The Holothuroidea Collected by the Royal Society Expedition to Southern Chile, 1958-1959

D.1. P AWSO N l

ABSTRACT : The holothurians collected by the Royal Society Expedi tion to south­ ern Chile, totalling 180 specimens, are described . Ten genera (of which one is new) and ten species are represented. N eopsolidium n.g., type species Psolidium con­ vergens (Herouard), is erected to accommodate those species in the genus Psolidium (sensu lato) in which the dorsal plates are reduced to a diameter of about 0.4 mm . Th e holorhurian fauna of southern Chile is generalised, containing few restricted species, and sharing many elements with distant subantarc tic islands and with Antarct ica.

DURINGlate 1958 and early 1959 an expedition and Deichmann (1947) have provided a clear sponsored-by. the Royal Society carri ed Out ma­ picture of the composition of the fauna. It is rin e and terrestrial observations and collections unlikely that many new shallow-water species in southern Chile. Stations were established in will be taken from this region. three separate areas, namely: I am grateful for the opportunity to study this most interesting collection, and I would like to 1. Isla Chiloe (approx. 42 ° S) thank Prof. G. A. Knox and the Royal Society 2. Puerto Eden to Punta Arenas (approx. forrnakingrhis materialavailable to me. 49 ° S to 52° S) 3. Isla Navarino and southern regions (a p­ prox. 55° S) . LIST OFSPE CIES COLLECTED Ord er Den drochirotida These three areas, considered together, were ex­ pected to provide a good picture of the changes Family Phyllophoridae in flora and fauna along the Patagonian coast­ Subfamily Thyonidiinae Heding and Pan­ line. Th e present paper contains an account of ning, 1954 Athyonidium chilensis (Semper ) the holothurians collected by the marine biol­ ogist to the expedition, Prof. G. A. Knox , of Family Cucumariidae Canterbury Uni versity, Christchurch. Subfamily Cucumariinae A total of 180 specimens of holothurians were Cladodactyla crocea ( Lesson) collected, and these comprise 10 genera and 10 Stereoderma laevigata ( Verrill) species. The collection represents a good cross­ Pseudocnus dubiosus (Semper) section of the known fauna, although a number Subfamily Colochirinae Panning, 1949 of species, hitherto regarded as common, were T rachyth yone lechleri (Lampert) not found . Family Psolidae The shallow-water holothurian fauna of south­ N eopsolidium conuergens (Herouard) ern Chile is not large in terms of species, al­ Psolus patagonicus ( Ekman) though a wide variety of genera are represented. Thi s' is particularly true for the order Den­ Ord er Apodida drochirotida. Previous auth ors such as Ludwig Family Chiridotidae (1898,a, 1898b), Perrier (1905 ), Ekman (1925), Chiridota pisanii ( Ludwig ) T rochodota purpurea (Lesson)

1 Department of Zoology, Victoria Un iversity of T aeniogyrus contortus ( Ludwig) Wellington, W ellington, N ew Zeala nd. Manuscript received September 27, 1963. Order Molpadida-none

453 454 PAOFIC SOENCE, Vol. XVIII, October 1964

Order Aspidochirotida-none Pseudocnus dubiosus (Semper ), 2 speci­ mens Order Elasipodida-none Psolus patagonicus Ekman, 1 specimen Area 3, Isla Na varino and South em R egions, MATERIAL EXAMINED Stations 50-78 Sta.50. Puerto Williams, Isla N avarin o, 54° 55' 180 specimens were collected from 12 of the 40" S, 67° 39' W ; intertidal boulder beach; 78 stations worked. Collecting was done by hand hand collecting; Jan. 7, 1959. in the intertidal zone, by diving, dredging, or Neopsolidium convergens (Herouard), seine netting. fragment Area 1, IslaCbiloe, Stations 1-1 8 Sta. 52. Puerto Robalo, Isla Navarino, 54° 55' 50" S, 67° 41' 40" W ; intertidal argillite rock; Sta. 4. Punta Gaviota, 42° 03' 50" S, 74° 02' 50" hand collecting and collecting from Macro­ W ' intertidal boulder beach, volcanic rocks; cystis fronds and holdfasts; Jan . 10, 16, 23, ha~d collecting; Oct. 4, 11, and 16, 1958. and 30, 1959. Athyonidium chilensis (Semper) , 4 speci­ Pseudocnus dubiosus (Semper), 10 speci­ mens mens Area 2, Puerto Eden to Punta Arenas, Stations N eopsolidium convergens (H erouard), 2 19-49 specimens Sta. 19. Puerto Eden, Isla Wellington, the point Sta. 54. Puerto Grandi, Isla Bertrand, 55° 12' S, to the north of the FACh base, 49° 08' 20" 67° 55' 30" W ; boulder beach; intertidal S, 74° 26' 55" W; intertidal granitic gneiss hand collecting; Jan. 12, 1959. rocks, sheltered; hand collecting, also collec­ Pseudocnus dubi osus (Semper), 4 speci­ tionfrom--Macrocystisfronds -and · holdfasts m ens and sublittoral collection by diving; Nov. 29 Trachythyone lecbleri (Lampert), 9 speci­ and 30, 195R mens Pseudocnus dub'iosus (Semper) , 1 specimen Sta, 73. Seno Grandi, small island opposite Sta. 27. Isla Carlos, 49° 09' 35" S, 74° 25 ~ 24" Puerto Grandi , 55° 15" S, 67° 56" W ; col­ W; collection from Macrocystis fronds and lection from Macrocystis fronds and holdfasts; holdfasts ; Dec. 5, 1958. Jan . 5, 1959. Pseudocnus dubjosus ~ Semper ) , 57 speci­ Cladodactyla crocea (Lesson), 24 specimens mens N eopsolidium convergens (Herouard), 1 Sta. 37. Caletta Lackawana, 49° 10' 32" S, 74° specimen ( juvenile) 25' 52" W; depth 18 m, sand, rock, shell; Chiridota pisanii (Ludwig), 4 specimens dredge; Dec. 9, 1958. T rochodota purpurea ( Lesson), 8 speci­ Stereoderma laevigata (Verrill) , 1 speci­ mens men Taeniogyrus contortus (Ludwig), 1 speci­ Sta. 39. Puerto Eden, west side of Canal Sur, men 49° 09' 52" S, 74° 26' 08" W; intertidal Sta.74. Seno Grandi, peninsula on Isla Navarino opposite Puerto Grandi, 55° 11' 20" S, 67 ° boulder beach of granitic gneiss; hand collect­ 56'W; collection from Macrocystis fronds ing and collection from Macrocystis fronds and holdfasrs; Jan. 5, 1959. and holdfasts; Dec. 9 and 11, 1958. Tro chodota purpurea (Lesson), 2 speci­ Pseudocnus dubiosus (Semper), 15 speci­ mens mens Sta. 77. Puerto Grandi, Isla Bertrand, to the west Psolus patagonicus Ekman, 30 specimens of the wharf; 55° 12' S, 67° 55' 30" W; Sra, 40. Isla Dulce and Isla Levinson, Puerto intertidal granitic rocks and boulder beach, Eden, 49 ° 09' 02" S, 74° 25' 10" W; inter­ semi-sheltered, hand collecting; Feb. 7, 1959; tidal and sublit toral granitic gneiss rocks; collection by diving among Macrocystis; Feb. hand collecting; Dec. 12 and 13, 1958. 8, 1959. Holothuroidea from Southern Chile-PAWSON 455

