°02.94*S, 153°20.()4'E. 24 m, (stti'

ASB/6/21). NMV J47237 (3 females, tl. 3.45 4.4$

mm); 27°02.34 S. I53^19.42'E. 11 m, (stn ASEi/6/26)

NMV J41725 (1 female, tl. 4.10 mm); 27°02.34'S'

153“19.47'B. 13 m, (stn ASB/6/28). NMV J41726 {\

female, tl. 3.0 mm). Queensland, Middle Bank.s. More,

ton Bay(27“15'S, 153°15'E)- 18Nov 1976(.stn6). QM

W8682 (12 females, ll. 3.104.75 mm, 9 males, tl.

4.10 4.80 mm).

Diagnosis. Rostrum constricted. Antenna 2,

article 4 without well developed dorsal sclation.

Right lacinia mobilis bifid, distal branch simple.

Pereopods 3-4 carpus with 2-3 proximoposterior

robust setae. Pcrcopod 5 dactyl fully formed.

Percopod 7 basis without long ventral setae. Coxti

I not expanded distally. Coxa 4 lacking long vem

tral setae. Epimeron 3 without tooth; with ventral

setae; without long posterior seta; with obliqut;

row of facial setae. Urosomitc 3 with large dorsal

hook. Uropod 1 with basofacial setae. Uropodi^

1-2 inner rami lacking accessory apical nails.

Uropod 3 unreduced, outer ramus longer than

peduncle.

Description of female. Head about 16% of total

body length, greatest width about 100% of length;

rostrum constricted, narrow, exceeding apex of

peduncular article 1 on antenna 1. Eyes large,

occluded with pigment. Antenna 1 peduncular

article 1 about 1.3 times as long as wide, about 2.1

times as wide as peduncular article 2, ventral

margin with 8 setulcs. unproduced dorsal apex

with 1 setuie; peduncular article 2 about 0.75

limes as long as peduncular article 1, with 6 ven¬

tral setae: primary flagellum with 12 articles,

about 0.6 times as long as peduncle, lacking

acsthetascs; accessory flagellum with 10 anicles.

Antenna 2, peduncular article 4 robust setae for¬

mula = 1-2-4-5, dorsal margin with notch bearing

1 robust seta and 1 slender seta, ventral margin

with 7-8 groups of 1-2 long to short setae, with

vcntrodistal robust seta; peduncular article 5

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

279

Figure 18. Bimbius wallisae sp. nov., holotype female, tl. 5.00 mm (m = male allotype, 4.80 mm).

280

J. TAYLOR AND C. B. POORE

Figure 19. Binibius wallisae sp. nov., holotype female, tl. 5.00 mm.

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

281

Figure 20. Biruhius wallisae sp. nov., holotype female, tl. 5.00 mm.

about 0.7 limes as long as peduncular article 4,

facial robust seta formula = 0-2, dorsal margin

naked, ventral margin with 3 sets of 1-3 long to

short setae, 2-3 vcntrodistal long to medium

robust setae; flagellum 1.15 times as long as

peduncular articles 4-5 combined, with 14

articles. Mandibles with medium palpar hump;

right incisor with 3 teeth; left incisor with 2

humps in 2 branches; right lacinia mobilis bifid,

distal branch shorter than proximal branch, sim¬

ple, pointed, proximal branch simple, pointed; left

lacinia mobilis with 5-6 teeth; right raker 8; left

rakers 9; molar in fonn of short protrusion demar¬

cated mainly by robust setae, right molar with 10

282

J. TAYLOR AND C. B. POORE

Figure 21. Binihius wallisae sp. nov., holotype female, tl. 5.00 mm.

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

283

Figure 22. Binihius wallisae sp. nov., hololype female, tl. 5.00 mm.

long robust setae, left molar with 8 long robust

setae, no seta disjunct; palp article I short, article

2 with 1 long inner apical seta, and 2 other

medium inner setae, article 3 about 0.91 limes

long as article 2, apex oblique with 11 robust to

slender setae, without basofacial setae. Maxilla I

inner plate naiTow, bearing 1 long apical seta, 1

shorter apicomedial seta plus 2 shorter apical

setae; palp article 2 with 1 apicomedial marginal

robust seta, 3 apicomedial setae and 3 sub¬

marginal setae. Maxilla 2 inner and outer plates

extended equally. Maxilliped inner plates with 1

large thick apical robust seta, 5 apicofacial setae,

I medial seta; outer plate with 6 medial and api-

284

J. TAYLOR AND C. B. POORE

cal robust setae; palp article 1 with 1 apicolatcral

seta, article 2 with 2 apicolatcral setae and 2 other

lateral seta, article 3 slightly protuberant, with 7

facial setae, nail of article 4 medium length, with

2 accessory setules. Coxa 1 unexpandcd distally;

main ventral setae of coxae 1-4 = 9-8-7-0, pos-

teriormosl seta of coxae 1-3 shortened; anterior

and posterior margins of coxa 4 divergent, pos-

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

285

Figure 24. Birubius wallisae sp. nov., allotype male, tl. 4.80 mm.

terior margin oblique, posterodorsal comer

rounded, posterodorsal margin medium, straight,

width-length ratio of coxa 4 almost = 5:6. Long

posterior setae on basis of gnathopods 1-2 and

pcrcopods 3-4 = 6-10-11-10, short posteriors =

O-O-O-O , long anteriors = 6-6-0-0, short anteriors

= 2-1-4-2.

Gnathopods, width ratios of carpus-propodus

on gnathopods 1-2 = 8:11 and 2:3, length ratios =

12:11 and 1:1; palmar humps ordinary, palms

oblique; gnathopods 1-2 carpus elongate. Pere-

opods 3-4 similar, facial setae on merus = 7 and

5, on carpus = 5 and 7; main spine of carpus

extending to M. 100+ on propodus, carpus with I

proximopostcrior robust seta; robust setae for¬

mula of propodus = 6 + 7; acclivity on inner

margin of dactyls of pcrcopods 3-4 weak, mid¬

facial seta short. Coxae 5-7 posteroventral seta

286

J. TAYLOR AND C. B. POORE

fomiula = 6-3-4; mcrus-cariHis of pcrcopods 5-6

medium to narrow, facial robust setae rows

poorly developed, facial ridge fonnula on basis of

pereopods 5-7 = 0-2-2. anterior ridge of pereopod

7 short; width ratios of basis, mcrus, carpus,

propodus of pereopod 5 = 26:28:25:11, of pereo¬

pod 6 = 41:31:19:8, of pereopod 7 = 50:15:12:6,

length ratios of pereopod 5 = 47:19:23:25, of

pereopod 6 = 27:15:13:16, of pereopod 7 =

60:16:15:18; pereopod 7 basis reaching apex of

merus, with 1 medium ventral seta. Pleopods 1-3

with 2 coupling hooks; plcopod 1 with 3 mid and

3 proximal facial setae, pleopod 2 with 2 proxi¬

mal. 3 mid and 2 distal facial setae, pleopod 3

with 5 proximal and 5 distal facial setae; articles

on outer rami = 14-16-15, inner rami = 9-11-10.

Epinicron 1 posleroveniral comer rounded,

anteroventral margin with 3 setae, posterovcntral

face with 2 medium to long setae; cpimeron 2

posterovcntral comer rounded, with 9 facia! setae,

posteriomiost pair set vertically; cpimeron 3 pos-

teroventral comer without tooth, posterior margin

rounded, naked, midface with oblique row ol 9

setae near posterior margin, ventral margin with 6

long setae. Urosomitc 1 naked, articulation line

barely present; urosomite 3 with large hook dor-

sally. Uropods 1-2 rami with articulate enlarged

apical nails, uropod I outer ramus with 6 dorsal

robust setae, inner with 1 dorsomcdial robust seta,

uropod 2 outer ramus with 4 dorsal robust setae,

inner with 1 robust seta; uropod I peduncle with

3 apicolateral robust setae. 3 smaller setae, and 3

basofacial slender setae, medially with few

marginal setae plus apical enlarged robust seta:

uropod 2 peduncle with 9 dorsal robust setae; api¬

colateral comers of peduncles on uropods 1-2

without comb. Uropod 3 unreduced, outer ramus

longer than peduncle. Uropod 3 peduncle with 5

ventral robust setae, dorsally with 1 lateral seta;

rami feminine, inner extending to M. 76 on article

1 of outer ramus, apex with 2 setae, medial

margin naked, lateral margin with 3 long setae,

article 2 of outer ramus short, 0.15, bearing 2 long

setae, apicomcdial margin of article I with 2

setae, lateral margin with 4 acclivities, robust

setal fonnula = l-l-l-l-l, slender setal formula =

Telson, length-width ratio = 1:1, not

fully cleft, each apex wide, rounded, lateral

acclivity broad, shallow, bearing ordinary lateral

setule, robust setae next medial shorter than

setule. single midlateral setule.

Description of male. Similar to female but eyes

larger. Antenna I like female but with dense

medial setation on peduncular article I; primary

flagellum bearing calceoli. Antenna 2 elongate.

peduncular articles 3-4 with dense dorsal seta¬

tion, peduncular article 5 about as long as article

4, dorsal margin bearing 1 calccolus and 2 groups

of male setae, flagellum at least 19-articulate

(both broken) bearing calceoli. Maxilliped and

maxillae 1-2 similar to female. Right lacinia

mobilis bifid, distal branch much shorter than

proximal branch, simple; proximal branch simple,

pointed; left lacinia mobilis with 5 teeth; right

raker 6; left rakers 9; right molar with 4 long

robust setae, left molar with 6 long robust setae,

palp similar to female, article 3 with 2 basofacial

setae. Main ventral setae of coxae 1^ = 6-7-7-0.

Gnathopods 1-2 similar to female. Urosomitc 3

with large hook dorsally. Uropod 1 outer ramus

with 4 dorsal robust setae, inner with 1 dorso-

medial robust seta, uropod 2 outer ramus with 3

dorsal robust setae, inner with 1 dorsal robust

seta; uropod I peduncle with 4 apicolateral robust

setae, without basofacial slender seta; uropod 2

peduncle with 10 dorsal robust setae. Uropod 3

with inner ramus elongate, reaching article 1 on

outer ramus. Telson elongate, length-width ratio

= 11:9.

Etymology’. For Dr Elycia Wallis, Museum Vic¬

toria, in appreciation of her support during the

first author's PhD candidature.

Remarks. The following variations from the holo-

type were observed in the paratypes and material

examined. The main ventral setae of coxae 1^ =

(5-8)-(4-8)-(5-8)-0. Uropod 1 outer ramus with

2-5 dorsal robust setae, inner ramus with 1 dorsal

robust seta. Uropod 2 outer ramus with 1-3 dor¬

sal robust setae, inner ramus with 1 dorsal robust

seta.

Biruhius wallisae conforms well to Barnard

and Drummond's (1978) diagnosis of Biruhius

except that it exhibits a dorsal hook on urosomite

3 as seen in Tickalerus and Kulgaphoxus. It

differs from these genera in the lack of a short¬

ened outer ramus on uropod 3 but shares with

Kulgaphoxus the proximal vs widely spread

placement of ventral setae on antennae 1 pedun¬

cular article 2. Biruhius wallisae differs from the

other new species in the combination of char¬

acters listed in the diagnoses. The species is

number MoV37l6 in Museum Victoria's TAXA

database.

Biruhius wilsoui sp. nov.

Figures 25-30

Material examined. Holotype. Papua New Guinea, NW

comer of Pig 1. (05°9.98'S. 145°50.45'E), 21 m, J. 1).

Thomas. 4 Feb 1990 (stn PNG 33K), AM P56149 (1

female, tl. 4.20 mm).

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

287

Figure 25. Birubius mlsoni sp. nov., holotype female, tl. 4.20 mm (m = male allotype, 3.55 mm).

288

J. TAYLOR AND C. B. POORE

Figure 26. Birubius wilsoni sp. nov., holotype female, tl. 4.20 mm.

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

289

Figure 27. Biruhius wilsoni sp. nov., holotype female, tl. 4.20 mm (m = male allotype, 3.55 mm).

290

J. TAYLOR AND C. B. POORE

Figure 28. Bintbiiis wilsoni sp. nov., holotype female, tl. 4.20 mm (m = male allotype, 3.55 mm).

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

291

Figure 29. Biruhius wilsoni sp. nov., holotype female, tl. 4.20 mm (m = male allotype, 3.55 mm).

292

J. TAYLOR AND C. B. POORE

Allotype. Papua New Guinea, Madang (05°9.57'S,

I45°59.93'E), 4 m, J. D. Thomas, 20 Feb 1990 (stn

PNG 54), AM P56150 (1 male, tl. 3.55 mm).

Paratypes. Same data as holotype, AM P60002 (I

male, tl. 3.7 mm). Papua New Guinea, Madang

(05°9.57'S, 145°59.93’E), 4 m, J. D. Thomas, 20 Feb

1990 (stn PNG 54), AM P60003 (2 females, tl. 4.2 4.8

mm. 53 males, tl. 2.7-3.3 mm).

Diagnosis. Rostrum constricted. Antenna 2, arti¬

cle 4 without well developed dorsal setation.

Right lacinia mobilis bifid, distal branch simple.

Pereopods 3-4 carpus without proximopostcrior

robust setae. Pereopod 5 dactyl fully formed.

Pereopod 7 basis with long ventral setae. Coxa I

strongly expanded distally. Coxa 4 with long

ventral setae. Epimeron 3 with small tooth; with

ventral setae; without long posterior seta; without

facial setae. Urosomite 3 with small dorsal hook.

Uropod I with basofacial setae. Uropods 1-2

inner rami lacking accessory apical nails. Uropod

3 unreduced, outer ramus longer than peduncle.

Description of female. Head about 19% of total

body length, greatest width about equal to length;

rostrum constricted, narrow, elongate, reaching

middle of peduncular article 2 on antenna 1. Eyes

medium, pigmented. Antenna 1 peduncular

article 1 about 1.6 times as long as wide, about 2.1

times as wide as peduncular article 2, ventral

margin with 9 setules, unproduced dorsal apex

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

293

without setule; peduncular article 2 about 0.66

times as long as peduncular article I, with 9 ven¬

tral setae; primary flagellum with 9 articles, about

0.6 times as long as peduncle, lacking aeslhetascs;

accessory flagellum with 9 articles. Antenna 2,

peduncular article 4 robust setae formula = 1 -2-4-

3, dorsal margin with notch bearing 3 setae, ven¬

tral margin with 7-8 groups of 1-2 long to short

setae, 1 long venlrodislal robust seta; peduncular

article 5 about 0.68 times as long as peduncular

article 4, facial robust seta fomuila = 1-2, dorsal

margin naked, ventral margin with 3 sets of 1-2

long to short setae, 2 ventrodistal long to medium

robust setae; flagellum 0.85 times as long as

peduncular articles 4-5 combined, with 9 articles.

Mandibles with medium palpar hump; right

incisor with 3 teeth; left incisor with 2 humps in 2

branches; right lacinia mobilis bifid, distal branch

much shorter than proximal branch, simple,

pointed, proximal branch simple, pointed; left

lacinia mobilis with 4—5 teeth; right raker 6; left

rakers 8; molar in form of short protiusion demar¬

cated mainly by robust setae, right molar with 6

long robust setae, left molar with 6 long robust

setae, no seta disjunct; palp article 1 slighllyclon-

gate. article 2 with 3 long-medium inner apical

setae and 1 other medium inner seta, article 3

about 0.93 limes long as article 2, apex oblique

with 8 robust to slender setae, without basofacial

setae. Maxilla 1 inner plate narrow, bearing 1

long apical seta, I shorter apicomedial seta, 2

apicolateral much shorter seta; palp article 2 with

oncapicomedial marginal robust seta, 3 api¬

comedial setae and 3 submarginal setae. Maxilla

2 inner plate shorter and broader than outer.

Maxillipcd inner plate with 1 large thick apical

robust seta, 2 apicofacial setae, 3 medial setae;

outer plate with 5 medial and apical robust setae;

palp articles 1 and 2 with 2 and 3 apicolateral seta,

article 3 unprotuberant, with 2 facial setae, nail of

article 4 long, with 2 accessory sctules. Coxa 1

strongly expanded distally; main ventral setae of

coxae 1 -4 = 9-9-10-6, posteriomiost seta of coxae

1- 2 medium, of coxa 3 elongate; anterior and

posterior margins of coxa 4 strongly divergent,

posterior margin oblique, posterodorsal comer

rounded, posterodorsal margin medium, undulcnt,

width-length ratio of coxa 4 almost = 29:31. Long

posterior setae on basis of gnathopods 1-2 and

pereopods 3—4 = 3-8-8-8, short posteriors =

2- 2-4-3 , long antcriors = 4-6-()-0, shoil antcriors

= 2-I-I-0.

Gnathopods, width ratios of carpus-propodus

on gnathopods I-2 "" 2I:24 and 18:25, length

ratios = 8:11 and 8:13; palmar humps ordinary,

palms oblique; gnathopods 1-2 carpus of medium

length. Pereopods 3^ similar, facial setae on

merus = 5 and 5, on carpus = 6 and 5; main spine

of carpus extending to M. 87 on propodus, carpus

without proximoposterior robust setae; robust

setae formula of propodus = 2-^4 and 2 + 5;

acclivity on inner margin of dactyls of pereopods

3-4 weak, midfacial seta short. Coxae 5-7 pos-

terovenlral seta formula = 4-3-6; merus-carpus of

pereopods 5-6 medium to narrow', facial robust

setae row's poorly developed, facial ridge fomiula

on basis of pereopods 5-7 = 0-2-2, anterior ridge

of pcrcopod 7 long; width ratios of basis, merus,

carpus, propodus of pereopod 5 = 24:20:14:9, of

pcrcopod 6 = 34:23:14:6, of pereopod 7 =

50:13:9:5, length ratios of pcrcopod 5 =

38:23:22:27, of pcrcopod 6 = 24:15:17:18, of

pereopod 7 = 58:17:17:18; basis of pereopod 7

reaching apex of merus, heavily setose ventral ly.

Plcopods 1-3 with 2 coupling hooks; plcopod 1

with 4 distal facial setae, plcopod 2 with 2 distal

facial setae, pleopod 3 with 6 mid facial setae;

articles on outer rami = 14-13-15, inner rami =

9-8-11.

Epinieron I postcrovcntral comer rounded,

anteroventral margin with 4 setae, posteroventral

face with 3 medium setae; epimeron 2 pos-

tcroventral comer rounded, with 6 facial setae,

posteriomiost pair set vertically; epimeron 3 pos-

leroventral comer with small to medium tooth,

posterior margin almost straight w'ith 2 medium

setae, ventral margin with 6 medium setae. Uro-

somite 1 naked, articulation line absent; uro-

somite 3 with small hook dorsally. Uropods 1-2

rami with articulate enlarged apical nails, uropod

1 outer ramus with 4 dorsal robust setae, inner

with 1 dorsomedial robust seta, uropod 2 outer

ramus with 2 dorsal robust setae, inner without

robust seta; uropod 1 peduncle with 4 apicolateral

robust setae, and 4 basofacial slender setae, medi¬

ally with many marginal setae plus apical

enlarged robust seta; uropod 2 peduncle with

7 dorsal robust setae; apicolateral corners of

peduncles on uropods 1-2 with comb. Uropod 3

unreduced, outer ramus longer than peduncle.

Uropod 3 peduncle with 6 ventral robust setae,

dorsally with 1 lateral seta; rami feminine, inner

extending to M. 46 on article I of outer ramus,

apex with 1 seta, medial and lateral margins

naked, article 2 of outer ramus short, 0.23, bear¬

ing 2 long setae, apicomedial margin of article I

with 2 setae, lateral margin with 4 acclivities,

robust setal formula = 1-1-1-1-0, slender

setal fonmila = l-l-l-l-l. Tclson, length-width

ratio = 55:54, not fully cleft, each apex wide,

rounded, setose, lateral acclivity broad, shallow,

bearing ordinary lateral setule, robust setae next

294

J. TAYLOR AND C. B. POORE

medial shorter than setule, midlateral setules

diverse.

Description of male. Similar to female but eyes

larger. Antenna I like female but with dense

medial setaiion on peduncular article I; primary

flagellum bearing calceoli. Antenna 2 elongate,

peduncular articles 3^ with dense dorsal seta-

tion, peduncular article 5 about as long as article

4, dorsal margin lacking calceoli, bearing 2

groups of male setae, flagellum 24-articulate

bearing calceoli. Maxilliped and maxillae 1-2

similar to female. Right mandible damaged, left

lacinia mobilis with 5 spines; left rakers 7; molar

in form of bulbous hump, left molar with 4 long

robust setae, palp similar to female, article 3 with

1 basofacial seta. Main ventral setae of coxae 1^

= 8-7-8-5, Gnathopods 1-2 similar to female.

Pereopod 7 basis narrower than female. Uro-

somile 3 without hook dorsally Uropod 1 outer

ramus with 3 dorsal robust setae, inner with 1 dor¬

sal robust seta, uropod 2 outer ramus with 2 dor¬

sal robust setae, inner without robust seta; uropod

1 peduncle with 2 apicolatcral robust setae, with 4

basofacial slender seta; uropod 2 peduncle with 9

dorsal robust setae. Uropod 3 with inner ramus

falling short of article I on outer ramus. Tcison

elongate, length-width ratio = 6:5.

Etymology'. For Dr Robin Wilson, Museum Vic¬

toria, a good friend and colleague of both authors.

Remarks. The following variations from the holo-

type were observed in the paratypes. The main

ventral setae of coxae 1-4 = (7-9H7-9)-(8-l 1 )-

(6-9). Uropod 1 outer ramus with 1-4 dorsal

robust setae, inner ramus with 1 dorsal robust

seta. Uropod 2 outer ramus with 2 dorsal robust

setae, inner ramus without dorsal robust setae.

Binibius wilsoni conforms well to Barnard and

Drummond's (1978) diagnosis of Biruhius except

that it exhibits a dorsal hook on urosomite 3 as

seen in Tickalerus and Kidgaphoxus. It varies

from these genera in the lack of a shortened outer

ramus on uropod 3 but shares with Kidgaphoxus

the proximal vs widely spread placement of ven¬

tral setae on antennae 1 peduncular article 2. Biru¬

hius wilsoni dilTers from the other new species

described herein by the combination of characters

listed in the diagnoses. It can be distinguished

from B. lowryi, the only other species described

from Papua New Guinea, in the presence of pos¬

terior setae on coxae 1-3, long ventral setae on

coxa 4 and the absence of proximoposterior setae

on the carpus of pereopods 3-4. The species is

number MoV3666 in Museum Victoria's TAXA

database.

Discussion

Biruhius is by lar the largest genus of phoxo-

cephalid amphipods, now comprising 38 species

from Australia (Barnard and Drummond, 1978),

three species from Indonesia (Dana, 1853, Ortiz

and Lalana, 1997, 1999) and two species from

Papua New Guinea. Species of Biruhius occur

intertidally to 70 m in benthic sandy to muddy

sediments. The new species expand the depth and

geographic range of the genus from that pre¬

viously known. The biogcographic relationships

between the species from Australian waters and

those from Papua New Guinea and Indonesia arc

as yet unknown but could be elucidated only by

cladistic analysis of species of Biruhius,

Kidgaphoxus, Tickalerus and Van.

.Acknowledgements

We acknowledge the assistance and support

throughout the first author’s PhD candidature of

those people named in the etymologies. Examina¬

tion of Australian Museum collections was made

possible through financial assistance awarded to

the first author in an Australian Museum Post¬

graduate Award. This project was supported by

an Australian Research Council grant to G. C. B.

Poore and R. S. Wilson.

References

Alonso de Pina, 1993. Linca pinafa. a new phoxo-

ccphalid genus and species (Crustacea:

Amphipoda) from the Argeiiline continental shelf

Proceedings of the Biological Society of Washing¬

ton 106(3): 497-507.

Barnard, J.L., 1960. The amplhpod family Phoxo-

cephalidae in the ea.slcm Pacific Ocean, with anal¬

yses of other species and notes for a revision of the

family. Allan Hancock Pacific Expeditions 18:

175-368.

Barnard, J.L., 1979. Littoral gammaridean Amphipoda

from the Gulf of California and the Galapagos

Islands. Smithsonian Contributions to Zoology'

271: vi+ 149 pp. 74 figs.

Barnard, J.L. and Drummond, M.M.. 1976. Clarifica¬

tion of five genera of the Phoxocephalidac (marine

Amphipoda). Proceedings of the Biological Soci¬

ety of Washington 88: 515-547.

Barnard, J.L. and Daimmond, M.M., 1978. Gam-

maridcan Amphipoda of Australia. Part III: The

Phoxocephalidac. Smithsonian Contrihuiions to

Zoology’2A5: I 551.

Barnard. J.L. and Karaman, G.S.. 1991. The families

and genera of marine gammaridean Amphipoda

(except marine gammaroids). Parts I and 2.

Records of the Australian Museum. Supplement

13: 1 866!

NEW SPECIES OF PHOXOCEPHALID AMPHIPODS

295

Dana, J.D., 1853. Crustacea. Part II. United States

Exploring Expedition during the years IH38. 1839,

1840, 1841, 1842 under the command of Charles

Wilkes, US.N. 13: 691-618, with a folio atlas of96

plates.

Haswell, W.A., 1879. On Australian Amphipocla. Pro-

ceedings of the Linnean Society of New South

Wales 4; 245-79.

Jarrctt, N.E. and BousEcld. E.L., 1994. The amphipod

superfaniily Pho.\ocephaIoidea on the Pacific

Coast of North America. Family Phoxoccphalidac.

Parti. Melharpiniinae. new subfamily. Amphipaci-

fica 1(1): 58-140.

Ortiz, M. and Lalana, R., 1997. Results of the Zoologi¬

cal Expedition organised by “Grigore Antipa”

Museum, in the Indonesian Archipelago (1991). 1.

Peracarida (Crustacea). Amphipoda. Travaux du

Museum dllistoire Noturelle "Grigore Antipa”

38: 29-113.

Ortiz, M. and Lalana, R. 1999. Amphipoda (Crustacea)

from Indonesia collected by the Expedition of

“Grigore Antipa” Museum from Bucharest.

Travaux du Museum National D 'Histoire

Naturelle "Grigore Antipa" A\\ 155-198.

Pirlot, .I.M., 1932. Lcs amphipodes de Pexpedition du

Siboga. Deuxieme partie. Les amphipodes gam-

maridcs. I. — Les amphipodes fouisseurs. Phoxo-

cephalidae, Ocdiceroidae. Siboga-Expeditie 33b:

57-113.

Schellenberg, A., 1931. Gammariden und Caprclliden

des Magellangebietes, Sudgeorgicns und der Wes-

tantarklis. Further Zoological Results of the

Swedish Antarctic Expedition 1901-1903 2(6):

1-290.

fvlemoirs of Museum Victoria 58(2): 297-333 (2001)

BATHYAL JOEROPSIDIDAE (ISOPODA: ASELLOTA) FROM SOUTH-EASTERN

AUSTRALIA. WITH DESCRIPTION OF TWO NEW GENERA

Jean Just

Museum of Tropical Queensland, 78-102 Flinders Street, Townsville, Queensland 4810, Australia

Abstract

Just, J., 2001. Bathyal Joeropsididae (Isopoda: Asellota) from south-eastern Australia, with

description of two new genera. Memoirs of Museum Victoria 58(2): 297-333.

Fifty-three species of Joeropsis. listed here, have been previously described in this, until

now, the only genus of the family. Characters in the Joeropsididae are tabulated and discussed

to ascertain which may be diagnositic at family and genus \c\c;\. Joeropsis bicarinata sp. nov.,

Rugojoeropsis rugosa gen. nov., sp. nov., Scaphojoeropsis midticarmata gen. nov., sp. nov.,

and S. kimblao sp. nov. are described from the lower shelf and upper bathyal of south-eastern

Australia. All are strongly sculptured dorsally.

Introduction

Menibers of the Joeropsididae Nordcnslam have

been found around all continents and in all oceans

from the North Atlantic and Pacific boreal to the

subantarctic. Depths range from the intertidal to

the midcontinental slope, with most species

occurring on the mid to uppermost shelf. Up until

now, the family included a single genus. Between

them, Menzies (1962) and Wolff (1962) listed 14

species of Joeropsis Koehler; subsequently, 39

species have been described (Table 1).

Hale (1937) reported J. patagoniensis Richard¬

son from subantarctic Macquarie I. No further

records of Joeropsididae from Australia have

been made, although many undescribed species

exist in museum collections. The four new

species described herein were collected on the

lower shelf and upper slope (100-600 m, pri¬

marily below 200 m) off south-eastern Australia

from south of Sydney to the east and west coasts

of Tasmania. The four species differ from all

other jocropsidids in having a strongly sculptured

dorsum. Of the 53 species previously recognised,

only J. waltervadi Kensley has some dorsal

sculpture in the fomi of two low, broad, rounded,

parallel dorsal ridges on the anterior pereonites.

One of the new species is placed in Joeropsis',

one in a new genus, Rugojoeropsis, and two in a

.second new genus, Scaphojoeropsis.

Length of specimens was measured from the

tip of the pseudorostmm (see definition below)

along the dorsum to the apex of the pleotelson.

Analysis of characters

The new forms of Joeropsididae encountered in

this study suggested the possibility that new gen¬

era could be differentiated from Joeropsis. A

comparative analysis of the distribution of

characters in the known and new species was

undertaken to ascertain what might constitute

diagnostic characters at the family and genus

levels (Table 2).

Most known species of Joeropsis have been

incompletely described, which renders inter¬

specific comparisons difficult. Exceptions are

J. mediterranea Amar, and the redcscription, by

Boequet and Lemercier (1958), of./, brevicornis

Koehler. The description of./ vibicaria Barnard

contains few useful details. For most species

(more than 30), the following aspects are illus¬

trated in the literature: dorsal habitus view,

mandible(s), maxilla 1, maxilliped, male picopod

1, male and female pleopod 2, pleopod 3 and uro-

pod. Other characters arc occasionally illustrated

and/or described in the text. In many cases, illus¬

trations are difficult to interpret in detail. In sev¬

eral instances text and illustrations do not match.

Where infonnation could be found, the follow¬

ing characteristics were consistent in species of

Joeropsis and the four new joeropsidid species:

general configuration of antennae 1 and 2; pres¬

ence ol a pseudorostrum (sec discussion below);

no coxae visible in dorsal view; no free piconites;

all pcrcopods similar, ambulatory; shape of

mandibular molar; number of articles in

mandibular and maxillipcdal palps; general shape

of maxillae 1 and 2 (except Scaphojoeropsis

kimblae sp. nov., maxilla 2 is strongly reduced);

general shape of male and female plcopods;

general shape of uropods. These characters arc

considered here to be diagnostic of Joeropsididae.

Table 2 compares another 21 characters poten¬

tially useful in discriminating between joeropsi-

did genera. Where 30 (an arbitrary selection

reflecting availability of information) or more of

297

298

J. JUST

Table 1. Species of Joeropsis with area of type locality.

J. qtjinis Kussakin, 1961 (Sea of Okhotsk, Kurile Is)

J. antarctica Mcnzics and Schultz, 1968 (South Shetland Is)

J. antillensis Miiller, 1993 (Martinique)

J. beuroisi Kensley, 1975 (South Africa)

J. bicarinata sp. nov.(soulh-castcm Australia)

J. bifasciata Kensley, 1984a (Belize)

J. bourboni Muller, 1990a (Reunion)

J. brevicornis Koehler, 1885 (North-east Atlantic)

J. caboverdensis Miiller, 1988 (Cape Verde Is)

J. ceylonensLs Miiller, 1991a (Sri Lanka)

J. concava Schultz. 1966 (Santa Cruz Canyon)

J. coralicola Schultz and McCloskcy, 1967 (North Carolina)

J. curvicornis (Nicolet. 1849) (Chile)

J. doUfusi Norman. 1899 (Mediterranean)

J. ditbia Menzies, 1951 (California)

J.faurei Muller, 1990a (Reunion)

J. gertrudae Muller, 1989a (Society Is)

J. hawaiiensis MiWer^ 1941 (Hawaii)

J. indica Muller, 1991a (Sri Lanka)

J. integra Kensley, 1984b (South Africa)

J. intermedia Nordenstam, 1933 (Argentina)

J. juvenilis Kensley, Ortiz and Schottc, 1997 (Cuba)

J. lata Kussakin, 1961 (Sea of Okhotsk)

- J. legrandi }uc\iiiu\U 1962 (Mediterranean)

J. letourneiiri Muller, 1990a (Reunion)

J. lobata Richardson, 1899 ((Talifomia)

./. marionis Beddard, 1886 (Marion 1.)

J. mediterranea Amar, 1961 (Mediterranean)

J. meteor Muller, 1991b (Gulf of Aden)

./. mimita Muller, 1989a (Society Is)

J. monsmariniis Kensley, 1980 (Veina Seamount)

./. montalentii Fresi, 1968 (Mediterranean)

J. neozealanica Chilton, 1892 (New Zealand)

J. nigricapitis Kensley, 1994 (Bermuda)

J. palliseri Hurley, 1957 (New Zealand)

y. paradubia Muller, 1989b (Columbia)

J, paulensis Vanhoffen, 1914 (Antarctica)

J. patagoniensis Richardson. 1909 (Patagonia)

J. personata Kensley, 1984a (Belize)

J. polynesiensis Muller, 1989a (Society Is)

./. rathbunae Richardson, 1902 (Bermuda)

J. salvati Muller, 1989a (Society Is)

J, sanciipauli Kensley, 1989 (St Paul I.)

J. schoelcheri Muller, 1993 (Martinique)

J, serrtilus Kensley, 1984b (South Africa)

J. setosa George and Stromberg, 1968 (San Juan Archipelago)

J. stebbingi Kensley, 1975 (South Africa)

J. tayrona Muller, 1989b (Columbia)

J. tobagoensis Kensley & Schotte, 1994 (Tobago)

J. unidentata Kensley, Ortiz and Schottc, 1997 (Cuba)

/ vibicaria Barnard, 1965 (Gough 1.)

J. waltervadi Kensley, 1975 (South Africa)

./■ uWffi Muller, i991b (Gulf of Aden)

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

299

Table 2. Comparison of characters in Joeropsis and the four new species described herein.

