Memoirs of Museum Victoria 61(1): 1-40 (2004)

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

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

A molecular and morphological revision of genera of Asterinidae

(Echinodermata: Asteroidea)

P. Mark O’Loughlin 1 and Jonathan M. Waters 2

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

(pmo@bigpond.net.au)

department of Zoology, University of Otago, PO Box 56, Dunedin, New Zealand

(j onathan. waters @ stonebo w. otago. ac .nz)

Abstract O’Loughlin, P.M., and Waters, J.M., 2004. A molecular and morphological revision of genera of Asterinidae

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

A molecular phylogeny has inspired a reappraisal of the systematics of the Asterinidae. New morphological charac-

ters are defined and illustrated and used to diagnose all genera. A table of the distribution of morphological characters

among genera and key for genera of Asterinidae are provided. New genera of Asterinidae are erected: Aquilonastra

O’Loughlin, Indianastra O’Loughlin, Parvulastra O’Loughlin and Pseudopatiria O’Loughlin. Patiria is raised out of

synonymy with Asterina. Allopatiria is a junior synonym of Asterina , Manasterina is a junior synonym of Disasterina,

and Paxillasterina is a junior synonym of Asterinides. The genus Asterinopsis and the genus and species Desmopatiria

flexilis are nomina nuda. Patiriella tangribensis is a nomen dubium. Genera reviewed are: Anseropoda, Asterina ,

Asterinides, Callopatiria, Cryptasterina, Disasterina, Kampylaster, Meridiastra, Nepanthia, Paranepanthia, Patiria,

Patiriella , Pseudasterina , Pseudonepanthia, Stegnaster, Tegulaster and Tremaster. Asterina cephea var. iranica is raised

to species status. Enoplopatiria siderea is a junior synonym of Asterina stellifera. Disasterina leptalacantha var. africana

is no longer recognised as a subspecies. Disasterina spinulifera is a junior synonym of Disasterina praesignis. A syn-

onymy of Tremaster novaecaledoniae with Tremaster mirabilis is formalised. Asterinid species reassigned on the basis

of molecular evidence and morphological congruity are new combinations: Aquilonastra anomala, A. batheri, A. burtoni,

A. coronata, A. minor, A. scobinata, Meridiastra calcar, M. gunnii, M. medius, M. mortenseni, M. occidens, M. oriens,

Paranepanthia aucklandensis, Parvulastra calcarata, P. exigua, P. parvivipara, P. vivipara, Patiria chilensis, P. miniata,

P. pectinifera. Species reassigned on the basis of morphological evidence are new combinations: Aquilonastra cepheus,

A. corallicola, A. heteractis, A. iranica, A. limboonkengi, A. rosea, Asterina ocellifera, Asterinides hartmeyeri, A. pilosa,

A. pompom, Disasterina ceylanica, D. longispina, Indianastra inopinata, I. sarasini, Nepanthia pedicellaris, Parvulastra

dyscrita, Pseudonepanthia briareus, P. gracilis, P. grangei, P. nigrobrunnea, P. reinga, P. troughtoni, Pseudopatiria

obtusa, Tegulaster alba, T. leptalacantha, T. praesignis. Three species remain incertae sedis: Asterina lorioli, Asterina

novaezelandiae and Nepanthia brachiata. A table of asterinid species is provided, with original and current combinations.

It is concluded that Asterinidae is a cosmopolitan family, mainly of shallow-water narrow-range genera but including

some more widespread in deeper waters of all oceans.

Keywords Echinodermata, Asteroidea, Asterinidae, new genera, taxonomy, molecular, morphology

Introduction

Traditional systematic studies of asteroids have been con-

founded, to an extent, by morphological characters that are of

dubious phylogenetic value and limited by observable size and

historical choice (Clark and Downey, 1992). Some morpho-

logical characters currently used in asteroid systematics, for

instance, may be subject to strong selection and thus remain

stable while evolutionary divergence occurs in other characters

(e.g. molecular divergence in living fossils; Avise et al., 1994).

Hence taxa that are superficially similar, and perhaps con-

generic on traditional morphological criteria, may prove to be

unrelated (O’Loughlin et al., 2002; Waters et al., 2004). Some

other morphological characters may be plastic and readily

diverge among taxa that are closely related (e.g. O’Loughlin et

al., 2003). Current taxonomic treatments of asteroids may also

be hindered by the possibility that certain morphological char-

acters are phylogenetically informative for some clades but

homoplasious and unreliable for others (Mah, 2000). The abun-

dant phylogenetic information provided by DNA sequence data

(Avise, 2000) presents a means of reassessing taxonomic

relationships and stimulates reappraisal of morphological,

behavioural, physiological and ecological traits.

P. Mark O’Loughlin and Jonathan M. Waters

The Asterinidae comprise 21 genera and about 116 species

according to the comprehensive index of taxa by A.M. Clark

(1993) and the subsequent work of Rowe (in Rowe and Gates,

1995), Campbell and Rowe (1997), O’Hara (1998),

VandenSpiegel et al. (1998), A.M. Clark and Mah (2001),

H.E.S. Clark and McKnight (2001), O’Loughlin (2002),

O’Loughlin et al. (2002, 2003) and Dartnall et al. (2003). The

need for a systematic revision was first commented on by

Verrill (1913). Recently, Rowe (in Rowe and Gates, 1995),

Campbell and Rowe (1997), O’Loughlin (2002) and

O’Loughlin et al. (2002) noted continuing conflicting opinions

on the systematic status of, and assignment of species to, the

genera Asterina Nardo, 1834, Asterinides Verrill, 1913 and

Patiriella Verrill, 1913.

O’Loughlin (2002) provided a restricted morphological

review of Asterinidae and erected a new genus ( Meridiastra )

with three new species (M. fissura, M. nigranota, M. rapa ).

Hart et al. (1997) reported the first molecular phylogeny for 12

asterinid species assigned to the genera Asterina and Patiriella.

The phylogeny indicated that neither Asterina nor Patiriella is

monophyletic. O’Loughlin et al. (2002, 2003) reported two

molecular phylogenies and the erection of four new

Australasian asterinid species ( Patiriella medius, P. mortenseni,

P occidens, P. oriens), provisionally assigned to Patiriella in

the absence of molecular data from other genera. Hart et al.

(2003) reported a molecular phylogeny on which Dartnall et al.

(2003) erected an additional genus ( Cryptasterina ) and species

(C. hystera Dartnall and Byrne, 2003). Waters et al. (2004)

reported a molecular phylogeny (adapted in Fig. 1) for 31

asterinid species, predominantly Australasian. Their phylogeny

showed strong resolution at shallow levels, with six well-sup-

ported clades. With additional species included, the polyphylet-

ic assemblages of Asterina and Patiriella illustrated by Hart et

al. (1997) were further elucidated. Some genera, defined on the

basis of morphology, have molecular support. Other well-

defined clades may deserve generic rank but need morphologi-

cal support. Some genera as presently conceived appear poly-

phyletic. They invite reappraisal to explore whether morpholo-

gy will support division into smaller unrelated genera.

This paper seeks congruence between morphological and

molecular data: we evaluate the phylogenetic reliability of

characters traditionally used to differentiate asterinid genera

and explore the utility of new characters. Ideally, genera are

monophyletic groups of species diagnosed by shared character

states — morphological, behavioural and molecular — that

reflect common descent (De Queiroz and Gauthier, 1992). Here

we apply phylogenetically informative morphological charac-

ters to define genera anew and recognise new genera. The mor-

phological analysis is extended to many asterinid genera and

species for which there are currently no molecular data. Future

molecular analyses will help to evaluate the decisions made

here on the basis of morphology alone.

In the cladogram of Waters et al. (2004) basal relationships

were unclear. It did not demonstrate the monophyly of the

family as presently construed. Dermasterias (Asteropseidae)

was placed within the asterinid clade. It should be noted that

this relationship received minimal phylogenetic support. The

unresolved affinities of Demasterias have little bearing on the

objectives of this paper, which are to redefine genera currently

assigned to the Asterinidae. On morphological evidence, the

genus Tremaster Verrill, 1880 appears to be inappropriately

included in Asterinidae. But a reassessment of what genera

belong in Asterinidae, such as the inclusion or not of Cycethra

Bell, 1881 (currently in Ganeriidae), and retention or not of

Tremaster, should await appropriate molecular phylogeny data.

In this revision all of the species of all of the genera were

reviewed, except for Anseropoda where a sample of species

only was examined. Material for three asterinid species, for

which the morphological descriptions raise doubts about

their generic assignment, could not be found. They are placed

incertae sedis.

Terminology follows that defined in the glossary and illus-

trated by Clark and Downey (1992, figs 2, 3), except that

“papular space” (“restricted area with papular pores”, fig. 15f)

is used for “papular area”, and “papulate areas” is used to refer

to the parts of the abactinal surface where papulae occur (as

defined by O’Loughlin, 2002). The form of spines and spinelets

is of ten broad types (form frequently revealed by clearing with

bleach): long and thin, needle-like, “acicular” (fig. 5c); round

base and apical point, cone-shaped, “conical” (fig. lOg); long,

thick, finger-like, “digitiform” (fig. 5e); short and thick, “gran-

uliform” (fig. 5a), which may be subspherical, “globose” (fig.

5b), or “short columnar” (figs 15d, 16h); thin, few prominent

points laterally and distally, “splay-pointed” (figs 5d, 6c, 9d);

distal pointed glassy tip, “thorn-tipped” (fig. 17d); sac-like base

and tapering distally to narrowly rounded end or point, “sacci-

form” (figs 5f, 8b, 8g, 9c); combination of “sacciform” and

“splay-pointed” (fig. 5d). The armature of abactinal plates is

consistently referred to as “spinelets”, irrespective of size. The

armature of actinal plates is consistently referred to as “spines”,

irrespective of size. Specimen size refers to preserved material,

and is categorized as “small” (up to R = 25 mm), “medium”

(R = 26-65 mm), and “large” (R greater than 66 mm). All

measurements refer to preserved material.

Abbreviations for institutions are: AM, Australian Museum,

Sydney; BMNH The Natural History Museum, London;

MNHN, Museum National d’Histoire Naturelle, Paris; MNZ,

Museum of New Zealand, Te Papa Tongarewa, Wellington;

NMV, Museum Victoria, Melbourne, Australia; TM, Tasmanian

Museum, Hobart; UF, University of Florida, USA; USNM,

United States Museum of Natural History, Smithsonian

Institution, Washington; WAM, Western Australian Museum,

Perth; YPM, Peabody Museum of Natural History, Yale

University, New Haven; ZMUC, Zoological Museum,

University of Copenhagen, Denmark. Photography for figures

4-18 was performed using a Leica MZ16 stereomicroscope,

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

composition of images.

Invalid and junior synonyms of genera and species (indexed

by A.M. Clark, 1993) are not repeated in this work unless their

systematic status is reviewed. The summary distributional data

given for the genera below are in part based on more detailed

ranges given by A.M. Clark (1993), Rowe (in Rowe and Gates,

1995), O’Loughlin (2002), O’Loughlin et al. (2002, 2003), and

Dartnall et al. (2003). Throughout the paper, “Rowe (1995)”

refers to “Rowe (in Rowe and Gates, 1995)”. The erection of

Genera of asterinid sea-stars

Dermasterias imbricata (Pacific United States)

Nepanthia troughtoni (temperate Australia)

Stegnaster inflatus (New Zealand)

asterinid sp (New Zealand)

Asterina aucklandensis (Auckland Is)

Paranepanthia grandis (temperate Australia)

100

I Asterina chilensis (Chile)

I Asterina pectinifera (Japan)

100

Asterina miniata (Pacific United States)

— Patiriella gunnii (temperate Australia)

Patiriella oriens (temperate Australia)

Patirieila medius (temperate Australia)

Patiriella occidens (temperate Australia)

Patiriella calcar ( temperate Australia)

- Meridiastra fissura (temperate Australia)

- Meridiastra nigranota (temperate Australia)

100 1 — Meridiastra atyphoida (temperate Australia)

Patiriella mortenseni (New Zealand)

Asterina minor ( Japan)

92_

14 ioo \ Asterina anomala (Samoa)

100

49 1 Asterina anomala (L. Howe)

i — Asterina coronata (Japan)

Asterina batheri (Japan)

Asterina burtoni (Red Sea)

Asterina scobinata (temperate Australia)

10Q p Cryptasterina pacifica (Japan)

24 L Cryptasterina pentagona (tropical Australia)

Patiriella regularis (New Zealand)

Patiriella exigua (temperate Australia)

Patiriella exigua (S. Africa)

Patiriella parvivipara (temperate Australia)

Patiriella vivipara (temperate Australia)

Patiriella calcarata (Juan Fernandez)

ioo

Asterina gibbosa (E. Atlantic, Mediterranean)

- Asterina phylactica (E. Atlantic, Mediterranean)

Pseudonepanthia

Stegmaster

I Paranepanthia

II Patiria

III Meridiastra

IV Aquilonastra

Cryptasterina

Patiriella

V Parvulastra

VI Asterina

Figure 1. Phylogenetic relationships of asterinid mtDNA sequences (after Waters et al., 2004). Support for particular reconstructions is indicated

by bootstrap values (>50%; in bold) and by decay indices (in italics). MtDNA Clades I-VI are identified on the basis of high bootstrap support.

Generic assignments made in this paper appear on the right.

P. Mark O’Loughlin and Jonathan M. Waters

new genera and systematic decisions in this work are made by

the first author (O’Loughlin).

Morphological diagnostic characters (see Table 1)

The molecular phylogeny of Waters et al. (2004) has provided

evidence of relationships between species. Morphological

examination of molecular clades has been a starting point for

seeking and confirming characters which are useful for generic

diagnoses. A significant outcome has been the recognition of

internal skeletal plates as good diagnostic characters. Their

absence and presence in different combinations vary across

genera and are objective criteria.

Clark and Downey (1992) recognised these plates, and

defined them collectively as “superambulacral plates”. These

internal support plates are distinguished here as three inde-

pendent types, which are each reliable diagnostically. The

Clark and Downey (1992) definition of “superambulacral

plate” is emended to: an internal plate articulating between the

actinal end of an ambulacral plate, and actinal or marginal or

abactinal plates (figs 4a, e, 14e, f; illustrated in A.M. Clark,

1983: fig. 5b). Two additional types of plates are defined. A

“superactinal plate” is: an internal interradial plate articulating

between abactinal plates, and actinal or other superactinal or

superambulacral plates (figs 4b, d, lOd, 15c). “Transactinal

plates” are: internal plates in transverse series across rays above

the actinal plates (fig. 4f; illustrated but not defined in A.M.

Clark, 1983: figs 5d, e [ brevis a junior synonym of belcheri ]).

As with many diagnostic characters, the development and pres-

ence of internal plates may be related to size. They have not

been observed in the pedomorphic species of some genera.

Body form, in terms of the degree to which rays are discrete

and the form of the rays, is useful diagnostically. But both are

subject to variation within a species and because of the habit of

the animal and preservation artefact. The presence of noticeable

body integument is consistent for some genera, but varies in the

degree to which it is evident. Whether abactinal plates imbri-

cate, or form regular series, or are irregularly arranged, are use-

ful criteria, but also vary. The presence of single large papulae

or numerous small papulae in papular spaces distinguishes

some genera but the distinction is not absolute. Whether actinal

plates are in longitudinal series parallel to the furrow, or in

oblique series from the furrow to the margin, are useful charac-

ters, but sometimes a combination of plate arrangements is evi-

dent and sometimes there is variation within a species (in one

species apparently changing with size). The form of spinelets

and spines is broadly useful for generic diagnoses, but there is

variation in detail of form among and within species. Whether

spinelets are judged to be “opaque” or “glassy” is attempted,

but when denuded by bleach all spinelets appear crystalline

and thus glassy. There is subjectivity in making diagnostic

statements using the above characters.

The occurrence of pedicellariae is diagnostically reliable for

some genera but in others they are present in only some of the

species. Glassy convexities occur on the plates of many

species. They are present on all species of some genera and

absent in one genus, but within one genus they are present on

one species and absent on another.

Some characters were found to vary within genera and

species and are unhelpful for generic diagnostic purposes:

extent of development of a carinal series of plates; presence

of a border of five large radial and five small interradial

plates around the disc; precise numbers of papulae and

secondary plates per papular space; numbers of abactinal

spinelets and actinal spines per plate; size of plates and

spinelets and spines; fissiparity; and gonopore occurrence

abactinally or actinally.

We acknowledge the limitations we face in diagnosing

genera using morphological characters. These result from

examination of few specimens of each species, consequent

limited exposure to intraspecific variations and developmental

variations, character varability such as noted above, and the

subjectivity of decision-making when addressing continuous

characters. Table 1 lists some morphological characters judged

to be useful diagnostically and their occurrence in the asterinid

genera confirmed or erected in this study.

Key to genera of Asterinidae

1. Furrow spines in longitudinal arrangement on

adambulacral plates (fig. 16d); 2 series of tube feet in

each ambulacrum; lacking internal plated brood

chambers 2

— Furrow spines in vertical series on adambulacral plates

(fig. 17f); 4 series of tube feet; internal plated brood

chambers with abactinal and actinal interradial openings .

Tremaster Verrill

2. Rays narrow at base, to varying degrees subcylindrical or

digitiform (fig. 2j) 3

— Rays broad at base or not discrete, not subcylindrical or

digitiform (figs 2b, i) 6

3. Superambulacral series of plates present (fig. 4a) 4

— Lacking superambulacral, transactinal and superactinal

plates (fig. 16g) . . . Pseudopatiria O’Loughlin gen. nov.

4. Superactinal plates present (fig. 4b) 5

— Superactinal plates absent (fig. 14e)

Pseudonepanthia A.H. Clark

5. Rays narrowly flat actinally, marginal edge weakly

angular; pedicellariae present; transactinal plates present

(fig. 4f) Nepanthia Gray

— Rays broadly flat actinally, marginal edge strongly angu-

lar (fig. lOd); lacking pedicellariae; lacking transactinal

plates Callopatiria Verrill

6. Interradii very thin; abactinal and actinal plates interiorly

contiguous throughout the interradii (fig. 7) 7

— Interradii not very thin; abactinal and actinal interradial

plates contiguous or connected by superactinal plates dis-

tally only (fig. 15c) 9

7. Body arched; furrow spines project actinally, in continuous

series, with integument and granule cover (fig. 17c);

actinal spines glassy thorn-tipped (fig. 17d)

Stegnaster Sladen

— Body not arched; furrow spines not projecting actinally, in

webbed groups on adambulacral plates, not covered with

integument and granules; actinal spines not glassy thorn-

tipped 8

Genera of asterinid sea-stars

8. Abactinal spinelets sacciform; interradii extensively sup-

ported by long thin articulating interior projections from

abactinal and actinal plates (fig. 7) . . Anseropoda Nardo

— Abactinal spinelets granuliform; interradii extensively sup-

ported interiorly by contiguous imbricating abactinal and

actinal plates Pseudasterina Aziz and Jangoux

9. Lacking superambulacral, transactinal and superactinal

plates 10

— One or both of superambulacral and superactinal plates

present 13

10. Abactinal body covered by thick integument with subgran-

ular spinelets; body typically in high arched (domed) shape

(fig. 17e); abactinal interradial plate arrangement irregular

Kampylaster Koehler

— Body not covered by thick integument with subgranular

spinelets; body rarely in high arched (domed) shape;

abactinal interradial plate arrangement regular 11

11. Abactinal spinelets glassy, acicular or subsacciform, in

tufts (fig. 8f) or paxilliform (fig. 8i); inferomarginal plates

with distal tuft of spinelets (fig. 8h) . . Asterinides Verrill

— Abactinal spinelets opaque, granuliform or digitiform, not

in tufts or paxilliform; inferomarginal spinelets not in

distal tufts 12

12. Rays discrete; abactinal spinelets columnar to digitiform

(fig. 6a) (Atlantic, Mediterranean) Asterina Nardo

— Rays not discrete, margin straight or slightly incurved;

abactinal spinelets granuliform (fig. 5a) (temperate Indo-

Pacific) Meridiastra O’Loughlin

13. Abactinal and actinal plates with dense subpaxilliform

tufts of glassy acicular subsacciform spinelets and spines

(fig. 5c); lacking superambulacral plates (fig. 15c)

Paranepanthia Fisher

— Abactinal spinelets and actinal spines not in dense subpax-

illiform tufts; superambulacral plates present 14

14. Abactinal spinelets glassy, acicular or sacciform or splay-

tipped sacciform or long thin conical (fig. 13c) 15

— Abactinal spinelets opaque, granuliform or digitiform, or

short thin to thick columnar (fig. 15d) 18

15. Abactinal plates with numerous firmly attached, glassy

spinelets (fig. 9f); superomarginal spinelets same as those

on abactinal plates (fig. 9d)

Aquilonastra O’Loughlin gen. nov.

— Abactinal plates with very fine glassy spinelets, numerous

to none, weakly attached (fig. 13c); superomarginal

spinelets few to none (fig. 13d) 16

16. Abactinal plates loosely contiguous leaving non-plated

spaces (fig. 12); distal abactinal plates in series perpendic-

ular or zig-zag to margin (fig. 12a); superomarginal plates

small, not in regular series; inferomarginal plates with

stout sacciform spinelets (fig. lOe) . . Disasterina Perrier

— Abactinal plates imbricate, always contiguous; distal

abactinal plates not in perpendicular series to margin;

superomarginal plates in distinct regular series; infero-

marginal spinelets not stout sacciform 17

17. Body low; abactinal plates small, thin, deeply notched, up

to 6 series along each side of rays (fig. 13b); infero-

marginal spinelets acicular, in dense integument-covered

tufts (fig. 13d) Indianastra O’Loughlin gen. nov.

— Body high; abactinal plates large, thick, shallow notches or

crescentiform, up to 3 series along side of rays (fig. 18a);

inferomarginal spinelets discrete, not in dense integument-

covered tufts Tegulaster Livingstone

18. Form subpentagonal to medium-rayed stellate; large

papular spaces; numerous papulae and secondary plates

per space (fig. 15f) 19

— Form pentagonal to subpentagonal; small papular spaces;

few papulae and secondary plates per space (fig. lib). .20

19. Abactinal plates with close subpaxilliform cover of

spinelets; spinelets thick to thin, short columnar or

sub-globose (figs 15d, e); more than 3 spines on mid-

interradial actinal plates Patiria Gray

— Spinelets on abactinal plates not in close subpaxilliform

cover; spinelets granuliform or digitiform (fig. 16e); up to

3 spines on mid-interradial actinal plates Patiriella Verrill

20. Superomarginal plates typically in prominent series, longi-

tudinally subrectangular; inferomarginal plates project nar-

rowly to define the margin; abactinal papular spaces with

predominantly 1 papula (fig. 11a); midray and distal

actinal interradial plates frequently with 1 spine

Cryptasterina Dartnall et al.

— Superomarginal and inferomarginal plates typically sub-

equal; inferomarginal plates project prominently to define

margin; abactinal papular spaces frequently with more than

1 papula (fig. lib); midray and distal actinal interradial

plates frequently with 2 spines (fig. 16c)

Parvulastra O’Loughlin gen. nov.

Asterinidae Gray, 1840

Synonymy. See Clark and Downey, 1992.

Diagnosis. See Clark and Downey, 1992.

Genera. See Table 2.

Remarks. We do not revise the family diagnosis of Clark and

Downey (1992). Changes to the list of genera in A.M. Clark

(1993) are: Cryptasterina Dartnall et al., 2003, Meridiastra

O’Loughlin, 2002, Aquilonastra O’Loughlin gen. nov.,

Indianastra O’Loughlin gen. nov., Parvulastra O’Loughlin

gen. nov. and Pseudopatiria O’Loughlin gen. nov. are added;

Allopatiria Verrill, 1913 is a junior synonym of Asterina Nardo,

1834; Patiria Gray, 1840 is raised out of synonymy with

Asterina', Manasterina H.L. Clark, 1938 is a junior synonym of

Disasterina Perrier, 1875; Paxillasterina A.M. Clark, 1983 is a

junior synonym of Asterinides Verrill, 1913, which is main-

tained as a valid genus; and Asterinopsis Verrill, 1913 and

Desmopatiria Verrill, 1913 are rejected.

We agree with H.L. Clark (1938) that Asterinopsis Verrill,

1913 is nomen dubium. Verrill (1913) designated A sterias peni-

cillaris Lamarck, 1816 (Indo-Pacific) the type species, and

assigned Asterina pilosa Perrier, 1881 (West Indies) and

Asterina lymani Perrier, 1881 (synonymous according to Clark

and Downey, 1992) to Asterinopsis. Fisher (1919) added

Nepanthia pedicellaris Fisher, 1913 to Asterinopsis. Mortensen

(1933) judged that the type of A. penicillaris was lost and that

no reliable specimens or image were available. H.L. Clark

(1938, 1946) pointed out that Mortensen’s case for the dubious

P. Mark O’Loughlin and Jonathan M. Waters

Figure 2. Abactinal views of type species of Asterinidae genera, a, Anseropoda placenta (R = 31 mm, AM J 107 14). b, Aquilonastra cepheus (R

= 12 mm, AM J8072). c, Asterina gibbosa (R = 16 mm, AM G11524). d, Asterinides folium (R = 19 mm, USNM E28573). e, Callopatiria gran-

ifera (R = 50 mm, NMV F98049). f, Cryptasterina pentagona (R = 11 mm, NMV F95959). g, Disasterina abnormalis (R = 22 mm, WAM Z6754).

h, Indianastra sarasini (R = 20 mm, NMV F95802). i, Meridiastra atyphoida (R = 12 mm, AM J9909). j, Nepanthia maculata (R = 73 mm, AM

J13918). k, Paranepanthia platydisca (R = 57 mm, holotype, USNM 32644). 1, Parvulastra exigua (R = 11 mm, neotype, TM H508).

Genera of asterinid sea-stars

Figure 3. Abactinal views of type species of Asterinidae genera (continued; Tremaster mirabilis is figure 17e). a, Patiria miniata (R = 67 mm,

NMV F98040). b, Patiriella regularis (R = 31 mm, AM J5594). c, Pseudonepanthia gotoi (R = 74 mm, holotype, USNM 36899). d, Pseudopatiria

obtusa (R = 54 mm, holotype, BMNH 1938.6.23.24). e, Stegnaster inflatus (R = 45 mm, NMV F95675). f, Tegulaster emburyi (R = 19 mm, holo-

type, AM J5605).

state of penicillaris would render Aster inop sis also dubious. He

further pointed out that Asterinopsis included both West Indian

and Indo-Pacific species. A.M. Clark (1993) awaited reassess-

ment, and listed N. pedicellaris and A. pilosa under

Asterinopsis. In this work N. pedicellaris is referred to

Nepanthia and A. pilosa is assigned to Asterinides.

A.M. Clark (1993) listed Desmopatiria Verrill, 1913, but

noted “validity doubtful since identity of type species doubt-

ful”. The genus was erected for the new species D. flexilis

Verrill, 1913. It was described from a single dry specimen,

without a label, found in the YPM collections with some aster-

oids from Chile. A recent search of the YPM collections by Eric

Lazo-Wasem (pers. comm.) failed to find the specimen. No

subsequent specimen has been referred to the species or genus.

In the absence of both type specimen and record of type local-

ity, the species D. flexilis and genus Desmopatiria are judged to

be nomina nuda.

Anseropoda Nardo

Figures 2a, 7, 8a-b

Anseropoda Nardo, 1834: 716.— Bell, 1891: 234-235.— Fisher,

1906: 1088.— Clark and Downey, 1992: 174-175.— A.M. Clark, 1993:

204-205.— Liao and Clark, 1995: 128.— Rowe, 1995: 32-33.—

McKnight (in Clark and McKnight), 2001: 161. (For complete

synonymy and discussion see Clark and Downey, 1992).

Diagnosis. Rays 5-18; body very thin, margin variably curved,

rays short or not discrete, broadly rounded or pointed; narrow

raised radial areas with single papulae, in scattered or single

longitudinal series, sometimes ringed by secondary plates;

abactinal plates thin, in longitudinal and oblique series, not

notched, each with subpaxilliform glassy, sacciform spinelets,

few or tuft; pedicellariae over papulae; actinal plates in longi-

tudinal and oblique series; actinal spines few to numerous

per plate, glassy, sacciform; lacking superambulacral plates,

superactinal plates present or absent; interradial plates meet

internally by long thin articulating projections, extensive in

interradii.

Type species. Asterias placenta Pennant, 1777 (subsequent

designation by Bell, 1891) (junior synonym: Asterias

membranacea Retzius, 1783, by Bell, 1891).

Other species. A. antarctica Fisher, 1940; A. aotearoa McKnight,

1973; A. diaphana (Sladen, 1889); A. fisheri Aziz and Jangoux, 1985;

A. ( Palmipes ) grandis Mortensen, 1933; A. habracantha H.L. Clark,

1923; A. insignis Fisher, 1906; A. lobiancoi (Ludwig, 1897);

A. ludovici (Koehler, 1909); A. macropora Fisher, 1913; A. novemra-

diata (Bell, 1905); A. pellucida (Alcock, 1893); A. petaloides (Goto,

1914); A. rosacea (Lamarck, 1816); A. tenuis (Goto, 1914).

Material examined. A. placenta. NE Atlantic, off Great Britain, NMV

F98043 (2).

A. rosacea. NW Australia, Broome, NMV F95811 (1); AM J7718

(1); Queensland, AM El 8 14 (1); Tasmania, AM E5037 (1).

Description with some species variations. Body very thin; rays

5 ( grandis , placenta ) or 6 ( insignis ) or 15-18 (rosacea), margin

variably curved; rays not discrete or short, broad basally,

P. Mark O’Loughlin and Jonathan M. Waters

Figure 4. Abactinal views, with plates removed and cleared with bleach to show internal morphology, a-b, Parvulastra dyscrita (R = 24 mm, TM

H854): a, series of superambulacral plates linking ambulacral and actinal plates; b, superactinal plates linking abactinal and actinal plates to sup-

port the interradial margin, c, Meridiastra oriens (R = 28 mm, NMV F73468): margin supported by internal contiguous projections of abactinal

and actinal plates (arrow), absence of superambulacral and superactinal plates, d, Patiria miniata (R = 65 mm, NMV F97444): multiple super-

actinal plate stmts supporting interradial margin, e, Pseudonepanthia troughtoni (R = 85 mm, NMV F73013): series of superambulacral plates,

and thick internal tisue lining, f, Nepanthia belcheri (R = 40 mm, NMV F95806): transactinal plates extending transversely across actinal inner

ray, and thick internal tissue lining.

Genera of asterinid sea-stars

Figure 5. Forms of spinelets and spines, a, Meridiastra gunnii (NMV F73248): granuliform abactinal spinelets. b, Parvulastra dyscrita (TM

H854): globose abactinal spinelets. c, Paranepanthia grandis (NMV F73976): acicular abactinal spinelets. d, Pseudonepanthia gotoi (USNM

36899): splay-pointed abactinal spinelets. e, Parvulastra dyscrita (TM H854): digitiform actinal spines, f, Disasterina longispina (WAM Z6760):

sacciform actinal spines.

Figure 6. Forms of abactinal fasciculate pedicellariae. a, Asterina ocellifera (BMNH 1969.12.16.13). b, Indianastra sarasini (WAM Z6836). c,

Nepanthia belcheri (NMV F95805).

rounded ( grandis , insignis, placenta ) or pointed ( rosacea );

rays slightly elevated as low ridges; size large {grandis up to

R = 130 mm); pedicellariae present {grandis, placenta,

rosacea ); none fissiparous.

Abactinal plates thin, imbricate, narrow raised radial papul-

ate area, extensive interradial non-papulate area; radial plates in

regular {grandis) or irregular {placenta ) distribution on narrow

upper ray; single papulae scattered on upper ray {rosacea, pla-

centa) or in 1-3 longitudinal series along each side of radial

midline {grandis)-, papulae may be each surrounded by ring of

up to 5 secondary plates {placenta , rosacea)-, interradial plates

imbricate, rhombic to fan-shaped, in both longitudinal and

oblique series, may be distally perpendicular to margin {pla-

centa ); disc not bordered; abactinal plates sometimes with

proximal low {placenta ) to high {rosacea) spinelet-bearing ele-

vation or dome; subpaxilliform tufts of spinelets (up to about

25 in placenta, up to 3 in grandis), sometimes multiple tufts per

plate {insignis); spinelets glassy, sacciform, pointed {placenta,

rosacea), sometimes more stout over papulae {grandis, pla-

centa, rosacea; presumed to be pedicellariae); cleared abactinal

plates sometimes with reticulate glassy appearance {rosacea),

lacking glassy convexities; superomarginal and infero-

marginal plates with abactinal-type spinelets, plates slightly

larger than adjacent abactinals; margin defined by projecting

inferomarginal plates. Actinal plates in longitudinal and oblique

series; may be distally perpendicular to margin {placenta ).

Actinal spines per plate: oral 5-8; suboral 3-10; furrow 4-9

proximally, webbed; subambulacral 2-7; actinal 3-12, webbed,

P. Mark O’Loughlin and Jonathan M. Waters

Table 1 . Selection of morphological characters present (•) in the genera of the Asterinidae

Genera of Asterinidae

Morphological characters

■W

f/m/w/m,

$ /*§

noticeable integument cover (fig. lOe)

interradii thin, inner surfaces connected (figs 2a, 7b)

habit arched, not flat (figs 3e, 17e)

rays usually 5 only

rays 5 and more

margins straight to slightly curved (figs 2a, i, 3e)

rays discrete, flat actinally, wide basally (figs 2b, c)

rays discrete, flat actinally, narrow basally (fig. 2e)

rays discrete, subcylindrical (figs 2j, 3c, d, 14e)

superambulacral plates present (figs 4a, 14e)

transactinal plates present (fig. 4f)

superactinal plates present (figs 4b, d, 15c)

interradial internal articulating supports (figs 7a, b)

internal resinous body lining (figs 4e, f)

abactinal plates imbricate (figs 8e, lOi, 15a)

abactinal plates loosely contiguous (figs 12a, b)

ray plates mostly in series (figs 2b, c, i, 9a, 13b)

ray plates mostly not in series (figs 3b, 12a, b)

few to many papulae per space (figs 10a, 15f)

mostly single papula per space (figs 9f, 13b)

spinelets opaque, stout, granuliform (figs 5a, b, lib)

spinelets opaque, stout, elongate (figs 6a, 16h)

spinelets glassy, fine, acicular (figs 5c, 13d)

spinelets glassy, fine, not acicular (figs 9c-f)

spinelets absent, or glassy, very fine, often lost (fig. 13c)

body with integument/granule cover (figs 17a, c, d)

pedicellariae always present (fig. 6c)

pedicellariae sometimes present (figs 6a, b)

pedicellariae never present

glassy convexities always present (fig. 9c)

glassy convexities never present

superomarginal plates in regular series (figs 10c, 13d)

superomarginal plates irregular (fig. lOe)

superomarginal plates absent

inferomarginals project narrowly, or not at all (fig. 16g)

inferomarginal plates loosely contiguous (fig. lOe)

inferomarginal spinelets large, sacciform (fig. lOe)

inferomarginal spinelets acicular, dense tufts (fig. 1 3d

furrow spines in longitudinal series (figs 5e, lOh, 16d)

furrow spines in vertical series (fig. 17f)

tube feet rows biserial (fig. 15b)

tube feet rows quadriserial

actinal plates in distinct oblique series (fig. 16c)

Genera of asterinid sea-stars

Table 2. List of genera and species of Asterinidae, including new synonymies and combinations, and species nomen dubium, nomen nudum, and

incertae sedis. Type species with asterisk.

Combination, this paper, or senior synonym Original combination

Anseropoda antarctica Fisher, 1940

Anseropoda aotearoa Me Knight, 1973

Anseropoda diaphana (Sladen, 1889)

Anseropoda fisheri Aziz and Jangoux, 1985

Anseropoda grandis Mortensen, 1933

Anseropoda habracantha H.L. Clark, 1923

Anseropoda insignis Fisher, 1906

Anseropoda lobiancoi (Ludwig, 1897)

Anseropoda ludovici (Koehler, 1909)

Anseropoda macropora Fisher, 1913

Anseropoda novemradiata (Bell, 1905)

Anseropoda pellucida (Alcock, 1893)

Anseropoda petaloides (Goto, 1914)

* Anseropoda placenta (Pennant, 1777)

Anseropoda rosacea (Lamarck, 1816)

Anseropoda tenuis (Goto, 1914)

Aquilonastra anomala (H.L. Clark, 1921)

Aquilonastra batheri (Goto, 1914)

Aquilonastra burtoni (Gray, 1 840)

* Aquilonastra cepheus (Muller and Troschel, 1 842)

Aquilonastra corallicola (Marsh, 1977)

Aquilonastra coronata (Martens, 1866)

Aquilonastra coronata (Martens, 1886)

Aquilonastra heteractis (H.L. Clark, 1938)

Aquilonastra iranica (Mortensen, 1940)

Aquilonastra limboonkengi (Smith, 1927)

Aquilonastra minor (Hayashi, 1974)

Aquilonastra rosea (H.L. Clark, 1938)

Aquilonastra scobinata (Livingstone, 1933)

Asterina fimbriata Perrier, 1875

Asterina gibbosa (Pennant, 1777)

*Asterina gibbosa (Pennant, 1777)

Asterina gracilispina H.L. Clark, 1923

Asterina ocellifera (Gray, 1 847)

Asterina pancerii (Gasco, 1870)

Asterina phylactica Emson and Crump, 1979

Asterina stellifera (Mobius, 1859)

incertae sedis

*Asterinides folium (Liitken, 1860)

Asterinides hartmeyeri (Doderlein, 1910)

Asterinides pilosa (Perrier, 1881)

Asterinides pompom (A.M. Clark, 1983)

Callopatiria formosa (Mortensen, 1933)

^Callopatiria granifera (Gray, 1847)

Callopatiria granifera (Gray, 1847)

Cryptasterina hystera Dartnall and Byrne, 2003

Cryptasterina pacifica (Hayashi, 1977)

Cryptasterina pentagona (Muller and Troschel, 1 842)

* Cryptasterina pentagona (Muller and Troschel, 1842)

Cryptasterina pentagona (Muller and Troschel, 1842)

antarctica, Anseropoda, Fisher, 1940

aotearoa, Anseropoda , McKnight, 1973

diaphana, Palmipes, Sladen, 1889

fisheri, Anseropoda, Aziz and Jangoux, 1985

grandis, Anseropoda (Palmipes), Mortensen, 193 3

habracantha, Anseropoda, H.L. Clark, 1923

insignis, Anseropoda, Fisher, 1906

lobiancoi, Palmipes, Ludwig, 1897

ludovici, Palmipes, Koehler, 1909

macropora, Anseropoda, Fisher, 1913

novemradiata, Palmipes, Bell, 1905

pellucida, Palmipes, Alcock, 1893

petaloides, Palmipes, Goto, 1914

placenta, Asterias, Pennant, 1777

rosaceus, Asterias, Lamarck, 1816

tenuis, Palmipes, Goto, 1914

anomala, Asterina, H.L. Clark, 1921

batheri, Asterina, Goto, 1914

burtoni, Asterina, Gray, 1840

cepheus, Asteriscus, Muller and Troschel, 1842

corallicola, Asterina, Marsh, 1977

coronata, Asterina, Martens, 1866

coronata fascicularis, Asterina, Fisher, 1918

spinigera, Asterina, Koehler, 1911

heteractis, Asterina, H.L. Clark, 1938

cephea var. iranica, Asterina, Mortensen, 1940

limboonkengi, Asterina, Smith, 1927

minor, Asterina, Hayashi, 1974

rosea, Paranepanthia, H.L. Clark, 1938

scobinata, Asterina, Livingstone, 1933

fimbriata, Asterina, Perrier, 1875

crassispina, Asterina, H.L. Clark, 1928

gibbosa, Asterias, Pennant, 1777

gracilispina, Asterina, H.L. Clark, 1923

ocellifera, Patiria, Gray, 1847

pancerii, Asteriscus, Gasco, 1870

phylactica, Asterina, Emson and Crump, 1979

siderea, Enoplopatiria, Verrill, 1913

stellifer, Asteriscus, Mobius, 1859

lorioli, Asterina, Koehler, 1910

novaezelandiae, Asterina, Perrier, 1875

folium, Asteriscus, Liitken, 1 860

hartmeyeri, Asterina, Doderlein, 1910

lymani, Asterina, Perrier, 1881

pilosa, Asterina, Perrier, 1881

pompom, Paxillasterina, A.M. Clark, 1983

formosa, Parasterina, Mortensen, 1933

bellula, Patiria, Sladen, 1889

granifera, Patiria, Gray, 1847

hystera, Cryptasterina, Dartnall and Byrne, 2003

pseudoexigua pacifica, Asterina, Hayashi, 1977

obscura, Patiriella, Dartnall, 1971

pentagonus, Asteriscus, Muller and Troschel, 1842

pseudoexigua, Patiriella, Dartnall, 1971

flexilis, Desmopatiria, Verrill, 1913

nomen nudum (this work)

P. Mark O’Loughlin and Jonathan M. Waters

*Disasterina abnormalis Perrier, 1875

Disasterina abnormalis Perrier, 1875

Disasterina ceylanica Doderlein, 1888

Disasterina longispina (H.L. Clark, 1938)

Disasterina odontacantha Liao, 1980

Disasterina spinosa Koehler, 1910

Indianastra inopinata (Livingstone, 1933)

Indianastra sarasini (de Loriol, 1 897)

Indianastra sarasini (de Loriol, 1897)

* Indianastra sarasini (de Loriol, 1897)

*Kampylaster incurvatus Koehler, 1920

^Meridiastra atyphoida (H.L. Clark, 1916)

Meridiastra calcar (Lamarck, 1816)

Meridiastra fissura O’Loughlin, 2002

Meridiastra gunnii (Gray, 1 840)

Meridiastra gunnii (Gray, 1840)

Meridiastra medius (O’Loughlin et al., 2003)

Meridiastra modesta (Verrill, 1870)

Meridiastra mortenseni (O’Loughlin et al., 2002)

Meridiastra nigranota O’Loughlin, 2002

Meridiastra occidens (O’Loughlin et al., 2003)

Meridiastra oriens (O’Loughlin et al., 2003)

Meridiastra rapa O’Loughlin, 2002

Nepanthia belcheri (Perrier, 1875)

Nepanthia crassa (Gray, 1847)

Nepanthia fisheri Rowe and Marsh, 1982

*Nepanthia maculata Gray, 1840

Nepanthia pedicellaris Fisher, 1913

incertae sedis

Paranepanthia aucklandensis (Koehler, 1920)

Paranepanthia grandis (H.L. Clark, 1928)

* Paranepanthia platydisca (Fisher, 1913)

Parvulastra calcarata (Perrier, 1869)

Parvulastra dyscrita (H.L. Clark, 1923)

* Parvulastra exigua (Lamarck, 1816)

Parvulastra parvivipara (Keough and Dartnall, 1978)

Parvulastra vivipara (Dartnall, 1969)

Patiria chilensis (Liitken, 1859)

*Patiria miniata (Brandt, 1835)

Patiria miniata (Brandt, 1835)

Patiria pectinifera (Muller and Troschel, 1842)

Patiriella inornata Livingstone, 1933

Patiriella oliveri (Benham, 1911)

Patiriella paradoxa Campbell and Rowe, 1997

* Patiriella regularis (Verrill, 1867)

nomen dubium (this work)

*Pseudasterina delicata Aziz and Jangoux, 1985

Pseudasterina granulosa Aziz and Jangoux, 1985

Pseudonepanthia briareus (Bell, 1894)

^Pseudonepanthia gotoi A.H. Clark, 1916

Pseudonepanthia gracilis (Rowe and Marsh, 1982)

Pseudonepanthia grangei (McKnight, 2001)

abnormalis, Disasterina, Perrier, 1875

pulchella, Habroporina, H.L. Clark, 1921

ceylanica, Disasterina, Doderlein, 1888

longispina, Manasterina, H.L. Clark, 1938

odontacantha, Disasterina, Liao, 1980

spinosa, Disasterina, Koehler, 1910

inopinata, Asterina, Livingstone, 1933

perplexa, Asterina, H.L. Clark, 1938

lutea, Asterina, H.L. Clark, 1938

nuda, Asterina, H.L. Clark, 1921

orthodon, Asterina, Fisher, 1922

sarasini, Palmipes, de Loriol, 1897

incurvatus, Kampylaster, Koehler, 1920

atyphoida, Asterina, H.L. Cark, 1916

calcar, Asterias, Lamarck, 1816

fissura, Meridiastra, O’Loughlin, 2002

brevispina, Patiriella, H.L. Clark, 1938

gunnii, Asterina, Gray, 1840

medius, Patiriella, O’Loughlin et al., 2003

agustincasoi, Asterina, Caso, 1977

modesta, Asterina ( Asteriscus ), Verrill, 1870

mortenseni, Patiriella, O’Loughlin et al., 2002

nigranota, Meridiastra, O’Loughlin, 2002

occidens, Patiriella, O’Loughlin et al., 2003

oriens, Patiriella, O’Loughlin et al., 2003

rapa, Meridiastra, O’Loughlin, 2002

belcheri, Asterina (Nepanthia), Perrier, 1875

crassa, Patiria, Gray, 1847

fisheri, Nepanthia, Rowe and Marsh, 1982

maculata, Nepanthia, Gray, 1840

pedicellaris, Nepanthia, Fisher, 1913

brachiata, Nepanthia, Koehler, 1910

aucklandensis, Asterina, Koehler, 1920

grandis, Nepanthia, H.L. Clark, 1928

praetermissa, Asterinopsis, Livingstone 1933

platydisca, Nepanthia, Fisher, 1913

calcarata, Asteriscus, Perrier, 1869

dyscrita, Asterina, H.L. Clark, 1923

1, Lamarck, 1816

parvivipara, Patiriella, Keough and Dartnall, 1978

vivipara, Patiriella, Dartnall, 1969

chilensis, Asterina, Liitken, 1859

coccinea, Patiria, Gray, 1840

miniata, Asterias, Brandt, 1835

pectinifera, Asteriscus, Muller and Troschel, 1 842

inornata, Patiriella, Livingstone, 1933

nigra, Patiriella, H.L. Clark, 1938

oliveri, Asterina, Benham, 1911

paradoxa, Patiriella, Campbell and Rowe, 1997

regularis, Asterina (Asteriscus), Verrill, 1867

tangribensis, Patiriella, Domantay and Acosta, 1970

delicata, Pseudasterina, Aziz and Jangoux, 1985

granulosa, Pseudasterina, Aziz and Jangoux, 1985

briareus, Patiria, Bell, 1894

gotoi, Pseudonepanthia, AM. Clark, 1916

gracilis, Nepanthia, Rowe and Marsh, 1982

grangei, Nepanthia, McKnight, 2001

Genera of asterinid sea-stars

Pseudonepanthia nigrobrunnea (Rowe and Marsh, 1982)

Pseudonepanthia reinga (McKnight, 2001)

Pseudonepanthia troughtoni (Livingstone, 1934)

*Pseudopatiria obtusa (Gray, 1847)

*Stegnaster inflatus (Hutton, 1872)

Stegnaster wesseli (Perrier, 1875)

Tegulaster alba (H.L. Clark, 1938)

^Tegulaster emburyi Livingstone, 1933

Tegulaster leptalacantha (H.L. Clark, 1916)

Tegulaster praesignis (Livingstone, 1933)

nigrobrunnea, Nepanthia, Rowe and Marsh, 1982

reinga, Nepanthia, McKnight, 2001

troughtoni, Parasterina, Livingstone, 1934

obtusa, Patiria, Gray, 1847

inflatus, Pte raster, Hutton, 1872

wesseli, Asterina, Perrier, 1875

alba, Asterina, H.L. Clark, 1938

emburyi, Tegulaster, Livingstone, 1933

leptalacantha, Asterina, H.L. Clark, 1916

leptalacantha var. africana, Disasterina, Mortensen, 1933

praesignis, Disasterina, Livingstone, 1933

spinulifera, Disasterina, H.L. Clark, 1938

*Tremaster mirabilis Verrill, 1879 mirabilis, Tre master, Verrill, 1879

Tremaster mirabilis Verrill, 1879 novaecaledoniae, Tremaster, Jangoux, 1982

sometimes in fans ( grandis , insignis, placenta)', adradial row of

actinal plates with complete series of spines; interradial actinal

spines glassy, sacciform.

Superambulacral plates absent; superactinal plates absent

( rosacea ) or present proximally in multiple plate struts (pla-

centa ); abactinal and actinal interradial plates meet extensively

internally by long thin articulating projections.

Distribution. Indo-West Pacific, South Africa, Australia, New

Zealand, Antarctic Peninsula, North-East Atlantic,

Mediterranean; 10-600 m.

Remarks. Molecular data are not available for any Anseropoda

species. This restricted generic review is based on examination

of specimens of A. placenta (type species) and A. rosacea, and

original descriptions of A. grandis by Mortensen (1933) and

A. insignis by Fisher (1906). Most of the species assigned to

Anseropoda have not been examined and their generic place-

ment has not been reviewed. The very thin body, and presence

of extensive interradial internal thin projecting articulating sup-

ports distinguish Anseropoda from all other asterinid genera.

The contrasting internal plate structures and presence or

absence of superactinal plates in A. placenta and A. rosacea

(Fig. 7) suggest that Anseropoda may not be monophyletic.

Clark and Downey (1992) regarded the very small Anseropoda

lobiancoi (Ludwig, 1897) from the Mediterranean (R about 8

mm) as being incompatible with Anseropoda.

Aquilonastra O’Loughlin gen. nov.

Figures 1 (clade IV, part), 2b, 9a-f

Diagnosis. Rays 5 (6-8 in fissiparous species); interradial

margin deeply incurved, form stellate; rays discrete, broad at

base, tapering, rounded distally; flat actinally, high convex

abactinally; abactinal plates in longitudinal series, not perpen-

dicular to margin; papulate areas extensive; papulae predomin-

antly single, large, in longitudinal series along sides of rays;

abactinal plates with glassy convexities; abactinal spinelets and

actinal spines predominantly fine, glassy, conical or sacciform

or splay-pointed sacciform, in bands or tufts, numerous (10-40

per plate); actinal plates in longitudinal, not oblique, series;

superambulacral plates present for all of ray, sometimes for part

of ray or absent in pedomorphic species; superactinal plates

present.

Type species. Asteriscus cepheus Muller and Troschel, 1842.

Other species. A. anomala (H.L. Clark, 1921); A. batheri (Goto, 1914);

A. burtoni (Gray, 1840); A. corallicola (Marsh, 1977); A. coronata

(Martens, 1866) (junior synonyms: Asterina coronata fascicularis

Fisher, 1918, possible synonymy raised by H.L. Clark, 1928, for-

malised by Rowe, 1995; and Asterina spinigera Koehler, 1911, by

VandenSpiegel et al., 1998); A. heteractis (H.L. Clark, 1938);

A. iranica (Mortensen, 1940) (raised to species status, this work);

A. limboonkengi (Smith, 1927); A. minor (Hayashi, 1974); A. rosea

(H.L. Clark, 1938); A. scobinata (Livingstone, 1933).

Material examined. A. anomala. Caroline Is, WAM Z6845 (2);

Christmas L, Z6851 (1); Lord Howe L, AM J6169 (21); NMV F95593

(6); F97690 (6); F96699 (1); Maidive Is, Z6854 (1); Samoa, F96698

(2); Torres Strait, Damley I., Z6849 (2); Western Australia, Kimberley,

Z6843 (1).

A. batheri. Japan, Toyama Bay, NMV F97441 (1); AM J11564 (2);

Oman, UF 70 (7).

A. burtoni. Gulf of Suez, AM J17892 (2); TM H1815 (2); Red Sea,

Egyptian coast, H1814(l).

A. cephea iranica. Syntype. Persian Gulf, S of Bushire, coral reef,

AM J17891 (1).

A. cepheus. Queensland, NMV F95594 (2), AM J23331 (1);

Western Australia, Abrolhos Is, WAM Z6778 (7); J8321 (5); New

Guinea, Trobriand Group, J22934 (1); Hong Kong, BMNH

1981.2.6.25 (1; previously identified as Asterina limboonkengi).

A. corallicola. Paratypes. Caroline Is, WAM Z1704 (3).

A. coronata. Northern Territory, Darwin, NMV F95796 (4); AM

J6613 (1); J8206 (2); Caroline Is, J13660 (1); Bombay, BMNH

1960.10.4.11-16 (2; previously identified as Asterina lorioli)', Taiwan,

J19956 (1); Japan, J1 1563 (2).

A. heteractis. Queensland, Townsville, AM J9541 (2); Heron I.,

J 19449 (1).

A. limboonkengi. Oman, UF 1645 (2); UF 246 (1); UF 68 (6).

A. minor. Japan, NMV F96697 (2).

Paranepanthia rosea. Paratypes. Western Australia, RottnestL, AM

J6171 (3). Other material. Western Australia, Jurien Bay, AM J7437

(4).

A. scobinata. Holotype. Tasmania, AM J1241. Other material.

Tasmania, Eaglehawk Neck, J9060 (3); Victoria, Phillip I., NMV

F72998 (1); Port Fairy, F72985 (2); Killarney, F72997 (10).

P. Mark O’Loughlin and Jonathan M. Waters

Figure 7. Longitudinal sections through interradii of Anseropoda species, showing internal contiguous projections from abactinal and actinal

plates, and absence (b) of superambulacral plates, a, Anseropoda placenta (R = 56 mm, NMV F98043). b, Anseropoda rosacea (R = 82 mm, NMV

F95811).

Description with species variations. Rays predominantly 5, or

6 (fissiparous corallicola, heteractis ), or 6-7 (fissiparous

burtoni ), or 7-8 (fissiparous anomala)-, interradial margin

deeply incurved, form stellate; rays discrete, broad at base,

tapering, rounded distally; flat actinally, high convex abactin-

ally; size medium ( iranica up to R = 35 mm) to small (pedo-

morphic species anomala, burtoni, corallicola, heteractis,

minor with R up to 10 mm); pedicellariae sometimes present,

scattered ( iranica ) or few on proximal lateral rays above

papulae ( batheri , coronata)-, gonopores abactinal or actinal

( corallicola , scobinata)\ fissiparity in 4 species.

Abactinal surface even, or very uneven with some paxilli-

form plates ( coronata , rosea)-, plates in longitudinal series, not

series perpendicular to margin; papulate areas extensive, to

near distal end of rays, to near interradial margin; papular

spaces small with few secondary plates and predominantly 1

large papula (rarely up to 3); 2-6 distinct longitudinal series of

large papulae along each side of rays, series variably evident

carinally; disc variably bordered by continuous series of 5

radial and 5 interradial plates (not in fissiparous species);

abactinal plates predominantly singly notched for papulae,

crescentiform on rays and interradially to near margin, round

distally, with transverse or round spinelet-bearing proximal ele-

vations; carinal series of plates absent, or proximal series of

singly or doubly notched plates not widely separated by sec-

ondary plates, or doubly papulate to end of ray ( heteractis );

cleared abactinal plates with glassy convexities; abactinal

spinelets glassy, conical or sacciform or splay-pointed sacci-

form; abactinal spinelets in transverse bands or tufts of 8-40

spinelets per plate, tufts sometimes multiple per plate {batheri,

cepheus), spinelets sometimes 2 distinct forms (coronata),

spinelets sometimes paxilliform (coronata, rosea)-, marginal

plates with abactinal spinelets; superomarginal plates not

larger than inferomarginals; acute margin formed by project-

ing inferomarginal plates (rounded margin in pedomorphic

corallicola).

Actinal plates in longitudinal series, sometimes also oblique

(coronata), or oblique (pedomorphic heteractis).

Actinal spines per plate: oral 5-9; suboral 3-12; furrow

3-9 proximally; subambulacral 2 up to tuft; adradial actinal

plates fully spinous; actinal interradial spines similarly var-

iable forms as abactinally, predominantly in tufts, up to 20

per plate.

Superambulacral plates present for all of ray, or distal half

(pedomorphic minor), or absent (pedomorphic corallicola)-,

superactinal plates present.

Distribution. Eastern Mediterranean, northern Indian and west-

ern Pacific Oceans, Gulf of Suez, Red Sea, Persian Gulf,

Arabian Sea, southern Australia, Japan; 0-15 m.

Etymology. From the Latin aquilonalis (northern) and astrum

(star), referring to the predominantly northern Indo-Pacific

occurrence of the species of this genus (feminine).

Remarks. The new genus Aquilonastra is represented on the

molecular clade of Waters et al. (2004, clade IV, part) by six

species: A. anomala, A. batheri, A. burtoni, A. coronata,

A. minor, A. scobinata. These share a consistent morphology:

discrete high rays; series of single large papulae along rays;

fine glassy spinelets; presence of superambulacral and super-

actinal plates. Other species share this morphology and are

included in the genus: cepheus, corallicola, heteractis, lim-

boonkengi and rosea. The new genus shares some characters

with Cryptasterina Dartnall et al., 2003: extensive papulate

areas; small papular spaces with few secondary plates and

single papulae per space; up to four longitudinal series of

papulae along each side of the rays; doubly papulate carinal

plates variably present, as short or long series or absent; margin

formed by projecting inferomarginal plates; superactinal plates

support the interradial margin. This similarity is supported by a

sister taxon relationship on the molecular tree (clade IV, part).

