MEMOIRS OF THE QUEENSLAND MUSEUM BRISBANE | VOLUME 43 30 JUNE 1999 PART 1 SILURIAN AND DEVONIAN CRINOIDS FROM CENTRAL VICTORIA PETER A. JELL Jell, Р.А. 1999 06 30: Silunan and Devonian erinoids from central Victoria. Memoirs of the Queensland Museum 43(1): 1-114, Brisbane, ISSNQU79-3835. Crinoids have long been known trom middle Palaeozoic clastics in central Victoria but this is the first comprehensive attempt to document the fauna; 54 taxa are described; 25 are camerates (са), 7 disparids (d), 19 cladids (cl) and 3 flexibles (T). New taxa are the genera Hoallawayerinus calvus (ca), Duncanicrinus calvariolus (са), Frankocrinus lholmesi, F. enidae (cà), Pterihoerinidae gen. nov. (ca), Darragherinus tomi (d), Kroppocrinus heatheotensis, K. mathiesonensis (d), Stewhrecrinus terevi (cl), Holmesocrinus enidae, Н. idaensis (cl) and Quadritaxocrinus websteri (Гү, camerate species Ophiocrinus metlae, Dimerocrinites bispinosa, Eucrinus clurkae, Eudimerocrinus ekardti, E. gilli, Nexocrinus wallanensis, Hexaerinites chirnsidensis, QOehlertierinus lemenni, O. jeani, Alisocrinus lineatus, Clematocrinus perforatus and С, argylensis: disparid species Trichocrinus morleyi and Phimocrinus hanscli; cladid species Codiacrinus secundus, Cupulecrinus ausirogracilis, Dendroerinus akFugius, Plicadendroerints australis, Shiniocrinus cometensis, 5. richi, Antihomocrinus chapman, Nassoviocrimis corcoraut, Dictenocrinus ibaeypus and D. remotus: and the [lexible Meristocrinus qualrirumts. D Crinoids, Silurian, Devonian, Victoria. Peter A. Jell, Queensland Museum, PO Bax 3300, South Brisbane 4101, Australia; received 28 March 1998. Middle Palaeozoic sequences in the Melbourne Trough of central Victoria have yielded rich fossil faunas since the middle of last century but the diverse crinoid component has been largely ignored among echinoderm groups (Withers & Keble, 1934a and b; Jell, 1983). Possibly the first record and certainly the first illustration of crinoids from central Victoria was that of Blandowski (1855) but he figured only stem columnals and did not follow with the descriptive paper promised. Bather ( 1897) first described an articulated crinoid, Hapalocrinus victoriae, with a line drawing that he acknowledged in a footnote as being somewhat uncertain, Photographs of Bather's (1897) specimen and its counterpart (Bather claimed it was lost) are provided below and show that the cup is poorly known. Chapman (1903) described Helicocrinus plumosus and Botryoerinus longibruchiaius from the Silurian of the Melbourne area. Both are reviewed below with the latter re-assigned to Mussoviocrinus Jaekel. 1918. More recent reports of erinoids in clastic sequences of central Victoria are restricted to : 1. Talent (1965-17, pl. 4, fig. 2) noted crinoid stem ossicles as a common element throughout the Heathcote sequence and crowns at localities 41 (=Hapalucrinites argylensis sp. nov., below) and 42 from where he figured the unidentifiable external mould of a crown; 2, Crotalocrinites from Heathcote (Jell, 1982) and; 3. Kooptoona- erinus nulti, Codiacrinus rarus, Dendrocrinus suundersi and a new genus of Dimerocrinitidae (=Duncanicrinus calvariolus below) from near the Silurian/Devonian boundary east of Melbourne (Jell in Jell & Holloway, 1983). Numerous other records of crinoids in these sequences (e.g, Williams, 1964; Gill & Caster, 1960) simply noted them among wider faunal listings without naming or describing taxa, Other described crin- oids (rom Victoria oceur in limestones at Lilydale (Bates, 1972), Mansfield (Jell et al.. 1988) and Toongabbie (Philip, 1961). In the rest of Australia, Devonian crinoids have been described from limestones in Queensland and NSW (Jell et al., 1988) and Pisocrinus (referred то Parapisocrinus by Rozhnov, 1981) and Lecanocrinus have been reported from the Silurian at Yass (Etheridge, 1904; Chapman, 1934). BIOSTRATIGRAPHY The sections from which these erinoids have been collected span the Ludlow and Pridoli of the Silurian and the Lochkov of the Devonian, However. soil cover, urban development and metamorphism around intrusions limit available sections. Virtually all the eehinoderms known have been collected from quarries, road cuttings, creek beds or other man-made exposure. to MEMOIRS OF THE QUEENSLAND MUSEUM Wenlock Ludlow E Lochkov Pragian Aegiria thomasi Pi Nor , Boucotia janeae Boucotia australis d rs entiensis 1 a 3 4 5 6 7 8 9 10 11 Dendrocrinus sp. x О О [9] Trichocrinus morleyi Dendrocrinus arrugius X X X X O |Hollowaycrinus calvus Nassoviocrinus corcorani X X O O |Ctenocrinus signatus Hapalocrinus victoriae X О О О |Ctenocrinus paucidactylus Helicocrinus plumosus X [9] Dimerocrinites bispinosus Nassovio- longibrachiatus E | O Eucrinus clarkae Phimocrinus americanus x H X Clematocrinus argylensis X О O Clematocrinus perforatus Alisocrinus lineatus x О | Frankocrinus holmesi Phimocrinus hanschi X (Ө) | Frankocrinus enidae Shintocrinus cometensis X о R) О О |Duncanicrinus calvariolus Antihomocrinus chapmani X X X X Quadritaxocrinus websteri X [9] Nexocrinus sp. Holmesocrinus enidae X О Plicodendro- australis Nexocrinus wallanensis X O | Stewbrecrinus terryi | Кғорро- heathcotensis Xx | О X |Codiacrinus secundus Cupulo- austrogracilis X [9] Holmesocrinus idaensis Hexacrinites chirnsidensis X О Darraghcrinus tomi Ophiocrinus nnettae х О | Kroppo- mathiesonensis Eudimerocrinus gilli х O |Meristocrinus quatriramus Ctenocrinus stellifer X х х О |Decacrinus sp. Dictenocrinus remotus X O | Aneyrocrinus sp Dictenocrinus ibaeypus X О Crotalocrinites pulcher Kooptoonocrinus nutti x о О |Eudimerocrinus eckardti Geroldicrinus sp. X [9] О | Oehlerticrinus lemenni Shintocrinus richi X [9] O |Oehlerticrinus jeani Pterinocrinidae gen. nov Е х О |Myelodactylid indet. Dendrocrinus saundersi | х Codiacrinus rarus X FIG. 1. Distribution of crinoids in the Silurian-Devonian clastic sequences of central Victoria; all species listed are dealt with in this paper, Jell & Holloway (1983) or Jell (1982). The international scale is correlated to the local Brachiopod Assemblage Zone scheme of Garratt (1983) in the top two rows, following Garratt & Wright (1988). The numbering sequence of columns in the third row refers to aggregates of localities as listed in the Appendix; detailed relative levels of widely distributed localities is uncertain and these must be seen in many cases as best approximations. Occurrence of taxa listed on the left of the figure are indicated with *X" and those on the right by ‘O’. Generic names ending in a hyphen require addition of the suffix crinus and are shortened to fit a single line. Structure within the Melbourne Trough consists of tight folding with long approximately N-S axes, also limiting measurable sections. Fossils occur in profusion in a few horizons; they are relatively rare in many other horizons; but most of the sedimentary sequence is unfossiliferous. Brachiopods are the most common group among the shelly fauna and have been used to establish a preliminary biozonation (Fig. 1) against which the shelly faunas have been placed in sequence (Garratt, 1983). The crinoids described herein come mainly from the few horizons where fossils are prolific and preservation is superb; very large numbers of articulated echinoderms occur there. For example, NMVPL252 provides the greatest diversity of crinoids and of echinoderms in the Melbourne Trough. This locality was originally a small road metal quarry but most available CRINOIDS FROM CENTRAL VICTORIA 3 material came from a new excavation of the quarry in the summers of 1980-1983 undertaken by the author with the sustained help of Frank and Enid Holmes and Steve Eckardt and occasional help of several others. Those localities that yield only one or 2 species mostly represent localities where fossils are rare and their age determined by stratigraphic position. The crinoid localities (Appendix) are placed in approximate relative stratigraphic sequence (Fig. 1) using these best available data. The brachiopod zonation of Garratt is related to the international stage scale with graptolites that occur at a few horizons through the sequence (Garratt & Wright, 1988). The crinoids described herein are not signif- icantly useful biostratigraphically as they occur relatively rarely in terms of both localities and horizons. Nevertheless there are clear differences between the Ludlow and Lochkov faunas for example and the continued study of this fossil group with other elements ofthe shelly fauna will continue to refine what is at present a very preliminary brachiopod zonation. PALAEOECOLOGY Palaeoecological settings in the Melbourne Trough were discussed in a number of papers (Garratt, 1983; Cas, 1983; Vandenberg & Wilkinson, 1982; Vandenberg, 1988, 1992). Cas (1983, fig. 17) provided a schematic diagram of regional palaeogeography for SE Australia showing the Melbourne Trough as a long narrow embayment of the ocean with ample neighbour- ing relief and continued subsidence to provide the thick pile of clastic sediments. Localities rich in articulated echinoderms occur in distal turbidites (PL252) or extensive thin sand units (within a finer grained sequence) (PL300, 229, 1924). Both these settings indicate catastrophic events causing the death and fast burial of these animals (Jell, 1983) but the cause or causes of such events remain unclear. PALAEOBIOGEOGRAPHY These crinoid faunas provide few useful data for palaeobiogeographic reconstruction as they contain taxa otherwise restricted to North America or to Europe and other taxa common to both those areas. Few other parts of the world have well enough described crinoid faunas ofthis age for meaningful comparison. At the generic level there are slightly more representatives of the European fauna as is the case with brachio- pods (Boucot et al., 1969) but among camerates there are more American affinities. SYSTEMATIC PALAEONTOLOGY Material described is housed in the Museum of Victoria (NMVP) and the localities are registered in the same Museum (NMVPL). Specimens are preserved in decalcified clastics so are found in the form of internal and external moulds often partially infilled with iron oxides making the mould less than faithfull. Morphological detail of the moulds improves with diminishing grain size of the matrix. All illustrations are of latex casts taken from these moulds and whitened with a sublimate of ammonium chloride. Terminology follows Moore & Teichert (1978). Measurements are given as: length, parallel to the central axis; width, transverse to, but never cutting or meeting the central axis; and depth, normal to, and may join the central axis. Class CRINOIDEA Miller, 1821 Subclass CAMERATA Wachsmuth & Springer, 1885 Order DIPLOBATHRIDA Moore & Laudon, 1943 Suborder EUDIPLOBATHRINA Ubaghs, 1953 Superfamily RHODOCRINITOIDEA Roemer, 1855 Family OPSIOCRINIDAE Kier, 1952 Frest & Strimple (1981) and Ausich (19862) recognised the close similarity of Opsiocrinus Kier, 1952 and Ophiocrinus Salter, 1856 and assigned them both to this family; they are considered synonymous by Jell & Theron, 1999. Ausich (1986a) also erected 2 new genera from the Llandovery of Ohio in this family and discussed the group in detail. Inclusion of the Australian Hollowaycrinus nov. is discussed under that heading below. This addition necessitates widening the family concept to include a form with radials in lateral contact; this difference has been sufficient to separate taxa at superfamily level in the past (Ubaghs, 1978b). To take an even more heretical view I question whether Stelidiocrinus Angelin, 1878 from the Upper Silurian of Gotland should not be included in this family. It has been classified in the Monobathrida because it lacks the second circlet of plates below the radials but it (particularly S. laevis Angelin, 1878 as figured by Ubaghs (1978b, fig. 307, le)) resembles the Opsiocrinidae very much in cup shape, median column of anal plates in CD interray, 10 biserial arms, few slightly depressed interradial plates and presence of an intersecundibrach, Its first primibrach is shorter and wider than in most Opsiocrinidae but is comparable with that of O. 4 MEMOIRS OF THE QUEENSLAND MUSEUM Eifelian Ophiocrinus mariae | | Ophiocrinus sp. cf. O. mariae | Emsian | n 2 == | Ophiocrinus stangeri —— — — m — “melds мА ——— Е, — | Pragian Г | Hollowaycrinus Lochkov | Е ya Ophiocrinus nnettae NE — E | Pridoli а pr sr m | Ludlow | | = Stelidiocrimus | н er | Wenlock = n m Llandovery Rhachicrinus Silfonocrinus | a | i — LL ——— Ordovician | Gaurocrinus - Tike ancestor FIG. 2. Sketch of phylogeny of the Opsiocrinidae following Ausich (1986a) and discussion herein. nnettae described below. Throughout the Opsio- crinidae the infrabasals are concealed by the stem attachment and Opsiocrinus itself was originally described as monocyclic; I suggest therefore, that a careful examination of the Swedish material of Stelidiocrinus should be made to determine this feature. Stelidiocrinus would join Holloway- crinus as the second Opsiocrinidae with radials in lateral contact. Witzke & Strimple (1981) sug- gested that evolution from rhodocrinitid to dimerocrinitid stage (i.e. withdrawl of the first interbrachial to allow lateral contact of radials) occurred several times. They postulated 3 separate occasions from different Ptychocrinus stocks so my suggestion of 2 further occasions above is in line with their thinking. Current knowledge of this family suggests that only small sections of the lineages involved are known (Fig. 2) and I agree with Witzke & Strimple (1981) that a great deal more basic data about Silurian forms in particular will be necessary to refine our understanding. Ophiocrinus Salter, 1856 TYPE SPECIES. Ophiocrinus stangeri Salter, 1856 from the Lower Devonian Bokkeveld Series, South Africa. REMARKS. This genus is discussed in detail elsewhere (Jell & Theron, 1999). Its occurrence in the Lower Devonian of Victoria in close proximity geographically and temporally with Hollowaycrinus gen. nov. makes it quite feasible that one gave rise to the other as remarked under that genus below. Ophiocrinus nnettae sp. nov. (Fig. 3) ETYMOLOGY. An anagram from Annette, MATERIAL. NMVP149344 from NMVPL1990. DIAGNOSIS Interrays narrow; primanal in posterior interray contacting C and D radials and Ist primibrachs, supporting 3 anal plates distally; central column of larger hexagonal plates in anal sac. Arms 10, with triangular axillary 2nd primibrach, distinctly biserial above distal to secundibrach. Stem circular, with small but distinct marginal projections on nodals and internodals. DESCRIPTION. Crown 740mm long, 74 times as long as wide at cup, with arms spreading gently (tips not preserved). Cup low conical, approximately 5mm long, wider than long; plates smooth, convex so sutural margins depressed. Infrabasals presumably concealed by stem. Basals longer than radials, hexagonal, longer than wide. Radials pentagonal, wider than long, separated from each adjoining radial by basals and 15 interprimibrach, with distinctly margined outer ledge across distal margin matched by similar ledge on Ist primibrach, similar matching ledges on succeeding interplate sutures along the arm gradually diminishing in distinctness and size; lst primibrach rectangular, as wide as radial, short; 2nd primibrach axillary, triangular, as wide as Ist primibrach. Arms 10, subquadrate in section, with flattened outer face; each ray with 2 main rami, uniserial as far as 6th secundibrach, becoming biserial abuptly (3rd in one arm observed) distally, with long pinnule of 5 or 6 pinnulars on each free brachial; only primibrachs fixed in cup; interprimibrachs small and few; 1st largest, contacting basal, 2 radials and Ist primibrachs, supporting 2 long narrow plates distally. Primanal hexagonal though contacting 8 plates, almost as large as basals but not quite as long, supporting 3 anals distally; anal plates numerous, decreasing in size distally, with average size larger than that of interbrachials, distinct central column of decreasing hexagonal plates. Stem circular in section, heteromorphic with slightly different sized (length and diameter) columnals alternating, noditaxis N1, with epifacet on each columnal having an outer circlet of small but distinct tubercles or pseudocirri. REMARKS. This Victorian species differs from the South African genotype (Jell & Theron, CRINOIDS FROM CENTRAL VICTORIA 3 FIG. 3. Ophiocrinus nnettae sp. nov., incomplete holotype crown in С ray view, NMVP149344 from NMVPL1990, х3. 1999) in having fewer interprimibrachs, no intersecundibrachs, distinct ledges on each plate on each interplate suture from radials up to about 3rd secundibrach, its arms becoming biserial much earlier (secundibrach 3 in some arms) and in the stem with tubercles on internodals. From the North American O. mariae Kier, 1952 and O. benderi (Kesling, 1968). if those are separate species as suggested by Frest & Strimple (1981) contrary to Kesling & Chilman (1975), the Victorian species is distinguished by the same features as separate the genotype as well as a lack of ornament on the interprimibrachs and very short Ist primibrach. Hollowaycrinus gen. nov. TYPE SPECIES. Hollowayerinus calvus sp. nov. ETYMOLOGY. For David Holloway who greatly assisted with fieldwork, curation and scientific advice. DIAGNOSIS. Infrabasals 5, concealed by stem attachment. Кафа! large. heptagonal, in contact with each other laterally except across CD inter- ray; Ist primibrach hexagonal; 2nd primibrach axillary. Primanal in radial circlet, supporting 3 plates distally, with median column of larger anal plates involving 4 and probably more plates. Arms 10-20, with each ray having 2 main arms becoming biserial distal to the 6th or 8th secundibrach, with some arms branching irreg- ularly once or twice distal to primaxil even though other rami in the same individual are unbranched. REMARKS. This genus is placed in the Opsio- crinidae based on the similarities in cup shape, 5 concealed infrabasals, median column of anal plates in CD interray. primanal supporting 3 plates, large Ist interprimibrach supporting 2 plates followed by large number of very small irregular plates, 10-20 arms, uniserial to biserial. Conventional classification following Ubaghs (1978) would place this species in Dimero- crinites because of the radials being in lateral contact. However, the arms being uniserial proximally then becoming cuneate and finally biserial, the interradial plates being many small ones above a few large ones and the radials being barely in lateral contact all point to an origin from the Opsiocrinidae and it is classified accordingly. Hollowaycrinus calvus sp. nov. (Fig. 4) ETYMOLOGY. Latin calvus, bald; referring to the lack of median ray ridges and ornament. MATERIAL. HOLOTYPE: NMVP100158. PARATYPES: NMVP100150, 100153, 107099-107104, all from NMVPL229. DIAGNOSIS. As for genus. DESCRIPTION. Crown subcylindrical, averag- ing 30mm long, at least 4 times as long as wide. Cup low to medium conical, 2-6mm long, just wider than long. of unornamented plates. Infrabasals 5, concealed by stem. Basals 5. longer than radials. hexagonal except for heptagonal posterior one, longer than wide. Radials hept- agonal, wider than long, with longer sides contacting basals and 1st primibrach, with short sides contacting adjoining radials or primanal in CD interray: 1st primibrach hexagonal. not as wide as radial and little more than half as long; MEMOIRS OF THE QUEENSLAND MUSEUM CRINOIDS FROM CENTRAL VICTORIA 7 2nd primibrach axillary, pentagonal, as wide as Ist primibrach; each ray with 2 main arms, some without further branching, others with 1 or 2 further branchings irregularly, uniserial as far as 6th or 8th secundibrach, becoming biserial distally, with long pinnule of 5-6 pinnulars on each free brachial; arms free distal to 2nd secundibrach; interprimibrachs numerous, with large heptagonal proximal one supporting 3 plates in a row distally, with size decreasing rapidly distally, with distal medial plates depressed. Primanal hexagonal, almost as large as basals, supporting 3 anals; anal plates num- erous, decreasing distally, with average size larger than interprimibrachs, Stem circular in section, with slightly different sized (length and diameter) nodals and internodals alternating proximally but distally with noditaxis N212, may be long (no complete stem known but an incomplete one in excess of 9cm known), distal termination unknown. All plates smooth, without ornament, REMARKS. This species is distinguished from Ophiocrinus by most radials just making lateral contact and by variable arm branching. Its separation from dimerocrinitids is discussed above. The arms are highly variable in their branching pattern and also in the arrangement of brachials. Most observable rays divide once and have only 2 arms whereas others may have one undivided arm from the first branching while the second may divide twice more. Most subsequent divisions appear to be at fairly high and variable angles to growth direction; there is a question of whether these could represent a regrowth response to breakage or predation to which no answer is readily available. Brachials begin to become cuneate just distal to the cup and in most cases become biserial cuneate by about the 8th secundibrach; a few remain cuneate farther distally and none become rectilinear biserial. Superfamily DIMEROCRINITOIDEA Zittel, 1879 Family DIMEROCRINITIDAE Zittel, 1879 This family is a regular component of crinoid faunas from Ordovician to Devonian and has been discussed by many authors (Ausich, 1986a; Brower, 1973; Breimer, 1962; Frest & Strimple, 1981; MacIntosh, 1981, 1987; Witzke & Strimple, 1981 among others). Since most that is known about this family comes from the Northern Hemisphere it is outside my aims to review its phylogeny. However, in adding the several Australian forms that follow I have had to make some inferences about Northern Hemi- sphere taxa. The simplest way to recognise members of the family until recently had been to identify that a crinoid was dicyclic and that its radials were all in contact except for interrupion by the primanal in the CD interray. Witzke & Strimple (1981) questioned the universality of this single feature and І agree with their discussion of evolution between the Rhodocrini- toidea and Dimerocrinitoidea, In assigning Hollowaycrinus to the Opsiocrinidae and suggesting that Stelidiocrinus may also belong to that family 1 follow their line of thought. I also follow Witzke & Strimple (1981) in the generic divisions of the family based on numbers of arms (Fig. 5) although there is good reason to believe that numbers of arms may have changed on several parallel lineages within the family. Moving along Witzke & Stimple's (1981) ideas we now have available generic names for dimerocrinitoids with uniserial arms that divided once (Duncanicrinus gen. nov.), twice (Nexo- crinus Eckert, 1984) or more than twice 1n each ray (Ptychocrinus) and for biserial represent- atives with 10 (Dimerocrinites), 20 (Eucrinus) or more than 20 arms (Eudimerocrinites (=Ambicocrinus), Griphocrinus). Whether latest Ordovician Ptychocrinus fimbriatus, Llandovery P. longibrachialis and Early Devonian Duncanicrinus constitute a lineage or not will only be answered by gaining further knowledge of Silurian taxa and thus detailed phylogenies but at present this is a reasonable hypothesis worth further testing (Fig. 5). These generic groupings accommodate Brower's (1973) idea that the 3 latest Ordovician species of Ptychocrinus represented 3 separate lineages and should be separated generically and Witzke & Strimple's (1981) ideas on the existence of 3 lineages in the Dimerocrinitidae based primarily on numbers of arms. There are no doubt numerous exceptions to these general groupings and the most obvious is the high probability that the transition from uniserial to biserial arms occurred more than once in each lineage. However, it is necessary to continue assimilating new data into an hypothesis FIG. 4. Hollowaycrinus calvus gen. et sp. nov., all crowns with or without stem attached, from NMVPL229, A, NMVP107101, *2.5. B, holotype, NMVP100158, х4. С, NMVPI00150, х5. D, NMVP100153, х4. E, NMVP107102, x3.5. Е, NMVP107104, x3.5. G, NMVPI07103, х3. Н, NMVP107099, x5. MEMOIRS OF THE QUEENSLAND MUSEUM A Biserial; »20 Arms Uniserial; >20 Arms Biserial; 20 Arms Uniserial; 20 Arms Eifelian E. a A. arborescens | Emsian | Ргар1ап | | Ен. eckardti | E. clarkae Nexocrinus sp. Lochkov de 27 Eu*gilli L | = | A X | Pridoli [Г iE | | | N. vases Ludlow | 1 | EUCRINUS NEXOCRINUS E. laevis, nodibasis, К А Eu. multibrachiatus DEREN N. occidentalis Wenlock | кА da bs E. icosidactylus | N H = \ E. pentlandicus | N SUN. | Llandovery \ УММУ № Adamsensis P. medinensis | . \ p — N. delicatulus [ee Ordovician ^P. splendens N. parvus C Biserial; 10 Arms Uniserial; 10 Arms Eifelian Emsian Pragian Lochkov D. bispinosus D. calvariolus Pridoli DIMEROCRINITES | DUNCANICRINUS — Ludlow D. longimanus | D. planus, milligani, roemeri, | lilliformis, brachiatus Wenlock | | FIG. 5. A, sketch of phylogeny of the D. decadactylus, inornatus | Dimerocrinitidae with more than 20 arms | including Ptychocrinus, Eudimerocrinus and D. sculptus, | T Ambicocrinus. B, phylogeny of the Llandovery popkintonensis, elegans | D. СИ Dimerocrinitidae with 20 arms including [rd Nexocrinus and Eucrinus. C, phylogeny of Ordovician x D. fimbriatus the Dimerocrinitidae with 10 arms including Duncanicrinus and Dimerocrinites. CRINOIDS FROM CENTRAL VICTORIA 9 FIG. 6. Dimerocrinites bispinosus sp. nov. from NMVPL252. A,B, D ray view and enlargement of posterior interray of holotype NMVP108581a, *2.5 and x5, respectively. C, A ray view of counterpart (В ray on left, Е on right), NMVPIOSS81b. х2.5. D, splayed crown in basal view, NMVP108615. «2.5. 10 MEMOIRS OF THE QUEENSLAND MUSEUM until all the details are worked out. To that end I offer a possible phylogeny (Fig. 5) for known species of Dimerocrinitidae for future testing and modification where necessary. Just how many lineages and which features remained constant to be able to recognise separate lineages remain largely unanswered for the dimerocrinitids today as when Witzke & Strimple (1981) raised many questions on the origins of their groupings. Dimerocrinites Phillips in Murchison, 1839 ГҮРЕ SPECIES. Dimerocrinites decadactylus Phillips in Murchison, 1839 from the Wenlock of England; by subsequent designation of Roemer, 1855. REMARKS. I apply the concept of Witzke & Strimple (1981) to Dimerocrinites (Dimero- crinites) which is to restrict the genus to those forms ofthe broader generic group with 10 arms. Dimerocrinites bispinosus sp. nov. (Fig. 6) ETYMOLOGY. Latin bi-, two and spinosus, spine, for the pairs of spines on the outer sides of arms. MATERIAL. HOLOTYPE: NMVP108581. PARATYPE: NMVP108615 from NMVPL252. DIAGNOSIS. Ten arms, with prominent reg- ularly spaced pairs of spines every fifth brachial on outer face, with hexagonal intersecundibrach, with low but distinct anitaxis of hexagonal plates. DESCRIPTION. Crown subcylindrical, up to 45mm long, up to 4 times as long as wide. Cup subconical, about Іст long, longer than wide, with convex tegmen; cup plates thin, smooth. Infrabasals concealed by stem. Basals pent- agonal, except for hexagonal posterior one, with narrowest proximal margin forming part of circular rim to basal concavity, about as long as wide, with marked proximal medial projection (suggesting a pentagonal stem with the angles of the stem in the centres of basals beneath this projection), with Y-shaped median ray ridge dividing about midlength and extending onto adjacent radials. Radials heptagonal, much wider than long, with inverted Y-shaped ray ridge, with arms ofthis Y running onto adjacent basals. First 2 secundibrachs fixed in cup. Arms 10, of uniform cross section through available arm length; 1st primibrach hexagonal, not as wide as radial and little more than half as long; 2nd primibrach axillary, heptagonal; each arm uniserial with cuneate brachials as far as 6th or 8th secundibrach, biserial distally, with pair of strong solid spines on outer surface of rami placed about every 5 brachials; pinnules on each free brachial (i.e. distal to 2nd secundibrach) on each side of ramus, long, of 8-12 pinnulars. Interprimibrachs numerous, with heptagonal proximal one supporting 2 plates of about the same size, with size decreasing rapidly distally. First intersecundibrach hexagonal, subsequent plates not clear but presumably part of tegmen. Posterior interray with central anitaxis of hex- agonal plates; hexagonal primanal supporting 3 anals in next row; anal plates numerous, decreasing distally, with average size smaller than that of interbrachials. Stem unknown. REMARKS. With 10 biserial arms this species is placed in Dimerocrinites of Witzke & Strimple (1981) and is distinguished within that taxon by the spines on the outer arm faces, long hexagonal intersecundibrachs and the heptagonal axillary 2nd primibrach. Eucrinus Angelin, 1878 TYPE SPECIES. Eucrinus laevis Angelin, 1878 from the Wenlock of Gotland; by subsequent designation of Wachsmuth & Springer, 1881. REMARKS. This taxon is used in the sense of Witzke & Strimple (1981) to include species of Dimerocrinites with 20 arms; there seems little point in maintaining subgeneric distinction since we have no clear idea of phylogeny among these many species and when lineages are established it seems likely that both Dimerocrinites and Eucrinus will prove to be polyphyletic. The Victorian species is assigned on the basis that all rays countable have 4 arms except for one which has 5 and is considered aberrant; thus a total of 20 arms would be expected if branching is uniform. Eucrinus clarkae sp. nov. (Fig. 7) ETYMOLOGY. For Penny Clark who helped greatly with photography, curation and fieldwork. MATERIAL. HOLOTYPE: NMVP108642. PARA- TYPES: NMVP108643, 108645 all from NMVPL252. DIAGNOSIS. Basals, radials and radianal with distinctive coarse ornament of short fat radial FIG. 7. Eucrinus clarkae sp. nov. all from NMVPL252. A,C, lateral view of crown with stem and enlargement of distal cup and proximal arms of NMVP108645a, x2.5 and х4, respectively. B, lateral view of NMVP108643, x2, D, lateral view of NMVP108645b, x2.5. E, CD interray view of holotype NMVP108642, х4. CRINOIDS FROM CENTRAL VICTORIA 11 t MEMOIRS OF THE QUEENSLAND MUSEUM ridges normal to sutures and not joining centrally on plates. Crown with fine granulose ornament throughout. Intersecundibrachs few, in narrow vertical column, one in each row. Arms free distal io about 3rd tertibrach, uniserial proximally, secundibrachs becoming irregularly cuneate distally, tertibrachs