Joeropsis Joeropsis Rugojoeropsis Scaphojocropsis

Character spp.‘ bicarinata rugosa multicarimta kimhlae

Body

1. Body with subparallel margins

one or more of pereonites 2-4

occasionally slightly broader

than 1 and succeeding ones,

particularly in brooding females

(45y

— Body fusiform, distinctly tapering

from pereonite 2/3 to apex of

plcotelson

2. Cuticle smooth

(45)

(setae not considered)

— Cuticle roughly calcified

3. Without dorsal ornamentation on

(44)

pereon

— With two broad, rounded

(1)

longitudinal dorsal ridges on

pereonites 1-4(5)

— With two parallel, sharp

longitudinal dorsal keels along

entire pereon and onto pleotelson

— With variety of humps and short

ridges

Cephalon

4. Without dorsal ornamentation

(45)

— With middorsal humps

— With anterior and dorsal ridges

5. Anterior margin between antennae

not depressed (Fig. 2)

— Anterior margin between antennae

(45)

strongly depressed relative to

dorsal side of cephalon (Fig. 14c)

6. Pseudorostrum in lateral view

(43)

with apical projection (Fig. 2)

— Pseudorostrum in lateral view

without apical projection (Fig. 9cl)

— Pseudorostrum attached to

depressed anterior margin of

cephalon in nearly vertical position

(Fig. 14c)

Eves

7. With dorsolateral eyes on low bulge (45)

— Blind, without remnants of eye bulge

Pereopods

8. Dactylus with 2 claws

(11)

— Dactylus with 3 claws at least on

(28)

pereopods 2-7

300

J. JUST

Table 2. Continued.

Joeropsis Joeropsis Rugojoeropsis Scaphojoeropsis

Character spp.' bicarinata rugosa multicarinata kimblae

Mouthparts

9. Upper lip evenly rounded, less than (6)

twice as wide as long (Fig. 4 ul)

— Upper lip weakly convex, more than

twice as wide as long (Fig. 20 ul)

10. Mandible cutting edge with strong (40)

teeth in curved, regular row

(Fig. 4 md)

— Mandible cutting edge divided into

two parts, large terminal tooth

pointing forward, posterior teeth

on broad flange pointing mediad

to backwards (Fig. 16 md)

11. Mandibular raker setae long, in (40)

regular row (Fig. 4 md)

— Mandibular raker setae short,

sparse, in irregular or unusual

configuration (Fig. 16 md)

12. Mandibular palp article i with apical(31)

and subapical setae (Fig. 4 md)

— Mandibular palp article 3 with apical

setae only (Fig. 20 md)

13. Lower lip, lobes longer than wide, (6)

distally tapering, pointed (Fig 4 II)

— Lower lip, lobes wider than long,

rectangular, lateral comer

projecting, pointed (eg. Fig. 20 11)

14. Maxilla 1 outer plate with 12 (17)

dentate setae

— Maxilla I outer plate with 11 (11)

dentate setae

— Maxilla I outer plate w\\\\ 10 (4)

dentate setae

— Maxilla I outer plate with 9 (1)

dentate setae

— Maxilla 1 outer plate with 8 (2)

dentate setae

15. Maxillipeds covering at least 2/3 (37)

of ventral width of cephalon,

overreaching other mouthparts

(Fig. 3 c)

— Maxillipeds covering about 1 13

of ventral width of cephalon; do

not reach anterior margin of

cephalon to fully cover other

mouthparts (Fig. 15 c)

16. Maxilliped basis reaching to apex (37)

of or beyond palp ar. 3 (Fig. 3 mp)

— Maxilliped basis reaching to middle

of palp ar.2 (Fig. 20 mp)

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

301

Table 2. Continued.

Joeropsis Joeropsis Rugojoeropsis Scaphojoeropsis

Character

spp.'

bicarinata

nigosa

multicarinata

kimblae

17. Maxillipedal palp about 1/2 length

(38)

of basis (Fig. 11 mp)

— Maxillipedal palp about 80-90%

length of basis (Fig. 16 mp)

18. Maxillipedal palp articles

(39

unproduced medially

— Maxillipedal palp article 3 strongly

produced medially

19. Maxillipedal palp article 4 longer

(40)

than 3

— Maxillipedal palp article 4 shorter

than 3

Pleopods

20. Female pleopod 2 without long

(29)

apical setae (Fig. I i pl2)

— Female pleopod 2 with long

apical setae (Fig. 19 pl2)

21. Pleopod 3 exopod articulation

(33)

strongly oblique (Fig. 6, 3)

— Pleopod 3 exopod articulation

transverse or nearly so (Fig. 21 pI3)

' Species of Joeropsis described prior to this study.

2 Number in parentheses: number of species in which character condition could been confirmed.

the species of Joeropsis were found to share a

character, with none found to differ (characters I,

2,4, 5, 6, 7, 10, 11, 12, 15, 16, 17, 19,21), that

character is considered to be shared by all species.

Comments are presented below for remaining

characters.

Character 3 (dorsal sculpture). Joeropsis \ml-

tervadi and the new species ./. bicarinata differ

from all other species o\'Joeropsis in having two

longitudinal ridges running along the dorsum,

broadly rounded and confined to the pereon in the

former, and sharp keels extending to the pleotel-

son in the latter. The homology of these two

sculptural expressions is not clear. The dorsal

sculpture on the pereon and pleotelson of the

other three new species differs distinctly from that

in the two species of Joeropsis.

Character 8 (number of pereopodal claws).

Eleven species of Joeropsis are described and/or

illustrated as having two dactylar claws on all

percopods, 28 as having three claws at least on

pereopods 2-7. Generally only one or two pereo-

pods are illustrated, and in several instances text

and illustrations do not match. Where three claws

are illustrated, the accessory claw is distinct and

of similar shape to the two terminal ones. Boequet

and Lemercier (1958), in their redescriplion of the

type species J. hrevicornis Koehler, state that

pereopod I carries two dactylar claws, all other

pereopods carry three claws, as shown in their

illustrations of pereopods 1 and 7. Similarly,

several species, e.g. J. intermedins Nordenstam

and J. patagoniensis Richardson are described

and illustrated (Nordenstam, 1933) as having two

claws on pereopod 1 and three on the rest of the

pereopods. Amar (1961) described and illustrated

J. mediterranea as having two claws on all pereo¬

pods (pereopods 1, 2 and 6 illustrated). The four

species described in this study have two claws

only on all percopods../. mediterranea, as well as

the four new species, carry small, simple setae on

the lateral and medial surfaces of the dactylus (not

known or difficult to interpret in all other species

with two claws only). Homology between one

such seta and a distinctive accessory claw cannot

be ruled out (Wilson, 1994). While two claws on

pereopod I and three on pereopods 2-7 may be

more common in Joeropsis than two claws on all

pereopods, the character is useful only at the

species level pending further comparative studies.

302

J. JUST

I am not aware that a similar dichotomy in the

number of fully developed pcreopodal claws has

been reported in other janiroid genera.

Character 9 (upper lip). The upper lip in J. bre-

vicornis (see Boequet and Lcmercicr, 1958), five

other species of Joeropsis and two of the new

species is evenly rounded, about 1.5 times as wide

as long. The upper lip has not been described or

illustrated for other species o\' Joeropsis. In the

other two new species, the upper lip is less

rounded and about 2.5 times wider than long.

More information is required to fully evaluate the

generic significance of these differences, but the

character has been included tentatively in the

diagnoses of genera.

Character 13 (lower lip). In six species, includ¬

ing J. brevicornis, plus in two of the new species,

the lobes of the lower lip are longer than wide,

cur\ang mediad and tapering to a more or less

acute apex (no infonnation is available for other

species of Joeropsis). This shape is common to

most janiroids and is tentatively considered to be

shared by all species in Joeropsis. In two of the

new species, the lower lip differs significantly,

lobes being rectangular, broader than long, with

projecting lateral comers.

Character 14 (maxilla 1 outer plate). In 34

species of Joeropsis the number of apical spine-

like setae on the outer plate of maxilla I is given

in text and/or illustration. The numbers given are

12 (17 species), 11 (II), 10 (4), 9 (I) and 8 (2). It

is not clear, however, whether numbers have been

enumerated in a consistent manner, i.c., large den¬

ticulate setae only, or smaller simple setae

included, where these occur, (cf. Fig. 4 ml: 12

dentate + 2 small simple and Fig. 16 ml: 10 + 3).

In several cases, the number given in the text and

the number illustrated do not agree, c.g. J. huwui-

iensis Miller (text: 6, illustration: 8). Clear evi¬

dence of considerable interspecific variation in

the number of denticulate setae occurs within

Joeropsis, but further comparative studies arc

required before its significance can be ascer¬

tained. Numbers given in descriptions of the new

species do not include the small simple setae.

Character 18 (maxillipedal palp article .3.)- In 40

species Joeropsis, the third article of the max-

illipcdal palp is unproduced medially (see Fig. 3

mp). J. saniipaiili Kensley has a slight expansion

medially on article 3. The two species referred to

Scapbqjoeropsis gen. nov. have distinctive

medial lobes on article 3. This character may be

useful in separating genera.

Character 20 (female pleopod 2). In 20 species

Joeropsis illustrations show that the distal fifth

to four-fifths of the lateral margins of pleopod 2

in females are densely fringed with what has been

described as ‘sctules' or 'setae' (sec below: cutic*

ular scales). No long true setae occur apically,

another nine species where the female pleopod 2

is not illustrated, pleopod 2 of the male is shown

to carry similar lateral ‘sclulcs/sclac’. Evidence

from Joeropsis bicarinata sp. nov. (Fig. 5) sug¬

gests that the ‘setaf fringe is similar in male and

female in species oi' Joeropsis. Presence of this

fringe, and absence of long apical setae on the

female operculum, (a few short simple setae may

be present), may be characteristic for all species

Joeropsis. Rugojoeropsis rugosa gen. nov., sp,

nov. has a similar fringe to Joeropsis on male and

female plcopods 2. Scaphojoerop.sis multi,

carinata sp. nov. has an almost invisible fringe in

the distalmost tenth of both female and male plco-

pod 2. Pleopod 2 of male S. kimblae sp. nov. has

a fringe along nearly the entire lateral margin (not

verified in females). Both species of Scaphojo-

eropsis carry a row of long simple setae apically

on pleopod 2. The presence of lateral ‘sctulcs/

setae' appeal's to be ubiquitous in Joeropsididae,

whereas the presence of long apical .setae appears

to be confined to Scaphojoeropsis.

Additional remarks on morphology

Rostral projection. Wilson (1989: 130) defined

the ascllote rostrum as 'a projection of tlie

cephalic frons that may also include the dorsal

surface of the ccphalon.’ Serov and Wilson

(1995: 41) proposed the following definition 'any

anterior extension from the frons or vertex

[midanterior dorsal cephalic margin] of the

ccphalon.' While a rostrum derived from the

vertex generally will incorporate the upper frons

along its ventral side, a projection derived from

the frons normally does not involve an extension

of the vertex. Whether the dorsoposterior margin

of a frons-derived projection could fuse with the

cephalic vertex, thus obscuring the origin of the

rostrum, is not clear. For the purposes of phylo¬

genetic analyses, discriminating between a true

rostrum, being an extension of the cephalic

vertex, and a frons-derived pseudorostrum may

be useful. All species in Joeropsididae possess a

pscudorostrum, and that term is used throughout

this study. All species in Joeropsis and Rugojo-

eropsis have the cephalic vertex concavely

recessed, with the pseudorostrum fitting into, and

projecting from, the recession. In Joeropsis, the

pscudorostrum projects straight forward over the

inferior parts of the frons (sec Fig. 2 habitus,

lateral view ). In Rugojoeropsis the pscudorostrum

forms a box-like hump without an apical pro¬

jection (Fig. 9 cl). In Scaphojoeropsis the

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

303

pseudorostriim borders the vertex along a straight

line, is broadly rounded apically, and angled

downwards (sec Fig. 14 c).

Cuticular scales. In most instances where plco-

pod 3 has been illustrated in species o\' Jocropsis.,

the exopod is shown with a dense fringe of thin,

simple setae. The four species described here

have complex, llattcned, often overlapping cutic¬

ular scales (arising as outgrowths from the

cuticle, not inserted into it like setae, sec, c.g..

Fig. 6, 3) along the entire margin of the exopod.

This character is not confined to these four

species because it occurs in J. mecUterranea

(Amar, 1961: fig. 3)andJ. sa I va t i {MuWer. 1989a:

fig. 75). The scales are so transparent that nor¬

mally they can be seen in their entirety only with

the aid of Nomarski or phase contrast. With ordi¬

nary light microscopy, only the slightly strength¬

ened posterior edge of the individual scale usually

is visible (as a thin, simple setule). These scales

vary in fomi and size between species as evi¬

denced by the four species described below. They

are likely to be present in all species of Jocro-

psididae. The fringe of reported ‘setules/sctac’

on the lateral margin of plcopod 2 in females

and males in Joeropsididae consists of similar

cuticular scales (c.g. Figs 51J and I Ipt).

Female reproductive structures. With the

exception of male plcopods 1 and 2, the literature

does not provide infonnation on reproductive

structures in Joeropsididae. A preparatory female

of the new species Scaphojoeropsis multicarinata

allows some preliminaiy comments to be made

(Fig. 1). The elongate ovaries reach from the ante-

riormost part of perconitc 1 through to perconite

6. The oviduct attaches to the ventral side of the

ovary in the middle of perconite 3, curves laterad

and posteriad, and opens vcntrally on perconite 5

mediad to the coxa. The spemialhecal duct

appears to open on the anterior surface of perco¬

nite 5, from where it curves into perconite 4 and

runs straight to the middle of perconite 3 to join

the oviduct in a common opening to the ovary. In

the specimen illustrated, the anteriormost portion

of the spermalhecal duct is slightly inflated and

filled with denser material. A single female of the

new species Rugojoeropsis rugosa has a similar

configuration.

Conclusion

Joeropsis hicariuata sp. nov. ditTers from other

species in the genus as currently conceived only

in the presence of two sharp, longitudinal dorsal

keels. A possibly intermediate condition between

those keels and the nomially smooth dorsum in

Joeropsis is seen in the low, rounded ridges in J.

waltervadi. J. hicarinata is referred Xo Joeropsis.

The second new species shares with Joeropsis

all characters analysed in Table 2 except for the

rough, calcified cuticle, dorsal sculpture in the

fonn of humps and transverse ridges, and the box¬

shaped, unproduced pscudorostmm. Since all

species of Joeropsis possess a smooth cuticle and

straight, overhanging pscudoroslnim, a new

genus Rugojoeropsis is justified. The remaining

two new species differ in many characters

from Joeropsis and Rugojoeropsis (Table 2),

notably in the shape of the cephalon and pseu-

dorosti-um, the shape of the entire body, and

details of mouthparts. They too warrant a new

genus, Scaphojoeropsis.

Joeropsididae Nordenstam

Jaeropsini Nordenstam. 1933: 190.

Jaeropsidae.—Menzies, 1962: 63.—Menzies and

Kruezynski, 1983: 94.

Joeropsididae.—Sivertsen and Holthuis, 1980: 96

(correction of spelling).—Kussakin, 1999; 10.

Diagnosis. Janiroidea with flattened body; lateral

margins nomially parallel, occasionally tapering

posteriorly. Perconites of subequal length, with

truncate, entire or at most finely serrate, lateral

margins covering coxae. All pereopods similar,

slender, ambulatory, with 2 or 3 dactylar claws.

Sessile eyes nonnally present in dorsolateral posi¬

tion on cephalon. Anterior margin of cephalon

with strong concavity, or rarely without concav¬

ity. Pseudorostnim present, inserted into cephalic

concavity, rarely joined along straight line.

Picotclson subcqual in width to pereonite 7, with

no free pleonites. Antenna I shorter than

cephalon, peduncular article I expanded, longer

than articles 2 and 3 combined; Hagellum shorter

than peduncle, with 2 or 3 articles, rarely 4 (one

species herein) or 5 (Joeropsis sanctipauli and

one species herein). Antenna 2 geniculate, with

peduncular article 6 and flagellum folding laterad

and backwards under lateral expansion of pedun¬

cular article 5; first 4 peduncular articles short,

article 4 more or less telescoped into 3, article 5

longer than 1-4 combined, greatly expanded lat¬

erally, article 6 much shorter than 5, generally

widening distally; antennal scale absent; fiagel-

lum with enlarged, nonnally conjoint article 1.

Mandible molar a long, slender, pointed projec¬

tion; incisor of 5-6 large teeth, (occasional small

accessory' denticles not counted); lacinia mobilis

absent; palp with 3 articles. Maxillipcds, palp

with 5 articles, at least article 2 medially

expanded. Plcopod 2 of male and female with

304

J. JUST

longer or shorter lateral fringe of modified cutic-

ular scales; pleopod 3 cxopod 2-articulatc, longer

than endopod, with lateral fringe of modified

culicular scales, endopod with 3 plumose setae;

pleopod 4 cxopod vestigial. Uropods biramous,

inserted ventrally on pleotclson nomially within

distinctive insinuation in pleotelson margin;

peduncle usually broader than long and medially

expanded; rami shorter than peduncle. Anus out¬

side plcopodal chamber, between bases of uro-

podal peduncles, partly or entirely covered by

male pleopod 1 or female pleopod 2 respectively.

Oostegites on pereopods 1-5. Female spennathe-

cal duct opening on anterior surface of pcrconite

5, oviduct opening ventral on perconite 5 mediad

to coxa.

Included genera: Joeropsis Koehler, 1885,

Rugojoeropsis gen. nov., Scaphojoeropsis gen.

nov.

Joeropsis Koehler

Koehler, 1885: !.—Sivertsen and Holthuis,

1980: 96.—Kussakin, 1999: 12.

Joeropsis. —Nordenstam. 1933: 191.—Menzies and

Barnard, 1959; 10.—Menzies, 1962: 63. (unjustified

emendation).

laeropsis. —Nierstrasz, 1941: 288 (unjustified

emendation).

Type species. Joeropsis hrevicornis Koehler,

1885 (by monotypy).

Diagnosis. Jocropsididae with lateral body

margins parallel, rarely with dorsal sculpture.

Cuticle smooth. Cephalon with anterior margin

strongly concave. Pseudorosirum inserted into

concavity, tlush with dorsal surface of cephalon,

with overhanging apex. Dorsolateral eyes present.

Upper lip evenly rounded, less than twice as wide

as long. Mandibles with incisor of 5 or 6 strong

subequal teeth in regular, curved row; spine row

setae long, in regular row. Lower lip, lobes longer

than wide, distally tapering, pointed. Maxillipcds

in ventral view covering at lease two-thirds of

cephalic width, and covering other mouthparts

except for mcdioapical concavity; enditc reaching

to end of or beyond palp article 3; palp about

half length of enditc; palp article 3 without

medial lobe, (J. sanctipatdi, with a small medial

lobe, appears to be an exception), article 4 much

longer than 3. Pleopod 2 of female with at

most a few short simple setae apically. Pleopod 3

exopod with strongly oblique articulation

between articles.

Included species. See Table 1.

Joeropsis hicarinata sp. nov.

Figures 2-7

Material e.xamined. Holotypc. Vic. Bass Strait, S of

Point Hicks. 38°17.70'S I49°11.30'E, 400 m, coarse

sand, gravel, mud. many sponges, WIIOI cpibcnihic

sled. M.F. Gomon et al., 24 .lul 1986, ORV Franklin

(stn SLOPE-40). NMV J18685 (female, 4.8 mm, \vith

including 6 slides).

Paratypes (43 specimens). NSW. E of Newcastle

32°53'S I52®35’E, 146-175 m. 15 Aug 1985. pRv

Kapala (.stn K85-12-23). AM P38889 (5). Off Ecl^n

36°57.40 S 150°18.80'E, 220 m. muddy shell, Wt^Q]

epibenlhic sled, G.C.B. Poore et al.. 20 July 1986, ORV

Franklin (sin SLOPE-21), NMV J18681 (14) oty

Eden. 37°0.60'S 150°2().70'E. 363 m, coarse shell

WHOI epibenthic sled, G.C.B. Poore cl al., 21 July

1986, ORV Franklin (stn SLOPE-22), NMV Jl868^

( 1 ).

Vic. Bass Strait. S of Point Micks, 38®17.70‘S

149°11.30'E, 400 m. coarse sand, gravel, mud, many

sponges, WHOI epibenlhic sled, M.F. Gomon el al 24

Jul 1986, ORV Franklin (stn SLOPE-40), NMV J18683

(I), NMV J27641 (1 male, pleotelson and pleopods 1 -5

illustrated, 2 slides), NMV J18684 (2). 50 km S of

Mallacoota, 38^06.2*S 149°45.5'E, 188 m, WHOI

epibenlhic sled, R. Wilson, 14 Oct 1984, FRV Soela

(stn 505/84/30), NMV J18687 (5).

Tas. 15 km E of Mistaken Cape. Maria L, 42°37's

I48°20'E, 102 m, WHOI epibenthic sled, R. Wilson 9

Oct 1984, FRV Soela (sin S05/84/1), NMV J18686 (3)

30 km NNW of Cape Sorcll, 42“I0.9'S I44®48.9 'e!

160 m, WHOI epibenthic sled, R. Wilson. 20 Oct 1984*

FRV Soela (sin S05/84/54), NMV J18688 (4). 25 km W

of Port Davey. 45°23.3'S 145“39.8'E. 160 m, WHOI

epibenthic sled. R. Wilson, 21 Oct 1984, FRV Soela

(stn S05/84/60), NMV J18689 (6).

Description. Cephalon about third wider than

midline length; pscudorostrum as long as wide,

projecting to right angled point. Body without

setae, dorsally with 2 sharp, parallel, longitudinal

keels ninning on to proximal third of pleotelson.

Cephalon and percon margins smooth. Pleotelson

width about 1.5 length, with medioproximal keels

in continuation of pcrconite keels, low sinuous

keels in distal half terminating laterad of uropods.

lateral margins evenly curv'cd, minutely serrate,

distal margin strongly insinuated for uropods,

apex between uropods rounded right angled.

Antenna I of up to 5 articles; article 1 rectan¬

gular, distomedial comer with small teeth, width

0.75 length, about twice as long and wide as arti¬

cle 2 which is twice as long and wide as 3, article

5 as long as 3, both longer than 4, article 5 with 2

acsthetascs; lateral margins of articles I and 2

lined with cuticular outgrowth. Antenna 2, pedun¬

cle articles 1-4 of subcqual length, article 3

widest, article 4 third embedded into 3, article 5

ovate, nearly twice as long as 1^ combined.

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

305

Figure 1. Scaphojoeropsis multicarinata gen. nov., sp. nov. Reproductive structures of preparatory female

{paratype, NMV J18690), dorsal view; dorsal sculpture of body omitted; od, oviduct; ov, ovary; sd, spermathecal

duct. Scale bar: 1 mm.

306

J. JUST

Figure 2. Joeropsis bicahnata sp. nov., holotype, except pt: paratype, male, (NMV J27641). pt, male pleotelson,

ventral view, setae omitted; up, right uropod, ventral view. Habitus scale bar: 1 mm.

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

307

Figure 3. Joeropsis bicarinata sp. nov., holotype. a, right antennae 1 and 2, dorsal view; c, ventral view of cephalon,

setae omitted; mp, maxilliped, with enlarged ventral (v) and dorsal (d) view of mediodistal part of endite.

308

J. JUST

Figure 4. Joeropsis hicarinata sp. nov., holotype. II, lower lip; md, mandible; m, maxilla (1 and 2); ul, upper lip.

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

309

Figure 5. Joeropsis bicarinata sp. nov. M, paralype, male, (NMV J27641). H, holotype. pi. pleopod (1 and 2); d,

dorsal and v, ventral view.

310

J. JUST

Figure 6. Joeropsis bicarinata sp. nov., paratypc, male, (NMV J2764I). Pleopocis 3-5.

width two-thirds length, lateral margin fringed

with cuticular outgrowth, article 6 as long as 1-4

combined, distally expanded; flagellum about

fifth longer than peduncle article 6. with up to 10

articles, article 1 inflated, conjoint, twice length

of remaining articles combined.

Mouthparts: Mandibles, palp reaching apex of

5-denlate incisor, articles of subcqual length,

article 2 with distal tuft of long pectinate setae,

article 3 with terminal long pectinate seta and

subapical row of successively shorter similar

setae along distal half of article. Maxilla I outer

plate with 12 coarsely dentate setae and 2 shorter,

slender, simple setae; inner plate third width of

outer plate, reaching to mediodistal comer of

outer plate, with 3 long, simple setae and many

long setules distally. Maxilla 2, all 3 plates with 4

long, curved, finely pectinate setae, outer and

middle plates of similar shape and length, over¬

reaching inner plate with about third length, inner

plate with many long setules medially. Maxil-

lipeds covering about 0.6 ventral width of

cephalon. reaching forward beyond upper lip,

basis twice as long as greatest width, endite reach¬

ing to proximal third of palp article 4, with scat¬

tered simple setae on ventral surface and dorsal

field of slender, plumose setae inside medial

margin, mediodistal concavity of endite with 1

medial lump-shaped seta and a few short spear-

head-shaped setae, apical and lateral margins

evenly rounded, finely serrate; palp article 2 three

times longer than I, with bluntly triangular, for¬

ward pointing medial lobe reaching apex of

article 3, article 3 0.4 length of distally tapering

article 4, medial surface of article 4 covered with

short setules, article 5 minute; cpipod about as

long as palp, narrow, tapering to acute point.

Pereopods slender, similar, with 2 claws; basis

about 4 times longer than wide, twice length of

ischium. 2-7 with two penicillatc setae in proxi¬

mal half; mems about 0.8 length of ischium, dor-

sodistally expanded; carpus slightly wider than

and of subequal length to propodus, ventrodistal

comer with group of acute, forw'ard pointing

spines; propodus with 3 to 4 slender, robust setae

along ventral margin, single penicillatc seta dor-

soapically; dactylus of pereopod I about quarter

length of propodus, twice as long as wide, of

pereopods 2-7 about third length of propodus,

about 2.5 times longer than wide, dorsal and

ventral claws of equal length, dorsal claw more

slender.

Pleopods: Male plcopod 1 2.5 times longer than

greatest width, strongly tapering towards

midlength, margins only slightly diverging distal

to midpoint, apical lobes broadly rounded, set off

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

Figure 7. Joeropsis hicarinata sp. nov., holotype. Pereopods 1-5 and 7.

312

J. JUST

from lateral margins by pointed right angle, apical

lobes fringed with short, simple setae; dorsal sur¬

face with simple stylet guides running from sperm

duct opening to right-angled lateral comers; mid¬

dorsal surface with oval field of tiny setules,

ventral surface with 2 longitudinal rows of short,

simple setae. Male pleopod 2, protopod 2.5 times

longer than midwidth, proximal two-thirds of

lateral margin faintly convex, naked, distal third

curved mediad, slightly concave, with dense

fringe of long, thin cuticular scales, apex pointed;

stylet evenly curved, in retracted position reach¬

ing to apex of protopod. Female pleopod 2,

shaped like combined protopods of male pleopod

2, distal third triangular, lateral fringe of cuticular

scales about 1.2 times longer than in male, apex

rounded, ventral surface with scattered short,

simple setae including a couple at apex. Pleopod

3, endopod rounded rectangular, midsurface

length 1.5 times midwidth; exopod article 1 about

5 times longer than midwidlh, reaching just

beyond apex of endopod. lateral margin densely

fringed with cuticular scales, short and broad in

proximal two-thirds, longer and more slender dis-

tally; exopod article 2 articulating along medial

margin of article 1, barely half length of article 1,

distally tapering to blunt apex, lateral margin

fringed with long, slender cuticular scales, medial

margin with a few selules. Pleopod 4, endopod

subcqual in size and shape to endopod pleopod 3,

without setae; exopod a vestigial, rounded lobe

with apical setules, length about 1.3 width. Pleo¬

pod 5, similar to pleopod 4 in shape but about 1.2

times longer.

Uropods: Peduncles not reaching beyond

margin of pleotcison. broadly expanded medially,

mediodistal comer produced, acutely pointed,

ventrodistally with row of simple submarginal

setae, medial and lateral margins serrate; rami of

equal length, about quarter length of lateral mar¬

gin of peduncle, outer ramus twice as long as

wide, with apical row of long, simple setae, inner

ramus length 1.5 times width, with midventral

row of long, simple setae and several apical peni-

cillate setae.

Size. Largest male 5.5 mm; largest female 4.8

mm.

DistributiotL Eastern Australia south of 33®S,

eastern Bass Strait, east and west coasts of

Tasmania; 102-400 m.

Et}molog\\ The species is named for the two

prominent dorsal keels.

Remarks. The dorsal keels are present in speci¬

mens as small as 2 mm, although barely raised

from the dorsal surface, but they cannot be seen in

specimens of 1 mm.

Rugojoeropsis gen. nov.

Type species. Rugojoeropsis rugosa sp. nov.

Diagnosis. Joeropsididae with lateral body

margins parallel, with humps and ridges dorsally.

Cuticle roughly calcified. Cephalon with anterior

margin strongly concave. Pscudorostrum inserted

into concavity, box shaped, without overhanging

apical projection. Dorsolateral eye bulges without

ommatidia. Upper lip evenly rounded, less than

twice as wide as long. Mandibles with incisor of

5 strong subequal teeth in regular, curved row;

spine row setae long, in regular row. Lower lip,

lobes longer than wide distally tapering, pointed.

Maxillipeds in ventral view covering at least two-

thirds of cephalic width, and covering other

mouthparts except for medioapical concavity;

endite reaching beyond palp article 3; palp about

half length of endite: palp article 3 without medial

lobe, article 4 much longer than 3. Pleopod 2 of

female with a few tiny simple setae apically. Pleo¬

pod 3 exopod with strongly oblique articulation

between articles.

Included species. Rugojoeropsis rugosa sp. nov.

Etymology’. From the Latin rugosus = uneven,

rough, alluding to the texture of the cuticle.

Rugojoeropsis rugosa sp. nov.

Figures 8-13

Material examined. Holotype, Tas., off Frcycinet

Peninsula, 4l'’57.50'S 148°37.90'E, 400 m, coarse

shell, WHOI epibenthic sled. M.F. Gomon et al.. 27 Jul

1986. ORV Franklin (stn SLOPE-48), NMV J18678

(preparatory female, 2.8 mm. with 5 slides).

Paratypes (29 specimens). NSW. Off Eden

37‘^7.30'S 150°20.20'E. 550 m, grey coarse shell!

WHOI epibenthic .sled, G.C.B. Poore et al., 20 Ju! 1986

ORV Franklin (stn SLOPE-19). NMV J18676 (1).

Tas. Off Frcycinet Peninsula, 41°58.60'S

I48®38.80‘E, 500-600 m, coarse shell, WHOI epiben¬

thic sled, M.F. Gomon et al.. 27 Jul 1986. ORV

Franklin (stn SLOPE47), NMV JI8677 (15, maxil-

liped of 1 female illustrated. 1 slide). Off Frcycinet

Peninsula, 41°57.50'S 148''37.90'E, 400 m. coarse

shell. WHOI epibenthic sled, M.F. Gomon cl al., 27 Jul

1986, ORV Franklin (stn SLOPE-48), NMV J18679 (1

male, pleotcison and pleopods 1 and 2 illustrated, 2

slides), NMV J18680 (12, ventral view of spent female

illustrated).

Description. Cephalon twice as wide as midline

length, with serrate lateral margins, with a

rounded hump on each side medial to eyes; lobes

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

313

Figure 8. Rugojoeropsis rugosa gen. nov., sp. nov., holotype. a, antennae (1 and 2, right, dorsal view); up, uropod,

left, ventral view. Habitus scale bar: I mm.

314

J. JUST

Figure 9. Rugojoeropsis nigosa gen. nov., sp. nov. cl, cephalon left lateral view (holotype); pr, pseudorostrum, cv^

cephalon ventral view (paratype, spent female, NMV J18680).

surrounding midanlerior sinus raised to broad,

rounded keels; pseudorostrum in dorsal view with

evenly convex, finely serrate anterior margin,

width 1.3 length, barely reaching beyond

surrounding cephalic lobes; in lateral view

pseudorostrum vertically truncated, without over¬

hanging projection.

Pcrconitcs dorsally with 2 low, transverse,

rounded ridges, lateral parts of ridges more pro¬

nounced than central parts, ridges on pereonites 1

and 2 approximately half width of segment, suc¬

cessively shorter on more posterior pereonites,

lateral and lateroposlerior margins of pereonites

serrate and carrying scattered short, simple setae.

Plcotclson width about 1.1 times length, lateral

margins evenly curv-ed, coarsely serrate, distal

margin strongly insinuated for uropods, apex

between uropods rounded, slightly less than right

angled.

Antenna 1 article 1 rectangular, reaching about

halfway along cephalic sinus lobes, width two-

thirds length, about twice as long as and half as

wide again as article 2 which is twice as long and

wide as 3; tiagellum of up to 5 articles, article I

as long as peduncular article 3, article 2 twice

length of article 1 and as long as 3-5 combined,

articles 3-5 each with 1 long acsthetasc. Antenna

2, peduncle articles 1-3 of increasing length,

article 2 rectangular, article 3 medial margin

twice as long as lateral margin resulting in

strongly oblique apical margin, article 4 fully

embedded into 3, subcqual in size to article 2

article 5 elongate, poorly expanded laterally, as

long as 1 -4 combined along dorsal midline, width

0.5 length, margins fringed with culicular out¬

growth, article 6 three quarters length of 5, dis-

tally expanding; flagellum as long as peduncle

articles 4 and 5 combined, with up to 10 articles,

article 1 moderately inflated, conjoint, as long as

remaining articles combined.

