Memoirs of Museum Victoria 68:1-28 (2011)

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

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

A revision of the Australian fossil species of Zoila (Gastropoda: Cypraeidae)

Thomas A. Darragh

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

Abstract Darragh, Thomas A. 2011. A revision of the Australian fossil species of Zoila (Gastropoda: Cypraeidae). Memoirs of

Museum Victoria 68: 1-28.

Zoila Jousseaume, 1884 is a cowry genus, the species of which are now confined to the coastal waters of Western

Australia and southern Australia. Six fossil species are known from southeast Australia, of which three are new, ranging

in age from late Oligocene to middle Miocene, known from the Murray, Otway and Bass basins. Three fossil species are

known from Western Australia, of which three are new, ranging in age from late Eocene to late Pliocene, known from the

Carnarvon, Bremer and Eucla basins. Umbilia (Gigantocypraea) Schilder 1927 (type species Cypraea gigas McCoy) is

regarded as a synonym of Zoila. The earliest record of the genus is Z. chathamensis (Cemohorsky, 1971), Paleocene/early

Eocene, Chatham Islands, probably ancestral to the Australian fossils. Zoila is closely related to Cypraeorbis Conrad,

1865, of which Bernaya Jousseaume, 1884 is regarded as a synonym. Living species are known to have no planktotrophic

larval stage, so there is considerable variation in species morphology. Such direct development arose in these cowries at

least as early as the late Eocene. Fossils described here are Zoila viathomsoni sp. nov., Z. didymorhyncha sp. nov., Z.

glomerabilis sp. nov., Z. dolichorhyncha sp. nov., Z. mulderi (Tate, 1892), Z. platypyga (McCoy, 1876) (= consobrina

McCoy, toxorhyncha Tate, platypyga simplicior Schilder), Z. gigas (McCoy, 1867) (= dorsata, gabrieli ), Z. campestris n.

sp. and Z.fodinata n. sp.

Keywords Gastropoda, Cypraeidae, cowries, Australia, Tertiary, new species, taxonomy

Introduction

Zoila is a member of a group of Australian cowries that

includes Notocypraea, Umbilia and Austrocypraea, the

members of which all seem to have direct larval development

rather than having a planktotrophic larval stage as in most

species of cowry (Wilson, 1985; Wilson and Clarkson, 2004).

This paper reports research results that continue from a study

of the Australian endemic genus Umbilia (Darragh, 2002).

Subfamily classification

The genus Zoila is usually placed in the subfamily Bernayinae

Schilder, 1927 erected for the fossil genus Bernaya Jousseaume,

1884, of which Zoilinae Iredale, 1935 is a synonym (Wilson

and Clarkson, 2004). Meyer (2003), in his work on molecular

systematics of cowries, retained most of the subfamilies

previously used by many authors on the family, but did not

place the genus Zoila in any subfamily (although his Table 3

could imply a placement in the subfamily Bernayinae).

However, following further molecular work, he subsequently

(Meyer, 2004) placed the genera Zoila and Barycypraea

within the subfamily Bernayinae. Cypraeorbinae Schilder,

1927 was erected for the fossil genus Cypraeorbis Conrad,

1865.1 consider that Bernaya is a synonym of Cypraeorbis, as

argued below. Therefore, if subfamilies are regarded as useful

in classification, Cypraeorbinae should be used instead.

Bouchet and Rocroi (2005) included the subfamilies

Bernayinae and Cypraeorbinae, along with three other

subfamilies, within the synonymy of Gisortinae Schilder,

1927, erected for the genus Gisortia Jousseaume, 1884. Species

of Gisortia have an obsolete fossula, quite different from those

of species of Cypraeorbis and Zoila, so this synonymy is

unlikely.

Nomenclatural history of Zoila fossil species

The genus Zoila was established by Jousseaume (1884a) for

three species of large living cowries from Western Australia.

The first fossil species later assigned to this genus were

described by Frederick McCoy from the Tertiary of Victoria.

Cypraea gigas was described by McCoy in 1867 without

illustration and in 1875, it was redescribed, figured and placed

in the subgenus Aricia. Two more species, Cypraea (Aricia)

platypyga and C. (A.) consobrina were described and figured

by McCoy in 1876 and 1877, respectively. In 1890, Tate revised

these three species and added another two species, C.

toxorhyncha, regarded here as a synonym of Z. platypyga, and

Cypraea dorsata, regarded here as a synonym of Z. gigas. In

1892, Tate figured a new species C. mulderi and described it in

1893, pointing out its affinities with C. platypyga. Tate

assigned all these species to Cypraea. Harris (1897) assigned

T.A. Darragh

C. gigas and C. platypyga to the subgenus Erosaria, which he

regarded as a senior synonym of Aricia. In 1912, Chapman

described Cypraea gabrieli, regarded here as a synonym of Z.

gigas.

Schilder (1927) erected Gigantocypraea as a subgenus of

Umbilia with C. gigas McCoy as the type species. He also

placed the other species mentioned by Tate in Umbilia.

However, in 1930, Schilder raised Gigantocypraea to generic

status and referred the other species to Zoila, but subsequently

(Schilder, 1935) reduced Gigantocypaea to a subgenus of

Zoila and added anew subspecies, Z. (Z.)platypyga simplicior,

regarded here as a synonym of Z.platypyga.

Time range and distribution

Living species of Zoila are known only from the southern and

western coasts of Australia. There are seven recognised

species, with distribution ranging from Apollo Bay in Victoria

westwards across the southern coast and up the west coast of

Western Australia as far north as the West Kimberley district

(Wilson and Clarkson, 2004). Fossil species of Zoila are more

widespread. The earliest known species, recognised by this

author, occurred in the Paleocene to early Eocene of the

Chatham Islands, New Zealand. This species may have given

rise to younger Australian species. The earliest Australian

species Z. viathomsoniae sp. nov., recorded from the upper

Eocene of southwest Western Australia, is probably an

ancestor of the younger Australian species of Zoila, of which

there are two groups: an eastern and a western group. The

eastern group is known from southeast Australia, and ranged

in age from late Oligocene to middle Miocene. This group is

characterised by having prominently developed anterior and

posterior canals and does not seem to be ancestral to any

living species. The western group is known from Victoria,

Western Australia, Indonesia and India, and ranged in age

from middle Miocene to Recent.

The oldest known member of the western group occurred

in the middle Miocene of Western Australia. It seems to be

ancestral to Z. campestris sp. nov. from the Pliocene of

Western Australia. This species is probably ancestral to some

of the living species, in particular Z. venusta (Sowerby, 1846).

Species somewhat similar to the Western Australian species

are found in the Neogene of Indonesia and India. A single

species, known from one fragmentary specimen, occurred in

the late Miocene of Victoria.

Affinities

Generic distinction among cowries is subject to much controversy.

Few features of the shell can be used to characterise species

groups, particularly without knowing their anatomy and

molecular biology. Even when this information is available, it is

not always possible to apply this knowledge to include

unequivocally fossil species in groups composed of living

species. The fossula is one shell structure that does seem to vary

sufficiently to be able to use it to some extent in supraspecific

classification, and link fossil and living species.

The Australian fossil cowries treated here have a type of

fossular morphology similar to that found in species assigned to

four nominal genera: Cypraeorbis Conrad, 1865; Bernaya

Jousseaume, 1884; Zoila Jousseaume, 1884; and Barycypraea

Schilder, 1927. Of these four, only Zoila is based on a living

species, and hence can be characterised on the basis of anatomy

and genetics, although Barycypraea — based on a fossil species

— has living representatives. The close morphological

relationship between the first three genera was first recognised by

Schilder (1926), who placed Zoila as a subgenus of Cypraeorbis

and synonomised Bernaya with Cypraeorbis. Later, as his

subdivisions of the Cypraeidae became smaller, and more genera

were recognised on the basis of minor differences, Schilder

(1941) placed the three genera in his subfamily Cypraeorbinae,

with Zoila and Bernaya in the Tribe Bemayini, and Cypaeorbis

in the Tribe Cypraeorbini. Pol in (1991) regarded Zoila as a

possible subgenus of Cypraeorbis in the subfamily Bemayinae.

In the present study, specimens of the type species of each of

these genera, as well as some other species assigned to them,

have been examined to determine possible affinities.

Cypraeorbis, type species Cypraea sphaeroides Conrad,

1848, was based on a species occurring in the lower Oligocene of

the southwest United States of America. The genus ranges in age

from late Eocene to at least early Miocene. The fossula (fig. IB)

of a C. sphaeroides specimen from Byram, Mississippi (USNM

498351) is deeply excavated and bounded by a prominent lateral

ridge on the anterior side. Posterior to this lateral ridge is a slight

notch in the fossula margin. The lateral ridge merges anteriorly

with a very weak terminal ridge. Between the first columellar

tooth and the terminal ridge there is a wide depression, which

extends as a sulcus parallel with the terminal ridge into the

fossula. The fossula (figs 1C,3E-F) of a Cypraeorbis ventripotens

(Cossmann, 1903) specimen — a species closely related to, if not

synonymous with C. sphaeroides (MacNeil and Dockery, 1984)

— from the upper Eocene, Town Creek, Jackson, Mississippi

(Mississippi Geological Survey collection) is large,

subrectangular, concave, with a prominent notch at the anterior

end of the thickened interior edge. The terminal ridge, like that of

C. sphaeroides, is weakly developed and runs into the interior of

the aperture forming the anterior edge of the fossula as a

prominent ridge. A well developed groove or sulcus between the

first columellar tooth and the terminal ridge runs parallel to the

terminal ridge into the fossula. Specimens of Cypraea willcoxi

(Dali, 1890) and Zoila arlettae Dolin, 1991 from the Early

Miocene Chipola Formation of Farley Creek, Florida (MNHN,

Paris) were also examined. Z. arlettae Dolin has a fossula (fig.

3N) very similar to C. sphaeroides and C. ventripotens, though

larger and with a broader notch. Cypraea wilcoxi Dali is very

close in morphology to Z. arletti and also has a fossula (fig. 30)

similar in structure to Cypraeorbis sphaeroides and C.

ventripotens, hence both Cypraea arlettae and Zoila wilcoxi

seem to be better placed in Cypraeorbis rather than in Zoila as

suggested by Dolin (1991).

Petuch (2004) erected two new genera in the subfamily

Cypraeorbinae, Floradusta (type species Cypraea heilprini Dali,

1890, early Miocene, Florida) and Loxacypraea (type species

Cypreaea chilona Dali, 1900, early Miocene, Florida). He

included in these genera several species that had been included

by Dolin (1991) in Cypraeorbis, Zoila, Siphocypraea and

Erronea (Adusta). Petuch stated that a fossula was absent in

A revision of the Australian fossil species of Zoila (Gastropoda: Cypraeidae)

species of the two genera. This statement is incorrect, as shown

by the descriptions and illustrations in Dolin and by my own

observations of two of the species, Cypraeorbis arlettae and C.

wilcoxi (see figs 3H-J, N, K, O). Revision of American taxa is

beyond the scope of this paper, but I consider that Floradusta and

Loxacypraea are synonyms of Cypraeaorbis, as the type species

of each have a fossula similar to C .sphaeroides and C.

ventripotens. Some of the other species included in Floradusta

by Petuch (2004) are unlike species of Cypraeorbis in that the

fossula is crossed by extensions of the columellar teeth and

therefore cannot belong in the Cypraeorbinae. These species are

best left in the genera to which they were assigned by Dolin

(1991).

Bernaya, type species Cypraea media Deshayes, 1835, was

based on a species from Valmondois in the Paris Basin, Sables

Moyens, Bartonian (late Eocene). The differences between

Cypraeorbis and Bernaya are very slight. The fossula of

specimens of Bernaya media (Deshayes) (figs 1A, 4B, F) from

Bois du Roi, Auvers (MNHN), is large, subrectangular and

concave. On the interior side, it is bounded by a prominent lateral

ridge, in which there is a prominent anterior notch. The terminal

ridge of the anterior canal extends into the interior of the aperture

and merges with the lateral ridge of the fossula. A well-developed

groove adjacent to the terminal ridge runs down into the fossula.

This groove is partly obstructed by a columellar tooth, which

merges with the terminal ridge. Specimens of Cypraea

bartonensis Edwards, 1854 from Barton, Hampshire, England

(NMV P310165-6) have a similarly structured fossula, but there

are no columellar teeth obstructing the groove. In this respect, it

closely resembles Cypraeorbis ventripotens. A specimen of

Cypraea splendens Grateloup, 1827, from the Rupelian of Gaas,

Landes, France (NMV P310168) also has a fossula identical to C.

ventripotens.

MacNeil and Dockery (1984) described the protoconch of C.

sphaeroides as having five whorls, some of which are sculptured

with a fenestrate pattern. Dolin and Dolin (1983) have described

and figured (figs 3 and 4) the protoconch of Bernaya media as

having four whorls, which have a fenestrate sculpture on the last

two whorls. This form of protoconch suggests a planktotrophic

larval stage. As there are no significant distinguishing features

between the two genera, it seems appropriate to synonymise

Bernaya with Cypraeorbis.

The type species of Zoila is Cypraea friendii Gray, 1831,

from the waters off Fremantle, Western Australia. Specimens of

this species have a large spoon-shaped, deeply concave fossula

(Wilson and Clarkson, 2004, pi. 47). The terminal ridge is not

prominent, if at all present (on some specimens it is merely

represented by a blunt thickening), and there is no prominent

ridge bounding the edge of the fossula. There is also no groove

running from the ventral surface down into the fossula, though

there may be a gap between the labial teeth representing such a

groove. There is no notch on the internal edge of the fossula. In

these respects, it is unlike the American and European species;

however, other species of Australian Zoila do have these features,

but the edge of the fossula is not markedly notched, rather,

merely indented (figs 1E-G).

The fossula in Zoila chathamensis (figs 1H, 5I-J) is weakly

developed, smooth, broad, very slightly concave, slightly

projecting, and bounded by a very weak terminal ridge. It

resembles the fossula in Z. didymorhyncha sp. nov. Specimens of

Zoila viathomsoni sp. nov. have a well-developed fossula (figs

ID, 5L) bounded by a prominent ridge and with a very weak

indentation towards the anterior end. There is a very shallow

sulcus running into the fossula. Specimens of Zoila platypyga

(McCoy) have a fossula somewhat similar to Z. friendii but the

edge of the fossula is more ridge-like. There is no sulcus

extending into the fossula parallel to the terminal ridge.

Specimens of Z. didymorhyncha sp. nov. (figs IE, 6C) have a

weak terminal ridge merging with the edge of the fossula and a

shallow sulcus running parallel to it into the fossula. None of the

Australian fossils have the well-defined fossular margin or the

well-developed notch behind it that is present in the European

and American species of Cypraeorbis.

Unlike the protoconchs of Cypraeorbis species, the

protoconchs of species of Zoila, where known, are paucispiral,

slightly tilted and smooth (Ranson, 1967, pi. 6, figs. 1-3; Wilson,

1998). The protoconch of Z. platypyga consists of one whorl

slightly tilted from the axis of the shell. The protoconch of Z.

gigas (McCoy) consists of two whorls, the first of which is

irregular and granulated (figs 13E-F). In cross-section (figs

2A-C), the protoconchs of Z. viathomsoni, Z. platypyga and Z.

gigas resemble cross-sections of protoconchs of Umbilia species

(Darragh, 2002, fig. 1). This morphology suggests a protoconch

formed from a partly chitinous whorl. Wilson (1985) has shown

that living species of Zoila have direct development. The

protoconchs of the fossils suggest that the fossils also had direct

development and that the change from planktotrophic

development to direct development had taken place by the late

Eocene. Unfortunately, the protoconch of the earliest known

species, Z. chathamensis, is unknown. Zoila species have direct

larval development like the other Australian cowries referred to

the genera Austrocypraea Cossmann, 1903, Umbilia Jousseaume,

1884 and Notocypraea Schilder, 1927. The last three genera are

endemic to Australia and do not have a distribution in shallow

tropical seas, whereas species of Zoila are known from tropical

as well as temperate waters and also have been found in the lower

Tertiary of New Zealand, and the upper Tertiary of Indonesia and

India. Austrocypraea, Umbilia and Notocypraea seem to be

representatives of an ancient southern group, whereas the

distribution of Zoila suggests that its origin may have been

Tethyan. Most tropical cowries are known to have planktotrophic

larvae, which accounts for their widespread distribution in the

Indo-Pacific realm. Zoila seems to have evolved from the

widespread genus Cypraeorbis, whose species almost certainly

had planktotrophic larvae, although the occurrence of a species

of Zoila in the Chatham Islands as early as the Paleocene to early

Eocene may suggest a southern origin.

Though the differences between the genera are slight, Zoila

seems sufficiently different from Cypraeorbis to be maintained

for the present. Wilson and Clarkson (2005) suggested that Zoila

evolved in Asian waters from a directly developing Tethyan

ancestor of the subfamily Bernayinae and migrated to Australia

in Miocene time. However, the occurrence of species in Australia

in the late Eocene and in the New Zealand region in the

Paleocene/early Eocene suggests that Zoila evolved much

earlier, probably from Cypraeorbis or some other closely related

T.A. Darragh

taxon, such as Protocypraea Schilder, 1927, in the earliest

Tertiary or even in the late Cretaceous. Species of Cypraeorbis

(recorded as Bernaya ) and Protocypraea have been recorded

from the upper Cretaceous of North America by Groves (1990,

2004) and Protocypraea is also known from the upper

Cretaceous of India and the Paleocene of Pakistan (Cox, 1930).

Schilder (1927) erected Barycypraea as a sub genus of Zoila

with type species Cypraea (Aricia) caputviperae Martin, 1899

(probably a synonym of Cypraea murisimilis Martin, 1879) for

a group of cowries known from the upper Tertiary of Indonesia

and living in the Indian Ocean. Molecular biological studies of

cowries by Meyer (2004) showed that the nearest living relatives

of the Australian living Zoila species are the western Indian

Ocean species of Barycypraea and he grouped both genera in

the subfamily Bemayinae. He included no other living taxa in

this subfamily. Species of Barycypraea have a similar fossula

(fig. 3L) to Zoila species and they are known to be spongivores

like Zoila (Liltved, 1989). Species of Barycypraea occur in

South Africa, Pakistan, India and Indonesia, and range in age

from Miocene to Recent (Kay, 1990). The morphology and

biogeography of Barycypraea suggests an origin similar to that

of Zoila and a probable common ancestry.

Terminology and repositories

Measurements are given as follows: L = total length of shell;

W = width of shell; H = height of shell. Tooth counts are cited

as LT = labral teeth and CT = columellar teeth. The terms left

and right refer to the animal’s true left and right sides,

respectively.

All material used in this study, unless otherwise stated, is

held in the collections of Museum Victoria, registration

numbers with prefixes P (invertebrate palaeontology

collection) and F (living mollusca collection). Localities are

cited where possible using the Museum Victoria fossil locality

Museum acronyms used for other material are as follows:

BM(NH) = Natural History Museum, London; SAM T = Tate

Collection, South Australian Museum, Adelaide; AMNZ =

Auckland Museum, New Zealand; AM C = Australian

Museum, Sydney, New South Wales; WAM = Western

Australian Museum; USNM = Smithsonian Institution,

National Museum of Natural History; MNHN = Museum

national d’Histoire naturelle, Paris; GNS TM = GNS Science

(New Zealand).

All specimens figured were coated with ammonium

chloride for photography, unless stated otherwise.

Class Gastropoda

Family Cypraeidae Rafinesque, 1815

Subfamily Cypraeorbinae Schilder, 1927

Zoila Jousseaume, 1884

Zoila Jousseaume, 1884a: 14.— Jousseaume, 1884b:89.—

Cossmann, 1903:146,149.- Thiele, 1929:275.- Schilder, 1935:336.-

Schilder and Schilder, 1939:173.—Wenz, 1941:971.— Dolin, 1991:11

(synonomised Gigantocypraea Schilder, 1927). — Wilson and

Clarkson, 2004:44.

Cypraeorbis (Zoila) Jousseaume. Schilder, 1926:378.

Umbilia (Gigantocypraea) Schilder, 1927:86 (type species,

original designation, Cypraea gigas McCoy, 1867, Miocene, Victoria).

Zoila (Zoila) Jousseaume. Schilder, 1935:337.— Schilder,

1939:177.

Zoila (Gigantocypraea) Schilder. Schilder, 1935:337.— Schilder,

1939:177.- Wenz, 1941:971.

Cypraea (Zoila) Jousseaume. Wilson and McComb, 1967:469. —

Wilson, 1993:192.

Type species. Cypraea scottii Broderip, 1832 (= Cypraea

friendii Gray, 1831) by subsequent designation Jousseaume

(1884b, p. 89), western and southern Australia.

Diagnosis. Shell small (35 mm) to very large (247 mm) in size,

highly glazed, varying in shape, elongate-ovate or pyriform or

globose, ventrally flattened. Spire whorls usually covered in

callus, rarely protruding. Protoconch, where known, consisting

of one to two whorls, the first whorl large, somewhat irregular

in shape, deviated slightly from shell axis. Anterior and

posterior canals deeply incised, usually short, but on some

species produced and bounded by weak to prominent lateral

flanges. Aperture narrow, sinuous with weakly to strongly

developed short to elongate teeth not extending into aperture or

onto fossula. Fossula weakly to strongly developed, broad,

slightly to deeply concave and bounded anteriorly by a weak to

strong terminal ridge; on some species with very weak sulcus

parallel and immediately posterior to terminal ridge.

Time range. Late Paleocene/early Eocene-Recent.

Distribution. India (early Miocene), Indonesia (Pliocene-

Pleistocene), Western Australia (late Eocene, Miocene-

Recent), South Australia (Miocene, Recent), Victoria (late

Oligocene-late Miocene, Recent), Tasmania (early Miocene),

New Zealand (late Paleocene/early Eocene).

Remarks. Specimens of Zoila itoigawa Tomida, 1989 from the

late Miocene of Japan are poorly preserved and have not been

prepared sufficiently to enable generic assignment.

As I have recognised two groups of species, an eastern and

a western group, it could be argued that these should be

accorded taxonomic status if there are significant

morphological differences to separate them, in which case

Gigantocypraea could be used for the eastern group. However,

I regard these groups as more geographic entities. There are

some morphological differences, but I regard them as minor

and not of sufficient importance to use as generic characters.

Such variations in morphology are to be expected in species

that have no pelagic larval stages.

Some species of the eastern group have considerably

produced anterior and posterior canals — for example, Zoila

platypyga (McCoy) and Z. mulderi (Tate) — but other species

have canals similar to the living species of the western group

— for example, Z. glomerabilis n. sp. is similar to Z. venusta

(Sowerby, 1846) or Z. gendinganensis (Martin, 1899).

Columellar dentition varies considerably from species to

species. In the living species, columellar dentition can vary

from strong to weak even within the one species; for example,

A revision of the Australian fossil species of Zoila (Gastropoda: Cypraeidae)

Zoila friendi (Wilson and Clarkson, 2004, pp. 79-80). Fossil

species of the western group all have prominent columellar

dentition. In Z.fodinata, the columellar teeth are elongated to

form short ridges. In the eastern group, columellar dentition is

more highly variable. In Z.platypyga, the columellar dentition

is in the form of strong, elongate ridges, whereas in Z. mulderi

(Tate) (almost certainly ancestral to the former), the columellar

dentition is more like that of the living species, as is the

dentition of Z. glomerabilis sp. nov. and Z. dolichorhyncha sp.

nov. In Z. gigas, there is no columellar dentition. I do not

consider that columellar dentition can be used to provide a

consistent taxonomic character to separate the eastern and

western group of species of Zoila as implied by Wilson and

Clarkson (2004, p. 49).

Fossular morphology also varies somewhat, though the

basic pattern throughout all the species is the same — that is,

broad, concave and bounded by the terminal ridge. Even

within a species, fossular morphology can vary. Compare, for

example, the fossulae of specimens of Z. venusta figured on

plates 265-276 of Wilson and Clarkson (2004).

Zoila chathamensis (Cernohorsky, 1971)

Figures 1H, 5A, D-G, I-K

Bernaya chathamensis Cernohorsky, 1971, p. 117, fig. 13.

Description. Shell of small size for genus (36 mm), solid,

somewhat globose to subpyriform, dorsal surface uniformly

convex, ventral surface weakly convex, almost flat. Posterior

canal narrow, barely projecting, deeply incised; anterior canal

very short, deeply incised. Spire not visible. Outer lip with 14-

lb teeth, columella lip with 12-18 teeth. Fossula well developed,

smooth, broad, very slightly concave, slightly projecting,

bounded by very weak terminal ridge.

Dimensions.

Holotype AMNZ TM-1325

35.8

27.3

21.1

18.0

14.0

Figured specimen GNS TM8792

36.5

21.0

12.0

16.0

Type locality. Flowerpot Harbour, Pitt Island, Chatham Islands,

New Zealand, Red Bluff Tuff, late Paleocene/early Eocene.

Occurrence and material. Pitt Island, New Zealand: Red Bluff

Tuff, Flowerpot Harbour (holotype AMNZ TM-1325), (GNS

GS12159, one specimen); Red Bluff Tuff, GS12173 Rocky side,

Tarawhenua Peninsula (GNS TM8792, one specimen).

Remarks. Cernohorsky (1971) assigned his new species to the

genus Bernaya, apparently not realising that the fossula of

species assigned to that genus has a deep groove at the anterior

end. His description of the fossula is quite accurate, but his figure

does not show the morphology of the fossula very well, as it was

not fully prepared. The holotype is refigured here, but the fossula

is not very clear in the illustration (fig. 51). Another specimen

kindly provided by Dr Alan Beu has been more fully prepared

and although the aperture is narrow, making photography,

difficult the simple nature of the fossula can be seen (fig. 5J). The

fossula is somewhat similar to that in Z. viathomsoni n. sp. and Z.

decipiens (EA. Smith, 1880), but is not as concave.

Although much smaller, the overall morphology of this

species is somewhat similar to that of Zoila friendii thersites

(Gaskoin, 1849). The fossula is relatively shorter and is not as

deeply concave as that of the latter species, but is otherwise

similar.

Zoila viathomsoni sp. nov.

Figures ID, 2C, 5B-C, H, L

Description. Shell of small size for genus (19-28 mm in length),

pyriform. Spire not visible on most specimens, projecting on

one specimen. Posterior canal very short, slightly bent to left.

Anterior canal very short, slightly deflected to right. Aperture

slightly sinuous; outer lip with about 23-25 teeth present along

entire lip; columella with about 23-26 teeth present along

entire lip. Fossula well-developed, deep, elongate, projecting,

bounded anteriorly by thickened ridge; very weak notch present

in inner edge just posterior to anterior ridge; weak terminal

ridge joining edge of fossula. First columellar tooth adjacent to

terminal ridge, almost blocking shallow sulcus that extends

into fossula parallel to terminal ridge.

Dimensions.

Holotype

WAM

72.296

24.0

16.0

12.5

25.0

26.0

undistorted

Paratype

WAM

72.253

27.0

15.0 est.

14.0

est.

c24.0

24.0

distorted

Paratype

P310193

28.0

15.0

13.0

est.

24.0

23.0

crushed

laterally

Paratype

P310194

17.0

11.7 fragment showing fossula

est. = estimated

Type locality. PL3171 gravel scrape beside Thomson Highway,

23.5 km north of Highway 1, north of Walpole, Western

Australia. Grid ref. Deep River (1:100 000 series) 743487.

Pallinup Siltstone, Aldingan, Late Eocene.

Type material. Western Australian Museum: Holotype WAM

72.296, collected T.A. Darragh and G.W. Kendrick, 12-14

March 1969; paratype WAM 72.253, collected G.W. and W.E.

Kendrick, 25 January 1969. Museum Victoria: Paratype

P310193, collected T.A. Darragh & G.W. Kendrick, 12-14

March 1969.

Time range. Aldingan, Late Eocene.

Occurrence and material. Type locality only (one undistorted

specimen, four complete distorted specimens, nine fragments).

Remarks. This is one of the two oldest true cowries recorded

from Australia and occurs only in southwest Australia,

associated with Willungia ovulatella (Tate) and Semitrivia

pompholugota (Tate). Specimens from the type locality are

preserved as silica replacements weathered out of the Pallinup

T.A. Darragh

Figure 1. A, Cypraeorbis medius (Deshayes, 1835), x 5; B, Cypraeorbis sphaeroides (Conrad, 1848), x 5; C, Cypraeorbis ventripotens (Cossmann,

1903), x 5; D, Cypraeorbis viathomsoni sp. nov. WAM 72.296, x 5; E, Zoila didymorhyncha sp. nov. P302687, x 2.6; F, Zoila carnpestris sp. nov.

WAM 89.437, x 2.6; G, Zoila fodinata sp. nov. WAM 89.637, x 3; H, Zoila chathamensis (Cernohorsky, 1971) GNSTM 87922, x 2.3. Arrows

indicate the location of the notch.

A revision of the Australian fossil species of Zoila (Gastropoda: Cypraeidae)

Figure 2. Protoconchs. A, Zoilaplatypyga (McCoy, 1876), P308781, x 7; B, Zoila gigas (McCoy, 1867), P308792, x 3.6; C, Zoila viathomsoni sp.

nov., WAM x 10.