becoming rectilinear biserial distal to about the 9th or 10th, Proximal brachials with distinctive fine ridge along proximal and distal margins, Stem round in section. DESCRIPTION. Crown flaring gently distally, up to 60mm Jong, Cup high bowl-shaped, of large thin plates, up to 15mm long. Infrabasals concealed by stem (fragments evident in holo- type between basals and stem but arrangement or number not available), Basals 5, with 4 of them pentagonal and posterior one larger and hexago- nal, with proximal margin raised into prominent rim to basal cavity and stem, with conspicuous nodes representing ray ridges 2 per infrabasal directed towards middle of sutures with contiguous basals but 3 on posterior basal (central one vertical in middle of posterior interray). Radials 5, heptagonal except fot hexagonal posterior one, largest plates in cup. with inverted Y-shaped ray ridges continuing from basals, with secondary radialing ridges horizontally onto adjacent radials. Primanal hexagonal, in radial circlen supporting 3 anal plates distally, with prominent radial ridges near and normal to margins bul not reaching centre of plate, lower central ridge from CD basal most prominent. Anal interray with large number of polygonal plates decreasing in size distally and only roughly in rows. First primibrach hexagonal, with broad low median ray ridge; 2nd primibrach axillary, heptagonal, with median ray ridge in Y-shape to enter 2 arms; [st secundi- brachs hexagonal, in sutural contact up middle of each ray, in contact with [st intersecundibrach. Secundibrachs fixed in cup but protruding laterally, with 6th axillary. becoming cuneate distally. Arms free distal to about 3rd tertibrach, cuneate uniserial up to 6th tertibrach then biserral and pinnulate, tapering gently, longer than theca: pinnules long, slender, of at least 10 or more pinnulars, attached to each free brachial on both sides of each arm; interprimibrachs numerous, slightly depressed between fixed arms, with single large plate in contact with 2 radials and Ist primibrachs and supporting next row of 2 interbrachials, with more distal rows becoming much smaller and less regularly arranged; intet- secundibrachs similar size to interprimibrachs at same height. Tegmen of many small polygonal plates (height of tegmen and anal opening nol available). Stem circular, heteromorphic, with nodals higher and of greater diameter than internodals, noditaxis N1. REMARKS, This species is distinguished within Ihe genus by the distinctive ornament on the proximal theca, the large Ist secundibrach, the low ray ridges not occupying full width of fixed ray plates and the large flat hexagonal Ist inter- primibrach. One half ray (Fig. 7A, C) has a normal branching on the 5th secundibrach then another branching on the 2nd tertibrach on the left division: this last branching is not seen anywhere else and is considered aberrant. The axillary tertibrach is cuneate with its long side in the Y of the previous division giving off the abnormal arm which then fills the Y of the previous division with 8-10 quite irregular, mostly cuneate brachials. The reason for this extra arm is not known. Eudimerocrinus Springer, 1926 TYPE SPECIES. Eudimerocrinus multibrachiarus Springer, 1926 from the Middle Silurian of Tennessee; hy monotypy. DIAGNOSIS, Cup subconical to deep bowl-shaped; basals with strong proximal projections. Arms 40, biserial in free section, with narrow гау ridges. Stem subpentagonal or at least not per- fectly round in section. REMARKS. In erecting this genus Springer (1926) identified the multiple arm branchings as the most significant feature, Ubaghs (1978b) maintained this feature and added a subpent- agonal stem. In view of the new Australian species the projections of the basals may be another feature to add to the generic complex. Witzke & Strimple (1981) placed Eudimerm- erinus with Ambicocrinus Kirk, 1945 and Griplioerinus Kirk, 1945 in a group of dimero- crinitids with more than 20) biserial arms. The latter genus is generally distinguished by some if not all its radials being separated by FIG. 8. Eudimeracrinus eckardti sp. nov. ull from NMVPL229. A, incomplete specimen NMVPTDS931, «1.5. B, partial cup and stem NMVP108963, «2,5. C. D, part and counterpart of fattened erown NMVP1091632 and b, *1.5. E, holotype crown NMVP]109113, <2 NMYVP109090, *2,5, G, crown NMV P0910, * 2,5. ME parag cup and inner side of arms from opposite side of cup с^, CRINOIDS FROM CENTRAL VICTORIA 14 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 9. Eudimerocrinus eckandti sp. nov. juveniles from NMVPL229. A, NMVP100492, x4. B, NMVP110645, x4.5. C, NMVP110647, x2.5. D, NMVP149393, x3. interbrachials thus placing it intermediate between Rhodocrinitoidea and Dimero- crinitoidea and allowing separation from the other 2 genera; how a species of Griphocrinus with all its radials in contact could be distinguished from Eudimerocrinus is not clear. On the other hand distinction of Ambicocrinus is less certain: it has 40 biserial arms, subpentangular stem and identical cup plating arrangement. Goldring (1923:84) commented on the highly crushed preservation of and difficulty of illustrating the type. 4. arborescens (Talbot, FIG. 10. Eudimerocrinus eckardti sp. nov. A, partial cup and arms NMVP109767 from NMVPL232, х2. B, partial cup and arms NMVP108689 from NMVPL229, х2.5. C, partial cup and arms NMVP109768 from NMVPL232, х2. D, distorted crown in posterior view NMVP108962 from NMVPL229, x2.5. un CRINOIDS FROM CENTRAL VICTORIA MEMOIRS OF THE QUEENSLAND MUSEUM 10 CRINOIDS FROM CENTRAL VICTORIA 17 1905), from the Lower Devonian (Lochkovian) of New York; she also noted its small size and indicated its depiction in her work was a re- constructionby the artist. Only the ray ridges and lack of projecting basals separate 4. arborescens from Eudimerocrinus so given the circumstances noted by Goldring these could well be due to preservation or the artist's licence: I think it highly likely that.4mbicocrinus 15 a junior syno- nym of Eudimerocrinus and that Griphocrinus may also prove to be synonymous when fully understood. Eudimerocrinus eckardti sp. nov. (Figs 8-11) ETYMOLOGY, For Steve Eckardt who has rendered enormous field assistance and many specimens from his private collection. MATERIAL. HOLOTYPE: NMVP109113. PARA- TYPES: NMVP100166, 100492, 108689, 108950, 108951, 108962, 108963, 109090, 109125, 109153, 109163, 109168, 109204, 110645, 110647, 149355, 149393 all from NMVPL229. Other specimens NMVP109221 and 109755 from NMVPL1924 and NMVP109767 and 109768 from NMVPL232. DIAGNOSIS Basal plates with prominent proximally directed projections near lower margin; median ray ridges variably developed, often prominent; Ist primibrach hexagonal; 2nd primibrach axillary, heptagonal; arms 40, free distal to 4th-6th secundibrach: stem with distinctive decalobate cross section. DESCRIPTION. Crown up to 55mm long. with cup and arms each about same length; cup up to 10mm long. high bowl-shaped, of large thin plates: infrabasals small, pentagonal, with conspicuous nodes on proximal margin (3 per infrabasal) in line with columns of lobes on stem. Basals 5. 4 hexagonal and CD basal heptagonal. largest plates in cup, with prominent proximally directed projections or spines medially near proximal margin, with spine extending beyond and concealing sutures between adjacent infrabasals, with faint median ray ridges in form of Y (Fig. 8E). Radials heptagonal, with inverted Y-shaped median ray ridge on most specimens. forming contiguous circlet around cup except in CD interray where hexagonal radianal intervenes incirclet: 1st primibrach hexagonal, with median ray ridge running in vertical line; 2nd primibrach axillary, heptagonal, with Y-shaped ray ridge into arms; Ist secundibrachs hexagonal. in sutural contact up middle of each ray, in contact with Ist intersecundibrach: first 6 secundibrachs fixed in cup; free arms biserial and pinnulate distal to 1st free secundibrach, branching approx- imately at secundibrach 10-11 and tertibrach 11-12, but branching often not at same level even in one ray, of relatively small diameter, tapering gently, approximately as long as theca; adoral groove wide and deep; pinnules long. slender, of at least 4-5 pinnulars, attached to each free brachial on both sides of each arm. Inter- primibrachs numerous, not depressed between fixed arms. with single large plate resting on shoulders of 2 adjacent radials, with 2nd row of 2 plates, 3rd row of 3 plates level with top of primary axil, distally becoming much smaller and less regularly arranged with rows having 3-6 plates. Intersecundibrachs with Ist same size as interprimibrachs at same level. Anal interray wider than others, with primanal supporting 3 anals distally, subsequent anal plates without obvious order, decreasing in size to same extent as in interprimibrach series. Tegmen of many small vertically elongate plates, apparently quite long (almost as long as cup): anal opening not seen. Stem decagonal in section, heteromorphic. with noditaxis N3231323. with lobes aligned vertically in 10 columns to give stem appearance of fluted column with differential horizontal layering. no complete stem available. MORPHOGENY. Small crowns (Fig. 11) have most of the cup occupied by the basals, radials. and first interbrachials, the diameter of the free arms compared to cup diameter is larger than the same ratio in large individuals and free arms are cuneate uniserial. These specimens are assigned to 4. eckardti mainly because of the basal plate projections and similanty in every other feature available. These differences indicate that through growth the arms change from cuneate uniscrial to cuneate biserial and then to rectilinear biserial; the arms do not increase in diameter at the same rate as the cup: and the cup increases in size in more distal parts with addition of interbrachial plates and greater growth there than in lower parts of the cup. FIG. 11. Kudimerocrinus eckardti sp. nov. ^, partial specimen with inner side of arms from opposite side of cup NMVP109221 from NMVPL 1924, «2.5. B, crown NMVP149355 from NMVPL229, 2.5. C. cup and some arms NMVP109755 from NMVPL 1924, x3.5. D, cup and partial arms NMVP 109204 [rom NMVPL 229, 2.5. E, cup and stem NMVP100166 from NMVPL229, 18 MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 12. Eudimerocrinus gilli sp. nov., holotype crown ММУР 149345 from NMVPL1990, x3. REMARKS. This species is distinct within the genus in the detail of its stem. more subtle ray ridges and narrower arms relative to cup plates. It most closely resembles 4. arborescens (type species of mbicocrinus Kirk, 1945) from the Lower Devonian of New York which is distin- guished by lacking the basal plate projections. Two specimens from NMVPL232 (Fig. 10A, B)are assigned to this species but differ in having à subdued radial ornament on interprimibrachs, radials, primibrachs and secundibrachs and in having the second arm branching farther from the primaxil by about 4-6 brachials. They do have the projections on the basals and so are grouped with the material from NMVPL229 knowing that future material may show them to be part of a separate taxon. Eudimerocrinus gilli sp. nov. (Fig. 12) ETYMOLOGY. For the late Edmund Gill for his geological work around Lilydale. MATERIAL. NMVPL1990, DIAGNOSIS. Infrabasals barely evident in lateral view; basals with prominent rounded proximal projections; secundibrachs all fixed in сир; arms branching at least 3 times; stem subcircular to subpentagonal. DESCRIPTION. Crown 32mm long. sub- cylindrical. Cup deep bowl-shaped, about 12mm long; cup plates thin, smooth except for prom- inent ray ridges. Infrabasals 5, mostly concealed by stem, upper tips just visible at top of stem between basals. Basals 5, hexagonal. except for pentagonal posterior one, with proximal margin forming part of circular rim to stem attachment, about as long as wide, with prominent well- rounded proximal projection. with median ray ridges emanating from the projections and running onto adjacent radials at right angles to interplate suture. Radials 5, heptagonal, wider than long. with inverted Y-shaped ray ridge. Arms apparently 40, with all primibrachs and secundibrachs fixed in cup. with free arms biserial, tapering distally: Ist primibrach hexagonal, not as wide or as long as radial, with median ray ridge across it; 2nd primibrach axillary, heptagonal, with Y-shaped ray ridge: secundibrachs 6, with median ray ridges, subquadrate or hexagonal; 6th axillary, with 2 round laterally (but not outwardly) declined facets: branching in free arms asy mmetrical, with 6th tertibrach axillary in free arms of C and D rays immediately adjacent to CD interray but with next adjacent free arm in C ray having 12th tertibrach axillary and in D ray having 9th tertibrach axillary ( branching in A, B and E rays unknown). Interprimibrachs large, numerous, with proximal one supporting 2 plates of about the same size, with size decreasing rapidly distally: CD interray with Ist interprimibrach supporting 3 plates. wider than other interradial areas. First intersecundibrach hexagonal. sup- porting another 6-8 plates in 3 or 4 rows. Stem circular in section. noditaxis N212, with many low ossicles having wide epifacets (slightly wider in nodals than internodals). REMARKS. Ubaghs (1978b) distinguished Eudimerocrinus from Dimerocrinites by its arms HOLOTYPE: NMVP149345 from CRINOIDS FROM CENTRAL VICTORIA 19 FIG. 13. Nexocrinus wallanensis sp. nov. from NMVPL 1923. A, lateral view of holotype crown NMVP100116, х3. B.C, posterior view of crushed crown (and enlargement of cup) NMVP107111, x2 and х5, repectively. dividing several (presumably more than twice) times and its subpentagonal stem; К. gilli is thus assigned to Eudimerocrinus. It is distinguished from the genotype by the axillary secundibrachs being fixed in the cup. the cup being more bowl-shaped and the arms tapering more rapidly distally. Nexocrinus Eckert, 1984 TYPE SPECIES. Nexocrinus delicatulus Eckert, 1984 from the Llandovery of Ontario; by original designation. OTHER SPECIES. Ptychocrinus parvus Hall, 1872 trom the Upper Ordovician of North America; Nexocrinus wallanensis sp. nov. Ludlow of Victoria; Dimerocrinites occidentalis (Hall, 1863) from the Middle Silurian Waldron Shale of USA: Nexocrinus sp. Lochkovian of Victoria (herein), Prtychocrinus adamensis Ausich & Dravage, 1988 from the Llandovery of Ohio. DIAGNOSIS. Cup conical; CD interray slightly wider than others; ray ridges prominent, may be subdued on proximal cup plates: infrabasals concealed by stem to long in lateral view. Posterior basal hexagonal. supporting primanal that separates C and D radials; arms 20, 4 per ray, uniserial, pinnulate, much longer than cup. REMARKS. Brower (1973) dealt in detail with Ptychocrinus noting that the 3 Upper Ordovician species assigned are probably generically distinct. Eckert (1984) agreed that the genus was in need of revision and suggested that P. parvus should be removed because it had 20 arms unlike any other species included. He noted the similar- ity of his new genus to P. parvus but separated them on the number of fixed secundibrachs. width of the CD interray. distinctness of the anal ridge and plating of the posterior interray. Of these features the new Victorian species shares 20 MEMOIRS OF THE QUEENSLAND MUSEUM with N. delicatulus the 4 fixed secundibrachs, the distinct anal ridge and the plating arrangement of the posterior interray. The only major distinction of the Victorian species is that its infrabasals are fully in view laterally and the ray ridges become less distinct proximally. I consider these to be interspecific differences and not of generic significance in the same way that the differences between P. parvus and N. delicatulus noted by Eckert (1984) distinguish those species within one genus. Ptychocrinus adamsensis Ausich & Dravage (1988) from the Llandovery of Ohio could be assigned to Eucrinus since it has 20 biserial (partially) arms. However, Ausich & Dravage (1988) made a good case for assigning it to the previously exclusively uniserial Ptychocrinus; they did not consider comparison with Nexocrinus which is a ptychocrinid genus with 20 uniserial arms distinguished by axillary 4th secundibrachs and some features of the post- erior interradius. Since the posterior interradius of adamsensis is not available its features cannot be compared. However, the second division of the arms is at the 4th or 5th secundibrachs in adamsensis and I consider this species should be assigned to Nexocrinus. Ausich & Dravage (1988) argument that a species with uniserial proximal arms becoming biserial distally can belong to an essentially uniserial genus com- mands re-assessment of species with such arms. A single specimen from the Lower Devonian of Victoria with such arms, though they become biserial a little more proximally than in adamsensis and with 6th secundibrachs axillary, is assigned to this genus herein. That specimen is comparable to Dimerocrinites occidentalis (Hall, 1863) from the Silurian Waldron Shale of central USA on features of the cup but since arms are unknown for the latter species its assignment is equivocal. Nexocrinus wallanensis sp.nov. (Fig. 13) ETYMOLOGY, From near the town of Wallen. MATERIAL. HOLOTYPE: NMVP100116 and paratype NMVP107111 from NMVPL1923, DIAGNOSIS Infrabasals long in lateral view; ray ridges prominent above the radials, becoming less distinct proximally. Primibrach 2 and secundibrach 4 axillary; arms fixed in cup up to about tertibrach 4 or 5. Arms 20, 4 per ray, uniserial, with primibrach 2 and secundibrach 4 axillary. Interrays with large number of polygon- al plates (1 in first row followed distally by rows of 2, 3, 4 then irregular). Posterior interray with large basal longer than other basals; primanal directly distal, hexagonal, supporting 3 plates distally; prominent anal ridge medially on central column of vertically elongate hexagonal anal plates. Surface of plates smooth. DESCRIPTION. Crown more than 60mm long, flaring distally. Cup high conical, with most of length in tegmen, with some tertibrachs fixed in cup. Infrabasals longer than radials, pentagonal, with highly obtuse angle distally. Basals hexagonal, large, largest plates in cup, with very faint broad ridges in X-shape crossing proximal and distal sides normal to margins. Radials heptagonal, in contact with adjacent radials except presumably in CD interray, with subtle ridges from basals continuing to meet medially near upper margin and continue as more prominent single ridge into brachials. First primibrach hexagonal, more convex in arm section distally; 2nd primibrach axillary, hept- agonal, in contact with 2nd and 3rd rows of interbrachial plates; 4th secundibrach axillary; all secundibrachs and 2-3 tertibrachs fixed in cup. Free arms 4 per ray, with short fine pinnules, longer than cup, uniserial, with length of brachials decreasing gradually distally; pinnules one per brachial, alternating from side to side along arm. Interprimibrachs numerous, in regular rows of 1, 2, 3, 4, with Ist resting on shoulders of radials, with distally large number of irregular polygonal plates forming large but presumably not very competent tegmen; intersecundibrachs and intertertibrachs not clearly evident. Anal interray with basal longer than other basals, with distinct median ridge from infrabasal to tegmen; primanal and subsequent median plates long, hexagonal, with greatest width above midlength; other anal plates large, polygonal. Stem sub- circular to subpentagonal in section, with pentagonal lumen. REMARKS. This species is assigned to Nexo- crinus on cup shape, ray ridges, shape of plates in the median anal column, 20 uniserial arms and depressed interprimibrachs. It is distinguished within the genus by the length of the infrabasals in lateral view. This species may ultimately prove to be generically distinct but in the present state of knowledge this is the most likely placement. Nexocrinus sp. (Fig. 14) MATERIAL. NMVP149343 from NMVPL252. CRINOIDS FROM CENTRAL VICTORIA 21 FIG. 14. Nexocrinus sp. incomplete crown in lateral view external (A) and internal (B) NMVP149343 from NMVPL232, х4. DESCRIPTION. Cup high conical, of thin plates. with median ray ridges well-developed, with radial ornament on interbrachial areas. Infrabasals 5, small, visible in lateral view, with ray ridges from adjacent basals meeting medially on the lower margin. Basals 5, hexagonal (post- erior one presumably heptagonal and supporting 3 anal plates distally), smaller than radials. with cross of broad ray ridges from adjacent radials and infrabasals. Radials 5. heptagonal, much wider than long, with inverted Y-shaped ridges forking at midpoint of plate. with much thinner and lower ridges radiating from the midpoint to the middle of the other 4 sides. First primibrach hexagonal, wider than long, crossed vertically by prominent ray ridge, with faint lateral ridges from centre to sutures with interbrachials. Second primibrach pentagonal. axillary, with ray ridges occupying most of plate and dividing into both rami, with lateral marginal flange and upper tip depressed to interray areas. First secundibrachs and tertibrachs in contact laterally. Secundi- brachs uniserial, cuneate. fixed, 6th axillary, with number of small long narrow intersecundibrachs up to level of Ist tertibrach. Tertibrachs cuncate for proximal 3 or 4 then biserial, with long pinnules on each brachial on each side of arm in biserial part (full length of arms not available). Interprimibrachs numerous, 1 in first row more distal rows with 2, 3. 4, then becoming more elongate and less regular, with fine radial ornament. Anal interray unknown. Internal mould of cup with a groove in cup plates matching the positionofthe ray ridges, beginning on the radials, becoming deeper and narrower upwards onto the arms, suggesting that similar though shallower and less distinct grooves branched from the 3rd secundibrach out and up across lateral interprimibrachs. Stem circular in section, noditaxis N1, of very low ossicles of uniform length but with epifacet of nodals of greater diameter than that of internodals. REMARKS. This specimen resembles Dimero- crinites occidentalis (Hall, 1863) from the Silurian Waldron Shale of Indiana with which it shares the infrabasals well exposed in lateral view, narrow ray ridges, fine radial ornament on interbrachials and fine longitudinal ornament on the ray ridges. However, it may be distinguished by its larger diameter stem (relative to thecal diameter). the ray ridges on basals in an X rather than a Y as in occidentalis and its 20 arms as opposed to the 10 surmised in occidentalis. No illustrated specimen of D. occidentalis is complete in the arms distal to about the 2nd 22 MEMOIRS OF THE QUEENSLAND MUSEUM VIG. 15. Duncanicrinus calvariolus gen. et sp. nov. A-C, NMVP110649 trom the rubbish tip on Watson's Road, Pheasant Creek, Kinglake. A, E ray enlarged view of cup. «10. B, enlarged basal view of cup with posterior basal at 12 o'clock, «10, C, crown in E ray view with distal arms incomplete, *5. D,E, part and counterpart of slightly disarticulated NMVP108647 from NMVPL252, x4. secundibrach. This Australian specimen illustrates that the second division of the arms is on the 5th or 6th secundibrach so it is quite possible that the arms of D. occidentalis may have divided a second time to produce 20 arms. Until the arms of this North American species are known the comparison must remain incomplete but I suggest it is more likely that occidentalis belongs to Nexocrinus than to Dimerocrinites or Euerinus, CRINOIDS FROM CENTRAL VICTORIA L2 К] sp г, a: - E =a d pæ FIG: 16, Duncanicrinus calvariolus gen. et sp. nov., all crowns from NMVPL252. A.B. NMVP 108640. <4 and x6, respectively. C. NMVP108634. «3, D.E. NMVP108617, x6. F, NMVP108633. «2.5. Duncanicrinus gen. nov. ETYMOLOGY. For Peter Duncan of the Whittlesea district. a benefactor of Victorian palaeontology. TYPE SPECIES. Duncanicrinus calvariolus sp. nov. 24 MEMOIRS OF THE QUEENSLAND MUSEUM OTHER SPECIES. Ptychocrinus fimbriatus (Shumard, 1855) from the Ordovician of USA; Ptvchocrinus longibrachialis Brower, 1975 from the Silurian of Scotland. DIAGNOSIS. Cup conical, of smooth or tub- erculate plates, with or without ray ridges. Infrabasals 5, pentagonal, concealed by stem FIG. 17. A.B, Duncanicrinus calvariolus gen. et sp. nov., basal views of incomplete crowns from NMVPL1924. A, NMVP109579, x4. B, NMVP109761, x5. С, Pterinocrinidae new genus, splayed crown in proximal view, with posterior interray at 1 o'clock, NMVP149389 from NMVPL 1990, x1.1. attachment or partially visible in lateral view. Basals 5, hexagonal; posterior basal longer than others, reaching more than halfway up radials. Radials in lateral contact except in posterior interray. Interprimibrachs few. beginning with one resting on radials. Tegmen of small polygonal plates. Arms 10, uniserial, of cuneate brachials, with long pinnules on prominent facets CRINOIDS FROM CENTRAL VICTORIA 25 alrermating along arm. Stem circular, noditaxis N3231323, some with cirral spines at noditaxis bise. REMARKS. Brower ( 1973) noted that within the Dimeroennitidae the 3 Ordovician species of Pivehocrinus could be regarded as belonging to separate genera. In discussing phylogeny he suggested that the formis with 10 arms may belong to one lineage and could have given rise to the 10 armed Dimeracrinites, Accepting Brower’s (1973) phylogeny the Victorian species is placed with Pachocrinus fimbriatus in à genus characterised by 10 uniserial arms that was foreshadowed by Jell (in Jell & Holloway, 1983) in discussion of the holotype, then the only known specimen, Macurocrinus Jaekel, 1895 is a related genus but its type, M. springeri Jackel, 1895 has 15 arms and is considered generically separate. Species of Macarocrimuis with 10 arms should probably be transferred to Diaicanicrinus but detailed study of those species is outside the scope of this paper. Duncanicrinus calvariolus sp, nov, (Figs 15-17) Dimerovrinilidae gen. el sp. nov. Jell in tell & Holloway. 1983712. Поя TOK 8 ETYMOLOGY. Latin colvariola, small cup. MATERIAL. HOLOTYPE: NMVP74246 from the floor of Winncke Reservoir, Christmas Hills, E of Melbourne described and figured by Tell (in Jell & Holloway, 1983:12, figs 7C-b, 8) as Dimerocrinitidae gen. et sp. nov. PARATYPES: NMVPILL0649 from the rubbish tip on Watson's Road, Pheasant Creek, Kinglake and NMVP 108617. 108633, 108634, 108640, 108647 all from NMVPL252, Other material NMVP109579, 109761 and 109763 from NMVPI. 1924. DIAGNOSIS. Cup very small, of smooth plates. without ray ridges. Infrabasals may or may not be visible in lateral view. Interprimibrachs. few, maximum of 6-8 per interray. Arms (тес distal to primaxil, Stem cirriferous. DESCRIPTION. Crown up to 30mm long, strongly flared in proximal part then sub- cylindrical. with arms more than 5 times as long as cup. Cup very small (about 2min across and 1.5mm long), conical, of smooth thin plates, with shallow sharply rimmed stem facet. Infrabasals 5. completely concealed by stem (Fig. 15B), just visible laterally as wide low triangle (Jell & Holloway, 1983, fig. 7C) or fully visible and pentagonal (Fig. 150). Basals 5, usually hexagonal, maybe pentagonal with proximal margin slightly curved and forming sharp margin іо basal concavity. occupying up to 60% of cup length, with acute distal tip at pomt where racials begin to descend into interradial depressions: posterior basal heptagonal, supporting primanal above horizontal suture. remainder of anal sac uncertain but apparently with median anitaxis of hexagonal plates, Radials 5. heptagonal. contacting cach other laterally through lateral flanges in interradial depressions: radial neel реперіепагу (c.0.8 of radial width), horizontal, subeirewlar in section, Each primibrach: with latera] flanges in interradial depression in line with those on radials. Arms 10. with axillary 2nd primibrach. tree distal to primaxil, of cuneate secundibrachs, with long pinnules alternating side to side along each arm; each pinnule of 3 pinnulars. Interprimibvachs convex, in regular rows 1, 2, 3 distally, Stem circular in section. noditaxis N3231323, with nodals each hearing stout cirri. REMARKS. The specimen from Winneke Res- ervoiris now fully interpretable with discovery of the several specimens from different localities al| just to the NE of Melbourne. This species is distinguished from D. fimbriatus by its smaller cup. fewer smaller interprimibrachs and cirriferous stem, There 1s some variation within the species it the amount of the infrabasals visible laterally, in the interbrachials and in the shape of the brachials but none of this variation can be considered interspecific. The discussion of the holotype provided by Jell (in Jell & Holloway, 1983) indicated close relationship to Ptychocrinus and this is confirmed here with closest relationship to Prychocrinus finbriatus now assigned to Duncanicrinus, Family PTERINOCRINIDAE Melntosh, 1987 Pterinocrinid gen. nov. (Fig. 17C) MATERIAL. NMVP149389 from NMVPL1990. DESCRIPTION. Crown splayed on beddiüg plane, approximately 50mm in diameter. Cup low bowl-shaped (probably more conical originally and now shortened by compaction), with plate margins not discernible: ornament of prominent ray ridges. Infrabasals not discernible. Basals presumably 5, not discernible except posterior one intervening between C and D radials; posterior basal 5-sided, with strong central boss and 5 radiating ridges lo centre of each side. Radials 5, 5-sided, with 2 ray ridges from centre 26 MEMOIRS OF THE QUEENSLAND MUSEUM of2 proximal sides meeting at centre of plate then running to centre of distal síde; C and D radials with extra ridges to anal X. Posterior interray wider than others; anal X large, resting on basal, separating C and D radials, supporting 3 anal plates, with ray ridges running vertically and 2 running diagonally from adjacent radials; distal anal plates not clearly defined but with strong tubercles or short spines. Arms 40 (if branching is regular; E ray appears to have one branch missing the 3rd division and another with a 4th division - maintaining the 8 branches for the ray but irregularly; C and D rays branching regularly); primibrachs 2, with Ist hexagonal, with 2nd pentagonal and axillary; secundibrachs not readily countable due to preservation but approximately 5 or 6, 3 or 4 fixed in cup: free brachials with raised proximal and distal margin- al rims leaving a distinct groove at midlength, each with 2 long slender pinnules, one on each side of arm; pinnules with more than 6 pinnulars each. Stem pentagonal in section, with angles of stem aligned with ray ridges bisecting proximal margin of each basal, heteromorphic, with lateral extremities of each columnal at 5 stem angles appearing to be curved distally (possibly due to preservation). REMARKS. Although infrabasals are not clear there are many dimerocrinitoids in which they are concealed by the stem and the plate arrangement is not characteristic of monocyclic camerates. I therefore, infer concealed infrabasals. The com- pound biserial brachials assign the taxon to the Pterinocrinidae within this superfamily and the prominent ray ridges separate it from all other members of the family except Apurocrinus McIntosh (1981) from which it is distinguished by its different branching pattern. McIntosh (1987) described cornice-shaped rims along distal lips of free brachials in the other South American Devonian pterinocrinid Bogotacrinus scheibei; its brachials are thus very similar to those of the Victorian taxon where a similar rim also occurs on the proximal lip. Bogotacrinus is also comparable in having a pentagonal stem in the same relationship to the basal circlet. How- ever, Bogotacrinus lacks strong ray ridges and has a different arm branching pattern. Relatively poor preservation of the single specimen avail- able precludes application of a formal name so I retain the new taxon in open nomenclature. Although McIntosh (1979) described the columnal in the stem facet as round, his figure (McIntosh, 1979, fig. 5.1) clearly shows it to be subpentagonal with each angle at the centre ofthe proximal edge of a basal. Order MONOBATHRIDA Moore & Laudon, 1943 Suborder COMPSOCRININA Ubaghs, 1978 Superfamily HEXACRINITOIDEA Wachsmuth & Springer, 1885 Family HEXACRINITIDAE Wachsmuth & Springer, 1885 Hexacrinites Austin & Austin, 1843 TYPE SPECIES. Platycrinites interscapularis Phillips, 1841 from the Middle Devonian of England; by monotypy. REMARKS. This genus is widespread through- out the Devonian of Australia (Philip, 1961 - L. Dev.; Jell et al., 1988 - M. Dev.) This is the first species with upper arms preserved and with unweathered plate surfaces. Hexacrinites chirnsidensis sp. nov. (Fig. 18) ETYMOLOGY. From Chirnside Park, an outer suburb of Melbourne, near Lilydale. MATERIAL. HOLOTYPE: NMVP107097. PARATYPES: NMVP107095, 107096, 107098, 107100, 110644 all from NMVPL1922. DIAGNOSIS. Low but distinct median ray ridges on all cup plates except primanal; Ist primibrach and 4th secundibrach axillary; 4 arms per ray. Stem circular; columnals almost as high as diameter, with protruding flange near midlength. DESCRIPTION. Crown up to 25mm long, con- ical proximally, less flared distally. Cup high conical, of unornamented plates except for ray ridges. Basals 3, equal, symmetrically beneath A, C and D radials, with a median ray ridge centrally on A basal leading onto A radial, with 2 other ray ridges from A basal continuing onto B and E radials, with 2 median ray ridges on each of other 2 basals continuing onto B, C and D, E radials, respectively (Fig. 18A), with distinct rim around stem attachment giving rise to ray ridges. Radials 5, up to twice as long as wide, each with distally decreasing median ray ridge; radial facet about 1/2 radial width, declivate, semicircular to horseshoe-shaped on outer but only weakly con- vex on interior. Primanal same size as radials, smooth, without ray ridge or facet. Arms 20, 4 per ray, uniserial, with deep groove on inner side of each. First primibrach axillary, pentagonal; sec- undibrachs 4 per arm, 4th axillary, subquadrate in lateral view, almost circular in section except for CRINOIDS FROM CENTRAL VICTORIA 27 FIG. 18. Hexacriniteschirnsidensissp.nov.. allincomplete crowns from NM VPL 1922. А.