JOURNAL OF THE ROYAL SOCIETY OF WESTERN AUSTRALIA VOLUME 53 PART 4 1970 REGISTERED AT THE G.P.O., PERTH FOR TRANSMISSION BY POST AS A PERIODICAL THE ROYAL SOCIETY OF WESTERN AUSTRALIA PATRON Her Majesty the Queen VICE-PATRON His Excellency Major-General Sir Douglas Kendrew, K.C.M.G., C.B., C.B.E., D.S.O., Governor of Western Australia COUNCIL 1970-1971 President Vice-Presidents Past President Joint Hon. Secretaries Hon. Treasurer Hon. Librarian Hon. Editor B. J. Grieve, M.Sc., Ph.D., D.I.C., F.L.S. G. M. Storr, B.Sc., Ph.D. R. M. Berndt, M.A., Dip.Anth. (Sydney), Ph.D. (London) , F.R.A.I., F.F.A.A.A. P. E. Playford, B.Sc., Ph.D. B. Ingram, B.Sc. (Hons.) P. G. Wilson, M.Sc. R. N. Hilton, M.A. Ariadna Neumann, B.A. A. S. George, B.A. S. D. Bradsha\v, B.Sc. (Hons.), Ph.D. S. J. Curry, B.Sc. A. B. Hatch, M.Sc., Dip. For. J. H. Lord, B.Sc. D. C. Lowry, M.Sc. A, J. McComb, M.Sc., Ph.D. D. Merrilees, B.Sc., Ph.D. R. T. Prider, B.Sc., Ph.D., M.Aust.I.M.M., F.G.S. 10. — The Australian Majid spider crabs of the genus Achaeus (Crustacea, Brachyura) by D. J. G. Griffin* Manuscript received and accepted 17 February, 1970 Abstract The genus Achaeus is represented in Austra- lia by eight species. A. brevirostris (Haswell) and A. lacertosus Stirapson. widespread around western, northern and eastern Australia, and the Iiido-west Pacific are redescrlbed and figured. A. fissifrons, also widely distributed in the Indo-west Pacific, is recorded from West- ern Australia for the first time and A. pugnax (De Man), previously known only from Japan, is recorded from Western Australia. Three new species are described, two from eastern Aus- tralia and one from Western Australia; the identity of three small specimens of an addi- tional species are discussed — they are not con- specific with any Australian species known at present. A. brevifalcatus Rathbun. known from the Seychelles, western Indian Ocean, and from Hawaii, is figured and additional descrip- tive notes are given. Introduction The small, long-legged spider crabs of the genus Achaeus were reviewed by Griffin &; Yaldwyn (1965) who provided a key to the Australian species knowm at that time and re- described A. fissifrons (Haswell) and commented in detail on the synonymy of the species. The key was repeated by Gi'iffin (1966a, 1966b) and the description by Griffin (1966a). Apart from A. fissifrons (see Griffin & Yaldwyn 1965: 45) show’s that there is good agreement and Iranian Gulf material is con- sidered to be conspecific wdth Australian. The characteristic features of A, lacertosus are the smooth carapace and the spinulate ros- trum bearing curled hairs. Distribution: Western, northern and eastern Australia from Broome (WA) to Port Jackson (NSW); Indo-west Pacific from South Africa to Japan; 3-45 fms. Buitendijk (1950) recorded the species from Paw’ai Island near Singapore. Achaeus paradicei n.sp. Figs. 3b. 6. 15b, c. Achaeus sp. Griffin & Yaldwyn, 1965: 33. Griffin. 1966a: 38: 1966b: 276. Material examined: A total of 10 specimens as follows : Holotype — Male, 5.4 mm, southern Coral Sea. 26°33'S.. 153'’31'E.. dredged, 86 metres, gravel. 5 11 1951, “Galathea” Expedition 1950-52 Sta. 539 (CM). Paratypes — Two males, six female (5 ovig.). 4. 0-5.0 mm, smallest ovig. 9 4.5 mm (many with legs detached), same data as for holotype *1 male. 1 female now reg. AM P.17517; remain- ing paratypes from this series CM). One female (ovig.), 4.5 mm, Gibson Reef, off Cairns, Queensland, 28 fms, before 1925. W. E. J. Paradice (AM P.7985 — specimen in spirit, some legs detached but in same tube as rest of animal). Description: Carapace moderately elongate, length almost 1^ times greatest breadth, mar- gins with a few stout, blunt spines or tubercles, dorsal surface with 11-12 blunt tubercles, other- wise smooth. Branchial and cai’diac regions well demarcated by broad groove from surround- ing regions, regions otherwise ill-defined. Culled hairs grouped along base of rostral spines and laterally on hepatic and branchial regions. Rostral spines very short, separated by broad V-shaped notch in distal third, rounded apically. up to four blunt spinules on margins, most apical the largest and sharpest. Supraorbital eave bearing along outer edge, at least posteriorly, many minute blunt spinules; some spinules immediately behind eave. A short, slender, curved spine at posterolateral corners of epistome partly visible in dorsal view behind eave. Eyestalks long and stout, a prominent flattened triangular process midway along anterior edge, a small blunt tubercle opposite on ventral surface and a prominent tubercle above cornea at distal extremity of eyestalk; cornea large, circular, obliquely sub- terminal. Region betw'een eave and hepatic region broadening regularly, unconstricted. Hepatic regions weakly inflated, laterally acute, with a group of two to four broad, stout spines. Pterygostomian regions with a large tubercle posteiolaterally and visible in dorsal view behind hepatic region. Branchial regions, sw’ollen, three or four acute spines just forw’ard of wddest part of carapace on margin, first tw^o larger than others and a few small tubercles sometimes behind these. Pos- terolateral and posterior margins bordered by minute, sharp spinules. Dorsal surface of carapace with a low’, broad- based, conical spine far back on mesogastric region, tumid cardiac region w'ith two blunt tubercles side by side and a small medial in- testinal tubercle sometimes on posterior slope. Two large tubercles on protogastric regions laterally forw'ard of mesogastric spine. Six similar tubei'cles laterally on branchial regions, three on each side in a semi-circle, one an- teriorly, one opposite cardiac tubercle, and one near posterior margin above base of last am- bulatory leg. Antennular fossae large, anterolaterally somewhat outwardly splayed. Basal segment of each antennule bearing three or four small, sharp spinules near distal edge, Interantennular partition a narrow’ compressed triangular lobe. Basal antennal articles terminating in a large, sharp spine directed anteriorly, lateral edge bearing six to eight small, sharp spines, three more spines along medial edge. Antennae as long as carapace, first segment with three or four spinules ventrally. Epistome slightly longer than wide, a spine laterally just outside and forward of opening of green gland with a smaller spine sometimes immediately in front of it. Ischium of third maxillipeds with two distally divergent, oblique, ill-defined rows of spines on each side of a shallow longitudinal groove: medial edge minutely toothed. Merus with tw'o subparallel, longitudinal rows of spines on each side of shallow' groove which extends 2/3 of length tow'ards distal edge; distal edge irregu- larly crenulate and lobed, a few small spines slightly laterally. Palp long and stout, later- ally, medially and apically fringed by long hairs; carpus bearing a strong, slender spine medially towards distal edge, propodus with a similar ventral spine about midway along its length. 