MEMOIRS

OF THE

EUM

MUS

QUEENSLAND

VOLUME 42

BRISBANE

PART 1

30 JUNE 1997

MEMOIRS

OF THE

(QUEENSLAND MUSEUM

BRISBANE

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SOME RECENT SPECIES OF THE GENUS ANASKOPORA WASS, 1975 (BRYOZOA:

CRIBRIOMORPHA) FROM QUEENSLAND P.W. ARNOLD AND P.L. COOK

Arnold, P.W. & Cook, P.L. 1997 06 30: Some Recent species of the genus Anaskopora Wass, 1975 (Bryozoa: Cribriomorpha) from Queensland. Memoirs of the Queensland Museum 42(1): 1-11. Brisbane. ISSN 0079-8835.

Several species of the genus Anaskopora have small, globular colonies, composed principally of frontally budded autozooids, kenozooids and avicularia. Hitherto, all known species have been Tertiary fossils from Victoria. A. parkeri sp. nov. is a Recent species here described from the Queensland continental slope, Colonies possess long rhizoids, which are inferred to have also occurred in fossil species with the same mode of growth and colony form. Cellepora doliaris Maplestone, 1909, a rare conical species with more regular frontal budding, originally found from New South Wales, hàs been referred in the past to the genus Reginella. It is. here redescribed from specimens collected on the continental slope of Queensland, and is reassigned to Anaskapora. The characters of the genus Reginella, which have been somewhat misunderstood, are discussed and the type species, R, furcata (Hincks, 1882), is redescribed. CT Bryozoa, Anaskopora, Reginella, taxonomy, morphology.

Peter W. Arnold, Museum of Tropical Queensland, 70-84 Flinders Street, Townsville, Queensland, 4810, Australia; Patricia L.Cook, Associate, Museum of Victoria, Melbourne,

Victoria, 3000, Australia: 6 March, 1997.

Colonies of cribriomorph Bryozoa tend to be encrusting and unilaminar, although a few fossil spe- cies were inferred to have had a ‘free-living’ habit.

In this paper, we report the discovery of a new species of deep-sea free-living cribriomorph an- chored by rootlets. We place the new species in the genus Anaskopora. Wass (1975) revised the genus Corbulipora MacGillivray (1895) and de- scribed several fossil species from the Victorian Tertiary, some of which he referred to a new subgenus Anaskopora. Recent revision of Corbulipora has shown that its species occur in multiple growth phases (see Bock & Cook, 1994), and that the species of Anaskopora are generically distinct (see Bock & Cook, in press).

The discovery of a Recent species of An- askopara makes a significant contribution to the interpretation of the colony structure of fossil species, several of which can now confidently be inferred to have been anchored by rhizoid sys- tems, Understanding of this mode of growth also allows the inference that the fossil species lived in and upon the upper layers of particulate sea- bottoms, and were part of a 'sand-fauna' (see Cook, 1981) of similarly adapted bryozoan species.

METHODS

Specimens were bleached in sodium hypochlo- rite solution and coated with AuPd or carbon for SEM. Abbreviations used: AM, Australian Mu- seum, Sydney: BMNH, Natural History Mu-

seum, London; MOV, Museum of Victoria, Mel- bourne; MTQ, Museum of Tropical Queensland, Townsville; QMG, Queensland Museum, Bris- bane,

Morphological terms used in the descriptions are defined in Bassler (1953) and Boardman & Cheetham (1983).

SYSTEMATICS

Class GYMNOLAEMATA Allman, 1856 Order CHEILOSTOMATIDA Busk, 1852 Infraorder CRIBRIOMORPHA Harmer, 1926 Family CRIBRILINIDAE Hincks, 1879

Anaskopora Wass, 1975

Anaskopora Wass, 1975:170. Bock & Cook 1996: in press.

TYPE SPECIES. Cribrilina elevata MacGillivray, 1895,

DESCRIPTION. Colony encrusting, originating, on very small substrata, often globular-to-coni- cal. Autozooids with septulae in the vertical walls, some or all of which become surrounded by à partially calcified chamber with an upper, cuticle-covered window. Zooids budded laterally and distally, or from chambered pores (see below) as interzooidal frontal buds. Interzooidal kenozooids and avicularia, together with rhizoids in some species, produced in the same way. Au-

2 MEMOIRS OF THE QUEENSLAND MUSEUM

lozooids with a costate pericyst, with lacunae: pelmatidia rare or ubsent. Secondary, calcified orifice with paired spines and a raised, distal, fimbriated plate. Avicularia distal or distolateral; occasionally proximal, rostrum rounded or sub- triangular, with paired condyles. Ovicells un- known.

The globular species of Anaskopora, in pårtic- ular, possess a kind of pore chamber, here called à ‘chambered pore’ to distinguish it from other forms of dictellae. This originates as a calcified chamber surrounding one or more septulae in the vertical walls ol zooids. The chamber is uncalci- fied on the frontal side, and the cuticle covering this window is able to expand intussusceptively so that the chamber may enlarge to form inter- zooial Kenozooids, autozooids, aviculana and rhizoids. Chambered pores occur in Cerbulipora, but are not the regular source of frontally budded zooids, as they are in the globular species of Anaxkopora, Some specimens of species de- scribed by Bock & Cook (in press), and a few of the colonies of A. parkeri (see below), originate upon very small shell substrata. These are rapidly covered by encrusting zooids with chambered pores al their comers. These pores produce an interzooidal, frontally budded series of zooids 1n subsequent astogeneti¢ generations, which com- pletely supersede any further encrusting growth. In Anaskopora doliaris. frontal budding of this kind seems to form the entire colony, and is very regular, so that the conical form is produced.

Anaskopora parkeri sp. nov. (Figs 1 A.B, 2A,B, 3)

MATERIAL. EXAMINED. HOLOTYPE: QMG- 21282, in alcohol, 'Cidaris UU Sin 42.2, 17721778. 146"48,52'E, [5.v, 1986, 296-202m, epibenthic sledge LL/2 inch inner liner), at MTQ. PARATYPES: QMG21283, one colony mounted on SEM stub, same data as holotype; QMG21284, one colony mounted on SEM stub, same data as holotype; QMG21285, 55 colonies in alcohol, same data as holotype: MOV F80820, 9 colonies in alcohol, same data as ho- lotype. ADDITIONAL MATERIAL: QMG212B8, 71 colonies in alcohol, same data as holotype; QNIG21289, 2 colonies in alcohol, "Cidaris 1" Stn 43.2, VI*34,58'S, 146°53,21°E, 15,v.1986, 458-500m, epibenthic sledge (1/2 inch liner), at MIO.

ETYMOLOGY. For the late Shane Parker, of the South Australian Museum,

DESCRIPTION, Colonies 0.50-5. 0mm in diam- gler, encrusting small foraminiferans or com- pletely free-living (Fig. 1A), anchored by large rhizoids (Fig. 1B). Autozootds with frontal shield

of 13-24 costae alternating with rows of 7-8 lacu- nac (Fig. 2A). Calcified orifice with one pair of flattened, sometimes terminally hifid, lateral-oral spines (Fig. 2A), paired lateral condyles and a raised distal plate with 5-8 fimhriations (Fig. 2B) und a pit at the base of the outer surface (Fig. 3). Operculum golden-brown, with a marginal scle- rite, filling the secondary orifice, Septulae 2-3 distal, 3-4 lateral, becoming surrounded by cham- hered pores (Fig. 3) which develop frontally as a series of 4-6 small kenozooids with cuticular frontals, surrounding each zooid. Avicularia disto- lateral (Figs 2A, 3), developed from chambered pores; occasionally paired or even proximal, Sub- rostral chambers raised, rounded (Fig. 3); mandi- bles with a marginal sclerite, oriented laterally. Zooids budded in alternating interzooidal series from chambered pores, oriented irregularly, col- ony forming an ectoproctolith (Figs LA, 2A, 3). Interzooidal kenozooids becoming extrazooidal.

Colony and zooid dimensions are given in Table 1.

REMARKS. Å. parker! closely resembles A, comuta (MacGillivray, 1895), which was re- ferred to Corbulipora by Wass (1975) but reas- signed to Anaskopora by Bock & Cook (in press), The two species have similar colony structures, and the autozooids of A. cornuta have numerous straight rows of costae, alternating with small lacunae, like those of A. parkeri. A. cornuta dil- fers in the distal position of the avicularium, which is not nearly as prominent as that of A, parkeri, The two species are so alike that it seems certain that the Victorian Tertiary species was a direct ancestor of the Recent form from Queens- land. 1t may also be inferred with a high degree of confidence that the colonies of A. cornuta possessed rhizoids and lived in a similar micro- environment to (hat inhabited by A. parkert.

Of 75 colonies of A. parkeri from Stn 42.2, 20 had well developed rhizoids which were 2-3mm long and, when turgid, 0.33-0.38mm wide. The colonies exhibited a developmental series; the

smallest had 6 zooids, the largest approximately

50 autozooids visible at the surface. Where the youngest stages encrusted small foraminifera there was a large, cuticle-covered cavity next to the primary zooid; in two cases, with a large thizoid beside it. Whether the cuticle-covered cavity is the ancestrula is not known, The rhizoids themselves terminated in numerous rootlets which had shell fragments and foraminiferans adhering to the cuticle. One colony was the sub- stratum for a small colony of a tubuliporid cyclo-

RECENT SPECIES OF ANASKOPORA 3

FIG. 1. A-B, Anaskopora parkeri sp. nov. A, QMG21283, frontally budded colony, x 32; B, Sketch of colony with rhizoids, x 15. C-D, A. doliaris (Maplestone). C, QMG21286, colony from frontal side, x 21; D, QMG21287, colony from basal side, x 31.

4 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2. A-B, Anaskopora parkeri sp. nov. A, QMG21284, general view of zooids, x 58; B, QMG21283, enlarged view of one zooid, x 100. C-D, A. doliaris (Maplestone). C, QMG21286, general view of zooids, x 42; D, QMG21287, enlarged view of one zooid, x 66.

RECENT SPECIES OF ANASKOPORA 3

FIG. 3, Anaskopora parkeri sp. nov, QMG2]283. Au tazooids showing details of orifices and frontal buds, x 37.

stome. Two others had the horny skeleton of Stephanoscyplius (SCyphozoa: Coronatae) grow- ing on them; Lagaiij (1963) described a similar case in which å hydroid grew on the globular colonies of Fedora nodosa.

The specimens of A. parkeri were accompanied by other rooted species associated with sand» fau- nas. These included Parmidaria sp., Parasticho- pora vanna (see Cook & Chimomdes, 1981) and Conescharellina sp.

Anaskopora doliaris (Maplestone) (Figs 1C,D, 20,D, 4A,B)

Cellepara daliaris Maplestone, 1909: 272, Pl. 77, Figs 10 A,B.

Reginella doliaris Brown 1958: 53, Hastings 1964; 254, Pl. 1, Figs 1-3. Pls 2 and 3

LECTOTYPE (indicated here); AMU201, 22 miles (34.4km) East of Port Jackson, New South Wales, 80 fath. (146m). HMCS ‘Miner’, labelled ‘TYPE’. PARALECTOTYPE: BMNH1909.11.12.14, as above.

MATERIAL. EXAMINED. QMG21286, G21287, ‘Cidaris 1' Stn 432, 17734.58' S, 14675321 E, 15.v.1986, 458-500m, epibenthic sledge (1/2 inch inner liner). 2 colonies on SEM stubs; at MTQ,

DESCRIPTION. Colony conical, one autozooid thick (Fig. 1D), inferred to be anchored by basal rhizoids, Autozooids erect, deeply immersed and oriented with the distal part of the orifice towards the ancestrula and the centre ot rhe colony (Fig. IC). Each autozooid with a series of simple septulae in the lateral, distal and proximal walls (Fig. 4B). all septulae becoming included in a senes of chambered pores which develop fron- tally inlo new uutozooids or aviculana (Fig. 4B).

The budding of autozooids is regular and radial, each astogenetic generation of autozooids alter- nating with one of kenozooids (Fig. 2C). Au- tozooid frontal shield with about 25 closely spaced costae, the distal costae thickened as an apertural bar which may be weakly bifid or thick- ened to form a median ridge (Figs 2C,D). Costae alternate with regular rows of 7-8 fine lacunae on each side of the mid-line. Secondary calcified orifice rounded, with a pair of flattened, curved bifid to trifid oral spines (Fig. 2D), a raised. fimbriated distal plate with 3 denticles (Fig. 4A) and distal pit (Fig. 4A), and a pair of condyles. Avicularia usually paired, oral (Fig. 2C), or sin- gle, proximal (Figs 1D, 4B); rostra raised, with paired condyles (Fig. 2C). Basal surface of cone hollow, with laree paired foramina surrounded by kenozooids and avicularia budded round the proximal ends of the successive generations of erect autozooids (Fig. 1 D).

Colony and zooid dimensions are given in Table 1.

REMARKS. Maplestone (1909) described the colony as encrusting, but there was no sign of any substratum in specimens we examined. Of the 39 species listed by Maplestone (1909) associated with A, doliaris from New South Wales, 1] were new Io science and 15 had a sand-fauna (see Cook, 1981) adapted colony form. Of these, 10 had rhizoids (7 species of Conescharellina, | of Zeuglopora, 1 of Sphaerapora, and | of An- askopora), and 5 were free-living, lunulitiforra species (1 of Lunularia, 3 of Selenaria and 1 of Orionella spp.).

Hastings (1964) gave a very full description and interpretation of colony growth and relation ships in C. daliaris. She noted the development of kenozooids from septulae, and figured and described the distal fimbriated plate. However, her account was written before the explanation of frontal budding by Banta (1972) and of reversed frontal budding by Cook & Lagaaij (1976). Has- tings used the structure of the ascophoran genus Conescharellina in her comparisons. The bud- ding pattern is analogous in the two exainples, although the type of frontal bud produced is quite different in Conescharellina, Hastings (1964: 258; 1966: 68) noted the similarities in develop- ment of intercalated avicularia and kenozooids in C. doliaris and Membraniporella agassizii Smitt (1873: 11, Pl. 5, Figs 103-106), from Florida. M. agassizii is ereet and quadriserial; the autozooids have costae without either pelmatidia or lateral costal fusions (see also Ristedt, 1979, Pl, 2, Figs

h MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. Anaskopora doliaris (Maplestone). A, QMG21286, details of orifices, x 90; B, QMG21287, details of kenozooids, x 55.

1,2). The gymnocystis distinct earl y in ontogeny, but later changes produce avicularia and kenozooids surrounding the autozooids which eventually obscure all but the orifices on the frontal surface of branches. The origin of the kenozooids and avicularia has not been de- scribed, but may be inferred ta be from cham- bered pores similar to those in Anaskopora. They appear to strengthen the delicate, erect branches. Hastings (1964) also emphasized the similarities in interzooidal budding from the pore-chambers of Hipporhoa with the budding of kenozooids in C. doliaris, The similarities in pore-chamber and chambered-pore structure are discussed by Bock and Cook (in press).

Anaskopora doliaris is known from only 4 col- onies. The lectotype and paralectotype were both from the *Miner' collections, one specimen hav- ing been sent by Maplestone to the then British Museum (Natural History) as a ‘cotype’. We infer that the two colonies of Anaskopora doliaris from the ‘Cidaris 1' collection were anchored hy rhi- zoids like those of A. parkeri, which probably emanate from the large, paired foramina on the

basal surface (Fig. 2C). It differs from A. parkeri and the other globular, fossil species, A. elevara, A. cornuta and A. rotundata (see Bock & Cook, in press), in the regularity of its reversed frontal budding and hollow basal surface, It resembles other species of Anaskopora in the mode of de- velopment and distribution of interzooidal kenozooids and avicularia, and the presence of distal fimbriated plate in the autozooids. The reasons for its removal from the genus Reginella are related to the history of the concepts and the characters of that genus which are discussed below.

Reginella Jullien, 1886

The attribution of Cellepora doliaris Maplestone, 1909 to the genus Reginella by Has- tings (1964) seems to have been the result of a series of misconceptions which have had several repercussions in the later interpretation of Aus- iralasian and other cribriomorph species.

Reginella was introduced by Jullien (1886: 605) ås a genus of Costulidae (=Cribrilinidae), for C. furcata Hincks (1882: 470, PI. 20, Fig. 5), a Recent species from the coast of British Colum- bia. No other species was included. Jullien's def- inition was not based upon the examination of specimens, but was an interpretation of Hincks's description and figure. Canu & Bassler (1929: 243) provided a literal English translation of Jullien's generic description, which mentioned that the frontal shield was composed of ‘velumi- nous ribs much in relief, and that the ‘intercostal furrows traverse entirely the zooecium and sepa- rate completely each pair of transverse ribs'. Pelmatidia were described as decreasing in size from the margin along each costa, and the lacunae were noted as each occupying ‘the middle of a calcareous polygonal cell. Osburn (1950: 179, Pl. 28, Fig. 3) redescribed R. furcata from Cali- fornian specimens, and noted that septulae were present, that the brooding zooid orifices were dimorphic, that the large ovicells were punctate and had a median keel, and that the oral spines were often absent. Hastings (1964: 253) also re- described K. furcata from Hincks's British Co- lumbian material and noted that Jullien’s ‘polygonal cells’ were *no more than an effect of light and shade’, There are, howeyer, other dis- crepancies between Hincks’s (1882) description and Jullien's (1886) definition. Hincks empha- sized that the costae were only ‘slightly raised", and that the intercostal *erooves! were ‘shallow’ and were arranged ‘radiating ta the median line’.

RECENT SPECIES OF ANASKOPORA

TABLE 1, Comparative measurements of colony, autozooid and avicularium in Anaskopora parkeri and A. doliaris, Original measurements calculated from scanning electron micrographs. Width of lacuna in microns; all other measurements in millimetres. The frontal shield length marked by an asterisk is a probable misprint in

Hastings (1964). Autozooid length is considered the same as frontal shield width.

Colony dimensions

Auiozooid length

1.06, 1.06, 1.08 0.69, 0.72, 0:76, 0.68

Fromal shield length, width 0.1* 4 0.5

Aperture antozooid

0.95. 1.01 L, 0.66, 0.70 W 0.21, 0.22, 0.22 L, 0.22, 0.24, 0.24 W

0.49, 0.60 L. 0.41 W 0.14, 0.15 L, 0.17, 0.18,0.21 W

Apertural bar width

Number ordenticleson | of denticles on fi (Number ordenticleson | late

0.15 03,04.04,04L.02,02 W

0.05, 0.07 0.1, 0.1 L, 0.06, 0.06 W

0.07, 0.07, 0,10

Width of costae

ih of pit at base of fimbriated Number of costae 9-13 (one side only)

Number ot lacunae per side Width of lacunae

Height of oral àvicularium

Width of basal kenozooidal

chambers

10-25

0,03. 0.03, 0.03, 0.04, 0.06, 0,06, 0,06,0.09

CO — |

0.07. 0.07, 0.09,0 09

5-8

ET |= 0.18,0.19 =E aiw

Aperture ot oral avicularium 0.16, 0.15 L, 0.12. 0.10 W

0.17, 0.17, 0.17 L, 0.13, 0.13, 0.13 W

D, 16, 0.19, 0,29

nor applicyble

This is in contrast to Jullien's prominent costae, and furrows which traversed the zooid horizon- tally. Cook (1985: 123) also examined Hincks's specimens of R. furcata and noted that the costae were broad and flattened, with pelmatidia and small lacunae, and remarked ‘the relationship of R. furcata with the various species subsequently assigned to Reginella requires investigation’. The type and other specimens of R. furcata have been re-examined, and a short description is given below,

Reginella furcata (Hincks)

Cribrilina furcata Hincks, 1882: 470, PI. 20, Fig. 5.

Reginella furcara O'Donoghue & O'Donoghue 1926: 52; Osburn 1950: 179, PI. 28, Fig. 3: Hastings 1964: 253; Soule et al, 1995: 123, Fig. 42 A-C,

MATERIAL EXAMINED. BMNH1886.3.16,18, Queen Charlotte Islands. Hincks Coll. on 4 shell frag- ments; 1921. [1.17.12 and 1968,1.18,100, Departure Bay, Vancouver Island, O'Donoghue Coll; 1986.9.10.3 Cannon Beach. Western Washington, L: Pitt Coll.

RETRO. Autozooids encrusting, colony

riginating from a membraniporiform ancestrula with long marginal spines. Zooidal gymnocyst very narrow, vertical walls shallow. Frontal

shield of 11-17 flattened costae, each with 3-4 pelmatidia, Suboral bar variously thickened, sometimes with å minute central mucro. Costae irregularly fused with a shallow central suture; the distal pairs of costae extending horizontally, the proximal 5-7 pairs converging distally and medially, Intercostal furrows narrow, with 4-6 rounded lacunae. Six evanescent oral spines, one lateral pair often remaining, and becoming thick- ened, or extended frontally and bifurcated termi- nally, Zooids communicating through 2 distal and 3 jateral groups of septulae, each enclosed in a very shallow, slit-like chamber at the base of the gymnocyst. Ovicells large, hyperstomial, with a median suture line and 16-20 small, rounded or irregular pores scattered over the surface; fertile orifices very slightly wider than autozooid ori- fices. Avicularia absent.

REMARKS. Osburn (1950) illustrated a single zooid of his Californian material of R. furcata. The drawing is misleading, as it resembles Jullien's concept of Reginella rather more than Hincks's description of R. fureata. The zooid appears to be at a late ontogenetic stage, with thickened, but not prominent, costae and the lines of lacunae traverse the frontal shield horizontally, with no obvious median suture line. Pelmatidia were reported to be present but were not

8 MEMOIRS OF THE QUEENSLAND MUSEUM

illustrated. Osburn (1950: 180, Pl. 28, Fig. 4 and PI. 29, Fig. 3) also illustrated another Californian species as Reginella mucronata, which had orig- inally been introduced for Pleistocene material from the same region by Canu & Bassler (1923: 92, PI. 35, Fig. 4). Canu & Bassler (1923) referred it to the genus Metracolposa Canu & Bassler (1917). Canu & Bassler's (1923) retouched pho- tographs show zooids very similar to that illustrated by Osburn for R. furcata, rather than the rounded zooids with large lacunae he illustrated for R. mucronata. Hastings (1964: 253) considered that these two species were iden- tical, although Soule (1959: 46), who had exam- ined both fossil and Recent material, had regarded R. mucronata as distinct. The genus Metracolposa was originally introduced for the Eocene species M. robusta Canu & Bassler (1917: 35, Pl. 3, Fig. 6; and 1920: 308, Pl. 43, Figs 1-7), from North Carolina. M. robusta and the other Eocene species described by Canu & Bassler were all illustrated by heavily retouched photographs, many of elongated zooids with un- interrupted lines of lacunae extending across the frontal shield, with little or no indication of a median suture line. M. robusta has large, erect, bilaminar colonies and zooids with paired oral avicularia. The ovicells are very large and ridged centrally, but not punctate like those of R. furcata. Generally, M. robusta, the type species, and the other Eocene forms assigned to Metracolposa, have little in common with, and are distinct from, Reginella. We accept that the Pleistocene-to-Re- cent M. mucronata is congeneric with R. furcata, but this provides no justification for regarding the genus Metracolposa as a synonym of Reginella. That idea had been very tentatively suggested by Osburn (1950) but was later greatly extended by Hastings (1964) in order to include other species with avicularia within the generic description (see below).

Jullien's (1886) definition and the studies just cited so modified the generic concept of Reginella that it now included characters quite unlike those of its type species. This would ex- plain the implicit referral of Cribrilina vas Brown, 1954, a Pliocene species from New Zea- land, to Reginella by Brown (1958), when dis- cussing a somewhat similar species from the Tertiary of Victoria, Reginella maplestonei Brown, 1958. Neither species has avicularia, but small, peristomial ovicells were illustrated in R. maplestonei. Powell (1967: 221, Pl. 2, Fig. 6, Fig. 4) redescribed R. vas from Recent New Zealand specimens, noting its elongated zooids, absence

of pelmatidia, and small, imperforate, peristomial ovicells; all characters completely unlike those of R. furcata. Brown (1958) also included the Re- cent, Antarctic Cribrilina projecta Waters, 1904, which has small oral avicularia and distinct pelmatidia, in his concept of Reginella. Hastings (1964) considered that C. projecta was not con- generic with R. furcata, but gave no evidence for her conclusion. Moyano (1985) introduced the genus Dendroperistoma for C. projecta.

The characters of Cellepora doliaris Maplestone, 1909 also became involved in the Reginella problem, when Brown (1958:53) men- tioned that Hastings (presumably in litt.), "has also pointed out a very close resemblance in the nature of the frontal shield between C. vas and Cellepora doliaris Maplestone'. Hastings, how- ever, did not redescribe C. doliaris for a further 6 years. When she did, she remarked (1964: 258, footnote 1), with a reference to Brown (1958), that Brown had told her in 1955 that he had ‘referred Cribrilina alcicornis and Cellepora doliaris to Reginella’ when, in fact, Brown (1958) had never mentioned C. alcicornis at all (see below).

C. alcicornis Jullien (1883: 508, Pl. 14, Figs 23-25) is a deep-water, encrusting species from the Northeastern Atlantic. Autozooids have cos- tae with no pelmatidia, and four, large, branched oral spines. Interzooidal avicularia and large interzooidal kenozooids are budded among the autozooids, and the elongated ovicells are not punctate. C. alcicornis is therefore unlike R. furcata, and it is significant that Jullien (1886) did not include C. alcicornis when introducing Reginella, and that later, Calvet (1907:399) and Prenant & Bobin (1966: 578, Fig. 200), continued to refer the species to Cribrilina. More recently, both d'Hondt (1974: 47, Fig. 7) and Harmelin (1978: 178, Pl. 1, Fig. 3, Figs 3-4), who examined additional Atlantic material, also maintained C. alcicornis in Cribrilina. Harmelin noted that one of Hastings's ‘points of resemblance’ between C. doliaris and C. alcicornis was based on the mis- taken assumption that Jullien's (1886) phrase ‘grande ponctuations referred to the interzooidal kenozooids, whereas in fact, it referred to the intercostal lacunae. He remarked "Le changement de genera ne semble pas justifié pour cette espèce’.

Quite apart from this confusion, it is unfortun- ate that Hastings (1964: 252) extended Osburn’s (1950) tentative synonymy of Metracolposa with Reginella, including some of the characters of the Eocene M. robusta, such as avicularia, in her

RECENT SPECIES OF ANASKOPORA 9

concept of. Reginella. Thus the presence of nu- merous avicularia in C. doliaris, which are totally absent in Å. furcara, did not preclude its referral to Reginella, because avicularia had become å "generic character". In the same way, the presence of kenozooids with the avicularia in C. doliaris, was mitigated by the tacit assignment of Cribrilina alcicornis, which has both, ta Reginella, although neither were present in R. furcata. The presence of avicularia and kenozooids in both C. alcicornis and C. doliarts is interesting. but the structures differ completely in their origins and general morphology between the 1Ww0 species. In any case, neither has anything 10 do with the characters of the genus Reginella, Two North American species which have been referred tu Keginella also require further investi- gaton. Reginella floridana (Srnitl, 1873) has been described by Winston (1982) from Florida, and by Cook (1985) from West Africa. No ovicells have ever been found in any specimen of (bis species. R. repangulata Winston & Håkansson, 1986, from an interstitial Floridan fauna, has ovicells somewhat similar to those of R. furcara.