Pseudocnus dubiosus (Semper), 1 specimen rosettes in younger indi viduals. Colour greyish N eopsolidium convergens ( Herouard), 2 mottled to almost black, ventrum paler, tentacles specimens dark. Shallow-water forms. ( After Deichmann , T rochodota purpurea (Lesson ), 1 specimen 1941.) MATERIAL EXAMINED: Sra. 4, four specimens from Macrocystis zone. SYSTEMATIC ACCOUNT REMARKS: Th e four specimens in the present Order DENDROCHIROTIDA collection range in length between 140 mm and 185 mm. The body is covered in tube feet, which Family PHYLLOPHORIDAE are more numerous ventrally. Colour in alcohol Subfamily THYONIDIINAE Heding and is dark brown to blackish dorsally, fading to Panning, 1954 light brownish-grey on the ventral surface. There are 20 black tentacles arranged in two ring s. Th e Genus Athyonidium Deichmann , 1941 outer ring has 10 regularly spaced, profusely branched larger tentacles, averaging about 25 Eucyclus Lamp ert, 1885, p. 920; Th eel, 1886a, mm in length. The inner ring lies close around p. 268; Ludwig, 1887, p. 1239; Heding and the mouth, and comprises five radially placed Panning, 1954, p. 36: name preoccupied. pairs of short, sparsely branched tentacles of 8 TYPE SPECIES: Athyonidium chilensis (Sem­ mm average length. The oral field is about 20 per ) . mm in diameter. Immediately outside the ring REMARKS : This genus is rnonotypic. Deich­ of large tentacles, in the middorsal interradius, mann ( 1941) pointed out that the generic lies a small approximately circular mound of name Eucyclus was preoccupied, having been tissue about 1 mm high and 2 mm broad, which claimed -sorne- years-earlier - for -the Mollusca. supportsrhegeniralaperrure. Heding and Panning ( 1954) unfortunately per­ The mouth is large, and examination of sisted with the old generic name. the intestinal contents in a dissected specimen showed that hard pieces of Macrocystis stipe up Athyo,!1-idium cbilensis (Semper) to 42 mm long and 6 mm wide, or 25 mm X 10 mm can be ingested. The intestine also contains T hyone (Sto lus) cbilensis Semper, 1868, p. 241, fragments of green algae, both filamentous and pI. 40, figs. 3-6; Lampert, 1885, p. 156. rhalloid, brown algae (predominantly Macro­ Eucyclus duplicatus Lampert, 1885, p. 250; cystis) , appendages of small crustaceans, and Th eel, 1886a, p. 268. hydroids. This species seems to be primarily a T hyone cbilensis Theel, 1886a, p. 139. vegetarian browser. Phyllophorus chilensis Ludwig, 1886, p. 24. The calcareous ring -is large, and has- been Athyonidium chilensis Deichmann, 1941, p. 127. illustrated by Heding and Panning ( 1954). Eucyclus chilensis Heding and Panning, 1954, The gut is thin-walled, and takes a large p. 36, fig. 2. S-shaped loop. The cloaca is broad, thick-walled , DIAGNOSIS : Large forms (25 ern ) with thick, attached to the body wall by very numerous soft skin and numerous stout feet. Tentacles, five muscle strands. Respiratory trees take the form large external pairs and five small inner pairs. of two broad flat tubes with scattered, profusely Calcareous ring with five stout, almost rectangu­ branch ed tufts of respiratory tubul es. The trees lar radials and five insignificant interradials, arise from the anterior end of the cloaca in the often completely concealed in surrounding tis­ lateral dorsal interradii, and extend anteriorly sue. One large dorsal stone canal, often branched, for about half the length of the body cavity. and one or two tufts of smaller stone canals with A tuft of numerous Polian vesicles arises from minute heads. Numerous tufts of branched the ventral side of the water-vascular ring. The Polian vesicles. Inner anatomy otherwise typical stone canal stems from the dorsal side of the of the family. ring, and gives off smaller canals which lie to Deposits large, well-developed end plates, a each side of the strong dorsal mesentery, and few perforated spi nous rods; tentacles with few terminate in nodular madreporites. 456 PAOFIC SOENCE, Vol. XVIII, October 1964

The gonad almost fills the rest of the body Cladodactyla crocea (Lesson) cavity, and consists of two bunches of sparsely Cucumaria crocea Lesson, 1830, p. 153, pI. 52, branched genital caeca, lying one to each side of figs. 1, l a; Theel, 1886a, pp. 58, 110, pl. 12, the dorsal mesentery. The caeca are orange in figs. 1, 2; Ludwig, 1898a, p. 15, pI. 1, figs. colour, with white thickenings scattered along 6-13 (l ist of references); Bell, 1908, p. 2; their length. Ekman, 1925, p. 75, figs. 15, 16; Deich­ Radial muscles are represented as broad flat mann, 1947, p. 331. straps. The retractor muscles are each split .inro Cladodactyla orocea Panning, 1940, p. 172; Pan­ two or three narrow bands, joined by a thin web ning, 1949, p. 4 13; Panning, 1957, p. 27, of tissue for most of their length. figs. 12, 13. In the specimen dissected a commensal pinno­ DIAGNOSIS : Thin-skinned, small (up to 100 therid crab was found in the last branch of the mm ), dendrochirotes with equal-sized tentacles. intestine, half way along the body cavity. The Tube feet restricted to the ambulacra, larger crab appeared to have caused little damage to ventrally, smaller and more numerous dorsally adjacent tissues, apart from a slight rupturing of (when present); usually arranged into double the wall of the intestine. This rupture might rows. Calcareous ring simple, with no posterior conceivably have been caused during preserva­ processes. Deposits perforated rods or platel ets, tion or transport of the specimens. reduced or lacking in older specimens. Tube feet I found no calcareous deposits in the body with end plates. Brood-protecting. wall or in the tube feet, although end plates are MATERIAL EXAMINED: Sta, 73, 24 specimens. known to occur in the tube feet of this species. REMARKS: Cladodactyla crocea.is one of the Due to the work of Deichmann ( 1941) and best-known of the holothurians from southern Heding and Panning (195 4 ) , the systematic waters, and I have little to add to the thorough positionofrhis species is nowquite'clear:--.- accountsgiven by the workers listed inthes yn­ DISTRIBUTION : Deichmann (1941) notes that onymy above. The specimens in the present col­ Athyonidium chilensis is probably "the most lection range in total length from 10 mm to 27 common phyllophorid known from Chile." The mm. Colour in alcohol is light brown to dark species is also found along the coast of Peru. greyish-brown. In some specimens, tube feet The present locality record does not affect the are entirely lacking from the dorsal radii. Theel known distribution pattern, and it is apparent (1886) noted that in the 20-40 mm specimens, that this species is reasonably common in the dorsal tube feet are wanting, but this is not the Macrocystis zone, imm ediately below low-tide case in all of the specimens in the present col­ mark. lection. I found no juveniles on the dorsal sur­ face of any specimens, although Theel's (1886) Family CUCUMARIIDAE material, which was collected at about the same time of the year, included some specimens carry­ Subfamily CUCUMARIINAE ing broods of juveniles. Examination of the Genus Cladodactyla Brandt, 1835 emend. gonad in the larger specimens in my collection Panning, 1940 indicated that sexual maturity had not as yet been reached. DIAGNOSIS : Tentacles 10. Calcareous ring Calcareous deposits proved to be very rare, small, without bifurCfltingpro:::esses. Skin de­ the tentacles someti mes containing isolated per­ posits merely plates. The plates are thin, smooth , forated rods in the smallest specimens ( Fig. 1, developed from forked rods. (After Panning, l ) . In larger specimens the tentacle rods were 1949.) mainly absent. With the exception of the end TYPE SPECIES : Cladodactyla crocea ( Lesson) . plates in the tub e feet ( Fig. 1, 2), body wall REMARKS: The genus Cladodactyla contains deposits tend to be very scarce or lacking al­ thr ee species at the present time. C. senegal­ together. In a single 14 mm specimen I found ensis Panning and C. monodi Cherbonnier are but thr ee deposits in the skin of the dorsal side known from Dakar and the Cameroons coast, ( Fig. 1, 3 ). These took the form of perforated respectively. platelets. Holothuroidea from South ern Chile--PAWSON 457

