PAPERS AND PROCEEDINGS OF THE ROYAL SOCIETY OF TASMANIA. VOLUME 104

[Manuscript received 4 September 1969

THE ASTERINID SEA STARS OF TASMANIA

By

A. J. DARTNALL The Tasmanian Museum, Hobart

(With one plate and one table)

SUMMARY Remarks: No recent material of the species from The ten species of sea stars of the family Tasmania has been seen by this author. Asterinidae occurring in Tasmania are considered. Asterina scobinata Livingstone, 1933 Synonymies, locality lists and distributional data A. A. Livingstone, 1933. Rec. Austr. Mus. vol. 19, are given. Patiriella gunnii (Gray) is redescribed no. 1, p. 1, PI. 5, figs 9-12. 15. and lectotypes designated. Some account of characters distinguishing P. gunnii and Patiriella M~terial examined: A total of twenty-four brevispina H. L. Clark is provided. A key to the speCImens. species is presented and some account of the zoo­ Localities: Boat Harbour; Greens Beach' Cape geography of the Tasmanian species given, atten­ Portland; Maria Island; Dunalley; Eagiehawk tion being drawn to the existence of pairs of Neck; Carlton; Blackmans Bay, Kingston; Variety sibling species in the group. Bay, Bruny Island; Bond Bay, Port Davey. Habitat: Under rocks and crevices in the lower INTRODUCTION littoral zone. Ten species of three genera of the sea star Distribution: Tasmania and Victoria. Extension family Asterinidae are known from the littoral of the known range to Victoria has been reported and shallow waters around Tasmania. In this in another place (Dartnall, in press). paper abbreviated synonymies are given as neces­ sary and details of localities, habitat and distri­ Genus PATIRIELLA Verrill, 1913 bution provided. In one case (viz. Patiriella gunnii (Gray» it has been found necessary to redescribe , regularis' group the species. A key to the Asterinidae has been Patiriella regularis (Verrill, 1867) constructed and is presented here. Finally the zoogeography of the Tasmanian asterinids is con­ Restricted synonymy sidered. Asterina regularis Verrill, 1867. Trans. Conn. Acad., vol. 1, p. 250. Family ASTERINIDAE Patiriella regularis Verrill, 1913. Amer. Jour, Genus ASTERINA Nardo, 1834 Sci., ser. 4, vol. 35, p, 480, Asterina atyphoida H. L. Clark, 1916 Patiriella mimica Livingstone, 1933. Ree. Austr. Mus. Sydney vol. 19. H. L. Clark, 1916. Endeavour Rept, p. 57; PI. 17, figs 1, 2. Asterina regularis Fell, 1959. Tuatara, vol. 7. (3), p. 127-142. Material examined: A total of three specimens. Patiriella regularis Dartnall, 1969. Proc. Roy. Localities: Greens Beach, near the mouth of the Soc. Tasm. vol. 103, p. 54. estuary of the River Tamar. Material examined: A total of 420 specimens. Habitat: Under rocks and in crevices. Low water to 40 metres, perhaps found at greater Localities: Oyster Cove, Kettering; Coningham; depths (Shepherd, 1968), Margate; Blackmans Bay, Kingston; Tinderbox; Taroona; Sandy Bay, Hobart; Wrest Point, Hobart; Distribution: The north coast of Tasmania, Hobart wharves; Ralphs Bay; Roches Beach; Victoria and South Australia. Midway Point; Slopen Main; Variety Bay, Bruny Asterina inopinata Livingstone, 1933 Island. A. A. Livingstone, 1933. Rec. Austr. Mus. vol. 19, Habitat: From the mid-littoral to 15 metres on no. 1, p. 3, PI. 5, figs 1-8. 14. rock, sand and mUd. Habitat: Very shallow water (H. L. Clark, Distribution: New Zealand and S.E. Tasmania 1946) . martnall, 1969). Distribution: S.E. Australia from New South Remarks: Evidence demonstrating that this Wales to Northern Tasmania. species may have been imported into Tasmania