108 Expedite bearing a longitudinal row of small tubercles along its outer surface. Thoracic sternum in male densely covered by small tubercles. Chelipeds in male long, about li times cara- pace length, granular and densely tuberculate, chelae robust. Ischium subtrigonal. merus sub- trigonal to subcylindrical. carpus subcylindrical, chela compressed. Ill-defined ventromedial and ventrolateral edges of merus with rows of irreg- ularly sized blunt and shai'p spines. Carpus with several small spines medially and laterally and a group of about 4 larger tubeicles proxim- ally on dorsal surface. Distal articulating pro- cesses dorsally and ventrally spinulate. Chela with palm little longer than high, bearing dorsally and ventrally irregularly sized spinules or tubercles which are larger dorsally; spinules extending along proximal part of fingers dors- ally and ventrally. Fingers as long as palm, in- curved distally and acute; inner edges toothed; fixed finger with a large tooth at base and two more teeth, the first the smaller and sometimes obsolete, close together midway along and separated from basal tooth by a broad hiatus; dactyl with two large teeth not far from base and fitting into hiatus between basal and re- maining teeth of fixed finger; distal parts of both fingers with irregular small teeth along inner edges which meet apically. Ventromedial edge of merus, medial surface of carpus and ventral edge of chela with large and small straight hairs. Cheliped of female no longer than carapace, slender, merus subtrigonal: spines of merus, carpus and propodus much longer than in male, in particular a comb-like row of long spines along ventrolateral edge of merus: fingers as long as palm, almost meeting along cutting edges, weakly toothed, teeth larger and closer together distally. Ambulatory legs very long and slender, first the longest, about 3^ times carapace length, remaining legs decreasing regularly in length, fourth the shortest, about twice carapace length; curled hairs singly along carpus and propodus dorsally- Dactyls of first two legs long and almost straight, weakly curved distally only and unarmed, bearing long straight hairs: dactyls of third and fourth legs falcate, bearing sharp, recurved spines ventrally for almost entire length, spinules very small or occasionally obsolete in proximal half, larger in distal half except last one or two which are slightly smaller. Male abdomen with all segments, except last, wider than long, last segment the longest, as long as wide. Third segment, with strongly convex lateral edges, last segment subpentagonal distally subtruncate. Surface elevated in mid- line, bearing a tubercle on first segment, a wide elevation distally on third to fifth segments and on the last a central tubercle and a transverse pair of smaller tubercles not far from distal edge; third segment laterally inflated with spinate surface. Female abdomen broad, elongate subovate, elevated in midline. Male first pleopod moderately slender except for more bulbous base, tip expanded, curved medially and subtruncate ; aperture subterminal, a distal slit at end of groove extending along lateral surface for distal half; a fringe of short hairs distally along medial surface, a few short hairs midway along medial surface, and a few long plumose hairs at base medially and later- ally. Measurements : Male ( holotype ) — -carapace length 5.4 mm, carapace width 3.8 mm, rostrum length 0.8 mm, rostrum width 0.8 mm, cheliped length 6.2 mm. chela length 3.5 mm. chela height 1.3 mm, dactyl length 2.0 mm. Female (paratype) — carapace length 5.0 mm, cheliped length 6.5 mm. chela length 2.9 mm, chela height 0.8 mm. Female (paratype) — carapace length 4.8 mm. first ambulatory leg length 15.5 mm. Remarks: In general shape and ornamenta- tion of the carapace and arrangement of teeth on the chela in the males, A. paradicei closely resembles A. hrevirostris. However, there are a number of important differences. A. paradicei is a much smaller species, reaching maturity las judged by expansion of male chelae and female abdomen) at a cai'apace length of 4.5 mm 16.8 mm minimum in A. hrevirostris) , the males have a much shorter and less constricted postorbital ‘neck’, the rostral lobes are separated by a V-shaped notch (usually U-shaped in A. hrevirostris > and are apically armed with spinules (absent in A. hrevirostris), the edges of the supraorbital eaves bear numerous spinules (absent in A. hrevirostris), the exopodite of the third maxilliped bears a row of spinules (smooth in A. hrevirostris), there is a spine and a tubercle on the epistome close to the opening of the green gland (a single small tubercle in A. hrevirostris) , there are three or four prominent tubercles in a row immediately below the branchial margin anterioi'ly that this previously unde- scribed species existed in Australia. Distribution: Eastern Australia from off Cairns to off Noosa Head (Qld), 28-44 fms. Achaeus podocheloides n.sp. Figs 7a, 8. 14e, f. Material examined: A total of four specimens as follows: Holotype— M3.\e, 8.1 mm, N.W. of Jurien Bay Western Australia, 30°00'S., 114°22'E., 70-75 fms 28/1/1964, HMAS “Diamantina” Cruise 1/64 CSIRO Sta. 3 (WAM 95-67). 109 Paratypes — 1 female, 6.5 mm, S.W. of Point Cloates, W.A., 23°39'S., 113°11'E., 73 fms, 7 10 1963, HMAS “Diamantina” Cruise 6/63, CSIRO Sta. 187 (WAM 331-67). 1 female, 4.8 mm, N.W. of Carnarvon, W.A., 24°59'S., 112°27'E., trawled, 71 fms, 8/10 1963, HMAS “Diamantina” Cruise 6/63, CSIRO Sta. 197 (AM P.16779— ex WAM 276-67 part). 1 male, 7.8 mm. N.W. of C. Naturaliste, W.A., 33^40'S., 114°28'E.. 75 fms, 27-28/7/1963. HMAS “Diamantina” Cruise 4/63, CSIRO Sta. 134 (AM P.16774— ex WAM 332-67). Description: Carapace elongate, length slightly more than 1| greatest breadth, lateral margins with a few sharp spines and spinules, dorsal surface with 11 blunt tubercles. Branchial and cardiac regions well demarcated by broad grooves from surrounding regions, other regions ill- defined. Curled hairs scattered over lateral parts of carapace. Rostral spines short, separated by very broad V-shaped notch in distal third, each terminating in a small weakly upturned spine. S'upraorbital eave unarmed. Eyestalks long and stout, two spinules about midway along anterior edge, more distal the larger, proximal one somewhat ventral, a prominent tubei’cle above coi*nea at distal extremity of eyestalk; cornea large, circular, obliquely subterminal. Region between eave and hepatic region broadening regularly, weakly constricted, two to four spinules not far behind eave. Figure 7. — a, Achaeus podocheloides, n.sp., HOLOTYPE, male, carapace, dorsal view; b, Achaeus pugnax (De Man), male, 5.5 mm, S.W. of Pt. Cloates, W.A. (WAM 125-67), carapace, dorsal view. 110 Hepatic regions inflated, laterally acute, with a group of two to four stout spines and spinules. Pterygostomian regions with a small