Siltstone. The associated fauna comprises large numbers of

siliceous sponges. The protoconch observed in cross-section

(fig. 2C), is paucispiral and slightly deviated from the axis of the

teleoconch, unlike the multispiral protoconchs of Cypraeorbis

species. The general size and shape of the shell resembles that

of Cypraeorbis medius, C. bartonensis and C. ventripotens;

however, the fossula (fig. 5L) is not the same as that present in

these species. The fossula is very wide and prominent, projecting

into the aperture. It is more elongate than that of these three

species, does not have a prominent notch and the anterior border

is not thickened. In general shape and concavity, it resembles

those in Zoila friendii and Z. decipiens. Zoila viathomsoni

bears a close resemblance to Z. chathamensis (Cernohorsky),

late Paleocene to early Eocene, Chatham Islands, New Zealand,

but it is not as tumid, has much finer and more numerous

apertural teeth, the posterior labral border of the posterior canal

is more produced posteriorly and it is only about two-thirds the

size. Given the similarities, Z. chathamensis is a possible

ancestor of Z. viathomsoni sp. nov.; Z. chathamensis is the

earliest known representative of the genus.

Zoila viathomsoni bears some resemblance in size and

shape to Lyncina (Austrocypraea) onkastoma Yates, 2009,

recorded from the early Oligocene of South Australia, but

lacks the well-defined anterior rostrum of L. (A.) onkastoma,

as well as the prominent notch on the fossula.

Etymology. Latin via road.

Zoila didymorhyncha sp. nov.

Figures IE, 6A-E, I

Description. Shell solid, small for genus, elongate, subfusiform;

base relatively flat, but rounded on either side of aperture. Spire

not visible. Posterior canal long, with rounded sides, sunk into

massive rounded posterior rostrum formed by extensions of

inner and outer lips. Anterior canal long, with rounded sides,

sunk into a thick, rather flat rostrum. Aperture somewhat

sinuous; outer lip with 10-24 well-developed teeth extending

along entire lip; teeth completely obsolete on one specimen.

Columella lip with 3-26 well-developed teeth extending along

entire lip on most specimens; teeth obsolete on one specimen.

Fossula moderately developed, very shallow, bounded

anteriorly by a small, weak, terminal ridge.

Dimensions.

Holotype

P302687

Paratype

P302685

P302686

64+

22+

25 anterior

canal broken

Type locality. PL3022 cliff section at Addiscot Beach, beds

B109-107, southwest of small gully, clay overlying Demons

Bluff Formation, Victoria. AMG Torquay BT619490. Jan Juc

Formation. Early Janjukian.

Type material. Holotype P302687, collected T.A. Darragh, 4

December 1985; paratype P302685, collected T.A. Darragh 8

May 1990.

Time range. Early Janjukian, Late Oligocene.

Occurrence and material. PL3022 Addiscott Beach (three

specimens).

T.A. Darragh

Figure 3. A-G, Cypraeorbis ventripotens Moodys Branch Formation, late Eocene, Town Creek, Jackson, Mississippi, United States of America;

A-C, x 2; F, x 4; D-E, G, x 2; H-J, N, Cypraeorbis arlettae MNHN, Chipola Formation, Burdigalian, early Miocene, Farley Creek, Calhoun

County, Florida, USA; K, O, Cypraeorbis wilcoxi MNFIN, Chipola Formation, Burdigalian, early Miocene, Farley Creek, Calhoun County,

Florida, United States of America; L-M, Barycypraea zietsmani Liltved and Le Roux, 1988, P31664, Alexandria Formation, late Neogene,

Eastern Cape, South Africa, x 1.

A revision of the Australian fossil species of Zoila (Gastropoda: Cypraeidae)

Figure 4. A-F, Cypraeorbis medius MNHN, Bartonian, late Eocene, Bois du Rois, Auvers sur Oise, France; A-C, D-E, x 1.5; F, x 2.

Remarks. The small size and the massive development of the

anterior and posterior rostra separate this species from all others

in the genus. In morphology, it comes closest to Zoila mulderi

(Tate), but it has a massive anterior rostrum lacking in Z. mulderi

and the posterior rostrum is not notched as it is in Z. mulderi.

Etymology. Greek, didymos, double; rhynchos, snout.

Zoila glomerabilis sp. nov.

Figures 6F-H, 7A-F

Description. Shell of large size for genus, solid, globosely

pyriform, abruptly contracted anteriorly to short, narrow

anterior canal. Spire projecting slightly, almost entirely

enveloped by last whorl, covered with thick callus. Posterior

canal short, deeply incised, reflexed to the left. Anterior canal

deeply incised, with rounded sides, reflexed dorsally, supported

laterally by very short, narrow flanges. Aperture sinuous,

slightly widened and sloping steeply inwards before anterior

canal. Outer lip with 28-36 teeth extending along entire lip;

inner lip with 15-32 teeth extending along entire lip on most

specimens; weaker posteriorly on some specimens and on few

specimens extending only along half of lip. Fossula well

developed, broad, relatively deeply depressed, bounded

anteriorly by weak terminal ridge.

Dimensions.

Holotype PI4515

Paratype P308740

Paratype P315526

Type locality. Lower beds of section south of Fishermen’s

Steps, Bird Rock Cliffs, Torquay, Victoria. Jan Juc Formation.

T.A. Darragh

Figure 5. A, D-G, I-K, Zoila chathamensis\ A, E, I, holotype, AIM 71325, Flower Pot Harbour, Chatham Islands; A, E, x 1.5; I, x 2; D, F-G,

J-K, GNS TM8792, Taruwhenua Peninsula, Chatham Islands; D, F-G, K, x 1.5; J, x 2; B-C, H, L, Z. viathomsonv, B-C, H, holotype, WAM

72.296, Thomson Highway, Western Australia; L, fossula, P310194, Thomson Highway, x 3.

Type material. Holotype P14515, F.A. Cudmore collection;

paratype P308740, presented S. F. Colliver, 16 January 1962;

paratype P315526, presented C. Goudey, 2009.

Time range. Janjukian, Late Oligocene-Longfordian, Early

Miocene.

Occurrence and material. Janjukian: Jan Juc Formation. At sea

level, Bird Rock, Torquay (P304422, one specimen); Half

Moon Bay, Torquay (P315526, one specimen); Torquay

(P308706, P304432, P304423, seven specimens); Geological

Survey of Victoria locality Ad22, Bird Rock, Torquay (P308717,

one specimen).

Longfordian: Fishing Point Marl. ‘Picnic bed’, Horden

Vale (P308741-2, two specimens).

Remarks. This species is very similar in morphology to

A revision of the Australian fossil species of Zoila (Gastropoda: Cypraeidae)

Figure 6. A-E, I, Zoila didymorhyncha\ A-C, paratype, P302685, PL3022, x 1; D-E, I, holotype, P302687, PL 3022, x 1; F-H, Z. glomerabilis,

paratype, P315526, Half Moon Bay, Torquay, x 1.

Z. dolichorhyncha sp. nov. but differs in that it is more globose,

has a shorter anterior canal and tapers more abruptly towards

the anterior. Zoila glomerabilis bears some resemblance to

Z. venusta (Sowerby), Recent, Western Australia, but is more

globose and has a prominent anterior canal lacking in Z.

venusta. The fossula is very similar to that in Z. venusta, but

not quite as elongate.

Etymology. Latin, glomerabilis, round.

Zoila dolichorhyncha sp. nov.

Figures 8A-F, 9C-E

Description. Shell of large size for genus, solid, pyriform,

somewhat humped posteriorly and tapering anteriorly, of six

teleoconch whorls. Spire projecting slightly, almost enveloped

by last whorl, covered with thick callus. Protoconch of one

smooth whorl, not differentiated from teleoconch whorls,

T.A. Darragh

Figure 7. A-F, Zoila glomerabilis ; A-C, paratype, P308740, Torquay, x 1; D-F, holotype, P14515, Bird Rock cliffs, Torquay, x 1.

A revision of the Australian fossil species of Zoila (Gastropoda: Cypraeidae)

Figure 8. A-F, Zoila dolichorhyncha-. A, C, E, paratype, P304418, PL3028, x 1; B, D, F, holotype, P14514, PL 3028, x 1.

T.A. Darragh

coiled with the axis of the shell. Posterior canal short, deeply

incised, reflexed to left. Anterior canal deeply incised, with

rounded sides, reflexed dorsally, supported laterally by short,

narrow flanges. Aperture sinuous, slightly widened and sloping

steeply inwards before anterior canal; outer lip with 23-33

teeth extending along entire lip, on some specimens becoming

very weak posteriorly; inner lip with 3-11 teeth (generally

fewer than 10) present on anterior third of lip. Fossula well

developed, broad, relatively deeply depressed, bounded

anteriorly by weak terminal ridge.

Dimensions.

Holotype P14514

101

Paratype P304417

Paratype P304418

incised, sides rounded, reflexed dorsally, supported by very

thick lateral flanges extending from base. Aperture somewhat

sinuous, narrow; outer lip with 24-32 teeth, markedly

decreasing in strength at posterior two-thirds. Columella with

9-29 weak, short teeth, present on anterior two-thirds. Fossula

weakly depressed, subcircular, bounded anteriorly by weak

terminal ridge. Dorsum coarsely dimpled on some specimens.

Dimensions.

Holotype SAM

T822

102

Figured specimen

P13374

11 topotype

Figured specimen

P3044471

106

23+

14 Curlewis

Type locality. Lower bed, Table Cape, Wynyard, Tasmania, (i.e.

PL3028 lower bed in cliff between Fossil Bluff and 1.5 km

northwest towards Table Cape, Wynyard, Tasmania, Table

Cape 930630). Freestone Cove Sandstone, Early Miocene,

early Longfordian.

Type material. Holotype P14514, purchased R.N. Atkinson, 8

May 1911; paratypes P304417-8, purchased R.N. Atkinson, 15

January 1910.

Time range. Longfordian, Early Miocene

Occurrence and material. Table Cape (P2644, P14618, P2766-

2770, P14595-6, P304417, P304449-50, 11 specimens);

Crassatella bed, Fossil Bluff (P304440, one specimen); lower

bed, Table Cape (P308708, P304431, P304433, seven specimens).

Remarks. This species has been mistaken for Zoila platypyga

(McCoy) and recorded as such in lists of fossils from Table

Cape published by Johnston (1888, p. 262) and Pritchard (1896,

p. 106). It probably evolved from Z. glomerabilis sp. nov., from

which it differs by being pyriform with a tapering anterior

rather than being globose. Zoila glomerabilis has a uniformly

rounded dorsum, whereas Z. dolichorhyncha sp. nov. has a

dorsum humped posteriorly.

Etymology. Greek, dolichos, long; rhynchos, snout.

Zoila mulderi (Tate, 1892)

Figures 9A-B, 10A-F

Cypraea mulderi Tate, 1892, pi. 9, fig. 2.— Tate, 1893, p. 316

(description).

Gistortia (Palliocypraea) mulderi (Tate).— Vredenburg, 1927, p.

60.

Zoila (Zoila) mulderi (Tate).— Schilder, 1935, p. 337.

Zoila (Gigantocypraea) mulderi (Tate).— Wilson and Clarkson,

2004:52, pi. 56.

Description. Shell solid, of medium to large size for genus,

subglobose, covered with glaze. Spire scarcely visible, covered

with thick callus. Posterior canal wide, sides rounded, deeply

notched dorsally, sides extended posteriorly into prominent

thick flanges extending from base. Anterior canal deeply

Type locality. ‘In a well sinking at Belmont’. A note by Mulder,

dated 1891, with a photograph of the specimen sent to Professor

Tate (i.e. the holotype), states that only two specimens were

known, one in Mulder’s possession (i.e. P13374) and the other

sent to Tate. They were ‘taken from the bottom of a shaft about

60 feet below the surface. The shaft was sunk at Belmont near

Geelong’. The shaft was sunk in about 1891 in an effort to find

fire clay in a paddock close to the junction of the Colac Road

(now Princess Highway) and the Germantown (now Grovedale)

Road (now Corio Street), probably in allotment 9, Parish of

Barrabool. The site was described by Mulder (1897, p. 23).

Type material. Holotype SAM T822, R. Tate collection.

Figured specimen P13374, collected J.F. Mulder 1891,

purchased 1921; figured specimen P3044471, collected F.A.

Cudmore, 21 April 1940.

Time range. Longfordian-Batesfordian, Early Miocene.

Occurrence and material. Longfordian. Fishing Point Marl:

PL3035 southeast of Fischers Point (P308770, one fragment).

Batesfordian. Fyansford Formation: PL3040 Belmont

Shaft (P13374, one specimen). Curlewis (P304416, P304447,

P308743, four specimens and three fragments).

Gellibrand Formation: PL3048 Bornong Road cutting

(P304421-2, P308767, one specimen and two fragments);

PL3163 Williams Road cutting (P310103, one fragment of

anterior).

Horizon unknown. Fishing Point Marl? ‘Cape Otway’

(probably Aire River) (P302701, one specimen).

Fishing Point Marl: Fishing Point (P308769, one fragment).

Remarks. The Bornong cutting specimens have the general

shape of specimens from Belmont and Curlewis, but have some

characters intermediate with Zoila platypyga (McCoy) in that

the teeth are more strongly developed. The labral teeth extend

along the entire lip and the columellar teeth are strongly

developed ridges extending along the entire columella as in Z.

platypyga. This suggests that Z. mulderi is ancestral to Z.

platypyga. The specimen labelled Cape Otway probably came

from the upper part of the Fishing Point Marl (Batesfordian)

outcropping in the cliffs along the Aire River at Horden Vale.

A revision of the Australian fossil species of Zoila (Gastropoda: Cypraeidae)

Figure 9. A-B, Zoila mulderi holotype, SAM T822, Belmont, x 1; C-E, Z. dolichorhyncha, paratype, P304417, PL 3028, x 1.

T.A. Darragh

Figure 10. A-F, Zoila mulderv, A-C, E, P13374, Belmont, x 1; D, F, P304416, Curlewis, x 1.

A revision of the Australian fossil species of Zoila (Gastropoda: Cypraeidae)

Zoila platypyga (McCoy, 1876)

Figures 2A, 11A-H, 12A-G, 13A-C

Cypraea (Aricia)platypyga McCoy, 1876, p. 39, fig. 1-lc.

Cypraea (Aricia) consobrina McCoy, 1877, p. 36, pi. 49, fig. 2.

Cypraea toxorhyncha Tate, 1890, p. 210.— Tate, 1892, p. 5, fig. 6; pi.

6, fig. 5.

Cypraea platypyga McCoy.— Tate, 1890, p. 211.

Cypraea (Erosaria)platypyga McCoy.— Harris, 1897, p. 209.

Gisortiaplatypyga (McCoy).— Vredenburg, 1927, p. 43.

Gisortia consobrina (McCoy).— Vredenburg, 1927, p. 43.

Zoila (Zoila) platypyga platypyga (McCoy). — Schilder, 1935, p.

338.

Zoila (Zoila) platypyga simplicior Schilder, 1935, p. 338.

Zoila (Zoila) consobrina (McCoy).— Schilder, 1935, p. 338.

Zoila (Zoila) toxorhyncha (Tate).— Schilder, 1935, p. 338.

Zoilaplatygyra [,v/c] (McCoy). — Wilson and Clarkson, 2004:51, pi.

52, pi 53 ( toxorhyncha ) form.

Zoila consobrina (McCoy). — Wilson and Clarkson, 2004:51, pi. 54.

Description. Shell of five to six whorls, solid, of medium size for

genus, globose to subpyriform, last whorl covered with glaze,

enveloping the previous whorls, somewhat humped posteriorly

with posterior dorsal slope steeper than anterior; smooth dorsal

surface. Spire slightly projecting, of 4—5 whorls, covered with

thick callus on most specimens. Protoconch smooth, of one whorl

merging imperceptibly with teleoconch whorls; beginning of

protoconch slightly immersed. Posterior canal with rounded sides,

wide, notched, projecting posteriorly on most specimens with

prominent solid lateral projections, longer on columellar side.

Anterior canal subcylindrical, narrow, deeply incised, reflexed

dorsally and supported by thick lateral flanges on most specimens;

on large specimens canal considerably extended. Aperture narrow,

very slightly curved, slightly wider and steeply sloping inwards at

anterior end before anterior canal; outer lip with 24—33 well-

developed teeth extending along entire lip. Columella with 24-33

strongly developed ridges, extending along entire columella,

bifurcating on some specimens, not extending onto fossula.

Fossula well developed, concave, broad and spoon-shaped,

bounded anteriorly by well-developed single terminal ridge.

Dimensions.

Holotype P12137

P12134

28 holotype of

consobrina

T823

26 holotype of

toxorhyncha

Figured specimen

P304414

100

Figured specimen

P304415

Figured specimen

PI4594

100

Type locality. ‘Tertiary clays of shore near foot of Mount Eliza’.

Type material. Holotype PI2137, collected Geological Survey of

Victoria. Holotype of Cypraea (Aricia) consobrina PI2134,

Moorabool River. Holotype of Cypraea toxorhyncha Tate, 1890,

SAM T832, Muddy Creek, Ralph Tate collection. Holotype of

Zoila (Z.) platypyga simplicior Schilder, 1935, BM(NH), Muddy

Creek, E.O. Teale collection (specimen missing). Figured

specimen P304414, FA. Cudmore collection, collected 1941.

Figured specimen P304415, presented Mr J. T. Cunningham, 22

February 1966. Figured specimen P14954, collected FA.

Cudmore, February 1932.

Time range. Balcombian-Baimsdalian, Middle Miocene.

Occurrence and material. Balcombian. Muddy Creek Formation:

Muddy Creek (P14639, P61587, P304426, P308692-4, P308696,

P308748, P308798, P308796, 21 specimens); PL3082 Clifton

Bank (P58641, P304419, P304434, P304446, P308691, P308695,

P308697, 16 specimens).

Gellibrand Formation: Gellibrand River (P304424, P304448,

P308734, four specimens); northwest of Princetown (P304441,

one specimen); Princetown (P5347, one specimen); PL3060 dam

on lot 393 (P304429, one specimen); 4 km north of Port Campbell

(P304439, one specimen); between Pt. Ronald and Gibsons Beach

(P304444, one specimen); Cowley Creek (P304438, one

specimen); dam at Clyne’s house (P308698, one specimen); north

of Port Campbell (P308699, one specimen); Curdies (P59228,

one specimen); PL3068 southwest end of Gibson Beach

(P304443, one specimen).

Fyansford Formation: PL3078 Fossil Beach (P304427-8,

P304437, P304415, ten specimens); Balcombe Bay (P308751-3,

three specimens); Schnapper Point (P308755, two specimens);

Braeside Tunnel (P30445, one specimen); Campbells Point

(P304436, one specimen); PL3069 Red Bluff, Shelford (P304435,

one specimen); Shelford (P304420, one specimen); Overburden,

Batesford Limestone Quarry (P308811, P309021-3, P310316,

P310313, six specimens).

Cadell Marl: PL3084 south of Morgan, South Australia

(P304430, one specimen).

Baimsdalian

Fyansford Formation: Grices Creek (P14594, P304425, five

specimens); PL3103 downstream Grices Creek (P308754, one

specimen); Grices Creek, middle beds (P304414, one specimen);

PL3100 Murgheboluc 4A (P308735, one specimen).

Remarks. The holotype of Cypraea consobrina McCoy is a

slightly immature, water-worn specimen found in the Moorabool

River near Geelong, washed out of the Fyansford Formation. The

posterior and anterior canals are not so highly callused, even when

allowing for wear, as typical specimens of Z. platypyga. However,

these forms intergrade and it is synonymised with the latter.

The holotype of Cypreaea toxorhyncha Tate is a large

specimen, with the extensions of the posterior canal very well

developed. The anterior canal is very long and reflexed dorsally.

Similar specimens are associated with and intergrade with typical

Z. platypyga specimens in the Muddy Creek Formation and the

Fyansford Formation. This name is also synonymised with Zoila

platypyga. The holotype of Zoila (Zoila) platypyga simplicior

Schilder, 1935, said to be in the Teale collection, Natural History

Museum, London, cannot be located. Schilder indicated that it

came from Muddy Creek and that it differed from Z. platypyga

sensu stricto by having less produced and less rounded extremities,

T.A. Darragh

Figure 11. A-H, Zoila platypyga\ A-B, D, H, holotype, P12137, Mount Eliza, x 1; C, E, holotype of Cypraea consobrina McCoy, P12134,

Mooroobool River, x 1; F-G, holotype of Cypraea toxorhyncha Tate, SAM T 823, Muddy Creek, x 1.

Figure 12. A-H, Zoilaplatypyga\ A-D, G, P14594, Grices Creek, x 1; E-F, P304414, Grices Creek, x 1.

T.A. Darragh

Figure 13. A-C, Zoilaplatypyga, P304415, PL 3078, x 1; E-H, Z. gigas\ E, protoconch, P308804, Muddy Creek, x 2.5; F, protoconch, P308807,

Batesford Quarry, x 2; G-H, holotype of Cypraea dorsata Tate, SAM T849, Muddy Creek, x 1.

A revision of the Australian fossil species of Zoila (Gastropoda: Cypraeidae)

less developed lateral flanges and a less projecting spire. The

posterior extremity was bent to the left. These differences in callus

development are minor intraspecific variations, as the specimens

illustrated in figs. 11-13 show. Schilder stated that it bore a

resemblance to Z. consobrina and that it came from a younger

horizon than the latter. In fact, the holotype of consobrina comes

from the same stratigraphical horizon.

Zoila gigas McCoy, 1867

Figures 2B, 13E-H, 14A-H, 15A-E

Cypraea gigas McCoy, 1867a: 18.— McCoy, 1867b: 194.— McCoy,

1867c:438.

Cypraea (Aricia) gigas McCoy.— McCoy, 1875:19, pi. 15; pi. 16,

fig. 2; pis. 17-18, fig. 1; — McCoy, 1876, p. 35, pis. 28-29, fig. 1.

Cypraea dorsata Tate, 1890:212.— Tate, 1892, pi. 10, fig. 4; pi. 11,

fig .6.

Cypraea (Erosaria) gigas McCoy.— Harris, 1897:208.

Cypraea gabrieli Chapman, 1912:190, pi. 13, figs 1-3.

Gisortia gigas (McCoy).— Vredenburg, 1927:58.

Gisortia dorsata (Tate).— Vredenburg, 1927:58.

? Gigantocypraea gigas (McCoy).— Schilder, 1930:126, fig. 34-5

(cast).

Zoila (Gigantocypraea) gigas (McCoy).— Schilder, 1935:337, 338,

fig. 17. — Wilson and Clarkson, 2004:53, pi. 57.

Zoila (Gigantocypraea) dorsata (Tate).— Schilder, 1935:337,338.

Description. Shell solid, very large for genus, globose, covered

with glaze, humped at dorsal midpoint of shell. Ventral surface

flat to slightly concave on most specimens. Spire scarcely

projecting, covered with thick callus. Four teleoconch whorls.

Protoconch of two whorls, the first depressed, large, irregular,

with finely granulated surface, coiled at an angle to axis of

teleoconch whorls. Posterior canal bounded by very thick walls,

deeply incised, strongly reflexed dorsally, deeply notched.

Anterior canal short, about same length as posterior canal, narrow,

deeply incised, strongly reflexed dorsally. Aperture very sinuous,

widest at beginning of anterior canal; outer lip with 2-28 teeth, on

most specimens only extending to midpoint of lip. Columellar lip

edentulous. Fossula scarcely differentiated from columellar lip,

relatively small, slightly depressed, somewhat rectangular,

extending only slightly into aperture, bounded by very weak

terminal ridge.

Dimensions.

Lectotype P12139

212

143

109

SAMT849

- holotype C.

dorsata

P12366

135

- holotype C.

gabrieli

Figured specimen

P13060

167

108

- Grices Creek

Figured specimen

P308774

197

138

P308679

145

102

P12969

247

174

126 largest specimen in

collection.

Type locality. 'Blue clay of Muddy Creek, 10 miles south of

Hamilton’, (i.e. PL3082 Clifton Bank, Muddy Creek, 7 km west

of Hamilton, Victoria). AMG Coleraine WD 818225. Muddy

Creek Formation. Balcombian, Middle Miocene.

Type material. Lectotype PI2139, presented Lindsay Clarke.

McCoy’s figured specimen chosen as lectotype.

Holotype of Cypraea dorsata Tate, SAM T849, Muddy

Creek, Ralph Tate collection.

Holotype of Cypraea gabrieli Chapman, P12366, Bird

Rock cliffs, presented by C J. Gabriel, 3 April 1912.

Figured specimen P13060, presented W.H. Gregson 14

April 1919; figured specimen P308774, collected F.P. Spry

1922.

Time range. Janjukian-Baimsdalian, Late Oligocene-Middle

Miocene.

Occurrence and material. Janjukian. Jan Juc Formation: Bird

Rock cliffs, Torquay (P308679, P308801, one specimen and one

fragment); Deadmans Gully, Torquay (P315577, one specimen).

Batesfordian.

Gellibrand Formation: PL3047 cutting Princetown-

Simpson Road (P308786, one fragment).

Balcombian

Muddy Creek Formation: Muddy Creek (PI2969, P219370,

P61265, P308774, P308794, P308780, P308772-3, 11

specimens); PL3082 Clifton Bank (P308776, P308795, two

specimens).

Gellibrand Formation: Southeast of Gibson Beach (P54362,

P308682, two specimens).

Fyansford Formation: PL3078 Fossil Beach (P308677,

308779 two specimens); Balcombe Bay (P308683-5, three

specimens); Momington (P308782, one specimen); Schnapper

Point (P24866, P30777-8, three specimens); Port Phillip

(P308781, one specimen); Altona Bay Coal Shaft (P308787,

one fragment); Batesford (P308790, one specimen);

Overburden, Batesford Quarry (P303315, one specimen and

one fragment); Orphanage Hill (P308784, one specimen).

Morgan Limestone: Broken Cliffs opposite Waikerie,

Murray River Cliffs, South Australia (private collection,

Mannum).

Bairnsdalian

Gellibrand Formation: Glenample Steps (P308771, one

specimen); Princetown (P5288, one specimen).

Fyansford Formation: Grices Creek (P5286, PI 2968,

P12970, P13060, P308681, five specimens); Middle beds,

Grices Creek (P3 08680, one specimen); Murgheboluc

(P308678, one specimen).

Casts of a large cowry — which, due to their size, are

almost certainly this species — have been found as follows:

Longfordian?

Gambier Limestone: Mount Gambier (P61262, one

specimen, BM(NH) one specimen).

Batesfordian.

Curlewis Limestone: Curlewis (P308806, one specimen).

Balcombian

Nullarbor Limestone: 5 km south of Watson Station, South

Australia (P316748, one specimen).

T.A. Darragh

Figure 14. A-H, Zoila gigas\ A-B, holotype of Cypraea gabrieli Chapman, P12366, Bird Rock cliffs, Torquay, x 0.5; C-E, P308679, Bird Rock

cliffs, x 0.5; F-H, P13060, Grices Creek, 0.5.

A revision of the Australian fossil species of Zoila (Gastropoda: Cypraeidae)

Figure 15. A-E, Zoila gigas\ A-B, P308774, Clifton Bank, x 0.5; C-E, lectotype, P12139, Muddy Creek, x 0.5.

T.A. Darragh

Fyansford Formation: Keilor (P58646, one specimen);

sewerage excavation, Belmont (P308688, one specimen).

Gippsland Limestone: Newmerella Railway cutting

(P308686, one specimen).

Morgan Limestone: Blanchetown (P316749, one specimen);

‘Brittan’, Murray River cliff 4.8 km downstream of Morgan

(P316751, one specimen).

Remarks. This is the largest known Australian cowry, living or

fossil. Specimens over 190 mm in length are not uncommon.

Specimens are quite common in the middle Miocene, but because

of their size, complete or undamaged specimens are uncommon.

Cypraea dorsata Tate is merely a small specimen of Zoila gigas.

The general shape, fossula and aperture are the same. The main

difference is that the anterior and posterior canals are more

strongly developed in most large specimens of Z. gigas. Small

specimens, like the type of C. dorsata, occur with larger

specimens at Fossil Beach, Grices Creek, Muddy Creek and

Batesford Quarry, and there are also gradations between the

smallest and the largest specimens. For these reasons, C. dorsata

is synonymised with Z. gigas.

Cypraea gabrieli Chapman was based on a crushed specimen

from the Jan Juc Formation, which is very difficult to compare

with other material. The main differences between it and the

lectotype of Z. gigas McCoy are a depressed spire, the lack of

callus covering the spire and the presence of a dimpled dorsum.

Only one other specimen from the Jan Juc Formation is available.

This is complete and uncrushed, but has a very chalky shell.

There are well-developed teeth on the outer lip extending to

about midway on the lip. On the columella there are some very

low but distinct teeth situated over the fossula and about midway

along the aperture. The dorsum is smooth and the base is

rounded rather than flat or convex. Apart from the presence of

the columellar teeth, this specimen closely matches a similar¬

sized specimen from the Fyansford Formation at Grices Creek

in all features including the spire and fossula. McCoy pointed

out there were weak teeth just visible on the columella of the

holotype of Z. gigas. The significance, if any, of the presence of

the dimpled dorsum cannot be assessed as there are so few

specimens. Because there are no consistent differences between

the two specimens from the Jan Juc Formation and specimens of

Z. gigas from the Fyansford Formation, Cypraea gabrielli is

regarded as a further synonym of Z. gigas.

The protoconch of this species (figs 2B, 13E-F) is different

from those of other species of Zoila and also from those of

species of Umbilia. There are fewer whorls and the first is

irregular in shape, suggesting that it might originally have been

chitinous and subsequently calcified. Other species of Zoila are

similar to species of Umbilia in that they have a smooth, regular

protoconch of three whorls, and a coiling axis at an angle to that

of the teleoconch.

Zoila sp.

Figures 16C-D, M

Zoila sp. McNamara and Kendrick, 1994:34.