73 74 THE ASTERINID SEA STARS OF TASMANIA

with cargoes of commercial oysters was presented Remarks: This author has now examined in a previous paper (Dartnall, 1969). In the ~ame material from South Africa and from the New paper it was suggested that Patiriella mimica Guinea archipelago. Even though this material (Livingstone) is a synonym of Patiriella regularis. has been attributed to Patiriella exigua on external Patiriella calcar (Lamarck, 1816) morphology some differences are observable. South African material, for instance, appears to Restricted synonymy attain a larger size than Australian P. exigua. Asterias calcar Lamarck, 1816. Anim. sans Vert., More consistent sampling throughout the Indo­ vol. 2, p. 557. Pacific area will be needed to examine the pro­ Asterina calcar Gray, 1840. Ann. Mag. Nat. position that the species known as Patiriella Hist., vol. 6, p. 290. exigua is a polymorphic assemblage of forms. Patiriella calcar Verrill, 1913. Amer. JOlLr. Sci., Patiriella vivipara Dartnall, 1969 ser. 4, vol. 35, p. 484; H. L. Clark, 1946. Dartnall, 1969. Proc. Linn. Soc. N.S.W. 93 (3f: Carnegie Institution of Washington. Publi­ 294-196, pI. XXIX cation 566, p. 134. Localities: Roches Beach; Midway Point; Material examined: A total of 199 specimens. Lewisham; Tesselated Pavement; Blowhole, Eagle­ Localities: Trial Harbour; Granville Harbour; hawk Neck. Boat Harbour; Wynyard; Cape Portland; Plinders Island; Bay of Fires; Binnalong Bay; 2 miles Habitat: Mid-littoral, gently sloping rocky north of Scamander; Shelly Beach, Orford; shores. Dodges Ferry; Roches Beach; Primrose Sands; Distribution: Restricted to S.E. Tasmania. Taroona; Coningham; Adventure Bay, Bruny Remarks: It is suggested that Patiriella vivipara Island; Bond Bay, Port Davey. is a sibling species to Patiriella exigua. Habitat: Mid-littoral to 10 metres. Rocky , gunnii' group shores and rock pools. Patiriella gunni (Gray, 1840) Distribution: Tasmania, south and east coasts of Australia from South Australia to Queensland. Restricted synonymy Remarks: H. L. Clark (946) employed the Asterina gunnii Gray, 1840. Ann. Mag. Nat. absence of suboral spines in this species to dis­ Hist., vol. 6, p. 289. tinguish it from other members of the genus. '1 he Patiriella gunnii Verrill, 1913. Amer. Jour. Sci., presence or absence of suboral spines is r.ot a ser. 4, vol. 35; p. 484; H. L. Clark, 1946. reliable characteristic of Tasmanian material and Carnegie Institution of Washington. Publi­ tends, also, to impute close relationships between cation 566, p. 135. P. calcar and the six-rayed' gunnii' forms. Patiriella yunnii A. M. Clark, 1966. Mem. Nat. The distribution of suboral spines in a .>ample of Mus. Viet., No. 27, p. 320; Shepherd, 1968. 199 specimens of Tasmanian Patiriella calcar is Rec. S. Aust. MUS., vol. 15 (4), p. 747. expressed in Table 1. The genitive of modern patronymics would give , exigua' group , gunni' as the specific name of this species. No Patiriella exigua (Lamarck, 1816) evidence has been given of typographical errors, lapsus calami or errors of transcription. Thus the Restricted synonymy rulings concerning customary usage must apply Asterias exigua Lamarck, 1816. Anim. sans and' gunnii' retained (see Article 19 and Summary Vert., vol. 2, p. 554. 8 of Opinions Rendered, Schenk and McMasters, Asterina exigua Perrier, 1876. Arch Zool Exp., 1936) . vol. 5, p. 222. Material examined: A total of seventeen Tas­ Patiriella exigua Verrill, 1913. Amer. Jour. manian specimens. Sci., ser. 4, vol. 35, p. 484; H. L. Clark, Localities: Greens Beach; Cape Portland; 1946. Carnegie Institution of Washington. Maria Island; Primrose Sands; Variety Bay, Bruny Publication 566, p. 136. Island. Material examined: A total of 257 specimens. Habitat: Mid- and lower-littoral to 30 metres, Localities: Trial Harbour; Granville Harbour; under rocks and ledges. Marrawah; Rocky Cape; Boat Harbour; Greens Distribution: The north and east coasts of Tas­ Beach, East Sandy Cape; Cape Portland, Swan mania, and the south and east coasts of Australia Island; Flinders Island; Shelly Beach, Orford; from Western Australia to New South Wales. Maria Island; Port Arthur; Nubeena; Murdunna; Also recorded from Lord Howe Island. Dunalley; Bellerive; Hobart; Blackmans Bay, Kingston; Coningham; Policemans Point; Simp­ Remarks: Gray's 0840, 1866) description of 'Asterina yunnii' from material collected by R. C. sons Bay, Bruny Island; Roaring Beach, Port Gunn in Van Diemens Land was concise and barely Davey. informative. Preserved material of ' gunnii' Habitat: Upper and mid-littoral zones. Often fascies (I.e., of Patiriella gunnii and Patiriella found associated with the alga Hormosira banksii. brevispina H. L. Clark) has proven difficult to Distribution: Tasmania, south and eastern distinguish. Through the offices of Miss A. M. Australia, Lord Howe Island. Said to occur Clark, the Director and Trustees of the British throughout the Indo-Pacific region. Museum (Natural History) made available to me A. ;r. DARTNALL 75 specimens, determined by Gray, to redescribe the Diagnosis: A flattened, six-rayed species of species. Patiriella closely related to Patiriella brevispina A single specimen from a group of four .from H. L. Clark. In Tasmanian material paired sub­ Sandy Bay, Hobart Town and registered in 1840 ambulacral spines distinguish P. gunnii from P. is designated the lectotype and three specimens brevispina which most often has single sub­ from a series collected at George Town, Northern ambulacral spines. A discussion of specific Tasmania and registered in 1849, paralectotypes. characteristics is given in a following section of this account. Lectotype: British Museum Reg. No. 40.3.9.