С. part and counterpart of holotype NMVP107097, «3.5 and 74, respectively, B_G, part and counterpart of NMVP107098. х4 D, internal mould (not a latex cast) of NMVP107095. «5. E, NMVPI07100, х6. Е, NMVP107096. 52.5, groove on inner side. apparently not pinnulate: tertibrachs becoming cuneate distally. with large well-developed facet for attachment of pinnules: pinnules one per brachial. alternating from side to side up each arm (distal extent of arms not available). Single large plate of tegmen adjoining radials between radial facets. Stem circular in section, with strongly crenulate sutures between columnals throughout, heteromorphic proximal- ly; proximal section of short columnals with lateral flange at midlength. with flange slightly wider on nodals: distal section of uniform columnals. long (as long as 4 proximal columnals), with lateral midlength flange occupying only small part of length of columnal. REMARKS. Most species of /exacrinites have some obvious and distinctive ornament on the basal and radial plates: a few are smooth (some illustrated specimens may be smooth through postmortem weathering) but none of the smooth species have the subtle ray ridges of H. chirnsid- ensis. A similar species is Arthroacantha vega Prokop. 1982 which shares the very long basals, 2» MEMOIRS OF THE QUEENSLAND MUSEUM subtle median ridge (‘considerably convex, ts- pecially in their middle parts’ Prokop, 1982) on radials but not on radianal and horizontal depres- sion of radial plates just proximal to distal margin. It 15 distinguished by the tuberculate ornament which is the only feature that could be used toassign the Czech species to /Irrrogeantha However, numerous species of evacrinites have tuberculate ornament and T suggest that 4, vega probably belongs to Hexacrinites closely allied to AL chiensidensis. Similar also is the specimen of exacrinites deseribed by Philip (1961) which is available as an interior mould having а distinct median convexity, a horizontal groove just proximal їо the distal margin and long conical shape all the same as in // chirnsidensts; it is quite possible that Philip's specimen and the Chirnside Park material are conspecific, Oehlerticrinus LeMenn, 1975 CYPE SPECIES, Ochlerticrimus. selllotensis: LeMenn, 1975 from the Lower Devonian ol France: by original destgnation. REMARKS. LeMenn (1975) erected this genus for 5 species for hexacrinitids with strongly and distinctively ornamented cup plates, with 2 primibrachs, with organised interbrachial plating, with compound brachials and with a cirriferous stem. The last 2 features are evident in only one species cach and their occurrence in the other species are inferred. Although the 2 species added herein have uniserial arms and non- cirriferous stems they are assigned to this genus because 1) basic cup plate ornament is evident though less striking; 2) there are 2 primibrachs; 3) the primanal supports 3 plates; and 4) a single large interbrachial rests on adjacent radials and is surrounded distally by small polygonal plates. OF other hexacrinitids with 2 primibrachs drthroacantha Williams, 1883 from the Dev- ошап of Europe and North America has spines articulated on tubercles on the cup and biserial arms. that branch 3 times and lacks a linear cup plate ornament, /"Iatyhexaerinus Schmidt, 1913 trom the Devonian of Europe and Siberia is similarly distinguished. except that it lacks the сир spines and has a very inflated tegmen, while Prohexacrinus Yakovlev, 1946 from the Silurian of the Urals also has a subglobose cup and almost equidimensional radials. When the stem and arms are known for all species the variation here inferred may suggest further subdivision but at present assignment to Qehlerficrinus is consider- ed most hkely- Oehlerticrinus lemenni sp. nov. (Figs 19, 20) ETYMOLOGY, For Jean Le Мела, Ше author of the genus, MATERIAL. HOLOTYPE: NMVP109213. PARA- TYPES: NMVPLO9098, 109116, 109164, 109166, all from NMVPL229, NMVPI08672, 108679 and 108682 from NMVPL252. DIAGNOSIS. Cup high conical. Ornament of strong narrow ray ridges single on A, C, and D radials. double on B and Е radials and primanal, with slightly less prominent sharp horizontal ridges around the cup at height of radial facet and just proximal to it, with background ornament of fine tubercles over entire cup. Tegmen with 5 large subtriangular 1nterambulaerals surrounded orally by small irregular ambulacrals. Arms 20, rectilinear uniserial Stem circular in section. heteromorphie with strongly crenulate inter- columnal suture, with columnals becoming longer distally. DESCRIPTION, Crown up to 48mm long, with arms flaring distally and more than twice as long as cup. Cup high conical, consisting of basals, radials and. primanal. Basals 3, equal, forming hexagonal circlet, symmetrically beneath A, С and D radials, occupying about 30% of cup length, each with 3 ridges radiating from distinct rim around stem attachment and crossing each of 3 distal edges. Radials 5, up to twice as long as wide, about 70% of cup length, with narrow sharp ridges continumg from basals, with similarly narrow and sharp horizontal ridges running around the cup at height of facet and just proximal (о jt, horizontal ridges fading out on approach to vertical ridges, with background of fine tubercles over entire сир; A, С and D radials with single median vertical ridge; B and E radials and primanal each with 2 ridges quite close together and converging on base of radial facet; radial facet about 1/4 radial width, declivate, semicircular to horseshoe-shaped. Primanal sante size as radials, smooth, without facet bul with strong tuberele at point where 2 vertical ridges meet, with single ridge continuing. from this tubercle to distal margin, with distal margin FIC. 19. Ochlerticrinus lemenni sp, nov, all from NMVPL229, А.В,Е, holotype cup and stem NMVP 109213. A, lateral C ray view, v4. B, lateral A ray view, х4. F, oblique posterior tegminal view. х7. C, lateral C ray view NMVPIS098, x3, D, partial crown NMVPTO09164, <4, E, crown NMVPI09116. *2,5, CRINOIDS FROM CENTRAL VICTORIA 30 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 20. Oehlerticrinuslemenni sp. nov. all from NMVPL232.A. crown with indistinct preservation of cup plates NMVP108682a, »2.5. B,D, enlargement of cup and crown NMVP108679. x6 and x4, respectively. C. crown NMVP108672, x5. 32. А, 2.5. E; FIG. 21. Qehlerticrinus Jeani sp. nov., all crowns (В and F with arms missing) from NMVPL2 NMVP108613. x2.5. В, NMVPI08664, «3.5, C. holotype NMVP108626a, «2.5. D. NMVP108614, «2.5. NMVP108660, «4. Е. NMVP108618. хб. CRINOIDS FROM CENTRAL VICTORIA ny e. Lote OTE OM A Meu Met rea ti "A MEMOIRS OF THE QUEENSLAND MUSEUM CRINOIDS FROM CENTRAL VICTORIA of 2 straight sections, supporting 2 large anal plates. Arms 20, 4 per ray, rectilinear uniserial, usually with fine close-spaced longitudinal ridges, more than twice as long as cup. Second primibrach axillary, pentagonal; secundibrachs 4 per arm, 4th axillary, subquadrate in lateral view, almost circular, apparently not pinnulate; tertibrachs becoming cuneate distally, with large well-developed facet for attachment of pinnules; pinnules one per brachial, alternating from side to side along each arm. Tegmen with 5 large sub- triangular interambulacral plates each resting symmetrically on 2 radials, with small irregular ambulacrals. Stem circular in section, with strongly crenulate sutures between columnals throughout; proximal section of low columnals with lateral flange at midlength, with flange slightly wider on nodals, noditaxis N212; distal section with longer (as long as 2 proximal columnals) columnals, with lateral midlength flange occupying only small part of length of nodals; internodals without lateral flange; noditaxis NI. REMARKS. This species is distinguished from European members of the genus by its ornament, uniserial arms and noncirriferous stem. It is separated from O. jeani sp. nov. by its ornament and by its 20 rather than 10 arms. Oehlerticrinus jeani sp. nov. (Figs 21, 22) ETYMOLOGY. For Jean Le Menn, the author of the genus. MATERIAL. HOLOTYPE: NMVP108626. PARA- TYPES: NMVP 108613, 108614, 108618, 108664, 108666, 108667, 110641, 149375, 149376 all from NMVPL252; NMVP109150 from NMVPL229. DIAGNOSIS. Cup ornament of low narrow ray ridges single on A, C, and D radials, double on B and E radials and radianal, with slightly more prominent horizontal ridge around the cup just proximal to radial facet and incomplete at each facet, with secondary ornament of fine broken wavy linear ridges parallel to the main ridges. Primanal supporting 3 anal plates. Arms 10, uniserial, of cuneate brachials. Stem circular in section, heteromorphic with strongly crenulate intercolumnal suture, with columnals becoming longer distally. „Өө 2 DESCRIPTION. Crown up to 35mm long, sub- cylindrical, arms at least 3 times as long as cup. Cup high conical, consisting of basals, radials and primanal; cup plates with ornament of fine, often wavy ridges, occasionally anastomosing, sometimes incomplete and then almost tubercular, subparallel to main ray ridges and thus forming more or less triangular patterns between midlines of adjacent radials and line joining adjacent radial facets. Basals 3, equal, forming hexagonal circlet, symmetrically proximal to A, C and D radials, occupying about 1/3 of cup length, each with 3 ridges radiating from distinct rim around stem attachment. Radials 5, up to twice as long as wide, occupying about 2/3 of cup length, with narrow ray ridges continuing up from basals, with narrow horizontal ridge running around the cup just proximal to facet but usually incomplete at the midline of radials: A, C and D radials with single central ridge; B and E radials and primanal each with 2 ridges converging on base of radial facet; radial facet about 1/3 radial width, declivate, semicircular to horseshoe-shaped. Primanal same size as radials, without facet, with single vertical ridge continuing distal to level of radial facets to distal margin medially, supporting 3 anal plates distally. Arms 10, 2 per ray, uniserial; 2nd primibrach axillary, pentagonal; secundibrachs becoming cuneate, with large well-developed facet for attachment of pinnules; pinnules | per brachial, alternating from side to side along each arm, of 6-10 pinnulars. Tegmen with 5 large subtriangular interambulacrals each resting symmetrically on 2 radials, with small polygonal ambulacrals. Stem circular in section, with strongly crenulate sutures between columnals; proximally low columnals with lateral flange at midlength, with flange slightly wider on nodals, noditaxis N212; distal section of uniform columnals, longer (as long as 2 proximal columnals), with lateral midlength flange occupying only small part of length. REMARKS. Oehlerticrinus jeani is disting- uished from European species by its ornament, its 10 uniserial arms and its noncirriferous stem. It is distinguished from the closely related and co- occurring О. lemenni under that species above. Available specimens have been crushed so that many plates have disslocated at sutures while FIG. 22, Oehlerticrinus jeani sp. nov. A, crown NMVP108667 from NMVPL252, х3. B, juvenile crown NMVP149375, from NMVPL252, x4. C,D, external and internal of NMVP149376, x3. E, crown NMVP109150 from NMVPL229, x4. F, cup with some arm fragments (counterpart of Fig. 21C), holotype NMVP108626 from NMVPL232, x4. 34 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 23. Frankocrinus holmesi gen. et sp. nov. crowns from NMVPL229. A,B, holotype in D ray view NMVP109099, x4 and x1.4. С. NMVP109109, x3. others have fractured across the plate itself; this is particularly so near the upper margins of radial plates suggesting numerous reentrants into the radials. However, І have been unable to discern any Clear pattern and maintain that the upper margins of the radials formed a smooth circle above which the tegmen was sutured. Frankocrinus gen. nov. TYPE SPECIES. Frankocrinus holmesi sp. nov. ETYMOLOGY. For Frank Holmes of Melbourne. DIAGNOSIS. Cup high conical, of smooth or finely ornamented plates. Radials with broad low ray ridges and horizontal flexure at level of radial facets, distal to which radials depressed between arms. Second primibrach axillary. Arms fixed in cup up to about 2nd or 3rd secundibrach. Single large interprimibrach extending proximally to level of radial facet in large excavation of distal corners of adjacent radials. Arms 20, uniserial. pinnulate. Stem circular in section, with crenulate intercolumnal sutures. REMARKS. This genus is assigned to the Hexa- crinitidae on the basic cup pattern of plates proximal to the radial facets. However, distal to the radial facets the arms are fixed in the cup which extends distally 71/2 length of the radials even though depressed between the arms. I suggest that this genus represents a further development from Oehlerticrinus in which the radials extend distal to the radial facets and there is a large interprimibrach resting on the straight upper margin of the radials: extension of the CRINOIDS FROM CENTRAL VICTORIA 35 FIG. 24. Frankocrinusenidae gen. et sp. nov. splayed holotype crown NMVP108571 from NMVPL 252. A, distal view of crown showing inner sides of arms, x2. В, basal view with A ray at 11 o'clock, х4. C, lateral view with cup inverted; D ray crushed under cup in forground, x4. D, basal view with A ray at 12 o'clock, x2. interprimibrach proximally between the upper corners of the radials, radials smaller relative to size of cup than in other hexacrinitids and arms fixed in cup at least to the 2nd secundibrach are distinctive features of this genus. A comparable genus is Cerasmocrinus Strimple & Levorsen. 1973 from the Upper Devonian of Iowa in which the cup is typically hexacrinitid up to the radial facets and the arms are fixed in the cup and separated by a few large interprimibrachs up to at least the 1st secundibrach; that genus is separated from Frankocrinus by its flat wide primibrachs and 3 or more plates rather than one separating the fixed arms. Frankocrinus holmesi sp. nov. (Fig. 23) MATERIAL. HOLOTYPE: NMVP109099. PARATYPE: NMVP109109 from NMVPL229. DIAGNOSIS. Cup of smooth plates. with broad low angulation of the cup in position of ray ridges. with marked flexure around the cup at level of radial facet. Arms 20, fixed in cup up to 2nd secundibrach, uniserial, pinnulate, of 36 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 25. Frankocrinusenidae gen. et sp. nov. part and counterpart of paratype cup and stem in posterior view, with disarticulated arms above the cup NMVP108622 from NMVPL232, x3.5. cuneate brachials. Fixed arms separated by de- pressed areas each occupied by a single large interprimibrach, with a number of smaller teg- menal plates above that. Stem circular in section. heteromorphic. with crenulate intercolumnal sutures. DESCRIPTION. Crown up to 45mm long, conical to subcylindrical. Cup high conical, consisting of basals, radials and primanal: ray ridges low. broad, indistinct. Basals 3. equal. forming hexagonal circlet, symmetrically beneath A. C and D radials, occupying about 1/3 of cup height, each with vertical median ridge. Radials 5. only slightly longer than wide. occupying about 2/3 of cup length, with broad low ridges continuing from basals; A, C and D radials with single central ridge: B and E radials and primanal each with 2 ridges converging on base of radial facet: radial facet a little more than 1/3 radial width. almost horizontal, subsemicircular, proximal to the distal margin of radial. Primanal same size as radials, smooth, supporting 3 large plates. Arms 20 or more, uniserial, more than 4 times as long as cup. Second primibrach axillary. pentagonal: secundibrachs 6-10 per arm, cuneate, almost circular in section. pinnulate; tertibrachs cuneate. pinnulate, with large well- developed facet for attachment of pinnule on longer side; pinnules one per brachial, alternating from side to side along each arm. long, well spaced. Tegmen of small polygonal plates. Stem circular in section. with strongly crenulate sutures between columnals, heteromorphic: proximal section of low columnals with epifacet slightly wider on nodals, noditaxis N1. REMARKS. This species is distinguished from Е enidae by its smooth plates. longer radials and greater number of secundibrachs per arm. Frankocrinus enidae sp. nov. (Figs 24. 25) MATERIAL. HOLOTYPE: NMVP108571. PARATYPE: NMVP108622 from NMVPL252. DIAGNOSIS. Crown with external finely punctate ornament. Radials smaller relative to cup size than in other hexacrinitids. Arms 20). fixed in cup up to 2nd secundibrach, uniserial, pinnulate, of cuneate brachials. with strongly crenulate interbrachial sutures. Fixed arms sep- arated by depressed areas occupied by single large interprimibrach. Stem circular in section. with crenulate attachment suture. CRINOIDS FROM CENTRAL VICTORIA 37 VIG. 26. Alisocrinus lineatus sp. nov. incomplete crowns from NMVPL300. A, C, part and counterpart of paratype NMVP109791, х3. В, С ray view of holotype NMVP110632, х3. DESCRIPTION. Crown estimated to be 35mm long. conical. Cup high conical. consisting of basals, radials and primanal, with ray ridges low broad indistinct. Basals 3. equal. hexagonal, symmetrically proximal to A, C and D radials, each with vertical median ridge. Radials 5, only slightly longer than wide, with broad low ridges continuing from basals; radial facet about 2/3 radial width. declivate, subsemicircular. Pri- manal same size as radials, smooth, without facet. Arms 20, uniserial, with deep groove on inner side; 2nd primibrach and secundibrach axillary, pentagonal: tertibrachs cuneate. almost circular in section, pinnulate, with large well developed facet for attachment of pinnule: pinnules one perbrachial, alternating from side to side along each arm, long, well spaced. Tegmen of small irregular poly gonal plates. Stem circular in section, with strongly crenulate sutures between columnals, noditaxis N1 with lateral flange at midlength of latus slightly wider on nodals, with columnals increasing gradually in length distally, with lateral midlength flange occupying only small part of length, with large circular central lumen. REMARKS. This species is distinguished from Е holmesi under that species above. The basal circlet crushed up into the calyx of the holotype makes determination of proportions impossible and has broken and concealed the primanal (at 5 o'clock Fig. 24D) masking features of the posterior of the cup. Ornament is a little more linear on the paratype but nevertheless it is still punctate as in the holotype. Crown length or shape is not available due to presevational features. Suborder GLYPTOCRININA Moore, 1952 Superfamily MELOCRINITOIDEA d'Orbigny, 1852 Family MELOCRINITIDAE d'Orbigny, 1852 The evolutionary lineage outlined by Kirk (1929), Ubaghs (1958) and Brower(1976) begin- ning in the Ordovician Glyptocrinidae, through Alisocrinus, Ctenocrinus, Melocrinites and to 38 MEMOIRS OF THE QUEENSLAND MUSEUM Trichotocrinus in the Middle Devonian is accepted, as are the generic concepts of those authors. However, Kesling's (1964) comment that the concepts of several type species are so poorly understood that he considered Creno- crinus among other genera to be synonymous with Melocrinites is equally applicable to the myriad of species assigned to these genera. More than 40 specific names are currently included in Ctenocrinus and doubtless this is a figure inflated by intraspecific variation and preservation and failure on the part of several authors to recognise growth series. It is not the aim of this paper to assess these essentially European and North American taxa; I have illustrated many spec- imens of this very common element of the Victorian fauna to facilitate future species level revisions of the family. Assignment to estab- lished species from the Northern Hemisphere is a deliberate effort to avoid introducing new names into an already overcrowded arena. These are the closest known species available but future work on the status of known species could change the concept of any or all ofthe 3 and could necessitate reassignment of the Australian material. My assignments are made by comparisons with published illustrations, not all of them photo- graphic, which may require reassessment. Alisocrinus Kirk, 1929 TYPE SPECIES. Mariacrinus warreni Ringueberg, 1888 from the Middle Silurian of New York; by original designation. DIAGNOSIS. See Ubaghs (1978). Alisocrinus lineatus sp. nov. (Fig. 26) ETYMOLOGY. Latin lineatus, linear, for the ornament. MATERIAL. HOLOTYPE: NMVP110632. PARATYPE: NMVP109791 from NMVPL300. DIAGNOSIS. Cup plate ornament of narrow low ray ridges and fine radial ridges on a background of fine tubercles. Arms fixed in cup up to Ist tertibrach, 4 per ray, uniserial, of cuneate brachials; 2nd primibrach and 3rd secundibrach axillary. DESCRIPTION. Crown up to 35mm long, subcylindrical. Cup high, conical, up to 15mm long; cup plates large, thin, with ornament of radial ridges (including ray ridges) and fine raised incomplete and irregular comarginal growth bands (so incomplete as to be simply a fine tuberculate ornament on most plates). Basals 4, with 3 of them pentagonal, with one larger and hexagonal, with proximal margin as distinct rim to stem attachment, with fine continuous ray ridges diverging distally from midpoint of base to cross 2 distal sides at right angles, with extra central vertical ridge bisecting CD interray on posterior basal. Radials 5, heptagonal, largest plates in cup, with inverted Y-shaped ray ridges continuing from basals and onto Ist primibrach, with secondary radiating ridges horizontally onto adjacent radials and sloping onto Ist interprimibrachs, all ridges at right angles to sutures. Primanal heptagonal, resting on shoulders of 2 adjacent radials, supporting 3 plates above, with 7 fine radial ridges radiating from centre to middle of each side and beyond to contiguous plates. Anal interray wider than others, with large polygonal plates decreasing in size upwards (arrangement unclear on only avail- able specimen). First primibrach hexagonal, with broad low median ray ridge running in vertical line, with fine ridges running diagonally to other 4 lateral sutured margins; 2nd primibrach axillary, heptagonal, with median ray ridge in Y-shape to enter 2 arms. Secundibrachs fixed in cup, hexagonal, as large as primibrachs, 3rd axillary. Arms 20, uniserial, free above 2nd div- ision, tapering gently, with longitudinal grooves on outer side of arms; free brachials cuneate, with pinnules one per brachial alternating from side to side along each arm. Interprimibrachs numerous, not depressed, with single large plate in contact with 2 radials and | st primibrachs and supporting next row of 2 interprimibrachs, with plates in subsequent rows becoming smaller and less regularly arranged; intersecundibrachs tiny by comparison, few in number, depressed, poly- gonal, irregularly arranged. Tegmen unknown. Stem circular, heteromorphic, with wider epi- facet on nodals but all columnals of about same height. REMARKS. This species is distinguished from the genotype by the finer ray ridges and 3 rather than 2 secundibrachs. It is separated from the other species of the genus by its generally finer plate ornament and 3 rather than 2 secundibrachs. FIG, 27. Ctenocrinus paucidaetylus (Hall, 1859), all crowns in lateral view from NMVPL229 except C from NMVPL232. A, NMVP149377, x3.5. B, NMVP109158, x2.5. C, NMVP108583, x3. D, NMVP110646, x2. E, NMVP108961, x3. F, NMVP109154, x1.5. б, NMVP149378, x2. 39 CRINOIDS FROM CENTRAL VICTORIA 40 MEMOIRS OF THE QUEENSLAND MUSEUM S b FIG, 28. Crenocrinus paucidactylus (Hall, 1859), all incomplete crowns in lateral view from NMVPL229, A. NMVP149379. x3. B. NMVP108968. x4. C, NMVP149380, «2. р, NMVP109165, *2. E, ММУРІ00165, :2.5. Е, NMVP109117, «2. G, NMVP14938]. x2. CRINOIDS FROM CENTRAL VICTORIA 41 FIG, 29. Ctenocrinus paucidactvlus (Hall, 1859), all incomplete crowns in lateral view from NMVPL229 except C from NMVPL232. A, NMVPI09111. «2.5. B, ММУРІ00168, x2.5. C, NMVP108596, «2.5, D, NMVP108610, «2.5. E, NMVP108956, *2.5. F, NMVP149382, «2.5. 42 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 30. Ctenocrinus paucidactylus (Hall, 1859), both from Locality 25, N of Heathcote (Talent, 1965). A, large set of arms with partial upper cup NMVP149356. x1. B, large partial cup and arms NMVP149383, x1.5. Ctenocrinus Bronn, 1840 TYPE SPECIES. Crenocrinus typus Bronn, 1840 from the Lower Devonian of Germany. REMARKS. In light of the discussion under the family heading above | have not attempted to diagnose these species erected last century but have attempted to illustrate their features in the descriptions provided below. The description of the first species appliesto all 3 in most features so descriptions of succeeding species are provided only where they differ from that of C. pauci- dactylus. The species to which the Australian material is assigned belong to what Brower (1976) termed primitive Crenocrinus. Ctenocrinus paucidactylus (Hall, 1859) (Figs 27-30, 31 A-E) MATERIAL. NMVP100165, 100168, 108685, 108956, 108961, 108964, 108968, 109111, 109117, 109127, 109154, 109158, 109165. 109168, 110646. 149377-149382 from NMVPL229; NMVP108583, 108596, 108610, 110636 from NMVPL252; NMVP149356 and 149383 (тот locality 25 in the Mt Ida Formation of Talent (1965, fig. 1). DESCRIPTION. Crown up to 170mm long (NMVP148625), averaging 60-80mm long at NMVPL229 and 252. Cup high conical: surface of plates with strong radial ornament of narrow strongly convex ridges of uniform width throughout except for ray ridges being wider and more prominent than the rest. Basals 4, with interplate sutures at A, C. D and E rays, at least 4mm long. Radials 5, large, in contact laterally. 9mm long and 8mm wide at the widest point 3mm from the top, with 7 sides including horizontal distal margin and broadly chevron shaped proximal margin. First primibrach hexagonal, with horizontal proximal and distal margins, 7mm long by 6mm at greatest width near midlength. Second primibrach axillary, variously hexagonal or heptagonal, with strong dividing median ray ridge and markedly thinner and weaker radiating ridges laterally: 2nd secundibrach axillary, giving rise to larger inner arm trunk and smaller outer ramule. Arms 10, up to 4 times as long as cup. fixed in cup to about 3rd or 4th tertibrach; 2 arm trunks per ray laterally sutured into pseudobiserial structure, essentially a single arm, sharing a single groove down the inner side. with every 6-8th tertibrach axillary, 5-sided, with upper obtuse angle about midwidth of arm, branching off long uniserial pinnulate ramules at each axillary. Interprimibrachs CRINOIDS FROM CENTRAL VICTORIA 43 FIG. 31. A-E, Ctenocrinus paucidactylus (Hall, 1859), juvenile crowns all from NMVPL229 except C from NMVPL252. A, NMVP109127,x3.5. B. NMVP 108964, x4.5. С, NMVP110636, x3.5. D.E, exterior and inner side of arms, respectively NMVP108685, х5. F, Ctenocrinus signatus Follmann, 1887, crown NMVP109152 from NMVPL229, x2. numerous, with large proximal one resting on shoulders of 2 radials. with striking ornament of high narrow radial ridges of uniform width (not expanded at centre of plate); 2nd row of 2 plates except in CD interray with 3 in 2nd row; subsequent rows less regular. of smaller plates. merging into the tegmen just above the fixed 2nd secundibrach. Intersecundibrachs 2, small, pentagonal, enclosed by large central arms in each ray. with central tubercle and weak radial ridges. Intertertibrachs between larger inner arm and first outer arm very small. irregular. up to 8, with central prominence on each and lateral ridges running between the inner and outer arms. Stem circular in section, heteromorphic, higher and wider nodals alternating with internodals. Ctenocrinus stellifer Follman, 1887 (Figs 32-34) MATERIAL. Syntypes of Follmann (1887, pl. 2, fig. 2, 2a,b) in the Bonn University Museum. Australian material assigned NMVP100175, 100188, 108584, 108589, 108597, 108603, 108658, 108684. 