Th e features of the internal anatomy have Herouard, 1901, p. 44; Perrier, 1905 , p. 22; already been described. Herouard, 1906, p. 12, pI. 2, figs. 5, 6; Bell, DISTRIBUTION: Cladodactyla crocea is most 1908, p. 2; Helfer, 1917, p. 164; Ekman , commonly found at the southern tip of South 1925, p. 56, text fig. 11; Deichmann , 1947. America, about the Straits of Magellan, and Cucumaria serrata Theel, 1886a, p. 73. along the eastern coast of southern South Amer­ Stereoderma laevigata Panning, 1949, p. 422. ica, as far north as the mouth of the Rio de la DIAGNOSIS: Tentacles of equal size, feet re­ Plata. The species is also known from Kerguelen, stricted to the ambulacra, calcareous ring simple. South Georgia, and Antarctica (Coulman Is., Spicules as oblong plates with one end denti cu­ Hut Point, and Franklin Is.)(Bell, 1908 ) . C. late, sometimes knobbed with a reticul ated net­ crocea has been taken between low-tide mark work ; they are numerous in smaller specimens, and 4,300 rn, but is most common to depths of more scattered in larger ones. Tube feet have about 30 rn, often associated with seaweed. end plates and three-armed rods. Tentacles wit h perforated rods; introvert with four-hol ed but­ Genus StereodermaA yres, 1851 emend. tons sometimes with an external reticulum. Panning, 1949 Total length up to 120 mm. ( After Deichmann , 1947.) Pentactella Verrill, 1876. MATERIAL EXAMINED: Sra, 37, one specimen. DIAGNOSIS: Tentacles 10. Calcareous ring REMARKS : The single specimen is small (total simple, without forked processes. Body wall de­ length 8 rom), and is strongly contracted, with posits are knobbed plates, all of the same shape, many deep transverse wrinkles. Colour in alco­ and arranged in one layer; no tables, no rosettes, hol is light brown: Th e calcareous deposits are no cups.( After Panning, 1949.) typical of this species and need no.furrher dis­ TYPEsPEcIEs':Sier;ioderma 'unisemita (S timp ~ cussiOn nere:Stiri oderma laevigata is"a species son ) . which is readily recognised because of its dis­ REMARKS: Th e genus Stereoderma contains tinctive calcareous deposits. about 12 species at the present sime. Four of the DISTRIBUTION: Deichmann (1947 ) notes that species have a circumpolar distribution when S. laevigata is known from the south ern tip of considered together. South America, the Chile coast, Falkland Is., S. leoninoid es (Mortensen), Auckland and Kerguelen Is., the Crozets, Marion Is. Bell Campbell islands. (1908) reported specimens from the vicinity of S. godefJroyi (Semper), west coast of South McMurdo Bay, Antarctica, to depths of about America 20°-40° S (Deichmann, 1947). 41 fathoms. S. laevigata (Verrill), southern end of South America, Falkland Is., Kerguelen Is., Ma­ Genus Pseud ocnus Panning, 1949 rion Is., the Crozets, Antarctica. DIAGNOSIS : Tentacles 10. Calcareous ring S. perrieri (Ekman) , southern Chile, Falkland simple, without posterior processes. In the body Is., South Georgia. wall are knobbed plates of two types in two Th ese four species seem to be closely related layers. (After Panning, 1949.) to each other, and their distribution parallels TYPESPECIES : Pseudo cnus ko llikeri (Semper). the case of certain species of T rachythyone (Pawson, 1962). Pseudocnus du biosus (Semper)

Stereoderma laevigata ( Verrill) For synonymy see Pann ing ( 1950) . DIAGNOSIS: Medium-sized, length up to 100 Pentactella laevigata Verrill, 1876, p. 68; Studer, mm . Tube feet in five bands, numerous in the 1876; Studer, 1879 ; Smith, 1879, p. 271. dorsal ambulacra. Tentacles bushy, of unequal Cucumaria laevigata Th eel, 1886a, p. 57, pI. III, size. Calcareous ring simple. Deposits numerous fig. 5, pI. IV, fig. 13; Lamp ert, 1886, p. oblong knobbed plates (of average length 0.12 828; Ludwig, 1898a, p. 32, pI. II, fig. 25; mm ) with one end denticulate, and four-holed 458 PACIFIC SCIENCE, Vol. XVIII, October 1964

p ·1mm. • O·1mm.

O·1mm.

O·1mm· 1 figs.4 ,6.

O·1mm. , figs.5,T Holorhuroidea from Southern Chile-PAWSON 459 knobbed buttons (0.9 mm average length). Tube Subfamily COLOCHIRINAE feet with rudimentary end plates or none at Genus Trachythyone Studer, 1876 all, and numerous perforated supporting rods, mostly three-armed. Introvert and tentacles have For synonymy, see Panning (1949) . perforated plates and rods. (After Deichmann, DIAGNOSIS: Calcareous ring simple, without 1941.) forked processes. In the skin are cups and smooth MATERIAL EXAMINED : Sta. 19, 1 specimen; plates, the plates imbricating in some species; Sra, 27, 57 specimens ; Sta, 39, 15 specimens ; ( After Panning, 1949. ) Sta, 40, 2 specimens; Sra. 52, 10 specimens ; Sra. 1YPE SPECIES: Trachythyone muricata (Stu­ 54, 4 specimens; Sta. 77, 1 specimen. der ) ~ REMARKS : The 90 specimens range in length REMARKS : This genus comprises about 20 from 10 mm to 50 mm. The colour varies be­ species, of which 11 are distributed in the south­ tween white and pink. This species is one of the ern Pacific Ocean from Australia to South Amer­ best-known from Chile and has been described ica and also Kerguelen Is. The remaining species by many workers. Panning (1950, 1952) has are found in the Indo-west-Pacific and Medi­ suggested that there are three "forms" of this terranean regions. species. They are P. dubiosus-leonina, from the southern end of South America. (all of the pres­ T rachythyone lecbleri (Lampert) ent specimens are of this type); P. dubiosus­ dubiosa ranges the Atlantic and eastern Pacific Thyone lecbleri Lampert, 1885, p. 253, fig. 64; shores, as far south as Chile; P. dubiosus-jageri Theel, 1886a, p. 267; Ludwig 1898a, p. 44, is known from South Africa. pIs. 2, 3, figs. 26--33; Perrier, 1905, p. 35; The deposits of the body wall in the present Ekman, 1925, p. 101, fig. 22; Deichmann, material ( Fig.-1,- 5, 7) are those of the leonina­ 1947, p.335. --- form. Small specimens may show stages in the Thyone hassleri Theel, 1886b, pp. 11-12. development of knobbed buttons (Fig. 1, 8). Trachythyone lecbleri Panning, 1949, p. 426, The tube foot deposits (Fig. 1, 4) are perforated figs. 12-14. plates, often three-armed, about 0.22 mm in DIAGNOSIS: Medium-sized (up to 15 ern total average length. The tentacles contain perforated length ); colour in alcohol, brown. Tube feet rods and small plates up to 0.32-mm long ( Fig. numerous, scattered . Deposits thick oval plates, 1,6) . sparsely perforated, length 0.07-0.2 mm, over­ A thorough investigation of all of the species lain by rudimentary "cups" of varying shapes. End plates in tube feet surrounded by perforated in the genus Pseudocnus is urgently needed, par­ supporting plates of 0.2 mm average length. ticularly those species which have the body wall MATERIAL EXAMINED : Sta. 54, nine speci­ deposits as plates with one end denticulate, and mens. buttons. For a successful attempt at a revision, REMARKS : These are typical specimens of this a representative range of specimens of each unusual species. The total length ranges from species would be required, and unfortunately 47 mm (a strongly contracted specimen) to 130 these are nor available to the writer at the pres­ mm . The colour in alcohol is mottled light ent time. brown to dark orange-brown. The tube feet are DISTRIBUTION : The leonina-form of this spe­ lighter in colour than the rest of the body. In cies is known from southern Peru in the west of most cases the tentacles are completely retracted. South America to the Rio de la Plata in the east, The calcareous ring is simple, with no pos­ and the Falkland Is., from the intertidal zone to terior processes, and is solid, with thefive radials approximately 100 rn, and five interradials firmly fused together. The