-10. Sandy, Bay, Hobart Town (type locality). Pre­ Patiriella brevispina H. L. Clark, 1938 sented by R. C. Gunn. H. L. Clark, 1938. Mem. Mus. Comp, Zool., vol. 55, Pamlectotypes: Three specimens. British p. 166, pI. 22, figs 2, 3 Mmeum Reg. Nos 49.11.19.-10-14-33. George Town, Material examined: A total of twelve Tasmanian Tasmania. Presented by R. C. Gunn. specimens. Description of lectotype: Plate I, figs (i) and Localities: Burnie; Greens Beach; Cape Port­ Oil-specimen 40.3.9.-10; R = 22 mm; r = 16 mm; land; Flinders Island. r:R = 1:1.37; vb = 6 mm. Habitat: Mid- and lower-littoral to 10 metres A six-rayed, asterinid sea star, markedly fiattened, on a rock substrate. Most often found under rocks. the actinal surface plain and with a very acute Distribution: The north coast of Tasmania and marginal angle. the coasts of Western Australia, South Australia The abactinal surface is paved with imbricating Victoria, New South Wales and Southern Queens~ plates of two kinds. Those of the rays, the disc land. and most of the abactinal intermediate areas are Remarks: Patiriella gunnii and Patiriella brevi­ crescentic at their free edge allowing the passage spina have long been considered distinct forms by of papulae. The carinal row of papular plates of many Australian workers. H. L. Clark (1938, 1946) each ray are doubly notched allowing the passage and A. M. Clark (966) have both commented upon of two rows of papulae, one on each side of the the validity of P. brevispina. A. M, Clark's com­ midline of the ray. ments and the brief description of P. gunnii The plates of the interradial, abactinal areas offered by Gray 0840 and 1866) prompted an near to the margin are smaller and non-papulate. investigation into the characteristics of the Small secondary plates are present between the material showing the six-rayed' gunnii' fascies. larger papulate plates. Including the lectotype of Patiriella gunnii 250 The exposed surfaces of the abactinal plates are specimens were available for stUdy and the follow­ crowned with groups of small, blunt, thorny ing characteristics were considered reliable enough spinelets, some twenty-eight spinelets being present to distinguish between the species:- on the larger plates near the centre of the disc and (1) Size and shape: P. brevispina may attain six to eight spinelets on the plates near to the a size of 52 mm R as opposed to about margin. 39 mm R in P. gunnii. P. gunnii is more flattened than P. brevispina. The The madreporite is rounded, channelled and range of vh:R for the two species was perforate and situated interradially some 3 mm 0.21-0.26:1 in P. gunnii and 0.35-0.57:1 from the centre of the disc. The anus is not in P. brevispina, I.e., the latter is a visible in the lectotype. more arched form. The superomarginal plates are not distinct from (2) Furrow spinulation: P. gunnii carries more the succeeding rows of abactinal plates. The furrow spines on more adambulacral inferomarginal plates form the edge of the body plates than P. brevispina. and each carries a fringing group of seven to nine (3) Subambulacral spinulation: P. gunnii, not­ spinelets. withstanding the description of the The actinal surface between the ambulacral lectotype, most often carries paired sub­ grooves is paved with imbricate plates. The ambulacral spines whilst the subambula­ largest chevron of plates, reaching from the tip cral spines of P. brevispina are most of each ray to the mouth plates, carries one spine often monacanthid. on each plate. The succeeding three inner chevrons (4) Colour when alive: In life Patiriella of plates carry one spine on each plate for about brevispina is consistent in its rich purple the proximal half of their length and the remainder colouration on both surfaces of the body of the actinal intermediate plates two spines and never possesses the mottled and except towards the disc margin where some plates varied colours of Patiriella gunni. The may carry three spines. tube feet are pale straw coloured in P. gunnii and bright orange in P. brevi­ In the lectotype each adambulacral plate bears spina. a single subambulacral spine and a pair of furrow Examination of a series of specimens from spines. Furrow spines may be grouped in threes Western Australia suggests that intermediate or towards the mouth and singly towards the tip of hybrid forms probably exist on the south-western the rays. seaboard of Australia and until more material is Each oral plate bears five oral spines, the inner­ available the characteristics recorded here can most being the largest. The lectotype bears no only be applied to Patiriella gunnii and Patiriella suboral spines. Pedicellariae are absent. brevispina from the Maugean Province of Australia. 76 THE ASTERINID SEA STARS OF TASMANIA