Description. Shell solid, small for genus, somewhat globose.

Spire covered with callus, not projecting. Posterior canal short,

notched. Anterior canal missing; slight trace of anterior basal

extension on left flank. Aperture narrow, very slightly curved;

outer lip with well-developed, elongate teeth (15 teeth

preserved). Columellar lip with well-developed teeth extending

along the whole aperture (18 teeth preserved). Base of shell

rounded on both sides of aperture. Fossula well developed,

concave, broad and spoon-shaped, bounded anteriorly by well-

developed, single terminal ridge.

Dimensions.

Figured specimen WAM 82.549

48+

Figured material. WAM 82.549, collected K. J. McNamara and

G. W. Kendrick, September 1981.

Occurrence. Locality 12, Latitude Point, from large boulders

of pink limestone fallen from upper level of cliff, Barrow

Island, Western Australia. Poivre Formation, middle Miocene.

Remarks. This species is known only from a single specimen,

which has a small piece of the left side of the posterior canal

broken off, as well as a large portion of the right side of the

anterior end, including the posterior canal. As a consequence,

detailed comparison with other species is not possible; however,

there is enough of the specimen preserved to indicate that it is

a species of Zoila and almost certainly ancestral to the younger

species occurring in Western Australia. Both aperture and

fossula are very similar to those of Z. campestris sp. nov. from

the late Pliocene, Roe Calcarenite. It differs from Z. campestris

in that its outline is more rounded and it does not have a flat

base. The dorsum is also not humped anteriorly as in Z.

campestris sp. nov. It bears no close resemblance to any of the

fossil species known from southeast Australia.

Zoila sp. is somewhat similar to the middle Pliocene

species Zoila gendingensis (Martin, 1899) from the Upper

Kalibeng Formation of Sonde, Java, Indonesia, but is not so

high and does not have the flat base of that species. The dorsum

is not as humped as in Zoila kendengensis Schilder, 1941 from

the Pleistocene Putiangan Formation of Java and neither does

it have the flat base of that species. Of the living species of the

genus, it is most similar to Zoila venusta (Sowerby, 1846),

known from the Great Australian Bight to Shark Bay, Western

Australia.

Zoila campestris sp. nov.

Figures 16A-B, E-J, O

Cypraea (Zoila) sp. Ludbrook, 1978, p. 129, pi. 13, fig. 19.

Zoila sp. Wilson and Clarkson, 2004: 52, pi. 55, figs a, b.

Description. Shell solid, polished, of average size for genus,

globose, surface on some specimens malleated with rectangular

depressions, with sides about 1.5 mm long; ventral surface

flattened. Spire barely protruding beyond last whorl, covered

with thick callus. Posterior canal short, notched, sides

thickened. Anterior canal very short, abruptly truncated, deeply

incised. Aperture sinuous, widened above fossula; outer lip

with 17-24 well-developed teeth, extending along entire lip;

Figure 16. A-B, E-J, O, Zoila campestris\ A-B, paratype, WAM 89.437, Roe Plain, Western Australia, uncoated to show colour pattern, x 1; E,

J, O, paratype, P308704, Roe Plain, x 1; F-H, holotype, WAM 89.177, Roe Plain, x 1; C-D, M, Z. sp., WAM 82.549, Barrow Island, Western

Australia, x 1; K-L, N, Z.fodinata, holotype, WAM 89.637, Roe Plain, Western Australia, x 1.

T.A. Darragh

inner lip with 17-21 well-developed teeth, extending along

entire lip. Fossula well developed, deeply depressed, bounded

anteriorly by sharp terminal ridge. Colour pattern present on

some specimens, of brown ground with darker brown patches.

Dimensions.

Holotype WAM 89.177

Paratype WAM 89.437

Paratype NMV P308704

Type locality. Quarry 2.5 km north of Hampton microwave

repeater tower, Roe Plains, Western Australia. AMG Eucla

1:250,000 sheet CK365465. Roe Calcarenite.

Type material. Holotype WAM 89.177, collected G.W. Kendrick

27-30 October 1988; paratype WAM 89.437, collected Sam

Rowe, January 1989; paratype NMV P308704, collected T. A.

Darragh, 9 August 1973.

Time range. Pliocene.

Occurrence and material. Roe Calcarenite: PL3172 Hampton

Tower (P308704-5, WAM 69.494, 70.17, six specimens);

PL3167 1.5 km north of Hampton Tower (P308703, WAM

80.109, three specimens); PL3166 2.5 km north of Hampton

Tower (P121293, WAM 89.178, 89.437, 89.637, four specimens).

Remarks. At first glance, small specimens of this species may

be confused with the common Austrocypraea amae Fehse and

Kendrick of the Roe Calcarenite, but they are readily

distinguished by their smooth fossula. In Austrocypraea, the

columellar teeth are produced into thin ribs that continue

across the fossula. This species is most closely related to the

living species Zoila decipiens (Smith, 1880), Recent, Western

Australia, from which it differs by having stronger teeth and

teeth present along the entire columella. It is also globose in

shape rather than pyriform as in Z. decipiens and lacks the

prominent protruding spire of that species. The fossula is very

similar to that of the Z. decipiens. It is not closely similar to the

fossil species of Zoila known from the Oligocene and Miocene

of southeast Australia, with the exception of a species known

from a single broken specimen from the upper Miocene of

Victoria. Z. campestris bears some resemblence to Zoila

kendengensis Schilder, 1941 from the Pleistocene Putiangan

Formation of Java, but that species has relatively prominent

anterior and posterior canals.

Etymology. Latin campester, pertaining to a plain.

Zoila fodinata sp. nov.

Figures 16K-L, N

Description. Shell solid, polished, of average size for the genus,

globose, pyriform, ventral surface rounded. Spire slightly

protruding beyond last whorl, covered with thick callus.

Posterior canal short, notched, sides thickened. Anterior canal

very short, abruptly truncated, deeply incised. Aperture

sinuous, widened above fossula; outer lip with 25-28 well-

developed teeth, extending along entire lip; inner lip with 21-

25 well-developed teeth, extending along entire lip. Fossula

well developed, subrectangular, concave, bounded on inner

side by low ridge and anteriorly by sharp terminal ridge;

terminal ridge rather broad, extending down into aperture as

sharp ridge forming anterior edge of fossula.

Dimensions.

Holotype WAM 89.637

Type locality. Quarry 2.5 km north of Hampton microwave

repeater tower, Roe Plains, Western Australia. AMG Eucla

1:250,000 sheet CK365465. Roe Calcarenite.

Type material. Holotype, WAM89.637, collected G.W.

Kendrick, 27-30 October 1988.

Time range. Pliocene

Occurrence and material. Roe Calcarenite: PL3166 2.5 km

north of Hampton Tower (one specimen).

Remarks. This species is most closely similar to the living

species Zoila venusta, from which it differs by having stronger

teeth and teeth present along the entire columella. The fossula

is also shallower but deeper within the aperture than in Z.

venusta. It is not as globose, being more pyriform. It does not

seem to be closely similar to the fossil species of Zoila known

from the Oligocene and Miocene of southeast Australia.

Etymology. Latin, fodina, a quarry.

Acknowledgments

The following persons have assisted me by providing

information, and the donation or loan of specimens, and are

gratefully acknowledged: Chris J. Goudey, Lara, Victoria;

Peter Hunt, Adelaide, South Australia; David T. Dockery,

Mississippi Bureau of Geology; the late Warren Blow, United

States National Museum; Wolfgang Grulke, Dorset, England;

Ben McHenry, South Australian Museum; George Kendrick

and Helen Gore, Western Australian Museum; Ian Loch,

Australian Museum; John Cooper, Natural History Museum,

London; Barry Wilson, Perth, Western Australia; Pierre

Lozouet, Philippe Bouchet and Virginie Heros, Museum

national d’Histoire naturelle, Paris; Alan Beu and John Simes,

GNS Science, Lower Hutt, New Zealand. I am grateful to

Rodney Start, Museum Victoria and Mark Darragh for

considerable assistance with photography. I thank Alan Beu

and Barry Wilson, who provided significant helpful comments

and suggestions that have considerably improved the

manuscript.

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Memoirs of Museum Victoria 68: 29-35 (2011)

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

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

A new species of Peribrissus (Echinoidea, Spatangoida) from the middle Miocene

of South Australia

Francis C. Holmes

Honorary Associate, Invertebrate Palaeontology, Museum Victoria, GPO Box 666, Melbourne, Victoria 3001, Australia,

and 15 Kenbry Road, Heathmont, Victoria 3135, Australia (fholmes@bigpond.net.au)

Abstract Holmes, F. C. 2011. A new species of Peribrissus (Echinoidea, Spatangoida) from the middle Miocene of South Australia.

Memoirs of Museum Victoria 68: 29-35.

A new species of spatangoid echinoid from the middle Miocene Glenforslan Formation cropping out in the Murray

River cliffs near Blanchetown, South Australia, is described and assigned to the genus Peribrissus. Peribrissus janiceae

sp. nov. is only the third species of this genus to be recorded, and the first to occur outside the Mediterranean area of

Europe and North Africa. Brief references are made to the similarity of certain features in Prenaster, Pericosmus and

Peribrissus, which have caused confusion with identification in the past.

Keywords Echinoidea, Spatangoida, Peribrissus, new taxa, middle Miocene, South Australia

Introduction

In the Miocene stratigraphic sequences along the Murray River

and elsewhere in Australia, species belonging to the Spatangoida

constitute approximately 50 per cent of the recorded taxa of

irregular echinoids (Holmes et al., 2005). The discovery of yet

another new species of spatangoid, albeit a single specimen,

should come as no surprise considering the vast extent of these

generally poorly examined outcrops in South Australia.

However, what is intriguing is that the new species belongs to a

genus that, so far, has been recorded in the literature from only

the Mediterranean area of Europe and North Africa. The

specimen was found by Chris Ah Yee and Janice Krause in

2007 at Museum Victoria locality PL3203 (see fig.l), the same

location as the three specimens of Murraypneustes biannulatus

Holmes et al., 2005, discovered in 2003.

Materials and methods

The specimen number prefixed ‘P\ on which this study is

based, is housed in the Invertebrate Palaeontology Collection,

Museum Victoria (NMV). Wherever possible, measurements

where made with a dial calliper to an accuracy of 0.1 mm.

Parameters are expressed as a percentage of test length (%TL),

test width (%TW) or test height (%TH).

Age and stratigraphy

The Glenforslan Formation, in which the specimen was found,

is synonymous with the Lower Morgan limestone, which

conformably overlies the Finniss Formation and is of early

middle Miocene (Batesfordian, Langian) age. The thickness of

the unit is relatively consistent at 13-15 m, although this is

reduced in southern exposures due to post-middle Miocene

uplift and subsequent erosion. Echinoids tend to be found at or

above the floatstone-rudstone contact at the base of cycles

composed of mollusc-bryozoan floatstone grading upward

into Celleporaria rudstone tops (Lukasik and James, 1998).

Sediments are pervasively mottled, obscuring all physical

sedimentary textures. The middle Glenforslan Formation is

interpreted as being deposited in relatively shallow waters,

possibly less than 10 m, based on the presence of calcareous

algae and mixotrophic foraminifers (Dr Jeff Lukasik, Petro-

Canada Oil and Gas, Calgary, pers. com., 2005). This section

of the formation forms part of the richest warm-water biotic

record from southern Australia at a time of maximum

transgression of the sea across the continental shelf

(McGowran and Li, 1994, and papers cited therein).

Associated fauna

Refer to Holmes et al. (2005) for a table of echinoid species

recorded from the Glenforslan Formation.

Systematic palaeontology

Order Spatangoida L. Agassiz, 1840

Suborder Paleopneustina Markov and Solov’ev, 2001

Family Prenasteridae Lambert, 1905

Remarks. The family Paleopneustidae A. Agassiz, 1904,

together with the families Pericosmidae, Schizasteridae and

Prenasteridae, initially established as tribes within the family

F.C. Holmes

Figure 1. A and B, general location maps; C, map of Murray River between Waikerie and Swan Reach, South Australia, showing locality of NMV

PL3203, north of Blanchetown.

Brissidae by Lambert (1905, p. 153), allowed numerous

spatangoid genera to be divided into groups based primarily on

the distinctive path followed by their fascioles. Lambert and

Thiery (1925, pp. 514-515) listed Peribrissus Pomel, 1883 as a

subgenus of PrenasterDesor, 1853 within the tribe Prenasterinae.

However, in subsequent classifications of the Order Spatangoida

by Mortensen (1951), Termier and Termier (1953), Durham and

Melville (1957), Fischer (1966) and Smith (1984), the family

Prenasteridae was not recognised, and Peribrissus and

Prenaster were placed within the Schizasteridae. Not until

Smith et al. (2005) and Smith and Stockley (2005) did the

family Prenasteridae reappear in any subdivision of the

Spatangoida. Finally, Kroh and Smith (2010) presented a

primary framework for the classification of post-Palaeozoic

echinoids based on extant taxa into which fossil taxa have been

incorporated. In this classification, Prenasteridae, Schizasteridae

and Periasteridae form the Suborder Paleopneustina.

Genus Peribrissus Pomel, 1869

Type species. Peribrissus saheliensis Pomel, 1883, by subsequent

monotypy.

Other species. P. sotgiai Giorgio, 1923.

Diagnosis. Modified from Smith et al. (2005). Test medium to

large and cordiform with distinct anterior sulcus, posterior face

oblique to vertically truncate, profile depressed to moderately

domed. Apical disk well anterior of centre, ethmolytic with

three gonopores. Ambulacrum III sunken aborally, the groove

increasing in width and depth to ambitus, with rows of enlarged

tubercules occurring just outside adradial sutures, pores small.

Petals straight, narrow and depressed, cruciform, the anterior

pair longer than the posterior pair. Peristome and plastron

plating of type species unknown. Periproct high on posterior

truncate face. Semipetalous fasciole band combines with

continuous marginal fasciole immediately behind and below

anterior petals.

Remarks. There has been confusion regarding the designation

of the type species of Peribrissus. Fischer (1966, p. U576) and

Smith et al. (2005) stated that P. saheliensis is the type species

by original designation, but Pomel (1869, p. 13) did not name

any species he assigned to his genus, for which he gave only a

very brief diagnosis and made comparisons with Prenaster.

Pomel later (1883, p. 36) gave a slightly more detailed diagnosis

followed by the statement ‘P. saheliensis est du miocene

superieur’. As saheliensis was the only named species assigned

A new species of Peribrissus (Echinoidea, Spatangoida) from the middle Miocene of South Australia

to Peribrissus, the diagnosis given for the genus applies also to

the species, thus satisfying the criteria for availability (ICZN,

Article 12.2.6) and making P. saheliensis the type species by

subsequent monotypy (ICZN, Article 68.3).

Two species of Pericosmus described by McNamara and

Philip (1964) from the Miocene of Australia — P. celsus and

P. quasimodo — were reassigned by Smith et al. (2005) to

Peribrissus. Though Pericosmus and Peribrissus are

superficially alike, the path of the peripetalous fasciole in the

two Australian species is clearly different from that in the

Prenasteridae and, consequently, in Peribrissus. Smith et al.

(2005) stated that in the Prenasteridae, ‘marginal and

peripetalous fasciole combine anteriorly, the combined band

passing several plates below the end of the anterior petals’. In

contrast, the peripetalous fasciole in Pericosmus celsus and P.

quasimodo — as well as in P. torus, also erected by McNamara

and Philip in the same paper — follow a distinctly different

path. These three species have the peripetalous fasciole closely

bounding the distal end of the anterior petals, then transversely

crossing interambulacral plates in columns 2a and 3b before

taking a longitudinal path (sometimes irregular and/or

intermittent) towards the marginal fasciole in interambulacral

columns 2b and 3a. Due to the state of preservation of the

numerous Pericosmus specimens inspected in Museum

Victoria and private collections, it is not possible to determine

whether the peripetalous fasciole always reaches the marginal

fasciole on either side of the anterior sulcus. Nevertheless, in

all Australian species assigned to Pericosmus, including P.

compressus Duncan, 1877 and P. maccoyi Gregory, 1890, the

peripetalous fasciole closely bounds the distal end of the

anterior petals and continues transversely onto interambulacra

2 and 3, clearly negating any reassignment to Peribrissus.

However, whether the five Australian fossil species listed

above strictly belong in the genus Pericosmus is a matter of

conjecture, considering the type species Pericosmus latus

Desor in Agassiz and Desor, 1847, has separate and continuous

marginal and peripetalous fascioles, the latter crossing

ambulacrum III well above the anterior margin.

Stefanini (1911, p. 86) reassigned Prenaster excentricus

(Wright, 1855) to Peribrissus in the belief that the two genera

overlap based on the similarity of their upper test profile with

highly eccentric anterior apex and four ethmolitic genital

pores. Pomel’s statement (1887, p. 63) — that the number of

genital pores in Peribrissus is unknown — seems to have been

ignored by Stefanini, whose reference to four genital pores

may have been based on details of Wright’s species. Giorgio

(1923, p. 125), in describing Peribrissus sotgiai from Sardinia,

accepted Stefanini’s finding that Wright’s Prenaster from

Malta was a Peribrissus ; noting that P. sotgiai has four

gonopores, but that the right anterior one is poor and almost

atrophied. These statements appear to have resulted in

Mortensen (1951) and Fischer (1966) listing both genera as

having four genital pores. However, of the eight genera now

included in the family Prenasteridae by Smith et al. (2005),

only Peribrissus and Tripylus Philippi, 1845 are listed as

having three genital pores. Although both of these genera have

a well-defined anterior sulcus, Peribrissus is easily

distinguished from Tripylus by the markedly anterior location

of its apical disk compared to the central position in the latter.

The lack of a sulcus and the presence of four genital pores in

species of Prenaster clearly refute Stefanini’s reassignment of

Prenaster excentricus to Peribrissus.

Peribrissus janiceae sp. nov.

Figures 2A-E, 3A-I, Table 1

Type material. Holotype and only known specimen, NMV P316528,

from the early middle Miocene Glenforslan Formation (Batesfordian,

Langian), Morgan Group, 7 km north-northeast of Murray River Lock

1, Blanchetown , South Australia [NMV locality PL3203].

Description. Test moderately large, ovate in outline with well-

formed anterior sulcus; only known specimen 58.0 mm long,

with maximum width of 52.0 mm (89.7%TL) occurring

posterior of centre at 54.3%TL from anterior ambitus.

Maximum test height 38.5 mm (66.4%TL) anterior of centre,

but posterior of apical disk at 44.8%TL from anterior ambitus.

Adapical surface inflated with high, vertically convex

anterior, gently curved ridge along interradial suture of

interambulacrum 5 and prominent vertically truncated

posterior. Laterally, sides gently curved at approximately 40°

to the horizontal between dorsal ridge and well-rounded

ambitus situated about one-third test height above the

underside. Adoral surface posterior of peristome flat along

centre line of labrum and plaston (fig. 2C-E).

Small, very closely spaced tubercles cover nearly all the

test; smallest around ambitus and largest towards peristome.

Figure 2. Peribrissus janiceae sp. nov. A-E, outline drawings of

adapical, adoral, left lateral, anterior and posterior views of holotype

NMV P316528, showing paths followed by marginal (mf) and

semipetalous (spf) fascioles, and position of peristome (pse) and

periproct (ppt).

F.C. Holmes

Figure 3. Peribrissus janiceae sp. nov. holotype NMV P316528. A-D and F, anterior, right lateral, posterior, adapical and adoral views; E, detail

of apical disk; G, oblique left lateral view of semipetalious fasciole crossing interambulacrum 4, plates 7a, 8a, 9a and 9b; H, detail of peristome,

labrum and phylodal plates; I, detail of junction between marginal and semipetalous fascioles on interambulacrum 1, plate 4b. Scale bars = 10 mm

unless otherwise shown.

Tubercles in ambulacra II and IV first appear on plates 3a and

b, and in I and V on plates 4a and b. By plates 5a and b, the size

and spacing generally matches that of adjacent interambulacra.

Largest tubercles with an approximate areole diameter of 1.0

mm occur on interambulacra 1 and 4 adjacent to adoral edge of

plates 2a and b, aborally on plate 1, and along the adradial

suture line between ambulacrum III and interambulacra 2 and

3 from the marginal fasciole to the apical disk. These tubercles

have a perforate mamelon and crenulate platform but appear to

lack a scrobular ring. Because of the very close spacing of these

tubercles, miliary granules occur mainly towards the peristome

and between the periproct and marginal fasciole in

interambulacrum 5 where the spacing between the larger

tubercles increases. They also occur around the apical disk.

A well-defined marginal fasciole occurs just above the

sloping ambitus, dipping sharply below the periproct posteriorly

but crossing ambulacrum III anteriorly slightly below the

ambitus at about 25%TH (see fig. 2). The semipetalous fasciole

is only marginally indented between the posterior paired petals

and crosses interambulacra 1 and 4 on plates 8/9 before

descending transversely to join the marginal fasciole at right

angles, posterior to the angle of the anterior paired petals (see

fig. 31). Although continuous, fasciole widths vary but maintain

a fine tubercule (granule) density of about 100-120 per mm 2 .

A new species of Peribrissus (Echinoidea, Spatangoida) from the middle Miocene of South Australia

Table 1. Comparison of diagnostic features of Peribrissus janiceae sp. nov. with those of the type species of the genus, P. saheliensis Pomel, 1883,

and P. sotgiai Giorgio, 1923.

Diagnostic feature

Peribrissus janiceae sp. nov.

Peribrissus saheliensis Pomel

Peribrissus sotgiai Giorgio

Width/length ratio of test

89.7%TL

Similar

Maximum width location

Marginally posterior at

54.3%TL

Marginally anterior, approx.

45%TL

Similar to P. saheliensis

Height//length ratio of test

66.4%TL

Not known

Approx. 54%TL

Maximum height location

Slightly anterior at 44.8%

Well anterior but posterior of

apical disk, approx. 30%TL

Well posterior, approx.

68%TL

Anterior lateral profile

High, vertically convex

Slopes forward from apex at

approx. 35°

Similar to P. janiceae

Posterior lateral profile

High vertical truncation

Oblique truncation

Adoral surface lateral profile

Flat, posterior of peristome

Unknown (type specimen

compressed)

Slightly swollen posterior of

centre

Sulcus

Max. depth/width ratio approx.

1:4.6, occurs below ambitus

Figured much deeper with a

depth/width ratio of approx. 1:2.2

Figured far shallower and

wider than P. janiceae

Apical system location and type

21%TL, ethmolitic, 3

gonopores (none in plate G2)

Approx. 33%TL, detail of apical

system unknown (Pomel 1887).

Stefenini (1911) incorrectly

assumes 4 gonopores. This

repeated by Mortensen (1950)

and Fisher (1966).

Approx, average of 2

specimens 25%TL,

ethmolitic. Giorgio’s

description refers to 4

gonopores but states the pore

in G2 almost atrophied

Ambulacmm III, marginal

tubercles

Not unduly prominent, situated

just outside adradial sutures

Similar to P. janiceae

Larger, far more prominent

with rows further apart

Detail of petals

Straight, parallel sided and

sunken

Similar, but probably shallower

Similar

Length differentiation paired

petals

Anterior petals 138% longer

than posterior ones

Similar to P. janiceae

More equal, but posterior

petals still shorter than

anterior ones

Anterior paired petals

divergence angle

175°

Approx. 13 5°

Described as 140°. Giorgio’s

figures, however, suggest

divergence wider

Posterior paired petals

divergence angle

315°

Approx. 295°

Similar to P. janiceae

Peristome

Reniform and slightly sunken

Insufficient information for

comparison

Insufficient information for

comparison

Periproct

Vertically elliptical at top of

posterior truncation

Semicircular, assumed high on

posterior truncation

Elliptical (axis not clear),

high on posterior truncation

Fascioles, marginal and

semipetalous

Marginal fasciole occurs just

above sloping ambitus and is

joined by semipetalous fasciole

at right angles behind and

below anterior paired petals

Similar, but with semipetalous

fasciole shown angled forward at

junction with marginal fasciole,

apparently due to less oblique

divergence of anterior petals

Insufficient information for

comparison, as stated to be

only visible in some places

Apical system situated well anterior of centre at 21.0% TL

from anterior ambitus to centre of disk and is level with

proximal end of paired petals. Ethymolitic with three

gonopores, no gonopore in plate G2, and approximately 60

hydropores fairly evenly spaced over the latter’s length.

Paired petals straight, parallel sided, sunken, open distally

and devoid of tubercles. Anterior paired petals 138% longer

than posterior pair, extending 50% of the radius (28.0%TL)

measured along the surface of the perradial suture from centre

of ocular to ambitus. Anterior paired petals diverge at 175° and

contain 23/24 pore pairs, posterior petals 315° and 20/21 pairs.

Outer pores elliptical, inner pores slightly smaller and more

tear shaped. Zone between inner and outer pores approximately

equal in width to outer pores, pairs not conjugate. Interporiferous

zone marginally narrower than poriferous zones.

Ambulacrurum III depressed for its full length below

F.C. Holmes

adjacent interambulacra, reaching a maximum depth of 3 mm

(5.2%TL) below the anterior ambitus. Pore pairs are visible

adapically between the ocular plate and approximately one-

third of the radius to the anterior ambitus. Adapically, the pore

pairs are angled inwards at approximately 45° to the perradial

suture but gradually become monoserial halfway towards the

anterior ambitus. The ambulacrum is covered with closely

spaced small tubercles and miliary granules, the former

gradually increasing in diameter adorally.

Peristome reniform and slightly sunken, longitudinal

dimension 4 mm (6.9%TL), width 8.6 mm (14.8%TL), anterior

edge situated 12.4 mm (21.4% TL) from ambitus. Phyllodes

unipored with periporal areas protuberant. Basicoronal plates

amphiplaceous.

Labrum small, wider than long, covered with small

tubercles and flared anteriorly where bordered by a smooth

raised rim (fig. 3H). Curved anterior edge projects over the

peristome for about one-third of the latter’s length. Posterior

edge does not extend beyond the first adjacent ambulacral

plates. Plastron wide, long, and covered with rows of closely

spaced angular tubercles without interstices. Maximum width

of plastron (45%TW) occurs about three-quarters of the test

length from the anterior ambitus.

Periproct elliptical shaped with slightly pointed upper and

lower junction with interradial suture, height 8.0 mm

(13.8%TL), width 5.0 mm (5.6%TL). Underside of vertical

opening situated high above base of test (44.2%TH) on

truncated posterior surface. Subanal surface slightly depressed.

Etymology. Named for Janice Krause of Hamilton, Victoria, an

exceptionally dedicated fossil echinoid collector.

Remarks. Comparison of Peribrissus janiceae sp. nov. with the

type species P. saheliensis from Algeria and P. sotgiai from

Sardinia is complicated by the lack of detailed descriptions,

comparative measurements and illustrations of many of the

important diagnostic features of the latter two species. The

difficulty is compounded by the excellent preservation of detail

found on the single specimen of P. janiceae and the large

difference in size between specimens of the three species, with

P. saheliensis approximately twice the length and width of P.

janiceae and four times that of P. sotgiai. Where possible,

diagnostic features of the three species are compared in table 1,

based on the descriptions of Pomel (1887), Giorgio (1923) and

Stefanini (1911), together with approximate measurements

taken from their illustrations of the partial and poorly preserved

type specimens.

Acknowledgements

I am indebted to Christopher Ah Yee and Janice Krause

(Hamilton, Victoria) for collecting and donating the holotype,

and to David Holloway (Invertebrate Palaeontology, Museum

Victoria) for valuable advice and support during the

preparation of this manuscript. Kenneth McNamara

(Department of Earth Sciences, Cambridge University) and

Andrew Smith (Department of Palaeontology, Natural History

Museum, London) are thanked for suggesting improvements

to the manuscript. Peter Aucote of Sunlands, South Australia,

kindly gave approval to collect on his property and Museum

Victoria Library staff and Eric Poirot (www.echinologia.com)

were extremely helpful in obtaining references. Michael Gatt

(Rabat, Malta) is also thanked for providing photographs for

comparative purposes.

References

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Museum of Comparative Zoology at Harvard College 31: 1-243.

Agassiz, L. 1840. Catalogus systematicus Ectyporum Echinodermatum

fossilium Musei Neocomiensis, secundum ordinem zoologicum

dispositus; adjectis synonymis recentioribus, nec non stratis et locis

in quibus reperiuntur. Sequuntur characteres diagnostici generum

novorum vel minus cognitorum. Petitpierre, Neuchatel, 20 pp.

Agassiz, L. and Desor, E., 1847. Catalogue raisonne des families, des

generes et des especes de la classe des echinodermes. Annales des

Sciences naturelles, series 3, Zoologie 8: 5-35.

Desor, E. 1853. Notice sur les echinides du terrain nummulitique des

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279.

Duncan, P. M. 1877. On the Echinodermata of the Australian Cainozoic

(Tertiary) deposits. Quarterly Journal of the Geological Society of

London 33(1): 42-73, pis 3-4.

Durham, J. W. and Melville, R. V. 1957. A classification of echinoids.

Journal of Paleontology 31: 242-272.

Fischer, A. G. 1966. Spatangoids. Pp. U543-U628 in Moore R. C.

(ed.). Treatise on Invertebrate Paleontology, Part UEchinodermata

3(2). The Geological Society of America Inc. and The University

of Kansas Press.

Giorgio, A. Di 1923. Echinidi Miocenici della Sardegna. Atti della

Societa Toscana di Scienze Naturali residente in Pisa, 35: 116-

BO, pi.2(1).