However, there are significant morphological differences

Genera of asterinid sea-stars

exhibited by Cryptasterina : form pentagonal or subpentagonal;

abactinal spinelets granuliform; at most very low raised

rounded spinelet-bearing elevations along the proximal project-

ing edge of abactinal plates; superomarginal plates larger

than inferomarginals; actinal spines fewer (oral 4-5, suboral

1-2, furrow 2-3, subambulacral 1-2 proximally, actinal

interradial 0-1 proximally 1-2 distally), conical to digitiform;

adradial actinal series of plates with none to few spines; super-

ambulacral plates not in complete series, few present.

In an electrophoretic study of five Asterina species from

Japan, Matsuoka (1981) found them to be a heterogeneous phy-

logenetic group with A. batheri and A. coronata japonica

closely related, A. pseudoexigua pacifica fairly closely related,

and A. pectinifera and A. minor each probably belonging to

separate lineages. This result is generally congruent with the

molecular phylogeny of Waters et al. (2004), except that in their

clades A. minor was not separate but in the same lineage as

A. batheri and A. coronata.

H.L. Clark (1938) described juvenile specimens of

Paranepanthia rosea (up to R = 8 mm). Material examined in

this work from Jurien Bay was significantly larger (up to

R = 17 mm) and characterised by: absence of two distinct

‘fields’ of abactinal plates; splay-pointed sacciform spinelets

and spines; predominantly large single papulae in papular

spaces; numerous longitudinal series of papulae along the sides

of rays; actinal plates not in oblique series; presence of series

of superambulacral plates; and single superactinal plate supports.

P. rosea is removed from Paranepanthia. The inferomarginal

plates define the margin in P. rosea, not superomarginal plates

as stated by H.L. Clark (1938).

Mortensen (1940) distinguished a variety iranica for the

species A. cepheus on the basis of having 5-6 suboral spines

per plate and A. cepheus two (up to four seen in this work).

Further to this observation, the variety is distinguished from

A. cepheus (characters in brackets) by having: up to eight stout

sacciform spinelets in single series across proximal abactinal

plates (up to 20 fine spinelets in bands across plates); up to ten

stout inferomarginal spinelets per plate (more than, 20);

typically two actinal spines per plate (4-8). Asterina cepheus

iranica is raised to species status.

Marsh (1977) compared the three similar small fissiparous

Indo-Pacific asterinids A. anomala, A corallicola and A. heter-

actis, and her confirmation of valid species status for all three

is supported here. A. heteractis has some characters which are

exceptional to those shared by other Aquilonastra species: reg-

ular doubly papulate carinal series for length of ray; actinal

plates in oblique series; fewer actinal spines per plate, with up

to four oral, sometimes no suboral, up to four furrow, up to

three actinal. These differences are not sufficient to justify

assignment to another genus.

Smith (1927), Liao (1980) and Liao and Clark (1995)

observed that A. limboonkengi was close to A. cepheus (Smith

as close to A. burtoni from the Philippines, A. burtoni presum-

ably A. cepheus ). Liao and Clark (1995) described the differ-

ences between A. cepheus and A. limboonkengi. Our observa-

tions confirm that the abactinal spinelets in A. cepheus are long

and fine, and those in A. limboonkengi short and stout. Rowe

(1995) rejected the subspecies A. coronata fascicularis Fisher,

1918. The possible species status of the other subspecies of

A. coronata (< cristata Fisher, 1916 and euerces Fisher, 1917)

has not been examined.

Pairs of abactinal spinelets on A. anomala, A. cepheus,

A. corallicola and A. minor are sometimes angled and contigu-

ous, suggesting that they may act as pedicellariae. But the

spinelets are not differentiated in form.

Asterina Nardo

Figures 1 (clade VI), 2c, 6a, 8c-e

Asterina Nardo, 1834:716. — Clark and Downey, 1992: 177-181. —

A.M. Clark, 1993: 206-214.— Liao and Clark, 1995: 129.— Rowe,

1995: 33-35.— O’Loughlin, 2002: 278.— Waters et al., 2004: 874, 875,

877 (part). (For complete synonymy and discussion see Clark and

Downey, 1992).

Allopatiria Verrill, 1913: 480. — Verrill, 1914: 273. — Tortonese,

1965: 174.— Spencer and Wright, 1966: U69.— A.M. Clark, 1983:

372-373.— Clark and Downey, 1992: 172.— A.M. Clark, 1993: 204.

New synonym.

Diagnosis. Rays 5; body integument noticeable; interradial

margin incurved to varying extents; rays discrete, short, dis-

tally broadly or narrowly rounded or pointed; flat actinally,

convex abactinally; papular spaces with numerous small

papulae and secondary plates; abactinal spinelets opaque, digi-

tiform to conical, rounded distally; actinal plates in oblique and

sometimes longitudinal series; actinal interradial spines digiti-

form to conical, up to 5 per plate; internal contiguous projec-

tions from abactinal and actinal plates support margin; lacking

superambulacral and superactinal plates.

Type species. Asterias gibbosa Pennant, 1777 (subsequent des-

ignation by Fisher, 1906; full treatment by A.M. Clark, 1983)

(junior synonym: Asterina crassispina H.L. Clark, 1928, by

Rowe, 1995).

Other species. A. fimbriata Perrier, 1875; A. gracilispina H.L. Clark,

1923; A. ocellifera (Gray, 1847); A. pancerii (Gasco, 1870); A. phy-

lactica Emson and Crump, 1979; A. stellifera (Mobius, 1859) (junior

synonym: Enoplopatiria siderea Verrill, 1913, this work).

Material examined. A. fimbriata. Argentina, Comodoro Rivadavia, AM

J6411 (3).

A. gibbosa. Mediterranean, Tunisia, USNM E42476 (1); see

O’Loughlin (2002).

A. gracilispina. South Africa, Cape Agulhas, BMNH 1975.10.29.47

( 1 ).

A. ocellifera. Mauritania, Cape Lever, BMNH 1969.12.16.13 (1).

A. stellifer. Argentina, AM J6410 (1).

Enoplopatiria siderea. Holotype. Panama (assumed here to be

mistaken locality), YPM No. 9830. New synonym.

Description with species variations. Rays 5; interradial margin

incurved to varying extents, rays discrete, distally narrowly

( gibbosa ) or broadly (fimbriata, gracilispina) rounded, or

pointed (ocellifera, stellifera ); size large (ocellifera up to

R = 81 mm) to small (phylactica up to R = 7.5 mm); pedicel-

lariae numerous abactinally, predominantly 2 valves, fasciculate

(gibbosa, ocellifera, stellifera), or absent (fimbriata , gracil-

ispina); gonopores abactinal (fimbriata , gracilispina, ocellifera,

stellifera ) or actinal (gibbosa, phylactica)', not fissiparous.

P. Mark O’Loughlin and Jonathan M. Waters

Extensive abactinal papulate area of weakly notched or shal-

low crescentiform or rhombic ( ocellifera ) plates, narrow inter-

radial non-papulate margin of proximally rounded plates; upper

rays with variably developed carinal series of doubly notched

plates (fimbriata, gibbosa, gracilispina ), or irregular plates

(stellifera), or rhombic plates ( ocellifera ); up to 4 longitudinal

series of plates along each side of rays; papular spaces large,

secondary plates and papulae numerous, small secondary plates

may partly obscure ( ocellifera ) or encroach on ( stellifera )

primary plates; disc bordered by series of 5 radial and 5 inter-

radial plates (gibbosa, gracilispina) or not (fimbriata, ocellif-

era, stellifera ); abactinal plates with spinelet-bearing ridges and

domes; denuded plates with glassy convexities (gibbosa,

gracilispina, ocellifera, stellifera) or reticulations (fimbriata );

abactinal spinelets digitiform or conical, rounded dis-

tally; spinelets spaced on plates closely (stellifera) or widely

(fimbriata, gibbosa, gracilispina, ocellifera), not in tufts; inter-

radial plates in short series perpendicular to margin (ocellifera,

stellifera) or not (fimbriata, gibbosa, gracilispina)', supero-

marginal and inferomarginal plates in distinct series, covered

with abactinal-type spinelets; projecting inferomarginal plates

form marginal edge.

Actinal plates in oblique series (small specimens of A.

gibbosa; longitudinal series in large specimens of A. gibbosa).

Actinal spines per plate: oral 4-9; suboral 1-4; furrow 2-5

proximally, webbed; subambulacral 1-3; adradial row of

actinal plates with complete series of spines; actinal interradial

predominantly 2-3 in mid-interradius; interradial actinal spines

digitiform to conical.

Lacking superambulacral and superactinal plates; inter-

radial margin supported by contiguous internal tongue-like

extensions of abactinal and actinal plates.

Distribution. Atlantic Ocean, Europe, SW and S Africa, SE and

S South America; Mediterranean Sea; 0-250 m.

Remarks. The principal characters in this new diagnosis of

Asterina are: papular spaces with numerous small papulae and

secondary plates, not predominantly single large papulae;

abactinal spinelets opaque, digitiform to conical, not glassy and

fine; actinal plates in oblique and longitudinal series; internal

contiguous projections from abactinal and actinal plates sup-

port the margin; lacking superambulacral and superactinal

plates.

Molecular evidence (Waters et al., 2004, clade VI) places

A. phylactica as sister species of A. gibbosa. On the basis of the

diagnosis the following species are removed from Asterina to

other genera: A. alba, A. anomala, A. aucklandensis, A. batheri,

A. burtoni, A. cepheus, A. chilensis, A. corallicola, A. coronata,

A. hartmeyeri, A. heteractis, A. inopinata, A. miniata, A. minor,

A. pectinifera, A. sarasini, A. scobinata.

A.M. Clark (1983) considered that the rhombic and non-

crescentic form of the large primary plates in the radial field,

and the proliferation of small secondary plates to the extent that

abactinal plates become obscured peripherally, distinguish

Allopatiria from Asterina. Clark and Downey (1992) noted

superficial similarity of similarly sized specimens of

Allopatiria and Asterina. We have found that the form of

abactinal plates and number of secondary plates are not

generically significant. In diagnosing Allopatiria, Clark and

Downey (1992) referred to “additional internal plates” support-

ing the margin. If such superactinal plates were present they

might justify the retention of Allopatiria. But we did not con-

firm their presence. What could appear to be superactinal plates

are long internal projections of distal abactinal and actinal

plates. This is a character shared with Asterina, and Allopatiria

is judged to be a junior synonym. A.M. Clark (1993) listed

Patiria Gray, 1840 and Asterinides Verrill, 1913 as junior syn-

onyms of Asterina. Rowe (1995) and Campbell and Rowe

(1997) supported the validity of Asterinides. Patiria is raised

out of synonymy in this work.

O’Hara (1998) synonymised Asterina hamiltoni Koehler,

1920 and Cycethra macquariensis Koehler, 1920 with Asterina

frigida Koehler, 1917 and assigned A. frigida to Cycethra Bell,

1881 in the Ganeriidae. Rowe (1995) synonymised Asterina

crassispina H.L. Clark, 1928 with Asterina gibbosa, and judged

the northern Australia locality data to be incorrect. Based on an

examination of the type of Asterina spinigera, VandenSpiegel

et al. (1998) synonymised it with Asterina coronata.

Clark and Downey (1992) summarised the characters

thought to distinguish A. pancerii from A. gibbosa as: flatter

and near pentagonal shape; fewer and larger skeletal plates;

plates with coarse crystal bodies; and more numerous, usually

three, suboral spines. O’Loughlin (2002) noted in A. gibbosa

material: some conspicuous glassy convexities around the

margin of abactinal plates; and some plates with up to three

suboral spines. The near pentagonal body form and relative size

of plates have been found to show variation within species in

this study. A. pancerii is probably conspecific with A. gibbosa.

In the absence of molecular evidence, pancerii is retained here

as a species of Asterina.

The holotype of Enoplopatiria siderea was examined and

found to conform in all morphological features with Asterina

stellifera. The Panama type locality must be an error, since

A. stellifera occurs on the east and west coasts of the South

Atlantic.

Specimens of Asterina lorioli Koehler, 1910 and Asterina

novaezelandiae Perrier, 1875 were not examined. Based on the

descriptions, assignment to Asterina could not be confirmed.

These species are placed incertae sedis, and discussed below.

All species placed here in Asterina are from the Atlantic and

Mediterranean.

Asterinides Verrill

Figures 2d, 8f-i

Asterinides Verrill, 1913: 479.—' Verrill, 1914: 263.— Verrill, 1915:

58.— Tommasi, 1970: 14-15.— A.M. Clark, 1983: 364.— Rowe, 1995:

33. — Campbell and Rowe, 1997: 131. — Clark and Mah, 2001: 335. —

O’Loughlin, 2002: 291, 293.

Paxillasterina A.M. Clark, 1983: 373. — Clark and Downey, 1992:

193. — A.M. Clark, 1993: 227. New synonym.

Diagnosis. Rays 5 or 6; form subpentagonal or subhexagonal or

discrete rays; abactinal plates in longitudinal series; extensive

papulate areas, predominantly single papulae; abactinal

plates singly notched; abactinal spinelets glassy, acicular to

Genera of asterinid sea-stars

subsacciform, in tufts or subpaxilliform or on paxilliform

columns; lacking pedicellariae; superomarginal plates with tuft

of spinelets; inferomarginal plates with distal dense subpaxilli-

form tuft of larger spinelets; actinal spines subsacciform, thin;

lacking superambulacral and superactinal plates, distal inter-

radial margin supported by inward projecting tongues from

abactinal and sometimes actinal plates.

Type species. Asteriscus folium Liitken, 1860 (original

designation).

Other species. A. hartmeyeri (Doderlein, 1910); A. pilosa (Perrier,

1881) (junior synonym: A. lymani Perrier, 1881, by Clark and Downey,

1992); A. pompom (A.M. Clark, 1983).

Material examined. A. folium. Holotype. Atlantic Ocean, Virgin Is

(ZMUC). Other material. Atlantic Ocean, Caribbean Sea, USNM

E28573 (1); Gulf of Mexico, USNM 38811 (3); Bermuda, USNM

38236 (4).

A. hartmeyeri. Atlantic Ocean, Caribbean Sea, USNM E49050 (2).

A. pilosa. Atlantic Ocean, Caribbean Sea, USNM E17973 (2); Gulf

of Mexico, USNM E13707 (1); Venezuela, USNM E28411 (1).

A. pompom. Atlantic Ocean, Caribbean Sea, USNM E47755 (4);

E47866 (1).

Description with species variations. Rays predominantly 5

(folium, hartmeyeri, pompom) or 6 (pilosa ), interradial margin

straight to slightly to deeply (pilosa ) incurved, subpentagonal

or discrete rays (pilosa ); body thick, sides of rays steep (folium,

hartmeyeri, pilosa ), or thin ( pompom ); size small (folium up to

R = 20 mm) to very small (hartmeyeri up to R = 7 mm); lack-

ing pedicellariae; gonopores abactinal; one fissiparous (pilosa,

R up to 10 mm).

Abactinal surface uneven due to raised proximal edge of

thick imbricating plates (folium ) or paxilliform columns

( pompom ); abactinal plates in longitudinal series; denuded

appearance dominated by singly notched plates, heart-shaped

to crescentiform; papulate areas extensive; papulae pre-

dominantly single, large, up to 4 longitudinal series along each

side of rays; doubly notched and papulate carinal series

variably present for most of ray, or upper ray plates irregular

(pilosa)-, papular spaces fairly large, sometimes up to 3 papulae

and 3 secondary plates per space; disc bordered by series of

5 radial 5 interradial plates (folium ), or not (hartmeyeri,

pilosa)-, denuded plates with glassy convexities (folium , hart-

meyeri) or irregular texturing (pompom,)-, plates with apical

ridge or dome; spination variable, up to 3 subpaxilliform tufts

across each plate proximally, each tuft with up to 20 spine-

lets, single subpaxilliform tufts distally (folium), or tufts of

spinelets (hartmeyeri, pilosa), or soft paxillar columns and

rounded domes, each with up to 50 spinelets (pompom );

spinelets glassy, acicular to subsacciform; superomarginal

plates with central tuft or pompom of spinelets; inferomarginal

series projects to form margin, plates bare proximally, promi-

nent marginal tuft of larger spinelets, sometimes on elevations

(folium, pompom).

Actinal interradial plates in longitudinal series (folium ,

pompom), sometimes oblique (folium), or oblique (hartmeyeri).

Actinal spines per plate: oral 4-7; suboral 1-10; furrow 3-5,

webbed; subambulacral 2-4 or clusters (pilosa ); adradial acti-

nal plates with complete series of spines; actinal 1-4 or clusters

(pilosa), webbed if more than 1; actinal interradial spines

subsacciform, thin.

Lacking superambulacral and superactinal plates; distal

interradial margin supported by internal contiguous tongue-like

projections from abactinal and actinal plates (folium, pompom)

or from abactinal plates only (hartmeyeri).

Distribution. W Atlantic, Florida, Bahamas, West Indies,

Caribbean, Brazil; 0-256 m.

Remarks. Molecular data are not available for any species of

Asterinides, and this review is based on morphology. Rowe

(1995) and Rowe and Campbell (1997) considered Asterinides

to be a valid genus. Verrill (1913) referred Asteriscus folium

Liitken, Asteriscus cepheus Muller and Troschel, 1842 and

Asterina (Asteriscus) modesta Verrill, 1870 to his new genus

Asterinides. O’Loughlin (2002) removed A. modesta to his new

genus Meridiastra. A. cepheus is removed here from

Asterinides to Aquilonastra. The genera Aquilonastra and

Asterinides share many morphological similarities, such as the

multiple spinelet tufts per plate in A. batheri (A. folium also)

and paxillar colu mn s in A. coronata (A. pompom also). But the

generic morphological difference evident in this study is the

presence of superambulacral and superactinal internal skeletal

plates in Aquilonastra and absence in Asterinides.

A.M. Clark (1983) considered the paxillar columns on many

abactinal plates of P. pompom to be of generic significance, but

Clark and Downey (1992) noted some similarity between

A. folium and P. pompom. Both papers considered the paxillar

columns to be otherwise unknown among Asterinidae but

Aquilonastra coronata has similar columns. Their presence in

two otherwise well-defined genera, Asterinides and

Aquilonastra argues against use of paxillar columns as a

generic character. Paxillasterina is placed in synonymy with

Asterinides.

The description of A. folium by O’Loughlin (2002) was of

specimens up to R = 15 mm, and is superceded by the descrip-

tive details here based on a specimen with R = 19 mm (USNM

E28573). The earlier description referred to ad-disc interradial

papular spaces, each with two lateral papulae separated by one

or two secondary plates. In the larger specimen these spaces

have up to three papulae and three secondary plates. Multiple

spinelet tufts are evident on the abactinal plates of the larger

specimen, but were not evident on smaller specimens. Clark

and Downey (1992) discussed differences between A. folium

and A. hartmeyeri recognising separate but similar species.

Asterina hartmeyeri also belongs to Asterinides. Asterinopsis

pilosa (Perrier, 1881) is assigned here to Asterinides since it

shares the generic diagnostic morphological characters.

Callopatiria Verrill

Figures 2e, lOa-d

Callopatiria Verrill, 1913: 480.— A.M. Clark, 1983: 367-372.—

Clark and Downey, 1992: 190-191.— AM. Clark, 1993: 217.

Diagnosis. Rays 5, discrete, narrow basally, long rayed stellate;

flat actinally, high convex abactinally, sides of rays close t

o perpendicular above angular margin; abactinal plate

P. Mark O’Loughlin and Jonathan M. Waters

arrangement irregular on upper rays, plates crescentiform;

plates closely covered by numerous glassy spinelets, digitiform

to sacciform on primary plates, thin conical pointed on second-

ary plates; lacking pedicellariae; papular spaces large, numer-

ous papulae and secondary plates; inferomarginal series of

plates project only slightly; actinal plates in oblique series;

actinal spines digitiform, numerous per plate; interior

resinous body lining; series of irregular superambulacral plates;

superactinal plates fill interradial angular margin.

Type species. Patiria bellula Sladen, 1889 (original designa-

tion) (junior synonym of Patiria granifera Gray, 1847, by A.M.

Clark, 1956).

Other species. C. formosa (Mortensen, 1933).

Material examined. C. granifera. South Africa, Western Cape

Province, NMV F98049 (1).

Description with species variations. Rays 5, interradial margin

deeply incurved, rounded proximally, rays discrete, narrow

base tapering to rounded end, form long-rayed stellate; body

flat actinally, wider actinally than breadth of upper ray, high

convex abactinally, sides of rays close to perpendicular above

angular margin; size medium, up to R = 60 mm ( granifera );

lacking pedicellariae; not fissiparous.

Abactinal surface coarse, irregularly arranged crescentiform

plates only ( granifera ); sometimes also enlarged rounded plates

distally, mostly bare of spinelets (formosa ); lacking carinal

series of plates; papulate areas extensive; papular spaces large,

not clearly bordered, up to about 10 large to small secondary

plates and up to about 10 papulae per space; abactinal plates

irregularly notched for papulae, each plate with low rounded

elevation, some subpaxilliform, closely covered with up to

about 60 spinelets, glassy, digitiform to sacciform on primary

plates, thin conical pointed on secondary plates; bare plate sur-

face with large glassy convexities; sometimes longitudinal series

of plates and papulae evident on sides of rays; series of large

subequal superomarginal and inferomarginal plates, covered

closely with digitiform spinelets, not in marginal tufts; infero-

marginals projecting only slightly at almost right-angular margin.

Actinal plates in oblique series.

Actinal spines per plate: oral 6; suboral 5-6; furrow 4 prox-

imally; subambulacral 3-4; actinal interradial up to 10 (fans

proximally, clusters distally); adradial actinal plates with com-

plete series of spines; actinal plates with spine-bearing low

domes; interradial spines digitiform.

Superambulacral plates present as irregular series, variable

size, sometimes paired or absent, contiguous with superactinals

for most of ray; marginal angle filled with numerous super-

actinal plates; abactinal and actinal plates near margin lacking

interior projections, meet at angle; superambulacral and super-

actinal plates embedded in resinous interior lining.

Distribution. Southern Africa; 0-82 m.

Remarks. Molecular data are not available for either species of

Callopatiria, and this review is based on a morphological

examination of a specimen of C. granifera. and the description

of C. formosa by Clark and Downey (1992). Patiria bellula

Sladen, 1889 was designated by Verrill (1913) as type species

of his new genus but it was subsequently assigned to

Parasterina by H.L. Clark (1923) and Mortensen (1933).

However, Parasterina is a junior synonym of Nepanthia.

Patiria granifera Gray, 1847 was reassigned to Asterina by

Perrier (1875), H.L. Clark (1923) and Mortensen (1933).

Mortensen (1933) thought it highly probable that P. bellula and

P. granifera were synonyms and A.M. Clark (1956) was more

confident. Fisher (1940, 1941) rejected Callopatiria as a junior

synonym of Patiria but it was restored to generic status by A.M.

Clark (1983), who noted similarities between Callopatiria and

Nepanthia.

Callopatiria, Nepanthia, Pseudonepanthia and Pseudo-

patiria are unusual asterinid genera in having discrete narrow

rays which are rounded abactinally. They also have irregularly

arranged abactinal plates on the upper rays and narrowly or not

projecting inferomarginal plates, characters shared with some

other genera. Callopatiria is separated diagnostically by having

rays that are not subcylindrical, with a flat actinal surface signifi-

cantly wider at mid-ray than upper ray breadth, and by a

marginal angle supported internally by many superactinal plates.

Cryptasterina Dartnall, Byrne, Collins and Hart

Figures 1 (clade IV, part), 2f, 11a

Cryptasterina Dartnall et al., 2003: 3-4.

Diagnosis. Rays 5, form pentagonal to short rays rounded dis-

tally; body with noticeable integument; low convex abactin-

ally; abactinal plates in longitudinal series, deeply notched for

papulae, many U-shaped more than crescentiform; longitudinal

series of predominantly single papulae along sides of rays; non-

papulate interradial areas extensive; few (1-3) papulae per

papular space; abactinal spinelets granuliform, spaced over

plates; lacking pedicellariae; lacking glassy convexities; super-

omarginal series prominent, longitudinally subrectangular,

inferomarginals predominantly project narrowly; non-plated

actinal area distal to oral plates frequently present, not exten-

sive; actinal interradial plates in oblique series; actinal spines

short conical, predominantly 1 per plate, rarely 2-3 distally;

very small superambulacral plates few, never as series; super-

actinal plates present, sometimes in multiple plate struts.

Type species. Asteriscus pentagonus Muller and Troschel, 1842

(original designation) (junior synonyms: Patiriella pseudo-

exigua Dartnall, 1971 and Patiriella obscura Dartnall, 1971, by

Dartnall et al., 2003).

Other species. C. hystera Dartnall and Byrne, 2003; C. pacifica

(Hayashi, 1977).

Material examined. C. hystera. Paratypes. Queensland, Yeppoon

(Statue Bay), NMV F96255 (3). Other material. Statue Bay, F98457

(2); F97057 (6).

C. pentagona. Queensland, Cairns, NMV F96702 (2); Townsville,

F97056 (2); F98452 (8); F98453 (1); Bowen (Airlie Beach), F97055

(4), F98454 (2); Mackay, F95959 (7); Hervey Bay, F98456 (2);

Kurrimine, F95961 (2); Bingil Bay, F98455 (2).

Description with species variations. Rays 5 (rarely 4 or 6);

interradial margin straight to slightly incurved, form pen-

tagonal to short tapered rays rounded distally; body with

Genera of asterinid sea-stars

noticeable integument; flat actinally, low convex abactinally,

sides of rays not steep; size small (pentagona up to R = 17 mm;

hystera up to R = 12 mm); lacking pedicellariae; lacking

actinal gonopores; not fissiparous.

Abactinal surface even; plates low, imbricate, deeply

notched for papulae, many U-shaped more than crescentiform;

plates in longitudinal series; papular spaces fairly large; papu-

lae predominantly single, large (pentagona) or small (hystera),

up to 3 per space; secondary plates 0-3 per papular space;

carinal series of doubly or singly notched plates variably pres-

ent; longitudinal series of papular spaces along sides of rays;

disc variably bordered by 5 radial and 5 smaller interradial

plates; abactinal spinelets granuliform, short columnar, spaced

over plates; denuded abactinal plates with reticulate

glassy ridges, not convexities; marginal plates with abactinal-

type spinelets, slightly thinner and longer on infero-

marginals; superomarginal series distinctive, longitudinally

subrectangular; inferomarginal plates less conspicuous,

predominantly project narrowly to form margin.

Actinal interradial plates in oblique series; non-plated

actinal area distal to oral plates variably present, not extensive.

Actinal spines per plate: oral 5-6 (proximal separated from

distal frequently in hystera)', suboral 1-2; furrow 2-3 proxi-

mally; subambulacral 1; adradial actinal 0-1; actinal inter-

radial predominantly 1 (rarely 2-3 distally); actinal spines short

conical.

Small superambulacral plates few, especially distally, lack-

ing series; superactinal plates present, sometimes in multiple

plate struts.

Distribution. Tropical Indo-Pacific; NE Australia, Indonesia,

Philippines, Solomon Is, Taiwan, Japan; littoral and shallow

sublittoral.

Remarks. In the molecular clade IV of Waters et al. (2004),

the label P pseudoexigua (Japan) refers to C. pacifica and

P. pseudoexigua (Australia) to C. pentagona (following

Dartnall et al., 2003). These sister taxa are on a clade separate

from Aquilonastra. Hart et al. (2003) added two additional line-

ages to this clade (C. hystera and an undescribed species from

Taiwan). Dartnall et al. (2003) erected Cryptasterina and pro-

vided a morphological diagnosis for the three species. The

principal diagnostic characters of Cryptasterina seen here are:

integument cover; pentagonal to short-rayed form; longitudinal

series of plates on rays; granuliform spinelets; predominantly

single papulae per space; absence of glassy convexities; promi-

nent longitudinally elongate superomarginal plates; narrowly

projecting inferomarginal plates; actinal interradial spines pre-

dominantly single in mid-ray; some superambulacral plates

present; superactinal plates present. The similarities and differ-

ences for Cryptasterina and Aquilonastra were discussed under

the latter.

Disasterina Perrier

Figures 2g, 5f, lOe-f, 12a-b

Disasterina Perrier, 1875: 289. — Livingstone, 1933: 5-7. — H.L.

Clark, 1946: 138.— Spencer and Wright, 1966: U69. — A.M. Clark,

1993: 218.— Liao and Clark, 1995: 131-132.— Rowe, 1995: 35.

Manasterina H.L. Clark, 1938: 157-158.— H.L. Clark, 1946:

139. — A.M. Clark, 1993: 220. — Rowe, 1995: 36. New synonym.

Diagnosis. Rays 5; body thin, covered by thick integument;

form medium to long-rayed stellate, rays with wide base, taper-

ing strongly, flat actinally, height low convex; abactinal plates

thin, on broad upper rays predominantly irregular in shape, size

and arrangement, loosely imbricate or contiguous or not con-

tiguous leaving non-plated spaces; plates on lower rays in few

irregular longitudinal series, sometimes weakly notched and

crescentiform; interradii thin distally, small non-papulate plates

in perpendicular or zig-zag series to margin; papulae few, irreg-

ular on rays; abactinal spinelets lacking or rare or few, glassy,

sacciform, short or long; superomarginal plates small, not in

distinct series; inferomarginal plates project widely, loosely

contiguous, with distal fringe of few large sacciform spinelets;

actinal interradial plates in oblique series; proximal actinal

interradial areas frequently not plated; interradial actinal plates

with 1-2 long sacciform spines; superambulacral plates irregu-

larly present mid-ray and distally, lacking complete series;

superactinal plates present as single plate struts.

Type species. Disasterina abnormalis Perrier, 1875 (monotypy)

(junior synonym: Habroporina pulchella H.L. Clark, 1921, by

Livingstone, 1933).

Other species. D. ceylanica Doderlein, 1888; D. longispina (H.L.

Clark, 1938); D. odontacantha Liao, 1980; D. spinosa Koehler, 1910.

Material examined. D. abnormalis. New Caledonia, AM J5042;

Queensland, Cairns, J4947 (2); Hayman I., J5699 (1); J5904 (4); J5957

(1); J7315 (1); Torres Strait, Murray I., Z6748 (1); N Western

Australia, Rowley Shoals, WAM Z6749 (2); Scott Reef, Z6753-5 (3).

D. longispina. Western Australia, Houtman Abrolhos, WAM Z6758

(1); Z6760 (2).

D. odontacantha. Fiji, UF 2391 (1); UF 1116 (1); UF, 1874 (1);

Guam, UF 1253 (1).

Description with species variations. Body thin, rays 5 (abnor-

malis 5-6); interradial margin deeply incurved, concave or

acute proximally, form medium to long-rayed stellate; rays

with wide base, tapering to narrowly or widely rounded end;

body flat actinally, low convex abactinally; distal interradii

wide and thin (abnormalis), or narrow and thin, rays rise dis-

tinctly from thin margin; body covered with thick integument;

size medium (abnormalis up to R = 38 mm) to small (spinosa

up to R = 14 mm); pedicellariae possibly present (longispina)',

none fissiparous.

Abactinal appearance irregular; abactinal plates on broad

upper rays thin, irregular in form, size, arrangement; plates not

distinguishable as primary and secondary, intergrade from large

to small; plates weakly imbricating or contiguous or not con-

tiguous leaving non-plated spaces on rays; lacking carinal

series; plates on lower rays in few irregular longitudinal series;

distal thin interradii with small rounded non-papulate plates in

irregular series perpendicular or zig-zag to margin; plates on

rays sometimes weakly notched for papulae and crescentiform

(spinosa)', papulae on rays only, single, few and sparse, large or

small, irregular; disc not regularly bordered (abnormalis) or

almost regular border (longispina, odontacantha)', low domes

sometimes present on abactinal plates (longispina)', abactinal

P. Mark O’Loughlin and Jonathan M. Waters

plates frequently bare, or with few sacciform spinelets around

disc ( abnormalis ), or sometimes with 1-2 spinelets, short prox-

imally, long distally ( odontacantha ), or with 1-4 spinelets vari-

able in form from digitiform to long, pointed or rounded, prin-

cipally on ends of rays ( longispina ), or with 1-3 long sacciform

spinelets proximally and on rays ( spinosa ); clusters of 2^4-

undifferentiated spinelets possibly act as pedicellariae inter-

radially ( longispina ); glassy convexities present on abactinal

plates ( abnormalis , longispina, odontacantha)’, superomarginal

plates small, not in distinct series, bare ( abnormalis ,

longispina ) or with 1 ( odontacantha ) or few long spinelets

(, spinosa ); inferomarginal plates project prominently, some

elongate and almost entirely projecting ( abnormalis ,

longispina, odontacantha), each plate with transverse distal

fringe of 1-6 integument-covered long sacciform spinelets.

Actinal interradial plates in oblique series; proximal actinal

interradial areas frequently not plated.

Actinal spines per plate: oral 4-8; suboral 1-3 ( odontacan-

tha 0); furrow 3-6 proximally, webbed; subambulacral 1-3;

adradial bare ( abnormalis , odontacantha) or with spines;

actinal 1 or 2 (spinosa); interradial actinal spines long

sacciform.

Superambulacral plates irregularly present mid-ray and dis-

tally (abnormalis, longispina), lacking complete series; super-

actinal plates present as single plate struts (abnormalis,

longispina).

Distribution. Indo-West Pacific, South China Sea (Xisha Is),

Ceylon, Bay of Bengal (Andaman Is), Indonesia, New

Caledonia, Guam, Fiji, northern Australia, western Australia.

Remarks. Molecular data have not been obtained for species of

Disasterina and the genus is reviewed on morphological

grounds. Specimens of D. ceylanica and D. spinosa. were not

seen and morphological data were obtained from the original

descriptions and figures by Doderlein (1888) and Koehler

(1910).

Species previously assigned to Disasterina fall into two

morphological groups, those close to the type D. abnormalis

and retained here in Disasterina, and D. leptalacantha and

D. praesignis (with junior synonym D. spinulifera) which are

close to Tegulaster emburyi and are removed to Tegulaster.

Disasterina differs from Tegulaster in having: body thin and

low, not thick and high; abactinal plates thin and loosely con-

tiguous, not thick and imbricating; non-plated abactinal spaces

present, not absent; median band of irregular abactinal plates

on rays wide, not narrow; distal interradial plates at margin in

irregular perpendicular series, not longitudinal and thus crossed

angled series; superomarginal plates small and irregular, not

large and in distinct series; covering integument thick, not thin.

The form and patterns of spinelets and spines are similar in the

two genera. Disasterina also shares some morphological char-

acters with Indianastra gen. nov. below, in particular spination,

limited presence of superambulacral plates and presence of

series of superactinal plates. The morphological differences

between Disasterina and Indianastra are detailed under the

latter.

Manasterina longispina H.L. Clark, 1938 was described

from a single specimen and the genus distinguished from

Disasterina by size, single minute spinelets on some abactinal

plates, long spinelets on rays distally, openly spaced abactinal

plates, and absence of non-plated proximal actinal areas. None

of these characters is of generic value. The long spinelets on

distal ray plates is unique to the species. M. longispina is inter-

mediate in size amongst species of Disasterina; species of

Disasterina sometimes have spinelets on abactinal plates; and

the extent of non-plated abactinal and actinal body wall varies

amongst specimens of M. longispina and amongst species of

Disasterina. Manasterina is synonymised here with

Disasterina. Clusters of 2-4 undifferentiated interradial

spinelets with their distal ends angled together and contiguous

were observed on preserved D. longispina material, and

possibly act as pedicellariae.

Livingstone (1933) considered D. spinosa to be inappropri-

ately assigned to Disasterina but gave no reason. The single

long sacciform spinelets on abactinal plates distinguish the

species but are not grounds for rejecting its generic placement.

Disasterina ceylanica Doderlein (1888) was reassigned to

Tegulaster by Livingstone (1933). Disasterina and Tegul-

aster have some common characters but the reassignment is

inappropriate.

Material from the University of Florida extends the

distribution of D. odontacantha from China to Guam and Fiji.

Indianastra O’Loughlin gen. nov.

Figures 2h, 6b, 13a-f

Diagnosis. Rays 5, petaloid to subpentagonal; size small; body

thin, integument-covered; abactinal and actinal interradial

plates in longitudinal series; abactinal plates deeply notched for

papulae, short crescentiform; numerous regular longitudinal

series of plates with papulae along sides of rays; abactinal

spinelets fine, glassy, inconspicuous, acicular to subsacciform,

few to numerous per plate, in tufts or cover over plate, not on

high raised ridges or domes, fragile, readily lost; pedicellariae

sometimes present over papulae; superomarginal plates with

rare spinelets or bare; inferomarginal plates with distal sub-

paxilliform dense tufts of acicular spinelets, covered by

integument; actinal interradial plates in longitudinal series;

plates with small clusters of webbed short sacciform spines;

lacking superambulacral series of plates, rare single plates

distally; superactinal plates as single plate struts.

Type species. Palmipes sarasini de Loriol, 1897 (junior syn-

onyms: Asterina lutea H.L. Clark, 1938, Asterina nuda H.L.

Clark, 1921, and Asterina orthodon Fisher, 1922 by Rowe, 1995).

Other species. I. inopinata (Livingstone, 1933) (junior synonym:

Asterina perplexa H.L. Clark, 1938, by Rowe, 1995).

Material examined. I. inopinata. Holotype, 12 paratypes. Australia,

New South Wales, AM J3077. Other material. New South Wales, NMV

F93460 (3); Byron Bay, AM J15254 (6);

I. lutea. Paratypes. NW Australia, Broome, AM J6167 (4).

I. orthodon. Hong Kong, BMNH 1983.2.15.116 (1).

I. sarasini. NW Australia, WAM Z6833 (1); Broome, AM J6640 (5);

NMV F95802 (2); Queensland, J4123 (1).

Description with species variations. Rays 5, petaloid (rays wide

basally, rounded distally, subacute to narrowly rounded junc-

tions) to narrowing rounded to subpentagonal (interradial

margin straight to shallow incurved); body integument-

Genera of asterinid sea-stars

Figure 8. a, Anseropoda placenta (R = 56 mm, NMV F98043): abactinal plates, spinelets, papulae, b, Anseropoda rosacea (R = 82 mm, NMV

F95811): adambulacral and actinal plates and sacciform spines, c-d, Asterina gibbosa (R = 19 mm, NMV F45108): c, actinal interradius with

gonopores; d, margin supported by internal contiguous projections from abactinal and actinal plates, absence of superambulacral plates,

e, Asterina ocellifera (R = 37 mm, BMNFI 1969.12.16.13): margin supported by internal contiguous plate projections, absence of superambulacral

and superactinal plates, f-h, Asterinides folium (R = 20 mm, USNM 28573): f, abactinal spinelet clusters and papulae; g, actinal interradius

with sacciform spines; h, marginal plates with spinelets. i, Asterinides pompom (R = 14 mm, USNM 47866): abactinal plates with paxilliform

elevations.

covered, flat actinally, low convex abactinally; size small (both

species up to R = 20 mm); simple pedicellariae present over

papulae, valves short, conical to sacciform, up to 6 valves

(sarasini only); not fissiparous.

Abactinal appearance dominated by numerous regular long-

itudinal series of papulae along sides of rays; plates imbricate,

deeply notched for papulae, interradially in longitudinal series

to margin; papulate areas extensive, papular spaces fairly large,

predominantly single large papula per space, up to 4 small papu-

lae and 5 secondary plates per space; up to 6 longitudinal series

of papulae along each side of rays; plates on narrow median

upper rays irregular, carinal series of plates variably present,

doubly notched; abactinal plates with low spinelet-bearing

elevations, on lower rays and proximal interradii predomin-

antly singly notched for papulae, crescentiform only ( in -

opinata ), additional deep to almost closed pedicellarial notch

proximally ( sarasini ); fine, glassy, fragile, inconspicuous, aci-

cular to subsacciform spinelets, sparsely present in apical tufts

or numerous in crescentiform cover on projecting proximal

edge of plates, spinelets readily lost; glassy convexities ( sarsini )

or reticulations ( inopinata ) on plates; disc variably bordered by

continuous series of 5 large radial 5 small interradial plates;

superomarginal plates in series, rare spinelets ( inopinata ) or

bare (, sarasini ); inferomarginal plates in series, project to form

margin, bare proximally, distal subpaxilliform dense tuft of

acicular spinelets projects laterally, integument-covered.

P. Mark O’Loughlin and Jonathan M. Waters

Actinal plates in longitudinal series.

Actinal spines per plate (lower numbers in inopinata ): oral

8-11; suboral 4-9, webbed fan; furrow 5-9 proximally,

webbed; subambulacral 3-9; adradial actinal plates with com-

plete series; actinal 2-8 in mid interradius, webbed transverse

clusters; actinal interradial spines short sacciform.

Lacking series of superambulacral plates, rare single plates

distally (seen in sarasini only); margin supported by series of

single superactinal plates, and internal tongue-like projections

of abactinal plates contiguous with actinal plates.

Distribution. Indian West Pacific, Sri Lanka to southern China,

N and E Australia, 0-25 m.

Etymology. From the first word of “Indian West Pacific” and

the Latin astrum (star), referring to the region of occurrence of

this genus (feminine).

Remarks. Molecular data are not available for either species

and this review is based on morphological evidence.

Indianastra shares some morphological characters with both

Disasterina and Tegulaster, in particular the spination, limited

presence of superambulacral plates and presence of series of

superactinal plates. Morphological differences between

Indianastra and Disasterina are: form petaloid to subpentag-

onal, or rays discrete; abactinal plates small, of uniform size

and imbricate, or irregular in size, form and arrangement and

loosely contiguous; abactinal plates deeply, or slightly notched;

longitudinal series of papulate plates numerous and regular on

rays, or few and irregular on lower sides of rays; abactinal

interradial plates in longitudinal series, or series perpendicular

to margin; superomarginal series of plates distinct and regular,

or reduced and irregular; inferomarginal spinelets acicular, in

dense tufts, or few, sacciform and discrete; actinal series of

plates longitudinal, or oblique; actinal interradial spines short,

in clusters, or long, 1-2. Morphological differences with

Tegulaster are given under Tegulaster below. These morpho-

logical differences are the basis for the erection of the new

genus.

Jangoux (1985) established a lectotype and paralectotype

for Palmipes sarasini. Rowe (1995) formalised the synonymies

of I. lutea, I. nuda and I. orthodon with I. sarasini, which had

been suggested by Clark and Rowe (1971). Dartnall (1970,

1980) recorded A. inopinata from northern Tasmania, but

records were based on material subsequently determined by

O’Loughlin as Meridiastra nigranota O’Loughlin, 2002 (TM

H1330), M. atyphoida (H.L. Clark, 1916) (TM H841) and

Asterina scobinata Livingstone, 1933 (TM H1746). The pedi-

cellariae on I. sarasini are frequently difficult to detect because

the valves are lost or close over the notch and are integument-

covered.

Kampylaster Koehler

Kampylaster Koehler, 1920: 136-137. — Fisher, 1940: 250-252.—

Bernasconi, 1973: 344.— A.M. Clark, 1993: 220.

Diagnosis. Rays 5; body small, frequently arched, abactinally

covered by integument and subgranular spinelets; rays 5,

short petaloid; abactinal plates thin, not notched, irregular

arrangement; actinal plates in oblique series; actinal spines per

plate few, subglobose; lacking superambulacral and super-

actinal plates; abactinal and actinal distal interradial plates

lacking contiguous internal projections.

Type species. Kampylaster incurvatus Koehler, 1920 (original

designation).

Material examined. K. incurvatus. Syntypes. Antarctica, Adelie Land,

AM J3581 (2, A and B).

Description. Rays 5, petaloid to tapered from wide base;

margin incurved and subacute interradially; body variably

arched; size small, up to R = 15 mm; lacking pedicellariae;

brood-protects under body.

Abactinal plates obscured by integument and close uniform

cover of coarse subgranular spinelets; abactinal plates on rays

large, thin, imbricating, not notched, irregular arrangement;

papulae small, inconspicuous, single, irregular over rays (seen

from coelomic side by Fisher, 1940; not seen here); abactinal

spinelets subgranular, wide base, narrowed waist, convex and

spinous distally; superomaginal plates similar to adjacent

abactinal plates, covered with granular spinelets, not aligned

with inferomarginals; series of thick rounded projecting infero-

marginal plates define margin, covered with granular spinelets.

Actinal interradial areas small, series of plates oblique.

Actinal spines per plate: oral 3; suboral 0; furrow 2-3 proxi-

mally, webbed; subambulacral 1-2; actinal 1-3; adradial acti-

nal plates with series of shorter spines; actinal spines short,

thick, rounded, subglobose distally.

Lacking superambulacral and superactinal plates; abactinal

and actinal distal interradial plates meet at an angle, lacking

contiguous internal projections.

Distribution. Antarctica, Enderby Land to Scotia Sea; 93-750 m.

Remarks. Molecular data are not available for incurvatus, and

this review is based on a morphological examination of syn-

types and the observations of Fisher (1940). Distinctive diag-

nostic characters of Kampylaster are the integument and gran-

uliform spinelet cover, irregular arrangement of thin abactinal

plates, inconspicuous papulae, absence of a series of supero-

marginal plates, and absence of superambulacral and super-

actinal plates.

Koehler (1920) considered Kampylaster to be similar to

Tremaster Verrill, 1880 and referrable to the subfamily

Tremasterinae. Fisher (1940) rejected the similarity because of

the distinctive interradial perforations of Tremaster, and con-

sidered Kampylaster to be closer to Stegnaster Sladen, 1889.

Additional reasons for rejecting a similarity with Tremaster are

the quadriserial arrangement of tube feet and vertical furrow

spines on adambulacral plates in Tremaster.

Meridiastra O’Loughlin

Figures 1 (clade III), 2i, 4c, 5a, lOg-i

Meridiastra O’Loughlin, 2002: 280. — Waters et al., 2004: 874, 875,

877.

Diagnosis. Rays predominantly 5 or 6 or 7 or 8; interradial

margin straight or incurved, rays not discrete; flat actinally, low

Genera of asterinid sea-stars

to high convex abactinally; abactinal plates in longitudinal

series along rays, carinal series present for at least part ray

length; papulae small, lacking longitudinal series of large papu-

lae along sides of rays; abactinal spinelets granuliform; lacking

pedicellariae; abactinal plates with glassy convexities; actinal

plates in longitudinal series; actinal spines in mid-interradius

digitiform or tapering or conical, 1-3; lacking superambulacral

and superactinal plates, distal interradial margin supported by

contiguous interior projections from abactinal and actinal plates.

Type species. Asterina atyphoida H.L. Cark, 1916 (original

designation).

Other species. M. calcar (Lamarck, 1816); M. fissura O’Loughlin,

2002; M. gunnii (Gray, 1840) (junior synonym: Patiriella brevispina

H.L. Clark, 1938, by O’Loughlin et al., 2003); M. medius (O’Loughlin

et al., 2003); M. modesta (Verrill, 1870) (junior synonym: Asterina

agustincasoi Caso, 1977 by O’Loughlin, 2002); M. mortenseni

(O’Loughlin et al., 2002); M. nigranota O’Loughlin, 2002; M.

occidens (O’Loughlin et al., 2003); M. oriens (O’Loughlin et al.,

2003); M. rapa O’Loughlin, 2002.

Material examined. See O’Loughlin (2002), O’Loughlin et al. (2002,

2003).

M. calcar. Australia, Victoria, Cape Paterson, NMV F73126 (4).

Description with species variations. Rays predominantly 5

( atyphoida , modesta, mortenseni, nigranota, rapa), or 6 ( gun-

nii , medius, occidens, oriens ), or 7 (fissura ), or 8 (calcar)’,

interradial margin straight to incurved, rays sometimes distinct,

narrowly rounded to pointed distally; integument noticeable;

size medium ( gunnii up to R = 56 mm) to very small (rapa up

to R = 6.0 mm); pedicellariae absent; gonopores actinal

(nigranota) or abactinal; fissiparity in one species (fissura ).

Abactinal plates in longitudinal series; carinal series present

to variable extent; abactinal appearance of two types, with

papulate areas extensive, papular spaces large with numerous

small papulae and secondary plates, plates distinctly doubly or

singly notched with projecting proximal edge crescentiform

except in distal plates (calcar, gunnii, medius, mortenseni, occi-

dens, oriens), or with papulate areas small, papular spaces

small with few small papulae and secondary plates, plates not

distinctly notched and projecting proximal edge not crescenti-

form (atyphoida, fissura, modesta, nigranota, rapa)', lacking

distinct longitudinal series of large papulae along sides of rays;

disc variably distinct with border of continuous radial and inter-

radial plates; abactinal plates sometimes with low spinelet-

bearing elevation; cleared abactinal plates with glassy convex-

ities; abactinal spinelets over projecting anterior edge of plates,

not in tufts, granuliform, and stout, opaque and firmly attached

(calcar group above) or fine, glassy and weakly attached

(atyphoida group above); series of subequal, spinelet-covered,

superomarginal and inferomarginal plates; margin formed by

projecting inferomarginal plates.

Actinal plates in longitudinal series, sometimes oblique

(atyphoida).

Actinal spines per plate: oral 4-7 ; suboral 0-2; furrow 2-5

proximally, webbed; subambulacral 1-3; actinal 1-3 in mid-

interradius, up to 4 distally; adradial row of actinal plates with

variably complete series of spines; interradial actinal spines

thick, bulbous to short tapered to conical to digitiform.

Lacking superactinal and superambulacral plates; inter-

radial margin supported by contiguous internal projections of

the abactinal and actinal plates.

Distribution. Southern Australia, New Zealand, central and

eastern South Pacific, Mexico, Panama; 0-59 m.

Remarks. Meridiastra, erected by O’Loughlin (2002) on mor-

phological evidence only, is supported by the molecular phylo-

genetic relationships of the nine species included in clade HI of

Waters et al. (2004). The key morphological features of these

nine species are: straight to shallow incurved interradial mar-

gins, rays not discrete; longitudinal series of abactinal plates;

granuliform abactinal spinelets; lack pedicellariae; presence of

conspicuous glassy convexities on plates; actinal plates in lon-

gitudinal series; low numbers of thick, tapering actinal spines;

lack superambulacral and superactinal plates; have internal

contiguous projections of distal interradial abactinal and actinal

plates supporting the margin. Most of the species of

Meridiastra were previously assigned to Patiriella which has

irregularly arranged abactinal plates, actinal plates in predomi-

nantly oblique series, and superambulacral and superactinal

plates present. The molecular phylogeny supports the separ-

ation of Meridiastra and Patiriella. Meridiastra is closest

morphologically to Asterina which also lacks internal skeletal

plates but which has more discrete rays, opaque spinelets which

are elongate, and actinal plates in predominantly oblique series.

The molecular phylogeny also supports the separation of

Meridiastra (Indo-Pacific) from Asterina (NE Atlantic). The

morphological similarities of these two genera could reflect

convergent evolution, or the retention of ancestral character

states, or a true sister relationship; the phylogeny of Waters et

al., 2004 was uninformative in this regard.

Molecular clade III provides evidence of two groups of

closely related species within the clade: M. gunnii, M. oriens,

M. medius and M. occidens’, and M. fissura, M. nigranota

and M. atyphoida. These closer molecular relationships are

reflected in the morphological description above, where these

two sets fall within the M. calcar and M. atyphoida groups. Our

decision to combine the two groups into a single genus is

subjective but supported by the significant internal skeletal

similarities.

Nepanthia Gray

Figures 2j, 4f, 6c, 14a-c

Nepanthia Gray, 1840: 287.— Sladen, 1889: 386-387.— Verrill,

1913: 480.— Fisher, 1940: 270-271.— Fisher, 1941: 451-155, figs,

20-22, pi. 70 fig. 2. — Spencer and Wright, 1966: U69. — Rowe and

Marsh, 1982: 93.— A.M. Clark, 1983: 370.— A.M. Clark, 1993:

220.— Liao and Clark, 1995: 132-133.— Rowe, 1995: 36-37.—

McKnight (in Clark and McKnight), 2001: 158.

Asterina (Nepanthia) Perrier, 1875: 320.

Diagnosis. Rays 4-7, subcylindrical, to varying degrees flat

actinally with distinct to slight marginal edge; plates on upper

rays irregular in arrangement; secondary plates present; abacti-

nal and actinal interradial plates with dense clusters of thick or

thin glassy spinelets, frequently on spinelet-bearing elevations;

pedicellariae present; glassy convexities on plates; infero-

P. Mark O’Loughlin and Jonathan M. Waters

Figure 9. a-d, Aquilonastra cepheus : a, cleared ray (R = 17 mm, NMV F95793); b, internal superambulacral (left arrow) and superactinal (right

arrow) plates (R = 22 mm, WAM Z6778); c, cleared abactinal plates with some sacciform spinelets (R = 17 mm, NMV F95793); d, abactinal

margin (R = 22 mm, WAM Z6778). e, Aquilonastra batheri (R = 19 mm, NMV F97441): abactinal carinal plates with tufts of spinelets.

f, Aquilonastra coronata (R = 22 mm, NMV F95796): side of ray with paxilliform elevations on upper ray.

marginal plates projecting slightly; oblique series of actinal

plates variably evident; furrow spines 6 and more per plate;

actinal spines predominantly thin, glassy, sometimes sacci-

form; superambulacral, transactinal and superactinal plates

present, embedded in interior resinous lining in most species.

Type species. Nepanthia maculata Gray, 1840 (restriction by

Perrier, 1875; subsequent designation by Verrill, 1913).

Other species. N. belcheri (Perrier, 1875) (junior synonym: N. vari-

abilis H.L. Clark, 1938 by Rowe and Marsh, 1982); N. crassa (Gray,

1847); N.fisheri Rowe and Marsh, 1982; N. pedicellaris Fisher, 1913.

Material examined. N. belcheri. Queensland, Magnetic I., NMV

F97721 (1); Western Australia, Dampier, F95806 (1); Exmouth Gulf,

F95805 (1).

N. crassa. Western Australia, Fremantle, AM J6165 (1); Ludlow

Reef, J7418 (1).

N.fisheri. Timor Sea, AM J12649 (1).

N. maculata. Australia, Timor Sea, AM J 139 18 (2); Gulf of

Carpentaria, J7404 (2); J10536 (1); J13063 (1).

N. pedicellaris. Holotype. Philippines, USNM 32643.

N. variabilis. Paratypes. Western Australia, Broome, AM J6187 (4).

Description with species variations. Rays 4-7, elongate, sub-

cylindrical, equal or unequal lengths, tapering strongly (fisheri,

pedicellaris) or slightly; large ( maculata up to R = 94 mm) to

small {pedicellaris up to R = 23 mm); integument variably

evident; flat actinally, distinct to slight edge at margin; pedi-

cellariae present, 2-valve (fisheri, maculata, pedicellaris ) or

multi- valve (belcheri, crassa)-, fissiparous (belcheri) or not.

Abactinal plates strongly imbricating, projecting proximal

edge creating uneven surface, irregularly arranged on upper

rays, in regular longitudinal sloping series on sides of rays;

plates predominantly irregular in shape ( crassa ) or notched

(belcheri, fisheri, maculata, pedicellaris), with spinelet-bearing

curved elevations above notch (belcheri, pedicellaris) or raised

ridges (fisheri, maculata) or domes (crassa), lower surface with

glassy convexities; disc variably bordered; papulate areas

extend to near margin; papular spaces with 1-2 large papulae

and 1-8 secondary plates per space; spinelets thin, glassy, vari-

ably sacciform (fisheri , maculata, pedicellaris), some splay-

pointed distally (belcheri, crassa), up to more than 40 per plate;

variably regular superomarginal plates, inferomarginal series of

plates project slightly at margin, covered with spinelets.

Actinal plates with spine-bearing elevations, variably in

oblique series.

Actinal spines per plate: oral 9-10; suboral about 10-26, tall

and short; furrow 6-10; subambulacral 6-20; complete series

of adradial actinal spines; interradial actinal 4-6 or dense sub-

paxilliform clusters, glassy, thin (belcheri), thick and thin

(crassa), sometimes sacciform (fisheri, maculata, pedicellaris).

Superambulacral, transactinal and superactinal plates

present, embedded in internal resinous lining or not (pedicel-

laris).

Distribution. N, S, E and W Australia, Lord Howe I., Indonesia,

Timor Sea, New Guinea, W Indonesia, Philippines, Vietnam,

Burma, Mergui Archipelago, 0-123 m.