KEY TO THE TASMANIAN ASTERINIDAE 1. Abactinal surface clearly divided into two areas: one on the disc and along the crests of the rays of crescentic plates; and other, comprising the rest of the upper surface covered by close set, small, rounded plates Genus PARANEPANTHIA Tasmanian Species P. grandis H. L. Clark Abactinal surface not clearly divided into two areas (2) 2. Carinal abactinal row of plates doubly notched for papuIae Genus PATIRIELLA Carinal abactinal row of plates with a single papuIar notch or none Genus ASTERINA (8) 3. Rays normally five in number (4) Rays normally more than five (6) 4. Large, >15 mm. Gonoduct directed aborally Patiriella regularis (Verrill) Small, R<15 mm. Gonoduct directed orally or absent (5) 5. Gonoduct absent; coelomic incubation of young. Actinal surface orange yellow Patiriella vivipara Dartnall Gonoduct directed orally. No internal incubation. Actinal surface blue-green Patiriella exigua (Lamarck) 6. Rays six; actinal intermediate spines paired (7) Rays seven to eleven, usually eight; actinal intermediate spines single Patiriella calcar (Lamarck) 7. Flattened; subambulacral spines usually diplacanthid. Colour varied Patiriella gunnii (Gray) Arched; subambulacral spines usually monacanthid. Colour in life consistently deep purple Patiriella calcar H. L. Clark 8. Form stellate Asterina scobinata Livingstone Form pentagonal, though margins may be concave (9) 9. Actinal intermediate plates with single spine or none Asterina atyphoida H. L. Clark Actinal intermediate plates with fine spines in groups of two to four Asterina inopinata Livingstone