Gregory, J. W. 1890. Some additions to the Australian Tertiary

Echinoidea. The Geological Magazine 27 (new series, decade 3,

vol. 7, no. 11): 481^-92, pis 1314.

Holmes, F. C., Ah Yee, C. and Krause, J. 2005. Two new Middle

Miocene spatangoids (Echinoidea) from the Murray Basin, South

Australia. Memoirs of Museum Victoria 62(1): 91-99.

ICZN (International Commission on Zoological Nomenclature) 1999.

International Code of Zoological Nomenclature, 4th edn.

International Trust for Zoological Nomenclature, London, 306 pp.

Kroh, A. and Smith, A. B. 2010. The phylogeny and classification of

post-Palaeozoic echinoids. Journal of Sy sterna tic Palaeontology 8

(2): 141-212.

Lambert, J. 1905. Notes sur quelques Echinides eoceniques de l’Aude

et de PHerault. In: L. Doncieux (ed.). Catalogue descriptif des

fossiles nummulitiques de PAude et de PHerault. Annales de

TUniversite de Lyon, Nouvelle Serie, 1. Sciences, Medicine 17:

129-164.

Lambert, J. and Thiery, P. 1925. Essai de nomenclature raisonee des

echinides. Libraire L. Ferriere, Charmont, fasc. 8-9, 513-607, pis

12-13, 15.

Lukasik, J. J. and James, N. P. 1998. Lithostratigraphic revision and

correlation of the Oligo-Miocene Murray Supergroup, western

Murray Basin, South Australia. Australian Journal of Earth

Sciences 45: 889-902.

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Australia. Records of the South Australian Museum 27: 197-212.

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echinoid Pericosmus from the Tertiary of Australia. Records of the

Western Australian Museum 11(40): 319-356.

A new species of Peribrissus (Echinoidea, Spatangoida) from the middle Miocene of South Australia

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Paleopneustidae (Echinoidea, Spatangoida) morfologiya, sistema,

filogeniya. Rossiyskaya Akademiya nauk, Trudy

Paleontologicheskogo lnstituta 280: 1-109.

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2. Amphisternata 2. Spatangidae, Loveniidae, Pericosmidae,

Schizasteridae, Brissidae. C. A. Reitzel, Copenhagen, 593 pp.

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kritischen Bemerckungen uber einige weniger bekannte Arten.

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servir d’introduction a Vetude des fossils. Deyrolle, Paris: 67 pp.

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vivants etfossiles. Adolphe Jordan, Alger, 131 pp., 1 pi.

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(Wright sp.). Rivista Italiana di Paleontologia e Stratigrafia

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Memoirs of Museum Victoria 68: 37-65 (2011)

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

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

The paracaudinid sea cucumbers of Australia and New Zealand (Echinodermata:

Holothuroidea: Molpadida: Caudinidae)

P Mark O’Loughlin 1 , Shari Barmos 2 and Didier VandenSpiegel 3

1 Marine Biology Section, Museum Victoria, GPO Box 666, Melbourne, Victoria 3001, Australia (pmo@bigpond.net.au)

2 (shari_barmos@hotmail.com)

3 Musee royal de l’Afrique centrale. Section invertebres non-insects, B-3080, Tervuren, Belgium (dvdspiegel@

africamuseum.be)

Abstract O’Loughlin, P. M., Barmos, S. and VandenSpiegel, D. 2011. The paracaudinid sea cucumbers of Australia and New

Zealand (Echinodermata: Holothuroidea: Molpadida: Caudinidae). Memoirs of Museum Victoria 68: 37-65

The four Paracaudina Heding species reported in Australia are reviewed: Paracaudina australis (Semper);

Paracaudina chilensis (Muller); Paracaudina luticola Hickman; Paracaudina tetrapora (H. L. Clark). The New Zealand

species Paracaudina coriacea (Hutton) is raised out of synonymy with the Chilean species Paracaudina chilensis

(Muller). Both Paracaudina chilensis (Miiller) and Paracaudina coriacea (Hutton) occur in New Zealand. The synonymy

of the Chinese and Japanese species Paracaudina ransonnetii (Marenzeller) with Paracaudina chilensis (Muller) is

maintained. Five new Caudinidae species are erected for Australia, with authors O’Loughlin and Barmos: Paracaudina

ambigua, Paracaudina bacillis, Paracaudina cuprea, Paracaudina keablei, Paracaudina tripoda. A key is provided for

Paracaudina species in Australia and New Zealand.

Keywords Sea cucumber, Molpadida, Caudinidae, Paracaudina , new species, synonymies, Australia, New Zealand, key.

Introduction

Museum Victoria holds numerous specimens of Caudinidae

sea cucumbers, most specimens collected from Victorian

beaches after storms. Rowe 1982 recorded a “ Paracaudina

sp.” from Port Phillip Bay in Victoria and southern Western

Australia. This species is represented by numerous specimens

from southern Australian waters, and we erect a new species.

We have examined collections of Paracaudina Heding, 1932

from the Australian, South Australian, Western Australian

and New Zealand Museums, and we review all of the species

of this genus in Australian and New Zealand waters. As an

outcome of this review we erect five new species of

Paracaudina. Our systematic study is based on traditional

morphological characters as there are to date no adequate

molecular genetic data.

Four potential Australian and New Zealand species of

Paracaudina have been intensively studied: Paracaudina

australis (Semper, 1868) with type locality Australia;

Paracaudina chilensis (Muller, 1850) with type locality Chile;

Paracaudina coriacea (Hutton, 1872) with type locality New

Zealand; Paracaudina ransonnetii (Marenzeller, 1881) with

type locality China. Their morphology and systematic status

have been researched and debated by numerous authors: Theel

1886; H. L. Clark 1908, 1935; Mortensen 1925; Hozawa 1928;

Ohshima 1929; Heding 1931, 1933; Pawson 1963; Pawson and

Liao 1992. Most recently Pawson 1963 has summarised what

has been debated, and agreed with the authors who considered

P. coriacea (Hutton) to be a junior synonym of P. chilensis

(Muller). We reject this synonymy, judging that Paracaudina

coriacea (Hutton) is a good species. But at the same time we

have found that Paracaudina chilensis (Muller) also occurs in

New Zealand waters, and one or possibly more undescribed

Paracaudina species. A comprehensive review of the many

Paracaudina specimens held in New Zealand will follow this

work. In reporting on the molpadid sea cucumbers of China,

Pawson and Liao 1992 agreed with the synonymy of the China

and Japan species P. ransonnetii (Marenzeller) with P.

chilensis (Muller). We agree with this synonymy. However, we

note that some of these species distributions are such that

molecular genetic data may reveal that some of these species

are paraphyletic.

Our experience of looking at a range of material leads us

to agree with H. L. Clark 1935 that there is considerable

variation in morphological form amongst Paracaudina

specimens that are conspecific, including ossicle form. And

we agree also that, although there is considerable variation in

ossicle form in the same and conspecific specimens,

predominant ossicle form does provide a reliable guide to

species identity.

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Methods

For scanning electron microscope (SEM) observations by

Didier VandenSpiegel ossicles were cleared of associated soft

tissue in commercial bleach, air-dried, mounted on aluminium

stubs, and coated with gold. Observations were made using a

JEOL JSM-6480LV SEM. Measurements were made with

Smile view software. Photos of preserved specimens by Shari

Barmos were taken with a Nikon D70 DSLR camera, using a

Nikon micro 60 mm lens.

Abbreviations

AM Australian Museum (with registration prefix J).

MRG Marine Research Group of the Field Naturalists Club of

Victoria.

NMV Museum Victoria (with registration prefix F).

NIWA New Zealand Institute of Water and Atmospheric

Research.

SAM South Australian Museum (with registration prefix K).

WAM Western Australian Museum (with registration prefix Z).

Numbers in brackets after registrations refer to the number

of specimens in a registered lot.

Spelling correction

Some authors have misspelled the species name ransonnetii,

and the correct spelling only is used throughout our work.

Key to the Australian and New Zealand species of

Paracaudina Heding

1. Ossicles in body wall small irregular rods only.

Paracaudina bacillis O’Loughlin and Barmos sp. nov.

(southern Australia)

— Ossicles in body wall include small buttons or perforated

plates or thick cups.2

2. Ossicles in mid-body predominantly small rods and loops,

and in caudal end predominantly spinous perforated

plates. Paracaudina ambigua O’Loughlin and Barmos

sp. nov. (central Western Australia)

— Ossicles similar in mid-body and caudally.3

3. Ossicles in body wall predominantly octagonal plates

with central perforation over which there is a bridging box

on the basal side and cross on the outer side.4

— Ossicles in body wall rarely octagonal plates with box and

cross bridges, or such ossicles completely absent.6

4. Octagonal plates predominantly with marginal bluntly

pointed projections and knobs; perforations not

significantly smaller in collective area than the surface

area of the plate; body with long thin tail. Paracaudina

chilensis (Muller, 1850) (circum-Pacific, north-western

Australia)

— Octagonal plates predominantly with rounded margin,

margin lacking bluntly pointed projections and knobs ... 5

5. Ossicles predominantly thick and button-like, perforations

significantly smaller in collective area than the plate

surface area; body with long thin tail. Paracaudina

coriacea (Hutton, 1872) (New Zealand)

— Ossicles predominantly thin-walled and open mesh-like,

perforations not significantly smaller in collective area

than the plate surface area; body with short thin tail.

Paracaudina keablei O’Loughlin and Barmos sp. nov.

(north-eastern Australia)

6. Ossicles in body wall predominantly thick-walled with 4

perforations.7

— Ossicles in body wall include irregular plates and buttons

with irregular perforations, with marginal and surface

knobs and pointed projections variably present; plates

with up to 12 perforations.8

7. Ossicles in body wall predominantly thick, shallow

concave crossed cups Paracaudina luticola Hickman,

1962 (southern Australia)

— Ossicles in body wall predominantly thick, knobbed and

irregularly oval flat plates. Paracaudina tetrapora (H.

L. Clark, 1914) (southern Australia)

8. Ossicles in body wall predominantly buttons with smooth

rounded lateral margin, short thick blunt surface spines,

frequently with central perforation bridged by a tripod of

rods ... Paracaudina tripoda O’Loughlin and Barmos, sp.

nov. (north-eastern Australia)

— Ossicles in body wall plates, most with lateral and surface

rounded marginal spines and knobs.9

9. Deeper mid-body wall with irregular rods, less than 50

pm long; colour never yellowish-red (rusty).

Paracaudina australis (Semper, 1868) (north-eastern

Australia)

— Deeper mid-body wall lacking irregular rods; larger

specimens increasingly yellowish-red (rusty) in colour.

Paracaudina cuprea O’Loughlin and Barmos sp. nov.

(southern Australia)

Remarks. The remaining subspecies and species of

Paracaudina Heding, 1932 that are not included in this key are:

Paracaudina chilensis obesacauda (H. L. Clark, 1908), a

central American east Pacific and west Atlantic subspecies,

retained with subspecific status solely on geographical grounds

by Pawson et al. 2001; Paracaudina delicata Pawson and Liao,

1992, from the Gulf of Tonkin, has thin-walled chilensis- like

ossicles, with box and cross bridging a central perforation, and

fine digitiform projections around the ossicle margin.

Order Molpadida Haeckel, 1896

Diagnosis (emended from Pawson and Liao 1992). Tentacles

15, digitate; body stout, lacking tube feet, usually with an

evident tail; anal papillae, tentacle ampullae and respiratory

trees present; ossicles may include tables, cups, rods, perforated

plates and modified anchors; phosphatic bodies often present.

The paracaudinid sea cucumbers of Australia and New Zealand (Echinodermata: Holothuroidea: Molpadida: Caudinidae)

Remarks. Rods may be present with perforated plates, as in

Paracaudina australis (Semper, 1868), and rods are not

necessarily fusiform (spindle-shaped). We have emended the

diagnosis of Pawson and Liao 1992 in these two respects.

Family Caudinidae Heding, 1931

Diagnosis (emended from Pawson and Liao 1992). Tentacles

without a terminal digit, and with one or two pairs of lateral

digits; tail sometimes inconspicuous; ossicles may include

large tables, crossed cups, perforated plates and irregular rods;

phosphatic bodies usually absent.

Remarks. We have emended the statement concerning which

ossicles may be present, but we have not changed the ossicle

types listed.

Paracaudina Heding, 1932

Figures If, 5c-e, 7, 8, 12a, b

Pseudocaudina Heding, 1931: 283.

Paracaudina Heding, 1932: 455-456.

Type species. Molpadia chilensis Muller, 1850 (subsequent

designation by H. L. Clark 1935).

Other included species. Paracaudina ambigua O’Loughlin

and Barmos sp. nov.; P. australis (Semper, 1868); P. bacillis

O’Loughlin and Barmos sp. nov.; P. coriacea (Hutton, 1872); P.

cuprea O’Loughlin and Barmos sp. nov.; P. delicata Pawson

and Liao, 1992; P. keablei O’Loughlin and Barmos sp. nov.; P.

luticola Hickman, 1962; P. chilensis obesacauda (H. L. Clark,

1908); P. tetrapora (H. L. Clark, 1914); P. tripoda O’Loughlin

and Barmos sp. nov.

Diagnosis. Cylindrical form, smooth or wrinkled body wall;

tentacles 15, each with 2 pairs of digits (figs If, 5c); posterior end

of body with caudal taper or discrete thin tail; tail may be short or

long; 5 radial triangular non-calcareous anal valves, each with up

to 4—5 pairs of marginal digitiform papillae, terminal ones longest

(fig. 5d); radial plates of calcareous ring with two anterior lateral

low blunt projections, one with small notch, posterior digitiform

prolongation about half the length of the plate, prolongation

divided by terminal notch of variable depth or deeper division

(fig. 12a); interradial plates with anterior central blunt point,

posterior end truncated (fig. 12a); dorsal short to long tubular

stone canal with terminal madreporite, free in coelom or attached

to pyloric mesentery (specimen NMV F174894); single ventral,

elongate, tubular to globular polian vesicle, usually with dark

reddish-brown colouration; longitudinal muscles broad, flat, with

distinct longitudinal division (fig. 5e); retractor muscles formed

by pair of in-tumed outer margins of divided longitudinal muscles

(fig. 5e); gonad tubules usually branched, in tufts on each side of

dorsal mesentery (fig. 5e); right branch of respiratory tree

extending in the coelom to the calcareous ring; ossicles may be

concave or flat, thick crossed and knobbed cups, thick knobbed

perforated plates, octagonal plates with large central perforation

and cross or tripod bridging one side and sometimes square the

other side (figs 7, 8, 12b), perforated smooth and knobbed and

spinous plates with variably developed secondary layering, and

irregular rods; ossicles never tables; phosphatising of ossicles

and calcareous ring may occur, and a red to brown to yellow

pigment may occur in the body wall.

Remarks. A comprehensive diagnosis of genus Paracaudina

Heding, 1932 is provided to avoid diagnoses of species with

repetition of characters that all have in common. We recognise

that some of the characters listed in this diagnosis of

Paracaudina are shared with other genera and at family and

possibly order level.

Paracaudina ambigua O’Loughlin and Barmos sp. nov.

Figure la, 2

Material examined. Holotype. Western Australia, Shark Bay, FWA-

WAM RV Naturaliste Shark Bay Survey II Feb / Mar 2003, stn

13/173/P, 24°47.02'S 113°21.97'E to 24°46.48'S 113°22.08'E, 24 m, 6

Mar 2003, S. Morrison and S. M. Slack-Smith, WAM Z29767.

Paratype. Western Australia, Ningaloo Marine Park, AIMS RV

Solander , stn RVS 4545-D069, 23°48’S 113°30'E, 33 m, 1 Feb 2008,

M. Salotti and S. M. Slack-Smith, WAM Z23331 (1).

Diagnosis. Paracaudina species up to 45 mm long (holotype,

caudal taper but anal end and valves missing), body up to 15

mm high (body flattened laterally); body wall thick, soft to

semi-gelatinous, white (preserved); posterior body with caudal

taper to short discrete tail (evident in paratype); ossicles

different mid-body and caudally; mid-body ossicles

predominantly small irregular rods, frequently forming a

single loop, or small plates with up to 4 perforations, these

ossicles up to 40 pm long; rare mid-body irregularly oval

perforated plates with spinous margin and some secondary

bridging, about 64 pm long; caudal ossicles irregularly oval to

round to octagonal perforated plates, long pointed spinous

margin, surface spines and bridges and secondary development,

some plates chilensis- like with box and cross bridges over

central perforation, caudal ossicles typically 56-64 pm long.

Type locality. Central Western Australia, Shark Bay.

Distribution. Central Western Australia, Shark Bay, Ningaloo

Marine Park; 24-33 m.

Etymology. From the Latin ambiguus (of double meaning,

uncertain), feminine ambigua , referring to the generic

uncertainty created by the presence of Acaudina- like ossicles

mid-body and Paracaudina- like ossicles posteriorly.

Remarks. Paracaudina ambigua O’Loughlin and Barmos sp.

nov. is distinguished diagnostically amongst Paracaudina

species by having fine irregular rod and loop ossicles in the

mid-body wall and marginally spinous chilensis- like plate

ossicles in the caudal region, some with the characteristic cross

and box bridges over a central perforation. Both specimens are

in poor condition, but the ossicles are in excellent condition and

characterise the new species. Distal pairs of digits are evident

on a few tentacles of the holotype, but the condition of the

tentacles is such that a second pair is not clearly evident on any

tentacle. The caudal part is missing from the holotype, but a

distinct narrow tail and soft anal valves are present on the

paratype (18 mm long).

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 1. Photos of preserved specimens of species of Paracaudina Heding, 1932: a, holotype of P. ambigua O’Loughlin and Barmos sp. nov.

(insert with drawings of rod and loop ossicles from the mid-body wall; WAM Z29767); b, P. australis (Semper, 1868) (insert with drawings of

rod ossicles from the mid-body wall; AM J13583); c, P. chilensis (Muller, 1850) (WAM Z5638); d, P. coriacea (Hutton, 1872) (NIWA 70955); e,

P. coriacea (Hutton, 1872) (AM J12290); f, mouth and tentacles of P. coriacea (Hutton, 1872) (NIWA 70954).

The paracaudinid sea cucumbers of Australia and New Zealand (Echinodermata: Holothuroidea: Molpadida: Caudinidae)

Figure 2. SEM images of ossicles from the holotype of Paracaudina ambigua O’Loughlin and Barmos sp. nov. from Shark Bay, central Western

Australia (WAM Z29767): upper, forms of predominant ossicles from the mid-body wall; lower, forms of ossicles from the caudal body wall

(rarely occurring chilensis-hke ossicles with cross and box bridging shown lower top right and lower bottom left).

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 3. SEM images of ossicles from the mid-body wall of a specimen of Paracaudina australis (Semper, 1868) from Dunwich, Moreton Bay,

south-east Queensland (AM J13583). Rarely occurring chilensis -like ossicle with cross and box bridging shown top right.

The paracaudinid sea cucumbers of Australia and New Zealand (Echinodermata: Holothuroidea: Molpadida: Caudinidae)

Figure 4. SEM images showing variation in form of ossicles from the mid-body wall of a specimen of Paracaudina australis (Semper, 1868)

from Dunwich, Moreton Bay, south-east Queensland (AM J13583). Rarely occurring chilensis-Wke ossicle with cross and box bridging shown

bottom right.

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 5. Photos of Paracaudina bacillis O’Loughlin and Barmos sp. nov.: a, preserved holotype specimen (insert with drawings of rods from

the mid-body wall; NMV F151853); b, photo of live holotype (oral end right; photo by L. Altoff and A. Falconer); c, mouth and tentacles of

paratype (NMV F174894); d, anal valves and papillae (holotype); e, dorsal mesentery with tufts of gonad tubules on both sides, divided flat

longitudinal muscle with thin retractor muscle (holotype); f, live specimen spawning. Port Phillip Bay, Rye Pier, 4 m, 10 Nov 2007 (photo by D.

McKenzie).

The paracaudinid sea cucumbers of Australia and New Zealand (Echinodermata: Holothuroidea: Molpadida: Caudinidae)

Figure 6. Paracaudina bacillis O’Loughlin and Barmos sp. nov. a, b, SEM images of rod ossicles from the mid-body wall of a paratype from

Westernport Bay, Victoria (NMV F174893); c, photo of live specimen in Blairgowrie Marina, Port Phillip Bay, at 5 m depth (photo by J. Finn, 18

June 2011).

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 7. SEM images of ossicles from the mid-body wall of a specimen of Paracaudina chilensis (Muller, 1850) from Eighty Mile Beach, NW

Australia (WAM Z5640).

The paracaudinid sea cucumbers of Australia and New Zealand (Echinodermata: Holothuroidea: Molpadida: Caudinidae)

Figure 8. SEM images of ossicles from the mid-body wall of a small specimen of Paracaudina chilensis (Muller, 1850) from Tasman Bay, South

Island, New Zealand (NIWA 70956).

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 9. SEM images of ossicles from the mid-body wall of a specimen of Paracaudina coriacea (Hutton, 1872) from the west coast of the South

Island, New Zealand (AM J12290).

The paracaudinid sea cucumbers of Australia and New Zealand (Echinodermata: Holothuroidea: Molpadida: Caudinidae)

Figure 10. SEM images of ossicles from the mid-body wall of a specimen of Paracaudina coriacea (Hutton, 1872) from Cook Strait, New

Zealand (NIWA 70954).

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 11. Photos of preserved specimens of species of Paracaudina (a, c-f): a, P. cuprea O’Loughlin and Barmos sp. nov. holotype (NMV

F157396); b, photo of live specimens of P. cuprea including holotype (photo by P. Vafiadis); c, P. keablei O’Loughlin and Barmos sp. nov.

holotype (AM J13579); d, P. luticola Hickman, 1962 (NMV F169342); e, P. tetrapora (H. L. Clark, 1914) (from Merricks, Westernport Bay,

Victoria; NMV F76565); f, P. tetrapora (H. L. Clark, 1914) (from off Glenelg, South Australia; AM J24918).

The paracaudinid sea cucumbers of Australia and New Zealand (Echinodermata: Holothuroidea: Molpadida: Caudinidae)

Figure 12. a. Calcareous ring of a specimen of Paracaudina cuprea O’Loughlin and Barmos sp. nov. from Portland Bay, Victoria (NMV

F174890) (insert with drawing of two radial plates and one interradial plate typical of Paracaudina species); b, SEM images of ossicles from a

specimen of Paracaudina chilensis obesacauda (H. L. Clark, 1908), judged here to be typical of Paracaudina chilensis (Muller, 1850) (copied

from Pawson et al. 2001).

Paracaudina australis (Semper, 1868)

Figures lb, 3, 4

Molpadia australis Semper, 1868: 233-234, 268, pi. 39 fig 14.—

Theel, 1886: 55.

Caudina chilensis— H. L. Clark, 1908: 175—176. (part non

Molpadia chilensis Muller, 1850).

Pseudocaudina australis.— Heding, 1931: 283.

Paracaudina australis.— Heding, 1932: 455.—Heding, 1933: 127-

142, pi. 6 figs 5-7, pi. 7 figs 8-9, pi. 8 fig. 6.-H. L. Clark, 1935: 267-

284.—H. L. Clark, 1946: 45 (part).-A. M. Clark and Rowe, 1971:

193.—Pawson, 1977: 119 (part).-Rowe, 1982: 472 (part), fig. 10.35b.-

Cannon and Silver, 1986: 40, figs 8f, 10f.—Rowe and Gates, 1995: 264

(part).

Material examined. Queensland, Moreton Bay, Stradbroke Island,

Dunwich, half buried on sandbar, 2 Dec 1978, AM J13583 (1); Port

Denison District, AM J4145 (5).

Diagnosis. Paracaudina species up to 135 mm long, up to 35

mm diameter (preserved); thin, pliable, soft to firm smooth

body wall; colour translucent pink to white live, off-white to

pale brown preserved, lacking yellow colouration, at most

slight yellowing anteriorly; posterior body with caudal taper to

narrow rounded end, sometimes short tail; mid-body ossicles

predominantly plates with bluntly spinous margin and surface,

smooth plates, rods in deeper body wall; ossicles not thick

buttons; spinous plates irregular, pointed marginal projections,

surface knobs / blunt spines frequently joined by rods creating

secondary layering, rods sometimes bridging a large central

perforation as single rod or tripod or cross, rare chilensis- like

plates with large central perforation bridged by box on one side

and cross on the other side, up to 12 perforations, spinous plates

up to 56 pm long; smooth plates irregular, up to 12 perforations,

margin smooth, lacking marginal and surface spines and

knobs, smooth plates up to 48 pm long; rods sparse, irregular,

variably straight, bent, Y-shaped, C-shaped, J-shaped, some

with node, rods frequently 24 pm long, up to 48 pm long.

Type locality. Rockhampton, Queensland.

Distribution. Northern Australia, Queensland, south-east

coast; Singapore (?).

Remarks. Paracaudina australis (Semper, 1868) is

distinguished diagnostically by having in the body wall both

irregular rods and spinous perforated plates with secondary

layering, a body form with tapered caudal end but not long

discrete tail, and an absence of distinct yellow or reddish-

yellow colour. H. L. Clark 1908 included Paracaudina australis

in his synonymy of Caudina chilensis, but subsequently (1935,

1946) rejected his own synonymy. Southern Australian

specimens judged to be Paracaudina australis by H. L. Clark

1946, Rowe 1982 and Rowe and Gates 1995 are our new species

Paracaudina cuprea O’Loughlin and Barmos (below).

Mortensen 1925 based his discussion of Paracaudina australis

on SAM specimens. Based on Mortensen’s figures we judge

that these specimens were our new species Paracaudina

cuprea O’Loughlin and Barmos (below). We note that Heding

1933 did not indicate what specimens he used to illustrate

ossicles of Paracaudina australis.

David Lane (pers. comm, by Ria Tan) identified the

common “See-through sea cucumber” in Singapore waters as

Paracaudina australis. Ria Tan (pers. comm.) has observed

the species only on the estuarine northern shores of Singapore

near the mouth of the Johor River, floating or partly buried on

sand bars near seagrass meadows, as shallow as the intertidal

zone at low spring tide. We have to date not been able to

confirm the determination as Paracaudina australis.

Paracaudina bacillis O’Loughlin and Barmos sp. nov.

Figures 5a-f, 6a-c

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Paracaudina sp. Rowe, 1982: 470, fig. 10.34c.

Paracaudina australis. —Gowlett-Holmes, 2008: 263.—

Saunders, 2009: 99, fig. 5.19. (non Molpadia australis Semper, 1868)

Material examined. Holotype. Victoria, Westernport Bay, Phillip

Island, Ventnor, McHaffie Point, MRG, 16 Feb 2008, NMV F151853.

Paratypes. Westernport Bay, San Remo, Griffith Point, 2 Jan 1999,

NMV F89700 (1): Merricks beach, 5 Nov 1967, NMV F45254 (1); 27

Jul 1969, NMV F45240 (1); 4 Jun 2011, NMV F174893 (1); 10 Jun

2011, NMV F174894 (1); Shoreham, 26Aug 1978, NMV F76071 (2);

30 Mar 1902, NMV F173250 (1) (removed from Shoreham lot 60669-

71 / H19 (3) examined in part by Joshua 1914).

Other material. Victoria, Westernport Bay, Crawfish Rock, 12 m,

13 Oct 1968, NMV F169343 (1); Somers, 28 Sep 1968, NMV F45273

(1); Port Phillip Bay, Brighton beach, NMV F45020 (1); Rosebud

beach, NMV F173247 (1); Rye pier, 6 Apr 2011, NMV F173272 (1);

Blairgowrie Marina, 5 m, 18 Jun 2011 (photo J. Finn, NMV);

Tootgarook beach, 20 Jun 2011, NMV F174896 (1). South Australia,

Yorke Peninsula, Browns Beach, SAM K2493 (6); Edithburgh, sand,

3-4 m, 12 May 1999, SAM K2486 (1); 2-3 m, 11 Jun 2005, SAM

K2487 (1); sand, low tide, 26 Oct 2007 SAM K2490 (1); 8 Nov 2003,

SAM K2491 (1). Western Australia, east of Duke of Orleans Bay,

Niminup beach after storm, Nov 1990, WAM Z31888 (1); Albany, 5

Dec 1983, WAM Z31881 (1).

Diagnosis. Paracaudina species up to 180 mm total length, up

55 mm diameter, caudal taper 45 mm long (SAM K2491,

preserved); thin, parchment-like to soft leathery body wall

(preserved); live colour off-white, preserved colour off-white to

pale yellow or brown; posterior body with caudal taper to

narrow rounded end, sometimes short discrete tail, lacking a

discrete long thin tail; body wall ossicles small irregular rods

only, straight, bent, wavy, J-shaped, sometimes with nodes,

rarely with short branches, up to 64 pm long.

Type locality. Victoria, Westernport Bay, Phillip Island,

Ventnor, McHaffie Point.

Distribution. Victoria (Westernport Bay, Port Phillip Bay),

South Australia Gulfs, Eyre Peninsula, to southern Western

Australia (Albany); 0-145 m (Rowe 1982); 0-230 m (Gowlett-

Holmes 2008).

Etymology. From the Latin baculus (rod), and its diminutive

bacillus, referring to the very small rod ossicles only in the

body wall.