Mouthparts: Mandible palp slightly overreach¬

ing apex of 5-dentatc incisor, articles 1 and 2 of

equal length, article 3 two-thirds length of 2,

article 2 with mediodistal row of pectinate setae,

article 3 with tenuinal long pectinate seta and

subapical row of successively shorter similar

setae along distal half of article. Maxilla 1 outer

plate with 12 coarsely dentate setae and 1 short,

stout simple seta at midfacc; inner plate 0.5 width

of outer plate, barely reaching mediodistal comer

of outer plate, with 3 long, mcdioapical setae and

many apical and distolateral long setules. Maxilla

2, outer and middle plates of subcqual size and

length about 1.3 times longer than inner plate,

with 4 unequally long, straight to moderately

curved, finely pectinate setae; inner plate with 3

similar setae and long medial setules. Maxillipcds

covering about two-thirds of ventral width of

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

315

Figure 10. Rugojoeropsis riigosa gen. nov., sp. nov., holotype. II. lower lip; md, mandible; m, maxilla (1 and 2);

ul, upper lip (apical margin folded back artificially).

cephalon, reaching forward beyond upper lip,

basis twice as long as greatest width, endite reach¬

ing to about middle of palp article 4, with dorsal

Held of slender setulcs inside medial margin,

mediodistal concavity with row of 4 low, lump-

shaped setae; apical and lateral margins of endite

evenly rounded, llnely serrate, with single slender

sctule in each indentation and small peg-shaped

seta anterior to concavity; palp article 2 three

times longer than 1 along midsurface, with for¬

ward pointing medial lobe reaching about two-

thirds along article 3, apex of lobe with a few long

slender setae, article 3 about half length of distally

tapering article 4, medial margin of 2 and 3

fringed with culicular outgrowth, medial surface

of article 4 covered with short setulcs, article 5

0.2 length of 4, rounded, with tuft of long, simple

setae; epipod about as long as palp, width about

third length, tapering to acute point.

Pereopods similar, with 2 claws; basis about 4

316

J. JUST

Figure 11. Rugojoeropsis mgosa gen. nov., sp. nov. pt, pleotelson, ventral view (paratype, male, NMV J18679);

mp, maxilliped (paratype, female, NMV J18677); pl2, pleopod 2 (holotype).

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

317

Figure 12. Riigojoeropsis rugosa, gen. nov., sp. nov. Paratype, male, (NMV J18679): pll, pleopod 1, dorsal view;

pl2, pleopod 2, ventral view. Holotype, pl3-5, pleopods 3-5, (apex of pl4 artificially folded).

318

J. JUST

Figure 13. Rugojoeropsis nigosa gen. nov., sp. nov., holotype. Pereopods 1-5 and 7.

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

319

times longer than wide, twice length of ischium,

with 1 or more pcnicillale setae on proximal half

of dorsal margin (not verified in pereopod I);

meriis about 0.6 length of ischium, dorsodistally

expanded; carpus slightly wider than and of sub-

equal length to propodus, single bottlebrush seta

on dorsoapical margin, ventrodistal comer of

carpus of some percopods with group of acute,

forward pointing spines, (verified on percopods I,

2 and 3 only in several specimens); propodus with

3 slender, robust seta along ventral margin,

single penieillalc seta dorsoapically; dactylus of

pereopod 1 about third length of propodus, of

percopods 2-7 about 0.4 length of propodus, all

about twice as long as wide, dorsal and ventral

claws of subequal length, dorsal claw more

slender.

Pleopods: Male picopod 1 2.3 times longer than

greatCvSt width, tapering towards distal two-thirds,

margins only slightly diverging in distal third,

apical lobes broadly rounded, set oft' from lateral

margins by acute backward pointing projection

reaching about two-thirds along apical lobes,

apical lobes fringed with simple setae; dorsal sur¬

face with simple stylet guides running from spenn

duct opening to acute lateral comers, covered by

extended lateral edge of stylet guide; middorsal

surface with oval field of tiny setules. Male pico¬

pod 2, protopod length 2.25 times midwidth,

proximal 0.8 of lateral margin evenly convex,

distal fifth concave, apex pointed, entire lateral

margin with dense fringe of elongate cuticular

scales serrate along anterior edge (cf female. Fig.

II, pl2); stylet evenly cur\^ed, in retracted

position reaching to apex of protopod. Female

pleopod 2, shaped like combined protopods of

male pleopod 2 except distolateral concavity

about third length of lateral margin, lateral fringe

of cuticular scales as in male, apex rounded with

a few tiny simple setae. Pleopod 3, endopod

rounded rectangular, width about 0.7 midsurface

length; exopod article 1 about 0.9 length of endo¬

pod, barely reaching apex of endopod, lateral

margin densely fringed with long, narrow cuti¬

cular scales; exopod article 2 articulating along

medial margin of article 1, about half length

article 1, reaching beyond endopod by third

length, distally tapering to rounded apex, lateral

margin fringed with long slender cuticular scales,

medial margin fringed with short setules proxim-

ally and short simple setae distally. Picopod 4,

endopod of subcqual size and shape to endopod

pleopod 3; exopod a vestigial rounded lobe with

apical setules, about six times longer than wide.

Pleopod 5 similar to pleopod 4 endopod in size

and shape.

Uropods: Peduncle broadly expanded medially,

mediodistal comer reaching beyond apex of

pleotclson, vcntrodistally with a few simple

submarginal setae, distal half of medial margin

strongly serrate, mediodistal comer produced

mediad, acutely pointed; rami of unequal

length, outer ramus two-third length of inner

ramus, with apical tuft of long simple setae, inner

ramus about two-third length of lateral margin of

peduncle, width 0.4 length, with midventral row

of long, simple setae and several apical pcnicillatc

setae.

Size. Largest male 2.8 mm; largest female 2.8

mm.

Distribution. Eastern Australia south of 37°S,

eastern Bass Strait, east coast of Tasmania;

400-600 m.

Etymiology. From the Latin nigosus = uneven or

rough, alluding to the texture of the cuticle.

Scaphojoeropsis gen. nov.

Type species. Scaphojoeropsis multicarimita sp.

nov.

Diagnosis. Joeropsididac with body strongly

tapering in dorsal view from pereonitc 2 to apex

of picotelson, with variety of dorsal humps and

short keels. Cuticle roughly calcified. Cephalon

with midanterior dorsal surface strongly

depressed, depression surrounded by sharp ridge,

anterior margin straight. Pseudorostmm pointing

downward. Joining vertex along straight line,

without apical projection. Eyes absent. Upper lip

more than twice as wide as long. Mandibles with

incisor divided into 2 parts, with large tenninal

tooth pointing foi-ward, posterior 2-3 teeth on

broad flange pointing mediad to backwards; spine

row setae short, stubby, in irregular row. Lower

lip, lobes wider than long, rectangular, lateral

corners projecting, pointed. Maxillipeds in

ventral view covering about third of cephalic

width, not reaching forward to cover mandibles;

endite reaehing to middle of palp article 2; palp

0.8-0.9 length of endite; palp article 3 with dis¬

tinctive medial lobe; article 4 shorter than 3. Pico¬

pod 2 of female with long apical setae. Picopod 3

exopod with transverse articulation between

articles.

Included species. Scaphojoeropsis kimhlae sp.

nov., S. muhicarinata sp. nov,

Etymoiog}\ From the Greek skaphe = boat, allud¬

ing to the body shape in dorsal view.

320

J. JUST

Scaphojoeropsis multicarinata sp. nov.

Figures 14-18

Material examined. Uolotype. Vic. Bass Strait, S of

Point Hicks. 38°I4.80'S, 149°9.30'E, 200 m, coarse

sand and gravel, WHOl epibenthic sled, M.F. Gomon et

al., 24 Jul 1986, ORV Franklin (stn SLOPE-4]). NMV

J18691 (female, 2.2 mm, with 6 slides).

Paratypes (11 specimens). NSW. Off Nowra.

34”59.52’S, !51“5.94'E, 204 m, coarse shell, WHO!

epibenthic sled, G.C.B. Poore ct al., 14 Jul 1986, ORV

Franklin (stn SLOPE-I). NMV J18690 (4. 1 female

illustrated, see Fin. 1).

Vic. S of Point Hicks. 38^14.80 5, 149°9.30'E 200

m, coarse sand and gravel. WHOl epibenthic sled. M.F.

Gomon et al.. 24 Jul 1986. ORV Franklin (stn SLOPE-

41), NMV J277I9 (I male, pleotelson and pleopods 1

and 2 illustrated, 1 slide). 50 km S of Mallacoota.

38^06.2 8. 149°45.5 E, 188 m. WHOl epibenthic sled.

R. Wilson. 14 Oct 1984. FRV Soela (stn SO5/84/30),

NMV J18694 (3).

Tas. Bass Strait, 63 km E of North Point, Flinders L,

39°44.8'S, 148°40.6'E, 124 m, fine sand and mud. SM

grab. R. Wilson. 14 Nov 1981. RV Tangaroa (stn BSS

167), NMV JI8692 (2). Bass Strait, 100 km NE of

North Point. Flinders L, 38°5t.58'S 148'^26.5'E. 130 m.

fine sand. SM grab, R. Wilson, 15 Nov 1981. RV

Tangarua (stn BSS 170), NMV J18693 (1).

Description. Ccphalon 2.5 times wider than mid-

line length, midanterior margin broadly concave

in dorsal view, concavity rimmed by raised keel,

with 2 keels reaching backwards halfway along

cephalon on each side of midline, small keel pre¬

sent immediately lateral to posterior part of back¬

ward reaching keels, low keel present antcrolater-

ally on each side fonning recurved loop lateral to

insertion of antenna 1; pseudorostrum evenly

convex.

Body broadest at pereoniles 2 and 3. Pcrconilcs

with middorsal longitudinal keel, that ofpereonite

1 divided into short anterior and longer posterior

part, perconite 1 with small crescent shaped keels

lateral to each part of midkcci, pcrconitcs 2- 7

with single middorsal keel and crescent shaped

mid-lateral keels, anterior parts of midand lateral

keels linked by low serrate ridge; pereonites 1-4

of equal length, pcrconitcs 5-7 of equal length,

0.8 length of more anterior pereonites. Pleotelson

width about 1.1 length, with middorsai longitudi¬

nal keel reaching to base of apical lobe; lateral

margins smooth except for fringe of cuticular out¬

growth, evenly tapering towards broadly rounded

apex, with deep insinuation for uropods each side

of apex.

Antenna 1 of 6 articles; article 1 rectangular,

not reaching lateral comer of frontal sinus, width

two-thirds length, about twice as long as and third

as wide again as article 2 which is barely twice as

long and wide as 3, articles 2 and 3 with lateral

Bange of cuticular outgrowth; tlagellum of 3

articles, article 1 two-thirds length of peduncle

article 3, article 2 as long as peduncle article 3,

article 3 as long as and half width of 1, articles 2

and 3 each with I long aesthctasc. Antenna 2,

peduncle articles 1-3 of slightly increasing length

and width, article 3 dislally expanded, distal

width twice midlcngth, article 4 third length of 3,

about 5 times wider than long, halfway embedded

into 3, article 5 nearly twice as long as 1-4

combined, broadly expanded laterally, width,

excluding broad fringe of cuticular outgrowth,

two-thirds length, article 6 about 0.4 length of 5,

expanded dislally; nagcllum as long as peduncle

article 6, with up to 6 articles, article 1 not

inflated, apparently not conjoint, as long as

articles 2 and 3 combined.

Mouthparts: Upper lip nearly rectangular, distal

margin slightly convex, fringed with setulcs, dis-

tolaleral comers distinct, slightly projecting.

Mandible palp slender, not reaching to near apex

of forward pointing tooth of incisor, article 2 0.6

length of 1, article 3 six-sevenths length of 2,

article 2 with 2 mediodistal pectinate setae, article

3 with 2 temiinal pectinate seta; incisor with I

large forward pointing tooth and 3 mediad to

backward pointing teeth on broad flange, forward

pointing tooth of right mandible with lateral ser¬

ration along proximal two-thirds; spine row of

few short, stout, minutely pectinate setae, left

with row divided into 2 groups (distalmost

slightly stouter non-pectinatc seta possibly

homologous with lacinia mobilis). Lower lip,

about 3 times broader than long, body rectangu¬

lar, distolateral parts expanded to wing-like pro¬

jections with slightly acute anterior and posterior

comers, lateral margins convex. Maxilla 1 outer

plate with 10 unequally long finely dentate setae

and 3 short simple setae on middorsal margin;

inner plate with evenly rounded apex half width

of outer plate, reaching to middle of medial

margin of outer plate, with 2 apical simple setae

and several short apical seiules. Maxilla 2, inner

and middle plates of equal length and width, 0.75

length and 0.6 width of outer plate with 2 subc-

qual. straight, finely pectinate setae tenninally,

outer plate with 3 finely pectinate setae, apical

one as long as plate, both subapicals much shorter

and more slender. Maxillipeds covering about

third ventral width of cephalon, endites reaching

forward to cover bodies of maxillae but not

mandibular incisors or upper lip; basis twice as

long as midwidth, endile reaching to about middle

of palp article 2, mediodistal concavity vestigial

with single multidentate lump-shaped seta;

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

321

Figure 14. Scaphojoeropsis multicarinata gen. nov., sp. nov., holotype. c, cephalon, anterolateral view; ptd, apex

of pleotelson, dorsal view; ptv, apex of pleotelson, ventral view, setae omitted. Habitus scale bar: I mm.

322

J. JUST

Figure 15. Scaphojoeropsis midticarinata gen. nov., sp. nov., holotype. a, antennae (1 and 2); c, cephalon, ventral

view.

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

323

Figure 16. Scaphojoeropsis multicahnata gen. nov., sp. nov., holotype. II. lower lip; nid, mandible (I, left, r, right);

m, maxilla (1 and 2), mp, maxilliped; ul, upper lip.

324

J. JUST

Figure 17. Scaphojoeropsis midticarinata gen. nov., sp. nov., holotype, except M: paratype, male, (NMV J27719).

pi, pleopod (1-5); pt, pleotelson ventral view; x, apex of female pleopod 2.

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

325

Figure 18. Scaphojoeropsis miilticarinata gen. nov., sp. nov., holotype. Pereopods 1-3, 5-7.

326

J. JUST

distolateral margin of endite evenly rounded,

finely serrate, with single slender sctulc in each

indentation, apical margin somewhat truncate

with a few short simple setae; palp articles 2 and

3 as broad as or broader than endite, palp article 2

2.5 times longer than 1 along midsurfacc, with

acute, mediad pointing medial lobe and acute for¬

ward pointing distolateral lobe reaching halfway

along article 3; article 3 subequal to 2 in length

and width, medially expanded to large, forward

pointing, rounded lobe, medial margins of articles

2 and 3 with coarse fringe of cuticular outgrowth,

lateral margin w ith tiny etules; article 4 two-thirds

length of 3, medial surface with selulcs in middle

third; article 5 about third length of 4, rounded,

w'ith lufl of long, simple setae: cpipod as long as

palp articles 1,2 and half of 3 combined, tapering

to blunt point.

Pereopods similar, with 2 claws; basis about 5

limes longer than wide, twice length of ischium,

with 1 or more penicillate setae on proximal half

of dorsal margin; merits about two-thirds length

of ischium, with broadly rounded expansion dor-

sodistally: carpus and propodus length subequal

to ischium, w'ith 1 or 2 penicillate setae on dor-

soapical margin (not verified on pereopod 1);

propodus with 3 slender, robust setae along

ventral margin; dactylus of all pereopods about

two-thirds length of propodus, about twice as

long as wide; dorsal claw slightly longer than

ventral claw.

Plcopods; Male pleopod 1 2.3 times longer than

greatest width, tapering towards distolateral cor¬

ners, apical lobes evenly rounded joining lateral

margins at wide angle, apical lobes fringed with

short setulcs and medium to long simple setae;

dorsal surface with simple, uncovered stylet

guides running from sperm duct opening to lateral

comers. Male pleopod 2 protopod 2.5 limes

longer than midwidth, proximal two-thirds of lat¬

eral margin evenly convex, distal third concave,

apex pointed, distalmost part of lateral margin

with overlapping rectangular cuiicuUir scales;

stylet evenly curved, in retracted position reach¬

ing to apex of prolopod, with group of liny teeth

laterally one sixth length from apex. Female pleo¬

pod 2 shaped like combined protopods of male

pleopod 2 except apex broadly rounded with

fringe of long, simple setae; marginal cuticular

scales as in male. Pleopod 3 endopod rectangular,

width 0.6 length of medial margin: exopod barely

reaching apex of endopod, article 1 as long as

endopod, article 2 articulating with 1 at nearly

right angle, twice as long as w'idc, about third

length of article I, distally tapering to blunt apex;

lateral margin of both articles fringed with robust

elongate cuticular scales. Pleopod 4 endopod

subcqual in length to endopod pleopod 3, tapering

to rounded apex; exopod a tiny rounded lobe

without sctiiles. Pleopod 5 similar to pleopod 4

endopod in shape.

Uropods extending beyond apex of plcolclson

by third their length; peduncle with broad,

rounded medioapical, backward pointing projec¬

tion covering length of inner ramus, medial length

of peduncle including medial projection 1.5 mid¬

width; rami of unequal length, outer ramus third

to half length of inner ramus, with tuft of long,

simple apical seta; inner ramus width 0.5 length,

with single, simple seta apical ly and a few setae

on ventral surface.

Size. Largest male 1.6 mm; largest female 2.6

mm.

Distribution. Eastern Australia south of 35°S, to

eastern Bass Strait; 124-204 m.

Etymology. The specific epithet refers to the

many short dorsal keels.

Remarks. Scaphojoeropsis miilticarinata is most

easily distinguished from S. kimhiae sp. nov.

(condition in parentheses) by its dorsal carinae on

pereon and pleotelson (no carinae), iiropod

peduncle with mediodistal projection (no projec¬

tion), maxilla 2 with full complement of three

lobes (reduced to one lobe), and male pleopod 1

with rounded apex (apex with two acute lobes).

Scaphojoeropsis kimhiae sp. nov.

Figures 19-22

Material examined. Holotype. Tas. 60 km E of North

Point, Flinders I.. 39°41.7'S. 148°39.5’E, 115 m.

muddy sand, naturalists' dredge, G.C.B. Poore, 27 Mar

1979, UMAS Kimhla (stn HSS 32), NMV J18695

(male, 2.1 mm, with 7 slides).

Paratypes (4 specimens). NSW. OH* Eden, 37“0.60'S

150°2().7()'E, 363 m, coarse shell. WIlOl cpibcnthic

sled. G.C.B. Poore et al., 21 Jul 1986, ORV Franklin

(sin SLOPB-22). NMV J186% (3, one without

cephalon).

Tas. 60 km E of North Point, Flinders I., 39'’4I.7'S.

148^39.5'E, 115 m. muddy sand, naturalists' dredge.

G.C.B. Poore, 27 Mar 1979, UMAS Kimhla (stn BSS

32), NMV J27720(! female, with 1 slide).

De.scription. Cephalon about 4 limes wider than

dorsal midlength, midantcrior margin nearly

straight in dorsal view, depressed anterior part

rimmed by raised concave keel, with 2 straight

keels reaching backwards from lateral parts of

concave rim haUway along cephalon, small hump

present on both sides posterior to insertion of

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

327

Figure 19. Scaphojoeropsis kimhlae gen. nov., sp. nov., holotype, except F: paratype, female, (NMV J27720). a,

antennae (1 and 2), right, dorsal view; c, ccphalon, ventral view; pl2, female pleopod 2; up, uropods, ventral view;

pt. pleotelson, ventral view, setae omitted. Habitus scale bar: 1 mm.

328

J. JUST

Figure 20. Scaphojoeropsis kimblae gen. nov., sp. nov., holotype, except F: paratype, female, (NMV J27720). IF

lower lip; md, mandible (I, left, r, right); m, maxilla (1 and 2); mp, maxilliped; ul. upper lip.

antenna 1; pseudorostrum broad, convex. Body

broadest at pereonite 2. Pereonites 1, 3 and 4 of

subequal length, 2 about 1.2 times longer, 1 with

low midanterolatcral humps, 1^ with larger pos¬

terolateral humps; pereonites 5-7 of equal length,

three-quarters length of 4, posterolateral humps

joined across posterior half of pereonite to form

low, rounded ndge. Pleotclson length about 1.1

width, lateral margins finely, irregularly serrate,

convexly tapering to narrow point at three-

quarters length and further to broadly rounded

apex; apex without incisions foruropods in dorsal

view, Liropods barely visible in dorsal view.

Antenna 1 of 7 articles; article 1 distally taper¬

ing, reaching lateral comer of frontal ridge, width

at base 0,8 length, about 2.5 times longer than

article 2, articles 2 and 3 of subequal length, 3

about two-thirds width of 2; llagellum of 4

articles, article 2 length 1.25 times 1, articles 2

and 4 with 1 long acsthetasc (broken off in 3?)*

Antenna 2, (configuration of peduncle articles

1-4 not clarified, possibly as in S. multicarinata,

at least article 4 very short), article 5 moderately

expanded laterally, width, excluding broad cutic-

ular fringe, 2.3 times length, article 6 length 1.5

width, about 0.25 length of 5; flagellum barely 3

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

329

Figure 21. Scaphojoeropsis kimblae gen. nov., sp. nov., holotype. pi, pleopod (1-5); p, pereopod (I and 2); d. dor¬

sal view; v, ventral view.

330

J. JUST

Figure 22. Scaphojoeropsis kimhlae gen. nov., sp. nov., holotype. Pereopods 3-7.

JOEROPSIDIDAE FROM SOUTH-EASTERN AUSTRALIA

331

times longer than peduncle article 6, with up to 8

articles, article I not inilated, apparently not

conjoint, as long as articles 2-7 combined.

Mouthparts; Upper lip, length about 0,4 width,

anterior margin convex, notched in middle,

(ringed with long setules, lateral margins

rounded. Mandible palp slender, reaching to mid¬

dle of incisor, article 2 about iwo-ihirds length of

1, article 3 0.8 length of 2, article 2 with 2

mediodistal pectinate setae, article 3 with 3 ter¬

minal pectinate seta; incisor with 1 forward point¬

ing tooth and 3 mediad pointing teeth on broad

flange, forward pointing tooth of right mandible

with small accessory tooth laterally; spine row

composed of 4 short, peg-like setae on conical

projections. Lower lip about 3 times broader than

long, body rectangular, distolateral comers pro¬

jecting forward to rounded apices. Maxilla 1 outer

plate with 10 unequally long completely smooth

setae; inner plate a vestigia! rounded lobe without

setae. Maxilla 2 composed of single plate, about 3

times longer than wide, with 2 pectinate robust

setae apically. Maxillipeds covering about third

of ventral width of cephalon, endites reaching for¬

ward to cover bodies of maxillae but not

mandibular incisors and upper lip; basis length

2.5 times midwidth, reaching about two-thirds

along palp article 2, mediodistal concavity

vestigial with single peg like robust seta; lateral

margin ofendite evenly rounded, coarsely serrate,

apical margin somewhat truncate with 2 peg-like

robust setae and a few short simple setae; palp

article 2 broader than enditc; palp article 1 with

short distolateral projection; article 2 broadly

expanded medially, about 3 times longer than I

along midsurface; article 3 about half length and

width of 2, with medial rounded lobe; media!

margins of articles 2 and 3 and lateral margin of

1, 2 and 3 with coarse fringe of cuticular out¬

growth; articles 4 and 5 of subcqual length and

width, combined as long as 3; cpipod reaching to

insertion of palp, 2.3 times longer than wide,

rounded apex with fringe of cuticular outgrowth.

Percopods similar, with 2 claws; basis about

4.5 times longer than wide, twice length of

ischium; 2-7 with 1 or 2 small penicillatc setae

midway on dorsal margin; mcrus about two-thirds

length of ischium, with broadly rounded expan¬

sion dorsodistally; carpus and propodus of sub-

equal length, about 0.8 length of ischium, with

single large penicillatc seta on dorsoapical

margin; caqms and propodus with 1 or 2 slender,

robust setae along ventral margin; fringe of cutic¬

ular outgrowth along dorsal margin of basis and

ventral margin of mcrus, carpus and propodus;

dactylus of all pereopods about half length of

propodus, about 2.5 times longer than wide;

dorsal claw subequal in length to and more slen¬

der than ventral claw.

Plcopods; Male pleopod 1 midlength about 1.7

times greatest width, tapering towards midpoint,

slightly expanding in distal half towards right

angled lateral corners; apical lobes acutely

pointed, separated in midline at about 90 degrees,

lateral margins concave, laterally with small

simple setae, medially with long simple setae,

apices with fringe of small setules; dorsal surhice

with funnel shaped, uncovered stylet guides run¬

ning from sperm duct opening to lateral comers.

Male pleopod 2, prolopod 2.5 limes longer than

midwidth, apex acutely pointed, entire lateral

margin with fringe of cuticular scales (details not

available); stylet evenly curved, in retracted posi¬

tion reaching to about apex of prolopod. Female

pleopod 2, shaped like combined protopods of

male pleopod 2 except apex broadly rounded with

fringe of long, simple setae, (lateral cuticular

scales not observed). Pleopod 3, endopod rectan¬

gular, width about two-thirds length of medial

margin; exopod barely reaching apex of endopod;

article I three-quarters length of endopod; article

2 articulating with 1 at right angle, 3 times as long

as wide, about 0.6 length of article I, distally

tapering, apex rounded; lateral margin of both

articles fringed with short, overlapping rectan¬

gular cuticular scales. Pleopod 4, endopod subc¬

qual in length to endopod pleopod 3, ovoid, with¬

out setae but with a few apical cuticular combs;

exopod about quarter length of endopod, with

single long pectinate seta apically. Pleopod 5

similar to pleopod 4 endopod but more slender.

Uropods: Peduncle without medial projection,

medial margin as long as peduncle width at base,

with acute denticles in midlhird; rami of unequal

length, outer ramus third length and width of

inner ramus, with a few long simple setae; inner

ramus two-thirds length of peduncle inner

margin, width 0.6 length, with simple setae and

penicillatc setae apically.

Size. Largest male 2.1 mm; largest female 2.1

mm.

Distribution. Eastern Australia south of 37°S to

eastern Bass Strait; 115-363 m.

Etymolog\K The species is named after HMAS

Kimbla.

Remarks. See previous species for characters

to separate Scaphojoeropsis kimblae from S.

multicarinata.

332

J. JUST

Acknowledgments

The bulk of the material studied was obtained

during two cruises of the Australian National

Facility ORV Franklin in 1986 and 1988 as part

of the South-Eastern Australian Slopes program

jointly organised by Museum Victoria and the

Victorian Institute of Marine Sciences (VIMS, no

longer in existence), Melbourne. The program,

and my participation and subsequent research at

Museum Victoria, were supported by a grant from

the Australian Marine Sciences and Technology

Scheme/Australian Research Council. Most of the

remaining material was collected by Museum

Victoria during studies in Bass Strait. 1 thank Dr

Penny Berents, Australian Museum, Sydney, for

loan of additional material, and Dr Buz

Wilson, Australian Museum, Sydney, and Dr

Gary Poore, Museum Victoria, for valuable

discussions and technical support.

References

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(isopode asellote) ct les Jaeropsis mediter-

raneennes. Recueil de Travaivc dc la Station

Marine d'Endoume 23 (37):121-I29.

Barnard, K.ll. 1965. Isopoda and Aniphipoda collected

from Gough Island Scientific Survey. Annals of the

South African Museum 48 (3): 195-210.

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Memoirs of Museum Victoria 58(2): 335-345 (2001)

PLATYPROTUSPHYLLOSOMA, GEN. NOV., SP. NOV., FROM ENDERBY LAND,

ANTARCTICA, AN UNUSUAL MUNNOPSIDID WITHOUT NATATORY

PEREOPODS (CRUSTACEA: ISOPODA: ASELLOTA)

Jean Just

Museum of Tropical Queensland, 78—102 Flinders Street, Townsville, Queensland 4810, Australia

Abstract

Just, J., 2001. Platyprotus phyllosoma, gen. nov and sp. nov., from Enderby Land, Antarctica,

an unusual munnopsidid without natatory pereopods (Crustacea: Isopoda: A.sellota). Memoirs

of Museum Victoha 5%{2)\ 335-345.

A new genus and species. Platyprotus phyllosoma. of munnopsidid isopod is described Irom

Enderby Land, Antarctica. Platyprotus differs from all other miinnopsidids, except Micropro-

tiis Richardson. 1910 and some species in Storthyugurella Malyutina, 1999, in having fully

ambulatory pereopods 5-7 without natatory setae. Platyprotus differs from those genera pri¬

marily in body shape, lack of marginal spines on pereonites 5-7 and pleotelson, lack of a pre-

anal ridge, and in possessing highly derived spatulate setae on article 3 of the mandibular palp.

All three genera are part of the wider ^Storthyngura' clade. If the present concept of Storthyn-

gurella is confirmed, ambulatory pereopods 5-7 have been derived independently more than

once within that clade. The family name is emended to Munnopsididac Hansen, 1916.

Introduction

The Munnopsididac are a highly diverse family

occurring in all oceans primarily in deep water

from upper slope to hadal depths, as well as at

shelf depths in polar and subpolar regions. The

natatory pereopods 5-7 with expanded, flattened

carpi and propodi fringed with long plumose

natatory setae generally have been considered the

most inimediardy recognisable synapomon-jhy in

the Munnopsididac (Wilson, 1989).

Wilson ct al. (1989) transferred Microprotus

Richardson, 1910 from the Janiridae to the

Munnopsididac. The five known species in

Microprotus have ambulatory pereopods 5-7

with tubular carpi and propodi without plumose

natatory setae. Wilson (1989) recognised seven

subfamilies in the Munnopsididac, but left sev¬

eral genera, including Microprotus and Storthyn-

gura Vanhoffen, 1914 as incertac sedis. Wilson ct

al. (1989) considered Microprotus to be most

closely related to the heterogeneous Storthyngura

complex.

Malyutina (1999) described a new genus, Star-

thyngurella, with eight species from abyssal to

upper hadal depth, five of which were transfeiTcd

from the Storthyngura complex. Two of the

species referred to Storthyngiirella, S. menzies

Malyutina and S. triplispinosa Menzies, have

ambulatory pereopods 5-7. The remaining

species, including the type species of Storthyn-

gureUa, S. hirsuta Malyutina, 1999, have slender

natatory pereopods 5-7 with short plumose

setae along the posterior margin of carpus and

propodus.

This paper reports on a new munnopsidid

species from Antarctica, Platyprotus phyllosoma

gen. nov. and .sp. nov., with ambulatory pereo¬

pods 5-7 and no natatory setae. Platyprotus

shares with Storthyngura and Microprotus a suite

of characters listed by Wilson (1989; 343) as

strong indicators of a monophyletic taxon.

Malyutina (1999: 269) added Storthyugurella to

this clade. Platyprotus differs from the other

three genera mentioned, notably in body shape, in

the lack of body spines along the margins of pere¬

onites 5-7 and tlie pleotelson, in the complete

lack of a prcanal ridge on the pleotelson, and in

the uniquely spatulate setae on article 3 of the

mandibular palp (Table I). These differences are

considered aulapomorphic.

Wilson ct al. (1989) concluded that the ambu¬

latory pereopods 5-7 of Microprotus are an

apomorphic reversal within the Munnopsididac,

derived from the natatory pereopods of an ances¬

tral Storthyngura. The presence of a group of

species with ambulatory pereopods within Stor-

thyngurella., and the discovery of Platyprotus, do

not disprove this hypothesis. The question arises,

however, as to whether this revensal combines

Microprotus, Storthyngurella and Platyprotus in

a monophyletic taxon. For that hypothesis to be

accepted S. menzies and S. triplispinosa would

have to be, a priori, recognised as a separate

335

336

J. JUST

Table 1. Characters of presumed generic significance among Microprotus^ Storthyngurella (characters

in bold refer to species without natatory pereopods), Storthyngitra and Platyprotus gen. nov.

Character

Microprotus

Storthyngurella Storthyngura

Platyprotus

Body shape

subcylindrical

flat

Dorsal spines on cephalon

absent

present

present/absent

absent

Lateral spines on pereonites number

2-7

4-7

1/3/4-7

Lateral spines on pleotelson

present

(absent*)

absent

Apical spines on pleotelson

variable/absent

absent

Pleotelson articulation

fused

free/fused

fused/free

free

Preanal ridge

present

absent

Medial spine on article 1 of antenna 1

absent

present

present/absent

present

Mandibular palp article 3 setae

simple

complex,

spatulate

Maxilliped epipod lateral tooth

present

or absent

present

or absent

absent

Male pleopod 2, apex of basis

acute

rounded

acute

rounded

Male/female pleopod 2 setae

simple

variable; no

setae to short

plumose

long

hemiplumose

Plcopod 3, endopod setae

numerous

3-numerous

♦Two of the many Storthyngura species, S. eltaniae Georges and Menzies, 1968 and S. torbeni George,

1987 lack pleotelson spines; both have a simple pleotelson of EiaycopeAxkc. shape.

(new) genus, since the remaining species in

Storthyngurella have slender natatory pereopods

5-7. The alternative is to accept that ambulatory

pereopods 5-7 were derived more than once as

part of an evolutionary trend within the ''Storthyn¬

gura clade. This is the only possible hypothesis

if the present concept of Storthyngttrella is con¬

firmed. Table 1 does not provide answers to these

questions, which can only be addressed by a full

cladistic analysis of the entire "Storthyngura^

clade.

Munnopsididae Hansen, 1916, nom. emend.

Remarks. Most prior authors have referred to this

family as Munnopsidae (or erroneously as

Eurycopidae). G.D.F. Wilson (pers. comm.) has

drawn my attention to the fact that the genitive of

the latinised adjectival ending -apsis (Greek, like)

of Munnopsis is -opsidis and the family name is

here emended to Munnopsididae.

Platyprotus gen. nov.

Type species. Platyprotus phyllosoma sp. nov.

Diagnosis. Munnopsididae with pereopods 5-7

ambulatory, without natatory setae. Body flat¬

tened, widening from pereonite 1 to pleo¬

telson. Pleotelson free. Middorsal spines short.

triangular. Lateral spines (not coxal spines) on

pereonites 3 and 4 only. Pleotelson without

marginal spines. Pleotelson without preanal ridge.

Antenna I article 1 with strong medial spine.

Antenna 1 fiagellum with aesthetascs on distal-

most half of articles. Mandible with posterior pro¬

jection articulating with cephalon in short notch.

Mandibular palp article 3 with highly modified

spatulate setae. Mandibular incisor teeth distinct.

Maxilliped epipod lateral margin smooth. Male

plcopod 2 protopod with broadly rounded apex,

fringed with long hcmiplumosc setae. Plcopod 3

endopod with 3, exopod with 9 pappose setae.