149384-149386 all from NMVPL252; NMVP149346 from NMVPL 1990; NMVP149357 from NMVPL 1841. 44 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 32. Clenoerinus stellifer Follmann, 1887. all crowns in lateral view from NMVDPL252 except A from NMVPL1990. А. NMVP149346, x1. B, partial cup NMVPLOOLS88, «1.5. C, NMVP149384, x3. D. NMVP108389, «1.5. E, ММУРІ08584. «1.25, un CRINOIDS FROM CENTRAL VICTORIA 4 eB FIG. 33. Crenocrinus stellifer Follmann, 1887. A-C, cups from NMVPL252. А, NMVP108597, x2.5. B. NMVP 108603, +2. C, NMVP100175, х2. D. interorand exterior ofan arm NMVP149357 from NMVPL 1841. х ].5. 46 MEMOIRS OF THE QUEENSLAND MUSEUM VIG. 34. Ctenocrinus stellifer Follmann, 1887, all crowns in lateral view (A interior view) from NMVPL252 A. NMVP149385, x4. B. NMVP149386. x3. C, NMVP108684. «2. D. NMVP108658, «3. CRINOIDS FROM CENTRAL VICTORIA 47 VIG 35. Crenocrinus signatus Schmidt, 1941, all crowns in lateral view from NMVPL252 except E. from NMVPL229. A, NMVP108609, »2.75. В, NMVPI0S8608, «2.5. C. NMVP 108601, «2.5. D, NMVP109167. 22.5, К, NMVP109104, <3, Е, NMVP149387, х3. 4% MEMOIRS OF THE QUEENSLAND MUSEUM VIG, 36. Ctenocrinus sp. all from NMVP1924. A,B. part and counterpart of tegmen with some arm fragments NMVP109756, х3. C. crown NMVP109750. x2, D, crown NMVP109222. x4. DESCRIPTION. Crown up to 80mm long. Other- wise as for C. paucidactvlus except: 1, plate ornament including the ray ridges. is greatly reduced on proximal parts of the cup: 2. tertiary ridges occur especially in the proximal plates of the cup parallel to the radial ornament; 3. Ist intersecundibrach with radial ridges running onto adjacent plates and producing distinctive 5-rayed star not evident in other Australian species; 4. ramules with axillary brachials slightly longer on outer side but otherwise no modification to arm trunk, REMARKS. Reduced ornament on the proximal cup is clear on Schmidt's (1941, pl. 8, fig.4b) figure. Schmidt (1941. fig. 17) illustrated the tertiary. ridges parallel to the radial ridges on proximal plates and the distinctive radial ornament of the Ist intersecundibrach. Follmann (1887. fig. 2, 2a.b) showed the last mentioned fcature (fig. 2a) and the laterally longer axillary CRINOIDS FROM CENTRAL VICTORIA 49 FIG. 37. Hapalocrinus? victoriae Bather, 1897, part and counterpart of splayed holotype crown NMVP386. from Yarra Improvement works near Prince's Bridge, х2. brachials in the arm trunks; however, he showed a ramule arising about every 3rd or 4th brachial whereas the Australian species has upto 9 nonaxillary brachials in a row. averaging 6-8. Assignment to stellifer is made with slight reservation but it is a better match than with any other known species. Ctenocrinus signatus Schmidt. 1941 (Figs 31F. 35) MATERIAL. HOLOTYPE: in the Bonn Museum (Schmidt, 1941:73, fig 18a). Australian material assigned NMVP108601, 108608, 108609, 109167, 149387 from NMVPL252 and NMVP109104, 109152 from NMVPL229. DESCRIPTION. Crown up to 70mm long. Otherwise as for C. paucidactylus except in relation to plate ornament as discussed below and in the branching of ramules where the axillaries inthe arm trunks are quite asy mmetrical, the facet for the ramule being much smaller and more lateral than that for the continuing central arm. REMARKS. This species assignment is made principally on plate ornamentation which consists of a raised flat-topped central tubercle that is expanded laterally to occupy about half the plate area and from which the radial ridges radiate only a short distance so producing a stellate ornament. This ornament is unlike that of C. paucidactylus where the radial ridges maintain a uniform width even across the centre of the plate and even though it is subdued in some specimens (Fig. 35D) it is still much more pro- nounced than in C. rhenanus and it lacks tertiary ridges also. Ctenocrinus sp. (Fig. 36) MATERIAL. NMVP109222, 109750, 109756 from NMVPL1924. DESCRIPTION. As for C. paucidactylus except l, plate ornament is virtually absent, although one specimen (Fig. 36D) retains a very subdued radial ornament on basals, radials and primi- brachs. 2. the tegmen consists of a very large number of tiny polygonal plates, with a small diameter anal tube (broken base only on this specimen). 3, largest specimen with a fringe of tiny plates between bases of ramules adjacent to arm trunks. 4. brachials forming ramules with proximal and distal margins parallel in exterior view but cuneate in interior view. 50 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 38. Clematocrinusperforatus sp.nov., all crowns in lateral view from NMVPIL 300. A, NMVP109816, «4,5, В, NMVP149388. x4, C. NMVP109824, «2. D, NMVP109821, «4. Е, NMVP109782. «5, F, NMVP109785. x5. G: NMVP109811. х5. CRINOIDS FROM CENTRAL VICTORIA REMARKS. This material with reduced plate ornament could be placed with C. stellifer from NMVPL252 but features listed above, in par- ticular features of the mature arm and lack of ornament make such identification doubtful and it is best left in open nomenclature. Superfamily PLATYCRINITOIDEA Austin & Austin, 1842 Family HAPALOCRINIDAE Jackel, 1895 Hapalocrinus Jackel, 1895 TYPE SPECIES. Hapalocrinus elegans Jaekel, 1895 from the Lower Devonian of Germany. Hapalocrinus(?) victoriae Bather, 1897 (Fig. 37) MATERIAL. NMVP386a and b trom the Yarra Improvement Works near Prince’s Bridge, Melbourne, in the Ludlovian Melbourne Formation. DESCRIPTION. Crown splayed: form of cup uncertain, basal circlet not discernible. Radials 5, 7-sided (one being radial facet). smooth, widest at distal shoulders. Arms 10, up to 15mm long. with 3 primibrachs. 3rd axillary. uniserial, with well-developed facets for pinnule attachment giving zigzag appearance to arm: pinnules 1 per brachial alternating along each arm, widely separated, long (up to 5mm) and thin, with 4-6 long pinnulars: brachials rectangular, about twice as highas wide, with wide strongly zigzag groove on inner surface. REMARKS. The detail of this specimen is not certain as was acknowledged by Bather (1897: 337. footnote 3) when he said he could not vouch for each line in his drawing. However, I agree with his general conclusions and only the nature of the basal circlet described by Bather is not readily apparent to me. Since so few features are available a separate diagnosis is not provided; the species is principally distinguished by its long brachials in the arms and is deliberately isolated onthe type because of the difficulty of comparing other material with it. Ausich (1986b) discussed this species in relation to Ubaghs’ (1978b) tentative extension of the range of Clematocrinus to include Aus- tralia. І agree with Ausich's (1986b) conclusion supporting separation of the 2 genera and retain victoriae in Hapalocrinus based principally on the 3 primibrachs. Bather (1897) noted that the counterpart of the type specimen was lost but part and counterpart „л — FIG. 39. Clematocrinus perforatus sp. nov., crown NMVP100185 from NMVPL232, x3. have been reunited in the Museum of Victoria collection. Clematocrinus Jaekel, 1898 TYPE SPECIES. Aectinocrinites? retiarius Phillips in Murchison, 1839 from the Wenlock of England: by monotypy. DIAGNOSIS. See Ausich, 1986b: 894. REMARKS. Ubaghs (1978b:512) tentatively ex- tended the distribution of this genus to include North America and Australia but as noted above there are no published records in Australia and /7. victoriae cannot be considered a member of this genus. Ubaghs (1978b:518) suggested that Chapman's (1903) Helicocrinus, although of un- certain placement, might be a hapalocrinid but that genus is best isolated on the type specimen with features as noted below. Clematocrinus perforatus sp. nov. (Figs 38-40) ETYMOLOGY. Latin per-, through and foramen, hole; referring to the stem appearing perforate. MEMOIRS OF THE QUEENSLAND MUSEUM m r; perforatus sp. nov., all crowns from NMVPL300, A. NMVP10982 3. “4, B, 5. x 5. Г. NMVP109784, х4 F. NMVP10977 к 3, D. NMVP109777, х *5, С.ММУР109809. lematacrinus ,X3. 40 NMVP109819 0. C G. NMVP109826, x4. FIG. 4 CRINOIDS FROM CENTRAL VICTORIA 53 MATERIAL. HOLOTYPE: NMVP109782. PARA- TYPES: NMVP109775, 109777. 109782, 109784, 109785, 109805, 109809. 109811, 109816, 109819, 109821. 109824, 109825, 109826, 110624. 110627, 110628, 110629, 110631. 110633, 110642, 149388 all from ММУРІ.300; NMVP100185 from NMVPL232. DIAGNOSIS. Cup bowl-shaped. Basal circlet apparently fused. Posterior interray with vertical anal tube of large polygonal plates. Arms 10. uniserial, of low subquadrate (in lateral view) cuneate brachials. Stem noncirriferous, with crenulate intercolumnal sutures, with columnals becoming longerand epifacet breaking up into an annulus of tubercles distally. DESCRIPTION. Crown elliptical, 32mm long. Cup low bowl-shaped. about 5mm high, radially symmetrical except in posterior interray, without basal concavity. Cup plates smooth. thin, convex. Basals apparently fused into pentagonal unit, short in lateral view, with circular crenulate stem facet occupying central 1/2-2/3. Radials 5, forming complete circlet, largest plates in cup. occupying about 1/2 cup length, hexagonal, con- vex with distal corners depressed. Radial facet horizontal, just over half radial width. Arms 10. free distal to primaxil, uniserial; Ist primibrach subquadrate. with convex raised central part and depressed lateral flanges: 2nd primibrach pentagonal, axillary, fixed in cup. with depressed lateral flanges: brachials variable in proportions but usually wider than long, subquadrate prox- imally, rapidly becoming cuneate distally; pinnules | per brachial, alternating along each arm, long. becoming more slender distally. of long pinnulars (4-6 per pinnule). Proximal interprimibrach large. hexagonal. resting on depressed shoulders of adjacent radials, only interbrachial in cup; CD interray with row of 3 anal plates resting on shoulders of C and D radials. with interbrachials protruberant in elongate convex bulge: primanal central, pentagonal, smaller than proximal interbrachial of other interrays, flanked by smaller anal sac plates in contact with primibrachs. Tegmen with short stout anal tube at posterior. Stem circular in section, heteromorphic: proximally with alternating nodals and internodals of similar length but with epifacets or annular flange at midlength of latus being wider on the nodals: distally with uniform columnals, each with the epifacet at midlength of latus broken up into a circlet of small nodes: intercolumnal artic- ulations symplexial. with culmina from one columnal not fitting neatly into opposing crenellae on next columnal so producing the effect of two adjacent circlets of pores around stem. REMARKS. This species is distinguished within the genus by its distinctive stem, narrow radial facets. uniserial arms and posterior anal tube. Australian species do not closely resemble any of the other species and might ultimately prove separate generically but at present І cannot determine a clear diagnosis of such a genus and these species fit the current concept of Clemato- crinus. Moreover, phylogeny within the family as in Ubaghs (1978b) is unclear and unnecessary introduction of further names seems unhelpful. Clematocrinus argylensis sp. nov. (Fig. 41) ETYMOLOGY. From just east of the Argyle Railway Station near Heathcote. MATERIAL. HOLOTYPE: NMVP109188. PARA- TYPES: 109189, 109193, 109201 from ММУРІ 2259. DIAGNOSIS. Cup bowl-shaped. Posterior inter- ray with vertical anal tube. Radials about half cup length. with extremely narrow radial facets less than half plate width. Arms 10, uniserial, of sub- rectangular (in lateral view) to cuneate brachials. Stem noncirriferous. with crenulate inter- columnal sutures, with columnals becoming longer and epifacet not breaking up into tubercles distally. DESCRIPTION. Crown elliptical. 25mm long. Cup bowl-shaped. about 5mm long, radially symmetrical except for posterior interray, with- out basal concavity. Cup plates smooth, thin. Basals 3, fully exposed laterally. with circular crenulate stem attachment occupying central 1/3 of diameter. Radials 5. largest plates in cup. about 1/2 cup length, hexagonal: radial facet horizontal, about 1/3 radial width. Arms 10, free distal to primaxil, uniserial; Ist primibrach subquadrate, with convex raised central part and depressed lateral flanges: 2nd primibrach pentagonal. axil- lary, fixed in cup. with depressed lateral flanges: brachials variable in proportions but usually longer than wide, subpentagonal proximally. rapidly becoming cuneate distally: pinnules one per brachial. alternating along each arm, long. becoming more slender distally, of long pinnulars (4-8 per pinnule). Proximal interprimibrach large, hexagonal. resting on shoulders of adjacent radials, only interbrachial in cup: CD interray with row of 3 anal plates resting on shoulders of contiguous radials, with anals forming vertically MEMOIRS OF THE QUEENSLAND MUSEUM бте ае ча rpartof holotype х0. DF, part and 9. A. E. part and counte 225 . NMVP109201, х6. C. NMVP109193. eclively, B sargylensis sp. nov. all crowns from NM VPL and x4, resp NMVP109188. *4.5 FIG, 31. Clematocrinu counterpart of NMVP109189. «4. CRINOIDS FROM CENTRAL VICTORIA FIG. 42. Helicocrinus plumosus Chapman, 1903, holotype, NMVP384, from a quarry in West Brunswick Melbourne, х 1.7. elongate anal tube; primanal central, pentagonal, smaller than proximal interbrachial of normal interrays, flanked by smaller anal tube plates in contact with primibrachs. Stem circular in section, heteromorphic: proximally with alternating nodals and internodals of similar length but with epifacets or annular flange at midlength of latus being wider on the nodals: distally with uniform columnals. each with the epifacet at midlength of latus. REMARKS. This species is close to C. perforatus but has narrower radial facets and arms. different stem and longer unfused basal circlet. Helicocrinus Chapman, 1903 TYPE SPECIES. Helicocrinus plumosus Chapman, 1903 from the Ludlow of Melbourne; by monotypy. REMARKS. Ubaghs (1978b) suggested that this genus was ‘Possibly a hapalocrinid although placing it in the group of uncertain taxonomic un Un placement. Chapman (1903) had noted the same alliance originally and no contrary opinion has been offered. The present state of the holotype and only specimen makes definite assignment impossible but the discussion of that specimen below gives confidence to a family assignment in the Hapalocrinidae. І recommend that since critical features of the holotype are not now retrievable the generic name should be confined to the holotype and isolated on that specimen. Although it may seem likely that Clematocrinus perforatus sp. nov. should belong to the same genus there is no proof of this and it is better not to compound the problem by extending use of such a poorly defined name. Nevertheless, occurrence of 5. and probably more. species of Hapalo- crinidae in Victoria demonstrates that the family was successful in this region during the Late Silurian. Helicocrinus may be separated from known Hapalocrinidae by the distally coiled stem with pentagonal section. Helicocrinus plumosus Chapman, 1903 (Fig. 42) MATERIAL. HOLOTYPE: NMVP384 from a quarry at West Brunswick, between Albert and Victoria Streets. The matrix into which this individual was moulded is soft and friable; І have not made a latex cast because I consider it likely damage would be caused to the type. REMARKS. Chapman (1903. pl. 18. figs 1-5) provided drawings of various parts of the holotype that are not clear from his photographor from the holotype today. The holotype shows signs of having been damaged through efforts at preparation somewhere in the intervening 80 years since discovery but it may also be sig- nificant that in his description Chapman did not allude to any features of the cup plating. Even his drawing of the cup is incomplete just distal to the radials. Since the primibrachs are critical for species and genus recognition it is impossible to accommodate this specimen within current taxonomic concepts. Hapalocrinidae indet. (Fig. 43) MATERIAL. NMVP107091 from the South Yarra Brick- works dump originally from excavations in the Ludlovian Melbourne Formation at the eastern end of Melbourne City area; collected by Leo Stach of Melbourne High School. REMARKS. This 28mm long crownis preserved in a medium to coarse micaceous sandstone so that interplate sutures are not clear anywhere on the specimen. However. the cup is high conical 56 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 43. crown NMVP107091 from the South Yarra Brickworks Dump originally from excavations at the eastern end of Melbourne CBD. x3. Hapalocrinidae indet., and the general impression of radials and basals is discernible with a vertical (anal) tube beginning above the radials and extending up between the arms. There are 10 pinnulate arms; details of the stem are unclear except that there appear to be circlets of tubercles and some perforations distally. This meagre combination of features allows the specimen to be interpreted as a posterior view of a hapalocrinitid but generic determination appears impossible. posterior view of Subclass DISPARIDA Moore & Laudon, 1943 Superfamily CALCEOCRINOIDEA Meek & Worthen, 1869 Family CALCEOCRINIDAE Meek & Worthen. 1869 Darraghcrinus gen. nov. TYPE SPECIES. Darraghcrinus tomi sp. nov. ETYMOLOGY. For Tom Darragh of the Museum of Victoria. DIAGNOSIS. Cup elongate, waisted near mid- length, bilaterally symmetrical: E radial divided into small triangular supero- and inferoradials separated by long suture separating large lateral A and D radials; basal circlet semi-elliptical, of 3 plates, with two tiny plates and large triangular posterior plate, with stem attachment in contact with all 3 plates. E ray arm with or without a single division at primibrach 13; lateral arms with axillary 2nd primibrach, with subsequent isotomous divisions on both resulting branches (ramules and main axil series not evident). DISCUSSION. Moore (1962) reviewed the phylogeny of the Calceocrinidae indicating 2 Ordovician to Devonian lineages distinguished by the degree of symmetry of the crown and his phylogeny has largely been accepted by sub- sequent workers (Arendt. 1965; Prokop, 1970). Since this Australian species has perfect bilateral symmetry it must belong to Moore’s group arising from Calceocrinus. However, Moore (1962) also indicated the trend in this group to- wards development of main axils with ramules. In the Australian species main axils are not developed; A and D radials each support a lateral arm that divides isotomously twice. indicating that neither of the 2 branches from the first div- ision can be considered a ramule. These features are sufficient to isolate the Australian species at generic level and to throw into doubt its possible affinities. The cup of D. tomi is very similar to that of Kohalysiocrinus Prokop, 1970 from the Devonian of Bohemia. Prokop (1970) did not figure specimens with arms but nevertheless provided a reconstruction with a single lateral arm on the A radial (and presumably another, not seen, on the D radial) that branches on the 2nd primibrach and then each branches again on the 3rd secundibrach; he identified primaxil and FIG. 44. Darraghcrinus tomi gen. et sp. nov., all crowns from NMVPL232. A, lateral view ofNMVP108653, x4. B,D, partand counterpartofNMVP | 10637,x3. C, E ray view of NMVP1 10635, x4. E,F,partand counterpartof NMVP108654, x4. G, D ray view of NMVP110640, x7.5. Н, oblique E ray view of NMVP108625, х5. "^ CRINOIDS FROM CENTRAL VICTORIA 58 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 45. Darragherinus tomi gen. et sp. nov., sketches of plate arrangement of A, NMVP108654a (Fig. 44E); B. NMVP110635 (Fig. 44C); C, NMVP108654b (Fig. 44F); D, NMVP110637a (Fig. 44D). secundaxil arms and a ramule. While this branch- ing pattern has some similarity with D. tomi there are distinct differences: branches after divisions in D. tomi are of equal diameter, the primibrachis much longer and narrower in D. romi and the E ray arm is of similar diameter to the lateral arms. Brett (1981) indicated that Prokop's recon- struction of the arms is “completely hypothetical’ and showed an advanced arm branching pattern with main axil structure in Fo/ialysiocrinus typus from the Silurian of North America. Nevertheless. the cup of £. typus is very similar to that of D. tomi in observable features, particularly on the anterior side. Notably, however, Brett's species has a different arm branching pattern, anal tube completely enclosed by the arms and E ray brach- ials much larger than those of the lateral arms. In the context of Moore's phylogeny D. tomi cannot be easily linked to any of the known Devonian members of the family. Its arm branch- ing pattern seems akin to the most primitive members of the family or even the ancestral group, yet its cup resembles some of the most advanced Devonian taxa. | assign this genus to the Calceocrinidae with confidence but cannot suggest a reasonable ancestral lineage from among known forms. Darraghcrinus tomi sp. nov. (Figs 44-46) MATERIAL. HOLOTYPE: NMVP149347, PARA- TYPES: NMVP108625, 108651, 108653, 108654, 108656, 110634. 110635. 110637-110640, 149363 from NMVPL232. DIAGNOSIS. As for genus. DESCRIPTION. Crownup to 36mm long. Calyx subrectangularin E ray view (c. 20% longer than wide). slightly waisted near midheight: E ray side of cup flat to weakly concave, opposite (posterior) side convex. Plates with finely granulose surface. with stereom structure of plates clear in most specimens. Basal circlet not clear in any one specimen but inferred from parts of several; semi-elliptical, 3 plates, stem attachment straddling suture between two larger ones, third triangular much wider than long and occupying most of circlet. E ray (median) inferoradial and superoradial widely separated by A and D radials: separation distance more than 1/2 cup height. Small, triangular E-ray inferoradial occupying just over 1/3 basal cup width; lower margin with a rectangular ligament fossa, bounded on each side by dentate projections of inferoradial. E-ray superoradial also triangular, 3 times as broad as long, extending above the A and D radials; radial facet broad, nearly circular. Lateral (A and D) radials very large, enclosing cup laterally. extending onto posterior side: subanal plate (fused B and C radials) low, wide. only slightly recessed for anal X; anal X large. identical to more distal anal plates, bounded laterally by A and D radials and subanal. with rounded lower corners, occupying most of posterior side of cup. Anal tube projecting posteriorly from cup. curving anteriorly upwards io be enveloped by the lateral arms above midheight, of large laterally convex rectangular plates on posterior side, tapering slightly in upper half, rounded upper end, with anterior side of lower subrectangular plates. FIG. 46. Darragherinus tomi gen. et sp. nov., all crowns from NMVPL232. A, lateral view of NMVPI10639, х4.7. B.lateral view of damaged cup with stem NMVP108651, «4.7. C, small theca with stem NMVP108653b, х5.7. D.E, part and counterpart of crown attached to large Sphenothallus NMVP110638a and B, x3.3. F.G, lateral views of part and counterpart of holotype NMVP149347, «4.7. H, lateral view of NMVP110634, x5.7. 59 CRINOIDS FROM CENTRAL VICTORIA 60 Arms 3; E-ray arm branching isotomously once high above cup. with 12 primibrachs, 12th axillary: Ist primibrach subtrapezoidal. tapering distally: rest of arm more or less uniform in section; interplate sutures poorly defined. A and D radials with 2 primibrachs. 2nd axillary: on more anterior branch 3rd or 4th secundibrach axillary, with 4th or 5th tertibrach axillary above that on posterior branch and with anterior branch undivided: on posterior branch of original division 2nd secundibrach axillary on posterior branch and 3rd or 4th tertibrach on anterior division axillary, then 4th quartibrach axillary on posterior branch (Fig. 44D): each arm tapering near base but mostly of uniform diameter. Stem circular in section. of short laterally con- vex columnals. with columnals becoming slightly longer and less convex laterally away from cup: attachment simple cementation, ob- served in two cases to be cemented to large specimen of Sphenothallus. Superfamily PISOCRINOIDEA Angelin. 1878 Family PISOCRINIDAE Angelin, 1878 Trichocrinus Miiller, 1856 TYPE SPECIES. 7richocrinus altus Müller, 1856 from the Middle Devonian of Germany: by monotypy. REMARKS. Moore et al. (1978) considered Trichocrinus a junior synonym of 7riacrinus Munster, 1939 from the Late Silurian and Dev- onianof Europe. Rozhnov (1981) separated them and characterised 7richocrinus as having a longer, regularly conical, cup of smaller diameter as opposed to convex-conical shape with lobate profile in distal view, a shallow or no basal con- cavity for stem facet and a “different form of distal growth’. Rozhnov (1981, fig. 9) also provided a phy- logeny showing 7riacrinus and Trichocrinus evolving from different subgenera of Pisocrinus. I accept Rozhnov's generic concepts and phylogeny and assign this Victorian species to Trichocrinus with its tall conical cup lacking basal concavity. 7. morlevi from the Ludlow is among the oldest known species of the genus and greatly extends its geographical range. Its occur- rence in Australia may be significant in view of MEMOIRS OF THE QUEENSLAND MUSEUM the Upper Devonian, Western Australian Jaekelicrinus (Jell & Jell. 1999) which genus belongs to the same lineage as 7richocrinus. Trichocrinus morleyi sp. nov. (Figs 47. 48) ETYMOLOGY. For David Morley who drew my attent- ion to the occurrence of crinoids at the type locality. MATERIAL. HOLOTYPE: NMVP100111. PARATYPES: NMVP100103, 100105. 100110, 100113-100115, 107108-107110, 109827 all from NMVPL 1923. NMVP148608, 148611, 148612 from Moonee Ponds Creek; NMVP111845, 148624 [rom NMVPL299; NMVP391 from NMVPL1615. DIAGNOSIS Cup with 3 basals, long. with rounded distal margin to inferradial. First primi- brach short, tapering upwards. Arms 5, atomous. of long brachials, 5-10 times length of cup. Stem long, heteromorphic. of alternating columnals proximally becoming strongly beaded distally. DESCRIPTION Crown small, with very long arms up to 10 times as long as cup. Cup approx- imately twice as long as wide. smooth. Basals 3, forming circlet only marginally wider than long. Radials smooth, with A and D radials more than twice as long as wide. with E radial small tri- angular and taking part of distal corner from both Aand D radials (more from D than A): B super- radial and C radial approximately same size as E radial but apparently with concave proximal margin; B inferradial large, with rounded distal margin, rotated to right (theca oriented with stem down) 1.е. anticlockwise relative to superradial: all radials with vertical grooves on both plates at and normal to the radial to Ist primibrach suture. Anal X distal to cup, between the C and D primibrachs. Arms 5, atomous, 5-10 times as long as cup, nonpinnulate, laterally compressed. with deep adoral groove covered by 2 series of interlocking cover plates; Ist brachial subtri- angular. with broad base on radial facet. lower than more distal brachials, twice as wide proximally as distally; other brachials more than twice as long as wide, of approximately uniform length through arm, with 8 pairs of adoral groove cover plates in contact with each brachial. Stem long but full extent not known, circular in section: columnals uniform in length proximally VIG. 47. Trichocrinus morleyi sp. nov. all crowns in lateral view except for one cup (C and E) from NMVP1923. A,NMVP100103, x3.5. B.G, part and counterpart of holotype NMVP100111, х4.5.С Е, part and counterpart of NMVP100110, x4.5 and x7. D.F, NMVP100113 (to right in D) and part and counterpart of NMVP100114, x3 and x4.5, respectively. Н, NMVP100115, x3. I, NMVP107110. x4.5. J, NMVP107108, x4. К, NMVP107109, x5. 6l CRINOIDS FROM CENTRAL VICTORIA 62 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 48. Trichocrinus morleyi sp. novy., all crowns in lateral view. A,B, part and counterpart of NMVP 148608 from Moonee Ponds Creek, x3. C,E, part and counterpart of NMVP148624 from NMVPL299, x3. D, NMVP111845, х4. F, NMVP391 from NMVPL1615 (with type of Nassoviocrinus longibrachiatus), x3. G, NMVP148612 from Moonee Ponds Creek, х4. Н, NMVP148611 from Moonee Ponds Creek, х3. but alternating long and short distally. diameter decreasing uniformly and rapidly through 10 most proximal, then uniform for variable distance of between 30-200 columnals, then alternating large and small to give a beaded appearance to most of stem. REMARKS. This species is closely related to 7: elongatus Follmann, 1887 and T. altus Muller from the Lower Devonian Hunsrück Shale of Germany but differs in its shorter arms with longer more uniform brachials. its longer basals. rounder distal margin to the B inferradial, and more noticeably beaded distal stem. Several species of Pisocrinus grouped by Rhozhnov (1981)as Р (Pocillocrinus) and exemplifiedby P. (Р.) pocillum Angelin, 1878 from the Late Silurianof Gotland have similarly shaped cups as remarked by Bather (1893:23), are part of a line- age among long species from 5 basals to 3 and prompt the question of whether the number of basal plates is a watertight generic distinction. Since the same reduction occurs in other groups (e.g. the Synbathocrinidae — Phimocrinus to Synbathocrinus) it is clear that numerous lineages involved reduction in numbers of basals. Since Rozhnov (1981) identified the separate lineages leading from Pisocrinus to forms with 3 basals the question now arises whether he has identified all the lineages in the family or did the 5 to 3 basals transition occur more than twice in the Pisocrinidae? Could there have been a separate lineage in Australia evolving in parallel CRINOIDS FROM CENTRAL VICTORIA 63 to the European lineages outlined by Rozhnov (1981) or, as seems most likely at present, did the Australian pisocrinids migrate iteratively from Eurasia during the Ludlow, and the Upper Devonian? Several specimens (Fig, 48) assigned to this species from other than the type locality are pre- served in medium to coarse sandstone; in these, details of plate boundaries are not always clear so assignment is made on cup shape, on length of arms and on the anal X above the cup. Superfamily MYELODACTYLOIDEA S.A. Miller. 