Remarks. Paracaudina bacillis O’Loughlin and Barmos sp.

nov. is distinguished diagnostically amongst Paracaudina

species by having only small irregular rod ossicles in the body

wall. Rods of similar form occur also in Paracaudina australis

(Semper, 1868), a species that also has irregular marginally

spinous perforated plate ossicles in the body wall. Rowe 1982

recognised this “undescribed form” from Port Phillip Bay and

southern Western Australia, and illustrated (fig. 10.34c) the

diagnostically characteristic minute irregular rod ossicles from

the body wall. Gowlett-Holmes 2008 illustrated and described

a species from southern Australia as Paracaudina australis

that has the body form, size up to 20 cm long, and off-white

colour of Paracaudina bacillis O’Loughlin and Barmos sp.

nov. Gowlett-Holmes 2008 described the habit as “usually

completely buried 5-10 cm below the sediment surface; moves

slowly through the sand feeding on detritus, leaving a broad

furrow-like trail”. Saunders 2009 also illustrated a specimen as

P. australis at Coffin Bay on the Eyre Peninsula that we judge

to be P. bacillis based on size and form and colour. Joshua 1914

referred two specimens from Westernport Bay and “Mordialloc”

(Port Phillip Bay) to Caudina chilensis (Muller). We found

these two specimens with a third specimen in lot H19 / 60669-

71 / NMV F45019, and assigned (below) the two referred to by

Joshua 1914 to the new species Paracaudina cuprea O’Loughlin

and Barmos (F169344) and Paracaudina tetrapora (H. L.

Clark, 1914) (F45019, original registration). The label indicated

all three specimens were collected by J. A. Kershaw at

Shoreham. We judge that Joshua’s reference to “Mordialloc”

was a mistake. We found no specimens from Mordialloc in the

NMV collection. The third specimen in the lot, not commented

on in Joshua 1914, is the third new species (above) Paracaudina

bacillis O’Loughlin and Barmos (F173250).

Paracaudina chilensis (Muller, 1850)

Figures lc, 7, 8, 12b

Molpadia chilensis Muller, 1850: 139.—Muller, 1854: pi. 6 fig. 14,

pi. 9 fig. 1.-Semper, 1868: 233,-Theel, 1886: 55.

Microdactyla caudata Sluiter, 1880: 348-351, pi. 6 fig. 1, pi. 7 figs

1 - 6 .

Caudina ransonnetii Marenzeller, 1881: 126-127, pi. 4 figs 5,

5A.—Ludwig, 1883: 158-159.—Lampert, 1885: 210.—’Theel, 1886:

54.—Ludwig, 1891: 354,-Mitsukuri, 1912: 261-262, pi. 8 fig. 76.

Caudina caudata— Ludwig, 1883: 159. (synonymy with Caudina

chilensis (Muller) by H. L. Clark 1908)

Caudina coriacea.— Theel, 1886: 47, pi. 3 fig 4a-c. (non Caudina

coriacea Hutton, 1872)

Caudina rugosa R. Perrier, 1904: 16.—R. Perrier, 1905: pi. 4 figs

10-12. (synonymy with Caudina chilensis (Muller) by H. L. Clark

1908)

Caudina pigmentosa Perrier, 1904: 16-17.—Perrier, 1905: pi. 4

figs 1-9. (synonymy by H. L. Clark 1935)

Caudina contractacauda H. L. Clark, 1908: 38-39, 173, 177, 178,

pi. 9 figs 9-13. (synonymy by H. L. Clark 1935)

Caudina chilensis— H. L. Clark, 1908: 173, 175-176.—Hozawa,

1928: 361-378, pis 14-17.-Ohshima, 1929: 39-45.

Pseudocaudina chilensis.— Heding, 1931: 283.

Pseudocaudina ransonnetii.— Heding, 1931: 283.

Paracaudina chilensis— Heding, 1933: 127-142, pis 5-8.—H. L.

Clark, 1935: 267-284.-Deichmann, 1938: 383-384, fig. 15,-Pawson,

1969: 139-140.—A. M. Clark and Rowe, 1971: 184-185.—Pawson,

1977: 119 (part).—Cannon and Silver, 1986: 40.—Rowe and Gates,

1995: 264.—Liao and Clark, A. M„ 1995: 518-519, fig. 316,-Lane et

al., 2000: 491.

Paracaudina ransonnetii— Heding, 1933: 455.—Djakonov et al.,

1958: 377.

Paracaudina chilensis var. ransonnetii H. L. Clark, 1935: 281.—

H. L. Clark, 1938: 540-541 ,-H. L. Clark, 1946: 444.-A. M. Clark

and Rowe, 1971: 194-195, fig. 96a. (synonymy by Liao and Pawson

1992)

Material examined. Western Australia, Roebuck Bay, Broome, Sep

1929, from H. L. Clark collection, AM J6435 (1); Eighty Mile Beach,

19°20'00"S 121 o 21'00"E, Annabim Expedition 1999, mudflat, WAM

Z5637 (1); WAM Z5638 (2); WAM Z5639 (1); WAM Z5640 (1); WAM

Z5641 (1); WAM Z5642 (1); WAM Z5653 (1); WAM Z5654 (1); WAM

Z5655 (2 tails). New South Wales, Twofold Bay, Nullica Bay, 9.1 m, 22

Feb 1985 AM J19908 (one 3 mm fragment). New Zealand, South

The paracaudinid sea cucumbers of Australia and New Zealand (Echinodermata: Holothuroidea: Molpadida: Caudinidae)

Island, Tasman Bay, 41.17°S 173.17°E, 0 m, 24 Jan 1972, NIWA

70956 (2).

Diagnosis. Paracaudina species up to 150 mm total length,

diameter up to 20 mm, caudal taper and long discrete tail 80

mm long; posterior body with discrete long thin tail, up to more

than half the length of the body; body firm thin leathery to

parchment-like, finely wrinkled, variably transversely creased

(preserved), live colour off-white with slight purple colouration

(H. L. Clark 1938; Liao and A. M. Clark 1995), preserved

colour pale grey to pink-grey; mid-body ossicles predominantly

octagonal plates with large central perforation bridged by basal

box with 4 supporting arms on one side and cross with 4

supporting arms on the upper side, surface knobbed, margin

with 8 bluntly pointed projections, plates up to 56 pm across.

Type locality. Chile.

Distribution. Circum-Pacific, including northwest Australia

and New Zealand; to 1000 m (Pawson 1963).

Remarks. Ludwig 1883, Theel 1886, Heding 1931, 1933, and

Djakonov et al. 1958 judged the Chinese / Japanese species

Paracaudina ransonnetii (Marenzeller, 1881) to be a discrete

species. Type locality is Yantai (Cheefoo) on the Yellow Sea. H.

L. Clark 1935, 1938, 1946 (based on northern Australia

material) and A. M. Clark and Rowe 1971 considered the

species to be a variety of Paracaudina chilensis (Muller, 1850).

H. L. Clark 1908, Hozawa 1928, Ohshima 1929, Pawson and

Liao 1992, Rowe and Gates 1995, and Liao and Clark A. M.

1995 considered Paracaudina ransonnetii to be a junior

synonym of Paracaudina chilensis. We agree with this

synonymy. We judge that some north-western Australian

material is Paracaudina chilensis, and that some north-eastern

Australia material (from the Gulf of Carpentaria and Moreton

Bay) is a new species Paracaudina keablei O’Loughlin and

Barmos that we describe below.

Below we raise the New Zealand species Paracaudina

coriacea (Hutton, 1872) out of synonymy with the Chilean

species Paracaudina chilensis (Muller, 1850). Theel 1886 was

prompted to refer New Zealand specimens to Caudina

ransonnetii Marenzeller, 1881 but finally referred them to

Caudina coriacea (Hutton, 1872). But the ossicles illustrated by

Theel 1886 are the predominant ossicle form found in

Paracaudina chilensis. We examined two small New Zealand

specimens (NIWA 70956) from the shallows of Tasman Bay

and found the ossicles to be those of Paracaudina chilensis (fig.

8). We dismissed our consideration that this ossicle form might

be a juvenile developmental stage of Paracaudina coriacea on

the grounds that Hozawa 1928 found no such significant

development changes in his study of Paracaudina chilensis at

Asamushi. Both Paracaudina chilensis (Muller) and

Paracaudina coriacea (Hutton) occur in New Zealand waters.

We note that the distal tail of the specimen of Paracaudina

chilensis WAM Z5638 is not very thin (fig. lc), while the distal

tails of the two specimens of Paracaudina coriacea NIWA

70955 and AM J12290 are both very thin (figs Id, e).

We also note that some, but not the predominant, ossicles

from specimens judged to be Paracaudina chilensis from

northwest Australia are similar to the ossicles illustrated for

the single type specimen of Paracaudina delicata Pawson and

Liao, 1992 taken in the Gulf of Tonkin. And some, but not the

predominant, ossicles are similar to those in the new species

Paracaudina keablei O’Loughlin and Barmos (below).

Paracaudina coriacea (Hutton, 1872)

Figures ld-f, 9, 10

Molpadia coriacea Hutton, 1872: 17.—Hutton, 1879: 307.—

Lampert, 1885: 208-209.

Echinosoma (?) coriacea. —Hutton, 1879: 307. (synonymy with

Caudina coriacea (Hutton) by Theel 1886)

Caudina meridionalis Bell, 1883: 58-59, pi. 15 fig. 1.—Lampert,

1885: 210-211. (synonymy with Caudina coriacea (Hutton) by Theel

1886)

Caudina coriacea — Theel, 1886: 54-55.—Dendy, 1897: 28-32,

pi. 3 figs 9-18.—Dendy, 1898: 456-464, pi. 29,-Farquhar, 1898:

324.—Ludwig, 1898: 63-64.-Dendy and Hindle, 1907: 95, 108-110,

fig. B.—Mortensen, 1925: 364-366, figs 46b, 47b. (synonymy with

Caudina chilensis (Muller) by H. L. Clark 1908)

Caudina coriacea var. brevicauda R. Perrier, 1905: 121-123, fig.

N. (synonymy with Caudina Coriacea by Dendy and Hindle 1907;

with Paracaudina chilensis var. coriacea by H. L. Clark 1935)

Caudina pulchella R. Perrier, 1905: 117-120, pi. 5 figs 14-17.

(synonymy with Caudina coriacea by Dendy and Hindle 1907; with

Paracaudina chilensis var. coriacea by H. L. Clark 1935)

Caudina chilensis.— Benham, 1909: 110. (non Paracuadina

chilensis (Muller, 1850))

Paracaudina chilensis var. coriacea.— H. L. Clark, 1935: 267-

284.

Paracaudina chilensis.— Pawson, 1963: 18-21, pi. 4.—Pawson,

1965: 14.—Pawson, 1970: 49-50, pi. 2 fig. 2.-Pawson, 1977: 119

(part).—Mah et al., 2009: 382, 398, fig. p. 383. (non Paracuadina

chilensis (Muller, 1850))

Material examined. New Zealand, Cook Strait, 61 m, 26 May 1975,

NIWA 70954 (1); Cook Strait, 18 m, 15 Dec 1983, NIWA 70955 (1);

South Island, west coast, Arawhata River mouth, washed onto spit, 8

Jul 1969, AM J12290 (3).

Diagnosis. Paracaudina species up to 172 mm total length, up

to 28 mm diameter, caudal taper and narrow tail 102 mm long

(NIWA 70954); discrete thin tail, frequently longer than main

body; body wall parchment-like, smooth to wrinkled,

frequently with transverse creasing, preserved colour off-white

with patches of yellowish-red (rusty) colouration in largest

specimens; mid-body ossicles predominantly thick, button¬

like, octagonal, with central perforation bridged by a box

basally and cross on the upper side, perforations small, margin

predominantly rounded and lacking projecting knobs and

bluntly pointed projections, surface variably knobbed or with

short thick spines, ossicles typically 64 pm, up to 80 pm long.

Distribution. New Zealand; 0-61 m.

Remarks. In erecting the species Caudina meridionalis (junior

synonym of P. coriacea ) for two specimens from New Zealand

waters, Bell 1883 noted that in comparison with Caudina

ransonnetii Marenzeller, 1881 (junior synonym of P. chilensis )

the ossicles were “stout”, the perforations “small”, and the

marginal projections not as distinct. We agree that these are a

significant diagnostic difference between the predominant

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

ossicle forms of Paracaudina coriacea and Paracaudina

chilensis. We raise Paracudina coriacea (Hutton, 1872) out of

synonymy with Paracaudina chilensis (Muller, 1850).

Perrier 1905 illustrated (pi. 5 figs 16,17) what he considered

to be the predominant ossicle form in his Caudina pulchella

from New Zealand. Perrier 1905 also illustrated two ossicles

(fig. N) for his New Zealand variety Caudina coriacea var.

brevicauda. In both cases these are the thick ossicles with

rounded margin and small perforations that we have found and

illustrated as the predominant ossicles in New Zealand

specimens of Paracaudina coriacea examined in this work.

This form of ossicle was illustrated for New Zealand specimens

by Dendy 1897, Mortensen 1925, Heding 1933 (pi. 6 figs

10-13, New Brighton, Christchurch, specimen), and Pawson

1963 (cups from adult specimen). These are the predominant

ossicles of Paracaudina coriacea (Hutton).

Perrier 1905 illustrated (pi. 4 figs 11,12) what he considered

to be the predominant ossicle form in his Caudina rugosa

from Cape Horn, and they are the ossicles of medium thickness

with prominent blunt marginal projections that we have found

in the Australian and New Zealand specimens of Paracaudina

chilensis (Muller) examined in this work. This form of ossicle

was illustrated by Miiller 1854, Marenzeller 1881, Clark 1908,

Hozawa 1928, Heding 1933 (pi. 6 figs 1-4, type), Pawson and

Liao 1992, Liao and Clark 1995 and Pawson et al. 2001 for

material falling within the synonymy of Paracaudina

chilensis.

We note that Heding 1933 (pi. 6 figs 8-9) illustrated

ossicles from a New Zealand specimen from Tiritiri Matangi

(north of Auckland) that we judge to be typical of Paracaudina

chilensis. If this was the predominant ossicle form the

specimen was Paracaudina chilensis.

Our New Zealand colleagues were unsuccessful in

attempting to find the type material for Molpadia coriacea

Hutton, 1872.

Paracaudina cuprea O’Loughlin and Barmos sp. nov.

Figures 11a, b, 12a, 13

Caudina chilensis.— Joshua, 1914: 6 (part).—Joshua and Creed

1915: 21-22 (part), (non Molpadia chilensis Muller, 1850)

Caudina australis— Mortensen, 1925: 364-367, figs 46c, 47a.

(non Molpadia australis Semper, 1868)

Paracaudina australis.— Hickman, 1962: 63-64, figsl06-130, pi.

2 fig. 9—Rowe, 1982: 472 (part), pi. 32.3.—Rowe and Gates, 1995: 264

(part), (non Molpadia australis Semper, 1868)

Material examined. Holotype. Victoria, Corner Inlet, Sunday Island,

mudflat, MRG, 15 Mar 2004, NMV F157396.

Paratypes. Corner Inlet, Port Welshpool, in sediment, MRG, 5

Mar 2010, NMV F169322 (2).

Other material. Victoria, Seaspray, 8 Mar 1977, AM J10610 (1);

Westernport Bay, San Remo, 2 Apr 1972, NMV F169346 (1);

Shoreham, 30 Mar 1902, NMV F169344 (1) (removed from Shoreham

lot 60669-71 / H19 (3) examined in part by Joshua 1914); Portland

Bay, 27-35 m, 29 Aug 1975, NMV F76073 (1); NMV F174890 (1).

Tasmania, Seven-mile Beach, 13 Aug 1956, AM J7195 (1); Roches

Beach, 6 Mar 1974, AM J8437 (4). South Australia, 24 Jun 1924, SAM

K2504 (1); St Vincent Gulf, mixed localities, Aug 1886, SAM K1381

(13); Port Stanvac, dredged 16 m, 6 Feb 1991, SAM K2495 (12); SAM

K2498 (1); Apr 1991, 17 m, SAM K2503 (1); Brighton Beach, 3 Jul ?

1916, SAM K2506 (2); Port Lincoln, 5 m, 22 Aug 1975, SAM K2492

(1). Western Australia, AM J2341 (1; no additional data); AM J2342 (1;

no additional data); Rottnest I., 146-155 m, 15 Aug 1962, WAM Z8977

(1); 139-145 m, 12 Aug 1962, WAM Z8979 (1); 183-188 m, 14 Aug

1962, WAM Z8981 (1); 146 m, 10 Aug 1962, WAM Z8985 (1).

Diagnosis. Paracaudina species up to 153 mm long (F169346,

preserved), main body 144 mm long, width up to 55 mm, tail 9

mm long; cylindrical body sharply tapered at ends to pointed

oral end, discrete short narrow caudal end / tail; thick, firm,

leathery body wall, smooth, slight wrinkling at oral and anal

ends, variable transverse creasing; live and preserved colour

variably rusty, orange, copper, yellow, some off-white patches;

oval yellow phosphatic bodies present, up to 40 pm long; mid¬

body ossicles irregular, variable, round to oval small plates, flat

to slightly concave, margin and surface smooth or with pointed

spines or knobs, knobs sometimes joined to create secondary

layering, up to 12 irregular perforations, frequently with large

central perforation and or lacking surrounding perforations,

central perforation bridged by 1 or 3 or 4 arms, rarely chilensis-

like and bridged by cross on one side and box on other side,

ossicles up to about 60 pm long.

Type locality. Victoria, Corner Inlet, Sunday Island, intertidal

sediments.

Distribution. Southern Australia, from eastern Victoria

(Seaspray), south to Tasmania, and west to Rottnest Island (off

Perth); 0-188 m.

Etymology. From the Latin cupreus (copper), referring to the

coppery, rusty colour of live and preserved specimens

Remarks. The diagnostic characters that distinguish the new

species Paracaudina cuprea O’Loughlin and Barmos are the

distinctive body form with discrete, short narrow tail, the rusty

and orange colour, absence of mid-body wall rods, and the

predominance of irregular, perforated plate ossicles frequently

with irregularly bridged central perforation and with blunt

marginal and surface spines and knobs.

Joshua 1914 determined a 100 mm long specimen (seen

here, F169344) from “Mordialloc” on Port Phillip Bay as

Caudina chilensis (Muller), but reported that it tapered sharply

posteriorly and could not be described as caudate. Colour was

yellow, blotched with brownish pink. The cross in the ossicles

was frequently lost by fusion with the disc. He was referring to

a specimen of the new species Paracaudina cuprea O’Loughlin

and Barmos. As discussed in the Remarks under P. bacillis

(above) we judge from specimen labels that the specimen

came from Shoreham on Westernport Bay, not “Mordialloc”.

Although they determined specimens from South Australia as

Caudina chilensis (Muller), Joshua and Creed 1915 described

some of them as being up to 125 mm long, lacking the caudate

character of the species, and yellow with patches of rusty red

in colour. They were referring to specimens of the new species

Paracaudina cuprea. Hickman 1962 also described and

illustrated (as P. australis ) the new species Paracaudina

cuprea. Rowe 1982 (pi. 32.3) also illustrated this new species

(as P. australis), and in describing the colour of southern

Australian specimens as “rusty pink and brown” was referring

The paracaudinid sea cucumbers of Australia and New Zealand (Echinodermata: Holothuroidea: Molpadida: Caudinidae)

Figure 13. SEM images of ossicles from the mid-body wall of the holotype specimen of Paracaudina cuprea O’Loughlin and Barmos sp. nov.

from Sunday Island, Corner Inlet, Victoria (NMV F157396).

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

to the new species Paracaudina cuprea. Mortensen 1925

based his discussion of Paracaudina australis on SAM

specimens. Based on Mortensen’s figures we judge that these

specimens were our new species Paracaudina cuprea.

Paracaudina keablei O’Loughlin and Barmos sp. nov.

Figures lie, 14

Material examined. Holotype. Queensland, Moreton Bay, Stradbroke

Island, Dunwich, sandbar, H. Silver, 2 Dec 1978, AM J13579.

Other material. Queensland, Gulf of Carpentaria, SE corner,

17°24'55"S 140°42'35" E, 4.6 m, CSIRO Rama prawn survey trawl stn

555, 16 Jan 1964, AM J17085 (1); 17°30'15"S 140°40'10" E, 2.3 m,

CSIRO Rama prawn survey trawl stn 494, 18 Dec 1963, AM J17051

(3).

Diagnosis. Paracaudina species up to 75 mm total length,

diameter up to 18 mm, caudal taper and short discrete tail 25

mm long (preserved); body wall firm, not thin, smooth to

wrinkled, partly transversely creased, slightly rugose, off-white

(preserved); short caudal taper and discrete thin tail; no

evidence of phosphatising; mid-body ossicles predominantly

thin-walled octagonal plates with large central perforation,

basal box with 4 supporting arms bridging one side of ossicle,

cross with 4 supporting arms bridging outer side, short blunt

spines only around marginal surface and on cross, not on box,

lacking prominent marginal sub-digitiform projections and

knobs, ossicles up to 70 pm across.

Type locality. Northeast Australia, Moreton Bay.

Distribution. Northeast Australia, Moreton Bay, Stradbroke

Island, Gulf of Carpentaria; 0-5 m.

Etymology. Named for Dr Stephen Keable, Collection Manager,

Marine Invertebrates (Natural Science Collections), Australian

Museum, in appreciation of his prompt and gracious assistance

with loans from the Australian Museum for this and other

research projects.

Remarks. The holotype specimen of Paracaudina keablei

O’Loughlin and Barmos sp. nov. is damaged, but the ossicles

are in good condition. The ossicles in the additional material

are somewhat eroded, but the predominant ossicle form is

diagnostically distinguishable. Four Paracaudina Heding

species have a predominant ossicle form of octagonal plates

with a basal box with four supporting arms bridging the ossicle

on one side and a cross with four supporting arms bridging the

ossicle on the opposite side. The predominant ossicle form in

Paracaudina chilensis (Muller) is of moderate thickness with

prominent knobs and sub-digitiform projections around the

margin. The predominant ossicles in Paracaudina coriacea

(Hutton) are thick buttons with rounded margin and small

perforations. Parcaudina keablei sp. nov. is distinguished from

both these species by having a predominant ossicle form of

thin-walled plates with large perforations and lack of prominent

marginal projections. The ossicles in Paracaudina delicata

Pawson and Liao, 1992 are also thin-walled, but have numerous

fine digitiform projections around the margin. Paracaudina

keablei is further distinguished from the other three species

mentioned here by having a short thin tail.

Paracaudina luticola Hickman, 1962

Figures lid, 15

Caudina chilensis— Joshua and Creed 1915: 21-22 (part) (non

Molpadia chilensis Muller, 1850).

Paracaudina luticola Hickman, 1962: 65-66, figs 131—139.—

Hickman, 1978: 32, figs 25-44, pi. 2.—Pawson, 1977: 119—Rowe,

1982: 471, fig. 10.35a.—Rowe and Gates, 1995: 265.

Material examined. Syntypes. Tasmania, Derwent Estuary, Ralph’s

Bay, 13 m, 30 Jun 1959, AM J7205 (2).

Other material. Victoria, Shallow Inlet, mud/sand seagrass,

intertidal, 2 Feb 1990, NMV F169342 (2); Wilson’s Promontory,

Waratah Bay, Sandy Point, 31 Mar 1969, NMV F76072 (1). South

Australia, SAM K1379 (1); Aug 1886, SAM K1380 (1); St. Vincent

Gulf, donated by SAM, NMV F45018 (3; labelled as determination by

Joshua and Creed in 1915 as Caudina chilensis ); St Vincent Gulf,

mixed localities, Aug 1886, SAM K2484 (1); Adelaide Outer Harbour,

Feb 1935, SAM K2505 (1); Port Stanvac, 6 Feb 1991, SAM K2499 (1);

Henley Beach, SAM K2488 (1); Brighton to Semaphore, SAM K2494

(2); Largs Bay beach, 26 Aug 1971, SAM K2508 (1); Edithburgh,

Sultana Point, sand bar, 8 Nov 2003, SAM K2489 (1); Sir Joseph

Banks Group, Marum I., in Posidonia , 12 m, 11 Jan 1984, SAM K2497

(1); Port Lincoln, 22 Aug 1975, AM J9466 (1); Venus Bay, 1982, SAM

K2500 (4); Edward Bay, N of Streaky Bay, 23 Oct 1986, SAM K2483

(1). Western Australia, Bremer Bay, near Albany, on beach after storm,

5 Aug 1984, WAM Z31887 (1).

Diagnosis. Paracaudina species up to 137 mm total length, up

to 19 mm diameter, caudal taper and discrete thin tail 52 mm

long (up to 160 mm long live, Hickman 1978); long thin discrete

tail; body wall thin, firm, parchment-like, variably wrinkled

and transversely creased, preserved colour off-white to faint

pink to grey; ossicles variable in form, similar from mid-body

and tail, predominantly thick crossed cups, oval in form, large

central perforation, lacking peripheral perforations, about 10-

12 projecting marginal knobs, fewer marginal surface knobs,

central perforation spanned by discrete cross, each arm of cross

with rounded knob-like end; ossicles never with bridging cross

on one side and box on other side of central perforation; ossicles

up to about 56 pm long.

Type locality. Tasmania, Derwent Estuary, Ralph’s Bay, 13 m.

Distribution. Southern Australia; southern Tasmania, north to

Shallow Inlet (east of Wilson’s Promontory, Victoria), west to

Streaky Bay (west side of Eyre Peninsula in South Australia)

and Bremer Bay (near Albany in Western Australia); 0-13 m.

Remarks. Hickman 1962 erected this species for caudal part-

specimens only, but subsequently (1978) described whole

specimens from the type locality. Ossicle size and form do not

vary for body wall tissues taken from the main body and tail.

The predominant ossicle form of thick four-holed cups with

marginal and surface knobs and discrete cross is diagnostically

distinctive for Paracaudina luticola Hickman, 1962. Joshua

and Creed 1915 determined specimens from South Australia to

be Caudina chilensis (Muller) that showed a “very great

variation”. We judged above, from their description, that some

of the larger specimens were Paracaudina cuprea O’Loughlin

and Barmos. We judge here, from their description of “about 70

mm long, white colour, markedly caudate discrete tail about

The paracaudinid sea cucumbers of Australia and New Zealand (Echinodermata: Holothuroidea: Molpadida: Caudinidae)

Figure 14. SEM images of ossicles from the mid-body wall of the holotype specimen of Paracaudina keablei O’Loughlin and Barmos sp. nov.

from Dunwich, Stradbroke Island, Moreton Bay, Queensland (AM J13579).

half the length of the whole specimen”, that some of the

specimens were Paracaudina luticola Hickman.

Paracaudina tetrapora (H. L. Clark, 1914)

Figures lie, f, 16, 17

Caudina chilensis.— Joshua, 1914: 6 (part) (non Molpadia

chilensis Muller, 1850).

Caudina tetrapora H. L. Clark, 1914: 170, fig. 1.

Paracaudina tetrapora— H. L. Clark, 1935: 267-284.—H. L.

Clark, 1938: 541.-H. L. Clark, 1946: 445,-Pawson, 1977: 119.—

Rowe, 1982: 472.—Rowe and Gates, 1995: 265.

Material examined. Western Australia, Perth, Kwinana beach, Oct

1958, WAM Z31886 (1); South Cottesloe, 8 Nov 2008, WAM Z31883

(1). South Australia, Spencer Gulf, Sir Joseph Banks Group, Marum I.,

sand, 5 m, 8 Jan 1984, SAM K2496 (1); St. Vincent Gulf, North Haven

to Largs Jetty, seagrass, 1 Dec 1980, SAM K2501 (1); Brighton Beach,

14 Feb 1975, SAM K2507 (2); off Glenelg, 10 m, 15 Feb 1969, AM

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 15. SEM images of ossicles from the mid-body wall of a specimen of Paracaudina luticola Hickman, 1962 from Sandy Point, Waratah

Bay, Victoria (NMV F76072).

J24918 (1); mixed localities, Aug 1886, SAM K2485 (2). Victoria,

Westernport Bay, Merricks, 29 Jan 1983, NMV F76565 (1); Flinders,

beach, Jul 1967, NMV F45259 (1); Shoreham, 30 Mar 1902, NMV

F45019 (remaining specimen from Shoreham lot 60669-71 / H19 (3)

examined in part by Joshua 1914); Merricks, beach, Apr 1969, NMV

F45274 (1).

Diagnosis. Paracaudina species up to 116 mm total length, up

to 24 mm diameter, caudal taper and thin tail combined length

53 mm (preserved; WAM Z31886); firm leathery to parchment¬

like body wall with slight wrinkling; off-white to cream

preserved colour, some yellow to yellowish-red colouration,

some phosphatising with yellow colouration around eroding

ossicles; posterior body with distinct long caudal taper to

narrow rounded end, sometimes tapering to a discrete long thin

tail; mid-body ossicles predominantly 4-holed thick buttons,

also thick perforated plates and bluntly spinous cups; 4-holed

buttons irregularly oval to rectangular, some shallow concave,

frequently 4 perforations with 2 large central and 2 small distal,

variably thickened and marginally knobbed, up to 48 }i m long;

some thick irregularly oval to round perforated plates with up

to 15 small perforations, margin and surface variably lumpy

and knobbed and with blunt pointed projections, rare joining of

knobs to create secondary layering, thickened plates up to 72

//m long, more evident in larger specimens; shallow cups with

4 perforations, long blunt thick marginal spines, marginal

surface knobs, up to 72 pim long, more evident in larger

The paracaudinid sea cucumbers of Australia and New Zealand (Echinodermata: Holothuroidea: Molpadida: Caudinidae)

Figure 16. SEM images of ossicles from the mid-body wall of a specimen of Paracaudina tetrapora (H. L. Clark, 1914) from Merricks,

Westernport Bay, Victoria (NMV F76565).