It is obvious that by this time the original char- acteristics of the genus have become completely submerged by the accumulation of additional or alternative features derived from the diversity of species included. This is illustrated by Gordon (1984: 63, Pl. 20, Figs D-E), who redescribed Reginella vas, and introduced a new, unisenal encrusting species from New Zealand, R. stolonifera Gordon (1984: 63, Pl. 20, Figs A-C), which had large, branched oral spines like those of C. alcicornis. The concept of the genus was now extended to include all the disparate forms mentioned above, from the Arctic, Antarctic, At- lantic and Pacific Oceans, from deep and shallow, even interstitial waters, and from the Tertiaries of North America, Australia and New Zealand. The generic characters now encompassed: colonies encrusting, erect and bilaminar, and conical; zo- vids uniserial to contiguous and frontally budded, avicularia present or absent; ovicells hyperston- jal and peristomial with or without pores or fenes- trae; pore chambers present or absent, To these may be added, pelmatidia and kenozooids present or absent. The only consistent characters are 'slit- like intercostal lacunae’, and “costae arranged in straight rows across the zooid, often traversing the mid-line without interruption’. Although these features occur late in ontogeny in Å, vas and R. stolonifera, neither occurs in R. furcata.

In conclusion, it is obvious that Reginella should be restricted to Recent and Pleistocene species from the North American ànd Japanese Pacific coasts, and that Merracolposa should be retained only for the American Eocene species. Soule, Soule & Chaney (1995) give. notes on Reginella furcata and introduce å new genus. Reginelloides, for R. stolonifera Gordon, 1984. Cribrilina alcicornis, C. vas, Reginella maplestonei, R. floridana and R. repangulam should also he re-investigated, and new generic groups should be introduced for their reception where necessary.

The colonies of R. furcata from Vancouver Island examined by one of us (PLC) included about 20 with ancestrulae present. The an- cestrulac had small pore-chambers distally and laterally, and 12 lung spines surrounded the ope- sia. Mawatan (1988: 149, Figs 9-14) hus de- scribed specimens from Hokkaido, Japan, as Reginella furcata, and discussed previous Japan- ese records in detail. These include the species described as Lyrula multipora by Sakakura (1935: 109, PI, 8, Fig. 7), and as Figularia multi- parabySilén (1941: 117, Figs 178-180). All these Japanese records differ trom AA, furcata very slightly, in the complete lack of oral spines, even in early ontogeny, the raised mucros beside the orifice which give it å subtriangular appearance, and the lack of any dimorphism in the fertile orifices. Osburn (1950) described several nomi- nal taxa from the Arctic to the Mexican coasts of North America and referred them to Reginella. They all resemble R. furcata in possessing straight, converging rows of lacunae, paired oral spines and large ovicells with a central suture and punctate surface. Until the actual limits of varia- non of these species, especially those from Cali- fornia, like R. mucronata, have been analysed, it is probably best to maintain them, together with R. multipora, as distinct species of Regineffa.

ACKNOWLEDGEMENTS

We thank Dr PE. Bock (Royal Melbourne Institute of Technology) for his help and interest. Ms M. Spencer Jones (Natural History Museum, London), for the loan of specimens of Reginella

furcata and Dr P. Berents (Australian Museum)

for confirming details of Maplestone's type of C. doliaris. The 'Cidaris' expeditions on RN. ‘Franklin’ were partially funded by an Australian Manne Science and Technology Grant to Prof. M. Pichon,

10 MEMOIRS OF THE QUEENSLAND MUSEUM

LITERATURE CITED

ALLMAN, G.J. 1856. ‘A monograph of the freshwater Pulyzoa, including all the known species, both British and foreign’, (The Ray Society: London),

BANTA, WC 1972. The body wall of Cheilostome Bryozoa, V. Frontal budding in Schizoporella uni- cornis floridana, Marine Biology 14(1): 63-71.

BASSLER, RS. 1953, Bryozoa. In Moore, R.C, ed) “Treatise on Invertebrate Paleontology’, Part G. ( University ol Kansas Press: Lawrence, Kansas).

BOARDMAN, R.S, & CHEETHAM, A.H. 1983. Glos- sary of morphological terms. Pp. 304-320, In Boardman, RS et al, (eds) Bryozoa (revised), In Robison, R.A. et al. (eds) ‘Treatise on Inveriehrate Paleontology’, Part G. (University of Kansus Press: Lawrence, Kansas) .

BOCK, PE. & COOK, PL. 1994. Occurrence of three phases of growth with taxonomically distinct zooid morphologies. Pp 33-36, In Hayward, PI. Ryland, J.S, & Taylor, PD, (eds) "Biology and Palacobiology of Bryozoans’. (Olsen & Olsen: Fredensborg).

In press. A revision of the genus Anaskopora Wass (Bryozoa: Cribriomorpha). Memoirs of the Mu- seum of Victoria.

BROWN, DA. 1954. Polyzoa Irom a submerged Lime- stone off the Thrce Kings Islands, New Zealand. Annals and Magazine of Natural History (12)7: 415-437,

1958. Fossil cheilostomataus Polyzoa from South- west Victoria, Memoirs of the Geological Society of Victoria 10: 1-90.

BUSK, G, 1852, "Catalogue of marine Polyzoa in the collections of the British Museum". Vol.1, pp. 1-54, (Trustees of the British Museum: London),

CALVET, L. 1907. Bryozoaires. Expeditions scientifiques du ‘Travailleur et du ‘Talisman’ 1880-1883, 8: 355-495.

CANU, F. & BASSLER, R.S, 1917, A synopsis of American Early Tertiary cheilostome Bryozoa. Bulletin of the United States National Museum 96: 1-87,

1920, North American Early Tertiary Bryozoa. Bul- e: of the United States National Museum 106: 1-879

1923. North American Later Tertiary and Quater- nury Bryozoa. Bulletin of the United States Na- tional Museum 125: 1-302.

1929. Bryozoa of the Philippine region, Bulletin of the United States National Museum 100(9): 1- 685.

COOK, P.L. 1981. The potential of minute bryozoar colonies in the analysis of deep-sea sediments. Cahiers de Biologie marine 22: 89-106,

1985, Bryozoa from Ghana —a preliminary survey. Annales du Musée royale de l'Afrique central (Sciences zoologiques) 238: 1-315,

COOK, P.L. & CHIMONIDES, PJ, 1981. Morphology and systematics of some rooted cheilostome

Bryozoa. Joumal of Natural History 1501) 97- 134

COOK. P.L. & LAGAAN, R. 1976, Some Tertiary and Recent conescharelliniform Bryozoa. Bulletin of the British Museum (Natural History) Zoology 29/6): 317-376.

GORDON, DP 1984, The marine fauna of New Zens land: Bryozoa: Gymnolaemata from the Kermadec Ridge, New Zealand Oceanographic Institute Memoir 91: 1-198,

GORDON, D.P. & HASTINGS, A.B. 1979, The inter- zooidal communications of Hippathoa sensu lato (Bryozoa) and their value in classification, Journal of Natural History 13; 561-579.

HARMELIN. J.-G. 1978. Sur quelques Cribrimorphes (Bryozoa, Cheilostomata) de |’ Atlantique Orien- tal. Tethys 82): 173-192.

HASTINGS, A.B. 1964, The cheilostomatous Polyzoa Neoeathyris woosteri (MacGillivray) and Reginella doliaris (Maplestone). Bulletin of the British Museum (Natural History) Zoology 11 (3): 243-262,

1966, Observations on the type-material of some genera und species of Polyzoa. Bulletin of the British Museum (Natural History) Zoology 14(3): 55-78.

HINCKS, T, 1879, On the classification of the British Polyzoa. Annals and Magazine of the Natural History (5)3: 153-164.

1882, Report on the Polyzoa of the Queen Charlotte Islands. I, Annals and Magazine of the Natural History (5) 10: 459-471,

D'HONDT, J.L. 1974, Bryozoaires récoltés par la "Thalassa! dans le Golle de Gascogne (Campagnes de 1968 å 1972), Cahiers de Biologie marine 15: 27-50.

JULLIEN, J. 1833. Dragages du ‘Travailleur’. Bryozoaites. Espéces draguées dans l'Océan Atlantique en. 1881. Bulletin de la Société Zoologique de France 7(5): 479-529.

1886, Les Costulidécs, nouvelle Famille de Bryozbaires. Bulletin de la Société Zoologique de France 11(4): 601-620.

LAGAAL. R. 1963. New additions to the Bryozoan Fauna of the Gulf of Mexico. Publications of the Insticute of Marine Science, University of Texas 9: 162-236,

MACGILLIVRAY, P.H. 1895. A Monograph of the Tertiary Polyzoa of Victoria. Transactions of the Royal Society of Victoria n.s. 4: 1-166.

MAPLESTONE, C.M. 1909, The resulls of Deep-Sea Investigations in the Tasman Sea 1 (5) The Poly- 20a. Records ol the Australian Museum 7: 267- 273.

MAWATARI, S.F 1988. Two Cheilostomatous Bryozo- ans New to Hokkaido. Memoirs of the National Science Museum 21: 145-151.

MOYANO, H.LG. 1985. Briozoos marinos Chilenos. V. Taxa nuevos o poco conocidos, Boletin de la Sociedad Biologia de Concepcion 56: 79-114.

RECENT SPECIES OF ANASKOPORA 1

O'DONOGHUE, C.H. & O'DONOGHUE, E. 1926. A second list of Bryozoa(Polyzoa) from the Vancou- ver Island Region. Contributions to Canadian Bi- ology n.s. 3(3): 49-131.

OSBURN, R.C. 1950. Bryozoa of the Pacific Coast of America Part 1. Cheilostomata Anasca. Report of Allan Hancock Pacific Expeditions 14 (1): 1-269.

POWELL, N.A. 1967. Polyzoa (Bryozoa) - Ascophora — from North New Zealand. 'Discovery' Reports 34: 199-393.

PRENANT, M. & BOBIN, G. 1966. Bryozoaires, Part 2. Chilostomes Anasca. Faune de France 68: 1- 647.

RISTEDT, H. 1979, Skeletal ultrastructure and as- togenetic development of some Cribrimorph By- rozoa. Pp. 141-152. In Larwood, G.P. & Abbott, M.B. (eds), “Advances in Bryozoology'. (Aca- demic Press: London).

SAKAKURA, K. 1935. Bryozoa from Toyama Bay, Sea of Japan. Annotationes Zoologicae Japonenses 15: 106-119.

SILEN, L. 1941. Cheilostomata Anasca (Bryozoa) Col- lected by Prof. Dr Sixten Bock's expedition to Japan and The Bonin Islands 1914. Arkiv för Zoologi 33A(12): 1-130.

SMITT, F.A. 1873. Floridan Bryozoa, collected by Count L.F. de Pourtales. Part II. Kongliga Svenska Vetenskaps-Akademiens Handlingar 11(4): 1-83.

SOULE, J.D. 1959. Results of the Puritan-American Museum of Natural History Expedition to Western Mexico 6. Anascan Cheilostomata (Bryozoa) of the Gulf of California. American Museum Novitates 1969: 1-54.

SOULE, D.F., SOULE, J.D. & CHANEY, H.W. 1995, Taxonomic atlas of the benthic fauna of the Santa Maria basin and western Santa Barbara channel. Volume 13, The Bryozoa. Irene McCulloch Foun- dation Monograph Series, Number 2: vi + 344 pp. (University of Southern California Press: Los An- geles).

WASS, R.E. 1975. A Revision of the Bryozoan Genus Corbulipora MacGillivray. Proceedings of the Royal Society of Victoria 87: 167-173.

WATERS, A.W. 1904. Bryozoa. Expedition Antarctique Belge, Resultats Voyage S.Y. 'Belgica' 1897- 1899. Zoologie 4: 1-114. Anvers.

WINSTON, J.E. 1982. Marine Bryozoans (Ectoprocta) of the Indian River Area (Florida). Bulletin of the American Museum of Natural History 173(2): 99-176.

WINSTON, J.E. & HAKANSSON, E. 1986. The Inter- stitial Bryozoan Fauna from Capron Shoal, Flor- ida. American Museum Novitates 2865: 1-50.

12 MEMOIRS OF THE QUEENSLAND MUSEUM

SYNONYMY OF CTENOTUS MONTICOLA STORR, 1981 AND CTENOTUS HYPATIA INGRAM AND CZECHURA, 1990, Memoirs of the Queensland Museum 42(1): 12. 1997:- Cienotus monticola and C. hypatia are mediurn-sized. Both are classed "poorly known" (McDonald er al, 1991), The former has å ‘a very restricted distribution

.. with a maximum geographic distribution of less than

100kn". The latter is"... known only from the type collection", C. monticala was described from à series of specimens col- lected *77km W af Mareeba, Queensland (17°02'S, 145920 EI The type locality of C. hypatia is ' grantte gorge, 15km W of Mareeba, (17°00'S, 145" 1 7' EJ , These localities are both on Granite Ck, approximately 6km apart (J. Cov- acevich pers. comm. ). That, and the fact that the type descrip- tions of C. monticola and C. hypatia sre very similar. prompted a re-examination of the type material of each to determine whether or not they are distinct.

Re-examination of the (ype specimens and type descrip- lions indicates that C, monticola and C. hypatia are conspe- cific (Table 1). Almost without exception, the measurements and scale characters of C. hypatia fall well within the range nf values. provided by Storr (1981) for C. monticola. The differences in elongation, indicated by the chin-vent and

aravertebral counts are inconsequential. Greater variation is requently recorded in a single species (e.g, C. astiemis Horner, 1995 & C. stuartt Horner 1995). Additionally, both type specimens share the following characters: nasals separated; nasal groove absent, rostral and fronronasal in narrow to moderate contact; second loreal 1.2 times as wide as high; prefrontals large and separated, frontal long and narrow con- tacting the prefrontals, the frontonasul, the first 3 supra- oculars, and the frontopanetals, 4 supraoculars, 2nd the largest; Ist supraciliary the largest, 4th to penultimate consid- erably smaller than others; Sth supralabial subacular, ear aperture large; toes compressed,

Colour and pattern are known to be useful in distinguishing some species of Crenotus (e.g, C. areanus Czechura. E Wombey, 1982 and C. robustus Storr, 1970). However, this is not the case with C, monticola and C. hypatta, Colour and pattern. of the type specimens of C. monticola (5) and C. kypatia (1) have been detailed by Storr (1981) und Ingram & Czechura (1990). Variution in the former fully covers the fatter.

Thus, in every respect (meristics, sealation, colour/pattern) there are no significant differences between C, monricola and C. hyparia. Ctenotus mouticola Storr, 1981 is, therefore, a senior synonym of C. hyparia Ingram & Czechura, 1990,

Acknowledgements We thank Jeanette Covacevich for reading and suggesting Improvements to this nole,

Literature Cited

CZECHURA, ON E WOMBEY, J. 1982, Three new striped skinks, (Crenoms, Lacenilia, Scincidae) from Queens- land. Memoirs of the Queensland Museum 20(3): 639-43,

HORNER, P. 1995. Two new species of Crenotus (Reptilia. Scincidae) from the Northern Territory. The Beagle, Records of the Museums and Art Galleries of the North- em Territory 12:75-88.

TABLE |, Comparison of SE of C. monticola (QMJ39468) and C, hypatia (QM 42092). Values in paren theses are from the type descriptions.

Species

Character C. monticola

(QMJ39468) (QM142092) Snout-vent (mm) 53 (51-61) i (SNL) 44 (44) | 215 (215)

13 (NA) 12012)

20 (NA) 19 (NA)

KU 0r 8) 8 (9)

Upper ciliaries

[Presuboculars — | 2(2.rarely 3) |

ae canc ^

| Upper labials ZO) | Ear lobules 2 or 3 (3-5) 3 or 4 (3-4) E arg I or 2 (NA)

Largest ear lobule JL usually langest)| nuchal scales 26 (24-28) 28 (28)

Midbody scale rows 76 (NA) N2(753

No. of scales from chin-vent

Paravertebral scale rows

Subdigital lamellae 18/19 (17-19) 18/19 (20)

Structure of moderately wide | narrow callus subdigital lamellae | callus (narrow to (keeled) moderately wide callus)

&7(NA) T2(NA)

INGRAM, G. J. & CZECHURA, G.V. 1990. Four new spe- cies of striped skinks from Queensland. Memoirs of the Queensland Museum 29(2): 407-410,

McDONALD. K.R., COVACEVICH, J.A., INGRAM, G.J. & COUPER, P.J. 1991. The status of frogs and reptiles. Pp 338-345. In Ingram, GJ, & Raven, R.J. (eds), “An Atlas of Queensland's Frogs, Reptiles, Birds and Mammals", (Queensland Museum: Brisbane). 391 pp.

STORR, G.M. 1970. The genus Crenoius (Lacertilia : Scincidae) in the Northem Territory. Journal of the Royal Society of Western Australia 32: 97-108.

19831. Ten new Crenotus (Lacertilia: Scincidae) from Aus- tralia. Records of the Western Australian Museum 2): 125-146,

P.J, Couper, Queensland Museum, PO Box 3300, South Bris-

bane, Queensland 4107, Australla. Pianka, E.R., Department

af Zoology, The University of Texas at Austin, Austin, Texas,

78712-1064, USA: 23 May 1997

A NEW GENUS OF HIPPOLYTID SHRIMP (CRUSTACEA: DECAPODA; HIPPOLYTIDAE) FOR THOR MALDIVENSIS BORRADAILE

AJ. BRUCE

Bruce, A.J. 1997 06 30: A new genus of hippolytid shrimp for Thor maldivensis Borradaile. Memoirs of the Queensland Museum, 42(1) 13-23. Brisbane, ISSN 0079-8835,

Å new genus Thorina, is designated to accommodate the hippol ytid shrimp Thor maldivensix Borvadaile 1915, The new genus is distinguished from Thor particularly by the short, unidentate rostrum and the absence of an appendix masculina from the male second hate with marked sexual dimorphism of the first pereiopods. First recorded from thé

aldive Islands, the species is sparsely recorded throughout the Indo-West Pacific region and newly recorded from Tanganyika, Mauritius, Seychelle Islands, Western Australia, the Great Barrier Reef, Papua New Guinea, Tonga, Cook and Society Islands. [ ] Natantia, Hippolytidae, Thorina , Indian Ocean, Pacific Ocean.

AL Bruce, Crustacea Section, Queensland Museum; P.O. Box 3300, Sauth Brisbane,

Queensland, 4101, Australia: 17 May [996.

In his study of the hippolytid shrimps of the Albaiross Expedition, Chace (in press) indicated ihe anomalous systematic position of Thor maldivensis Borradaile 1915. He clearly defined the characters of the genus Thor Kingsley s. sit. and lists five major characters that distinguish T. maldivensis trom all other species of the genus Thor, omitting it from his key to the species of that genus. A new monospecific genus for the reception of this species is here formally estab- lished. The original description of Thor maldivensis provided by Borradaile (1915) con- sisted of a brief diagnosis only but a short more detailed account with an illustration of å d exam- ple was published in Borradaile (1917). This ac- count remains the most detailed available of this species, which has been only infrequently re- corded in the zoological literature. Abbreviations used: CL, postorbital carapace length; MNHN, Museum national d'Histoire Naturelle, Paris; NTM, Museums and Art Galler- ies of the Northern Territory, Darwin; QM, Queensland Museum, Brisbane.

SYSTEMATICS

Class CRUSTACEA Order DECAPODA Suborder NATANTIA Infraorder CARIDEA Family HIPPOLYTIDAE

Thorina gen. nov,

DIAGNOSIS. Small hippolytid shrimps of sub- cylindrical body form. Rostrum very short, slen- der. acute, not exceeding proximal segment of

antennular peduncle, with single small dorsal tooth only. Carapace without carinae; supraor- bital, non-articulate antennal spines present, he- palic and pterygostomial spines absent. Antennular peduncle with proximal segment with moveable plate distally, upper flagellum brush- like. Scaphocerite mid-laterally unarmed, Eyes well-developed, cornea hemispherical, With elongate acute median process anterior to lirst thoracic sternite. Mandible without palp, with incisor process, maxillipeds with epipods and well developed flagella on exopods. Pereiopods without arthrobranchs, epipods or exopods. First pereiopod with fingers greater than 1/3 of palm length: fingers without interlocking terminal spines, merus with ventral teeth, chelae usually greatly hypertrophied in some d d. Second pe- relopod with carpus with 6 segments. Propods of last 3 pereiopods not multiarticulate. Abdomen non-cannate, without dorsal teeth, with anterior 3 pleura rounded, posterior 2 posteroventrally acute, 6th abdominal segment without articulated posteroventral plate. d second pleopod without appendix masculina and appendix interna, Telson with 3 pairs of dorsal spines, 3 pairs of posterior spines.

TYPE SPECIES. Thor malilivensis Borradaile, 1915.

ETYMOLOGY. From Thor, a hippol ytid generic name first used by Kingsley, 1878, and -ina (Latin), diminu- tive. Gender, feminine.

SYSTEMATIC POSITION. Closely related 10 Thor Kingsley, with which most generic charac- ters are shared and emphasised by the presence of a triangular mobile plate on the distal segment of the antennular peduncle, a feature otherwise

Li MEMOIRS OF THE QUEENSLAND MUSEUM

found To eceur only in Thor, Distinguished fram Thor by the greatly reduced rostrum with only å single dorsal tooth, marked sexual dimorphisn of first pereiopods, reduced sexual dimorphism ot Id pereiopods, presence of 3 pairs of posterior telson spines instead of 1-2 pairs, and particularly, the complete absence of an appendix masculina and appendix interna on the à 2nd pleopod. The strongly distolaterally spinose meri of the hyper- trophied å Let pereiopods are also characteristic, but it may be noted that a single small articulated spine may be present in the West Atlantic species Thor dobkini Chace (1972). In Thorina, the spines are non-articulate denticular processes.

Thorina maldivensis comb, nov, (Figs 1-6]

ror maldiversis. Borradaile, 1915: 208-209; 1917: 301-407, på. 56 fig, 6; Kemp, 1916: 391; Edmond- sen, 1925: 6; 1946: 252, 253, fig. 153d; Holthuls, 1953: 53-54; Bruce, 1976: 51; Kumezaki, et al., 1988: 8], col, pl.; Chace, in press.

MATERIAL EXAMINED. QMW 19914, 13. ] ovig. 9, stn DF,37, Heron Island, Capricorn Islands, Queensland, 3.0m, 16 October 1976, coll, D.F. Fisk. QMW21438, 13, 1 ovig. 9, stn AJB/162, Jadini, Kenya, 22 December 1972. c. 0.4m, lagoon, in Acropora, coll. A.J. Bruce. QMW21439, Id, Aldabra, Seychelle Islands, 3 Navember 1964, from coral in channel reef, intertidal, coll. AJ. Bruce. QM W21 440, 2 spms, macerated, Ras Iwaline, Mombasa, Kenya, 8 February 1972, lagoon, in Millepora, coll. AJ. Bruce. QMW21441, 1, 1 ovig, 9, sin 140, Kirwetu, Kenya, 346,7 S 39?50.9'E, low water spring tide level, 6 November 1971, coll, A.J, Bruce, reef flat, under dead coral, QMW21442, 4 ovig. ?, Astove Island, Seychelle Islands, R.V, Manihine, Cr. 312, 20 August 1970, reef flat, under dead coral, coll. A.J. Bruce. QOMW21443, 34, 30vig. 2 Farquhar Island, Seychelle Islands, R.V. Manihine, Cr.336, sin AJB/60, 26 Febru- ary 1972, coll. AJ, Bruce. QM W21444, 2 spms (1 ovig. 9), stå AJB/107, Ras Iwatine, Mombasa, Kenya, 4"01,3'5 39*44.0' E, 1m, 27 February 1971, lagoon, in Srylophora, coll. A.J. Bruce. QM W21445, 1 spm (3 7), sin AJB/99, Ras Twatine, Mombasa. Kenya, 4*04,0'5 19944, YE, 0,5m, | January 1971, lagoon, In Pacillopara, coll. A.J. Bruce. QM W21446, I ovig. 9, sin AJB/157, Jadini, Kenya, 4"19,0'5 39935,5'E, 2.001, 19 March 1972, in corals, coll, AJ. Bruce. OM W21447, 1d, 1 ovig. F, stn AIB/I66, Bamburi, Mombasa, Kenya, 4900,5'5 39995.0'E, outer reef creat, 18 August 1973, coll. A.J, Bruce, QM W21448, HV, 2 juvs, stn AJB/119, Ras Iwatine, Kenya, 47 D1.15'839*43.8' E, 2m, 27 July 1971, edge of deepreet channel, in Pavona, coll, AJ, Bruce: QMW21449, | 9, stn AJB/I38, Jadini. Kenya, 4°21.5'S 39*34,5 E, 0- 2m, nter lagoon, in coral. 3 November 1971, coll, al Bruce. (author's collection), I spm, sm ATB/181a,

Tamannd, Mauritius, intertidal pools, 24 May 1974, coll. AJ. Bruce, QMW21540, 1d, 3 op, ?, sth AJB/140, Vipengo, Kenya, 6 November 197 1 reef Nat, coll. AJ. Bruce. (author's collection), 2 spms, stn AJB/209, South Patch, Motupore Island, Papua New Guinea, 9°34.27'S 147*12,65' E, 10-20m, seaward reef slope, scuba, 6 November 1980, coll. J.M. Lowry. QMW?21451. (og 2, stn AJB/141, Tutia Reef, Mafia Island, Tanganyika, 2m, 14 November 1971, voll, AJ. Bruce, NTMCr 009156. 14, stn RH 92 12, Cartier Reef, Western Australia, 127226'$ 123?32,2"E, 10-15m, 9 May 1992, coll. J.R, Hanley, BC. Russell, NTMCr004374, 13, 1 ovig, 2,2 juv., Masausi Bay, Tutuila, American Samoa, reef l. N.4m. 22 July 1986, ‘Operation Raleigh", coll. M. Ricli- mond. NTMCr.004375, 13, Omuta Landing, Mitiaro, Cook Islands, reet lagoon, 2-3m, 11 July 1986, "Omer: ation Raleigh’, coll. M. Richmond. NTMCr 004376, I ovig. "d, Oholei, Tongatapu, Tonga, reef lagoon, 1m, 14 August 1986, ‘Operation Raleigh’, coll. M. Rich- mond, MNHNNa-6627, 29 (1 ovig.), sin S-20, Tahiti, Society Islands, coll. O, Odinetz, 1982.

DESCRIPTION. Small hippolytid shrimp ol moderately slender, subeylindrical body form. Carapace smooth, with short acute rostrum, reaching to midlength of proximal segment of antennular peduncle in dorsal view, without dis- unci dorsal carina, with single small acute toath at about level of tip of inferior orbital angle in lateral view, ventral carina shallow, concave, lat- eral carinae broadly divergent, continuous with supraorbital margin, giving rostrum triangular appearance in dorsal view, with small acute sti- praorbital teeth, posterior 10 level of dorsal tooth; inferior orbital angle slightly produced, rounded, with small acute antennal spine ventrolaterally; hepatic and pterygostomial spines absent, branchiostegite with anterolateral angle broadly rounded.