FIG. 1. 1- 3, Cladodactyla crocea (Lesson ) . 1, Tentacle deposits . 2, Dorsal rube foot end plate. 3, Dorsal skin deposits. 4-8, Pseudocnus dub iosus (Semper). 4, Tube foot deposits. 5, Knobbed plates from the skin . 6, Tentacle deposits. 7, Kn obbed buttons. 8, Developing knobbed buttons. 3·0mm.

, O·1mm. ,

0 ·2 rnrn,

FIG. 2. 1-5, Trecb ytbyone lecbleri ( Lampert) . 1, Radial and interradial pieces of calcareous rin g. 2, Pla tes from the body wall. 3, Tube foot deposits. 4, Plares from the posterior extr emity of the body. 5, Rudimentary cups. 6-10, N eopsolidiam conoergens (H erouard ). 6, Cups from the dor sal skin. 7, Tentacle deposits. 8, Plates from the dorsal body wall. 9, Buttons from the ventral body wall. 10, Plares from the ventral body wall. Holothuroidea from Southern Chile-PAWSON 461 radials each have an anterior notch for insertion have assigned to species lechleri are in fact has­ of retractor muscles (Fig. 2, 1). There is a single sleri) , or Lampert's type specimen of lechleri tubular Polian vesicle about 40 mm in length, may have been unusual in possessing posterior which arises from the water-vascular ring"in the processes. The calcareous deposits of species left dorsal interradius; the rnadreporite is a very lecbleri and hassleri are apparently identical and small nodule about 1.5 mm in diameter, lying are so unique in character that it seems logical near the posterior edge of the calcareous ring in to leave the two species as synonyms. The prob­ the middorsal interradius. The gonad comprises lem cannot finally be resolved until Lampert's two bunches of elongate unbranched orange­ type material can be re-examined. coloured caeca, which attach to the long genital DISTRIBUTION: Trachythyone lecbleri is duct near the middle of the body in the mid­ known from the vicinity of Magellan Straits and dorsal interradius. Tierra del Fuego in depths up to 30 m. A single The skin is packed with great numbers of cal­ holothurian from Heard Is., resting in the col­ careous deposits, which chiefly take the form of lection of the Dominion Museum, Wellington, thick, oval plates, with few perforations or none, was identified by the author as Trach ythyone and a length varying between 0.07 mm and 0.2 lecbleri, and thus the range of this species is now mm (Fig. 1, 2). Overlying the plates are a small considerably extended. number of rudimentary "cups" which vary con­ siderably in shape (Fig. 2, 5 ), and occasionally Family PSOLIDAE carry short, blunt knobs, and may have up to three perforations. At the extreme posterior end Genus Neopsolidium n. gen. of the body the plates are larger, and more com­ DIAGNOSIS: Small forms, with sole not sharply plex, with a great number of perforations, and , distinguished from the rest of the body. Dorsal a tendency to become knobbed or two-layered deposits small (up to 0.4 mm) smooth per­ (Fig. 2, 4). These plates are up to 0.3 mm long: forated plates, and cups. Ventral deposits plates The tube feet contain stout end plates which are similar to dorsal plates and perforated buttons; surrounded by perforated supporting deposits no cups ventrally. (Fig. 2,3), having an average length of about TYPE SPECIES: Psolidium convergens (He­ 0.2mm. rouard). Lampert ( 1895) apparently stated that this REMARKS: Deichmann (1941, 1947 ) stated species has a calcareous ring with posterior pro­ that the species Psolidium convergens deserves longations, as Theel (1886a) , after examining " a separate genus because of the nature of its dor­ the "Challenger" specimens of Tbyone lechleri, sal deposits, which differ from those in most stated that "each piece (of the calcareous ring) other Psolidium species. This opinion was based . . . has a bifurcate projection posteriorly." Then chiefly on the thorough description and figures Theel (1886b) described a new species, Thyone given by Perrier (1905) of this species. Clark bassleri, which resembled Lampert's Tbyone (1946) agreed with Deichmann, but neither of lechleri in most respects, but lacks posterior these workers elected to propose a new generic processes on the calcareous ring. Ludwig (1898a) name. Examination of specimens of Psolidium illustrated the calcareous ring of what he be­ convergens in the Royal Society collection lieved to be T hyone lechleri, and it is similar in has convinced me that this species should be every way to the ring illustrated in this paper assigned to a separate genus. Although N eop­ (Fig. 2, 1), and lacks forked posterior processes. solidium is monotypic at present, it may accom­ Panning (1949 ) also states that the calcareous modate some other Psolidium species, and future ring in species lechleri lacks posterior processes. work will show whether or not this is possible. Either Theel (1886b) was quite correct in erect­ The family Psolidae contains but a small num­ ing the species hassleri for those specimens with ber of genera, and the genus Psolus itself is in no posterior processes on the calcareous ring (in urgent need of revision. Such a revision is be­ which case some specimens previous workers yond the scope of this work. 462 PACIFIC SCIENCE, Vol. XVIII, October 1964