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 17. SEM images of ossicles from the mid-body wall of a specimen of Paracaudina tetrapora (H. L. Clark, 1914) from off Glenelg, St.

Vincent Gulf, South Australia (AM J24918).

The paracaudinid sea cucumbers of Australia and New Zealand (Echinodermata: Holothuroidea: Molpadida: Caudinidae)

Figure 18. Photos of Paracaudina tripoda O’Loughlin and Barmos, sp. nov. (photos by L. Altoff, MRG). a, holotype from Queensland, east of

Lady Elliot Island, 230 m (AM J24922); b, small paratype, north of Fraser Island, 137 m, (AM J11150 (7)).

specimens; ossicles never chilensis-like with central perforation

with box and cross bridges.

Type locality. Western Australia, Perth, Cottesloe beach.

Distribution. Southern Australia, from Abrolhos Islands,

Western Australia, to Westernport Bay, Victoria; to 10 m.

Remarks. Paracaudina tetrapora (H. L. Clark, 1914) is

distinguished amongst Australian species of Paracaudina by

having ossicles that are thick knobbed irregularly oval buttons

with predominantly 2 large central and 2 distal smaller holes.

Joshua 1914 reported two specimens from Victoria as Caudina

chilensis (Muller). Details provided for the larger one (and

confirmed here, NMV F169344) indicate that it was

Paracaudina cuprea O’Loughlin and Barmos sp. nov. (above).

The second specimen (seen here, NMV F45019) was

Paracaudina tetrapora (H. L. Clark). Yellow body wall

colouration with phosphatising of ossicles and phosphatic body

wall residues is evident in larger specimens. In the largest

specimen (SAM K2507) there is dark red phosphatising of the

calcareous ring, and the reddish-brown colouration of the

polian vesicle is presumed to be due to phosphatising.

Paracaudina tripoda O’Loughlin and Barmos sp. nov.

Figures 18, 19

Material examined. Holotype. Queensland, east of Lady Elliot Island,

24°07'S 152°52'E, 230 m, 7 Jul 1984, AM J24922.

Paratypes. Type locality and date, AM J18814 (3); east of Lady

Elliot Island, 24°04'S 152°48'E, 192 m, 7 Jul 1984, AM J18813 (1);

north of Fraser Island, 24°23'S 153°17'30"E, 137 m, 15 Dec 1977, AM

Jill50 (7).

Diagnosis. Paracaudina species up to 75 mm total length,

diameter up to 27 mm; oral and caudal tapers, lacking long thin

tail, sometimes short discrete tail; body wall firm, leathery,

slightly rugose; small preserved specimens reddish-yellow

(rusty), larger specimens off-white; reddish-yellow pigment

may be present; mid-body with abundant irregularly round to

oval, thick button-like ossicles, some slightly concave, up to 64

pm long; majority of ossicles with rounded edge, short thick

surface spines, ossicles frequently with large central perforation

with tripod or single rod or cross bridge, up to 11 variably sized

perforations; many australis- like plates with horizontal

marginal projections; some chilensis- like plates with bridging

box and cross over central perforation, some with bridging

tripod; rare ambigua- like mid-body plate ossicles with 2-4

perforations, up to 32 pm long.

Type locality. Northeast Australia, Queensland, E of Lady

Elliot Island, 230 m.

Distribution. Northeast Australia, off Lady Elliot and Fraser

Islands; 137-230 m.

Etymology. From the Greek trion (three) and podos (foot),

referring to the frequency of a tripod bridge over the central

perforation in the ossicles.

Remarks. The holotype specimen of Paracaudina tripoda

O’Loughlin and Barmos sp. nov. is damaged, but the ossicles

are in good condition and the body form is evident. Three

Paracaudina Heding, 1932 species lack a long thin tail and

have irregular small plate or button ossicles in the mid-body

wall. In two of these species, Paracaudina australis (Semper,

1868) and Paracuadina cuprea O’Loughlin and Barmos sp.

nov., the predominant ossicles from the mid-body are plates

with lateral rounded marginal spines and surface spines and

knobs. In the new species Paracaudina tripoda O’Loughlin

and Barmos the predominant ossicles lack lateral rounded

marginal spines, and have surface spines. Paracaudina

australis specimens show rare slight yellowing, and have rods

in the mid-body wall. Paracaudina cuprea specimens show

strong reddish-yellow colouration with increasing size.

Paracaudina tripoda specimens show decreasing reddish-

yellow colouration with increasing size.

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

Figure 19. SEM images of ossicles from the mid-body wall of the holotype of Paracaudina tripoda O’Loughlin and Barmos sp. nov. (AM

J24922).

The paracaudinid sea cucumbers of Australia and New Zealand (Echinodermata: Holothuroidea: Molpadida: Caudinidae)

Figure 20. SEM images of ossicles from the mid-body wall of a 35 mm long specimen of a species of Paracaudina Heding 1932 from the

Chatham Rise (NIWA 70957).

Paracaudina species (not described)

Figure 20

Material examined. East of New Zealand, Chatham Rise, 43.00°S

179.00°W, 546 m, 12 Sep 1989, NIWA 70957 (1); outer east Cook

Strait, 41.70°S 175.65°E, 1040-1080 m, 15 Apr 2010, NIWA 63034 (1);

41.51°S 175.72°E, 1076-1104 m, 17 Apr 2010, NIWA 63195 (1);41.95°S

174.62°E, 964-1005 m, 23 Apr 2010, NIWA 63806 (5).

Remarks. Our colleagues in NIWA (Wellington) inform us of

many Paracaudina specimens in their collections. We have had

the opportunity to examine only eight. Mid-body ossicles from

one small specimen (35 mm long; NIWA 70957) are illustrated

here. They are not those of Paracaudina coriacea (Hutton, 1972)

or Paracaudina chilensis (Muller, 1850). Nor do the ossicles in

the eight specimens appear to be the ossicles of only one species,

those from the Chatham Rise specimen differing from those

from the outer east Cook Strait specimens. A comprehensive

examination of the NIWA Paracaudina specimens will be

undertaken after this work, presumably in Wellington.

Acknowledgments

We are grateful for the generous assistance of the following:

Leon Altoff and Audrey Falconer (MRG, photos of specimens);

John and Sue Barmos (collection of specimens); Ben Boonen

(photoshop and format of figures); Niki Davey, Sadie Mills

and Kareen Schnabel (NIWA, facilitation of loans, and search

for Hutton type material); Julian Finn (NMV, collection and

photos of live specimens); Jane Fromont and Mark Salotti

(WAM, facilitation of loans); Glenys Greenwood (photos of

live specimens); OlgaHionis (NMV, assistance with literature);

Stephen Keable, Helen Stoddart and Roger Springthorpe (AM,

facilitation of loans); David Lane, Peter Ng and Ria Tan

(Singapore, dialogue and photos of relevant Singapore species);

Thierry Laperousaz (SAM, facilitation of loans); Darryn

McKenzie (Reef Watch Victoria, photo of spawning specimen);

Mark Norman (NMV, photos of live specimens); Mike Reich

(University of Gottingen, assistance with literature); Wendy

Roberts (Reef Watch Victoria, collection of specimens); Chris

Rowley (NMV, assistance with sending tissue samples and

P.M. O’Loughlin, S. Barmos & D. VandenSpiegel

with curation of specimens);Yves Samyn (Royal Belgian

Institute of Natural Science, assistance with literature); David

Staples (NMV, assistance with photography); Platon Vafiadis

(MRG, photo of live specimens); Rick Webber (Te Papa

Tongarewa, New Zealand, search for Hutton type material).

We are grateful for the supportive and helpful reviews by

David Pawson (Smithsonian Institution) and Frank Rowe

(formerly of the Australian Museum).

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Memoirs of Museum Victoria 68: 67-70 (2011)

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

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

What is Alloxysta fulviceps (Curtis, 1838) (Hymenoptera: Cynipoidea: Figitidae:

Charipinae)?

JULI PUJADE-VILLAR 1 , MAR FERRER-SUAY 1 , JESUS SELFA 2 AND MlGUEL A. ALONSO-ZARAZAGA 3

1 Universitat de Barcelona, Facultat de Biologia, Departament de Biologia Animal, Avda. Diagonal, 645,08028-Barcelona

(Spain) (jpujade@ub.edu; mar.ferrer.suay@gmail.com)

2 Universitat de Valencia, Facultat de Ciencies Biologiques, Departament de Zoologia, Campus de Burjassot-Paterna, Dr.

Moliner 50,46100-Burjassot (Valencia, Spain) (jesus.selfa@uv.es)

3 Museo Nacional de Ciencias Naturales, Departamento de Biodiversidad y Biologia Evolutiva, Jose Gutierrez Abascal, 2.

28006-Madrid (Spain) (zarazaga@mncn.csic.es)

Abstract Pujade-Villar, J., Ferrer-Suay, M., Selfa, J. and Alonso-Zarazaga, M.A. 2011. What is Alloxysta fulviceps (Curtis, 1838)

(Hymenoptera: Cynipoidea: Figitidae: Charipinae)?. Memoirs of Museum Victoria 68: 67-70.

The validity of Alloxysta fulviceps (Curtis, 1838) and its synonymies have been examined. After studying the type

series and the taxonomic history of this species, we conclude that the lectotype was wrongly designated. A new lectotype

is designated and A. fulviceps is synonymised with A. victrix (Westwood, 1833).

Keywords Hymenoptera, Figitidae, Charipinae, Alloxysta,fulviceps

Introduction

The Charipinae (Hym., Figitidae) species Alloxysta fulviceps

(Curtis, 1838) has been a troublesome species since its

description. Curtis (1838, p. 688) described it as a species of

Cynips with the following features: ‘27. fulviceps Curt. Shining

black, head and legs bright ochre; antennae longer than the

body, fuscous, base ochreous, 2 basal joints ovate, 3rd slender,

scarcely longer than the following, wings very ample: 2/3 long.

Bred from female Aphides by the late Mr. T. Carpenter’.

Later, it was transferred to Allotria by Kieffer (1900, p.

114). According to Kieffer (1902, p. 76), this species was

considered as doubtful and insufficiently described. It was

kept in Allotria by Dalla Torre and Kieffer (1902, p. 41), and

finally it was transferred by Dalla Torre and Kieffer (1910, p.

288) to Charips.

Quinlan and Fergusson (1981) mentioned that the type

series deposited in Museum Victoria (Australia) was studied

by Kerrich in 1948. According to these authors, this type

series consisted of three specimens, and Kerrich defined one

of them as ‘type of Cynips fulviceps = Alloxysta fulviceps ’.

These data were not published and Fitton (1978, p. 65) was the

first to consider Curtis’s species in Alloxysta.

According to Quinlan and Fergusson (1981), the type

series of A. fulviceps grouped two distinct morphologies: 1 9

with Kerrich’s notes (with open radial cell) and 2 9 without

Kerrich’s notes (with closed radial cell). These two specimens

were labelled by Quinlan with the following notes: Alloxysta

minutal (Hrt.) 9 det. Quinlan 1986’ and ‘this specimen has not

type-status’. Since then, Curtis’s species has been a species

with an open radial cell (a feature not mentioned by Curtis in

his description).

In the same study, Quinlan and Fergusson (1981)

synonymised Alloxysta erythrothorax (Hartig, 1840) with A.

fulviceps. This synonymy has two problems: (1) the choice as

lectotype of the specimen with an open radial cell, when Curtis

did not mention the morphology of the radial cell, and (2) the

distinct chromatic features that are mentioned in the original

description of Alloxysta fulviceps (Curtis) and A. erythrothorax

(Hartig). Hartig (1840, p. 200) defined Xystus erythrothorax as

follows: ‘5) X. erythrothorax m.: niger, capite rufo; facie flava,

pectore rufo; antennis pedibusquie rufis. Male’.

Results and discussion

None of the aforementioned authors did a redescription of Curtis’s

species, so we borrowed the type material from Museum Victoria.

Only one of the three original specimens remains (Catriona

McPhee, Collection Manager Terrestrial Invertebrates (Mon-

Wed) of the Museum Victoria, pers. com. on 21 Feb. 2011): one

of the two of the original series (before Kerrich’s study) that has a

closed radial cell. We are not able to determine how Kerrich chose

a lectotype of Cynips fulviceps, but after Quinlan’s study and the

synonymy proposed by Quinlan and Fergusson (1981), this

specimen would have to have a reddish yellow mesoscutum.

Therefore, it cannot be the lectotype of A. fulviceps because Curtis

J.Pujade-Villar, M. Ferrer-Suay, J. Selfa & M.A. Alonso-Zarazaga

Figure 1 . Lateral view of the designated lectotype of Alloxysta fulviceps (Curtis).

stated that the mesoscutum is completely black. For these reasons,

we must consider that the specimen that was chosen as lectotype

does not fit the original description by Curtis. On the other hand,

we have examined the one remaining specimen of Curtis’s type

series, which corresponds exactly with Curtis’s description. In

summary, we designate here a new lectotype for Alloxysta

fulviceps (= Cynips fulviceps Curtis), which has the following

labels: ‘ Alloxysta minutal (Hrt.) 9 det. Quinlan 1986’ (white

label), ‘this specimen has not type-status’ (white label), ‘MUS.

VIC. ENTO 2011-4-L’ (green label), ‘this specimen has type

status JP-V, 2011’, ‘27 Cynips fulviceps bred from Aphides female

by Aphidius? T.C. 13 July 26 — [unreadable] Dorset. Bred from

aphids of Willows and Cow Parsnip Hal’ (photocopy of Curtis

original annotations of his notebook), ‘lectotype of Cynips

fulviceps Curtis, 1838 9 , designed J.P-V 2011’ (red label),

‘Alloxysta victrix (Westwood, 1833) Ferrer-Suay det-2011 (white

label).

Our study of this material also shows that A. fulviceps is

the same species as A. victrix. Therefore, in this study, the

synonymy of A. fulviceps and A. erythrothorax is removed, a

new lectotype for A. fulviceps is established, this species is

synonymised with A. victrix and the validity of A. erythrothorax

(Hartig) is re-established.

The taxonomic changes we propose restructure the list of

synonyms and valid species implied, so we present it here:

Alloxysta erythrothorax (Hartig, 1840) Dalla Torre and

Kieffer, 1902

Xystus erythrothorax (Hartig, 1840, p. 200). Synonymised by

Quinlan and Fergusson (1981, p. 254). Type: ZSBS (according to

Evenhuis, 1982, p. 23).

Xystus defectus (Hartig, 1841, p. 352). Synonymised by Fergusson

(1986, p. 10). Type: ZSBS (according to Evenhuis, 1982, p. 22).

Allotria nigriventris (Thomson, 1862, p. 409). Synonymised by

Fergusson (1986, p. 10). Type: MZLU (according to Andrews, 1978, p.

87).

Comments : Hellen (1963) redescribed A. erythrothorax and A.

defecta. The latter was synonymised with the former by

Fergusson (1986). The two species match chromatically.

Fergusson (1986) also synonymised A. nigriventris (Thomson)

What \sAlloxysta fulviceps (Curtis, 1838) (Hymenoptera: Cynipoidea: Figitidae: Charipinae)?

with A. erythrothomx, and according to Kieffer (1902), the

mesosoma of A. nigriventris is reddish; this also matches A.

erythrothomx.

Alloxysta victrix (Westwood, 1833) Hellen, 1963

Allotria victrix (Westwood, 1833, p. 495). Type: OXUM (according

to Andrews, 1978, p. 92).

Cynips fulviceps (Curtis, 1838, p. 688). Type: NMVM n. syn.

Cynips ruficeps (Zetterstedt, 1838, p. 410). Synonymised by Giraud

(1860, p. 127). Type: MZLU (according to Evenhuis and Kiriak, 1985, p.

16).

Xystus erythrocephalus (Hartig, 1840, p. 199). Synonymised by

Giraud (1860, p. 127). Type: ZSBS (according to Evenhuis, 1972, p. 211).

Allotria tritici (Fitch, 1861, p. 841). Synonymised by Menke and

Evenhuis (1991, p. 147). Type: USNM (according to Menke and

Evenhuis, 1991, p. 147).

Allotria ( Allotria ) macrocera (Thomson, 1877, p. 814). Synonymised

by Dalla Torre and Kieffer (1910, p. 285). Type: MZLU (according to

Hellen, 1963, p. 4).

Allotria curvicornis (Cameron, 1883, p. 366). Synonymised by

Fergusson (1986, p. 11). Type: BMNH (according to Quinlan, 1978, p.

124).

Allotria ( Allotria ) luteicornis (Kieffer, 1902, p. 15). Synonymised by

Evenhuis and Barbotin (1987:, p. 217). Type: NHM, Amiens (according

to Dessart, 1969, p. 193).

Allotria ( Allotria ) victrix var luteiceps (Kieffer, 1902, p. 16.

Synonymised by Evenhuis and Barbotin (1987, p. 217). Type: NHM,

Amiens (according to Dessart, 1969, p. 194).

Allotria ( Allotria) luteicornis var lateralis (Kieffer, 1902, p. 70).

Synonymised by Evenhuis and Barbotin (1987, p. 216). Type: NHM,

Amiens (according to Dessart, 1969, p. 193).

Charips areolata (Kieffer, 1909, p. 481). Synonymised by Menke

and Evenhuis (1991, p. 145). Type: USNM (according to Menke and

Evenhuis, 1991, p. 145).

Sarothrus io (Girault, 1932, p. 3). Synonymised by Paretas-Martmez

and Pujade-Villar (2010, p. 355). Type: QM (according to Paretas-

Martmez and Pujade-Villar, 2010, p. 355).

Acknowledgements

We want to thank sincerely K. Walker and C. McPhee (Museum

Victoria, Australia) for the comments about the Curtis collection

and for sending us the type material of Cynips fulviceps treated

in this study. We also thank J. LaSalle (Commonwealth

Scientific and Industrial Research Organisation, Australia) and

J. Paretas-Martmez (UB, Catalunya) for providing us with the

names of those responsible for the Museum Victoria collections.

References

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importance. Entomologische Berichten 32: 210-217.

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of Entomology 49: 354-358.

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Memoirs of Museum Victoria 68:71-91 (2011)

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

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

Review of the genus Monotheca (Hydrozoa: Leptolida) from Australia with

description of a new species and a note on Monothecella Stechow, 1923

Jeanette E. Watson

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

Australia (hydroid@bigpond.com)

Abstract Watson, J.E. 2011. Review of the genus Monotheca (Hydrozoa: Leptolida) from Australia with description of a new species

and a note on Monothecella Stechow, 1923. Memoirs of Museum Victoria 68: 71-91.

Monotheca Nutting, 1900 is a genus with few known species worldwide. It is an artificial genus, as some species of

the Plumulariidae sometimes have one or two hydrocladial hydrothecae and are thus borderline between Monotheca and

Plumularia. This review considers only Australian species with consistently monothecate hydrocladia; species with more

than one hydrotheca on the hydrocladium are considered to belong to Plumularia. Ten species referrable to Monotheca are

reported from southeast Australia; three are synonymised here in other species, four are presently known to be endemic to

Australia and one species. Monotheca amphibola, is described as new. The gonosome of Monotheca togata is described

for the first time. Monothecella is synonymised in Monotheca. A key to the Australian species of Monotheca is given.

Keywords Monotheca, artificial genus, eight Australian species, key to species; Monothecella synonymised.

Introduction

Monotheca Nutting, 1900 (type species Monotheca margaretta

Nutting, 1900) is an artificial genus; Vervoort and Watson

(2003) list 10 species and Bouillon et al. (2006) list eight

species worldwide. Other authors, such as Millard (1975) and

Hirohito (1995), synonymised Monotheca in Plumularia. In

their phylogeny, Leclere et al. (2007) demonstrated that

Monotheca with monohydrothecate hydrocladia ( M.pulchella,

M. margaretta, M. obliqua) form a distinct clade within the

Plumulariidae.

Species of Monotheca reported from Australia are

Monotheca obliqua (Johnston, 1847), Monotheca australis

(Kirchenpauer, 1876), Monotheca compressa (Bale, 1882),

Monotheca hyalina (Bale, 1882), Monotheca pulchella (Bale,

1882), Monotheca spinulosa (Bale, 1882), Monotheca aurita

(Bale, 1888), Monotheca flexuosa (Bale, 1894), Monotheca

obesa (Blackburn, 1938) and Monotheca togata (Watson,

1973). Four of these species — M. hyalina, M. pulchella, M.

spinulosa and M. togata — are also reported from New

Zealand (Vervoort and Watson 2003). A new species,

Monotheca amphibola sp. nov., from seagrass habitat in

southeast Australia, is described here. Some species assigned

to Monotheca — for example, Plumularia excavata (Mulder

and Trebilcock, 1910), Plumularia crateriformis (Mulder and

Trebilcock, 1910), Plumularia epibracteolosa Watson, 1973

and Plumularia meretricia Watson, 1973 sometimes have two

hydrothecae on the hydrocladium and are thus borderline

between Monotheca and Plumularia as defined here. In this

review, only strictly monohydrothecate species are considered,

and species that sometimes have more than one hydrotheca on

the hydrocladium are referred to Plumularia.

Although several species of Monotheca from Australia

discussed in this paper as presently known are endemic to

Australia there are some exceptions. These include M. obliqua

known from the Posidonia oceanica seagrass meadows of the

Mediterranean Sea (Boero 1981a, Boero et al. 1985; Fresi et al.

1982, Bouillon et al. 2004), the eastern Atlantic (Calder 1997)

and Japan (Hirohito 1983); M. spinulosa is known from South

Africa, the South Atlantic (Millard 1975) and Japan (Yamada

1959, Hirohito 1995); and M. flexuosa (as M. pulchella, see

later discussion) is reported from the Mediterranean Sea and

South Africa (Millard 1975). A key to species of Monotheca is

given.

With the exception of Monotheca flexuosa, which is an

opportunistic species occurring on many substrates, all

Australian species of Monotheca are epiphytic — M. obliqua,

M. spinulosa and M. hyalina are associated with algae, while

M. australis, M. compressa, M. obesa and M. amphibola sp.

nov. occur on seagrasses. New Zealand species recorded as

M. hyalina and M. flexuosa (see later discussion concerning

their identity) are from algae.

Monothecella Stechow, 1923.

Stechow (1923a) erected the genus Monothecella for three

species {Monotheca australis Kirchenpauer, 1876; Monotheca,

aurita Bale, 1888; Monotheca compressa Bale, 1882), all of

J.E. Watson

which have a hydrocladial median inferior nematotheca on a

stout immovable base. The term monothalamic is usually

applied to this structure, assuming that the distal cup is openly

contiguous with its supporting peduncle. Detailed examination

of fresh, preserved and mounted material of this suite of

species (synonymised in M. australis, see discussion later)

clearly shows that the cup is seldom completely open, and

there is usually a faint line of demarcation between the cup

and its base. The term monothalamic is therefore abandoned

in the present context and the nematothecae regarded as

incipiently bithalamic. For this reason, the genus Monothecella

is untenable and is here synonymised in Monotheca.

Material.

In the following account, a large amount of material collected

over four decades by the author and others around Australia is

examined. Material is held in the collections of Museum

Victoria, Melbourne, Australia (NMV), the Australian

Museum, Sydney (AM), the South Australian Museum (SAM),

the Western Australian Museum (WAM), the National

Museum of New Zealand (NMNZ) and in the author’s personal

collection.

Family Plumulariidae McCrady, 1859

Monotheca Nutting, 1900

Colony small, caulus erect, monosiphonic, simple or sparingly

branched, divided into internodes without hydrothecae.

Hydrocladia alternate, consisting of two internodes, the distal

one bearing a hydrotheca with one median inferior nematotheca

and two superior lateral nematothecae seated on a distal

enlargement or a bifurcation of the internode. Gonangia fixed

sporosacs, usually borne on proximal part of the stem, ovate or

sac shaped, without protective appendages.

Monotheca amphibola sp. nov.

Fig. 1A-G

Material examined. NMV F147479, holotype, microslide (malinol

mounted), fertile colony from leaves of the seagrass Amphibolis

antarctica, 200 m offshore from Queenscliff, Victoria, depth 2 m,

coll: J. Watson 23 March 2008. NMV F147481, remainder of holotype

colony, alcohol preserved. Paratype, NMV F147480, microslide

(malinol mounted), fertile colony on leaves of the seagrass Amphibolis

antarctica, 200 m offshore from Queenscliff, Victoria, depth 2 m,

coll: J. Watson 23 March 2008.

Description from holotype and paratype. Stems to 3.5 mm

long, monosiphonic, arising at regular intervals from hydrorhiza

running more or less straight up seagrass leaf; stolons wide,

ribbon-like with a narrow perisarcal flange with numerous

internal flexion joints. Proximal stem with four or five strong

transverse joints, succeeding cauline internodes variable in

length, longer on taller stems, perisarc smooth and thick with

several strong, more or less equally spaced transverse internal

septa, nodes broad V-shaped overlapping joints.

Hydrocladia alternate, distal on cauline internode, on a

short, strong apophysis, directed almost perpendicularly

outward from stem, distal node of apophysis slightly oblique

to transverse; proximal hydrocladial internode short, athecate,

distal node strongly oblique.

Hydrothecate internode moderately long and deep with

two distinct internal opposing septa; hydrotheca distal on

internode, facing upward at an angle of c. 45° to hydrocladium;

cup shaped, slightly broader than deep, adcauline wall weakly

concave to straight, abcauline wall distinctly concave, margin

circular, entire, rim not everted, perisarc thin, a large sinuous

abcauline perisarcal flange passing from margin downwards

to beyond base of hydrotheca; a smaller triangular adcauline

flange passing from hydrothecal margin to hydrocladium.

Nematothecae small, bithalamic, one on cauline internode,

about one-third distance up internode and on same side as

hydrocladium, base moderately long, adpressed to internode,

cup slightly adcaudally excavated, one axillar, on front of stem

beside apophysis, cup slightly elliptical, one hydrocladial

median inferior on hydrothecate internode, not moveable, base

long, stout and bent, cup small, adcaudally excavated and

adpressed to flange of internode, not quite reaching base of

hydrotheca; twin laterals standing erect, distal on hydrocladium

below hydrothecal flange but not reaching hydrothecal margin,

cups slightly adcaudally shortened; stolonal nematothecae

scattered along hydrorhiza, base long, slightly tapering, cup

circular, shallow.

Male and female gonothecae often borne on opposite sides

of same caulus, large, similar in shape, irregularly ovoid,

borne on a pedicel inserted below proximal stem apophysis;

some gonothecae recumbent to substrate, others standing out

perpendicular to stem; walls of gonotheca smooth, no distinct

operculum, a thin diagonal apical fold usually marking site of

future rupture; female gonophore packed with many

moderately large ova.

Monotheca amphibola, measurements (/tm)

Hydrorhiza, width

160-170

Stem internode

length

272-360

width at node

36-40

Apophysis, adcauline length

32-56

diameter at distal node

40-52

Hydrocladium

length adcauline athecate segment

32-36

basal length of thecate internode

176-200

Hydrotheca

length, base to margin along axis

100-108

diameter at margin

116-128

Nematotheca

cauline, length of base

40-50

cauline, depth of cup

28-20

cauline, diameter of cup

28-32

axillar, length of base

30-36

axillar, depth of cup

12-14

axillar, diameter of cup

28-34

Review of the genus Monotheca (Hydrozoa: Leptolida) from Australia with description of a new species and a note on

Monothecella Stechow, 1923

Figure 1. A-G, Monotheca amphibola sp. nov. (drawn from holotype and paratypes). A, fertile stem with male and female gonothecae; B,

hydrocladium and hydrotheca; C, median inferior nematotheca; D, cauline nematotheca; E, one of twin lateral nematothecae; F, axillar

nematotheca; G, hydrorhizal nematotheca. H-N, Monotheca australis. H, infertile stem; I, hydrocladium and hydrotheca; J, female gonotheca

with developing larva; K, median inferior nematotheca. Note clear demarcation between cup and peduncle (I) and its absence (K); L, axillar

nematotheca; M, one of twin lateral nematothecae; N, axillar hydrostatic pore (axillar nematotheca not shown). Scale bar: A, 1 mm; B, I, H, 0.2

mm; C-G, K-N, 0.1 mm; J, 0.5 mm.

J.E. Watson

median, length of base

34-40

median, depth of cup

12-16

median, diameter of cup

30-34

lateral, length of base

38-42

lateral, depth of cup

10-16

lateral, diameter of cup

36-40

hydrorhizal, length of base

98-100

hydrorhizal, depth of cup

30-32

hydrorhizal, diameter of cup

28-36

Remarks. The Mulder and Trebilcock collection in Museum

Victoria contains five microslides (Canada Balsam-mounted)

labelled ‘ Plumularia setaceoides var. crateriformis, 1910’. Two

of these slides are labelled ‘type’ by Mulder and Trebilcock

(see Mulder and Trebilcock 1910, p. 118). The locality of the

specimens is Bream Creek on the central Victorian coast.

Unfortunately, the specimens are so badly fragmented that it is

impossible to determine whether they were originally

Plumularia or Monotheca. These, and a third slide from the

same series (NMV F57987, F57988, F57989), were designated

syntypes by Stranks (1993). I select the best of these (F57987)

as lectotype of Plumularia setaceoides var. crateriformis.

Stechow (1925) raised the var. crateriformis to specific

rank, recording the species from brown algae at 12-14 m

depth in Warnbro Sound, Western Australia. Watson (2005)

also reported P. crateriformis from algae at 16 m depth in the

nearby Archipelago of the Recherche, Western Australia.

Plumularia nodosa Stechow, 1924, also from Western

Australia, is probably conspecific with P. crateriformis ;

however as neither species is referrable to Monotheca as

defined here, they are not considered any further.