tubercle posterolaterally and visible in dorsal view be- hind hepatic region. Branchial regions swollen, margins with a small spine anteriorly: posterolateral margins bordered by spinules. Dorsal surface of carapace with a low, broad- based, conical tubercle far back on mesogastric region, cardiac region with two blunt tubercles submedially surmounting prominent elevation and a very obscure medial tubercle on posterior slope. Two blunt tubercles submedially on pro- togastric region forward of mesogastric tubercle; three tubercles laterally on branchial regions in a curved row. first anteriorly, second opposite cardiac elevation, third tubercle near posterior margin above base of each ambulatory leg. Antennular fossae large. Basal segment of each antennule bearing four or five spinules near base. Interantennular partition narrow. Basal antennal articles with two or three spinules distally. Antennae almost as long as carapace, first segment with two or three spinules distally. Epistome slightly longer than wide. Ischium of third maxillipeds with two or three sharp spines on lateral side of a shallow longitudinal groove and up to five similar spines medially: medial edge minutely toothed. Sur- face of merus unarmed, two to four spines on medial edge, distal edge with a few small spines slightly laterally. Palp long and stout, laterally, medially and apically fringed by long hairs; carpus bearing a strong, slender spine medially towards distal edge, propodus with a similar spine distally. Exopodite smooth. Thoracic sternum in male with a few small tubercles laterally and several around anterior edge of abdominal fossa. Chelipeds in male long, chelae moderately robust. Ischium subtrigonal, merus subtrigonal to subcylindrical, chela compressed. Ill-defined Figure ^.—Achaeus podocheloides n.sp., HOLOTYPE, male, a, front of carapace, ventral view; b abdomen- c, right chela; d, left fourth ambulatory dactyl, posterior view; e. left third maxilliped. Ill ventromedial and ventrolateral edges of merus with a few blunt and shai’p spines proximally and distally on ventrolateral edge and proxim- ally on ventromedial edge; three slightly longer spines equally spaced along dorsal surface. Carpus with several small spines laterally, some denticles medially and a group of about four larger tubercles proximally on dorsal surface. Chela with palm twice as long as high, bearing three or four irregularly sized spinules dorsally. Fingers as long as palm, incurved distally and acute; inner edges adjacent, coarsely toothed: dactyl with a large tooth near base. Ventro- medial edge of merus. dorsomedial surface of carpus and ventral edge of chela with large and small straight hairs. Chelipeds of female no longer than carapace, extremely slender, weakly spinous, ventrally and dorsally hairy, chela of uniform width throughout, fingers as long as palm, meeting along weakly toothed cutting edges. Ambulatory legs very long and slender, first the longest, almost four times carapace length, remaining legs decreasing regularly in length, fourth the shortest, almost three times cara- pace length: long straight hairs and curled hairs singly along merus, carpus and propodus dorsally. Dactyls of first two legs long and almost straight, weakly curved distally and un- armed. bearing long straight hairs: dactyls of third and fourth legs faclate. bearing sharp recurved spines ventrally for almost entire length. Male abdomen with all segments wider than long, last the longest, almost as long as wide. Third segment with strongly convex lateral edges, last segment subtriangular, distally rounded. Surface elevated in midline, bearing a short spine on first segment, a wide elevation distally on third to fifth segments and on the last a central tubercle; third segment laterally inflated, smooth. Female abdomen broad, elongate subovate, elevated in midline. Male first pleopod bulbous basally, more slen- der midway along, weakly expanded in distal half and weakly curved, tip blunt: aperture subterminal, a long slit on medial surface at end of groove extending along medial surface and partly covered by a transpai-ent projection on sternal surface: a few long plumose hairs at base laterally, surface otherwise naked. M easurements : Male ( holotype ) — carapace length 7.9 mm, carapace width 4.6 mm, ros- trum length 1.0 mm. rostrum width 1.2 mm cheliped length 12.3 mm, chela length 5.6 mm, chela height 1.9 mm, dactyl length 3.3 mm, first ambulatory leg length 32.0 mm. fourth ambulatory leg length 24.5 mm. Female fparatype> — carapace length 6.5 mm. cheliped length 8.0 mm, chela length 4.4 mm. chela height 0.9 mm. dactyl length 2.2 mm. Remarks: This species resembles A. brevifal’ catus in several features including pointed ros- tral spines, cardiac region surmounted by a pair of tubercles, postorbital region with spinules. posterolateral border of carapace with spinules. eyestalk with two small tubercles or spines, merus of third maxillipeds with spines along medial edge, dactyls of last ambulatory legs falcate and ventrally spinulate for more than half their length and abdomen of the male with a single tubercle on the last segment. However, the two species differ markedly in several features, the most important of which is the form of the male first pleopod. In A, brevifalcatus the general shape and position of the aperture is much the same as in the majority of Achaeus species. In A. podoche- loides, on the other hand, the aperture is sub- terminal and pai'tly surrounded by a flap of tissue; such a pleopod is typical of American inachines such as Podochela species (where the flap is greatly developed). Other differences between A. brevifalcatus and A. podocheloides are the presence of spinules on the supraorbital eave in A. brevifalcatus but not in A. podoche- loides, the more pronounced development of spinules behind the eave in the former, the presence of protogastric and branchial tubercles in A. podocheloides, the paucity of spinules on the basal antennal article and the absence of spinules close to the posterolateral margin of the carapace in A. podocheloides whereas A. brevifalcatus has numei'ous spinules on the basal antennal article and on the posterolateral regions of the carapace near the border in addition to those on the border. Finally, in A. brevifalcatus the lateral margins of the third segment of the male abdomen are spinate proximally (smooth in A. podocheloides f and the fingers of the chela in the male gape widely in the proximal ^ to 2 3 and two prominent teeth project into the gape (in A. podocheloides the fingers gape only slightly). A. podocheloides is similar to a new species described elsewhere by M. Takeda (pers. comm.) from a single male taken in 200 metres at Ogasawara I.. Japan. The resemblances are particularly noticeable in carapace shape and ornamentation and in the form of the first pleopod of the male. However, the Japanese species differs in having more acuminate rostral spines, a strong spine on each eyestalk. spinules on the supraorbital eave anteriorly, more spines on the third maxilliped. a longitudinal elevation on the outer surface of the palm of the chela and the terminal segment of the abdomen in the male is sharp. There are possibly other more minor differences also. Distribution: Western Australia from off Point Cloates to just north of Cape Naturaliste, 70-75 fms. Achaeus brevifalcatus Rathbun, 1906 Figs 9, 10. 