KEY TO THE GENERA IN TH E FAMILY PSOLIDAE is 4 mm in length. Colour in alcohol is dirt y white to light brown. The tentacles are orange 1 ( 10). Tentacles 10. to light brown, darker in colour than the rest of 2 (7). Dorsal surface with tube feet-. the body. Tube feet are restricted to the radii 3 (4 ) . Dorsal deposits include hourglass­ ventrally, where they are arranged in a double shaped bodies, but lack cups or bas- row in each radius. Near the extreme anterior kers., _. T hyonepsolus Oark and posterior ends of the body, the feet decrease 4(3). Hourglass-shaped bodies lacking, in numbers and adopt a biserial arrangement. cups or baskets present. Dorsally the feet are scattered over both radii 5 (6) . Sole sharply set off; dorsal deposits and inrerradii , The mouth and anus are slightly conspicuous scales, and small cups. upturned. Th ere are about 10 anal teeth or Dorsal tube feet may pass through papillae. some of the scales. Sole deposits Dep osits in the dorsal skin are thick per­ plates or buttons, usually with an ex- forated plates 0.2-0.4 mm in length (Fig. 2, 8). ternal layer of small deep cups . The plates are closely aggregated together and ...... _ _ .PsolidiumLudwig overlain by cups, which are 0.14 mm in di­ 6 ( 5) . Sole not sharply set off; dorsal de­ ameter (Fig. 2, 6 ). The cups are lacking in posits small (up to 0.4 mm), smooth the juvenile specimen. Ventrally the plates are perforated plates, and cups. Ventral slightly more irregular in outline and are not so deposits plates (similar to dorsal closely crowded together ( Fig. 2, 10 ). Inter­ plates), and buttons (no cups)...._..... mingled with the ventral plates, and more ___ ..___ Neopsolidium n. gen. numerous than they are irregular perforated but­ 7 (2 ) . Dorsal surface with out tub e feet. tons (Fig. 2, 9). The tentacles contain perforated 8 (9 ) : Dorsal surface -with imbricating plates and burron-like deposits in great numbers scales. Mouth and anus dorsaL . (Fig. 2, 7).Some of these deposits are curved ...... _.._Psolus Oken and carry small knobs. 9 (8 ). Dorsal surface smooth, deposits spar­ This species differs from the others known in ingly scattered oval perforated plates, the region ( Psolidium dorsipes and P. disci­ sometimes with warty surfaces, of farmis) in possessing the characteristic small average diameter 0.1 mm. Mouth dorsal plates, which have warranted erection of terminal, anus subdorsaL .. a new genus to accommodate it...... Pseudopsolus Ludwig DISTRIBUTION : N eopsolidium conv ergens has 10 ( 1) . Tentacles 15 .Stolin«s Selenka been taken from the waters about Cape Horn, Magellan Straits, Falkland Is., and South Georgia, N eopsolidiuni convergens ( Herouard ) in depths ranging to about 15 m, and seems to favour life in the Macrocystis zone, where Cucumaria convergens H erouard, 1901, p. 30. Herouard ( 1901) first discovered the species. Psolidium convergens Perrier, 1904, p. 15; Per­ rier, 1905, p. 38, text figs. D-F, pI. 2, figs. Genus Psolus Oken, 1815 2-4; Herouard, 1906, p. 12, pI. 1, figs. 7,8; pI. 2, figs. 10-12; Ekman , 1925, p. 111; DIAGNOSIS: Tentacles 10. Dorsal surface lack­ Deichmann, 1947, p. 336. ing tube feet, covered by imbricating scales. Sole DIAGNOSIS : As for the genus. sharply defined, deposits smooth or knobbed MATERIAL EXAMINED: Sta. 50, fragment of plates or but tons. anterior end; Sta, 52, 2 specimens; Sta. 73, 1 TYP E SPECIES : Psolus phantapus (Strussen­ specimen; Sta, 77, 2 specimens. feldr). REMARKS : The material examined agrees well REMARKS: This genus now contains over 30 in most respects with the excellent descriptions species, which are spread widely over the Arcti c and figures given by Perrier ( 1905) and Herou­ and Antarctic regions and in the trop ics. Psolas ard (1906) . The total length ranges between species are most common in shallow water, but 18 mm and 22 mm while the juvenile specimen some types are known from considerable depths . Holothuroidea from Southern Chile--PAWSON 463

Psolus patagonicus Ekman The calcareous ring comprises five radials and Psolus patagonicus Ekman, 1925, p. 140, text five interradials fused together. Each radial piece figs. 35, 36; Deichmann, 1941, p. 148, pI. has an anterior process with a deep, narrow 30, fig. 8; Deichmann, 1947, p. 339, figs. notch. Interradials each have an anterior process 1, 2. with no notch. There are no posterior processes DIAGNOSIS: A small psolid, up to 20 mm in (Fig. 3, 6) .T he ring is turned so as to lie parallel total length, with oral and anal valves, and radial with the dorsal surface of the body to correspond teeth between them. Few scales between mouth with the dorsal position of the mouth. Therefore and anus. Sole distinct with one or two margi­ the mid-ventral radial piece is the most anterior nal rows of tube feet. Mid-ventral radius naked. portion of the ring. Sale deposits mostly four-holed knobbed buttons Th e thin-walled intestine is coiled into three (average length 0.1 nun), together with a few loops, and the mesentery of the posterior loop small plates. Tentacles invested in smooth or of the intestine lies as usual in the right ventral knobbed perforated plates (0.07- 0.17 mm in interradius. Overall, the intestine is a dark length). orange-brown in colour. A single bulbous Polian MATERIAL EXAMINED: Sta, 39,3 0, specimens; vesicle arises from the water vascular ring in the Sta. 40, 1 specimen. left ventral radius. Th e short stone canal and DESCRIPTION: The specimens are small (total nodular madreporite lie in the midd orsal inter­ length ranges from 2.5 mm to 11 mm ), dorso­ radius. ventrally flattened, oval in outline, broadest near The gonads in the larger specimens are well the anterior end. The dorsal surface carries developed as two bunches of light brown tubular mouth and anus, and is invested in overlapping unbranched caeca, which extend for the length plates. Th e ventral sale is soft. Colour in life of the body cavity. The genital duct proceeds "salmon-pink'tjin alcohol; lightyellowish-brown along the dorsal side ofthe calcareous ring (or dorsally, orange-brown ventrally. The tentacles more correctly, the "posterior" side ) , and opens are orange-yellow, with some small brown spots. to the exterior immed iately behind the tentacles, The dorsal plates are about 0.8 mm broad and but apparently within the oral valves. tend to overlap towards the midline, while the Apart from the dorsal surface, calcareous de­ dorsal surface is bordered by one to three rows posits are present in the sale, the tentacles, and of smaller marginal plates (Fig. 3, 1) . The plates the tube feet. are thick, reticulated, and beset with minute 1. Sole deposits: Th e thin sole contains four­ knobs. The plates also carry a small number of holed button ( Fig. 3, 3 ), and some slightly tiny pearl-like grains, which are less common in larger knobbed or smooth plates ( Fig. 3, 3). smaller specimens. There are about five plates These deposits are sparingly scattered in the between mouth and anus. sole. The dendri tic tentacles are extended in most 2. T ube foot deposits: Each tube foot has an specimens, and the ventral pair are somewhat end plate ( Fig. 3, 7) of average diameter 0.23 smaller than the rest, unbranched or weakly mm, which is surrounded by some curved per­ branched. Th ere are five conspicuous triangular forated rods and plates, which may carry knob­ interradial oral valves ( Fig. 3, 1), between and like pro jections ( Fig. 3, 5 ), The length of these below which lie five radial valves, which are in deposits varies between 0.06 mm and 0.2 mm. the form of elongate isosceles triangles. In a 3.Tentacle deposits: The tentacles are in­ small number of specimens, one to four smaller vested in a network of perforated rods and plates lie near the base of the oral valves, but plates, some of which carry knobs ( Fig. 3,4). these are not always present. The anal aperture REM ARKS: After having examined th ese speci­ is also covered by five radial valves and five anal mens, I thought that I had found a new species, valves ( Fig. 3, 1). as they differ in some respects from Ekman's The soft sole is semitransparent and is bor­ (1925) type specimen. Both Ekman (1925) and dered by a ring of tube feet in a single or some­ Deichmann (1941) state that Psolus patagonicus times double row. The mid-ventral radius is can have small intercalary plates between the naked ( Fig. 3,2). larger dorsal plates. The photograph of Ekman's 464 PACIFIC SCIENCE, Vol. XVIII, October 1964