Genera of asterinid sea-stars

Figure 10. a-d, Callopatiria granifera (R = 50 mm, NMV F98049); a, mid-ray abactinal primary and secondary plates, spinelets and papulae;

b, internal superambulacral (left arrow) and superactinal (right arrow) plates; c, cleared lower lateral ray, with marginal plates, glassy convexities,

papulae and secondary plates; d, section through ray showing superactinal (left arrow) and superambulacral (right arrow) plates, e, Disasterina

abnormalis (R = 38 mm, WAM Z6749): integument-covered inferomarginal plate (arrow) with two distal sacciform spinelets. f, Disasterina

longispina (R = 20 mm, WAM Z6760): abactinal tip of ray, with margin and long upper ray spinelet (arrow), g, Meridiastra atyphoida (R = 12

mm, NMV F87166): minute abactinal conical spinelets. h, Meridiastra occidens (R = 22 mm, NMV F73186): proximal actinal interradius, with

oral spines, i, Meridiastra gunnii (R = 40 mm, NMV F73248): interradial margin supported by contiguous projections from abactinal and actinal

plates, absence of superambulacral and superactinal plates.

Remarks. Molecular data are not available for any species of

Nepanthia and this review is based on a morphological exami-

nation. Six species previously assigned to Nepanthia are

removed to Pseudonepanthia. The morphological characters

which distinguish Nepanthia from Pseudonepanthia are: rays

flat actinally, with a marginal edge; presence of secondary

plates, pedicellariae and glassy convexities; furrow spines six

and more per plate; presence of transactinal and superactinal

plates.

Nepanthia pedicellaris Fisher, 1913 was referred by Fisher

(1919) to Asterinopsis Verrill, 1913 (a nomen dubium). We

return it to Nepanthia.

Paranepanthia Fisher

Figures 1 (clade I), 2k, 5c, 15a-c

Paranepanthia Fisher, 1917: 172. — Fisher, 1919: 419. — H.L.

Clark, 1946: 136.— Spencer and Wright, 1966: U69. — A.M. Clark,

1993: 222. — Rowe, 1995: 39. — McKnight (in Clark and McKnight),

2001: 160.— Waters et al„ 2004: 874, 875, 877.

Diagnosis. Rays 5, medium length, wide basally, pointed or

rounded distally; flat actinally, rays elevated to low; abactinal

and actinal interradial plates with subpaxilliform dense clusters

of acicular subsacciform spinelets; abactinal plates mostly

irregular on upper rays, mostly series perpendicular to

P. Mark O’Loughlin and Jonathan M. Waters

Figure 11. a, Cryptasterina hystera (R = 11 mm, NMV F98457): abactinal surface with small papulae and granuliform spinelets. b, Parvulastra

exigua (R = 10 mm, NMV F98062): abactinal surface with larger papulae and granuliform spinelets.

Figure 12. Abactinal views of Disasterina species, a, Disasterina abnormalis (R = 11 mm, WAM Z6753). b, Disasterina longispina (R = 25 mm,

WAM Z6760).

margin in interradii; marginal plates subequal, in regular series,

with dense tufts of spinelets, inferomarginals project promi-

nently; actinal interradial plates in oblique series; super-

ambulacral plates absent; superactinal plates present as

multiple struts.

Type species. Nepanthia platydisca Fisher, 1913 (original

designation).

Other species. P. aucklandensis (Koehler, 1920); P. grandis (H.L.

Clark, 1928) (junior synonym Asterinopsis praetermissa Livingstone,

1933, by Rowe, 1995).

Material examined. P. aucklandensis. New Zealand, Auckland I., AM

J5160 (2); J6023 (1).

Asterinopsis praetermissa. Holotype. Australia, New South Wales,

Port Jackson, AM J4793.

Nepanthia platydisca. Holotype. Indonesia, 377 m, USNM 32644.

P. grandis. Australia, Victoria, NMV F72887 (1); Bass Strait,

F98044 (1); South Australia, F73976 (4).

Description with species variations. Rays 5, interradial margin

deeply incurved, rays distinct, ends of rays rounded ( aucklan-

densis, grandis ) or pointed (platydisca ); body flat actinally,

rays elevated or low (platydisca ); distal interradial body thin

(actinal to abactinal) or not (aucklandensis), area of non-papu-

late plates wide or not (aucklandensis)', size medium (grandis

up to R = 65 mm) to small (aucklandensis up to R = 22 mm);

lacking pedicellariae; none fissiparous; gonopores abactinal.

Abactinal plates imbricate, distinct radial and interradial

‘fields’, plates on upper rays irregular in size, shape, arrange-

ment (except aucklandensis with regular proximal carinal and

upper ray series, and crescentiform plates), up to 3 fairly irreg-

ular longitudinal series along each side of rays; papular spaces

large, with numbers of small papulae, variable numbers of sec-

ondary plates; interradial series of small plates perpendicular to

margin (except longitudinal series in aucklandensis)', disc not

bordered or variably bordered (aucklandensis)', abactinal plates

with high (grandis) or low (aucklandensis, platydisca) spinelet-

Genera of asterinid sea-stars

bearing domes or ridges; glassy convexities on plates present

( aucklandensis ) or not ( grandis , platydisca ); subpaxilliform

tufts of many acicular subsacciform spinelets; superomarginal

and inferomarginal plates in distinct series, subequal; both

series with spinelet-bearing domes and subpaxilliform dense

tufts of acicular spinelets; inferomarginals define acute margin,

projecting plates with narrowed waist ( aucklandensis , grandis)

or not ( platydisca ).

Actinal plates in oblique series; plates with high ( grandis ,

platydisca) or low ( aucklandensis ) spine-bearing domes.

Actinal spines per plate: oral 8-12; suboral tufts or trans-

verse series; furrow 7-10, webbed; subambulacral dense tufts;

adradial and actinal interradial dense subpaxilliform tufts (up to

about 40 spines); actinal interradial spines acicular subsacci-

form.

Lacking superambulacral plates; numerous super-

actinal plates in multiple plate struts; distal interradial margin

supported by angled contiguous abactinal and actinal plates

(grandis, platydisca), lacking internal projections (except auck-

landensis with internal tongue-like projections from abactinal

plates).

Distribution. E and S Australia, Indonesia, subantarctic New

Zealand; 0-377 m.

Remarks. Three species cluster together in clade I of Waters et

al. (2004). One of these is a species that has been assigned to

Paranepanthia in the past and one is undescribed. The molec-

ular evidence that Asterina aucklandensis is also a member of

this clade is supported by morphological criteria. All three

species of Paranepanthia were examined and the mor-

phological integrity of the genus confirmed. Paranepanthia is

characterised by the unique combination of subpaxilliform

dense clusters of acicular spines and spinelets on the plates,

absence of superambulacral plates, and presence of super-

actinal plates. Paranepanthia rosea has been removed to

Aquilonastra.

Parvulastra O’Loughlin gen. nov.

Figures 1 (clade V), 21, 4a-b, 5b, e, lib, 16a-d

Diagnosis. Rays 5, pentagonal to subpentagonal; noticeable

integument; carinal series variably present; abactinal plates

broadly notched for papulae, crescentiform more than U-

shaped; papulate areas extensive, papulae large or small, a few

per space; secondary plates a few per space; abactinal spinelets

clustered or spread, not paxilliform; spinelets granuliform or

digitiform or thin pointed; lack pedicellariae; superomarginal

and inferomarginal plate series predominantly subequal; fre-

quently extensive non-plated actinal area distal to oral plates;

actinal interradial plates in oblique series; actinal interradial

plates with frequently 2 spines mid-ray to distally, spines digi-

tiform to conical; superambulacral plates present to varying

extents, superactinal plates always present.

Type species. Asterias exigua Lamarck, 1816.

Other species. P. calcarata (Perrier, 1869); P. dyscrita (H.L. Clark,

1923); P. parvivipara (Keough and Dartnall, 1978); P. vivipara

(Dartnall, 1969).

Material examined. P. calcarata. Juan Fernandez I., NMV F96703 (1);

F96704 (1); F97445 (1); F97449 (1).

P. dyscrita. South Africa, False Bay, TM H800 (1); H854 (1);

Jeffrey’s Bay, NMV F98059 (3).

Patiriella exigua (Lamarck, 1816). Neotype (by Dartnall, 1971).

South Africa, False Bay, TM H508. Other material. Cape Town, NMV

F98062 (4); Durban, F98061 (4); Jeffrey’s Bay, F97450 (5); Saint

Helena, F98060 (4); Amsterdam I., F98063 (2); Australia, Victoria,

Apollo Bay, F97054 (6); Gabo I., F73079 (9); Port Arthur, F97451

( 12 ).

P. vivipara. Australia, SE Tasmania, NMV F77984 (3).

P. parvivipara. South Australia, Eyre Peninsula, NMV F97720

(3).

Description with species variations. Rays predominantly 5;

interradial margin straight to slightly incurved, form pentago-

nal to subpentagonal; noticeable integument; flat actinally, low

convex abactinally, size small (dyscrita up to R = 24 mm) to

very small (parvivipara up to R = 3 mm); lacking pedicell-

ariae; 2 smallest viviparous (parvivipara, vivipara)', none fissi-

parous; gonopores abactinal (calcarata, dyscrita) or actinal

(exigua).

Abactinal surface appearance granular, upper ray plates with

regular carinal series except proximally (calcarata, dyscrita,

exigua), or irregular (parvivipara, vivipara)', plates in longitu-

dinal series; plates broadly notched for papulae, crescentiform

more than U-shaped; papulate areas extensive, non-papulate

interradial areas not extensive, papular spaces large, a few

papulae and secondary plates per space; papulae large (exigua,

parvivipara, vivipara) or small (calcarata, dyscrita)', disc vari-

ably bordered by series of 5 large radial and 5 small interradial

plates; abactinal plates with raised spinelet-bearing elevations

(calcarata) or not; abactinal spinelets in splayed clusters (cal-

carata) or fairly close cover over plates; spinelets both digiti-

form on primary plates and pointed on secondary and distal

interradial plates (calcarata), or granuliform globose (dys-

crita), or short columnar (exigua, parvivipara, vivipara);

glassy convexities and reticulations variably evident on

denuded plates; superomarginal and inferomarginal series

of predominantly subequal plates; typically widely project-

ing inferomarginal plates define margin, with proximal

abactinal-type spinelets, distal fringe of thinner and longer

spinelets.

Actinal interradial plates in oblique series; actinal proximal

areas of non-plated body wall frequently extensive.

Actinal spines per plate: oral 2 tall, 3-5 short sometimes

grouped separately with distalmost frequently longest; suboral

1-2, thick; furrow 2 short, thin; subambulacral 1 tall, thick;

adradial actinal spines present (dyscrita) or few (vivipara)

or absent (calcarata, exigua); actinal interradial frequently

2 midray to distally; actinal interradial spines digitiform

(dyscrita) or pointed or short conical.

Superambulacral plates present as series (in large dyscrita),

or rare single, or distal only plates; superactinal plates present

as single or multiple-plate series (neither seen in the pedo-

morphic parvivipara).

Distribution. Southern Pacific Ocean, southern Australia,

southern Indian Ocean, southern Africa, SE Atlantic Ocean;

0-10 m.

P. Mark O’Loughlin and Jonathan M. Waters

Etymology. From the Latin parvulus (very small) and astrum

(star), referring to the small to very small species of the genus

(feminine).

Remarks. Four species previously included in Patiriella belong

to a monophyletic clade V in Waters et al.’s (2004) analysis.

They share a consistent morphology characterised by: subpen-

tagonal form; longitudinal series of abactinal plates on rays;

oblique series of actinal plates; granuliform and digitiform

spinelets; a few papulae per space; and superambulacral and

superactinal plates. The clade warrants generic rank. Asterina

dyscrita shares these morphological characters and is assigned

to the genus on this basis.

Morphologically, Parvulastra is similar to Cryptasterina

and Patiriella. Waters et al.’s cladogram suggests that these

three genera are not closely related on molecular grounds.

Either the similar morphological characters have remained

stable while divergence has occurred in characters not consid-

ered, or there is strong morphological convergence among the

three unrelated genera, or the molecular data are unreliable at

levels more basal than inter- species. Nevertheless, the genera

can be differentiated by morphological characters.

Cryptasterina is differentiated by predominantly single papulae

per space, elongate superomarginal plates, narrowly projecting

inferomarginal plates, and single mid-actinal spines. The

characters which differentiate Patiriella are listed under that

genus.

Patiria Gray

Figures 1 (clade II), 3a, 4d, 15d-f

Patiria Gray, 1840: 290. — Gray, 1847: 82-83. — Gray, 1866: 16. —

Sladen, 1889: 384.— Verrill, 1913: 480, 482.— Verrill, 1914:

263-264.— Fisher, 1919: 410.— Fisher, 1940: 269.— Fisher, 1941:

451-455, pi. 70 fig. 1.— Spencer and Wright, 1966: U69.—

Bernasconi, 1973: 336. — Clark and Courtman-Stock, 1976: 78. —

Campbell and Rowe, 1997: 131, 135. — Dartnall et al., 2003: 12.

Diagnosis. Rays 5, medium to short-rayed stellate, not pen-

tagonal, rays pointed distally; noticeable integument; flat actin-

ally, convex abactinally; primary abactinal plates strongly or

weakly crescentiform, close subpaxilliform cover of short,

thick or thin, columnar or subglobose spinelets, not acicular or

sacciform, secondary plates with smaller spinelets; lacking

pedicellariae; papulate areas extensive, papular spaces large,

numerous secondary plates and papulae per papular space;

series of spinelet-covered superomarginal and inferomarginal

plates, spinelets on thick rounded inferomarginals extend onto

actinal surface; actinal plates in oblique series; actinal spines in

mid-interradius thick, combs of 4-6; lacking complete super-

ambulacral series of plates, present distally; superactinal plates

present, multiple plate struts in larger specimens.

Type species. Patiria coccinea Gray, 1840 (original designa-

tion) (junior synonym of Asterias miniata Brandt, 1835 mis-

takenly recorded from South Africa, synonymy by Mortensen,

1933).

Other species. P. chilensis (Liitken, 1859); P. pectinifera (Muller and

Troschel, 1842).

Material examined. P. chilensis. Chile, Arica, NMV F95674 (3); Peru,

NMV F97442 (1); F97443 (1).

P. miniata. California, NMV F97444 (1); F97448 (1); F98040 (1);

F98041 (1); F98042 (1).

P. pectinifera. Japan, Toyama Bay, NMV F95672 (7); F95673 (2).

Description with species variations. Rays 5 (sometimes 6);

noticeable integument; interradial margin incurved, medium to

short-rayed stellate, not pentagonal, rays distally pointed to nar-

rowly rounded; flat actinally, convex abactinally, body thick,

sides of rays steep; size large {miniata up to R = 85 mm) to

small {chilensis up to R = 21 mm); lacking pedicellariae;

gonopores abactinal.

Abactinal surface uneven; appearance dominated by sub-

paxilliform spinous primary plates, variably crescentiform, and

large papular spaces with smaller spinous secondary plates;

plates in irregular longitudinal series; papulate areas extensive,

to near distal end of rays and near margin; papular spaces with

numerous secondary plates and small papulae (up to about 25

of each); lacking longitudinal series of large single papulae

along sides of rays; disc weakly bordered by discontinuous

series of larger radial and smaller interradial plates; abactinal

primary plates variably crescentiform or oval or round or linear,

with spinelet-bearing elevations, variably with 1 or more notch-

es for papulae; carinal series variably present, plates separated

by large papular spaces; cleared plates with glassy convexities

prominent ( chilensis ) or not; abactinal spinelets opaque, short

columnar, thick {miniata), or subglobose ( pectinifera ), or thin

{chilensis), in section round {chilensis, pectinifera) or square

{miniataf, spinelets on primary plates in subpaxilliform trans-

verse or round clusters, very close and palisade-like {miniata)

or slightly more spaced {chilensis, pectinifera), on secondary

plates shorter, splayed; series of subequal superomarginal and

inferomarginal plates, covered with spinelets; spinelets on thick

rounded inferomarginals extend onto actinal surface; projecting

inferomarginals form acute margin.

Actinal plates in oblique series.

Actinal spines per plate: oral 5-7; suboral 2-7; furrow 3-4

proximally; subambulacral 2^4; adradial actinal plates fully

spinous; actinal interradial in combs of 3-8 proximally, 4-6

distally; actinal interradial spines conical to digitiform to

spatulate.

Superambulacral plates not present as series, sometimes

present in mid-ray {chilensis) or absent (miniata, pectinifera),

present distally linking with actinal or superactinal or abactinal

plates; superactinal plates present, multiple plate struts in

larger specimens.

Distribution. N and E Pacific Ocean, Japan, Alaska, California,

Peru, Chile; 0-300 m.

Remarks. Fisher (1908, 1911), Mortensen (1933), Hayashi

(1940) and A.M. Clark (1983) considered Patiria to be a junior

synonym of Asterina, while Verrill (1913), Fisher (1919, 1940,

1941) and Clark and Courtman-Stock (1976) maintained

Patiria as a valid genus. Verrill (1914) considered it to be sig-

nificant that Patiria lacked pedicellariae. Spencer and Wright

(1966) listed Callopatiria and Enoplopatiria as junior

synonyms of Patiria. Hayashi (1973) considered Patiria and

Patiriella to be junior synonyms of Asterina. A.M. Clark

Genera of asterinid sea-stars

Figure 13. a, Indianastra inopinata (R = 19 mm, AM J15254): cleared abactinal surface, b-f, Indianastra sarasini (R = 14 mm, WAM Z6835): b,

abactinal ray; c, abactinal elongate subsacciform spinelets, and papulae; d, bare superomarginal plates, and tufts of inferomarginal spinelets; e,

oral and proximal actinal spines; f, actinal plates and spines.

(1983), Clark and Downey (1992) and A.M. Clark (1993) con-

sidered Enoplopatiria and Patiria to be junior synonyms of

Asterina. Campbell and Rowe (1997) “accepted the separate

generic status [of Patiria ] until the matter is more clearly

resolved”.

Waters et al. (2004) found that the three species of Patiria,

including the type species, constituted a single clade (II). They

confirmed an earlier result, based also on molecular data (Hart

et al., 1997), that confirmed the type and one of the species to

be sister taxa. Together there is strong molecular evidence for

the generic status of Patiria, remote from Asterina and

Patiriella. Morphologically, Patiria is distinguished from

Asterina by the presence of superambulacral and superactinal

plates, and from Patiriella by the discrete rays, dense sub-

paxilliform clusters of granuliform spinelets, and more

numerous actinal interradial spines.

Patiriella Verrill

Figures 1, 3b, 16e-f

Patiriella Verrill, 1913: 480, 483^184.— Verrill, 1914: 263.—

Fisher, 1919: 410.— H.L. Clark, 1946: 134.— Spencer and Wright,

1966: U69.— Shepherd, 1968: 745.— Dartnall, 1970: 73-74.—

Dartnall, 1971: 39-40. — Bernasconi, 1973: 341. — Clark, A.M. and

Courtman-Stock, 1976: 80.— Clark, A.M. , 1983: 364-367, 378, fig. 3c,

4.— Clark, A.M. and Downey, 1992: 178, 192.— Clark, A.M., 1993:

224.— Rowe, 1995: 39.— Campbell and Rowe, 1997: 129-131.—

McKnight (in Clark and McKnight), 2001: 155. — O’Loughlin et al.,

2002: 701.— Dartnall et al., 2003: 11-12.— Waters et al., 2004: 874,

875, 877. (Part).

Diagnosis. Rays 5, interradial margin straight to incurved, sub-

pentagonal to short discrete rays, ends pointed or broadly or

narrowly rounded; noticeable integument; plates on rays irreg-

ularly arranged; abactinal spinelets granuliform to digitiform,

not webbed, in close to spaced distribution over projecting sur-

face of plates; regular series of granuliform or digitiform

spinelet-covered superomarginal and inferomarginal plates;

actinal plates in oblique series; actinal adradial spines in incom-

plete series; actinal spines digitiform to short conical, no more

than 3 per plate; superambulacral plates present from midray or

distally only; superactinal plates present as single and multiple-

plate supports.

Type species. Asterina ( Asteriscus ) regularis Verrill, 1867

(original designation).

Other species. P. inomata Livingstone, 1933; P. oliveri (Benham,

1911) (junior synonym: Patiriella nigra H.L. Clark, 1938, by Rowe,

1995); Patiriella paradoxa Campbell and Rowe, 1997.

Material examined. P. inomata Livingstone, 1933. Holotype. Western

Australia, AM J3198.

P. nigra. Paratype. Lord Howe I., AM J4439 (1).

P. oliveri. Kermadec Is, MNZ EC4805 (2 of 7); Lord Howe I., AM

G11519 (2); G2247 (1).

P. paradoxa. Holotype. Oman, BMNH 1997.1016. Paratypes.

BMNH 1997.1017 (1); BMNH 1997.1018 (2).

A. regularis. Syntypes. New Zealand, Auckland, H. Edwards, 1866,

YPM 988A (1), B (1, partially cleared), C (2). Other material. See

O’Loughlin et al. (2002).

Description with species variations. Rays 5, rarely 6; inter-

radial margin straight to incurved, form subpentagonal to

P. Mark O’Loughlin and Jonathan M. Waters

Figure 14. a-c, Nepanthia pedicellaris (R = 23 mm, holotype, USNM 32643): a, cleared lower lateral ray, and distinct margin; b, abactinal

spinelets; c, adambulacral and actinal plates and spines, d, Pseudonepanthia troughtoni (R = 55 mm, NMV F73977): cleared proximal lateral

lower ray, margin, and ambulacrum, e, Pseudonepanthia nigrobrunnea (R = 85 mm, NMV F95810): cleared section of ray with single series of

superambulacral plates (arrow), and absence of transactinal and superactinal plates, f, Pseudonepanthia gotoi (R = 70 mm, holotype, USNM

36899): internal cleared view of ambulacral plates, with series of superambulacral plates (arrow).

discrete rays with ends broadly rounded to pointed; integu-

ment cover sometimes evident ( regularis)', body flat

actinally, low convex abactinally; rays elevated, sides

steep; size medium ( regularis up to R = 39 mm); lacking

pedicellariae; rare evidence of fissiparity ( regularis ); abactinal

gonopores.

Abactinal plates on upper rays in longitudinally series

(oliveri) or irregular ( inornata , paradoxa, regularis ); carinal

plate series ray length ( oliveri ) or short proximal ( regularis ) or

not evident ( inornata, paradoxa ), plates doubly notched; vari-

ably regular longitudinal series of crescentiform plates on

lower sides of rays; papulate areas extensive, non-papulate

interradial area extensive ( inornata , oliveri) or narrow

( paradoxa , regularis)', papular spaces large, 1-4 large (in-

ornata, paradoxa) or numerous small papulae and irregular

secondary plates per space ( oliveri , regularis)', proximal

primary abactinal plates irregular (inornata, paradoxa) or cres-

centiform (oliveri, regularis), with spinelet-bearing ridges

(oliveri, regularis) or not; glassy convexities below

elevations on denuded plates (oliveri, paradoxa, regularis), or

reticulations (inornata)', abactinal spinelets granuliform,

subglobose to subconical (inornata, paradoxa), or proxim-

ally short columnar, distally short subsacciform (regularis),

or digitiform, many slightly widened terminally ( oliveri );

spinelets not webbed, in close arrangement on plates (oliveri,

paradoxa, regularis) or spread (inornata)', disc variably

bordered by 5 radial and 5 interradial plates; supero-

marginal and inferomarginal plates in regular series, with

typical abactinal spinelets; margin formed by projecting infer-

omarginals.

Actinal plates in oblique series (irregular proximally in inor-

nata, paradoxa)', proximal actinal interradial areas sometimes

not plated.

Actinal spines per plate: oral 4-6; suboral 0-4; furrow 2-4

proximally, short, webbed; subambulacral 1-3, tall; adradial

actinal 0-1, short, incomplete series; actinal interradial 1-3

proximally, 1-2 short distally; actinal interradial spines digiti-

form, slightly tapered (regularis), slightly swollen (oliveri), or

short conical (inornata), or short thick columnar (paradoxa ).

Superambulacral plates present distally (inornata not

examined distally); superactinal plates present as single and

multiple-plate struts.

Distribution. New Zealand, North and South Is, Stewart and

Chatham Is; Australia, Lord Howe I., New South Wales,

Tasmania; Western Australia; Oman; 0-92 m.

Remarks. O’Loughlin et al. (2002) did not report on the four

syntypes of Asterina regularis Verrill, 1867 in their systematic

review of Patiriella. They have now been examined and are

consistent with O’Loughlin et al.’s (2002) redescription. Waters

et al. (2004) presented a molecular phylogeny in which

P. regularis fell on a clade remote from those on which all other

species of Patiriella were included (clades III part, IV part, and

V, in Fig. 1). The species in clade III (part) were removed to

Meridiastra, those in clade IV (part) to Cryptasterina by

Dartnall et al. (2003), and those in clade V to Parvulastra.

Genera of asterinid sea-stars

The diagnostic characters of P. regularis are: subpentagonal

to very short-rayed form; abactinal plates on rays not in

longitudinal series; spinlets ganuliform, short columnar; large

papular spaces with numerous papulae and secondary plates;

actinal plates in oblique series; few superambulacral plates;

superactinal plates present. Patiriella is distinguished from

Meridiastra by lacking longitudinal series of plates on the rays,

by oblique series of actinal plates, and by having superactinal

plates. Patiriella is distinguished from Cryptasterina and

Parvulastra by lacking longitudinal series of plates on the rays,

and by having large papular spaces with numerous papulae and

secondary plates.

On morphological grounds P. inornata and P. paradoxa are

retained in Patiriella. Rowe (1995) transferred Asterina oliveri

to Patiriella. P. oliveri has many of the characters of Patiriella,

but is retained in Patiriella with reservation because of the reg-

ular carinal and longitudinal series of plates on the rays. On

morphological grounds, Asterina dyscrita H.L. Clark, 1923 is

removed from Patiriella to Parvulastra.

Patiriella tangribensis Domantay and Acosta, 1970 is

judged to be nomen dubium. Domantay and Acosta (1970)

established the species without type material being designated,

and the description and figures are not adequate to diagnose the

material (reference was made to an absence of marginal plates).

It was distinguished only from Patiriella exigua, which does

not occur in the type locality (Philippines).

Pseudasterina Aziz and Jangoux

Pseudasterina Aziz and Jangoux, 1985: 283-284. — A.M. Clark,

1993: 227.

Diagnosis. Small thin body; rays 5, discrete, broad base, round-

ed or pointed distally, medium-rayed stellate; abactinal upper

ray plates variably regular in form and arrangement; interradi-

al plates small, long imbricating oblique series; papulae along

upper rays only, sparse, single; short columnar granuliform

abactinal spinelets; superomarginal series of large plates with

granuliform spinelets; inferomarginal plates project narrowly,

with glassy subsacciform spinelets; actinal plates in oblique

series; actinal interradial plates with numerous glassy subsacci-

form spines principally on proximal edge, few distally; lacking

superambulacral and superactinal plates; very thin interradii,

with abactinal and actinal imbricate plates angled and meeting

internally throughout the interradii.

Type species. Pseudasterina delicata Aziz and Jangoux, 1985

(original designation).

Other species. P. granulosa Aziz and Jangoux, 1985.

Material examined. P. delicata. Tonga, 149-157 m, MNHN

EcAs 11704 (2); Wallis and Futuna Is, 250 m, MNHN EcAsll706

( 1 ).

Description with species variations. Rays 5, discrete, broad

base, distally rounded ( delicata ) or pointed ( granulosa ),

medium-rayed stellate; body flat, thin; size small ( granulosa up

to R = 18 mm); lacking pedicellariae; neither fissiparous.

Abactinal appearance dominated by imbricating small

interradial plates in oblique series from upper rays to

margin; papulate areas confined to narrow upper rays; papular

spaces small, papulae single, rare secondary plates; abactinal

plates on narrow upper rays variably regular in form and

arrangement, some notched, some indented for papulae, proxi-

mal doubly or singly papulate carinal series sometimes

present; disc distinctly bordered; abactinal plates with short

columnar stout granuliform spinelets, weakly attached,

spaced not clustered, up to 10 on proximal edge or covering

plate, closer and slightly larger over some papulae; glassy

reticulations on plates, not convexities; superomarginal plates

larger than adjacent abactinals, regular series, longitudinally

subrectangular, covered with granuliform spinelets; margin

formed by narrowly projecting inferomarginal plates, each with

numerous conical to subsacciform to splay-pointed sacciform

spinelets.

Actinal interradial plates in distinct imbricating oblique

series.

Actinal spines per plate: oral up to about 10, webbed; subo-

ral up to about 15, short, glassy conical; furrow up to about 10

on furrow edge, webbed, up to about 10 on surface of plate;

subambulacral, adradial actinal, and actinal interradial about

10, predominantly on proximal edge of plates; actinal spines

glassy, short, pointed, sacciform.

Lacking superambulacral and superactinal plates; thin inter-

radii with abactinal and actinal plates imbricating and angled

interiorly to meet throughout the interradii; no tongue-like

inner projections from abactinal or actinal plates.

Distribution. Philippines, Wallis, Futuna and Tonga Is,

130-250 m.

Remarks. No molecular data are available and this review is

based on morphology. Until recently the two species were

known only from the type material from the Philippine Is

( Pseudasterina delicata holotype, MNHN EcAs 10065;

paratype, MNHN EcAs 10066. P. granulosa holotype, MNHN

EcAs 10067). None of the type material could be located.

Recently A J. Dartnall determined material in the MNHN col-

lections as P. delicata. These specimens have been examined in

this study. The distinctive characters of Pseudasterina are:

abactinal long imbricating oblique series of small plates from

narrow upper radii to margin, and similar series from furrow to

margin actinally; very thin interradii, with abactinal and actinal

imbricate plates angled and meeting internally throughout the

interradii; short columnar, granuliform, readily-detached

spinelets on abactinal and superomarginal plates, but glassy,

subsacciform, sometimes splay-pointed, spinelets on the infer-

omarginal plates. The spination characters given by Aziz and

Jangoux (1985) for distinguishing P. delicata and P. granulosa

appeared to be all variably present on the P. delicata specimens

observed here, suggesting that P. granulosa may be a

junior synonym. Aziz and Jangoux (1985) distinguished

Pseudasterina from all other asterinid genera by the distinctive

series of large superomarginal plates. This diagnostic char-

acter is a feature of Cryptasterina also, from which

Pseudasterina differs in abactinal and actinal plate arrangement

and spination, and in the absence of superambulacral and

superactinal plates.

P. Mark O’Loughlin and Jonathan M. Waters

Figure 15. a-b, Paranepanthia platydisca (R = 55 mm, holotype, USNM 32644): a, cleared abactinal ray from upper ray to margin; b, ambulacmm

with adambulacral and actinal plates and spines, c, Paranepanthia grandis (R = 50 mm, NMV F73976): section of ray showing superactinal plates

supporting margin, and absence of superambulacral plates, d, Patiria miniata (R = 65 mm, NMV F97444): cleared abactinal surface with sub-

paxilliform clusters of granuliform spinelets. e-f, Patiria pectinifera (R = 40 mm, NMV F95672): e, cleared abactinal surface, with subpaxilli-

form globose spinelets; f, cleared abactinal surface, with papular spaces, secondary plates and papulae.

Pseudonepanthia A.H. Clark

Figures 1, 3c, 4e, 5d, 14d-f

Pseudonepanthia A.H. Clark, 1916: 118. — A.M. Clark, 1993: 227.

Diagnosis. Rays 4-10, subcylindrical, not flat actinally, lacking

marginal edge; integument variably noticeable; lacking second-

ary plates; abactinal spinelets thick or thin or subsacciform,

covering plates or in tufts; lacking pedicellariae; plates lacking

glassy convexities; inferomarginal plates not projecting; furrow

spines up to 5 per plate; actinal interradial spines digitiform, up

to 7-30 per plate; series of superambulacral plates present, rare

transactinal plates; lacking superactinal plates.

Type species. Pseudonepanthia gotoi A.H. Clark, 1916

(original designation).

Other species. P. briareus (Bell, 1894); P. gracilis (Rowe and Marsh,

1982); P. grangei (Me Knight, 2001) (in Clark and Me Knight);

P. nigrobrunnea (Rowe and Marsh, 1982); P. reinga (McKnight, 2001)

(in Clark and McKnight); P. troughtoni (Livingstone, 1934).

Material examined. P. briareus. Mariana Is, Guam, 118-152 m, UF

226 (1).

Pseudonepanthia gotoi. Holotype. Japan, Sagami Bay, 90 m,

USNM 36899.

P. gracilis. Australia, New South Wales, AM J 12874 (2); Jll 880 (2).

P. nigrobrunnea. New South Wales, Coffs Harbour, NMV F95810

(2); Byron Bay, F95804 (1).

Parasterina troughtoni. Holotype. Western Australia, Albany, AM

J3978. Other material. Victoria, Cape Woolamai, NMV F73013 (1);

Wilsons Promontory, F73017 (1); South Australia, Cape Jervis,

F73977 (4).

Description with species variations. Rays 4-10, distinct,

elongate, subcylindrical, tapered slightly {nigrobrunnea,

troughtoni ) or strongly {briareus, gotoi, gracilis, grangei, rein-

ga ), some irregular {briareus, gotoi, nigrobrunnea ); integument

variably noticeable; margin rounded, rays not flat actinally,

lacking marginal edge; size large {troughtoni up to R = 87 mm)

to medium {grangei up to R = 33 mm); lacking pedicellariae;

gonopores sometimes actinal {troughtoni)-, one fissiparous

{briareus).

Abactinal plates imbricating, projecting proximal edge

creating uneven surface, irregular in form and arrangement on

upper rays, oblique longitudinal series along sides of rays; lack-

ing secondary plates; papulate areas extending to near margin;

papular spaces with single large papula or a few {nigro-

brunnea)-, abactinal plates with spinelet-bearing elevations or

not {troughtoni)-, spinelets glassy, subsacciform with splayed

points distally {briareus, gotoi), or subsacciform {gracilis),

or fine columnar {nigrobrunnea, troughtoni)-, spinelets cover-

ing plates closely {gotoi, gracilis, nigrobrunnea, troughtoni)

or in splayed groups {briareus)-, disc not bordered; cleared

plates lacking glassy convexities; superomarginal plates in

regular or irregular series; marginal plates covered with

abactinal type spinelets; inferomarginal plates in predomin-

antly regular series, frequently longitudinally elongate, not

projecting.

Genera of asterinid sea-stars

Figure 16. a-d, Parvulastra exigua : a, cleared ray (R = 10 mm, NMV F98062); b, denuded abactinal granuliform spinelets (NMV F98062); c,

actinal interradius (R = 9 mm, NMV F98062); d, oral spines (R = 10 mm, NMV F97450). e-f, Patiriella oliveri (R = 28 mm, MNZ EC4805): e,

abactinal disc and spinelets; f, absence of superambulacral plates (left arrow), presence of superactinal plates (right arrow), g-i, Pseudopatiria

obtusa (R = 54 mm, holotype, BMNH 1938.6.23.24): g, section of ray with absence of both superambulacral and superactinal plates; h, abactinal

subpaxilliform spinelets, and pedicellaria (lower left); i, actinal spines.

Actinal plates with spine-bearing elevations or not

(troughtoni)-, not in oblique series.

Actinal spines per plate: oral 3-10; suboral 4-12; furrow

3-5; subambulacral 6-10; actinal 7-30; adradial row of actinal

plates with complete series of spines; interradial actinal spines

digitiform.

Superambulacral plates present as predominantly single

series along ambulacrum, sometimes contiguous with infero-

marginal plates across floor of ray ( briareus , troughtoni) or

with abactinal plates ( nigrobrunnea ); rare transactinal plates

(sometimes irregularly present in gracilis)-, lacking superactinal

plates; interior of rays with resinous lining.

Distribution. Mariana Is, SE Japan, South China Sea,

Philippines, E Indonesia, E and S Australia, New Zealand;

0-540 m.

Remarks. Molecular data are available for P. troughtoni only.

This review is based on morphology. Six species are removed

from Nepanthia to Pseudonepanthia which is distinguished by:

rays not flat actinally, lacking a marginal edge; absence of sec-

ondary plates, pedicellariae and glassy convexities; furrow

spines fewer than 6 per plate; absence of transactinal and super-

actinal plates.

McKnight (in Clark and McKnight, 2001) referred

Nepanthia grangei and N. reinga “provisionally” to Nepanthia.

Material has not been examined, and internal skeletal plates not

observed. Based on the descriptions, the absence of secondary

plates and pedicellariae, and presence of up to five furrow

spines per plate, fit the diagnosis of Pseudonepanthia to which

both species are reassigned.

P. Mark O’Loughlin and Jonathan M. Waters

Pseudopatiria O’Loughlin gen. nov.

Figures 3d, 16g-i

Diagnosis. Rays 5; interradial margin deeply incurved, rays

distinct, unequal in size, subcylindrical, narrowly flat actinally;

margin weakly defined by irregular series of transversely elon-

gate inferomarginal plates; abactinal plates irregular; abactinal

spinelets opaque, digitiform to short columnar, dense subpaxil-

liform clusters; pedicellariae present in papular spaces; papu-

late areas extensive; papular spaces large, numerous secondary

plates and small papulae; actinal interradial plates irregular in

size and arrangement; actinal spines short columnar, dense sub-

paxilliform clusters; lacking superambulacral and superactinal

plates; lacking interior contiguous projections of abactinal and

actinal plates.

Type species. Patiria obtusa Gray, 1847.

Material examined. P. obtusa. Holotype. Panama, 11-18 m, BMNH

1938.6.23.24 (dry; poor condition).

Description. Rays 5, subcylindrical, unequal in length, up to R

= 54 mm; rays narrowly flat actinally, poorly defined margin;

fasciculate pedicellariae between abactinal plates, 2-3 curved

digitiform valves; not fissiparous.

Abactinal surface uneven, with abactinal plates domed and

irregular in size and arrangement on upper rays, in angled series

on sides of rays; papulate areas extensive, to near margin; papu-

lar spaces large, numerous large secondary plates intergrading

with primary plates, numerous papulae; single large

madreporite; disc not bordered by regular series of radial and

interradial plates; abactinal plates crescentiform or elongate or

rounded, with high dome, lacking glassy convexities; spinelets

thick, digitiform to short columnar, distally minutely spinous,

typically 0.4 mm long, in close subpaxilliform clusters of about

40 spinelets per plate; superomarginal and inferomarginal

plates in irregular series, inferomarginals frequently trans-

versely elongate, projecting slightly to define margin.

Actinal interradial plates irregular in size and arrangement.

Actinal spines per plate: oral 6, some thick subspatulate;

suboral, cluster, some thick subspatulate; furrow 3, subspatu-

late; subambulacral and actinal dense clusters of up to 20 per

plate; spines short columnar, typically 0.5 mm long.

Superambulacral, transactinal and superactinal plates

absent; margin not supported by contiguous internal projections

of abactinal and actinal plates; internal resinous body lining.

Distribution. Panama; 11-18 m.

Etymology. From the Latin pseudo (false), with Patiria, refer-

ring to the significant internal skeletal differences with the

genus Patiria.

Remarks. Gray (1866) and Verrill (1870) retained obtusa in

Patiria, but Perrier (1875) referred the species to Asterina.

Verrill (1913) referred it to Callopatiria. A.M. Clark (1983 figs

5a, 7) also discussed the generic placement and noted the

absence of what are referred to in this work as superambulacral

and superactinal plates. A.M. Clark (1993) noted the need for a

reassessment of generic status and opined that the single spec-

imen was “probably from the Pacific side of Panama”. No

molecular data are available, and a morphological reassessment

is undertaken here. The unique characters of this asterinid

genus are: subcylindrical rays; irregular abactinal and actinal

plate arrangements; plates with subpaxilliform dense clusters of

opaque, thick, short columnar spinelets and spines; presence of

pedicellariae; and absence of superambulacral, transactinal and

superactinal plates.

Stegnaster Sladen

Figures 1, 3e, 17a-d

Stegnaster Sladen, 1889: xxxiv, 375, 376 (key). — Fisher, 1911: 254

(key). — Verrill, 1913: 481 (key). — Verrill, 1915: 65. — Spencer and

Wright, 1966: U69. — Clark and Downey, 1992: 194. — A.M. Clark,

1993: 227.— McKnight (in Clark and McKnight), 2001: 161-162.

Diagnosis. Rays 5, subpentagonal, frequently arched, integu-

ment cover, rays with median ridge, body very thin interradial-

ly; abactinal cover of fine granules only; glassy convexities

present; projecting inferomarginal plates lack fringe of

spinelets; projecting furrow spines covered by continuous

granule-covered integument; actinal interradial plates with

thorn-tipped short spines; lacking superambulacral plates,

proximal interradius supported by multiple superactinal plate

struts, wide thin interradius supported internally by downward

projecting abactinal plate extensions meeting actinal plates.

Type species. Pteraster inflatus Hutton, 1872 (original

designation).

Other species. S. wesseli (Penier, 1875).

Material examined. S. inflatus. New Zealand, North Island, AM J 1856

(1); South Island, NMV F95675 (1); no locality data, AM G2030 (1).

Description with species variations. Rays 5, narrowly elevated

median ridge, interradii very thin, form subpentagonal to

rounded, body frequently in high domed (arched) shape, body

covered by integument; size medium ( inflatus up to R = 53

mm) to small ( wesseli up to R = 20 mm); lacking pedice-

llariae; not fissiparous; gonopores abactinal (inflatus), actinal

(wesseli).

Abactinal surface even, continuous cover of granuliform

spinelets partly obscured by integument, larger globose

spinelets on crown of plates (inflatus), spinelets slightly

enlarged around pores (wesseli)', cleared plates with prominent

glassy convexities; plates of radii in longitudinal series, not to

end of ray; carinal series in midray raised, longitudinally

oblong, not notched; plates on upper sides of rays slightly

notched for papulae; secondary plates rare to absent; papulate

areas not extensive, confined to narrow raised radii; papular

spaces large (inflatus) with up to 6 small papulae per space, to

small (wesseli) with single papulae in longitudinal series; dis-

tinct pentagonal disc bordered by 5 raised radial and 5 small

interradial plates; interradial areas extensive, not papulate,

small plates in regular series perpendicular to margin; supero-

marginal and inferomarginal plates subequal, regular series,

granule and integument cover; margin formed by infero-

marginals, without a fringe of spinelets, continuous with actinal

series of plates.

Actinal interradial plates in oblique series.

Genera of asterinid sea-stars

Actinal spines per plate: oral 3; suboral 0; furrow 2-3,

project actinally, continuous integument and granule cover;

sub-ambulacral none to rare; adradial series present; actinal 1

proximally, up to 6 distally, partly obscured by granule-covered

integument; actinal spines short, thick basally, tapering distally

to needle-like glassy point (thorn).

Lacking superambulacral plates; superactinal plates numer-

ous proximally, multiple plate struts between actinal plates and

inward projecting extensions of abactinal plates; superactinals

absent distally, wide thin interradius supported internally by

downward projecting abactinal plate extensions meeting actinal

plates.

Distribution. West Indies, Venezuela, New Zealand, Chatham

Is; 0-183 m.

Remarks. This morphological review is based on material

examined and description of S. wesseli by Clark and Downey

(1992). The phylogenetic trees of Waters et al. (2004) placed

S. inflatus on a clade separate from three species placed in

Paranepanthia but this distinction received only weak support.

Both genera lack superambulacral plates but significant mor-

phological differences reflect the clade separation, such as the

dense subpaxilliform tufts of acicular spines and spinelets in

Paranepanthia. Distinctive characters of Stegnaster are: notice-

able integument and granuliform spinelet cover; furrow spines

projecting actinally and covered by continuous granule-

covered integument; thorn- tipped spines actinally; lacking

superambulacral plates; superactinal plates proximally; wide

thin interradii supported internally by downward projecting

abactinal plate extensions meeting actinal plates.

Tegulaster Livingstone

Figures 3f, 18a-b

Tegulaster Livingstone, 1933: 11-12. — H.L. Clark, 1946: 143. —

Spencer and Wright, 1966: U69.— A.M. Clark, 1993: 228.— Rowe,

1995: 41.

Diagnosis. Rays 5, size small, body thick, covered by thin

integument; rays discrete, form medium to long-rayed stellate,

flat actinally, rays elevated, narrow or broad basally; abactinal

plates thick, imbricating, shallow notch and crescentiform,

irregular in form and arrangement on narrow median band of

upper rays; plates and papulae in few series along sides of rays;

papulae predominantly single per space; secondary plates rare;

abactinal spinelets few to none on plates, glassy, sacciform or

conical or acicular; glassy convexities present; superomarginal

plates in regular series; inferomarginal plates each with up to 7

discrete short spinelets or tuft of acicular spinelets; actinal

interradial plates in oblique series; actinal interradial plates

each with 1-2, long or short, sacciform spines; superambul-

acral plates rare, or in distal series, not in complete series;

superactinal plates present as single plate struts.

Type species. Tegulaster emburyi Livingstone, 1933 (original

designation).

Other species. T. alba (H.L. Clark, 1938); T. leptalacantha (H.L. Clark,

1916) (unrecognised subspecies: Disasterina leptalacantha africana

Mortensen, 1933, this work); T. praesignis (Livingstone, 1933) (junior

synonym: Disasterina spinulifera H.L. Clark, 1938, this work).

Material examined. Asterina alba. Paratypes. Lord Howe I., AM J6170

(5); NMV F45118 (2). Other material. J16575 (1).

T. emburyi. Holotype. Capricorn Group, Queensland, shallow

sublittoral, AM J5605.

Asterina leptalacantha. Holotype. Queensland, Capricorn

Group, AM J3082. Other material. J6097 (1); Mauritius, 24 m, UF

2499 (1).

Disasterina praesignis. Holotype. Queensland, Port Curtis,

AM J5059. Other material. J19314 (3); NMV F94009 (1); F94010

( 1 ).

D. spinulifera. N Western Australia, Lacepede I., WAM Z6773

( 1 ).

Description with species variations. Body thick, covered by

thin integument; rays 5, discrete, flat actinally, high convex

abactinally, narrow at base ( emburyi ) or broad basally tapering

distally to widely (alba) or narrowly (leptalacantha, prae-

signis) rounded; size small (leptalacantha up to R = 25 mm;

alba up to R = 7 mm); pedicellariae present over papulae

(emburyi, 2 conical valves), or not; none fissiparous; gonopores

abactinal or actinal (alba).

Abactinal surface uneven, dominated by projecting proxi-

mal edges of imbricating thick plates; narrow median band of

upper rays plates irregular in form and arrangement; part

carinal series variably evident on upper rays (alba, emburyi,

praesignis)', secondary plates rare (emburyi) or absent (alba), or

primary plates grading from large to small (leptalacantha,

praesignis)', up to 3 series of papulate plates with shallow notch

and crescentiform along each side of rays; papular spaces

small, predominantly single large papulae per space, irregular

on upper rays, series on sides of rays; disc bordered by 5

radial and 5 interradial plates, variably regular; abactinal plates

with spinelet-bearing raised rounded proximal edge (alba) or

domes (leptalacantha)', abactinal spinelets frequently lacking,

or sometimes up to 12, glassy, sacciform to conical, spinelets

across proximal edge of plates (alba), or present distally as tufts

of small sacciform spinelets (emburyi), or with tufts of acicular

sacciform spinelets on distal interradial plates (leptalacantha),

or rarely a few short sacciform spinelets on apex of plates

(praesignis); glassy convexities on bare plates; superomarginal

and inferomarginal plates predominantly subequal, in regular

series; superomarginal plates with a few abactinal-type

spinelets (alba), or with spinelets only in mid-interradius

(emburyi), or bare ( leptalacantha ), or with rare spinelets (prae-

signis); projecting inferomarginal plates with cover of up to

7 short acicular spinelets per plate (alba), or series of up to 6

short spinelets (emburyi, praesignis), or tuft of long acicular

sacciform spinelets (leptalacantha).

Actinal interradial plates in oblique series; proximal inter-

radial non-plated areas sometimes present (leptalacantha,

praesignis).

Actinal spines per plate: oral 6-9; suboral 1 (alba, lep-

talacantha, praesignis) or 2 large, 4 small (emburyi); furrow

4-7, webbed; subambulacral 1-5; adradial actinal spines

present; actinal predominantly 2 (emburyi) or 1; actinal inter-

radial spines sacciform, long or short (alba).

Superambulacral plates rare (alba, emburyi) or series dis-

tally (leptalacantha, praesignis), not in complete series; super-

actinal plates present as single plate struts.

P. Mark O’Loughlin and Jonathan M. Waters

Figure 17. a-d, Stegnaster inflatus (cleared, R = 45 mm, NMV F95675): a, abactinal globose spinelets and granules; b, internal adradial interra-

dius, showing absence of superambulacral plates (left) and presence of superactinal plates (right); c, actinal furrow spines, and granule-covered

integument; d, actinal glassy thom-tipped spines, e-f, Tremaster mirabilis (R = 60 mm, NMV F67741): e, arched body, with abactinal brood cham-

ber pore (arrow); f, ambulacrum showing vertical series of furrow spines.

Distribution. Northern Australia, Lord Howe I., Norfolk I.,

Mauritius, South Africa; 0-366 m.

Remarks. Molecular data are not available and this review is

based on morphological evidence. Two species have been

removed from Disasterina to Tegulaster and the reasons are

discussed under the former. Tegulaster also shares some mor-

phological characters with Indianastra gen. nov. above, in par-

ticular the spination, limited presence of superambulacral

plates and presence of series of superactinal plates. The features

distinguishing Tegulaster from Indianastra are: body thick with

discrete rays, not thin with petaloid to subpentagonal form;

abactinal plates large thick in few irregular series along sides of

rays, not small thin in numerous regular series along rays;

abactinal plates at most slightly notched and crescentiform, not

deeply notched; inferomarginal spinelets discrete, not acicular

in dense integument-covered tufts; actinal interradial spines per

plate up to 2, not small webbed fans; actinal interradial plates

in oblique series, not longitudinal series.

Livingstone (1933) established D. praesignis for a single

specimen (R = 14 mm) from Port Curtis (NE Australia). H.L.

Clark (1938) established D. spinulifera for a single specimen

(R = 8 mm) from Broome (NW Australia), and did not distin-

guish it from D. praesignis. Specimens of both species were

compared and found to be identical; D. spinulifera is a junior

synonym of D. praesignis. Asterina alba has all the diagnostic

characters of Tegulaster. Livingstone (1933) assigned

Disasterina ceylanica to his new genus Tegulaster but it has the

characters of Disasterina. D. leptalacantha var. africana was

established by Mortensen (1933) on the basis of its geo-

graphical separation (South Africa) from the D. leptalacantha

type locality (NE Australia). Material from Mauritius has been

determined as T. leptalacantha and no evidence found to

justify the retention of variety africana.

Tremaster Verrill

Figures 17e-f

Tremaster Verrill, 1879: 201.— Sladen, 1889: xxxiv, 375, 394.—

Spencer and Wright, 1966: U69.— Jangoux, 1982: 155-158,

161-162.— Leeling, 1984: 263-274.— Clark and Downey, 1992:

195-196.— A.M. Clark, 1993: 228.— McKnight (in Clark and

McKnight), 2001: 163.

Diagnosis. Rays 5, subpentagonal, frequently strongly arched;

5 plated ducts between proximal abactinal and proximal actinal

conspicuous interradial openings; abactinal plates with integu-

ment cover and large conical (proximal) and small granuliform

(overall) spinelets; lacking series of superomarginal plates; 4

rows of tube feet in each ambulacrum; furrow spines in vertical

series on adambulacral plates; actinal spines single, tall, slen-

der, spatulate, in oblique series; lacking superambulacral and

superactinal plates, wide thin interradial body supported by

interior distal keels on actinal plates meeting abactinal plates.

Type species. Tremaster mirabilis Verrill, 1879 (original

designation) (junior synonym: Tremaster novaecaledoniae

Jangoux, 1982 proposed by Leeling, 1984 and confirmed in this

work).

Genera of asterinid sea-stars

Figure, 18. Tegulaster alba (R = 5 mm, paratype, NMV F45118): a, abactinal view; b, actinal view.

Material examined. T. mirabilis. Southern Ocean, Heard I., 226-252

m„ NMV F67741 (6), F87362 (1), F67363 (1).

Description. Rays 5; interradial margin straight or slightly con-

vex or concave; body frequently strongly arched; conspicuous

proximal abactinal interradial openings to plate-lined ducts and

chambers, opening actinally distal to oral plates; wide very thin

body distally; size large (up to R = 85 mm); 4 rows of tube feet

in each ambulacrum; lacking pedicellariae; ducts act as brood

chambers.

Appearance dominated by domed form of body, 5 proximal

duct openings, regular fish-scale plates, large conical spinelets

on free edge of proximal plates, small granuliform spinelets on

plates; abactinal proximal plates large, rhombic, thin, irregular

on upper rays, singly notched for papula on upper sides of rays,

distal plates small, covered by integument and weakly attached

small granuliform spinelets; papular spaces small, predomi-

nantly single papula, secondary plates absent; 3-4 longitudinal

series of small papulae along each upper side of rays; proximal

abactinal plates with series of conical spinelets on proximal

edge over papular space; cleared plates lacking conspicuous

glassy convexities; abactinal interradial plates in longitudinal

series, irregular distally, not in series perpendicular to margin;

lacking series of superomarginal plates; acute margin formed

by projecting series of inferomarginal plates; margin fragile,

frequently broken off.

Actinal plates in oblique series, each plate with prominent

proximal spine-bearing dome, series separated by decalcified

body wall.

Actinal spines per plate: oral 1 tall proximally, adjacent pair,

few upper small conical; suboral 0; furrow vertical series of

4-5, short to tall actinally; subambulacral 1 tall; adradial

actinal series present; actinal 1 (rarely 2); actinal spines tall,

slender, spatulate, subsacciform.

Lacking superambulacral plates; wide thin interradial

margin supported by interior distal keels on actinal plates

meeting abactinal plates; lacking superactinal plates.

Distribution. Atlantic, Pacific and Southern Oceans;

150-1060 m.

Remarks. No molecular data have been obtained for

T. mirabilis, and this review is based on morphology. Leeling

(1984) questioned the specific status of T. novaecaledoniae as

its characters fell within the variation seen in T. mirabilis. We

agree with her proposed synonymy. The significantly different

characters of Tremaster from other asterinid genera are: abac-

tinal and actinal duct openings, and internal plated brood

chambers; absence of superomarginal series of plates; four

rows of tube feet in each ambulacrum; furrow spines in vertical

series on adambulacral plates; and extensive thin interradial

margin supported by internal keels on actinal plates.

Incertae sedis

Asterina lorioli Koehler

Asterina lorioli Koehler, 1910: 129-131, pi., 19 figs 5-8. — A.M.

Clark, 1993: 211.

Remarks. The description and illustrations by Koehler (1910)

of abactinal granuliform spinelets and actinal spination suggest

that A. lorioli is not a species of Asterina. Without material we

have not been able to examine the internal skeleton.

Asterina novaezelandiae Perrier

Asterina novaezelandiae Perrier, 1875: 308. — Koehler, 1920:

135-136, pi. 35 figs 8-9.

Patiria novaezelandiae. — Verrill, 1913: 482.

Remarks. The single type specimen of A. novaezelandiae was

reported by Perrier (1875) from New Zealand but A.M. Clark

(1993) noted “holotype originally with specimens of Patiriella

gunnii supposedly from New Zealand but almost certainly from

southern Australia”. If this is the case, it may be a five-rayed

form of a species of Meridiastra. The description of three fur-

row spines and three subambulacral spines suggests

Meridiastra medius (O’Loughlin et al., 2003). The specimen

could not be found.

Nepanthia brachiata Koehler

Nepanthia brachiata Koehler, 1910: 133-135, pi., 19 figs 14-15. —

Fisher, 1917: 173. — Fisher, 1919: 420. — Clark and Rowe, 1971: 70. —

A.M. Clark, 1993: 222.

P. Mark O’Loughlin and Jonathan M. Waters

Remarks. Fisher (1917, 1919) and Clark and Rowe (1971)

referred N. brachiata to Paranepanthia. The type specimen

could not be found. Based on the description and figures by

Koehler (1910), N. brachiata is characterised by: discrete nar-

row rays, flat actinally with narrow thin margins, not sub-

cylindrical; absence of two distinct ‘fields’ of abactinal plates;

predominantly large single papulae in papular spaces; few

suboral, subambulacral and actinal interradial spines; webbed

combs of digitiform actinal spines. This combination of char-

acters is not found in Nepanthia or Paranepanthia or

Pseudonepanthia. Without material we have not been able to

examine the internal skeleton.

Biogeography

Waters et al. (2004) noted a link between the phylogeny sug-

gested by molecular evidence and biogeographic patterns. The

shallow marine genera Paranepanthia (clade I) in southern

Australia and New Zealand, Patiria (clade II) in the northern

and eastern Pacific, Meridiastra (clade III) in southern

Australia and New Zealand, Aquilonastra and Cryptasterina

(clade IV) in the northern Indo-West Pacific, Parvulastra (clade

V) in the southern temperate waters from South Africa to South

America, and Asterina (clade VI) in the north-eastern and

southern coastal Atlantic and Mediterranean, all have localised

distributions. Additional species added to these genera on mor-

phological grounds do not alter this pattern. Other genera, not

included in the molecular phylogeny, Asterinides in the West

Indies, Calloptiria in southern Africa, Disasterina and

Indianastra and Pseudasterina and Tegulaster in the northern

Indo-West Pacific, Nepanthia and Pseudonepanthia around

Australia and the Indo-West Pacific, and Patiriella in southern

Australasia and the north-west Indian Ocean, are likewise

regionally localised.