1883 Family MYELODACTYLIDAE Miller, 1883 Myelodactylid indet. (Fig. 49) MATERIAL. NMVP149350 from NMVPL 1924. = = = s, = @ FIG. 49. Myelodactylid indet. A, stem fragment with arm brachials evident beneath cirri at top of photo indicating the cup was against the outer whorl of stem at about 2 o'clock on specimen NMVP149350 from NMVPL1924, x3. B, Sketch showing probable position of cup enclosed in cirri (from position of brachials beneath cirri) and probable configuration of stem coiling (from available stem ). DESCRIPTION. Proximal or evolute part of stem composed of many short columnals compressed in the plane of coiling and with a shallow narrow groove along the curved visible edge of the stem. Midsectionor 2 inner involute whorls with long- est columnals of whole stem, also compressed in the plane of coiling, with stout cirri on each columnal. Distal section or outer involute whorl apparently less compressed, stongly convex on outer side with groove on side (presumably one on other side as well) of central peak. with long cirri on each columnal. REMARKS. Details of members of this family are available from a number of taxa well- preserved as original carbonate (Bather, 1893; Springer. 1926) with crowns well displayed. In the available Victorian external mould the crown is indicated by a number of arms beneath the cirri of the outer whorl; taking this indication with the direction of the proximal stem the crown is 64 MEMOIRS OF THE QUEENSLAND MUSEUM inferred to have been adjacent to the outer whorl about 3/4 of the way around. Without the crown I do not attempt generic assignment as advocated by Eckert & Brett (1985) but the strongly coiled stem with differentiated proximal part, strong development of cirri, and indicated position of the crownall confirm the family assignment. The specimen is included here as an indication of the family's occurrence in Australia and warning to future collectors to seck out this interesting but relatively rare group of crinoids. Superfamily BELEMNOCRINOIDEA S.A. Miller. 1883 This superfamily as constituted by Moore et al. (1978) is a heterogeneous grouping of widely differing. mostly very small, crinoids whose affinities are often unclear. Rozhnov (1981) re- assigned Quiniocrinus Schmidt, 1941 from the Perissocrinidae to the Pisocrinidae and Prokop & Petr (1997) suggested that Storthingocrinus is à camerate related to the Platycrinitidae: reassign- ment of other taxa grouped in this superfamily in 1978 seems likely. Family PYGMAEOCRINIDAE Strimple, 1963 This family is monotypic and has been re- viewed in detail by Prokop & Petr (1997). The new genus described here initially looks quite different from the Czech type species but com- parison with Prokop & Petr’s (1997) emended diagnosis reveals that the only difference be- tween Czech and Australian taxa is in the number of brachials in each arm. Pygmaeocrinus Bouska. 1947 must be considered a derived form in the structure of its arms which consist of just 2 brachials, a small 1st and large widened 2nd that fully enclose the cup distally and Kroppocrinus canbe considered a reasonable intermediate from a synbathocrinid like Phimacrinus hanschi sp. nov. which has 5 basals and an anal plate almost expelled from the cup. Aroppocrinus has the 5 basals but fewer brachials per arm and no anal plate. Thus I suggest that as a probable ancestor this new genus should be placed in the Py gmaeo- crinidae. Kroppocrinus gen. nov. TYPE SPECIES. Aroppocrinus heatheotensis sp. nov. ETYMOLOGY. An anagram from Prokop plus the usual ending -crinus: for Rudi Prokop, Prague who has contributed to knowledge of Bohemian echinoderms. DIAGNOSIS. Crown more than 3 times as long as cup. Arms inclose lateral contact forming a tall FIG. 50. Kroppocrinus heathcotensis gen. et sp. nov., all crowns in lateral view from locality 36 of Talent (1965, fig. 1) at Heathcote except A from NMVPL229. A, NMVP107106, x6. В, NMVP- 107093, x7. C.E. part and counterpart of holotype NMVP107094, x10. Р.Е, NMVP107092, x9. conical enclosed space. with 5 or 6 brachials; Ist small, subquadrate, set into outer notch of radial facet; 2nd very large. up to 4 times as long as wide. with widest point above upper limit of radial's interradial peak, with series of horizontal ridges laterally on inner side; more distal brachials less than half as long as 2nd, of different lengths in different arms. Stem either attached to CRINOIDS FROM CENTRAL VICTORIA 65 rimmed facet in centre of flat cup base or same diameteras cup base but tapering rapidly distally; in latter case columnals low. uniform proximally becoming longer and heteromophic distally. REMARKS. These 2 species are quite different in the size of the stem relative to the cup prox- imally and recessed basals laterally on cup on one but not other. However, cup shape and details of the arms are so similar that they are confidently assigned to the same genus herein. А. math- iesonensis has cup features linking it to Pygmaeocrinus notabilis Prokop & Petr, 1997 namely. the flat cup base with small rimmed stem attachment facet, depressed triangular distal extremities of the basals in view laterally and shape and projection of radials: however, it is the only specimen available and it has one branching 2nd brachial making it probably an aberrant specimen and therefore. unsuitable for type genus status. A. heathcotensis has less distinctive link to Pygmaeocrinus having a similar cup shape to Р fabulosus Prokop & Petr. 1997 but without unique characters. Nevertheless, it is a far more suitable species to stand as type since it is represented by several specimens and has the atomous arms considered typical of the genus. Kroppocrinus heathcotensis sp. nov. (Fig. 50) ETYMOLOGY. From near Heathcote, central Victoria. MATERIAL. HOLOTYPE: NMVP107094, PARA- TYPES: NMVP107092, 107093 all from Locality 36 of Thomas (Talent, 1965, fig. 1), paratype NMVP148560 from locality 41 of Thomas (Talent, 1965, fig. 1); paratype NMVP107106 from NMVPL229. DIAGNOSIS. Cup evenly flaring in proximal part. same diameteras stem at their junction, with concave lateral cup margin due to extended radial facets. DESCRIPTION. Crown bipyramidal, penta- lobate in section through radials or more distal. up to 12mm long. Cup 2-4mm long and wide, flaring uniformly distally in basal part but increasing in distal part of radials producing concave outer profile. Basals 5. circlet occupying about 1/4 cup length, not well defined, with low obtuse peaks at interradial sutures. Radials 5, subrectangular. longer than wide, expanding strongly laterally to facet, with sharply pointed interray peaks: radial facet deep narrow notch. slightly declined outwards. Arms 5, atomous, in lateral contact throughout forming tightly closed space within, with horizontal ridges laterally on FIG. 51. Kroppocrinus mathiesonensis gen. et sp. nov.. part and counterpart of holotype crown NMVP- 109752 from NMVPL1924. x4 and x6 respectively. inner surface of brachials interlocking with adjacent brachials; brachials 5 per arm; Ist short; 2nd very long; distal ones of variable lengths within and between arms so adjacent arms may have distal interbracial articulations at different levels. Stem same diameter as base of cup at at- tachment: proximal part tapering strongly. of very short columnals of uniform length; distally heteromorphic, slightly wider and longer nodals alternating with very short internodals. REMARKS. This species is distinguished from K. mathiesonensis by nature of cup base and nature of stem. The available specimens have some hardened limonitic infills in the moulds which result in the slightly incomplete latexes illustrated but taken in combination available specimens provide a clear species concept. Al- though not figured. features of distal parts of the stem are available on NMVP148560. Kroppocrinus mathiesonensis sp. nov. (Fig. 51) ETYMOLOGY. From Mathieson' s Creek, near Kinglake. MATERIAL. HOLOTYPE: NMVP109752 from NMVPL1924. 66 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 52. Phimocrinus americanus Springer, 1923, all crowns in lateral view. A, NMVP109206 from NMVPL1615, x5. В, NMVP100151 from NMVPL229, x3. С, NMVP109205 from NMVPL229, x4. DIAGNOSIS. Cup with flat base, much greaterin basal diameter than proximal stem, with straight distally flaring outer wall. Basals as depressed interradial triangles in lateral view extending distally from cup base. DESCRIPTION. Crown bipyramidal, penta- stellate in section through radials or more distal, up to 7mm long. Cup 2mm long and wide. flaring uniformly distally giving straight outer profile. Basals 5, occupying most of flat cup base and depressed triangles interradially in side walls of cup. Radials 5, subrectangular but with prom- inent, strongly constricted base between basals, longer than wide, expanding strongly in midline to facet, with sharply pointed interray peaks; radial facet deep narrow notch, declined out- wards. Arms 5, atomous, in lateral contact throughout, forming tightly closed space within, with horizontal ridges laterally on inner surface of brachials interlocking with adjacent brachials. Brachials 5 per arm, Ist short, 2nd very long. distal ones of variable lengths within and be- tween arms so adjacent arms may have distal interbracial articulations at different levels. Stem of much smaller diameter than cup base, of uni- form diameter except for very small expansion towards cup in most proximal part; columnals uniform. short, separated by crenulate sutures. REMARKS. The most obvious comment on this specimen has to be on the bifurcate arm (Fig. 51B). In view of the related type species K. heathcotensis having its arms tightly closed and the other 4 arms of this specimen appearing the same, this bifurcate arm is here considered aber- rant, probably the result of damage and regrowth. How the animal lived without being able to seal the space within the arms is not clear because that seems to have been a functional requirement of this group of crinoids, judging from other avail- able material. I have not, therefore. included this in features of the species even though I do not have available a specimen to prove that this is an aberrant specimen. CRINOIDS FROM CENTRAL VICTORIA 67 Family SYNBATHOCRINIDAE S.A. Miller, 1883 Phimocrinus Schultze, 1867 TYPE SPECIES. Phimocrimus laevis Schultze, 1867 from the Middle Devonian of Germany. REMARKS. This genus is separated from Syn- bathocrinus principally on its 5 rather than 3 basal plates. However, some species of the latter genus have the basals fused so this feature is not clearcut. The 2 Australian species described here have much longer brachials than in any Syn- bathocrinus where the brachials are usually wider than long or subquadrate in lateral view: in the absence of arms for the European species this feature cannot be applied universally. Phimocrinus americanus Springer. 1923 (Fig. 52) MATERIAL. HOLOTYPE: Springer, 1923, pl. 5, fies 17-19 from the Lower Devonian Linden Formation in Benton County, Tennesee; NMVPI00151. 109205 from NMVPL229: NMVP109206 from NMVPL1615. DIAGNOSIS. Cup evenly tapering, high conical, of smooth plates. Anal X notched into tops of adjoining C and D radials. Arms long, of elongate brachials. DESCRIPTION. Cup small. up to 4mm long, high conical, smooth. Basals 5, short, with obtuse upper angle at interradial sutures. Radials long, forming most of cup, almost twice as long as wide. with horizontal distal suture; articular facet full width of radial; 151 primibrach short, tapering distally, sloping inwards so that bases of 2nd primibrachs remain in contact laterally but around a circle of smaller diameter than that of radials. Arms 5, atomous, straight. (probably lat- erally in contact throughout life), with broad shallow ambulacral furrow, cuneate in section; brachials of variable length, with some twice as long as others. Anal X small, triangular, nestled between C and D radials distally; anal tube ex- tending up inside arms, full extent unknown. Stem circular in section, diameter small in re- lation to cup, with nodals just larger and more prominent than internodals: length unknown. REMARKS. The single North American spec- imen lacks arms and stem but is matched in every available feature of the cup by the Australian specimens. There is no basis to separate these specimens from different parts of the world but future discovery of a complete specimen in North FIG. 53. Phimocrinus hanschi sp. nov., holotype crown in posterior view, NMVP100648 from NMVPL300, x2.5. America could necessitate a new specific name for the Australian material. Phimocrinus hanschi sp. nov. (Fig. 53) ETYMOLOGY. For David Hansch of Melboume who contributed specimens to this study. MATERIAL. HOLOTYPE: NMVP100648 from NMVPL300. DIAGNOSIS. Cup high conical, slender. Anal X strongly notched into radials. Arms long, that of C ray longer than others, of long brachials. Stem circular in section, slender, heteromorphic. with nodals longer than internodals. DESCRIPTION. Cup small, up to 4mm long. high conical, very slender, smooth. Basals 5. short, about 1/4 of cup length. Radials long. 68 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 54. Crotalocrinites sp., all distal stem elements of fused columnals with canals leading into cirri in 5 vertical columns from locality 54 of Talent(1965. fig. 1), x1.2. A, NMVP149349, В, NMVP16789.C, NMVP149348. forming most of cup. almost twice as long as wide, with straight distal suture; articular facet full width of radial. horizontal: 151 primibrach short. tapering distally and sloping inwards so that bases of 2nd primibrachs remain in contact laterally but around a circle of smaller diameter than that of radials. Arms 5. atomous. straight, (probably laterally in contact throughout life), with that in C ray about 1/3 again longer than other 4. with broad shallow ambulacral furrow. cuneate in section; brachials of variable length. with some twice as long as others. Anal X small, triangular, notched into distal corners of C and D radials; anal tube inside arms, full extent unknown. Stem circular in section, diameter small in relation to cup. with nodals just longer than internodals; length unknown. REMARKS. In this species the basal part of the cup is very slender because the flaring of the cup occurs near the distal margins of the radials whereas in other species the cup flares near the base to be more bell-shaped. Arms are unknown forany Northern Hemisphere specimens so com- parison is impossible but the elongate C ray arm distinguishes this species from P. americanus as interpreted from the Australian material described above. Subclass CLADIDA Moore & Laudon, 1943 Order CYATHOCRINIDA Bather, 1899 Family CROTALOCRINITIDAE Bassler, 1938 Crotalocrinites Austin & Austin, 1843 TYPE SPECIES. Cyathocrinites rugosus Miller, 1821 from the Upper Silurian of England; by original designation. Crotalocrinites sp. indet. (Fig. 54) MATERIAL. NMVP16789, 149348 and 149349 from locality 54 (Talent, 1965, fig. 1) in Unit 3 ofthe Mount Ida Formation, N of Heathcote. REMARKS. These specimens are large (up to 20mmindiameterand 80mm long) disarticulated distal stem terminations. They are subpentagonal in section, have a line of perforations leading into stout cirri or rootlets along each angle and are ornamented with close spaced horizontal grooves arond the stem. This long section is almost cer- tainly formed by fusion of numerous columnals with the horizontal grooves indicating the former columnal boundaries. Distal stems of this type were described from the Silurian of the Urals by Militsina (1980. pl. 2, fig. 4: pl. 3. figs 3. 4) for Crotalocrinites and Syndetocrinus of the Crotalocrinitidae. The Russian material has a round rather than CRINOIDS FROM CENTRAL VICTORIA 69 FIG. 55. Codiacrinus secundus sp. nov., all crownsin lateral view from NMVPL252. A,B, part and counterpart of NMVP108574, x2.5. С, NMVP108576, x2. D, NMVP108573, x2. subpentagonal section and the cirri are less regularly placed in Crotalocrinites and less nodular in Syndetocrinus. Nevertheless. there are striking resemblances in size, alignment of cirri in 5 vertical columns and external ornament of close-spaced horizontal grooves: on this basis it seems reasonable to make a family assignment. A crown of Crotalocrinites pulcher Hisinger. 1840 was recorded from the same rock unit in a white porous sandstone (Jell, 1982): it was collected froma gully where it had been placed by man and may have come from the same locality as these stem sections. On this basis I assign these distal stem segments to Crotalocrinites with the high probability that they belong to C. pulcher. Family CODIACRINIDAE Bather, 1890 Codiacrinus Schultze.1867 TYPE SPECIES. Codiacrinus granulatus Schultze, 1867 from the Eifelian of Germany near Prum: by original designation. Codiacrinus secundus sp.nov. (Figs 55. 56) ETYMOLOGY. Latin secundus, second; the second species of the genus recognised in Victoria. 70 MATERIAL. HOLOTYPE: NMVP108570. PARA- TYPES: NMVP108573, 108574, 108576, 108578, 108580, 108660, 109208, 109209, 149364 all from NMVPL2352. DIAGNOSIS. Cup finely granulose, with only the faintest suggestion of median ray ridges; radial facet occupying 1/2 radial width; first branching variable — 2nd, 3rd, 4th or 5th primibrach axillary;or peaks on the midline ofthe brachials absent; Ist secundibrachs and tertibrachs sutured against each other just distal to peak of axillary; stem tapering in proximal part, of short columnals separated by highly crenulate sutures. DESCRIPTION. Crown subelliptical, 40mm long. Cup probably (always found flattened) of medium length, globose, with convex base; infrabasals 3, 2 large and | small, about half length of basals, forming pentagonal base to cup. Basals pentagonal, largest plates in cup, with extremely faint low median ray ridges forming central prominence and apparently in high cross meeting proximal and distal sutures at right angles. Rradials pentagonal, with low ray ridges continuing from basals to meet at midwidth of radial facet; radial facet less than half width of radial, almost circular (small tangential sector missing on oral side), sloping down and out only very gently (compression prevents accurate assessment but least disrupted specimen suggests virtually no slope), flat surface except for extremely weak transverse ridge almost dis- appearing medially, with fine canal emerging near outermost point on margin, with slightly raised rim around margin. Arms branching isotomously at least 3 times probably more; primibrachs usually with 4th axillary, one ray (Fig. 56F, left) with 2 primibrachs, quite short (diameter more than twice length), with same barely discernible transverse ridge as radial facet, with angular (70°-90°) projections orally beside wide shallow adoral groove; secundibrachs 5-7; tertibrachs 5; with Ist brachials in each arm above a fork sutured against each other for all or most oftheir length, with low nodes developed on the aboral midline of many secondary and distal brachials, with brachials increasing in length relative to diameter up the arms, with distal parts of arms curved back axially. Stem with very short, wide, highly crenulate columnals at base of MEMOIRS OF THE QUEENSLAND MUSEUM cup, with columnal length increasing and columnal diameter decreasing away from cup, with long columnals in distal part of stem bearing circlet of stout rootlets; distal end of stem not preserved. MORPHOGENY. One small individual (Fig. 56C) suggests that during growth the low ridges on cup plates became much less obvious, length of the basals increased relative to length of the radials from being less than in the small in- dividual to being more than in large specimens, ratio of width to length of radials increased so making the cup more globose without intro- duction of new plates, and ratio of width to length of primibrachs and stem columnals increased dramatically. REMARKS. This species is close to C. schultzei Follmann, 1887 from the Early Devonian Hunsrück Schiefer near Bundenbach, Germany, but is distinguished by its larger radial facets relative to size of radial plate, its variation in which primibrach is axillary, the suture between the Ist pair of brachials after an arm fork, by its nodes (if anything at all) rather than spines aborally on the midline of the brachials, and by the more tapering stem with shorter columnals. Codiacrinus rarus Jell in Jell & Holloway 1983 from the ?Early Devonian Humevale Formation at Winneke Reservoir may be distinguished by its narrower radials and consequently more conical cup, by its relatively higher infrabasals, and by its Ist brachials above arm forks not being sutured to each other. Czechoslovakian species C. procerus (Prokop,1973) and C. ornatus (Prokop, 1973) and the type species are all distinguished by their coarser granulose ornament among other features. The variation in number of primibrachs in different rays of the same individual is unusual (Fig. 56F) and the ray with 2 primibrachs having 7 secundibrachs so that the 2nd forking takes place at the same level as in the adjacent ray with 4 primibrachs and 5-6 secundibrachs suggests some compensatory adjustment to arm growth. This species is relatively common at NMVPL- 252 but virtually every specimen is disrupted by plate dislocations making its shape difficult to establish and preventing any useful biometrics. FIG. 56. Codiacrinus secundus sp. nov., all crowns in lateral view from NMVPL252. A, NMVP108660, x1.9. B, NMVP108576, x2.8. C, juvenile NMVP149364, x4. D, NMVP108578, x3.3. E, holotype NMVP108570, x2.8. Е, enlargement of distal articulating face of brachial NMVP108570, x4.7. 71 CRINOIDS FROM CENTRAL VICTORIA 72 MEMOIRS OF THE QUEENSLAND MUSEUM Family CUPULOCRINIDAE Moore & Laudon, 1943 Cupulocrinus d'Orbigny, 1850 TYPE SPECIES. Sevphocrinus heterocostalis Hall, 1847 from the Upper Ordovician of Canada; by monotypy. REMARKS. Springer (1911) clarified the distinction between Dendrocrinus and Cupulocrinus based principally on the radial facet occupying all of the width of the radial in the latter but angustary in the former, Ramsbottom (1961) indicated a division within Cupulocrinus by noting that his Ashgillian species from Scotland are more similar to the type species than to North American species referred to the genus by Springer (1911). However, he did not indicate what features he used to make this division and no subsequent authors appear to have taken up this remark. Brower (1992) maintained the content of the genus with species as assigned except for C. sepulchrum Ramsbottom, 1961 (moved to Dendrocrinus) and C. conjugans for which he erected Praecupulocrinus. Despite this well expressed generic concept the 2 new Australian species which comply with the generic diagnosis are so different, that assessment of Ramsbottom's (1961) inference for further subdivision is necessary. The other problem with the Australian species is their geographic and stratigraphic separation. In the absence of any Silurian forms І am very dubious about assignment to this previously Ordovician genus. However, C. austrogracilis is so similar to C. drummuckensis Kolata, 1975 (=C. gracilis of Ramsbottom, 1961) that it is difficult to diagnose them effectively. Given the isolation of this Australian species I am very loathe to credit a genus with such an unusual range and distribution, but there is no morphological basis for excluding it from Cupulocrinus. The second Australian cupulocrinid is separated as Stewbrecrinus gen. nov. below. FIG. 57. Cupulocrinus austrogracilis sp. nov., holotype crown NMVP109194 from NMVPL1841, x6. A, A ray view. B.C. posterior views. D, sketch of plate arrangement in posterior view. Cupulocrinus austrogracilis sp. nov. (Fig. 57) ETYMOLOGY. Latin australis, southem and gracilis slender, also referring to it being southem version of C. gracilis. MATERIAL. HOLOTYPE: NMVP109194 from NMVPL 1841. DIAGNOSIS, Crown 15mm long, subcylindrical. Cup low conical, moderately flared distally, with straight to slightly concave sides. Infrabasals short (1/4 or less of cup length), tapering prox- imally. Arms long and slender. Stem very slender, of uniform, relatively long, barrel-shaped columnals. CRINOIDS FROM CENTRAL VICTORIA 73 DESCRIPTION. Crown small, 14mm long. Cup 1.4mm long, conical, maximum width distally on radials 2mm. Thecal plates smooth. Infrabasals pentagonal. very short. upflared (not well defined on the specimen). Basals 5, hexagonal except for heptagonal CD basal. about as long as wide. Radials 5, pentagonal, as long as wide, largest plates in cup; radial facet plenary. Radianal pentagonal, in inferradial position proximal to C radial; anal X resting directly on CD basal. reaching almost to top of D radial and about halfway up C radial. supporting single large hexagonal anal leading into vertical column of similarly shaped plates decreasing in size disially. Arms isotomous, narrow, dividing 3 times on 4th primibrach, 3rd to 5th secundibrach and on various tertibrachs, 3rd divisions in same arm at same height, uniserial, nonpinnulate, with rectangular brachials throughout. Stem circular in section, of uniform barrel-shaped columnals. REMARKS. This species is very similar to C. gracilis from the Middle Ordovician of North America (Kolata, 1975) particularly in structure of the posterior interray and even to relative size shape and plating of the anal tube. but differing only in the stem, height to width ratio of brachials and ratio of stem diameter to maximum cup diameter. Stewbrecrinus gen. nov. TYPE SPECIES. Stewbrecrinus terryi sp. nov. ETYMOLOGY. An anagram from Webster plus the usual suffix for crinoids, for Gary Webster who has contributed greatly to Australian crinoid studies. DIAGNOSIS. Cup dicyclic. usually expanding distally. with base of small to medium diameter. Radials usually widerthan long. with radial facet plenary. Radianal in inferradial position proximal to C radial. Anal X very large, sup- porting a single large anal plate, Interprimibrachs small. irregular. numerous. Arms 5. uniserial. nonpinnulate, dividing isotomously at least 3 times, with primibrachs of more or less uniform width, with distinct patelloid processes at least in primibrachs. Stem heteromorphic, with nodals circular in section, with internodals decagonal. REMARKS. Although this species satisfies the criteria for Cupulocrinus and is very closely VIG. 58, Stewbrecrinus terryi gen. et sp. nov., crowns from NMVPL1924. A,B, part and counterpart of juvenile NMVP109766, x1. С.Ю, part and counterpart of holotype NMVP109215, x2.2. 74 MEMOIRS OF THE QUEENSLAND MUSEUM allied to a few species ofthat genus its geographic and stratigraphic separation prompts me to allo- cate generic status. If in the future a lineage is found to link these species at least 2 species now assigned to Cupulocrinus could move to Stewbrecrinus. The new Australian Devonian genus may be distinguished from Cupulocrinus and Praecupulocrinus by its stem structure, by its curved rather than straight horizontal suture between radial and 1st primibrach and by the juvenile radial facet being much narrower than the radial plate. Brower (1992) showed in the species he erected that the radial facet occupies the full width of the radial even in the juvenile stage. However, growth series are not available for most species. The small interbrachial plates in the holotype of S. terryi may be a feature that indicates a generic grouping and 1f species with interbrachials are found in the Silurian the content of Stewbrecrinus may include Ordov- ician C. humilis, and C. jewetti (Springer, 1911). Stewbrecrinus terryi sp. nov. (Fig. 58) ETYMOLOGY. For Terry Brady who helped collect crinoids at Kinglake West. MATERIAL. HOLOTYPE: NMVP109215. PARATYPE: NMVP109766 from NMVPL1924. DIAGNOSIS. As for genus. DESCRIPTION. Crown subcylindrical, 30mm long, with arms more than twice as long as cup. Cup high conical, with basal diameter more than half summit diameter, dicyclic; plates smooth. Infrabasals 5, pentagonal, slightly longer than wide, with very high obtuse angle between 2 upper sides. Basals 5, hexagonal, as wide as long; CD basal larger than others, with 7 sides, sup- porting anal X symmetrically distal to it; anal X large, extending above radials, supporting one large anal plate medially over most of width of anal X and many tiny plates lateral to it, Кайта] 5, pentagonal except C radial subquadrate; radial facets angustary in small individual becoming plenary in large specimen, declivate, with evenly downcurved suture to Ist primibrach. Radianal pentagonal, in inferradial position proximal to C radial and separated from it by straight horizontal suture. Arms 5, branching isotomously at least 3 times, uniserial, nonpinnulate, with patelloid processes; primibrach 4 axillary except in D ray where 5th primibrach axillary; secundibrach 4 axillary; tertibrach 5 or 6 axillary. Stem heter- omorphic, tapering slightly distally in most proximal part, circular in overall section, noditaxis N212, with internodals having serrated latus, tapering slightly over proximal Іст. REMARKS. This species is distinguished by its stem, its curved suture between radial and Ist primibrach and by its interbrachial plates. The holotype is slightly crushed so that inter- radial areas between A, B and C rays are obliterated and anal X has ridden out over C radial. The interbrachial plates are only clear in the EA interradius; some are evident deep within the DE interradius. The edge ofthe anal X against the D radial is projected straight out ofthe surface of the latex just above the D radial and between it and the 1st primibrach of the D ray are a number of tiny plates suggesting strongly that a median column of large plates is laterally attended by fields of tiny plates; such plating of the anal interarea would align perfectly with that of Cup- ulocrinus humilis and C. jewetti as illustrated by Springer (1911). The numerous pits randomly distributed over the holotype cup and proximal primibrachs are interpreted as borings effected by another unknown organism. Order DENDROCRINIDA Wachsmuth & Springer, 1886 Family DENDROCRINIDAE Wachsmuth & Springer, 1886 Dendrocrinus Hall, 1852 TYPE SPECIES. D. /ongidactylus Hall, 1852 from the Wenlock of New York; by original designation. Dendrocrinus arrugius sp. nov. (Figs 59-63) ETYMOLOGY. Latin arrugia, a mine; from the vicinity of Comet Creek Mine. MATERIAL. HOLOTYPE: NMVP109774. PARA- TYPES: NMVP109119, 109781, 109786, 109788- 109790, 109802, 109813, 109823, 110630, 149365 all from NMVPL300 (Ludlow). OTHER MATERIAL: NMVP109187 from SW side of Bald Hills, NE of Kilmore; NMVP112133, 112137, 112138, 112140 and 112141 from NMVPLI925 (Ludlow). DOUBTFUL MATERIAL: NMVP108619, 149358 from NMVPL252 (Lochkovian). FIG. 59. Dendrocrinus arrugius sp. nov., all crowns in lateral view from NMVPL300. A, anterior view of NMVP109813, x3.5. B, NMVP109823, x4. С, NMVP109790, x4. D, posterior view of NMVP149365, x1.5. E, posterior view of NMVP109781, x3. F, C ray view of NMVP110630, x3. Ww. CRINOIDS FROM CENTRAL VICTORIA 76 MEMOIRS OF THE QUEENSLAND MUSEUM VIG. 60. Dendrocrinus arrugius sp. nov. A, crown in anterior lateral view of NMVP109788 [rom NMYVPL 300, x2.5. B, posterior view of NMVP109779 from NMVPL300, «3. C, large specimen from SW side of Bald Hills. E of Kilmore NMVP109187, «2.5. D. cup NMVP109789 [rom NMVPL300. «5. PIG. 61. A, C. D, ?Dendroerinus arrugius sp. nov. A, crown in anterior view NMVP 108619 from NMVPL252. 