Abdomen smooth, with 3rd segment slightly produced posterodorsally, non-carinale; 6th seg- ment c. 1.33 times longer than maximal depth, c. 0.7 of carapace length in d, 0.55 in 2, with posterolateral and posteroventral angles acute, posteroventral angle non-articulate; first 3 pleura broadly rounded, not markedly enlarged in ovigerous ? 2, 4th and Sub posteriorly produced, posteroventral angles broadly acute.

Telson c. 1.5 times length of 6th abdominal segment in d, L3 times in 2, subequal (0 cara: pace length in g, c. 0.73 in H. 2.6 times longer (han anterior width, lateral margins sublineur, posteriorly convergent, posterint margin angular with acute median process, willy å pairs of small dorsal spines at 0,4, 0.6 and 0.8 of telson length, posterior margin. wilh 3 pirs of spines, lateral spines small, subequal to dorsal spines, interme-

A NEW GENUS OF HIPPOLYTID SHRIMP 15

wr

JNQA E

m

Si Kaes

)

FIG. |. Thorina maldivensis (Borradaile). d. Aldabra Island (upper) and ovigerous ?, Heron Island (lower).

Scale in mm.

diate spines c. 3.0 times as long as lateral spines, submedian spines slender, plumose, twice length of lateral spines.

Antennule with proximal segment broad, about as wide as long, with stout acute tooth at 0,6 of ventromedial margin, stylocerite elongate, acute, exceeding length of peduncle, with small an- teroverted process proximodorsally; statocyst ob- solete; intermediate segment short, broad, with acute non-articulated lateral process; distal seg-

ment short, broad, with articulated triangular plate dorsolaterally: upper flagellum with proxi- mal 12-13 segments stout with dense tufts of aesthetascs dorsolaterally, distal flagellum with c. 5slender segments; lower flagellum slender, with c. 20 segments.

Antenna with basicerite stout, with broad acute, ventrolateral tooth; carpocerite stout, sub- cylindrical, extending to about end of antennular peduncle, flagellum short, filiform c. 3.5 times

16 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2. Thorina maldivensis (Borradaile). Ovigerous 9, Heron Island. A, anterior carapace and appendages, lateral; B, same, dorsal; C, rostrum and anterior carapace, lateral; D, antennule, dorsal; E, same, peduncle, ventral; F, antenna; G, eye, dorsal; H, 2nd pereiopod; I, 3rd pereiopod; J, uropod; K, telson; L, same, posterior spines, dorsal spine (inset).

A NEW GENUS OF HIPPOLYTID SHRIMP 17

M erok — Ye

be f

C

In m

FIG. 3. Thorina maldivensis (Borradaile). Ovigerous Ẹ , Heron Island. A, mandible; B, same, molar process; C, same, incisor process; D, maxillula; E, same, palp; F, maxilla; G, same, palp; H, Ist maxilliped, median process of anterior sternite stippled; I, 2nd maxilliped; J, 3rd maxilliped; K, same, terminal spines of distal segment of

endopod.

18 MEMOIRS OF THE QUEENSLAND MUSEUM

carapace length, Seaphocerite exleniing well he- vend stout part of upper antennular flagellum, c. 2,5 times longer than maximal width, situated at c. 0,3 of length, tapering distally to rounded distal lamella, distinctly exceeding tip of stout dis- tolateral tooth, situated at c. 0.8 of straight lateral margin length.

Eye well developed, with large well pigmented globular cornea, diameter c. 0.33 of carapace length in måle, 0.4 in 9, with conspicuous dorsal ocellus; stalk stout, compressed.

Mouthparts generally similar to those of Thor species. Large acute compressed transverse wian- gular median plate, anterior to Ist thoracic sler- mie, occluding space between coxal endites of first maxilliped, Mandible without palp; molar process stout, obliquely truncate distally, with marginal setae and small denticles; incisor pro- cess slender, distally oblique, with f small acute teeth. Maxillula with short, feebly bilobed palp, upper lube with slender simple seta, lower lobe with stouter spiniform seta; upper lacinia larger, oval, with numerous short spines distoventrally. lower lacinia small, short, with several long spines distally. Maxilla with slender, tapering palp, with single spiniform terminal seta, short preterminal dorsal seta and medial sela; distal endie well developed, deeply bilohed, densely sciase medially; proximal endite feebly devel- oped, slightly bilobed, with few long setae, Scaphognathite well developed. c. 3.0 times longer Than central width. posterior lobe small, anterior lobe large, with median margin slightly voneave. First maxilliped with 2-segmented palp, distal segment c, 2.5 times longer than wide, sparsely setose medially, proximal segment broader than distal, medial border convex, sparsely selose; basal endite broad, angular, me- thal margin with dense short setae; coxal endite convex, medial Margin with sparse. coarse, long plumose setae; exopod with well developed fa- gellum. with numerous plumose setae distally, caridean lobe small, with numerous plumose marginal setae; epipod large, triangular. feebly tnlobed. Second maxilliped with dactylar seg- ment short, narrow, with numerous short stout spines, propodal segment large, broad, distomed- ial horder with numerous long spines; carus and ischiomerus normal; basis with medial margin excavate, dorsal and ventral medial borders with numerous long slender setae, exopod normal, coxa medially produced, laterally with small elongate epipod, bearing small podobranch ante- riorly. Third maxilliped robust, exceeding car- pocerite by about terminal and half penultimate

segment in d and terminal segment only in 2. In ?, terminal segment c. 73 times longer than central width, subcylindrical, slightly flattened ventrally with numerous transverse rows of short spines, distally obliquely truncate, with 6 stoul spines, penultimale segment c, Då of Terminal segment length, twice as long as wide, with groups ol short spiniform setae medially, long setae laterally; ischiomeral segment fused with basis, combined segment subequal to terminal segment length, moderately bowed, compressed proximally, expanded distally, lateral margin with c, 9 spiniform setae, distolateral angle with single long Straight spine with short acute tooth medially; hasis short. e. 0.2 of antepenultimate segment length, medial margin slightly bilobed, lateral border with small robust rounded epipod; without arthrobranch. 3 third maxilliped similar ta 9, exceeding carpocerite by penultimate and terminal segments,

First perciopods showing marked sexual di- morphism, small in 9 2, greatly hypertrophied in some å 3, In 9 9, exceeding basicerite by about carpus and chela; chela c. 0.5 of carapace length, with palm subcylindrical, slightly swollen proxi- mally, with few short pectinate cleaning setae proxrmoventrally, feebly compressed distally, c. 2.4 times longer than proximal depth, smooth, fingers about 1/2 palm length, stout, tapering distally, with sharp medial culting edges, dactyl with 4 stour non-cornified terminal leeth, fixed finger with 3, central tooth enlarged with laminar lateral expansion; carpus c. 0.4 of chela length, 2.5 limes longer than distal width, tapered proxi- mally, smooth, unarmed, with several cleaning setae distoventrally; merus c, 0,95 of chela length, widest at midlength, 2.8 times longer than central width, with acute ventrolateral tooth dis- lally, proximal ventral margin with single small spiniferous tubercle; ischium c. 0.55 of chela length, compressed, 2.2 times longer than wide, slightly broader distally than proximally, with single small preterminal distoventral spinule; busis and coxa without special features; without exopod. Hypertrophic male chelae slightly un- equal, similar, c. 0.25 of carapace length, with palm subevlindrical, without cleaning setae pro- ximoventrally, subuniform, €. 5.5 tines lunger than central depth, ventromedially tuberculate, fingers c. 0.3 of palm length, stout, deflexed, with numerous long stilt setae, tapering distally, with sharp strongly concave culling edges. with stout hooked Lips crossing distally, dactylus with single large acute tooth proximally, fixed finger with smaller acute tooth more distally; carpus c. 0.5 of

A NEW GENUS OF HIPPOLYTID SHRIMP 19

X] N i ` 4

LA mm

|

An

FIG. 4. Thorina maldivensis (Borradaile). First pereiopods. A, ovigerous 9, Heron Island, CL 2.2 mm; B, d, Heron Island, CL 2.3mm; C, å, Aldabra Island, CL 2.2mm; D, d, Astove Island, CL 2.4mm.

chela length, 3.3 times longer than distal width, tapered proximally, feebly tuberculate ventrome- dially, unarmed, without cleaning setae dis- loventrally; merus c. 0.9 of chela length, subcylindrical, 6.0 times longer than central width, ventrolateral surface tuberculate, with nu- merous small acute distally directed tubercles, with several (3-6) large stout acule preterminal teeth distolaterally, distoventral angle rounded; ischium obliquely articulated with merus, c. 0.27 of chela length, compressed, distally expanded, 2.0 times longer than greatest width, dorsally carinate proximally with c. 8-9 denticles or tuber- cles and long stiff erect spiniform setae; basis and coxa robust, without special features; without exopod. Intermediate males with similar but smaller and less robust chelae, less spinose and tuberculate, more closely resembling 9 chelae. Second pereiopods slender, exceeding car- pocerite by carpus and chela in 9, by chela and 3 distal segments of carpus in d. chela with palm

subcylindrical, c. 2.3 times longer that central width, with few setal tufts distally, fingers c. 0.6 of palm length, slender tapering, 3.0 times longer than proximal depth, cutting edges sharp, medial, tips with 3 acute spines on dactyl, 2 spines and short tooth on fixed finger; carpus 2.5 times chela length, 12 times longer than wide, 6-segmented, segments in ratio of 2: 1: 3,4: 1.9: 1.1: 1.6, first 2 segments poorly separated, distal segment with transverse row of long serrulate setae distoventr- ally. merus c. 1.75 times chela length, 6.3 times longer than central width, 6.0 times longer than wide, simple; ischium 0.9 of merus length, 5.3 times longer than wide, with 2 long simple spini- form setae proximoventrally; basis and coxa without special features.

Ambulatory pereiopods moderately robust, third pereiopods with slight sexual dimorphism. 9 third pereiopod exceeding carpocerite by pro- pod and dactyl; dactyl c. 0.27 of propod length, 2.6 times longer than proximal depth, stout, com-

20 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. Thorina maldivensis (Borradaile). Ovigerous 9 , Heron Island. A, first pereiopod chela and distal carpus; B, fingers of chela, lateral; C, same, medial; D, same, tip of fixed finger; E, same, medial ischiomeral spines; F, 2nd periopod, chela; G, same, fingers; H, same, distal fingers, dactyl above; I, same, medial ischial spine; J, 2nd pereiopod, chela. K, third pereiopod, propod and dactyl. d. Heron Island. M, 3rd pereiopod, propod and dactyl. N, major 2nd pereiopod, fingers. O, minor 2nd pereiopod, same.

pressed, distal end c. 0.6 of proximal depth, with ` propod c. 0.8 of carapace length, 7.5 times longer smaller dorsal ungual spine and larger, stouter than wide, uniform, slightly bowed, ventral bor- distoventral spine, each c. 0.3 of length of dorsal der with 2 larger distoventral spines, 5 ventral border of corpus, ventral margin with two spines; spines, decreasing in size proximally, dorsal mar-

A NEW GENUS OF HIPPOLYTID SHRIMP 21

FIG. 6. Thorina maldivensis (Borradaile). Ovigerous 9 , Heron Island. A, 3rd pereiopod, dactyl. d. CL 2,3mm, Heron Island. B, 3rd pereiopod, dactyl. C, Ist pleopod. D, same, endopod. E, 2nd pleopod. F, 3rd pleopod, endopod. d, CL 2.4mm, Aldabra Island. G, 3rd pereiopod, dactyl. H, Ist pleopod. 1, 2nd pleopod.

gin sparsely setose; carpus c. 0.36 of propod length, unarmed; merus robust, c. 0.72 of propod length, 3.2 times longer than central width, with articulated distolateral spine; ischium c. 0.68 of meral length, 2.5 times longer than distal width, narrower proximally, unarmed; basis and coxa robust, without special features; without exopod. 4th and 5th pereiopods similar to 3rd. propods slightly longer than 3rd pereiopod propod length, meri shorter and more slender, 4th c. 0.95 and 5th

0.85 of 3rd merus length, Male 3rd pereiopod exceeding carpocerite by propod and dactyl, with slight sexual dimorphism; propod slightly ex- panded distally in smaller specimen, with 3 pairs of distoventral spines, dactylus as in 2, with 2 ventral spines; in larger d, propod scarcely ex- panded distally, with 2 groups of 4 and 5 dis- toventral spines, dactylus with 3 additional spines on proximal ventral margin.

22 MEMOIRS OF THE QUEENSLAND MUSEUM

Ovigerous 9 pleopods without special fea- tures, basipodite enlarged and expanded, rami short, broadly expanded, 2nd to 5th endopods with appendix interna. Male Ist pleopod with basipodite robust, broad, ventromedially con- cave, coxopodite with uncinate distoventral pro- cess; endopod sublanceolate, c. 0.8 of exopod length, 1.6 times longer than proximal width, medial margin straight, setose, lateral margin convex, with short plumose setae; exopod c. 3.3 times longer than wide, with plumose marginal setae; 2nd pleopod larger, endopod c. 0.8 of exo- pod length, with plumose marginal setae, lacking appendix masculina and appendix interna, exo- pod normal; third pleopod normal, with appendix interna at c. 0.4 of medial margin length.

Uropod with protopodite with large acute lat- eral tooth; exopod c. 2.5 times longer than wide, lateral margin straight, sparsely setose, with small acute distal tooth, with large mobile spine medi- ally; endopod 0.95 of exopod length, 3.0 times longer than wide.

TYPES. The type material is deposited in the collection of the Zoology Museum, Cambridge, United Kingdom. 3 lots of specimens are held (pers. comm., R.C. Preece, 16 August 1995), consisting of the following: (i) 1 specimen in reasonable condition, from Minikoi, Laccadive Islands; (ii) 1 specimen in reasonable condition, from Salomon Island. (iii) 2 specimens, with loose appendages, from Holulé Island, Malé, Maldive Islands. 2 lots of specimens, (i) and (iii), are noted as co-types. The specimens from Minikoi and Holulé, are noted on their record cards with "Gardiner Colln. Ann. Mag. Nat. Hist. (8) XX, p. 208. Percy Sladen Trust. Exp. XXII pt 3. p 400'. The Salomon Island specimen is anno- tated only “Gardiner Colln.* All have the acces- sion number AR 3.1920.

MEASUREMENTS (mm). å, Astove Island: postorbital carapace length, 2.3; carapace and rostrum, 3.3; total body length, 12.5; major first pereiopod chela, 5.5; minor first pereiopod chela, 4.8; 3rd pereiopod propod, 1.8; same, merus, 2.3. Ovigerous 9 , Heron Island: postorbital carapace length, 2.2; carapace and rostrum 3.1; total body length, 11.8; Let pereiopod chela, 1.2, 3rd perei- opod propod, 1.7; same, merus, 1.9.

COLOURATION. The Jadini, Kenya, specimen was noted to be an opaque white, except for the transparent posterior 4th, 5th and 6th abdominal segments and caudal fan; pereiopod bases also white, meri banded with white; antennule and

antenna white, filiform flagella transparent. The Mitiaro specimens were noted by the collector as "translucent, with pink and white mottled head and legs, green tinge to lower abdomen and tail and ‘translucent, with pink predominating, mot- tled white head and legs', the Tutuila specimens as 'transparent, with green tinge on lower abdo- men, mottled green- pink head and legs', and the Tongatapu specimens as 'transparent, with pink tinge and pink mouthparts, red/white eggs, dark body organs'. A good colour photograph is pro- vided by Kamizake et al. (1988). This shows a semi-translucent pale yellowish shrimp, heavily mottled with fine red speckling and scattered whitish patches, particularly at bases of pleopods and base and tips of caudal fan.

REMARKS. The exact habitat of Thorina maldivensis has not been established but most personally collected specimens have been ob- tained from shallow intertidal coral reef pools with a wide variety of corals and other coelenter- ates and algae. As many were collected by the use of poison, the precise niches occupied were not observed. Several specimens were collected from coral heads but these associations may have been accidental. Hayashi (1986) records this species from under coral blocks. The species appears to be generally uncommon, but Edmondson (1946) reported that in Hawaiian waters the species ‘is common among brown seaweeds near the shore'. The colouration of the Jadini specimen does not appear cryptic and suggests a commensal life- style. Where several specimens were collected together the association is more likely to be genu- ine. The Papua New Guinea specimens from 10- 20m represent the maximum bathymetric range for this species.

The d specimen from Cartier Reef had only un-hypertrophied first pereiopods but the distal propod and dactylus of the 3rd pereiopods were distinctly sub-prehensile, the ventral dactylus with accessory spines. The d from Tutuila pos- sessed only part of a single first pereiopod, lack- ing the chela. The merus was rather feebly spinulate, with blunt spines, and the spiniform setae along the dorsal margin of the carpus were feebly developed, mostly short and slender, with only a single distal long spiniform seta.

The transverse triangular median sternal pro- cess that appears to lie anteriorly to the sternite of the first maxilliped appears unusual but it is rather difficult to discern its exact relationships. No similar feature seems to have been reported in other hippolytids but a similar, though much

A NEW GENUS OF HIPPOLYTID SHRIMP 23

smaller and narrower process, more antero-pos- teriorly orientated, is also present in Thor amboinensis (De Man).

DISTRIBUTION. Throughout most of the Indo- West Pacific region. Type localities: Malé Atoll, Maldive Islands; Minikoi, Laccadive Islands, and Salomon Island. Also known from Kenya, Tan- ganyika*, Mauritius*, Seychelle Islands*, Mald- ive Islands, Andaman Islands, Cartier Reef*, Great Barrier Reef*, Papua New Guinea*, Ryukyu Islands, Marianas Islands, Marshall Is- lands, Tonga*, Cook Islands*, Kiribati, Society Islands*, and Hawaiian Islands. (* — new localities). Tanganyika is used in its zoogeographical mean- ing and not in a political sense.

LITERATURE CITED

BORRADAILE, L. A. 1915. Notes on Carides, Annals and Magazine of Natural History, (8)15: 205-213. 1917, On Carides from the western Indian Ocean. Transactions of the Linnean Society of London, Zoology, (2)17: 397-412,

BRUCE, A.J. 1976. A Report on a small Collection of Shrimps from the Kenya National Marine Parks at Malindi, with Remarks on selected Species. Zoologische Verhandlingen, Leiden, 154; 1-72.

CHACE, E.A. 1972. The Shrimps of the Smithsonian- Bredin Caribbean Expeditions with a Summary of ihe West Indian Shallow-water Species (Crustacea: Decapoda: Natantia). Smithsonian Contributions to Zoology, 98: i-x, 1-179.

In press. The Caridean Shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedi- tion 1907-1910, Part 7: Atyidae, Eugonat- onotidae, Rhynchocinetidae, Bathypalaemonidae, Processidae and Hippolytidae. Smithsonian Con- tributions to Zoology.

EDMONDSON, C.H. 1925. Crustacea of Central Trop- ical Pacific. Marine Zoology of Tropical Central Pacific. (Tanager Expedition Publication No. 1, 1915), Bulletin of the Bernice P. Bishop Museum, Honolulu, 27: 3-62.

1946. Reef and Shore Fauna of Hawaii. Special Publication, Bernice P. Bishop Museum, Hono- lulu, 22: i-iii, 1-381.

HAYASHI, K. 1986. An Annotated List of Shrimps (Alpheidae and Palaemonidae Excluded) Col- lected from the Gilbert and Solomon Islands. Pro- ceedings of the Japanese Society of Systematic Zoology, 32: 17-29,

HOLTHUIS, LB 1953. Enumeration of the Decapod and Stomatopod Crustacea from Pacific Coral Islands. Atoll Research Bulletin, 24: 1-66.

KEMP, S. 1916. Notes on Crustacea Decapoda in the Indian Museum, VII. Further Notes on Hippolytidae. Records of the Indian Museum, 12: 385-405.

KINGSLEY, J.S. 1878. Notes on North American Car- idea in the Museum of the Peabody Academy of Sciences at Salem, Massachusetts. Proceedings of the Academy of Sciences of Philadelphia, 1878: 89-98.

KAMEZAKI, N., NOMURA, K., HAMANO, T. & MISAKI, H 1988, A Guide Book of Marine Ani- mals and Plants of Okinawa, Crustacea (Macrura and Anomura), 8: 1-232, col. pls. Shinsei Guide Book Series. (Southern Press: Okinawa, Japan).

74 MEMOIRS OF THE QUEENSLAND MUSEUM

NEW DATA ON LERISTA INGRAMI, A RARE SKINK FROM SOUTHERN CAPE YORK PENINSULA, AUS- TRALIA. Memoirs of the Queensland Museum 42(1): 24 (997:- Lerista ingrami Storr, 1991 is poorly known. It & treated as ‘rare’ (McDonald etal., 1991) arid was kriown, prior to Sept., 1996, from 10 specimens (the holotype and nine

ralypes) from what amounts to a single locality — the oredune of the beach hetween the fondi of the Mclvor R, und Cape Flattery. on southern Cape York Peninsula. L ingrami is one of å few species of Australian skinks (4/320 species) not photographed in life. The type locality of this species has been visited recently (13 Sept, 1996), As s result, we can clarify some minor discrepancies with collection daia af the type specimens; present phorographs of a live specimen of L ingrami and of iis type locality: describe the colour of L ingrami m lifes and comment on prey of L. ingrii.

Published data on the type locality and the collection tocality of the pararypes of L. ingrami is at Variance shghtly with 1nformation recorded in the Queensland Museum regis- ter. Storr (1991) records:

"Holotype 132396 in Queensland Museum collected on 27 July 1976 by G.J. Ingram near beach north of Melver River, Queensland, in 15°07'S, 1457] 5E",

'Paratypes Queensland: 7km N of mouth of Melver River (QM20644-5 1) and 5km N (QM20653)".

The QM register entries for these specimens are: 9QMI32396 (holotype) op mission road near beach, north of Melvor R., 13 July, 1976, G.J. Ingram’, (Date of registration and entry 27 July, 1976). ^QMJ20644-51 (paratypes) approx. din N of Melvar R. mouth Cooktown 1:250000 328089, 20 Nov., 1970, JAC, T Pi Teb- ble, C. Tanner; ‘buried in sand under log on first dune’. QMJ20653 (paratype) approx. Am. N of Melvor R. mouth. Cooktown 1:250000 318090, 20 Nov., 1970, JAC, T.P. Teb- lle, C. Tanner: "buried in sand of first dune - underlogs'. (Date of registration and entry 6 Jan., 1971).

For the last-mentioned, sometime in or about 1986, lati- tude/longifude were calculated from an atlas and added to the register in pencil, 15905'5 145?]4" E.

Following advice from Gå, Ingram (collector of the holo- type of L. ingrami) and using å ‘Magellan Global Positioning System (GPS), we have calculated the type locality for L. imgrami( Fig. D) at L57070]"S 145*14'42"E, very close to the locality calculated for/by Storr (but nor entered in the QM register) at the Ome of his description of the species - 15°07 145915".

We collected topotypic specimens QMJ62430-1, QMJ62443, One has heen photographed (Fig. 2). All were found in the middle of the day, ‘at rest’, on slightly moist sand under dead coconuts, Once disturbed, they invariably sought escape into the sand, Both mid body (20 x3) and supraciliary

FIG. 1 The type locality of Lerissa grani Starr, 1991. (Photograph, JAC)

= NE d

FIG. 2 Leriste ingrami. (Photograph, Jell Wright, OM),

counts (3x2, 4x1) for these specimens fall within the range given for the species by Starr ( 1991).

L. ingrami was described from old (15-21 years) spirit Material, so its colour in life was not known. Storr's descrip- non (1991) can be augmented: body upper surface a shiny pink or grey-beige, profusely marked by tiny dark brown dors; head scales dark-edged, Å sharp-edged dark brown lateral stripe runs from the rostral to the base of the tail, tapering posteri- orly. Upper surfaces of limbs the same colour as dorsum, also with dark brown (minute) dots, Venter (SV) pale grey-white, Tail pale to very bright orange ventrally and dorsally,

There is little information on the prey of Lerista spp. Wilson & Knowles (1988) report `-. Small arthropods, their eggs und larvae probably constitute the bulk of their diets ..." L. bipes and L. muelleri are the only species from sandy localities for which prey have been reported, and there 15 only one other reference ta prey of Lerista (a Tasmanian record of Hewer & Mollison, 1974). Smith (1976) records orthopterans from L bipes, and proturans and collembolans from 4. muelleri. Faecal pellets from the recently-collected L ingrami have been examined by Dr G, Monteith, Senior Curator (Ento- mology) atthe Queensland Museum. They contain remains of two smäll scorpions (Arachnida: Scorpionida) and two small seed bugs (Insecta! Lygaeidae). Both, he advises, are common and active amongst leaf litter,

Tissue samples were taken from QMJ62430 und QM16243 1 and sent to the tissue library of the South Austra- lian Museum, Adelaide. Only minor external morphological characters (e.g., mid-body scales, lamellae ot the third (oe, supracilary scales and colour) separate L. ingrami from L. orientalis (De Vis, 1889) and L. senulata Storr, 1991,

Literature Cited

Hewer, A. & Mollinson, B.C. 1974. Reptiles and amphibians of Tasmania, Tasmanian Yearbook No.8 (1974): 50-60),

McDonald, K.R Covacevich, J,A., Ingram, G.J. & Couper, PJ. 199]. The status of frogs and reptiles. Pp. 338-345, In Ingram. G.J, & Raven, R.J. (eds) An atlas of Queensland's frogs, reptiles, birds and mammals. (Board of Trustees, Queensland Museum, Brisbane),

Smith, L.A, 1976, The reptiles of Barrow Island. Western Australian Naturalist 13(6): 125-36,

Storr, G.M, 1991. Revision of Lerista orientalis (Lacertilia: Srincidae )ofnorthem Australia Records of the Western Australian Museum 15(2); 413-417

Wilson, S.K. & Knowles, D.G. 1988, Ausiralia's reptiles, A photographic reference to rhe terrestrial repriles of Aus- tralia. (William Collins Pty Lid: Sydney).

LA, Covacevielt, PJ Couper & L Roberts, Queensland Mu-

seum, PO Box 3300, South Brisbane, Queensland 4101, Aus-

traller 3 April 1997

TWO NEW SPECIES OF FRESHWATER ATYID SHRIMPS (CRUSTACEA: DECAPODA: ATYIDAE) FROM NORTHERN QUEENSLAND AND THE DISTRIBUTIONAL ECOLOGY

OF THE CARIDINA TYPUS SPECIES-GROUP IN AUSTRALIA SATISH CHOY AND JONATHAN MARSHALL

Choy, 8. & Marshall, J. 1997 06 30: Two new species of freshwater atyid shrimps (Crustacea:

Decapoda. Atyidae) from northern Queensland and the distributional ecology of the Caridina

topus [o ed in australia, Memolrs of rhe Queensland Muxeun AY I): 25-36. Brisbane. ISSN 79-8835.