Two of the three remaining Mulder and Trebilcock

microslides are labelled ‘ Plumularia setaceoides var.

crateriformis, Bream Creek, January 1914’ and are presumably

those on which Mulder and Trebilcock (1915) based their

augmented description of the var. crateriformis. The third

slide is labelled ‘ Plumularia setaceoides var crateriformis,

Torquay, February 1915’. The Bream Creek specimens have

only one hydrotheca on the hydrocladium and are referrable to

Monotheca , whereas the Torquay specimen has more than one

hydrocladial hydrotheca and thus belongs to Plumularia. The

1914 Bream Creek specimens are identical to Monotheca

amphibola from Queenscliff.

The conspicuous anterior and posterior hydrothecal flanges

of Monotheca amphibola resemble those in some of the smaller

and more variable forms of the Plumularia setaceoides species

group. These structures, together with the strong internal cauline

segmentation and the wide, flat hydrorhiza with flexion joints are

adaptations to strengthen the hydrocaulus in the high-energy

Amphibolis seagrass habitat. The tall hydrorhizal nematothecae

are probably for defence against the many grazing predators in

the seagrass habitat. The smooth texture of the gonothecal wall

is visible only in fresh material, as the gonothecae tend to

crumple and become corrugated in mounting.

Etymology. The species is named for the seagrass Amphibolis

antarctica upon which it grows.

Monotheca australis (Kirchenpauer, 1876)

Fig. 1H-N

Plumularia obliqua var. australis Kirchenpauer, 1876: 49, pi. 6,

fig. 10.— Von Lendenfeld, 1885a: 474.— Stranks, 1993: 8.

Plumularia australis Bale, 1884: 143, pi. 12, figs 7, 8.— Von

Lendenfeld, 1885a: 475.— Bartlett, 1907: 42.— Mulder and Trebilcock,

1916: 77, pi. 10, figs 1-lb.— Bedot, 1921: 26.— Stechow, 1921: 260.—

Blackburn, 1938: 316.— Blackburn, 1942: 108.— Watson, 1973: 189.—

Staples and Watson, 1987: 218.— Watson, 1992: 220.

Plumularia compressa Bale, 1882: 31, pi. 15, fig. 5.— Bale, 1884:

142, pi. 12, figs 9, 10, pi. 19, figs 39, 40.- Von Lendenfeld, 1885a:

475.- Stranks, 1993: 9.

Plumularia aurita Bale, 1888: 784, pi. 19, figs 16-19.

Monothecella australis— Stechow, 1923a: 13.— Stechow, 1923b:

221.- Leloup, 1932: 160.

Monotheca australis.— Watson, 2003: 252.

Material examined. The Kirchenpauer dry hydroid collection in

Museum Victoria holds material (NMV F58239) labelled (presumably

in Kirchenpauer’s handwriting) ‘ Monopyxis australis Port Phillip

(Australien) on Zostera sp. c. 1865’.

Material in author’s collection-, i) from the seagrasses Amphibolis

antarctica and Amphibolis griffithv, Torquay, Victoria, depth 3 m, 7

June 1970. Queenscliff, Victoria, depth 3 m, coll: J. Watson, 15

October 1986. Portland, Victoria, depth 3 m, 14 May 1969. Tipara

Reef, Gulf St Vincent, South Australia, depth 5 m, coll: J. Watson, 24

November 1970. Starvation Bay, Western Australia, depth 2 m, coll: J.

Watson, 11 January 1986. Point Peron, Western Australia, depth 3 m,

coll: J. Watson, 26 January 1986. Whitfords Reef, Marmion, Western

Australia, depth 4 m, coll: J. Watson, 6 February 1986. Cockburn

Sound, Western Australia, depth 6 m, 12 October 1983. Starvation

Bay, Western Australia, depth 6 m, coll: J. Watson, 13 January 1986.

ii) from seagrasses Posidonia australis and Posidonia sinuosa-. Gulf

St Vincent, South Australia, depth 10 m, coll: J. Watson, 14 December

1968. Gulf St Vincent, South Australia, depth 12 m, 10 November

1968. Whitfords Reef, Marmion, Western Australia, depth 3 m, coll: J.

Watson, 22 November 1983. Whitfords Reef, Marmion, Western

Australia, depth 24 m, coll: J. Watson, 30 January 1986.

Description (from mounted and preserved material). Stems to

7 mm high, arising from a hydrorhiza reptant on seagrass leaf,

stolons broad and flat with flexion joints. Stems monosiphonic,

unbranched, sometimes with a short athecate basal internode,

following internodes all of same length, cylindrical, smooth,

widening to distal apophysis; node above apophysis transverse,

narrow.

Hydrocladia alternate, in one plane, one distal on internode,

given off below node on an upwardly directed apophysis with

slightly oblique distal node. Hydrocladium with a short

proximal athecate internode with thick walls and a strong,

almost transverse distal node, hydrothecate internode

considerably longer than athecate internode, base slightly

convex, distal end blunt, ending below hydrothecal margin, no

intranodal septa.

Hydrotheca seated in a concavity of internode, occupying

almost entire internode, slightly laterally compressed,

posterior quarter of abcauline wall strongly convex, then

becoming straight to margin, base of hydrotheca divided into

two segments by a strong intrathecal septum passing obliquely

backwards into hydrotheca from margin, a small circular

foramen in floor of hydrotheca, location variable from mid-

Review of the genus Monotheca (Hydrozoa: Leptolida) from Australia with description of a new species and a note on

Monothecella Stechow, 1923

base to posterior of hydrotheca; margin perpendicular to or

slightly oblique to hydrocladial axis, rim weakly to noticeably

sinuous, margin slightly everted with thickened perisarc.

Nematothecae of two different shapes and sizes: one

cauline similar to laterals in axil of hydrocladial apophysis, a

small cylindrical hydrostatic pore (mamelon of former authors)

on apophysis beside base of nematotheca; one hydrocladial

median inferior on hydrothecate internode between base of

hydrotheca and node, base robust, cup incipiently bithalamic,

scoop shaped, sometimes without a line of demarcation

between cup and base, closely adpressed to wall of hydrotheca,

a distinct flange connecting abcauline wall with internode;

twin lateral nematothecae bithalamic, directed outwards or

downwards in front of hydrotheca, base large, cup excavated

adcaudally to base.

One to three gonothecae borne on lower stem, male and

female often on same stem; gonothecae large, turgid, inserted

on a short, smooth pedicel below apophysis, standing out

perpendicularly from stem or recumbent to substrate,

abcauline (upper) wall inflated, adcauline wall straight to

weakly convex, body narrowing distally to an upwardly bent

cylindrical neck with a wide circular thickened aperture with

broadly everted rim. Immature female gonophore with several

ova, reduced to one large planula at maturity.

Perisarc of stems thick, translucent white when young,

becoming pale-straw coloured with age; gonotheca brown,

female gonophore cream.

Monotheca australis , measurements (/<m)

Hydrorhiza, width

~ 289

Stem

length of internode

230-260

width at node

20-40

width at level of apophysis

90-149

Hydro cl adium

length of athecate internode

50-110

length of thecate internode

140-200

Hydrotheca

diagonal length, margin to base

160-172

depth, margin to floor

90-180

diameter of rim

80-140

Nematotheca

total length of median inferior

30-60

length of lateral including base

40-60

width of cup

20-50

width of axillar pore

14-20

Gonotheca

length, including pedicel

650-1000

maximum width

200-350

diameter of neck

100-220

diameter of aperture

110-250

Remarks. The material in the dry Kirchenpauer collection of

Museum Victoria is probably that on which Kirchenpauer

(1876) described Plumularia ( Monopyxis ) obliqua var.

australis. The specimen (NMV F58239) comprises many

stems, some fertile, on dried leaves of the seagrass Amphibolis

antarctica, formerly known as Zostera. Stranks (1993)

nominated this material as possible syntype. Because of its

unusual hydrocladial median inferior nematotheca, Bale (1884)

raised Kirchenpauer’s var. australis to specific rank as

Plumularia australis. I designate F58239 as lectotype of

Monotheca australis Kirchenpauer, 1876.

Compared with the height of the stems, the hydrorhizal

stolons of M. australis are very wide, the strongly developed

flexion joints permitting movement of the hydrorhiza on the

Amphibolis leaves in strong water movement. Although the

nematothecae are all essentially bithalamic, in some colonies

the cup of the hydrocladial median inferior nematotheca is so

adcaudally reduced that the line of demarcation between

peduncle and cup is either very faint or altogether lost.

The small hydrostatic pore on the apophysis of the stem is

obscured by the axillar nematotheca and is only visible in

cleared specimens. The pore was described by Mulder and

Trebilcock (1916) as similar in shape to a ‘steamship ventilator’;

in fresh material, however, it is cylindrical and likely to bend

under pressure of a cover slip.

Monotheca australis is an obligate epiphyte of seagrasses,

growing on Amphibolis antarctica in cool temperate south¬

eastern Australia and on Posidonia australis in warm

temperate waters of southern and south-western Australia. In

the rigorous Amphibolis habitat, the hydroid colonises the

inner, sheltered leaves of the seagrass, while in the quieter

Posidonia habitat it occurs on all but the outermost leaves of

the plants. Colony size and cauline dimensions of M. australis

on Amphibolis are usually smaller than those on Posidonia.

Boero (1981a, 1984) reported an increase in cauline length

and loss of the intrathecal septum in M. obliqua from

Posidonia seagrasses with increasing depth and decreasing

water movement in the Mediterranean Sea. Because M.

australis also grows on Posidonia in southern Australia,

specimens from Posidonia habitat from 0-25 m depth were

examined to investigate whether the same relationship exists

among Australian Monotheca seagrass epiphytes. No such

gradient was found; any tendency to loss of the intrathecal

septum probably being lost in the more rigorous Australian

Posidonia habitat.

Type locality. Port Phillip, Australia.

Known distribution. South-eastern and southern Australia to

depth of 25 m.

Status of Monotheca compressa (Bale, 1882), Monotheca

aurita (Bale, 1888) and Monotheca obesa (Blackburn, 1938).

Bale (1882) described Plumularia compressa from fertile

material collected by Mr T. B. Smeaton at Robe, South

Australia. (The gonotheca was figured in 1884). The collection

in Museum Victoria contains three microslides labelled by

Bale ‘ Plumularia compressa Bale, 1884’. These are presumably

from Bale’s 1882 collection and relabelled in 1884. The

J.E. Watson

specimens are probably from the shallow water seagrass

Posidonia australis common at Robe. As Bale did not designate

a holotype I designate a microslide NMV F59056 of a fertile

colony as lectotype of Monotheca compressa.

Bale (1884, 1888, 1894) recognised the close relationship

between M. compressa and M. australis, but nevertheless kept

the two species separate — the main points of difference were

length of stem, height and shape of the hydrotheca, position of

the intrathecal septum and minor differences in shape of the

neck of the gonotheca. Length of stem is an unreliable

character as it may vary considerably between the same or

different colonies according to environmental conditions and

rate of growth. Height of the hydrotheca is quite variable over

the range of material examined in the present study and is

actually related to the degree of torsion around the intrathecal

septum; shape of the base of the hydrotheca is also quite

variable, some australis morphotypes being almost

hemispherical in shape. The position of the foramen in the

floor of the hydrotheca is also variable and depends on the

degree of torsion of the hydrotheca around the intrathecal

septum. It is more or less central in 75% of the australis

morphotypes examined but is displaced towards the rear in the

compressa form. Hydrothecae with a long, backwardly curved

septum typical of Kirchenpauer’s australis occur in colonies

from the cool temperate Amphibolis antarctica habitat, the

septum in these hydrothecae penetrating at least halfway into

the hydrotheca, providing a pivotal point for torsion of the

hydrotheca. In the warm temperate Posidonia habitat, the

hydrothecae are taller and the septum is often directed forward

as a rudimentary intrathecal ridge rather than a deep inflexion.

Gonothecae vary in size and shape from almost cylindrical to

turgid, and no relationship between sexual dimorphism,

habitat or geographical distribution could be found to account

for these variations.

Bale (1888) described Plumularia aurita from Botany,

New South Wales. A microslide (NMV F58776) in the

collection of Museum Victoria is the only known material of

the species and was nominated syntype by Stranks (1993). As

this is the only known specimen, by monotypy it is the holotype

of the species. Bale’s description and figure of the hydrotheca

of M. aurita shows no intrathecal ridge, yet examination of the

microslide clearly shows a backwardly oblique ridge passing

almost halfway into hydrotheca. The incipiently monothalamic

hydrocladial median inferior nematotheca is the same as that of

M. australis, and the single small gonotheca on the slide

although somewhat damaged confirms the species as a

morphotype of M. australis. Although Bale provided no

ecological notes, the species was almost certainly collected

from Posidonia seagrass in Botany Bay.

The holotype microslide of Plumularia obesa Blackburn,

1938, in the collection of Museum Victoria, consists of three

small, infertile stems labelled ‘holotype NMV F70661,

Plumularia obesa Blackburn, 1938, Reevesby Island, South

Australia, December, 1936’. No other mounted or preserved

material is known to exist. Since the microslide specimen was

not cleared of tissue before mounting many critical structures

are obscured, making it difficult to accurately measure

important features.

In his description of Plumularia obesa, Blackburn

surprisingly compared the species with Plumularia spinulosa

rather than comparing it with Plumularia australis, which he also

recorded from the same locality. The hydrotheca of Monotheca

obesa closely resembles the compressa form of M. australis from

Posidonia seagrass, the major difference being the shorter and

more robust hydrocaulus resembling Bale’s M. aurita. Blackburn

described and figured the nematothecae of P. obesa as bithalamic

with slender bases; however, examination of the type shows that

the hydrocladial median inferior nematothecae have sturdy bases

similar to those of P. australis. The bases of the twin lateral

nematothecae are not slender as figured by Blackburn, nor are the

cauline nematothecae of the holotype similar to the laterals and

the bases are not so slender as to be considered truly moveable.

Apart from the much smaller and more robust hydrocaulus, and

taller hydrothecae, P. obesa is indistinguishable from the extreme

compressa form of M. australis. Although the gonosome of P.

obesa is unknown, I am confident that the species is a

morphological variant of M. australis and, accordingly, I include

it in the synonymy of M. australis.

Monotheca australis is a highly variable species, the size

of the colony and structural dimensions over the geographic

range are closely related to substrate type and environmental

conditions.

Monotheca flexuosa (Bale, 1894)

Fig. 2 A-G

Plumularia flexuosa Bale, 1894: 115, pi. 5, figs 6-10.— Bartlett,

1907: 42.- Shepherd and Watson, 1970: 140.- Watson, 1973: 187.-

Watson, 1982: 106, fig.4.11e.- Watson, 1992: 220.- Stranks, 1993:

10 .

?Plumulariaflexuosa — Bedot, 1921: 27.— Blackburn, 1938: 315.

Monotheca flexuosa — Stechow, 1921: 260.— Stechow, 1925:

246.- Hirohito 1974: 37, fig. 17.-Watson, 1996: 78.- Watson, 2000:

48, fig. 37A, B.— Vervoort and Watson, 2003: 369.— Watson, 2003:

252.- Watson, 2005: 540.- Bouillon et al„ 2006: 368.

Monotheca pulchella — Medel and Vervoort, 1995: 58, fig. 25.—

Medel and Lopez-Gonzalez, 1996: 202.— Watson and Mclnnes, 1999:

111.— Vervoort and Watson, 2003: 369, 373, fig. 90J-L.— Bouillon et

al„ 2004: 172, fig. 92G-K.

Material examined. Microslides in the Bale collection. Museum Victoria,

labelled in Bale’s handwriting are: NMV F58754, ‘Catalogue number

206, Plumularia flexuosa, mouth of Snowy River, Dr Macgillivray,

1889’ and three other slides, NMV F58755, all labelled ‘ Plumularia

flexuosa. Snowy River, Dr Macgillivray, 1892’ (Bale’s catalogue number

205).

Material in author’s collection'. Popes Eye reef. Port Phillip, fertile

colonies on invertebrate and algal substrates, coll: J. Watson, depth 10 m,

20 September 2008. St Leonards pier. Port Phillip , depth 3 m, i nf ertile

colonies on red alga, coll: J. Watson, 10 August 2008. Clifton Springs,

Port Phillip, jetty piles, depth 1.5 m, fertile colonies on algae, coll: J.

Watson, 30 November 2005. Clifton Springs, Port Phillip, jetty piles,

depth 1 m, fertile colonies on green alga Caulerpa scalpelliformis, coll:

J.Watson, 16 September 2006. Clifton Springs, Port Phillip Jetty piles,

depth 1.5 m, fertile colonies on green alga Caulerpa scalpelliformis, coll:

J. Watson, 23 October 2008. Pearson Island, South Australia, infertile

stems on red alga Mychodea carnosa, depth 37—45 m, coll: J. Watson,

January 1969. Channel Island, Darwin, Northern Territory, depth 6 m,

coll: J. Watson, July 2006.

Review of the genus Monotheca (Hydrozoa: Leptolida) from Australia with description of a new species and a note on

Monothecella Stechow, 1923

Figure 2. A-G, Monotheca flexuosa. A, part of stem; B, hydrocladium, lateral view; C, hydrocladium, ventral view; D, male and female

gonothecae on same stem; E, median inferior nematotheca; F, cauline nematotheca; G, one of twin lateral nematothecae. Scale bar: A, D, 0.5 mm;

B,C, 0.2 mm; E-G, 0.1 mm.

Description. Hydrorhiza of thin tubular stolons reptant on

invertebrate substrate, sometimes intersecting in tangled knots, on

algal substrate stolons often flattened with flexion joints. Caulus

of same diameter as stolon, monosiphonic, flexuous, variable in

length, up to 20 mm, base of stem smooth with a few irregular

transverse joints; short stems unbranched, some longer stems with

one or two orders of short side branches, stem intemodes long.

Hydrocladia alternate, in one plane, borne on a strong

apophysis below cauline node, apophysis curving smoothly

away from internode, distal node transverse with rounded

shoulder. Hydrocladial athecate internode long, cylindrical,

straight to slightly curved, walls smooth, increasing slightly in

diameter distally; distal node transverse, deep, a transverse

septum behind node, typically several transverse internal septa

in intemode.

Hydrothecate intemode about same length as athecate

intemode, almost entirely occupied by hydrotheca, proximal

node narrow, deep, inserted into shoulder of athecate intemode;

intemode slightly distally inflated, terminating behind margin of

hydrotheca, a faint intranodal septum sometimes in intemode

below hydrotheca. Hydrotheca cup shaped, adcauline wall

convex, set deeply into internode, abcauline wall weakly

concave, margin circular, transverse to hydrothecal axis, rim

delicate, a slight thickening below margin.

J.E. Watson

Nematothecae all of same size and shape, bithalamic,

moveable, fairly robust, base long, cup circular, slightly

adcaudally shortened, three nematothecae on cauline

internode, one on apophysis, the other axillar and one, variable

in position, about one-third to mid-distance up internode,

hydrocladial median inferior reaching one-quarter distance

along hydrotheca, twin laterals extending well beyond rim of

hydrotheca.

Gonothecae of both sexes borne thickly in a single row

along lower stem or sparsely along branches. Male and female

gonothecae the same shape, top-shaped when young, ovoid to

barrel shaped at maturity, distally truncate, pedicel inserted

beside apophysis of stem, upwardly bent, tapering into base of

gonotheca; walls smooth to faintly undulated, aperture distal,

transverse, circular, produced into a short thick neck of

variable height closed by a low dome or shallow concavity.

Male gonophore with rod-shaped spadix, mature female

packed with large ova.

Cnidome:

i) large microbasic euryteles capsule bean shaped 15-16 x

6 pm, abundant in coenosarc and nematothecae;

discharges sideways.

ii) small ?isorhizas, capsule elongate, 5-7 x 2 pm, tubule

short; abundant in coenosarc and tentacles.

Perisarc of hydrocaulus and gonothecae fairly thin, thicker

at base of stems. Colonies transparent white, basal stem region

brown in older colonies, male gonophores white, often with a

shining bluish tinge, mature ova in female gonophore yellow.

Monotheca flexuosa, measurements (p m)

Hydrorhiza, width

50-80

Stem

internode length

344-400

diameter at node

36-44

length of apophysis (adcauline wall)

48-52

Hydrocladium

length of athecate internode

116-128

width at distal node (shoulder)

50-58

length of thecate internode (measured along

base)

164-200

Hydrotheca

length of abcauline wall

132-148

length of adcauline wall

136-60

diameter of margin

140-152

Gonotheca

length (excluding pedicel)

560-696

maxiumum diameter

280-320

Nematotheca

length of base

45-55

diameter of cup

40-50

Remarks. All microslides of Monotheca flexuosa in the Bale

collection of Museum Victoria are designated syntypes by

Stranks (1993). I designate NMV F58754 as lectotype of

Monotheca flexuosa.

The systematic importance of the presence, absence or

position of the cauline nematotheca has been much discussed

by authors (e.g. Medel and Vervoort, 1995). In many southern

Australian specimens of M. flexuosa, cauline nematothecae

are present in the proximal one-third to mid-cauline internode,

suggesting precise position is of little or no diagnostic value

for identification of the species.

Stems of M. flexuosa from sheltered habitat are usually

much longer, and more flexuous and silky in appearance than

those from more rigorous habitats. The longer stems may bear

a single row of up to 15 female gonothecae whereas shorter

stems have fewer gonothecae, usually near the base of the

stem. No correlation with environmental conditions or habitat

could be found to explain the presence, absence or length of

the terminal neck of the gonotheca.

M. flexuosa is the most abundant and widespread species

of Monotheca in Australia. In southern Australia, it is fertile

from late spring to autumn (November to March) when

colonies occur on many invertebrate substrates (mussels,

ascidians, polychaete tubes), on some flabellate red algae and

the green algae Caulerpa scalpelliformis and Caulerpa

remotifolia.

An unusual endoparastic association between M. flexuosa

and the red alga Mychodea carnosa was reported by Watson

(1973). The stolons of the hydroid penetrate the outer medulla

of young Mychodea fronds, producing external hydrocauli at

intervals. As the alga grows, its tissue is gradually broken

down by the hydroid stolons so that eventually the alga is

attached to the substrate only by the hydrorhiza of the hydroid.

Hydrocauli emerging from the algal medulla differ somewhat

from the normal epiphytic form, comprising one or two short

cylindrical basal internodes bearing one or two large

nematothecae, followed by normal hydrocauline internodes.

Type locality. Mouth of Snowy River, Victoria, Australia.

Known distribution. Temperate and tropical Australia, New

Zealand, Strait of Gibraltar. The species may prove to be

cosmopolitan.

Monotheca hyalina (Bale, 1882)

Fig. 3A-I

Plumularia hyalina Bale, 1882: 41, pi. 15, fig. 9.— Bale, 1884: 141,

pi. 12, figs 4, 5.— Bartlett, 1907: 422 — Bedot, 1921: 28.— Trebilcock,

1928: 24, pi. 6, fig. 6.- Watson, 1975: 170, fig. 29.- Stranks, 1993: 11.

Monotheca hyalina — Stechow, 1921: 260.— Watson, 1996: 78.—

Watson, 1997: 529.- Bouillon et al„ 2006: 368.

Not Plumularia hyalina — Ralph, 1961b: 41, fig. 5a, b.— Ralph,

1961c: 109.

Not Monotheca hyalina.— Vervoort and Watson, 2003: 371, fig.

90F-I.

Material examined. NMV F59052, microslide; NMV F59053,

microslide. Bale collection. Museum Victoria, locality Queenscliff,

Victoria.

Material in author’s collection : Floutman Abrolhos Islands,

Western Australia, infertile colony on Sargassum and red alga, depth

6-10 m, coll: J. Watson, 12 October 1986; Hopetoun, Western

Australia, female colony on brown alga, depth 2 m, under ledge, coll:

J. Watson, 13 January 1986. Boatswains Rocks, South Australia,

Review of the genus Monotheca (Hydrozoa: Leptolida) from Australia with description of a new species and a note on

Monothecella Stechow, 1923

Figure 3. A-H, Monotheca hyalina. A, part of robust stem from cool temperate waters with intranodal septa; B, part of slender stem from

temperate waters with no intranodal septa; C, hydrocladium and hydrotheca, lateral view; D, hydrocladium and hydrotheca, ventral view; E,

gonotheca with female gonophore; F, apical stolonal plate; G, axillar hydrostatic pore; H, median inferior nematotheca; I, one of twin lateral

nematothecae. Scale bar: A, B, 1 mm; C, D, 0.2 mm; E, 0.5 mm; G-I, 0.1 mm.

J.E. Watson

colony detached from substrate, coll: J. Watson, 27 January 1969.

Fluted Cape, Bruny Island, Tasmania, female colony, depth 16 m, on

red alga, coll: J. Watson, 10 February 1972. Lawrence Rocks, Portland,

Victoria, infertile colony on red alga, depth 16 m, coll: J. Watson, 14

May 1969. Walkerville, Victoria, infertile colony detached from

substrate, depth 3 m, coll: J. Watson, 1 March 1968. Nambucca Heads,

New South Wales, infertile colony on coralline alga and ascidian,

depth 13 m, coll: J. Watson, 11 January 1972.

Description. Hydrorhiza of tubular rugose stolons reptant on

substrate; stolons usually with internal flexion joints. Stems

3-10 mm high, monosiphonic, straight to weakly sympodial,

up to four short basal internodes with transverse joints above

hydrorhiza. Succeeding stem internodes variable in length,

longer ones typically on taller stems, smooth, expanding a little

distally with a strong oblique V-shaped proximal node and a

transverse distal node, usually an internal transverse septum

below distal node at level of apophysis.

Shorter stems with one, rarely two, short proximal athecate

internodes, stem thereafter hydrocladiate; cauline internodes

deeply and more or less irregularly septate with strong

V-shaped proximal nodes and transverse distal nodes, and

three or four strong transverse internal septa. Hydrocladia

alternate, fairly long, given off in one plane from a short,

strong apophysis distal on internode, apophysis curving

smoothly out from internode, adcauline wall a shoulder with

slightly oblique deep node, a cylindrical axillar hydrostatic

pore on adcauline shoulder.

Athecate hydrocladial internode long, deeply inserted in

shoulder of apophysis, with deep transverse distal node and

several complete internal septa. Hydrothecate internode longer

than athecate internode, terminating in a protuberance

separating the twin lateral nematothecae, hydrotheca set

deeply in internode, a large prehydrothecal chamber expanding

from proximal node to floor of hydrotheca; hydrotheca deep

bowl shaped, base convex with a large foramen connecting

with prehydrothecal chamber; in lateral view abcauline wall

convex, adcauline wall completely adnate to internode, in

ventral view walls expanding in a smooth curve from base,

narrowing just below margin, rim slightly thickened but not

everted; margin hemispherical, wide in ventral view, abcauline

side straight, adcauline side deeply concave, in lateral view,

wall a broad sinous curve down to internode.

Three nematothecae on hydrothecate internode,

hydrocladial median inferior about halfway along

prehydrothecal chamber, small, bithalamic, moveable,

adpressed to internode, base tapering, cup slightly adcaudally

shortened, twin laterals of same shape as median, standing

erect on each side of strong terminal protuberance of internode,

tucked within hydrothecal margin, cup circular to slightly

laterally compressed but not excavated; no cauline

nematothecae.

Gonotheca large, one to several in a row on lower stem,

conical, widening from a short indefinite pedicel inserted beside

apophysis, walls of gonotheca smooth to obscurely undulated,

truncated distally by a transverse, weakly concave aperture

without operculum; female gonophore with 15-20 large ova.

Perisarc of slender stems thin, much thicker in shorter,

robust stems.

Colour of live colonies where known, buff to yellow,

gonophores yellow.

Monotheca hyalina, measurements (/tm)

Slender group

(warm

temperate)

Robust group

(cool

temperate)

Hydrorhiza, width

112-120

Stem

length of internode

456-640

304-400

width at node

80-120

80-112

distal width of apophysis

80-100

92-112

Hydrocladium

length of athecate

internode

80-220

80-120

length of hydrothecate

internode

200-296

184—212

Hydrotheca

length of abcauline wall

193-220

152-160

length of adcauline wall

160-186

80-116

width of margin front view

160-192

128-168

Nematotheca

basal length of median

inferior

35^12

16-26

diameter of cup

42-52

40-60

basal length of lateral

40-50

30^10

diameter of cup

40^16

44—50

Gonotheca

length, including pedicel

960-1400

920

maximum width (margin)

720-900

680

Remarks. There is no known type material of Monotheca

hyalina ; a microslide NMV F59052 in the Museum Victoria

Bale collection labelled in Bale’s handwriting ‘ Plumularia

hyalina Queenscliff, February 1881’ was nominated as

probable syntype by Stranks (1993). I select this microslide as

lectotype of Monotheca hyalina.

Monotheca hyalina is very variable in height of the stem,

and slenderness and robustness of the hydrocaulus, in the

presence or absence of intranodal septa and size of the

hydrotheca. In some specimens, the prehydrothecal chamber

is so reduced that the hydrocladial median inferior

nematotheca extends one-third the length of the hydrotheca.

Specimens from warm-temperate southern Australian

Review of the genus Monotheca (Hydrozoa: Leptolida) from Australia with description of a new species and a note on

Monothecella Stechow, 1923

waters are distinguished by their slender, flexuous stems with

longer internodes, while those from the cool-temperate

waters have sturdier stems with shorter internodes and

smaller hydrothecae. Dimensions of these two groups are

compared in the above table.