Genus PARANEPANTHIA Fischer, 1917 and comment on the possible status of this form was made earlier in this account. Paranepanthia grandis (H. L. Clark, 1928) Two phenomena are of interest- Synonymy 1. The distributions of Asterina scobinata, Nepanthia grandis H. L. Clark, 1928. Rec. S. Asterina atyphoida and Asterina inopinata Aust. Mus. vol. 3, p. 393, fig. 113. mesh in the crossroads of the Bass Strait. Paranepanthia grandis H. L. Clark, 1938. Mem. The distribution of these species is addi­ Mus. Compo Zool., vol. 55, p. 159. tional evidence for the south-eastern Material examined: A total of four specimens. Australian 'Maugean' marine province as defined by Bennett and Pope (960) and Localities: Tasmania; D'Entrecasteaux Channel; the overlapping of warm and cool tem­ Piersons Point, Derwent Estuary. perate areas also described by these Habitat: All the Tasmanian material was col­ authors. lected between 4 and 6 fathoms. Shepherd (968) 2. Pairs of sibling species occur whose distribu­ gives the vertical range of this species as between tion lies along various radii from the focal 2 and 40 metres in South Australia. point of Bass Strait. These pairs are Distribution: West Australia to Sydney Harbour Asterina atyphoida/inopinata, Asterina and southwards to Tasmania. burtoni/scobinata, Patiriella exigua/vzvi­ para and Patiriella gunnii/brevispina. In ZOOGEOGRAPHY OF THE TASMANIAN two cases, those of A. scobinata and P. ASTERINID FAUNA vivipara hermaphrodism occurs and A. atyphoida possesses orally directed gono­ Of the ten species of Tasmanian asterinids one, ducts in contrast to A. inopinata where Paranepanthia grandis, cannot be included in any the gonoducts are orientated aborally. discussion of the zoogeography of the family. The Thus each form of a species pair is isolated material available only refiects localised collecting by a reproductive specialisation. It and although that species may be expected along remains to be seen by what mechanism the eastern and northern coasts of Tasmania dis­ the Patiriella gunnii/brevispina species cussion must await further evidence. remain distinct, but investigation of the Patiriella regularis must also be excluded from ecology and the temporal sequence of discussion because evidence has been provided that reproduction in the two species may pro­ the species is a recent recruit to the Tasmanian vide a fruitful field of investigation. sea star fauna (Dartnall, 1969). The relationships of the Tasmanian asterinid Only one species can be said to be endeInic to fauna lie with Australia and the Indo-West-Pacific. Tasmania, viz. Patiriella vivipara, and only The 'regularis' forms of Patiriella have been Patiriella exigua is distributed outside Australia successful in extending their distribution around A. J. DARTNALL 77 the Southern Hemisphere, perhaps by means of REFERENCES the West Wind Drift. The sequence of species BENNETT, 1. & POPE, E. C., 1960.-Intertidal Zonation of the eastwards in this case is (a) Patiriella calcar in Exposed Rocky Shores of Tasmania and its Relationship with the Rest of Australia. Aust. J. Mar. Freshw. Res., Australia, (b) Patiriella nigra H. L. r::lark, Lord n, pp. 182-219. Howe Island, (c) Patiriella regularis and l-'atiriella CLARK, A. M., 1966.-The Port Philip Survey 1957-63. Echino- oliveri (Benham) in New Zealand and (d) Paiiriella dermata. Mem. Nat. Mus. Vic., 27, pp. 289-384. calcarata (Perrier) from the Juan E'el'nandez CLARK, H. L., 1938.-Echinoderms from Australia. Mem. Islands and possibly the eastern coasts of South Mus. Compo Zool., Harvard, 55, pp. 1-596. America. This author does not accept that Perrier's ------, 1946.-The Echinoderm Fauna of Australia Patiriella jimbriata belongs to that genus and that its composition and its origin. Publ. Carnegie Instn.: form is not included here. No. 566, 567 pp. DARTNALL, A. J., 1969.-New Zealand Sea Stars in Tasmania. ACKNOWLEDGMENTS Proc. Roy. Soc. Tasm., 103, pp. 53-55. My thanks for advice, information and material ------, (in press) .-Some species of Asterina from are due to the following: Miss A. M. Clark (Bri.tish Flinders, Victoria. Victorian Naturalist. Museum (Natural History», Miss M. E. Downey GRAY, J. E., 1840.-A Synopsis of the Genera and Species of the Class Hypostoma (Asterias Linnaeus). Ann. Mag. (Smithsonian Institution), Mr R. H. Green (Queen Nat. Hist., 6, pp. 175-184, 275-290. Victoria Museum, Launceston), Dr E. P. Hodgkin -----, 1866.-Synopsis of the Species of Starfish in and Mrs L. Marsh (University of Western Australia), !he British Museum (witlh figures of some of the species). Miss E. C. Pope (Australian Museum), Dr B. J. IV + 17 pp" 16 pIs. (London). Smith (National Museum of Victoo:i.a'), and Mr D. SCHENK, E., & McMASTERS, J., 1936.-Procedure in Taxonomy. Wolfe (Fisheries Division, Department of Agricul­ Stanford University Press. ture, Hobart). I also wish to thaak Emeritus SHEPHERD, S. A., 1968.-The Shallow Water Echinoderm Fauna Professor V. V. Hickman for his kind help in of South Australia. I. The Asteroids. Reo. S. Aust. initiating me to this field of study. MUB., 15, (4), pp. 729-756.

TABLE I Distribution of suboral spines in a sample of 199 specimens of Patiriella calcw' from Tasmania. (a) ---~-,-_.. 00- _.~- ,--_._-- _._.'------~---- Number of Number of Number of oral plates Rays Specimens 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22

8 56 6 7 2 2 4 5 2 1 2 2 4 3 4 12 9 6 1 1 1 1 2 ------10 3 1 2

11 1 1

Total ., 66 Number of specimens carrying single suboral spines

-- (b)

Number of Number of Number of oral plates Rays Specimens 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22

8 8 1 2 2 3

10 1 1 Total .. 9 Number of specimens carrying paired suboral spines

Total number of specimens in sample 199 Number carrying suboral spines 75 Number without suboral spines 124 Number of specimens with one and two suboral spines on some plates 9

( i )

( ii )

PLATn 1.-LeetotYlw PaJ£riclla rllOHu'j (Cl'fty). British MU3e1.Hl1 (Natural Histonr ) register No. 40.:~.H.~1(}. (n Actina} surface. Iii) Abuctinal surface J Phoiogra].lha: C. t..-;, HUl'riS0fl FP 7§