Two new species of freshwater atyid shrimps are described froni tropical Queensland and the four members of the Caridina typus-group now known from Australia are reviewed. Caridina confusa sp, nov. is a slender animal with a relatively long, dorsoventrally flattened, naked rostrum, found predominantly in open grassland streams of the Atherton Tableland. Caridina spinula sp. nov., distinguished by its spiniform pterygostomian angle. is found in small secondary rainforest streams on northeastern Cape York Peninsula. Although both these new species look superficially like C. zebra Short. 1993, they can be distinguished by à Combination of characteristics such as rostrum length, shape of the prerygostotniau angle, Jength-depth ratio of the sixth abdominal segment and the shape of the protopod of the uropod. C. zebra is found predominantly in primary rainforest streams of thé Atherton Tableland and C. rypus is found in coastal tropical streams. All species are allopatric, except for slight overlap in the distribution of C. zebra and C. confusa in some tnthroncipenically disturbed streams of the upper Barron and the upper North Johnstone catchments. [| Crustacea, Atyidae, Caridina, Queensland, distriburion,

S, Choy, Resource Sciences Centre, Department of Natural Resources, 1345 Ipswich Road, Rocklea, Queensland 4106, Australia; J. Marshall, Faculty of Environmental Sciences,

Griffith University, Nathan, Queensland 4111, Australia: 11 December 1996.

Caridina zebra Short, 1993 is a tropical mon- tane species belonging to the Caridina rypus spe- cies-group (Short, 1993), This group is characterised by a short, dorsally unarmed ros- trum, the presence of epipods on the 1st four pairs of pereiopods and the presence ol an appendix interna on the endopod of the Ist pleopod of d d. Short (1993) reported C. zebra from the monrane streams of [he wet tropical rainforest areas of the Tully, Herbert and Johnstone River catchments, at altitudes of 400-900m. A smaller population of C. zebra, is also known 1o occur in the lower Koolmoon Creek (alt. 150m). à tributary of the Tully River (Hughes et al, 1996). Caridina typus Milne Edwards, 1837 has been reported from coastal lowland streams at Cooktown and on Dunk Island (Roux, 1926; Riek, 1953; Short, 1993),

Re-examination of Short’s material from the Wet Tropics-Atherton Tableland area (Short, 1993: 62) indicated the presence of two distinct morphological groups, one group possessing a longer rostrum, from anthropogenically dis- turbed grassland areas and the other possessing a shorter rostrum, generally from relatively undis- turbed rainforest areas, This was confirmed by examination of recently collected material trom

the same area (17-I8^S, 145-146°E). Although the distribution of these two groups tend to over- lap slightly, they are morphologically and ecolog- ically distinct. They each warrant species status. Since the short rostrum form is the holotype of C. zebra Short, 1993, its status is maintained. The long rostrum form is described as a new species, Caridina confusa.

A third morphological and geographically iso- lated group was recently collected from the streams in the Mcflwraith Range, Cape York Pen- insula (13°35-44°S, 143?20' E). This is also de- scribed as a new species, Caridina spinula,

MATERIAL AND METHODS

Specimens from the Queensland Museum are provided with catalogue numbers with the prefix OM. Unless otherwise stated all material was collected by the first author and various col- leagues from Griffith University and the Queens- land Department of Natural Resources (formerly part Department of Primary Industries). All ex- amined specimens will be deposited in the Queensland Museum. Collection was made using a standard pond net and all samples immediately preserved. The abbreviations used are: 68. sixth abdominal segment length; A1P, antennular pe-

26 MEMOIRS OF THE QUEENSLAND MUSEUM

dunele length; AZP, antennal peduncle length; CL, post-urbital carapace length; RL, rostrum length; SC, scaphocerite length; SL, body length from the post-orbital margin of the carapace to the tip of the telson; ST, stylocente length; T, telson length.

The format of the description and morphologi- cal terms follow Choy (1991). Although prob- lems in the terminology of cuticle spination and setanon still exist, we have followed the termi- nology of Felgenhauer (1992),

SYSTEMATICS

Caridina typus Milne Edwards, | 837

Cundinu typus Milne Edwards, 1837: p, 263, pl. 25615, figs. 4, 5; Holthuis, 1965, p, 10, fig. 3.

Candina typus typica Bouvier, 1925, p.250, figs. 272- 295

Caridina typa Roux, 1926, p. 201; Rick, 1953, p. 117.

MATERIAL EXAMINED, Proserpine River below Proserpine (20?24,2'S, 148 31.1 E), 19.10.94, LR. Paterson, 15 d d 12.54. 111m CL), 7 9 9 (4.8-5-1mm CL): QM W4795, Brandy Creek near Proserpine (207208, 148"38 E), 214.75. R Monroe; P. File- wood, ovigerous v (7.81mm CL), non-ovigerous 9 (5.82mm CL); QM W 14241, Lindeman 13., west side, small creek near goll course (20278, 149?02' E), 27,3,83, 1, Short, 33 (4.03-4.09mm CL), 9 9 15.5- 7.2mm CL).

DIAGNOSIS. Rostrum short, laterally cam- pressed, with 1-5 teeth of ventral margin; diaer- esis with more than 20 hamate setae; intermediate setae on posterior telsoni¢ margin longer than lateral pair, plumose, with sclerotinous plug; egg small (length «0.52mm).

REMARKS. Caridina typus has å wide ranging distribution, extending from eastern and southern Africa through the Indian Ocean islands, South- cast Asia jo Japan, Australia and through to French Polynesia, In Australia it has been col-

tected only from the northeastern coastal areas of

the mainland and from the nearby islands, be- tween latitudes 15-2175. Although it seems to be confined to the lower reaches of coastal and is- land Australian streams, C. typus has been col- fected trom altitudes of 300m elsewhere (Choy, 1991).

Caridina zebra Short. 1993 Curidina zebra Short, 1993. p. 62 (in part],

MATERIAL EXAMINED. All SNE listed hy Short. 1995, p. 62. ADDITIONAL MATERIAL.

Upper Tully Catchment: QMW1I7118, Tully River above Koombooloomba Dam (17°49"S, 145?35' E. 720m), 23.10,91, numerous specimens: Koombooloomba Creek (17?51.5'S, 145°35.9'E, 790m), 30.10.93, 25 specimens; Echo Creek (17°59,5'S, 145?38,3 E, 830m), 30.10.93, 50 speci- mens; Carpentar Creek (17°53.3°S, 145?35 Y'E, 750m), 30.10.93, 23 specimens; Costigan Creek (17°56°S, 145"37'E, 770m), 3,12.94, 11 specimens (1.8-4,2mm CL); QMW17119, Koolmoon Creek (179445, 145734" E), 25.7.90, 4 specimens; Koolmoon Creek nt Walter's Waterhole (17°44,11°S, 145*34' E. 760m), 30,10.93, 7 specimens; Koolmoon Creek near Tully River confluence (17^44.9'S, 145?37.1'E, 150m), 31.10.93, 6 specimens, Upper Herbert Catch- ment: QMW17116, Blunder Creek (17736'S, 145?33'E), 28.11.90, 3 specimens; Blunder Creek (17947.5'8, 145*32.2'E, 750m), 3.12.94, 1 4, 2 ovigerous NN (4,08-5.6mm CL); Rocky Creek (1744,7'S, 145931,3'E, 760m), 30.10,93, 34 spéci- mens; Cameroon Creek, 3.12.94, 14 specimens (1.4- 4.2mm CL);millstream River al Diversion Weir (V7?40'S, 145*26' E, 720m), 12.10.94; 2 specimens; Upper North Johnstone Catchment: QMW19261, Malanda Falls, (17°20.2°S, 145?43,8'E, 750m), 211,93, 30 specimens (including ovigerous ? 2 1; QMW 19285, Raspberry Creek, 3-4 km above Malanda Falls (17°22.75'S, 145E33.6 E), 14.11.91, J. Short, P. Davie, A. Humpherys, silt & para. grass (Urochloa mutica) infested stream; Raspberry Creek (17?25,5' S, 145728, 8&' E), 31.10.93, 100 specimens; North Johnstone River (17?40' S, 145739" E, 650m), 5.8.93, 2 9 9. I g; Whaca Creek (177248, 145°38'E, 650m), 7.11.94, 2 9 9; Ithaca Creek at Bauld Rd (17°23'S, 145*38'E, 685m), 13.10.94, 50 specimens; Thiaki Creek at edge of rainforest (17725'S, 145?32' E, 795m), 5.12.94, 9 specimens (3.4-4, 8mm CL); Thiaki Creek in rainforest, 13.10.94, 15 specimens; North Beatrice River at Palmerston Highway (17°32'S, 145*36'F, 720m), 13,10.94, 25 specimens; Henrieta Creek at Palmerston Highway (17?36'S, 145°45°E, 360m), 14.10.94, 18 specimens; QMW18722, Goolugan Creek, 23,10,91, numerous specimens; Goolagan Creek (17936.3'8, 145°45.5'E, 370m), 32 specimens; Upper Barron Catchment: QMW3078, Atherton lap water (179168, 145?29'Ej, 14.5.62, QDPI, 2 d d; Barron River at The Crater (199175, 145?29 E), 7.12.94, 10d G, 8 ? 9 (Zovigerous) (2.0-5.0mm CL); Barron River at Hemmings Road, 28.11.94, 3 juve- niles; Wright's Creek (179048, 145?45'E), 4.12.94, 4 ?9 LI ovigerous); Peterson Creek (17"17'S, 145"36 E), 1994-95, numerous specimens; Kauri Creek (169545, 145738 B), 1994-95, numerous spec- imens; Prior's Creek, I 9; Upper Gwynne Creek in small pocket of rainforest (17^23.3' 8, 145%30.3'E), 5.12.94,2 F 9, 19 d 8 (3.6-4.0mm CL).

DIAGNOSIS. Body (Fig. 1b) rotund, may have black and white transverse banding; rostrum (Fig. 2a-f} short, extending to base of 3rd antennular peduncle (RL«0,5CL), dorsoventrally com-

NEW FRESHWATER SHRIMPS 27

FIG, |. a, Caridina confusa sp. nov,, 3; b, C. zebra Short 1993, d c, C. spinula sp. nova d. Scale I mm.

pressed, may have an apical tooth (Fig. 2e); pterygostomian angle acute but not spiniform; dorsal telsonic spines (Fig. 3a) on posterior 0.66 of telson; posterior telsonic margin (Fig. 3f, g) usually with median spine, sub-lateral pair of setae sigmoid, longer than intermediate setae, selalion numerically variable; protopod of uro- pod (Fig. 3a, d) elongate, spinate; eggs large (length «1.32mm); found mainly in primary rainforest streams on the Atherton Tableland along the Lamb-Francis-Card well Ranges.

REMARKS, Specimens of C. zebra from some locations (streams in the upper Barron Catch- ment) may seem to have a fairly long rostrum (Fig. 2a, d, f), thus resembling C. confusa sp. nov. However, the rostrum length relative to the cara- pace length as well as other features (Table 2), such as the relative lengths of the antennular peduncle, scaphocerite, sixth abdominal segment and the telson are characteristic of this species. Caridina zebra is one of only two species of atyid shrimps in the primary rainforest streams of the upper Tully (the other being Australatya striolata), where it is very abundant. In other catchments (Barron, N Johnstone and Herbert),

it is found mainly in the rainforest reaches of streams. However, it may be common in some disturbed streams such as Raspberry, Ithaca und Prior's Creeks. Unlike the upper Tully, which has an extensive relatively undisturbed forested area, the upper reaches of the Barron and N. Johnstone Rivers have been generally cleared and converted to pasture. Streams running through these open prassland are inhabited predominantly by C. con- fusa sp. nov. Only a small area of the upper Herbert catchment is in the wet (simple notophyll vine to complex mesophyll vine) forest; the rest is in dry sclerophyll forest. The abundance ol C. zebra in these dry-zone streams is low; these streams are inhabited by another atyid, Paratya australiensis. Ti is interesting Io note that only specimens from some of the rainforest streams of the upper Tully have the black and white trans- verse banding on their bodies (see Short, 1993), In all other areas these animals are translucent brown, with scattered tiny reddish and bluish- green chromatophores, similar to individuals of the IWO new species.

Caridina confusa sp. nov.

Caridina zebra Short, 1993, p. $2 (in part): QMW18841, QMW18720.

MATERIAL EXAMINED. HOLOTYPE. QMW- 21906 ovigerous H. 4.8mm CL, 2.56mm RL, 19.2mm SL, Thiaki Creek al Seamark Road crossing (17°23.5'S, 145°32.5°E, 750m), stream flowing through open grazing land, fringing para grass, water depth 0.3- 0.5m, velocity Q.3ms `, silty substrate, hand-netted amongst edge para grass, 26.8.94, S. Choy, M. Hopper. ALLOTYPE, QMW21907 adult c, 3.4mm CL. 2.0mm RL, 14.2mm SL, same locality data as holotype, PARATYPES, Upper North Johnstone Catchment: QMW21908 upper North Johnstone River near Brom- field Swamp (17722.5'S, 145°31.3'B, 700m). open grazing land, fringing para grass, water depth 0,3-0,5m, hand-netled, 4.12.94, 21 3 d, 50 9 ' (13 ovigerous), (2.6-5.0mm CL); QMWI18841], Thiaki Creek (17°24.9'S, 145?35.3'E, 750m), water depth 0.2m, electro-fished, 1992, Queensland DPI Fisheries Johnstone Rivers Survey, I d (4.5mm CL); QMW18720, small creek about 6 km SW of Malanda (17722. S, 145°33.6°E, 750m), fringing para grass, waler depth 0.2-1.5m, electro-fished, 1992, Queens- land DPI Fisheries Johnstone Rivers Survey, å (3.9mm CL), 2 9 9 (4.3, 4.5mm CL); QMW 18725. Thiaki Creek, macrophyte area, sume data as OMW 18841, [3 3 d. (3.2-4.8mm CL), 16 9 9 (3.8- 6.1mm CL); QMW21909 Thiaki Creek at Seamark Road crossing (17°23.5'S, 145932.5'E, 750m), open grazing land, [ringing para grass, water depth 0.3-(1. 5m, hand-netted, 26.8.94, c, 100 specimens; QMW21910 Thiaki Creek at downstream edge of rainforest, Ining-

28 MEMOIRS OF THE QUEENSLAND MUSEUM

ing para grass, depth 0.3-0.5m, hand-netted, 5.12.94, c. 50 spec.; QMW21917 Ithaca Creek in rainforest pocket (17?24.7'S, 145?30.3' E), leaf litter, water depth 0.3-0.5m, hand-netted, 5.12.94, 6 33,7 9 ?(2 oviger- ous). Upper Barron Catchment: QMW21911 Gwynne Creek at Gillies Road crossing (17°20.3’S, 145°31.1°E, 750m), open grazing land with small pocket of riparian rainforest upstream, fringing para grass, water depth 0.3-0.6m, hand- netted, 5.12.94, c. 200 specimens (1.28-5.84mm CL); QMW21912 upper Gwynne Creek in rainforest pocket (17?23.3'S, 145?30.3'E, 720m), hand-netted, leaf litter, water depth 0.2-0.4m, 5.12.94, 14 specimens (4.0-5.4mm CL).

DIAGNOSIS. Body slender; rostrum long, reaching tip of antennular peduncle (RL >0.5CL), dorso-ventrally com- pressed, may be armed with one dorsal tooth; antennular peduncle long (A1P>0.5CL), antennal peduncle long (A2P>0.6CL), stylocerite long (ST>0.4CL), scaphocerite long (SC>0.8CL), sixth abdominal segment long (6S>0.5CL) with acute postero-ventral margin; telson long and slender (T>0.6CL); dorsal telsonic spines confined to posterior half of telson, median spine on posterior telsonic margin ab- sent; protopod of uropods acute but not spinose.

DESCRIPTION. Body (Fig. la) small, sub- cylindrical; d d in collection up to 4.9mm CL, 9 9 up to 6.2mm CL.

Cephalothorax (Figs. 2i, j; 4a, b) rotund, gla- brous, breadth c. 0.7 CL, depth c. 0.7 CL; rostrum long, 0.43-0.76 CL, length 12-16 X height, curv- ing downward or sigmoid, reaching base to tip of distal segment of antennular peduncle, asetose, dorsoventrally compressed, a dorsal tooth may be present. Antennal spine short, strong, placed on lower orbital angle; pterygostomian angle obtuse, pterygostomian spine absent. Eyes large, c. 0.25 CL, corneal diameter c. equals eyestalk length, retinal pigmentation present. Antennular pedun- cle shorter than scaphocerite, 0.6-0.7 CL;

FIG. 2. a, C. zebra, from Wright's Ck, ovig. 9; b, C. zebra, from Kauri Ck, ovig. 9;c, C. zebra, from upper Gwynne Ck, 9 ; d, C. zebra, from Peterson Ck, d; e, C. zebra, from upper Tully River, 9; f, C. zebra, from upper Gwynne Ck, 9; g, C. spinula, 9; h, C. spinula, 3; i, C. confusa, from Gywnne Ck, ovig. 9; j, C. confusa, from Gwynne Ck, d. Scale = 1mm.

stylocerite length 0.7 X proximal antennular seg- ment length; anterolateral angle of proximal seg- ment acute, reaching to about 0.15 X intermediate segment length; intermediate segment 0.7 X proximal segment length, about 1.7 X distal seg- ment length; all segments with submarginal plu- mose setae; distal segment fringed laterally and apically with plumose setae. Antennal peduncle 0.5-0.6 X scaphocerite length; scaphocerite slightly longer than antennular peduncle, 0.8-1.0 CL, outer margin straight to slightly concave, asetose, ending in strong subapical spine, length 3.5 X width, distal lamella and inner margin with plumose setae. Branchial formula typical for genus.

Mandibles dimorphic, without palp; right man- dible with 6-8 strong, sharp incisor teeth laterally;

NEW FRESHWATER SHRIMPS 29

FIG. 3. a, C. zebra, telson and protopod of uropod, 3, 4.8mm CL; b, C. confusa, telson and protopod, d , 432mm CL; c, C. confusa, posterior body; d, C. zebra, posterior body; e, C, zebra, dactylus of 3rd pereiopod of 9 ; f, C. zebra, dactylus of 3rd pereiopod of dg, h, C. zebra, posterior margin of telson. Scales = Imm (a-d), 0.1 mm (e-h).

medially two groups of setae. one group with bent hamate setae, other group with finer straight plu- mose setae; molar process ridged; left mandible with 6-8 strong teeth; medially three groups of setae, molar process ridged.

Maxillula with simple palp. slightly expanded distally, with long plumose setae distally, few simple setae proximally; lower lacinia with broadly rounded margin, bearing several rows of plumose and simple setae; upper lacinia broadly elongate, inner edge straight, with several rows af strong spiniform, hamate, denticulate and plu- mose setae, outer and lower inner margins with plumose setae.

Maxilla with slender tapering palp, shorter than upper endite cleft, setose; margin and submargin of upper and middle endite with simple, hamate, plumose and denticulate setae; lower endite with hamate setae; scaphognathite with regular row of long plumose setae on distal margin, with shorter hamate ones continuing down proximal iriangu-

lar process which has e. 11 long simple setae, some with prominent dilation at base.

First maxilliped with broadly triangular lamel- lar palp, ending in pointed tip, margins with plu- mose setae; ultimate and penultimate segments of endites indistinctly divided; inner margin of ulti- mate segment with long denticulate setae, long rows of plumose. simple and hamate setae sub- marginally, transverse rows of plumose setae proximally; exopod flagellum distinct, well-de- veloped, with submarginal and marginal plumose setae; caridean lobe narrow, with marginal and submarginal plumose setae.

Second maailliped with dactylar and propodal segments of endopod fused; inner margins of ull three proximal segments with long simple, ha- mate and plumose setae; exopod long, narrow with marginal long plumose setae distally and shorter ones proximally.

Third maxilliped reaching beyond tip of an- tennular peduncle; endopod three-segmented, basal segment length c. 7 X width; penultimate segment length c. 7 X width, c. 0.9 X basal segment length, with transverse rows of spini- form hamate setae; distal segment c. 0,9 X as long as penullimate segment, ending in large claw-like apical hamate seta surrounded by simple and plumose ones, behind which there are 7-9 hamate setae on distal 3rd of posterior margin, clump of serrate and pappose setae proximally; exopod reaching about 0.5 of 2nd endopod segment, dis- tal margin with long plumose setae.

First pereiopod (Fig. 4c) reaching tip of basal antennular segment; chela length 1.7-2.2 X width, movable finger 1,! X as long as palm, length 2.9-3.1 X width; finger tips rounded, with- oul hooks, setal brushes well developed. Carpus attached to chela venirally, excavated distodors- ally, length 1.3-1.6 X width, 0.6-0.8 X chela length, 0.94 X merus length. Merus compressed, 0.6 X as wide as carpus. Ischium length 0.41 X merus length. Epipod present.

Second pereiopod (Fig. 4d) reaching up of 2nd segment of antennular peduncle, more slender and longer than Ist pereiopod. Chela length 2..4- 2.6 X width; movable finger length 4.9-5.1 X width, 1.5 X as long as palm; finger tips without hooks, setal brushes well developed. Carpus sub- conical, length 3.5-4.8 X width, 1.0-1.3 X chela length, 1.2 X merus length. Ischium length 0.67 X merus length. Epipod present.

Third pereiopod over-reaching antennular pe- duncle tip by about 0.33 distal propodus. Dacty- lus sexually dimorphic in adults (cf. Figs, 3e and 3f), length c. 3.8 X width, c. 0.2 X propodus

30 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. a-e, Caridina confusa sp. nov.; a, cephalothorax; b, anterior cephalothorax, dorsal view; c, Ist pereiopod; d, 2nd pereiopod; e, margin of abdomen and telson, lateral view; f-q, C. spinula sp. nov.; f, margin of abdomen and telson, lateral view; g, cephalothorax; h, rostrum and orbital margin, dorsal view; i, Ist pereiopod; j, 2nd pereiopod; k, 3rd pereiopod; I, dactylus of 3rd pereiopod; m, 5th pereiopod; n, dactylus of Sth pereiopod; o, posterior margin of telson; p, Ist pleopod of adult d ` q, endopod of 2nd pleopod of adult 3. Scales = Imm (a, g), 0.5mm (b-f, h-k, m), 0.1 mm (l, n, o-q).

NEW FRESHWATER SHRIMPS H

length, ending in prominent claw-hke hamate sela surrounded by simple setae, behind which posterior margin bears 4-6 shorter spiniform ha» mate setae, these being more robust and upright 1n adult d d. Propodus length 9-11 X width, posierior margin and lateral surface bearing two rows of small spiniform hamate setae. Carpus length 0.8 X propodus length, distal projectian feebly developed, posterior and lateral surfaces with up to 10 small hamate setae, more spiniform setation in adult d 3. Merus 1.1-1.4 X length of carpus, with 2-3 strong, movable spiniform hå- maie setae along posterior margin. Ischium 0.2 X length of merus. Epipod present.

Fourth pereiopod reaching tip of 2nd segment 10 Up of 3rd segment of antennular peduncle, Morphologically similar to 3rd pereropod.

Fifth pereiopod reaching tip of 2nd segment to tip of 3rd segment of antennular peduncle, Dac- tylus unguiculate, compressed, length c. 3.8 X width, ending in claw-like apical hamate seta, bearing comb-like row of 55-65 hamale setae gradually increasing in length distally on poste- nor margin, Propodus length 8-10 X width, 3.6 X dactylus length, bearing two rows of 15-20 short hamate setae on posterior margin. Carpus length 0.5-0.6 X propodus length, bearing 2-7 short ha- mate setae, distal projection. well-developed. Merus distinctly shorter (0.6-0.8 X) but broader (1.8 X) than propodus, bearing 2-4 large spini- form hamate setae. Ischium c. 0.4X length of merus, with simple setae. Epipod absent,

Abdomen (la) well developed, rotund, gla- brous, c. 2.8 X CL; sixth abdominal segment clongate, c. 0.6 X CL, length-depth ratio c.1.7; protopod of uropod (Figs. 3c, 2e) acute, aspinose; telson (Fig. 3b) narrow, length c. 0.7 X CL, dorsal spination (3-5 pairs) confined to posterior half af telson; posterior telsonic margin rounded, 3-4 pairs of spine-like setae, decreasing in size inte- riorly. median spine absent, diaeresis on telsonie exopod 10-16,

First 3 pleopodal endopod with well developed appendix interna arising sub-distally. Appendix interna of 2nd. Å pleopodul endopod reaching beyond middle of appendix masculina, with many retinaeulae distally. Appendix masculina subcylindrical, long hamate setae distally and on inner lateral margin,

Live colour, translucent brown,

ETYMOLOGY. The specific epithet ischosen to high- light the fact that this new species can be confused wilh Caridina zebra

REMARKS. Caridina confusa sp, nov. possesses & longer rostrum and is a much more slender and elongate animal than C. zebra. Wis found primar- ily in open grassland streams, flowing through pastural land of the upper Barron (Gwynne Creek) and the upper North Johnstone (Ithaca and Thiaki Creeks) catchments, Smaller numbers are found, together with C. zebra, in the small, rem- nant rainforest areas of these streams. Despite extensive sampling, C. confusa sp. nov, has not been found anvwhere else.

Caridina spinula sp. nov,

MATERIAL EXAMINED. HOLOTYPE. QMNW- 21913 adult $, 4, 6mm CL, | 2mm RL, 17,6mm SI... eust-flowing first order tributary of Leo Creek, near crossing Of Leo Creek Mine Road (13"44.6'5. 143921 S E]. Nesbit River catchment, Mcllwraith Range, Cape Yurk, alt. c. 450m, 12.7.95, J, Marshall, N, Phillips. ALLOTYPE. QMW21914 adult 4, 3.2mmCL, 0.8mm RL, 12.4mm SL, same locality data as holotype, PARATY PES. QMW219155d 2,7 2? (2 OVigerOus), same data as holotype; QMW21422, west-[lowing headwaters of Peach Creek (143*20'E, 13244" 8), Archer River catchment, Mellwraith Range. Cape York Peninsula, alt, c. 500m, 15.11.94, K, Mc- Donald, 7 33 (3,8-4.6mm CL), 34 9 (1 oun" (2.6-5.4mm CL. ovig. ? 4.9mm).

DIAGNOSIS. Body rotund, rostrum short (RL «0.4CL), dorsoventrally compressed but slightly elevated towards dorsal carina, may have å tooth on the ventral margin, reaching ta tip ol basal antennular segment; pterygostomian angle broadly angular to acute and spiniform; dorsal telsonic spines. confined to posterior half of tel- son, posterior median spine may be present; eges large (length >1.32mm).