43.C.D. D ray view of incomplete crown with successive latex pulls from same specimen showing exterior (C ) and interior (D) of anal tube of NMVP149358 from NMVPL232, x3. B. Dendrocrinus arrugius sp, nov.crown in anterior view NMVP109786 from NMVPL300, x3. CRINOIDS FROM CENTRAL VICTORIA 78 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 62. Dendrocrinus arrugius sp. nov. A.C, part and counterpart of holotype crown with small asterozoan at base of NMVP109774 from NMVPL300, x2.5. B, crownin С ray viewof NMVP109790 from NMVPL 300, х3. DIAGNOSIS. Crown subcylindrical to evenly subconical, 75mm long. with anal tube longer than arms. Cup plates thin, smooth except for ray ridges variably expressed from specimen to specimen. Radianal pentagonal, in inferoradial position beneath C radial. Radial facet occupying 3/4 or more of radial width, circular, gently declined. Anal X supporting 3 anals becoming plicate and succeeded by very tall plicate anal tube. Arms uniserial, isotomous, branching 5 or more times, nonpinnulate. Stem very narrow, circular in section, noditaxis N1, with longer nodals projecting slightly laterally. DESCRIPTION. Crown very long, almost 10 times as long as maximum cup width, with anal tube extending well beyond arms. Cup conical, moderately long and flared, length about equal to maximum width. Thecal plates thin, smooth, with depressed corners in some specimens, with ray ridges well-developed on some specimens (particularly on radials). Infrabasals 5, pentagonal, longer than wide, tapering strongly proximally, with high obtuse angle between upper 2 sides, fully visible laterally except for basal flange against top of stem. Basals 5, hex- agonal, longer than wide; posterior basal 7-sided, supporting anal X distally. Radials 5, hexagonal, with distal 2 sides angling up inside the radial facet: radial facet occupying 3/4 or more of radial width, circular, gently declined outwards, with FIG. 63. Dendrocrinus arrugius sp. nov. all crowns from NMVPL1925. A, NMVP112140, х1.8. B, NMVP112137, x1.8. C, NMVP112133, x1.8. D, NMVP149359, x1.8. E, NMVPI12138, x1.8. F, NMVP112141, x0.9. 79 CRINOIDS FROM CENTRAL VICTORIA 80 MEMOIRS OF THE QUEENSLAND MUSEUM weak transverse ridge. Anal plates in cup 2: anal X large, 6-sided, as long as wide, in radial circlet, onsame level as C radial but slightly longer than D radial: radianal. pentagonal, in inferradial position proximal to C radial from which it is separated by a horizontal suture. Anal sac longer than arms. straight, of 10 vertical columns of plicate plates. Arms narrow, circular to deep U-shaped in section. with deep V-sectioned furrow on inside covered by small cuneate coverplates, isotomous, dividing 5 or more times, with primibrach 5 axillary (6 in A ray). secundibrach 8 axillary. Stem circular in section, heteromorphic, with nodals longer and of slightly greater diameter than internodals. REMARKS. The only record of Dendrocrinus from Australia (D. saundersi Jell in Jell & Holloway, 1983) is possibly the youngest record of this essentially Ordovician genus globally, coming from near the Silurian/Devonian bound- ary. Ludlow species are apparently unknown but the type species is from the Wenlock of New York. This new species from the Ludlow helps to link saundersi back to the bulk of the genus. As with my discussion of D. saundersi many of the Northern Hemisphere species remain difficult to compare because they have not been revised this century. Nevertheless, it appears the plicate ornament on the very long anal tube beginning on the Ist anal distal to anal X, broader radial facets, radianal in inferradial position and low ray ridge ornament separate this from known species of the genus including the type which also has quite large lower anal plates filling the entire interray. D. saundersi has a smooth slender anal tube. fewer branches in the arms and more obviously beaded stem. Material assigned to this species from NMVPL1925, in Dry Creek adjacent to the rail- way line underthe bridge on Saunders Road, E of Kilmore, has a markedly different appearance to that from NMVPL300 to the SE. The NMVPL- 1925 matrix has a fine crinkly cleavage and as the locality is close to a fold axis (Vandenberg, 1992) within the Kilmore East Synclinorium I suggest that a lot of the sharp angle bends in some cup plates are due to this cleavage. Depending on their orientation to the tectonic forces some specimens have been shortened others have been elongated and narrowed. However, the very long plicate anal tube (Fig. 63A,E) and the radianal symmetrically below C radial (Fig. 63D) along with the nature of the stem and numerous arm FIG. 64. Dendrocrinus sp. B ray view of cup NMVP100491 from NMVPL 1927, x3. branchings are sufficient to be confident of the identification. Two specimens from NMVPL252 in the Loch- kovian are very tentatively referred to this specieseventhoughonly the anterior is available. The more complete specimen (Fig. 62A) matches extremely closely. typical specimens from NMVPL300 (Fig. 62B). In the Devonian specimen the stem appears wider relative to the cup and what appears to be the anteriorof the anal tube running from between the A and B radials to about 11 o'clock beneath the arms lacks the plicate ornament. However, the second specimen from the same locality clearly shows the plicate ornament. There are no other known species from NMVPL252 to which these specimens could be assigned unless they represents a new taxon. Therefore we make very doubtful assignment to D. arrugius fully aware that this would infer an extremly longranging species. Ludlow to Lochkovian. A number of other species do have this range in the Melbourne Trough sedimentary structure. Dendrocrinus sp. (Fig. 64) MATERIAL. NMVP100491 from NMVPL1927 at the crossing of Broadhurst Creek by the Kilmore to Wandong Road. DESCRIPTION. One side of cup (A-C rays only) preserved: cup low, conical, of smooth thin plates with broad low median ray ridges. Infrabasals 5, pentagonal. more or less equidimensional, with proximal margin projecting out medially. Basals CRINOIDS FROM CENTRAL VICTORIA BI 5, hexagonal, longer than wide; A-B basal with straight horizontal suture dividing it into 2 pentagonal plates. Radials 5, pentagonal except B radial with an extra side against radianal and C radia! with one less side because of horizontal base against radianal, with low indistinct radial ridges emanating from centre just below radial facet angustary, only gently declivate. Radianal only anal plate available, pentagonal, in inferradial position proximal to C radial. separated by horizontal suture, with low. ridge running proximally from © radial and dividing near midlength and continuing onto basals. Arms 5. deep U-shaped in section, with deep V-shaped groove on inner side, apparently isotornous, with 3rd primibrach axillary. Stem unknown. REMARKS. Thi$.specinien 15 tao incomplete to provide a full species concept but it has some features that make it unique. It is the only crinoid from this horizon. The radianal! in inferradial position proximal to the C radial identifies it as Dendrocrinus. The divided AB basal is unknown in amy other cladid: it could be ап aberrant specimen, which l consider most likely orit could be the last remnant of a 4th circlet of cup plates that may have been in its ancestry. The latter possibility suggests derivation from 4ethacrinus as suggested by Melntosh (1983), Family PLICODENDROCRINIDAE nov. DIAGNOSIS. Cup low to hrgh conical; thecal plates with narrow, radial ridges; intrabasals 5; radial facets poorly developed, transverse ridge usually absent but when present bisected by prominent V-shaped ambulacral groove; 2 or 3 anal plates in cup: rarely with inferradianal, radi- anal either pentagonal and in inferradial position or quadrangular: arms usually isotomous, rarely heterotomous; outer faces of proximal brachials mostly angular: anal sac straight, inflated, of thin plicate plates, GENERA ASSIGNED. Plicadenidracrinus Brower. 1995 IUpper Ordovician (Ashgill); central LISA). Compagierinus Jobson & Paul, 1979 (Lower Ordovician LArenig); northem Greenland). Holnesaerinus gen. nav. (Silunan (Ludlow); central Victoria), REMARKS. The concept of this family was first documented in an unpublished thesis by George McIntosh (1983) and when he refereed this paper he insisted that | name the family from this date upon which I insisted that I acknowledge the origin of the concept. McIntosh (1983) separated D. casei from Dendrocrinus as type of a new genus and used thal genus as the type of a subfamily of the Deudrocrinidae including Compagicrinus and a new genus based on Botryocrinus reimanni Goldring, 1934 from the Middle Devonian of New York. Thus when Brower (1995) established Plicadendroorinus with D. casei as type species he endorsed Melntosh’s generic concept but with a dilferent name. Tt was the group of species now asigtied to Plicodendrocrinus that Jobson & Paul (1979) had in mind when they showed Camipagicrinus giving rise ta Dendrocrinus. Their figure would now be modified to show the Plicodendrocrini- dac continuing on beyond the Middle Silurian and a parallel arrow for the Dendroennidae which has very similar stratigraphic range. MelIntosh^s subfamily concept remains jn tact and now takes its name from Brower’s genus. In this paper 1 employ Moelntosh's subfanily concept at family level because with the description below of Ludlow and Lochkov members it is clear that the lineage continued from the Lower Ordovician through the muddle Palaeozoic independent of Dendrocrinus and although it may contain the ancestor of the Dendrocrinidae that is à recognisably distinct lineage that should be treated as such. The diagnosis used here is Melntosh's with minor modification to aecept the Australian genus Plicodendrocrinus Brower. 1995 TYPE SPECIES, Porerjacrinites (Dendracrinus) easet Meek, 187! trom the Upper Ordovician of USA: by original designation. DIAGNOSIS. Two anal plates in cup; inferradi- anal absent; superradianal pentagonal, in inferradial position proximal to C radial, Anal sac inflated, elongate, of staggered columns of strongly plicate plates proximally, with nuimerous (20-30) randomly arranged sets of sac plates distally. Arms isotomous, non-pinnulate; proximal brachials angular or cuneiform, with distinct cornice-like faring in distal direction- Stem pentastellate or decagonal to round, lacking pentameres, INCLUDED SPECIES. The type, Dendrucrinus grandimhus Ramsbollom, 1961, D, rugocyaihlius Ramisbottom, 1961. Dendhoctinns proboseidiarns Billings, 1857 and P australis sp. nov. Plicodendrocrinus australis sp. nov. (Figs 65B-E, 66B) ETYMOLOGY. Latin australis. southem. MATERIAL. HOLOTYPE: NMVP1001I91. PARATYPE: NMVP112129 from NMYPL252. 82 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 65. A, Shintocrinus richi sp. nov., posterior view of crushed crown of holotype NMVP149390 from NMVPL1990, x2.5. B-E, Plicodendrocrinus australis sp. nov. B,C, part and counterpart of crown (B stem and infrabasal circlet only) NMVP112129, x2. D.E, D ray and posterior views of holotype crown NMVP100191, x3. DIAGNOSIS. Thecal plates with low, narrow radial ornament and fine subsidiary interradial ridges parallel to the radial ridges: radial facet narrow relative to radial width: 4th primibrach axillary; arms isotomous, dividing at least 3 times. Stem decagonal, round proximally, with distinct nodals and internodals becoming less differentiated distally. DESCRIPTION. Crown subcylindrical, 15mm long. Cup low, conical, wider than long. Thecal plates, thin, prominently ornamented with vertical (on radials and posterior interray ), horizontal (on radials and basals) and diagonal (on all plates) ridges continuous from plate to plate; major ridges at right angles to interplate sutures, radiating from just below facet on radials, from middle of basals, radianal and anal X and from midpoint of proximal margin against stem on infrabasals; secondary ridges between main ridges running to corners of plates. Infra- basals 5, pentagonal, as wide as long, smallest plates in cup, upflared, visible in lateral view except for bases resting on stem. Each infrabasal with 3 ridges originating from centre of base; 2 run diagonally onto adjacent basals, 3rd vertical to upper corner not extending further. Basals 5, hexagonal, largest plates in cup. longer than wide, with central elevated point as origin of 6 major ridges continuous onto contiguous plates CRINOIDS FROM CENTRAL VICTORIA 83 FIG. 66. Sketches of holotypes to show posterior plating (C radial heavily outlined). A, Shintocrinus richisp.nov., B, Plicodendrocrinus australis sp. nov. at right angles to interplate sutures, with second- ary ridges running to plate corners not always originating at centre; posterior basal 7-sided and with 7 major ridges, with 3 distal riges running to D radial, anal X and radianal. Radials 5, pent- agonal, width about equal to length, with 6 major ridges radiating from centre just proximal to facet, with 2 horizontally to adjacent radials, 2 diagonally to basals and 2 diagonally to distal corners of radial distal to facets. Radial facets angustary, U-shaped, just distal to midlength of radial. Two anal plates in cup: radianal pent- agonal, in inferradial position proximal to C radial, with 5 main ridges radiating from centre to contiguous plates; anal X large, symmetrically distal to CD basal in radial circlet, with 7 main ridges radiating from the centre, with asymmetrical ridge diagonally to radianal. Anal sac long. composed of columns of plicate plates, with strong vertical ridge on posterior midline. Arms isotomous, non-pinnulate, narrow and deep giving high U-shaped section with angular outer margin, bifurcating on 4th primibrach, 4th secundibrach and 7th tertibrach. Stem decagonal (most noticeable in internodals) in section, consisting of alternating nodals and internodals, markedly heteromorphic in proximal part of stem but almost uniform distally. REMARKS. This species is most simply dis- tinguished from the other 5 species, all from the Ordovician of the Northern Hemisphere, by its decagonal to rounded stem but also by its finer. more elaborate thecal plate ornament. Although the holotype cup is crushed the central peak of the radianal and the C radial are in a vertical line to confirm this dendrocinid feature. Although the upper extension of the anal sac is not available it does not seem likely, from the arms closing around the sac in the holotype, that it is so promi- nently expanded as it is in P. granditubus. P. rugocyathus is distinguished by the coarse radial ornament on thecal plates. Shintocrinus gen. nov. TYPE SPECIES. Shintocrinus cometensis sp. nov. ETYMOLOGY. An anagram from Intosh, for George McIntosh of the Rochester Museum. DIAGNOSIS. Two anal plates in cup: radianal subquadrate to pentagonal; anal sac inflated, of 8 staggered rows of plicate sac plates; arms isotomous; proximal brachials angular, with distinct cornice-like flaring: stem round. INCLUDED SPECIES. The type, S. richi sp. nov. and possibly S. costatus (Angelin, 1878) (i.e. Pycnosaccus scrobiculatus in part of Springer, 1926, pl. 11, fig. 10). REMARKS. This genus shares with P/ico- dendrocrinus its isotomous arms, its conservative radial facets, thin cup plates and straight anal tube but differs from it principally by the position of its radianal, by its arms branching only 3 times and round stem. Shintocrinus differs from Holmesocrinus in the nature of the stem, slightly different position of the radianal (more to left of C radial than inferoradial) and in type of ornament although the latter is variable within Holmesocrinus. Shintocrinus cometensis sp. nov. (Figs 67, 68) ETYMOLOGY. From the vicinity of Comet Creek Mine. MATERIAL. HOLOTYPE: NMVP109778. PARA- TYPES: NMVP109771, 109801, 109803, 109808, 109812, 109818, 149366-149368 all from NMVPL300. DIAGNOSIS. Cup low conical; thecal plates with many radiating rays (i.e. more than 1 per side) particularly in basal circlet; radials about as long as wide; radial facet very narrow, circular: MEMOIRS OF THE QUEENSLAND MUSEUM 84 CRINOIDS FROM CENTRAL VICTORIA 85 two anal plates in cup; radianal subquadrate, below and just left of C radial; anal X large with dominant radial ornament; anal sac longer than arms, with 4 strong vertical ridges joined by finer diagonal ridges, with distal portion of irregular polygonal plates; stem circular. DESCRIPTION. Crown conical, 35mm long, with arms 6 times as long as cup. Cup low con- ical, slightly wider than long. Thecal plates thin, ornamented with broad low radial ridges continuing across sutures between basal and radial circlets and anal plates. Infrabasals 5, pentagonal, about as wide as long, smallest plates of cup, upflared, completely visible in lateral view except for proximal flange resting on stem, smooth except for broad low ridge on distal sutural margins connecting to similar ridges on adjacent basals. Basals 5, hexagonal, about as wide as long, about same size as radials, with low wide main ridges forming vertically elongate cross and continuing to radials and infrabasals, with horizontal sec- ondary ridges (2 or 4) continuing horizontally onto adjacent basals; posterior basal heptagonal, supporting anal X distally, with strong ridge from centre continuing onto anal X and distally along anal sac, with strong upper ridge running onto radianal rather than C radial. Radials 5, pent- agonal, as wide as long. with 4 prominent ridges radiating from centre just proximal to radial facet and continuous onto adjacent basals, radials, anal X or radianal; radial facet angustary, U-shaped, with distal 1/2 of radial extending distally around facet. Anal plates in cup 2: anal X pentagonal, in radial circlet, with 5 strong radial ridges to posterior basal, anal sac, adjacent radials and radianal; radianal subquadrate, proximal and left of C radial, with broad low crossed ridges continuing to contiguous plates. Anal sac longer than arms, consisting of columns of plicate anals proximall y becoming less regular and polygonal distally, with strong low wide ridge along posterior side on proximal portion. Arms isotomous, narrow, deep U-shaped sectional shape, with angular outer edge, dividing 4 times at 4th primibrach, 4th secundibrach, 7th tertibrach and at a similar distance again (no specimen well enough preserved to be able to count distal brachials). Stem circular in section, of uniform diameter over available length, irregularly dimorphic with some sections of stem having 3 or 4 identical columnals adjacent to each other but most of stem with nodals of larger diameter alternating with smaller internodals in noditaxis pattern N212, with subtle expression of decagonal section in some internodals in some specimens. REMARKS. This species is distinguished from S. richi by its cup plate ornament with secondary ridges, probably by the shape of the radianal and by the slightly different stem structure. The Swedish S. costatus has some secondary ridges on the thecal plates but the major ridges are still narrow and discrete by comparison with the Australian species. Shintocrinus richi sp. nov. (Figs 65A, 66A) ETYMOLOGY. For Tom Rich, my former colleague at the Museum of Victoria. MATERIAL. HOLOTYPE: NMVP149390 from NMVPL1990. DIAGNOSIS. Radianal large, pentagonal, proximal and left of C radial. Thecal plate orna- ment of fine radial ridges continuing distally on anal tube. DESCRIPTION. Crown conical, estimated 50mm long. Cup low conical. Thecal plates thin, ornamented with narrow wire-like radial ridges continuing across sutures between all cup and anal tube plates. Infrabasals and basals collapsed, indistinct except for part of BC basal showing ornament. Radials 5, pentagonal, as wide as long, with 4 prominent ridges radiating from centre just prox- imal to radial facet and continuous onto adjacent basals, radials, anal X or radianal; radial facet angustary, U-shaped, with distal 1/2 of radial extending distally around facet. Anal plates in cup 2: anal X 7-sided, in radial circlet, large with 7 radial ridges to posterior basal, anal sac, adjacent radials and radianal, with vertical ones stronger than others; radianal proximal and left of C radial, with 5 fine radial ridges suggesting pentagonal shape (outline not clear). Anal sac long, of columns of plicate anals, with strong ridge along posterior side proximally. Arms isotomous, narrow, deep U-shaped sectional shape, with angular outer edge, dividing at least 3 FIG. 67. Shintocrinus cometensis sp. nov., all crowns from NMVPL300. A, NMYP149366, х3, B, NMVP109818, x3. C,F, D ray view of NMVP109801, x3 and x6. D, C ray view of NMVP149367, x3. E, NMVP109803, x3. G, NMVP109812, x2.75. MEMOIRS OF THE QUEENSLAND MUSEUM 86 CRINOIDS FROM CENTRAL VICTORIA 87 (and probably 4) times. first division at 4th primi- brach. Stem circular in section, of uniform diameter over available length, heteromorphic, columnals long, of uniform diameter, with latus near midlength of differing widths. REMARKS. This species is based on a single crushed and poorly preserved specimen preserved in friable silistone so its assignment could be considered tentative. It is distinguished from other species of the genus by its sharp fine radial ridge ornament and probably pentagonal radianal. Its crown resembles that of Plicodendrocrinus australis sp. nov. but that species has its radianal directly beneath the C radial and a distinctive stem although the proximal part of its stem resembles that of S. richi. Doubtless these species of Plicodendro- crinus and Shintocrinus are closely related but knowledge of the Australian species is poor so I refrain from any speculation on relationships. Holmesocrinus gen. nov. TYPE SPECIES. Holmesocrinus enidae sp. nov. ETYMOLOGY. For Frank and Enid Holmes of Melbourne for their unstinting assistance in the field and donation of specimens. The specific epithet is for Enid who collected the holotype. DIAGNOSIS. Cup long, slender, conical. Thecal plates with prominent radial ridge ornament or smooth. Infrabasals 5, pentagonal, with ridges crossing close to proximal margin and running into different lobes of stem. Basals 5, hexagonal except for heptagonal CD basal. with radial ridge ornament. Radials 5. with horseshoe-shaped facets more than 1/2 radial width: C radial with only 5 radiating ridges, smaller and further distally in cup than others. Arms uniserial, narrowly horseshoe-shaped in section, branching isotomously 4 times: primibrachs, secundibrachs and tertibrachs 4 or 5 axillary. Radianal pentagonal, proximal and slightly left of C radial; anal X heptagonal, in radial circlet. with 5 radiating ridges, supporting single anal plate above: anal tube as long as arms, of plicate plates. Stem pentalobate, heteromorphic. with large circular nodals separated by 3 pentagonal internodals the middle one being slightly wider than the other 2, noditaxis N212. REMARKS. The prominent radial ridge ornament is distinctive of earliest cladids. Aethocrinus Ubaghs. 1969 (type species A. moorei Ubaghs.1969 from the Lower Ordovician of southern France) is distinguished by the extra circlet of plates in the cup, pentameric stem and wider radial facet. Holmesocrinus is assigned to the Plicodendrocrinidae because of the width and horseshoe-shape of radial facets, long anal tube of plicate plates, uniserial non-pinnulate arms, 2 circlets of plates proximal to the radials and radianal situated proximal and only slightly left of the C radial. It differs from other members of the family principally in its distinctive stem structure of pentagonal internodals between greatly expanded circular nodals. It is further separated from P/icodendrocrinus by its radianal being proximal and left of the C radial and from Compagicrinus by having | radianal. It most closely resembles Shintocrinus but that genus has anal X in the radial circlet with its distal margin at the same level as the distal margin of the radials and a more elaborately ornamented and longer anal tube. Shintocrinus and Holmesocrinus almost certainly evolved from a common ancestor among Ordovician or Llandovery plico- dendrocrinids but represent separate lineages clearly defined by the different stem structures, Holmesocrinus enidae sp.nov. (Figs 69, 70) MATERIAL. HOLOTYPE: NMVP100112. PARA- TYPES: NMVP100102, 100104, 100107, 100108. 107107 all from NMVPL1923. DIAGNOSIS. Cup strongly ornamented with prominent narrow radial ridges on all plates. Radianal with 5 radiating ridges. DESCRIPTION. Crown subcylindrical. 45mm long. with arms more than 8 times as long as cup. Cup low conical. Infrabasals 5, each with midline running into a longitudinal groove on stem, with rounded ridges emanating from near bottom corners then crossing near proximal edge and ex- tending across distal sutures at right angles, without transverse circular ridge. Basals 5, hex- agonal. each with 3 ridges normal to margins and crossing each otherin middle of plate, with ridges continuous with others on adjacent circlets and third with annular ridge on other basals; CD basal heptagonal, with extra ridge beginning at FIG. 68. Shintocrinus cometensis sp. nov., all crowns in lateral view from NMVPL300. A, posterior view of NMVP109771, x2.75. B.E. posterior view of holotype ММУР 109778, x6 and x3, respectively. C, posterior view of NMVP109808, х4. D, NMVP149368, х3. MEMOIRS OF THE QUEENSLAND MUSEUM 88 CRINOIDS FROM CENTRAL VICTORIA 89 midpoint and running distally across anal X onto anal tube, otherwise ridges same as on basals. Radials in contact except in posterior interray where anal X intervenes in circlet, hexagonal, with horseshoe-shaped radial facet almost horizontal to slightly downsloping and occupying approximately 1/2 width of plate, with same 3 ridges as on basals but crossing just proximal to radial facet, with separate ridge beginning at point of crossing of other 3 and running distally along arm, with 2 diagonal ridges finishing at distal margin of radial; C radial smaller than others, further distally in cup than others, with B radial proximal and right of it so that 1 ridge does not continue and only 5 plus the arm meet in the crossing of the ridges, in contact with radianal and anal X radianal pentagonal, with some variation in ridge pattern across it but with ridges crossing each suture at right angles to continue 2 diagonal ridges but introducing a section of transverse ridge that is between the basal and radial circular ridges; anal X 7-sided, supporting very long anal tube of strongly plicate plates. Arms narrowly horseshoe-shaped in section, with primibrach 4 or 5 axillary with apparent consistency in each individual, with brachials becoming shorter distally, with Ist sec- undibrachs and Ist tertibrachs in contact for a short distance (« height of the columnal) distal to the midline of the axillary. Stem pentagonal to pentalobate in section, becoming circular distally, heteromorphic; nodals large, circular, twice diameter of rest of stem, almost twice length of other columnals; noditaxis N212; full length of stem not known. REMARKS. There is some variation in cup shape with those that are more narrowly conical also having almost knife-edged outer faces to arms. This suggests that there has been some degree of tectonic compression of those speci- mens but such distortion is not uniform in the echinoderms or the co-occurring brachiopods. FIG. 70. Holmesocrinus enidae gen. et sp. nov., sketches of plate arrangements with radial ridge ornament shown on cup plates and radials black except for the radial ridges. A,B, NMVP100112 (Fig. 69C) and NMVP100102 (Fig. 691), respectively, in posterior view showing pentagonal radianal. C, D, NMVP100107 (Fig. 69E) and NMVP100108 (Fig. 69H), respectively, in anterior view. There has been a certain amount of dislocation of cup plates in some specimens and the sutures are not clear on several specimens as they are deep in the recesses between ridges. Nevertheless sutures are readily determined by projecting the suture from where it crosses the ridge. There is some intraspecific variation in which brachials are axillary and also in the structure of the stem (order of columnals) but in general it is a uniform species. It is distinguished from all other plico- dendrocrinids by the nature of its stem except for H. idaensis which has an unornamented cup. Holmeseocrinus idaensis sp. nov. (Fig. 71) ETYMOLOGY. For Mount Ida, north of Heathcote, in the vicinity where the species occurs. MATERIAL. HOLOTYPE: NMVP59256 (counterpart = NMVP148625). PARATYPE: 148623 from locality 52 of Thomas (see Talent, 1965, fig. 1), Parish of Redcastle, N of Heathcote, in clean orthoquartzite. FIG. 69. Holmesocrinus enidae gen. et sp. nov., all crowns from NMVPL 1923. A,F, C,D, part and counterpart of holotype NMVP100112, x2, x4, x2 and x4, respectively. B. NMVP100104, x5. E, NMVP100107, x2.5. G, NMVP107107, х3. Н, NMVP100108, x5. I, NMVPI00102, x3.5. 90 MEMOIRS OF THE QUEENSLAND MUSEUM VIG. 71. Holmesocrinus idaensis gen. et sp. nov. A.B.C.E, holotype NMVP59256. A. part of crown, x3.4. В, proximal stem and cup, x2.1. C. crown. 