14b, c. Achaeus affinis: Rathbun. 1906:877, Not Achaeus affinis Miers, 1884 t — A. brevirostis (Haswell) ). Achaeus brevifalcatus Rathbun. 1911: 244-246. fig. 2. Material examined: At the time of the descrip- tion of this species Rathbun had available a male and a female taken by the “Sealark" and The material from Hawaii, which she considered to be conspecific with the “Sealark” specimens, reported uDon previously (Rathbun. 1906) as Achaeus affinis. No statement was made as to which was the holotype. I choose as LECTO- TYPE the male. c.l. 6.6 mm (USNAI 41380), 112 Figure 9 . — Achaeus brevifalcatus Rathtaurn, LECTO- TYPE, male, carapace dorsal view. taken by the “Sealark” from the Seychelles and figured by her; details are as follows: Seychelles, western Indian Ocean, 44 fms, H.M.S’. '‘Sealark’’, 20/10/1905, Percy Sladen Trust Expedition Sta. F5. The female (USNM 41397) from Sta. F4, Seychelles, 39 fms, 20/10 1905 is designated PARALECTOTYPE. The Hawaiian specimens have also been re- examined: details are as follows: Hawaiian Islands as weakly curved distally. This species shows considerable variation in spinulation. The spinules on the supraorbital eave are sometimes absent, the sides of the large spines of the carapace are sometimes smooth and the spines above the last leg are sometimes topped by a group of spinules. The carapace has a shorter and less constrict- ed “neck” than the male as is usual in the genus. Distribution: Western Australia: previously known from south-eastern Japan from Sagami Bay to Koshiki Islands near Kuysyu; 43-75 fms. Achaeus sp. Pigs 12, 13e. f. Material examined: A total of three specimens as follows: Westerji Australia: N.W. of Point Cloates. 22°52'S., 113°29'E., triangle dredge, 73 fms. 6 10 1963, HMAS “Diamantina” Cruise 6/63. CSIRO Sta. 178, 1 c5, 4.5 mm, 1 $ , 3.8 mm 'WAM 71-67). N.W. of Bluff Point, 27'^18'S.. 113°16'E., triangle dredge, 54 fms, 9/10/1963 HMAS “Diamantina” Cruise 6/63, CSIRO Sta 204, 1 9 (ovig.), 4.6 mm (WAM 130-67) Remarks: A detailed description of these specimens would be premature in view of the few specimens and the absence of chelipeds from the male and of most legs from all speci- mens. The three specimens are generally similar to A. lacertosus but differ in the following features: (1) The carapace is covered by close set spinules except anteriorly and there is a metagastric tubercle or spine plus a transverse cardiac pair of tubercles about the same size; (2) the rostral spines are without spinules; (3) the supraorbital eaves bear close set spinules along their outer margins and there are a few spinules behind: (4) the eyestalks each bear three sharp spines on the anterior surface; (5) the basal segment of each antennule bears four spines in an oblique row: (6) the basal antennal articles bear a row of spinules along both the lateral and medial edges and terminate in a strong anterolateral spine; (7) the ischium of the third maxillipeds bears a longitudinal row of spinules on each side of the longitudinal central groove and the merus bears long spines along the medial edge; (8) the thoracic sternum in the male is spinulous; (9) the abdomen of the male has stronger medial tubercles, with two prominent ones in the midline on the last seg- ment: (10) the first pleopod of the male is more expanded distally than is that of A. lacertosus and there are only short hairs near the tip; the aperture is pro- tected by a small lateral flap. The smaller female possesses chelipeds. These are slender but short and the ischium, merus and propodus are provided with long and short spines along the dorsal and ventral surfaces and there are some short spines on the cai'pus dorsally. The male has the tip of the dactyl of one ambulatory leg caught among the hairs of the carapace. It is weakly curved with a few short spines ventrally towards the tip. Two ambula- tory legs, probably anterior ones, are associated with the specimens. They are provided with long hairs and the dactyl is weakly curved and unarmed. It is possible that these specimens may belong to A. rohustus Yokoya to which there are gen- eral resemblances in carapace shape and orna- mentation and also in the first pleopod of the male (M. Takeda, pers. com.). ( Figure 12.— Achaeus sp.; male, 4.5 mm, N.W. of Point Cloates, W.A. (WAM 71-67). a, carapace, dorsal view; b, front of carapace, ventral view. 117 Figure 13. — Male left first pleopods of Achaeus galu- theae, n.sp., A. fissifrons (Haswell) and Achaeus species, a. d. A. galatheae, PARATYPE, 3.3 mm (CM): b, c. A. fissifrons, 10.0 mm, Port Jackson, N.S.W. (AM P.1442); e, f. Achaeus sp., 4.5 mm (WAM 71-67). a, c. e, abdominal surface; b. d. f. sternal surface. Discussion The genus Achaeus appears to comprise two or possibly three subgroups. In the majority of the species the first pleopod of the male is weakly curved distally and the aperture is a simple subterminal or sometimes terminal slit. In a second group. A. podocheloides, a new species from Japan and the unindentified species discussed in this report, the subterminal aper- ture is protected by a membranous flap, a feature not commonly found in Indo-Pacific machines but known in a number of Atlantic and east Pacific forms and several Indo-Pacific majines. In a third gz'oup, comprising A. sulu- ensis and A. galatheae the pleopod is strongly twisted and quite different from that of any other knowm majid. These differences generally are not well correlated with other morphological features, however. The seven named Australian species fall into three distributional groups. One, comprising A. lacertosus and A. hrevirostris, is distributed around western, northern and eastern Australia and also has a widespread Indo-West Pacific 118 distribution. Another, comprising only A. fissi- frons, occurs off south-western and south- eastern Australia but is not known from north- ern Australia. This is also widespread in the Indo-West Pacific. The third group comprises species known from either north-western Aus- tralia Dickins (1963) separated the upper Byro faunas of the Norton Greywacke and Baker Formation from the D 1 faunas of older hori- zons, and referred the faunas of the overlying Coolkilya Greywacke at the base of the Ken- nedy Group to the same D 2 substage. The writer examined the collections at the Bureau of Mineral Resources, Canberra, and fully agrees with this distinction, though preferring to see the difference upgraded to stage rank. At the D 1-D 2 boundary Taeniothaerus and many Strophalosiids disappeared, together with molluscs such as Platyceras*, Kuculanella* . * Asterisked species reappeared in the Coolkilya Greywacke. 