Etymology. The name combines references to the

flat body and to the similarity of pereopods 5-7 to

Microprotus.

Platyprotus phyllosoma sp. nov.

Figures 1-8

Material examined. Holotype. Off Endcrby Land.

Antarctica (65°5().rs, 50°34.30'E-65®50.1()'S,

50°34.90'E), 540 m-2°C, mainly rocky bottom with

mud/clay patches, dominant biota; crustaceans, ophi-

uroids, bryozoans, hard coral, 20 Nov 1985, M. Norman

on hlella Dun^ WHOI cpibcnthic sled (stn. HRD-011).

Museum Victoria, J47017 (male, 5 mm, with 5 slides —

pleopod 1 missing, presumably lost during collecting).

AN UNUSUAL MUNNOPSIDID ISOPOD FROM ANTARCTICA

337

Figure 1. Platyprotus phyllosoma sp. nov., holotype. Scalebar: 1 mm.

Paratypc. Same data as holotype. Museum Victoria,

J47I35 (female, 4.1 mni).

Description (holotype, male). Body broad, flat¬

tened; width increasing from pcrconitc 1 to 7,

pereonite 7 about 2.3 times wider than 1.

Cephalon without rostrum, 2.5 times wider than

long, 20% wider than pereonite 1; with strong

frontal ridge, epistome broadly rounded in lateral

view.

Pereon: Pereonites 1-4 with I short, dorsal, tri¬

angular, forward pointing spine on midanterior

margin; pereonite 1 with single long coxal spine;

pereonite 2-4 with bifurcate broad coxal spine,

anterior part more than twice length and width of

338

J. JUST

Figure 2. Platyprotus phyllosoma sp. nov., holotype, except ul; paratype female, al, antenna 1; a2, peduncle of

antenna 2; II, lower lip.

posterior part; perconites 3 and 4 with single

anterolateral spine. Perconite I about 75% length

of 2, perconites of equal length; pereonites

5-7 fused, 5 with midantcrior sinus flanked by 2

short, pointed spines. 6 and 7 with 2 broad,

apically rounded spines near anterior margin at

point of pereonite fusion, lateral margins

widely over-reaching coxae, without spines. Pere¬

onites 1-4 combined about same length as 5-7

combined.

Plcotclson free, straight and ventrally flat in lat¬

eral view, as long as perconites 4-7 combined; as

wide at base as perconite 7. semicircular, except

for triangular notches at insertion of uropods,

length 60% greatest width; without preanal ridge;

without lateral or apical spines; anterior dorsal

margin with raised transverse keel in area of artic¬

ulation with perconite 7; dorsal surface with

broad, rounded midlongiludinal ridge and

rounded inflations on each side, 2 short rounded

spines at midlcngth between middle ridge and

lateral inflations; anus external to respiratory

chamber, not covered by pleopods.

Antennae: Antenna I as long as perconites M

and half of 5 combined; article 1 with rounded

projection antcrolaterally and long, slender spine

medially, article 2 inserted dorsally on I, article 3

slightly shorter than 2. 4 quarter length of 3, 5

subequal to 3, with an additional 19 short articles,

distal ones with aesthetascs. Antenna 2 article 1

AN UNUSUAL MUNNOPSIDID ISOPOD FROM ANTARCTICA

339

Figure 3. Platvprotusphyllosoma sp. nov., holotype. Mandibles: r, right; I, left.

with small distolateral spine, article 2 without

spines, article 3 with 1 short and 1 long disto-

mcdial spines and 1 distolateral spine, all spines

lipped by short broad seta; (flagellum not known).

Mouthparts: Upper lip evenly rounded, with 2

fields of distal setae.

Mandible with elongate cuticular projection

articulating with cephalon in shallow groove (Fig.

7c); left mandible with broad, right with more

narrow incisor of 3 rounded (presumably worn)

teeth, left with broad, irregular lacinia mobilis

with 4 rounded teeth and row of more than 10

slender setae, right with similar number but

broader, microdentate setae; molar cylindrical,

hollowed truncate, with short row of microdentate

setae on posterior margin; palp exceeding length

340

J. JUST

Figure 4. Platyprotusphyllosoma sp. nov., holotype. mp, maxilliped; ml, m2, maxillae I, 2.

AN UNUSUAL MUNNOPSIDID ISOPOD FROM ANTARCTICA

341

Figure 5. Platyprotus phyllosoma sp. nov., holotype. Pereopods 1, 3 and 4.

342

J. JUST

Figure 6. Platyprotus phyllosoma sp. nov., holotype. Pereopods 5, 6 and 7.

AN UNUSUAL MUNNOPSIDID ISOPOD FROM ANTARCTICA

343

of mandibular body by entire article 3, article 2

2.5 times longer than 1, with 2 tiny, curx'ed, acute

setae distolatcrally, article 3 broad, of twisted

spoon-shape, lateral and apical margins with

highly modified spalulate setae, concave surface

with densely set transverse rows of microsctules.

Lower lip outer lobes with medial row of robust

setae and apical and lateral cover of sctules, inner

lobes rounded with lateral and apical setules.

Maxilla I outer lobe with 11 nearly straight

spine-like setae, posterior 6 microdentate, inner

lobe apically rounded, reaching to mediodistal

comer of outer lobe, with 1 apical spine-like seta

and many long setules. Maxilla 2 middle lobe

slightly shorter than subequal outer and inner

lobes, inner lobe 1.5 times width of equal width

middle and outer lobes, outer and middle lobes

with 2 long and 2 shorter smooth setae, inner lobe

with apical row of short, pectinate setae and dense

fringe of medial setules.

Maxiliiped basis with 4 coupling hooks, trun¬

cate apex of endite with 6 slender fan setae, palp

article 2 broadest distally, 1.5 times midwidth of

endite, epipod reaching to distal end of palp

344

J. JUST

Figure 8. Platyprotus phyllosoma sp. nov., holotype. Pleopods 2-5 (2 viewed from above).

article 2, apex rounded, lateral margin smooth,

convex in posterior half, slightly concave in distal

half.

Pereopods: Pereopod I with scattered simple

setae, much shorter and more slender than suc¬

ceeding pereopods, about half length of cephalon

and pereon combined; relative length of pereo¬

pods; 1 (1.0), 2 (not measured), 3 (1.9) 4 (2.5), 5

0-7), 6 (1.7), 7 (broken); pereopods 2-3 carpus

and propodus of equal length, carpus twice width

of propodus, posterior margin of carpus and

propodus with short simple setae and a few more

robust setae; pereopod 4 propodus approximately

third longer than carpus, carpus nearly 3 times

AN UNUSUAL MUNNOPSIDID ISOPOD FROM ANTARCTICA

345

wider than propodus with 6 robust setae mainly in

distal half, propodus with a few robust setae

among small simple setae in distal half of pos¬

terior margin. Pereopods 5-7 fully ambulatory;

propodus slightly shorter than carpus, carpus little

wider than propodus; pereopods 5-6 (and pre¬

sumably 7) carpus anterior margin with a few

robust setae, propodus with row of 6-7 robust

setae. Dactylar unguis of all pereopods with mid¬

point tooth on concave surface, pereopods 1-4

with 2 stiff setae arising from postcrodistal comer

of dactylus and resting against unguis tooth, pere¬

opods 5-6 (presumably 7) with single strong,

spatulate seta in same position.

Plcopods: (pleopod 1 not known); picopod 2

protopod about half length of pleotelson, width of

combined pleopods 2 about half midwidth of

pleotelson, medial margin concave, lateral margin

evenly convex, apex broadly rounded, distal part

of lateral margin and apex with row of long,

slender hemiplumose setae; exopod a short cone

with setose apex; endopod inserted about 0.6

along protopod margin, stylet in retracted position

overreaching protopod apex with about third

stylet length. Pleopods 3-5 endopod of similar

shape and size, broad, scmirectangular, 3 with 3

apical pappose setae, 4 and 5 without setae, exo¬

pod of 3 as long as and third width of endopod,

apex rounded with 9 long pappose setae, entire

lateral margin of exopod with dense fringe of long

setulcs; exopod of 4 reaching about two-thirds

along endopod, tapering to pointed apex carrying

I pappose seta; 5 without exopod.

Uropods 0.4 length of pleotelson, protopod and

inner ramus of equal length, outer ramus 0.6

length of inner ramus, both rami with tuft of

apical setae.

Female. Generally as male. Operculum nearly cir¬

cular with slightly concave apex, just covering

respiratory chamber, fringed with long hemi¬

plumose setae in distal half.

Etymology. The epithet is derived from the Greek

phyllos = leaf and soma = body.

Acknowledgments

I thank Dr Gary Poore, Museum Victoria, for

making the material available, and Dr G.D.F.

‘Buz’ Wilson, Australian Museum, for helpful

discussions.

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Scripps Institution of Oceanography 27: i-xii,

1-138.

Wilson, G.D.F., Kussakin, O.G. and Vasina, G.S.,

1989, A revision of the genus Microprotus

Richardson with descriptions of two new species,

M. acutispinatus and M. lobispinatus (Asellota,

Isopoda, Crustacea). Proceedings of the Biological

Society of Washington 102 (2): 339-361.

Memoirs of Museum Victoria 58(2); 347-364 (2001)

THREE NEW SPECIES OF CIROLANA LEACH, 1818 (CRUSTACEA:

ISOPODA: CIROLANIDAE) FROM AUSTRALIA

Stephen J. Keable

Crustacea Section, Australian Museum, 6 College Street, Sydney, NSW 2010, Australia

Abstract

Keable, S.J., 2001. Three new species of Cirolana Leach, 1818 (Crustacea: Isopoda:

Cirolanidae) from Australia. Memoirs of Museum Victoria 58 (2): 347-364.

Three new species are described from Australia, Cirolana australis sp. nov. off eastern

and southem coasts of Tasmania, C. comato sp. nov. off north-eastern Queensland, and

C dissimilis sp. nov. off northern Western Australia. Northern Territory and northern

Queensland. All appear to be associated primarily with coral or rocky reef habitat. Cirolana

coma/a and C. dissimilis occur on the continental shelf, C. australis on the continental slope

at depths exceeded by few other species of Cirolana. The species arc abundant in collections

made using baited traps, indicating that they are scavengers and potentially important pests of

tlsheries. They have nodular sculpting on the pereon. pleon and pleolelson but difller from

other Australian species with similar ornamentation in having the setae on the lateral margin

of the uropod endopod in a continuous row, not in discrete widely spaced groups. Cirolana

comata and C. dissimilis are sexually dimorphic and have a highly sclerolized robust seta on

the posterodistal angle of the basis of pereopods 4-5. These and other shared characters

suggest that the two are related.

Introduction

Twenty-nine named and one unnamed species of

Cirolana Leach, 1818 were included in the most

recent key to the Australian fauna (Bruce, 1986).

Cirolana schioedtei Miers, 1884 has subse¬

quently been transferred to Aatolana Bruce, 1993

(Bruce, 1993). The juvenile specimens included

in this key and discussed as Cirolana sp. appar¬

ently represent an undcscribed species of

Plakolana Bruce, 1993, and C. hinyana Bruce,

1991 is also a species of Plakolana (Bruce,

1993). Brusca et al. (1995) listed 84 species of

Cirolana worldwide. Cirolana ohtrimcata

Richardson, 1901 was moved to Neocirolana

Hale, 1925 by Javed and Yasmeen (1990),

C fornicata (Mezhov, 1981) is regarded as a

species oV hdetacirolana Kussakin, 1979 (Baicc,

1996), and Keable (1999) Iransfered C parcel-

lana Barnard, 1936 and C. alhicauda Nunomura,

1985 (not listed by Brusca ct al., 1995) to

Doiicholana Bruce, 1986 as suggested by Bruce

(1986). These changes, and the species described

by Kwon (1988), Weider and Feldmann (1992),

Bruce (1995), Javed and Yasmeen (1995) and

Botosaneanu and Iliffe (1997), brings the recog¬

nised species to 90 (including one species

inquirenda, three inccrtac sedis and one fossil

species). Botosaneanu and Iliffe (1997) discussed

the similarities of Anopsilana Paulian and

Delmare Deboulteville, 1956 and Cirolana and

suggested that Anopsilana is a subgenus of

Cirolana. They recognised that this is an

artificial solution and it is not followed here.

Scavenging cirolanid isopods play an impor¬

tant role in marine foodwebs (Keable, 1995) and

are significant pests of fisheries (Bird, 1981;

Stepien and Brusca, 1985; Berrow, 1994; Miz-

zan, 1995). This study describes three new scav¬

enging species of Cirolana from Australia col¬

lected with baited traps.

The terminology and procedures used follow

Keable (1997). Abbreviations are: AM, Aus¬

tralian Museum, Sydney, Australia; BMNH, The

Natural History Museum, London, United King¬

dom; NMV, Museum Victoria, Melbourne, Aus¬

tralia; NTM, Northern Territory Museum of Arts

and Sciences, Darwin, Australia; TM, Tasmanian

Museum and Art Gallery, Hobart, Australia;

USNM, National Museum of National History,

Smithsonian Institution, Washington, D.C., USA;

n, number of specimens; CE, cephalon; Al,

anlcnnule; A2, antenna; CL, clypeal region; FL,

frontal lamina; MD, mandible; MP, maxillipcd;

MXl, maxillulc; MX2, maxilla; PE, penes; PN,

pleon; PI-7, pereopods 1-7; U, uropod; PL 1-5,

plcopods 1-5; PT, plcotclson.

Cirolana Lc'dch, 1818

Remarks. For useful recent synonymies, diag¬

noses, desciiptions and discussions see Bruce

(1986: 139), Kenslcy and Schottc (1989: 132),

Bruce (1993: 2), Bruce (1995: 376) and Brusca et

347

348

S. J. KEABLE

ai. (1995: 17). Revisionary work has provided

increasingly narrower and more practical mor¬

phological limits to Cirolana with the transfer of

many species to other genera (e.g. Bruce, 1981a;

1993). Three informal subgroups of Cirolana

have also been proposed by Bruce (1986, 1995):

(1) the “C parva group"; (2) the “C southern

group": and (3) the ‘‘C. tuberculate group". The

new species described here (and several others)

cannot be unambiguously placed in any group:

the rostrum does not meet the frontal lamina and

there is some sculpting on the body segments of

all three species eliminating them from the

“C. parxxC group; and they cannot be members of

the “C. soulhem'" or the “C. tuberculate" groups

because they have continuous setae on the lateral

margin of the uropod exopod. This, combined

with the character conflict which exists between

these infonnal groups, suggests that they may be

based on plesiomorphic or homoplastic charac¬

ters. A phylogenetic analysis to resolve this issue

is needed.

All species of Australian Cirolana with nodular

sculpting in the form of tubercles, and that are

otherwise similar to the three new species, have

the uropod exopod lateral margin with discon¬

tinuous plumose setae (i.e. occurring in

widely spaced groups) and fewer robust setae, or

the rostrum meeting the frontal lamina.

A close relationship between Cirolana comata

sp. nov. and C clissimilis sp. nov. is suggested by

many similarities, in particular corresponding

sexually dimorphic characters rare in species of

Cirolana (Bruce, 1986: 140). Both have a highly

sclerotized robu.st seta on the postcrodistai angle

of the basis of pereopods 4-5 (4-6 in C. comata).

This character docs not occur in C. australis

sp. nov., nor in C. ausiraliense Hale, 1925,

C. capricornica Bruce. 1986 or C. similis Bruce.

1981b (specimens AM P47660, P47668, P47669

examined). Its distribution is otherwise undocu¬

mented. Additionally, an iridescent brush fonned

by the aesthetascs of the antcnnule (Kcablc, 1998;

Parker, in prep.) was found in both C. comata and

C. clissimilis but not C australis.

The three new species appear to be associated

primarily with coral or rocky reefs. Cirolana

comata and C. clissimilis occur on the continental

shelf, C. australis on the continental slope. Of the

species currently placed in Cirolana or incertac

sedis, only C bisulcata Hobbins and Jones, 1993,

C. hougaarclii Kenslcy, 1984. C. mesecla Hobbins

and Jones, 1993. C. vanlwejfeni Nierstrasz, 1931

and C stebhingi Nierstrasz, 1931 are known from

similar or greater depths than C australis.

Cirolana australis sp. nov.

Figures 1-3

Cirolana new species.—Koslow and Gowlett-

Holmes, 1998: 41.

Cirolana n.sp. 4.—Lowry, 1998: 63, 64.

Material examined. Holotype. Main Pedra Seamount,

off southern Tasmania (44°15.6'S 147°7.8'E), baited

trap, 1,312 m, 21-24 Jan 1997 (CSIRO Cruise

SSOl/97, stn 8), AM P59351 (male, 23 mm).

Paratypes. All same data as holotype. AM P59352

(977 specimens); BMNH 2()()().24()8-2409 (male,

female); NMV J47153 (male, female); TM G3588 (2

females); USNM 296460 (male, female).

Additional material. E of Fortescue Bay, Tasmania

(43°08.96'S 145°15.36'E), baited trap, 5.I°C, 1000 m,

J. Lowry and P. Freewater, 16-17 Apr 1993 (stn SEAS

T.AS-365). AM P59353 (2 females, 1 male, 6 mancas);

stn SEAS TAS-367. AM P59354 (7 females, 1 male, 5

mancas).

Diagnosis. Ccphalon: rostrum not extending to

frontal lamina, not dividing antcnnulcs; anterior

margin not overriding antennulcs. Eyes: well

developed, round. Frontal lamina: anterior margin

angled. Pereonites: 1-7 with 2 transverse carinae;

4 7 with tubercles. Pleonites: 3-5 with tubercles.

Pleonitc 4: ventral margin free of plconitc 3: pos-

lerodorsal margin apex broadly rounded dorsally

but meeting convex ventral margin at a point.

Pleotelson: dorsal surface with paired tubercles in

rows, (3 in each row, fomiing an indistinct ridge

parallel to each lateral margin), conspicuous fine

setae absent; anterolateral margins convex; pos¬

terolateral margins concave; apex imncate; 2-6

(usually 2 or 4) robust setae across apex. Penes:

present. Pereopod 1: propodus without plumose

setae. Pereopods 4-6: basis postcrodistai angle

robust setae absent. Pleopod 2 appendix mas-

culina: arising subbasally; extending beyond tip

of endopod, 1.46 length of endopod from inser¬

tion point; margins sinuate, tapering along entire

length; slender; apex not at angle to margins,

bluntly rounded. Uropods: endopod not dimor¬

phic; lateral margin straight for proximal two-

thirds, convex at distal third: exopod not dimor¬

phic; lateral margin straight, robust and plumose

setae continuous along margin.

Additional descriptive characters based on holotype.

Body; length approximately 2.6 greatest width; white in

alcohol: chromalophores absent; cuticular surfiiccs

scale-like.

Cephalon: tubercles absent. Eyes: visible in ventral

view; black in alcohol: partially overlapped by pereo-

nite 1: ommatidia in rows, 7 ommatidia in horizontal

diameter, 7 ommatidia in vertical diameter. Interocular

furrow: distinct, not extending across cephalon. Frontal

lamina: length approximately 1.9 basal width; penta-

THREE NEW SPECIES OF CIROLANA

349

Figure 1. Cirolana australis sp. nov., holotype. Scales = 0,5 mm.

350

S. J. KEABLE

gonal; lateral margins divergent: apex not projecting,

not visible in dorsal view, not expanded, in I plane (not

stepped). Clypeus; triangular, not produced.

Perconites: 4-7 length subequal and longest, 1-3

subequal; tubercles small, fine, subequal, continuous

across entire posterior margin,

Pleonites: 1-5 equally visible along dorsal margin;

tubercles small, subcqual, continuous across entire

posterior margin.

Pleotelson: length 0.85 basal width; anterodorsal

uropodal sutures present; 4 robust setae across apex;

plumose setae restricted to posterolateral margins,

numerous proximal to robust setae.

Antennulc: just reaching perconite 1. Peduncular

bases touching; articles I -2 free; article 1 length less

than width, subcqual to article 2: article 2 longer than

wide, with a few scattered slender and penicillate setae;

article 3 longer than combined lengths of articles 1-2,

length greater than width. Flagellum shorter than

peduncle; articles not compressed (lengths of most

greater than half width); 8-articuIate; aesthetascs not iri¬

descent. Antenna; 0.3 length of body, when extended

against body reaching to posterior of pereonitc 3.

Peduncular article 2 shorter than article 3; article 4

much longer than article 3, posterodistal angle with 4

slender setae, antcrodistal angle with 5 slender setae:

article 5 subcqual in length to article 4, posterodistal

angle with 2 penicillate and 2 slender setae, anterodistal

THREE NEW SPECIES OF CIROLANA

351

angle with 1 pcnicillalc and 7 slender setae. Flagellum

l9-ailiculatc; sctal brush absent.

Mandible: molar medial surface covered with short

fine slender setae, cluster of long slender setae proxi-

mally present, long slender setae in submarginal row

across 0.75 length of anterior margin; marginal robust

setae close set. Setal row with 15 robust setae; medial

surface covered in fine setae. Maxillule: medial lobe

lateral margin protuberance absent; lateral lobe with

10 robust setae on distal surface and 2 slender setae.

Maxilla: lateral lobe with 4 plumose and 1 slender

setae; medial lobe with 14 plumose setae, medial

plumose setae subcqual to proximal setae; middle lobe

with 4 plumose and 8 slender setae. Maxilliped: right

enditc with 1 coupling hook, left endite with 2 coupling

hooks.

352

S. J. KEABLE

Pereopods: 1-3 menus posterior margin robust setae

strongly molariform on I only; 2-7 coxal furrows com¬

plete. Pereopod 1: posterior margin setose fringe

absent; propodus robust; daclylus shorter than 0.5

propodus length.

Penes; separated by more than width of both penes;

forming flattened lobes; length approximately 1.5 basal

width.

Pleopods: exopod suture complete on 3-5. Pleopod

I: exopod medial margin tapering evenly, proximolal-

cral robust seta present; endopod shorter than exopod,

lateral margin slightly concave.

Uropods: extending beyond pleoteLson. Peduncle

ventrolateral angle without robust setae, with 13

plumose setae; lateral margin robust seta absent; disto-

lateral angle rounded. Endopod medial margin convex,

with 5 robust setae, plumose setae along etttire length;

apex subbifid with lateral spine largest, with 1 minute

medial robust seta, setal cluster formed by plumose and

slender setae; lateral margin with 4 robust setae,

plumose setae along entire length. Exopod 0.94 length

of endopod; medial margin convex, with 4 robu.st setae,

plumose setae along entire length; apex subequally

bifid, without robust setae, .setal cluster formed by

slender setae; lateral margin with 7 large robust setae,

plumose setae along entire length.

Sexual dimorphism. Females differ from males

only in the primary sexual characters.

Variation. Plcolelson and uropod robust setal

counts from margins (N = 20, subsample of 10

males and 10 females from AM P59352): Pleolel-

son: 1:1 (30%), 2:1 (40%), 2:2 (25%), 3:3 (5%).

Endopod: (medial) 5 (10%), 6 (70%), 7 (20%);

(lateral) 3 (65%), 4 (35%). Exopod: (medial) 4

(75%), 5 (25%); (lateral) 5 (10%), 6 (40%), 7

(45%), 8 (5%). Subadull males (19 mm, AM

P59352) have the vas deferens opening almost

flush to the surlace of the stemile and the

appendix masculina inserted submedially.

Size range. Mancas approximately 7 mm, adults

to approximately 25 mm.

Etymology’. Australis, Latin, southern, referring to

the position of the type locality.

Distribution. Eastern and southern Tasmania:

1000-1312 metres.

Remarks. Cirolana australis is most readily dif¬

ferentiated by the ornamentation of the somites,

coupled with the sinuate lateral margins and trun¬

cate apex of the plcotclson, low number of robust

setae on the pleotelson apex, straight lateral mar¬

gins of the uropod rami, and continuous distribu¬

tion of the robust and plumose setae along the

uropod exopod lateral margin. Other species of

Cirolana in which robust setae occur on the mar¬

gins of the pleotelson usually have six or more,

but four have also been recorded in C. rugicauda

Heller, 1861. Cirolana rugicauda is distinguished

from C. australis by (in C. rugicauda): the

rounded apex of the frontal lamina; the projecting

clypeus (a character which has been used to diag¬

nose genera and groups of genera excluding

Cirolana, e.g. Bruce (1986)); and pleonitc 3

enclosing pleonitc 4 (VanhofTen, 1914; Barnard,

1940; Kensley, 1978), Cirolana sulcata Hansen,

1890 and Cirolana franscostafa Barnard, 1959

may be difficult to distinguish from C. australis

but have a frontal lamina with a rounded apex,

and uropods with lateral margins that arc more

convex. Cirolana tuberculata (Richardson, 1910)

is also similar but has more prominent tubercles

on the pleotelson and uropods with convex lateral

margins (Delaney, 1986).

Cirolana comata sp. nov.

Figures 4-6

Material examined. Ilololype. Portlock Reef, Coral

Sea, Queensland (9°42.10'S 144°50.17'E), bailed trap,

unknown substrate, 65 m, S. Keable, 28 Jan 1993 (stn

QLD-775), AM P59355 (male, 13 mm).

Paratypes. All same data as holotype, AM P39356

(298 specimens); BMNII 2000.2410-2411 (male,

female); NMV J47152 (male, female); USNM 296461

(male, female).

Additional material. Off Flynn Reef. Queensland

(16°41.32'S 146°I8.26'E), bailed trap, unknown sub¬

strate, 100 m, J. Lowry, P. Freewater and W. Vader, 7

Jun 1993 (stn SEAS QLD-937), AM P47678 (10 spec¬

imens); E of Fitzroy Reef, Queensland (23°32.53'S

152°16.45'E), bailed trap, unknown substrate, 105 m, J.

Lowry, P. Freewater and R. Springthorpe, 16 Jun 1993

(stn SEAS QLD-956), AM P47675 (13 specimens).

Diagnosis. Ccphalon: rostrum not extending to

frontal lamina, not dividing antcnnulcs; anterior

margin not overriding antcnnulcs. Eyes: well

developed, round with upper margin flat. Frontal

lamina: anterior margin angled. Pereonites: with¬

out transverse carina; tubercles absent (except for

6 indistinct tubercles on submarginal anterior

ridge of pereonite 1 in mature males). Pleonites:

3-5 with tubercles. Pleonitc 4: ventral margin free

of pleonitc 3; posterodorsal margin apex broadly

rounded dorsally but meeting convex ventral

margin at a point. Pleotelson: dorsal surface with

2 paired tubercles, conspicuous fine setae fonning

dense patch in males; anterolateral margins

almost straight and angling posteriorly toward

midlinc; posterolateral margins straight, contigu¬

ous with anterolateral margins; apex rounded;

8-11 (usually 10) robust setae on margins. Penes:

present. Pereopod I: propodus without plumose

THREE NEW SPECIES OF CIROLANA

353

Figure 4. Cirolana comata sp. nov., holotype. Scales = 0.2 mm.

354

S. J. KEABLE

setae. Pereopods 4-6: basis posterodistal angle

with 1 strongly sclcrotizcd robust seta. Pleopod 2

appendix masculina: arising subbasally; extend¬

ing beyond tip of endopod, 1.22 length of

endopod from insertion point; margins straight,

parallel along most of length, but tapering toward

apex; slender; apex not at angle to margins,

tapered to finely acute point. Uropods: endopod

dimorphic, males with dorsal setae; lateral margin

slightly convex. Exopod dimorphic, males with

dorsal setae; lateral margin convex, robust and

plumose setae continuous along margin.

AMtional descriptive characters based on hohtype.

Body: length approximately 3.35 greatest width; cream

in alcohol; chromatophores, small, brown in alcohol,

scattered over body; Cuticular surfaces scale-like.

Ccphalon: with 4 indi.slinct tubercles, 2 medially and

2 along posterior margin. Eyes: visible in ventral view;

orange-tan in alcohol; partially overlapped by pcrconite

1; ommatidia in rows, 10 ommalidia in horizontal

diameter, 9 ommatidia in vertical diameter. Inlerocular

furrow: distinct, extending across ccphalon, smoothly

convex. Frontal lamina: length approximately 2 basal

width; pentagonal: lateral margins parallel; apex

not projecting, not visible in dorsal view, not expanded,

in 1 plane (not stepped). Clypeus: triangular, not

produced.

Pereonites: 1 longest, 4-6 length subequal and longer

than 2-3 and 7 which are subequal.

Pleonites: 1-5 visible but 1 almost completely

THREE NEW SPECIES OF CIROLANA

Figure 6. Cirolana comata sp. nov., holotype, except female = paratype, 17.5 mm, AM

P59356. Scales = 0.5 mm.

356

S. J. KEABLE

concealed along dorsal margin by pcreonite 7; tubercles

not continuous across entire posterior margin, plconite

3 tubercles small, subequal, pleonitc 4 median tubercles

prominent, pleonite 5 lateral tubercles prominent.

Pleotelson: length 0.88 basal width; anterodorsal

uropodal sutures present; 5 robust setae on each

posterolateral margin; plumose setae restricted to

posterolateral margins, numerous proximal to robust

setae.

Antennule: just reaching pcreonite 1. Peduncular

bases touching; articles 1-2 free; article I length sub¬

equal to width, greater than article 2; article 2 wider

than long, anteromedial margin with 1 penicillatc seta,

postcrodistal angle with 2 penicillatc setae; article 3

shorter than combined lengths of articles 1-2. longer

than article 1, length greater than width. Flagellum

longer than peduncle; articles not compressed (lengths

of most greater than half width); 16-articulate;

aesthelascs iridescent. Antenna: 0.41 length of body,

when extended against body reaching to posterior of

pcreonite 4. Peduncular article 2 and article 3 subequal

in length; article 4 much longer than article 3, with 4

slender setae at anterodislal angle, 1 penicillatc seta on

posteromedial margin, 2 slender setae at postcrodistal

angle; article 5 longer than article 4 and all other

articles, with 1 slender and 2 penicillatc setae at

anterodistal angle, 2 penicillatc and 1 slender .setae at

postcrodistal angle. Flagellum 35-articulate; selal brush

present.

Mandible: molar medial surface covered with short

fine slender setae, cluster of long slender setae proxi-

mally present, long slender setae submarginal to anter¬

ior margin absent; marginal robust setae close set. Setal

row with 11 robust setae: medial surface without setae.

Maxillulc: medial lobe lateral margin protuberance well

developed: lateral lobe with 11 robust setae on distal

surface. Maxilla: lateral lobe with 6 slender setae;

media! lobe with 12 slender and 7 plumose setae, with

2 medial plumose setae longest and bent; middle lobe

with 13 slender setae. Maxilliped: right and lel\ endite

with 2 coupling hooks.

Pereopods: 1-3 menis posterior margin robust setae

strongly molariform on 1 only; 2-7 coxal furrows com¬

plete. Pereopod I: posterior margin setose fringe

absent; propodus robust; dactylus long, 0.5-1 propodus

length.

Penes: separated by more than width of both penes;

forming flattened lobes; length approximately 1.5 basal

width.

Plcopods: exopod suture complete on 3-5 (but indis¬

tinct). Pleopod 1: exopod medial margin tapering

evenly, proximolateral robust seta present; endopod

length subequal to exopod. lateral margin slightly

concave.

Uropods: extending beyond pleotelson. Peduncle

vcmrolaieral angle with 2 robust setae and 1 plumose

seta; lateral margin robust seta present; dislolaleral

angle rounded. Endopod medial margin convex, with 6

robust setae, plumose setae along entire length; apex

subbifld with lateral spine largest, without robust setae,

setal cluster formed by plumose and slender setae; lat¬

eral margin with 2 robust setae, plumose setae along

entire length. Exopod 0.86 length of endopod; medial

margin convex, with 3 robust setae, plumose setae on

distal two-thirds: apex subbifid with lateral spine

largest, without robust setae, setal cluster formed by

slender setae; lateral margin with 6 large robust setae

plumose setae along entire length.

Sexual dimorphism. Females differ from males in

the primary sexual characters and do not develop

the dense patches of setae found on the dorsal

surface of the pleotelson. uropod endopod and

uropod exopod, or the prominent lateral tubercles

on plconite 5 and dense brush of setae formed on

the proximal articles of the antennal tlagelluni.

The tubercles on the cephalon, pcreonite i

plconiles 3-4 and on the pleotelson arc also

absent in females.

Variation. Pleotelson and uropod robust setal

counts from margins (N = 20, siibsample of iQ

males and 10 females from AM P39356): Pleotcl-

son: 4:4 (5%), 4:5 (5%). 5:5 (85%) , 5:6 (5%).

Endopod: (medial) 5 (70%), 6 (30%); (lateral) i

(5%), 2 (95%). Exopod: (medial) 2 (5%l 3

(95%); (lateral): 5 (65%), 6 (35%). Males approx¬

imately 6 mm long lack tubercles, and the dorsal

setae on the pleotelson and uropods.

Size range. Mancas approximately 3 mm, adults

approximately 5-17.5 mm.

Etymology’. ComatOy Latin, hairy, referring to the

dense setae found on the dorsal surfaces of the

pleotelson and uropod rami in mature males.

Distribution. Off north-east Queensland; 65-105

metres.

Remarks. The prominent lateral tubercles on

pleonitc 5 and the dense patch of setae on the

pleotelson distinguish mature males of Cirolana

comata and C. dissimilis from those of other

species in the genus. Cirolana comata differs

from C dissimilis in having a dense patch of setae

on the dorsal surface of the uropod rami, a setal

brush on the antenna in mature males and a

strongly sclcrotized robust seta at the posterodis-

tal angle of the basis of pereopods 4-6 (4-5 in

C. dissimilis). Females and immature males of

C comata and C. dissimilis arc difficult to distin¬

guish from other species but the lack of chro-

malophores, lack of strong tubercles or other

sculpting on the somites, rostrum not extending to

the frontal lamina, relatively high number of

robust setae on the pleotelson, shape of the pleo¬

telson, presence of a strongly sclerotizcd robust

seta at the postcrodistal angle of the basis of pere¬

opods 4-5 (C. dissimilis)^ or 4-6 (C comata), and

the continuous distribution of the robust and

THREE NEW SPECIES OF CIROLANA

357

plumose setae along the uropod exopod lateral

margin are diagnostic. Cirolana indica Nicr-

strasz, 1931, which is only known from a single

female which lacks tubercles or ridges, may be

similar to C comata and C. dissimilis. However,

C indica is described as having a row of plumose

setae along the postcrodislal angle of the pro-

podus of pcrcopod 1 which is lacking in these

species. Cirolana meinerd Barnard, 1920 appar¬

ently has a patch of setae on the dorsal surface of

the picotelson and may also be similar to

C comata and C. dissimilis in other respects.