The remaining genera with more widespread distributions,

Anseropoda in the Atlantic and Indo-Pacific, Stegnaster in the

West Indies and New Zealand, and Tremaster in the Atlantic

and Pacific and Southern Oceans, show significant morpho-

logical differences from the other asterinid genera and all occur

in deeper water. Anseropoda also shows morphological

evidence of possibly being paraphyletic.

It can be concluded that Asterinidae is a cosmopolitan

family, mainly of shallow-water narrow-range genera but

including some more widespread in deeper waters of all

oceans.

Acknowledgements

We have drawn extensively on the work of Ailsa M. Clark, and

express our gratitude for her invaluable contribution to asteroid

systematics. We were graciously given access to an unpub-

lished draft manuscript on a morphological review of the

genera of Asterinidae at the Australian Museum during the

1980s and are grateful to Frank Rowe and Penny Berents for

this informative work and permission to use some of the pho-

tos. And we are grateful to the many persons who generously

assisted us with collecting, donations, loans, advice and tech-

nical skill: Cynthia Ahearn (USNM); Nadia Ameziane and

Marc Eleaume (MNHN); Penny Berents (AM); Ben Boonen

(composition of the tables and figures); Maria Byrne

(University of Sydney); Andrew Cabrinovic and Miranda Lowe

(BMNH); Mieko Komatsu (Toyama University); Eric Lazo-

Wasem (YPM); Yulin Liao (formerly of the Institute of

Oceanology, Qingdao); Susie Lockhart and Rich Mooi

(California Academy of Sciences); Chris Mah (University of

Illinois Urbana-Champaign); Loisette Marsh, Mark Salotti and

Jane Fromont (WAM); Claus Nielsen (ZMUC); Mark Norman

and Chris Rowley (photography, NMV); Tim O’Hara and Gary

Poore (NMV); Gustav Paulay (UF); Frank Rowe (Research

Associate of the Australian Museum); Ahmed Thandar

(University of Durban- Westville, South Africa); Liz Turner

(TM); and Rick Webber (MNZ).

References

Avise, J.C. 2000. Phylogeography. The history and formation of

species. Harvard University Press: Cambridge.

Avise, J. C., Nelson, W. S., and Sugita, H. 1994. A speciational history

of “living fossils”: molecular evolutionary patterns in horseshoe

crabs. Evolution 48: 1986-2001.

Aziz, A., and Jangoux, M. 1985. Description de six asterides nouveaux

(Echinodermata) de la region des Philippines. Indo-Malayan

Zoology 2: 281-291, pis 1-8.

Bell, F.J. 1891. Stray notes on the nomenclature etc. of some British

starfishes. Annals and Magazine of Natural History (6) 7: 233-235.

Bemasconi, I. 1973. Asteroideos Argentinos. VI. Familia Asteridinae.

Revista del museo Argentino de ciencias naturales

Bernardino Rivadavia e instituto nacional de investigacion de las

ciencias naturales 3(4): 335-346, 2 pis.

Campbell, A.C., and Rowe, F.W.E. 1997. A new species in the asterinid

genus Patiriella (Echinodermata, Asteroidea) from Dhofar, south-

ern Oman: a temperate taxon in a tropical locality. Bulletin of the

British Museum, (Natural History) ( Zoology ) 63(2): 129-136.

Clark, A.H. 1916. Seven new genera of echinoderms. Journal of the

Washington Academy of Science 6: 115-122.

Clark, A.M. 1956. A note on some species of the family Asterinidae

(class Asteroidea). Annals and Magazine of Natural History (12) 9:

374-383, 4 figs, 2 pis.

Clark, A.M. 1983. Notes on Atlantic and other Asteroidea. 3. The

families Ganeriidae and Asterinidae, with description of a new

asterinid genus. Bulletin of the British Museum (Natural History)

(Zoology) 45(7): 359-380.

Clark, A.M. 1993. An index of names of recent Asteroidea — Part 2:

Valvatida. Echinoderm Studies 4: 187-366. Balkema: Rotterdam.

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

ern Africa. British Museum (Natural History) Publication No. 776.

London. 277 pp.

Clark, A.M., and Downey, M.E. 1992. Starfishes of the Atlantic.

Chapman and Hall (Natural History Museum Publications):

London. 794 pp, 75 figs, 113 pis.

Clark, A.M., and Mah, C. 2001. An index of names of recent

Asteroidea — Part 4: Forcipulatida and Brisingida. Echinoderm

Studies 6: 229-347. Balkema: Rotterdam.

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

Indo-west Pacific echinoderms. British Museum (Natural History):

London, vii+238 pp, 100 figs, 31 pis.

Clark, H.E.S., and McKnight, D.G. 2001. The marine fauna of New

Zealand: Echinodermata: Asteroidea (sea-stars). NfWA Biodiversity

Memoir 117: 1-269, 89 pis.

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

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

Genera of asterinid sea-stars

Clark, H.L. 1928. The sea-lilies, sea-stars, brittle stars and sea-urchins

of the South Australian Museum. Records of the South Australian

Museum 3(4): 361-482, figs 108-142.

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

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

Comparative Zoology at Harvard College 55: 1-596, 28 pis, 63

figs.

Clark, H.L. 1946. The echinoderm fauna of Australia. Its composition

and its origin. Carnegie Institution of Washington Publication 566:

iv+567 pp.

Dartnall, AJ. 1970. The asterinid sea stars of Tasmania. Papers and

Proceedings of the Royal Society of Tasmania 104: 73-77, 1 pi.

Dartnall, A.J. 1971. Australian sea stars of the genus Patiriella

(Asteroidea, Asterinidae). Proceedings of the Linnean Society of

New South Wales 96(1): 39-49, fig. 1, pis 3-4.

Dartnall, A. 1980. Fauna of Tasmania Handbook 3. Tasmanian

echinoderms. University of Tasmania: Hobart. 82 pp., 36 figs, 5 pis,

18 maps.

Dartnall, A.J., Byrne, M., Collins, J., and Hart, M.W. 2003. A new

viviparous species of asterinid (Echinodermata, Asteroidea,

Asterinidae) and a new genus to accommodate the species of

pan-tropical exiguoid sea stars. Zootaxa 359: 1-14.

De Queiroz, K., and Gauthier, J. 1992. Phylogenetic taxonomy. Annual

Review of Ecology and Systematics 23: 449-480.

Doderlein, L. 1888. Echinodermen von Ceylon. Zoologische

Jahrbucher Supplement 3(6): 822-846, pis 31-33.

Domantay, J.S., and Acosta, T.E. 1970. The littoral echinoderm fauna

of Ilocos Sur between Candon and Vigan. Acta Manilana A5(10):

49-103.

Fisher, W.K. 1906. The starfishes of the Hawaiian Islands. Bulletin of

the United States Fish Commission 23: 987-1130, 49 pis.

Fisher, W.K. 1908. Necessary changes in the nomenclature of star-

fishes. Smithsonian Miscellaneous Collections 52: 87-93.

Fisher, W.K. 1911. Asteroidea of the North Pacific and adjacent waters.

1. Phanerozonia and Spinulosa. Bulletin of the United States

National Museum 76: vi+419 pp, 120 pis.

Fisher, W.K. 1917. A new genus and subgenus of East-Indian seastars.

Annals and Magazine of Natural History 20 (8) 116: 172-173.

Fisher, W.K. 1919. Starfishes of the Philippine seas and adjacent

waters. Bulletin of the United States National Museum 100(3): i-xi,

1-712, 156 pis.

Fisher, W.K. 1940. Asteroidea. Discovery Reports 20: 69-306, 23 pis.

Fisher, W.K. 1941. A new genus of sea-stars ( Plazaster ) from Japan,

with a note on the genus Parasterina. Proceedings of the United

States National Museum 90: 447-456, figs 20-22, pis 66-70.

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

Hypostoma (Asterias Linnaeus ). Annals and Magazine of Natural

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

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

Asteriadae. Proceedings of the Zoological Society of London 1847

(15): 72-83.

Gray, J.E. 1866. Synopsis of the species of starfish in the British

Museum. John van Voorst: London, iv+17 pp., 16 pis.

Hart, M.W., Byme, M., and Johnson, S.L. 2003. Patiriella pseudo-

exigua (Asteroidea: Asterinidae): a cryptic species complex

revealed by molecular and embryological analyses. Journal of the

Marine Biological Association of the United Kingdom 83:

1109-1116.

Hart, M.W., Byrne, M. and Smith, M.J. 1997. Molecular phylogenetic

analysis of life-history evolution in asterinid starfish. Evolution 51:

1846-1859.

Hayashi, R. 1940. Contributions to the classification of the sea-stars of

Japan. 1. Spinulosa. Journal of the Faculty of Science Hokkaido

Imperial University ( Zoology ) (6)7(3): 107-204, pis 7-13, 63 figs.

Hayashi, R. 1973. The sea-stars ofSagami Bay. Biological Laboratory,

Imperial Household: Japan, viii+114 pp., 18 pis.

Jangoux, M. 1982. On Tremaster Verrill 1879, an odd genus of recent

starfish (Echinodermata: Asteroidea). Pp. 155-163, 4 pis in:

Lawrence, J.M. (ed.), Echinoderms: Proceedings of the

International Conference, Tampa Bay. Balkema: Rotterdam.

Jangoux, M. 1985. Catalogue commente des types d’Echinodermes

actuels conserves dans les collections nationales suisses, suivi

d’une notice sur la contribution de Louis Agassiz a la connaissance

des echinodermes actuels. Museum d’Histoire Naturelle: Geneve.

67 pp.

Koehler, R. 1910. An account of the shallow-water Asteroidea.

Echinoderma of the Indian Museum. 1-183, pis 1-20. Calcutta:

Trustees of the Indian Museum.

Koehler, R. 1911. Description de quelques Asteries nouvelles. Revue

Suisse de Zoologie 19(1): 1-21, pi. 1.

Koehler, R. 1920. Echinodermata: Asteroidea. Scientific Reports of the

Australasian Antarctic Expedition. C 8(1): 308 pp., 75 pis.

Leeling, B. 1984. Ziir Morphologie und Systematik der Gattung

Tremaster Verrill 1879 (Echinodermata, Asteroidea). Mitteilungen

aus dem Hamburgischen Zoologischen Museum und Institut 81:

261-276.

Liao, Y. 1980. The echinoderms of Xisha Islands, Guangdong

Province, China. 4. Asteroidea. Studia Marina Sinica 17: 153-171,

6 pis. (In Chinese, with English summary.)

Liao, Y. and Clark, A.M. 1995. The echinoderms of southern China.

614 pp, 23 pis. Science Press: Beijing.

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

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

Mah C. L. 2000. Preliminary phylogeny of the forcipulatacean

Asteroidea. American Zoologist 14: 375-381.

Marsh, L.M. 1977. Coral reef asteroids of Palau, Caroline Islands.

Micronesica 13(2): 251-281.

Matsuoka, N. 1981. Phylogenetic relationships among five species of

starfish of the genus, Asterina: an electrophoretic study. Journal of

Comparative Biochemistry and Physiology 70B: 739-743.

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

Ophiuroidea). Videnskabelige Meddelelser fra Dansk naturhistorisk

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

Mortensen, Th. 1940. Echinoderms from the Iranian Gulf. Danish

Scientific Investigations in Iran 2: 55-112, pis 1, 2. Ejnar

Munksgaard: Copenhagen.

Nardo, J.D. 1834. De Asteriis. Pp. 716-717 in: Oken, L. (ed.), Isis 1.

O’Hara, T.D. 1998. Systematics and biology of Macquarie Island

echinoderms. Memoirs of Museum Victoria 57(2): 167-223.

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

and Pacific Asterinidae (Echinodermata, Asteroidea). Memoirs of

Museum Victoria 59(2): 277-296.

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

new species of Patiriella from New Zealand, and review of

Patiriella regularis (Echinodermata, Asteroidea) based on morpho-

logical and molecular data. Journal of the Royal Society of New

Zealand 32(4): 697-7 11.

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

morphological review of the asterinid, Patiriella gunnii (Gray)

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

181-195.

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

d’Histoire Naturelle de Paris. Paris. 384 pp.

Rowe, F.W.E., and Gates, J. 1995. Echinodermata. Pp. xiii+510 in:

Wells, A. (ed.), Zoological Catalogue of Australia 33. CSIRO:

Melbourne, [as “Rowe (1995)” throughout paper]

Rowe, F.W.E., and Marsh, L.M. 1982. A revision of the asterinid genus

Nepanthia Gray 1840 (Echinodermata: Asteroidea), with the

P. Mark O’Loughlin and Jonathan M. Waters

description of three new species. Australian Museum Memoir 16:

89-120, figs 1-6.

Shepherd, S.A. 1968. The shallow water echinoderm fauna of South

Australia. 1. The asteroids. Records of the South Australian

Museum 15(4): 729-756.

Sladen, W.P. 1889. Asteroidea. Report on the scientific results of the

voyage of the H.M.S. Challenger during the years 1873-76.

Zoology 30: 1-935, 118 pis.

Smith, G.A. 1927. A collection of echinoderms from China. Annals

and Magazine of Natural History 9 (20) 34: 272-279.

Spencer, W.K., and Wright, C.W. 1966. Asterozoans. Pp. U4-U107, 89

figs in: R.C. Moore (ed.), Treatise on Invertebrate Paleontology.

The Geological Society of America and The University of Kansas

Press.

Tommasi, L.R. 1970. Lista dos asteroides recentes do Brasil.

Contrgoes Instituto Oceanografico da Universidade Sao Paulo 18:

1-61, 60 figs.

Tortonese, E. 1965. Echinodermata. Fauna d’ltalia 6: xv+422 pp.

Bologna.

VandenSpiegel, D., Lane, D.J.W., Stampanato, S. and Jangoux, M.

1998. The asteroid fauna (Echinodermata) of Singapore, with a dis-

tribution table and an illustrated identification to the species. Raffles

Bulletin of Zoology 46(2): 431-470.

Verrill, A.E. 1870. Notes on the Radiata in the Museum of Yale

College. 2. Notes on the echinoderms of Panama and west coast of

America, with descriptions of new genera and species. Transactions

of the Connecticut Academy of Arts and Sciences [1867 to 1871]

1(2): 251-322.

Verrill, A.E. 1879. Notice of recent additions to the marine

Invertebrata, of the northeastern coast of America, with descriptions

of new genera and species and critical remarks on others.

Echinodermata. Proceedings of the United States National Museum

2: 201-204.

Verrill, A.E. 1913. Revision of the genera of starfishes of the subfami-

ly Asterininae. American Journal of Science (4) 35 (209): 477^85.

Verrill, A.E. 1914. Monograph of the shallow- water starfishes of the

North Pacific coast from the Arctic Ocean to California. Harriman

Alaska Series, Smithsonian Institution 14: 1 — 408, 110 pis.

Verrill, A.E. 1915. Report on the starfishes of the West Indies, Florida

and Brazil. Bulletins from the Laboratories of Natural History,

University, Iowa City, Iowa 7(1): 1-232, 29 pis.

Waters, J.M., O’Loughlin, P.M., and Roy, M.S. 2004. Molecular sys-

tematics of some Indo-Pacific asterinids (Echinodermata,

Asteroidea): does taxonomy reflect phylogeny? Molecular

Phylogenetics and Evolution 30: 872-878.

Memoirs of Museum Victoria 61(1): 41-45 (2004)

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

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

A new genus of millipedes (Diplopoda: Polydesmida: Dalodesmidae) from wet

forests in southern Victoria, with brief remarks on the Victorian Polydesmida

Robert Mesibov

Queen Victoria Museum and Art Gallery, Launceston, Tasmania 7250, Australia (mesibov@southcom.com.au)

Abstract Mesibov, R. 2004. A new genus of millipedes (Diplopoda: Polydesmida: Dalodesmidae) from wet forests in southern

Victoria, with brief remarks on the Victorian Polydesmida. Memoirs of Museum Victoria 61(1): 41-45.

Victoriombrus gen. nov. is erected for Victoriombrus acanthus sp. nov. from the Otway Ranges and V. seminudus sp.

nov. from the Mt Donna Buang area. Four species of Dalodesmidae and 15 species of Paradoxosomatidae have now been

described from Victoria. A diverse fauna of non-paradoxosomatid Polydesmida remains to be described from Victorian

wet forests.

Keywords Diplopoda, Polydesmida, Dalodesmidae, millipede, Australia, Victoria

Introduction

At the time of writing, there are 18 valid names for native poly-

desmidan millipedes known or believed to occur in Victoria

(Table 1). Sixteen of these taxa are in Paradoxosomatidae, a

family with numerous representatives in the dry forests and

woodlands of eastern and western Australia. Paradoxoso-

matids also occur in Victoria’s wetter forests, where they

share moist microhabitats with Polydesmida most closely

related to forms found in the wet forests of Tasmania and New

Zealand. This non-paradoxosomatid, wet-forest polydesmidan

fauna is very poorly known, yet collections by the author and

others have shown it to be diverse and to include undescribed

species in the genera Asphalidesmus Silvestri, 1910,

Lissodesmus Chamberlin, 1920 and Paredrodesmus Mesibov,

2003.

In this paper I describe two wet-forest species in a

genus with a novel gonopod structure. Victorian species of

Lissodesmus will be reviewed in a forthcoming paper.

All material examined has been preserved in 75-80%

ethanol. Preliminary drawings on graph paper were made

using an eyepiece graticule. Gonopods were cleared and tem-

porarily mounted in 60% lactic acid; other body parts were

temporarily mounted in glycerine jelly. A Philips Electroscan

ESEM 2020 operated in high-vacuum mode was used to

examine material which had been air-dried before sputter-

coating with gold. SEM images were acquired digitally.

Abbreviations are as follows: NMV, Museum Victoria,

Melbourne, Vic.; QVM, Queen Victoria Museum and Art

Gallery, Launceston, Tas.

Order Polydesmida Leach, 1815

Suborder Dalodesmidea Hoffman, 1980

Dalodesmidae Cook, 1896

Victoriombrus gen. nov.

Type species. Victoriombrus acanthus sp. nov., by present

designation.

Diagnosis. Dalodesmids c. 15 mm long with head + 20 seg-

ments, rounded and somewhat swollen paranota and pore

formula 5, 7, 9-19. Telopodite of gonopod short, wide, divided

by a transverse constriction into a setose, posteromesally exca-

vated basal portion and a bare distal portion bearing a small

mesal branch comprising a long, tapering solenomere accom-

panied by a distally flattened and roughened process, a much

larger lateral branch, and between the two a narrow, more or

less rod-like process arising posteromesally.

Description. Head with moderately deep impression lateral to

antennal base; bases separated by about twice a socket dia-

meter; vertigial sulcus faintly indicated; head moderately setose

with moderately long setae on clypeus and frons, a few long

setae on the vertex. Antenna (Fig. 1) short, slender; anten-

nomere 2 the longest, 6 the widest. Collum about as wide as

head and second somite; sparsely setose with moderately long

setae; posterior margin straight, anterior margin gently convex,

corners rounded. Somite 2 with lateral margin of paranotum

well below collum corner, somite 3 margin at about the level of

the collum comer. Somites with well-defined waist (Fig. 1); a

Robert Mesibov

Table 1. Native Polydesmida from Victoria.

Taxon Published localities

Dalodesmidae

Gephyrodesmus cineraceus Jeekel, 1983

Lissodesmus martini (Carl, 1902)

Paradoxosomatidae

Archicladosoma magnum Jeekel, 1984

Australiosoma laminatum Jeekel, 1984

Cladethosoma forceps (Verhoeff, 1941)

Dicranogonus pix Jeekel, 1982

Hoplatessara nigrocingulata Jeekel, 1984

H. pugiona Verhoeff, 1941

Hoplatria clavigera Verhoeff, 1941

Isocladosoma guttatum Jeekel, 1984

I. maculatum Jeekel, 1984

I. pallidulum Jeekel, 1984

Pogonosternum adrianae Jeekel, 1982

P. coniferum Jeekel, 1965

P. laetificum Jeekel, 1982

P nigrovirgatum infuscum Jeekel, 1982

P. nigrovirgatum nigrovirgatum (Carl, 1902)

Somethus biramus Jeekel, 1984

Drummer State Forest, 15 km E of Cann River (Jeekel, 1983)

Melbourne (Carl, 1902); Femtree Gully National Park, 18 km NNE of Dandenong

(Jeekel, 1983); Cockatoo Creek (Johns, 1964).

Australia (Jeekel, 1984; 1 ); Ferntree Gully National Park, 18 km NNE of Dandenong

(Jeekel, 1984; tentative identification)

Australia (Jeekel, 1984)1

Gippsland (Verhoeff, 1941)

13 km SE of Buchan; 4 km ESE of Bmthen; Mt Taylor, 1 1 km NNW of Baimsdale

(Jeekel, 1982)

Australia (Jeekel, 1984)1

Whittlesea (Verhoeff, 1941) 2 ; Belgrave (Jeekel, 1984)

Australia (Jeekel, 1984)1; Gippsland (Verhoeff, 1941)

Glenaladale National Park, 28 km WNW of Baimsdale (Jeekel, 1984)

Australia (Jeekel, 1984)1

Gunyah Gunyah, 32 km SSW of Morwell (Jeekel, 1984)

Toongabbie, 18 km NNE of Traralgon (Jeekel, 1982)

Australia (Jeekel, 1965, 1982)1

Femtree Gully National Park, 18 km NNE of Dandenong (Jeekel, 1982)

Mt Taylor, 11 km NNW of Baimsdale (Jeekel, 1982)

Melbourne (Carl, 1902)

Drummer State Forest, 15 km E of Cann River; 13 km SE of Buchan; also NSW and SA

(Jeekel, 1984)

1 Jeekel (1982, 1984) believed that the type specimens of these species are from Victoria because they were collected by Victorian naturalist

Charles Barrett and, in the case of Isocladosoma and Pogonosternum spp., because congeners are known from Victoria.

2 Verhoeff (1941: 14) gives “Whettlerea bei Melbourne”, probably an error for ‘Whittlesea’, c. 40 km north of Melbourne.

few setae on somite 2, remaining somites bare and apparently

smooth (very small seta-like processes scattered over dorsal

surface of metazonite, only visible under high magnification);

paranota on most so mi tes just above midlateral (Fig. 1), some-

what swollen, laterally with a very narrow, upwardly concave

groove bordered by a slightly thickened margin; from above,

anterior margin of paranotum gently curved, lateral margin

very slightly convex, posterior margin slightly concave on

anterior somites, more so on posterior somites, where the pos-

terior corner projects slightly and bluntly; limbus a comb of

fine, straight teeth. Ozopores small, opening dorsolaterally just

above and near the posterior end of the paranotal groove on

somites 5, 7 and 9-19. Stemites slightly longer than wide, cross

impressions well-defined. Anterior legs (Fig. 1) with swollen

podomeres, prefemur strongly arched dorsally; posterior legs

less swollen; tarsus about as long as next longest podomere, the

femur. Sphaerotrichomes on tibia and tarsus only, from leg 3

rearwards, diminishing in number on the more posterior legs

and absent from the last leg. Dense ‘brushes’ of setae ventrally

on prefemur, femur and postfemur of most legs from 3 rear-

wards. Leg 2 coxa somewhat extended mesally, the genital

opening a simple pit on this extension. Legpairs 4 and 5 not

separated at bases, legpairs 6 and 7 well-separated; flexed

gonopod telopodites reaching to legpair 6. Spiracles simple in

structure with slightly raised rims; on diplosegments the anter-

ior spiracle just above and anterior to the anterior leg, the pos-

terior spiracle just anterior to the midway point between the leg

bases. Preanal ring moderately setose; epiproct tapering to

blunt, rounded and depressed tip extending well past anal

valves; hypoproct paraboloid in outline.

Gonopod aperture ovoid (Figs 2, 4), wider than long, about

one-half width of somite 7 prozonite, with posterior margin in

the form of a flattened ‘V’and posterolateral portions of rim

projecting ventrally. Gonocoxa in overall form a truncated

cone, tapering distally, with moderately long setae on pos-

teromesal surface; gonocoxae fairly firmly joined along mid-

line but not quite fused into a syncoxite. Cannula prominent.

Gonopod telopodites (Figs 2, 4), short, wide, closely appressed

but not joined; partly fused with gonocoxa on anterior side of

base; basal portion clearly separated from branched distal

portion by a transverse constriction; basal portion excavated

posteromesally, sparsely setose, mainly posterolaterally; distal

portion bare, divided into a small mesal branch with a long,

tapering solenomere joined basally to a distally flattened and

roughened process, a much larger lateral branch, and a narrow,

more or less rod-like posterior process arising posteromesally.

Females somewhat larger than males, with the same colour-

ing; legs smaller than corresponding male legs and not swollen;

posterior rim of epigynum projecting slightly, more so in

centre; cyphopods not examined.

Distribution. Southern Victoria.

Etymology. Victoria + Greek ombros (“rainshower”),

masculine.

New genus of millipedes

Remarks. It is not clear how Victoriombrus is related to other

Australian dalodesmid genera. The new genus shares with

Tasmanian Bromodesmus (Mesibov, 2004) a long, tapering,

acutely pointed solenomere and telopodite elements fringed

with teeth, but the gonocoxae in Victoriombrus are more

nearly fused into a syncoxite than in the Tasmanian genus, the

telopodites are much shorter and wider, and the pore formula is

5, 7, 9-19 instead of 5, 7, 9, 10, 12, 13, 15-19.

Victoriombrus acanthus sp. nov.

Figures 1, 2, 3, 6 (map)

Material examined. Holotype. Male, Australia, Victoria. Young Creek

Road, 0.2 km NE of Ciancio Creek crossing, 38°40’S, 143°29'E,

pitfall 15.xi. 1994-313.1995, G. Milledge, Nothofagus forest, sample

NOH-1079, NMV K-8842.

Paratypes. 1 stadium 7 juvenile male, same details as holotype,

NMV K-8843; 2 females, Aire Crossing Track, 0.5 km N of Aire River

crossing, 38°40'S, 143°29’E, pitfall 15ju.1994-31.U995, G. Milledge,

Nothofagus forest, sample NOH-1084, NMV K-8844, K-8857; 1 male,

same details but pitfall 31.i-ll.iv.1995, NMV K-8845; 2 males,

1 female, Maits Rest, 10 km W of Apollo Bay, 22.x. 1991, K. Walker,

Nothofagus litter and moss, NMV K-8846, K-8874, K-8875; 1 male,

Beauchamp Falls, 38°39'S, 143°36'E, pitfall 6.ix.-15.xi.l994,

G. Milledge, Nothofagus forest, sample NOH-1089, NMV K-8847;

2 females, same details but pitfall 15.xi. 1994-313. 1995, sample

NOH-1090, NMV K-8848, K-8858; 5 males, Turtons Pass,

38°38'39"S, 143 o 41'20"E, 12.xii.2003, R. Mesibov and T. Moule, wet

eucalypt forest litter, NMV K-8849, K-8859-K8862; 2 females, 9

stadium 7 juvenile males, 1 stadium 7 juvenile female, same details,

NMV K-8851, K-8863-K8873; 1 stadium 7 juvenile male, same

details but 38°38'43"S, 143°40'36 M E, NMV K-8850.

Diagnosis. Differs from V. seminudus sp. nov. in the form of the

gonopod, notably the posterior curvature of the solenomere and

the presence of small, branched, spiny processes on the lateral

division of the telopodite.

Description. Males c. 15 mm long, c. 1.2 mm in maximum

vertical diameter. In alcohol, freshly collected specimens with

pale yellow head and antennae; ground colour of body pale

yellow, patterned with brown patches: prozonite with a patch

laterally and a pair of paramedian patches dorsally, metazonite

with a triangular mid-dorsal patch anteriorly, a quadrangular

mid-dorsal patch posteriorly, a small patch anteroventrally on

paranotum; posterior margin of metazonite and distal

podomeres brown.

Gonopod telopodites (Figs 2, 3) short, wide, closely

appressed basally but not joined. Basal portion of telopodite

excavated posteromesally, with variably long setae on postero-

lateral surface and surrounding the excavate area. Telopodite

with a transverse constriction at between one-third to one-half

its length, separating setose basal portion from bare distal

portion, the latter divided basally into mesal, posteromesal and

lateral branches. Mesal branch divides basally into a long,

tapering, acutely pointed solenomere and a long, flattened

process arising posterior to solenomere. Solenomere curves

first anteriorly and basally, then turns sharply distally, curving

slightly posteriorly but remaining parallel to long axis of

telopodite, terminating at about two-thirds telopodite length.

Prostatic groove running along mesal surface of telopodite,

following curvature of the solenomere and terminating at its

tip. Laterally flattened process accompanying the solenomere

curves first mesally, then anteriorly, terminating just distal to

solenomere; tip of process and a somewhat expanded portion at

about mid-length with numerous marginal teeth and a rough-

ened surface. Just distal and posterior to solenomere origin,

posteromesal process arising as a narrow, rod-like, tapering

process, laterally flattened from about half its length, curving

slightly mesally, then laterally. Lateral branch of telopodite by

far the largest, complicated in structure (Figs 2, 4), basically a

laterally flattened process, curving slightly laterally and some-

what concave mesally, with marginal teeth of varying sizes;

posterior edge giving rise to a mesally curving process ending

in a rounded, somewhat expanded tip resembling a spanner or

wrench midway between solenomere origin and tip; lateral and

anterior surfaces of main process armed at about one-third the

process length with small, short, branched, spiny structures

resembling the tips of a thorny shrub.

A B

Figure 1. Victoriombrus spp. details. A, V. acanthus sp. nov., male

paratype from NMV K-8849; B, V. seminudus sp. nov., male paratype

from NMV K-8856. 1, Antenna; 2, posterior leg on somite 6; 3, poster-

ior leg on somite 12; 4, dorsal outline view of somite 12; 5, posterior

outline view of somite 12. Scale bar =1.0 mm; setation and antennal

cones not shown.

Robert Mesibov

Figure 2. Victoriombrus acanthus sp. nov., male, paratype. Posterior

view of gonopods in situ (specimen from NMV K-8849). Scale

bar = 0.5 mm.

Figure 3. Victoriombrus acanthus sp. nov., male, paratype. Anterior

(left) and mesal (right) views of right gonopod (specimen from NMV

K-8849 ). Dashed line indicates course of prostatic groove; for clarity,

the continuation of the groove to the solenomere tip is not indicated.

Figure 5. Victoriombrus seminudus sp. nov., male, paratype. Anterior

(left) and mesal (right) views of right gonopod (specimen from NMV

K-8856). Dashed line indicates course of prostatic groove.

New genus of millipedes

Figure 6. Localities in Victoria of Victoriombrus acanthus sp. nov.

(crosses) and V. seminudus sp. nov. (squares).

Distribution and microhabitat. In wet eucalypt and Nothofagus

forest litter in the Otway Ranges (Fig. 6).

Etymology. Latin acanthus (“prickly shrub”), noun in

apposition, from the appearance of the gonopod telopodite.

Victoriombrus seminudus sp. nov.

Figures 1, 4, 5, 6 (map)

Material examined. Holotype. Male, Australia, Victoria. Myrtle Gully

Reserve, 3.4 km WSW of Mt Donna Buang, 37°43'S, 145°38'30"E,

pitfall 29.xi. 1994-203.1995, G. Milledge, Nothofagus forest, sample

NOH-1840, NMV K-8852.

Paratypes. 1 male, 7 females, 1 stadium 7 juvenile female, same

details as holotype, K-8853, K-8876- K-8883; 1 male, same details but

pitfall 2Li.-7.iv.1995, sample NOH-1841, NMV K-8854; 2 males,

2 females, Road 26, 0.2 km WNW of Donna Buang Road junction,

37°43'S, 145°39'30"E, pitfall 29.xi.1994 to 20L1995, G. Milledge,

Eucalyptus forest, sample NOH-1846, NMV K-8855, K-8884-

K-8886; 1 male, Acheron Gap, 6 km NE of Mt Donna Buang,

37°40'43"S, 145°44'20"E, pitfall 28.xii.1995-21.ii.1996, G. Milledge,

Nothofagus forest, sample NOH-1853, NMV K-8856.

Diagnosis. Differs from V. acanthus sp. nov. in the form of the

gonopod, notably the anterior curvature of the solenomere and

the absence of small, branched, spiny processes on the lateral

division of the telopodite.

Description. Males c. 15 mm long, 1.2 mm in maximum ver-

tical diameter. In alcohol, well-coloured specimens with pale

yellow antennae and yellow-orange head, legs and paranota;

prozonite and metazonite ground colour pinkish-brown; meta-

zonite reddish-brown below paranota and with mid-dorsal, tri-

angular reddish-brown patch; posterior margin of metazonite

and distal podomeres reddish-brown.

Gonopod telopodites (Figs 4, 5) short, wide, closely

appressed basally but not joined. Basal portion of telopodite

excavated posteromesally, with variably long setae on postero-

lateral surface. Telopodite with a transverse constriction at

between one-third to one-half its length, angled distad from lat-

eral to mesal, separating setose basal portion from bare distal

portion, latter divided basally into mesal, posteromesal and

lateral branches. Mesal branch divides basally into a long,

tapering, acutely pointed solenomere and a long, flattened

process arising posterior to solenomere. Solenomere extends

distally, then turns abruptly anteriorly at about three-quarters

the length of the telopodite before terminating in a short, dis-

tally directed tip. Prostatic groove running along mesal surface

of telopodite, following curvature of the solenomere and

terminating at its tip. Laterally flattened process accompany-

ing the solenomere directed anterodistally and slightly

mesally, expanding in its distal half into a large flattened, fan-

like structure with numerous marginal teeth and a roughened

surface. Just distal and posterior to solenomere origin, pos-

teromesal process arising as a narrow, rod-like, tapering

process, curving mesally and anteriorly and terminating just

distal to the most distal teeth of the process accompanying the

solenomere. Lateral branch of telopodite by far the largest,

greatly inflated basally, terminating in toothed crests anteriorly

and posteriorly.

Distribution and microhabitat. In wet eucalypt and Nothofagus

forest litter in the Mt Donna Buang area (Fig. 6).

Etymology. Latin semi- (half) + nudus (naked), adjective,

because the setose basal portion of the gonopod telopodite is so

clearly separated by a transverse constriction from the bare

distal portion.

Acknowledgements

This study was supported by a grant from the Plomley

Foundation. I am grateful to David Steele (University of

Tasmania) for acquiring the SEM images, and to Peter

Lilly white (Museum Victoria) for the loan of specimens and for

his hospitality during my visit to Museum Victoria in March

2004. A draft of this paper was improved by suggestions from

Sergei Golovatch.

References

Carl, J. 1902. Exotische Polydesmiden. Revue Suisse de Zoologie 10:

563-679, pis. 10-12.

Jeekel, C.A.W. 1965. A new genus and a new species of the family

Paradoxosomatidae from Australia (Diplopoda, Polydesmida).

Entomologische Berichten 25: 7-14.

Jeekel, C.A.W. 1982. Millipedes from Australia, 2: Antichiropodini

from Victoria (Diplopoda, Polydesmida, Paradoxosomatidae).

Bulletin Zoologisch Museum, Universiteit van Amsterdam 8(24):

201 - 212 .

Jeekel, C.A.W. 1983. Millipedes from Australia, 8: A new genus and

species of the family Dalodesmidae from Victoria (Diplopoda,

Polydesmida). Bulletin Zoologisch Museum, Universiteit van

Amsterdam 9(16): 145-152.

Jeekel, C.A.W. 1984. Millipedes from Australia, 6: Ausfraliosomatini

from Victoria (Diplopoda: Polydesmida: Paradoxosomatidae).

Records of the Australian Museum 36: 19-44.

Johns, PM. 1964. The Sphaerotrichopidae (Diplopoda) of New

Zealand. 1. Introduction, revision of some known species and

description of new species. Records of the Canterbury Museum

8(1): 1-49.

Mesibov, R. 2004. A new genus and four new species of millipedes

from Tasmania, Australia (Diplopoda: Polydesmida:

Dalodesmidae), with notes on male leg setae in some Tasmanian

dalodes mi ds. Zootaxa 558: 1-19.

Verhoeff, K.W. 1941. Zur Kenntnis australischer Strongylosomiden

und einiger anderer Diplopoden. Lunds Universitets Arsskrift n.s. 2,

36(17): 1-25.

Memoirs of Museum Victoria 61(1): 47-55 (2004)

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

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

Biosystematics of Australian mygalomorph spiders: descriptions of three new

species of Teyl from Victoria (Araneae: Nemesiidae)

Barbara York Main

School of Animal Biology M092, University of Western Australia, Crawley, WA 6009, Australia

(by main @ cyllene .u wa. edu . au)

Abstract Main, B.Y. 2004. Biosystematics of Australian mygalomorph spiders: descriptions of three new species of Teyl from

Victoria (Araneae: Nemesiidae). Memoirs of Museum Victoria 61(1): 47-55.

Revised diagnostic notes on the tribe Teylini and the genus Teyl Main are presented. Three new species of Teyl from

Victoria are described: T. harveyi, T. walkeri and T. yeni. The biogeography of the genus is briefly discussed.

Keywords Araneae, Mygalomorphae, Nemesiidae, Teyl, Teylini, new species, biogeography, taxonomy

Introduction

The genus Teyl Main, 1975 is one of four genera of the tribe

Teylini (Main, 1985b). Teyl luculentus Main, 1975 is the only

named species attributed to the genus. However, many

undescribed species are recognised in south-western Western

Australia and further north in the Carnarvon Basin (Main,

1985b, 1996, 1999, Main et al., 2000). While it was formerly

stated that the genus was confined to south-western Western

Australia (Main, 1985b) examination of specimens in the South

Australian Museum and Museum Victoria shows that it also

occurs in Eyre Peninsula and western Victoria (Main, 1996:

168, 1997: 119, 1999: 237). Main (1999) suggested that the dis-

tribution of Teyl was fragmented in southern Australia during

the Cretaceous inundation of central Australia and that the

radiation of the genus in southern Western Australia has

resulted from the continuing isolation in relictual habitats

which have retained aspects of Gondwanan conditions.

Similarly, occurrence of the genus in Eyre Peninsula and

western Victoria may be related to persistence of microhabitats

simulating Gondwanan characteristics.

Specimens collected from the Little Desert in Victoria by

Museum Victoria are attributed to three new species described

below. All specimens are lodged in Museum Victoria,

Melbourne (NMV). Measurements are in millimetres.

Abbreviations are as follows: ALE, anterior lateral eyes; AME,

anterior median eyes; PLE, posterior lateral eyes; PME

posterior median eyes; d, dorsal; pd, prolaterodorsal; rd, retro-

laterodorsal; v, ventral; pv, prolateroventral; rv, retrolatero-

ventral; PMS, posterior median spinnerets; PLS, posterior

lateral spinnerets; WA, Western Australia; SA, South Australia;

WAM, Western Australian Museum.

In the species descriptions the leg formula is obtained by

dividing the length of the leg by the length of the carapace. The

patella width is measured across the dorsal proximal base (=

knee); an indication of the relative thickness of the legs is given

by the tibial index which = (100 x width of patella) divided by

(length of tibia + patella) (Petrunkevitch, 1942).

Tribe Teylini Main

Teylini Main 1982: 274.— Main, 1983: 925.— Main, 1985b: 744.—

Raven 1985: 82, 87.

Diagnosis. The main diagnostic features of the Teylini (and

which distinguish the tribe from the Anamini) are: the narrow

band of cuspules on the maxillae (covering no more than a third

of the length of the maxillae) in contrast to the broader distri-

bution in Anamini ( Chenistonia Hogg, 2001 and some possibly

misplaced species of Aname Koch, 1872 are exceptions);

absence of a spine-bearing spur on tibia I of males except in

Teyloides and the spherical, subspherical or gourd shaped bulb

of the male palp with the embolus arising equatorially or from

the apex; embolus sometimes reflexed, usually tapering but

may be flanged proximal to or at tip. Inclusion of Merredinia

Main, 1983 (which lacks a tibial spur) in the Anamini has

already been noted as tentative (Main, 1985b).

Remarks. Main (1982) erected the tribe Teylini (to include at

that time only the genus Teyl) as distinct from the Anamini.

However it was not fully described until 1985 (Main, 1985b)

when the four genera Teyl Main, 1975, Namea Raven, 1984,

Teyloides Main, 1985 and Pseudoteyl Main, 1985 were attrib-

uted to it. Remarks by Raven (1985: 5 December 1985) that

Teylini was a nomen nudum were redundant in that they were

Barbara York Main

predated by publication of a description of the tribe (Main,

1985b: 17 September 1985).

Representatives of the tribe occur in southwestern Western

Australia, southern South Australia, western Victoria and the

border country of southeastern Queensland and northeastern

New South Wales.

Teyl Main

Teyl Main, 1975: 74.

Type species. Teyl luculentus Main, 1975 by monotypy (female

holotype from 14.5 km N of Bruce Rock, WA, (WAM 75-

944).— Raven, 1981: 341.— Main, 1982: 73.— Main, 1983:

925.— 1985a: 39.— 1985b: 746.— Raven, 1985: 82, 87, 88.

Diagnosis. As in Main (1985b) but with the following modifi-

cations/additions. Small to medium-sized spiders (carapace

length 2.0 to 10.0 mm). Scopula sparse or absent on tarsi of

palp and anterior legs of female. Scopula of male complete (i.e.

covering whole ventral face) on tarsi I and II, on apical part

only (if present) on anterior metatarsi. Metatarsus I of males

straight (unmodified) or with “elbow” i.e. “bowed”; with many,

few or no spines. Palpal bulb and embolus of male variable, as

for tribe and with embolus short or long (based on described

and undescribed species). Differs from Namea by the broad

sternum which is convex behind labium and the short ovoid ter-

minal segment of the posterior lateral spinnerets; from

Teyloides by lacking a spined spur on tibia I in the male and

lobate, not coiled, internal genitalia in the female; and from

Pseudoteyl by absence of dark smudges on the legs.

Remarks. Raven (1985: 88) incorrectly stated that the holotype

of Teyl luculentus was a “male”. The holotype is a female

(Main, 1975: 74). The comment by Raven (1985: 88)

“cymbium . . . aspinose” requires modification in that some

undescribed species and one species described here have spines

on the male palpal tarsus. The first part of the third couplet of

the key (Main, 1985b) requires modification as follows

“Southwest WA, Eyre Peninsula, SA and western Victoria

(instead of “Southwest WA only”). Species (including many

undescribed) occur in southwestern Western Australia, Eyre

Peninsula South Australia and western Victoria.

Key to males of species of Teyl

1 . Spines on palp tarsus T. harveyi

— Palp tarsus without spines 2

2. Embolus tip tapering; metatarsus I with spines 3

— Embolus tip flanged; metatarsus I without spines

T. walkeri

3. Metatarsus I with few spines T. yeni

— Metatarsus I with many spines T. luculentus

Teyl harveyi sp. nov.

Figures 1-9, table 1

Material examined. Holotype, male, Victoria, 24.9 km SE of

Murrayville, 35°24'S, 141°24'E (site 79), drift fence pitfall trap, Jun

1986, A.L. Yen (NMV T-3011). Paratype, male, Victoria, 20.3 km SE

of Murray ville 35°24'S, 141°20'E (site 76), drift fence pitfall, Jun 1986,

A.L. Yen (NMV T-3012).

Table 1 . Teyl harveyi sp. nov. Holotype, male. Leg measurements and

leg formulae

Total

2.8

1.8

2.2

2.1

1.4

10.3

2.5

1.3

2.0

1.0

8.8

2.2

1.2

1.6

2.1

1.3

8.4

3.0

1.3

2.7

2.8

1.7

11.5

Palp

1.8

1.1

1.3

1.1

5.3

Width of patella I at knee = 0.4. Tibial index =10.00.

Width of patella IV at knee = 0.4. Tibial index = 10.00.

Leg formulae: 4/3.7, 1/3.32, 2/2.83, 3/2.7.

Diagnosis. Male. Light tan brown, glabrous carapace with

sparse and delicate posterolateral spines. Abdomen with dorsal

dark reticulated pattern, ventrally yellow with a few speckles

anterior to spinnerets. Retrolateral spines of palp tibia: 1 heavy

apical, 2 heavy proximal and 2 long, tapering spines on retro-

ventral margin. A group of heavy apical, dorsal spines on palp

tarsus. Embolus tapering. Metatarsus I with pair of thin apical

ventral spines, moderate “elbow”. Tibia I with spines on all

faces; a line of 4 megaspines on retroventral angle.

The group of dorsal apical spines on palp tarsus distin-

guishes the species from T. luculentus and the other two species

(T. walkeri and T. yeni ) described here; the presence of a pair of

ventral apical spines on metatarsus I further distiguishes it from

T. walkeri which lacks spines and yeni which has additionally

several prolaterodorsal spines.

Female. Unknown.

Description of male (holotype). Carapace glabrous, light tan

brown with darker margin and dark radial lines; abdomen

with dorsal pattern of dark , mottled reticulations, venter pale

yellowish with a few dark speckles in front of spinnerets.

Carapace with few marginal spines, all delicate, median lateral

group and posterior marginal group. Fovea straight. Carapace

length 3.1, width 2.6mm; abdomen length 3.7, width 2.3. Eye

group on mound 0.5 long; eye group 0.3 mm long, 0.7 wide;

diameters of eyes: ALE 0.2, AME 0.2, PLE 0.15, PME 0.5.

Sternum slightly convex, length 1.9, width 1.6; evenly covered

with fine bristles; sigilla small, faint, submarginal; labium

length 0.3. Chelicerae with pseudorastellum of about 8 thick

spines on apical inner edge of paturon; groove with 8 pro-

marginal teeth, proximal two large and separated by a gap

from others; a few basal granular teeth on retromargin.

Maxillae with 33 cuspules on right maxilla; about 30 on left,

some damaged.

Legs with scopula on tarsi I and II (thin on II), and distal

half of metatarsi I and II. Tarsal claws with double combs of

superior claws with about 15, fine, close-set teeth decreasing in

size proximally; median claw long and curved. Spination of

legs - 1, femur d 6, patella rv 1, tibia rv 3 megaspines on slight

mounds in a line followed by 1 heavy proximal spine, r 1-1, v

2-2, pv 1-1-1, metatarsus pv 1 apical, rv 1 apical, tarsus 0. II,

femur d 6, patella 0, tibia v 1-2-2, pd 1-1, metatarsus v 2-2, pd

1-1. Ill, femur d 6, patella pd 1-1, tibia v 2-2-4, pd 1-1, rd 1-1,

metatarsus v 2-2-3, pd 1-1-1, rd 1-1-1, tarsus 0. IV, femur d 6,

Barbara York Main

patella, pd 1 apical, rd 1, tibia v 2-2-3, pd 1-1-1, rd 1-1-1,

metatarsus v 2-1-2-3, pd 1-1-1, rd 1-1-1, tarsus 0.

Palp (Left palp missing). Tarsus with apical group of about

10 thick spines. Tibia with 2 heavy terminally hooked and 2

long thin proximal retrolateral spines, prolaterally with 2 taper-

ing heavy apical spines and 1 finer spine. Bulb spherical but

viewed dorsally, gourd shaped with embolus arising at sharp

angle (reflexed) ; embolus finely tapering.

Variation. Paratype carapace length 3.0, width 2.5.

Etymology. The species is named as a tribute to M. S. Harvey

for his vast contribution to arachnology.

Teyl walkeri sp. nov

Figures 10-30, table 2

Material examined. Holotype, male, Victoria, 17.8 km SE of

Murrayville, 35°22'S, 141°19'E (site 74), drift fence pitfall, Jun 1986,

A.L. Yen (NMV T-3008). Paratypes. Allotype female, 27.2 km SE of

Murrayville, 35°25'S, 141°irE, drift net fence pitfall trap (site 79), Jun

1986, A.L. Yen (NMV T-3009). Male, 20.3 km SE of Murrayville,

35°24'S, 141°20'E (site 76), drift fence pitfall , Jun 1986, A.L. Yen

(NMV T-3010).

Diagnosis. Male. Generally pale yellow, abdomen with irreg-

ular dark brown, median dorsal pattern. Carapace non hirsute,

heavy marginal bristles. Palp tarsus without spines; tibia retro-

laterally with 2 hooked, proximal spines, 1 long tapering spine

and 1 apical spine. Embolus slightly reflexed, flanged tip; bulb

large, spherical but in some views gourd-shaped. Legs moder-

ately hirsute, heavily spinose. Metatarsus I without spines.

Tibia I with 4 large spines in retroventral line (3 megaspines

terminally hooked, distal 2 on low mounds).

Absence of spines on metatarsus I and the flanged embolus tip

distinguish the species from T. luculentus, T. harveyi and T. yeni.

Female. Small (< 4 mm), uniformly pale yellow except

abdomen greyish with dorsal median darker streak; carapace

glabrous. Legs with dark spines and sparse hairs and bristles.

The presence of thin, sparse scopula on tarsi of palps and

legs I and II differs from T. luculentus which lacks scopula in

female. Tibia I with fewer heavy spines than T. luculentus.

Internal genitalia lobate with short, thick stems.

Description of male (holotype). Uniformly pale yellow except

for ocular area black, abdomen dorsally with irregular dark

brown median pattern. Carapace non hirsute, with heavy mar-

ginal bristles. Legs not glabrous, moderately hirsute, heavily

spinose. Fovea straight. Carapace length 3.2, width 2.5;

abdomen length 3.5. Eye tubercle slightly wider than edges of

ALE/PLE, tubercle 0.4 high; length of eye group 0.3, width 0.7.

Diameters of eyes: ALE 0.15, AME 0.12, PLE 0.1, PME 0.1.

Sternum length 1.5, width 1.3; labium 0.2. Chelicerae with

heavy apical bristles intermixed with a few reduced spine-like

bristles. Fang groove with 7 teeth on promargin and 6 or 7

small granule-like teeth retrobasally. Maxillae with narrow

band of 18-20 cuspules (left and right).

Legs. Scopula present on tarsi and apical half of metatarsi

I and II. Tarsal claws with 11 or 12 teeth in each line of all

claws. Spination I, femur d 1- 1-3-1 (pd apical), patella pd 1

(apical), tibia rd 1-1, pv 1-1-1, pd 1-1-1, r 1-1-1- 1 (distal 3

megaspines with slightly hooked tip), metatarsus 0. II, femur

1-3- 1-1 (pd apical), patella pd 1 apical, tibia pd 1-1, v 2-2,

metatarsus pd midregion 1, rv midregion 1. Ill (right missing)

left femur 1-1-3-3-2, patella pd 1 apical, rd 1 mid region, tibia

d 1-3-2, metatarsus about 3 spines in a line on each face. IV,

femur d 3 median, pd 3, rd 3, patella 0, tibia 2 or 3 spines in a

line on each face, metatarsus about 3 spines in a line on each

face (difficult to determine exact ‘side’). (Note: also a half

circle of dorsal curved bristles on trochanters). Metatarsus I

with slight proximal ventral depression and slight “elbow”

(Figs 14-16).

Palp. Tarsus without spines; tibia retrolaterally with 2 prox-

imal, terminally hooked, stout spines and 1 long, bristle-like

tapering spine, 1 midretrodorsal spine and 1 retrolateral apical

spine, prolaterally with 3 stout spines and 2 prolateral-ventral

proximal tapering bristle-like spines. Bulb subspherical to

gourd-shaped, height 0.6 mm, width 0.5 mm; embolus arising

mid (equatorial) region (Figs 19-21). Embolus not distinctly

reflexed, tip flanged.

Comments on paratype. Carapace damaged on posterior edge

(length approximately 3.0, width 2.5); retrolateral spination of

right palp tibia as for holotype; retrolateral aspect of tibia I

lacks median spine; abdomen missing.

Description of female (paratype). Uniformly pale yellow

except abdomen greyish with dorsal median dark brown streak

reaching about two-thirds from anterior margin. Legs with

sparse hairs and bristles; brown spines. Carapace glabrous and

Table 2. Teyl walkeri sp. nov. Holotype, male and paratype, female. Leg measurements and leg formulae, male with female measurements in

parentheses.

Total

2.8 (2.7)

1.5 (1.7)

2.1 (2.0)

1.6 (1.9)

1.4 (1.2)

9.4 (9.5)

2.6 (2.5)

1.2 (1.5)

1.9 (1.8)

1.5 (1.4)

9.1 (9.0)

2.4 (2.5)

1.1 (1.4)

1.7 (1.7)

2.1 (2.1)

1.3 (1.3)

8.6 (9.0)

3.0 (3.1)

1.3 (1.6)

2.6 (2.7)

2.6 (2.9)

1.6 (1.6)

11.1(11.9)

Palp

1.8 (1.8)

0.8 (1.1)

1.2 (1.3)

1.0 (1.0)

4.8 (5.2)

Width of patella I at knee = 0.45 (0.45). Tibial index = 12.5 (12.16).

Width of patella IV at knee = 0.40 (0.5). Tibial index = 10.2 (11.62).

Leg formulae: 4/3.0 (3.6), 1/2.9 (2.9), 2/2.8 (2.72), 3/2.7(2.72).

Australian mygalomorph spiders

Figures 10-21. Teyl walked sp. nov. Holotype, male. 10, carapace, chelicerae and abdomen, dorsal view; 11, sternal area and maxillae; 12, eye

group; 13, left chelicera, pro- view; 14, leg I, tarsus, metatarsus, tibia, retrolateral view; 15, right leg I, tarsus, metatarsus, tibia, retroventral view;

16, right leg I, metatarsus and tibia prolateral view; 17-21, right palp; 17, tibia, tarsus and palpal organ, retrolateral; 18, same view as for 17,

palpal tip enlarged; 19, tibia, tarsus and palpal organ, prolateral view; 20, tarsus and palpal organ, prolaterodorsal ; 21, palpal organ (bulb and

embolus) dorsal. Scale =1.0 mm. (figs 10, 14-17, 19); 0.5 mm. (figs 11, 12); 13 and 18 not to scale.

Barbara York Main

Figures 22-30. Teyl walked sp. nov. Paratype (allotype), female. 22, carapace and chelicerae; 23, abdomen, dorsal view; 24, sternal area and

maxillae; 25, right maxilla, proximal with cuspules; 26, right leg I ventral, tarsus, metatarsus and tibia; 27, right leg III dorsal, tarsus, metatarsus

and tibia and patella; 28, right leg I, tarsal claws, p, prolateral, r, retrolateral; 29, genital lobe/aperture; 30, internal genitalia (spemathecae).

Scale =1.0 mm (figs 22-24, 26, 27); 0.5 mm (fig. 30); figs 25, 28, 29 not to scale.

Australian mygalomorph spiders

with black ocular area. Fovea straight. Carapace length 3.3,

width 2.7, caput width 1.9; abdomen length 6.2. Carapace lacks

marginal spines. Eye group 0.85 long, 0.4 wide. Diameters of

eyes: ALE 0.2, AME 0.15, PLE 0.15, PME 0.5; ALE apart 0.55.

Sternum length 1.7, width 1.5. Labium 0.3 long. Chelicerae

with pseudorastellum of stout bristles and long thin spines.

Fang groove with 7 (left), 8 (right) promarginal teeth and 5

basal retromarginal teeth. Maxillae with cuspules in narrow

band around incurved edge (about 22 on left maxilla, 35 on

right), small rounded heel.

Legs. Scopula thin on palp tarsus and tarsus I; a few proxi-

mal scopula hairs on tarsus II; a few thin apical hairs on

metatarsus I. Tarsal claws. Palp claw with 6 prolateral teeth

(left) (right claw detached); legs with 6-9 teeth in each row of

all claws. Spination I, femur pd 1 apical, rd a few, patella pd

1- 1-1 bristle-like, tibia rv 1-1, v 2 bristle-like, metatarsus v 2-1

(rv) -2. II, femur pd 1 apical, patella pd 1 apical, tibia v 1-1-2

(bristles), pd 1-1, metatarsus v 2-2-2, pd 1. Ill, femur pd 1

apical, patella d 2-1, tibia v 1-2-2 (bristles), d 3- 2, metatarsus

v 2-2-3, d 8 or 9 spines and heavy bristles. IV, femur pd 1

apical, patella rd 1, tibia v 2-2-2, rd 1-1, metatarsus v 2-2-3, d

2 - 1 - 1 - 2 - 1 .

Palp, tibia pv 4 distal, pd 1 proximal, tarsus pv 1 proximal.

Internal genitalia lobate with short, thick stems, terminally

bulbous (Fig. 30).

Etymology. The species is named in honour of Dr Ken Walker

in recognition of his entomological and arachnological work.

Teyl yeni sp. nov.

Figures 31—40, table 3

Material examined. Holotype, male, Victoria, 22.2 km SE of

Murrayville, 35°25'S, 141°21'E (site 77), drift fence pitfall trap, Jun

1986, A.L. Yen (NMV T-3013). Paratype, male, Victoria, 17.8 km SE

of Murrayville, 35°23'S, 141°19'E (site 74), drift fence pitfall trap, Jun

1986, A.L. Yen. (NMV T-3014).