43.4. E. counterpart with long stem section, x3.4. D. C ray view of NMVP148623. x2.5. CRINOIDS FROM CENTRAL VICTORIA 91 DIAGNOSIS. Crown conical, 45mm long. Cup smooth, without radiating ridges. Stem with nodals barely wider than internodals proximally, becoming greatly enlarged distally with rounded latus. DESCRIPTION. Cup low conical, as long as wide. Thecal plates thick, smooth, slightly depressed at corners. Infrabasals 5, pentagonal, as long as wide, upflared, fully visible laterally except for lower flange resting on stem, with central basal projection in line with angle of stem. Basals 5, about as long as wide; posterior basal 7-sided, supporting the anal X distally across a horizontal suture. Radials 5, pentagonal, as long as wide, largest plates in cup; radial facet, pene- plenary to plenary, round, subhorizontal. Two anal plates in cup: radianal quadrate to rect- angular, proximal and left of C radial; anal X in radial circlet, same shape as radials. Anal sac poorly preserved, with central posterior column of smooth plates, may be short slim and straight, mostly unknown. Arms isotomous, narrow, deep U-shaped cross section, dividing 3 times on 4th-6th brachial each time; brachials rectangular, nonpinnulate. Stem pentagonal in section proximally, heteromorphic, noditaxis N1, with nodals up to and sometimes more than twice as long as and slightly wider than internodals; distally internodals becoming exceptionally expanded, with rounded latus. REMARKS. Generic assignment of this taxon depends heavily on stem structure which matches that of H. enidae very closely in having pent- agonal internodals and greatly expanded circular nodals with priminternodals becoming circular distally. The difference in cup ornament could be considered a generic feature but this disparity is observed between congeneric species in other dendrocrinoid genera and so Holmesocrinus is considered best identified by the stem structure. As in H. enidae the radianal is proximal and left of the C radial, the radial facets are peneplenary and the arms branch isotomously. Family THALAMOCRINIDAE Miller & Gurley, 1895 The family was broadly interpreted by McIntosh (1979) to include 13 genera but he (1983) later redefined the family more closely with 7 genera including 3 genera not included in 1979. In 1988 McIntosh & Brett recognised the priority of Thalamocrinidae and reorganised the family content once more. We acknowledge that classification of primitive cladids is far from settled at both familial and generic levels and thus retain the broader concept of the family. Antihomocrinus Schmidt, 1934 TYPE SPECIES. Homocrinus tenuis Bather, 1893 from the Wenlock of Gotland, Sweden. DIAGNOSIS. Cup high conical; thecal plates smooth, thick; 2 anal plates in cup; radianal quad- rangular; anal sac straight, long, nonplicate; radial facets peneplenary; arms branching isotomously on first 2 divisions, third divisions may be at different heights in same ray; stem circular in section, heteromorphic proximally. REMARKS. The type species has a round stem whereas other species assigned by Schmidt (1934, 1941) have pentagonal. stems and may not belong to the same lineage. There is little to choose between Bactrocrinites and Antihomocrinus and indeed Schmidt's (1934) original diagnosis mentioned only that the infrabasals and basals of Antihomocrinus were more normally proportioned i.e. not elongate as in Bactrocrinites. The Australian species is assigned here on the basis that its cup plates are not nearly as elongate as in Bactrocrinites and its stem is round in section as opposed to pentagonal in Bactrocrinites (Schultze, 1867, pl. 5, fig. 1g). McIntosh (1983) noted the only significant distinction between the type species of Anti- homocrinus and Lasiocrinus Kirk, 1914 as the isotomous vs. heterotomous arm branching, respectively. The new Australian species has isotomous arm branching in the lower 2 divisions but the 3rd divisions, in many cases, are at dif- ferent heights in the same arm. In at least one specimen (Fig. 75A-C) the arms branch isotom- ously but the branches nearest to the central axis ofthe ray form 2 straight vertical columns and all lateral branches move asymmetrically away from the central axis. These features could be regarded as beginning to evolve from isotomous towards heterotomous arms and thus intermediate between the 2 genera. I place the new species in Antihomocrinus because the arm branching comes closer to the isotomous condition. Antihomocrinus chapmani sp. nov. (Figs 72-77) ETYMOLOGY. For Frederick Chapman, for his con- tribution to Victorian Palaeozoic faunal studies. MATERIAL. HOLOTYPE: NMVP108630. PARA- TYPES: NMVP100170, 108606, 108612, 108616, 108631, 108632, 108639, 108648, 108687, 109769, S OF THE QUEENSLAND MUSEUM MEMOIR 92 CRINOIDS FROM CENTRAL VICTORIA FIG 73. Antihomocrinus chapmani sp. nov.. all incomplete crowns [rom NM VPL 1924. А.Е. part and counterpart of NMVP109757,x3 (Е with radianal central). B.C. part and counterpart of NMVP109225. «3, D.H, part and counterpart of NMVP109217, x3. E, NMVP109227. x3. б, NMVP109751,. х3. FIG. 72. Antihomocrinus chapmani sp. nov., all crowns in lateral view from NMVPL300. A. NMVP149369 x2.7. B. NMVP109797,x2.7. C.NMVP109792, x2.7. D, C ray view ofNMVP109770.x2.7. E. NM VP110625, 7. Е. NMVP149370, x2.7. G. NMVP149371, x2.7. Н. NMVP 109799, x2.7, х2. х2, MEMOIRS OF THE QUEENSLAND MUSEUM 94 VIG. 74. Antihomocrinus chapmani sp. nov., all lateral views of incomplete crowns from NMVPL252 except F from Chirnside Park (NMVPL 1922). A, anterior view of crown NMVP109769, x2.4. B,D, part and counterpart of cup NMVP108616, x2.4. C, posterior view including part of anal tube of NMVP 149360, x2.4. E, anterior view of crown with anal tube evident NMVP149372, x2.4. F, cup NMVP107090, x2.4. G, D ray view of cup NMVP108687, x2.4. L anterior view of crown with anal tube but no arms preserved NMVP149361, x3.2. H, Dictenocrinus ibaeypus sp. nov., anterior view of crown NMVP108612, x2.5. CRINOIDS FROM CENTRAL VICTORIA 95 VIG 73. Antilomocrinuschapmani sp. nov.. all crownsin lateral view from NMVPL252, A-C. Е, holotype crown NMVP108630.A.B.C. posterior view, х5, «2 and x5, respectively. C. distal anal tube. F, anterior view, х5. D,E. D ray view of NMVP108632. x2.5 and «4, respectively. E showing crushed anal tube. 96 MEMOIRS OF THE QUEENSLAND MUSEUM + E i We “gece FIG. 76, Antihomocrinus chapmani sp. nov., all crowns from NMVPL252. A, NMVP 108632. х3. B.C, part and counterpart of. NMVP108639, х3. D. NMVP108631, х3. E, NMVP108648, x3. Е, NMVPII2131. «3. G, NMVP100170, *3. H, NMVP 108606, *3. CRINOIDS FROM CENTRAL VICTORIA 97 VIG. 77. Antihomocrinus chapmani sp. nov., all crowns or partial crowns from NMVPL229, A, NMVP108688, x3. B, NMVPI09157, x3. С.р, part and counterpart of NMVP100167, x3 and x2, respectively, E-G NMVP 100154, x5, «5 and «3, respectively. Н, NMVPI08953. x3. 98 MEMOIRS OF THE QUEENSLAND MUSEUM 112131, 149360, 149361, 149372 from NMVPL252. NMVP109217, 109225, 109227, 109751, 109757 from NMVPL1924;: NMVP107090 from Chirnside Park Estate, Lilydale. NMVP100154, 100167, 108688, 108953, 109157 from NMVPL229; NMVP109770, 109792, 109797, 109799, 110625, 149369-149271 from NMVPL300. DIAGNOSIS. Cup high conical. Infrabasals almost as long as basals. Radial facets angustary. Arms uniserial, branching isotomously up to 4 times. Anal tube long, with curved tip, of vertical columns of smooth (smaller specimens) to radial- ly ornamented polygonal plates. Stem circular in section, heteromorphic, becoming uniform dis- tally; nodals longer but only slightly wider than alternating internodals. DESCRIPTION. Crown 30mm long, sub- cylindrical to subconical with distal arms flaring slightly. Cup conical, longer than wide. Thecal plates thick, smooth. Infrabasals 5, pentagonal, from 0.5 up to twice as long as wide, upflared, fully visible laterally except for lower flange resting on stem. Basals 5, same size or larger than radials, longer than wide; posterior basal 7-sided, supporting the radianal distally and slightly to the right across an oblique suture, with distal sutural margin supporting anal X not quite horizontal. Radials 5, pentagonal, as long as wide; radial facet, peneplenary, round, subhorizontal, not declined. Anal plates in cup 3; radianal quadrate, proximal and left of C radial; anal X in radial circlet, same shape as but slightly smaller (i.e. not extending as far distally) than radials; proximal part of 3rd anal plate below distal margin of C radial. Anal sac long, of vertical columns of small smooth hexagonal plates, with plates in alternate columns bearing stellate ornament in larger in- dividualals (merely depressed corners in smaller individuals), with curved tip. Arms isotomous, almost circular in cross section, dividing 3 or 4 times on 3rd-5th primibrach, 3rd or 5th sec- undibrach and on 7th tertibrach, uniserial with rectangular brachials throughout, with deep V-shaped furrow on inner side, nonpinnulate, with Ist and 2nd divisions always at same level but with other divisions sometimes at different levels in same arm. Stem circular in section, noditaxis N1, with nodals up to and sometimes more than twice as long as and slightly wider than internodals, with fine central canal. REMARKS. I initially separated this material into 3 species based on relative sizes of infra- basals to basals, numbers of primibrachs per arm, width of the radial facet and relative sizes of the radianal. Although there is some consistency of the variation in these features between localities there are numerous exceptions and I think it more likely the collections from different localities represent variable populations (the variants in different proportions) of a long ranging (Ludlow to Lochkovian) species. A large number of illus- trations are provided to show the variation as much as possible. It differs from the German species in its round rather than pentagonal stem and from the type species by the stellate ornament on anal tube and heteromorphic stem but it is close to the type. Ancyrocrinus Hall, 1862 TYPE SPECIES. Ancyrocrinus bulbosus Hall, 1862 from the Middle Devonian of New York. Ancyrocrinus sp. (Fig. 78) MATERIAL. NMVP107105 from NMVPL229. DESCRIPTION. Cup cylindrical, as long as wide. Thecal plates smooth. Infrabasals 5, pentagonal, wider than long, upflared, fully visible laterally except for lower flange resting on stem. Basals 5, largest plates in cup, only slightly longer than wide; posterior basal 7-sided, supporting the anal X distally across a horizontal suture. Radials 5, pentagonal, as long as wide; radial facet peneplenary, round, declivate. Two anal plates in cup: radianal quadrate, proximal and left of C radial; anal X in radial circlet, similar size and shape to radials. Anal sac tall, of smooth poly- gonal plates. Arms almost circular in cross section, with deep U-shaped furrow on inner side, with 3rd primibrach axillary, uniserial with rectangular brachials throughout. Stem rounded subquadrate or subpentagonal in section, hetero- morphic, with nodals much longer and slightly wider than internodals. REMARKS. This incomplete specimen is assigned to Ancyrocrinus based on cup shape, shape position and size of radianal and shape of the stem. The most distinctive feature of the genus, the grapnelhook termination to the stem has not been found at this locality but other features are sufficient for assignment. It is left in open nomenclature because it is incomplete and a single specimen. Nassoviocrinus Jaekel, 1918 TYPE SPECIES. Heterocrinus pachydactylus Sandberger & Sandberger, 1855 from the Lower Devonian of Germany. CRINOIDS FROM CENTRAL VICTORIA FIG. 78. Ancyrocrinus sp. A, crown with incomplete arms NMVP107105 from NMVPL229, x4. В. sketch posterior plate arragement (C radial heavily outlined). DIAGNOSIS. Cup low conical; thecal plates with or without ornament: radianal quadrangular: radial facets angustary to peneplenary; anal sac long, straight. of regular columns of small plicate plates: arms isotomous, branching 3 or more times: stem pentagonal. heteromorphic. REMARKS. McIntosh (1983) reviewed this genus assigning 6 species from the Lower and Middle Devonian of Europe and North America. These Australian Ludlow species are assigned by comparison with the type species with which they share the pentagonal heteromorphic stem. quadrangular radianal, isotomous arms and general shape and proportions of the cup. Although N. /ongibrachiatus has perfectly smooth thecal plates unlike European representatives. the depressed corners of thecal 99 plates in №. corcorani are comparable with other members of the genus. Nassoviocrinus longibrachiatus (Chapman, 1903) (Fig. 79) Botryocrinus longibrachiatus Chapman, 1903:108. pi. 18, figs 6-8. MATERIAL. LECTOTYPE: (designated here) NMVP390 & 392 (part and counterpart) from NMVPL1615. Other material NMVP109174, 109176 from NMVPLI1615 and NMVP109177-109179 from the Upper Yarra. DIAGNOSIS. Cup low conical: thecal plates smooth, unornamented: radial facet peneplenary:; radianal quadrangular: anal X supporting 3 anal plates distally: anal tube long. straight. of vertical columns of plicate plates: arms branching isotomously, at least 3 times: stem pentastellate in section, heteromorphic. DESCRIPTION. Crown subcylindrical. 35mm long. Cup low conical, wider than long. Thecal plates smooth, without ridges. Infrabasals 5. pentagonal. wider than long, with lateral basal corners projecting slightly where they extend onto the projections in the stem. Basals 5. hexagonal. longer than wide, largest plates in cup: posterior basal with 7th side on distal margin, supporting anal X distally. Radials 5, pentagonal, wider than long: radial facet angustary to peneplenary. subcircular, with deep ventral groove. Anal plates in cup 2: radianal quadrangular, proximal and left of C radial, with diagonals horizontal and vertical: anal X heptagonal, wider than long, in radial circlet. supporting 3 plates at base of anal sac. Anal sac long, narrow, straight. of plates with radial ridges (2 in each direction laterally and 1 each distoproximally) arranged in vertical columns, with larger proximal central plate having ridges only near upper margin, with lateral basal plates much smaller than central one. Arms isotomous, branching at least 3 times, deep U-shaped in section (i.e. laterally compressed). with deep groove on inner side, primibrach 4 axillary except in A ray where 5th primibrach is axillary: axillary secundibrach variable 6-8. Stem pentastellate, heteromorphic. noditaxis N212. showing REMARKS. This species is assigned to Vassov- iocrinus on the basis of its quadrangular radianal. 100 MEMOIRS OF THE QUEENSLAND MUSEUM b. - yw b # lx s & FIG. 79. Nassoviocrinus longibrachiatussp, nov; all lateral views of crowns; A,B.F from Upper Yarra: С.Е, from ММУРІ.1615. A, NMVP109177. #4, B. D ray view of NMVP109179, х3. C.D. posterior view of NMVP109176.*4 and x2,5. respectively. E, posterior view of NMVP 109174, x4. F. two crowns overlying each other NMVP109178, «2.5. G, posterior view of holotype with inner side of A and B ray arms exhibiting deep groove. NMVP390, «3 CRINOIDS FROM CENTRAL VICTORIA 101 FIG. 80. Nassovioerinus corcorani sp, nov all crowns in lateral view; A.GH Irom NMVPL299: B-E, from NMVPL2259. A. NMVP109207. «3. B, NMVD59498, «7. C.D, part and counterpart of holotype NMVP109202. «4 and x5, respectively. E. NMVP149363.F. NMVP109199, «7. GI IL. NMVP1483575.*2 and «4, respectively. pentastellate stem, isotomous arm branching and peneplenary radial facets. It is distinguished from the type and other species of the genus by having unornamented thecal plates. This, along with N. corcorani, is the first record of the genus from the Silurian and the depressed corners of thecal plates in the latter species indicate that the main lineage with thecal ornament began before the Ludlow and that №. longibrachiatus was an early offshoot that evolved a smooth theca. Nassoviocrinus corcorani sp. nov. (Fig. 80) Class Crinoidea Talent. 1965:17. pl. 4. fig. 2. ETYMOLOGY. For Peter Corcoran of Sandringham, Melbourne who contributed material to this study. MATERIAL. HOLOTYPE: NMVP109202 from NMVPL2259. PARATYPES: NMVP59498, 109199, 149363 from NMVPL2259; NMVP 109207, 148575 from NMVPL299, DIAGNOSIS. Cup high conical; thecal plates with broad low ridge ornament leaving corners of plates depressed; radial facet peneplenary; radi- anal quadrangular; arms branching isotomously, at least 3 times; stem pentagonal in section, heteromorphic. DESCRIPTION. Crown 20mm long, subcylind- rical. Cup high conical, longer than wide. Thecal plates with 2 broad low ridges running diagonally distally from the basal corners of infrabasals to cross on infrabasals and basals and then meet another such ridge at the base of the radial facet. Infrabasals 5, pentagonal, longer than wide, with lateral proximal corners projecting slightly where they extend onto the projections in the stem. Basals 5, hexagonal, longer than wide, largest plates in cup. Radials 5, pentagonal, longer than wide; radial facet peneplenary, sub- circular, horizontal or almost so. Anal plates in cup 2; radianal quadrangular, proximal and left of C radial; anal X longer than wide, in radial circlet. Anal sac unknown. Arms isotomous, branching at least 3 times, deep U-shaped in section (i.e. laterally compressed), with deep groove on inner side; primibrach 5 or 6 axillary; 2nd and 3rd branchings may be at different levels in same arm. Stem pentagonal, noditaxis N212, with alternating long nodals and short internodals. REMARKS. This species is distinguished by its high conical cup although plate boundaries are poorly defined in some specimens, its broad low ridge ornament, its quadrangular radianal, MEMOIRS OF THE QUEENSLAND MUSEUM peneplenary horizontal radial facets and its pentagonal stem of very small diameter. It is a much smaller species than others in the genus and has a taller cup but its stem and posterior interray are quite compatible with Nassoviocrinus. Dictenocrinus Jaekel, 1918 TYPE SPECIES. Botryocrinus decadactylus Bather, 1891 from the Upper Silurian of England. Dictenocrinus ibaeypus sp. nov. (Fig. 81) ETYMOLOGY. Ib = infrabasal and Greek aipys, tall. MATERIAL. HOLOTYPE: NMVP108612. PARA- TYPES: NMVP108624, 108635, 108649, 108657, 108680, 108682, 149373 from NMVPL2352. DIAGNOSIS. Cup high conical; thecal plates smooth; infrabasals long, longer than wide; radial facets peneplenary, horizontal; radianal quad- rangular; anal tube tall, straight, of smooth hexagonal plates; arms 10, isotomous, pinnulate, with 3rd or 4th primibrach axillary; stem circular in section, heteromorphic. DESCRIPTION. Crown subcylindrical, 35mm long. Cup high conical, longer than wide. Thecal plates smooth. Infrabasals 5, pentagonal, upto twice as long as wide, only gently upflared, fully visible laterally except for lower flange resting on stem. Basals 5, largest plates in cup, twice as long as wide; posterior basal 7-sided, supporting the anal X distally and slightly to the right across a slightly oblique suture. Radials 5, pentagonal, as long as wide; radial facet, peneplenary, round, subhorizontal, not declined. Two anal plates in cup: radianal quadrangular, proximal and left of C radial; anal X in radial circlet, same size and shape as radials, distal edge distal to top of D radial but same level as top of C radial. Anal sac long, of alternating vertical columns of small smooth plates, tip unknown. Arms isotomous, deep, narrow, U-shaped cross section, dividing only once on 3rd or 4th primibrach, uniserial with rectangular brachials in proximal parts becoming slightly cuneate distally, with deep V-shaped furrow on inner side, with long fine pinnules one per brachial alternating from side to side up each arm, pinnules apparently absent from primibrachs including axillary. Stem circular in section, noditaxis N1, with nodals at least twice as long as and slightly wider than internodals, with fine central canal. REMARKS. Within the genus the length of the infrabasals coupled with smooth anal plates and CRINOIDS FROM CENTRAL VICTORIA 103 N. х» E d o» a 23 АК) a FIG 81. Dictenocrinus ibaeypus sp. nov.. all lateral views of crowns from NMVPL252. A. NMVP 1080657. х3. В, NMVP149373. x3. C. NMVP108624, 43. D, posterior view of holotype NMVP 108612. «3. E, NMVP108682b. *3. F, NMVP108680. х3, б, NMVP108635, x3. Н, NMVPIORO649, х3, 104 round stem separate all except D. cvathiformis (Haarmann, 1921), D. evathiformis is only distinguished by its stem having uniform columnals and expanding proximally to base of cup. by its anal X being 6-sided and by its shorter pinnules. Dictenocrinus remotus sp. nov. (Fig. 82B) ETYMOLOGY, Latin remotus, distant. the material was collected a long way from where it occurred because the malenal was excavated al Coldstream West Road. and taken down Mooroolbark Road to (ll a large depression, MATERIAL. HOLOTYPE: NMVPI49374 from NMVPL 1990. DIAGNOSIS. Low conical cup, of convex plates: radial facets peneplenary: anal tube tall, of plicate plates: radianal subquadrate. proximal and left of C radial; arms heterotomous, with one main diy- ision in each ray on axillary 3rd primibrach. with long nonpinnulate ramules on every 3rd or 4th brachial distal to the firsi division: stem pent- agonal in section, heteromorphic. DESCRIPTION. Crown subcylindrical, 35mm long. Cup low conical. as long as wide. Thecal plates convex. smooth. Infrabasals 5. pentagonal, short (slightly wider than long), fully visible lat- erally, Basals 5, large, hexagonal: posterior basal 7-sided. supporting the anal X distally. Radials 5. pentagonal, as long as wide: radial facet, penc- plenary. round, declivate at very low angle. Two anal plates in cup: radianal subquadrate. proximal and left of C radial: anal X in radial circlet. same size and shape as radials. distal margin in line with distal margin of C and D radials, Anal sac tall. of vertical columns of small plicate plates, tip unknown. Arms heterotomous, with narrow, U-shaped cross section, with one main division at 3rd primibrach, unisecial with loug rectangular brachials proximally becoming slightly cuneate distally. with long nonpinnulate ramules on every 3rd or 4th brachial distally. Stem pentagonal in section, heteromorphic, with noditaxis N323 1323, REMARKS. Although known from only one specimen this species has the single main division of ramulate arms and pentagonal stem found in the type species (Bather.1891) from which it is distinguished by the plicate anal tube and peneplenary radial facets. MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 82. A, Dierenocrinus sp. el. D. rhaeypus sp, nov... unterior lateral view of crown NM VP 107058 trom а road cutting on the road to Wonga Park. east of Melbourne, *3. B, Dietenoerinus remotus sp. nov. posterior view of crown NMVP149374 from NMVPI. 1990 (although collected from spoil heap on Mooroolbark Road), +3, Dictenocrinus sp. cf D. ibaeypus sp. nov. (Fig. 82) MATERIAL. NMVP107058 from a road cutting on the toad to Wonga Park in eastem Melbourne, DESCRIPTION. Crown subcylindrical, 40mm long. Cup high conical, longerthan wide. Thecal plates smooth, Basals largest plates in cup. twice as longas wide. Radials 5. pentagonal, as long as wide: radial facet angustary. round, subhorizontal, Arms isotomous. U-sliaped cross section, divid- ing once on 3rd or 4th primibrach, uniserial, with reciangular brachials proximally becoming cuneate distally, with long fine pinnules 1 per CRINOIDS FROM CENTRAL VICTORIA brachial alternating from side to side along each arm, pinnules apparently absent from primi- brachs including axillary. Stem not known. REMARKS. This specimen differs from D. ibae- ypus in its radial facet being more angustary than peneplenary and in its secundibrachs being cuneate. Without knowledge of the posterior inter- radius it is not possible to assign this specimen with certainty but if the posterior was identical with D. ibaeypus the variations noted above might well be considered to be intraspecific variation particularly since the 2 populations represented occurred some 40-50km apart. Subclass FLEXIBILIA Zittel, 1895 Order TAXOCRINIDA Springer, 1913 Family TAXOCRINIDAE Angelin, 1878 Meristocrinus Springer, 1906 TYPE SPECIES. Zaxocrinus loveni Wachsmuth & Springer, 1880 (=Cyathocrinus interbrachiatus Angelin, 1878) from the Upper Silurian of Gotland, Sweden; by original designation. Meristocrinus quatriramus sp. nov. (Fig. 83) ETYMOLOGY. Latin quatri-, four and ramus, branch; for the 4 branchings of each arm. MATERIAL. HOLOTYPE: NMVP109765 from NMVPL1924. DIAGNOSIS. Infrabasals evident laterally anteriorly but not posteriorly. C ray with radianal in inferoradial position. Arms branching 4 times. Stem of uniform diameter, proximally of short uniform plates becoming heteromorphic distally. DESCRIPTION. Crown inverted pear-shaped, 60mm long. Cup small, low bowl-shaped, flaring strongly. Infrabasals visible in lateral view as short circlet with low peaks at 3 way junctions with 2 adjoining basals, number not determinable, may be fused. Basals 5, hexagonal, in AB and probably CD and DE interrays, pentagonal in AE and probably BC interrays, with short vertical sides, with highly obtuse or no basal angle and upper angle of 90°; CD basal irregular, with projection extending distally between D radial and radianal, with radianal in inferoradial position beneath C ray. Radials pentagonal, with plenary facets. Arms long, isotomous, expanding above radials, branching on primibrach 3, secundibrach 4, tertibrach 5 and quaternibrach 6, without interprimibrachs but with single pentagonal intersecundibrach 105 (visible from inner side of arms), wide shallow groove on inner surface becoming deeper and occupying more of the arm width distally. Stem circular in section, widest proximally, of short, strongly crenulate columnals proximally, becoming heteromorphic distally with nodals and internodals alternating in length but uniform in diameter. REMARKS. The posterior interray is not well preserved and is apparently distorted. The radianal is interpreted to be beneath the C radial and is drawn (Fig. 83B) as it appears as 2 separate pieces but this is probably due to preservation. More distal anal plating is very unclear. Among the 5 species recognised in this genus only M. tuberosus Springer, 1920 from the Wenlock of Gotland has arms that divide 4 times but it has only 3 secundibrachs as opposed to 4 in this Australian species. Quadritaxocrinus gen nov. TYPE SPECIES. Quadritaxocrinus websteri sp. nov. ETYMOLOGY. Latin quadri-, four and generic name Taxocrinus. DIAGNOSIS. Cup small; infrabasals not visible in lateral view; primibrachs 4; arms branching isotomously at 151 division, with 3rd divisions on inner branches at same level and those on outer branches at same level but more distal than on inner branches; stem circular in section, of short columnals proximally. REMARKS. This genus compares closely with Taxocrinus in the infrabasals concealed by the stem, the wide posterior basal extending up between the C and D radials, shape of the axillary brachials and the lipped interbrachial articulations. Among Taxocrinidae the number of primibrachs is consistent at the generic level. Taxocrinus, with which Quadritaxocrinus is most closely allied, has 3 primibrachs and I suggest that evolution from the former to the latter in the Lower Devonian involved introduction of an extra primibrach. Ubaghs (1978a) pointed out that all Ordovician and Silurian, most Devonian and some Carboniferous genera have 2 primibrachs and from the Carboniferous on 3 is the standard number. More than 3 primibrachs are known in some species of Onychocrinus (Synerocrinidae) but that genus has entirely different arms from the new Australian taxon. The obvious inference is that the number of primibrachs increases with evolution in the flexible crinoids so evolution to 4 106 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 83. Meristocrinus quatriramus sp. nov.. large crown NMVP109765. from NMVPL 1924. А. closeup of cup in posterior view, *4. B, sketch of plate arrangement of A. C, posterior view of crown. «1.5. D, interior of arms, *4. E, anterior view of crown, х2 CRINOIDS FROM CENTRAL VICTORIA 107 FIG. 84. A.B, Quadritaxocrinus websteri gen. et sp. nov., holotype crown ММУР 149354 from NMVPL6601.A. E ray view, x3. B, С ray view with circular columnal concealing CD interray above anal X, x5. C,D, Geroldicrinus sp., crown NMVP149362 from NMVPL 1990, x3. C, lateral anterior view. D, distal view. primibrachs in Quadritaxocrinus is not unexpected even if it is the first record of such increase in the Taxocrinidae. Quadritaxocrinus websteri sp. nov. (Fig. 84A, B) ETYMOLOGY. For Gary D. Webster, Washington State University who provided many useful discussions on crinoids. MATERIAL. HOLOTYPE: NMVP149354 from NMVPL6601. DIAGNOSIS. As for genus. DESCRIPTION. Crown 35mm long. Infrabasals concealed by stem. Basals not clearly defined but probably 5, widerthan long; CD basal irregular in shape, with projection extending up between C and D radials. Radials pentagonal, with plenary facets. Arms long, dividing isotomously on 4th primibrach, with next division at secundibrach 4 producing unequal branches. with 3rd and 4th divisions at different levels, without interprimi- brachs. Stem circular in section, widest proximally. of short, strongly crenulate columnals proximally. REMARKS. The only available specimen is not clear in definition of the cup plates but the prox- imal and distal margins of the basals, with the posterior basal extending up betweenthe C and D radials and outline of the radials are clear. Order SAGENOCRINIDA Springer, 1913 Family LECANOCRINIDAE Springer, 1913 Geroldicrinus Jackel, 1918 TYPE SPECIES. Lecanocrinus roemeri Schultze, 1867 from the Middle Devonian of Germany. 