DESCRIPTION. Body (Fig, le) small, sub- cylindrical; 3 å in collection up tà 2.7mm CL, ? 9 upto $.4mm CL,

Cephalothorax (Figs. Ic; 2g, hy 4g, h) round, glabrous, breadth c, 0,8 CL, depth 0.6-0.8 CL, rostrum short 0.20-0.33 CL, length 5-10 X height, curving downward, may be elevated medially, reaching tip of basal segment of antennular pe- duncle, may be setose dorsodistally, dorsoven- trally compressed, a rostral tooth may be present on the ventral margin. Antennal spine short, placed on orbital angle; pterygostomian angle broadly angular (in Peuch Creek specimens) or acure, spiniform (in Leo Creek specimens), Eyes large, c. 0.2 CL. corneal diameter c. equal eve- stalk length, retinal pigmentation present, At- tennular pedunele shorter than scaphocerite, 0.6-0,7 CL; styloverite length 0,7 X proximal

32 MEMOIRS OF THE QUEENSLAND MUSEUM

antennular segment length; anterolateral angle of proximal antennular segment acute, reaching lo about 0.15 X intermediate segment length; inter- mediate segment 0.7 X proximal segment length, about 1.7 X distal segment length; all segments with submarginal plumose setae; distal segment fringed laterally and apically with plumose setae. Antennal peduncle 0.5-0.6 X scaphucerite length; scaphocerite slightly longer than antennular pe- duncle, 0.6-0.7 CL. outer margin straight to slightly concave, asetose, ending in strong sub- apical spine, length 2.9-3.0 X width, distal la- mella and inner margin with plumose setae. Branchial formula typical for genus.

Mandibles dimorphic, without palp; right man- dible with 5-6 strong, sharp incisor teeth laterally; medially two groups of setae, one group with bent hamate setae, other group with finer straight plu- mose setae; molar process ridged; teft mandible with 5-6 strong teeth; medially three groups of setae, molar process ndged

Maxillula with simple palp, slightly expanded distally, with long plumose setae distally, few simple setae proximally; lower lacinia with broadly rounded margin, bearing several rows of plumose and simple setae; upper lacinia broadly elongate, inner edge straight, with several rows of strong spiniform, hamate, denticulate and plu- mose setae, outer and lower inner margins with plumose setae,

Maxilla with slender tapering palp, shorter than upper endite cleft, setose; margin and submargin of upper and middle endite with simple, hamate, plumose and denticulate setae; lower endite with hamate setae; scaphognathite with regular row of long plumose setae on distal margin, with shorter hamate ones continuing down proximal triangu- lar process which has c. I1 long simple setae, some with prominent dilation at base.

First maxilliped with broadly triangular lamel- lar palp, ending in pointed tip, margins with plu- mose setae; ultimate and penultimate segments of endites indistinctly divided; inner margin of ulti- mate segment with long denticulate setae, long rows of plumose, simple and hamate setae sub- marginally, transverse rows of plumose setae proximally; exopod flagellum distinct, well- developed, with submarginal and marginal plu- mose setae; caridcan lobe narrow, with marginal and submarginal plumose setae,

Second maxilliped with dactylar and propodal segments of endopod fused; inner margins of all three proximal segments with long simple harnate and plumose setae; exopod long, narrow with

TABLE |. Morphometric ratios (range) of pereiopods ot Caridina zebra Short, 1993, C, confusu sp. nov. and C. spinula sp. nov. D, dactylus; P, propodus; C, carpus; M, merus; L, length; W, width. 1, 2, 3 and 5 refer to the corresponding pereiopods.

[øm ivret! reen "rg CLEN

KEE

M3L/P3 11-13 LIU

Sars

PSL/PSW 115-145 RO-11.0

CSL/CSW 42357 4.146 2,9-5,8

MSL/M5W 5.7-7.5 5.6-6.0

CSL/PSL äng — [0505 0405 |

DES 1805

o EE — [pm A

marginal long plumose setae distally and shorter ones proximally.

Third maxilliped reaching beyond tip of an- iennular peduncle; endopod three-segmented, basal segment length c. 6.5 X width; penultimate segment length c. 8 X width, c. 0.9 X basal segment length, with transverse rows of spiniform hamate setae; distal segment c, 0.9 X as long as penulti- male segment, ending in large claw-like apical hamate seta surrounded by simple and plumose ones, behind which there are 7-9 hamate setae on distal third of posterior margin, clump of serrate and pappose setae proximally; exopod reaching about 0.5 of 2nd endopod segment, distal margin with long plumose setae. l

First pereiopod (Fig. 4i) reaching tip of basal antennular segment; chela length 1,9-2.2 X width, movable finger 1,1 X as long as palm, length 2.0-3.4 X width; finger tips rounded, with- out hooks, setal brushes well-developed. Carpus attached to chela ventrally, excavated distodors- ally. length 1.6-2.5 X width, 0.7-1.0 X chela length, 1,0 X merus length, Merus compressed,

NEW FRESHWATER SHRIMPS 33

TABLE 2. Distinguishing characteristics of Caridina zebra Short, 1993, rounded by simple setae, behind C. confusa sp. nov. and C, spinul sp. nov. CL, post-orbital carapace which posterior margin bears 4-5

length.

mer ZW

Thstrihitian mainly in rainforese covered streams of the upper Tully, Herbert, Johnstone and Barron catchments

Rostnun tà tip of Znd segment of antennat peduncte, length. 0.21-0.50 CL

0.58 + 0,01 CL.

lengilr Antennal peduncle , length

Stylocerive aptennular peduncle ‘segment, length 0,30 +0,04CL

Lenyth of ft abdominal segruent

Length-depth ratio of | 1.26 + 0.07 Gih abdom. segment

DAR t LOU

short, broad, 0.53 + 0.02 CL. anterior telsonic width 0.44 + 11.01 1alsonic length, posterior (elsonic width 0.19 4 0.01 Telsonic length

Dorsal telsonit to pasterior wo

Dev. epp size (mm). | 0,70-0.80.x 1.22-1.32

Scaphocerite length | 0.61 + 0,02 CL. 0,94 + 0,03 CT. 0,62+ 002 CL

Plerygostamian angle | bluntly angular bluntly angular bhunthy angular to acutá, spiniform

Protopod of uropod

Undey, egg (mm) 0.63-0.81 x 1.02-1.21 | U,63-0,85 x 0.98-1.36 | (,8O-.NS x 1,30-1,44 (1.73-0.88 x 1.22-1.44 | not avatlable Number oggs/ female | 38-57, mean 45 28-74, inean 51,5

shorter spiniform hamate setae,

these being more robust and up- e edad cu right in adult d 3. Propodus length peche? af streams ol" | streams of Nisbet und 10,5- 13.0 X width, posterior mar- the upper Johnstone | Archer egtetiments in gin and lateral surface bearing and Barron the Melllwraith rows of small spiniform hamate catchment: Range, Cape Vork setae. Carpus length 0.6-0.8 X pro- fo beyond tip of 2nd | la tip of Ist segrnent podus length, distal projection fee- segment of of antenfular bly developed, posterior and meire) epe pm lateral surfaces with 1 large and up 1.692 0.011 CL 055+ 001 CT. lo 5 small hamate setae, more

spiniform setation in adult d d. Merus 1.6-1.9 X length of carpus, with 2-4 strong, movable spini- Form hamare setae along posterior margin. Ischium 0.3 X length of metus. Epipod present.

Fourth pereiopod reaching tip of 2nd segment to tip of 3rd segment of antennular peduncle, morpho- logically similar to 3rd pereiopod-

Fifth pereiopod (Fig. 4m, n) reaching tip of 2nd segment tå tip of 3rd segment of antennular pe-

mot tn tip of Ist not to tip of lat antennular peduncle | antznnular peduncle segment, length O45 | segment, Inget 0,32 200505 s001CL

0.50 0.11 CL

LOR + (1:03

1.65 = 0.08

elongate, spinose

elongate, narrow. short, broad, 0,54 £

0,75 -- 0.04 CL, 0.05 CL, anyenor 4 ull anterior telsanic telsonic width DA): duncle. Dacty lus un guicu late, widih(39 001 ` | 001 telsonic lengib. compressed, length c. 4.0 X widti,

delsomte length. posterior telsonic width L5 500! telsonic length confined to posterior

posterior telsenic width 1.20 + 0.01 islsonre lengir

ending in claw-like apical hamate seta, bearing comb-like row of 45- 55 hamate setae gradually increas- ing in length distally on posterior

Jo postener puo

spination ihiras of telson half of telson Hints of telson å 3. F Telsonic margin methan sping may be | median spine absent | median spinte may be ina Tii. Propodus length 12-18 X

present »-— width, 3.3 X dactylus length, hear- 1622 ing rows of 10-15 short hamate

setae on posterior margin. Carpus length 0.4-0.6 X propodus length, bearing 2-7 short hamate setae, distal projection well-developed.

17-18, mean 17,5

0.6 X as wide as carpus. Ischium length 0.41 X — Merus shorter (0.8-0,9 X) but broader (1.5 X)

merus length. Epipod present.

Second pereiopod (Fig. 4j) reaching tip of 2nd segment of antennular peduncle, more slender

than propodus, bearing 2-4 large spinifurm ha- male setae. Ischium c. 0.3 X length of merus, with simple setae. Epipod absent.

and longer than Ist pereiopod, Chela length 2.4- Abdomen (1c) well-developed, rotund, gla-

2.9 X width; movable finger

length 3.2-4,5 X brous, c. 3 X CL; 6th abdominal segment elon-

width, 1.6 X as long as palm; finger tips without pate, c. 0.5 X CL, length-depth ratio c.1,8; hooks, setal brushes well-developed. Carpussub- ` protopod of uropod acute, aspinose; telson (Fig. conical, length 5.8-6.1 X width, 1.4-2.0 X chela — 40) broad, length c. 0.5 X CL, dorsal spinåtion length, 1.1 X merus length. Ischium length 0.48 — (3-5 pairs) confined to posterior 0.66 of telson; X merus length. Epipod present. posterior telsonic margin rounded, 4-5 pairs of

Third pereiopod (Fig. 4k, 1) over-reaching an- tennular peduncle tip by about 0.33 distal pro- podus. Dactylus sexually dimorphic in adults,

spine-like setae, decreasing in size anteriorly, me- dian spine may be present; diaeresis on telsonie exopod 16-22.

length c. 2.6 X width, c. 0.25 X propodus length, First å pleopodal endopod (Fig. 4p) with well-

ending in prominent claw-like

hamate seta sur» developed appendix interna arising sub-distally.

34 MEMOIRS OF THE QUEENSLAND MUSEUM

GE interna of 2nd 3 ver X2 endopod 4q) reaching beyond middle of appendix e with many retinaculae distally. Ap- pendix masculina subcylindncal, long hamate setae distally and on inner lateral margin. Live colour, translucent brown.

ETYMOLOGY, The specific name refers to the dis- linctive, spiniform pterygostomian angle which may be present in some specimens. No other Australian Car- idina species exhibits this spiniform plerygostomian angle.

REMARKS. Although C. spinula sp. nov. looks very much like C. zebra, there are distinct mor- phological differences (Tables | and 2). The spiniform pterygostornian angle m specimens from Leo Creek is ulso very distinctive. This species is currenily known only from the Mellwraith Range in the Cape York Peninsula und, despite extensive sampling, has not been found on the Atherton Tablelands around the Lamb-Francis-Cardwell Ranges (where C. zebra and C. confusa sp. nov. are found), The only other likely area of its occurrence between the Atherton Tableland and the McIlwraith Range that we have not sampled is the Cape Tribulation/Daintree area. However, none were found here during other sampling trips (J. Short and B. Herbert. pers. comm.), The Leo Creek tributary from where Caridina spinula sp. nov. was collected consists of a series of riffles and pools no more than 3m long, Im wide and 0.3m deep, flowing over a substrate of bedrock with some sand and gravel, Discharge at time of sampling was ap- proximately 1 Ls. Vegetation in the area is trop- ical mesophyll rainforest with a closed canopy at only 6-8m and some emergent vegelation. The low canopy suggests that this area may have recently been disturbed (David Hanger, pers. comm.). Density of the shrimp was low (1-2 D Other ånimals recorded from the collection site include the prawn Macrobrachium tolmerum (Decapoda: Palaemonidae) and the frogs Litoria genimaculata, L. longirestrus (Anura: Hylidac), Sphenophryne gracilipes (Anura: Microhylidae) and Rana daemeli (Anura: Ranidae).

DISCUSSION

The Caridina typus species-group. characterised by its short, dorsally unarmed ros trum, can be identified using the key of Choy and Horwitz (1995) (to couplet 6, p. 52), The four species can then be identified using the following key.

]. Rostrum long, extending beyond tip of second segment of antennular peduncle, 0.4-0.8 times carapace length; stylocerite long, 0.4-0.5 times carapice length; sixth abdominal segment long, 0.55-0.59 times carapace length

die CP A Caridina confusa sp.nov Rosin rum short, not reaching tip of second seg- ment of antennular peduncle, 0.2-0.5 times cara- pace length; stylocerite and sixth abdominal segment short, «0,4 and «0.5 times carapace length respectively

2. Rostrum somewhat laterally flattened, with 1-5 teeth on ventral margin; posterior telson margit angular, median plumose setae on posterior lel- son margin with selerotinous plug: eggs small, <0.6mm long C. rop

Rostrum somewhat dorsoventrally flattened, ven- tral teeth usually absent although one may be present; posterior telson margin rounded, me- dian plumose setae without sclerotinous plug; eggslarge,20.8mmlong .....

3. Rostrum short, not extending beyond up of first segment of antennular peduncle; length to depth ratio of sixth abdominal segment >1.4, number of eggs curried by female «25 C. spinula sp. nov.

Rostrum relatively long, usually extending be- yond tip of first segment of antennular pedun- cle, length 10 depth ratio of sixth abdominal scgment «1.4, number of eggs carried by fe- male 230 . ea omms rms Ca EDU

Caridina adr Short, 1993, C. confusa sp. nov. and C. spinula sp, nov, may also be separated on the basis of distinguishing characters given in Tables 1 und 2. It is emphasised that individual characters may be highly variable ånd so a com- bination of characters should be used to confirm the identity of keyed out specimens. The relation- ship between carapace length, rostrum, and the sixth abdominal segment lengths are linear for all three species (Fig. 5). The slopes and intercepts of the regression lines are significantly different for C. confusa when compared to the other two species (P>0.05). All three species tend to be allopatric. It i$ only in and near the short rainfor- est reaches of Gwynne and Thiaki Crecks that C. zebra and C. confusa are sympatric. Although C. zebra has been collected from some open, para grass infested and anthropogenically disturbed reaches of streams (e.g., Raspberry, Ithaca und Prior's Creeks) it does not appear 10 be very tolerant of these conditions. It seems to prefer the rainforest reaches of stream. C. zebra is a rotund animal, more commonly found on sandy, silty and/or leaf litter beds of riparian-covered rainfor- est streams and is particularly abundant in the more elevated, cooler areas (c. 800-950m), where fish predators and crustacean competitors are ah-

NEW FRESHWATER SHRIMPS 35

4 T——T T T T 4 T T T T T Caridina zebra C. zebra 3r 4 € "e y=0.084+0.317x, r-0.718, P«0.05 E £ £ e 2 2 e n e E 5 oO 3 i 1 £ E D [0] L 1 1 1 a 1 2 3 4 5 6 7 e 1 1 1 L

andi 1 2 3 4 5 6 7

Carapace Length (mm) Carapace Length (mm)

4 [——T T T

Caridina confusa C. confusa Ü

Rostrum Length (mm) bv] Sixth Abd. Seg. Length (mm) N T 1

^ y=-0.124+0.607x, r-0.842, P<0.05

[9] L L L L l 1 2 3 4 5 6 7 [0] 1 Ge 4 i - 2 3 4 5 6 7

Carapace Length (mm) Carapace Length (mm)

4 T T T T T 4 T T 1 T Caridina spinula C. spinula

L J € € d E E y=0.177+0.196x, r=0.648, P<0.05 ro £ 2 a, vod ger J ? E o 2 3 E Ed a PA 1 x [n]

[e] L L L 1 L [6] l L L L 1

1 2 3 4 5 6 7 1 2 3 4 5 6 7 Carapace Length (mm) Carapace Length (mm)

FIG. 5. Correlation between carapace length, rostrum length and sixth abdominal segment length of Caridina zebra, C. confusa sp. nov. and C. spinula sp. nov. All regressions are significant (P<0.05). The slopes and intercepts of the regression lines of the appropriate variables between C. confusa and C. zebra and between C. confusa and C. spinula are significantly different (ANOCOVA, P>0.05).

36 MEMOIRS OF THE QUEENSLAND MUSEUM

sent, C. confusa sp. nov. is a more elongate, slender animal more commonly found amongst the bank vegetation (particularly para grass) of streams flowing through open grassland areas of the Atherton Tableland.

It is likely that both the species were present in these previously rainforested areas. The modified environment may be favouring C. confusa, hence its predominance in these areas.

ACKNOWLEDGEMENTS

Stuart Bunn, Chris Marshall and David Hur- wood of Griffith University, Brett Herbert of the Department of Primary Industries, Walkamin and Mark Hopper of the Department of Natural Re- sources, Mareeba assisted in collecting the sam- ples. Peter Davie allowed access to the Queensland Museum collection and John Short assisted in many ways, including helpful discus- sion. Funding from the Co-operative Research Centre for Tropical Rainforest Ecology and Man- agement and the Wet Tropics Management Au- thority (through Griffith Universily), the Monitoring River Health Initiative Program and the Department of Natural Resources made pos- sible the expeditions to collect samples. Stuart Bunn, Jane Hughes and John Short provided con- structive criticism of the draft manuscript.

LITERATURE CITED

BOUVIER, EL 1925. Recherches sur la morphologie, les variations, la distribution géographique des Crevettes de la famille des Atyidés. Encyclopédie Entomologique, Sér. A 4: 1-370.

CHOY, S, 1991. The atyid shrimps of Fiji with descrip- tion of a new species. Zoologische Medelingen 65(27): 343-362.

CHOY, S. & HORWITZ, P. (1995). Preliminary key to the species of Australian shrimps (Atyidae) found in inland waters. Pp. 51-54. In Horwitz, P, "A preliminary key to the species of Decapoda (Crustacea: Malacostraca) found in Australian in- land waters. Identification Guide No. 5.' Co-oper- ative Research Centre for Freshwater Ecology, Albury.

FELGENHAEUR, B.E. (1992). External anatomy and integumentary structure. Pp. 7-43. In Harrison, EW E Humes, A.G. (eds), ‘Microscopic anatomy of invertebrates’ Vol. 10. (Wiley-Liss: New York).

HOLTHUIS, L.B. 1965. The Atyidae of Madagascar. Memoires du Museum national d'Histoire Naturelle (N.S.) (A), Zoologie 33(1): 1-48.

HUGHES, J.M., BUNN, S.E, HURWOOD, D.A., CHOY, S.C. & PEARSON, R.G. 1996. Genetic differentiation among populations of Caridina zebra (Decapoda; Atyidae) in the tropical rainfor- est streams, northern Australia. Freshwater Biol- ogy 36: 287-296.

MILNE EDWARDS, H. 1937. Histoire naturelle des Crustaces, comprenant l'anatomie, la physiologie et la classification de ces animaux 2: 1-532, Atlas, pp. 1-532, pls. 1-42.

RIEK, E.F. 1953. The Australian freshwater prawns of the family Atyidae, Records of the Australian Mu- seum 23: 111-121.

ROUX, J. 1926. An account of the Australian Atyidae. Records of the Australian Museum 15(3): 237- 254.

SHORT, J.W. 1993. Caridina zebra, a new species of freshwater atyid shrimp (Crustacea: Decapoda) from northeastern Queensland rainforest. Mem- oirs of the Queensland Museum 34(1): 61-67.

GASTROPODS FROM THE BURDEKIN FORMATION, MIDDLE DEVONIAN, NORTH QUEENSLAND

ALEX G. COOK

Cook, Alex G. 1997 06 30: Gastropods from the Burdekin Formation, Middle Devonian, north Queensland. Memoirs of rhe Queensland Museum 42(1): 37-49. Brisbane, ISSN

0079-8835.

Twenty one taxa of gastropods are described from the Middle Devonian Burdekin Formation, north Queensland. New amongst these are Burdekinostoma burdekininensis gen. et. sp. nov., Euryzone burdekinensis sp.nov., Anomphalus pajelli sp. nov., Didymalgia bartholomai gen. el sp. nov., Murchisonia (Murchisonia) jackjelli sp. nov., Palaeozygopleura machenryi sp. nov., and Trinema heideckeri gen. et sp. nov. The fauna contains typical Old World Realm genera but is specifically strongly endemic. [ ] Gastropods, Devonian, Queensland, Burde-

kin Formation,

Alex G, Cook. Queensland Museum, PO Box 3300, South Brisbane, Queensland 410],

Australia; 10 March 1997.

Burdekin Formation is å shallow marine car- bonate sequence exposed within the Burdekin Subprovince (Wyatt & Jell, 1967: Henderson, 1980; ). The formation consists of, inter alia, a reefal and biostromal complex, with associated lagoonal. inter-reef and offshore facies (Cook, 1995). It contains diverse fossil assemblages which include rugose corals (Zhen 1991, 1994), tabulate corals, stromatoporoids (Cook, 1994), sponges, molluses (Cook, 1993a) and conodonts (Talent & Mawson, 1994). Large, thick-shelled gastropods from the underlying and interdigitat- ing. siliciclastic-dominated, Big Bend Arkose and basal units of the Burdekin Formation are Burdikinia burdekinensis (Etheridge, 1917), Amphelissa carinatum (Heidecker, 1959), Labrocuspis nodosa Heidecker, 1959, and Fletcherviewta septata Cook, 1993b.

This paper is concerned with a diverse gastro- pod faunule from a stratigraphically higher unit in the Burdekin Formation in coralline packstones interpreted to represent shallow- water coralline thickets seaward of a biohermal structure (Cook, 1995), The fauna has been silicified and was recovered by bulk dissolution in acelic acid. Both large and small gastropods were retrieved, in addition to corals, brachiopods and bivalves. All material comes from QML1094. near Little Rocks, 27km NW of Char- ters Towers.

SYSTEMATIC PALAEONTOLOGY

Phylum MOLLUSCA Class GASTROPODA Order ARCHAEOGASTROPODA Superfamily BELLEROPHONTOIDEA Family BELLEROPHONTIDAE McCoy, 1851 Subfamily TROPIDODISCINAE Knight, 1956

Tropidodiscus Meck & Worthen, 1866 Tropidodiscus sp. (Fig. 1A-C)

MATERIAL EXAMINED. QMF35513, QMF35514.

DESCRIPTION. Shell minute, approximately 2.5mm in diameter, 0.8mm wide; involute with moderately wide umbilicus. Whorl profile sharply angular, without obvious crest. Relics of fine, numerous growth lines present.

Bellerophon de Montfort, 1810 Bellerophon (Bellerophon) de Montfort, 1810

Bellerophon (Bellerophon) sp. Å (Fig. 1D,E) MATERIAL EXAMINED. QMF34916, QMF34902,

DESCRIPTION. Small to medium-sized, spher- ical, involute, 5.5mm wide; narrow deep umbili- cus. Whorl profile evenly rounded; margin bearing weak crest. Aperture unknown. Orna- ment consists of many fine growth lines,

REMARKS. The few specimens possess similar characteristics to the type, B.(B.) vasulites de Montfort, 1810 from the Middle Devonian of

38 MEMOIRS OF THE QUEENSLAND MUSEUM

Germany as figured by Knight (1941). The nu- merous growth lines and overall form are similar, but the Burdekin form is much smaller (Knight, 1941: pl. 11) and the selenizone-bearing crest is weaker, possibly due to imperfect silicification.

Bellerophon (Bellerophon) sp. B (Fig. IEG)

MATERIAL EXAMINED. QMF34908.

DESCRIPTION. Spherical, medium-sized, ap- proximately 10mm in diameter, 9mm high; invo- lute with deep, wide umbilicus. Whorl profile wide and rounded with weak crest; slight flatten- ing of inner whorl surface adjacent to penultimate whorl, extending approximately one third across the inner whorl profile.

REMARKS. The specimen is larger and has a wider whorl profile than Bellerophon (Bellerophon) sp. A, which is a result of the flattened area on the whorl profile abutting the penultimate whorl.

Superfamily EUOMPHALOIDEA de Koninck, 1881 Family EUOMPHALIDAE de Koninck, 1881

Straparollus de Montfort, 1810 Straparollus (Euomphalus) Sowerby 1814

Straparollus (Euomphalus) sp. (Fig. 1Q)

MATERIAL EXAMINED. QMF34965.

DESCRIPTION. Medium-sized, planispiral, 21.4mm wide, 8.2mm high. Suture in a channel. Whorl profile square with prominent angulation above and below gently convex midwhorl sur- face. Weak spiral line adumbilically to the upper angulation which may mark the position of the UD Aperture rounded, but quadrate. Base flat- tened.

Straparollus (Serpulospira) Cossman, 1916 Straparollus (Serpulospira) sp. (Fig. IR)

MATERIAL EXAMINED. QMF34966.

DESCRIPTION. Shell planispiral, disjunct, 13.6mm wide, 4.6mm high. Whorl profile rounded, aperture rounded. No growth lines pre- served.

Suborder PLEUROTOMARIINA Cox & Knight, 1960 Superfamily PLEUROTOMARIOIDEA Swainson, 1840 Family RAPHIOSTOMATIDAE Koken, 1986 Subfamily OPHILETINAE Knight, 1956

Burdekinostoma gen. nov.

TYPE SPECIES. Burdekinostoma burdekinensis sp. nov.

ETYMOLOGY. For the Burdekin River, and alluding to the Raphiostomatidae.

DIAGNOSIS. Minute, low-spired to lenticular, gradate shell, widely phaneromphalous, having a commonly channeled suture and prominent, gently inclined, flattened area on the upper whorl surface bordered by 2 strong cords, inferred to contain the selenizone and also commonly con- taining weak threads; midwhorl profile generally subrounded but stepped with several cords.

REMARKS. Placement of this striking if minute genus is difficult, as it superficially resembles members of the Ophiletinae, Raphiosomatinae and Euomphalidae. Placement depends on the interpretation of the inclined surface on the upper whorl face, which is inferred to contain the selenizone. In particular the raphiostomine Wisconsonella Blodgett (1988) from the Eifelian of North America is a similar taxon to the Burde- kin material, sharing the gradate shell form, but lacking the additional cords and the channeled suture, Placement within the Raphiostominae is restricted by the widely phaneromphalus nature of this snail. Alternatively the channeled suture may be interpreted as containing a sinus, which would ally the genus to the Euomphalidae, of which Poleumita Clarke & Ruedemann and Cen- tifugus Bronn are similar cord-bearing forms. Neither of these placements is satisfactory. Place- ment in the Ophiletinae best satisfies the interpre- tation of the selenizone, accomodates the channelled suture and the phaneromphalous con- dition.

Burdekinostoma burdekinensis sp. nov. (Fig. 2J-P)

MATERIAL EXAMINED. HOLOTYPE: QMF34961. PARATYPES: QMF35522-QMF35530.

ETYMOLOGY. From the Burdekin River. DIAGNOSIS. As for genus.