Diagnosis. Male. Generally pale yellowish brown with dark

median pattern on abdomen dorsum. Palp tibia retrolaterally

with 2 basal and 1 apical stout, heavy spines and 2 long thin

tapering spines. Tarsus without apical spines. Embolus base

arising equatorially and slightly reflexed, tapering. Metatarsus

I slightly bowed, 2 retrolateral spines in proximal half. Tibia I

retrolaterally with 3 megaspines in proximal line.

Differs from T. harveyi by having 2 retrolateral spines

(instead of 1) on metatarsus I, and further from T. harveyi and

T. walkeri by having more metatarsal spines (including a pro-

laterodorsal line) on metatarsus I. Differs from T. luculentus by

having a more strongly “bowed” metatarsus I, the strongly

hooked retrolateral spines on the palp, smaller size and delicate

marginal spines on carapace and from all the other species by

the more widely spread maxillary cuspules.

Female. Unknown.

Description of male (holotype). Carapace, sternum, legs yel-

lowish brown with faint smudge of darker brown. Abdomen

pale yellowish brown with reticulated dark brown pattern on

median dorsal area. Legs with short evenly distributed hairs

and sparsely scattered strong spines. Carapace with delicate

Table 3. Teyl yeni sp. nov. Holotype, male. Leg measurements and leg

formulae.

Total

4.4

2.0

3.4

3.3

2.3

15.4

4.2

1.9

3.1

3.2

2.3

14.7

III

4.3

1.4

2.5

3.0

2.1

13.3

3.9

1.8

3.7

4.5

2.3

16.2

Palp

2.7

1.5

2.1

1.0

7.3

Width of patella I at knee = 0.4. Tibial index = 7.4.

Width of patella IV at knee = 0.4. Tibial index = 7.27.

Leg formulae: 4/4.26, 1/4.05, 2/3.86. 3/3.5

marginal spines. Caput low. Height of carapace at fovea 0.6, at

eyes 1.0. Fovea straight, slightly reflected at edges. Carapace

length 3.8, width 3.5, caput width 2.0. Abdomen length 4.2,

width 2.4. Eye group broad on low tubercle 0.4 long, 0.9 wide.

Diameters of eyes: ALE 0.25, AME 0.2, PLE 0.2, PME 0.1.

Sternum length 2.3, width 2.0; with long fine bristles uniform-

ly scattered. Labium 0.3 long. Sternal sigilla small, round,

indistinct, marginal although posterior pair separated by a mar-

ginal line of bristles. Chelicerae long and narrow with a

pseudorastellum of short apical spines on inner edge.

Promargin of cheliceral groove with 8 teeth, 5 basal retromar-

ginal teeth. Maxillae with broad band of about 50 cuspules

around inner edge (extending farther from inner edge than is

usual for Teyl specimens).

Legs. I, metatarsus “bowed” i.e. with a distinct “elbow”.

Scopula complete (that is covering full extent of segment) on

tarsus I, thin on II, sparse and divided by a line of bristles on III

and IV; present on metatarsi I and II only (apical half ).

Trichobothria. Palp tarsus at least 8. Tarsus I with about 9, plus

four rigid, vertical hairs, metatarsus about 12 trichobothria.

Tarsal claws 8-12 teeth on each comb of paired claws, basal

ones may be fused. Spination, I, femur d 14, patella pd 2, tibia

d 2- 1-2-2 (including pd and rd), rv 3 megaspines in apical half,

1 in lower third (Fig. 35), metatarsus pd l-l-l-l-l (left 1-1-1-

0- 1), rv 2 distal to elbow, pv 1 apical. II, femur dll, patella pd

2, tibia d 7, v 2-2-3, metatarsus d 5, v 2-2-2. Ill, femur dll,

patella pd2, rd 1, tibia d 3-3-3, v 2-2-2, metatarsus d 9, v 2-2-3.

IV, femur dll, patella d 0, tibia d 6, v 2-2-2, metatarsus d 2-1-

1- 2-3.

Spinnerets. PMS length 0.4, PLS, basal segment 0.6, medi-

an 0.4, terminal 0.5.

Comments on paratype. Specimen larger than holotype; cara-

pace length 4.4, width 4.0. Left palp with 3 basal retrolateral

heavy spines (instead of 2). Right legs II, III, IV and left HI, IV

missing. Tibia I spination (right) d 2- 1-1-2, rv 5 in a line (2

proximal, 3 distal megaspines), pv 2-1, metatarsus pd 1-1

(proximal) - 1 (distal/apical), rv 1 distal to elbow and 1 very

small apical spine; (left) d 2-1-2-2, rv 3 in line (1-1-1 distal

megaspines), pv 1-1-1, metatarsus pd 1-1-1, rv 1-1 distal to

elbow.

Etymology. The species is named in recognition of Dr Alan

Yen’s untiring activities in entomology and arachnology.

Barbara York Main

Figures 31-40. Teyl yeni sp. nov. Holotype, male. 31, dorsal view, carapace, chelicerae and abdomen; 32, sternal area and left maxilla; 33, eye

group; 34, right tarsus I, prolaterodorsal; 35, right leg I, retrolateral, tibia, metatarsus, tarsus; 36, schematic dorsal spination right tibia I; 37, right

metatarsus I, prolaterodorsal ; 38, right palp, retrolateral; 39, right palp, prolatero ventral; 40, schematic spination of right palp tibia, prolateral.

Scale = 0.1 mm (figs 31, 32, 34, 35, 37-39); 0.5 mm (fig. 33); figs 36, 40 not to scale.

Australian mygalomorph spiders

Acknowledgements

I thank Drs K. Walker and A.L. Yen of Museum Victoria for

making available the specimens from the biological survey of

the Little Desert.

References

Main, B.Y. 1975. The citrine spider: a new genus of trapdoor spider

(Mygalomorphae: Dipluridae). The Western Australian Naturalist

13: 73-78.

Main, B.Y., 1982. Adaptations to arid habitats by mygalomorph spi-

ders. Pp. 273-283 in: Barker, W.R. and Greenslade, PJ.M. (eds),

Evolution of the Flora and fauna of Arid Australia. Peacock

Publications: Frewville.

Main, B.Y. 1983. Further studies on the systematics of Australian

Diplurinae (Chelicerata: Mygalomorphae: Dipluridae): two new

genera from south western Australia. Journal of Natural History 17:

923-949.

Main, B.Y. 1985a. Mygalomorphae. Pp. 1-48 in: Walton, D.E. (ed.),

Zoological Catalogue of Australia, Arachnida. Vol.3. Australian

Government Publishing Service: Canberra.

Main, B.Y. 1985b. Further studies on Australian Diplurinae: a review

of the genera of the Teylini (Araneae: Mygalomorphae:

Diplurinae). Australian Journal of Zoology 33: 743-759.

Main, B.Y. 1996. Microcosmic biogeography: trapdoor spiders in a

time warp at Durokoppin. Pp. 163-167 in: Hopper, S.D., Chappill,

J.A., Harvey, M.S., and George, A.S. (eds), Gondwanan heritage.

Past, present and future of the Western Australian biota. Surrey

Beatty and Sons: Chipping Norton.

Main, B.Y. 1997. Granite outcrops: A collective ecosystem. Journal of

the Royal Society of Western Australia 80: 113-122.

Main, B.Y. 1999. Biological anachronisms among trapdoor spiders

reflect Australia’s environmental changes since the Mesozoic. Pp.

236-245 in: Ponder, W., and Lunney, D. (eds), The other 99%. The

conservation and biodiversity of invertebrates. Royal Zoological

Society of New South Wales: Mosman.

Main, B.Y., Sampey, A., and West, PL.J. 2000. Mygalomorph spiders

of the southern Carnarvon Basin, Western Australia. Records of the

Western Australian Museum Supplement 61: 281-293.

Petrunkevitch, A. 1942. A study of amber spiders. Transactions of the

Connecticut Academy of Arts and Sciences 34: 1 19—464.

Raven, R.J. 1981. A review of the Australian genera of the mygalo-

morph spider subfamily Diplurinae (Dipluridae: Chelicerata).

Australian Journal of Zoology 29: 321-363.

Raven, R, J. 1985. The spider infraorder Mygalomorphae (Araneae):

cladistics and systematics. Bulletin of the American Museum of

Natural History 182(1): 1-180.

Memoirs of Museum Victoria 61(1): 57-64 (2004)

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

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

Chirostylidae from north-western Australia (Crustacea: Decapoda: Anomura)

Shane T. Ahyong 1 and Keiji Baba 2

'Department of Marine Invertebrates, Australian Museum, 6 College St, Sydney, NSW 2010, Australia

( shanea @ austmus .go v. au)

2 Faculty of Education, Kumamoto University, 2-40-1 Kurokami, Kumamoto 860-8555, Japan

(keiji@gpo.kumamotu-u.ac.jp)

Abstract Ahyong, S.T., and Baba, K. 2004. Chirostylidae from north-western Australia (Crustacea: Decapoda: Anomura).

Memoirs of Museum Victoria 61(1): 57-64.

New records and a new species of Chirostylidae from north-western Australia are reported. Prior to the present study,

two chirostylids were known from north-western Australia. The present collection comprises seven species arrayed in

three genera: Chiro stylus, Eumunida and Uroptychus. Five species are newly recorded from Australian waters: Eumunida

ampliata, E. funambulus, E. pacifica, Uroptychus joloensis and U. nigricapillis. Chirostylus dolichopus is newly

reported from the Northern Territory. One species from the North West Shelf, U. oxymerus is new to science. A total of

40 species of Chirostylidae are now known from Australian waters.

Keywords Cmstacea, Decapoda, Anomura, Chirostylidae, Uroptychus, new species

Introduction

The Australian deep-water squat lobsters of the family

Chirostylidae are known from relatively few studies

(Henderson, 1885, 1888; Haig, 1974; Baba, 1986, 2000; de

Saint Laurent and Macpherson, 1990; de Saint Laurent and

Poupin, 1996; Ahyong and Poore, 2004). Of the 34 chirostylids

recorded from Australia (Ahyong and Poore, 2004), only two

are known from north-western Australia: Chirostylus doli-

chopus Ortmann, 1892 and Uroptychus brucei Baba, 1986. The

present study is based on miscellaneous chirostylid collections

from north-western Australia in the collections of the

Australian Museum, Museum Victoria, and the South

Australian Museum. Of the seven species reported below, five

represent range extensions into Australian waters, and one is

new to science. A total of 40 species of Chirostylidae is now

known from Australian waters.

Measurements of specimens, given in millimetres (mm),

indicate the carapace length including the rostrum unless indi-

cated otherwise. Specimens are deposited in the Australian

Museum, Sydney (AM), Natural History Museum, London

(BMNH), Museum Victoria, Melbourne (NMV), and the South

Australian Museum, Sydney (SAM).

Chirostylidae Ortmann, 1892

Chirostylus Ortmann, 1892

Chirostylus dolichopus Ortmann, 1892

Chirostylus dolichopus Ortmann, 1892: 246, pi. 11, figs 2, 2b, 2c,

2e, 2i, 2z [type locality: Kadsiyama (= Katsuyama), Sagami Bay,

Japan]. — Haig, 1974: 447. — Davie, 2002: 30.

Material examined. AM P25058, 1 male (4.4 mm), Timor Sea,

Northern Territory, 9°30'S, 132°34'E, 124 m, MV San Pedro Sound,

P. Colman, 9 Nov 1969.

Remarks. Chirostylus dolichopus was reported from tropical

Western Australia by Haig (1974) without mention of a

specific locality. The present specimen represents a new record

for the Northern Territory.

Distribution. Western Indian Ocean, Sulu Archipelago, Japan,

Western Australia and now the Northern Territory; 35-140 m

(Baba, 1988).

Eumunida Smith, 1883

Eumunida ( Eumunida ) funambulus Gordon, 1930

Eumunida funambulus Gordon, 1930: 744, figs lc, 2a, b, 3b, 4b, 5

[type locality: Gulf of Aden],

Eumunida (Eumunida) funambulus. — de Saint Laurent and Poupin,

1996: 350-352.

Material examined. SAM C725, 1 female (38.8 mm), Timor Sea, off

Palmerston [now Darwin], Northern Territory, P. Foelske, Nov 1890.

Remarks. The single specimen agrees well with published

accounts. Unfortunately, the depth of capture was not recorded

for this and other specimens of Eumunida reported herein.

Distribution. Gulf of Aden to Indonesia, the Philippines, Japan

(de Saint Laurent and Poupin, 1996) and now from Timor Sea,

Northern Territory; 130-732 m.

Shane T. Ahyong and Keiji Baba

Eumunida (Eumunida) pacifica Gordon, 1930

Eumunida pacifica Gordon, 1930: 746, figs 6-7 [type locality:

Timor Sea, Indonesia].

Eumunida ( Eumunida ) pacifica. — de Saint Laurent and Poupin,

1996: 359-362, figs 4a, b, 12a.

Material examined. SAM C724, 1 female (43.0 mm), Timor Sea, off

Palmerston [now Darwin], Northern Territory, P. Foelske, Nov 1890;

SAM C725, 4 males (16.9-32.1 mm), 1 female (20.0 mm), Timor Sea,

off Palmerston [now Darwin], Northern Territory, P. Foelske, Nov

1890.

Remarks. The present specimens of E. pacifica are all in dry

condition. One lot (SAM C725) forms part of a larger series of

Eumunida that includes E. funambulus and E. ampliata (see

below) registered as a single lot.

Distribution. Indonesia (Timor Sea, Moluccas, Kai Islands) and

now from northern Australia; 204-304 m (de Saint Laurent

and Poupin, 1996); 293-605 m (de Saint Laurent and Poupin,

1996).

Eumunida (Eumunidopsis) ampliata de Saint Laurent and

Poupin, 1996

Eumunida (Eumunidopsis) ampliata de Saint Laurent and Poupin,

1996: 368-371, figs 7a-e, 8a-e, 12c [type locality: Timor Sea].

Material examined. SAM C725, 2 males (21.8-25.9 mm), 2 females

(23.1-28.1 mm), Timor Sea, off Palmerston [now Darwin], Northern

Territory, P. Foelske, Nov 1890.

Remarks. The specimens are fragmented but nevertheless

identifiable as E. ampliata.

Distribution. Indonesia (Kai Islands, Sahul Bank) and now

from northern Australia; 204-304 m (de Saint Laurent and

Poupin, 1996).

Uroptychus Henderson, 1888

Uroptychus joloensis Van Dam, 1939

Figure 1

Uroptychus joloensis Van Dam, 1939: 395-398, figs 2, 2a, 2b, 2c

[type locality: Jolo, Indonesia].

Material examined. NMV J21046, 2 males (5.6-7.3 mm), ovigerous

female (7.8 mm), North West Shelf between Port Hedland and

Dampier, 18°45'S, 118°24'E, 142 m, trawl, G. Poore and H. Lew Ton,

5 Jun 1983 (stn NWA-23).

Diagnosis. Carapace lateral margins convex; dorsum unarmed;

with anteriorly directed anterolateral spine and 2 lateral spines

in advance of midlength; with low, narrow, marginal ridge

extending from orbital to posterior margin. Sternal plastron

with faintly concave anterior margin, and median notch.

Cornea not dilated, about one-third length of remaining stalk.

Antennal basal segment with small lateral spine; ultimate and

penultimate segments of peduncle each with long distal spine.

Antennal scale not exceeding apex of ultimate peduncle seg-

ment. Chelipeds smooth but sparsely setose; ischium with

2 dorsal and 1 ventral spine. Walking legs similar, slightly

decreasing in length posteriorly, sparsely setose, segments

smooth; propodus with paired movable spines on distal flexor

margin; dactylus with 6 or 7 strong, widely spaced, perpen-

dicularly (or near perpendicularly) directed corneous teeth on

flexor margin.

Description. Carapace: Length excluding rostrum slightly less

than breadth. Lateral margins convex; with anteriorly directed

anterolateral spine and 2 lateral spines in advance of midlength;

with low, narrow, marginal ridge extending from orbital to pos-

terior margin. Rostrum narrow, triangular, dorsally excavate.

Outer orbital angle produced to small acute spine. Dorsum

smooth, with scattered setae. Pterygostomian flap visible in

dorsal view; with strong anterior spine, otherwise unarmed.

Sternum: Plastron about as long as wide, subquadrate.

Stemite 3 (at base of maxilliped 3) with faintly concave or

irregular anterior margin; with narrow, U-shaped median notch;

anterolateral angle with blunt tubercle; anterior margin level

with or very slightly exceeding sternite 4. Stemite 4 (at base of

pereopod 1) with anterolateral margins angular.

Abdomen: Segments smooth, sparsely setose. Telson about

one-third as long as broad; distal portion broadly convex

posteriorly, shorter than proximal portion.

Eye: Cornea not dilated, about one-third length of remaining

stalk; reaching approximately to midlength of rostrum.

Antenna: Basal segment with distolateral spine. Peduncle

extending almost to apex of rostmm. Ultimate segment longer

than penultimate segment, both with long distomesial spine.

Antennal scale slightly wider than opposite peduncular

segments, extending almost to apex of ultimate segment

(excluding distal spine).

Maxilliped 3: Dactylus and propodus unarmed. Carpus with

proximal spine on extensor margin. Mems with distalolateral

spine and 2 small flexor spines. Ischium with small distal

spine lateral to rounded flexor distal margin. Crista dentata

denticulate for length of ischium.

Pereopod 1 (cheliped): About 4 times carapace length;

smooth but sparsely setose. Palm 5 times as long as high.

Fingers crossing, opposable margins dentate and each with low

process proximally; slightly longer than one-third palm-length.

Carpus about as long as palm. Merus with 2 or 3 small spines

on lower proximal margin. Ischium with 2 dorsal spines;

ventrally with short subterminal spine on mesial margin.

Pereopods 2^4 ( walking legs): Similar, slightly decreasing

in length posteriorly, sparsely setose; segments smooth. Merus

shorter than carapace. Propodus more than twice as long as

carpus, not broadened distally, with paired movable spines on

distal flexor margin. Dactylus with 6 or 7 strong, widely

spaced, perpendicularly (or near perpendicularly) directed

corneous teeth on flexor margin.

Remarks. The present specimens generally agree with the type

account of U. joloensis Van Dam, 1939, based on a specimen

from Jolo, Indonesia, in carapace shape and lateral spination,

the armature of the dactyli of the walking legs, and small eyes.

The present specimens differ from the type account of U.

joloensis chiefly in the following features: the rostrum lacks

distalolateral spines, the fingers of the cheliped are longer than

one-third instead of about one-quarter the length of the pro-

podal palm, the anterior margin of the sternal plastron (sternite

Chirostylid squat lobsters from north-western Australia

Figure 1. Uroptychus joloensis Van Dam, 1939, ovigerous female (7.8 mm), NMV J21046. A, dorsum. B, anterior carapace, right lateral view.

C, right cheliped, proximal mesial view. D, right antenna, ventral view. E, right maxilliped 3, lateral view. F, crista dentata, right. G, sternal

plastron. H, telson. Scale A-C, G = 3.0 mm, D-F = 0.7 mm, H = 1.5 mm.

Shane T. Ahyong and Keiji Baba

3) does not distinctly exceed that of sternite 4, and the median

notch of the sternal plastron is distinctly narrowed rather than a

broad U-shape. The difference in rostral spination, whether

apically trifid (as in the type) or singular (as in present mater-

ial) is variable in this species as also in U. tridentatus

(Henderson, 1885) and U. zezuensis Kim, 1972 (Baba, unpub-

lished). The last mentioned differences appear to be within the

range of expected morphological variation. As only the second

record of the species, a description and figures of Australian

material are provided.

Distribution. Jolo, Indonesia, and now from north-western

Australia at depths between 37.8-56.7 m and 142 m.

Uroptychus nigricapillis Alcock, 1901

Figures 2A-I

Uroptychus nigricapillis Alcock, 1901: 282-284, pi. 3 fig. 3, 3a

[type locality: Andaman Islands]. — Alcock and McArdle, 1902, pi. 56

fig. 3.

Material examined. AM P67832, 1 ovigerous female (11.0 mm), North

West Shelf, 240 km NW of Port Hedland, Western Australia, 500 m, J.

Paxton, 7 Apr 1982 (stn S02/82/31).

Diagnosis. Carapace excluding rostrum distinctly longer than

broad; lateral margins slightly divergent; with 3 or 4 small

spines behind base of cervical groove; with anteriorly directed

anterolateral spine; posterior quarter with low ridge. Rostrum

sharply triangular, exceeding one-third length of remaining

carapace. Outer orbital angle produced to small spine extending

to level of anterolateral spines. Dorsum with pair of distinct

epigastric spines. Sternite 3 anterior margin deeply emarginate,

with pair of median spines separated by narrow notch. Antennal

basal segment with small outer spine; ultimate and penultimate

segments unarmed. Antennal scale extending slightly beyond

midlength of ultimate peduncular segment. Chelipeds smooth,

fingers setose. Pereopod 4 slightly shorter than pereopods 2 and

3; sparsely setose; propodi with 6-9 movable spines on distal

flexor margin, none paired; distalmost flexor spine remote from

distal margin; dactyli with 8-10 small spines on flexor margin,

oriented oblique to dactylar margin, distal 2 appressed.

Description. Carapace: Length excluding rostrum distinctly

greater than breadth. Lateral margins slightly divergent; with 3

or 4 small spines behind base of cervical groove, anterior-

most spine with small additional dorsal spine; with anteriorly

directed anterolateral spine; posterior quarter with low carina.

Rostrum sharply triangular, exceeding one-third length of

remaining carapace; margins unarmed. Outer orbital angle pro-

duced to small spine extending to level of anterolateral spines.

Dorsum smooth, with pair of distinct epigastric spines; cervical

groove distinct medially. Pterygostomian flap with small

anterior spine.

Sternum: Plastron broader than long, widening posteriorly.

Sternite 3 (at base of maxilliped 3) depressed, anterior margin

narrow, deeply emarginate, with pair of median spines separ-

ated by narrow notch; with distinct anterolateral tooth. Sternite

4 (at base of pereopod 1) with distinct anterolateral tooth

extending anteriorly to level of base of emargination of sternite

3; margins dentate, irregular; demarcation between stemites

4 and 5 dentate.

Abdomen: Abdominal segments glabrous. Telson length

about half breadth; distal portion posteriorly emarginate, about

twice length of proximal portion.

Eye: Cornea moderately dilated, subequal to length of

peduncle; reaching to distal quarter of rostrum;

Antenna: Basal segment with small outer spine. Peduncle

extending to distal third of rostrum. Flagellum about 3 times as

long as peduncle. Ultimate and penultimate segments unarmed;

ultimate segment about 2.5 times length of penultimate seg-

ment. Antennal scale slightly wider than opposite peduncular

segments, extending slightly beyond midlength of ultimate

peduncular segment.

Maxilliped 3: Dactylus, propodus, carpus and merus

unarmed. Crista dentata distinctly serrate on proximal two-

thirds of ischium, extending onto basis.

Pereopod 1 (cheliped): Slender, cylindrical, about 3.5 times

carapace length; glabrous dorsally; fingers setose. Palm about 4

times as long as high, about 1.5 times as long as fixed finger;

lower margin with row of low granules. Fingers crossing,

opposable margins dentate; occlusal margin of movable finger

with obtuse process proximally; occlusal margin of fixed finger

without distinct prominence. Carpus longer than merus and

propodal palm; glabrous. Merus usually with several low gran-

ules on inner proximal margin. Ischium with short, slender

spine on dorsolateral margin.

Pereopods 2-4 (walking legs): Similar; sparsely setose;

decreasing in length posteriorly. Pereopod 1 merus longer than

carapace. Merus extensor margin smooth. Carpus about 0.5

merus length, about two-thirds propodus length. Propodi not

broadened distally, with 6-9 movable spines on distal flexor

margin; distalmost flexor spine remote from distal margin.

Dactyli setose marginally, with 8-10 small spines on flexor

margin, oriented oblique to dactylar margin, distal 2 appressed.

Pereopod 4 slightly shorter than pereopods 2 and 3.

Ova: Diameter 0.8 mm.

Remarks. The single Australian specimen agrees well with the

original account and figures of U. nigricapillis in most respects

(Alcock, 1901; Alcock and McArdle, 1902) and represents the

first record from Australian waters. It differs from figures of the

holotype in having relatively longer legs, such that the merus of

the first walking leg in the Australian specimen is longer than,

instead of shorter than, the carapace length (excluding ros-

trum). In a series of Indian Ocean specimens (from several

localities) attributed to U. nigricapillis, Tirmizi (1964) noted

considerable variation in the ornamentation of the stemites, the

distinctness of the epigastric and lateral spines on the carapace,

and length of the antennal scale - features normally considered

diagnostic in many species. Unfortunately, the sternum of the

holotype of U. nigricapillis was neither figured nor described

by Alcock (1901). A Java Sea specimen figured by Van Dam

(1940: fig. 2), however, bears a transverse row of tubercles

behind the dentate anterior margin of sternite 4 in contrast to

the Australian specimen which lacks tubercles. Tirmizi (1964)

also noted variation in the sternal armature, in which the ster-

num of only one specimen resembled Van Dam’s (1940) figure.

Chirostylid squat lobsters from north-western Australia

Figure 2. A-I, Uroptychus nigricapillis Alcock, 1901, ovigerous female (11.0 mm), AM P67832. J, Uroptychus gracilimanus (Henderson, 1885),

female (9.4 mm), AM P65825. A, dorsum. B, carapace, right lateral view. C, cheliped, proximal ventral view. D, cheliped, distal ventral view.

E, right antenna, ventral view. F, right maxilliped 3, lateral view. G, crista dentata, right. H, sternal plastron. I, telson. J, left pereopod 2, dactylus

and distal propodus. Scale A-D = 3.0 mm, E-F = 0.7 mm, G-J =1.5 mm.

Shane T. Ahyong and Keiji Baba

The Australian specimen bears well-developed epigastric

spines, and small but distinct lateral carapace spines as in

Alcock’s (1901) account. Variation in the presence and degree

of epigastric ornamentation is also known in other congeners

(e.g. U. australis (Henderson, 1885), U. gracilimanus

(Henderson, 1885), U. vandamae Baba, 1990). However, vari-

ation in other characters mentioned by Tirmizi (1964) and other

studies (e.g. Baba, 1981, 1988), namely, the lateral carapace

spination and ornamentation and morphology of the sternal

plastron, suggests that more than one species could be confused

under U. nigricapillis. In view of morphological variation in

U. nigricapillis reported in previous studies, the Australian

specimen is described and illustrated.

Uroptychus nigricapillis closely resembles U. gracilimanus,

described from off Port Jackson, in general habitus, armature of

the dactyli of the walking legs, and in variation in the presence

and size of the epigastric spines. Uroptychus nigricapillis dif-

fers, however, in the presence of lateral spines on the carapace

margins, in bearing a short slender spine instead of a low tooth

on the cheliped ischium, and in the spination of the propodi of

the walking legs. Thus, in U. nigricapillis and the distal flexor

spine on the propodi of the walking legs is positioned more

remotely from the distal margin than in U. gracilimanus (Fig.

2A). The distal-most of the flexor spines is located close to the

articul-ation with the dactylus, a fact confirmed by examination

of the holotype of U. gracilimanus (BMNH 1888: 33, off Port

Jackson). In the specimen of U. gracilimanus figured by

Ahyong and Poore (2004: fig. 10, AM P65825, off Port

Jackson), some of the movable propodal spines on the walking

legs are damaged or lost. The dactylus and distal portion of the

propodus of the left first walking leg of the same specimen of

U. gracilimanus is shown (Fig. 2J) for comparison with the

condition in U. nigricapillis.

Distribution. Western Indian Ocean, Andaman Sea, Indonesia,

the Philippines and now from north-western Australia at

66-1930 m depth.

Uroptychus oxymerus sp. nov.

Figure 3

Material examined. Holotype: NMV J21040, male (>6.5 mm), North

West Shelf, between Port Hedland and Dampier, 1 84 m, G. Poore and

H. Lew Ton, 6 Jun 1983 (stn NWA-24).

Diagnosis. Carapace lateral margins convex; dorsum unarmed;

with strong, anteriorly directed anterolateral spine and 5 or 6

stout lateral spines. Rostrum apically trifid; dorsally excavate.

Sternal plastron with faintly concave anterior margin, and

median notch. Cornea not dilated, about one-third length of

remaining stalk. Antennal basal segment with outer spine; ulti-

mate and penultimate segments of peduncle each with long dis-

tal spine. Antennal scale exceeding apex of ultimate peduncle

segment. Chelipeds rugose, setose; merus with row of strong,

stout spines on mesial ventral margin. Walking legs similar,

slightly decreasing in length posteriorly, setose, segments

rugose; propodus distal flexor margin with 2 or 3 movable

spines, distal most paired; dactylus with 6 or 7 strong, widely

spaced, perpendicularly (or near perpendicularly) directed

corneous teeth on flexor margin.

Description. Carapace: Length excluding rostrum subequal to

breadth. Lateral margins convex; with anterolateral spine and 5

lateral spines and spinule above third lateral spine; antero-

lateral spine, strong, directed anteriorly; first and second

lateral spine slender; third and fourth lateral spines stout, larg-

er than others; fifth lateral spine low, blunt. Rostrum apically

trifid; dorsally excavate. Outer orbital angle produced to small

acute spine. Dorsum smooth, with scattered setae.

Pterygostomian flap with strong anterior spine and field of

spines and tubercles on anterior half.

Sternum: Plastron about as long as wide, subquadrate.

Stemite 3 (at base of maxilliped 3) anterior margin shallowly

concave, with narrow U-shaped median notch, anterolateral

angle blunt. Stemite 4 (at base of pereopod 1) with antero-

lateral margins pointed, not produced beyond anterior

concavity of sternite 3.

Abdomen: Segments glabrous. Telson about half as long

as broad; distal portion posteriorly emarginate, longer than

proximal portion.

Eye: Cornea not dilated, about half as long remaining stalk;

not reaching beyond midlength of rostmm;

Antenna: Basal segment with distolateral spine. Peduncle

extending beyond distal half of rostmm. Ultimate segment

about twice as long as penultimate segment, both with long

distomesial spine; ultimate also with 2 small marginal spines.

Antennal scale slightly wider than opposite peduncular

segments, extending slightly beyond apex of ultimate segment.

Maxilliped 3: Dactylus and propodus unarmed. Carpus with

distal and 2 proximal spines on extensor margin. Mems

with distal extensor spine and 2 distal flexor spines. Ischium

with small spine lateral to distal end of flexor margin. Crista

dentata denticulate for length of ischium.

Pereopod 1 (cheliped): About 4 times carapace length; all

segments rugose and setose. Palm 3 times as long as high,

about 2.5 times as long as fixed finger; surface with short,

transverse, depressions. Fingers crossing, opposable margins

dentate and each with low process proximally. Carpus with

small lower distal spine adjacent to each articular condyle of

propodus. Merus with row of 5 stout spines on mesial ventral

margin and group of 3 spines proximally. Ischium with 2

dorsal spines and 1 ventral spine; distal dorsal spine slender,

depressed.

Pereopods 2-4: Similar, slightly decreasing in length poster-

iorly, strongly setose; segments rugose, setose. Merus shorter

than carapace. Propodus more than twice length of carpus, not

broadened distally, with 2 or 3 movable spines on distal flexor

margin, distalmost paired. Dactylus with 6 or 7 strong, widely

spaced, perpendicularly (or near perpendicularly) directed

corneous teeth on flexor margin.

Etymology. From oxy, sharp, and merus, the fourth limb seg-

ment, in reference to the sharp, stout spines on the merus of the

cheliped that distinguish the new species from U. tridentatus

and U. zezuensis.

Remarks. Uroptychus oxymerus sp. nov. closely resembles

U. tridentatus (Henderson, 1885), described from Indonesia,

Chirostylid squat lobsters from north-western Australia

Figure 3. Uroptychus oxymerus sp. nov., holotype male (+6.5 mm), NMV J21046. A, dorsum. B, carapace, right lateral view. C, right cheliped,

proximal mesial view. D, left antenna, ventral view. E, right maxilliped 3, lateral view. F, crista dentata, right. G, sternal plastron. H, telson. Scale

A-C = 2.0 mm, D-H =1.0 mm.

Shane T. Ahyong and Keiji Baba

and U. zezuensis Kim, 1972 from Korea, in almost all respects

including the spinose carapace margins, distally spinose

segments of the antennal peduncle, armature of pereopods 2-4,

and shape of the sternal plastron. The new species is readily

distinguished from both U. tridentatus and U. zezuensis by

having a row of large, stout spines instead of a cluster of spines

on the mesial ventral margin of the merus of the cheliped, in

having three instead of two spines on the extensor margin of the

carpus of the third maxilliped, and in having spines along

the lower margin of the ultimate segment of the antennal

peduncle in addition to the distal spine.

Four species of Uroptychus are now known from the North

West Shelf. Uroptychus oxy merus is readily distinguished from

U. joloensis by having distinctly more spinose lateral carapace

margins (5 or 6 instead of 2); from U. nigricapillis by lacking a

pair of epigastric spines on the carapace and in having a distal

spine on the ultimate and penultimate segments of the antennal

peduncle; and from U. hrucei by having lateral carapace spines.

Distribution. Known only from the type locality at 184 m

depth.

Acknowledgments

We are grateful to Jo Taylor (Museum Victoria) and Thierry

Laperousaz (South Australian Museum) for the loan of

specimens.

References

Ahyong, S.T. and Poore, G.C.B. 2004. The Chirostylidae of southern

Australia (Cmstacea: Decapoda: Anomura). Zootaxa 436: 1-88.

Alcock, A. 1901. Descriptive catalogue of the Indian Deep Sea

Crustacea Decapoda Macrura and Anomura in the Indian Museum.

Being a revised account of the deep-sea species collected by the

Royal Indian Marine Survey Ship Investigator. Trustees of the

Indian Museum: Calcutta. 286 pp.

Alcock, A., and McArdle, A.F. 1902. Illustrations of the Zoology of the

Royal Indian Marine Survey Steamer “Investigator”, Crustacea.

Part 10, pis 56-67. Trustees of the Indian Museum: Calcutta.

Baba, K. 1981. Deep-sea galatheidean Crustacea (Decapoda,

Anomura) taken by the R/V Soyo-Mam in Japanese waters. I.

Family Chirostylidae. Bulletin of the National Science Museum,

Tokyo, series A 7: 111-134.

Baba, K. 1986. Two new anomuran Cmstacea (Decapoda: Anomura)

from North-West Australia. The Beagle, Occasional Papers of the

Northern Territory Museum of Arts and Sciences 3: 1-5.

Baba, K. 1988. Chirostylid and galatheid Cmstaceans (Decapoda,

Anomura) of the “Albatross” Philippine expedition, 1907-1910.

Researches on Crustacea, Special Number 2: 1-203.

Baba, K. 1990. Chirostylid and galatheid cmstaceans of Madagascar

(Decapoda, Anomura). Bulletin du Museum national d’Histoire

naturelle Paris, series 4, section A 11: 921-975. [Dated 1989].

Baba, K. 2000. Two new species of chirostylids (Decapoda: Anomura:

Chirostylidae) from Tasmania. Journal of Crustacean Biology,

Special Number 2: 246-252.

Davie, P.J.F. 2002. Cmstacea: Malacostraca: Eucarida (Part 2). In:

Wells, A. and Houston, W.W.K. (eds) Zoological Catalogue of

Australia. Vol. 19.3B. CSIRO Publishing: Melbourne, xii + 551 pp.

Gordon, I. 1930. On the species of the galatheid genus Eumunida

(Cmstacea, Decapoda). Proceedings of the General Meetings for

Scientific Business of the Zoological Society of London 1929:

741-753.

Haig, J. 1974. The anomuran crabs of Western Australia: their distri-

bution in the Indian Ocean and adjacent seas. Journal of the Marine

Biological Association of India 14(2): 443-451.

Henderson, J.R. 1885. Diagnoses of the new species of Galatheidea

collected during the “Challenger” Expedition. Annals and

Magazine of Natural History (ser. 5) 16: 407-421.

Henderson, J.R. 1888. Report on the Anomura collected by H.M.S.

Challenger during the years 1873-76. Report on the Scientific

Results of the Voyage of H.M.S. Challenger during the years

1873-76, Zoology , 27, i-vi+1-221, pis 1-21.

Kim, H. S. 1972. Anew species of the family Chirostylidae (Cmstacea:

Anomura) from Jeju Island, Korea. Korean Journal of Zoology 15:

53-56.

Ortmann, A. 1892. Die Decapoden-Krebse des Strassburger Museums

IV. Die Abtheilungen Galatheidea und Paguridea. Zoologischen

Jahrbuchem, Abtheilung fur Systematik, Geographie und Biologie

der Tiere 6: 241-326, pis 11, 12.

Saint Laurent, M. de, and Macpherson, E. 1990. Cmstacea Decapoda:

le genre Eumunida Smith, 1883 (Chirostylidae) dans le eaux neo-

caledoniennes. In: A. Crosnier (ed.), Resultats des Campagnes

MUSORSTOM, Vol. 6. Memoires du Museum national d’Histoire

naturelle, Paris A 145: 227-288, figs 1-17.

Saint Laurent, M. de, and Poupin, J. 1996. Cmstacea Anomura: Les

especes indo-ouest pacifiques du genre Eumunida Smith, 1880

(Chirostylidae). Description de six especes nouvelles. In: A.

Crosnier (ed.), Resultats des Campagnes MUSORSTOM, Vol. 15.

Memoires du Museum national d’Histoire naturelle, Paris A 168:

337-385, figs 1-13.

Smith, S.L 1883. Preliminary report on the Brachyura and Anomura

dredged in deep water off the south coast of New England by the

United States Fish Commission in 1880, 1881, and 1882.

Proceedings of the United States National Museum 6: 1-57.

Tirmizi, N.M. 1964. Cmstacea: Chirostylidae (Galatheidea). The John

Murray Expedition 1933-34, Scientific Reports 11: 167-234.

Van Dam, A.J. 1939. Uber einige U roptychus- arten des museums zu

Kopenhagen. Bidrajen tot de Dierkunde Uitgegeven door het konin-

klijk zoologisch genootschap “Natur Artis Magistra” de

Amsterdam 27: 392-407.

Van Dam, A.J. 1940. Anomura, gesammelt vom Dampfer “Gier” in der

Java-See. Zoologischer Anzeiger, Leipzig 129: 95-104.

Memoirs of Museum Victoria 61(1): 65-73 (2004)

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

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

Sicafodiidae, fam. nov. for Sicafodia stylos , gen. nov., sp. nov., from the marine

bathyal of south-eastern Australia (Crustacea: Amphipoda: Gammaridea)

Jean Just

Abstract

Keywords

Museum of Tropical Queensland, 78-102 Flinders Street, Townsville Queensland 4810, Australia

(jeanjust@bigpond.com.au)

Honorary Associate, Museum Victoria, PO Box 666E, Melbourne, Victoria 3000, Australia

Just, J. 2004. Sicafodiidae, fam. nov. for Sicafodia stylos , gen. nov., sp. nov., from the marine bathyal of south-eastern

Australia (Cmstacea: Amphipoda: Gammaridea). Memoirs of Museum Victoria 61(1): 65-73.

Anew genus and species, Sicafodia stylos, is described from bathyal depths (400-1277 m) off south-eastern Australia.

Following a discussion of its possible placement in existing families, a new family, Sicafodiidae, a potential sister taxon

to the Pardaliscidae, is established. The new species is characterised by the extreme elongation of its conically bundled

mouthparts including the inner plate of the maxillipeds, a completely smooth body, short, broadly rounded coxal plates,

slender, simple but strongly dissimilar gnathopods and elongate pereopods. An analysis of the functional morphology of

the mouthparts, including muscles in the upper lip and epistome, indicates that Sicafodia stylos feeds by piercing its prey

and sucking up macerated/liquefied food. The actual food items are not kn own.

Cmstacea, Amphipoda, Sicafodiidae, fam. nov., Sicafodia, deep-sea, Australia

Introduction

The new species described below was collected as part of the

South-Eastern Australian Slopes Program of Museum Victoria

and the Victorian Institute of Marine Sciences. The program

commenced in 1986 with sampling at bathyal depths (approxi-

mately 200 to 2900 metres) off southern New South Wales,

Bass Strait and eastern Tasmania, followed by a similar sam-

pling program in 1988. A highly diverse and novel fauna of

Peracarida was brought to light. A preliminary analysis of the

composition and diversity of Isopoda and comments on other

groups of Crustacea, together with environmental data from

this stretch of continental margin towards the Tasman Sea, was

provided by Poore et al. (1994).

The length of specimens was measured from the tip of the

rostrum along the dorsal curvature to the tip of the telson. The

specimens are deposited in the collections of Museum Victoria,

Melbourne, Australia.

Classification of the new species

Using Barnard and Karaman’s (1991) key to families, the new

species falls among Ochlesidae (as Ochlesinae in their broad

concept of the Iphimediidae) and Lafystiidae on account of its

lack of a mandibular molar and the reduced number of articles

(three) in the maxillipedal palp. Using the key in Coleman and

Barnard (1991) the species keys out in the vicinity of

Lafystiidae and Laphystiopsidae on account of its simple

gnathopod 1, rounded, non-elongate coxae 1-3, and lack of

mandibular raker setae. None of these characters is unique to

those families. Bousfield (1983), Barnard and Karaman (1991)

and, by implication, Coleman and Barnard (1991) assumed that

at least Lafystiidae, and for the last two works possibly also

Laphystiopsidae, are basically related to the iphimedioids. A

recent phylogenetic analysis of amphipod families (Berge et al.,

2000), however, places those two families in separate clades,

neither with iphimedioid families. The Lafystiidae do, how-

ever, show some similarities with the iphimedioids (see discus-

sion below) in the elongate, acuminate coxa 4 and short, robust

pereopods with curved dactyli, the conically pointed, but

weakly modified mouthparts, simple gnathopod 1 and sub-

chelate gnathopod 2. For these reasons, the family is implicitly

included in the general discussion of the iphimedioids below.

Laphystiopsidae have simple, identical gnathopods similar to,

but smaller than, pereopods 3 and 4, and quadrately bundled

unmodified mouthparts. Laphystiopsidae bear little resem-

blance to the new species and will not be discussed further. For

Ochlesidae see discussion below.

The most immediately obvious characteristics of Sicafodia

stylos is the extreme elongation and acuteness of the down-

ward, backward pointing mouthpart bundle. Conically bundled,

pointed mouthparts are uncommon in gammaridean

Jean Just

Amphipoda. They are found primarily in the conicostomatid

group of genera in the Lysianassoidea (see Lowry and Stoddart,

1983), in monotypic Didymocheliidae, in the iphimedioids, and

in two genera of the Pardaliscidae.

Sicafodia stylos clearly does not belong in the

Lysianssoidea, species of which have a characteristic body

shape and an elongate ischium and mitten shaped propodus of

gnathopod 2. Whilst there are some similarity in the mouth-

parts, notably in the enlarged, folded outer plate and the elon-

gate inner plate of the maxillipeds in some conicostomatid

genera, these similarities must have be independently derived

to serve a similar function (see section on mouthpart function

below).

The three known species of Didymochelia (see Bellan-

Santini and Ledoyer, 1986, Lowry and Stoddart, 1995) have

moderately conical mouthpart bundles, but little or no modifi-

cation of mouthparts with regard to elongation or reductions.

Furthermore, didymocheliids have gnathopods 1 and 2 chelate,

pereopods 3-7 short and strong with curved dactyli, uropod 3

reduced to a stubby ramus only, and the telson shorter than

wide, entire. Sicafodia stylos cannot be referred to the

Didymocheliidae.

Coleman and Barnard (1991) divided the non-eusiroid

iphimedioids into six families: Iphimediidae, Ochlesidae,

Acanthonotozomatidae, Acanthonotozomellidae, Dikwidae and

Odiidae. Berge et al.(1999) merged Ochlesidae and Odiidae

under the former name. Species in these families have conic-

ally bundled, more or less elongate mouthparts with a variety of

strong or less strong reductions. The strongest elongations and

reductions are found in species of Ochlesidae s.s. None, how-

ever, exhibits the extremes found in Sicafodia stylos.

Iphimedioids have moderately compressed bodies (broadly

rounded dorsally in S. stylos), dorsal spines and/or carina

(smooth), one or more of coxae 1-4 elongate, tapering and

acuminate, or elongate and oddly shaped (Ochlesidae s.s.)

(approximately as long as broad, non-tapering, apex broad,

rounded), one or more of pleonal sideplates 1-3 with acute pos-

teroventral corner and often with additional cusp on posterior

margin of sideplate 3 (broadly rounded, no cusps), pereo-

pods 3-7 short, strong, ‘clinging’ with strong, curved dactylus

(slender, elongate, dactylus slender, nearly straight). Some

iphimedioid families have simple, similar or dissimilar

gnathopods 1 and 2; in some families both gnathopods are sub-

chelate or chelate; in others a combination; while they are

carpo- or merochelate in Ochlesidae s.s. Simple gnathopods

occur in many gammaridean families. Thus, there are no char-

acters that unequivocally places S. stylos within the iphi-

medioids, while its smooth dorsum, elongate, slender pereo-

pods 3-7 and short, rounded coxae 1-4 in particular preclude

such an allocation.

Sicafodia stylos shares with Pardaliscidae a smooth body,

short simple coxae 1-4, simple gnathopods 1-2, slender pereo-

pods 3-7 (5 and 7 assumed), and lanceolate uropodal rami.

However, the gnathopods in S. stylos are highly dissimilar,

whereas in the Pardaliscidae they are generally of similar shape

and size. Pardaliscids normally have quadrately bundled

mouthparts, with broad molar incisors and strongly reduced (or

absent) inner plate of the maxillipeds (prominent and much

elongate in S. stylos ). The exceptions to quadrate mouthparts in

pardaliscids are Halicella Schellenberg (monotypic) and

Rhynohalicella Karaman (monotypic), in both of which mouth-

parts form a small downward or forward pointed cone respec-

tively, but the structure and reductions of individual mouthparts

differ significantly from those of S. stylos.

Conical bundling with some elongations and reductions in

mouthparts clearly has been independently derived in several

non-related families, and therefore does not in itself place

S. stylos within any of the groups discussed. Disregarding the

mouthparts, S. stylos appears to be closer to the Pardaliscidae

than to any of the other families mentioned. The prominent

inner plate of the maxillipeds and the dissimilar gnathopods

would appear to preclude the inclusion of the new species in

that family, although it could be conceived as representing a

separate subfamily taxon in the Pardaliscidae with extremely

apomorphic mouthparts, but having retained a plesiomorphic

inner plate of the maxillipeds, reduced or lost in all other mem-

bers of the family (the sister clade). However, because of the

lack of males, and lack of complete antennae 1 and 2 and pereo-

pods 5 and 7 in all specimens, which might provide firmer clues

to the placement of this species, I hesitate to place Sicafodia

stylos in the Pardaliscidae. Instead, I refer it to a new family,

Sicafodiidae, a potential sister taxon of Pardaliscidae.

Sicafodiidae fam. nov.

Diagnosis. Body broad, smooth; cuticle smooth. Cephalon

short with well developed rostrum. Pereonite 1 enlarged, partly

overlapping cephalon. Pleonal sideplates broadly rounded.

Primary flagellum of antennae multiarticulate (not confirmed

for antenna 2), accessory flagellum present, 1 -articulate, antenna

1 with callynophore commencing on first flagellar article.

Coxal plates 1—4 apically rounded, barely longer than broad,

overlapping, forming an even ventral curve, coxa 4 barely

excavate posteriorly. Mouthparts elongate, grouped into

sharply pointed downward-backward pointing cone. Upper lip

apically notched. Mandibles without molar and spine row, with

3-articulate palp, both mandibles with rod-shaped lacinia

mobilis. Lower lip with mandibular lobes, inner lobes absent.

Maxilla 1 without palp, inner plate digitiform, outer plate with

9 distal spine-like setae. Maxilla 2 plates slender, with apical

and subapical setae only. Maxilliped with 3-articulate palp,

outer and inner plates of subequal length. Gnathopods 1 and 2

simple, strongly dissimilar. Pereopods 3-7 slender, dactyls

slender, nearly straight, pereopods 5-7 with article 2 expanded.

Urosomites free. Uropods with full complement of slender,

lanceolate rami, peduncle of uropod 3 short. Telson short, lam-

inate, apically incised. Coxal gills simple sacks, on pereopods

2-7. Oostegites on pereopods 2-5, 2-4 broad, sparsely setose,

5 linear.

Type genus . Sicafodia gen. nov.

Sicafodia gen. nov.

Type and only species. Sicafodia stylos sp. nov.

Diagnosis. With the characters of the family.

A new bathyal amphipod family

Etymology. The name of the genus is derived from Latin, sica,

dagger, and fodio, to stab, hence ‘the one that stabs with a

dagger’, alluding to the shape of the mouthpart bundle.

Feminine.

Sicafodia stylos sp. nov.

Figures 1-4

Material examined. Holotype. Female with developing ovaries, 3.7

mm. Australia, eastern Tasmania, off Freycinet Peninsula, 41° 57.50'S,

148° 37.90'E, 400 m, coarse biogenic rubble, mud, WHOI epibenthic

sled, M.F. Gomon et al., RV Franklin, 27 Jul 1986 (stn SLOPE-48),

NMV J23915 (condition: antennae, some pereopods and tip of some

uropods broken).

Paratypes (5 specimens, all from Australia). Same data as holotype,

NMV J23916 (female A, with apparent traces of broken oostegites;

condition: antennae and most pereopods and uropods broken, telson

missing). Eastern slope of Bass Strait, 38° 21.90'S, 149° 20.00’E, 1000

m, WHOI epibenthic sled, G.C.B. Poore et al., RV Franklin, 23 July

1986 (stn SLOPE-32), NMV J23917 (female B, with fully developed

oostegites, 4.6 mm; condition: antennae, some pereopods and uropods

broken), NMV J23918 (female C, with fully developed oostegites;

condition: half specimen, everything posterior to pereonite 6 missing,

antennae and most pereopods broken), NMV J23919 (female D, with

fully developed oostegites; condition: half specimen, everything poste-

rior to pereonite 7 missing, antennae and most pereopods broken).

Eastern Bass Strait slope, 67 km S of Point Hicks, 38° 23.78'S, 149°

17.02'E, 1119-1277 m, fine mud, WHOI epibenthic sled, G.C.B. Poore

et al., RV Franklin, 25 Oct 1988 (stn SLOPE-67), NMV J23920

(1 specimen, condition: antennae, all pereopods and most of uropods

broken).

Description. Cephalon depth approximately 2.5 times mid-

lateral length. In lateral view cephalon tapering towards right

angled ventral comer. Ocular lobes broadly rounded, continu-

ing in nearly straight line to ventral comer. Eyes absent.

Rostrum about as long as deep in lateral view, apex rounded tri-

angular, approximately half length of non-rostral part of

cephalon, ventrally forming a narrow angle with elongate,

convex frons part of cephalon. Epistome short, unproduced.

Pereonite 1 as long as its anterior depth, as long as 2 and 3

combined. Pleosome dorsal length equalling combined length

of pereonites 2-7, pleonites of equal length.

Coxal plates 1-4 with smoothly rounded ventral margin;

plates 1-3 of subequal length when flattened, (appear to

increase in length from 1 to 3 in Fig.l habitus, due to 1 and 2

curving slightly inwards), of rounded rectangular shape but

increasing in width from 1 to 3; coxa 1 length: width 1.25, coxa

2 length: width 1.20, coxa 3 length: width 1.0, coxa 4

length: width 0.70, with rounded angular posterior projection

and weak concavity posterior to projection; coxa 5 with lobes

equally deep, anterior lobe wider, evenly rounded, posterior

lobe rounded rectangular; coxa 6 similar to 5, but posterior

lobe approximately third deeper than anterior lobe; coxa 7 an

evenly rounded semicircle. Pleonal sideplates with smooth

margin; plate 1 with weakly angular apex, 2 and 3 rounded.

Antenna 1, peduncular article 1 as long as 2 and 3 combined,

approximately 10% longer than lateral width, articles 2 and 3 of

subequal length, both somewhat telescoped into preceding

article, article 3 with lateral apical margin extended beyond

insertion of accessory flagellum; accessory flagellum approx-

imately half length of flagellar article 1, distally tapering, with

1 apical and 1 subapical setae; article 1 of flagellum approx-

imately two-fifths as wide as long, with single-field callyno-

phore composed of 4-5 irregular rows of aesthetascs, article 2

and 3 with 2 distal rows of similar aesthetascs. Antenna 2 with

broad, apically pointed antennal cone; peduncular article 4

approximately twice as long as wide, reaching to apex of fla-

gellar article 1 of antenna 1. (Antennae otherwise broken in all

specimens, but the shape of the proximal parts known suggests

that the flagellum of both antennae is simple, multiarticulate).

Upper lip approximately 2.5 times longer than wide at base

(anterior view), narrowing in middle, tapering towards bilobed

apex, apical notch forming a narrow angle. Mandibles styliform

distal to insertion of palp, reaching to apex of upper lip; left

mandible with irregularly notched apex, lacinia mobilis a long

slender rod terminating in strongly oblique cutting edge with 6

or 7 small denticles; right mandible with 3 blunt teeth apically,

lacinia mobilis quarter to half length of left lacinia mobilis,

with bifid apex; strong palp inserted in proximal third of

mandibular body, reaching slightly beyond apex of mandible,

article 1 as long as broad, article 2 approximately 4 times

longer than broad with 2-3 distal setae, article 3 approxim-

ately three-quarters length of 2, with up to 4 setae on posterior

margin and approximately 4 apical setae, lateral surface partly

covered with fine setules. Lower lip with styliform lobes, lobes

finely setose apically not quite reaching apex of mandibles;

mandibular lobes short, slender, pointing directly backwards.

Maxilla 1 outer plate approximately 1 1 times longer than great-

est width; distal 9 spine-like setae falling into three groups:

2 apical setae (one coarsely serrate along entire length, one

finely serrate in distal quarter), 5 setae closely set on strongly

oblique margin (regularly denticulate in distal half), 2 poorly

articulating, flattened more proximal setae closely adpressed to

medial surface of plate, coarsely and irregularly serrate along

entire margin; inner plate approximately third length of outer,

slender, cylindrical, tapering towards rounded apex, without

setae. Maxilla 2 outer plate approximately 9 times longer than

greatest width, with tuft of approximately 6 unequally long

apical setae; inner plate approximately 80% length of outer,

with 4 apical simple setae and 3 short, subapical medial bi- or

trifid setae. Maxilliped palp article 1 ovoid, approximately 50%

longer than wide, article 2 approximately 80% length of 1, of

similar shape, article 3 as long as 1 and 2 combined, third width

of 2, approximately 9 times longer than wide with 3 or 4 groups

of 2 or 3 setae on midlateral surface and tufts of apical and sub-

apical setae; outer plates broad in proximal third, tapering

towards truncate apex, apex with row of at least 8 blunt, rod-

like setae evenly decreasing in length from medial to penulti-

mate lateral seta, lateralmost seta stouter than the rest, approxi-

mately as long as medialmost one, curved mediad, with

denticles in distal quarter, (because of the natural forward fold-

ing of outer plates around remaining mouthparts, Fig. 2C mxp

dv, the lateralmost, denticulate seta, Fig. 2A mxp op, is func-

tionally in a medial position, cf. above); inner plates nearly as

long as outer plates, each plate slender with nearly parallel mar-

gins, terminating in bluntly pointed apex with what appears to

be glandular tissue in terminal part (Fig. 2C mxp ip), plates

Jean Just

Figure 1. Sicafodia stylos, gen. nov, sp. nov. Holotype, except B, paratype, female B (NMV J23917). al, antenna 1; a2, antenna 2; af, accessory

flagellum; c, head; lv, lateral view; or, outer ramus up3; t, telson; up3p, uropod 3 peduncle; us, urosome. Habitus scale bar: 1 mm.

A new bathyal amphipod family

Figure 2. Sicafodia stylos, gen. nov., sp. nov. A, paratype, female A (NMV J23916); C, paratype, female C (NMV J23918); D, paratype, female

D (NMV J23919); H, holotype. av, anterior view; dv, dorsal view; ep, epistome; ip, inner plate; 1, left; 11, lower lip; 1, r md, mandible; mxl, 2,

maxilla 1, 2; mxp, maxilliped; op, outer plate; r, right; ul, upper lip; vv, ventral view; asterisk: tip of mandible and lacinia mobilis broken.

Jean Just

Figure 3. Sicafodia stylos, gen. nov., sp. nov. Holotype, except B, paratype, female B (NMV J23917). ex, coxa; gp, gnathopod; o, oostegite;

p, pereopod; pi, pleopod.

A new bathyal amphipod family

with small opposing notch in medial margin proximal to

‘gland’, each plate laterally, between notch and ‘gland’, with

tuft of 4 or 5 inward curving setae decreasing in length from

lateral to medial.

Gnathopod 1 basis approximately 4 times longer than broad,

as long as merus and carpus combined, with anterior longi-

tudinal concavity to receive article 5 when fully folded back;

ischium approximately twice as long as lateral width; merus tri-

angular, anteriorly pointed, approximately as long as ischium,

with carpus inserted on anterior margin; carpus slender, twice

as long as merus, with parallel margins, approximately 6 times

longer than wide, unarmed; propodus approximately 65%

longer than carpus, 12 times longer than broad at base, slightly

curved and faintly tapering distally, with a few short posterior

setae; dactylus weakly curved, approximately sixth length of

propodus, with posterior tooth near base of stout, curved

unguis, 1 straight, simple seta level with tooth and 2 curved,

blunt, simple setae at posterior base of and adpressed to unguis.