However, C meinerti is described as having

medial tubercles on pleonite 5 which are more

prominent than the lateral tubercles. This is not

the case in C. comata or C dissimilis. Mature

males of C. pleonastica Siebbing, 1900 develop a

dense patch of setae on the dorsal surface of the

uropod exopod, as in C. comata. However, they

do not develop a similar patch on the uropod

endopod or the pleotelson (Bruce, 1995).

Cirolana comata and C pleonastica are also dis¬

tinct in many other features including the sculpt¬

ing of the somites, and the shape and sclation of

the pleotelson.

Cirolana dissimilis sp. nov.

Figures 7-9

Cirolana sp. 1.—Kcablc, 1997: 251.

Material examined. Holotype. Just off West Point. Dar¬

win Harbour, Northern Territory (12°26.3'S

I29°46.3’E), baited trap, unknown substrate but proba¬

bly rock reef, 8 m, S. Keable, 8-9 Jul 1993 (stn

NT-123), NTM CrO 12796 (male, 18 mm).

Paratypes. All same data as holotype, AM P44797

(40 males, 40 females); BMNH 2000.2412-2413 (male,

female); NMV J47I51 (male, female); NTM CrO 12797

(female); USNM 296462 (male, female).

Additional material. Bet Reef, Torres Strait. Queens¬

land (10'’I0.54'S 142®56.0rE), bailed trap, grey clay

mud with shell grit, 20 m, S. Keable, 30 31 Jan 1993,

AM P44799 (many specimens). Ngalaguru (High

Cliffy) 1., Western Australia (15°54.77'S 124°20.68'E),

baited trap, unknown substrate, unknown depth, F.

Wells, 22-23 Nov 1994, AM P59357 (19 specimens).

Diagnosis. Cephalon: rostrum not extending to

frontal lamina, not dividing antennules; anterior

margin not overriding antennules. Eyes: well

developed, round with upper margin flat. Frontal

lamina; anterior margin angled. Pcrconitcs: with¬

out transverse carina; tubercles absent (except for

4 indistinct tubercles on submarginal anterior

ridge of pcrconite 1 in mature males). PIconites:

3-5 with tubercles. Pleonite 4: ventral margin free

of pleonite 3; postcrodorsal margin apex broadly

rounded dorsally but meeting convex ventral mar¬

gin at a point. Pleotelson: dorsal surface with 2

paired tubercles, conspicuous line setae fonning

dense patch in males; anterolateral margins con¬

vex; posterolateral margins convex; apex

rounded; 8-11 (usually 10) robust setae on mar¬

gins. Pereopod 1: propodus without plumose

setae. Pereopods 4-6: basis posterodistal angle

with 1 strongly sclerotizcd robust seta on pere¬

opods 4-5 only. Penes: present. Pleopod 2

appendix masculina: arising basally; extending

beyond tip of endopod, 1.21 length of endopod

from insertion point; margins straight, parallel

along most of length, but tapering toward apex;

slender; apex not at angle to margins, tapered to

finely acute point. Uropods: endopod not dimor¬

phic; lateral margin slightly convex; exopod not

dimorphic; lateral margin straight, robust and

plumose setae continuous along margin.

Additional descriptive characters based on holotype.

Body: length approximately 3.6 greatest width; cream

in alcohol; chromatophores absent; cuticular surfaces

scale-like.

Cephalon: with tubercles, 4 indistinct along posterior

margin. Eyes: visible in ventral view; black in alcohol;

partially overlapped by pereonite 1; ommalidia in rows,

9 ommatidia in horizontal diameter, 8 ommatidia in ver¬

tical diameter. Interocular fuiTow: distinct, extending

across cephalon, smoothly convex. Frontal lamina:

length approximately 2 basal width; pentagonal; lateral

margins concave; apex not projecting, not visible in

dorsal view, not e.xpanded, in 1 plane (not stepped).

Clypcus: triangular, not produced.

Pereonites: 1, 4-6 length subequal and longest, 2-3

and 7 subequal.

PIconites: 1-5 visible but 1 almost completely

concealed along dorsal margin by pereonite 7; tubercles

not continuous across entire posterior margin, pleonites

3-4 tubercles small and subequal, pleonite 5 lateral

tubercles prominent.

Pleotelson: length 0.79 basal width; anlerodorsal

uropodal sutures present; 5 robust setae on each

posterolateral margin; plumose setae restricted to pos¬

terolateral margins, numerous proximal to robust setae.

Antennule: just reaching pereonite 1. Peduncular

bases touching: articles 1-2 free; article I length sub¬

equal to width, subequal to article 2; article 2 longer

than wide, with I penicillate seta at posterodistal angle

and on anterolateral margin; article 3 shorter than com¬

bined lengths of articles 1-2. longer than article I,

length greater than width. Flagellum longer than pedun¬

cle; articles not compressed (lengths of most greater

than half width); 19-articulale: aesthctascs iridescent.

Antenna: 0.38 length of body, when extended against

body reaching to posterior of pereonite 4. Peduncular

article 2 shorter than article 3; article 4 much longer

than article 3, wath 1 penicillate seta on posterolateral

margin, 3 slender setae at posterodistal angle; article 5

358

S. J. KEABLE

Figure 7. Cirolana dissimilis sp. nov., holotype. Scales = 0.5 mm.

THREE NEW SPECIES OF CIROLANA

359

longer than article 4 and all other articles, with 2

pcnicillatc setae at anterodistal angle, 4 slender and 1

penicillatc setae at posterodislal angle. Flagellum

4!-articulate; setal brush absent.

Mandible: molar medial surface covered with short

fine slender setae, cluster of long slender setae proxi-

mally present, long slender setae submarginal to anter¬

ior margin absent; marginal robust setae close set. Setal

row with 15 robust setae; medial surface without setae.

Maxillule: medial lobe lateral margin protuberance well

developed; lateral lobe with 10 robust setae on distal

surface. Maxilla: lateral lobe with 4 slender setae;

medial lobe with 4 slender and 14 plumose setae, with

medial plumose seta longest and bent; middle lobe with

lateral row of 10 long slender setae and medial row of

10 short slender setae. Maxilliped: right and left endite

with 2 coupling hooks.

Pereopods: 1-3 menis posterior margin robust setae

strongly molarifomi on 1 only; 2-7 coxal furrows com¬

plete. Pcrcopod 1: posterior margin setose fringe

absent; propodus robust; dactylus long, 0.5-1 propodus

length.

Penes: separated by more than width of both penes;

forming flattened lobes; length approximately 1.5 basal

width.

Pleopods: exopod suture complete on 3-5 (but

360

S. J. KEABLE

Figure 9. Cirolana dissimilis sp. nov., holotype, except female - paratype, 21 mm, AM P44797. Scales - 0.5 mm-

THREE NEW SPECIES OF CIROLANA

361

indistinct). Pleopod 1: exopod medial margin tapering

evenly, proximolateral robust seta present; endopod

length subequal to exopod. lateral margin straight.

Uropods: extending beyond pleotelson. Peduncle

ventrolateral angle with 2 robust setae and 2 plumose

setae; lateral margin robust seta absent; distolateral

angle rounded. Endopod medial margin convex, with 7

robust setae, plumose setae along entire length; apex

subbifid with lateral spine largest, without robust setae,

setal cluster formed by slender setae; lateral margin

with 2 robust setae, plumose setae along entire length.

Exopod 0.83 length of endopod; medial margin convex,

with 3 robust setae, plumose setae on distal two-thirds;

apex subequally bifid, without robust setae, setal clmstcr

formed by slender setae; lateral margin with 7 large

robust setae, plumose setae along entire length.

Sexual dimorphism. Females differ from males in

the primary sexual characters and do not develop

the dense patch of setae found on the dorsal

surface of the pleotelson or the prominent lateral

tubercles found on pleonite 5. The tubercles on

pleonites 3-4 and on the pleotelson arc also less

conspicuous in females and are absent in speci¬

mens smaller than approximately 20 mm. The

indistinct tubercles on the ccphalon and perconite

1 are also absent in females.

Variation. Pleotelson and uropod robust setal

counts from margins (N = 20, subsample of 10

males and 10 females from AM P44797): Pleotel¬

son: 4:4 (15%), 4:5 (25%), 5:5 (55%). 5:6 (5%).

Endopod: (medial) 5 (20%), 6 (55%), 7 (25%);

(lateral) 2 (100%). Exopod: (medial) 3 (95%), 4

(5%); (lateral) 5 (15%), 6 (45%). 7 (40%). In

males, development of tubercles on the somites

and the patch of setae on the pleotelson appear

to be associated with maturity. Males approxi¬

mately 9 mm long lack tubercles, while males

approximately !2 mm long have tubercles

on the ccphalon and pereonitc 1, and weakly

developed on the pleonites. Males of 12 mm in

length, and shorter, lack the patch of setae on the

pleotelson.

Size range. Adults 7.5-21 mm.

Etymiolog}’. Dissimilis^ Latin, dissimilar, referring

to the sexually dimorphic pleotelson and pleon.

Distribution. Off Kimberley region. Western

Australia; Darwin Harbour, Northern Territory;

Torres Strait, Queensland; 8-20 m.

Remarks. See Cirolana comata. Kcable (1997)

recorded that in Darwin Harbour Cirolana dis-

similis (as C sp. I) was one of the numerically

dominant scavengers collected among subtidal

rock and coral reef habitats, and was also col¬

lected in lower numbers from the scour zone of

the main channels where gravel sediments pre¬

dominate.

Acknowledgements

I thank Drs J. Lowry and P. Berents for making

the material available for study, and Dr F. Wells

for collecting and donating the specimens from

Western Australia. Drs N. Bruce and R. Brusca

kindly provided unpublished character lists for

cirolanid isopods which were useful in writing the

species descriptions. 1 am also grateful to Dr P.

Berents, Dr G. Poore and an anonymous reviewer

for comments on drafts of the manuscript, and to

Mr R. Springthorpe who composed and inked my

illustrations. This study was undertaken while in

receipt of an Australian Museum Collection

Visiting Fellowship.

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Memoirs of Museum Victoria 58(2): 365-371 (2001)

REDESCRIPTION OF THE TROPICAL AUSTRALIAN ISOPOD, LYIDOTEA

NODATA HALE, 1929 (CRUSTACEA: IDOTEIDAE)

Rachael A. King'-^ and Gary C. B. Poore'

' Museum Victoria, GPO Box 666E, Melbourne, Vic. 3001, Australia

^ Zoology Department, The University of Melbourne, Vic. 3010, Australia

(rking@muscum.vic.gov.au; gpoorc@museum.vic.gov.au)

Abstract

King, R. and Poore, G.C.B., 2001. Redescription of the tropical Australian isopod, Lyidotea

nodata Hale, 1929 (Crustacea: Idoteidac). Memoirs of Museum Victoria 58(2): 365-371.

Lyidotea Hale, 1929 is rediagnoscd and its only species, L. nodata Hale, 1929 rcdescribcd.

Membership of Idoteidac is confirmed but its relationship to other genera remains unresolved.

Introduction

Lyidotea nodata was described by Hale (1929)

from a mature female (SAM Cl699) and several

smaller (possibly immature) specimens from

Flinders I., northern Queensland. He placed the

species in the valviferan isopod family Idoteidac

Samouellc and believed the distinctive flagella of

both pairs of antennae and the fusion of pereonite

7 with the pleotclson justified a new genus. Poore

and Lew Ton (1990, 1993) omitted the genus

Lyidotea from their reviews of the related fami¬

lies Holognathidac Thomson and Idoteidac of

Australia and New Zealand and in the latter paper

explicitly removed it from Idoteidac. They

believed the species’ “habitus, fusion of body

segments, pereopod I, antenna 2 and oostegite 5

are all arcturid-like.” They could not decide on its

family placement but their error has since been

realised (Poore, 2001).

In Australia the Idoteidac are most diverse in

southern latitudes with 22 species recorded south

of 30°S. The only taxa recorded north of this lat¬

itude are rare: Euidotea bakeri (Collinge) and

Paridotea miersi Poore and Lew Ton extending

from the southern coast as far north as 22°S in

WA, Idotea hrevicorna Milne Edwards in Shark

Bay, WA, and Indonesia, and Lyidotea nodata.

Only the last two of these are essentially tropical.

Here, this rather arcturid-like valviferan is

rcdescribcd from new material in the collections

of Museum Victoria and Hale’s (1929) family

placement confirmed. The rediagnosis is neces¬

sary as part of a wider phylogenetic study on the

Arcturidae Dana (where the genus is used as an

outgroup in the analysis) and to correct inaccura¬

cies in Hale’s original description. The generic

diagnosis uses the characters and follows the

format used by Poore and Lew Ton (1993) for

other genera of Idoteidae.

The new material was collected by CSIRO and

Museum Victoria on the North-west Australian

shelf between Dampierand Port Hedland in 1982

and 1983 and from as far south as Roltnest 1. No

new material from Queensland was available.

The taxonomic drawings were prepared with a

camera lucida. The limbs arc drawn to the same

scale. The following abbreviations are used:

NMV, Museum Victoria, Melbourne; WAM,

Western Australian Museum, Perth; SAM, South

Australian Museum, Adelaide.

Lyidotea Hale

Lyidotea Hale, 1929: 35-36.

Type species. Lyidotea nodata Hale, 1929 by

original designation.

Diagnosis. Body semicylindrical; all pereonites

with paired dorsal elevations;. Head as wide as

pereonite I; pereonite 1 fused to head with fusion

indicated by a groove; pereonites 1 to 7 parallel¬

sided; pereonite 7 fused to pleon with no distinct

suture indicated. Pleon with all pleonites fused;

pleotclson apically blunt. Antenna I with single

short flagellar article. Antenna 2 flagellum of 2

articles plus claw. Mandible with well developed

truncate molar process, spine row, lacinia mobilis

and toothed incisor. Maxilla 1 with 2 typical

lobes. Maxilla 2 with 3 typical lobes. Maxil-

lipedal enditc with distal setae; palp with articles

2+3 and 4+5 fused. Coxa completely fused to

pereonal tergites. Pereopod 1 reduced; pereopods

2 to 7 with few spiniform setae; dactylus with

secondary unguis almost as large as primary

unguis. Penes separate, simple and straight. Oost-

egites on maxilliped and pereopods 1 to 5; not

thickened, oostegites 2 to 4 forming the majority

of the marsupium. on fully mature female with

supportive extensions on coxae 2, 3 and 4.

365

366

R. A. KING AND G. C. IB. POORE

Remarks. The ambulatory pereopods, mouthparts

with fused maxilliped palp articles, separate

penes, and absence of uropoda! exopods place

Lyidotea within the Idotcidae. In Arcturidae and

similar families (Poore, 2001) maxillipcdal palp

articles always number five, and there is a single

penial plate. Pcrcopods are usually differentiated

into different functional groups and the uropodai

exopod is rarely absent.

The genus is unique among Idotcidae in fusion

of all plconites to pereonite 7. Fusion of the

head to pereonite 1 is found elsewhere in this

family only in Crahy'zos Bate (sole species,

C longicaudatus\ Poore and Lew Ton. 1993).

There arc many differences in body shape, pereo-

pods and mouthparts between tliese two genera

and the fusion is undoubtedly homoplasious. The

short, compact pereopods. almost gnathopod-like

percopod 1, dorsal body scuplurc, and fusion of

maxillipcdal palp articles 2+3 and 4+5 resemble

the situation seen in species Sytudotea 1 larger

(e.g.. Richardson, 1905; Poore and Lew Ton,

1990). Again, profound differences arc apparent

between Lyidotea and Synidotea, notably in the

possession of a fused penial plate in males and the

absence of oostcgile 5 in females of Synidotea.

The flagellum of antenna 2 of most genera of

Idotcidae is multiarliculate; the short ‘clavate'

condition seen in Lyidotea is distinct yet similar

to that seen in Parasymmerus, Cleantiella and

Ehchsoneila (Brusca, 1984) from America.

These genera have only slight similarities to

Lyidotea in other morphological comparisons. A

further unique condition in Lyidotea is the pres¬

ence of coxal supports under oostcgiles. This is

parallelled in Antarciuridac Poore, llolidoteidae

Wagcle, Austrarcturellidac Poore and Bardslcy

and Rcctarcturidac Poore and may be correlated

with cylindrical body shape (Poore, 2001).

Relationships of Lyidotea to other idotcid

genera remain unresolved.

Lyidotea nodata I laic

Figures 1-4

Lyidotea nodata Bale, 1929: 35-36, fig. 1.

Type material. Flinders I.. Princess Charlotte Bay. Qld,

SAM C1699 (female holotype), SAMA C1845

(paratype) — not examined.

Material examined. Western Australia. North-west

Shelf, between Dampier and Port Medland. CSIRO

Division of Fisheries, RV SoeUu WHOI epibcnthic

sled. 31 NWA .stations within rectangle defined by

18“56.6S, 119^^2.4'E and 20'^1.2'S. 116°57.5E.

30-142 m. 7 Dec 1982-30 Oct 1983, (93 specimens

including males, females, juveniles and mancas. 5 13

mm), NMV collections.

NW of Bluff Point (27^28'S, 133°16'F:). 97 m, 9 Oct

1963. WAM 636-86 (2 males, 9 mm, 10.5 mm); W of

Rottnest I.. (32°00'S. 115“08'E). 135 m, 28 Aug 1973,

WAM 548-73 (1 female, 13 mm).

Illustrated specimens. North-west Shelf, between

Dampier and Port Hedland. I9“27.2'S. 118°58.6'E

36-46 m. 8 Dec 1982 (.stn NWA 346), NMV .123702 (l'

male, 8 mm); 19°29.0'S, 118^S3.5'E. 40-40 m, 12 Feb

1983 (stn NWA 81). NMV J23699 (1 female, 7.5 mm)

W of Rottnest I.. (32^’00'S, I I5”08 E), 135 m, 28 Aug

1973, WAM 548-73 (1 female. 13 mm).

Description. Male. Head with 2 rounded dorsal

elevations between eyes; with setae on elevations,

anterolateral margins rounded, rostral point

absent. Fusion of head and pereonite 1 indicated

by a shallow dorsolateral groove not incised later¬

ally. Pereonite I as long as head, with 2 rounded

dorsal elevations. Pereonites 2 to 4 progressively

longer, each with 2 rounded dorsal elevations at

midlength and 2 smaller posterior dorsal eleva¬

tions. Pleotelson and pereonite 7 fused; pereonite

7 with small dorsal elevations; the whole widest

two-thirds along, tapering to bluntly rounded

apex.

Eyes lateral, prominent. Antenna 1 extending

midway along peduncular article 3 of antenna 2;

flagellum a single hemispherical article with 5

aesthetascs. Antenna 2, 0.46 times body length;

fiagcllum of 2 articles and claw, article 1 with

numerous short setae, 1.3 limes as long as last

peduncular article, article 2 minute, claw longer,

almost straight.

Mandible with truncate triluritive molar; spine

row of 4 spines; left lacinia mobilis with 2 teeth;

incisor 4-toothcd. Maxilla 1 inner lobe with 2

terminal setae; outer lobe with 10 distal robust

setae. Maxilla 2 inner lobe with 10 plumose setae;

middle lobe with 3 setae; outer lobe with 4 setae.

Maxillipcdal endite with 5 long setae; 1 coupling

hook; small oostegite present; palp oval, twice as

long as wide, twice as wide as article I; articles 2

and 3 fused, with dislomesial row of 11 setae,

with oblique distal suture between this and more

distal articles; articles 4 and 5 fused, with mesial

row of 9 seatc and 5 more scattered setae disially.

Percopod 1 ambulatory, compact; basis-rnerus

with few setae; carf)us cylindrical, with 2 mesial

setae near posterior margin; propodus with semi¬

circular blade produced from palm, mesial face

with curved row of 13 barbed setae; dactylus

almost as long as propodus, with mesial setae,

with primary and secondary unguis. Pereopods 2

to 7 similar, with scattered setae; propodi cylin¬

drical; dactylus tapering, with rounded denticles

along posterior margin, setose, with primai 7

secondary unguis.

REDESCRIPTION OF THE ISOPOD, LYIDOTEA NODATA

367

Figure 1. Lyidotea nodata male (NMV J23702): a, dorsal view; b, lateral view. Female (NMV J23699): c, dorsal

view; d. lateral view. Scale = 1.0 mm.

368

R. A. KING AND G. C. B. POORE

Figure 2. Lyidotea nodata female (NMV J23699): antennae 1 and 2, left maxilliped, left mandible, left maxillae 1

and 2, pereopods 1 to 5. Scales = 0.5 mm: a, A2; b, Al; c, MD, MXl, MX2, MP; d, P1-P5.

REDESCRIPTION OF THE ISOPOD, LYIDOTEA NODATA

369

Figure 3. Lyidotea nodata female (NMV J23699): pereopods 6 and 7, uropod, ventral view detailing oostegites.

Male (NMV J23702): pleopods 1 and 2. Scales = 0.5 mm: a, P6, P7; b, PLl, PL2; c, U. Scale = 1 mm: d, ventral

view of female.

370

R. A. KING AND G. C. B. POORE

Figure 4. Lyidotea nodata male (NMV J23702): pleopods 3, 4 and 5, ventral view of pereonite 7 and pleon show¬

ing position of penes in situ. Female (NMV J23699): ventral view of fully mature female detailing oostegites.

Female (WAM 548-73): ventral view. Scale = 0.5 mm: a, PL3-PL5. Scales = 1.0 mm; b, ventral views.

REDESCRIPTION OF THE ISOPOD, LYIDOTEA NODATA

371

Pleopod 1 peduncle longer than wide, with

4 coupling hooks; cndopod with 33 mesial and

distal long plumose setae, longest as long as

endopod; exopod with 27 long plumose setae

concentrated distally. Pleopod 2 peduncle little

longer than wide; cndopod with 15 distal plumose

setae, as long as endopod; appendix masculina

simple, straight and blunt, reaching beyond the

apex of endopod; exopod with 40 marginal

plumose setae. Pleopod 3 to 5 progressively

larger, without long marginal setae. Uropodal

cndopod 1.5 times as long as wide at base,

tapering to rounded-truncate apex.

Female. Differs from the male in smaller size

and wider pereonites. Small ostegite on maxil-

liped; ooslcgitc on pereopod I small, posteriorly

directed; oostegites on pcrcopods 2 to 4 elongated

along body, with distal transverse sutures; pereo-

pods 2 to 4 with L-shaped coxal extensions fomi-

ing thickened lateral edges supporting oostegites,

oostegites 5 small, meeting in midlinc (Fig. 3D,

4C).

Distrihiition. North Queensland, Western

Australia; subtidal to 140 m depth.

Remarks. This redescription is the first illustration

of a male, unavailable to Male (1929), and also

illustrates the oostegites for the first time. A

female with slightly more developed oostegites

(WAM 548-73) was found after the description of

the female (NMV J23699) was completed. Both

possess functional oostegites and all other

appendages arc the same but it was decided that

the larger female be illustrated to show the

slightly more developed condition. Hale failed to

note the second article of the antenna 2 flagellum.

Acknowledgements

Provision and collection of these specimens was

possible through the courtesy of CSIRO Marine

Science Laboratories, Hobart. Field work was

funded by a grant to G.C.B. Poore from the Aus¬

tralian Biological Resources Study, Environment

Australia, Canberra.

References

Brusca, R.C., 1984. Pliylogeny, evolution and biogeog¬

raphy of the marine isopod subfamily Idoteinae

(Crustacea: Isopoda: Idoleidac). Transactions of

the San Diego Society of Natural History 20:

99-134.

Hale, H.M., 1929. Crustacea from Princess Charlotte

Bay, north Queensland. The Isopoda and Stom-

atopoda. Transactions of the Royal Society of

South Australia 53: 33-36.

Poore, G.C.B.. 2001.1.sopoda Valvifcra: diagnoses and

relationships of the families. Journal of Crus¬

tacean Biology 21:213-238.

Poore, G.C.B. and Lew Ton, H.M, 1990. The Holog-

nathidae (Crustacea: l.sopoda: Valvifera) expanded

and redefined on the basis of body-plan. Inverte¬

brate Taxonomy 4: 55-80.

Poore, G.C.B. and Lew Ton, H.M., 1993. Idoteidae of

Australia and New Zealand (Crustaeea: Isopoda:

Valvifera). Invertebrate Taxonomy 7: 197-278.

Richardson, H.. 1905. A monograph on the isopods of

North America. United States National Museum

Bulletin 5A: VILLIII, 1-727.

Memoirs of Museum Victoria 58(2): 373-382 (2001)

PLAKARTHRIUMAUSTRALIENSE, A THIRD SPECIES OF

PLAKARTHRIIDAE (CRUSTACEA: ISOPODA)

Gary C. B. Poore' and Angelika Brandt^

'Museum Victoria, GPO Box 666E, Melbourne, Vic. 3001, Australia

(gpoore(^museum.vic.gov.au)

^Zoological Institute and Museum, Martin-Lulher-King Platz 3, 20146 Hamburg, Germany

(abrandt@zoologie.uni-hamburg.de)

Abstract

Poore. G.C.B. and Brandt, A., 2001. P/akarthnum australiense^ a third species of Plak-

arthriidae (Crustacea: Isopoda). Memoirs of Museum Victoria 58(2): 373-382.

The Plakarlhriidae share with four genera of Sphacromatidae a flat habitus and a marginal

row of setae on antenna 1, coxae and uropods, modified to maintain contact with the substrate.

They do not possess areas of scales on pleopod 5, a probable sphacromatid synapomorphy. The

possibility exists that Plakarlhriidae arc a highly derived offshoot of a clade that also gave

rise to the flattened “cassidiniine” sphaeromatids. Plakarthriiim australiense sp. nov. is

described from the southern Australian marine shelf and compared with the only two other

species of the genus and family. The new species differs from P. typicum from New Zealand

and P. punctatissimum from subantarctic islands of the South Atlantic and Antarctic Peninsula

in having pleonite 1 fully indicated dorsally, only pleonite 2 indicated laterally and pleotelson

tapering to 80% of its anterior width posteriorly.

Introduction

The Plakarthriidae Hansen, 1905 are previously

known from only two species, Plakarthrium

typicum Chilton, 1883 and P. punctatissimum

(Pfeffer, 1887). The family is confined to shallow

marine shores in cool-tcmpcratc and cold

environments of the Southern Hemisphere.

Plakarthrium typicum was described from New

Zealand and has been reported as far south as The

Snares islands (48‘’S) (Hurley, 1961; Poore,

1981) but not from Macquarie Island (GCBP col¬

lections in Museum Victoria). It has also been

reported from southern Chile (Jaramillo. 1977)

but this record is more likely to be the second

species, P. punctatissimum, if not an undescribed

Chilean endemic. Plakarthrium punctatissimum

is reported from subantarctic islands of the

South Atlantic and Antarctic Peninsula, namely

South Georgia (type locality). South Orkneys,

Petcmiann I., Booth-Wandel I. and Anvers I.

(Richardson, 1906, 1913; Tattersall, 1921; Pcsta,

1928; Wilson ct al., 1976). A third species from

southern Australia is here reported.

The taxonomic affinities of Plakarthrium

Chilton, 1883 to other '‘flabellifcraiT* isopods has

remained enigmatic. Hansen (1916) erected the

Plakarthriinac as a subfamily of Sphacromatidae

and Menzies (1962) believed them to belong in

the “platybranchiata” group of this family, a

group more or less equivalent to Cassidininae

Hansen, 1905 (Iverson, 1982; Harrison and Ellis,

1991; Bruce, 1994). The monophyly of this and

other sphacromatid subfamilies remains doubtful

(N.L. Bruce, pers. comm.). The affinities of

Plakarthriidae were reviewed with a redescription

of P. typicum by Wilson et al. (1976) who con¬

cluded “that the Plakarthriidae are unique with no

close affinities with any other family.'’ Wagele

(1989) placed the family within his suborder

Sphaeromatidca, sister taxon to a clade including

Serolidae, Balhynataliidae and Sphacromatidae.

Brusca and Wilson (1991) also assessed their

position, placing the family as sister taxon to

Serolidae, and possibly Keuphyliidae and Bathy-

nataliidae. This paper, describing a new Aus¬

tralian species, also addresses some newly recog¬

nised characters which may have a bearing on

plakarthriid relationships to Sphacromatidae in

particular.

Pfeffer’s (1887) generic name Chelonidium is a

junior synonym of Plakarthrium Chilton, 1883, a

fact realised by Richardson (1904: 6; 1906: 6)

who synonymised the two. Plakarthriidae

(erected by Hansen (1905) as Plakarthriinac, a

subfamily of Sphacromatidae Latreille, 1825) is

the family name to have been used by most

authors and, under ICZN (1999) Article 40,

replaces Pfeffer’s (1887) family name, Cheloni-

diidac. Both dales should be cited for the family.

Wilson et al. (1976) who incorrectly attributed

373

374

G. C. B. POORE AND A. BRANDT

the family name to Richardson (1904) reviewed

later accounts of plakarthriids by Richardson

(1906, 1913), Tattersall (1921), Pesta (1928),

Nierstrasz (1931), Menzies (1962), Glynn (1970)

and Kussakin (1973).

Material is lodged in Museum Victoria,

Melbourne (NMV) and the Zoological Institute

and Museum, Hamburg (ZMH).

Plakarthriidac Hansen, 1905 (1887)

Chelonidiidac Pfeffer, 1887: 85.—Pesta. 1928.

Plakarthriinae Hansen, 1905: 100.—Nierstrasz,

1931: 192.

Plakarthriidac.—Richardson, 1906: 6.—Richardson.

1913: 7.—Wilson el al., 1976: 334 (see above for other

references).

Diagnosis. First coxa free and articulating.

Margin of the body being defined by peduncular

articles 1 and 2 of antenna 1, peduncular articles

3 and 4 of antenna 2, all coxae, uropodal

rami. Plcotclson completely fused and without

articulation.

Composition. There is only one genus.

Remarks. The family was rediagnosed by Wilson

el al. (1976) and is recognised by the combination

of characters given in the diagnosis. Only the sec¬

ond character, the arrangement of the marginal

segmentation, is unique. Similar patterns involv¬

ing an expanded peduncle of antenna 1 and

uropods are seen in some genera of "cassidiniinc"

Sphacromatidac, in particular, Amphoroidella

Baker, 1908, Leptosphaeroma Hilgendorf, 1885,

Paraieptosphaeroma Buss and Iverson, 1981. and

Platysphaera Holdich and Harrison, 1981. In all

four genera the body is flattened as in Plakarthri¬

idac but antenna 2 docs not contribute to the body

margin and the uropodal staicturc is typically

sphaeroniatid with peduncle and endopod fused.

Further, all possess the cuticular scale patches on

the end of the exopod of pleopod 5 which may be

a sphaeromatid synapomorphy absent in

Plakarthriidac (N.L. bruce, 1993, fig. IF; pers.

comm.). The four genera arc unquestionably

sphaeromatids and similarity to Plakarthriidac

seems on this evidence to be convergent. Another

similarity, an adaptation to maintaining a smooth

profile and minimum resistance to water move¬

ment, is what Buss and Iverson (1981) have

called the membrana cingula, a fringe around the

edge of the body of webbed marginal setae. The

membrana cingula of Paraieptosphaeroma glynni

was illustrated in SEMs by Buss and Iverson

(1981: fig. 4) and of the two other species of this

genus in line drawings by Muller (1990) and

Kussakin and Malyutina (1993). Bruce (1994: fig.

50B) figured the membrana cingula for

Platysphaera membranata Holdich and Harrison,

1981. Kwon (1990) figured a similar structure in

Leptosphaeroma gottschei Hilgendorf and we

have observed the same in Museum Victoria

material of Amphoroidella elliptica Baker. While

there is a strong similarity between the membrana

cingula in the four sphaeromatid genera, the

equivalent structure in Plakarthriidac comprises

broad separate fringed overlapping setae and is

complemented by a free skirt-like flange of

webbed setae underneath and parallel to the

margin of the body.

Within Flabcilifcra the pIcon fully fused to the

plcotclson is seen in at least one species of

Sphaeromatidae, Maricocens hrucei Poore, 1994.

This and similar genera, Juletta Bruce, 1993 and

Margueritta Bruce, 1993 (both with free pleonitc

1), are placed in the Dynamcninae and any

similarity to Plakarthrium is coincidental.

The possibility exists that Plakarthriidac arc a

highly derived early offshoot of a clade which

also gave rise to fiattened “cassidiniinc'’ sphaero¬

matids. In this scenario some reversals may have

occurred (e.g. free coxa 1) as well as numerous

synapomorphics in limbs and plcotclson.

Plakarthrium Chilton, 1883

Plakarthrium Chilton, 1883: 74. - Hansen, 1905:

115,—Menzies, 1962: 128.—Wilson ct al., 1976:

334 -335. (type species: Plakarthrium typicum Chilton,

1883 by monotypy).

Chelonidium Pfeffer, 1887: 86 (type species Chcloni-

dium pxmetatissimum Pfeffer, 1887 by monotypy).

Remarks. The genus has the characters of the

family. Only Chilton (1883) gave a diagnosis.

Plakarthrium australiense sp. nov.

Figures 1-5

Material examined. Holotype. Australia, Tasmania,

Biclieno, granite reef 50 m offshore, N end of “The

Gulch" (41°53'S, 147®18'E), 7 m. encrusting foliose

alga, SCUBA, G.C.B. Poore and 11.M. Lew Ton, 22

Mar 1988 (stn TAS-87), NMV J47067 (1 ovigerous

female, 3.2 mm, with 3 slides).

Paralypes. Collected with holotype: NMV J27472 (1

male. 2.2 mm: 8 females, 1.9 2.8 mm. 1.9 3.2 mm);

NMV J47021 (I male, 2.4 mm); NMV J47068 (2

females, 3.2, 2.3 mm): ZMH K-39833 (3 females).