GASTROPODS FROM THE BURDEKIN FORMATION 39

FIG. 1. A-C. Tropidodiscus sp. SEM images. A,B, QMF35512, side and oblique views respectively, x 16; C, QMF35513, side view, x 16. D-E. Bellerophon (Bellerophon) sp. A. QMF34916, apertural and side views respectively. F, G. Bellerophon (Bellerophon) sp. B. QMF34908, side and broken apertural views respectively. H-P, Euryzone burdekinensis sp. nov. H-J, Holotype QMF34917, apical apertural and basal views respectively, x 4,4; K,L, Paratype QMF 34969, basal and apical views, x 3.2; M,N, Paratype QMF34964 apical and basal views x 6.3: O, Paratype, QMF35536 oblique side view, x 22.5, SEM image; P, Paratype QMF35536 apical view x 22,5, SEM image. Q, Straparollus (Euomphalus) sp. QMF34965 apical view, x 1.8. R, Straparollus (Serpulospira) sp. QMF34966, apical view x 2.3.

40 MEMOIRS OF THE QUEENSLAND MUSEUM

DESCRIPTION. Shell small to minute, less than 4.5mm wide and 1.5mm high, very low-spired, somewhat gradate, widely phaneromphalous, with deep umbilicus. Whorl profile series ot an- gular surfaces but giving an overall rounded pro- file. Suture with adjacent, wide channel. Upper whorl face bears numerous spiral threads, of at least 2 orders of intensity. Three prominent threads on smaller specimens; bordering ihe su- tural channel, another high on the whorl profile and another at the edge of the upper whorl face giving way to the steeper mid whorl surfaces, the last 2 inferred to contain the selenizone. At least 2 threads on the midwhorl, creating å vertical surface at the midwhorl periphery: at least 3 revolving cords on the lower whorl face. The upper whorl face bears up to 7 minor spiral threads, in addition to the major cords.

REMARKS. This striking taxon is easily differ- entiated by Euryzone burdekinensis sp. nov. by the numerous spiral threads and more angular whorl profile.

Family GOSSELETINIDAE Wenz, 1938 Sublumily COELOZONINAE Knight, 1956

Euryzoue Koken, 1896 Euryzone burdekininesis sp. nov. (Fig. (HP

MATERIAL EXAMINED. HOLOTYPE: QMF24917. PARATYPES: QMF34918-QMF34927.QMF34964, QMF34969.

DIAGNOSIS. Small to minute member of genus with sinus on upper whorl profile bouded by fine ihreads,

DESCRIPTION, Shell discoidal, minute to small, up to 11.9mm wide, 3,6mm high, generally smaller (Table 1). Whorls abut, with impressed sulure. Base with moderately deep umbilicus. Whorl profile rounded, bearing two fine cords which border the sinus. Growth lines fine, numer- ous. Shell repair in evidence on holotype.

REMARKS, The species is characterised by its discoidal shape and the striking presence of the sinus on the upper whorl face and its bordering cords. The most similar member of the genus to the Burdekin material is Euryzone perilitornara Linsley 1968 from the Middle Devonian An- derdon Limestone, Michigan, but its selenizone is slightly lower on the whorl profile, and it is à less planispiral than the Burdekin species. The type species E. delphinuloides (Schotheim) from

TABLE |. Measurements lor Euryzene burdekinensis Sp. DON,

QMF34969

QMF34964 20 urn il

the Middle Devonian Stringocephalus Lime- stone, Germany, has a turbiniform shell and a wider selenizone.

Suborder TROCHINA Cox & Knight, 1960 Superfamily ANOMPHALOIDEA Wenz, 1938 Family ANOMPHALIDAE Wenz, 1938

Anomphalus Meek & Worthen, 1567

TYPESPECIES. Anomphalus rotulus trom the Middle Pennsly vanian, St Davids Limestone, Illinois, USA, by original designation,

Anomphalus pajelli sp. nov. (Fig. 2A-T)

MATERIAL EXAMINED. HOLOTYPE: QMF34850, PARATY PES: OMF34851-OMF343858.

ETYMOLOGY, For Peter A. Jell.

DIAGNOSIS. Smooth-shelled, medium-sized member of genus with very weak sutures and prominent thickening of inner lip which creates a hemiomphalus base.

DESCRIPTION. Medium-sized, low-spired, rotelliform hemiomphalus gastropod, up tà 11.5 mm high, 13.5 mm wide (Table 2). Sutures flush, commonly difficult to detect, whorls embrace high on upper whorl surface. Umbilicus narrow obscured or occluded by thickening at the inner labrum, otherwise base rounded, Whorl profile rounded with periphery at midwhorl. Aperture rounded. Shell thick, inner labrum particularly thickened forming partial umbilical plug. Shell smooth, very faint growth lines on the base of QMF34851. There is some variation in the taxon with respect to where the whorls embrace. On QMF34851 the whorls embrace further down the whorl face, producing a slightly more naticiform shell shape. Despite silicà replacement the

smooth shell has an almost polished appearunce

on some specimens.

GASTROPODS FROM THE BURDEKIN FORMATION 4l

FIG. 2. A-I. Anomphalus pajelli sp. nov. A-C, Holotype QMF34850, apertural, apical and basal views, x 9. D-F, Paratype QMF34852, apertural, basal and apical views, x 2.2. G-I, Paratype QMF34851 apical, apertural and basal views, x 2.5. J-P, Burdekinostoma burdekinensis gen. et sp. nov. J,K, Paratype QMF35523, basal and apical views, SEM images, x13.5; L.M, Paratype QMF 35522, oblique apertural and apical views, SEM images, x 15; N, QMF 35525 side view, x 13.5; O,P, Holotype QMF34961, x 5.4, apical and side views respectively.

42 MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 2. Measurments for Anomphalus pajelli sp. nov,

| — QMF34850 13.1 OMFMRSI | us |

REMARKS. Both Knight (1941) and Knight et al. (1960) noted the variable umbilicus in this genus. The Burdekin material is assigned to the genus on the basis of gross shape, lack of orna- ment and the hemiomphalus condition, The type species Å. rotulus Meek & Worthen as figured by Knight (1941) has a slightly lower whorl profile and more prominently incised sutures. An- omphalus umbilicoliratus Batten (1966) from the Lower Carboniferous Hotwells Limestone, En- gland is clearly phaneromphalous and has spiral ornament on the base. Anomphalus helicinaeformes (Schlotheim) from the Middle Devonian Stringocephalus Limestones near Gladbach, Germany has a fully developed umbil- ical plug (see Knight 1941: pl 63), but in overall aspect is most similar to the Burdekin species, and is closely related. Specimens illustrated by Knight (1941) show minor variation in spire height for A. helicanaeformis (Schlotheim). Naticopsis margheriti Mansuy (1912), may also be an Anomphalus, albeit with å slightly more naticiform shell.

Suborder NERITOPSINAE Cox & Knight, 1960 Superfamily NERITOIDEA Rafinesque, 1815 Family NERITOPSIDAE Gray, 1847

Naticopsis M'Coy, 1844

Naticopsis (Naticopsis) sp. (Fig. 4A)

MATERIAL EXAMINED. QMF34913,

DESCRIPTION. Very small, 4.9mm wide, Smm high, naticiform, anomphalous; suture adpressed to slightly impressed; whorl profile rounded, with periphery well above midwhorl. Upper whorl surface flatter that lower whorl face. Final whorl dominated by strong collabral cords, increasing in intensity towards the aperture. Early whorls lack this ornament. Parietal and inner lip heavily thickened; aperture ovate. Base rounded.

REMARKS. The specimen strongly resembles Natica nexicosta (Phillips) of Whidbome (1892),

from the Middle Devonian at Lummaton, En- gland, but the latter possesses strong collahral cords on early whorls. Naticopsis ( ?Naticopsis) sp. (Rollins, Eldridge & Spiller, 1971), from the Middle Devanian Marcellus Formation, New- York, shows a similar ornament, but the threads are fine and inclined on the whorl face, Straparollus corrugatus (Stauffer, 1909), which Linsley (1968) placed within ”Ysonema, and Rol- lins, Eldridge & Spiller (1971) assigned to Naticopsis (Naticopsis), from the Middle Devon: jan of Ohio also shows strong collabral ornament, but not restricted to the mature whorls. ?/sarte me corrugatus (Stautfer) from the Middle Devonian Anderdon Limestone (Linsley, 1968) is a mi- nutely illustrated taxon. Linsley (1968) describes the early whorls as smooth suggesting affinity lo the Burdekin specimen. Positive generic identifi- cation of Linsley's taxon is impossible without viewing his material. 1 place Natica nexicosta (Philips) of Whidborne and Srraparollus corrugatus Stauffer both within Naticopsys (Naticopsis) alongside this distinct, if poorly pre- served and unresolved, taxon from the Burdekin Forinaton.

Superfamily ORIOSTOMATOIDEA Wenz, 1938 Family ORIOSTOMATIDAE Wenz, 1938

Didymalgia gen nov. TYPE SPECIES. Didvmalgia bartholamai sp. nav.

ETYMOLOGY, For ‘Didymalgia’ hill, informal name of laterite profile near the Fletcherview locality,

Didymalgia bartholomai gen. et sp. nov. (Fig. 3)

MATERIAL EXAMINED. HOLOTYPE; QMF34896. PARATYPES: QMF34876-OMF34895, QMF34915.

ETYMOLOGY. For Alan Bartholomai.

DIAGNOSIS. Turbiniform, widely phan- eromphalous shell with numerous revolving cords and strong comarginal cords forming and cancellate structure, whorl profile angular, formed by series of inclined surfaces.

DESCRIPTION. Medium-sized, turbinilorm, widely phaneromphalus shell, up to 9.5mm high and 11.1 mm wide (Table 3), Whorl face adorned with strong revolving cords and slightly less strong comarginal ribs, producing a cancellate shell decoration, Whorl profile series of angu- lar surfaces with a subsutural shelf, steeply inclined upper whorl face, angular periphery,

GASTROPODS FROM THE BURDEKIN FORMATION 43

FIG, 3, Didymalgia bartholomai gen. et sp. nov. A-D, Holotype QMF34896, x 3 apertural, basal (x 2.8), apical and side views (x 2.8). E-G, Paratype QMF34875. E, apertural view x 5; F, apical views x 5; G, side view x 4.

LJ, Paratypes QMF34876, x 3 apical and side views.

inclined lower surface leading to the phan- eromphalous base, sutures embrace just below periphery which is situated slightly below mid- whorl. Slight subsutural shelf, 4 major revolving cords on the upper whorl face, a major cord at the periphery, and 7 to 8 cords on the lower whorl face and base. Two orders of cords on the lower whorl face. Aperture rounded to subrounded, slightly wider than high. Thick colabral cords are numerous almost orthocline, and are not de- flected by a sinus or selenizone.

REMARKS. Oriostoma sp. aff O. gerbaulti Oehlert of Blodgett & Johnson (1992) from the Eifelian of Nevada is grossly similarto the Burde- kin material but has a channeled subsutural zone. Gyronema multinodosa Blodgeu & Johnson from the Bifelian of Nevada, which could be accomodated in Kitikamispira, has fewer cords and more prominent nodes on the shell surface. Placement of the genus within the Ori-

ostomatidae is problematic, in that it extends the range of the family beyond their supposed Eifelian demise (Blodgett et al. 1990), but the strong ornament, lack of selenizone on the whorl face and trochiform, phaneromphalus shell form make the placement appropriate, if contro- versial.

TABLE3. Data for Didymalgia bartholomai gen. et sp.

nov. Height (mm) 9 i ia

Specimen QMF34896

A QMF34876 72

H MEMOIRS OF THE QUEENSLAND MUSEUM

Suborder MURCHISONIINA Cox & Kmght, 1960 Superfamily MURCHISONIOIDEA Koken, 1896 Family MURCHISONIIDAE KOKEN, 1896

Murchisonia d' Archaic & de Verneuil 1841

Murchisonia (Murchisonia) jackjelli sp. nov. (Fig. 4B-H)

MATERIAL EXAMINED. HOLOTYPE: QMF34905. PARATYPES; QMF34897-QMF34900, QMF34902, QMF34903, QMF34929, QMF34932, QMF34933, QMF34935-QMF34940, QMF34942-QMF34943, QMF34946-QMF34948, QMF34952-QMF34953.

ETYMOLOGY. For J.S. Jell.

DIAGNOSIS. Medium-sized member of subge- ius witharelatively wide selenizone, subrounded whorl profile and up to 5 threads on the adult lower whorl surface,

DESCRIPTION. Medium-sized to large, high- spired shell, up to 41.5mm long and |1.6mm wide at base. Suture impressed, whorls embrace at approximately lower one third ot the whorl height. Whorl profile subrounded but with 2 weak peripheral cords bordering a wide selenizone, Up to 5 additional weaker spiral threads on the lower whorl face with 2 extremely faint threads on the upper surface. In juvenile specimens cords bor- dering the selenizone are stronger on early whorls, and the lower whorl face lacks the multi- ple threads of the adult whorls, instead 1 or 2 threads are more prominent. Aperture ovate with minor thickening of the inner lip.

REMARKS. Poor preservation of the material significantly impedes the study of this species. Clearly the presence of the wide midwhorl selenizone denotes the subgenus, and the numer- ous weak threads on adult whorls distinguish a new taxon. The width of the selenizone and the numerous threads may suggest Carboniferous Stegecoelia, butthe selenizone is at the periphery negating that assignment. In the Middle Devon- ian Murchisonia (Murchisonia) underwent sig- nificant radiation into a multitude of unusual shell forms (Knight et al. 1960; Blodgett et al., 1990) and the included species described to date are numerous (e.g., Whidborne, 1892). Taxa with similar multiple threads on the whorl face are. M. (M.) vicariana Whidborne, M. (M.) loxanemoides Whidborne, both from the Middle Devonian of Southern England, but the former hàs nodose threads and the latter a thinner, lower set selenizone.

Murchisonia (Murchisonia) sp, Å, (Fig. AI-J)

MATERIAL EXAMINED. QMF34930.

DESCRIPTION. Small, 10.5mm high, 5.3mm wide, high-spired, with eight whorls present. Su- ture strongly impressed, whorls embrace well below midwhorl; angular whorl profile with 2 prominent spiral cords bordering a selenizone. Other decoration not preserved.

REMARKS. A selenizone on the midwhorl, bor- dered by 2 cords is the classic murchisoniid con- dition and the specimen is here allocated to the subgenus on this basis. The species resembles M. (M.) fermioni Tassell, 1982, from the Emsian ot New South Wales, and also known from the Givetian in the Broken River Province, but the material is too poorly preserved to be confident of any specific assignment.

Murchisonia (Murchisonia) sp. B. (Fig. 4K)

MATERIAL. EXAMINED. QMF35520.

DESCRIPTION, Shell small, 6mm high, 2mm basal width, high-spired, with 7 whorls present. Sutures impressed, whorls embrace just below periphery. Whorl profile angular with 2 strong cords bordering a selenizone at the periphery. the upper cord slightly stronger. Upper whorl face gently sloped with a weak spiral cord close to the main omament on the margin. Lower whorl pro- file obscured, aperture and base unknown.

REMARKS. The bordered selemzone identifies the subgenus. It is higher-spited than Murchisonia (Murchisonia.) sp. A. and has.a significantly dif- ferent whorl profile than Murchisonia (Murchisonia) jackjelli sp. nov.

Superfamily CRASPEDOSTOMIDOIDEA Wenz, 1938 | Family CODONOCHEILIDAE Miller. 1889

Mitchellia de Koninck, 1876 ?Mitchellia sp. (Fig. 4L,M) MATERIAL EXAMINED. QMF35515, QMF35516,

DESCRIPTION. Two fragmental specimens, neither of which have a gerontic stage or aperture preserved. Small high-spired gastropod with im- pressed suture; whorls embrace just below mid-

GASTROPODS FROM THE BURDEKIN FORMATION 45

FIG. 4. A, Naticopsis (Naticopsis) sp. QMF34913, oblique apical view, x 5. BH. Murchisonia (Murchisonia) Jackjelli sp. nov. B, C, Holotype QMF34905, side views x 2.8; D, Paratype QMF34900, side view x 1.5; E, Paratype QMF34897, side view x 2; F, Paratype QMF34903, side view x 2.6; G, Paratype QMF34899 apertural view x 1.6; H, Paratype QMF35520, side view x12. I,J, Murchisonia (Murchisonia) sp. A. QMF34930, x 3.5. I, side view; J, apertural view. K Murchisonia (Murchisonia) sp. B. side view x 10. L, M, ?Mitchellia sp. L, QMF35515, x 20; M, QMF35516 x 20.

whorl. Whorl profile rounded, face bearing nu- lian Emsian genus on the basis of gross shell merous, (up to 10) spiral threads of equal inten- shape. It lacks the characteristic aperture needed sity. Aperture not preserved, but inner whorl for more definite assignment to the taxon. profile ovate with long axis vertical.

REMARKS. Generic assignment is impossible without more material, but the material is ques- tionably assigned to the common eastern Austra-

46 MEMOIRS OF THE QUEENSLAND MUSEUM

Order CAENOGASTROPODA Cox, 1959 Superfamily LOXONOMATOIDEA Koken, 1889 Family PALAEOZYGOPLEURIDAE Horny 1955

Palaeozygopleura Horny, 1955

Palaeozygopleura machenryi sp. nov (Fig. 5A-F)

MATERIAL EXAMINED. HOLOTYPE: QMF35531. PARATYPES: QMF35532-QMF35525, QMF35539,

ETYMOLOGY. For Colin McHenry.

DIAGNOSIS. Small to minute member of genus with moderately impressed suture and only slightly sigmoidal ribs.

DESCRIPTION. Shell high-spired, many whorled, up to 8 present, up to 4mm long and 1.6mm wide. Sutures impressed, whorls embrace well below midwhorl; whorl profile rounded. Pe- riphery at midwhorl. Whorl face bears many slightly sigmoidal ribs, obvious on all adult whorls. Protoconch poorly preserved, but smooth and dextral.

REMARKS. The material is most similar to P. joanni Linsley from the Middle Devonian An- derdon Limestone of Michigan, but in the Burde- kin species the ribs are less numerous and the suture less impressed. It differs from the single specimen Palaeozygopleura sp. described below by the more impressed sutures and coarser orna- ment.

Palaeozygopleura sp. (Fig 5G)

MATERIAL EXAMINED. QMF35519,

DESCRIPTION. Minute, 3.6mm high, 1.5mm wide, high-spired, nearly pupiform with shal- lowly impressed suture, gently rounded whorl profile. Ornament consists of fine, prosocline threads on the whorl surface.

REMARKS. The single specimen has weaker sutures than P. machenryi sp. nov, and finer pro- socline ornament.

Family TURRITELLIDAE Woodward, 1851

Trinema gen. nov.

TYPE SPECIES. Trinema heideckeri sp. nov. from the Middle Devonian Burdekin Formation, north Queens- land.

ETYMOLOGY. Tri-, three, nema (Greek) thread.

DIAGNOSIS. Small to medium-sized, anompha- lous, high-spired, many whorled shell; whorl pro- file rounded, bearing three prominent midwhorl cords. Suture impressed; a subsutural surface is bounded below by a weak thread which forms the upper limit of the deep sinus, in turn bounded below by the highest of the three midwhorl cords. Finer threads are present on the lower whorl face. Growth lines are prosocline above the sinus and opisthocline below. Whorls embrace just below the higher of the weak threads on the lower whorl face.

REMARKS. Placement in the family is indicated by the overall shell form and the position of the sinus above the midwhorl cords (see Knight et al. 1960: I317). The taxon is differentiated by the only other Devonian turritelid (Linsley 1978), Acanthonema Sherzer & Grabau, 1908 by lack- ing the distinctive nodose cords. In the Late Car- boniferous Orthonema Meek & Worthen, 1861 (see Knight 1934, 1941, Knight et al., 1960) there are 2 prominent midwhorl cords, widely sepa- rated below the similarly positioned sinus. In Callispira Nelson,1947, the sinus is also posi- tioned high on the whorl face, but the whorl face is less rounded and there are many more stronger cords (5-6) on the whorl face. There is a consid- erable gap in the taxonomic record of the Turritellidae between the appearance of Acanthonema in the Early Devonian (Sherzer & Grabau, 1908), their Middle Devonian record (herein) and the numerous Carboniferous records of Orthonema (Knight, 1934). Such a discontinu- ity in the record may be resolved by analysis of Late Devonian age faunas.

Trinema heideckeri sp. nov. (Fig. SH-K)

MATERIAL EXAMINED. HOLOTYPE: QMF34954. PARATYPES: QMF34941, QMF34945, QMF34955.

DIAGNOSIS. As for genus.

DESCRIPTION. Material somewhat fragmental, Shell small- to medium-sized, anomphalous, high-spired, up to 12 whorls present. Whorl pro- file rounded overall, slightly more convex on lower whorl face; periphery at midwhorl. Suture impressed with subsutural face bordered below by weak spiral thread. Midwhorl face dominated by 3 strong spiral cords the middle of which is slightly weaker. Between the uppermost of the strong peripheral cords and the weak cord below

GASTROPODS FROM THE BURDEKIN FORMATION 47

FIG. 5. A-F. Palaeozygopleura machenryi sp. nov., all SEM images. A, Paratype QMF35532, x 19; B, QMF35533 x 17; C, QMF35534, x 21; D, QMF35535, x 19; E, QMF35538, x 15; F, Holotype QMF35531, x 17. G. Palaeozygopleura sp. QMF35519, x 15. H-K, Trinema heideckeri gen. et sp. nov. H, Paratype QMF34941, x 4.3; I, Holotype QMF34954, side view x 5; J, Paratype QMF34955, x 2.5; K, Paratype. L, Paratype QMF34945, small specimen, x 19 SEM image. L, Trinema sp. QMF34931, x 4,3. M, ?Subulites (Subulites) sp. QMF34934, x 4.2. N, ?Subulties (Fusispira) sp. A. QMF34912, x 6.1. O, P. ?Subulties ( Fusispira) sp. B, QMF35517, x 9, x 17 respectively.

48 MEMOIRS OF THE QUEENSLAND MUSEUM

the suture lies the trace of the deep sinus, indi- cated by the growth lines, Three weak spiral cords are present on the lower whorl face. Growth lines fine, numerous, prosocline above the sinus and opisthocline below. Whorls embrace just below the uppermost fine thread on the lower whorl face.

REMARKS. Trinema heideckeri is superficially resemblant of Murchisonia species described below, bul is clearly distinguished by the 3 mid- whorl cords and the placement of the sinus. Trinema sp. below is differentiated by the posi- tion of the suture.

Trinema sp. (Fig, 5L)

MATERIAL EXAMINED. QMF34931.

DESCRIPTION. Single fragmentary specimen, high-spired; sulure impressed, whorl face domi- nated by three prominent spiral cords; whorls embrace at lowermost of these; weak spiral thread on uppermost whorl face. Sinus positioned above the highest thick cord. Growth lines fine and numerous.

REMARKS. The specimen is distinct from Trinema heideckeri sp. nov. in that the whorls embrace higher on the whorl face, at the level of the lower of the three major midwhorl cords. Whether this 1s an abberrant form or a new taxon is unclear.

Superfamily SUBULITOIDEA Lindstrom 1884 Family SUBULTIDAE Lindstrom 1884

Subulites Emmons, 1842

?Subulites (Subulites) sp. (Fig. 5M)

MATERIAL EXAMINED. QMF34934.

DESCRIPTION. Small, high-spired, fusiform shell, | 1.3mm high, 4.2mm maximum width, su- tures slightly impressed; sutural angle moderate. Whorl profile gently curved. Weak, fine, prosocl- ine, growth lines. Aperture unknown, but shell form suggests presence of anterior notch; inner lip unknown.

REMARKS. Questionable assignment is made on the basis of the general shell form, but it cannot be conclusive without data on the columella and the aperture.

?Subulites (Fusispira) sp. A. (Fig. 5N)

MATERIAL EXAMINED. QMF34912.

DESCRIPTION, Fusiform, small, rapidly ex- panding shell, approximately 4mm high and 2mm wide. Whorls embrace just below periphery which is situated just below midwhorl. Whorl profile smoothly rounded. Growth lines, aperture and inner lip unknown.

REMARKS. Lack of material prevents accurate assignment of this taxon. It differs from the ma- terial described below in having no subsutural angulation,

?Subulites (Fusispira) sp. B. (Fig. 50,P)

MATERIAL EXAMINED. QMF35517, QMF35518.

DESCRIPTION. Small, fusiform, maximum height 4mm, width 2.2mm. Whorl profile rounded except for prominent subsuturål surface and angualtion high on upper whorl face. Sutures impressed, whorls embrace just below periphery which is slightly below the midwhorl.

REMARKS. Inadequate material and poor pres- ervation of lip, ornament and aperture prevents assignment of this material. It differs trom ?Sub- ulites ( Fusispira) sp. A. in having a prominent subsutural surface and angulation.

FAUNAL AFFINITIES

Presence of species of Murchisonia, Bellerophon and Palaeozygopleura, Euryzone and Anomphalus demonstrates Old World Realm affinities, but these are at generic level only. It is unclear whether there are links to Chinese and Russian faunas of this age, as they have been so sparingly described, and microgastropods of this age are little-recorded in the literature. There is à general taxonomic continuance with respect to Early Devonian (Emsian) faunas described by Tassell (1982 ), but this is only on a generic level with the genera represented in both being long- ranging and widespread. Thus there is à strong endemism to the fauna, even when compared to Giveuan gastropods from the adjacent Broken River Province (Cook & Camilleri, 1997), with no common species apart from Burdikinia burdekinensis and Labrocuspis nodosa, This may be a function of the strongly embayed palaco- geographic setting of the Burdekin Subprovince,

GASTROPODS FROM THE BURDEKIN FORMATION 49

in comparison to the open marine conditions of

the Broken River Province during the Givetian.

LITERATURE CITED

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BLODGETT, R.B. 1988. Wiscensonella, à new raphiostomatinid gastropod genus from the Mid- dle Devonian of Wisconson. Journal of Paleontol- ogy 6203); 442-444,

BLODGETT, R.B. & JOHNSON, J.G. 1992, Early Middle Devonian (Bifelian) gastropods of Central Nevada. Paliiontographiea Abt A. 222 (4-6): 85- 134,

BLODGETT, RR. ROHR, D.M. & BOUCOT, AJ, 1990. Early and Middle Devonian gastropod hio- geography. Pp. 277-284. In MeKerrow. WS & Scotese, C.R. (eds) Palaeozoic palacogeography and biogeography. Geological Society Memoir 12. (The Geological Society: London).

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ETHERIDGE, R. 1917. Descriptions of some Queens- land Palaeozoic and Mesozoic fossils. 3. A re- markable univalve from the Devonian limestone of the Burdekin River, Queensland. Geological Survey of Queensland Publication 260: 13-16.

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HENDERSON, R.A. 1980. Structural outline and sum- mary geological history for northeastem Aus- tralia, Pp. 1-26, In Henderson, R. A. & Stephenson, P. J, (eds) The geology and geophys- ics of northeastern Australia. (Geological Society of Australia, Queensland Division, Brisbane),

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TASSELL, C.B. 1982. Gastropods from (he Early Devonian "Receptaculites" Limestone, Taemas, New South Wales. Records of the Queen Victoria Museum Launceston 77.