Gnathopod 2 little more than half the length of 1, without

elongate elements; carpus approximately 80% the combined

length of ischium and merus, triangular, approximately 50%

longer than lateral distal width, posterodistal lobe setiferous,

not projecting along propodus; propodus approximately 10%

longer than carpus, moderately curved, tapering distally, with 4

or 5 posterior groups of stiff, pectinate setae, anterior margin

with group of similar setae at distal two-thirds; dactylus half

length of propodus, similar to that of gnathopod 1 but with 2

small midposterior teeth.

Pereopods 3 and 4 similar, slender, approximately 4 times

longer than respective coxal plates; merus and carpus subequal

in length, combined as long as basis; propodus twice as long as

carpus; merus, carpus and propodus with a few posterior and

anterior small setae, propodus with small posterodistal robust

seta; dactylus half length of propodus, nearly straight, length

approximately eight times width at base. Pereopods 5-7

apparently alike (pereopod 6 the only entire limb known, hence

Jean Just

relative lengths of 5-7 not known, but probably of subequal

length), with posteriorly expanded distally rounded basis,

posterior margin smooth with scattered setules, with midlateral

and anterolateral ridges; at least pereopod 6 approximately

third longer than pereopods 3 and 4, but otherwise of similar

shape and proportions, carpus posterodistally and propodus

posteriorly with groups of slender robust setae.

Pleopods strong, alike; peduncle approximately twice as

long as wide (posterior view), with 2 mediodistal coup-

ling hooks; subequal rami as long as peduncle, proximal part

of rami with opposing surfaces excavate, outer ramus with

posterior proximal locking projection.

Urosomite 1 as long as 2 and 3 combined; urosomite 2

dorsal length approximately quarter that of elongate 3; uro-

somite 3 dorsally flattened to slightly concave in transverse

section. Uropods 1 and 2, peduncles with lateral robust setae;

uropod 1 overreaching 2 and outer ramus of 3, (tip of uropod 1

rami broken in all specimens, but proportions between rami and

peduncle probably approximating those in uropod 2); uropod 2

shorter than 1, reaching nearly to tip of outer ramus of 3, outer

ramus as long as peduncle, inner ramus approximately 40%

longer than outer; uropod 3 biramous (inner ramus lost in all

specimens, but presence indicated by socket and musculature),

peduncle as long as wide, reaching to tip of telson, outer ramus

approximately 5 times longer than peduncle; rami of all

uropods slender lanceolate, without terminal seta(e), but appar-

ently all with a few small lateromarginal setae, margins finely

serrate. Telson approximately 10% broader than long, reaching

to distal margin of peduncle of uropod 3, quarter cleft in open

V-shape; lobes rounded, each with 2 distolateral marginal teeth,

2 or 3 small pappose setae level with teeth and 1 long similar

seta at mid-surface.

Size. Largest female, 4.7 mm. Male not known.

Etymology. The species name is derived from the Latin stylus

meaning pointed instrument or pen, and os meaning mouth,

alluding to the long, pointed mandibles. Noun in apposition.

Distribution. South-eastern Australia, 400-1277 m depth.

Function of the mouthparts

Conically bundled, elongate, pointed mouthparts are frequently

referred to as having piercing and possibly sucking functions.

In most cases the piercing action can be readily understood,

whereas the subsequent transport of food (particles, macerated

tissue, body fluids) has not been studied in detail. Dahl (1964)

discussed the probable function of the mouthparts and adjacent

parts of the digestive tract in Acidostoma neglecta Dahl, 1964

(Lysianassoidea; ectoparasite on sea anemones, Vader, 1983).

He found that muscles in the oesophagus and the anterior part

of the stomach may create the necessary negative pressure to

lift up food through the narrow passageways of the conically

bundled mouthparts.

In spite of many differences in mouthpart details, the gen-

eral outline of the bundle is quite similar between Acidostoma

and Sicafodia. Sicafodia also seems to possess the two pre-

requisites necessary for sucking up food through the mouth

cone, viz., a tight lumen, and a sucking mechanism. When the

maxillipeds are brought fully forward, a tight funnel may be

created between the forward-inward folded outer plates and the

elongate upper lip (as in Acidostoma)', the equally elongate

inner plates of the maxillipeds will help seal the funnel poster-

iorly. Once the apex of the mouthpart cone is embedded in the

food item tissue and a tight funnel has been established, a suck-

ing devise may be found in the proximal part of the upper lip,

where obliquely set transverse muscles (Fig. 2A ul, lv) will, on

contraction, pull the soft-walled posterior surface of the upper

lip forward thus drawing liquid or semi-liquid food upward

through the funnel. The obliquely transverse muscles in the

epistome may have a similar function, bringing food closer to

the oesophagus. Additional mechanisms, like those described

by Dahl (1964), have not been observed but may be present to

finally lift food into the stomach.

Following the initial piercing by the mandibles, the apical

and subapical spine-like setae located on the maxillipedal outer

plate and the outer plate of maxilla 1 may help in extending the

wound and shredding the tissue. If the tips of the maxillipedal

inner plates are indeed glanduliferous as suggested in the

description, such glands may release proteolytic enzymes into

the wound and thus help speeding up the maceration process

before suction transport.

The actual food items of Sicafodia stylos are not known.

However, the lack of grasping gnathopods or strongly dacty-

late, clinging pereopods suggest that Sicafodia is not a fish

parasite. In view of its slender epibenthic/pelagic pereopods

3-7 and the strongly developed pleosome the species is

probably a roaming predator on some group or groups of

sessile or sluggish benthic or epibenthic invertebrates.

Acknowledgments

The material reported was obtained during two cruises on board

the Australian National Facility ship the RV Franklin in 1986

and 1988. The South-Eastern Australian Slopes Program of

Museum Victoria, Melbourne, and the Victorian Institute of

Marine Sciences (no longer in existence), Melbourne, and my

participation in that program were supported by a grant from

the Australian Department of Science’s Marine Sciences and

Technology Scheme/Australian Research Council. I thank Jim

Lowry, Australian Museum, Sydney, for valuable discussions

and comments during the preparation of this paper. I also must

acknowledge the great pleasure of discussing this new amphi-

pod with Jerry Barnard on his last visit to Australia, and listen-

ing to his extraordinary ‘spiel’ of talking me through the entire

key to gammaridean fa mi lies from memory. In the end he

looked at the new species in wonder.

References

Barnard, J. L., and Karaman, G. S. 1991. The fa mi lies and genera of

marine gammaridean Amphipoda (except marine gammaroids).

Records of the Australian Museum, Supplement 13(1): 1-417.

Bellan-Santini, D., and Ledoyer M. 1986. Gammariens (Crustacea,

Amphipoda) des iles Marion et Prince Edward. Bolletino del Museo

Civico di Storia Naturale Verona 13: 349^435.

Berge, J., Boxshall, G., and Vader, W. 2000. Phylogenetic analysis of

the Amphipoda, with special emphasis on the origin of the

A new bathyal amphipod family

Stegocephalidae. Polskie Archiwum Hydrobiologii 47(3-4):

379-400.

Berge, J., Vader, W., and Coleman, O. 1999. A cladistic analysis of the

amphipod families Ochlesidae and Odiidae, with description of a

new species and genus. Crustaceans and the Biodiversity Crisis:

239-265, Koninklijke Brill NV: Leiden.

Bousfield, E.L. 1983. An updated phyletic classification and palaeo-

history of the Amphipoda. In: Schram, F.R. (ed.), Crustacean

Phylogeny. Crustacean Issues 1 : 257-277.

Coleman, C. O., and Barnard, J. L. 1991. Revision of Iphimediidae and

similar families (Amphipoda: Gammaridea). Proceedings of the

Biological Society of Washington 104(2): 253-268.

Dahl, E. 1964. The amphipod genus Acidostoma. Zoologische

Mededelingen 39: 48-58.

Lowry, J.K. and Stoddart, H.E., 1983. The shallow-water gammaridean

Amphipoda of the subantarctic islands of New Zealand and

Australia: Lysianassoidea. Journal of the Royal Society of New

Zealand 13(4): 279-394.

Lowry, J.K., and Stoddart, H.E. 1995. A new species of Didymochelia

from New Caledonia (Crustacea: Amphipoda: Didymocheliidae).

Bulletin du Museum national d’Histoire naturelle, Paris (4) 17

(1-2): 193-200.

Poore, G.C.B., Just, J., and Cohen, B.F. 1994. Composition and diver-

sity of Crustacea Isopoda of the southeastern Australian continental

slope. Deep-Sea Research 41(4): 677-693.

Vader, W. 1983. Associations between amphipods (Crustacea:

Amphipoda) and sea anemones (Anthozoa, Actiniaria). Memoirs of

the Australian Museum 18: 141-153.

Memoirs of Museum Victoria 61(1): 75-83 (2004)

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

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

Pseudidotheidae (Crustacea: Isopoda: Valvifera) reviewed with description of a

new species, first from Australia

Gary C. B. Poore and Tania M. Bardsley

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

Abstract Poore, G.C.B., and Bardsley, T.M. 2004. Pseudidotheidae (Cmstacea: Isopoda: Valvifera) reviewed with description of

a new species, first from Australia. Memoirs of Museum Victoria 61(1): 75-83.

The family Pseudidotheidae, comprising only four species of Pseudidothea, is reviewed. A new highly-ornamented

species from south-eastern Australia, Pseudidothea hoplites, is described. A key to species of the genus and family is

provided.

Keywords Isopoda, Valvifera, Pseudidotheidae, Pseudidothea , new species, south-west Pacific, Australia

Introduction

The family Pseudidotheidae Ohlin, 1901 was erected for the

east Patagonian genus and species Pseudidothea bonnieri

Ohlin, 1901 on the basis of the male first pleopods being

modified for a copulatory function, a character never before

recorded in the Isopoda. Ohlin also noted that the second to

seventh pereopods are of virtually the same size and form. He

regarded the Pseudidotheidae as an intermediate link between

the Idoteidae Samouelle, 1819 and Arcturidae Dana, 1849.

Barnard (1920) noted that the Pseudidotheidae have a flattened

body, pereonite 4 never elongate, pereopod 1 prehensile and

pereopods 2-4 stout. In Arcturidae the body is cylindrical,

pereonite 4 often elongate, pereopod 1 setiferous and pereo-

pods 2-4 slender and setiferous. Nordenstam (1933) added

another character: penial processes fused but distally cleft or

bilobate.

Pseudidotheidae are distinguished, with other arcturoid

valviferans, from Idoteidae and similar families by having the

head fused to pereonite 1 (with a few exceptions not so in

Idotedidae), penial processes fused but distally cleft, and pleo-

pod 1 with an elongate peduncle and modified exopod in the

male. Wagele (1989, 1991) treated the family as one of four

subfamilies of Arcturidae, the others being Arcturinae,

Xenarcturinae Sheppard, 1957 and Holidoteinae Wagele, 1989.

He included Arcturides Studer, 1882 with Pseudidothea in

Pseudidotheinae which, he believed, shared the synapomorphy

of all pleonites fused and not divided by furrows. This state is

true of all arcturoid families, with rare reversals in two genera

(Poore, 2001). Arcturididae Poore, 2001 ( Arcturides alone),

Pseudidotheidae and Xenarcturidae were treated as families by

Poore (2001), three members of an unresolved clade. Poore

(2001) separated pseudidotheids from other arcturoid families

by the uniquely undifferentiated pereopods 2^1, similar to more

posterior ambulatory pereopods. While it is true that all limbs

are ambulatory and 2—4 do not bear long filtering setae of the

arcturid type, pereopods 2 and 3 are more robust than 4-7. In

Xenarcturidae, only pereopods 2 and 3 are slender, setose and

arcturid-like while pereopods 4-7 are ambulatory. In

Arcturididae, pereopods 2-7 are all similar and ambulatory.

Holidoteidae was only remotely related in Poore’s (2001)

cladogram (see revision by Poore, 2003).

The family contains only Pseudidothea, the type species of

which is a junior synonym of an earlier described species,

“ Idothea Miersii ” Studer, 1884, an observation suggested by

Ohlin, suspected by later authors, and confirmed here.

Microarcturus scutatus Stephensen, 1947 from the South

Shetland Islands, was transferrred to Pseudidothea by

Sheppard (1957). Hurley (1957) described Pseudidothea

richardsoni from New Zealand. Here, a fourth species is

described from southern Australia.

All limbs are drawn from the left side unless otherwise

stated. The following abbreviations are used in figures: Al, A2,

antennae 1, 2; MD, mandible; MP, maxilliped; MX1, maxilla 1;

MX2, maxilla 2; P1-P7, pereopods 1-7; PL1-PL5, pleopods

1-5; U, uropod; 1, left; r, right. Material is lodged at Museum

Victoria, Melbourne (NMV); the Canterbury Museum,

Christchurch, New Zealand (CMNZ), Museum of New Zealand

(Te Papa Tongarewa) (MNZ), Zoological Institute and

Museum, Hamburg, and Museum fur Naturkunde, Berlin.

Gary C. B. Poore and Tania M. Bardsley

Pseudidotheidae Ohlin

Pseudidotheidae Ohlin, 1901: 274-276. — Stebbing, 1905: 43. —

Barnard, 1920: 381.— Nordenstam, 1933: 112-113.— Hale, 1946:

168.— Sheppard, 1957: 173-174.— Poore, 2001: 227.

Pseudidotheinae.— Wagele, 1989: 137-138.— Wagele, 1991: 80-81.

Diagnosis. Body strongly vaulted. Head and pereonite 1 fused.

Pereonite 4 of similar length to pereonite 3. All pleonites fused

into pleotelson. Body variously tuberculate or spinose; pleo-

telson without dorsolateral ridges ending in mediodorsal

posterior spine, never with posterior dorsolateral pair of strong

spines; limbs and most of surface covered with fine setae that

trap sediment. Dorsal coxal plates 2-7 obsolete, bases of pereo-

pods exposed. Mouthparts and pereopod 1 visible in lateral

view. Eyes well developed. Antenna 2 flagellum of 2 or 3

articles plus distal claw. Pereopod 1 a gnathopod, pereopods

2 and 3 differentiated from ambulatory pereopods 4-7.

Pereopod 1 dactylus evenly curved along anterior margin,

evenly tapering. Pereopods 2 and 3 with propodus able to close

on carpus, articles broad and with posterior robust setae; with

prominent dactylus, unguis short. Pereopods 4-7 similar and

ambulatory. Pereopods of males without dense mat of fine

setae. Uropodal exopod (smaller ramus) tapering (with terminal

setae only), more than half as long as endopod. Oostegites 1-4

functional, not supported by coxal lobes; oostegite 5 present or

absent. Penes fused as a single penial plate, apically simple or

barely slit. Pleopod 1 peduncle more elongate than on other

pleopods; with marginal setae on rami longer than or equal to

length of rami. Pleopod 1 exopod of male thickened and with

groove on posterior face, with few simple setae along straight

lateral margin; with groove on posterior face of exopod ending

on tapering distolateral apical extension. Pleopod 2 of male

with appendix masculina about as long as endopod, basally less

than half width of endopod.

Remarks. The diagnosis is rephrased from Poore (2001) to

better define limb differentiation. We note that the body is

covered with fine setae, not illustrated in the new species and

indicated or mentioned only in passing for other species by

Hurley (1957) and Brandt and Wagele (1990). Such setation is

not a typical valviferan characteristic. Poore (2001) defined the

family as lacking oostegites on pereopod 5; this is true of three

species but not of P. scutata. The presence of a fifth pair of

oostegites in one species is anomalous among arcturoid

families. It is seen elsewhere only in Austrarcturellidae Poore

and Bardsley, 1992 where oostegites 5 are vestigial and act as

egg guides rather than as part of the marsupium.

Wagele (1989) considered Arcturides a family member but

Poore (2001) erected a separate family for this genus.

Pseudidothea Ohlin, 1901

Pseudidothea Ohlin, 1901: 276. — Nordenstam, 1933: 113. — Hale,

1946: 168.— Sheppard, 1957: 174.— Hurley, 1957: 15.— Wagele,

1991: 84-87.

Type species. Pseudidothea bonnieri Ohlin, 1901 (by mono-

typy)-

Diagnosis. As for family.

Remarks. Ohlin (1901) was “almost convinced” that

Pseudidothea bonnieri was identical to Idotea miersii Studer,

1884 and noted that “the localities where they were dredged are

nearly the same.” He nevertheless, erected his new species. He

also expressed his “suspicion” that his new genus was identical

to Arcturides Studer, 1883. Hale (1946) listed three characters

that separated the two genera. In Arcturides, the coxae of pereo-

nites 2-7 are distinctly marked off, the antenna 2 flagellum is

of three articles and the uropodal exopod as long as endopod.

Hale’s view was confirmed by Poore (2001) whose phylo-

genetic analysis concluded the two genera belong in different

families.

Key to species of Pseudidothea

1. Pereonites 2 and 3 with forked dorsolateral spines; all

pereonites with lateral rows of blade-like ridges, each with

anteriorly and posteriorly directed spines; tergites pro-

duced laterally over coxae to form a shield with 3 points;

Pseudidothea hoplites

— Pereonites with low or high flat tubercles; tergites pro-

duced laterally as large tubercles or rounded or flattened

laterally; 2

2. Pereon with large high flat tubercles; pereonite 1 with dor-

sal pair, pereonites 2-4 with dorsal and lateral pair and

pereonites 5-7 with dorsal, dorsolateral and lateral pair;

male pleopod 2 with appendix masculina twice as long as

rami Pseudidothea scutata

— Pereon with low irregular tubercles; male pleopod 2 with

appendix masculina and rami subequal 3

3. Uropodal exopod with a single strong setae, endopod with

3 pappose setae; antenna 2 peduncle with long fine setae

on articles 3-5; pereopods without tubercles; male pleopod

1 endopod with 5 lateral spinules proximally, 5 apical

plumose setae; exopod with 15 spinules on lateral margin,

tapering distally to an obtuse apex

Pseudidothea richardsoni

— Uropod rami each with single seta; antenna 2 peduncle

with short setae on articles 3-5; pereopods with tubercles;

male pleopod 1 endopod with plumose setae marginally;

exopod with 16-17 spinules laterally, with acute apex bent

outwards Pseudidothea miersii

Pseudidothea hoplites sp. nov.

Figures 1-3

Material examined. Holotype. Western Bass Strait, 70 km W of Cape

Farewell, King Island, Tasmania (39°38.2’S, 143°07.2'E), 127 m, sand,

epibenthic sled, R. Wilson on RV Tangaroa, 21 Nov 1981 (stn BSS

195), NMV J8705 (male, 4.4 mm).

Paratypes. Type locality, NMV J8706 (1 female); 36 km SSW of

Stokes Point (40°26.7'S, 143°41.4'E), 85 m, rock dredge, 22 Nov 1981

(stn BSS 198), NMV J8709 (1); 59 km WNW of Cape Farewell

(39°28'S, 143°17'E), 103 m, Smith-Mclntyre grab/pipe dredge,

G.C.B. Poore on HMAS Kimbla, 10 Oct 1980 (stn BSS 81), NMV

J8703 (1).

Victoria, 80 km SSE of Cape Otway (39°26'S, 142°57'E), 113 m, 9

Oct 1980 (stn BSS 67), NMV J8701 (2); NMV J23186 (1 ovigerous

female, 5.1 mm , figured); 51 km SSW of Cape Otway, Victoria

Isopods of the family Pseudidotheidae

(39°16'S, 143°17'E), 90 m, 10 Oct 1980 (stn BSS 73), NMV J8702 (1);

45 km SSW of Cape Otway (39°15’S, 143°19’E), 94 m (stn BSS 74),

NMV J8704 (2 males); 55 km SW of Cape Otway (39°16.7'S,

143°06.7'E), 95 m, rock dredge, R. Wilson on RV Tangaroa, 21 Nov

1981 (stn BSS 193), NMV J8707 (male, female); 44 km SW of Cape

Otway (39°06.3'S, 142°55.6'E), 81 m (stn BSS 192), NMV J23077 (1);

60 km SW of Cape Otway (39°06.3'S, 142°55.6'E), 84 m, fine shell

(stn BSS 191), NMV J8708 (1).

Other material. Tasmania. Breaksea Island, Bathurst Harbour

(43°20’S, 145°57’E), 4 m, NMV J23085 (1 ovigerous female, SEM

examination). Isle des Phoques (42°25’S, 148°10'E), NMV J23084 (1).

Bicheno, eastern side of Waubs Bay, reef (41°53'S, 147°18'E), 7 m,

Macrocystis holdfasts (stn TAS 94), NMV J23081 (1); E side of Waubs

Bay, reef local name “Split Rock” (41°53'S, 147°18'E), 11m, red and

brown algae (stn TAS 102), NMV J23083 (2); granite reef 50 m off-

shore, N end of “The Gulch” (41°53'S, 147°18'E), 7 m, erect red algae

(stn TAS 88), NMV J53071 (1); reef close to base of “Split Rock”

(41°53'S, 147°18'E), 12 m, fine sand from base of reef (stn TAS 96),

NMV J23082 (1).

Victoria. “Harry’s Hole”, W side of Twin Reefs, Venus Bay

(38°41’S, 145°39'E), 9 m, rocky (stn CPA 8), NMV J23079 (1). 75 m

SW of Eagles Nest (38°40’S, 145°40’E), 8 m (stn CPA 3), NMV J23078

(1). Off Eagles Nest (38°40.67'S, 145°38.76'E), 10-11 m, mixed algae

(stn BUN 3), NMV J53073 (1). Aireys Inlet (38°28’S, 144°06'E), from

Sphacelaria, NMV J23080 (1).

Diagnosis. Head dorsally strongly elevated, with paired double

or single spines. Pereonite 1 with 1 pair of small dorsal spines,

2 small lateral spines. Tergites 2-7 each produced laterally in

form of a with 3 points, anteriorly, posteriorly and laterally.

Pereonites 2 and 3 with small paired middorsal spines, large

paired dorsolateral forked spines, and lateral ridges produced

acutely anteriorly and posteriorly; pereonites 4-7 with dorso-

lateral and lateral ridges, each produced acutely anteriorly and

posteriorly, and several anterodorsal and posterodorsal spines.

Pleotelson with anterolateral processes, paired anterodorsal

convexities, a series of 3 spines on each side (dorsal-lateral)

and pair of mediodorsal convexities, remaining pleotelson

tapers to an obtuse apex.

Male antenna 1 flagellum with 6 clusters of aesthetascs.

Antennae and pereopods with tubercles. Antenna 2 with short

setae on articles 3-5. Male pleopod 1 endopod about 1.5 times

as long as peduncle with plumose setae apically; exopod

longer, 8 spinules on lateral margin, tapering distally, thickened

and folded laterally to partially cover a groove that runs to the

apex. Male pleopod 2 appendix masculina styliform, slightly

longer than endopod. Uropodal exopod about two-thirds as

long as endopod; exopod with a strong apical seta; endo-

pod with 3 brush setae and 2 setules. Oostegite absent from

pereopod 5.

Description. Ornamentation. Head dorsally strongly ele-vated,

with paired double or single spines. Eyes prominent, arising

laterally. Lateral margin of head armed with about 7 small

teeth, extending downwards and outwards. Head fused to pere-

onite 1, partial suture visible laterally. Pereonite 1 with pair of

small dorsal spines, 2 small lateral spines, lateral margin with

about four small teeth. Tergites 2-7 produced laterally to form

a shield with 3 spines, laterally, anteriorly and posteriorly.

Pereonites 2 and 3 with small paired dorsal spines, large

paired dorsolateral forked spines, and lateral ridges finished

anteriorly and posteriorly with a small spine. Pereonite 4-7

with dorsolateral and lateral ridges, finished anteriorly and pos-

teriorly with a small spine. Pereonite 4 with 4 anterodorsal and

posterodorsal spines, pereonites 5 and 6 with 3 and pereonite 7

with 2 and a single dorsal denticle.

Sculpture of pleotelson from anterior to posterior as follows.

A central pair of dorsal convexities each with an anteriorly

directed spine, lateral to these a series of 3 anteriorly directed

spines on each side, followed by large lateral convexities on

each side, followed by a pair of central dorsal convexities,

remaining pleotelson tapers to an obtuse apex.

Antennae, mouthparts and limbs (from male). Antenna 1

peduncle articles with brush setae, articles rounded and becom-

ing successively smaller; flagellum article 1 very short; article

2 with 6 aesthetascs and setules. Antenna 2 peduncle articles

3-5 with blunt tubercles on lower margin, bearing robust setae,

especially on articles 4 and 5; flagellum almost as long as

peduncle article 5, articles becoming successively smaller, first

with distal robust seta and setules, second with setules, third a

short claw.

Mandible incisor with 4 uneven teeth; left lacinia mobilis

almost as wide as incisor, with 3 teeth; right lacinia mobilis an

unevenly toothed column; left molar process with concave face

rimmed by obscure teeth and bearing a setal cluster; right molar

process with face ending with row of blunt teeth and bearing

setal cluster. Maxilla 1 inner lobe with 2 long pappose setae;

outer lobe with 11 apical setae, some obscurely dentate.

Maxilla 2 inner lobe oblique margin with 6 pappose setae along

posterior edge, 5 setae on anterior edge; middle lobe with 2

longer pappose setae; outer lobe with 3. Maxilliped endite with

complex of thin pappose setae and rows of blunt tubercles; palp

with tubercles and long setae on mesial margins of articles 2-5;

articles 1 and 2 short, 3 and 4 of subequal length, 3 produced

mesially, article 5 one-fifth as long as 4, almost as long as wide;

epipod apex with small blunt tooth.

Pereopod 1 held close to the mouthparts; merus-propodus

with uneven posterior tubercles and stout pectinate setae;

propodus almost as wide as long, with rows of mesial pectinate

setae along anterodistal margin; proximal part of dactylus

linear, about 2.5 times as long as greatest width, complexly

setose with mesial pectinate setae, 1 spinule on posterior

margin, posterodistal comer of dactylus with a spinule, seta

and 2 setules; unguis a strong claw, less than half length of

dactylus.

Pereopod 2 basis-meras short, subequal, carpus-dactylus

longer; merus with complex tubercle on lower margin bearing

short setae; carpus longer than greatest width, with tubeculate

ridge on lower margin bearing 2 long robust setae; propodus

robust, about twice as long as wide, with 2 robust setae on

lower margin opposing carpus; proximal part of dactylus

almost 3 times as long as wide, unguis a short claw. Pereopod

3 similar to pereopod 2. Pereopods 4-7 basis-merus with blunt

tubercles on upper margin, most articles with well spaced setae

on lower margin; basis about 1.5 times as long as wide; ischium-

carpus subequal, about as wide as long; propodus about 2.5

times as long as wide, dactylus similar to pereopod 2.

Male pleopod 1 peduncle twice as long as wide, with 4

coupling hooks; endopod lamellar with 6 apical plumose setae;

Isopods of the family Pseudidotheidae

Figure 2. Pseudidothea hoplites. Holotype male, NMV J8705. Left and

of endite. Pleopods 1-5; penial process; uropod with detail of ra mi .

exopod longer than endopod with 8 spinules on lateral margin,

tapering distally, thickened and folded laterally to partially

cover a groove that runs to the apex. Male pleopod 2 rami

apically flattened, endopod with 9 apical plumose setae; exo-

pod with 21 marginal plumose setae; appendix masculina styli-

form, slightly longer than endopod. Pleopods 3-5 becoming

successively larger, rami apically rounded with single simple

seta on endopod.

Uropod unarmed, rounded anteriorly, tapering posteriorly;

exopod about two-thirds as long as endopod, conical, with

apical seta; endopod broader, apically rounded with 4 distal

setae and 3 lateral setae.

Sexual differentiation. Female differs from male in broader

right mandibles, maxillae 1 and 2, maxilliped with detail of anterior face

body, especially of pereonites 2-4; ornamentation more

developed; antenna 1 flagellum with 3 clusters of aesthetascs

on article 2; pereopods 1-4 with oostegites, pereopod 5 without

oostegite; penial process absent; pleopods 1 and 2 without male

modifications. Male with ventral terga separate on pereonites

1^4 and fused across midline of pereonites 5-7.

Etymology. Hoplites (Gr.), man in armour, in reference to the

elaborate spines and ridges.

Distribution. South-eastern Australia (Victoria and Tasmania),

4-127 m depth.

Remarks. Pseudidothea hoplites is distinguished from other

species of Pseudidothea by the complex ornamentation of the

Gary C. B. Poore and Tania M. Bardsley

Figure 3. Pseudidothea hoplites. Holotype male, NMV J8705. Antennnae 1 (not all aesthetascs drawn) and antenna 2; left pereopod 1 with detail

of dactylus; inner face of right pereopod 1 propodus. Pereopods 2, 4 and 7, with details of dactyli.

Isopods of the family Pseudidotheidae

pereon comprising forked projections on pereonites 2 and 3,

and dorsolateral and lateral lobes produced front and back. It is

most similar to P. richardsoni from New Zealand but is much

more ornamented. The lateral projections of pereonites 2 and 3

are more exaggerated but both species have anterior and

posterior spines.

Pseudidothea miersii (Studer)

Idothea Miersii Studer, 1884: 17, pi. 1 fig. 5. (lapsus for Idotea)

Pseudidothea bonnieri Ohlin, 1901: 276-281, fig. 6. — Nordenstam,

1933: 114, fig. 27.— Sheppard, 1957: 175-176.

Pseudidothea miersii. — Barnard, 1920: 380-381. — Nordenstam,

1933: 114.— Shepherd, 1957: 175-176, figs Id, 14a-f.— Kussakin,

1967: 267-269, figs 28, 29.

Arcturides miersii. — Nierstrasz, 1941: 262.

Diagnosis. Head dorsally convex; pereon with irregular

minute tubercles. Pleotelson with blunt anterolateral processes,

dorsally smooth, tapering to broadly truncate and slighty

upturned apex. Male antenna 1 flagellum with about 6 clusters

of aesthetascs. Antennae and pereopods without tubercles.

Antenna 2 peduncle with short setae on articles 3-5. Pereopods

minutely setose and with tubercles. Male pleopod 1 endopod

about 1.5 times as long as peduncle with marginal plumose

setae; exopod longer, 16-17 spinules on lateral margin, taper-

ing distally, to acute apex bent outwards, with oblique furrow

opening at apex. Male pleopod 2 appendix masculina tapering

to acute point, slightly longer than endopod. Uropodal exopod

with strong apical setae, endopod with 1 short seta; exopod

about two-thirds as long as endopod (Ohlin, 1901). Oostegite

absent from pereopod 5 (Sheppard, 1957).

Distribution. East Patagonia, Falkland Islands, 115-500 m depth.

Remarks. Studer (1884) based his new species Idothea

miersii on a specimen 9 mm long, collected by the Gazelle

Expedition off the east coast of South America at 47°1'6"S,

63°29'6"W at 63 fathoms (110 m). In the same paper he

redescribed his earlier named species, Arcturides cornutus.

Ohlin (1901) based Pseudidothea bonnieri on two males, 9 mm

long, in the Hamburg Museum. When Ohlin (1901) described

P. bonnieri he was almost convinced that his specimens were

identical with Idothea miersii (Studer) and in a footnote

reported how he had tried to borrow Studer’s material from the

Museum fiir Naturkunde in Berlin but “got the reply that, as

there were only two of them, it would be against the regulations

to send them away from the Museum.” Angelika Brandt com-

pared material from the museums in Hamburg and Berlin on

our behalf:

from Hamburg, a 4 mm manca and a 6.4 mm male (ZMH K-

1877) labelled and catalogued “ Pseudidothea bonnieri ,

Pisagua, Chile, 19°27'S, 70°10W, K. Kophamel 1877-1889”;

and

from Berlin, a 6.2 mm male (18804) labelled “Zool. Mus.

Berlin 18804 Pseudidothea bonnieri (Syntype) Ohlin, 1901

Leg. Kap. Kophamel, 3.VI.1888, 43°6’S, 60°W” and on

another label “ Pseudidothea bonnieri Ohlin, 1901 ( Idothea

miersi (Studer)”, and catalogued in Berlin with further

information, “Hamburger Museum ded. Pisagua”.

Brandt (pers. comm.) could find no differences between the

specimens and concluded that one of Ohlin’s two males had

been donated to the museum in Berlin. This seems certain. The

locality recorded by the two museums, but not the coordinates

and collecting date of the Berlin specimen, is at odds with the

type locality and more recent records of the species and is

clearly wrong. Ohlin must have included antennae in his total

length of 9 mm while Brandt’s measurements of 6.2 and 6.4

mm do not. The manca was not mentioned by Ohlin. Studer’s

material can not now be found although Ohlin’s footnote tells

that it existed in 1901. It is tempting to speculate that, being

unable to borrow Studer’s material and convinced of the syn-

onymy of his species bonnieri with Studer’s miersii, Ohlin sent

one of his syntypes to Berlin for comparison. This may explain

why the Berlin male has two species names but whoever

concluded this remains a mystery.

Sheppard (1957) examined many specimens from the

Falklands region, reported them as Pseudidothea bonnieri but

thought too that I. miersii was a synonym. Kussa ki n (1967) also

illustrated a species using the older species name, as

Pseudidothea miersii, and noted that P. bonnieri is probably a

synonym. He observed that slight differences exist: in

P. bonnieri the second article of the peduncle of antenna 1 bears

a rounded tubercle with four setae (referring to Sheppard,

1957); in Kussakin’s specimens there is a slight swelling with

five setae. The epipod of the maxilliped in Sheppard’s illus-

tration of P. bonnieri has slightly concave lateral margins, while

in Kussakin’s specimens it has a regular oval form with convex

lateral margins. We consider that these minor differences can be

attributed to intraspecific variation or mounting.

To add to the confusion, Nierstrasz (1941) synonymised

Studer’s two species, Pseudidothea bonnieri and Arcturides

cornutus, without explanation. It seems improbable that

Studer could confuse his own two species in one paper and

specimens of A. cornutus in our possession look nothing like a

pseudidotheid; in fact, Poore (2001) placed the two species in

different families.

We conclude, with Kussa ki n (1967), that P. bonnieri should

be treated as a junior synonym of P. miersii. We treat as

additional evidence the observation that all authors have

reported only one species like this off eastern South America;

the only other in the genus in the region, P. scutata Stephenson,

1947 is quite different.

Pseudidothea richardsoni Hurley

Figure 4

Pseudidothea richardsoni Hurley, 1957: 15-17, figs 74-91.

Material examined. New Zealand, Banks Peninsula region. Off

Lyttelton, 4 fm [7.3 m], H. Suter, CMNZ (4 females, 5. 1-6.1 mm; 4

males, 4.8-5. 1 mm, 1 figured); NMV J47116 (1 female, 1 male). Big

Bay, mud bottom, 12 m, MNZ CR-9846 (2). Beacon Rock, mud bot-

tom, 10-12 m, MNZ CR-9850 (1). E side of Port Levy, mud bottom,

MNZ CR-9855 (1).

Diagnosis. Head with anterior margin vaguely tuberculate,

dorsally with tubercles, pereon finely setose and vaguely tuber-

culate; tergites slightly laterally produced. Pleotelson with

blunt anterolateral processes, laterally tuberculate, dorsally

Gary C. B. Poore and Tania M. Bardsley

Figure 4. Pseudidothea richardsoni. Male, CMNZ. Pleopods 1 and 2;

penial process.

smooth, tapering to a blunt apex. Male antenna 1 flagellum

with about 7 clusters of aesthetascs. Antennae and pereopods

without tubercles. Pereopods finely setose. Antenna 2 peduncle

with long, fine setae on articles 3-5. Male pleopod 1 endopod

about as long as peduncle, with plumose setae apically; exopod

longer than endopod, with 15 spinules on lateral margin, taper-

ing distally, thickened and folded laterally to partially cover

groove that runs to apex. Male pleopod 2 appendix masculina

styliform, about as long as endopod. Uropodal exopod with

strong apical setae, endopod with 3 pappose setae; exopod

about two-thirds as long as endopod. Oostegite absent from

pereopod 5.

Descriptive notes. We examined a male specimen of

P. richardsoni and here figure and describe pleopods 1 and 2

and the penial process:

Male pleopod 1 peduncle twice as long as wide, with 8

coupling hooks mesially; endopod lamellar with 5 lateral spin-

ules proximally, tapering distally with 5 apical plumose setae;

exopod longer than endopod with 15 spinules on lateral margin,

tapering distally, thickened and folded laterally to partially

cover a groove that runs to the obtuse apex. Male pleopod 2

endopod with 13 apical plumose setae; exopod with 39 mar-

ginal plumose setae; appendix masculina styliform, about as

long as endopod. Penial process fused, distally bilobate.

Distribution. Cook Strait and Lyttelton Harbour, New Zealand,

7-146 m depth.

Remarks. Hurley (1957) observed that P. richardsoni is close to

“P. bonnieri Ohlin” (=P. miersii ) but he considered there are

sufficient differences to warrant a separate species; these

include uropodal endopod with three pappose setae, lack of

tubercles on the pereopods and antenna 2 peduncle with long

fine setae on articles 3-5.

The first male pleopods are also different from those of

P. miersii. Pseudidothea miersii has a male pleopod 1 peduncle

with 6-7 coupling hooks, endopod with plumose setae margin-

ally, exopod with 16 or 17 spinules laterally and with an acute

apex bent outwards, almost at right angles. In contrast,

P. richardsoni has a peduncle with eight coupling hooks, an

endopod with five lateral spinules proximally and five apical

plumose setae, and an exopod with 15 spinules on its lateral

margin, tapering distally to an obtuse apex.

Pseudidothea scutata (Stephensen)

Microarcturus scutatus Stephensen, 1947: 15-17, figs 5, 6.

Pseudidothea scutatus Sheppard, 1957: 176-180, figs 15, 16.

Pseudidothea scutatas . — Brandt and Wagele, 1990: 97-105, figs

1-3 (lapsus)

Material examined. Antarctica, Western Weddell Sea, A. Brandt on RV

Polarstem, Jan-Feb 2002 (ANDEEP stns): 61°09.82'S, 54°33.40'W,

302-306 m, NMV J47401 (1 male); 61°11.94’S, 54°37.37W, 302-306

m, NMV J47402 (1 male); 61°20.51'S, 55°28.66’W, 159-117 m, NMV

J47403 (1 female); 61°44.88'S, 58°1.54'W, 256-295 m, NMV J47404

(1 male); 59°52.21’S, 59°58.75'W), 3643-3622 m, NMV J47405 (1

specimen).

Diagnosis. Head smooth, pereonites with large, high, flat

tubercles: pereonite 1 with 1 pair of dorsal tubercles and 3 pairs

of shorter lateral processes; pereonites 2-4 with 1 pair of

dorsal and 1 pair of lateral tubercles and 1 or 2 pairs of shorter

dorsolateral processes; pereonites 5-7 with 1 pair each of

dorsal, dorsolateral and lateral tubercles. Pleotelson with sub-

acute anterolateral processes, 3 pairs of mediodorsal spines and

shorter and more irregular processes, apically acute and bent

dorsally. Male antenna 1 flagellum with about 25 clusters of

aesthetascs. Antenna 2 and pereopods 2 and 3 with tubercles.

Antenna 2 peduncle with long, fine setae on articles 3-5. Male

pleopod 1 endopod about as long as peduncle with plumose

setae laterally and apically; exopod longer than endopod, with

diagonal groove, apically tapering, bent outwards and terminat-

ing in an acute tooth, proximal half of lateral margin with short

setae, distal half with longer, plumose setae. Male pleopod 2

appendix masculina apically acute, about twice as long as rami

(Stephensen, 1947). Uropodal exopod about two-thirds as long

as endopod, each ramus with single seta (Sheppard, 1957).

Oostegite present on pereopod 5.

Distribution. South Shetland Islands, Antarctic Peninsula,

159-3622 m depth.

Remarks. Sheppard (1957) included Stephensen’s

Microarcturus scutatus in the synonymy of what she called

“ Pseudidothea scutatus sp. n.”. She admitted that Stephen-

sen’s “species appears to be identical with my specimens”

and that she received Stephensen’s paper after making

her own descriptions and figures. Her intention would appear to

Isopods of the family Pseudidotheidae

have been be to make a new combination rather than a new

species.

The species differs from all other species of Pseudidothea in

the presence of well developed oostegites on pereopods 5

(Sheppard’s 1947 observation confirmed in new material).

Although the pair of fifth oostegites meet in the middle, they

are flat and do not help in enclosing the eggs.

The dorsal and lateral pereonal tubercles of the holotype, a

20-mm long male, are separated by gaps smaller than the

tubercle diameters. In the same- sized female described by

Brandt and Wagele (1990) the tubercles are relatively smaller

and separated by gaps equal to their diameters. The uropodal

rami of the two specimens also differ: the endopod of the male

being shorter and narrower than the exopod while the two are

subequal in the female. In the absence of other material, we

assume these differences are sexual rather than specific. The

new material collected during the 2002 ANDEEP cruise is typ-

ical of this well-described species. The individual dredged from

3643-3622 m depth, much deeper than the usual depths of a

few hundred metres, could not be distinguished from the rest.

Acknowledgements

This contribution was made possible through a grant from the

Australian Biological Resources Study. We are grateful to

Graham Milledge who inked the figures. We tha nk Angelika

Brandt, Zoological Institute and Museum, Hamburg, for exam-

ining type material and for the donation of material from the

ANDEEP cruise. We also thank her and Oliver Coleman,

Museum fur Naturkunde, Berlin, for discussion about the type

status of material in their museums. We thank the Canterbury

Museum, New Zealand, and the Museum of New Zealand (Te

Papa Tongarewa) for the loan of material.

References

Barnard, K.H. 1920. Contributions to the crustacean fauna of South

Africa. No. 6. Further additions to the list of marine Isopoda.

Annals of the South African Museum 17: 319-438.

Brandt, A., and Wagele, J.W. 1990. Redescription of Pseudidothea

scutatas (Stephensen, 1947) (Isopoda, Valvifera) and adaptations to

a microphagous nutrition. Crustaceana 58: 97-105.

Dana, J.D. 1849. Conspectus crustaceorum quae in orbis terrarum cir-

cumnavigatione, Carolo Wilkes e classe Reipublicae, Foederate

Duce, lexit et descripsit (continued.). American Journal of Sciences

and Arts 8: 424-428.

Hale, H.M. 1946. Isopoda -Valvifera. British, Australian and New

Zealand Antarctic Research Expedition, 1929-1931. Reports-

Series B ( Zoology and Botany) 5: 161-212.

Hurley, D.E. 1957. Some Amphipoda, Isopoda and Tanaidacea from

Cook Strait. Zoology Publications from Victoria University College

21 : 1 - 20 .

Kussakin, O.G. 1967. Fauna of Isopoda and Tanaidacea in the coastal

zones of the Antarctic and Subantarctic waters. [Translation from

Russian by the Israel Program for Scientific Translations,

Jerusalem, 1968.]. Biological Reports of the Soviet Antarctic

Expedition (1955-1958) 3: 220-389.

Nierstrasz, H.F. 1941. Die Isopoden der Siboga-Expedition. IV.

Isopoda Genuina. III. Gnathiidea, Anthuridea, Valvifera, Asellota,

Phreatocoidea. Siboga Expeditie Monographie 19: 235-308.

Nordenstam, A. 1933. Marine Isopoda of the fa mi lies Serolidae,

Idotheidae, Pseudidotheidae, Arcturidae, Parasellidae and

Stenetriidae mainly from the South Atlantic. Further Zoological

Results of the Swedish Antarctic Expedition, 1901-1903 3: 1-284,

282 pis, errata.

Ohlin, A. 1901. Isopoda from Tierra del Fuego and Patagonia.

Wissenschaftliche Ergebnisse der Schwedischen Expedition in die

Magellanregion oder nach den Magellanslandem 1895-1897 2:

261-306, pis 220-225.

Poore, G.C.B. 2001. Isopoda Valvifera: diagnoses and relationships of

the families. Journal of Crustacean Biology 21: 213-238.

Poore, G.C.B. 2003. Revision of Holidoteidae, an endemic southern

African family of Cmstacea, and re-appraisal of taxa previously

included in its three genera (Isopoda: Valvifera). Journal of Natural

History 37: 1805-1846.

Poore, G.C.B., and Bardsley, T.M. 1992. Austrarcturellidae (Crustacea:

Isopoda: Valvifera), a new family from Australasia. Invertebrate

Taxonomy 6: 843-908.

Samouelle, G. 1819. The entomologists’ useful compendium; or an

introduction to the knowledge of British Insects, comprising the

best means of obtaining and preserving them, and a description of

the apparatus generally used; together with the genera of Linne,

and modern methods of arranging the Classes Crustacea,

Myriapoda, spiders, mites and insects, from their affinities and

structure, according to the views of Dr. Leach. Also an explanation

of the terms used in entomology; a calendar of the times of appear-

ance and usual situations of near 3,000 species of British Insects;

with instructions for collecting and fitting up objects for the micro-

scope. Thomas Boys: London. 496 pp, 412 pis.

Sheppard, E.M. 1957. Isopod Crustacea Part II. The sub-order

Valvifera. Families: Idoteidae, Pseudidotheidae and Xenarcturidae

fam. n. With a supplement to isopod Crustacea, Part 1 . The family

Serolidae. Discovery Reports 29: 141-197, pis 148, 149.

Stephensen, K. 1947. Tanaidacea, Isopoda, Amphipoda and

Pycnogonida. Scientific Results of the Norwegian Antarctic

Expeditions 1927-28 27 : 1-90.

Studer, T. 1882. fiber eine neue Art Arcturus und eine neue Gattung der

Idotheiden. Sitzungsberichte der Gesellschaft Naturforschender

Freunde zu Berlin 1882: 56-58.

Studer, T. 1884. Isopoden, gesammelt wahrend der Reise S.M.S.

Gazelle um die Erde 1874—76. Abhandlungen der Mathematisch-

Physikalischen Klasse der Koniglich Bayerischen Akademie der

Wissenschaften 1883: 1-28, pis 21, 22.

Wagele, J.-W. 1989. Evolution und phylogenetisches System der

Isopoda. Stand der Forschung und neue Erkenntnisse. Zoologica

(Stuttgart) 140: 1-262.

Wagele, J.-W. 1991. Antarctic Isopoda Valvifera. Koeltz Scientific

Books: Konigstein. 213 pp.

Memoirs of Museum Victoria 61(1): 85-110 (2004)

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

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

The long-horned caddisfly genus Oecetis (Trichoptera: Leptoceridae) in Australia:

two new species groups and 17 new species

Alice Wells

Australian Biological Resources Study, Department of the Environment and Heritage, Canberra, ACT 2600, Australia

(alice. wells @deh.gov.au)

Abstract Wells, A. 2004. The long-horned caddisfly genus Oecetis (Trichoptera: Leptoceridae) in Australia: two new species

groups and 17 new species. Memoirs of Museum Victoria 61(1): 85-110.

Among Australian caddisflies (Insecta: Trichoptera), the leptocerid genus Oecetis McLachlan is one of the most wide-

spread and diverse genera. This paper brings to 53 the number of described species recorded for Australia, and at least a

further 15 to 20 species remain to be described. Here 17 newly described species are assigned with 12 others to an infor-

mal laustra- group, defined by having the phallus simple and lacking parameres. Three other newly described species are

placed in another informal group, the longiterga-growp, based on the broad form of the forewing and strongly pronounced

venation. Distributions are plotted for most species, new records extending distributions considerably; of the 29 laustra-

group species included here, 15 are known in Australia only from the north of the continent. Keys are provided to the

informal species-groups recognised for Australia, and to males of the laustra- and longiterga- groups.

Keywords Taxonomy, Trichoptera, Leptoceridae, Oecetis, new species, Australia

Contents

Introduction 85

Key to males of Oecetis species groups in Australia 87

Key to males of the Oecetis laustra-growp in Australia 87

Key to males of the Oecetis longiterga- group in Australia 89

Species of the laustra-growp 89

Oecetis laustra Mosely, 1953 89

Oecetis pseudolaustra sp. nov. 89

Oecetis atarpa Mosely, 1953 90

Oecetis scirpicula Neboiss, 1977 91

Oecetis inscripta Kimmins, 1953 91

Oecetis brevidentata sp. nov. 91

Oecetis asmanista Mosely, 1953 92

Oecetis minasata Mosely, 1953 93

Oecetis erskinensis sp. nov. 94

Oecetis aeoloptera Kimmins, 1953 94

Oecetis multipunctata Ulmer, 1916 94

Oecetis cracenta sp. nov. 94

Oecetis parka Mosely, 1953 96

Oecetis arcada Mosely, 1953 97

Oecetis cymula Neboiss, 1982 97

Oecetis paracymula sp. nov. 97

Oecetis spicata sp. nov. 97

Oecetis crena sp. nov. 98

Oecetis quadrata sp. nov. 98

Oecetis dilata sp. nov. 98

Oecetis koobarra sp. nov. 99

Oecetis falcata sp. nov. 101

Oecetis terrania sp. nov. 101

Oecetis papposa sp. nov. 101

Oecetis curta sp. nov. 103

Oecetis aduncata sp. nov. 103

Oecetis ornata Kimmins, 1962 103

Oecetis cepaforma sp. nov. 104

Oecetis dostinei sp. nov. 105

Species of the longiterga-growp 105

Oecetis digitata sp. nov. 105

Oecetis ancala sp. nov. 106

Oecetis crosslandi sp. nov. 107

Acknowledgements 107

References 110

Introduction

Oecetis McLachlan, 1877 (Leptoceridae), a cosmopolitan

genus of long-homed caddisflies, is one of several genera of

Trichoptera represented Australia-wide, occurring even in the

arid inland. Light trap samples of caddisflies in Australia

seldom fail to include several species and often adults and

immatures of some species are abundant. Adults of most

Australian species are rather nondescript in appearance,

although some have distinctive wing markings, and many

members of one group have patches of scales on the forewing.

Alice Wells

On the basis of features of wing venation and male genitalia,

however, five distinct groups can be recognised among

Australian species of Oecetis. Two of these were dealt with

recently. Males of the reticulata- group (Neboiss, 1989), have

an amour-like, sculptured dorsal plate formed by the extension

of abdominal tergite VIII over the terminal segments of the

abdomen. In the complexa-group (Wells, 2000), males are char-

acterised by external spiny processes or parameres associated

with the phallus, and forewing fork 1 with a footstalk. Here,

two further groups are recognised, the laustra- and longiterga-

groups. A fifth group, the pechana- group, will be dealt with in

a subsequent paper.

The laustra-group is distinguished by having the phallus

very simple, lacking parameres or spines of any kind (Fig. 3).

This group, its name taken from a widespread and common

Australian species, comprises 29 species, 17 of them newly

described. Most of these have the forewing with a distinct foot-

stalk on fork 1, the feature upon which Chen (1992), in his

unpublished revision, based a subgroup in one of his subgenera.

Several species, however, have the footstalk very short

(indistinct), and several have fork 1 sessile.

The longiterga-growp, named for a New Guinea species,

share a wing form atypical in the Australian fauna (Figs 91, 94),

with pronounced veins and fork 1 sessile. Three new Australian

species are included: O. crosslandi sp. nov., O. ancala sp. nov.

and O. digitata sp. nov. Another unpublished New Guinean

species is known that shares this wing form (Chen, 1992: fig.

4.3).

Species of the laustra- group vary in tibial spur counts: spur

formulae may be 0, 2, 2 or 1, 2, 2. A count of 2, 2, 2 for O. scir-

picula Neboiss, 1977 was reported, but has not been verified in

material checked in this study. Variation is seen, too, in length

of wing setae, with some species having long downy setae

along veins while others have the vestiture uniformly short over

the wing laminae, although always longer on the distal margins.

Some wings are uniformly fuscous, yet others show distinctive

patterns, with darker brown to black markings produced by dif-

ferences in wing membrane colour, emphasised by the colour

of the setae. Wing shape varies. Forewings are generally slen-

der, having length to width ratios close to 4, but are broader in

some species. The position of transverse veins in the forewing,

especially the posterior anastomosis (Fig. 1, following ter-

minology of Ruiter, 2000, after Schmid, 1980), varies across

the group. The crossveins are denoted tl for that closing dis-

coidal cell, t2 for r-m, and t3 for the vein closing the thyridial

cell. In some species, tl, t2, and t3 are more or less contiguous,

or linear — as in O. scirpicula, in others no two transverse

veins are aligned — as in O. multipunctata Ulmer, 1916 and

O. parka Mosely, 1953. Variation within some species is appar-

ent. Thus, in O. laustra the posterior anastomosis is linear

in some populations, while the crossveins are slightly out of

alignment in others. None of the laustra- group species has

wing-scales such as occur in most of the pechana- group.

Assignment of males to the five Oecetis species groups

using the key provided here is reasonably straightforward.

Using the key to O. laustra- group species, however, may be

difficult. Where possible, wing features are used for differen-

tiation and several species are readily identified by their

characteristic wing markings. For others, the often more cryp-

tic genitalic features must be used. It is recommended that,

when identifications are being made, close attention be given to

the illustrations provided and to notes on variation within

species.

Extensive collections in Museum Victoria (NMV),

Australian National Insect Collection (ANIC) and Northern

Territory Museum and Art Galleries (NTM) were available for

this study; other depositories of types or new material are The

Natural History Museum, London (BMNH), Queensland

Museum (QM) and Waite Agricultural Research Institute

(WARI). For all established species, only a diagnosis, illustra-

tions and new distribution data are given. For the more com-

monly collected species, only distribution maps are given —

the detailed locality data are not included here, or are included

for type material only; these data are available from the author

or from NMV. Members of the laustra- group occur in all

Australian states and territories, and tend to be more or less

peripherally distributed on the mainland; several species are

found in Tasmania and one is described from Lord Howe

Island. Some species appear to be localised, and others to have

disjunct distributions (see Table 1); similar patterns were

reported for complexa- group species (Wells, 2000). Fifteen

laustra- group species are known only from northern Australia,

and the richest diversity — 16 species — is recorded for

Queensland (see Table 1), with most records being from along

the eastern seaboard; four of these species have been collected

from northern Queensland only, and one is known from

Queensland and New Guinea. Few species in this group are

recorded for south-western Western Australia or South

Australia. Two species assigned to the longiterga-gxoup are

recorded only from northern Australia; the third is widespread

— collected from northern Western Australia, the north of

the NT, south-eastern Qld, eastern NSW and south-central

Victoria.

As for species of the complexa- group, few females can be

associated with males with certainty. Thus, new species are

diagnosed on the basis of males only. Several observations on

biology are pertinent, some deduced from label data. For

example, locality data for O. laustra include a large number of

lacustrine sites. Larvae of this species build their case from

portions cut from stems of aquatic macrophytes and live

amongst and on these plants (Wells, 1991; St Clair, 1994); the

cases are very light and would readily be swept free of the sub-

stratum by currents. In contrast, it appears from label data that

O. arcada Mosely, 1953, O. asmanista Mosely, 1953 and

O. cymula Neboiss, 1982 are probably adapted to swiftly flow-

ing waters. I have found the sand grain-cased larvae of

O. digitata sp. nov. in sand in the direct flow of water under

small rocky falls, and those of O. koobarra sp. nov. amongst

sand on ledges in the bedrock of small permanent creeks that

have moderate flow, but are subject to occasional spates.

Larvae and pupae of O. erskinensis sp. nov., from Lord Howe

Island, were collected from crevices, ridges and ledges in the

main flow of a small steam. Like many other species,

O. brevidentata sp. nov. constructs sand-grain cases and is

found associated with boulders and cobbles in sand- or cobble-

based creeks or rivers.

Australian caddis flies of the genus Oecetis

Table 1 . Oecetis laustra-g roup species clustered according to state/territory of occurrence with the exception that a distinction is made between

the north-west and south-west of Western Australia, and Lord Howe Island is included (LHI). Arrangement designed to emphasis commonalities

between parts of Australia. (Abbreviations: Qld, Queensland; NSW, New South Wales; Vic., Victoria; Tas., Tasmania; SA, South Australia;

SWWA south-western Western Australia; NWWA, north-western Western Australia; NT, Northern Territory; LHI, Lord Howe Island).

SPECIES

Qld

NSW

Vic.

Tas.

LHI

Oecetis erskinensis sp. nov.

Oecetis scirpicula Neboiss, 1977

Oecetis crena sp. nov.

Oecetis arcada Mosely, 1953

Oecetis asmanista Mosely, 1953

Oecetis paracymula sp. nov.

Oecetis terania sp. nov.

Oecetis minasata Mosely, 1953

Oecetis parka Mosely, 1953

Oecetis aduncata sp. nov.

Oecetis curta sp. nov.

Oecetis inscripta Kimmins, 1953

Oecetis laustra Mosely, 1953

Oecetis atarpa Mosely, 1953

Oecetis aeoloptera Kimmins, 1953

Oecetis dostinei sp. nov.

Oecetis dilata sp. nov.