Other material. Tasmania. Waterhouse Point. NMV

J47066 (I juvenile, 1.9 mm). Western Bass Strait, 6 km

W of Currie, King I., (39'=54.7*S. I43''43.4'E), 49 ni.

NMV J23890(8specimens. I.0-!.6mm). PegIcgCove,

Deal I., between Pulpit Cove and Winter Cove

(39°28'S, 147°22'E), 6 m, NMV J23891 (1 male. 1.7

NEW AUSTRALIAN ISOPOD OF THE GENUS PLAKARTHRIUM

375

Figure 1. Plakcirthrium australiense sp. nov. Holotype, NMV J47067: a, b, dorsal and lateral views, c, pereonite 7, pleo-

telson and uropods.

Plakarthrium typicwn Chilton. Male, NMV J683: d, pereonite 7, pleotcison and uropods.

376

G. C. B. POORE AND A. BRANDT

Figure 2. Plakarthrium australiense sp. nov. Holotype, NMV J47067: Al, A2, antennae 1, 2; MDI, MD r, left and right

mandibles; MXl, MX2, maxillae 1, 2; MP, maxilliped. Paratype male, NMV J4702I: Aim, A2m, antennae 1, 2.

NEW AUSTRALIAN ISOPOD OF THE GENUS PLAKARTHRIVM

Figure 3. Plakarthrium australiense sp. nov. Holotype, NMV J47067: P1-P7, pereopods 1-7.

378

G. C. B. POORE AND A. BRANDT

Figure 4. Plakarthnum ciu.s(raliefne sp. nov. Holotype, NMV J47067: PLI-PL5, plcopods 1-5; U, right uropod; X, ven¬

tral view of margins of IcA uropod and coxa 7.

mm). Hogan 1., Landing Beach (39°13'S, I46®59*E),

ZMH K-39834(l male, 2.0 mm).

Victoria, 75 m SW of Eagles Nest, Venus Bay

(38°40*S, 145°40'E), 8 m. NMV J23893 (1 male. 3.0

mm; 1 female, 2.0 mm, 2.0-3.0 mm). 1 km E of

Harmers Haven, 500 m offshore (38°34'S, 145°40'E),

11 m, NMV J23896 (1 male, 2.1 mm; 1 female, 2.0 mm,

2.0-2.1 mm). Cape Paterson, E side (38°4i’S.

145°36'E), 6-3 m, NMV J23895 (1 male, 1 female, 4

immature). 50 m S of Twin Reefs, Venus Bay (38°4rS,

145'^39'E), 9 m, NMV J23894 (1 male. 2.0 mm).

Wilsons Promontory, Hobbs Head, NE shore (39°2'S,

NEW AUSTRALIAN ISOPOD OF THE GENUS PLAKARTHRIUM

379

Figure 5. Plakarthriwn australiense sp. nov. Paratypc male, NMV J47021: PI, P2, pereopods 1,2; PLl, PL2, pleopods 1,

2; P, penes; U, uropod.

146°28'E), NMV J47063 (1 female, 2.6 mm). Dutton

Way, Portland (38°I8.4 S, I4P36.4'E), 3 m. NMV

J47065 (1 ovigerous female, 3.0 mm).

South Australia, Beachport, Snapper Point

(37°29.3'S, I39°59.6'E). 6 ni. NMV J20434 (I female,

2.4 mm). NE side of Topgallant 1., Investigator Group

(33‘’43.00'S, 134°36.60‘E), 12 m, NMV J23892 (1

male, 1.8 mm).

Western Australia, Breaksea I., SW comer

(35“3.90'S, 118^2.50'E). 15 m. NMV J23887 (2 man-

cas. 1 male, I.0-1.9mm): NMVJ23888(1 immature; 1

female, 2.1 mm; 4 males, 1.9-2.0, 1.6-2.1 mm). King

George Sound. N of False I. (35°0.7()'S, 118°10.10'E),

27 m, NMV J23889 (4 males, 1.7-2.6 mm: 2 females,

1.9, 3.0 mm. 1.7-3.0 mm).

380

G. C. B. POORE AND A. BRANDT

Other species. P. typicum. New Zealand. Auckland

region, Tawharanui Peninsula. 6 m, Carpaphvihim Jlex-

uosum, R. Taylor. 3 Jun 1992, NMV J37I02 (14).

Opposite Edward Pcrcival Marine Station. Kaikoura

(42°25'S. 173°42'E). 2 m. algae on boulders, G.C.B

Poore et al.. 23 Mar 1997 (stn NZ97-022), NMV

J39561-J39654 (25). The Snares Islands. D.S. Homing.

NMV J682 (male, with 3 slides): NMV J683 (several).

P. punctatissimum. South Georgia. Royal Bight,

Mollke Harbour (54^30.58'S. 36"0.45'\V). leaves of

Macrocystis, Dr Karl von den Steinen. NMV J473II (3

syntypes donated by Zoological Institute and Museum,

Hamburg).

Diagnosis. Pleotelson with pleonitc 1 fully indi¬

cated dorsally (not articulating), only pleonite 2

indicated laterally; pleotelson tapering to 80% of

anterior width posteriorly. Distal margins of

uropods aligned with margin of coxae. Coxae

with acute posterior comers.

Description. Holotypc female. Body c. 1.35 times

as long as wide; ration of lengths of antenna I :

head : pereon : pleotelson : uropod in midline =

13 : 18 : 33 : 27 : 9; widest at coxa 4 where each

coxa comprises c. 27% of width. Head 1.9 times

as wide as long, evenly rounded laterally, with

excavated triangular front with triangular clypeus;

eyes with c. 10 ommatidia. Pereonite 1 narrower

than head; pereonite 3 widest; perconites 6 and 7

shorter than others. Pleotelson with pleonite I

fully indicated dorsally (not articulating), only

pleonite 2 indicated laterally; pleotelson tapering

to 80% of anterior width posteriorly; posterior

notch semicircular, about third width of posterior

margin, bordered by fine setae. Marginal articles

of antennae I and 2 and coxae of pereopods 1-7

each with small triangular lobe on anterodistal

and posterodistal comers; lower surface with

flattened marginal surface (c. fifth of coxal

length) defined proximally by a free skirt-like

flange parallel to margin.

Antenna 1 peduncle with broad article 1, tri¬

angular article 2 and smaller tapering article

3 inserted in notch on lateral margin of article

2; fiagellum of 2 articles, first ring-like with

2 pappose setae, second with 4 aesthctascs and

5 simple setae. Antenna 2 peduncle with small

articles 1 and 2; article 3 short, broad; article 4

4-sided; article 5 cylindrical, longer than wide,

inserting on underside of margin of article 4;

flagellum of 12 cylindrical articles.

Mandible incisor with 2-3 obscure blunt teeth;

spine row of 3 spines; without palp or molar.

Maxilla 1 with single lobe having 7 short apical

robust setae. Maxilla 2 with single lobe with 8

short apical setae. Maxilliped without cpipod;

endite 4 times as long as wide, with 3 apical

robust setae and 6 slender setae; palp with short

article 1, longer article 2, substantially longer and

broader article 3, convex and setose mesially,

article 4 0.6 times as wide as article 4, article 5 as

long as and shorter than article 4, with 9 mesial

and distal setae.

Pereopod 1 elongate, longer than similar pereo-

pod 2; pereopods 3 5 compact; pereopods 6-7

more elongate. Pereopod I merus with anterodis¬

tal row of complex setae; carpus half as long as

merus, with minute tubercles on posterior

margin; propodus narrow; dactylus compact with

3-toothcd unguis. Pereopod 2 c. 85% length of

peroopod 1; meats with 2 distal robust setae;

compact dactylus with 3-toothcd unguis. Perco-

pod 3 less than half length of pereopod 1; ischium

with posterodistal denticles; mems with

anterodistal row of complex setae; carpus almost

triangular, with posterodistal short robust seta;

propodus swollen; dactylus short, with simple

curved unguis. Pereopods 4 and 5 similar to pere¬

opod 3, 5 the most compact. Pereopod 6 about as

long as pereopod 2; merus with 1 very stout and 1

shorter complex anterodistal setae; propodus with

row of 4 simple posterior setae distally; short

dactylus with 3-loothcd unguis. Pereopod 7

longer but similar to pereopod 6.

Plcopod I peduncle twice as wide as long, with

2 coupling hooks on mesiodistal comer; rami

attached on lateral half, not overlapping; endopod

and exopod with 13 and 11 mesial and distal

plumose setae respectively. Pleopod 2 peduncle

wider than long, with 2 coupling hooks; rami not

overlapping; endopod and exopod with 10 and 12

distal plumose setae respectively. Pleopod 3 with

peduncle wider than long, with 2 coupling hooks;

endopod triangular, without setae; exopod adja¬

cent to endopod, with 7 distal plumose setae.

Plcopods 4 and 5 similar to plcopod 3 but smaller

and relatively wider; exopods with 8 and 9 distal

plumose setae respectively.

Uropod peduncle strongly tapering; endopod

attached mesially to peduncle, 2.5 times as long

as greatest width, triangular in lateral view, with

setose rounded margin anteriorly bordering

respiratory orifice strongly produced vertically,

with 6 pappo.se setae near articulation with pedun¬

cle, mesial margin straight, posterior margin

curved and with c. 5 overlapping shingle-like

scales; exopod c. 1.5 times as long as wide, with

curved mesial margin against endopod, straight

lateral margin, convex distal margin with c. 11

overlapping shinglc-likc .scales.

Male. Differing slightly from female in some

NEW AUSTRALIAN ISOPOD OF THE GENUS PLAKARTHRIUM

381

proportions. Pleotelson more elongate. Antenna

1 with article 2 of peduncle more rounded;

flagellum with 6 acsthetascs. Antenna 2 with flag¬

ellum of 10 articles. Pereopods 1 and 2 slightly

more robust. Pleopod 2 endopod with appendix

masculinis twice as long as endopod, attached

two-thirds way along. Uropod with narrower

peduncle and rami than in female. Penes oval,

fused basally, attached to posterior margin of

pereonite 7.

Oistrihiition. Southern Australia, from central

Victoria to southern Western Australia, including

Tasmania; 6^9 m, on Amphibolis, Eckionia

holdfasts, red algae. No records from New South

Wales (P, Berents, Australian Museum, pers.

comm.).

Remarks. The Australian species differs from the

other two species of Plakarthrium in having a

clear demarcation dorsally of pleonite 1. In

P. typicum (see Fig. Id) and P. punciatissimum

pleonite I is indicated only laterally. Plakar-

ihrium austvaliense differs from P. typicum,

which it most resembles, in having the pleotelson

tapering rather than more or less parallel-sided,

and the distal margins of the uropods aligned with

the margin of the coxae. In P. typicum, the New

Zealand species, the uropodal exopods arc more

oval and protnide beyond the end of coxae of

pereopods 7 (Fig. Id). Plakarthrium typicum

lacks the acute comers of the coxae seen in the

Australian species.

Plakarthrium australiense differs from P. pimc-

tatissimum more markedly. The latter has

pleonites 1-4 indicated laterally and uropods

produced more posteriorly.

The new species has a more pronounced lower

marginal contact surface on coxae, antennae and

uropods than the other species. This surface seals

the underside of the animal and is made more

effective by the skirt-like fringe along its inner

edge.

Acknowledgements

We thank R. Taylor, University of Auckland, G.

Fenwick, University of Canterbury, and G. Edgar,

University of Tasmania, for donating material of

the New Zealand and Australian species. We

thank too P. Berents for checking for the presence

of the new species in collections of the Australian

Museum. Field w'ork by GCBP was supported by

Australian Biological Resources Study grants.

We are grateful for comments on the manuscript

by Buz Wilson and Niel Bruce.

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Memoirs of Museum Victoria 58(2): 383-410 (2001)

A PHYLOGENY OF THE LEPTOSTRACA (CRUSTACEA)

WITH KEYS TO FAMILIES AND GENERA

Genefor K. Walker-Smith'-^ and Gary C. B. Poore'

'Museum Victoria, GPO Box 666E, Melbourne, Victoria 3001, Australia

-Department of Zoology, University of Melbourne. Victoria 3010, Australia

(gwsmith(S}museum.vic.gov.au, gpoore(^museum.vic.gov.au)

Abstract

Walker-Smith. G.K. and Poore, G.C.B., 2001. A phylogeny of the Lcptostraca (Crustacea)

with keys to families and genera. Memoirs of Museum Victoria 58(2): 383-410.

A phylogenetic analysis of the Lcptostraca Claus, 1880 is undertaken using 32 of the 41

known species (including 4 undescribed species). The value of outgroups for deriving a plau¬

sible phylogeny in a group whose affinities remain contentious is discussed. A hypothetical

ancestor is considered the best solution to the problem and states were scored based on gen¬

eral principles of crustacean evolution as evidenced by a wide variety of taxa. States of the 43

characters used in the analysis are detailed. The new phylogenetic hypothesis is compared with

those of Olcscn (1999). We conclude that a phylogeny based on species-level taxa and many

infonnative characters is more likely to represent true evolutionary reIation.ships than one

based solely on genera and few characters. A new classification based on the phylogeny is

derived with a new family, Paranebaliidae, being erected for Paranebalia Claus. 1880 and

Levinchalia Walker-Smith, 2000. Nebaliopsididac Hessler, 1984 is supported iov Nchaliopsis

Sars. 1887. A restricted Nebaliidae Samouelle, 1819 for the remaining genera {Speonebalia

Bowman, Yager and IlifTe, 1985, NebalieUa Thiele, 1904, Dahlella Hessler, 1984 and NebalUi

Leach. 1814). Sarsinebalia Dahl, 1985 is synonomised with vVc/w//a. New keys and family and

generic diagnoses are presented. All known species are listed with notes on distribution.

Introduction

Lcplostracans arc marine crustaceans of the mala-

costracan subclass Pliyliocarida. Leptostracans

have many derived features that separate tliem

from other malacostracans; the loss of ambula¬

tory function of the thoracic limbs, which now

only function in feeding, respiration and brood

protection (Hessler and Schram, 1984; Dahl,

1976); a movable rostrum (Schram, 1986; Olc¬

scn, 1999); the scale-like ramus of the first

antenna (Hessler and Schram, 1984; Olesen,

1999); uniramous antenna 2 (Hessler and

Schram, 1984; Olesen, 1999); reduction of

pleopods 5 and 6 (Hessler and Schram, 1984;

Olesen, 1999); and direct larval development

(Manton, 1934; Hessler and Schram, 1984).

The first species of the Order Lcptostraca

Claus, 1880 was described by Otto Fabriciiis in

1780, as Cancer bipes from east Greenland.

Herbsl {1796) later relegated this species to sub-

specific status. Cancer gamarelius bipes (cited in

Sars, 1896). Leach (1814) introduced the genus

Nehaiia for Cancer bipes and a new British

species N. herhstii Leach, 1814.

The genera of Leptostra are distributed differ¬

ently. Nebalia is cosmopolitan. In contrast,

Levinebalia Walker-Smith, 2000 has been

recorded only in Australia and New Zealand.

Paranebalia Claus, 1880 is found in central

America, Bcmiuda, New Caledonia and Aus¬

tralia. Nebalielhx Thiele, 1904 is confined to cold

waters, being found in Antarctica, southern Aus¬

tralia and the high latitudes of the Northern

Hemisphere. Speonebalia Bowman, Yager and

Iliffc, 1985 has been recorded from only marine

caves in the Turks and Caicos, and Dahlella

Hessler, 1984 was collected from hydrothermal

vents near the Galapagos. Nebaliopsis Sars, 1887

is a pelagic genus with a world-w'idc distribution,

Leptostracans have been recorded in waters

from I m deep (Modlin, 1991) to more than

2000 metres (Page, 1929). Most species occur in

less than 200 metres. Water temperatures

influence the length of time taken to reach

maturity, the size at maturity and the incubation

time of young (Macquart-Moulin and Casticbon,

1983).

Until now three families have been recognised:

Nebaliidae Samouelle, 1819; Nebaliopsididac

Hessler, 1984; and the Permian Rhabdouraeidae

Schram and Malzahn, 1984. Messier's (1984)

original spelling, Ncbaliopsidac, is incorrect as

family names based on genera ending in "‘-apsis”

should end “-opsididae” (c.g. Sivertsen and

Holthius, 1980). Seven nominal genera (Nehaiia;

383

384

G. K. WALKER-SMITH AND G. C. B. POORE

Paranebalia: Nebaliella: Dahlella; Sarsinebalia

Dahl, 1985; Speonebalia\ and Levinebalia) and

36 extant species are contained in the family

Nebaliidae. Nebaliopsididae consists of the

monotypic genus Nebaliopsis. Rhabdouraea

Malzahn, 1962 is the monotypic fossil genus of

Rhabdouraeidae.

Martin ct al. (1996) reviewed the morphology

and natural history of Nebalia hessleri Martin,

Vetter and Cash-Clark, 1996 and provided a key

to the extant families and genera of Leptostraca

then accepted.

Olcsen (1999) conducted a phylogenetic analy¬

sis of the seven extant lepioslracan genera then

known and described Nebalia bnicei Olesen,

1999. Olesen questioned the monophyly of

Nebalia and stated that as he could not find

unique characters for this genus he could not

exclude the possibility it is paraphylctic with

Sarsinebalia and/or Dahlella. However, he

maintained the status of these three genera.

The present study is a derived from an unpub¬

lished BSc(Hons) thesis (Walker-Smith, 1993)

and presents a detailed phylogenetic analysis of

the Leptostraca using 32 of the 41 known species

(including four undescribed species). The mono¬

phyly of genera is tested and relationships

between the included species deduced as far as

possible. The value of certain characters in lep-

tostracan systematics is discussed, as are the

results of the phylogenetic analyses. The value of

outgroups in deriving plausible phylogenies in a

group whose affinities remain contentious is also

discussed. A new classification based on phylo¬

genetic principles is derived and new keys and

new family and generic diagnoses are offered. All

known species are listed with distributional notes.

G/'oh7/; in Leptostraca

Transformations in the shape of the carapace,

both pairs of antennae, pleopods, furca and other

features are gradual from moult to moult in imma¬

ture and subadult males and deviate from the

female morphology, which generally remains

unchanged except when reproductive (Dahl,

1985).

In species of Nebaliidae the terminal article of

the juvenile thoracopod endopod is elliptic with

thin marginal setae (Figs 4d, 4c). In sexually

mature females carrying eggs or embryos, the

entire endopod becomes elongated and the ter¬

minal article becomes enlarged, generally sitting

at right angles to the thoracopod axis (Fig. 41).

The exterior and terminal edges of the temiinal

article possess a dense annaturc of plumose setae

that are long, strong and curved, and interlock

with those of opposite and neighbouring thora-

copods to form the floor of the brood chamber.

Embryos develop in the brood pouch and when

the Juveniles are ready to leave the long setae

forming the floor of the chamber drop off, leaving

behind a pattern of ridges and furrows that are the

scars of seud attachment (Dahl, 1985). At this

stage, the terminal article of the endopod differs

so markedly from those of males and immature

females that they could be presumed to belong to

a different species (Dahl, 1985; Figs 6-10). The

exopod and epipod do not change shape during

this metamorphosis.

The eggs of Nebaliopsis are thought to be shed

directly into the water (Cannon, 1931, 1960) but

Brahm and Geiger (1966) reported Nebaliopsis

with eggs developing under the carapace. These

eggs appeared to be contained in a “basket formed

by the large and setose posterior pair of thoracic

appendages, that extend anteriad to the area of the

mouth parts” (Brahm and Geiger, 1966; 41-42)

and were shed when the specimens were placed in

fixative.

Ta.xonomic confusion in Leptostraca

In the past, failure to recognise characters related

to the sex and maturity of leptostracans resulted in

taxonomic confusion. Thomson (1879) described

Nebalia longicornis without taking sexual dimor¬

phism into account and thus recognised the elon¬

gate flagellum of antenna 2 as a specific character

rather than one of sexually mature males. Claus

(1888) added to the taxonomic confusion by

basing his identification of species on few mor¬

phological characters, most of which were

growth- or sex-related and could not be used to

successfully distinguish between genera or

species. Claus's (1888) taxonomic concept of

Leptostraca was followed by subsequent tax¬

onomists (e.g., Thiele, 1904, 1905) and resulted

in the erroneous assumption that each genus

consisted of a few highly variable species. In par¬

ticular, Nebalia bipes (Fabricius, 1780) and

N. longicornis have been reported as geographi¬

cally widespread while, in fact, each comprises

several species. The subspecies described for

each arc likely to be separate species. Many

records of nominal species in areas remote from

their type locality probably refer to undescribed

species. Fortunately, Dahl (1985) recognised the

conservative nature of leptostracan morphology

and redefined many species of Nebalia using new

diagnostic characters. Dahl's assessment of the

European shelf and Southern Hemisphere species

incorporated the description of six new species of

Nebalia (Dahl, 1985, 1990).

PHYLOGENY AND KEYS TO LEPTOSTRACA (CRUSTACEA)

385

Analytical methods

Material for this study is deposited in Museum

Victoria, Melbourne and type specimens of

Nehalia capensis Barnard, 1914 were borrowed

from the South African Museum, Cape Town.

Museum Victoria collections include representa¬

tive species Nehalia, Nebaliella, Paranebalia.

Levinehalia and Nehaliopsis, For most described

species infonnation relating to character states

was obtained from the literature. Thirty-two

species, including four undcscribcd species from

southern Australia, were selected for phyloge¬

netic analysis. Literature relating to six species

and three subspecies of Nebaliidae was either not

obtainable or provided insufficient diagnostic

information; these taxa were omitted from the

analysis. Nebalia gerkenae Haney and Martin,

2000, published later, was not included nor was

the fossil family Rhabdouraeidac.

Cladistic analyses were used to generate trees

of monophylelic groups as hypotheses ol the

relationship between the selected taxa. The

relationships between genera were of ^eatest

interest. Forty-three characters (all parsimony-

infonnativc) were scored for each taxon (Table 1)

resulting in a data matrix of 32 Icptostracan taxa

plus a hypothetical ancestor described by 43

characters (Table 2). Characters were treated as

unordered and unweighted.

The program PAUP* 4.0 (Beta 3 version for

Windows) (Swofford, 1998 and updates) was

used to establish relationships between taxa and

produce a hypothesis from which a classification

might be derived, A heuristic search was made

using most of the default options in the PAUP

block, except for the following commands: OUT-

ROOT=MONOPHYL; ADDSEQ=RANDOM;

NREPS=1000; NCHUCK=3; CHUCKSCORE

= 1; RANDOMIZE=TREES. The two most dis¬

tant parsimony trees were calculated using the

FILTER command and the characters states

changes were mapped on one of these trees. A

50% majority-rule consensus tree of all trees was

generated. Stability of the cladcs was assessed by

bootstrap analysis (using the default settings) and

a 50% majority-rule con.sensus tree of all boot¬

strap trees was constructed. Bremer support

values were calculated for the two most distant

trees using Auto Decay 4.0 (Eriksson, 1998) to

assess the stability of the clades. Trees were

illustrated using Tree View (Page, 1996).

Outgroups

Selection of an outgroup is the major problem

encountered in the phylogenetic study of the Lep-

tostraca. Leptostraca have been considered the

most primitive subclass within the Malacostraca

because they have a primitive caudal furca and

polyramous phyllopodous (flattened, leaf-like)

thoracic limbs used in filter feeding (Claus, 1888;

Manton, 1934; Dahl, 1987, 1992). Hessler and

Newman (1975) believed the relatively high

number of segments and full complement of seg¬

mental appendages should also be regarded as

primitive features. Dahl (1976) supported Hessler

and Newman's (1975) view with the fact that

while Phyllocarida were represented in the Lower

Cambrian, no fossils of the other malacostracan

subclass Eumalacostraca are known until the

Devonian. However, Walossek (1999) disputed

the existence of fossil malacostracans appearing

in the Cambrian and stated that the only clear

record appears after this time.

Other authors have placed the Leptostraca as a

subclass in Phyllopoda with phyllopodous

(polyramous and foliaceous) thoracopods thought

to unite Branchiopoda. Leptostraca and Cephalo-

carida (in this class) (Milne Edwards, 1834;

Schram. 1986). However Dahl (1987: 722)

refuted this, slating that “polyramous thoracopods

constitute a basic feature of malcostracan mor¬

phology and are therefore not a phyllopod

synapomorphy.” Dahl (1987) also highlighted the

fact that while most genera of Leptostraca have

foliaceous thoracopods those of Paranebalia are

most similar to the stenopodous appendages of

caridoid Malacostraca (e.g. Euphausiacea).

Martin and Christiansen (1995) also detailed

many differences between the fourth thoracopod

of Nehalia (Leptostraca) and Leptestheria sp.

(Branchiopoda: Conchostraca) including the size

and arrangement of endites and the type and num¬

ber of setae and their function. They too believed

that the phyllopodous limb cannot be used as an

indicator of phylogenetic affinity. This view was

supported by independant evidence from an I8S

rDNA study of Branchiopoda, Ccphalocarida and

Phyllocarida, seven other crustacean taxa and

three arthropod outgroups (Spears and Abele,

1999). They concluded, with little doubt, that the

presence of foliacous limbs docs not define a

monophyletic cladc comprising branchiopods,

ccphalocarids and phyllocarids. They, like Dahl

(1987) and Martin and Christiansen (1995),

believed that foliaceous limbs have multiple

origins.

While Icptostracans appear to be the basal

malacostracans they differ significantly from all

other taxa in this class, making selection of an

outgroup difficult. One potential sister taxon —

the subclass Hoplocarida (Order Stomatopoda)

— are morphologically so highly derived that

386

G. K. WALKER-SMITH AND G. C. B. POORE

Table 1. Character Iransfonnalions used in phyogenctic analysis of 32 species of Lcptostraca. Each

character is terminated by a colon and states (0, 1 ...) separated by a semicolon. The 17 characters with

Cl=l in tree 711 are indicated by #, those where 0.5<CI<1 by *.

1*. Rostrum, sublemiinal spine: absent (Fig. la) (0); present (Fig. lb) (1).

2#. Rostral keel: absent {()); shorter than rostral flange (Figs la, d) (1); longer than rostral flange (Figs

Ic, e) (2).

3*. Eye length: shorter than rostrum (0); longer than rostrum (1).

4#. Eye, supraocular scale: absent (0); longer or equal to length of eye (1); shorter than eye (Fig. If)

( 2 ).

5*. Eye surface: smooth (0); denticulate (Figs Ih, m) (1).

6. Eye dorsal papilla: absent (0); present (Fig. 10(1)-

7*. Eye, ventral margin: not extremely cur\'ed (Fig. I f) (0); extremely curved (Fig. Ik) (1).

8. Eye, dorsal margin: not dorsally convex (Fig. 1 i) (0); dorsally convex (Fig. Ij) (1).

9#. Eye: not bilobed (Figs I f, g) (0); bilobed (Fig. ln)(l).

10. Eye: with ommatidia (Fig. Ig) (0); without ommatidia (Fig. li) (1).

11*. Antenna 1 antcrodcnticulatc fourth article: absent (0); present (Fig. Iq) (1).

12#. Antenna I article 4: without robust setae (0); with 1 or more robust setae (Fig. Ip) (1).

13*. Antenna 1 of male: not swollen or a callynophorc (0); a swollen callynophore (Figs Ic, Id) (1);

with dense field of acsthctascs but not swollen (Fig. 2f) (2).

14*. Antenna 2 peduncle articles 3 and 4: not fused (Fig. 2a) (0); fused (Fig. 2b) (1).

15#. Antenna 2 peduncle articles 3 and 4; without two large culicular outgrowths (0); with two large

cuticular outgrowths (Figs Ir, 5a. b) (1).

16#. Antenna 2 peduncle surface: without denticles or minute cuticular outgrowths (0); with minute

denticles or cuticular outgrowths (Figs 5a. b) (1).

17#. Antenna 2 of male; greatly elongate, reaching to the caudal furca (0); not greatly elongate, only

half length of specimen (1),

18*. Antenna 2 peduncle article 2, dorsal surface: without spine (0); with spine (Fig. 2b) (1).

19. Mandible, article 2 of palp: with more than 2 setae (0); with 2 setae (1); with 1 seta (2).

20. Mandible palp, relative lengths of articles 2 and 3: 2 longer than 3 (0); 2 equal to 3 (1); 2 shorter

than 3 (2).

21. Mandible, article 3 of palp: tapering distally (0): with parallel margins (1); expanded distally (2).

22. Mandible incisor teeth; 2 (Fig. 2h) (0); 1 (Fig. 2c) (1); absent (2).

23#. Molar accessory tooth/spinc: absent (0); present (Fig. 2i) (1).

24#. Molar large accessory process: absent (0); present (Fig. 2e) (1).

25#. Molar process, -setal brush: absent (0); present (Fig. 2e) (I).

26#. Maxilla 1 second endite: complex (Fig. 3a) (0); bilobed (Figs 3b, d) (1); elongate (Fig. 3g) (2);

simple (Fig. 3c) (3); reduced (4).

27. Maxilla 2 endopod: biarliculate (0); uniarliculate (1).

28*. Maxilla 2 exopod: greater than or equal to half length of endopod (0); less than half length of

endopod (I); absent (2).

29#. Thoracopod length: short, not extending well beyond the ventral margin of the carapace (Fig. 4a)

(0); long, extending well beyond ventral margin of carapace (Fig. II) (I).

30*. Thoracopod exopod: heavily setose (Figs 4b, 4e) (0); with few setae (Fig. 4d) (1); with no setae

(Fig. 4c) (2).

31#. Thoracopod exopod: without proximal lobe (Figs 4b'd) (0); with proximal lobe (Fig. 4e) (1).

32#. Thoracopod 2-5; epipod longer than exopod (Fig. 4d) (0); epipod shorter than exopod (Fig. 4b)

(1); epipod absent (Fig. 4e) (2).

33#. Pleonite 4 posterior margin: smooth (0); crenate (saw-tooth) (Fig. 4a) (1).

34#. Pleonite 5 posterior margin; smooth (0); crenate (Fig. 4a) (1).

35#. Crenations on pleoniles 6 and 7: absent (0); only on dorsal margin (Fig. 11) (1); over entire

margin (Fig. 4a) (2).

36. Pleonite crenation: absent (0); pointed (1); blunt (2).

37*. Pleonite size: pleonite 6 and pleonite 7, each equal in length to pleonite 5 (0); pleonite 6 and

pleonite 7, each longer than pleonite 5(1).

38. Plcopods 1-4 peduncles margins: smooth (0); crenate (Fig. 3i) (1).

PHYLOGENY AND KEYS TO LEPTOSTRACA (CRUSTACEA)

387

Table I. — continued.

39. PIcopod I, ratio of lengths of comb-row to exopod: comb-row absent (0); less than or equal to

half length of exopod (I); greater than half length of exopod (Fig. 4k) (2).

40*. Pleopod 2-4 cxopod lateral margin: with smooth setae not in pairs (Fig. 4i) (0); with smooth setae

in pairs (Fig. 4h) (I).

41 *. Pleopod 5: shorter than pleopod 6 (0); longer than pleopod 6(1).

42*. Pleopod 6: biarticulate (0); uniarticulate (1).

43*. Carapace with: posterodorsal marginal spines (Fig. 3e) (0); without posterodorsal marginal spines

( 1 ).^

Table 2. Character matrix used in phylogenetic analysis of the Leptostraca

Character numbers

1234567891

1234567892

1234567893

1234567894

123

Hypothetical ancestor

0000000000

0070000007

7077070070

0077770000

007

Nebaliopsis tvpica

0000000000

1070007000

0200041202

0000000000

101

Levinebalia forumata

1000000100

1011111020

7100021010

0100000001

Levinebalia maria

1000000100

1011111021

OI000210IO

0100000011

Paranebalia belizensis

1000100100

1011101020

0107120010

0177I17I21

Faranebalia hmgipes

1000100100

1011101000

0101120010

0100111121

Paranebalia sp. A

1000100100

1011101021

0101121010

0100111121

Speonebalia cannoni

0010000101

000000702?

7200031101

0011210101

000

Nebaliella antarctica

0210001101

0000000101

2010011000

1211211120

100

Nebaliella brevicarinata

0210001101

0000000121

0010010000

1211271170

100

Nebaliella caboti

0210001101

0000000117

7010011000

1211271720

100

Nebaliella declivatas

0210001101

0000000112

0010011000

12I12I10I0

100

Dahlella caldariensis

0012101001

0071007172

2000000101

0011271121

Nebalia antarctica

0102010100

0101000112

2000000001

0011217721

Nebalia bipes bipes

0102000100

0101000122

2000000001

0011227721

Nebalia borealis

0102000100

010I000I2!

1000000001

0011217721

Nebalia hritcei

0102000100

0101000110

2000000001

0011211021

Nebalia cannoni

0102010100

0101000102

2000000001

0011227721

Nebalia capensis

0102000100

0101000111

2000000001

0011221021

Nebalia clausi

0102000100

0101000121

2000000001

0011227721

Nebalia daytoni

OlOlOOOIlO

0121000011

2000000001

0011211721

Nebalia falklandensis

0102010100

oioiooono

2000000001

0011227721

Nebalia herbstii

0102000100

0101000122

2000000001

0011227721

Nebalia hessleri

0102000100

0101000121

2000000001

0011211121

Nebalia lagartensis

0107000100

01()100()1?0

1000000001

0011217721

Nebalia longicornis

0702010100

0101000112

I000000001

0011227721

Nebalia marerubri

0102000100

0101000112

1000000001

0011211121

Nebalia patagonica

0702010100

0I01000I12

2000000001

0011227721

Nebalia strausi

0102000100

010100072?

0000000001

001I21772I

Nebalia sp. A

0101000110

0121000012

1000000001

0011211001

Nebalia sp. B

1102000000

0121000112

2000000001

0011211001

Nebalia sp. C

0102010100

0101000122

2000000001

0011211121

Sarsinebalia typhlops

1102000001

0101000712

0000000101

007722770I

sufficient characters relevant to generic differen¬

tiation in Leptostraca do not exist. Similarly,

resorting to the fossil orders of Phyllocarida (e.g.

Archaeostraca) provide few characters of value.