WHIDBORNE, GF. 1889-92. A monograph of the Devonian fauna of the South of England, Volume 1, the fauna of the limestone of Lummuton, Wolborough, Chircombe Bridge and Chudleigh. (Palaeontographical Society, London).

WYATT. DH & JELL, 1.5. 1967. Devonian ol the Townsville Hinterland, Queensland, Australia, Pp. 99.105. In Oswald, D.H. (ed.) International symposium on lhe Devonian System, Calgary, 1967, (Alberta Society of Petroleum Geologists: Alberta).

ZHEN. Y.Y. 199]. Devonian rugose coral faunas and biostratigraphy of the Fanning River Group, north Queensland. Ph.D. Thesis (Unpubl.). University of Queensland.

1994, Givetian rugose corals from the northern mar- gin of the Burdekim Basin, north Queensland. Alcheringa 18: 301-343.

50 MEMOIRS OF THE QUEENSLAND MUSEUM

PROBABLE PREDATION SCAR ON A DEVONIAN BRACHIOPOD. Memoirs of the Queensland Museum 42(1): 50. 1997:- A silicified pedicle valve of the brachiopod Warrenella sp. was recovered from limestone samples of the Burdekin Formation by acetic acid dissolution. The specimen derived from Queensland Museum Locality QML 1094, near Little Rocks, Fletcherview Station, and is registered in the Queensland Museum Palaeontology collection QMF35128. Stratigraphy and data concerning the Givetian age of the Burdekin Formation are outlined by Cook (1995),

The valve (Fig. 1) bears a small depression fracture near the right anterior margin, 9mm wide and approximately 2mm deep. This fracture bears a strong resemblance to scars docu- mented in Carboniferous brachiopods by Alexander (1981) and Gutteridge (1989). Alexander (1981) attributes such crushed valves to seized but uningested individuals being grazed upon by sharks. Gutteridge (1989) similarly attributed such scars to predation by sharks,

Given the minimal amounts of vertebrate fossil material recovered from the entire Burdekin Formation (only four incomplete and unidentifiable bone fragments), it is unclear what likely predators could have been responsible for this scar.

Literature Cited

ALEXANDER, R.R. 1981. Predation scars preserved in Chesterian brachiopods: probable culprits and evolu- tionary consequences for the articulates. Journal of Paleontology 55(1): 192-203.

COOK, A.G. 1995. Sedimentology and depositional environ- ments of the Middle Devonian Big Bend Arkose and Burdekin Formation, Fanning River Group, Burdekin

FIG. 1. Warrenella sp., QMF35128, arrow indicates probable predation scar, x 1,3. Specimen blackened with colloidal graphite and whitened with ammonium chloride for photog- raphy.

Subprovince, north Queensland, Australia. Memoirs of the Queensland Museum 38(1): 53-91. GUTTERIDGE, P. 1989. Shark predaüon on Dinantian brachiopods from the Derbyshire Dome. Mercian Ge- ologist 11 (4): 237-244. A.G. Cook, Queensland Museum, PO Box 3300, South Bris- bane, Queensland 4101; 23 December 1996.

REVIEW OF FOSSIL GASTROPODS BURDIKINIA KNIGHT 1937 AND AMPHELISSA ETHERIDGE 1921

ALEX G. COOK

Cook, A.G, 1997 06 30: A review of the gastropod Burdikinia Knight 1937 and Amphelissa Etheridge, 1921. Memoirs of the Queensland Museum 4211): 51-54, Brisbane. ISSN 0079-

8835.

Burdikinia is here placed within the Helicolomidae. Burdikinia aviønoides (Etheridge), from the Middle Devonian of New South Wales is formally included in the genus. Austerum Heidecker 15 å junior synonym of Amphelissa Etheridge. [ ] Gastropods, Devonian, Helicotomidae.

Alex G. Cook. Queensland Museum. P.O. Box 3300 South Brisbane Queensland 4101,

Australia; | May 1997.

The Middle Devonian gastropod Burdikinia Knight (1937), with type species B. burdekinensis (Etheridge, 1917) is a distinctive, robust, nodose taxon regarded as endemic to northeastern Aus- tralia, Erected by Knight (1937) to correct the original assignment of the type species to Poly- amma, the generic name was a corruption or misspelling of *Burdekin' either River or Lime- sione. Despite several studies and reports of Burdikinia burdekinensis including taxonomy (Etheridge, 1917; Knight, 1937; Knight, 1941; Heidecker, 1959; Knight et al., 1960) and ecology (Cook, 1993) there has been no conclusion reached on its higher taxonomic placement. partly due to the almost ubiquitous poor preser- vation of Burdikinia specimens, Thus the genus has been relegated to the genera inquirenda (Knight et al., 1960). Discovery of a moderately well-preserved specimen of B. burdekinensis fi- nally allows this enigmatic genus to be assigned to a family.

The posthumous publication of Etheridge (1921) of 2 gastropods from the Timor Lime- stone, Upper Hunter Valley, NSW has been ig- nored by most studies of Devonian gastropods. Etheridge (1921) erected Polyamma axionoides on the basis of 2 worn specimens forwarded to him by the Geological Survey of New South Wales. The only subsequent reference to this taxon is that of Pedder, Jackson & Ellenor (1970) who listed the taxon, assigning the species to 'Burdekinia' (sic).

Etheridge (1921) also erected Amphelissa isisensis, Which Pedder, Jackson & Ellenor (1970) listed as Euomphalus isisensis. Austerum Heidecker bears striking similarity lo Amphelissa and is synonymised.

SYSTEMATIC PALAEONTOLOGY

Superfamily EUOMPHALIDAE de Koninck, 1881 Family HELICOTOMIDAE Wenz, 1938

Burdikinia Knight 1937

TYPE SPECIES. Polyamma burdekinensis Etheridge 1917 hy original designation, from the Middle Devon- ian (Givetian) Fanning River Group, probably lower Burdekin Formation or Big Bend Arkose.

SPECIES INCLUDED. Burdikinia burdekinesis (Etheridge, 1917), Middle Devonian (Givetian), north Queensland; Burdikinia axionoides (Etheridge. 1921) from the Middle Devonian (Givetian) Timor Limestone, New South Wales.

DISTRIBUTION. Burdekin Subprovince. N Queensland; Broken River Province, N Queens- land; Timor Area, Upper Hunter, New South Wales.

REMARKS, Poor preservation has been the sin- gle contributing factor to the lack of understand- ing of the systematic position of Burdikinia. Burdikinia burdekinensis occupied high-energy, nearshore environmments (Cook, 1993). Conse- quent abrasion of shell material obliterated all but gross shell detail. Subsequent recrystallisation of much of the available material also obscured taxonomic features, such as the position of the sinus (if any) or selenizone (if any). In addition specimens are usually not conducive to prepara- tion due to the lack of mechanical (and chemical) difference between surrounding matrix and the recrystallised shell.

Burdikinia axionoides (Etheridge, 1921) differs from the type species, being more squat, and lacking the prominent cords on the lower whorl face. The lectotype of B. axionoides, (Fig. 1D,E) here designated as MMF16014, shows the sinus 10 be on the flattened upper whorl face, between

52 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. A-C, Burdikinia burdekinesis (Etheridge), QMF35452. A, apical view, arrow shows sinus position, X1; B, side view, X1; C, basal view. D-E. Burdikinia axionoides (Etheridge), Lectotype MMF16014 X1. D, apical

view, arrow shows sinus; E, side view.

the suture and the nodose cord. The only other specimen of the original suite examined by Etheridge (1921), MMF16015 is poorly preserved and lacks any particular diagnostic features.

Burdikinia is here placed in the Helicotomidae because of the dominance of spiral ornament, and particularly the prominent nodose carina border- ing the flattened upper whorl surface, and the presence of the shallow sinus on the upper whorl surface between the suture and the bordering carina. The anomphalous base is unlike other Helicotomidae, but reflects great shell thickening as a probable ecological adaptation.

Burdikinia burdekinensis (Etheridge, 1917) (Fig. 1A-C)

MATERIAL EXAMINED. HOLOTYPE: GSQF926. NEW MATERIAL: QMF35452 from Big Bend, near Charters Towers, NQ.

ADDITIONAL DESCRIPTION. See also Etheridge (1917) and Heidecker (1959). Growth lines are fine and numerous; prosocline from the uppermost and peripheral spiral cord, strongly recurved about the base towards the axis; these lines are uninterrupted across the lower whorl surface. The weak sinus is located on the upper whorl face regarded here as a subsutural ramp. The nodes on the spiral cords are solid (as noted by Heidecker) in deference to Knight et al. (1960).

REMARKS. The new material places the sinus

accurately and refutes any presence of a selenizone.

Amphelissa Etheridge 1921

Amphelissa Etheridge 1921: 2; Knight 1941: 34. Austerum Heidecker 1959: 6.

REVIEW OF FOSSIL GASTROPODS 53

FIG. 2. Amphelissa isisensis Etheridge. A-C, Holotype MMF16011, X1, apical, basal and apertural views repsectively. D-E, MMF16024, x1. D, apical view; E, side view, note fine growth lines.

TYPE SPECIES. By monotypy. Amphelissa isisensis Etheridge 1921 from the Middle Devonian Timor Limestone, NSW.

OTHER SPECIES INCLUDED. Amphelissa carinatum (Heidecker) 1959, from the Middle Devonian Burdekin Formation, north Queens- land.

REMARKS. Amphelissa presented problems to Knight (1941) who was unable to access type materials, due to their misplacement. Conse- quently, and due to the poor quality of the original illustrations, Knight (1941) regarded the genus as probably widely phaneromphalous. On the holo- type of A. isisensis (Fig 2A-C) the base is flat- tened, and the umbilicus is very shallow, wide and

partially plugged by shell, thus hemiomphalus. This is an identical condition to the base of the holotype of Austerum carinatum Heidecker. In A. isisensis there are well-preserved, fine, numerous growth lines (Fig. 2D,E), which are not preserved on specimens of the north Queensland species. Heidecker (1959) may have been unaware of the taxon described by Etheridge (1921) from the Timor Limestone which has gone largely ignored. Amphelissa was suppressed by Knight et al. (1960), who synonymised it with Srraparollus (Euomphalus). Amphelissa Etheridge cannot be accommodated in Straparollus (Euomphalus) given the hemiomphalus condition. Amphelissa is thus resurrected as a valid genus. Amphelissa carinatum (Heidecker) differs from the type in

54 MEMOIRS OF THE QUEENSLAND MUSEUM

being higher-spired. The holotype of Amphelissa isisensis, designated by Knight (1941), is regis- tered in the Geological Survey of New South Wales Collection as MMF16011. The paratypes designated by Knight (1941) are registered under MMF16012.

ACKNOWLEDGEMENTS

I am grateful to Dr Ian Percival for loan of material from the Geological Survey of New South Wales collections, to Robert Blodgett for suggestions to the manuscript and to Rae Sheri- dan for allowing the specimen to leave the edu- cational collections under her control.

LITERATURE CITED

COOK, A.G. 1993 Fletcherviewia septata, a new, high- spired gastropod from the Devonian of north SE Journal of Paleontology 67 (5): 816-

1.

ETHERIDGE, R. 1917, Descriptions of some Queens- land Palaeozoic and Mesozoic fossils. 3. A re- markable univalve from the Devonian limestone

of the Burdekin River, Queensland. Geological Survey of Queensland Publication 260: 13-16. 1921. Palaeontologia Novae Cambriae Meridionalis- Occasional descriptions of New South Wales fossils, No. 8. Records of the Geo- logical Survey of New South Wales 10 (1): 1-11.

HEIDECKER, E. 1959. Middle Devonian molluscs from the Burdekin Formation of North Queens- land. University of Queensland Papers, Depart- ment of Geology 5(2): 1-11.

KNIGHT, J.B. 1937. Genotype designations and new names for invalid homonyms among Paleozoic gastropod genera. Journal of Paleontology 11:709-714.

1941. Paleozoic gastropod genotypes. Geological Society of America Special Papers 32.

KNIGHT, J.B., COX, L.R., KEEN, A.M., BATTEN, R.L., YOCHELSON, E.L. & ROBERTSON, R. 1960. Systematic Descriptions. Pp. 1169-1331. In Moore, R.C. (ed.) Treatise on invertebrate paleon- tology, Part I. Mollusca 1. (Geological Society of America and University of Kansas Press: Law- rence).

PEDDER, A.E.H., JACKSON, J.H., & ELLENOR, D.W. 1970. An interim account of the Middle Devonian Timor Limestone of northeastem New South Wales. Proceedings of the Linnean Society of New South Wales 94 (3): 242-272.

MIDDLE DEVONIAN GASTROPODS FROM THE BROKEN RIVER PROVINCE,

NORTH QUEENSLAND ALEX G, CODK AND NATALIE CAMILLERI

Cook, A.G. & Camilleri, N, 1997 06 30: Middle Devonian gastropods from the Broken River Province, north Queensland. Memoirs of the Queensland Museum. 4201): 55-79. Brisbane. ISSN 0078-8835.

Twenty seven taxa of gastropods are described from the Eifelian and Givetian sequences of the Broken River Province, north Queensland with four new genera and ten new species. New taxa are Denayella lomandraensis sp. nov., Gyronema simpsoni sp. nov.. Frillbeastia queenslandicus gen, el sp. nov. Brokenriveria pharlapensis gen. et sp.nov., Gemininodosa langi gen. et sp. nov., Murchisonia (Murchisonia) wandovalensis sp. nov., M. (M.) lawlessi sp. nov, Palaeozygopleura dodgeyi sp. nov., Australoxa tasselli gen. et sp. nov. and Leptogyma queenslandicus sp. nov. Three gastropod communities are recognised in the Givetian rocks of the province, the Murchisonia community, inhabiting a biostromal envi- ronment, the Labrocuspis community inhabiting high-energy coarse siliciclastie environ- mens and the Brokenriveria community inhabiting an open, muddy, carbonate shelf, [ ] Gasrropods, Givetian, Eifelian, Broken River Province, Queensland.

Alex G Cook & Natalie Camilleri. Queensland Museum, PO Box 3300 South Brisbune,

410], Australia; I April 1997.

The Broken River Province. located approxi- mately 200km W of Townsville, north Queens- land consists. of two subprovinces; Graveyard Creek Subprovince and Camel Creek Sub- province (Withnall & Lang, 1993). The Grave- yard Creck Subprovince contains inter alia, a thick, widely-outcropping sequence of Silurian to Middle Devonian, dominantly shallow marine units.

Middle Devonian sequences within the Grave- yard Creek Subprovince contain diverse fossil assemblages which have been the subject of in- tense taxonomic and biostratigraphic study (Jell etal, 1993), Detailed studies of conodonts (Maw- son, 1987; Mawson & Talent,1989; Mawson ct al., 1988), fish remains (Turner, 1982, 1995; de Pomeroy, 1994) and corals (Wyatt & Jell, 1967) have provided a substantial biostratigraphic database for tàxonomic studies within the prov- ince. We follow the summary biostratigraphic scheme presented by Jell et al. (1993).

Until now gastropods have remained unstudied within the Broken River Province. Previous work on Devonian gastropods from north Queensland åre Etheridge (1917), Heidecker (1959), and Cook (1993, 1995). Gastropod faunas in south- castern Australia have received substantially more attention, more recently through the works of Tassell (1976, 1977, 1978, 1980, 1982).

This paper is concemed with gastropods col- lected from Eifelian and Givetian units of the Broken River Group, namely the Burges Forma- pon, Dosey Limestone and. Papilio Mudstone,

Material collected over a number of years by staff at The University of Queensland, Macquarie Uni- versity and the Queensland Museum contained å small number of gastropods scattered over å large number of localities in the Broken River Prov- ince. Subsequent larger collections made by the authors in 1994 and 1995 have revealed å more diverse assemblage. All material is deposited in ihe Queensland Museum (prefix QMF), and lo

calities are presented in the appendix (QML).

List of Localities. QML541; ‘Calceola’ stop 6. 2nd creek upstream from Broken River Gorge. Broken River Province. Burges Formation. Eitel- ian. Collected P.A. Jell. QML1016: Hill above crossing, S side of river near 'PharLap' prospect, Broken River, near old GSQ camp site. 19*28' S, 144743" E. Papilio Mudstone, Givetian. Collected A. Cook, N, Camilleri. QML 1018: Low rise, Ikm S of Storm Dam, 200m W of road at [9732 92'5, 144°40.51'E. Papilio Mudstone, Givetian. Col- lected A, Cook, N. Camilleri. QML1019: Ridge of silicified (silerete)- replaced Dosey Limestone, 100m north of road 500m E of Storm Dam 190. QML1083. Nuggery Gully, E of Gorge-PharLap Rd, basal Papilio Mudstone, just above and adjacent to top of N most Dosey Limestone, 19°27.79'S, 144°44.86'B. Papilio Mudsiume, Giverian. Collected A, Cook, N. Camilleri, P Lawless, D. Case, S. Dodgey-Hocknull, P. Dodgey-Tierney, R, Lootsma, 1995: QML 1090. Above 'PharLàap' crossing on Broken River, N

side of River on washed oui track approx. 40m

NE of QML1016, 19998,718, 144744.05' E,

56 MEMOIRS OF THE QUEENSLAND MUSEUM

Papilio Mudstone, Givetian. Collected QM Party 1995. QML1092: 500m NW of type section creek for Dosey & Lomandra Limestones. 150m above Dosey Limestone. 19°29.98’S, 144?43.87'E. Papilio Mudstone, Givetian. Collected QM Party July 1995. A. Cook, P. Lawless, C. McHenry, September 1995. SD21: E side of Dosey Syncline in tributary of Dosey Creek. (see Mawson & Talent, 1989; 221) Papilio Mudstone, Givetian. SD43e: Storm Dam area, Papilio Mudstone, Givetian. SD108: Approx 1.5km NE of Spanner Hill, Papilio Mudstone or Lower Mytton Forma- tion. (see Mawson & Talent 1989: 211) Givetian.

GASTROPOD ASSEMBLAGES

Three Givetian gastropod communities are recognised; | Labrocuspis community, Brokenriveria community and Murchisonia com- munity (Table 1), corresponding to differing sed- imentologic regimes. The Labrocuspis community is found in conglomerates and coarse sandstones interpreted as high energy facies at the basal Papilio Mudstone near ‘Nuggetty Gully’ area and from similar sandstones in the upper- most Papilio Mudstone or lowermost Mytton Formation in the ‘Spanner Hill’ area. This occur- rence is strikingly similar to the Burdikinia- Labrocuspis-Austerum faunule noted by Cook (1993), which occurs in high energy nearshore facies ofthe Big Bend Arkose (Givetian, Burdein Subprovince).

Brokenriveria community occurs within car- bonate mudstone facies of the Papilio Formation, interpreted as shallow, low energy, open marine, impure carbonate shelf. Murchisonia community is derived from within the uppermost Dosey Limestone and is associated with corals, stromatoporoids, sponges from a biostromal en- vironment.

Eifelian gastropods are insufficiently repre- sented in the collected fauna to comment on their commmunity arrangement.

FAUNAL AFFINITIES

Comment has already been made on the simi- larity of Labrocuspis community in the Burdekin Subprovince and the Broken River Province. In both are also found the bivalve Tanaodon louderbacki, which is also known from the Givet- ian of Guangxi (Pojeta et al., 1985). The presence of Murchisonia species, Soleniscus, palaeo- zygopleurids and Plaryceras suggests affinity with Old World Realm faunas, but sufficiently removed to develop distinct endemism at the

TABLE 1. Constituent taxa of Givetian gastropod com- munities. r = rare, c = common, a = abundant, s = single occurrence.

——— ===>

Species

Brokenriveria

Murchisonia community Labrocuspis community community

wn

Bellerophon (Bellerophon) sp. A. Bellerophon (Bellerophon) sp. B. 5

Straparollus (Straparollus) sp. 8

Straparollus (Euomphalus) sp. A r Straparollus (Euomphalus) sp. B E Labrocuspis nodosa Heidecker | c Omphaoltrochid indet. E

Denayella lomandraensis sp. nov. r

Frillbeastia queenslandicus | gen. et. sp. nov.

Brokenriveria pharlapensis sp. nov. a

Gemininodosa langi gen. et sp. nov. a

Platyceras(Platyceras) sp. 5

| platyceratoid indet. 8 Burdikinia burdekinensis (Etheridge) r

Murchisonia ( Murchisonia) wandovalensis sp. nov.

Murchisonia (Murchisonia) lawlessi sp. nov.

Muchisonia (M.) sp. cf. M. (M.) fermioni Tassell

Murchisonia (M.) sp. s

murchisoniid indet. r

Stylonema? sp. s

Australoxa tasselli gen, et sp. nov. r

Palaeozygopleura dodgeyi sp. nov. r

Soleniscus sp. cf. S. subcostata Schlotheim

` Leptogyma queenslandicus sp. nov. c | Mitchellia striatula de Koninck s

species level. There are, however closely related species of Brokenriveria gen. nov. and possibly Frillbeastia gen. nov. known from Germany as argued below. There are other generic level affin- ities with Eifelian faunas in Nevada (Blodgett, 1992) and Alaska (Blodgett & Johnston, 1992) and Givetian faunas in Yunnan (Mansuy, 1912), Guangxi (Wei & Pan, 1988). Taxonomic hang- overs from southeastern Australian faunas of Emsian age are evidenced by the co-occurence of Mitchellia, Leptogyma, and Murchisonia (M. ) sp. cf. M. (M.) fermioni Tassell. A dearth of taxo- nomic work on southeast Asian and Russian gas- tropods of Middle Devonian age makes conclusive assessment of the taxonomic affinities impossible.

GASTROPODS FROM THE BROKEN RIVER PROVINCE 57

SYSTEMATIC PALAEONTOLOGY

Class GASTROPODA Cuvier, 1797 Order ARCHAEOGASTROPODA Thiele, 1925 Superfamily BELLEROPHONTOIDEA M'Coy, 1851 Family BELLEROPHONTIDAE M'Coy, 1851 Bellerophon Montfort, 1808 Belleropohon (Bellerophon) Montfort, 1808

Bellerophon (Bellerophon) sp. A. (Fig. LA, B)

MATERIAL EXAMINED. QMF32642 from SD21, QMF32644 trom QML1018.

DESCRIPTION. Medium-sized, isostrophic shell, up to 20mm wide and 20mm diameter; doubly phaneromphalous. Whorl profile gently rounded, with a weak ridge mid-whorl, Smooth, fine, growth lines, nearly straight; shell rapidly expanding: lip and aperture unknown.

REMARKS. Poor preservation of this material does not allow confident specific assignment to ane of the many species of Bellerophon (Bellerophon). It is assigned to this subgenus on the basis of the rounded whorl profile, absence of spiral ornament and simplicity of growth lines.

Bellerophon (Bellerophon) sp. B (Fig. IC, D)

MATERIAL EXAMINED. QMF33621 from QML1092.

DESCRIPTION. Small, isostrophic shell, 10mm wide, 10mm diameter; rapidly expanding. aper- ture flared. Selenizone, prominent upon weak ridge. No growth lines preserved.

REMARKS. The single, poorly preserved speci- men cannot be assigned to a species, It is more rapidly expanding ànd smaller than Bellerophon (Belleraphon) sp. A.

Superfamily EUOMPHALOIDEA de Koninck, 1881 Family EUOMPHALIDAE de Koninck, 1881 Straparollus de Montfort, 1810 Straparollus (Straparollus) de Montfort, 1810

Straparollus (Straparollus) sp. (Fig. 1E, F)

MATERIAL EXAMINED. QMF33103 from QML1019,

DESCRIPTION, Medium-sized, trochifrom shell, approximately 5.5mm high, 10mm wide; apical angle approximately 110°, sutures strongly impressed. Whorl profile strongly rounded with slight shoulder. Ornament consists of numerous fine growth lines, slightly prosocline. There is à very slight flexure in the growth lines on the weak shoulder suggesting the sinus. Just below mid- whorl near the aperture, there is an apparent shell repair.

REMARKS. The specimen is confidently as- signed to the subgenus on the basis of the whorl profile, overall shape and ornament. The number of species of this subgenus is large and confident assigment 10 species is unwise due to the lack of material, and the degree of variation present in known taxa (Linsley & Yochelson, 1973). The specimen differs from S.($.) ater (Spitz) of Jhaveri (1969) from the lower Devonian ol the Carnic Alps, which is much more squat. 5. (S\) kokeni (Spitz) of Jhaveri (1969) has a much more prominent shoulder. Of the taxa described by Linsley & Yochelson (1973), S. (5.) laevis (Archiac E Vemeuil), from the Middle Devonian of Germany, is more flattened, but their S. (S.) ?laevis is more resemblant of the Broken River specimen. S.(S.) cyclostomus (Hall) of Linsley & Yochelson (1973) from the Middle Devonian of North America has a more promi- nent sinus.

Straparollus (Euomphalus) Sowerby, 1814

Straparollus (Euomphalus) sp. A (Fig. 1G)

MATERIAL EXAMINED. QMF33104, QMF33352 from L1019.

DESCRIPTION. Small, planispiral shell up 10 12mm in diameter, 4mm maximum height. Both specimens are preserved as basal moulds. Whorl profile subrounded to subquadrate, with a weak angulation between the midwhorl and basal sur- faces obvious in the last whorl. No obvious growth lines preserved, but there is a faint hint of fine collabral lines on QMF33104. Protoconch unknown.

REMARKS. This material cannot be identifed further due 10 inadequate preservation.

58 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. A,B, Bellerophon (Bellerophon) sp. A., QMF32642 x 2.2. A, side view; B, apertural view. C,D, Bellerophon (Bellerophon) sp. B, QMF33621 x 2.7. C, side view; D, view of selenizone. E,F, Straparollus (Straparollus) sp., latex moulds from QMF34267 x 3.1. E, apertural view; F, apical view. G, Straparollus (Euomphalus) sp. A, QMF33352 x 3.1, basal view. H, Straparollus (Euomphalus) sp. B, QMF34752 x 2.2. All specimens (and those on subsequent figures) whitened with ammonium chloride for photography.

GASTROPODS FROM THE BROKEN RIVER PROVINCE 59

Straparollus (Euomphalus) sp. B. (Fig. 1H)

MATERIAL EXAMINED. QMF34752 from QMLI1018.

DESCRIPTION. Large, planispiral, 42mm in di- ameter; 4 whorls preserved in cross section and external mould, the first 2 preserved whorls are septate; septa gently concave; collabral growth lines preserved on the final whorl are fine and numerous.

REMARKS. The planispiral form, simplicity of growth lines and the septation identifies the sub- genus, but a species cannot be assigned.

Family OMPHALOTROCHIDAE Knight, 1945

Labrocuspis Heidecker,1959 Labrocuspis Heidecker, 1959: 6; Kase,1989:149.

TYPE SPECIES. Labrocuspis nodosa Heidecker, 1959, by original designation, from the Middle Devon- ian (?late Eifelian- Givetian), Big Bend Arkose, north Queensland.