120 TABLE 1 Correlation of Permian aeqneneex. The loner Texan xiaqex xhoaht he xnhiH, ami the Skinner Ranch imiaileit n ith the Lenox llill>i ''idetl. A minonoid Western Australia Cnited States, Texas Russian Standard Subdivisions in Russia Carnarvon liasin I’ormations Stattes Dickins (llKkD (Hass Mt^. Formations StaKOs New Zealand Stajxes I'atarian ; Ociioan 1 Makarewan M'aiitian Kennedy ((roup I- Capitan ; Capitan Purulianan K azrnian ]•: , Word ' Wordian liraxtonian rflniian (no aininonoifls) D. Road Canyon Roadian (Harrettian) Kmi TO (irou]> 1), Catliedral Mountain Leonard Man‘ia])irian RaigeiKizinian Artinskiaii Aktastinian ? Wooraine! (irou]) (' Skinner Raiicli Tel ford ian Sakniarian Sakmarian ('allytharra Form li Lenox Hills Wolfcamp Asselian I.yons (>roup A Nc‘al Rancli Horizon Horizon A Quadratonxicula, Peruvispira, Chaenomya. Lep- tomphalus*, Macrochilina* , Nuculopsis*, Paleo- solen*, Pseudobaylea* , Naticopsis, Plagiostroma* , Acanthopecten, Astartella, Allorisma, Hetero- pecten*. Megadesmus also dropped out accord- ing to the lists in Dickins (1963», but is probably represented as so-called Cardio?norpha hlatch- fordi. Genera such as Warthia. Schizodus and Cancrinella also persisted. Glenister and Furnish (1961t have sug- gested on the basis of ammonoids that the Cool- kilya Formation, including upper Byro as defined by Teichert. ranged from Baigendzinian into the lower Guadalupian (including basal Word). Dickins <1956, 1963) i*eferred the upper Byro to the upper Artinskian, and Coolkilya (restricted) to the Kungurian. Changes are now necessary to these ages, because of redefinitions and subdivisions within the standard sections of North America and Russia. In the Glass Mountains, Texas, the standard sequence for North Ameri- can Permian, the lower Guadalupian of Glenis- ter and Furnish has been replaced in the new Road Canyon Formation by Cooper and Grant (1964), and recognised as a possibly distinct brachiopod stage by Nassichuk et al (1965), called Roadian by Furnish (1966 Table 1, p. 269). The Russian world standard has also been reinterpreted. When Dickins (1963) pro- posed a Kungurian age for the Coolkilya Formation he accepted the views of Licharev (1959) and others that the Kungurian Stage was followed by the Kazanian Stage, and that an intervening so-called Ufimian Stage should not be recognised. However the Soviet Com- mission on Permian Stratigraphy has set aside this view, and officially recognised the Ufimian Stage (Licharev 1966). They have also low- ered the Kungurian boundary. Thus the Kun- gurian correlation needs to be readjusted to the new Soviet interpretation. Waterhouse <1969 b) showed that the Ufimian as now understood rather than Kungurian was prob- ably equivalent to the Road Canyon, upper Byro and other faunas. Certainly there is a very distinct pre-Kazanian fauna, with charac- teristic fusulinids, brachiopods and ammonoids, developed widely above Baigendzinian faunas in Siberia. Arctic Canada, China, Japan. Aus- tralia and New Zealand. This fauna is pres- ent in the upper Byro Group. It possibly in- cludes the Coolkilya fauna as well, though Dickins (1956) did compare several bivalves with species from the Basleo beds of Timor. The Basleo beds are of Wordian (Kazanian) age. but Dickins considered that the affinities indi- cated a post-Artinskian age, and that absence of certain key genera ruled out a Kazanian correlation (Dickins, 1956, p. 39). Systematic description Order Productida Family Marginiferidae Stehli 1954 Diagnosis Marginal i*idge well defined in both valves or dorsal valve, cardinal process sessile to erect, broad median shaft, narrow backleaning median lobe and broad lateral lobes, costellae of variable strength, spines in row along hinge and/or umbonal slopes, specialised into very sturdy regularly arranged halteroid spines in some genera, comparatively few over visceral disc and trail, present or absent on doi'sal valve. Muscle scars usually not dendritic or lobate. 121 Binthalya Fm Pig. 1 n. sp. KENNEDY GROUP BYRO GROUP WOORAMEL GROUP LYONS GROUP Mungadan Ss Coolkilya Gw Baker Fm Nalbia Gw (= Norton Gw) Wandagee Fm Quinnanie Shale Cundlego Fm Bulgadoo Shale Mallens Gw Coyrie Fm Billidee Fm Moogooloo Ss Callytharra Fm •.y.Vv';-: • •' • 'y’/.** ; ♦vVT. : Av : *. i i ■■ ■ •* • •5 V'-'/f ‘I* v-V/Xv *•»•*•*•*• *VV** 1725 ' D. K. perforata K. perforata K. perforata D K. perforata 250 - 400 ' 3700 ? C B A -Sequence of formations in Carnarvon Basin, showing occurrences of Retimarginifera perforata and subdivision of faunal stages by Dickins (1963), based on Dickins (1963, fig. 2). 122 Subfamily classification Muir-Wood and Cooper (I960), followed by Muir-Wood (1965), recognised four subfamilies in the Mai'giniferidae partly on shape, partly on the presence or absence and strength of the marginal ridges. Their classification is not always consistently applied. For instance the Marginiferinae was defined as having a con- tinuous externally crenulated marginal ridge around the dorsal valve, and smooth adductors, in contrast to the Costispiniferinae, with crenu- lated ridges across the ears of both valves, and prominent endospines and smooth or rai’ely dendritic adductors. However Marginifera and Hystriculina do have marginal ridges across the ears of both valves, so that the definition of Marginiferinae erred. Endospines occur in the genus Kozlowskia, yet this is classed in the Marginiferinae. Elliotella was described as hav- ing no marginal ridge in the ventral valve (Muir-Wood and Cooper, 1960, p. 224). and Liosotella as having no marginal ridge in the dorsal valve (Muir-Wood and Cooper, 1960, p. 228), yet both were classed in the Costispini- ferinae. In fact, examination of the type species at the Smithsonian Institution shows that marginal ridges are present in both valves of both Elliottella and Liosotella. The subfamily Paucispiniferinae was classed in the Linoproductidae by Muir-Wood and Cooper (I960) and Muir-Wood (1965). Paucispinifera, as clearly shown in illus- trations by Muir-Wood and Cooper (1960, pi. 122, figs. 1-16) and confirmed from examination of types at the Smithsonian Institution, Washington, has a marginal ridge in both valves, a characteristic car- dinal process with narrow median lobe, a peculiar structure called a zygidium, found also in the Marginiferid genus Kozlowskia, smooth adductor scars, and transversely ar- ranged large halteroid spines. These features are typical of the Marginiferidae. not of the Linoproductidae. and the subfamily should be classed with the Marginiferidae (Waterhouse. 