(1925) specimen clearly shows these plates. references ); Ludwig, 1898b, p. 445; Clark, None of the specimens I examined possess these 1907, p. 118; Ekman, 1925, p. 145, text fig. intercalary plates. Ekman's specimen was 20 mm 62; Heding, 1928, p. 297, text fig. 62; long, while the largest in the present collection Heding, 1931, p. 676; Deichmann, 1947, is 11 mm in total length. This size discrepancy p.347. could explain the absence of intercalary plates Chirodota purpurea Theel, 1886a, pp. 15,35, pl. in my material, as they may develop later in the 2, fig. 1; Lampert, 1889, p. 851. life of the animal. DIAGNOSIS: Tentacles 12, usually with five The present specimens closely resemble the pairs of digits each. Calcareous d eposits wheels, type of P. patagonicus when their calcareous de­ which are arranged in papillaeup to 1.5 mm in posits are considered, and thus there is little diameter. Papillae confined to the inrerradii, and doubt that they are in fact examples of Ekman 's are more numerous dorsally, where they are ar­ species. ranged in a single row in each interradius. No DISTRIBUTION : The type specimen (Ekman, other deposits in the skin. Radial muscles con­ 1925) was collected from the Paragonian bank, tain miliary granules. Tentacle rods bracket­ 46° S, at a depth of 110 m. Deichmann (1941, shaped, with branched ends, average length 0.05 1947) states that the species has been taken mm. from the type locality and various other localities MATERIAL EXAMINED: Sra. 73, four speci­ in the Straits of Magellan. Present records in­ mens. dicate that Psolus patagonicus may be quite REMARKS : The total length of the four speci­ common in some localities, its small size per­ mens ranges between 11 mm and .60.mm. Colour haps enabling it to escape notice by the collector. in alcohol dirty white to light brown. There are 12 tentacles, each with five pairs of digits. The ORDER APODIDX -­ wheelpapillae vary considerably in diameter, up to a maximum of 1 rom. Family CHIRIDOTIDAE Dissection of the largest specimens revealed Genus Chiridota Eschscholtz, 1829 the presence of seven Polian vesicles, the largest being 7 rom in length. Rows of closely aggre­ Dactylota Brandt, 1835; Liosoma Brandt, 1835; gated ciliated funnels lie in the middorsal and Tro chinus Ayres, 1852; Lioderma Bronn , left lateral interradii, the rows commencing 1860. about 6 mm from the anterior end of the body DIAGNOSIS: Tentacles 12, digits 3-10 on each cavity, and extending to the extreme posterior side, the terminal pair being the longest. Polian end. The genital tubules are long, sparsely vesicles numerous ( 3-20). Deposits six-spoked branched, and packed with eggs averaging 0.3 wheels collected into small papillae containing mm in diameter. Deichmann (1947 ) has com­ varying numbers of wheels of diverse sizes. No mented on the relationships of this species, and sigmoid rods, but small curved rods with en­ notes that deeper water forms are known which larged ends may be present. Minute miliary may perhaps be assigned to Chiridota purpurea granules often occur in the longitudinal muscles. Theel. TYPE SPECIES: Chiridota discolor Eschscholtz. DISTRIBUTION: Chiridotapisanii is known REMARKS : This is a well-defined genus of from both coasts of southern South America, and world-wide distribution, .eonraining about 25 the Falkland Is., from the intertidal zone to species, none of which have a very extensive about 100 m. geographic range. Most species occur in shallow waters , although some have been taken at depths Genus Trocbodot« Ludwig, 1892 in excess of 3,500 m. DIAGNOSIS: Tentacles 10, digits 2-6 on each side. One Polian vesicle and one stone canal. Chiridota pisanii Ludwig Calcareous ring comprises 10 pieces, the radial Chiridota pisanii Ludwig, 1886, p. 29, pl. 2, fig. unperforated. Calcareous deposits sigmoid hooks, 14; Ludwig, 1898a, p. 71 (complete list of scattered or arranged into groups; and wheels, 0° 0 0 D 0 o 0 o 00 o 0 c 0 -~~-o.v. o 0 o 0 o 0 o 0 o 0 o 0 o 0 o 0 o 0 o 0 e 0 o 0 o 0 o 0 o 0 o 0 00 0° o 0 0 0°0 0 0 ° O·5cm. figs.1,2.

O·1mm. figs. 3,4, 5.

O·5mm.

O·05mm. I '

FIG. 3. 1-7, Psolus patagonicus Ekman. 1, Dorsal view of 9 mm specimen. 2, Ventral view of same speci­ men, showin g disp osition of tube feet. 3, Plates and buttons. 4, Tentacle deposits. 5, Tube foot deposits. 6, Porti on of calcareous ring . 7, End plate of tub e foot. S, T rocbodota purpurea (Lesson) , tentacle deposits. 466 PACIFIC SCIENCE, Vol. XVIII, October 1964 scattered and never grouped into papillae. (After The tentacles in some of the specimens Clark, 1907.) contain a number of bracket-shaped rods with TYPE SP ECIES : T rochodota purpurea (Lesson) . dichotomously branching ends ( Fig. 3, 8) .Per­ REMARKS: This is a well-defined, cosmopoli­ rier ( 1905 ) stated that the tentacles in his tan genus, containing about a dozen species, material were "totally devoid of calcareous de­ some of which are inadequately described. There posits," while Deichmann (1947) , in diagnos­ are three species in Australia, two in N ew Zea­ ing this species, wrote ''. . . no spicules in the land, and one in southern South America. T. tentacles." Ludwig ( 1898a) illustrated a single d eenedinensis from N ew Zealand resembles T . tentacle rod, and it closely resembles those illus­ purpurea from South America in many respects, trated here. There is no doubt, then, that ten­ and the two species may be related. tacle deposits may be present, perhaps in young stages, and become rare or disappear as a speci­ T rochodota purpurea (Lesson) men grows, as can happen with the wheels and H olothuria (Fistularia) purpurea Lesson, 1830, hooks in the body wall. p. 155, pI. 53, fig. 1. DISTRIBUTION: T rochodota purpurea is re­ Chirodota purpurea Jager, 1833, p. 16; Duj ardin corded from the southern tip of South Amer­ and Hupe, 1862, p. 616. ica and the Falkland Is., to depths of about Cbiridota purpurea Brandt, 1835, p. 259. 50 m. Habitat includes sand, shelly bottom, Sigmodota purpurea Studer, 1876, p. 454 (par­ and holdfasts and fronds of seaweed, especially tim ). Macrocystis. Chirodota australiana Theel, '1886a, p. 16. Chirodota studeri Lampert, 1889, p. 839, pI. Genus Taeniogyrus Semper, 1868 XXIV, fig. 12. _S igr(tOdota_Sttlcler, .181 6._ T roChiJddtd jtiideri Ludwig; 1892,-15.359. Sigmodota studeri Oesrergren, 1898. . DIAGNOSIS : Tentacles peltaro-digi tate, 10 or T rochodota purpurea Ludwig, 1898a, p. 83, pI. 12. D igits five to seven pairs per tentacle, the III, figs. 43-45; Perrier, 1905, p. 76; Clark, terminal pair being the longest. Calcareous de­ 1907, p. 123; Clark, 1921, p. 166; Ekman, posits are wheels gathered into papillae, and sig­ 1925, p. 149; Deichmann , 1947, p. 351. moid hooks (about 0.2 mm long), scattered in the skin. N o miliary granules in radial longi­ DIAGNOSIS: Tentacles 10, each with 2-6 pairs tudinal muscles. of digi ts. Wheels (0.13-0.18 mm diameter) TYPE S PECIES: Taeniogyrus australianus scattered in the skin, together with sigmoid Stimpson. hooks (0.12- 0.13 mm long).Tentacle deposits, REMARKS: This is a small genus contai ning when present, average 0.078 mm in length, and about six species. Three species are known from are bracket-shaped, with dichotomously branch­ Austra lia, one from Japan, one from Hawaii, ing ends. Colour in life commonly purple. and one from the southeast Pacific Ocean. MATERIAL EXAMINED: Sta, 73, 8 specimens; Taeniogyrus lies intermediate between Cbiri­ Sta, 74,2 specimens; Sta. 77, 1 specimen. dota (wheels and occasionally curved rods, no REMARKS: The smallest specimen in the col­ sigmoid hooks) and Tro chodota (wheels, and lection is 4 mm in total length, and the largest is sigmoid hooks, the wheels not grouped into 100 mm. Colour in alcohol ranges from off whi te papillae ) . to light brown or violet. The number of tentacle digits varies. Clark (1921 ) in his key to the Taeniogyrus contortus (Ludwig) species of Tro chodota stated that species pur­ purea has tentacles with six digits each. In the Chiridota contorta Ludwig, 1874, p. 80, pI. VI, present collection, one specimen has 12 digits fig. 6; Lampert, 1885, p. 234; Theel, 1886a, per tentacle, and another has eight. Clearly the pp. 16, 33, pI. 2, fig. 2; Theel, 1886b, p. 20; number of tentacle digits is not a reliable diag­ Lampert, 1889, pp . 851,853. nostic character in this species. The calcareous Sigmodota purpurea Studer, 1876, p. 454; Stu­ deposits in the skin have been well described. der, 1879, p. 123. Holothuroidea from Southern Chile-PAWSON 467