Oecetis spicata sp. nov.

Oecetis cepaforma sp. nov.

Oecetis papposa sp. nov.

Oecetis cracenta sp. nov.

Oecetis brevidentata sp. nov.

Oecetis koobarra sp. nov.

Oecetis pseudolaustra sp. nov.

Oecetis falcata sp. nov.

Oecetis quadrata sp. nov.

Oecetis multipunctata Ulmer, 1916

Oecetis omata Kimmins, 1962

Oecetis cymula Neboiss, 1982

Total species

Generally, larval morphology of Oecetis species is remark-

ably uniform, a major difference being colour patterns on the

head and pro- and mesonota, and occasional differences are

apparent in ventral head and thoracic sternal sclerites, and in

setation (Floyd, 1995; St Clair, 2000). However, mandibles of

at least one laustra - group species, O. brevidentata sp. nov., are

short and stout (Wells, 1991), suggesting a different diet from

that of most species. The more characteristic mandible form is

slender and sharply pointed.

Key to males of Oecetis species groups in Australia

1. Abdominal tergite VIII sculptured, expanded and

extended distally, forming shield over terminal abdominal

segments and genitalia

reticulata - group (see Neboiss, 1989)

— Abdominal tergite VIII unmodified 2

2. Wings with veins strongly pronounced, fork 1 without a

footstalk (sessile) (Figs 91, 94) longiterga - group

— Wings with veins normal, not particularly well pro-

nounced, fork 1 with or without footstalk (Figs 1, 30, 53)

3. Phallus simple, lacking parameres (Figs 3, 38, 72)

laustra-growp (see below)

— Phallus with one or more internal or external parameres or

spines 4

4. Forewing fork 1 sessile, wing lamina often bearing

patches of scales (androconia); phallus usually with a

single internal paramere or spine pechana- group

— Forewing fork 1 stalked, wing never bearing patches of

scales; phallus with one or more external parameres ....

complexa- group (see Wells, 2001)

Key to males of the Oecetis laustra- group in Australia

1. Forewing with distinct footstalk on fork 1 (Fig. 1) .... 2

— Forewing with fork 1 sessile (Figs 80, 87, 88), or very

nearly so 25

Alice Wells

2. In hind wing, M unbranched (Fig. 73); in male genitalia in

lateral view, inferior appendages about same width basally

and apically, constricted mesially (Fig. 72) . . .O. papposa

— In hind wing, M branched; male genitalia not as above .3

3. In hind wing, Cul and Cu2 arise at cross vein M-Cu

(Fig. 53) O. quadrata

— In hind wing, Cul and Cu2 arise distal to crossvein M-Cu

4. In forewing, at least 2 crossveins contiguous (linear), or

nearly so, in posterior anastomosis (Figs 10, 12) 5

— In forewing, crossveins of posterior anastomosis all

clearly stepped (Figs 29, 30) 13

5. Forewing with distinctive dark lines marking parts of

veins, including diagonal mark linking tl, t2, t3 (Fig. 12)

O. inscripta

— Forewing not as above 6

6. In genitalia, inferior appendages in lateral view swollen

basally, distally reduced to at least one-third basal width

and strongly curved dorsad 7

— In genitalia, inferior appendages in lateral view not as

above (Figs 18, 39) 10

7 In genitalia, in ventral view, inferior appendages tapered

gradually towards apex, mesial margin slightly dentate or

with a single sharp mesial hook (Fig. 2) 8

— In genitalia, in ventral view, inferior appendages sharply

excavated at about one-third length, distally slender,

mesial margin smooth (Figs 8, 26) 9

8. In genitalia, in ventral view, inferior appendages with a

small mesial tooth (Fig. 2) O. laustra

— In genitalia, in ventral view, inferior appendages with a

sharp anteriorly directed mesial hook (Fig. 4)

O. pseudolaustra

9 Forewing with dark membrane marking posterior ana-

stomosis; genitalia in lateral view with peg-like process

ventral to preanal appendages (Fig. 9) O. atarpa

— Forewing without dark markings at posterior anastomosis

(Fig. 10); without peg-like process ventral to preanal

appendages (Fig. 11) O. scirpicula

10. In forewing t3 well proximal to tl and t2, with M3+4

arising distal to t3; wing lamina patterned with dark mark-

ing on crossveins tl, t2 and M forming a short dark distal

cross (Fig. 15) O. brevidentata

— In fore wing t3 only slightly proximal to tl and t2, with

M3 +4 arising at t3; wing lamina mottled, without distinct

marking 11

11. In genitalia in ventral view, inferior appendages broad in

basal two-thirds, mesial margin rounded then abruptly con-

stricted, curved inwards towards apices, not sharply angled

(Fig. 19); tergite X slim, digitiform O. asmanista

— In genitalia in ventral view, inferior appendages broad-

based with meso-ventral angle slightly lobed, mesial

margin strongly constricted to form a narrow lobe latero-

distally, with apex sharply angled inwards (Figs 21, 24);

tergite X a broad membrane 12

12. In genitalia in ventral view, phallus with apex sharply

delineated and deeply V-shaped (Fig. 24) . .O. erskinensis

— In genitalia in ventral view, phallus with apex rounded,

very shallowly cleft (Fig. 21) O. minasata

13. In forewing, tl more distal than t2 or t3 (Figs 64, 73) .14

— In forewing, t2 more distal than tl or t3 (Fig. 30) ... .17

14. Male genitalia with prominent, unequal sclerotised spines

dorsally (Figs 65-67) O. falcata

— Male genitalia without any sclerotised spines (Figs 60, 62)

15. Fore wing fork 1 and its footstalk about equal length; on

forewing a zig-zag pattern marks posterior anastomosis

(Fig. 63) O. koobarra

— Fore wing fork 1 longer than its footstalk; fore wing

posterior anastomosis not marked by zig-zag pattern . 16

16. Male genitalia, in ventral view, with inferior append-

ages stout throughout length, about 1.5 times width,

rounded mesially, apically a small curved lateral process

(Fig. 57); tergite X broad, apically truncate (Figs 58, 59) .

O. dilata

— Male genitalia, in ventral view, with inferior appendages

stout basally, gradually constricted to about half basal

width; tergite X excavated mesially to form a pair of

apically acute lateral processes (Figs 47, 48)

O. spicata (N Qld form)

17. Male genitalia, in lateral view, with a small digitate lobe

dorsal to base of inferior appendages (Fig. 38) . O. parka

— Male genitalia, in lateral view, without a small lobe as

above 18

18. Male genitalia, in ventral view, with a lobe on mesial side

of inferior appendages (Fig. 69) O. terania

— Male genitalia, in ventral view, without a lobe as above .19

19. Wings uniformly fuscous, vestiture short, dense; in male

genitalia, tergite X comprising a median, tapered lobe

between a pair of broadly rounded, membranous lateral

lobes (Figs 42, 45) 20

— Wings mottled or with some small markings, or spotted;

tergite X not as above 21

20. Male genitalia, in lateral view, with inferior appendages

about same width throughout length, apices not down-

turned (Fig. 43) O. cymula

— In genitalia, in lateral view, inferior appendages dilated

baso-ventrally, apices down-turned, rounded (Fig. 46) . . .

O. paracymula

21. In genitalia, in ventral view, inferior appendages broad-

based, abruptly narrowed, expanded slightly mesially,

distally about width of narrower basal section (Fig. 50); in

lateral view, inferior appendages with a broadly rounded

basodorsal lobe, distally tapered, slender (Fig. 52)

O. crena

— In genitalia, in ventral view, inferior appendages not as

above 22

22 In genitalia, in lateral view, inferior appendages truncate

apically, in distal half about half width at base (Fig. 40); in

ventral view, a distinct notch on mesial margin at about

two-thirds length (Fig. 39) O. arcada

— Not as above 23

23. In genitalia, inferior appendages in lateral view expanded

dorsally, narrow in distal two-thirds (Fig. 36); in ventral

view gently curved inwards and tapered to acuminate

apices (Fig. 34) O. cracenta

— Not as above 24

Australian caddis flies of the genus Oecetis

24. In genitalia, in ventral view, inferior appendages stout

basally gradually curved and constricted to narrowly

rounded apex; preanal appendages rounded (Figs 31-33) .

O. multipunctata

— In genitalia, in ventral view, inferior appendages stout at

base, abruptly narrowed to form slender disto-lateral lobes;

preanal appendages in dorsal view triangular, in lateral

view slender, tapered (Figs 26-28) O. aeoloptera

25. Forewing distinctly and coarsely patterned, dark brown

and fuscous (Fig. 80) O. ornata

— Forewing not as above, but may be spotted, moth-like 26

26. Male genitalia with, tergite X hairy, rounded, cleft

medially (Fig. 75) O. curta

— Tergite X not as above 27

27. In genitalia, tergite X membranous with apicolateral angles

acute (Fig. 48) O. spicata (NT form)

— In genitalia, tergite X not as above 28

28. Wings not spotted; in genitalia, in lateral view, preanal

appendages narrow, length 2x width (Figs 78, 79)

O. aduncata

— Wings spotted (Figs 87, 88); in genitalia, in lateral view,

preanal appendages broadly rounded or subtriangular (Figs

86,89) 29

29. In genitalia, tergite X in dorsal view minaret- shaped (Fig.

85); inferior appendages in ventral view with tiny hook at

mesial angle O. cepaforma

— Male genitalia with, tergite X in dorsal view broad,

apically truncate (Fig. 90); inferior appendages in

ventral view without tiny hook at mesial angle

O. dostinei

Key to males of Oecetis longiterga- group in Australia

1. Fore wing Cula straight (Fig. 94); in genitalia a single

paramere with apex spiked and toothed (Figs 98, 100) . .

O. crosslandi

— Forewing Cula with distinct curve (Fig. 91); male

genitalia not as above 2

2. Male genitalia with, in lateral view, a sclerotised process

dorsal to inferior appendages (Fig. 97); in ventral view,

forming a stout structure, angled mesially at midlength

(Fig. 95) O. ancala

— Male genitalia without any sclerotised processes (Figs 92,

93) O. digitata

Species of the laustra- group

Oecetis laustra Mosely

Figures 1-3, 101

Oecetis laustra Mosely in Mosely and Kimmins, 1953: 295, fig.

209. — Neboiss, 1977: 146, figs 788-791. — Neboiss, 1982: 321, figs

129-131.

Material examined. Holotype. Male, Yanchep, WA, 31°32.9'S

115°41.2'E (BMNH).

Other material. About 180 samples in ANIC, NTM, NMV and

WARI.

Diagnosis. Wings slightly mottled, fuscous-brown/grey, with

short hair on veins; forewing (Fig. 1) length 4 times maximum

width, footstalk of fork 1 slightly shorter than fork, posterior

anastomosis almost linear, to linear, very slightly oblique; hind-

wing (Fig. 1) with M branched; in male genitalia, inferior

appendages (Figs 2-3) broad and stout, in ventral view, a small

mesial tooth at about two-thirds length, in lateral view broadly

rounded above and below in basal half, distally about one-third

maximum basal width; preanal lobes short, rounded.

Distribution. Widespread in Australia, in lentic and slower

lotic waters, but not collected from arid central Australia (Fig.

101 ).

Remarks. The stouter inferior appendages distinguish O. laus-

tra from O. atarpa and O. aeoloptera, the absence of a sharp,

anteriorly directed mesial hook on the inferior appendages

distinguishes it from O. pseudolaustra sp. nov. Oecetis laustra

is one of the most widespread and abundant of Australian

Oecetis species, although unlike O. pechana it has not been

collected from the natural and artificial waterbodies of arid

inland Australia. In its present concept, the species is quite

variable across its range, and may eventually be demonstrated

to be several species. The posterior anastomosis of the forewing

is usually more or less linear, but may be slightly stepped; and

in the male genitalia, the form of the mesial margin of the

inferior appendages may be more or less dentate (Mosely and

Kimmins, 1953: fig. 209c, d; Neboiss, 1986: 269).

Oecetis pseudolaustra sp. nov.

Figures 4-6, 102

Material examined. Holotype, male, Qld, Lakefield NP, Sweetwater

Lagoon, 11 Oct 2002, G. Theischinger (ANIC).

Paratypes. 21 males, 1 female, data as for holotype (ANIC); 18

males, 3 females, Cape York Peninsula, Wenlock River crossing (to

Iron Range), 5 Oct 2002, G. Theischinger (ANIC, NMV, QM).

Diagnosis. As O. laustra except inferior appendages with a

sharp anteriorly directed mesial hook.

Description. Tibial spurs 1, 2, 2. Male forewing length 3.4-5.0

mm. Wings as for O. laustra, mottled fuscous/brown, densely

covered with short hair. Male genitalia as in Figs 4-6. Preanal

appendages in dorsal view rounded apically and about as long

as wide. Tergite X a single membranous plate, narrowly

truncate apically. Inferior appendages stout, in lateral view

broadly rounded above and below in basal half, distally about

one-third maximum basal width, in ventral view with a sharp

mesial hook on inner margin. Phallus length in lateral view

equal to width.

Distribution. Far north of Cape York Peninsula, northern Qld

(Fig. 102).

Remarks. At the Lakefield National Park site, this species was

collected together with several specimens of Oecetis laustra,

which could be distinguished by their slightly darker colour as

well as the distinctive difference in male genitalia. The

stouter inferior appendages distinguish O. pseudolaustra and

O. laustra from O. atarpa and O. aeoloptera.

Alice Wells

Figures 1-3, Oecetis laustra Mosely, 1953: 1, wings, labelled to indicate forks present and terminology applied to crossveins of posterior anas-

tomosis; 2-3, male genitalia, ventral and lateral views [after Mosely and Kimmins, 1953]. inf app, inferior appendages; ph, phallus; pre app, pre-

anal appendage; IX, X, abdominal segments IX and X; 1,5, forks in wing venation.

Figures 4-6, O. pseudolaustra sp. nov., male genitalia, ventral, dorsal and lateral views. X, abdominal segment X.

Figures 7-9, O. atarpa Mosely: 7, forewing; 8, 9, male genitalia, ventral and lateral views [from Mosely and Kimmins, 1953].

Figure 10, O. scirpicula Neboiss, forewing [after Neboiss, 1977]. ft st, foot stalk of fork 1.

Oecetis atarpa Mosely

Figures 7-9, 103

Oecetis atarpa Mosely and Kimmins, 1953: 287, fig. 202.

Material examined. Paratype. Male, National Park [Royal National

Park], NSW, 34°3.9'S 151°3.1’E (BMNH).

Other material. 27 samples in ANIC and NMV.

Diagnosis. Forewing (Fig. 7) length 3—4 times maximum

width, fork 1 with footstalk, posterior anastomosis with t2 and

t3 contiguous, tl more proximal than both t2 and t3; in the male

genitalia (Figs 8, 9) abdominal segment IX in lateral view pro-

duced distally in a down-turned peg ventral to preanal lobe; and

inferior appendages in lateral view long and slender distally,

rounded and swollen basally with basal third about 4 times

broader than distal two-thirds which is less than twice length of

basal section.

Distribution. Throughout Australia but only small scattered

collections taken in NT (Fig. 103).

Remarks. Closely resembling Oecetis scirpicula and

O. aeoloptera in form of male genitalia but distinguished by the

Australian caddis flies of the genus Oecetis

stepped arrangement of crossveins in the posterior anastomosis

in fore wing, and the dorsolateral ‘peg’ on abdominal seg-

ment IX. The very slight difference between O. atarpa and

O. scirpicula Neboiss — mainly the more strictly linear poster-

ior anastomosis in the forewing of the latter and in the male

genitalia a small difference in proportions in inferior

appendages — is well within the range of variation exhibited by

other Australian Oecetis, particularly those with temperate to

tropical distributions. For the present, however, the two names

are retained.

Oecetis scirpicula Neboiss

Figures 10, 11, 104

Oecetis scirpicula Neboiss, 1977: 149, figs 804-807.

Material examined. Tas.: 2 males, Canal at Interlaken, 42°8.8'S

147°10.5'E, 2 Feb 1966, G.E. Edmunds (ANIC); males, females,

Navarre River, 42°9.5'S 146°8.6’E, 12 Feb 1967, E.F. Riek (ANIC); 1

male, Olga-Hardwood River, Saddle buttongrass plain, 42°57.8'S

145°55.3'E, 4 Apr 1977, Allbrook, Richardson, Swain (NMV); 2

males, 12 km NNE Bronte Park, 42°02’S 146°33’E, 2 Feb 1983, J.C.

Cardale (ANIC); male, 2 females, Pelion Hut, 3 km S Mt Oakleigh,

41°50'S 146°03'E, Feb. 1990, E.S. Nielsen, It trap (ANIC).

Diagnosis. Forewing (Fig. 10) length about 4 times maximum

width, veins of the posterior anastomosis contiguous; in male

genitalia (Fig. 11), abdominal segment IX produced poster-

iorly to form a triangular process ventral to preanal appendages

and inferior appendages with distal slender distal portion twice

length of basal portion.

Distribution. Tas. (Fig. 104).

Remarks. Although males of this species closely resemble those

of O. atarpa, as noted above, the name scirpicula is retained

here for these Tasmanian specimens. The two species have only

slight difference in forewing venation and in the genitalia —

proportions of the inferior appendages, differences that are well

within the range of variability accepted for other species. Size

differences, however, are considerable, the anterior wing length

of around 9 mm for O. scirpicula being several millimetres

longer than in O. atarpa. In addition, females of O. scirpicula

lack the broadly rounded dorsal lobes seen in O. atarpa

(Mosely and Kimmins, 1953: fig. 203c), having in their place a

pair of almond-shaped lobes.

Oecetis inscripta Kimmins

Figures 12-14, 105.

Oecetis inscripta Kimmins in Mosely and Kim mins, 1953: 294, fig.

208. —Neboiss, 1977: 148, figs 801-803. —Neboiss, 1982: 323, figs

127, 128.

Material examined. Holotype. Male, Bathurst, NSW (BMNH).

52 samples in ANIC, NMV and WARI.

Diagnosis. Forewing (Fig. 12) length almost 5 times maximum

width, lamina marked with bold dark streaks, not the more

usual dark spots or patches; footstalk on fork 1, and tl and

t2 contiguous, t3 more proximal than the 2 other crossveins

of the posterior anastomosis. In the male genitalia (Figs 13,

14), a rounded basodorsal lobe on the inferior append-

ages and infer-ior appendages in ventral view slender with

length 3 times maximum width, irregularly tapered towards

apex.

Distribution. Eastern Australia from Tas. northwards almost to

Cairns, north-eastern Qld, probably widespread in

Murray-Darling basin (Fig. 105).

Oecetis brevidentata sp. nov.

Figures 15-18, 106

Oecetis sp. E. — Wells, 1991: figs 138, 172-176.

Material examined. Holotype, male, NT, Katherine River Gorge

National Park, 13°23.9'S 133°10.0E, 13 Aug 1979, J. Blyth (NMV T-

18524).

Paratypes. NT: 4 males, 3 females, same data as for holotype

(NMV); 1 male, Kakadu National Park, Jim Jim Creek, 3 km below

falls, 13°15.9'S 132°51.1’E, 1 Sep 1979, J. Blyth (NMV); 1 male,

ARRS, South Alligator River, at Gimbat OSS Station, 13°34.3'S

132°36.7'E, 24 May 1988, A. Wells and P. Suter (NMV); 9 males, 1

female, South Alligator River below BHP camp, 25 May 1988, P. Suter

and A. Wells (NMV); 2 males, ARRS, SAR site 1, 14 Jun 1988, P

Dostine (NTM); 1 male, 1 female, ARRS, South Alligator River, at

Gimbat OSS Station, 13°34.3’S 132°36.7'E, 28 Apr 1989, P. Dostine

(NTM); 2 males, 1 female, 12°48'S 132°49'E, Kakadu National Park,

Baroalba Springs, 4 Oct 1991, A. Wells (NTM).

Other material. NT: male, 3 females, ARRS South Alligator River

above Fisher Creek jcn, 13°34'S 132°34'E, 18-20 Apr 1989, Suter and

Wells (NTM); 2 males, female, Gunlom (as UDP Falls), 13°24.9'S

132°26.0'E, 18-19 Jul 1980, M.B. Malipatil (NTM).

Diagnosis. Fore wing with length about 5 times maximum

width, lamina marked by a distinctive cross formed by dark

membrane and setae on tl and t2 and their junction with R3-4;

tl and t2 contiguous, t3 well proximal of both. In male g

enitalia, inferior appendages in ventral view slender and

curving, and in lateral view with apices acute and directed

ventrally, rather than straight as in O. multipunctata.

Description. Tibial spurs 1, 2, 2. Male forewing length 6. 1-7.8

mm. Wings with vestiture of even length, without spots, but

forewing (Fig. 15) with distinctive dark cross marking the

junctions of tl and t2 with R, footstalk on fork 1 longer than

fork. Male genitalia as in Figs 16-18. Segment IX widest mid-

laterally; preanal appendages separate, ovoid. Segment X elon-

gate, membranous, dorsal process short, slender. Inferior

appendages narrow, tapered, slightly bowed, apices down-

turned in lateral view. Phallus with length about twice width,

beak-like apically.

Distribution. Alligator Rivers region of northern NT (Fig. 106).

Remarks. The larva of this species is unique among Australian

Oecetis species. Associated as ‘sp. E’ by Wells (1991), it is the

only Oecetis species of which I am aware that has short, blunt

mandibles. This feature suggests that O. brevidentata has

dietary habits quite different from other congeners. It may well

be a detritivore.

Etymology. Latin, descriptive of the larval mandibles.

Alice Wells

Figure 11, Oecetis scirpicula Neboiss, male genitalia, lateral view [from Neboiss, 1977].

Figures 12-14, O. inscripta Kimmins: 12, forewing; 13, 14, male inferior appendages in ventral view, genitalia in lateral view [from Mosely and

Kimmins, 1953].

Figures 15-18, O. brevidentata sp. nov.: 15, forewing; 16-18, male genitalia, ventral, dorsal and lateral views.

Figure 19, 20, O. asmanista Mosely, male genitalia, ventral and lateral views [from Mosely and Kimmins, 1953].

Oecetis asmanista Mosely

Figures 19, 20, 107

Oecetis asmanista Mosely and Kimmins, 1953: 282, fig. 199. —

Neboiss, 1977: 147, figs 792-795.

Oecetis ochracea Jaquemart, 1965: 23. [Junior homonym of Oecetis

ochracea (Curtis, 1825)], (synonymy by Neboiss, 1977: 147).

Oecetis geevestonia Neboiss, 1974: 15. [Unnecessary replacement

name for O. ochracea Jaquemart.]

Material examined. Holotype. male, Tas. (BMNH).

Tas.: Male, female, Derwent Bridge, 12 Feb 1967, E.F. Riek

(ANIC); male holotype (BMNH); 1 male, Olga Camp, Gordon-Smith

River junction, 42°39.1'S 145°48.0'E, 2 Feb 1976, Howard, Suter

(NMV); males, females, West Bay River, Margate, 43°1.7'S

Australian caddis flies of the genus Oecetis

Figures 21-23, Oecetis minasata Mosely, male genitalia, ventral, dorsal and lateral views [from Mosely and Kimmins, 1953].

Figures 24-25, O. erskinensis sp. nov., male genitalia, ventral, dorsal and lateral views.

Figures 26-29, O. aeoloptera Kimmins: 26-28, male genitalia, ventral, dorsal and lateral views; 29, forewing [from Mosely and Ki mmins, 1953].

147°15.7'E, 6 Jan 1977, Coleman, Neboiss, Allbrook (NMV); males,

females, Franklin River-Roaring Creek junction, 4 km above Gordon

River junction, 8 Jan 1977, Coleman, Neboiss, Allbrook (NMV); 1

female, Swamp nr Olga River, 19 km above Gordon River Junction,

42°42.8'S 145°46.8'E, 13 Jan 1977, Neboiss and Swain (NMV); 3

males, Pelion Hut, 3 km S Mt Oakleigh, 41°50'S 146°03’E, 6-11 Mar

1991 at light, M. Horak, R McQuillan (ANIC). Vic: male, Tanjil River

Jen, 10 km N of Willow Grove, 38°4.2'S 146°10.6'E, 18 Dec 1973,

A. Neboiss (NMV).

Diagnosis. Forewing length over 4 times maximum width, fork

1 with footstalk and tl and t2 contiguous, t3 only slightly more

proximal than tl or t2. In male genitalia (Figs 19, 20), inferior

appendages skittle-shaped in lateral view being roundly

constricted at two-thirds length, narrow distally; tergum X a

narrow process, almost twice as long as preanal appendages.

Distribution. Tas. and south-central Vic. (one specimen only)

(Fig. 107).

Oecetis minasata Mosely

Figures 21-23, 108

Oecetis minasata Mosely in Mosely and Kimmins, 1953: 282, fig.

198. —Neboiss, 1977: 146, figs 784-787.

Material examined. Holotype. Male, Tas. (BMNH).

21 samples in ANIC and NMV.

Alice Wells

Diagnosis. Wings dark brownish, unicolorous, with long

downy hair on the veins; forewing length 4 times maximum

width, posterior anastomosis contiguous. In male genitalia

(Figs 21-23) in ventral view, inferior appendages with basal

portion as long as wide, roundly produced ventromesially, and

constricted and sharply returned forming dorsolateral lobes;

preanal appendages about 3 times as long as wide.

Distribution. South-eastern Australia: from south-eastern NSW,

ACT, Vic. and Tas. (Fig. 108).

Remarks. Males of O. minasata are distinguished from those of

O. erskinensis by the rounded apex of the phallus, and from

O. asmanista by the angular ventral expansion of the inferior

appendages and the shorter median process of tergum X.

The female lacks the patches of bristles characteristic of

O. asmanista. In form of male genitalia, O. cymula and

O. paracymula also show resemblance, but O. minasata can

be distinguished from the two species by the crossveins of the

posterior anastomosis contiguous, not stepped.

Oecetis erskinensis sp. nov.

Figures 24, 25

Material examined. Holotype, male, Lord Howe Island, Erskine Valley,

Erskine Creek, 2 1-22 Nov 1 996, Wells and Mound (ANIC).

Paratype. Male, data as for holotype (ANIC).

Other material. Larvae, pupa, locality as for holotype, 24 Dec 2001,

Wells and Mound (ANIC).

Diagnosis. As for O. minasata , but in male genitalia (Figs 24,

25) setae on intumed apices of inferior appendages are all

about equal in size and length, and in ventral view the apex of

phallus is divided to form 2 apically acute, sclerotised lobes.

Description. Spurs 1, 2, 2. Male forewing length, 7.9 mm.

Wings of typical shape, forewing fork 1 with footstalk less than

half length of fork, posterior anastomosis with tl and t2 linear,

oblique, t3 slightly more proximal, angled. Male genitalia, Figs

22, 25. Segment IX narrow, preanal appendages more than

twice as long as wide, rounded apically. Segment X slender,

almost same width throughout length. Inferior appendages in

ventral view clasper-shaped, inturned subapically, apices

obliquely truncate. Phallus very short, with distal sharply

downtumed section almost twice length of basal portion,

rounded basally, bifurcate, lobes sclerotised, acute apically.

Remarks. Oecetis minasata and O. erskinensis are closely

similar, possibly sister species, distinguished by genitalic

character states as noted under O. minasata.

Etymology. Named for the collecting site.

Oecetis aeoloptera Kimmins

Figures 26-29, 109

Oecetis aeoloptera Kimmins in Mosely and Kimmins, 1953: 287,

fig. 201.

Material examined. Holotype. Male, Murwillumbah, NSW (BMNH).

21 samples in ANIC and NMV.

Diagnosis. Forewing (Fig. 29) length 4 times maximum width,

posterior anastomosis clearly non-linear, with t2 more distal

than tl and t3 which are both approximately equal; in male

genitalia (Figs 26-28) inferior appendages pincer-shaped in

ventral view, in lateral view long and slender distally; preanal

lobes discrete, triangular.

Distribution. Eastern Vic., more north-eastern NSW, south-

eastern and northern Qld, northern NT to northern WA (Fig.

109).

Remarks. In male genitalic features O. aoeloptera closely

resembles O. atarpa and O. scirpicula but is distinguished by

the more attenuate inferior appendages, and preanal lobes elon-

gate triangular in ventral view, in lateral view, slender, tapered,

rather than rounded.

Oecetis multipunctata Ulmer

Figures 30-33, 110

Oecetis multipunctata Ulmer, 1916: 19. — Mosely and Kimmins,

1953: 293, fig. 206.

Material examined. 48 samples in ANIC and NMV.

Diagnosis. Wings with pattern of dark spots (Fig. 30); fore wing

length about 4.5 times maximum width, fork 1 with footstalk

and veins of the posterior anastomosis clearly stepped. In

male genitalia (Figs 31-33) segment X simple, short and

slender; inferior appendages without any lobes or processes,

length about 3 times width, tapered distally; and phallus curved

downwards.

Distribution. Widespread in northern and eastern Qld, and also

collected from north-eastern NSW (Fig. 110).

Remarks. Oecetis cepaforma, O. dostinei and O. parka also

have spotted wings, but O. cepaforma and O. dostinei have

the forewings broad and ‘floppy’ compared to those of

O. multipunctata and O. parka. These last two species are dis-

tinguished by male genitalic features, particularly the presence

of the digitate laterobasal lobe on the inferior appendages of

O. parka, and the more rounded, rather than angular shape of

the inferior appendages of O. multipunctata. The male genitalia

of Oecetis multipunctata closely resemble those of the NT

O. brevidentata from which O. multipunctata is distinguished

by its spotted wings and shape of the phallus. Also in O. multi-

punctata., in lateral view, the inferior appendages are almost

straight, whereas in O. brevidentata sp. nov. the apices are

down-turned.

Oecetis cracenta sp. nov.

Figures 34-36, 111

Material examined. Holotype, male, WA, King Edward River, 4—5

May 1992, 14°52.57'S 126°12.08’E, P.S. Cranston (ANIC).

Paratypes. NT: 2 males, 3 females, Litchfield National Park,

Florence Falls, 13°03'S 130°47'E, 9 Apr 1991, Wells and Horak

(NMV).

Diagnosis. Wings mottled, not noticeably spotty; forewing

length about 4 times maximum width, fork 1 with footstalk, and

Australian caddis flies of the genus Oecetis

Figures 30-33, Oecetis multipunctata Ulmer: 30, forewing; 31-33, male genitalia, ventral, dorsal and lateral views [from Mosely and Kimmins,

1953].

Figures 34-36, O. cracenta sp. nov., male genitalia, ventral, dorsal and lateral views.

Figures 37, 38, O. parka Mosely, male genitalia, dorsal and lateral views [from Mosely and Kimmins, 1953].

crossveins of posterior anastomosis clearly stepped. In male

genitalia, in ventral view, the inferior appendages taper

smoothly from base to apex and lack a mesial process, and the

preanal appendages are rounded.

Description. Tibial spurs 0, 2, 2. Male forewing length 4. 9-5. 4

mm. Wings with vestiture of even length, without scales or

spots, but forewing with a distinctive dark line marking the

distal crossveins; forewing with footstalk on fork 1 and t2 more

distal than tl and t3. Male genitalia as in Figs 34-36. Segment

IX short; preanal appendages in lateral view narrow at base and

apically rounded, in ventral view stout, rounded. Inferior

appendages in lateral view slightly expanded basodorsally,

slender in distal two-thirds, in ventral view closely appressed

basomesially, tapered to narrow apices, curved mesially,

straighter distally. Phallus strongly downturned, sharply

‘beaked’ apically.

Distribution. Kimberley region of northern WA, and northern

NT (Fig. 111).

Remarks. Seen in lateral view, the male genitalia of this species

resemble closely those of O. crena from which it is distin-

guished by the smoothly rounded inner margin of the inferior

appendages.

Etymology. Latin, descriptive of the male inferior appendages.

Alice Wells

Figures 39, 40, Oecetis arcada Mosely, male genitalia, ventral and lateral views [from Mosely and Kimmins, 1953].

Figures 41-42, O. cymula Neboiss, male genitalia, ventral, dorsal and lateral views [from Neboiss, 1982].

Figures 43-46, O. paracymula, sp. nov., male genitalia, ventral, dorsal and lateral views.

Figures 47-49, O. spicata sp. nov., male genitalia, ventral, dorsal and lateral views.

Oecetis parka Mosely

Figures 37, 38, 112

Oecetis parka Mosely in Mosely and Ki mmins, 1953: 290, fig. 205.

Material examined. Holotype. Male, NSW, National Park [?the Royal

National Park] (BMNH).

15 samples in ANIC and NMV.

Diagnosis. Wings spotted; forewing length about 4 times max-

imum width, footstalk present, veins of posterior anastomosis

clearly stepped with t2 more distal than tl and t3. In male gen-

italia (Figs 37, 38) an unusual dorsal digitiform process arises

at the base of the inferior appendages, which are about 4 times

as long as wide, and almost rod-shaped and straight in distal

three-quarters of length.

Australian caddis flies of the genus Oecetis

Distribution. Eastern Vic., east-central NSW and north-eastern

Qld (Fig. 112).

Remarks. The dorsal process at the base of the inferior

appendages is unique among Australian Oecetis, but closely

resembles similar structures seen in some Setodes species

described by Schmid (1987), for example S. apitayati and

S. uttamavarna. This species is never common in collections.

Oecetis arcada Mosely

Figures 39, 40, 113

Oecetis arcada Mosely, in Mosely and Kimmins, 1953: 290, fig.

204. — Neboiss, 1977: 148, figs 796-800.

Oecetis albodecorata Jaquemart, 1965 (Synonymy by Neboiss,

1977: 148)

Material examined. Holotype of O. arcada. Male, Cradle Mountains,

Tas. (BMNH).

1 8 samples in NMV and ANIC.

Diagnosis. Male genitalia (Figs 39, 40), with the inferior

appendages curving upwards and apically truncate; preanal

appendages almost 3 times longer than wide. Forewing length

less than 4 times maximum width; footstalk on fork 1 and t2

more distal than tl and t3.

Distribution. Widespread in Tas. (Fig. 113).

Oecetis cymula Neboiss

Figures 41^-3, 114

Oecetis cymula Neboiss, 1982: 323, figs 132-136.

Material examined. WA: 3 paratype males, same data as for holotype

(ANIC); paratypes, females, same locality as holotype, 22 Nov 1978,

A. Neboiss (ANIC); males, females, 15 miles NW of Walpole, 15 Nov

1958, E.F. Riek (ANIC); males, females, Harvey River nr Harvey

Falls, 15 km E of Harvey, 33°5.5'S 116°2.0'E, 21 Nov 1978,

A. Neboiss (ANIC).

Diagnosis. Forewing length almost 5 times maximum width,

footstalk on fork 1 about half as long as the fork, long downy

hairs along wing veins, and t2 more distal than tl and t3. In

male genitalia (Figs 41-43), ventral view, tergum X tapers to an

attenuate apex; length of median process of tergum X exceeds

length of preanal appendages and inferior appendages are well

separated mesially, thus lacking (when viewed laterally) an

expanded ventral projection near the base. Females have on

each side on segment VIII an area of bristle-like setae.

Distribution. South-western WA (Fig. 114)

Remarks. Oecetis cymula closely resembles O. paracymula but

has a tibial spur count of 1, 2, 2, whereas the latter has 0, 2, 2,

and O. cymula has fine setae apically and on the mesial margin

on the inferior appendages whereas O. paracymula has shorter,

stout almost peg-like setae.

Oecetis paracymula sp. nov.

Figures 44^-6, 115.

Material examined. Holotype, male, NSW, Upper Manning River, 20

km NNW Rawdon Vale, 31°34'S 151°34’E, 19 Feb 1980, A A. Calder

(NMV T-18511).

Paratypes. NSW, 2 males, data as for holotype (NMV); male,

Barrington Tops, 16 Nov 1953, A. Neboiss (NMV).

Other material. 4 females, data as for holotype (NMV).

Diagnosis. Adults densely hairy; forewing shape as for

O. cymula, footstalk on fork 1, and t2 more distal than tl or t3.

In male genitalia, in ventral view, median process of tergum X

constricted close do its base, then swollen, narrowest distally;

inferior appendages in ventral view loosely turned inwards in

distal third and in lateral view down-turned apically.

Description. Spurs 0, 2, 2. Male forewing length 8.7 mm.

Wings with long downy hair along veins; forewing narrow

proximally expanded more distally, rounded apically, footstalk

on fork 1 about one-fifth length of fork, discoidal and thyridial

cells about equal length, anterior anastomosis stepped with tl

and t2 almost level, t3 well anterior to other 2 veins; hind wing

slender, narrow. Male genitalia (Figs 44—46). Segment IX

broadly convex in lateral view, preanal appendages elongate,

slender, more robust in dorsal view, about two-thirds length of

median process of tergum X. Tergum X with the membranous

ventral plate forming a pair of widely almond-shaped lobes,

median process constricted towards base, slightly swollen

medially, narrower distally, in lateral view, about twice length

of anal appendages. Inferior appendages slender in postero-

lateral half, in lateral view form resembling O. minasata,

down-turned apically and with short stout setae on terminal

third. Phallus stout, strongly down-turned distally.

Distribution. North-eastern NSW (Fig. 115).

Etymology. Showing close resemblance to O. cymula.

Oecetis spicata sp. nov.

Figures 47-49, 116.

Material examined. Holotype, male, Gunshot Creek at Telegraph

Crossing, 11°42.9’S 142°20.0’E, 14-15 Feb 1992, D. Cartwright and

A. Wells (ANIC).

Paratypes, Qld: males, 5 females, Palmer River, 20 Jun 1971, E.F.

Riek (ANIC: WTH-1390); 21 males, Laura, Cape York Peninsula,

15°33.9'S 144 0 27.1'E, 7 Oct 1979, M.S. and B.J. Moulds (NMV); 1

male, Qld, Seary’s Creek, Rainbow Beach, 25°52'S 153°04'E, 6 Dec

984, G. Theischinger (NMV);1 male, Bertie Creek, 1 km SE

Heathlands HS, 11°49.9’S 142°29.0’E, 5 Feb 1992, D. Cartwright and

A. Wells (QM); 6 females, Tributary of Bertie Creek, 250 m SW

Heathlands HS, 11°45'S 142°35'E, 11 Feb 1992, D. Cartwright and A.

Wells (QM); 1 male, Cape York Peninsula, Heathlands, Bertie Creek,

11°49.9'S 142°29.0'E, 23 Mar 1993, M. Crossland (ANIC).

Other material. Qld: male, Mossman, 12 Jun 1971, E.F. Riek

(ANIC); males, females, Palmer River, 20 Jun 1971, E.F. Riek (ANIC);

male, Peach Creek Crossing, 25 km NNE of Coen, 4-5 Jul 1976, G.B.

and S.R. Monteith (ANIC); male, Cockatoo Creek Crossing, 17 km

NW Heathlands, 11°39'S 142°27'E, 25.vii-19 Aug 1992, P. Zborowski,

J.C. Cardale, open forest, Malaise #5 (ANIC); male, Cape York

Peninsula, Heathlands, Bertie Creek, 23 Mar 1993, M. Crossland

(ANIC). NT: 1 male, South Alligator River, UDP Falls [Gunlom],

13°24.9'S 132°26.0'E, 7 Sep 1974, J. Blyth (NTM); 3 males, South

Alligator River, UDP Falls [Gunlom], 13°24.9'S 132°26.0'E, 18-19 Jul

1980, M.B. Malipatil (NTM); 2 males, ARRS, South Alligator River,

at Gimbat OSS Stn, 13°34.3'S 132°36.7’E, 28 Apr 1988, P. Dostine

(NTM); male, ARRS, 5 km W South Alligator River OSS Stn, 19 Apr

Alice Wells

1988, Wells and Suter, 13°33'S 132°34'E (NMV); male, Litchfield NP,

Walker Creek, 18-19 Apr 1992, Wells (NTM). WA: 2 males, 2 females,

Bell Gorge, Melaleuca Hole, 17 a 01'S 125°14’E, I. Edwards (NMY).

Diagnosis. Male genitalia , in ventral view, with inferior

appendages broad-based, slightly produced apicomedially,

lateral lobes almost twice basal section, and tergite X a simple,

broad membranous plate with acute apicolateral angles.

Description. Spurs 1, 2, 2. Forewing length 4.5-5. 1 mm. Wings

of typical slender shape, neuration variable with forewing fork

1 with a short footstalk (N Qld forms) or sessile (NT form), tl

slightly more distal than t3 (N Qld) or both tl and t3 well

distal of t2; wing membrane dark along cross veins of poster-

ior anastomosis. Male genitalia, Figs 47-49. Segment IX

narrow, preanal appendages short, broad-based, rounded api-

cally. Segment X broad at base and apex, a wide apicomesal

concavity resulting in acute apicolateral processes. Inferior

appendages in ventral view broad-based, extending apico

laterally as stout short lobes, in lateral view more or less

straight but with a basi-ventral notch. Phallus strongly arched

in lateral view.

Distribution. South-eastern and northern Qld, northern NT and

northern WA (Fig. 116).

Remarks. The forewing venation of O. spicata varies from foot-

stalk short on forewing fork 1 (N Qld form) to having fork 1

sessile (NT form), and crossveins of the posterior anasto-mosis

varied, as indicated above. However, the males from NE

Queensland, N Northern Territory and NW Western Australia

show such close similarities in genitalic structures that, tenta-

tively, the differing position of the fork on the forewing is

considered simply to be variation within a local population.

Etymology. From Latin spica — point, for the apicolateral

processes on segment X.

Oecetis crena sp. nov.

Figures 50-52, 117

Material examined. Holotype, male, Vic., Gibbo River at Exhibition

Creek junction, 20 km N Benambra, 16 Jan 1982, A. Wells (NMV

T-18512),

Paratypes, Vic.: 1 male, 5 females, Mitta Mitta river, 8 km NE of

Benambra, 5 Feb 1974, A. Neboiss (NMV); 1 male, Wellington River,

23 km NNE of Licola on Tamboritha Road, 21 Feb 1978, NMV Survey

Department (NMV); 1 male, Yarra River, East Warburton, ii. 1980, I

Campbell (NMV); 1 male, 2 females, Nongungarra and Crooked River

junction, 4 Feb 1981, J. Blyth (NMV); male, Cobungra River, at

Anglers Rest, 15 January 1982, A. Wells (NMV).

Other material. Vic.: male, 4 females, Porepunkah River, 26 Jan

1960, A. Neboiss (NMV).

Diagnosis. Male genitalia inferior appendages clasper-shaped,

with a median expansion on the inner margin and fringes of

short setae dorso-basally; in the fore wing a footstalk on fork 1,

and t2 more distal than tl and t3.

Description. Spurs 1, 2, 2. Forewing length 7.1-8. 1mm. Wings

with long downy setae along veins; fore wing fork 1 about twice

length of footstalk, anterior anastomosis with tl and t2 almost

level, more distal than t3. Male genitalia. Figs 50-52. Segment

IX short, a triangular projection midlaterally on apical margin,

preanal appendages small, ovoid, separated. Segment X simple,

membranous. Inferior appendages elongate, slender, in ventral

view with a medial notch on inner margin; in lateral view a

knob dorsally at base, slender and tapered distally. Phallus

short, down-turned medially.

Distribution. Eastern Vic. (Fig. 118).

Etymology. Latin, crena — notch or rounded projection.

Oecetis quadrata sp. nov.

Figures 53-56, 118

Material examined. Holotype, male, North Qld, Zarda Creek nr Mt

Misery, W of Mossman, 1200 m, 23 Nov 1974, M.S. Moulds (NMV

T- 185 13).

Other material. Qld: male, Birthday Creek, 6 km NWW Paluma,

18°59'S 146°10'E, 25 Sep 1980, J.C. Cardale (ANIC).

Diagnosis. Cul and Cu2 in the hind wing arising at crossvein

m-cu, and preanal appendages exceedingly large, rectangular,

free.

Description. Tibial spurs 1, 2, 2. Male forewing length 7.6 mm.

Wings (Fig. 53) slender, forewing length about 4 times maxi-

mum width, footstalk on fork 1 about as long as fork, posterior

anastomosis stepped with t2 more distal than tl, t3 distad of tl

and t2; in hind wing Cul and Cu2 arise at crossvein m-cu.

Male genitalia, Figs 54-56. Segment IX broad ventrally and

laterally, narrow middorsally, preanal appendages large, sub-

quadrate. Segment X membranous, bifid. Inferior appendages

broad in basal half, strongly constricted medially, narrow

distally. Phallus short, apex strongly down-turned.

Distribution. North-eastern Qld (Fig. 118).

Remarks. When viewed laterally, the genitalic features of

O. quadrata males generally resemble those of O. inscripta.

The species has been collected from two disjunct localities

only.

Etymology. Latin, quadrata — four-cornered, for the unusual

shape of the preanal appendages.

Oecetis dilata sp. nov.

Figures 57-59, 119

Material examined. Holotype, male,WA, 9 km N of Kununurra, 19 Sep

1979, J. Blyth (NMV T-18514).

Paratypes.WA: 3 males, 2 female, same data as for holotype

(NMV); 2 males, Geikie Gorge, 18°06'S 125°42’E, 5 Oct 1996, I.

Edwards (NMV). Qld: 2 males, female, McLeod River, 15 km W of

Mt Carbine, 22-23 Jun 1975, S.R. Monteith (ANIC); 2 males, Cape

York Peninsula, Laura, 15°33.9'S 144°27.1’E, 7 Oct 1979, M.S. and

B.J. Moulds (NMV); 2 males, 2 females, 11°08'S 142°21'E, Jardine

River, 19 Oct 1992, P Zborowski and T. Weir (ANIC). NT: 2 males,

Daly River, 13°45'S 132°42'E, 9-10 Aug 1980, M.B. Malipatil (NMV).

Diagnosis. Forewing length about 4 times maximum width,

footstalk present on fork 1, tl more distal than t2 or t3. Males

genitalic features, inferior appendages, in ventral view have

width about 0.75 times length, and are rounded towards the

Australian caddis flies of the genus Oecetis

Figures 50-52, Oecetis crena sp. nov., male genitalia, ventral, dorsal and lateral views.

Figures 53-56, O. quadrata sp. nov.: 53, fore- and hind wings; 54-56, male genitalia, ventral, dorsal and lateral views.

Figures 57-59, O. dilata sp. nov., male genitalia, ventral, dorsal and lateral views.

apex, with a small apicolateral lobe, and in lateral view, have

length about 4 times width.

Description. Spurs 1, 2, 2. Male forewing length 4.8-5.4 mm.

Forewing with footstalk on fork 1 slightly shorter than fork;

posterior anastomosis stepped with tl more distal than t2, t3

oblique, slightly distad of t2. Male genitalia, Figs 57-59.

Segment IX uniformly narrow, preanal appendages ovoid,

widely separated. Segment X simple broad, apically truncate.

Inferior appendages stout, apically rounded with short, thin

lobe laterally; narrow and straight in lateral view, rounded api-

cally. Phallus stout, rounded, strongly curved downwards.

Distribution. Far northern Qld, NT and WA (Fig. 119).

Etymology. Latin, dilata — enlarge, for the unusually broad

inferior appendages.

Oecetis koobarra sp. nov.

Figures 60-63, 120

Oecetis sp. ‘H’ —Wells 1991: fig. 139.

Material examined. Holotype, male, NT, Kakadu National Park, Radon

Creek, 3 Sep 1979, J. Blyth (NTM).

Paratypes. NT: 15 males, ARRS, Radon Springs, 13/14 Apr 1988,

Suter and Wells (NTM and NMV); male, Magela Creek, 7 Apr 1993,

P. Dostine (NTM).

Other material. NT: male, 3 females, 12°50'S 132°51'E, 16 km

EbyN of Mt Cahill, 13 Jun 1973, J.C. Cardale (ANIC); male, 12°48'S

132°49'E, Kakadu National Park, Baroalba Springs, 16 Jan

1992, Wells and Webber (NTM); pupa, Litchfield National

Park, Walker Creek, 18 Apr 1992, A. Wells (NMV); 2 males,

Litchfield National Park, Walker Creek, 18/19 Apr 1992, A. Wells

(NMV); 4 males, 12°48'S 132°49’E, Kakadu National Park,

100

Alice Wells

Figures 60-63, Oecetis koobarra sp. nov.: 60-62, male genitalia, ventral, dorsal and lateral views; 63, forewing.

Figures 64-67, O.falcata sp. nov.; 64, forewing; 65-67, male genitalia in ventral, dorsal and lateral views.

Figures 68, 69, O. terania sp. nov., male genitalia in lateral and ventral views.

Baroalba Springs, 25 Apr 1991, Wells and Webber (NTM); 3 males, 1

female, 12°48'S 132°49'E, Kakadu National Park, Baroalba Springs,

29 May 1991, Wells and Webber (NTM); male, Melville Island,

11°36'S 130°43'E, 4 Oct 1996, G.R. Brown (NTM); male, Melville

Island, nr Pickertaramoor, 11°46'S 130°53'E, 10 Oct 1996, G.R. Brown

(NTM).

Diagnosis. Dark zig-zag mark on forewing (Fig. 63) formed

by pattern on wing membrane around the stepped posterior

anastomosis; overall straight- sided male genitalia, phallus

narrow mesially, with apex obliquely truncate.

Description. Tibial spurs 1, 2, 2. Male forewing length 4. 6-5. 6

mm. Wing laminae with vestiture of equal length; forewing

(Fig. 63) with a zig-zag pattern marking posterior anastomosis,

a footstalk on fork 1, tl more distal than t2, t2 and t3 contigu-

ous. Male genitalia, Figs 60-62. Segment IX almost uniform in

length all round, preanal appendages well separated, ovoid.

Segment X with long, slender dorsal process. Inferior

appendages straight- sided, stepped in midlength on inner

margin, and with a slender dorsal lobe at midlength. Phallus in

lateral view slender medially, obliquely angled towards apex.

Australian caddis flies of the genus Oecetis

101

Distribution. Northern NT (the ‘Top End’), and offshore

Melville Island in the Tiwi Islands Group (Fig. 120).

Remarks. Oecetis koobarra is distinctive for both its wing

markings and its very angular male genitalia. It is probably

most closely allied to O. parka. An associated pupa is from a

cornucopia-shaped sand case.

Etymology. Koobarra, now the official name for Baroalba

Springs, one of the collecting localities.

Oecetis falcata sp. nov.

Figures 64-67, 121

Material examined. Holotype, male, Upper Jardine River, Cape York,

Qld, 11°14’S 142°36’E, 26 Oct 1979, M.S. and B.J. Moulds (NMV

T-18515).

Paratypes. Qld: 72 males, females, same data as for holotype (NMV,

ANIC).

Other material. Qld: male, female, Iron Range, Cape York Pen.,

2- Jun 1971, E.F. Riek (ANIC); male, female, Mt Molloy, 13 Jun 1971,

E.F. Riek (ANIC); male, 2 females, Iron Range, West Claudie River,

17 Sep 1974, Moulds (NMV); male, Iron Range, Middle Claudie

River, 2-9 Oct 1974, Moulds (NMV); male, Jardine River, Cape York,

11°09'S 142°27'E, 11 Oct 1979, M.S. and B.J. Moulds (NMV);

7 males, 4 females, Jardine River, Cape York, 11°08'S 142°35'E,

14 Oct 1979, M.S. and B.J. Moulds (NMV); 3 males, Jardine River,

Cape York, 11°17'S 142°35'E, 17 Oct 1979, M.S. and B.J. Moulds

(NMV); 8 males, 2 females, Jardine River, Cape York, 11°19'S

142°37'E, 22 Oct 1979, M.S. and B.J. Moulds (NMV); 2 males, Jardine

River, Cape York, 11°14'S 142°36’E, 24 Oct 1979, M.S. and B.J.

Moulds (NMV); male, Qld, Tributary of Bertie Creek, 250 m SW of

Heathlands HS, 11°45'S 142°35’E, 11 Feb 1992, Cartwright and Wells,

1 1°45'S 142°35'E (QM) (slide); 3 males, 1 female, Cape York Peninsula,

Pascoe River crossing (to Iron Range), 4.10.2002, G. Theischinger

(ANIC); 6 males, 7 females, Cape York Peninsula, Dul-hunty River

crossing at Telegraph rd, 7.10.2002, G. Theischinger (ANIC).

Diagnosis. Forewing length about 4 times maximum width,

footstalk on fork 1 almost equal in length to the fork. In the

male genitalia, a pair of asymmetrical, scythe-like, sclerotised

processes dorsally, one at each side of the phallus

Description. Spurs 1, 2, 2. Male forewing length 4. 1-5.2 mm.

Wings (Fig. 64): forewing with patches of dark membrane at

crossveins of posterior anastomosis and, more proximally,

triangles of dark membrane at several forks; narrow, tapered

distally; footstalk on fork 1 equal in length to fork; tl and t3 at

about same level in wing, t2 more proximal. Male genitalia,

Figs 65-67. Abdominal segment IX narrow, but broader than in

O. ancala. Preanal appendages about twice as long as wide.

Tergite X very slender, reaching almost to length of other

genitalic parts. Inferior appendages in ventral view broad-

based, laterally produced to about twice length of basal section,

in lateral view more slender. Phallus broad. Dorsolateral

processes unequal, the left one twisted in basal half, curved out-

wards in distal half, right one more or less that same shape, but

less twisted.

Distribution. Far northern Qld (Fig. 121).

Remarks. As with O. ancala, grouping of this species is

equivocal, as the homologies of the processes are uncertain.

Etymology. Latin, falcata — sickle-shaped, for the shape of the

lateral processes.

Oecetis terania sp. nov.

Figures 68, 69, 122

Material examined. Holotype, male, NSW, Terania Creek, N of

Lismore, 28°25'S 153°18'E, 21 Jan 1986, G. Theischinger (NMV

T-18516) (slide).

Paratypes. 2 males, 1 female, same data as holotype (NMV) (one

male on slide).

Diagnosis. Wings with dense vestiture of downy hair and male

genitalia with inferior appendages bifurcate in lateral half, the

ventral lobes curving and clasper-like, the dorsal lobes more

slender and straight.

Description. Spurs 1, 2, 2. Male forewing length 6.0-6. 1 mm.

Wings typical form, tapered distally; footstalk on fork 1 about

half length of fork; t2 more distal than tlor t3. Genitalia, Figs

68, 69. Abdominal segment IX narrow. Preanal appendages dis-

crete, about twice as long as wide in lateral view, more or less

conical in dorsal view. Tergite X a short, narrow membrane,

about length of preanal appendages. Inferior appendages in

ventral view broad-based, bilobed in distal half, with the ven-

tral lobes almost meeting mesially, dorsal arms slender and

straight; in lateral view stout and irregular in shape basally, the

dorsal arm almost straight, extending well beyond the ventral

arm. Phallus curved ventrally.

Distribution. North-eastern NSW (Fig. 122).

Remarks. In shape the male inferior appendages of this species

more closely resembles those of some complexa- group species,

but the phallus lacks spines or parameres of the kind seen in

that group. The species is known only from the type locality.

Etymology. For the creek at the collecting site.

Oecetis papposa sp. nov.

Figures 70-73, 123

Material examined. Holotype, male, NT, South Alligator River, UDP

Falls [Gunlom], 13°24.9'S 132°26.0'E, J. Blyth (NMV T-18517).

Paratypes. NT: male, Jim Jim Creek, 3 km below falls, Kakadu

National Park, 1 Sep 1979, J. Blyth (NMV); male, Katherine River

Gorge National Park, 13 Aug 1979, J. Blyth (NMV); male, ARRS

Creek 5 km W of OSS South Alligator field station, 19 Apr 1989, Wells

and Suter (NMV); 6 males, ARRS, Radon Springs, 13-14 Apr 1989,

Suter and Wells (NTM); 1 male, 12°42'S 132°57'E, Kakadu National

Park, Magela Creek, OSS Site 009, 15 Mar 1991, Wells and Webber

(NTM); male, 12°48’S 132°49'E, Baroalba Springs, 4 Apr 1991, Wells

and Webber (NTM); male, 12°48'S 132°49'E, Baroalba Springs, 16

Aug 1991, Wells and Webber (NTM); 4 males, 2 females, 12°48'S

132°49'E, Baroalba Springs, 25 Apr 1991, Wells and Webber (NTM);

2 males, female, 12°48'S 132°49'E, Baroalba Springs, 4 Oct 1991,

Wells and Webber (NTM).

Other material. NT: male, Litchfield National Park, Florence Falls,

13°03'S 130°47'E, 9 Apr 1991, Wells and Horak (NMV); 4 males,

12°48’S 132°49’E, Baroalba Springs, 16 Jan 1992, Wells, Webber and

Bickel (NTM); male, Litchfield National Park, Walker Creek, 18-19

Apr 1992, A. Wells (NTM).WA: 4 males, Drysdale River headwaters,

30 km NW Mt Elizabeth H.S., 30 Sep 1979, J. Blyth (NMV); 9 males,

102

Alice Wells

Figures 70-73, Oecetis papposa sp. nov.: 70-72, male genitalia in ventral, dorsal and lateral views; 73, fore- and hind wings.

Figures 74-76, O. curta sp. nov., male genitalia, ventral, dorsal and lateral views.