Olesen's (1999) phylogenetic anaylsis of seven

leptostracan genera resulted in two hypotheses. In

one, he used Anoslraca (Branchiopoda) and

Mysidacea (Malacostraca) as outgroups without

388

G. K. WALKER-SMITH AND G. C. B. POORE

justifying these choices. In the other, he used

Mysidacea alone. Specific outgroups such as

these pose real problems. The presumed shared

similarities may not be homologous so it is doubt¬

ful whether the same characters are being scored

for the in- and outgroups. Besides, they often do

not possess relevant characters.

Olesen’s (1999) use of Mysidacea as an out¬

group was based on Cannon’s (1927) view that

the thoracopods of Panmebalia link malacostra-

cans, such as mysids, with Nebalia. Numerous

authors have viewed phyllocarids as malacostra-

cans (27 papers cited by Spears and Abele, 1999).

Mysidaceans themselves arc a problematic group

of two distinct clades. Although mysidaceans

have until recently been treated as members of

Peracarida there is now increasing morphological

and molecular evidence that while one clade,

Lophogastrida, is a member of Peracarida the

other, Mysida, is a member of Eucarida (Watling,

1999; Jarman el al., 2000). Olesen did not

difTcrcnliate the two.

Curiously, Sars (1887) suggested that body

divisions, antennules, antennae, moulhparts,

pleopods, caudal limbs, and development of

Nebalia and Copepoda, especially Harpacticoida

Sars, 1903 were homologues. Sars’s similarities

could be further evidence of the high level of

convergence within the Crustacea or symplc-

siomorphies shared by these and possibly other

groups.

Faced with the conditions that the closest rela¬

tives of leptostracans do not have similar mor¬

phologies, and that similarities between leptostra¬

cans and other less related taxa arc most likely

due to convergences, we were disinclined to

chose any one or set of outgroups. The best alter¬

native for polarising character states seemed to be

to use a hypothetical ancestor. We used general

principles of cmstacean evolution as evidenced

by a wide variety of taxa and were able to score

31 of 43 characters for the hypothetical ancestor.

These included characters where the presence of a

structure is confined to some leptostracans, char¬

acters involving fusion or loss of articles from a

multiarticulate slate, characters involving loss of

teeth or setae which are generally numerous in the

other Crustacea, and characters involving

reduction in size or complexity.

Character descriptions

The 43 characters are examined in turn with its

reasoned state of the hypothetical ancestor. All

characters are unordered and of equal weight

(Tables 1, 2). Character descriptions and figures

are for females except where male characters are

individually specified.

Although Dahl (1985) defined many new char¬

acters for Nebalia, most of these have not been

used. Dahl’s (1985) ratio characters (e.g.

lenglh/width characters) could only be scored

from literature descriptions, generally of a single

specimen. The ratios varied continously across all

taxa and it was not possible to assign taxa to a few

distinct classes.

Setal characters. Dahl (1985) slated that charac¬

ters related to numbers of spines (= robust setae)

and setae arc not of primary importance as they

arc related to growth of the individual. He used

rearing experiments to show growth related vari¬

ation in the moults of six females and five males

of Nebalia pugettensis (Clark, 1932). While his

experiments show a correlation between the cara¬

pace length and the number of spines and setae on

four appendages (antenna I, pleopod 1 exopod,

pleopod 5 and furca) (Dahl, 1985: Table 1), we

believe that for phylogenetic purposes setae may

be useful if comparisons were made between

ovigerous or brooding females but none is used

here.

Rostrum. Dahl (1985) recognised the presence or

absence of a ventral subterminal rostral spine

(character 1) as taxonomically informative. We

hypothesise the possesion of a rostral spine is

apomorphic as it appears to be a character unique

to the Leptostraca. All species of Paranehalia and

Levinebalia and Sarsinehalia typhlops (Dahl,

1985) and Nebalia sp. B possess a rostral spine

(stale 1: Figs lb, d).

The presence of a keel on the ventral face of the

rostrum (the rostrum minus the keel is sometimes

referred to as the rostral fiange; Fig. Ic arrow

pointing to fiange) is a character unique to Icp-

toslracans (character 2) and so is considered apo¬

morphic. The keel is absent in Nebaliopsis,

Paranehalia, Levinebalia, Speonebalia and

Dahlella (state 0). Dahl (1985,1990) did not draw

the rostral keel for any of the species he described

but as all other species of Nebalia have a keel, we

assume this was an oversight. We have scored all

Nebalia as having a keel shorter than the rostrum

(state 1: Figs la. Id). The possession of a keel

longer than the rostrum is an aulapomorphy of the

genus Nehaliella (state 2: Figs Ic, c).

Eye. Speonebalia, Nehaliella and Dahlella have

eyes longer than the rostrum (character 3, state 1:

Fig. le).

The length of the supraocular scale was con¬

sidered diagnostic by Dahl (1985) (character 4).

Nebaliopsis, Paranebalia, Levinebalia, Speone¬

balia and Nebaliella all lack supraocular scales

(state 0). Dahlella and almost all species of

PHYLOGENY AND KEYS TO LEPTOSTRACA (CRUSTACEA)

389

Nehalia have a supraocular scale shorter than the

eye (state 2: Figs If, i). Nehalia daytoni Vetter,

1996 and Nehalia sp. A have supraocular scales

longer than the eye (state 1: Fig. In). Following

the ontogenetic precedence criterion, the absence

of the supraocular scale in juveniles of Dahlella

caldariensis Messier, 1984 suggests its presence

is apomorphic.

Small teeth or denticles over the surEice of the

eye appear in Paranehalia (Fig. Ih) and Dahlella

(character 5, state I: Fig. Ini). Messier (1984)

suggested the teeth may be used by Dahlella

caldariensis to scrape for food such as bacterial

encrustations. The teeth on the surface of the eye

may not be homologous in Dahlella and

Paranehalia. As the teeth do not appear in the

Juveniles of Dahlella they are thought to be

apomorphic.

The possession of a papilla or dorsal outgrowth

on the eyestalk is a feature found in approxi¬

mately one-third of species of Nehalia (character

6, state I; Figs If, Ij) and is thought to be

apomorphic.

Dahl (1985) recognised eye shape as a valuable

diagnostic feature (characters 7, 8 and 9). The

eyes of Nehaliella and Dahlella have an

extremely curved ventral margin (character 7,

state 1: Figs Ik, Im). The eye of the first instar

larva of Dahlella is almost square, thus the onto¬

genetic evidence suggests the elongate, curved

eye of Dahlella and Nehaliella is the derived

state.

The eyes of Sarsinebalia typhlops, Nehalia sp.

B, Dahlella and Nehaliopsis are not dorsal ly con¬

vex like those of all other Lcptostraca (character

8, state 0). The eyes of Sarsinebalia typhlops are

almost square (Fig. li) (but have also been

described as almost circular [Dahl, 1985]), and

the eyes of Nehalia sp. B are triangular (Fig. Ig).

The eyes of Dahlella are dorsally angular (Fig.

l m) , w'hile the eyes of Nehaliopsis are square to

rectangular. The eyes of Nehaliopsis are very

similar in shape to that of the first inslar larva of

Dahlella. Nehalia daytoni and Nehalia sp. A have

an unusual bilobed eye (character 9, state 1: Fig.

l n) shared with no other species.

The presence of ommatidia in the eye is com¬

mon to most species of Lcptostraca (character 10,

state 0). Speonehalia. Nehaliella. Dahlella and

Sarsinebalia typhlops all lack ommatidia (state

I). Embryos of the genus Nehalia possess dark

eye pigment (Sars, 1896; Manton, 1934) thus, fol¬

lowing the ontogenetic precedence criterion, the

presence of ommatidia is considered primitive.

Antenna /. Modlin (1991) referred to the antero-

denliculate fourth article of antenna 1 as a lateral

flange but our observations reveal the flange lies

mesially (character II, state 1: Fig. Iq). This

mesiodislal flange is found in Levinehalia,

Paranehalia and Nehaliopsis (state 1). The flange

is lacking in all other Lcptostraca and assumed so

for the hypothetical ancestor (state 0; Fig. Ip).

The first four articles arc referred to as the

peduncle. The last peduncle article (article 4)

bears the scale and flagellum The arrangement of

setae on the fourth peduncle article is usually

linear with 1-4 simple robust setae in the

anterodistal comer and a variable number of thin

plumose setae (Fig. Ip). In some species and the

hypothetical ancestor robust setae are absent

(character 12, stale 0; Fig. Iq). Rearing experi¬

ments (Dahl, 1985) showed that change in setal

fonnula is growth related but the presence or

absence of robust setae, not their number, is a

valid character. Only Nehalia has robust setae on

antenna 1 (state I).

Mature males of Levinehalia and Paranehalia

possess numerous aesthetascs on a swollen flagel¬

lum of the first antenna (character 13, state I:Fig.

2d), this chemoreceptive callynophorc is found on

many eucarid and pcracarid Crustacea (Lowry,

1986). Immature males of Levinehalia and

Paranehalia have a swollen flagellum with few

aesthetascs (Fig. 2c). Abundant aesthetascs on a

non-swollen flagellum (state 2: Fig. 20 occur in

Nehalia daytonL Nehalia spp. A and B. This is

similar to that found in Nehaliapugettensis (orig¬

inally described as Epinebalia pugettensis) but

this species was excluded from the analysis.

Because the callynophore is so widespread in

Cmstacea we were unable to score the hypotheti¬

cal ancestor; the structure may be independently

derived in many taxa.

Antenna 2. The peduncle of antenna 2 bears the

flagellum. The peduncle has a maximum of four

articles but fusion of articles does occur. The

fusion of articles 3 and 4 peduncle is an apomor¬

phic state found in Nehalia, Paranehalia,

Levinehalia and Dahlella (character 14, state I:

Fig. 2b). In Nehaliop.sis. Nehaliella and Speone-

balia the articles are not fused (stale 0: Fig. 2a).

Fused articles 3 and 4 of antenna 2 peduncle of

Paranehalia pos.sess protuberances or elongate

outgrowtiis (usually one or two) on the anterior

surface (character 15, state I: Figs Ir, 5a and 5b;

arrows point to protuberances).

Minute denticles or cuticular outgrowths

appear over the surface of fused peduncle articles

3 and 4 and the flagellum in Levinehalia (charac¬

ter 16. state 1: Fig. 5a). The flagellum of male

Nehalia and Nehaliella is greatly elongated, often

extending past the caudal furca (character 17,

390

G. K. WALKER-SMITH AND G. C. B. POORE

state 0). The flagellum length of male Speone-

balia, Dahlella ciMariensis and Nebaliopsis is

unknown as mature males have not been identi¬

fied. The length of the flagellum does not differ

between males and females in species of

Paranehalia and Levinebalia. The presence of a

dorsal spine on article 2 (character 18: Fig. 2b) is

recorded for most species of Nebalia and

Nebaliella.Thc hypothetical ancestor is assumed

to have a simple peduncle of 4 articles, without

cuticular outgrowths or a dorsal spine.

Mandible. The number of setae on article 2 of the

mandibular palp (character 19: Fig. 2g) can be

diagnostic (Hessler, 1984). Ilow'cvcr, Dahl (1985)

suggested that there is a growth-related increase

in the number of spines (robust setae) and setae

throughout the Leptostraca. Therefore, although

setal characters may be useftil for supplementing

other morphological features, leptostracan

‘"chaetotaxonomy" can never be of primary

importance. Levinebalia. most species of

Paranebalia, Speonebalia, Dahlella, Nebaliella

brevicarinata Kikuchi and Gamo, 1992 and half

of the species of Nebalia have one seta on article

2 of the mandible palp. Half of the species of

Nebalia have two setae as does Nebaliella caboti

Clark, 1932 and N. declivatas Walker-Smith,

1998. Paranehalia longipes (Willcmoes-Suhm.

1875) has more than two setae, as does Nebaliella

antarctica Thiele. 1904 and Nebalia cannoni

Dahl, 1990.

The length of article 2 of mandibular palp

relative to article 3 was recognised by Dahl

(1985) as a diagnositic character (character 20).

The piesiomorphic state is unknown.

The shape of article 3 of the mandibular palp

was considered by Dahl (1985) to be a diagnostic

feature of Icptostracans (character 21: Fig. 2g).

This is a variable character within Nebalia. which

displays all three states. Nebaliopsis. Levinebalia.

Paranehalia, Speonebalia and most species of

Nebaliella have palps that taper dislally (state 0:

e.g.. Fig. 2g). The palp of Nebaliella antarctica

and Dahlella is expanded distally (state 2).

The mandible incisor of Nebaliella has two

teeth (character 22, state 0; Fig. 2i). Dahl (1985,

1990) did not draw or mention the mandible

incisor in his descriptions of Nebalia. However,

as all other described species and the undcscribcd

species of Nebalia from Australia have tw'o teeth.

We have scored all species described by Dahl as

having two teeth (state 0: Fig, 2h). Levinebalia,

Paranehalia and Dahlella have one tooth (state I:

Fig. 2c). The incisor is absent in Speonebalia and

Nebaliopsis (state 2).

The molar of Nebaliella has an accessory

tooth or spine (character 23, state 1: Fig. 2i; see

arrow). No other leptostracan has this character

state. The molar of Paranehalia alone has a large

accessory process (character 24, slate 1: Fig. 2e;

see arrow). The presence of a sctal brush on the

molar process (character 25, state 1; Fig. 2e) is

recorded only for Paranehalia. The hypo¬

thetical ancestor is assumed to have a setose

mandibular palp, well-developed molar and

toothed incisor.

Maxilla 1. In all species of Nebaliidac, including

Paranebalia and Levinebalia. the palp of maxilla

1 is long and well-developed as in the hypo¬

thetical ancestor. In Ncbaliopsididac it is reduced

to a small stub.

There arc four different types of second endites

found on maxilla I (character 26); complex (state

0: Fig. 3a) found in Nebalia and Dahlella: bilobed

(state 1: Figs 3b, d) in Nebaliella; elongate (state

2: Fig. 3g) in Paranehalia and Levinebalia; sim¬

ple (state 3: Fig. 3c) only in Speonebalia. The sec¬

ond endite is reduced in Nebaliopsis (stale 4). The

slate in the hypothetical ancestor could not be

determined.

Maxilla 2. The maxilla 2 endopod is uniarticulate

(character 27; state 1) in Nebaliopsis. Nebaliella

(except N. brevicarinata). Paranehalia sp. A,

Levinebalia and Speonebalia. This is thought to

represent the derived state.

The length of the exopod of maxilla 2 relative

to the endopod is infonnativc (character 28). For

Speonebalia. Dahlella and Sarsinehalia typhlops

the maxilla 2 exopod is less than half the length of

the endopod (stale I). Nebaliopsis docs not have

an exopod (state 2). All other genera have an exo¬

pod more than half the length of the endopod

(state 0).

Thoracopods. The length of the thoracopods is

the most obvious feature diagnosing genera of

Icptostracans (character 29). All genera except

Paranehalia and Levinebalia have foliaceous

thoracopods that do not extend well beyond the

ventral margin of the carapace (slate 0: Fig. 4a).

The thoracopods of Paranebalia and Levinebalia

extend well beyond the ventral margin of the

carapace (state 1: Fig. 11).

Thoracopod exopods are densely setose (char¬

acter 30) in Paranehalia. Levinebalia and

Nebaliella (slate 0: Figs 4b, 4e). The exopods

have few setae in Nebalia (state I: Fig. 4d),

Dahlella and Speonebalia and no setae in

Nebaliopsis (state 2; Fig. 4c).

Thoracopod exopods of Nebaliella have a

PHYLOGENY AND KEYS TO LEPTOSTRACA (CRUSTACEA)

391

proximal lobe (character 31, state 1: Fig. 4c) not

seen in other Leptostraca.

All genera except Nebaliella possess thora-

copodal cpipods (character 32, state 2) whose

length relative to that of the exopod is informa¬

tive. Paranehalia and Levinebalia have relatively

small epipods (stale I; Fig. 4b) compared to those

of Nebadopsis. Speonebalki. Dahlella and

Nebalia which are longer than the thoracopodal

exopod (state 0: e.g. Fig. 4d).

Pleonites. All species Nebaliella. Speonebalia,

Dahlella and Nebalia have a crenatc posterior

margin on pleonites 4 and 5 (characters 33 and

34; Fig. 4a). In his descriptions of Nebalia species

Dahl (1985, 1990) did not mention the form of

pleonites 4 and 5 but as these pleonites are crenate

in all oxhex Nebalia species we have assumed this

is also the case in DahFs species. The posterior

margins of ail pleonites of Nebadopsis are

smooth.

Pleonites 6 and 7 of Paranehalia are crenatc

only along the dorsal margin (character 35, slate

I; Fig II). The pleonite margins of Nebadopsis

and Levinebalia are smooth (state 0). Pleonites 6

and 7 of Nebaliella. Speonenbalia. Dahlella and

Nebalia are crenatc along the entire margin (state

2 )-

The shape of pleonite crenations is a useful

species-level character (character 36). Species of

Nebalia may have cither crenations that arc

pointed or blunt. Nebalia sp. A has both blunt and

pointed crenations along the same pleonite

margin (Fig. 30* but as this is an autapomorphy

for the purpose of the analysis it has been scored

as having only pointed crenations.

For most Leptostraca pleonites 6 and 7 are

much longer than pleonite 5 (character 37, state

1). However for Levinebalia. Speonebalia and

Nebadopsis pleonite 5 is approximately the same

length as pleonites 6 and 7 (state 0).

Pleopods. The posterior margin of picopods 1-4

of Paranehalia. Speonebalia and some species of

Nebalia are crenate (character 38, state 1: Fig. 3i).

The comb-row or '‘spine-row", considered

by Dahl (1985: pp. 142, 163) to be a generic

character, consists of a row of short, pinnate setae

along the exterior margin of the exopod of pleo-

pod I (Figs 4k, 6a and 6b). As Dahl (1985)

created a new genus for Sarsinebalia typhlops

which does not possess a comb-row on its llrsl

pleopod. The length of the comb-row relative to

the exopod is diagnostic (character 39). For all

species of Nebalia except Nebalia spp. A and B

the comb-row is greater than half the length of the

exopod (state 2). Nebalia spp. A and B do not

possess a comb-row (slate 0: Fig. 4i). The comb-

row of Levinebalia Walker-Smith, 2000 and

Nebaliella declivatas is less than half the length

of the exopod (state 1: Fig. 4j).

All genera except Nebaliella and Nebadopsis

have pairs of smooth setae along the exterior

margin of the exopod of pleopods 2-4 (character

40, state 0 (not in pairs): Fig. 4g. state 1: Figs

3i, 4h).

For all genera except Nebaliella and Nehadop-

sis the ramus of pleopod 5 is longer than the

ramus of pleopod 6, measured along the midline

(character 41, stale 0). We have scored pleopod 5

longer than 6 as the picsiomorphic state

Pleopod 6 may be uni- or biarticulate, a char¬

acter first used by Olesen (1999). Nebadopsis,

Speonebalia xind Nebaliella all have a biarticulate

pleopod 6 (character 42, state 0; Fig. 41). All other

Leptostraca have a uniarticulate pleopod 6 (state

1). The biarticulate condition is thought to be

plesionioiphic.

Carapace. The posterodorsal margin of the cara¬

pace of Nebaliella has small denticles (character

43, state 0: Fig. 3c). This character state is not

seen in other Leptostraca and the plesiomorphic

condition is unknown.

Results

Cladogranis

The phylogenetic program PAUP* 4.0 revealed

1527 equally parsimonious trees of 114 steps.

Tree 711 and tree 340 were the two most distant

parsimony trees (found using the FILTER com¬

mand). Their statistics are: consistency index (Cl)

= 0.52; homoplasy index (HI) = 0.48; retention

index (RI) = 0.79; rescaled consistency index

(RC) = 0.41.

Bremer support values were calculated for tree

711 (Fig. 6) and tree 340. Branch lengths for tree

711 were calculated in PAUP* 4.0 and are pre¬

sented diagramatically (Fig. 7). Characters with

Cl=l are also plotted on this tree.

A 50% majority-rule with bootstrap values and

the percentage of parsimony trees retaining

nominal clades is also presented (Fig. 8).

Characters defining the clades of parsimony

tree 71J.

As trees 711 and 340 retain the same character

state changes at the major (generic level) nodes,

only tree 711 is discussed in detail (Figs. 6 and 7)

with character stale changes (Table 3 ).

Clade 63 contains all Recent Leptostraca and is

supported by three synapomorphies from the

characters used (plus those characters generally

stated to define the taxon).

392

G. K. WALKER-SMITH AND G. C. B. POORE

Table 3. Character transformations at all nodes in tree 711 (one of 1527 parsimonious trees). Character

numbers follow each clade labelled in Fig. 6. Character numbers alone indicate a change from state 0

to state I, - indicates a reversal from state 1 to 0, and superscripts indicate a change from one state

(default 0) to another. Characters in bold have Cl=l.

Clade number or taxon

Characters changing

Clade 63

11,22\ 27

Nebaliopsis typica

26\ 28% 302,41

clade 62

8, 19% 36, 38,40

clade 37 (Paranebaliidae)

1, 13, 14, 15, 17, 22^', 26S 29, 32, 42

clade 34 (Levinebalia)

16, -36, -38

Levinebalia maria

20, 39

clade 36 (Paranebalia)

5, 24, 25, 35, 37, 39*

clade 35

-27

Paranebalia longipes

19-’"

Paranebalia sp. A

clade 61 (Nebaliidae)

3, 10,-ll,20^ 30, 33, 34, 35^ ^3

Speonebalia cannoni

26\ 28

clade 60

7, 18, 22, 37, 39-

clade 40 (Nehaliella)

1\ 23, 26, 30, 31,32^^0, 41

clade 38

20^'’

Nehaliella antarctica

192'",2H

Nehaliella brevicarinata

-27

clade 39

19'”,-38

Nehaliella declivatas

39'”

clade 59

4^ 14,2^,-27,42,43

Dahlella calderiensis

5, -8, 28

clade 58 {Nebalia)

2,-3,-7,-10, 12

clade 51

19'”

clade 43

Nebalia sp, C

191>2

clade 41

36'^'

Nebalia cannoni

-19

Nebalia longicornis

2pi

Clade 49

-38

clade 45

20'X)

clade 44

36‘^'

Nebalia capensis

Nebalia falklandensis

clade 48

13', 39'”*

clade 46

4'”, 9,-18

Nebalia dayfoni

20'”, 39'

Nebalia sp. A

21'”

clade 47

1,-8

Sarsinebalia typhlops

10, 13'”>,21'”>, 28, 36’^'

Nebalia marerubri

21'”

clade 57

36'^'

clade 56

20'”

clade 55

36'”

clade 54

21'^

clade 53

Nebalia lagartensis

-20

PHYLOGENY AND KEYS TO LEPTOSTRACA (CRUSTACEA)

393

Nehaliopsis ty’pica Sars, 1887 (Nebaliopsidi-

dae) is defined by four apomorphics (from the

characters used): maxilla 1 second endite

reduced; maxilla 2 exopod absent; thoracopod

exopod with no setae; pleopod 6 shorter than

picopod 5. At least 14 more character stales

define NebuHopsis typica but these unique states

arc uninformative and had been excluded a priori:

molar process reduced; maxilla I palp reduced to

a small stub (but may terminate in long seta);

maxilla 1, second endite reduced; maxilla 2 with

endites 2-4 reduced in size and setation; maxilla

2 nearly as large as thoracopod 1; thoracopod I

differing greatly from thoracopods 2-7, some¬

what maxillipediform; thoracopod endopod not

articulate; thoracopods well spaced; pleopods 24

exopod paddle-like, outer margin strongly curved

with numerous small spinules; carapace not

emarginate; carapace with network pattern of

sculpturing; body cuticle and carapace thin, mem¬

branous; caudal furca leaf-Iike, broadest midway;

entire length of mature female greater than 20

mm.

Clade 62 contains all Lcptostraca except

Nebaliopsididac. This clade occurs in all shortest

trees and has a Bremer support value of 2 and

72% bootstrap support. Five synapomorphies

define this clade although none has CI=1.

Clade 37, Paranehalia plus Levinebalia, occurs

in all trees and has a Bremer support value of 3

and 85% bootstrap support. The clade is defined

by ten synapomorphies, five with CI=1: antenna 2

articles 3 and 4 with two large cuticular out¬

growths; antenna 2 of male not greatly elongate,

only half length of specimen; maxilla I second

endite elongate; thoracopods long, extending well

beyond the ventral margin of the carapace; thora¬

copod 2-5 with epipod shorter than exopod. This

clade is also defined by the character stale —

males with swollen callynophorc — but as this

character is multistate it has CI=0.67.

Clade 34 (Levinehalia)^ evident in 100% of

trees, has Bremer support of 1 and 77% bootstrap

support. It is defined by three synapomorphies:

antenna 2 peduncle and flagellum surface with

minute denticles or cuticular outgrowths (CI=1);

pleonitc margins smooth (reversal); pleopods 14

peduncles with margins smooth (reversal).

Clade 36 (Paranebalia)^ evident in 100% of

shortest trees, has Bremer support of 5 and and

99% bootstrap support. Six synapomorphies

including three with CI=I define this clade: molar

with large accessory process; molar process with

setal brush; and pleonites 6 and 7 with denticles

only over dorsal part of margin.

Clade 61 occurs in all trees and has Bremer

support of 2 and 67% bootstrap support. This

clade, all species of Speonebalia. Nebaliella,

Dahlella and Nehalia, is defined by nine synapo-

mophies including three with CI=1. Some of the

characters defining the clade are; antenna 1 with¬

out anterodenticulate fourth article (CI=0.5);

thoracopod exopod not heavily setose (CI=0.67)

pleonite 4 margin denticulate (CI=I); pleonite 5

margin denticulate (CI=1); with crenations over

entire pleonite margin (CI=1).

Speonebalia cannoni Bowman, Yager and

Iliffe, 1985 is defined by two apomorphies, one

with CT=I; maxilla 1 second endite simple.

Speonebalia cannoni also has an autapomorphy

that was excluded from the analysis; maxilla 2

with marginal organelles.

Clade 60 (Nebaliella. Dahlella and Nehalia)

occurs in all trees and has a Bremer support value

of 1 and bootstrap support <50%. It is defined by

five synapomorphies. None is unique.

Clade 40 (Nebaliella) is supported in all trees

with Bremer support value of 6 and 100% boot¬

strap support. It is defined by eight synapomor¬

phies, five with CI=I: presence of a rostral keel

longer than the rostral flange; molar with acces¬

sory tooth/spine; maxilla 1 second endite bilobed;

thoracopod exopod with proximal lobe; and tho¬

racopods 2-5 without epipod. Picopod 6 longer

than pleopod 5 and pleopod 6 uniarticulate are

also characters linking species in this clade.

Clades 38 and 39 relating the species of

Nebaliella occur in all shortest trees, each clade

has Bremer support of 1 and clade 39 has 50%

bootstrap support.

Clade 59 (Dahlella and Nebalia\ occurs in all

parsimony trees and has a Bremer support value

of 2 and 72% bootstrap .support. Six synapo¬

morphies define this clade but the presence of a

supraocular scale is the only unique character

(CI=I). All species in this clade also share:

antenna 2 peduncle articles 3 and 4 not fused

(CI=0.5); pleopod 6 shorter than pleopod 5

(CI=0.5) and pleopod 6 biarticulate (Cl=0.5)

Dahlella has three apomorphies among the

characters in this matrix but none is unique.

Clade 58 (Nebalia) was evident in all trees with

Bremer support of 1 and <50% bootstrap support.

It is defined by five synapomorphies. The pres¬

ence of a rostral keel shorter than the rostral

fiange and article 4 of antenna I with robust setae

are characters unique to Nehalia. Species in this

clade also have eyes shorter than the rostnim and

eyes with the ventral margin not extremely

curved.

394

G. K. WALKER-SMITH AND G. C. B. POORE

Only four clades, grouping three pairs and one

group of four species of Nehalia appear in the

50% majority-rule tree (Fig. 8). Clade 46 {N. day-

tom and Nehalia sp. A) was retained in all trees

and has a Bremer support of 2 and 69% bootstrap

support. Clade 47 (5. typhlops and Nehalia sp. B),

retained in all trees, has a Bremer support value of

1 and <50% bootstrap suppoii. Clade 52 (M hipes

and N. herhstii) was retained in 62% of parsi¬

mony trees but has no Bremer support and <50%

bootstrap support. The clade linking Nehalia can-

noni, Dahl, \990, N. falklandensis\ Dahl, 1990, *V.

longicornis and N. paragonica Dalil, 1990 did not

occur in tree 711 but occurred in 75% of all trees.

It has no Bremer support and <50% bootstrap

support. The relationships of the remaining

species of Nehalia could not be resolved.

Systematics and a new classificanon

Hour synapomorphies used in this analysis and at

least 14 other character states define Nehaliopsis

typica (Nebaliopsididae) and differentiate it from

all other Leplostraca. The sister group (clade 62)

is described by robust synapomorphies so there is

support for the existing family Nebaliopsididae.

All shortest trees contain a clade (clade 37),

Paranehalia plus Levinehalia^ sister taxon of all

other species. We believe that with a Bremer sup¬

port value of 3, bootstrap value of 85% and five

aulapomorphies for this clade, a new family can

be Justified for the two genera.

Clade 61. apparent in all trees, contains the

remaining genera of Nebaliidae {Speonehalia.

Dahlella. Nehaliella, and Nehalia). This clade is

supported by nine synapomorphies, three aulapo-

morphic for the clade (see above) and has Bremer

support of 2 and 67% bootstrap support. This

clade defines the restricted family, Nebaliidae.

The monophyly of Speonehalia. Nehaliella.

Dahlella, and Nehalia is supported by the analy¬

sis. Speonehalia and Dahlella are monotypic and

their status as genera is confimied by the synapo¬

morphies of their sister taxa.

Two aulapomorphies for clade 58 unite all

species of Nehalia (including Sarsinehalia

typhlops): the presence of a rostral keel shorter

than the rostrum and the presence of robust setae

in the fourth article of antenna I.

We were unable to find any characters which

suppon separate generic status for the monotypic

Sarsinehalia and the genus must be synomised

with Nehalia and its species, S. typhlops returns to

its original combination.

Table 4 lists all described species with their

distribution.

Comparison with Olesen’s (1999) trees

Olesen (1999) presented two equally parsi¬

monious hypotheses of the phylogeny of the

genera of Leplostraca. He used 27 mostly binary

characters. The four monotypic genera are clearly

monophyletic and he was convinced a priori of

the monophyly of Paranehalia and Nehaliella.

He entertained the possibility that Nehalia might

be paraphyletic with respect to Sarsinehalia or

Dahlella (or both). Our hypothesis differs from

his. Olesen’s first tree was rooted against two out¬

groups, Mysidacca and Anostraca, and placed

Nehaliopsis as a sister taxon to all other Lep-

loslraca, as in our tree. However, the position of

Nehaliella and Paranehalia was directly trans¬

posed compared to our tree and Sarsinehalia was

placed as a sister to Dahlella and Nehalia. This

3-laxon clade occuired in both of Olesen s trees

(the second tree having only Mysidacca as an out¬

group) and he suggested this indicated strong

support. However, two of the characters linking

Dahlella and Nehalia in OlcsciTs tree actually

vary within Nehalia and thus arc not useful (char¬

acter 9: antenna 2, spine on segment 2; character

14: mandible, shape of segment 3). The third

character, character 2 (the absence of a rostral

spine) occurs throughout the Leplostraca and was

a reversal. Our tree treats Dahlella as a sister

taxon to Nehalia and synonomiscs Sarsinehalia

with Nehalia.

Olesen’s second tree (with only Mysidacca as

an outgroup) suggested Paranehalia at the base of

the Leplostraca, with the remaining taxa split into

two cladcs. Speonehalia sits as a sister taxon to

Nehaliella and Nehaliopsis supported by a single

character (25: pleopod 6 biarticulate). A single

character unites Nehaliella and Nehaliop.us (char¬

acter 24: pleopod 6 longer than pleopod 5). These

characters are both useful characters but, in our

tree appear to have evolved twice. The second

clade in Olesen's tree was the Sarsinehalia-

Dahlella-Nehalia clade, supported by four char¬

acters, the three mentioned above and character

10 (antenna 2 with three articles), which occurred

twice in this tree but only once in Olesen’s other

tree.

Olesen’s (1999) trees lead him to question

the family status of Nebaliopsididae and the

monophyly of Nehalia. Our tree indicates the

validity of the Nebaliopsididae. and more signifi¬

cantly of Nebaliidae and a third family. Our tree

suggestes Nehalia is monophyletic, Dahlella is a

separate genus and Sarsinehalia is a synonym of

Nehalia.

PHYLOGENY AND KEYS TO LEPTOSTRACA (CRUSTACEA)

395

Table 4. Taxonomic list of all families, genera and species of Recent Leptostraca with reported distri¬

butions. * indicates species omitted from phylogenetic analysis.

Order Leptostraca Claus, 1880

NEBALIOPSIDIDAE Messier, 1984

Nehaliopsis Sars. 1887

N. tvpica Sars, 1887. West and south-east coast of South America, near Falkland Is, off coast of Ghana,

Ivory Coast, south-west Indian Ocean, South Pacific, Scotia Sea

PARANEBALIIDAE fam. nov.