Chirodota purpurea Bell, 1881, p. 101; Lampert, osus, Stereoderma laevigata, and Psolus pata­ 1885, p. 236; Lampert, 1886, p. 18, figs. gonicus. 17-20; Ludwig, 1886, p. 29. By far the most common species here was Chirodota studerii Theel, 1886a, p. 33. Pseudocnus dubiosus, which was collected in Chiridota contorta Ludwig, 1892, p. 359; Lud­ numbers at four of the five stations. This species, wig, 1897, p. 217; Ludwig, 1898a, p. 73, together with Cladodactyla crocea and Stereo­ pI. 1II, figs. 37-42. derma laevigata, seems to favour the Macrocystis Sigmodota contorta Ostergren, 1898; Sluiter, zone as a habitat. 1901, p. 134. Seven species were found at Isla N avarin o Taeniogyrus contortus Clark, 1907, p. 122, pI. and the southern regions, at six of the 29 sta­ VII, figs. 8-13; Clark, 1921, p. 165; Ek­ tions worked. They were Pseudocnus dubiosus, man, 1925, p. 147; Heding, 1928, p. 311, Cladodactyla crocea, T rachythyone lecbleri, N e­ text fig. 66, 1-9; Dei chmann, 1947, p. 348. opsolidium convergens, Chiridota pisanii, T ro­ chodota purpurea, and Taeniogyrus contortus. DIAGNOSIS: Tentacles 12, with five to seven Of these seven species, three are apodous pairs of digits. Wheels (diameter 0.042-0.13 forms. In an account of the marine work carried mm) gathered into well-defined papillae; sig­ Out by the Royal Society Expedition, Prof. Knox moid hooks large (0.14- 0.2 mm long ) scattered (personal communication) noted that "one of in the skin. Tentacle rods 0.17 mm in length . the salient features of the southern region is the MATERIAL EXAMINED: Sta. 73, one specimen. reduction which has occurred in the number of REMARKS: The single specimen is 50 mm in species present," and he cited the case of species total length and has only 11 tentacles. Colour in of Brachyura (Crustacea) , of which there were alcohol is a dull, dark brown. The systematic 15 at Chepu (Isla Chiloe ), and only two-in the .. position of T aeniogYrttscon~ortus is now quite southern regions. This .state" of "affairs clearly " clear as a result of the work of Clark (1907, does not apply to the holothurians, as at least 1921 ) and Heding (1928 ). 14 species are known from the south ern region, DISTRIBUTION : The species is known from while approximately seven have been recorded the south ern tip of South America, Falkland Is., from the Isla Chiloe area. South Georgia, Burwood Bank, and Kerguelen Between Isla Chiloe and Isla Navarino the Is., to depths of about 200 m. Sluiter ( 190 1) character of the holorhuri an fauna changes, and recorded Sigmodota contorta from the Java Sea the number of species gradually increases. Th ere at a depth of 82 m. Clark ( 1921) doubted the are no drastic changes which might indicate a accuracy of the identification, and it might well provincial pattern, although the number of spe­ be that Sluiter's material represents yet another cies shows a definite increase at about 52° S, species. Fisher (1907) found a close relative to immediately north of the western entrance to T. contortus in Hawaii. the Straits of Magellan. Apodous species (e.g., T rochodota purpurea, DISCUSSION Chiridota pisanii, and Taeniogyrus contortus) 1. Th e H olothurian Fauna of Southern Chile are more numerous in the southern regions than elsewhere on the Chile coast, but there is no Collections made at 18 stations in the Isla apparent reason why this should be the case. Chiloe area revealed only one species, namely Th e fauna of Chile contains a remarkably Athyonidium cbilensis, while at least seven shal­ high percentage of cucumariids (ca. 40% ), low-water species are known from this region. compared with that of N ew Zealand (33 %) Athyonidium chilensis is one of the most con­ and Australia ( 22% ). This is a striki ng feature spicuous holothurians in Peru and Chile, and Isla of the fauna, as also is the virtual absence of Chiloe represents the extreme southern limit of phyllophorids below about 42° S, whereas New the range of the species. Zealand has 15% , and Australia 14 %. From Puerto Eden to Punta Arenas, 31 sta­ tions were worked, and holothurians were taken 2. Relationships of the Fauna from five stations. They were Pseudocnus dubi- A. With Australia, N ew Zealand , and the 468 PACIFIC SCIENCE, Vol. XVIII, October 1964

Antipodean Province Islands: The holothurian Stereoderma laevigata is also known in the fauna of southern Chile bears no close relation­ fauna of Marion Island and the Crozets. ship to that of New Zealand and Australia. A When considered overall, these islands have single species, Paracaudina cbilensis, is common faunas which are similar to that of southern to New Zealand and southern Chile, and this Chile, and the similarity is at the specific level. species is a circum-Pacific eurybath form. Ocnus The dispersal of species to these widely sepa­ calcareus is known from New Zealand and Juan rated areas is probably effected epiplanktonically Fernandez, but has not as yet been recorded (perhaps on rafts of seaweed), with the aid of from the Chilean coast. The Antipodean Prov­ the westwind drift. This is quite conceivable ince Islands (Auckland Is., Campbell Is. in par­ for such species as Stereoderma laevigata and ticular) of New Zealand .suppOrt a holothurian Cladodactyla crocea, which commonly live on fauna of New Zealand character, and thus the the fronds and holdfasts of Macrocystis, a brown fauna differs fundamentally from that of other seaweed which is found on all of the islands southern islands such as Kerguelen, Macquarie, mentioned above. The westwind drift dispersal South Georgia. However, one Antipodean Prov­ mechanism for echinoderms has already been ince species, Stereoderma leoninoides (Mor­ discussed by Mortensen (1925), Fell (1953, tensen) is a close relative of Stereoderma 1962), and Ekman (1953), and others. laevigata from southern Chile, and it is quite The holothurian fauna of southern Chile is a possible that these two species may have been generalised fauna, containing few restricted spe­ derived from some common source. cies, and notable because of the number of spe­ B. With that of Antarctica: Some species, in­ cies which are shared with distant islands, and cluding Cladodactyla crocea, Stereoderma laevi­ the Antarctic continent. gata, Trachyth yone parva, and Psolus antarcticus, are shared with the -fauna of Antarctica. "Fell REFERENCES (1961), in discussing the Ophiuroidea of Ant­ arctica, stated that the Magellanic ophiuroid BELL, F. J. 1881. Echinodermata. In: Account fauna is predominantly of southern American of the Zoological Collections made during the type, mingled with a few Antarctic species Survey of the H.M.S. "Alert" in the Straits of which are eurythermal. The four species men ­ Magellan and on the coast of Patagonia. Proe. tioned above are probably eurythermal and can Zool. Soc. London., 1881: 100-101. readily survive the difference in temperature be­ --- 1908. Echinoderma. National Antarctic tween Antarctica and southern Chile. Expedition 1901-04. Nat. His!. IV. London. C. With that of the Subantarctic Islands: The BRANDT, J. FR. 1835. Prodromus Descriptionus Falkland Islands (Islas Malvinas) have a holo­ Animalium ab H. Mertensio Observatorum. rhurian fauna which is scarcely distinguishable Fase. 1. Petropoli. from that of southern Chile. This is understand­ CLARK, H. L. 1907. The Apodous Holothurians. able when one considers that the islands are but A Monograph of the Synaptidae and Mol­ 300 miles east of Tierra del Fuego, in a good padiidae. Smithson. Contr. Knowl., 35:1-231, position to accept species carried from southern pIs. I-XIII. South America by the westwind drift. - -- 1921. The Echinoderm Fauna of Torres Among the Kerguelen Island holorhurians Strait: Its Composition and Origin. Carnegie are Cladodactyla crocea, Stereoderma laevigata, Inst. Wash ., Pub: 214, Dept. Marine BioI. 10. Trachythyone parva, and Ta eniogyrus contortus. viii + 224 pp., 38 pIs. Thus the fauna here also bears a remarkable - -- 1946. The Echinoderm Fauna of Aus­ likeness to that of southern Chile. tralia : Its Composition and Origin. Carnegie South Georgia, regarded as a separate biotic Insr. W ash., Pub. 566. 567 pp. province by Knox (1960 ), shares Cladodactyla DEICHMANN, E. 1941. The Holothurioidea col­ crocea, T rachythyone parva, Stereoderma laevi­ lected by the "Velero III " during the years gata, N eopsolidium convergens, Taeniogyrus 1942 to 1948. Part I, Dendrochirota. Reprs , contortus, and (with doubt) Anapta fallax with Allan Hancock Pacific Exped. 8 ( 3) :61-194, southern Chile. pIs. 10-30. Holorhuroidea from Southern Chile-PAWSON 469