Despite the considerable structural and dimensional

differences between the two morphological extremes,

intergradation does occur, hence M. hyalina is recognised as

a single but somewhat variable species. It is often difficult to

obtain a good lateral view of the hydrotheca as they tend to

twist forward in microslide preparations. M. hyalina is an

algal epiphyte; some specimens from Tasmania have a large

apical stolonal plate to securely anchor stems in a rigorous

oceanic environment.

Examination of New Zealand material and a review of

descriptions of Ralph (1961b) and Vervoort and Watson

(2003) suggests that the New Zealand material is not M.

hyalina because: i) the hydrotheca is pitcher-shaped with a

short, concave free adcauline wall, ii) there is a partial

intranodal septum below the hydrotheca, iii) there is an

axillar nematotheca in the apophysis of the stem, and iv) as

the New Zealand specimens usually have more than one

hydrotheca on the hydrocladium it is excluded from

Monotheca as defined here. It is likely that the New Zealand

material is an undescribed species of Plumularia. It is

noteworthy that Leclere et al. (2007) in their molecular

analysis demonstrate that M. hyalina from New Zealand is

outside the group, thus supporting the conclusion that the

New Zealand species is not M. hyalina.

Type locality. Queenscliff, Victoria, Australia.

Known distribution. Southern Australia.

Monotheca obliqua (Johnston, 1847)

Fig. 4 A-E

Plumularia obliqua Johnston, 1847: 106, pi. 28, fig. 1.— Bale,

1884: 138, pi. 12, figs 1-3.- Bartlett, 1907: 43.- Mulder and

Trebilcock, 1910: 116. — Mulder and Trebilcock, 1916: 76, pi. 11, figs

1-le. - Jaderholm, 1919: 22, pi. 5, fig. 6.- Stechow, 1919: 113.-

Bedot, 1921: 28,-Billard, 1927: 342.- Broch, 1933: 31, fig. -

Leloup, 1934: 15.— Billard, 1936: 9.— Blackburn, 1938: 315.—

Blackburn, 1942: 108.- Hodgson, 1950: 39, fig. 68.- Rossi, 1950: 23,

fig. 4b.— Pennycuik 1959: 180.— Watson, 1973: 189.— Millard and

Bouillon, 1974: 9,34, fig. 8A-D.- Millard, 1975: 396, fig. 125A-B.-

Garcia-Corrales, Aguirre Inchaurbe and Gonzalez Mora, 1978: 56,

fig. 25.— Millard, 1978: 196 et seq.— Watson, 1979: 234.— Boero,

1981a: 197.- Hirohito, 1983: 69. - Boero, 1984: 103, fig. 8.- Boero

et al., 1985: 29 - Gili and Garcfa-Rubies, 1985: 48, fig. 5B.— Isasi and

Saiz, 1986: 70.— Roca, 1987: 151.—Gili, Vervoort and Pages, 1989: 89,

fig. 17A.— Boero and Bouillon, 1989: 39, fig. 1. Garcia Rubies, 1992:

265.— Park, 1992: 294 Cornelius, 1995: 142, fig. 33.— Bouillon et al„

1995: 61.— Hirohito, 1995: 275, fig. 94d-f.— Watson and Mclnnes,

1999: 111.- Bouillon et al., 2006: 368.

Monotheca obliqua .— Stechow, 1923b: 17.— Leloup, 1932:

160.— Yamada, 1959: 78.— Rho and Park, 1986: 99.— Ryland and

Gibbons, 1991: 538, fig. 9.— El Beshbeeshy, 1995: 404.— Medel and

Lopez-Gonzalez, 1996: 202.— Watson, 1973: 529.— Watson and

Mclnnes, 1999: 111 — Vervoort and Watson, 2003: 369.— Watson,

2005: 541, fig. 15.- Bouillon et al., 2006: 368.

Plumularia (Monotheca) obliqua.— Stechow and Uchida, 1931:

565.

Plumularia obliqua australis.— Stranks, 1993: 8.

Material examined. NMV F101686, infertile colony. New Island,

Archipelago of the Recherche, Western Australia, depth 20 m on red

alga, coll: J. Watson, February 2003. WAM Z29970, fertile colony.

New Island, Archipelago of the Recherche, Western Australia, depth

20 m, on brown alga, coll: J. Watson, February 2003.

Material in author’s collection: fertile colony. Popes Eye reef.

Port Phillip , Victoria, on stem of Macrorhynchia whiteleggei, depth

6 m, coll: J. Watson, 7 November 2009. Infertile colony. Popes Eye

reef. Port Phillip, Victoria, depth, 3 m, on thallus of Sargassum sp.,

coll: J. Watson 29 December 2009.

Description. Hydrorhiza reptant on substrate, tubular,

somewhat rugose. Stems to 7 mm long, monosiphonic, straight

to sympodial, internodes moderately long, widening to distal

node; node transverse to slightly oblique, often a distinct

shoulder in perisarc and an internal septum above, sometimes

one to three strong internal septa in proximal part of

internode. Apophysis of stem short, upwardly directed,

situated well below node, widening to a strong distal shoulder.

Hydrocladia alternate, short, in one plane, distal on

internode, athecate internode with one to three internal

septa, distal node transverse. Hydrothecate internode almost

completely occupied by hydrotheca, internode with one or

two faint septa below hydrotheca, sometimes a short, faint

septum passing into hydrotheca from distal septum; internode

distally inflated, pushing base of hydrotheca upward,

prehydrothecal chamber short, triangular with internal

triangular septum.

Hydrotheca scoop shaped, abcauline and adcauline walls

smoothly convex, margin cut obliquely back to internode,

rim slightly everted, a thickening behind rim.

Cauline and hydrocladial median inferior nematothecae

similar in shape and size, bithalamic, moveable, base

moderately long, cup fairly wide, cauline nematotheca about

one-third distance up internode on side opposite

hydrocladium, standing out from internode, base connecting

with a large foramen of internode, hydrocladial median

inferior nematotheca adpressed to prehydrothecal chamber,

just reaching floor of hydotheca; twin laterals directed

outwards from an indentation in distal protuberance of

internode, base short, stout, cup deeply adcaudally excavated;

a nematotheca in axil beside apophysis; a dome-shaped

axillar hydrostatic pore with small terminal aperture,

sometimes also a small secondary pore on adcauline shoulder

of apophysis.

Male and female gonothecae borne on the same colony

but not on same stems, of same shape and size, large, conical,

circular in section, perisarc fairly thick with tendency to

proximal rugosity, typically one borne on a short, thin

pedicel beside an apophysis near base of stem; aperture

transversely truncate, occupying entire width of gonotheca,

closed by a sheet of tissue. Female gonophore containing

many large ova.

Perisarc of stems pale brown basally, becoming colourless

distally, gonotheca pale brown, gonophores flesh-coloured.

J.E. Watson

Figure 4. A-E, Monotheca obliqua. A, part of stem; B, hydrocladium and hydrotheca; C, female gonotheca; D, twin lateral nematothecae; E,

axillar nematotheca and hydrostatic pore. Scale bar: A, C, 1 mm; B, 0.2 mm; D, E, 0.1 mm. F-L, Monotheca pulchella. F, part of stem; G,

hydrocladium and hydrotheca; H, female gonothotheca with ova; I, anterior view of gonothecal aperture with internal lobate submarginal cusps.

J, median inferior hydrocladial nematotheca. K, one of twin lateral nematothecae. L, axillar nematothecae. Scale bar: A,C, 1 mm.; B, 0.2 mm; F,

H, 0.5 mm; D-E, I-L, 0.1 mm.

Review of the genus Monotheca (Hydrozoa: Leptolida) from Australia with description of a new species and a note on

Monothecella Stechow, 1923

Monotheca obliqua, measurements (/tm)

Hydrorhiza, width

70-88

Stem

length of internode

200-320

width at distal node

40-60

Apophysis

length of adcauline wall

26-58

width at distal shoulder (node)

40-70

Hydrocladium

length of athecate internode

56-80

width at distal node

42-60

length of thecate internode

160-104

Hydrotheca

length of abcauline wall

140-200

length of of adcauline wall

80-148

depth, floor to margin tip

180-200

width at rim

160-180

Nematotheca

total length of lateral

52-60

diameter of cup

42-48

length of median

48-52

diameter of cup

26-28

Gonotheca

length, including pedicel

1240-1580

width at margin

640-1100

Remarks. Monotheca obliqua is a well-known species with

worldwide distribution. Mulder and Trebilcock (1910, 1916)

briefly described material collected from Bream Creek on the

Bass Strait coast and Corio Bay in Port Phillip, commenting on

the slenderness of the base of the median inferior nematotheca,

the small axillar hydrostatic pore and the abcauline flange

connecting the base (prehydrothecal chamber) of the hydrotheca

to the internode.

Although widely distributed across the south and east of

the continent, M. obliqua is not a common species in Australia.

Type locality. British Isles.

Known distribution. Cosmopolitan; southern Australia to

Queensland.

Monotheca pulchella (Bale, 1882)

Fig. 4 F-L

Plumularia pulchella Bale, 1882: 30, pi. 15, figs 6, 6a.— Bartlett,

1907: 43.- Stranks, 1993: 13. - Watson, 1994: 67.

Not Plumularia pulchella. — Bedot, 1921: 28.— Trebilcock, 1928:

24. — Totton, 1930: 221, fig. 58 a-d.— Hodgson, 1950: 41, fig. 71.—

Ralph, 1961b: 39, fig. 5 c-e.- Blanco, 1973: 73, figs 1-3.- Millard,

1975: 398, fig. 125 C-D.— Izquierdo et al., 1986: 54, fig. 5.— Genzano,

1990: 50, figs. 16-17.-Medel and Vervoort, 1995: 58, fig. 25.-

Vervoort and Watson, 2003: 369, 373, fig. 90J-L.— Bouillon, et al.,

2004: 172, fig. 92G-K.

IPlumularia pulchella — Blackburn, 1942: 108.— Day etal., 1952:

404.- Millard, 1957: 232.- Millard, 1962: 300.- Pennycuik, 1959:

180.- Ralph, 1961b: 109.- Millard, 1966: 493.- Berrisford, 1969:

394.- Day et al., 1970: 14.- Millard, 1978: 196.- Millard, 1980:

133.— Medel and Lopez-Gonzalez, 1996: 202.

Monotheca pulchella.— Stechow, 1921: 260.— Watson and

Mclnnes, 1999: 111.

Material examined. NMV F59054, Bale collection. Museum Victoria,

one microslide, labelled in Bale’s handwriting ‘ Plumularia pulchella

Bale, Queenscliff, November, 1878’.

Material in author’s collection : microslide and preserved

material, Knobbies Island, Western Port, on brown alga, coll: J.

Watson, depth 10 m, 27 Januaryl985.

Description. Hydrorhiza flat and ribbon-like with flexion

joints. Stems monosiphonic, straight to weakly sympodial,

basal athecate region with several transverse nodes often with

one or two nematothecae, succeeding cauline internodes

variable in length, cylindrical, robust, nodes deep, transverse to

oblique, internode typically with several internal septa

imparting a wrinkled appearance to stem.

Hydrocladia alternate, usually in one plane, but sometimes

one or two forwardly displaced, halfway to one-third distance

up internode, apophysis of stem upwardly directed away from

internode, long, stout, with a broad transverse distal shoulder,

usually an internal septum behind shoulder; hydrocladium

short, athecate internode variable in length, proximal end

narrow, inserted into shoulder of apophysis. Hydrothecate

internode variable in length, almost entirely occupied by

hydrotheca, proximal end of internode a narrow neck inserted

into athecate node; distal end terminating in a protuberance

behind hydrothecal margin.

Hydrotheca cup shaped, adcauline wall convex, immersed

in internode, abcauline wall weakly concave and thickened;

margin circular, transverse.

Nematothecae bithalamic, moveable, cauline and

hydrocladial median inferior of same size and shape, base long

and conical, cup deep, rim circular, slightly adcaudally

shortened; a nematotheca one-third distance up cauline

internode and two axillar; hydrocladial median inferior seated

on a prominence of internode, just reaching base of hydrotheca,

twin laterals seated almost at end of internode, base fairly

short, cup adcaudally excavated.

Male and female gonothecae borne on same stem. Sexes of

same shape, kidney shaped to ovoid, inserted on a smooth,

short, bent pedicel at base of apophysis, pedicel expanding and

merging into body of gonotheca; walls of gonotheca thick,

smooth to faintly rugose, aperture distal, circular, displaced to

one side, rim slightly thickened, a submarginal row of large,

irregularly shaped inwardly directed lobate cusps, usually one

or two larger than the others. Mature female gonophore

containing a large ovum.

J.E. Watson

Monotheca pulchella, measurements (/<m)

Stem

internode length

204-400

diameter at node

48-60

length of apophysis (adcauline wall)

52-64

Hydrocladium

length of athecate internode

52-128

width at distal node (shoulder)

50-58

basal length of hydrothecate internode

164-200

Hydrotheca

length of abcauline wall

100-148

diameter of margin

116-152

Gonotheca

length (excluding bent pedicel)

560-696

maximum width

280-344

Nematotheca

length of base

40-55

diameter of cup

32-50

Remarks. Stranks (1993) considered microslide NMV F59054

a probable syntype of Monotheca pulchella. I select this

microslide as lectotype of Monotheca pulchella.

The submarginal lobate cusps mentioned by Bale are

clearly visible in the gonotheca of the type. On present

evidence, M. pulchella is associated only with brown algae.

This is in contrast to M. flexuosa, which is a common

opportunistic species occurring on a wide variety of red and

green algal and invertebrate substrates.

The hydroid reported as Plumularia pulchella by Izquierdo

et al. (1986) from the Canary Islands is probably Monotheca

margaretta (see discussion in Calder, 1997), further supporting

the contention that M. pulchella is endemic to Australia.

Type locality. Queenscliff, Victoria, Australia.

Known distribution. Victorian coastal waters.

Distinction between Monotheca flexuosa and Monotheca

pulchella. Monotheca pulchella (Bale, 1882) has often been

confused by authors with Monotheca flexuosa (Bale, 1894).

Although Bale clearly distinguished between the two species

on the basis of M. pulchella having a robust septate stem, very

short cauline internodes and hydrocladia midway along the

internode, Totton (1930) reported infertile material from

northern New Zealand as M. pulchella, presumably following

Trebilcock (1928), who synonymised M. flexuosa in that

species. Totton’s opinion was based on i) the specimens being

twice the size of those described by Bale (for M. flexuosa ),

some being half an inch in height (12 mm) and bearing 48

hydrocladia, ii) the trophosome of M. flexuosa falling well

within the range of variation of M. pulchella and iii) there

being only slight apparent differences in the gonosome. His

figure (fig. 58, p. 221) is clearly that of M. flexuosa.

Later authors, such as Millard (1975) followed Totton but,

with the exception of Millard, they did not provide figures of

their specimens so the accuracy of their identifications cannot

be confirmed. Watson (1973) examined Bale’s microslide

specimens of both species in the collection of Museum Victoria,

reporting the species to be markedly different and concluded

that Totton was incorrect for the following reasons: i) height of

the hydrocaulus is an unreliable character, ii) M. flexuosa has a

more flexuous hydrocaulus than M. pulchella and iii) the

gonothecae of the two species are markedly different. Medel

and Vervoort (1995) misinterpreted Watson’s (1973) note on

the submarginal gonothecal cusps of M. pulchella, and their

description and figure is clearly that of M. flexuosa. Infertile

material on green algae reported from New Zealand by Vervoort

and Watson (2003) is also M. flexuosa.

On present evidence, M. pulchella is endemic to southern

Australia where it is associated only with brown algae.

Monotheca spinulosa (Bale, 1882)

Fig. 5A-G

Plumularia spinulosa Bale, 1882: 30, pi. 15, fig. 8 — Bale, 1884:

139, pi. 12, figs 11-12.- Bale, 1888: 783, pi. 19, figs 11-13.- Von

Lendenfeld, 1885a: 475.- Bartlett, 1907: 43.- Warren, 1908: 320.-

Mulder and Trebilcock, 1910: 123, pi. 3, fig. 9, 9a.— Mulder and

Trebilcock, 1916: 81, pi. 11, fig. 4.— Briggs, 1918: 34,43.— Bedot, 1921:

29.— Blackburn, 1937: 368— Blackburn, 1942: 116.— Pennycuik,

1959: 180.- Ralph, 1961c: 109.- Millard, 1962: 301.- Millard, 1966:

494.— Berrisford, 1968: 394.— Day, et al., 1970: 14,.— Watson, 1973:

188, figs 54, 55.- Millard, 1975: 401, fig. 125 E-J.- Millard, 1978:

196.— Watson, 1982: 107, fig. 4.11f-h.— Stranks 1993: 13.

Monotheca spinulosa Stechow, 1921: 260.— Stechow, 1923:

17.- Leloup, 1932: 160.- Millard, 1975: 401, fig. 125 E-J.- Millard,

1978: 196- Hirohito, 1995: 278, fig. 96a-c.- Watson, 1996: 78.-

Watson, 1997: 529.— Watson and Mclnnes, 1999: 111 — Watson 2003:

243, 252.- Vervoort and Watson, 2003: 369, 374. - Watson, 2005:

542.- Bouillon et al., 2006: 368.

Plumularia spinulosa var. obtusa — Millard, 1957: 232.

Monotheca spinulosa var. obtusa.— Stechow, 1923d: 225.—

Yamada, 1959: 78.

Material examined. NMV F59053, microslide, Queenscliff Victoria,

Bale collection. Museum Victoria, nominated as probable syntype by

Stranks (1993).

Material in author’s collection : Stradbroke Island, south

Queensland, on alga, depth 20 m, coll: J. Watson, 24 August 1975.

Coniston Bay, Port Kembla, New South Wales, on bryozoan, depth 18

m, coll: J. Watson, September 1975. Gabo Island, Victoria, on ascidian

Herdmania grandis and red algae under ledges, depth 12 m, coll: J.

Watson, 15 February 1973. Port Phillip Heads, Victoria, on red alga,

depth 10 m, coll: J. Watson, 9 January 1984. South (20 km) of Cape

Bridgewater, Victoria, on red alga, depth 100 m, coll: V. Johnstone,

from fishing trawl, 16 May 1969. Port Noarlunga, South Australia, on

red alga, depth 10 m, coll: J. Watson, 10 February 1976. Abrolhos

Islands, outer reef. Western Australia, on red alga, depth 15 m, coll: J.

Watson, 17 October 1986. Karapuki Island, New Zealand, coll: D. A.

Staples 14 September 1975 on red alga.

Description. Hydrorhizal stolons ribbon-like, flattened to

substrate, outer flange with flexion joints in perisarc. Stems

monosiphonic, to 5 mm high, basal stem region athecate,

sometimes with two or three transverse nodes. Cauline

internodes variable in length among colonies but typically

moderately long and slender, straight to slightly curved, walls

smooth, expanding slightly distally to apophysis, nodes

Review of the genus Monotheca (Hydrozoa: Leptolida) from Australia with description of a new species and a note on

Monothecella Stechow, 1923

Figure 5. A-G, Monotheca spinulosa. A, part of stem; B, hydrocladium with short terminal spine; C, hydrocladium with long terminal spine; D,

female gonotheca with ova; E, two views of twin lateral nematothecae; F, axillar nematotheca; G, median inferior nematotheca. Scale bar: A, 0.3

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

transverse to slightly oblique, deeply constricted, an internal

transverse septum below node.

Hydrocladia alternate, distal on internode, in one plane,

borne on an upwardly directed apophysis, abcauline wall of

apophysis a smooth extension of internode, distal node a deep

transverse shoulder with internal septum behind node, a

minute mound-shaped hydrostatic pore in axil beside a

nematotheca. Hydrocladium with a short proximal athecate

internode, distal node transverse, deeply constricted with an

internal septum, occasionally an identical secondary athecate

internode between the athecate and hydrothecate internode.

Hydrothecate internode narrow, short, ending in a distal spine

of variable length, projecting beyond hydrotheca.

Hydrotheca occupying most of internode; internode

expanding distally as a prehydrothecal chamber between node

and floor of hydrotheca, often a minute septum projecting

backwards into chamber from floor of hydrotheca. Hydrotheca

bonnet shaped, laterally compressed, adcauline wall rounded,

set deep in internode, abcauline wall curved, proximally

convex, straightening towards margin; margin delicate, deeply

divided by a long V-shaped septum passing deeply backwards

into hydrotheca, a minute triangular in septum in base of some

hydrothecae. Hydranth with 12-14 tentacles.

Nematothecae bithalamic, moveable, all approximately of

same size: one cauline, halfway along internode on opposite

side to apophysis, one on internode in axil, one median

inferior, base close to internode, twin laterals standing erect in

front of hydrotheca, one at each side of distal spine, bases

short, slightly inflated, cup deeply adcaudally excavated into a

butterfly shape.

J.E. Watson

Male and female gonothecae large, one to several borne

near base of stem, inserted on a short indistinct pedicel beside

apophysis; body narrowly conical, facing upwards, walls

smooth to weakly undulated, distally truncate, closed by a

sheet of tissue, immature female gonophore with many ova.

Perisarc of stem and gonotheca moderately thick, thin on

hydrothecal margin. Colonies colourless to white.

Monotheca spinulosa, measurements, (p m)

Hydrorhiza, width

96-120

Stem

internode length

228-320

diameter at node

30-50

length of apophysis (adcauline wall)

30-40

Hydrocladium

length of athecate internode

32-56

width at distal node (shoulder)

40-60

length of base of thecate internode

(excluding spine)

160-192

length of subhydrothecal chamber

199-200

length of terminal spine

18-64

Hydrotheca

length of abcauline wall (measured

diagonally from base)

156-200

width of margin

136-168

Gonotheca

length (excluding pedicel)

1200-1560

maximum width (at margin)

640-740

Nematotheca

length of base

22-32

diameter of cup

22-36

Remarks. A microslide (NMV F59053) in the Bale collection

of Museum Victoria labelled ‘ Plumularia spinulosa Bale 1882,

Queenscliff’ was nominated as a probable syntype by Stranks

(1993). As this is the only known specimen, by monotypy it is

the holotype of the species.

The position of the cauline nematotheca varies between

colonies, typically being about halfway along the internode

but sometimes only one-third the distance up from the

proximal node. The position of the cauline apophysis is also

somewhat variable, ranging from well below to close to the

distal node. The axillar hydrostatic pore is very small and is

only seen with careful searching.

Stechow (1923) erected the var. obtusa for morphotypes

with a blunt terminal hydrocladial spine and Ralph (1961b)

erected the var. spinulosa for material from New Zealand with

a long terminal spine. Later studies, including the present one,

suggest the length of the spine is very variable. Watson (2005)

suggested it may be a response to environmental conditions

and evidence from this study supports this conclusion,

specimens with the longest spines being from deeper, less

turbulent habitat.

M. spinulosa is a very small species with a preference for

a substrate of flabellate red algae.

Type locality. Queenscliff, Victoria, Australia.

Known distribution. Temperate southern Australia to southeast

Queensland, Lord Howe Island, New Zealand, Japan, South

Africa, south Atlantic. Littoral to 100 m.

Monotheca togata (Watson, 1973)

Fig. 6 A-I

Plumularia togata Watson, 1973: 191, figs 65-67.— Bouillon et

al„ 2006: 371.- Stranks, 1993: 14.

IMonotheca togata — Vervoort and Watson, 2003: 374, fig. 91A-

Material examined. NMV F42060, holotype, microslide (malinol

mounted), Pearson Island, South Australia, on red alga Metagoniolithon

charoides, depth 33 m, coll: S. Shepherd 8 January 1969. Paratypes:

microslides NMV F42061, F42062, F42063, F4G2064, F42065,

F42066,F42105;SAMH40, Pearson Island onred algaMetagoniolithon

charoides , coll: S. Shepherd, 8 January 1969.

Description. Hydrorhiza a network of broad, flat stolonal tubes

with internal flexion joints. Stems short, to 4 mm long,

monosiphonic, with one to three short basal internodes, nodes

transverse, succeeding internodes moderately long and slender,

expanding distally to a broad shoulder at apophysis, a strong

transverse septum above and below node at level of apophysis,

distal node V-shaped, sometimes a weak transverse septum

about halfway up internode near level of cauline nematotheca.

Apophysis short, robust, just below distal cauline node,

slightly frontal on stem, upwardly directed, abcauline wall a

continuation of internode, adcauline wall set close to internode,

distal end wide, thickened, node transverse, a weak oblique

internal septum below node.

Hydrocladia alternate, one on cauline internode, slightly

forwardly directed, athecate internode very short, corrugated,

a strong transverse internal septum, distal node transverse.

Hydrothecate internode much longer than athecate internode,

inserted into shoulder of athecate internode with a slender

V-shaped joint; internode downwardly curved below

hydrotheca.

Hydrotheca cowl shaped (lateral view), scoop shaped

(anterior view), with a distinctly peaked adcauline margin (i.e.

opposite base of hydrotheca), prehydrothecal chamber short,

inflated, upper (adcauline) wall strongly convex, sometimes a

faint septum passing from internode into base, hydrothecal

margin hemispherical to triangular (anterior view), in lateral

view adcauline wall extending in a peak over margin; rim

thickened and outrolled.

Nematothecae bithalamic, moveable, all similar in shape

but differing slightly in length of base, cup robust, wide and

fairly shallow, adcaudally excavated almost to base; one

cauline, about halfway along internode on same side as

apophysis, one axillar beside apophysis, one median inferior

on hump of prehydrothecal chamber, just reaching base of

hydrotheca, twin laterals standing close together at either side

of a low distal protuberance of internode, bases short, sides

facing inwards to hydrothecal margin; in lateral view,

nematothecae appear to be almost inside hydrothecal margin.

Male gonotheca large, inserted on a short, straight pedicel

Review of the genus Monotheca (Hydrozoa: Leptolida) from Australia with description of a new species and a note on

Monothecella Stechow, 1923

Figure 6. A-I, Monotheca togata. A, stem; B, part of stem showing internodes; C, slightly oblique ventral view of hydrotheca; D, full ventral view

of hydrotheca, both views showing twin lateral nematothecae; E, apophysis of stem with proximal hydrocladium; F, gonotheca; G, cauline

nematotheca; H, median inferior nematotheca; I, one of twin lateral nematothecae. Scale bar: A, F, 1 mm; B, 0.3 mm; C-I, 0.1 mm.

J.E. Watson

beside apophysis on first stem internode, body barrel shaped,

distally truncated, walls smooth to weakly undulated, aperture

terminal, transverse, closed by a sheet of tissue. Female

gonotheca unknown.

Monotheca togata, measurements (/tm)

Hydrorhiza, width

72-80

Stem

length of cauline internode

300-360

width at subnodal septum

76-80

length of apophysis (adcauline)

40-50

Hydrocladium

length of subhydrothecal chamber adcauline

100-130

length of athecate internode

30-40

width at distal shoulder

44-64

length of hydrothecate internode (along

base)

180-220

Hydrotheca

depth, floor to highest point on margin

192-224

width across margin (frontal view)

186-200

Nematotheca

length of base of cauline

36-52

width of cup

26-40

length of base of median inferior

28-32

width of cup

48-52

width of cup of lateral

30-44

Gonotheca

length of pedicel

35-40

length excluding pedicel

1040-1260

distal diameter

600

Remarks. The cauline and axillar nematothecae are often

absent. The frontal position of the cauline apophysis causes the

hydrocladia and hydrothecae to twist forward during microslide

mounting, which tends to obscure some structures.

This is first description of the gonotheca of Monotheca

togata. The species is closely related to M. hyalina in size of

stems and the cowl-shaped hydrotheca with a hemispherical

margin. The distinctive adcauline apertural peak of M. togata

distinguishes it from M. hyalina.

Monotheca togata (NMNZ BS834), was reported from

East Cape and Ranfurly Bank, New Zealand, by Vervoort and

Watson (2003). However, the material differs from the

Australian species in having i) no axial hydrostatic pore, ii) a

much smaller hydrocladial median inferior nematotheca, and

iii) shorter cauline and hydrothecate internodes. In contrast to

the epiphytic Australian species, the New Zealand material is

epizootic on Synthecium suventricosum and Halopteris

campanula. These structural and substrate preferences may

indicate the New Zealand material is a different species, but

until more is known, the material is here doubtfully referred to

M. togata.

Type locality. Pearson Island, Great Australian Bight.

Known distribution. Southern Australia to Western Australia,

?New Zealand.

Key to species of Monotheca in Australia

1 Hydrotheca cup or bowl shaped.3

- hydrotheca not this shape.2

2 Hydrotheca scoop, cowl or hood shaped.6

3 Hydrotheca with conspicuous abcauline flange.

. Monotheca amphibola

- hydrotheca with inconspicuous abcauline flange or no

flange.4

4 Hydrocaulus flexuous, gonotheca barrel shaped with

distal aperture. Monotheca flexuosa

- hydrocaulus robust, gonotheca not barrel shaped.5

5 Aperture of gonotheca oblique with submarginal lobes.

. Monotheca pulchella

6 Hydrotheca with adcauline intrathecal septum.7

- hydrotheca without intrathecal septum.9

7 Intrathecal septum inconspicuous. Monotheca obliqua

- intrathecal septum well developed.8

8 Hydrocladium with terminal spine.

. Monotheca spinulosa

- hydrocladium without terminal spine.

. Monotheca australis

9 Hydrotheca subhemisphaerical, margin hooded.10

10 Hydrotheca with distinct adcauline marginal peak.

. Monotheca togata

11 Hydrothecal margin without adcauline peak.

. Monotheca hyalina

Acknowledgements

I thank the many colleagues who generously spent time

underwater with me over the years and the much appreciated

helpful criticism of the reviewers of the manuscript.

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