1969a, p. 232). The genus is distinguished in part by its cardinal process, which at least in the type species P. auriculata Muir -Wood and Cooper examined at the Smithsonian Institu- tion has an anteriorly extended median lobe. Otherwise, its most characteristic feature lies in the symmetrically disposed large halteroid spines across the ventral valve. The same spine pattern is seen in Kozlowskia, and some other genera referred to the Marginiferinae and Costipiniferinae by Muir-Wood and Cooper, but is not present in either Marginifera, or Costispinifera. In view of this, and because of the fact that the development of marginal ridges in these forms does not accord well with the arrangement of genera in Muir-Wood and Cooper, the three subfamilies are redefined as follows, using spines as the chief guide for classification. A. Marginiferinae: Marginiferidae with spines restricted to ventral valve, not differ- entiated into transverse row of about six large halteroid spines, halteroid spines arranged in radial rows over umbonal slopes. Marginifera, Anemonaria, Hystriculina, Lioso- tella, ?Elliotella Anemonaria. B. Paucispiniferinae: Marginiferidae with spines restricted to ventral valve, characterised by three to six large halteroid bracing spines developed in one of three concentric rows across shell. Paucispinifera, Kozlowskia, ?Alifera, Eomar- ginifera, Paramarginifera, Retimarginifera, Probolionia closely allied, Sajakella. Yakovlsvia and Muirwoodia, referred to the Paucispiniferinae by Muir- Wood and Cooper (1960), and Duartia, referred to the Margini- ferinae by Muir-Wood (1965) are not Margi- niferid. C. Costispiniferinae: Marginiferidae charac- terised by spines on both valves, not differ- entiated into few large regularly arranged halteroid ones on ventral valve. Costispinifera, Desmoinesia, Echinauris, Pro- marginifera, Spinomarginifera. Infiata and Nudauris are not Marginiferid, as claimed by Muir- Wood and Cooper *1965) but are Dictyoclostid. Subfamily Paucispiniferinae Muir-Wood and Cooper Genus Retimarginifera n. gen. Type species. — Retimarginifera perforata n. sp. Diagnosis. — Transverse shells with large ears, deep ventral sulcus, well defined dorsal fold. Halteroid body spines, usually in one to three transvei’se rows of 4 to 8. as well as row of hinge spines, limited to pedicle valve. Posterior disc reticulate, costae and concentric rugae strong, sturdy marginal ridge in both valves, marginiferid cardinal process, smooth adductors, large endospines. Discussion Kozlowskia Frederiks, with type species Pro- ductus capacii D'Orbigny is more globular than the new genus, and has a shallower sulcus and less prominent concentric and radial ornament. The sulcus and fold are less defined, or absent. Fig. 2. — 1-11, Retimarginifera perforata n. gen. n. sp. 1-9 external aspects of ventral valves, x 2. 1, 5 speci- men 59281 from locality UWA 29401 under different lighting. 2. holotype specimen 59282 from locality 27185. showing spines. 3. specimens .59283 from locality UWA 27185. 4. 8, posterior and anterior aspects of specimen 63828 from WC 20.1. showing extensive ears, with single laige spine base. 6. specimen 56384 from locality UWA 27185c. 7. specimen 56385 from locality UWA 27185. 9, specimen 56356 from UWA 27185h. with halteroid spine base on each ear. 10, transverse thin section of pedicle valve 63829 from locality UWA 29401. across sulcus, with external surfaces on top. showing slightly wavy lamellae of secondary layer, penetrated by taleolae (t). Note suggestion of inner pustule (arrowed), x 3 approx. 11. transverse thin section of dorsal valve of same specimen, exterior to right. X 35. 12. "Productxis” himalayense Dlcner. transverse thin section of ventral valve showing lamellar secondary shell with scattered pustules (arrowed), and an inner prismatic layer. Specimen collected by Dr. Gerhard Fuchs, (^eologische Bundesanstalt, Vienna, from Kash- mir. X 25. 123 124 Paucispinijera Muir-Wood and Cooper 1960, based on P. auriculata Muir-Wood and Cooper, is closer in general shape, and has modei’ately well defined costae, but concentric ornament and sulcus are inconspicuous. The genus is possibly represented by Productus altimontanus Merla, P. rimuensis Merla, Mar- ginifera hoofti Renz in the Karakorum and Marginifera pusilla Schellwien in south-east Europe, all of Lower Permian age. Retimarginifera perjorata n. sp. Figs. 2, 1-11, 3 Dictyoclostus gratiosus Prendergast (not Waag- en) 1943: p. 17, pi. 2. figs. 5-7. Marginifera gratiodentalis (not Grabau) Cole- man 1957: p. 79, pi. 9, figs. 1-14. Kozlowskia n. sp. Waterhouse 1969 a: p. 232, Fig 41. Material. — Material examined by the writer comprises two specimens with valves conjoined and three ventral valves from UWA 29401, a figured dorsal valve UWA 28453a; seven ventral valves, and one specimen with valves conjoined (the dorsal one masked), from UWA 27185, and a block WC 20.1 with four ventral valves, all kept at University of Western Australia. The shell material is preserved, slightly worn and decorticated sufficiently to obscure external growth lamellae and interior pustulation. Many have lost the anterior trail, and cardinal ex- tremities. Coleman’s description was based on these and a further 35 or so specimens, kept at the University of Western Australia. Prender- gast based her description on five specimens, kept at the Australian Museum. Sydney. Localities, — UWA 27185 — Cundlego Formation. Calceolispongia stage horizon north-east side of syncline on north bank of Minilya River, west of Coolkilya Pool, WC 20.1, Dictyoclostus zone, same description. UWA 28453 — Cundlego For- mation, 350 yards west of fence between Barra- biddy and Weer Paddocks, 2220 yards south of gate in the fence near Ban*abiddy Creek south of Wandagee Station. UWA 29401, Cundlego Formation — locality 1, horizon 1, of C. Teichert, Wandagee area. Other localities for specimens not examined by the writer are recorded by Coleman <1957, p. 79), including occurrences in both the Wandagee Formation and Baker Formation, Western Australia. Holotype. — Specimen 59282 from locality UWA 27185, fig. 2, 2. Figured topotypes. — Specimens 59283-29286 from locality UWA 27185. Diagnosis. — Transverse alate shells with sulcus deep anteriorly, costellae and wrinkles well de- veloped, endospines elongated. Inner shell penetrated by large taleolae. External features. — The species has been care- fully described by Prendergast <1943) and Cole- man <1957). Salient features are the trans- verse outline and prominent ears, set off in the dorsal valve of the holotype by a low ridge. A deep ventral sulcus commences 3-5 mm in front of the umbonal tip. widens rapidly to the start of the trail, and then becomes parallel- sided with a reduced sinal angle. The fold is anteriorly placed, the trail high and geniculate. Five to seven costae occur in 5 mm, increasing by branching and implantation, with low well rounded crests. One or two pairs may converge within the sulcus. Ears are smooth. About 15 growth rugae lie over the visceral disc, better defined in the specimens from locality UWA 27185, dividing radial ornament into tubercles, as in Dictyoclostidae. The rugae are