Figures 77-79, O. aduncata sp. nov.: male genitalia, ventral, dorsal and lateral views.

King Edward River, 14°54’S 126°12’E, 3 Sep 1996, I. Edwards

(NMV); 6 males, Pearson’s Creek S edge of Prince Regent Res.,

16°01'S 125°35'E, 5 Sep 1996, I. Edwards (NMV); males, females,

Drysdale River upper reaches, 16°09'S 125°58'E, 7 Sep 1996, I.

Edwards (NMV); males, females, Manning River nr Mt Barnett,

16°40'S 125°56'E, 8 Sep 1996, 1. Edwards (NMV).

Diagnosis. Hindwing vein M unbranched; in male genitalia,

short, straight phallus and elongate inferior appendages with a

strong medial constriction.

Description. Spurs 0, 2, 2. Wings (Fig. 73) narrow, length over

4 times maximum width, with downy hair (readily lost in pre-

served specimens). Male forewing length 4. 2-5. 4 mm.

Forewing with footstalk on fork 1 of variable length, less than

or equal to length of fork; posterior anastomosis in part

obliquely linear, t3 more proximal than tl and t2. Hind wing

with venation strongly reduced, M unbranched. Maxillary palps

very long, all segments of equal length. Male genitalia, Figs

70-72. Segment IX excavated midventrally, preanal

Australian caddis flies of the genus Oecetis

103

appendages elongate-ovate, separated. Segment X in form of

a pair of stout, membranous lobes, with setate, digitiform

mesal process. Inferior appendages in ventral view broad-

based, slender distally, slightly dilated towards apices, in

lateral view, slender but constricted medially. Phallus short,

straight, with only a slight apicoventral lip.

Distribution. Northern NT and northern WA (Fig. 123).

Etymology. From Latin, pappus — hair, for the hairy wings.

Oecetis curta sp. nov.

Figures 74-76, 124

Material examined. Holotype. Male, Qld, Girraween National Park, nr

Wyberba, 10 Oct 1973, A. Neboiss (NMV T-18518).

Paratypes. Qld: male, same data as for holotype; 1 male, Mothar

Mtn, 32 km SE of Gympie, 29 Oct 1980, A. Neboiss (NMV); male,

Cape York Pen, Laura, 15°33.9'S 144°27.1'E, 7 Oct 1979, M.S. and

B.J. Moulds (NMV). NSW: 3 males, female, 48 km N of Singleton,

‘Tuglo’ alt. 760 m, 5 Oct 1975, M.S. Moulds; (NMV); 16 males, 4

females, Terania Creek, N of Lismore, 28°25'S 153°18'E, 21 Jan 1986,

G. Theischinger (NMV); 6 males, female, Wilson River, NW of

Wauchope, 31°14'S 152°34’E, 30 Oct 1981, Wells and Carter (NMV).

Diagnosis. Wings uniformly fuscous to mottled; forewing with-

out footstalk on fork 1 . In male genitalia, tergite X covered with

short setae and shallowly cleft apically; inferior appendages

elongate and straight to slightly curved in lateral view.

Description. Spurs 1, 2, 2. Male forewing length 6.6-8. 6 mm.

Wings rounded apically, hair along veins long, downy in

appearance, forewing fork 1 sessile or with a very short foot-

stalk; tl more distal than t2 and t3 which are almost contiguous,

all three crossveins marked by darkened membrane. Male

genitalia, Figs 74-76. Segment IX with lateral cleft, preanal

appendages small, rounded, widely separated. Segment X

robust and setate, cordate but with apex cleft. Inferior

appendages stout, in ventral view separated basally, rounded

apically, uniformly stout along length; in lateral view slender,

apically obliquely truncate. Phallus short, rounded in ventral

view, in lateral view down-turned distally.

Distribution. Eastern NSW, and south-eastern and far northern

Qld (Fig. 124).

Remarks. The unusual, hairy segment X may represent the

medial process of segment X such as is seen in O. aduncata or

O. minasata, but expanded. A similar plate is seen in O.

cepaforma, but that species has setae only distally, and is not

cleft apically.

Etymology. Latin, curta — short, for the phallus.

Oecetis aduncata sp. nov.

Figures 77-79, 125

Material examined. Holotype, male, NSW, Boonoo Boonoo River, 10

km upstream of falls, 28°37.9'S 152°15.1'E, 31 Oct 1975, A. Neboiss

(NMV T-18519) (slide).

Paratypes. NSW: 1 male, Bargo River, 10 km SW of Picton,

34°14.9'S 150°34.0"E, 30 Dec 1977, A. Neboiss (NMV). Vic: 1 male,

Watson’s Creek, 37°42'S 145°16'E, 16 Dec 1951, A. Neboiss (NMV);

1 male, Yae River, 7 km S of Glenburn, 37°25.4'S 145°25.3'E, 1 Dec

1972, A. Neboiss (NMV); 1 male, Belles Clearing, 6 km S of Aber-

feldy, 37°39.7'S 146°26.4'E, 8 Feb 1977, T131, A. A. Calder (NMV)

(slide); 1 male, Gibbo River, E xh ibition Creek, 20 km N of Benambra,

36°44.6'S 147°44.4'E, 16 Jan 1982, A. Wells (NMV) (slide).

Other material. Qld: 1 male, Tewah Creek, Tin Can Bay, 25°48.9'S

153°1.0'E, 17-18 Oct 1971, S.R. Monteith (ANIC); 2 males, The

Crater, nr Herberton, 25°2.9'S 148°24.1'E, 18 Dec 1974, M. Moulds

(slides) (NMV); 3 males, 15°17'S 145°10'E, 5 km WbyN Rounded Hill

nr Hope Vale Mission, 7 Oct 1980, J.C. Cardale (ANIC); 2 males,

15°47'S 145°17'E, Moses Creek, Finnigan, 14 Oct 1980, J.C. Cardale

(ANIC).

Diagnosis. Fork 1 in forewing sessile and wings mottled. In

forewing, tl more distal than t2 or t3; in male genitalia, infer-

ior appendages broad and strongly ridged along their length, in

lateral view with tips sharply down-turned and phallus in later-

al view appearing as if it has a triangular fold subapically.

Description. Spurs 1, 2, 2. Male forewing length 6. 1-7.1 mm.

Wings tapered apically; forewing fork 1 sessile, posterior

anastomosis obliquely stepped, tl more distal than other two

crossveins. Male genitalia, Figs 77-79. Segment IX with apical

margin excavated midventrally and midlaterally, preanal

appendages, small, well separated, ovate. Segment X broad,

membranous, in lateral view broad and down-curved, medial

process club-shaped. Inferior appendages appear to be slender

and strongly in-turned apically, but are actually broad and

deeply ridge, in lateral view broad-based, tapered distally

and down-turned apically. Phallus short, simple, down-turned,

in lateral view always with a more or less triangular area

dorsolaterally before apex.

Distribution. Eastern Australia, from eastern Vic. to south-

eastern Qld (Fig. 125).

Etymology. Latin, aduncata — bent inwards, for the shape of

the inferior appendages.

Oecetis ornata Kimmins, 1962

Figures 80-83, 126

Oecetis ornata Kimmins, 1962: 157-158.

Material examined. Holotype. Male, Humbolt Bay, Dutch New Guinea

[Irian Jaya] (BMNH).

Male, Qld, 3 km ENE Mt Tozer, 12°44’S 143°14’E, 2 Jul 1986, J.C.

Cardale (ANIC).

Diagnosis. Forewing (Fig. 80) strongly and coarsely patterned,

with dark brown markings on the proximal two-thirds. In

forewing fork 2 sessile; in male genitalia, phallus with a

sclerotised digitate process subapically on ventral side.

Remarks. The single male specimen available for study varies

slightly from the type as illustrated by Kimmins (1962).

However, given the proximity to New Guinea of the collecting

locality in far northern Qld and that the species was described

from a single male, the specimen is referred to O. ornata for the

present. Differences are mainly genitalic — shape of the pre-

anal lobes and of the inferior appendages in lateral view.

Figures drawn from the Australian specimen are provided for

comparisons (Figs 81-83).

104

Alice Wells

Figures 80, O. ornata Ki mmi ns, forewing (drawn from Australian specimen).

Figures 81-83, Oecetis ornata Kimmins: 79-81, male genitalia, ventral, dorsal and lateral views (drawn from Australian specimen).

Figures 84-87, O. cepaforma sp. nov.: 84-86, male genitalia, ventral, dorsal and lateral views; 87, forewing.

Figures 88-90, O. dostinei sp. nov.: 88, forewing; 89, 90, male genitalia, ventral, dorsal and lateral views.

Distribution. New Guinea and far northern Qld (Fig. 126).

Oecetis cepaforma sp. nov.

Figures 84-87, 127

Oecetis sp. D. — Wells, 1991: 61 (in key).

Material examined. Holotype. Male, Qld, Erwin Falls on Eliot Creek,

100 km S of Bamaga, 7 Nov 1988, K. Walker (NMV T-18520) (slide).

Paratypes. NT: male, Radon Springs, 13-14 Apr 1989, Suter and

Wells (NTM); 2 males, female, same locality and collectors, 18-19

May 1988 (NTM); 2 males, 1 female, same locality and collectors, 14

Apr 1989 (NTM); 1 male, 12°31'S 132°54'E, 9 km N by E of

Mudginberri HS, 10-11 Jun 1973, J.C. Cardale (ANIC);

Other material. 35 samples in ANIC, NMV, NTM and QM.

Diagnosis. Forewing broad, length scarcely 3 times maximum

width, spotted and moth-like, footstalk on fork 1 sessile. In

male genitalia inferior appendages with length about 4 times

width, a small spur on the baso-mesial angle; tergite X

extended to an elongate-triangular apex, tipped with several

short setae.

Australian caddis flies of the genus Oecetis

105

Description. Spurs 1, 2, 2. Male forewing length 5. 5-5. 7 mm.

Wings, broad, forewing (Fig. 87) with fork 1 sessile and with

large dark spots extending across membrane away from veins;

posterior anastomosis stepped, tl more distal than t3 which is

more distal than t2. Male genitalia, Figs 84-86. Segment IX

widest midlaterally, preanal appendages large, broadly rounded

laterally, fused medially. Segment X onion-shaped in outline in

dorsal view, without medial process. Inferior appendages

clasper- shaped, widely separated at bases, narrower distally

beyond a mesial angle bearing a spur and a cluster of short

setae; in lateral view with a medial notch dorsally. Phallus very

short.

Distribution. Qld, NT and WA (Fig. 127).

Remarks. The soft, broad, spotted wings give this species a

truly moth-like appearance. The male genitalia vary as

follows: specimens from northern Qld have the inferior

appendages more slender than those from NT, with the several

setae at the meso-basal angle shorter and stouter. Larvae build

cornucopia-shaped sand grain cases.

Etymology. Latin for onion- or minaret- shaped — cepaforma ,

being descriptive of the shape of segment X,

Oecetis dostinei sp. nov.

Figures 88-90, 128

Oecetis sp. G. — Wells, 1991: fig. 144.

Material examined. Holotype male, Qld, Mulgrave River W of

Gordonvale, 17°13.9'S 145°57.fE, 29 Apr 1979, A. Wells (NMV

T- 18521, slide).

Paratypes. Qld: male, data as for holotype (NMV) (slide); male,

Hann River, 73 km NWbyW Laura, 15°12’S 143°52’E, 27 Jun 1986, JC

Cardale (ANIC). NT: male, female, UDP Falls [Gunlom], 13°24.9’S

132°26.0E, 18-19 Jul 1980, MB Malipatil (NTM); male, ARRS South

Alligator River at Gimbat OSS Stn, 13°34.3’S 132°36.7E, 24 Mar

1988, Wells and Suter (NMV); 2 males, same loc., 28 Apr 1988,

P. Dostine (NTM); male, Litchfield National Park, 13°03’S 130°47E,

6 Jun 1991, Wells and Webber (NTM); male, Butterfly Gorge,

Katherine River Gorge National Park, 27 Jan 1977. M.S. and B.J.

Moulds (NMV).

Diagnosis. Forewing length scarcely 4 times maximum width,

spotted and moth-like, but with wing markings on the forewing

smaller than in O. cepaforma, fork 1 on forewing sessile. In

male genitalia inferior appendages with length about equal to 3

times width, without a small spur on the basomesial angle;

tergite X truncate apically.

Description. Spurs 1, 2, 2. Male forewing length 6. 1-6.4 mm.

Wings, broad and spotted, forewing (Fig. 88) with fork 1

sessile and posterior anastomosis stepped, with tl more distal

than t3 which is more distal than t2. Genitalia, Figs 89, 90.

Segment IX widest midlaterally, preanal appendages large,

rounded to broadly conical in dorsal view, in contact in mid-

line, but not fused. Segment X broad, apically truncate,

without medial process. Inferior appendages clasper- shaped,

widely separated at bases, curving outwards beyond a

mesial angle bearing a cluster of short stout setae. Phallus very

short.

Distribution. Northern NT and northern Qld (Fig. 128).

Species of the / ongiterga - g r ou p

Oecetis digitata sp. nov.

Figures 91-93, 129

Material examined. Holotype, male, NT, Jim Jim Waterhole, Kakadu

National Park, 5 Sepember 1979, J. Blyth (NMV T- 18522).

Paratypes. NT: 7 males, 3 females, Muirella Park, 12 Oct 1972, E.F.

Riek (ANIC); male, 12°06'S 133°04E, 19 km E by S of Mt Borrodaile,

5-6 Jun 1973, J.C. Cardale (ANIC); male, Jim Jim Creek, 19 km WSW

of Mt Cahill, 12°5’S 132°33E, 17 Jun 1973, J. Cardale (ANIC);

1 male, 1 female, Gulungul Billabong, East Jabiru, 23 Feb 1988,

P. Dostine (NTM); 1 male, 12°38'S 132°53E, Gulungul Billabong,

East Jabiru, 20 May 1989, A. Wells and P. Suter (NTM); 1 male,

ARRS, South Alligator River at Gimbat OSS Station, 13°34.3'S

132°36.7E, 24 May 1988, A. Wells and P. Suter (NTM); 3 males,

ARRS, Kambolgie Creek, 13°28.9'S 132°22.0E, 25 May 1988,

A. Wells and P. Suter (ANIC); 1 male, ARRS, East Alligator River at

Cahills Crossing, 27 May 1988, A. Wells and P. Suter (NTM); 2 males,

SAR site 1, 30 Sep 1988, P. Dostine (NTM); 5 males, 12°48’S

132°49E, Baroalba Springs, 25 Apr 1991, Wells and Webber (NMV);

1 male, 12°42'S 132°57E, Kakadu National Park, Magela Creek, OSS

Site 009, 8 Jun 1991, Wells and Webber (NTM); 1 male, 12°48'S

132°49E, Little Baroalba Creek, 10 Jul 1991, Wells and Webber

(NTM); 5 males, female, Alligator River, Two Mile Hole, 12°42'S

132°09E, 5 Aug 1996, 1 Edwards (NMV).

Other material.WA: Kimberley, Four Mile Creek, 2 Feb 1979, J.E.

Bishop (ANIC) (slide); male, female, 14°25’S 126°38E, CALM site

13/H 12 km S of Kalumbum Mission, 7-11 Jun 1988, T.A. Weir

(ANIC); 5 males, female, NT, Alligator River Two Mile Hole, 12°42'S

132°09E, 5 Aug 1996, 1. Edwards, NMV Qld: female, Palmer River,

20 Jun 1971, E.F. Riek (ANIC); male, 15°41'S 145°12E, Annan River,

3 km W by S Black Mountain, 17 Sep 1980, J.C. Cardale (ANIC);

male, 4 females, 15°25'S 141°53E, Hann River, 7 km NW by W Laura,

27 Jun 1986, J.C. Cardale (ANIC). NT: male, 12°57'S 132°33E, Jim

Jim Creek, 19 km WSW of Mt Cahill, 17 Jun 1973, J.C. Cardale,

ANIC; female, Howard Creek, 3 km E of Howard Springs, 12°27.5'S

131°3.1E, 17 Aug 1979, J. Blyth (NMV); 1 female, Magela Creek at

Ranger pipe outlet, 20 May 1988, P. Suter and A. Wells (NTM);

female, Magela Creek at Ranger Pipe Outlet, Suter and Wells, 23 May

1988 (NMV).

Diagnosis. Forewing length less than 4 times width, veins

sclerotised, appear very prominent, footstalk absent on fork 1;

an unusual forward bulge present at base of vein Cula. In male

genitalia, inferior appendages with mesial margin irregular, not

smoothly curved; phallus smoothly arched, length about 4

times width.

Description. Spurs 1, 2, 2. Male forewing length 6.6-8. 1 mm.

Wings strongly sclerotised with veins dark, hair short; in

forewing (Fig. 91) fork 1 sessile, anterior anastomosis with tl

and t2 almost linear, and well distad of t3, a forwardly directed

bulge on Cula. Male genitalia, Figs 92, 93. Segment IX almost

of uniform length on all sides, but excavated deeply mid-

ventrally; preanal appendages slender, elongate. Segment X

elongate in lateral view, slightly swollen towards base. Infer-

ior appendages in ventral view with small inner lobe at about

half length. Phallus, curved, slender with length about 4 times

width.

106

Alice Wells

Figures 91-93, Oecetis digitata sp. nov.: 91, forewing; 92, 93, male genitalia, ventral and lateral views.

Figure 94, O. crosslandi sp. nov., forewing.

Figures 95-97, O. ancala sp. nov., male genitalia, ventral, dorsal and lateral views.

Figures 98-100, O. crosslandi sp. nov., male genitalia, ventral, dorsal and lateral views.

Distribution. Qld, NT, Kimberley Region of northern WA (Fig.

129).

Remarks. Several of the New Guinean Oecetis species

resemble O. digitata in having broad wings with bold or promi-

nent venation, and may be allied to this species. However, none

has Cula as in O. digitata and O. ancala sp. nov.

Etymology. Latin, digitata — having fingers, for the finger-like

appearance of the male genitalia in lateral view.

Oecetis ancala sp. nov.

Figures 95-97, 130

Material examined. Holotype, male, SE Qld, Bulimba Creek, nr Bris-

bane Site Rl, near Kimmax Street riffle, 23 Oct 1979 (NMV T-18523).

Paratypes. Qld: 3 males, Camp Mountain, 31 Mar 1967, N.

Dobrotworsky (NMV); 2 males, female, Coondoo Creek, 30 km NE of

Gympie, Toolara State Forest, 28 Oct 1980, A. Neboiss (NMV); 3

males, female, Cooloola National Park, Freshwater Lake, 27 Nov

1985, D. Bickel and G. Cassis (NMV).

Australian caddis flies of the genus Oecetis

107

Other material. WA: male, N end of Lake Argyle nr Kununurra, 6

Feb 1977, M.S. and B. Moulds (NMV); 7 males, female, Geikie Gorge,

18°06'S 125°42'E, 5 Oct 1996, l Edwards (NMV). NT: 2 males,

2 females, Nourlangie Creek, 6 km E of Mt Cahill, 12°52'S 132°46E,

18 Nov 1972, J.C. Cardale (ANIC); male, Katherine River Gorge

National Park, 13 Aug 1979, J. Blyth (NMV); male, 2 females,

Adelaide River, 15 km E of Stuart Highway, 15 Aug 1979, J. Blyth

(NMV); male, Devil Devil Creek, 70 km SW of Daly River Mission,

23 Aug 1979, J. Blyth (NMV); 2 males, 2 females, 12°52’S 132°46E,

Nourlangie Creek, 6 km E of Mt Cahill, 18 Nov 1972, J.C. Cardale

(ANIC); male, ARRS, SAR at Gimbat OSS Stn, 13°34.3'S 132°36.7E,

Wells and Suter, 24 May 1988 (NTM); 3 males, ARRS, Kambolgie

Creek, 13°28.9’S 132°22.0E, 25 May 1988, Wells and Suter (NTM);

male, 12°36'S 132°53E, ARRS, Gulungul Creek, Inlet to Gulungul

Billabong, 20 Apr 1989, Wells and Suter (NTM); 2 males, SAR Site 1,

30 Sep 1988, P. Dostine (NTM); male, 3 females, 12°42’S 132°57E,

Kakadu National Park, Magela Creek, OSS Site 009, 15 Feb 1991,

Wells (NTM); male, 12°42’S 132°57E, Kakadu National Park, Magela

Creek, OSS Site 009, 8 Jul 1991, Wells and Webber (NTM). Qld:

males, female, Cape York Peninsula, Pascoe River crossing (to Iron

Range), 4 Oct 2002, G. Theischinger (ANIC); Cape York Peninsula,

Dulhunty River crossing at Telegraph rd, 7 Oct 2002, G. Theischinger

(ANIC). NSW: Barrington Tops, Barrington Tops Country Retreat,

Dam, 22 Dec 2000, A. Wells (ANIC). Vic: male, Tyers River, LRES,

24 Feb 1974, Site 22 (NMV) (slide); male, female, Yarra River, below

Upper Yarra Dam, 28 Feb 1976, A. Neboiss (NMV); male, female,

Yarra River, Diamond Creek junction, 14 Mar 1976, A. Neboiss

(NMV).

Diagnosis. Forewing as for O. digitata. In male genitalia,

paired, sharply angled, sclerotised processes ventral to tergite

X, in ventral view appearing to be lateral to the phallus.

Description. Spurs 1, 2, 2. Male forewing length 7.9-9.4 mm.

Wings, broad: forewing with fork 1 sessile, veins prominent;

posterior anastomosis almost linear, oblique, marked by dark

membrane forming a line across wing; Cula with a prox-

imally directed bulge. Male genitalia, Figs 95-97. Segment IX

narrow, preanal appendages discrete, slender, elongate, apices

obliquely truncate in lateral view. Tergite X slender, setose.

Inferior appendages stoutly clasper- shaped in ventral view, a

subapical notch mesially, in lateral view swollen baso-

ventrally, then rod-shaped to rounded apex. Phallus long for a

laustra- group species, arched ventrally. A pair of slender

processes occur ventral to tergite X, appearing to be lateral to

phallus in ventral view, sharply angled inwards at about half

their length (ankle- shaped), in lateral view broad-based,

tapered and slender distally.

Distribution. Northern WA, northern NT, south-eastern Qld,

eastern NSW and south-central Vic (Fig. 130).

Remarks. The homologies of the unusual lateral processes are

obscure, being impossible to determine from the prepared

slides. In lateral view they seem to be associated in some way

with tergite X, although in ventral view they appear to be

closely associated with the phallus. If they are derived from the

phallotheca, then this species should probably be placed in

the complexa- group in the Australian fauna, but for the present

they are dealt with as part of the laustra- group. Only few

specimens have been collected at any one time, which suggests

that the species may be far more widespread, but not often

collected. This is consistent with the very curious distribution.

Etymology. Latin, ancala — a bent arm, descriptive of the

structures lateral to the phallus.

Oecetis crosslandi sp. nov.

Figures 94, 98-100, 131

Material examined. Holotype male, Qld, Gunshot Creek at Telegraph

Crossing, 11°44'S 142°29E, 4-5 Apr 1992, M. Crossland (ANIC).

Paratypes. 2 males, Qld: Heathlands, 11°45'S 142°35E, T. Weir

(ANIC).

Diagnosis. Forewing broad, length about 3 times width; fork 2

sessile and wing veins strongly pronounced, but without bulge

in vein Cula. In male genitalia a single heavily sclerotised

dorsal paramere with a group of teeth and spikes subapically.

Description. Wings broad, veins pronounced; forewing (Fig.

94) apex rounded, fork 1 sessile, Cula almost straight. Male

forewing, 5.3 mm. In male genitalia (Figs 98-100), abdominal

segment IX ventrally about 3 times middorsal length; preanal

appendages elongate, length about 4 times width, apically

rounded. Tergite X comprising a slender, elongate median lobe

dorsal to a short rounded membranous plate. Inferior

appendages broad-based, in ventral view sharply constricted on

mesial margin, tapered to narrow apices, in lateral view with a

short dorsal lobe. Phallus slender, downcurved in distal third,

laterally on right, an elongate heavily sclerotised paramere with

a subapical twist below a cluster of teeth and spikes.

Distribution. Northern Cape York, Qld (Fig. 131).

Remarks. One can only speculate on which features of

O. digitata, O. ancala and O. crosslandi are homoplasious. I

have assumed here, tentatively, that the synapomorphy for the

group is the unusual form of the wing, which is shared by the

New Guinean O. longiterga and at least one other New

Guinean species (see Chen, 1992). However, Oecetis longi-

terga has a pair of internal parameres in the phallus, which

feature could otherwise place it in the O. pechana- group;

O. ancala and the New Guinea species of Chen’s unpublished

work have paired external parameres, somewhat similar to the

arrangement in the O. complexa- group species (Wells, 2000);

and O. digitata has the phallus simple, without parameres,

which is characteristic of the O. laustra- group.

Etymology. Named for Michael Crossland.

Acknowledgements

My thanks to all who collected and provided specimens on

which this study is based, and to the Museum Victoria and

Museums and Art Galleries of the Northern Territory for mak-

ing available the material in their collections. The Natural

History Museum, London, through Dr R. Vane Wright, gave

permission to reproduce figures from Mosely and Kimmins

(1953). CSIRO Publishing is acknowledged for permission to

use figures from the Australian Journal of Zoology and the

Australian Journal of Marine and Freshwater Research.

Museum Victoria is thanked for allowing use of figures from

Memoirs of the National Museum of Victoria. CSIRO

Entomology kindly provided laboratory facilities for this work.

108

Alice Wells

Figures 101-115, distribution of species within Australia:

101, Oecetis laustra Mosely; 102, O. pseudolaustra sp. nov.; 103, O. atarpa Mosely; 104, O. scirpicula Neboiss; 105, O. inscripta Kimmins;

106, O. brevidentata sp. nov.; 107, O. asmanista Mosely; 108, O. minasata Mosely; 109, O. aeoloptera Kimmins; 110, O. multipunctata

Ulmer; 111, O. cracenta sp. nov.; 112, O. parka Mosely; 113, O. arcada Mosely; 114, O. cymula Neboiss; 115, O. paracymula, sp. nov.

Figures 116-128, distribution of species within Australia:

116, Oecetis spicata sp. nov.; 117, O. crena sp. nov.; 118, O. quadrata sp. nov.; 119, O. dilata sp. nov.; 120, O. koobarra sp. nov.;

121, O.falcata sp. nov.; 122, O. terania sp. nov.; 123, O. papposa sp. nov.; 124, O. curta sp. nov.; 125, O. aduncata sp. nov.;

126, O. ornata Ki mmins; 127, O. cepaforma sp. nov.; 128, O. dostinei sp. nov.

110

Alice Wells

Figures 129-131, distribution of species within Australia: 129, O. digitata sp. nov., 130, O. ancala sp. nov.; 131, O. crosslandi sp. nov.

My colleagues at the Australian Biological Resources Study are

acknowledged for advice, particularly on handling of graphics,

and for help in the use of the mapping programme, Maplnfo,

used to produce the maps. Brigitte Kuhlmayr’s graphics skill

improved some of the scanned images of published figures.

References

Chen, Y.E. 1992. Revision of the Oecetis (Trichoptera: Leptoceridae)

of the World. Unpublished PhD Thesis, Clemson University, South

Carolina, USA.

Floyd, M.A. 1995. Larvae of the caddisfly genus Oecetis (Trichoptera:

Leptoceridae) in North America. Bulletin of the Ohio Biological

Survey 10(3): i-viii 1-85.

Kimmins, D.E. 1962. Miss L. E. Cheeseman’s expeditions to New

Guinea. Trichoptera. Bulletin of the British Museum of Natural

History ( Entomology ) 11: 99-187.

Mosely, M.E., and Ki mmins, D.E. 1953. The Trichoptera ( caddis-flies )

of Australia and New Zealand. British Museum (Natural History):

London. 550 pp.

Neboiss, A. 1977. A taxonomic and zoogeographic study of Tasmanian

caddis-flies (Insecta: Trichoptera). Memoirs of the National

Museum of Victoria 38: 1-208.

Neboiss, A. 1982. The caddis-flies (Trichoptera) of south-western

Australia. Australian Journal of Zoology 30: 271-325.

Neboiss, A. 1986. Atlas of Trichoptera of the SW Pacific — Australian

Region. Dr W. Junk Publishers : Dordrecht/Boston/Lancaster. 286

pp.

Neboiss, A. 1989. The Oecetis reticulata species-group from the

South-West Pacific area (Trichoptera: Leptoceridae). Bijdragen tot

de Dierkunde 59(4): 191-202.

Ruiter, D.E. 2000. Generic key to the adult ocellate Limnephiloidea

of the Western Hemisphere (Insecta: Trichoptera). Ohio Biological

Survey Miscellaneous Contributions 5. (Ohio Biological Survey:

Columbus).

Schmid, F. 1980. Les insectes et arachnides du Canada, Partie 7,

Genera des Trichopteres du Canada et des Etats adjacent.

Agriculture Canada Publication 1692: 1-296.

Schmid, F. 1987. Considerations di verses sur quelques genres lepto-

cerins (Trichoptera, Leptoceridae). Bulletin de Vlnstitut Royal des

Sciences Naturelles de Belgique. Entomologie Vol. 57 Supplement:

1-147.

St Clair, R.M. 1994. Some larval Leptoceridae (Trichoptera) from

south-eastern Australia. Records of the Australian Museum 46:

171-226.

St Clair, R.M. 2000. Preliminary keys for the identification of

Australian caddisfly larvae of the family Leptoceridae. Cooperative

Research Centre for Freshwater Ecology, Identification Guide 27:

1-83.

Ulmer, G. 1916. Results of Dr. E. Mjoberg’s Swedish scientific expe-

dition to Australia 1910-1913, ‘Trichoptera’. Arkiv for Zoo logi 10:

1-23.

Wells, A. 1991. A guide to the caddisflies (Trichoptera) of the Alligator

Rivers region, Northern Territory. Open File Record available from

Supervising Scientist for the Alligator Rivers Region, GPO Box

461, Darwin, NT. 105 pp.

Wells, A. 2000. New Australian species of Oecetis allied to O. com-

plexa Kimmins (Trichoptera: Leptoceridae). Memoirs of Museum

Victoria 58(1): 77-88.

Memoirs of Museum Victoria 61(1): 111-118 (2004)

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

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

Descriptions of new species and a new genus of leptophlebiid mayflies (Insecta:

Ephemeroptera) from the Northern Territory, Australia

J. C. Dean 1 and P. J. Suter 2

1 Environment Protection Authority, Freshwater Sciences, Ernest Jones Drive, Macleod, Victoria 3085

(j ohn.dean @ epa. vie .gov. au)

2 Department of Environmental Management and Ecology, La Trobe University, PO Box 821, Wodonga, Victoria 3689

(p.suter@latrobe.edu.au)

Abstract Dean, J.C., and Suter, P.J. 2004. Descriptions of new species and a new genus of leptophlebiid mayflies (Insecta:

Ephemeroptera) from the Northern Territory, Australia. Memoirs of Museum Victoria 61(1): 111-118.

Adults and nymphs of a new monotypic genus ( Manggabora gen. nov.) and three new species of leptophlebiid

mayflies ( Manggabora wapitja sp. nov., Atalophlebia gubara sp. nov. and Tillyardophlebia dostinei sp. nov.) are

described from Kakadu National Park in northern Australia.

Key words mayflies, Leptophlebiidae, taxonomy, new species, new genus, Australia, Kakadu

Introduction

Although the Ephemeroptera (mayflies) of the Northern

Territory have been reasonably well known since environ-

mental monitoring programs were commenced in the early

1980s, their taxonomy has been neglected and most species

remain undescribed. The first serious attempt to document the

fauna was by Suter (1992) who sampled extensively in the

Alligator Rivers Region, associated adults with immature

aquatic stages by rearing, and produced identification keys to

both adults and nymphs of recognised voucher species. This

study formed the basis for inclusion of Northern Territory taxa

in subsequent identification guides to Australian mayfly

nymphs (Suter, 1997, 1999; Dean, 1999).

Suter (1992) recorded nine species of the family

Leptophlebiidae placed in five genera, and Dean (1999) recog-

nised ten species in seven genera. A recent attempt to collect

additional material was only partially successful, primarily due

to unsuitable conditions at the end of the dry season. Three new

species are described below, and a new genus is erected to

accommodate one of them.

Material has been preserved in alcohol, with parts of

some specimens mounted on microscope slides. Terminology

follows Peters et al. (1978). All type material is lodged in the

Museum of Victoria, Melbourne (NMV). The following

abbreviations are used for material examined: MI male imago;

FI female imago; MSI male subimago; FSI female subimago;

N nymph.

Family Leptophlebiidae

Subfamily Atalophlebiinae

Atalophlebia gubara sp. nov.

Figures 1-13

Type material. Holotype: male imago, Gubara (Baroalba Springs),

Kakadu National Park, Northern Territory, 12°49'S 132°53'E, 14 Aug

1999, J.Dean (NMV T-l 8498).

Paratypes: collected with holotype, 8 male imagos (NMV

T-18499-T- 18506).

Other material examined. Northern Territory. 3MI, 2N, Gubara

(Baroalba Springs), 12°49'S 132°53'E, 14 Aug 1999, J. Dean; 1FSI,1N,

Gulungul Creek (Radon Springs), 12°45’S 132°55’E, 18 May 1988, P.

Suter; 4N, Gulungul Creek (Radon Springs), 12°45'S 132°55’E,13 Aug

1999, J. Dean; 3N, Manggabor Creek, Arnhem Land, 12°17'S

134°05'E, 26 Aug 1999, J. Dean; 1MSI, 1FI (both reared from

nymphs). Walker Creek, Litchfield National Park, 13°05'S 130°42'E,

31 Aug 1999, J. Dean; 2MSI,2FSI, (all reared from nymphs), 3FI, 2N,

Magela Creek, Bowerbird Billabong, 12°47'S 133°02'E, 30 June 1990,

P. Suter; IN, Magela Creek, 1.5 km d/s Bowerbird Billabong, 12°46'S

133°02'E, 28 May 1988, P. Suter; 7N, Magela Creek, d/s Bowerbird

Billabong, 12°47’S 133°02’E, 3 May 1990, D.Cartwright; IN, South

Alligator River, d/s Gimbat, 13°35'S 132°36'E, 2 May 1990, D.

Cartwright; IN, South Alligator River, Koolpin Crossing, 13°32'S

132°33'E, 18 Aug 1999, J. Dean; 1MI,1FI, South Alligator River,

Koolpin Crossing, 13°32'S 132°33'E, 14 Oct 1987, P.Dostine; 2N,

South Alligator River, Coronation Hill, 13°36'S 132°37'E, 20 Apr 1987,

P. Dostine.

New mayflies from the Northern Territory

113

Description. Imago. Length of male: body 6. 8-7. 9 mm,

forewing 6. 1-6.9 mm; Length of female: body 6.5-7. 9 mm,

forewing 6.0-7.0 mm. Head medium brown; antennae medium

brown; male eyes with upper lobes medium brown, lower lobes

grey, upper lobes separated by distance approximately equal to

diameter of median ocellus. Thorax with meso- and meta-

scutum medium brown; thorax laterally pale yellow, with small

patches of dark brown pigment. Forelegs medium brown,

femora with dark brown bands a little beyond midlength and at

apex, tibiae with dark band at base and more heavily pig-

mented in apical half. Middle and hind legs paler, yellowish,

femora with dark bands as in foreleg, tarsal segments also

darker brown. Tarsal claws similar, with apical hook and

opposing ventral flange (Fig. 3). Forewings hyaline, cross veins

in costal and subcostal spaces washed with brown pigment;

elsewhere in forewing many crossveins and cells washed with

pale brown (Fig. 1). Male abdomen dark brown, pattern of paler

yellow maculae, predominantly along midline (Fig. 4); abdom-

inal sterna pale, with some darker brown markings. Penes

narrow at midlength, broader in apical half, the two halves

fused almost to apex; a pair of large ventro-lateral lobes at

about three-quarters length, and closer to the apex a smaller

ventro-median lobe (Figs 5-7); median lobe underlying deep

pockets with internal spine-like setae (Fig. 8). Male imago with

3 apical filaments, dark brown in basal half and white in apical

half. Female abdominal colour pattern similar to male;

abdominal sternum 9 with posterior margin deeply excised

(Fig. 9); strongly projecting egg guide on posterior

margin of abdominal sternum 7. Subimago. Wings yellow-

pale brown, forewing with suffusion of darker brown

covering most cross veins. Abdominal colour pattern

similar to imago but lateral pale areas more strongly

developed. Nymph. General colour medium to dark brown,

paler markings on abdominal terga; legs yellowish with brown

bands on all segments. Labrum broad, width approximately 2.5

times length along median line (Fig. 10). Foretarsus with

about 10 ventral spine-like setae, each seta less than one-sixth

diameter of tarsus (Fig. 11). Abdominal segments 2-9 with

relatively short posterolateral spines, those on segment V

about one-eighth length of segment (Fig. 12). Gills on abdom-

inal segments 1-7; all gills with upper and lower lamella

tridigitate (Fig. 13).

Etymology. The species is named for the type locality.

Comments. Although previously undescribed, this species has

been included in keys as Atalophlebia sp.l (Suter, 1992) and

Atalophlebia sp.AV16 (Dean, 1999). The species is distin-

guished from all other species of the genus by the structure of

the male genitalia, and in the nymph by the shape of the gills,

the size and number of ventral spines on the foretarsus and the

short posterolateral abdominal spines.

Manggabora gen. nov.

Type species. Manggabora wapitja sp. nov.

Diagnosis. Imago. Forewing length-width ratio 2. 8-3.0 (Fig.

14); membrane hyaline, except costal and subcostal cells in api-

cal third of wing which are opaque, white; costal crossveins

basal to bulla very faint, difficult to see, crossveins elsewhere

weakly developed; IC^ usually linked to CuA-CuP cross vein,

terminating free in some individuals; ICU[ and ICu, parallel as

wing margin approached. Hindwing approximately 0.22 length

of forewing; costal margin with shallow concavity a little

beyond midlength; vein Sc joining costal margin at about 0.85

wing length (Fig. 15). Legs with tarsal claws dissimilar, one

with an apical hook and opposing ventral flange, the other

large, pad-like (Fig. 16). Male genitalia (Figs 19-22) with

claspers three-segmented, penes extending beyond narrowing

of claspers; penes lobes narrow, fused almost to apex, ventral

surface with a robust, medial projection; each lobe with a stout,

retractable apical spine. Female ninth sternum strongly project-

ing, posterior margin entire (Fig. 23). Subimago. Wings uni-

formly pale grey-yellow, without pattern. Mature nymph.

Labrum a little broader than clypeus; maximum width about

2.3 times length along median line (Fig. 25); anterior margin

overhanging medial notch; frontal setae sparse, arranged as

narrow band. Mandibles (Figs 26, 27) with outer incisors slen-

der; outer margin swollen at base of incisor. Maxillae with sub-

apical row of 12-15 pectinate setae on ventral surface (Fig. 28).

Labium with glossae slightly dorsal to paraglossae, not turned

under ventrally (Fig. 29); labial palp with terminal segment

about half length of middle segment, without spine-like setae

along inner margin. Femora with long spine-like setae and hair-

like setae along outer margin (Fig. 30); tarsi with all ventral

spines similar in length; tarsal claws with ventral teeth (Fig.

31). Abdominal segments with posterolateral spines on seg-

ments 4 or 5-9; posterior margins of abdominal terga with an

almost continuous series of very small triangular spines, about

10 pm long, interspersed with fewer long hair-like seta (Fig.

33); gills linear, lateral tracheae very weakly developed or

absent (Fig. 32). Caudal filaments (Fig. 34) with apical whorl

of flattened, triangular spines on each segment, and between

each spine a series of 4 or 5 fine setae about half segment

length.

Etymology. The genus is named after Manggabor Creek,

Arnhem Land, one of the collection localities for the species.

Feminine.

Remarks. Manggabora is a member of the Austrophlebioides

lineage, as evidenced by the following features: (1) adult tarsal

claws dissimilar; (2) forewing with ICu L usually linked to CuA-

CuP crossvein; (3) sternum 9 of female with apical margin

entire; (4) mouthparts of nymph with labrum broader than

clypeus; (5) mandibles with outer incisors slender; (6) terminal

segment of labial palp, short, about half length of middle

segment. The lineage is restricted to the Southern Hemisphere,

and includes the Australian genera Austrophlebioides ,

Tilly ardophlebia and Kirrara. Manggabora can be dis-

tinguished from all other genera in the lineage by the fol-

lowing combination of characters: (1) penes lobes relatively

slender, fused almost to apex, with large ventral projection

near apex; (2) anterior margin of labrum overhanging central

notch; (3) maxillae with only 12-15 pectinate setae in sub-

apical row; (4) absence of elongate ventral spines from all

tarsi; (5) posterolateral spines on abdominal segments 4 or 5-9

only.

114

J. C. Dean and P. J. Suter

Manggabora wapitja sp. nov.

Figures 14-34

Type material. Holotype: male imago (reared from nymph),

Coobanrbora Spring, Kakadu National Park, Northern Territory,

12°24'S 132°40'E, 21 Aug 1999, J. Dean (NMV T- 18491).

Paratype: male imago (reared from nymph; wings, legs and

nymphal exuvia mounted on slides), collected with holotype (NMV

T- 18492).

Other material examined. Northern Territory. ION, Coobanrbora

Spring, Kakadu National Park, Northern Territory, 12°24'S 132°40'E,

21 Aug 1999, J. Dean; 1MI, 17N, Koolpin Gorge, Kakadu National

Park, 13°30'S 132°35'E, 18 Aug 1999, J. Dean; 1MI, 1FSI, 11N,

Walker Creek, Litchfield National Park, 13°05’S 1 30°42E, 31 Aug

1999, J. Dean; 14N, Florence Falls, Litchfield National Park, 13°06'S

130°47'E, 31 Aug 1999, J. Dean; 1FSI, 21N, Tolmer Falls. Litchfield

National Park, 13°12'S 130°43'E, 31 Aug 1999, J. Dean; 2MI, 20N,

Mann River, Amhem Land, 12°22’S 134°08’E, 26 Aug 1999, J. Dean;

6N, Liverpool River, Amhem Land, 12°21''S 134°07'E, 26 Aug 1999,

J. Dean; 1 FI, 22N, Manggabor Creek, Arnhem Land, 12°17'S

134°05'E, 26 Aug 1999, J. Dean; 1FI, 56N, Kambolgie Creek, Kakadu

National Park, 13°31'S 132°23'E, 16-19 Aug 1999, J. Dean; 3N,

Kambolgie Creek, Kakadu National Park, 13°31'S 132°23'E, 1 May

1990, D. Cartwright; 13N, Barramundie Gorge, Kakadu National Park,

13°19'S 132°26'E, 22 Aug 1999, J. Dean; 1FI, 3N, Gerowie Creek, 7

km N of Bukbukluk Lookout, Kakadu Highway, 13°26’S 132°16’E, 19

Aug 1999, J. Dean; 32N, creek 2 km N of Bukbukluk Lookout, Kakadu

Highway, 13°29'S 132°15’E, 19 Aug 1999, J. Dean; 1FI, IN, South

Alligator River, Koolpin Crossing, 13°32'S 132°33'E, 18 Aug 1999, J.

Dean; 13N, South Alligator River, Gunlom Road Crossing, 13°30'S

132°29'E, 19 Aug 1999, J. Dean; 2N, South Alligator River, d/s

Gimbat, 13°35'S 132°36F, 1 May 1990, D. Cartwright; 1MI, 1MSI,

1FSI, Jim Jim Creek, 3 km d/s Falls, 1 Sep 1979, J. Blyth; 2N, Magela

Figs 14-23 Manggabora wapitja Male imago: 14, forewing and outline of hind-wing; 15, hind-wing enlarged; 16, foretarsal claws; 17, abdomi-

nal terga, dorsal; 18, abdominal segments 3-5, lateral; 19, genitalia, ventral; 20, penes lobes, ventral; 21, genitalia, lateral; 22, penes lobes, api-

cal. Female imago: 23, sternum, abdominal segment IX.

New mayflies from the Northern Territory

115

Figs 24-34 Manggabora wapitjci Nymph: 24, nymph; 25, labrum; 26, left mandible, dorsal; 27, right mandible, dorsal; 28, left maxilla, ventral;

29, labium, dorsal (left of midline) and ventral (right of midline); 30, foreleg; 31, foretarsal claw; 32, gill, abdominal segment IV; 33, spines,

posterior margin of abdominal tergum V; 34, terminal filament, midlength.

Creek, u/s Bowerbird Billabong, 3 May 1990, D. Cartwright; 16N,

Radon Springs, Kakadu National Park, 12°45'S 132°55'E, 23 Apr.

1990, D. Cartwright; 4N, Gulungul Creek, Radon Springs, Kakadu

National Park, 12°45'S 132°55'E, 13 Aug 1999, J. Dean; 25N, Baroalba

Creek, Kubarra Pools, 12°49’S 132°52’E, 28 Apr 1990, D. Cartwright;

11N, Baroalba Creek, Gubarra Pools, 12°49’S 132°52'E, 14 Aug 1999,

J. Dean; 6N, Harris Creek, u/s South Alligator River, 1 May 1990, D.

Cartwright. Western Australia. 27N, King Edward River, Mitchell

River Road, 25 Sep 1995, L. Metzeling.

Description. Imago. Length of male: body 4. 7-5.0 mm,

forewing 5. 1-5.2 mm; Length of female: body 4. 1-5.9 mm,

forewing 4. 9-6. 3 mm. Head predominantly medium brown;

antennae pale yellow; ocelli white, black at base; male eyes

with upper lobes pale orange-brown, in contact dorsally, lower

lobes grey. Thorax with meso- and metascutum golden-yellow;

lateral surfaces golden with some patches of medium brown.

Forelegs with femora reddish-brown, tibiae pale yellow with

apex medium brown, tarsi pale yellow; middle and hind legs

uniformly pale yellow; forelegs of male with ratios of segment

lengths 0.69-0.70; 1.00 (1.74-1.75 mm); 0.05; 0.27-0.31; 0.28;

0.18-0.21; 0.10-0.11; tarsal claws dissimilar, one claw with

apical hook and ventral flange and the other expanded and pad-

like, without terminal hook (Fig. 16). Fore wings hyaline except

for pterostigma, which is white and opaque; veins predomi-

nantly unpigmented (Fig. 14). Male abdomen reddish, dorsally

pale tending to hyaline; each segment with narrow band of

116

J. C. Dean and P. J. Suter

medium brown along posterior margin and pair of weakly

developed brown lateral markings (Figs 17, 18); abdominal

sterna pale, tending to hyaline. Penes lobes (Figs 19-22)

narrow, fused almost to apex, each lobe with a stout, retractable

apical spine; subapically with a robust, medial projection on the

ventral surface. Female abdominal colour pattern similar to

male, although generally a little darker, reddish-brown; abdom-

inal sternum 9 strongly projecting, without apical excision (Fig.

23). Subimago. Wings pale greyish-yellow, abdominal colour

pattern similar to imago. Mature nymph. General colour

yellow. Mouthparts as in Figs 25-29. Legs yellow, all segments

banded; foretarsus with 10-15 ventral spine-like setae, rela-

tively uniform in length; tarsal claws with ventral teeth (Figs

30, 31). Abdomen yellow, each segment with pair of brown

lateral markings (Fig. 24); gills linear, lateral tracheae very

weakly developed or absent (Fig. 32).

Etymology. The name wapitja is derived from the word for

‘digging stick’ in the Datiwuy language of eastern Arnhem

land (Ganambarr, 1999), and refers to the appearance of the

male genitalia in lateral view.

Remarks. The genus is monotypic. Suter (1992) included this

species in his keys as “Leptophlebiidae Genus A sp.l”, and

Dean (1999) designated the nymph “Genus V sp.AVl”.

Tilly ardophlebia dostinei sp. nov.

Figures 35-47

Type material. Holotype: male imago (reared from nymph), Rockhole

Mine Creek, Kakadu National Park, Northern Territory, 13°30'S

132°30'E, 24 Jun 1995, P. Dostine (NMV T- 18493).

Paratypes: male imago (reared from nymph; wings, legs and

nymphal exuvia mounted on slide, labelled specimen 01), Rockhole

Mine Creek, Kakadu National Park, Northern Territory, 13°30’S

132°30'E, 24 Jun 1995, P. Dostine (NMV T- 18494); male imago, 2

female imagos (all reared from nymphs), Rockhole Mine Creek,

Kakadu National Park, Northern Territory, 13°30'S 132°30'E, 31 May

1995, P. Dostine (NMV T-18495- T- 18497).

Other material examined. Northern Territory. 1MSI (reared from

nymph), Rockhole Mine Creek, Kakadu National Park, Northern

Territory, 13°30'S 132°30'E, 24 Jun 1995, P. Dostine; 3N, Rockhole

Mine Ck, 2 May 1990, D. Cartwright.

Description. Imago. Length of male: body 5. 6-6. 5 mm,

forewing 6.0-6.6 mm; Length of female: body 5.5-5. 6 mm,

forewing 6.0-6.4 mm. Eyes of male with upper lobes brownish-

pink, in contact dorsally, lower lobes black. Thorax orange-

brown. Forewing with membrane hyaline (Fig. 35); costal and

subcostal cells in apical third of wing translucent, whitish;

length-width ratio 2.8-3.0; costal crossveins absent or weakly

developed basal to the bulla, 10-15 distal to the bulla; MA

forked at 0.37-0.39 wing length; MP 2 attached by crossvein to

MP, at about 0.18 wing length; ICuj linked to CuA-CuP

crossvein, ICu, and ICu 2 weakly diverging as wing margin

approached. Hindwing 0.20-0.22 length of forewing; costal

margin convex at about midlength, relatively straight basal and

distal to midpoint (Fig. 36); vein Sc joining costal margin a

little less than 0.9 wing length; hindwing with 3-5 costal

crossveins and 5-6 subcostal crossveins, all weakly developed.

Legs pale, not banded, but slightly darker brown at apex of

femur and tibia; tarsal claws dissimilar, one claw with an apical

hook and without an opposing ventral flange, the other large,

pad-like (Fig. 39); forelegs of male with ratios of segment

lengths 0.72-0.73; 1.00 (2.4 mm); 0.07-0.08; 0.31-0.32;

0.29-0.30; 0.22; 0.08-0.09. Abdomen predominantly pale yel-

low, restricted dark brown markings on segments 1-6, more

strongly developed markings on segments 7-9 (Figs 37, 38).

Male genitalia with claspers three- segmented, narrowing grad-

ually at about one-third length (Fig. 40); penes extending

beyond narrowing of claspers, lobes moderately broad, widely

separated in apical two-thirds and fused in basal third; each

lobe with a stout, inwardly directed subapical spine and a ven-

tral longitudinal ridge (Figs 41^-3). Female ninth sternum with

posterior margin entire, strongly convex, protruding beyond

apex of segment 10 (Fig. 44). Mature nymph. Head prog-

nathous. Mouthparts: Clypeus with lateral margins slightly

diverging to anterior. Labrum clearly broader than clypeus,

width about 2.4 times length along median line; 2 setal fringes

close to anterior margin, each fringe extending across more

than half width of labrum; anterior margin with broad central

notch, the base of which is concealed beneath an overhanging

canopy (Fig. 45). Mandibles with incisors slender. Maxillae

with subapical row of about 25 pectinate setae. Legs with tibi-

ae and tarsi relatively slender (Fig. 46); tibiae with ventral

spine-like setae relatively dense right to apex; tarsi with 10 or

fewer ventral spine-like setae, relatively uniform in length;

tarsal claws with ventral teeth, progressively larger apically.

Abdominal segments without setae on lateral margins, strongly

developed posterolateral spines on segments 2-9. Gills present

on abdominal segments 1-7, each gill narrowly lanceolate

without lateral tracheae (Fig. 47).

Etymology. The species is named for Peter Dostine, who

collected the type material and associated nymphs with adults.

Remarks. This species was not included in the original keys to

mayflies of the Alligator Rivers region presented by Suter

(1992). The nymph has previously been designated

Tillyardophlebia sp.AV8 (Dean, 1999), albeit with a comment

that the species should perhaps be placed in a new genus. While

we have opted to retain the species in Tillyardophlebia pending

completion of further study of two undescribed Queensland

species and a detailed phylogenetic analysis, we are of the

opinion that these three northern Australia species will event-

ually be transferred to a new genus. The nymphs of the species

from northern and southern Australia are very similar, with no

obvious characters to justify generic separation. In the imagos,

however, there are clear differences in the structure of the male

genitalia. The species from south-eastern Australia form a

monophyletic group with long, narrow and widely separated

penes lobes, each lobe bearing a large curved ventral spine

( Tillyardophlebia sensu stricto). Species from northern

Australia have more robust penes lobes, which are fused either

in the basal third or along most of their length, and each lobe

has a single subapical spine.

Although confirmed material of this species has only been

collected from the type locality, it is probably more widely

distributed. We have examined nymphs from Manning Gorge

in north-western Australia which we believe are conspecific.

118

J. C. Dean and P. J. Suter

Acknowledgements

We would like to thank management and staff at the Alligator

Rivers Research Institute, who provided encouragement and

material assistance during several field trips to the region. In

particular we would like to single out Peter Dostine and Chris

Humphrey; we greatly appreciate their friendship, hospitality

and the valuable time and knowledge they were able to provide.

Alice Wells, Peter Cranston and John Hawking are thanked for

assistance in the field.

References

Dean, J.C. 1999. Preliminary keys for the identification of Australian

mayfly nymphs of the family Leptophlebiidae. Cooperative

Research Centre for Freshwater Ecology, Identification Guide No.

20. 91pp.

Ganambarr, M. 1999. Datiwuy. In: Thieberger N. and McGregor W.

(eds), The “Macquarie Aboriginal Words”. Macquarie Library:

Sydney.

Peters, W.L., Peters, J.G., and Edmunds, G.F. 1978. The

Leptophlebiidae of New Caledonia (Ephemeroptera). Part I

Introduction and systematics. Cahiers d’ ORSTOM, Serie

Hydrobiologie 12: 97-117

Suter, PJ. 1992. Taxonomic key to the Ephemeroptera (Mayflies) of

the Alligator Rivers Region, Northern Territory. Open File Record

No. 96, Supervising Scientist for the Alligator Rivers Region.

Suter, PJ. 1997. Preliminary guide to the identification of nymphs of

Australian baetid mayflies (Insecta: Ephemeroptera) found in flow-

ing waters. Cooperative Research Centre for Freshwater Ecology,

Identification Guide No. 14. 36 pp.

Suter, PJ. 1999. Illustrated key to the Australian caenid nymphs

(Ephemeroptera: Caenidae). Cooperative Research Centre for

Freshwater Ecology, Identification Guide No. 23. 36 pp.

129 > Dimorphic brooding zooids in the genus Adeona Lamouroux from Australia

(Bryozoa: Cheilostomata)

Philip E. Bock and Patricia L Cook

135 > A review of Australian Conescharellinidae (Bryozoa: Cheilostomata)

Philip E. Bock and Patricia L Cook

183 > A review of the Tertiary fossil Cetacea (Mammalia) localities in Australia

Erich M. G. Fitzgerald

209 > A new Late Eocene cassiduloid (Echinoidea) from Yorke Peninsula, South Australia

Francis C. Holmes

217 > A new species of Quinquelaophonte (Crustacea: Copepoda: Harpacticoida: Laophontidae) from

Port Phillip Bay, Victoria, Australia

Genefor K. Walker-Smith

Memoirs of Museum Victoria

Volume 61 Number 1 2004

1 > A molecular and morphological revision of genera of Asterinidae (Echinodermata: Asteroidea)

P. Mark O’Loughlin and Jonathan M. Waters

41 > A new genus of millipedes (Diplopoda: Polydesmida: Dalodesmidae) from wet forests in southern

Victoria, with brief remarks on the Victorian Polydesmida

Robert Mesibov

47 > Biosystematics of Australian mygalomorph spiders: descriptions of three new species of Teyl

from Victoria (Araneae: Nemesiidae)

Barbara York Main

57 > Chirostylidae from north-western Australia (Crustacea: Decapoda: Anomura)

Shane T. Ahyong and Keiji Baba

65 > Sicafodiidae, fam. nov. for Sicafodia stylos, gen. nov., from the marine bathyal of south-eastern

Australia (Crustacea: Amphipoda: Gammaridae)

Jean Just

75 > Pseudidotheidae (Crustacea: Isopoda: Valvifera) reviewed with description of a new species,

first from Australia

Gary C. B. Poore and Tania M. Bardsley

85 > The long-horned caddisfly genus Oecetis (Trichoptera: Leptoceridae) in Australia: two new species

groups and 1 7 new species

Alice Wells

111 > Descriptions of new species and a new genus of leptophlebiid mayflies (Insecta: Ephemeroptera)

from the Northern Territory, Australia

J. C. Dean and P. J. Suter

Volume 61 Number 2 2004

121 > Mitochondrial 1 2S rRNA sequences support the existence of a third species of freshwater blackfish

(Percicthyidae: Gadopsis) from South-eastern Australia

Adam D. Miller, Gretchen Waggy, Stephen G. Ryan and Christopher M. Austin

Continued inside back cover >

Contents