DIAGNOSIS. Large, anomphalous trochiform gastropod; suture deep and impressed, whorl pro- file rises from the suture to a keel and descends to a variably developed peripheral buttress.

REMARKS. Labrocuspis is presently restricted to two taxa; the type and L. kobayashii (Kase & Nishida, 1988), from the Eifelian Nakazato For- mation, north east Japan. Kase (1989) assigned the genus to Omphalotrochidae. Heidecker (1959) remarked on the differential development of the peipheral buttress, it only being fully ex- pressed in larger forms.

Occurrences of the genus recorded by Heidec- ker (1959), Kase (1989) and Cook (1993) high- light association of this genus with neashore to shoreline, often high energy siliciclastic deposi- tion. Cook (1993) suggested strong ecological control for the genus within the Big Bend Arkose and Burdekin Formation. Material here described is from coarse-grained siliciclastic units, includ- ing conglomerates and very coarse to granular sandstones which display cross and planar lami- nation. Thus a high-energy shallow water envi- ronment is suggested.

Labrocuspis nodosa Heidecker, 1959. (Fig. 2A-C)

Labrocuspis nodosa Heidecker, 1959: 6, Pl. 1 fig. 2a-d, Pl. 3, fig. 2a-b.

MATERIAL EXAMINED. QMF16547, QMF33316 locality imprecise, Broken River Province, collected M. Wade. QMF33315 from SD108, Upper Papilio Formation or Mytton Formation, collected J. Jell. QMF33582 from QML1089; QMF33580, QMF33581, QMF33583 from QML1083. There is some doubt as to the exact origin of specimens QMF16547, QMF33315 and QMF33316. All were collected at the same time, with J. Jell (pers. comm.) recording the origin as Locality SD108, within the Papilio Formation. However the coarser grained lithol- ogy would suggest the nearby outcrops of Mytton Formation.

DISTRIBUTION. Middle Devonian (?late Eifel- ian- Givetian), north Queensland. The species is apparently endemic to the region. Very poorly preserved material from the Laroona Formation, Burdekin Subprovince may belong to this taxon, which would extend the age range to the Ems- ian.

DESCRIPTION. Moderately large, thick- shelled, dextrally-coiled, trochiform gastropod up to 63mm high and 36mm wide, with an apical angle up to 120°; suture deep and impressed; sutural slope approximately 10°. Whorl profile rises sharply from the suture to a gently rounded carina, and slopes to a very gently convex profile. Upper whorl profile is gently convex, but breaks midwhorl to produce a gently concave profile below the midwhorl producing a buttress on the lowermost surface. The carina is conspicuously nodose particularly in final whorls. Base flat, with a wide callus pad. Aperture quadrate. Growth lines preserved on base, none preserved on whorl surface.

REMARKS. Labrocuspis nodosa was only pre- viously recorded in the nearby Burdekin Sub- province, Givetian, north Queensland (Heidecker, 1959). The Broken River material is relatively poorly preserved but is inseparable from L. nodosa Heidecker from the Big Bend Arkose and Burdekin Formation, having the no- dose carina, variable development of the buttress, and strongly developed callus pad. It differs from L. kobayashii (Kase & Nishida) in the degree of sutural impression, the nodose carina, and hence the upper whorl profile.

60 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG.2. A-C, Labrocuspis nodosa Heidecker, 1959, QMF33583 x 1, apertural, apical and basal views respectively. D, Omphalotrochid indet., latex mould from QMF33687 x 2, apical view. EF, Burdkininia burdekininnsis (Etheridge, 1917), QMF33579 x 2. E, apertural view, F, apical view,

GASTROPODS FROM THE BROKEN RIVER PROVINCE GI

Omphalotrochid indet, (Fig. 2D)

MATERIAL EXAMINED. QMF330687 from QML1019.

DESCRIPTION. Mould of an upper surface of å large, shallowly trochiform shell, 25mm in diam- eter. Apical angle apporximately 140°, Suture deep, impressed forming a channel in lust whorl. Growth lines fine, collabral.

REMARKS, The specimen superficially resem- bles Orecopia murrayi Tassell, 1978 from the Lower Devonian Bell Point Limestone, Victoria, but cannot be confidently assigned to a genus as no basal structures are known. The specimen lacks the nodose carina of Labrocuspis nodosa Heidecker. The specimen is left in open nomen- clature pending more material.

Superfamily PLEUROTOMARIOIDEA Swainson, 1840 Family RAPHISTOMATIDAE Koken, 1896 Subfamily RAPHISTOMATINAE Koken, 1896

Denayella Blodgett & Johnson, 1992

TYPE SPECIES. Denayella housei Blodgett & John- son, 1992. from the Eifelian Denay Limestone of Ne- vada, USA,

Denayella lomandraensis sp. nov. (Fig. 3 A,B)

MATERIAL EXAMINED. HOLOTYPE: QMF33650 from QML 1092, Papilio Mudstone, Givetian, Broken River Province. PARATYPES: QMF33651-33672, QMF34215-34222 from QML 1092, QMF33619 from QML1090.

DIAGNOSIS. Small member of genus with flattenedbase and weak inductural deposit.

DESCRIPTION. Small (Table 2) lenticular, an- omphalus. Whorl profile sharply rounded, upper surface gently sloping, lower surface steeper. Pe- riphery on upper 1/3 of whorl profile. sharply rounded bearing 2 indistinct cords delimiting a probable selenizone, within which the lunulae are nol preserved. Lower cord is more obscure than the upper and on many specimens they are not preserved due to abrasion, Aperture rhomboid to subrounded, with a v-shaped sinus on the labrum. Shell thick especially at columella. There is a distinct thickening of shell on the columella. Base convex, but not sharply so. Very weak, fine, col-

TABLE 2. Measurments for Denayella lomandraensis sp. nov.

Specimen

labral growth lines, otherwise surface smooth and unornamented.

REMARKS. The material is placed in Denayella due to its raphistomid-like appearance, indistinct, but present, seleneizone and weak inductural de- posit on the columellar lip. It differs from the type species in having a more flattened base and a weaker deposit on the columellar lip. It is more similar to Denayella sp. of Blodgett & Johnson, 1992 with respect to the flatter base. The species differs from members of Arizonella Stoyanow, 1948 by its lack of à prominent selenizone, and well developed collabral ornament. It differs from Buchelia and Raphistoma due to the lack of cords on the upper whorl face. Umbotropis mesoni Tassell 1982, from the Receptacutites Limestone of New South Wales is clearly phan- eromphalus, and has more prominent spiral cords on the upper whorl face rather than the periphery.

ETYMOLOGY. For Lomandra Creek. Subfamily LIOSPIRINAE Knight. 1956

Frillbeastia gen. nov.

DIAGNOSIS. Small, trochiform with flattened base; angular periphery with selenizone bounded below by frilled carina; upper whorl face with 2 strong threads, the lowermost of which is nodose; strong prosocline ornament; planispiral pro- toconch.

TYPE SPECIES. Frillbeasria queenslandicus sp. nov.

DISTRIBUTION, Middle Devonian (Givetian), uppermost Dosey Limestone, Broken River Province, north Queensland, ?Middle Devonian (Givetian), Germany (Sandberger & Sandberger, 1850-6).

REMARKS. The genus is similar to Arcsrra Stoyanow. 1948 from the early Late Devonian in Arizona, having a distinct frill, but is less lentic- ular in shape, possesses stronger collabral orna- ment, strong spiral cords with nodes, and the

62 MEMOIRS OF THE QUEENSLAND MUSEUM

proioconch is planispiral. Superficially similar is Astralites Whiteaves, 1892 from the Middle Devonian of Canada, which lacks the prominent selenizone and spiral ornament, Members of the Luciellidae Knight, 1956 such as Luciella de Koninck, 1883 and Epiptychia Perner 1907 have the frill above the selenizone, rather than below in Liospirinae, Zalozone Linsley, 1968 possesses à double frilled selenizone, and Tylozone Linsley, 1968 has a sclenizone bordered by an upper frill. Frillbeastia is placed in Liospirinae due to closer similarity to Arastra, but is by far the most trochi- form member of the subfamily. Linsley (1968) placed Arasira within the Raphiostomatinae, with Zalozone. The base of Frillbeastia is wide and hence js retained in Liospirinae with the wide-based Arastra.

Within the genus could be included Lirrorina alata Sandberger & Sandberger, 1850-1856 from the Middle Devonian (Givetian) of Germany. The taxon illustrated (Sandberger & Sandberger, 1850-56: pl. 24: fig. 14) shows the characteristic frilled keel, a similar base, and numerous cords on the upper surface. It differs from E queenslandi- cus by having numerous equal strength cords, it lacks the strong growth lines and the nodose cord above the selenizone whichis not identifiable on the illustrations of the German taxon,

ETYMOLOGY. For the informal name given in the field, reflecting the character of the selenizone.

Frillbeastia queenslandicus sp. nov. (Fig. 3C-G)

MATERIAL EXAMINED. HOLOTYPE: QMF33687, from QML1019 uppermost Dosey Lime- stone Givetian, Broken River Province. PARATYPES: QMF33695, QMF34265 from QML 1019,

DIAGNOSIS. As for genus.

DESCRIPTION, Small, trochiform, nearly coni- cal shell, complete specimen is 16mm wide, | Imm high with a spire angle of approximately 60°. Base flattened, bearing spiral cord, narrowly phaneromphalus or anomphalous, Whorl face or- nate and angular with suture just below frill on midwhorl. Upper whorl face possesses 2 major threads, | close to the periphery, and another weaker thread on the mid-upper whorl face, These increase in strength throughout growth. The lowermost thread possesses stout rounded nodes, the upper thread hus vestigal nodes pre- served on the holotype. The angular shell periph- ery is decorated by a simple, fine spiral thread which is the upper boundary for the selenizonc.

Below the selenizone is hordered by a rhythmi- cally folded carina, thus producing a frill. The lower whorl profile possesses a single thread, slightly nodose. Collabral ornament is prosocl- ine. Protoconch nearly planispiral, 2-3 whorls with simple comarginal ornament; a vestige of a selenizone is recognisable on the 3rd whorl,

REMARKS. Despite the small number of speci- mens the material is signficantly distinct to war- rant erection of à new genus. One of the specimens has less distinct nodes and cords on the upper whorl face, and shows signs of abrasion,

ETYMOLOGY. For the state of Queensland,

Suborder TROCHINA Cox & Knight, 1960 Supertamily PLATYCERATOIDEA Hall, 1859 Family HOLOPEIDAE Wenz, 1938 Subfamily GYRONEMATINAE Knight, 1956

Gyronema Ulrich in Ulrich & Scofield, 1897

TYPE SPECIES. Trochonemu (Gyronema) pulchellum Ulrich and Scofield, 1897 from the Middle Ordovician, Minnesota, United States of America.

DIAGNOSIS. See Knight et al, (1960),

Gyronema simpsoni sp. nov. (Fig. 4C,D)

MATERIAL EXAMINED, HOLOTYPE; QMF32082 from QML541. Burges Formation, Eifelian, Broken River Province. PARATYPES; QMF32055, QMF32058 from QML541.

DIAGNOSIS, Very large Gyronema with charac- teristic thickened spiral cords, flattened upper- most whorl surface forming a low shoulder, 2 prominent cords on upper whorl surface, the highest being an angular carina, 6 prominent cords on final whorl.

DESCRIPTION, Large, turbiniform, narrowly phaneromphalus gastropod, with prominent thick spiral ornament and impressed sutures, Shell height up to 47mm, and width up to 32mm. with an apical angle of approximately 20° External whorl profile is dominated by spiral cords, the uppermost forming à prominent angular carina high on the whorl profile which is divided into two surfaces. Uppermost third of the whorl pro- file is a gently concave shoulder, which slopes shallowly from thesuture to the carina; mid whorl is nearly vertical, and the lower whorl profile gently convex with both marked by thick spiral cords. Sutures formed at midwhorl, slightly

GASTROPODS FROM THE BROKEN RIVER PROVINCE 63

above the 3rd spiral cord, thus obscuring all but 2 uppermost cords on early whorls. Aperture generally rounded, slightly vertically extended with abax- ial angulation above midwhorl. Growth lines unknown. The 2 paratypes are somewhat crushed and distorted, but retain the characteristic spiral cords.

REMARKS. Material has the typical form of the genus, but differs from the type species by the more flattened shoulder, in addition this taxon is several times larger than the genotype figured hy Knight (1941). In size and form this species is

comparable to G. bellense of

Tassell from the Early Devon- ian of Victoria but G. bellense has far more numerous spiral cords. G. lirata (Hall) of Rol- lins, Eldredge & Spiller (1971), from the Middle Devonian Marcellus Formation, New York is smaller (17mm high), with more spiral cords on the upper whorl surface, however it does possess a similar shoul- der to G. simpsoni. Gyronema scaliforme Zytlenok,1976 from the Devonian of Belorus' possesses more cords than the Broken River species. The species is close to G. or- mistoni Blodgett, 1992 from the Eifelian of Alaska, but lacks the prominent basal cords of that taxon.

ETYMOLOGY. For Andrew Simpson,

Brokenriveria gen. nov. TYPE SPECIES. Brokenriveria pharlapensis sp. nov

DIAGNOSIS. Small turbini- form gyronematid with 2 or- ders of nodose spiral cords.

FIG. 3. A.B, Denayella lamandraensis sp. nov., Holotype QMF33650 x 6. A, apertural view; B, apical view. C-G, Frillbeastia queenslandicus gen. et sp. nov. C-E, latex mould of Holotype QMF33687 x 3.6. C, oblique side view; D, basal view; E, side view. F, latex mould of Paratype QMF34265, x 3,1, side view, G, latex mould of Paratype QMF33695, x 4.5, oblique view showing protoconch.

Queensland, Middle Devonian, Rhineland (Goldfuss, 1844).

DISTRIBUTION. Middle Devonian (Givetian) REMARKS. Lack of a selenezone precludes as- Papilio Mudstone, Broken River Province, north signment of this material to grossly similar genera

64 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. A,B. Murchisoniid indet. A, latex mould of QMF33100, side view x 1.8. B, latex mould of QMF34259, oblique side view x 2.7. C, D. Gyronema simpsoni sp. nov. Holotype QMF32082, x 1.3. C, apertural view; D, side view. E-G, Platyceras (Platyceras) sp. QMF32641, x 2.7. E, apical view; F, side view; G, basal view.

Bembexia Oehlert, Nodonema Linsley, Or other Pleurotomarioidea. The pro- socline growth lines across the shell, the strong cords and the thicker shell suggest that this material can be accommodated within the Gyronematidae. The genus differs from other gyronematids in the possession Of two orders of spiral cord, rather than one on Yunnania Mansuy, and Gyronema Ul- rich. Robert Blodgett (pers. comm) has kindly drawn my attention to a similarly or- namented taxon Turbo caelatus Goldfuss, 1844 from the Middle Devonian of Ger- many, which should be in- cluded in the genus, but differs in the number of cords on the upper whorl face.

ETYMOLOGY. For the Broken River.

Brokenriveria pharlapensis sp. nov. (Fig. 5)

MATERIAL EXAMINED. HO- LOTYPE: QMF32234 from QML1016, Papilio Mudstone, Givetian, Broken River Province. PARATYPES: QMF32235- 32301 from QML1016.

DIAGNOSIS. As for genus.

DESCRIPTION. Small to me- dium-sized, turbiniform gas- tropod, up to 17mm high and wide (Table 3) with an apical angle of approximately 105°. Whorl profile rounded, but dominated by two orders of nu- merous nodose spiral cords. Suture impressed, whorls em- brace at the midwhorl. Periph- ery situated at midwhorl. Six major threads on the whorl sur- face, 1 on the upper whorl face, another at midwhorl, the re- mainder on the lower whorl face. Major cords are inter- spaced with 1 less prominent nodose spiral thread, with the exception of the uppermost

GASTROPODS FROM THE BROKEN RIVER PROVINCE 65

FIG. 5. Brokenriveria pharlapensis sp. nov. A, B, Holotype QMF32234, x 3. A, apertural view; B, apical view. C-E, paratype, QMF32268, x 2.8. C, side view. D, apical view. E, apertural view. F, QMF32273, x 2.8, apertural view. G, H. Paratype QMF32272, x 2.7. G, side view; H, apertural view. I, paratype QMF32274, apertural view x 2,8.

major thread which has 2 minor threads between it and the suture. Growth lines of 2 orders; numer- ous, fine, prosocline, slightly coarser growth lines intesect with spiral cords to produce nodes. Ap-

erture rounded, outer lip not preserved, inner lip thickened, and in some specimens reflected slightly. Shell relatively thickened for size. Pro- toconch unknown.

66 MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 3. Measurements for Brokenriveria phar- lapensis sp. nov.

Specimen height (mm) Kueder A weng angle (°) QMF32234 14.6 70 QMF32237 112 o | QMF32239 14.0 70 QMF32272 143 70 QMF33625 12.5 75 | QMF33629 13.5 65 QMF33628 13.2 75 | QMP366 | 131

REMARKS. This species differs from Gemininodosa langi sp. nov. by its lack of very large nodes on the shell periphery. The specimen of indeterminite platyceratoid described below has a more rounded whorl profile, greater expan- sion rate and less prominent spiral ornament.

ETYMOLOGY. For ‘Pharlap’ Crossing of the Broken River.

Gemininodosa gen. nov.

TYPE SPECIES. Gemininodosa langi sp. nov. from the Middle Devonian Papilio Mudstone, Broken River Province, north Queensland.

DIAGNOSIS. Small to medium-sized, turbini- form, minutely phaneromphalus, with numerous spiral cords; upper whorl surface bears 2 spiral rows of large rounded nodes.

REMARKS. The lack of a sinus or selenizone, dominance of spiral ornament and the turbini- form shape indicates placement within the Holopeidae Wenz. Superficially the genus is like Oriostoma Munier-Chalmas but does not possess the wide umbilicus. The distinct nodose ornament renders the genus unlike any other members of the family but is reminiscent of the node- (and selenizone-) bearing Pleurotomariitoidea, such as Nodonema Linsley, 1968 and Glyptomaria Knight, 1945.

ETYMOLOGY. For the twin nodes adorning the shell.

Gemininodosa langi gen. et. sp. nov. (Fig. 6A-I)

MATERIAL EXAMINED. HOLOTYPE: QMF3361 1 from QML1092, Papilio Mudstone, Givetian, Broken River Province. PARATYPES: QMF33608-33610, QMF33612-33618, from QML1092, QML33638- 33644 from QML1090.

DIAGNOSIS. As for genus.

DESCRIPTION. Medium-sized, turbiniform, minutely phaneromphalus, up to 16mm wide and 16mm high (Table 4) with an average spire angle of 75-115°. Sutures impressed, whorls embrace at midwhorl. Whorl profile overall rounded, flat- tened adjacent to the suture, rounded at midwhorl and lower whorl surface. Midwhorl dominated by 2 rows of large nodes, one at the edge of the flattened upper whorl surface the other near the periphery. A third less distinct set of nodes occurs below the midwhorl. Whorl surfaces are or- namented with many spiral cords, with at least 5 major cords on the lower whorl surface, 4 on the mid whorl, and 3 on the upper whorl surface. Growth lines fine, numerous and opisthocline; continuing across the midwhorl undeflected by any selenizone. Protoconch unknown.

REMARKS. This species differs from Brokenriveria pharlapensis and the indeterminite platyceratoid described below by possessing the twin row of nodes on the upper surface. The grossly similar Nodonema granulatum Linsley from the Middle Devonian Anderdon Lime- stone, is much smaller (holotype 6mm high), and possesses a distinct selenizone. Kitikamispira ukalundensis Cook, 1995, from the Emsian Ukalunda Beds of Queensland, and Kitikamispira kanekoi Kase & Nishida, 1988 from the Eifelian, Nakazato Formation, Japan both have nodes on all the spiral cords, and these spiral cords are strong and of equal intensity.

ETYMOLOGY. For Simon Lang.

Family PLATYCERATIDAE Hall, 1859 Platyceras Conrad, 1840

Platyceras (Platyceras) Conrad, 1840

TYPE SPECIES. Pileopsis vetusta, from the Lower Carboniferous of Queens County, Ireland by subse- quent designation of Tate (1869).

Platyceras (Platyceras) sp. (Fig. 4E-G)

MATERIAL EXAMINED. QMF32641, from SD43e.

DESCRIPTION. Small, horn-shaped rapidly ex- panding shell, 5.1mm high, 11.5mm maximum width; first whorl in contact, second disjunct; whorl profile ovate, aperture broken, but basal lip deflected strongly downwards near apertural

GASTROPODS FROM THE BROKEN RIVER PROVINCE 67

margin. Ornamentconsistsoffine,faintcollabral growth linesanddiffusecoarser growth rugae.

REMARKS. Substantial variation in members of the subgenus render it unwise to nominate å spe- cies on the basis of a single specimen. Platyceras (P. sp. A.of Tassell (1982) from the Earl y Devon- ian of Taemas has more disjunct whorls whereas the Broken River specimen has the first whorl in contact. Platyceras (P.) mansfieldense Tassell, 1977 from the Early Devonian Loyola Limestone has similar ornament and has the first whorl in contact, bur is a significantly larger form

PLATYCERATOIDEA gen. e! sp. indet, (Fig. 6), K)

MATERIAL EXAMINED. QMF34224 from QML1092,

DESCRIPTION. Turhiniform, medium-sized, 13mm high and 1 3mm wide. average spire angle approximately 110°. Sutures impressed, with whorls embracing slightly above midwhorl, Whorl profile rounded. Ornament consists of nu- merous spiral cords of equal strength, and fine weaker, slightly prosocline collabral growth lines.

REMARKS. The single distinct specimen lacks the two orders of spiral threads characteristic of Brokenriveria pharlapensis and does not bear the twin cords of Gemininodosa langi. The turbini- form, shape, lack of sinus and simple ornament places the specimen in the superfamily, and it 18 possibly a platyceratid, but further assignment 18 impossible without å wider selection of material.

Superfamily and Family indet. Burdikinia Knight, 1937

Burdikinia burdekinensis (Etheridge, 1917) (Fig. 2E,F)

Polyamma burdekeninesis 1917 Etheridge: 16; PI. 3 figs 1,2.

Burdikinia burdekinesis (Etheridge) Knight 1937: 711 (Etheridge) Knight 1941: 63; PL 57, figs 3a-h ; Heidecker, 1959: 5, PI 2, fig. 2, PI, 3, fig. 3a,b: Knight et al. 1960: 1309; fig. 205, 4a,b.

MATERIAL EXAMINED. QMF33578, QMF33579 from OML1083.

DESCRIPTION. Large, low-trochiform shell up to 35mm high, 55mm wide; apical angle of 130°. Suture channelled with keel on abaxial margin, whorls embrace on upper whorl face. Upper

TABLE4. Measurements tor Gemininodosa langi gen. et sp. nov,

Specimen | Height imm) | Width(mm) app M est I i42 11.0 | QOME33608 | 14,6 164 ]

QMF33638

OMP33641

Ce

109

whorl surface sloping and slightly concave, lower whorl surface rounded, Peripheral angulation al slightly above midwhorl. Aperture subrectangu- lar. Base rounded, heavily abraded, with poorly preserved prosocline growth lines. Relicts of nodes preserved on periphery and on sutural keel.

REMARKS. The material is heavily abraded, lacking the hasal ornament characteristic of the taxon, but it is otherwise indistinguishable from the holotype, and other material collected from the Burdekin Subprovince. A review ofthe higher taxonomy of this distinct gastropod is needed.

Superfamily MURCHISONIOIDEA Koken, 1896 Family MURCHISONIIDAE Koken, 1896 Murchisonia D^ Archaic & De Verneuil, 1841 Murchisonia (Murchisonia)

Murchisonia (Murchisonia) wandovalensis sp. nov. (Fig. TA-G)

MATERIAL EXAMINED. HOLOTYPE: QMF33680 from QML.1092, Papilio Mudstone, Givetian, Broken River Province. PARATYPES: QMF33675- QMF33679, QMF33681-QMF33685, QMF34129 QMF34184, QMF34195-QMF34202 from L1092.

DESCRIPTION. Small, high-spired, turbiniform gastropod, up to 11mm high and IImm wide (Table 5), with an apical angle of c. 55°. Suture impressed. Whorl profile angular with wide, mid- whorl, peripherally placed selenizone bordered by prominent 2 spiral cords. Upper whorl face

steep, slightly concave. Lower whorl face rounded, bearing a spiral cord 1/3 below ihe selenizone. Whorls embrace at this lowermost cord, just below the mid-whorl, Growth lines fine, numerous, collabral; concave 1n the selenizone; somewhat sinusoidal between the lower bordering cord of the selenizone and the spiral cord on the lower whorl face, Base rounded aperture subrounded with slit at midwhorl.

68 MEMOIRS OF THE QUEENSLAND MUSEUM

GASTROPODS FROM THE BROKEN RIVER PROVINCE 69

TABLE 5. Measurments for Murchisonia (Murchisonia)

Height (mm)

Width(mm)

QMF33676 å 7.2

QMF33680

QMF33685 QMF34202

DIAGNOSIS. Small, somewhat turbiniform Murchisonia (Murchisonia), with additional spiral cord on lower whorl face, just below selenizone.

REMARKS. The material is similar to M.(M.) fermioni Tassell 1982 from the Early Devonian Receptaculites Limestone, New South Wales, however M. (M.) fermioni has a more impressed suture and lacks the spiral cord on the lower whorl face. From other species of the subgenus it differs in the less high-spired form, and generally, the presence of the lower whorl face cord.

ETYMOLOGY. For Wandovale Station.

Murchisonia (Murchisonia) lawlessi sp. nov. (Fig. 8 A-I)

MATERIAL EXAMINED. HOLOTYPE: QMF33704 from QML1019, uppermost Dosey Limestone, Givet- ian, Broken River Province. PARATYPES: QMF33089, QMF34258, QMF33096, QMF33345, QMF33347, QMF33393, QMF33700, QMF34263 from QML 1019.

DIAGNOSIS. Medium-sized, high-spired mem- ber of subgenus, with ridge on upper and lower whorl faces, more prominent on upper, but not defining a distinct cord.

DESCRIPTION. Medium-sized, high-spired an- omphalus gastropod, up to 18.4mm high, 12.0mm wide at base (Table 6) with an apical angle of approximately 22-30°. Whorl profile angular with peripheral selenizone bordered by two cords. Upper whorl face shallowly sloping with a low, rounded, spiral ridge, just below the suture. Suture impressed, whorls embrace well below midwhorl at lower spiral ridge. Selenizone narrow, located at midwhorl. Lower whorl face rounded and convex, with weaker ridge, slightly lower and wider. Base rounded, aperture rounded with slit at peripheral margin.

TABLE 6. Measurements for Murchisonia (Murchisonia)

approx. apical angle (°)

lawlessi sp. nov.

Specimen

QMF33700 QMF33345

REMARKS. The species is differentiated from other Murchisonia (Murchisonia) in the area by the ridge on the upper and lower whorl faces in addition to those bordering the selenizone.

Murchisonia (Murchisonia) turris de Koninck from the Early Devonian Receptaculites Lime- stone, New South Wales lacks the two additional