less pro- nounced over the anterior disc, and missing or faint over the trail. Growth laminae are also present, about 5 mm per millimeter. Spines arise abruptly from costal crests, reach 1 mm in diameter though this varies, and are restricted to ventral valve. They form three rows, one along the hinge, and one row of usually three or so spines each side of the umbo just inside the ear, and 2-5 spines in a more erratic row each side of the sulcus. A pair of large halteroid spines lies on the ears, a pair on the posterior umbonal flanks, and an anterior pair on the sulcal flanks (Fig. 3). Other spines are few, with some anteriorly as in a specimen from UWA 29401 (Fig. 2, 1). Internal features. — Coleman described the ventral adductor muscle scars as an un- differentiated non-dendritic pair. A low mar- ginal ridge crosses the inner ears and is visible anteriorly in the holotype. The dorsal valve has a sessile cardinal pro- cess with elevated median shaft, broad lateral sulci, and very low lateral lobes. A broad plat- form lies in front of the process, passing into a short median septum which is highest at its anterior end. The posterior adductor scars are obscure, the anterior adductors small, rounded, anteriorly placed, not dendritic. Brachial 1 ‘idges are well rounded, and not clearly con- nected with the septum. The marginal ridge lies just within the hinge, strongly pocked across the inner ears, and faintly defined in front. About 8 elongated, narrow crested endo- spines lie each side of the midline between the septum and brachial ridges, immediately be- hind the marginal ridge. Shell structure. — The decorticated shell of several specimens from localities UWA 27185 and 29401 is flecked with white taleolae, 0.3 mm or more apart, and matrix-filled pores of similar spacing, rare posteriorly, and more common anteriorly, presumably later plugged by the taleolae. Pores and taleolae are only 0.1 to 0.2 mm apart in a large worn shell from UWA 29401. A polished transverse surface of a pedicle valve 1 mm thick from UWA 29401 has about 20 thick laminae, with large taleolae normally about 0.3 mm apart, extending from the inner surface half-way into the shell. Thin sections Fig. 3. — Generalised sketch of pedicle valve of Reti- marginifera perforata n. sp. showing distribution of halteroid spines, functional in black, non functional and old in open circles, x 1 approx. 125 of another more complete specimen (Fig. 2, 10) show numerous fine lamellae parallel to outer surface, interrupted by the large calcareous rods (taleolae), usually 0.10 mm across, and larger pores with matrix-filled cores, and the lamellae bulging inwards to each side. An in- nermost zone of smooth tissue is present, with large pustules doming outwards, only 0.7 to 1 mm apart, seemingly too numerous for spine bases. There are no visible taleolae. A transverse thin section of the dorsal valve (Fig. 2, 11 1 0.5 mm thick, is made up of numer- ous parallel lamellae with a thin outer yellow- ish band 0.05 mm thick of 2 to 5 lamellae. The inner edge is embayed by large pustules, and rare taleolae are visible in the inner layer, with a couple extending into the outer layer, but not disturbing the laminate structure. Other Productida have an outer layer with dense taleolae. but no tubes and pores, examples including other Paucispiniferinae (Fig. 2, 12) Anidanihus 'Waterhouse, 1968a' B-nd Kuvelousia (Waterhouse, 1968b). (-See Waterhouse 1970, p. 47.) PrendergasVs svecimens. — These specimens, examined at the Australian Museum, Sydney, appear to be closely related and perhaps conspecific, but the types were not to hand for comparison. The figui’ed specimen P 37569 (Prendergast, 1943, pi. 2, figs. 5-7) has spines, whereas the unfigured F 37570 has more body spines, and no obvious hinge row. perhaps due to preservation. Three costae pass forward from a spine base over parts of the shell. P 37571 has a few spines along the hinge, um- bonal slopes and outer sulcus. Resemblances These shells were identified with Marginifera gratiodentalis (Grabau, 1934) by Coleman (1957). Productus gratiodentalis was proposed by Grabau for shells described by Schwellwien (1892, p. 24, pi. 3. figs. 6-9; pi. 8, fig. 25' as Productus gratiosus occidentalis from the Fusu- line beds of the Carnic Alps. Pointing out that the Carnic specimens belonged to a full species distinct from gratiosus Waagen, Grabau (1934, p. 36) assigned a new name to Schell- wien’s species, because occidentalis was pre- occupied. No type specimen has been designated, but the selection of a lectotype should be de- ferred until the preservation of Schellwden’s figured specimens can be ascertained- Com- pared with the specimens from Western Aus- tralia, they have a moi'e posterior sulcus and finer concentric wrinkles seemingly more re- stricted to the posterior part of the shell. The radial ornament is much stronger over the an- terior inner ears. Schellwden implied through his comparison with P. gratiosus Waagen that the Carnic shells are Dictyoclostid, as in Bran- son (1948. p. 334). It appears unlikely that the Australian shells are in any way related to specimens assigned to occidentalis Schellwien or gratiodentalis Grabau by Chao <1925, pi. 2. fig. 6; 1927, p. 47. pi. 4, figs. 11-16); Grabau (1934, p. 36, pi. 10, figs. 7-8; 1936, p. 118, pi. 12, figs. 2a-d, 3a, b, 4a, b, 5) and King (1931, p. 71, pi. 14, figs. 1-3). All of these shells are more or less easily distinguished, apparently lacking halteroid spines, and having finer con- centric ornament and a shallow sulcus except in the specimens described by Chao (1925). None of them was compared to Marginifera. though the genus was recognised and species described elsewhere by the various authors. Sestini (1965. p. 178, pi. 22, figs. 6, 7) assigned the species to Marginifera, but her Karakorum specimen resembles neither the Australian nor European shells. TABLE 2 I)imem<\onn in mm of Kotinmrtzinifera pt'rforata n. sp. Ventral valves Sinai I'm- Card- Sinai an8= 28 18-4 9-4 do” 25= 85° 29 d 18-7 11 115' 55' OO 40= 27 ") 1 7 • 5 8-1 102^ 70 24" 85= Locality I'W.A 27185 20-8 18-9 7 ■ (i 110^ 25= 40= 22-0 11 d 7 • s ?8()’ 55= 25= 8.S-’ -f21 ■') 18-8 7-4 ?()5- 20= 85 d-20-d 12-5 d-d 95' 30= 85° + 24 12d d-4 112“ 30= 40“ 0-) •) 18-8 8-8 24 .'> 1 8 - s 7-s 112 ()5= 19= 30= 24-4 1 8 • 8 1 ■ 1 95= 55= •)>iO 38° Locality W(’ 20-1 25 0 Id-d 10-d 115= 45 15= 88° Dorsal valvp TV'.\ 28488a Width Lcnjith Hcitjht Card- Fold an«lc at Sejduni inal an 1(1 amil<* measured at start of trail ami at anterior marjiin. Most specimens have lost a little of the cardi- nal extremities and something of the trail. Septum length shows the distance between the cardinal process and anterior end of the septum. None of the American species of Kozlowskia is particularly close. The type species Productus capacii as figured by D’Orbigny (1842, pi. 3, figs. 24-26), Kozlowski (1914. pi. 2, figs. 1-15; pi. 5, fig. 13; text-fig. 1, 2) and Muir-Wood and Cooper (1960, pi. 63, figs. 13-19) is less consist- ently transverse and more oval in outline, with feeble concentric ornament and less pronounced 126 TABLE 3 St(t(i.