Paranebalia 1880

P, belizensis Modlin, 1991. Belize

P. longipes (Willemoes-Suhm, 1875). Bermuda, Virgin Is, southern Florida (USA), Japan, Gulf of

Siam, Torres Strait (Australia)

P. sp. A. South Australia (Australia)

Levinehalia Walker-Smith, 2000

L Jbrtimata (Wakabara, 1976). Otago Peninsula (New Zealand)

L maria Walker-Smith, 2000. Tasman Sea, off E coast of Tasmania (Australia)

NEBALIIDAE Samoucllc, 1819

Nehalia Leach, 1814

N. antarctica Dahl. 1990. Wilhelm II Land, Adelie Land (Antarctica)

*A. bipes abyssicola Fagc, 1929. Monaco

N. bipes bipes (Fabricius, 1780). Greenland, Arctic North America, Svalbard to western Norway

*vV. bipes valida Thiele, 1904. Pribilof Is (Bering Sea)

N. borealis Dahl, 1985. Norway, Sweden, British Isles, Shetland Is, Sleat Sound (Scotland)

yV. hrucei Olesen, 1999. Unguja I., Zanzibar (Tanzania)

N. canmmi Dahl, 1990. South Georgia

N. capensis Barnard, 1914. South Africa

*jV. chilensis (Claus, 1888) nomen nudum. Chile

N. ckiusi Dahl, 1985. Adriatic Sea (Italy)

*A. daldi Kazmi and Tinnizi, 1989. Karachi (Pakistan)

N. daytoni Vetter, 1996. San Diego (.southern California, USA)

N. faiklandensis Dahl. 1990. Falkland Is

*N. gerkenae Haney and Martin, 2000. Monterey Bay, California (USA)

N. berbstii Leach, 1814. Shetland Is, western Bristish Isles, western France to Spanish border

N. hessleri Martin. Vetter and Cash-Clark, 1996. Southern California (USA)

*N. ilheoensis Kensley, 1976. South-western Africa

*N. japanensis (Claus, 1888). Japan

*N. lagartensis Escobar-Briones, 1995. Ria Largartos, Yucatan Peninsula (Mexico)

N. longicornis longicornis Thomson, 1879. South Island (New Zealand), New Britain (Papua New

Guinea), South Africa, Lifou (New Caledonia), Blanche Bay, Sandal Bay

*V. longicornis sororl\i\Q\e, 1904. Caribbean Sea, Cuba

N. marerubri Wiigele, 1983. Red Sea

A. patagonica Dahl, 1990. Magellan region

*A. pugetfensis {C\^vk, 1932). Friday Harbour (Washington, USA)

N. strausi Risso, 1826. Channel Is, Guenisey, France, Monaco, Italy including Sicily

A. typhlops Sars, 1870. Red Sea, Lofoten Is (Norway), Messina, Bay of Naples (Italy), North America

from Davis Strait to New Jersey, Australia

A. sp. A. Eastern Bass Strait (Australia)

A. sp. B. Tasmania and eastern Bass Strait (Australia)

A. sp. C. southern Western Australia (Australia )

396

G. K. WALKER-SMITH AND G. C. B. POORE

Table 4. continued

Nebaliella Thiele, 1904

K antarctica Thiele, 1904. Kerguelen I., Akaroa Harbour (New Zealand)

N. brevicarinata Kikuchi and Gamo, 1992. Princess Ragnhild Coast (Antarctica), bathyal

N. caboti Clark, 1932. Cabot Strait (between Newfoundland and Cape Breton L), New Jersey (USA),

Rockall Trough

N. declivatas Walker-Smith, 1998. E coast of Victoria, New South Wales, Tasmania (Australia)

*N. extrema Thiele, 1905. Kaiser Wilhelm II Land, Palmer Archipelago (Antarctica)

Dahlella Hcssler, 1984

D. caldariensis Hessler, 1984. Galapagos I., hydrothermal vents

Speonebalia Bowman, Yager and Iliffe, 1985

S. cannoni Bowman, Yager and lliffe, 1985. Turks and Caicos Is, marine caves

Key to families of Leptostraca

1. Maxilla 2 with endites 2-4 reduced in size and setation (Fig. 2j); thoracopods

well spaced (Fig. 3h); pleopods 2-4 exopod paddle-like, outer margin

strongly curved, with numerous small spinules (Fig. 4g); caudal furca Icaf-

likc, broadest midway (Fig. 4n)...Nebaliopsididae (monotypic)

— Maxilla 2 with at least first 3 endites well developed (Fig. 21); thoracopods

closely space (overlapping); pleopods 2-4 exopod slightly expanded mid¬

way and/or distally or outer margin parallel (Figs 4b, d, e); caudal furca

tapering evenly to lip (Fig. 4m)...2

2. Antenna I of mature male with swollen callynophore (Figs 2d, e); thora¬

copods long, extending well beyond the ventral margin of carapace (Fig. 11);

thoracopods 2-5 epipod shorter than exopod (Fig. 4b).Paranebaliidae

— Antenna 1 of mature male not swollen or with callynophore, may have a

dense field of aesthetascs (Fig. 2f); thoracopods short, not extending well

beyond ventral margin of carapace (Fig. 4a); thoracopod 2-5 epipod longer

than exopod or absent (Figs 4d, e).Nebaliidae

Nebaliopsididae Hessler

Nebaliopsidae Hessler, 1984: 656.

Type genus. Nebaliopsis Sars, 1887 (original des¬

ignation).

Diagnosis. Rostrum without spine or keel. Eye

shorter than rostnim and with visual elements;

without denticles; without dorsal papilla; ventral

margin not extremely convex. Supraocular scale

absent. Antenna 1 with anterodenticulate fourth

article; article 4 without robust setae. Antenna 2,

peduncle articles 3 and 4 not fused, without cutic-

ular outgrowths, or minute denticles; without

dorsal spine. Mandible without incisor process;

molar process reduced, with armature. Maxilla I

reduced to small stub. Maxilla 2 (Fig. 2j) nearly

as long as thoracopod I; endopod reduced to

small, blunt distal lobe, without organelles; exo¬

pod absent; proximal endite enormously enlarged,

well-armed with marginal setae; endites 2-4

reduced in size and setation. Thoracopods not

extending well beyond ventral margin of carpace;

well-spaced (Fig. 3h). Thoracopod I differenti¬

ated from thoracopod 2-8. Thoracopod (Fig. 4c)

endopod blunt, featureless lobe, not articulate;

exopod strongly reduced, poorly difTcrentiated,

without setae; epipod well developed, longer than

exopod, somewhat maxillipediform. Posterior

margins of pleonitcs smooth. PIcopod I exopod

without comb-row. Pleopods 2-4 exopods pad¬

dle-like (Fig. 4g), length less than 3 times width.

Pleopod 6 longer than pleopod 5 and biarticulate.

Caudal rami leaf-like, broadest midway (Fig. 4n).

Thorax inflated; body cuticle and carapace thin

and membranous. Carapace with a network

pattern of sculpturing; not emarginate; extending

furthest posteriad midsagillally, without carina on

anterolateral lower comer. Entire length of

mature female greater than 20 mm.

Composition. Nebaliopsis Sars, 1887.

PHYLOGENY AND KEYS TO LEPTOSTRACA (CRUSTACEA)

397

Nebaliopsis Sars

Nebaliopsis Sars, 1887: 21.

Diagnosis. With the characters of the family.

Remarks. This family, contains only the type

species N. typica Sars, 1887. Descriptions of N.

typica may be found in Thiele (1905), Cannon

(1931) and Linder (1943). Males have not been

reported.

Parancbaliidae fam. nov.

Type genus. Paranebalia 1880.

Diagnosis. Subterminal rostral spine present (Fig.

lb); keel absent. Eye shorter than rostrum; visual

elements present. Eye sometimes with denticles

(Fig. Ih); without dorsal papilla; ventral margin

not extremely convex. Supraocular scale ab.sent.

Antenna 1 with antcrodcnticulate fourth article

(Fig. Iq); article 4 without robust setae; male

flagellum modified, either swollen (juveniles) or

transformed into callynophore (Figs 2c, 2d).

Antenna 2 peduncle articles 3 and 4 fused; pedun¬

cle with 2 rounded cuticular outgrowths and

sometimes with minute cuticular denticles or

spines (Figs 5a, b); without dorsal spine. Antenna

2 of male not greatly elongate, only half body

length. Mandible incisor with I tooth (Fig. 2e).

Molar process well developed (Fig. 2e); with or

without seta! brush and sometimes with large

accessory process. Maxilla 1 palp (Fig. 2g), long,

well developed: second endite elongate. Maxilla 2

with at least first 3 endites well developed; much

smaller than Ihoracopod I; endopod without

organelles; exopod greater than half length of

endopod. Thoracopods long, extending beyond

ventral margin of the carapace; closely spaced

(Fig. II). Thoracopod I differing only slightly

from thoracopods 2-7. Thoracopod exopod heav¬

ily setose and without proximal lobe (Fig. 4b).

Thoracopods 2-5 epipod shorter than exopod;

endopods showing a degree of articulation.

Plconitcs 4 and 5 with smooth margins. Pleonites

6 and 7 dorsal margins sometimes crenate (Fig.

II). Pleopod 1 exopod comb-row present.

Pleopods 2-4 c.xopod with parallel margins; outer

margins with setae in pairs. Pleopod 5 longer than

pleopod 6. Pleopod 6 uniarticulale. Caudal rami

tapering evenly to tip. Thorax not inflated; body

cuticle and carapace firm. Carapace not sculp¬

tured; emarginate; without carina on anterolateral

lower comer. Entire length of female less than

20 mm.

Composition. Paranebalia Claus, 1880;

Levinebalia Walker-Smith, 2000.

Remarks. Six unique character states link the gen¬

era of Paranebaliidae: antenna 2, articles 3 and 4

with two large cuticular outgrowths; male antenna

1 flagellum with swollen callynophore; males

without greatly elongate antenna 2; maxilla 1 sec¬

ond endite elongate; slender thoracopods extend¬

ing well beyond the ventral margin of the cara¬

pace; thoracopods with reduced cpipods. The

subtenninal rostral spine is a synapomorphy of

the Paranebaliidae, shared with Sarsinebalia

typhlops and Nebalia sp. B. The presence of an

antcrodcnticulate fourth article on antenna I is

found in Paranebaliidae and Nebaliopsididae.

Key to genera of Paranebaliidae

1. Eye with denticles (Fig. Ih); mandible molar with sctal brush and large

accessory process (Fig. 2e); antenna 2, without minute denticles or spine

over ther surface of the peduncle and flagellum (Fig. 5b); pleopods 1-4,

peduncle margin crenate (Fig. 2i); pleonites 6 and 7 margin dorsally

denticulate (Fig. II). Paranebalia

— Eyes without denticles; mandible molar without setal bmsh and accessory

process; antenna 2, with minute denticles or spines over the surface of the

peduncle and flagellum (Fig. 5a); pleopods 1^, peduncle margin smooth;

pleonites 6 and 7 without clearly defined crenations. Levinebalia

Paranebalia Claus

Paranebalia Claus. 1880; 576.—Thiele, 1905;

] 4 _ 19 , 24-25.—Verrill. 1923: 206-207.—Wakabara,

1976: 297.

Type species. Nebalia longipes Willemoes-Suhm,

1875 (by monotypy).

Diagnosis. Eyes with denticles or cuticular out¬

growths (Fig. Ih). Mandible incisor with seta!

brush and molar large acessory process (Fig. 2e).

Antenna 2 without minute denticles or spines

over the surface of the peduncle and flagellum

(Fig. 5b). Pleopods 1-4, peduncle margin crenate

(Fig. 2i). Pleonites 6 and 7, margin dorsally

crenate (Fig. 11).

398

G. K. WALKER-SMITH AND G. C. B. POORE

Composition. P. longipes, P. helizensis Modlin,

1991.

Remarks. This genus is distinguished most easily

from Levinebalia by the denticulate eyes, antenna

2 without minute denticles or spines over the sur¬

face of the peduncle and flagellum, pleopods 1-4,

peduncle margin crenate and pleonite 6 and 7 dor-

sally crenate. Undescribed species are known

from Australia.

Levinebalia Walker-Smith

Levinebalia Walker-Smith, 2000: 138.

Type species. Levinebalia maria Walker-Smith,

2000 (original designation).

Diagnosis. Eyes without denticles or cuticular

outgrowths. Mandible incisor without setal brush

or accessory molar process. Antenna 2 peduncle

and flagella with patches of minute denticles or

spines (Fig. 5a). Pleopods 1^, peduncle margin

smooth. Pleonites 6 and 7, margins with ill-

defined crenations.

Composition. L. maria, L. fortunata (Wakabara,

1976). '

Remarks. This genus is distinguished from

Paranehalia by smooth eyes, minute denticles or

spines over the surface of antenna 2 peduncle and

flagella, smooth pleonites margins and smooth

pleopod peduncles. Pleonites may sometimes

have tiny, ill-defined crenations.

Nebaliidac Samouelle

Nebaliadae Samouelle, 1819: 100.

Nebaliidae Baird, 1850: 31-38.—Sars, 1887: 6-7.—

Verrill, 1923: 205-206.—Hessler, 1984: 656.

Type genus. Nehalia Leach, 1814 {by monotypy).

Diagnosis. Subtenninal rostral spine rarely pre¬

sent; keel sometimes present (Figs la, d). Visual

elements present or absent. Supraocular scale

sometimes present (Fig. If). Antenna 1, antero-

denticulate fourth article absent; male, flagellum

not swollen, but may have numerous aesthetascs

(Fig. 2f). Antenna 2, peduncle without cuticular

outgrowths or minute denticles; articles 3 and 4

sometimes ftised (Fig. 2b); male antenna 2 greatly

elongate, reaching to the caudal furca (unknown

for Dalilella). Mandible incisor, present (except

for Speonehaiia). Molar process well developed

(Fig. 2h), without setal brush; without accessory

process; sometimes with accessory tootlVspine

(Fig. 2i). Maxilla 1 palp long, well developed.

Maxilla 2 (Fig. 21) with at least first 3 endites well

developed; much smaller than thoracopod 1.

Thoracopods not extending well beyond ventral

margin of carapace; closely spaced (Fig. 4a).

Thoracopod 1 differing only slightly from thora-

copods 2-7. Thoracopods 2-5 cpipod longer than

exopod (Fig. 4d), or absent (Fig. 4e); endopods

showing a degree of articulation. Pleonites 4-7

crenate over entire margin (Fig. 4a). Pleopod 1

exopod generally with comb-row (Fig. 4k). Pleo¬

pod 2-4 exopod with parallel margins or slightly

expanded medially; outer margins with setae

sometimes in pairs. Pleopod 5 longer or shorter

than pleopod 6. Pleopod 6 uni- or biarticulatc.

Cauda! rami tapering evenly to lip. Thorax not

inflated, body cuticle firm. Carapace strongly

emarginate midsagittally. Entire body length less

than 20 mm.

Composition. Nebalia Leach, 1814; Nebaliella

Thiele, 1904; Dalilella Messier, 1984; Speone-

balia Bowman, Yager and Iliffe, 1985.

Remarks. The diagnosis for Nebaliidac has been

modified since Messier (1984) to include Speone-

balia and exclude Paranebalia and Levinebalia

(removed to Parancbaliidae). Sarsinebalia has

been synonymised with Nebalia. Authorship of

the family name has been attributed to Baird

(1850) by other authors but Samouelle's (1819)

name has precedence.

Key to genera of Nebaliidae

1. Rostrum with keel shorter than rostral llange (Figs la, d) or absent; molar

without accessory toolh/spine (Fig. 2h); thoracopod exopod without proxi¬

mal lobe (Fig. 4d); thoracopods 2-5 cpipod longer than exopod (Fig. 4d);

pleopods 2-4 outer margins parallel (Figs 4h-k).2

— Rostrum with keel longer than rostral flange (Figs Ic, e); molar with acces¬

sory tooth/spine (Fig. 2i); thoracopod exopod with proximal lobe (Fig. 4e);

thoracopods 2-5 cpipod absent (Fig. 4e); pleopods 2-4 outer margins

slightly expanded midway and/or distally (Fig. 4f). Nebaliella

PHYLOGENY AND KEYS TO LEPTOSTRACA (CRUSTACEA)

399

2. Rostrum without keel; eye longer than rostrum; antenna I article 4 without

robust setae.3

— Rostrum with keel shorter than rostral flange (Figs la. Id); eye shorter than

rostrum; antenna I, article 4 with robust setae (Fig. Ip). Nebaiia

3. Eye without denticles, narrow, tapering distally, surface smooth, without

visual elements, without supraocular scale; maxilla 1 second endite simple

(Fig. 3c), maxilla 2 with marginal organelles (Fig. 2k). Speonehalia

— Eye (Fig. Im) with denticles, strongly curved (banana shape), surface with

denticles, without visual elements, with supraocular scale; maxilla I second

endite complex (Fig. 3a); maxilla 2, marginal organelles absent. Dafilella

Nehalia Leach

Nebaiia Leach. 1814; 99.—Thomson, 1879:

418-419.—Sars. 1896: 7-8.— Thiele, 1904; 10-12.—

Thiele. 1905: 61.—Barnard, 1914: 443-446.—Page,

1929:41^2.—Cannon, 1931; 221-222.—Clark, 1932:

225-230.—Wagele, 1983: 127-138.— Dahl, 1985;

144-157.—Dahl. 1990: 73-91.

Epinehalia Clark, 1932: 225-230 (type .species

Epinebalia piigettensis Clark, 1932 by monotypy).

Sarsbwhalia Dahl, 1985: 160-163 (type species

Nehalia typhlops Sars. 1870 by original designation)

syn. nov.

Type species. Cancer hipes Fabricius. 1780 (by

monotypy).

Diagnosis. Rostnim with keel shorter than rostral

flange, commonly without sublerminal spine

(Fig. la). Eyes shorter than rostrum, generally

dorsally convex, sometimes with papilla (Figs If,

j); usually with ommatidia or visual elements;

surface without denticles, ventral margin not

extremely convex. Supraocular scale present

(Figs If, i). Maxilla 2 exopod at least half length

of endopod (except N. typhlops). Antenna 1

article 4 with I or more robust setae (Fig. Ip).

Antenna 2, article 2 with commonly with dorsal

spine, articles 3 and 4 fused (Fig. 2b). Mandible

palp article 3 tapering distally, with parallel mar¬

gins; incisor with 2 teeth (Fig. 2h); molar process

well developed without accessory tooth/.spine.

Maxilla 1 second endite complex (Fig. 3a).

Thoracopods exopod without proximal lobe; with

few setae (Fig. 4d); epipods large, well devel¬

oped. PIcopod 1, exopod generally with comb-

row (Fig. 4k). Pleopod 6 shorter than plcopod 5,

uniarticiilatc.

Remarks. The two characters distinguishing

Nebaiia from other Leptostraca are the presence

of a keel shorter than the rostral Range and the

presence of one or more robust setae on article 4

of antenna 1. Most species of Nehalia are very

alike and difficult to distinguish from one

another. However, four species are particularly

distinctive. Nebaiia daytoni and N. sp. A have an

unusual bilobed eye (Fig. In). Nebaiia sp. A also

has vcrticle striations on the anteroventral surface

of the carapace. Nebaiia sp. A and sp. B and N.

typhlops all lack the comb-row on the exopod of

plcopod 1. Nebaiia sp. B has an unusual triangu¬

lar shaped eye and N. typhlops has a rectangular

to circular eye that lacks pigment. The exopod of

maxilla 2 of N, typhlops is reduced to less than

half the length of the endopod.

Sarsinehalia Dahl, 1985 is a new junior

synonym.

Nehaliella Thiele

yVe6a//£?//fl Thiele, 1904:4-9, 24-25.—Cannon, 1931:

216-221.—Walker-Smith. 1998: 41.

Type species. Nehaliella antarctica Thiele, 1904

(by monotypy).

Diagnosis. Rostrum with keel longer than rostral

flange, subtenninal spine absent (Fig. Ic). Eyes

strongly curved, extending beyond the end of the

rostral keel, lacking visual elements (Fig. le).

Antenna 1 without robust setae on article 4.

Antenna 2, peduncle articles 3 and 4 not fused,

without culicular outgrowths (Fig. 2a). Mandible

incisor with 2 teeth. Molar with accessory

tooth/spine (Fig. 2i), witliout large accessory pro¬

cess (Fig. 2e). Maxilla 1 second endite bilobed

(Figs 3b, 3d). Maxilla 2 exopod greater than half

length of endopod, biarticulate, without

organelles. Thoracopods without epipods (Fig.

4c). Thoracopod exopod with proximal lobe,

heavily setose (Fig. 4e). Pleonites 2-7 posterior

margin crenate. Pleopods 2-4 with lateral setae

not in pairs (Fig. 4f). Pleopod 6 longer than pleo¬

pod 5, biarticulate. Carapace not sculptured, but

may have a carina on lower anterolateral surface,

posterodorsal margin with liny denticles.

Remarks. Nehaliella occurs at depths ranging

from 3 m to over 100 m. The eyes, like those of

Dahlella and Nebaiia typhlops, lack visual pig¬

ments; they are strongly curved and extend

beyond the end of the rostrum like those of

Dahlella but, unlike Dahlella., lack denticles. The

rostrum is unique. Antenna 2 articles 3 and 4 arc

not fused in Nehaliella, Nehaliopsis and

Speonehalia. Thoracopod epipods are absent in

400

G. K. WALKER-SMITH AND G. C. B. POORE

Nebaliella but present in all other leptostracans.

The posterior margin of the carapace of

Nebaliella has a series of close-set spines; similar

ornamentation is found in Speonebalia.

Dahtella Messier

Dahlella Messier, 1984: 656.

Type species. Dahlella caldariensis}:\Q^^\QX^ 1984

(original designation).

Diagnosis. Rostrum without keel or subterminal

spine. Eyestalks without visual elements, curved,

longer than rostrum, tapering gradually to point;

anterior margin denticulate (Fig. Im); supraocular

scale present. Antenna 1 without robust setae on

article 4. Antenna 2, peduncle articles 3 and 4

fused. Mandible incisor with 2 teeth; molar

process well developed, without accessory

tootli/spine; mandible palp, distal article with 2

rows of setae. Maxilla I second endite complex

(Fig. 3a). Maxilla 2 exopod small, less than

quarter length of endopod; endopod biarticu-

late, without organelles. Thoracopod exopod

without proximal lobe, with few setae; cpipod

large, approximately equat in size to exopod;

proximal lobe small. Pleonites 2-7 posterior

margin crenate. Pleopods 2-4 exopods with pairs

of lateral setae. PIcopod 6 shorter than pleopod 5,

uniarticulate. Carapace not sculptured.

Remarks. The most pronounced feature of

Dahlella is the large, blind, toothed eye, seen only

in this monotypic genus from deep-sea vent com¬

munities. Messier (1984) suggested the eyes may

be used in scraping surfaces to loosen food such

as bacterial encrustations. Dahlella is most simi¬

lar to Nehalia, differing in the structure of the eye,

the lack of rostral keel, the small size of the exo-

pod of maxilla 2 and the shape of the proximal

lobe of the thoracic epipod. Dahlella shares with

Nebalia the presence of a supraocular scale.

Speonebalia Bowman, Yager and Iliffe

Speonebalia Bowman et al., 1985: 439.

Type species. Speonebalia cannoni Bowman,

Yager and Iliffe, 1985 (original designation).

Diagnosis. Rostmm without keel and subtenninal

spine. Eyes long and narrow, tapering distally,

extending beyond tip of rostrum, without visual

elements, surface smooth. Antenna I article 4

without robust setae. Antenna 2 peduncle articles

3 and 4 not fused, without large cuticular out¬

growths or minute denticles. Mandible without

incisor. Maxilla 1 second endite, simple (Fig. 3c).

Maxilla 2 endopod, uniarticulate, with scries of

oval marginal organelles, exopod very small (Fig.

2k). Thoracopods exopod without proximal lobe,

with few setae; epipods large, well developed.

Pleopod peduncles with crenate lateral margin.

PIcopod 1 without comb-row on lateral margin of

exopod. Pleopod 2-4 exopod with parallel

margins, smooth setae in pairs. Pleopod 6 shorter

than pleopod 5, biarticulate. Caudal rami short

and broad, tapering distally, margins densely

setose, setae on medial margin very long. Cara¬

pace strongly compressed laterally, covering

pleopods 1-5, more than 8 times length of

rostrum, with series of closc-sct obtuse spines

along posterior margin.

Remarks. Visual elements are also absent in

Dahlella, Nebaliella and Nebalia typhlops. The

mandibular incisor, absent in Speonebalia. is pre¬

sent in all other Nebaliidae. The mandibular palp

is unusually large in Speonebalia, reaching the

distal segment of the peduncle of antenna 2;

article 3 is unusual in its slender tapered

shape and its annature of three rows of complex

setae. The shape of the maxilla 1 second endite

(Fig. 3c) of Speonebalia is unique. Speone¬

balia is the only leptostracan with glands on max¬

illa 2. The exopod of maxilla 2 is reduced in

Speonebalia and Dahlella. The posterior margin

of the carapace of Speonebalia has a series of

close-set obtuse spines; species of Nebaliella also

show similar ornamentation on the carapace mar¬

gin. The caudal rami of Speonebalia has a dense

annature of long setae along the medial margin.

The genus is monotypic, its only species

recorded from marine caves. It has been sug¬

gested the caudal setae prevent the animal from

sinking and indicate a pelagic rather than a

benthic life (Bowman et al., 1985). All other

species of Leptostraca except Nebaliopsis typica

are thought to be benthic.

Acknowledgements

We would like to thank the South African

Museum for the loan of type material. We also

thank Dr Buz Wikson, Sydney, and an anonymous

reviewer for helpful comments and criticisms.

Thanks must go to Melissa Storey for proof¬

reading the manuscript and for providing useful

comments. We also thank Joan Clarke for her

assistance with the scanning electron microsopy

and David Paul for putting the SEM plate

together. This project was supported in part by

grants from the Australian Research Council and

the Victorian Institute of Marine Sciences.

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PHYLOGENY AND KEYS TO LEPTOSTRACA (CRUSTACEA)

403

Figure I. a. rostrum, dorsal and part lateral view, Nchaiia sp. C. b, rostrum dorsal view, Paranebalia sp. A. c, rostrum

dorsal view, NehalicUa extrema, after Thiele, 1905. d, rostrum lateral view, Nehalia typhlops^ after Dahl, 1985. c, eye and

rostrum, Nehaliella extrema, after Thiele, 1905. f. eye and supraocular scale, Nehalia pata^nnicuy after Dalil, 1990, g, eye

and supraocular scale, Nehalia sp. B. h. eye, Paranehalia sp. A. i, eye and suprocular scale, Nehalia lyphiops„ after Dahl,

1985. j, eye with dorsal papillae, Nehalia longicornis, after Dahl, 1990. k, eye. Nehaliella dcclivatas. I. Paranehalia sp. A.

rn, eye and supraocular .scale, Dahlella caldarensiSy after Mes.sler, 1984, n. eye and supraocular scale, Nehalia sp. A.

0 , antenna 2, Levinehalia maria, p, antenna I, Nehalia bipes hipes, after Dahl, 1985. q, antenna 1, mesial view,

Paranehalia sp. A. r, antenna 2, Paranehalia sp. A.

404

G. K. WALKER-SMITH AND G. C. B. POORE

Figure 2. a, antenna 2, NehaUeUa decUvatas. b, antenna 2, Nebalia sp. C. c, immature male, Paranehalia tippara. d. mature

male, Paranehalia sp. A. e. mandible incisor and molar, Paranehalia sp. A. f, antenna 2, Nehalia sp. A. male, g, mandible

palp, Levinehalia maria, h, mandible incisor and molar, Nehalia sp. A. i, mandible incisor and molar, Nebaliella decli-

vatas. j, maxilla 2, Nebaliopsis typica after Sars, 1887. k, maxilla 2. Speonehalia cannoni, after Bowman et al. 1985.

I, maxilla 2, Nehalia sp. C.

PHYLOGENY AND KEYS TO LEPTOSTRACA (CRUSTACEA)

405

Figure 3. a, maxilla \,Nehalia sp. C. b, maxilla X.Nebaliella declivatas. c, maxilla 1. Speonebalia cannorti, after Bowman

et al. 1985. d, maxilla 1, Nebaliella declivatas, male, e, carapace, Nebaliopsis typica, after Sars, 1887. f, Nebalia sp. A.

g, maxilla 1 (without palp), Paranebalia sp. A. h, Nebaliopsis typica, after Linder (1943). i, pleopods 1-4, Paranebalia

sp. A.

406

G. K. WALKER-SMITH AND G. C. B. POORE

Figure 4. a, Nebalia sp. C. b, thoracopod 3, Levinehalia maria, c, Ihoracopod 7, Nchaliopsts typica^ after Sars, 1887.

d, thoracopod 3, Nebalia sp. C. e, ihoracopod 3, Nebaliella brevicarinata., after Kikuchi and Gamo 1992. f. plcopod 2,

Nebaliella declivatas. g, plcopod 4, Nebaliopsis typica^ after Thiele, 1904. h, plcopod 2, Nebalia sp. C. i, plcopod 1,

Nebalia sp. A. j, plcopod 1, Nebaliella dedivatas. k, pleopod I, Nebalia sp. C. I, end of thoracopod 3, Nebalia puget-

tensis, after Dahl, 1985. m, caudal furca, Nebaliella dedivatas. n, caudal furca, Nebaliopsis typica., after Thiele, 1904.

PHYLOGENY AND KEYS TO LEPTOSTRACA (CRUSTACEA)

407

Figure 5. Antenna 2 in situ (left), peduncle article 3 and 4 (these are fused) and first article of flagellum, a, Levinehalia

maria, note row of small spines. Arrows point to large cuticular outgrowths, b, Paranehalia sp. A, note absence of small

spines. Arrows indicate large cuticular outgrowths.

408

G. K. WALKER-SMITH AND G. C. B. POORE

Hypothetical ancestor

Nebaiiopsis typica

Levinebalia fortunata

Levinebalia maria

Paranebalia sp. A

Paranebalia belizensis

Paranebalia longipes

Speonebalia cannoni

I r " Nebaliella antarctica

38*“ Nebaliella brevicarinata

1 p Nebaliella caboti

39L- Nebaliella declivatas

Dahlella caldariensis

Nebalia sp. C

Nebalia antarctica

Nebalia cannoni

Nebalia longicornis

’— Nebalia patagonica

Nebalia marerubri

Nebalia brucei

Nebalia capensis

Nebalia falklandensis

2f- Nebalia daytoni

46l— Nebalia sp. A

48 Sarsinebalia typhlops

47^ Nebalia sp. B

Nebalia bipes

Nebalia herbstii

Nebalia clausi

Nebalia hessleri

Nebalia strausi

Nebalia borealis

Nebalia lagartensis

Figure 6. Hypothesis for phylogeny of Leptostraca, tree 711. Numbers above branches are Bremer values; numbers below

are node numbers.

PHYLOGENY AND KEYS TO LEPTOSTRACA (CRUSTACEA)

409

Hypothetical ancestor

— Nebaliopsis typica

16 I Levinebalia fortunata

15 , 17 , 26 ^ 29 ,

25 , 35 |

Levinebalia maria

~ Paranebalia sp. A

Paranebalia belizensis

Paranebalia longipes

, 2 \ 23 , 26 ,

33 , 34 , 35 ) 31 , 32 ^

Speonebalia cannoni

Nebaliella antarctica

Nebaliella brevicarinata

Nebaliella caboti

Nebaliella declivatas

Dahlella caldariensis

- Nebalia sp. C

Nebalia antarctica

r Nebalia cannoni

~ Nebalia longicornis

Nebalia patagonica

Nebalia marerubri

Nebalia brucei

_r Nebalia capensis

Nebalia falklandensis

2 , 12

4 , 91

Nebalia daytoni

Nebalia sp. A

• Sarsinebalia typhlops

Nebalia sp. B

Nebalia bipes

Nebalia herbstii

Nebalia clausi

Nebalia hessleri

Nebalia strausi

Nebalia borealis

Nebalia lagartensis

Figure 7. Tree 711 with branch lengths. Numbers are characters with CI=1 and superscripts are state changes from the

plesiomorphic condition.

410

G. K. WALKER-SMITH AND G. C. B. POORE

100

-HvDothetical ancestor

- Nebaliopsis typica

77 r " Levinebalia fortunata

100 “ Levinebalia maria

Paranebalia sp. A

Paranebalia belizensis

Paranebalia longipes

- Speonebalia cannoni

r~ Nebaliella antarctica

100 - Nebaliella brevicarinata

100 50

Nebaliella caboti

100

-T

100 Nebaliella declivatas

61'

100 '

100

Dahlella caldariensis

Nebalia antarctica

Nebalia borealis

Nebalia brucei

Nebalia capensis

' Nebalia clausi

' Nebalia hessleri

' Nebalia lagartensis

■ Nebalia marerubri

’ Nebalia strausi

■ Nebalia sp. C

■ Nebalia bipes

■ Nebalia herbstii

" Nebalia cannoni

" Nebalia falklandensis

’ Nebalia longicornis

" Nebalia patagonica

“ Nebalia daytoni

" Nebalia sp. A

“ Sarsinebalia typhlops

" Nebalia sp. B

Figure 8. 50% majority-rule tree. Number above branch lines are Bootstrap values; numbers below arc percentage of

parsimonious trees retaining each clade (only values above 50% are included).

CONTENTS

Revision of Tertiary species of Anaskopora Wass (Bryozoa; Cribrimorpha)

P. E. Bock and P. L Cook . 179

Revision of the multiphased genus Corhulipora MacGillivray (Bryozoa: Cribrimorpha)

P. E. Bock and P. L Cook . j 91

Notes on the genera Nordgaardia and Uschakovia (Bryozoa: Bugulidae)

P.LCook . 215

New species and a new record of Chimaira Stephens (Trichoptera: Philopotamidae) from

Bougainville Island, Papua New Guinea

D. /. Cartwright . 223

New species of Hydropsychidae (Insecta: Trichoptera) from northern Australia

J. C. Dean . 231

Redcscription of Bungona Barker with new synonyms in the Australian Baetidae

(Insecta: Ephcmeroptera)

P. J. Snter and M. J. Pearson .247

Descriptions of new species of Binibhis (Amphipoda: Phoxocephalidae) from Australia and

Papua New Guinea with comments on the Birubiiis-Kidgaphoxus-Tickalerus-Yan complex

J. Taylor and G. C. B. Poore . 255

Bathyal Joeropsididac (Isopoda: Asellota) from south-eastern Australia, with description of

two new genera

J, Just . 297

Platyprotus phyllosoma^ gen. nov and sp. nov., from Enderby Land, Antarctica, an unusual

munnopsidid without natatory pereopods (Crustacea: Isopoda: Asellota)

.335

Three new species of Cirolana Leach, 1818 (Crustacea: Isopoda: Cirolanidae) from Australia

S. J. Keable . 347

Redesciiption of the tropical Australian isopod, Lyidotea nodata Hale, 1929 (Crustacea: Idoteidae)

R. King and G. C. B. Poore . 355

Plakarthrium australiense, a third species of Plakarthriidae (Crustacea: Isopoda)

G. C. B. Poore and A. Brandt . 373

A phytogeny of the Leptostraca (Crustacea) with keys to families and genera

G. K. Walker-Smith and G. C. B. Poore . 333