--- 1947. Shallow water holothurians from LAMPERT, K. 1885. Die Seewalzen. Hol othuri­ the Cabo de Hornos and adjacent waters. An. oidea. Eine systernatische Monographie. W ies­ Mus. Argent. B. Aires. 42:325-351, figs. 1- 5. baden. DU] ARDIN, M. F., and M. H . H UPE. 1862. His­ --- 1886. Die Holothurien von Sud-Georg­ toire N aturelle des Zoophytes Echinodermes. ien nach der Ausbeute der Deutschen Polar­ Pp. 609-62 2, pI. viii. Paris. station in 1882, 1883. Jahrb. Hamb. W iss. EKMAN, S. 1925. Holothurien , Further Zool. Anst. 3. Results of the Swedish Antarctic Expedition, --- 1889. Die "Gazelle" Holothurien. ZooL 1901-03,1 ( 6 ) :1-194, 37 figs. Jb . 4:806---839, pI. 1. ---195 3. Zoogeography of the Sea. London: LESSON, R. P. 1830. Cenrurie Zoologique. Paris. Sidgwi ck and Jackson Ltd. LUDWIG, H. 1874. Beitrage zue Kennrnis der FELL, H. B. 1953. The origins and migrations of Holorhuri en. Arb. Zool.-Zoot. Inst. Wurz­ Australasian echinoderm faunas since the Me­ burg. 2:77- 120, pIs. 6, 7. sozoic. Trans. Roy. Soc. N. Z. 81 (l ) :245­ --- 1886. Die von G. Chierchia auf der 255. Fahrt der Kgl. Ital. Corvette "Vertor Pisani " --- 1961. The Fauna of the Ross Sea. Part gesammelten Holothurien, Zool. Jb. 2:1- 36, I, Ophiuroidea. N. Z. Oceanogr. Inst. Mem. pIs. 1,2. 18: 79, 9 figs., 19 pIs. --- 1887. Drei Mittheilungen tiber alte und --- 1962. W est-wind-drift dispersal of echi­ neue Holothurien-arten. Sitzungsber. Akad. noderm s in the Southern H emisphere. N ature Berlin: 1217-1244, pI. 15. 193 (4827) :759'-761. --- 1892. Echinodermen. Die Seewalzen. In: FISHER, W. K. 1907. The Holothurians of the Bronn, Klassen und Ordnungen des Thier­ Hawaiian Islands. Proc. U. S. N at. Mus. 32: Reichs 2 (3) :1-460, pIs. 1-17, Leipzig. 637=744, pls.: 6&:-82. --.-.--. 1897. Ein rieiierFall vonBrutpflege bei HEDING, S. G. 1928. Synapt idae . . . from Dr. Holorhurien. Zool. Anz. 20:217-219. Mortensen's Pacific Expedition. V i densk. --- 1898a. Holothurien. In: Ergebnisse der Medd. dansk. naturh. Foren. Kbh. 85:105­ Hamburger Magahlaensische Sammelreise, 98 323, pIs. 2, 3. pp., 3 pIs. --- 1931. Dber die Synaptiden des Zoolo­ --- 1898b. Die Holothurien der Sammlung gischen Museums zu Hamburg. Zool. Jb. 61 : Plate. Zool. Jb ., Suppl. IV (2 ) :4 31-454, pI. 637-696, pI. 2. XXVI. --- and A. PANNING. 1954. Phyllophoridae. MORTENSEN, TH. 1925. The echinoderms of Eine Bearbeitung der polytentaculen Holo­ N ew Zealand and the Auckland-Campbell thurien des Zoologischen Museums in Kopen­ islands. Part IV, Holothuroidea. Vidensk. hagen. Spolia Zool. Mus. Haun iensis XIII: Medd. dansk. naturh . Foren. Kbn. 79 :322­ 1-209, 102 figs. 386, figs. 22-62. HELFER, H. 1917. Beitrag zur Kenntnis von OSTERGREN, H]. 1898. Das System der Synap­ Cucumaria laevigata Verrill nebst Beschrei­ .tiden (Vorlaufige Mitteilung ). Ofv. Kong. bung einer neuen Abart dieser Holothurie. Vet. Ak. Forhandl. LV: 111-120. Sitz.-Ber. d.Ges. naturf. Freunde Berlin . Jg. PANNING, A. 1940. Dendrochirote Holothurien 1917. von Dakar. Vidensk. Medd. dansk. natu rh. H EROUARD, E. 1901. N ote prelimin aire sur Ies Foren. Kbh. 104:169- 178, figs. 1-7. Holothuries . . .Belgica. Arch. Zool. Exp., --- 1949. Versuch einer N euordnung der N otes, ser. 3 (9): pp. xxxix-xlviii. Famil ie Cucumariidae ( Ho lothurioidea, Den­ --- 1906. H olothuries. Expedit. Anrarc, drochirora ) .Zool. Jb . 78 (4) :404-470. BeIge. Res. Voy. S. Y. Belgica. --- 1950. Uber Pseudocnus leoninus (Sem­ JAGER, G. FR. 1833. De Holothuriis, Dissert, per) und verwa ndte Arten. Zool. Anz. 146: inaug. Pp. 1- 40, pIs. i-iii. Turici. 73-80, figs. 1-7. KNOX,G. A. 1960. Littoral ecology and bio­ --- 1952. Bemerkungen tiber Holothurien geography of the southern oceans. Proc. Roy. aus dem N atur-Museum Senckenberg. Sene­ Soc. ( B) , 152:577-624. kenbergiana 33: 123- 133, figs. 1-15. 470 PACIFIC SCIENCE, Vol. XVIII, October 1964

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