‘42 1 5 • 292.S s • .5942 9S-S3' 23 -S3 S 2 • r)577 2-S29S 1 - 5309 14-7920 3 - 9r>45 V U)<)7:i0 IS -.5041 17-S7.5t> 14 •9(571 1 (5 • 594(5 <7 X .... 7- width : 0-()S:ki {)• 7.5(i2 (W) 1 0-4092 (W) 0-.58(W 1 4-2701 7 rmb. ' 1 -UK) 0- 00(5(5 <*oin])lete sjtei-iinens only 11 8 1 8 8 X 24-8250 15-(5750 9 - 3000 s 2-2752 ' 3-10(53 1 - 44(55 9-1(549 1 9 - 81(59 1 5 - 5538 <7 X 0-8044 ' 1-0982 0-5114 7width : 0-7791 0(5312 n mimlior of sp(“(-inionsi X = iiicaii s = stiuiclanl drvintiou V - rvMdfiok-nf of variation <7 X = Standard error of the mean T width =- (M)rrelation eoeilUdent (width — ) 7 I'lnb.' - eon-elation <-oelMeipnt with und)onal angle. sulcus and fold. A specimen figured as P. longi- spinus aiot Sowerby) by Salter H861. p. 64, pi. 4, fig. 2t is more elongated, with a moderately well formed sulcus, and emphasized costellae. The scattered tubercles shown over the ventral valve, if they represent spine-bases, would rule out any close alliance, and such also seem to be represented by Kozlowski U914, pi. 2. fig. 9b) but Kozlowski (1914. p. 22) stated in his text that spines are rare. Specimens examined at the Smithsonian Institution from Apillipampa south of Capinote, Bi’azil, (USNM 124030a, b; Muir-Wood and Cooper. 1960, pi. 63, figs. 13- 19) have 6 halteroid spines, and a few others only. There is no hinge row. perhaps because they have been rubbed off, for the specimens are not well preserved. The ears have been lost from USNM 124030b, but growth lines show that they were lax’ge. A small ventral valve kindly made available by Dr. Richard E. Grant from the U.S. Geological Survey, Washington, D.C. has been sectioned. Unfortunately the shell is partly silicified and no taleolae or spine bases can be discerned. Laminations are less pronounced than in the Australian species. Marginifera hiiTialayensis Diener (1899) is also close in shape, but has larger ears and more numerous spines. The types have been exam- ined at the Geological Survey of India, Calcutta. Specimen F 6285 (Diener. 1899, pi. 6, fig. 2> is selected as lectotype. The species seems to have symmetrically disposed halteroid spines, and a row of hinge spines is preserved on some (e.g. P 6236— pi. 2, fig. 2, and P 6238— pi. 2. fig. 4), so that the specimens are not Marginifera, but possibly belong to the Paucispiniferinae. It is possibly a new genus, distinguished by the strong concentric ornament of the dorsal valve, and cluster of tubercles or spine bases on the outer ears (as in F 6238), and shell structure. The shell structure has been examined in specimens collected from the type locality in Kashmir by D. G. Fuchs, Geologische Bundesanstalt, Vienna. The outer shell structure in the ventral valve (Fig. 2. 12) differs considerably from that of per- forata, coming much closer to that of Anidan- thiis described by Waterhouse (1968a). The in- nermost band, 0.125 mm thick, consists of paral- lel prisms perpendicular to the surface, each about 0.13 mm thick. This layer is not preserved in K. perforata. The inner secondary layer has cloudy calcite lamellae less conspicuous than in K. perforata, but essentially the same in possessing large whorls due to ?pustules, spaced about 0.4 to 0.6 mm apart in a single row and thus too numerous to have been spine bases. The outer layer, just as thick, has small calcite and matrix filled pores, about 0.1 to 0.05 mm in diameter, possibly due to taleolae, but very obscure. The pores resemble those of Anidanthus and Kuvelousia, and ai*e much smaller than in K. perforata. Productus altwiontanus Merla < 1934, pi. 20, figs. 27-32, 36-41) and P. rimuensis Merla (1934, pi. 24, figs. 7-16, 20) from the Lower Permian of the Karakorum Range are close in outline and ornament, but their spine pattern is not certain. Another externally similar species was described as Marginifera hoofti Renz (1940. p. 27, pi. 4, figs. 12a-c> from Upper Uralian beds of locality 5. Shukpa Kuchang Glacier, of the Karakorum. Unfortunately its spinose orna- ment has not been described, but the illustrated specimen resembles the new form in outline and ornament. ''Marginifera’* pusilla Schellwien (1892, pi. 4, figs. 11-21) has a few large halteroid spines in a concentric row, and strong radial ornament and large ears, and so probably be- longs to the same plexus as the new form. It came from the Auernigg beds of east Europe. Concentric ornament is less defined. Marginifera reticulata King (1931. pi. 22, figs. 3. 10) might also prove to be allied. It comes from the Leonard of the Glass Mountains, and has similar ornament and deep sulcus, but is slightly more rectangular in outline, with smaller ears and longer visceral disc. Acknowledgements Dr. P. J. Coleman, Dpeartment of Geology, University of Western Australia, Perth, lent specimens identified as Marginifera gratioden- talis from Western Australia, and Dr. Gerhard Fuchs, Geologische Bundesanstalt, Vienna, sent specimens of "Productus” hivialayense from Kashmir. Particular thanks are due to Dr. G. A, Cooper, Smithsonian Institution, U.S. National Museum, Washington for his help in guiding me through collections and types in his care. Drs. C. Mac- Clintock and A. L. McAlester facilitated exam- ination of relevant types at the Peabody Museum, Yale University and Messrs M. V. Sastry and S. C. Shah provided facilities for examining types at the Geological Survey of India. Calcutta. Dr. R. E. Grant, U.S. Geological Survey, Wash- ington, D.C. gave specimens of Kozlowskia capacii for sectioning. 127 References Branson. C. (1948). — Bibliographic Index of Permian Invertetsrates. Geol. Soc. Amer. Mem. 26: 1050 pp. Chao. Y. T. (1925). — On the age of the Taiyuan Series of North China. 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( 1963 ) . — Permian pelecypods and gastro- pods from Western Australia. Bur. Mineral Resour. Geol. Geophys. Bull. Aust. 63: 1 - 202 . Diener, C. (1899). — Anthracolithic fossils of Kashmir and Spiti. Palaeont. Indica ser. 15. (2). D'Orbigny. A. (1842). — Voyages dans I’Amerique meri- dionale. Vol. 3. pt. 4: Paleontologie. P. Bert- rand. Paris. Frederiks, G. (1933). — Palaeontological Notes 4. On some upper Paleozoic Brachtopods of Eurasia. Mater. Cent. Sci. Geol. Prospect. Inst. 2: 24-33 (in Russian with English summary). Furnish. W. M. ( 1966).— Ammonolds of the Upper Permian Cyclolobus Zone. N. Jb. Geol. Palaont. Abh. 125: 265-296. Glenister. Brian F. and Furnish. W. M. (1961). — The Permian Ammonolds of Australia. J. PaLeont. 35(4): 673-736. pis. 78-86, 17 text figs. Grabau, A. W. (1934). — Early Permian fossils of China, 1 : Early Peimian brachiopods, pelecypods and gasteropoda of Krieichow. Palaeont. Sinica B v. 8: pt. 3. (1936). — Early Lower Permian fossils of China, Pt. 2. 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