108 Geroldicrinus sp. (Fig. 84C. D) MATERIAL. NMVP149362 from NMVPL1990. DESCRIPTION. Crown 15mm long. egg-shaped but truncated distally, crushed and therefore, not useful in comparing dimensions. Cup, conical, of smooth plates, but only the distal part of 1 anterior radial available. Infrabasals and basals not evident. Arms plenary, with 151 primibrach axillary, with 1 further division at different levels in same ray. closely abutting each other laterally, strongly incurved distally to fully enclose a space distal to the cup. Stem circular in section, of short columnals proximally, becoming longer and heteromorphic distally. REMARKS. Even though anal plating at the posterior and the lower cup are not available crown shape, arm branching and structure and particularly the axillary Ist primibrach are sufficient to identify this genus. Since Schultze (1867) described this genus it has not been recognised elsewhere and so this single specimen from Victoria is a significant range extension. STEM ONLY Group PENTAMERATA Stukalina, 1966 Order STRIALATA Stukalina, 1978 Family DECACRINIDAE Yeltyschewa, 1957 Decacrinus Yeltyschewa, 1957 TYPE SPECIES. Decacrinus pennatus Yeltyschewa, 1957 from the Lower Devonian of central Kazakhstan. Decacrinus sp. (Fig. 85) MATERIAL, NMVP149353 from NMVPL 1924. REMARKS. This form genus is distinctive in having 5 grooves radiating from the central canal towards the 5 outer angles of the stem as well as 5 shorter but wider grooves radiating from the central canal towards the middle of each side of the stem. Stukalina (1986) recorded this genus in the Early Devonian (Gedinnian and Siegennian) whichis the same age as the Australian specimen. Species identification of the Australian specimen is not attempted because it does not fit easily into a known species and I am reluctant to erect a new one on a single specimen without knowing the crown it supported. The projections at the 5 angles are very reminiscent of the stem of Pterinocrinidae gen. nov., described above. from the Lilydale area but preservation of that MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 85. Decacrinus sp.. length of stem NMVP149353 from NMVPL 1924. A, lateral view, x3. В, distal view of columnal, x6. specimen is not good enough for certain identification. ACKNOWLEDGEMENTS Numerous individuals collected and donated specimens described herein. Several are thanked by application of specific epithets but I cannot overestimate the acknowledgement due to Frank and Enid Holmes and Steve Eckardt for their unfailing support over several years of back- breaking excavation in Middendorp’s Quarry to secure the most outstanding collection of fossil echinoderms from any one Southern Hemisphere locality. І am grateful to Gary Webster, Wash- ington State University, and George McIntosh. Rochester Museum for much advice on crinoid taxonomy. I thank David Holloway, Museum of Victoria for curatorial and field support, Fons CRINOIDS FROM CENTRAL VICTORIA Vandenberg, Geological Survey of Victoria for help in the field and with stratigraphy, Tom Darragh, Museum of Victoria for encourage- ment, Charlotte Parker, for preparation of specimens, Penny Clark and Paul Avern for photographic support, Tom Guensburg and George McIntosh for constructive refereeing and my wife Annette who allowed me the many weekends taken up by this project and supported me throughout. This work was partially supported by an ARC Grant during 1983. LITERATURE CITED ANGELIN, N.P. 1878. Iconographia crinoideorum in stratis Sueciae Siluricus fossilium. (Samson & Wallin: Holmiae). 62p. ARENDT, Yu. A. 1965. K poznaniyu morskihh lilii kaltseokrinid [Contribution to the knowledge of crinoids from the Family Calceocrinidae ]. Paleontologicheskii Zhurnal 1965(1): 90-96. AUSICH, W.1. 1986a. Early Silurian Rhodocrinitacean crinoids (Brassfield Formation, Ohio). Journal of Paleontology 60: 84-106. 1986b. New camerate crinoids of the Suborder Glyptocrinina from the Lower Silurian Brass- field Formation (Southwestern Ohio). Journal of Paleontology 60: 887-897. 1987. Brassfield Compsocrinina (Lower Silurian crinoids) from Ohio. Journal of Paleontology 61: 552-562. AUSICH, W.1. & DRAVAGE, P. 1988. Crinoids from the Brassfield Formation of Adams County, Ohio. Journal of Paleontology 62: 285-289. AUSTIN, T. & AUSTIN, T. Jr 1843. XXXIII. Description of several new genera and species of Crinoidea. Annals and Magazine of Natural History, series 1, 11(69): 195-207. BATES, D.E.B. 1972. A new Devonian crinoid from Australia. Palaeontology 15: 326-335. BATHER, F.A. 1891. British fossil crinoids. 5. Bot- ryocrinus, Wenlock Limestone. Annals and Magazine of Natural History 67: 389-423. 1893. The Crinoidea of Gotland, Part 1, the Crin- oidea Inadunata. Kongliga Svenska Vetenskaps- akademiens Handlingar 25(2): 1-182. 1897. Hapalocrinus victoriae, n.s., Silurian, Melbourne, and its relation to the Platycrinidae. Geological Magazine, Decade 4, 4: 337-345. BILLINGS, E. 1857. On the Crinoidea or stone lilies of the Trenton Limestone, with a description of a new species. Canadian Naturalist and Geologist, series 1, 1: 48-57. BLANDOWSKI, W. 1855. A description of fossil animalculae in primitive rocks from the Upper Yarra district. Transactions of the Philosophical Society of Victoria 1: 221-223. BOUCOT, A.J., JOHNSON, J.G. & TALENT, J.A. 1969. Early Devonian brachiopod zoogeography. Special Papers of the Geological Society of America 119:1-107. 109 BOUSKA, J. 1947. Pygmaeocrinus, a new crinoid from the Devonian of Bohemia. Vestnik Kralovske ceske spolecnosti nauk, Trida 2 (mathematicko- prirodovedna) 1946: 1-4. BREIMER, A. 1962. A monograph of Spanish Palaeozoic Crinoidea. Leidse Geologische Mededelingen 27: 1-190. BRETT, C.E. 1981. Systematics and paleoecology of Late Silurian (Wenlockian) calceocrinid crinoids from New York and Ontario. Journal of Paleontology 55: 145-175. BRONN, Н.С. 1840. Стелосгіпих. ein neues Krinoiden-Geschlecht der Grauwacke. Neues Jahrbuch Mineralogie, Geologie, Paläontologie 542-548. BROWER, J.C. 1973. Crinoids from the Girardeau Limestone (Ordovician). Palaeontographica Americana 7: 261-499, 1976. Evolution of the Melocrinitidae. Thalassia Jugoslavica 12: 41-49. 1992, Cupulocrinid crinoids from the Middle Ordovician (Galena Group, Dunleith Formation) of northern Iowa and southern Minnesota. Journal of Paleontology 66: 99-128. 1995. Dendrocrinid crinoids from the Ordovician of northern Iowa and southern Minnesota. Journal of Paleontology 69: 939-960. CAS, R. 1983. Palaeogeographic and tectonic develop- ment of the Lachlan Fold Belt, southeastern Australia. Geological Society of Australia Special Publication 10: 1-104. CHAPMAN, F. 1903. New or little known fossils in the National Museum, Melbourne. Part 1. Some Palaeozoic species. Proceedings of the Royal Society of Victoria new series 15: 104-122. 1934. New species of a crinoid (Lecanocrinus) and a cephalopod (Ophidioceras), from the Silurian of Yass. Proceedings of the Royal Society of Victoria, new series 47: 190-195. ECKERT, J.D. 1984. Early Llandovery crinoids and stelleroids from the Cataract Group (Lower Silurian) in southern Ontario, Canada. Royal Ontario Museum Life Sciences Contributions 137: 1-83. ECKERT. J.D, & BRETT, C.E. 1985. Taxonomy and palaeoecology of the Silurian myelodactylid crinoid Crinobrachiatus brachiatus (Hall). Royal Ontario Museum Life Sciences Contributions 141: 1-15. ETHERIDGE, R. Jr 1904. The occurrence of Piso- crinus or an allied genus, in the Upper Silurian rocks of the Yass district. Records of the Aus- tralian Museum 5(5): 287-292. FOLLMANN, O. 1887. Unterdevonische Crinoiden. Verhandlungen des naturhistorischen Vereines der preussischen Rhinelande, Westfalens und des Reg.-Bezirks Osnabrück series 5 4: 113-138. FREST, T.J. & STRIMPLE, H.L. 1981. New camerate crinoids from the Silurian of North America. Journal of Paleontology 55: 639- 655. 110 GARRATT, М.Ј. 1983. Silurian and Devonian bio- stratigraphy of the Melbourne Trough, Victoria. Proceedings of the Royal Society of Victoria, new series 95: 77-98. GARRATT, M.J. & WRIGHT, A.J. 1988, Late Silurian to Early Devonian biostratigraphy of southeastern Australia. Pp. 647-662. In McMillan, N.J., Embrey, A.F. & Glass, D.J. (eds) Devonian of the World, Vol. 3. (Canadian Society of Petroleum Geologists: Calgary). GILL, E.D. & CASTER, K.E. 1960. Carpoid echinoderms from the Silurian and Devonian of Australia. Bulletins of American Paleontology 41: 1-71. GOLDRING, W. 1923. The Devonian crinoids of the State of New York. Memoirs of the New York State Museum 16: 1-670. HAARMANN, E. 1921. Die Botryocriniden und Lophocriniden des rheinischen Devons. Son- derabdruck aus dem Jahrbuch der Preussischen Geologischen Landesanstalt 41(1): 1-87. HALL, J. 1847. Palaeontology of New York, vol. 1, containing descriptions of the organic remains of the lower division of the New York System (equivalent of the Lower Silurian rocks of Europe). Natural History of New York 6; 1-338. 1852. Palaeontology of New York, vol. 2, con- taining descriptions of the organic remains of the lower middle division of the New York System. Natural History of New York 6: 1-362. 1863. Notice of some new species of fossils from a locality of the Niagara Group, in Indiana; with a list of identified species from the same place. Transactions of the Albany Institute 4: 195-228. 1872. Description of new species of Crinoidea and other fossils from strata of the age of the Hudson River Group and Trenton Limestone. Annual Report of the New York State Museum of Natural History 24: 205-224. HAUDE, R. 1995. Lower Devonian echinoderms from the Precordillera (Argentina). Neues Jahrbuch fiir Geologie und Paläontologie, Abhandlungen 197: 37-86. HISINGER, 1840. Lethaea Suecica, seu petrificata Sueciae, iconibus et characteribus illustrata. Supplementum secundum, pp. 1-11, pls 38, 39. JAEKEL, O. 1895. Bietrage zur kenntnis der palaeozoischen Crinoiden Deutschlands. Palaeontologisches Abhandlungen, new series 3: 1-116. 1898, Über einige paláontologische Gattungen von Crinoiden. Deutsche Geologisches Gesellshafte Zeitschrift Verhandlingen 49: 44-48. 1918. Phylogenie und System der Pelmatozoen. Palaeontologische Zeitschrift 3: 1-128, JELL, P.A. 1982. Crotalocrinites pulcher (Hisinger, 1840) from the Early Devonian of central Victoria. Alcheringa 6: 174. 1983. Early Devonian echinoderms from Victoria (Rhombifera, Blastoidea and Ophiocystioidea). MEMOIRS OF THE QUEENSLAND MUSEUM Memoirs of the Association of Australasian Palaeontologists 1: 209-235. JELL, P.A. & JELL, J.S. 1999, Crinoids, a blastoid and a cyclocystoid from the Upper Devonian reef complex ofthe Canning Basin, Western Australia. Memoirs ofthe Queensland Museum 43:201 -236. JELL, P.A. & HOLLOWAY, D.J. 1983. Devonian and ?Late Silurian palaeontology of the Winneke Reservoir site, Christmas Hills, Victoria. Proceedings of the Royal Society of Victoria 95: 1-21. JELL, P.A., JELL, J.S., JOHNSON, B.D., MAWSON, R. & TALENT, J.A. 1988. Crinoids from Dev- onian limestones of eastern Australia. Memoirs of the Queensland Museum 25: 355-402. JELL, Р.А. & THERON, I.N. 1999, Early Devonian echinoderms of South Africa. Memoirs of the Queensland Museum 43: 115-199. JOBSON, L. & PAUL, C.R.C. 1979. Compagicrinus Jenestratus, a new Lower Ordovician inadunate crinoid from North Greenland. Rapport Gronlands Geologic Undersag 91: 1-81, KESLING, R.V. 1964. A new species of Melocrinites from the Middle Devonian of Michigan. University of Michigan Contributions from the Museum of Paleontology 19: 89-103. 1966. Botryocrinus niemani, anew crinoid from the Middle Devonian Silica Formation of Ohio. University of Michigan Contributions from the Museum of Paleontology 20: 271-276. 1968. Ameliacrinus benderi, a new dicyclic camerate crinoid from the Middle Devonian Silica Formation in northwestern Ohio. University of Michigan Contributions from the Museum of Paleontology 22: 155-162. KESLING, R.V. & CHILMAN, R.B. 1975. Strata and megafossils of the Middle Devonian Silica Formation. University of Michigan, Museum of Paleontology, Papers on Paleontology 8: 1-408. KIER, P.M. 1952. Echinoderms of the Middle Devonian Silica Formation of Ohio. Contributions from the Museum of Paleontology at the University of Michigan 10: 59-81. KIRK, E. 1914. Notes on the fossil crinoid genus Homocrinus Hall. Proceedings of the United States National Museum 46: 473-483. 1929, The status of the genus Mariacrinus Hall. American Journal of Science 17: 337-346. 1945. Four new genera of camerate crinoids from the Devonian. American Journal of Science 243: 341-355. KOLATA, D.R. 1975. Middle Ordovician echinoderms from northern Illinois and southern Wisconsin. Memoirs of the Paleontological Society 7: 1-74. LEMENN, J. 1975. Un nouveau genre d' Hexacrinitidae (Crinoidea, Camerata). Annales de la Societé Géologique du Nord 95: 243-250. 1985. Les Crinóides du Dévonien Inférieur et Moyen du Massif Armoricain. Mémoires de la Societé Géologique et Minéralogique de Bretagne 30: 1-268. CRINOIDS FROM CENTRAL VICTORIA McINTOSH, С.С. 1979, Abnormal specimens of the Middle Devonian crinoid Bactrocrinites and their effect on the taxonomy of the genus. Journal of Paleontology 53: 18-28. 1981. Apurocrinus sucrei, a new genus and species of camerate crinoid from the Lower Devonian of Bolivia. Journal of Paleontology 55: 948-952. 1983. Review of the Devonian cladid inadunate crinoids: Suborder Dendrocrinina. PhD Thesis, Univ. Michigan, 521p. 1986. Phylogeny of the dicyclic inadunate crinoid Order Cladida. Abstracts from the 4th North American Paleontological Convention. 1987. Review of the Devonian camerate crinoid Bogotacrinus scheibei Schmidt from Colombia. Journal of Paleontology 61: 750-757. MEEK, F.B. & WORTHEN, A.H. 1869. Descriptions of new Crinoidea and Echinoidea from the Carboniferous rocks of the western states, with a note on the genus Onychaster. Proceedings of the Academy of Natural Sciences of Philadelphia 21: 67-83. MILITSINA, V.S. 1980. Cystoidea and Crinoidea from the Ordovician and Silurian of the Urals. Ezhegodnik Bsesoyoznogo Paleontologi- cheskogo Obshchestva 23: 198-215. MILLER, J.S. 1821. A natural history of the Crinoidea or lily-shaped animals, with observations on the genera Asteria, Euryale, Comatula, and Marsupites. (Bryan & Co.: Bristol). MILLER, S.A. & GURLEY, W.F.E. 1895 New and interesting species of Palaeozoic fossils. Bulletin of the Illinois State Museum of Natural History 7: 1-89. MOORE, R.C. 1962. Revision of Calceocrinidae. University of Kansas, Paleontological Contributions, Article 4: 1-40. MOORE, R.C., LANE, N.G. & STRIMPLE, H.L., 1978. Order Cladida. Pp. T578-T759. In Moore, R.C. & Teichert, C. (eds) Treatise on invertebrate paleontology, Part T, Echinodermata 2. (Geological Society of America & University of Kansas Press: Boulder, Colorado & Lawrence, Kansas). MOORE, R.C., LANE, N.G., STRIMPLE, H.L. & SPRINKLE, J., 1978. Order Disparida. Pp. T520-577. In Moore, R.C. & Teichert, C. (eds) Treatise on invertebrate paleontology, Part T, Echinodermata 2. (Geological Society of America & University of Kansas Press: Boulder, Colorado & Lawrence, Kansas). MOORE, R.C. & TEICHERT, C. (eds) 1978. Treatise on invertebrate paleontology, Part T. Echinodermata 2 (3 vols). (Geological Society of America & University of Kansas Press: Boulder, Colorado & Lawrence, Kansas). MULLER, J. 1856. Über neue Crinoiden aus dem Eifeler Kalk. Koniglich Akademie der Wissenschaft Berlin, Monatsbericht 1856: 353-356. 111 MUNSTER, G.G. 1839, Beschreibung einiger neuen Crinoideen aus der Uebergangs-formation. Beitrage zur Petrefaten-Kunde 1: 31-34. MURCHISON, R.I. 1839. The Silurian System, Part 1. Founded on researches in the counties of Salop, Hereford, Radnor, Montgomery, Caermarthen, Brecon, Pembroke, Monmonth, Gloucester, Worcester, and Stafford; with descriptions of the coalfields and overlying formations. 578p; Part 2. Organic remains. 579-768. (John Murray: London). d'ORBIGNY, 1850. Prodrome du paléontologie stratigraphique universelle des animaux mollusques et rayonnes faisant suite au cours elementaire de paleontologie et de geologie stratigraphique. Vol. 1. (Masson: Paris). PHILIP, G.M. 1961. Lower Devonian crinoids from Toongabbie, Victoria, Australia. Geological Magazine 98: 143-160. PHILIP, G.M. & STRIMPLE, H.L. 1971. An interpretation of the crinoid Aethocrinus moorei Ubaghs. Journal of Paleontology 45: 491-493. PHILLIPS, J. 1841. Figures and descriptions of the Palaeozoic fossils of Cornwall, Devon, and West Somerset. (Longman, Brown, Green & Long- mans: London). PROKOP, R.J. 1970. Family Calceocrinidae Meek & Worthen, 1869 (Crinoidea) in the Silurian and Devonian of Bohemia. Sbornik Geologickych Ved, Rada P 12: 79-134. 1982. Some new hexacrinitids (Crinoidea, Cam- erata) from the Lower Devonian of Bohemia. Vestnik Ustredniho Ustavu Geologickeho 57: 277-284. 1973. Elicrinus n. gen. from the Lower Devonian of Bohemia (Crinoidea). Vestnik Ustredniho Ustavu Geologickeho 48: 221-224. PROKOP, R.J. & PETR, V. 1997. The genus Pygmaco- crinus Bouska, 1947 (Crinoidea, Inadunata) in the Devonian of the Barrandian area (Czech Republic). Acta Musei Nationalis Pragae, Series B. Historia Naturalis 53: 1-10. RAMSBOTTOM, W.H.C. 1961. A monograph on British Ordovician Crinoidea. Palaeontographical Society Monograph 114: 1-37. RINGUEBERG, E.N.S. 1888. Some new species of fossils from the Niagara shales of western New York. Academy of Natural Sciences of Philadelphia Proceedings for 1888: 131-137. ROEMER, C.F. 1855. Erste Periode, Kohlen-Gebirge. In Bronn, H.G. Lethaea Geognostica, vol. 2 (3rd ed.) (E. Schweizerbart: Stuttgart). ROZHNOV, S.V. 1981. Crinoids of the Superfamily Pisocrinacea. Trudy Paleontological Institute 192: 1-127. SALTER, J.W. 1856. Description of Palaeozoic Crustacea and Radiata from South Africa. Transactions ofthe Geological Society of London 7: 215-224. SCHMIDT, W.E. 1913. Cultrijugatuszone und unteres Mitteldevon sudlich der Attendorn-Elsper Doppelmulde. K. Preussen Geologisches Landesanst. Jahrbuch 33(2): 265-318. 1934. Die Crinoideen des Rheinischen Devons. Teil 1. Abhandlungen der Preussischen Geologischen Landesanstalt, Neue Folge 163: 1-199. 1941. Die Crinoideen des Rheinischen Devons. Teil 2. Abhandlungen der Reichsstelle für Bodenforschung, Neue Folge 182: 1-253. SCHULTZE, L. 1867. Monographie der Echinodermen des Eifler Kalkes. Denkschriften der Kaiserlichen Akademie der Wissenschaften, mathematisch- naturwissenschaftliche Klasse 26: 113-230. SHUMARD, B.F. 1855. Description of new species of organic remains. Missouri Geological Survey 2: 185-208. SPRINGER, F. 1906. Discovery of the disk of Onycho- crinus, and further remarks on the Crinoidea Flexibilia. Journal of Geology 14: 467-523. 1911. On a Trenton echinoderm fauna at Kirkfield, Ontario. Memoirs of the Geological Survey of Canada 15: 1-69, 1920. The Crinoidea Flexibilia. Smithsonian Institution Publication 2501: 1-486. 1923. On the crinoid Family Catillocrinidae. Smith- sonian Miscellaneous Collections 76(3): 1-41. 1926. American Silurian crinoids. Publications of the United States National Museum 2871: 1-240. STEWART, G.A., 1940. Crinoids from the Silica Shale, Devonian, of Ohio. Ohio Journal of Science 40: 53-61. STRIMPLE, H.L. 1963. Crinoids of the Hunton Group (Devonian - Silurian) of Oklahoma. Bulletin of the Oklahoma Geological Survey 100: 1-169. STRIMPLE, H.L. & LEVORSON, C.O. 1973. Additional crinoid specimens from the Shellrock Formation (Upper Devonian) of Iowa. Proceed- ings ofthe lowa academy of Science 80: 182-184. STUKALINA, G.A., 1986. Zakonomernosti istoricheskogo razvitiya krinoidei b rannem i crednem paleozoe SSSR [Laws of historical development of crinoidea from the early and middle Paleozoic of the USSR.] (Akademiya Nauk USSR, Paleontological Institute: Moscow). TALBOT, M. 1905. Revision of the New York Helderbergian crinoids. American Journal of Science, series 4, 20: 17-34. TALENT, J.A. 1965. The Silurian and Early Devonian faunas of the Heathcote district, Victoria. Memoirs of the Geological Survey of Victoria 26: 1-55. UBAGHS, G. 1958, Recherches sur les Crinóides Camerata du Silurien de Gotland (Suéde). Partie III: Melocrinicae. Avec des remarques sur l'évolution des Melocrinidae. Arkiv fór Zoologi series 2, 11: 259-306. 1969, Aethocrinus moorei Ubaghs n. gen., n. sp., le plus Ancien Crinoide Dicyclique Connu. University of Kansas Paleontological Contributions, Paper 38: 1-25. 19782. Skeletal morphology of fossil crinoids. Pp. T58-T216. In Moore, R.C. & Teichert, C. (eds) MEMOIRS OF THE QUEENSLAND MUSEUM Treatise on invertebrate paleontology. (Geological Society of America & University of Kansas: Boulder, Colorado & Lawrence, Kansas). 1978b. Camerata Pp. T408-T519. In Moore, R.C. & Teichert, C. (eds) Treatise on invertebrate paleontology. (Geological Society of America & University of Kansas: Boulder, Colorado & Lawrence, Kansas). VANDENBERG, A.H.M., 1988. Silurian - Middle Devonian. Pp. 103- 146. In Douglas, J.G. & Ferguson, J.A. (eds) Geology of Victoria. (Victorian Division, Geological Society of Australia: Melbourne). 1992, Kilmore 1:50,000 map geological report. Geological Survey of Victoria Report 91: 1-86, VANDENBERG, A.H.M. & WILKINSON, H.E. 1982. Victoria. In Cooper, R.A. & Grindley, G.W. (eds) Late Proterozoic to Devonian sequences of southeastern Australia, Antarctica and New Zealand and their correlation. Special Publications of the Geological Society of Australia 9: 36-47. WACHSMUTH, C. & SPRINGER, F. 1880. Revision of the Palaeocrinoidea, part 1. The families Ichthyocrinidae and Cyathocrinidae. Proceedings of the Academy of Natural Sciences of Philadelphia for 1879: 226-378. 1881. Revision of the Palaeocrinoidea, part 2. Family Sphaeroidocrinidae, with the sub- families Platycrinidae, Rhodocrinidae, and Actinocrinidae. Proceedings of the Academy of Natural Sciences of Philadelphia for 1881: 175-411. 1885. Revision of the Palaeocrinoidea, part 3, section 1. Discussion of the classification and relations of the brachiate crinoids, and conclusion of the generic descriptions. Proceedings ofthe Academy of Natural Sciences of Philadelphia for 1885: 223-364. WILLIAMS, СЕ. 1964. The geology of the Kinglake district, central Victoria. Proceedings of the Royal Society of Victoria, new series 77: 273-328. WILLIAMS, H.S. 1883. On a crinoid with movable spines (Arthroacantha ithacensis). American Philosophical Society Proceedings 21: 81-88. WITHERS, R.B. & KEBLE, R.A. 1934a. The Palaeo- zoic starfish of Victoria, Proceedings of the Royal Society of Victoria 46: 220-249. 1934b. The Palaeozoic brittlestars of Victoria. Pro- ceedings of the Royal Society of Victoria 47: 196-212. WITZKE, B.J. & STRIMPLE, H.L. 1981. Early Silurian camerate crinoids of eastern Iowa. Iowa Academy of Science, Proceedings 88: 101-137. YAKOVLEV, N.N. 1946. Un Hexacrinidae du silurien supérieur. Compte Rendu (Doklady) de Academie des Sciences de l'URSS 51: 153-154. YELTYSCHEWA, R.S. 1957. O novom semeistwe paleosoiskih morskih lilii. [On a new family of Paleozoic crinoids]. Ezhegodnik Vsesoyuznogo Paleontologicheskogo Obshchestva 14: 218-234. CRINOIDS FROM CENTRAL VICTORIA 113 APPENDIX List of localities arranged in descending age and grouped as per the columns numbered 1-11 in Figure 1. Column 11. Late Boucotia australis Zone (Lale Lochkovian). NMVPL229 Collins Quarry, on access road to youth camp about 2 km N of Tommy's Hut, Kinglake (=X0 of Williams, 1964) Hollowaycrinus calvus Eudimerocrinus eckardti Oehlerticrinus lemenni Oehlerticrinus jeani Frankocrinus holmesi Ctenocrinus signatus Ctenocrinus paucidactylus Phimocrinus americanus Antihomocrinus chapmani Ancyracrinus sp. Kroppocrinus heathcotensis NMVPL1924 sandstone bar across Mathieson's Creek about Ikm S of Kinglake to Flowerdale Road and about 6km NE of Tommy's Hut (=T95 of Williams, 1964) Duncanicrinus calvariolus Eudimerocrinus eckardti Ctenocrinus sp. Myelodactylid sp. Kroppocrinus mathiesonensis Stewhrecrinus terryi Antihomocrinus chapmani Meristocrinus quatriramus Decacrinus sp. Column 10. Middle Boucoria australis Zone (Late Lochkovian). Locality R52 of Thomas (1940), Unit 3 Mt Ida Fmn, Parish of Redcastle, N of Heathcote; see Talent (1965, fig. 1) Holmesocrinus idaensis Locality R25 of Thomas (1940), Unit 3 Mt Ida Fmn, Parish of Redcastle, N of Heathcote; see Talent (1965, fig, 1) Crotalocrinites pulcher Ctenocrinus paucidactylus Column 9. Early Boucotia australis Zone (Late Lochkovian). NMVPL252 Middendorp's Quarry, 2km E of Tommy's Hut, Kinglake (=W3 of Williams, 1964). Dimerocrinites bispinosus Eucrinus clarkae Eudimerocrinus eckardti Nexocrinus sp. Duneanicrinus calvariolux OQehlerticrinus lemenni Oehlerticrinus jeani Frankocrinus enidae Степосгіпих paucidactylus Ctenocrinus stellifer Ctenacrinus signatus Clematocrinus perforatus Darraghcrinus tonii Codiacrinus secundus Dendrocrinus arrugius Plicodendrocrinus australis Antihomocrinus chapmani Dictenocrinus ihaevpus Rubbish tip on Watson's Road at Pheasant Creek, 6km E of Tommy's Hut, Kinglake West. Duncanicrinus calvariolus Column 8. Late Boucotia janeae Zone (Early Lochkovian). NMYPL1990, Road cutting just N of intersection ol Victoria Road and Coldstream West Road, 5km N of Lilydale, NE Melbourne. Ophiocrinus nnettae Eudimerocrinus gilli Ctenocrinus stellifer Pterinocrinidae gen. nov. Shintocrinus richi sp. nov. Dictenocrinus remotus sp. nov. Geroldicrinus sp. Roadcutting on road (Yarra Road) to Wonga Park. just 5 of Bryson's Road. Dictenocrinus sp. cf. D. ibaeypus Column 7. Middle Boucotia janeae Zone (Early Lochkovian). NMVPL1922 sewerage trench in Kimberley Drive, Chirnside Park, N of Maroondah Highway 3.5km W оГ Lilydale. Hexacrinites chirnsidensis Antihomocrinus chapmani Crenocrinus stellifer Column 6. Early Boucotia janeae Zone (Early Lochkovian). NMVPLI84I small excavation on ridge above Ruddock’s Quarry (now filled in) W of Edwards Road, 300m NW of its intersection with Switchback Road, Chirnside Park. Cupulocrinus austrogracilis Column 5. Late Notoparmella plentiensis Zone (Pridoli). NMVPL1923 large roadcutting just N of Merriang on Woodstock to Wallan Road, 9km N of Woodstock, 40km М of Melbourne. Holmesocrinus enidae 114 Nexocrinus wallanensis Trichocrinus marleyi NMVPL260 Excavations for Winneke Dam, 33km NE of Melbourne, (Jell & Holloway, 1983). Duncanicrinus calvariolus Kooptoonocrinus nutti Dendrocrinus saundersi Codiacrinus rarus Column 4. Early Notoparmella plentiensis Zone (Late Ludlow). NMVPL300 near disused mine оп Comet Creek, c. 4.6km SE of Clonbinane, 60km N of Melbourne; (=X64 in Clonbinane Sandstone of Williams, 1964). Clematocrinus perforatus Alisocrinus lineatus Phimocrinus hanschi Dendrocrinus arrugius Shintocrinus cometensis Antihomocrinus chapmani NMVPL6601 on Boundary Road spur above Comet Creek Mine (NMVPL300), 4.6km SE of Clonbinane, 60km N of Melbourne. Quadritaxocrinus websteri Column 3. Late Aegiria thomasi Zone (Ludlow). SW of Bald Hills, just E of railway, N of Sunday Creek Road, 5km E of Kilmore. Dendrocrinus arrugius NMVPL299 road cutting on Sunday Creek Road, 700m NE of junction with Saunders Lane, Kilmore East (—F31 of Williams, 1964). Nassoviocrinus corcorani Trichocrinus morleyi Column 2. Late Aegiria thomasi Zone (Ludlow). MEMOIRS OF THE QUEENSLAND MUSEUM Yarra Improvement Works (refers to works to straighten the river between the South Yarra Railway Bridge and Princes Bridge, at the end of last century). Hapalocrinus victoriae NMVP1615, excavation on ridge on W of railway cutting c. 800m WSW of Royal Park Railway Station, Parkville, Melbourne Phimocrinus americanus Nassoviocrinus longibrachiatus Trichocrinus morlevi Quarry (now in Clifton Park) at West Brunswick between Albert and Victoria Streets. Helicocrinus plumosus NMVPL2259 in creek bed about 250m N of Heathcote- Nagambie Road, E of Argyle Railway Station, Heathcote (=H41 of Thomas, 1940). Clematocrinus argylensis Kroppocrinus heathcotensis Nassoviocrinus corcorani NMVPL1925 in creek bed under bridge (on Sunday Creek Road) just west of railway line at Kilmore East. Dendrocrinus arrugius Upper Yarra (no further details; grouped with other localities having this species). Nassoviocrinus longibrachiatus Excavation at E end of Melbourne CBD (presumably in area of Spring Street, possibly at Flinders Street intersection; exact locality unknown)) Hapalocrinidae indet. Column 1. Middle Aegiria thomasi Zone (Ludlow). NMVPL!1927 at crossing of Broadhurst Creek by Kilmore to Wandong Road (Kilmore Siltstone). Dendrocrinus sp. EARLY DEVONIAN ECHINODERMS FROM SOUTH AFRICA PETER A. JELL AND JOHANNES N. THERON Jell, P.A. & Theron, J.N., 1999 06 30: Early Devonian echinoderms from South Africa. Memoirs of the Queensland Museum 43(1): 115-199. Brisbane. ISSN 0079-8835. Echinoderms of the Lower Devonian Bokkeveld Group in the Cape Province, South Africa, have played an important role in sedimentary studies ofthe region but their taxonomic status has been known from only a few cursory papers. We here provide taxonomic treatment of all available crinoid (15 species), asterozoan (11) and blastoid (2) species revising all previously described taxa and more than doubling the known diversity. Palaeobiogeographic affinities appear to be with Europe and the USA but known faunas of South America and Australia are small and almost certainly incompletely known. New taxa described are the crinoid genera Mandelacrinus nelsoni, Eckidocrinus interbrachiatus, Sacrinus gamkaensis, S. hexensis, Monaldicrinus johni and Othozecrinus royi and the asterozoan genera Aulacolatiaster breviramus and Hotchkissaster macrodentatus. Hexuraster is introduced as a replacement for preoccupied Hexura Spencer. New species are the crinoids Kopficrinus halbichi, Cradeocrinus plenarius and Thalamocrinus arenaceus and the asterozoans Marginura hilleri and Eugasterella africana. O South Africa, crinoids, starfish, ophiuroids, Early Devonian, Bokkeveld. Peter A. Jell, Queensland Museum, P.O. Box 3300, South Brisbane 4101, Australia; Johannes N. Theron, Geological Survey of South Africa, P.O. Box 572, Bellville 7530, South Africa; 20 September, 1998. Echinoderms were first recorded from the Lower Devonian Bokkeveld Group of South Africa as early as 1816 by Dr G. Thom, a clergy- man by occupation but also an ardent amateur fossil collector. He gathered ‘endless numbers of specimens of shells, trilobites and encrinites or stone lilies'( Thom, 1830). Thus crinoids were the first echinoderms described (Salter, 1856). The Devonian age for the Bokkeveld fossils was assigned about the same time (Sandberger, 1852; Sharpe & Salter, 1856) but in the second half of the 19th century very little was attempted on the identification of these fossils. Rogers and Schwarz mapped these rocks in detail beginning in 1895 and assembled extensive fossil collections that formed the subject of numerous studies illuminating the fossil distribution and stratigraphical setting. Their Devonian age was also confirmed by comparison with faunas from South America and Europe (Corstorphine, 1897; Schwarz, 1906; Reed, 1904, 1906; Knod, 1908; Clarke, 1913; Kozlowski, 1923). In a comprehensive review of the Bokkeveld fauna, Reed (1925) recorded the first carpoid, blastoid and starfish. Echinoderms comprise only a minor element in the fauna however, and detailed descriptions of the various echinoderms in several collections have been rare. Ophiuroids have been mentioned by Rossouw (1933), Spencer (1930, 19502) and Rilett (1971). Rennie (1936) described carpoids from Gamkapoort, Breimer & Macurda (1972) described blastoids from the Ceres/De Doorns area and Ruta & Theron (1997) described carpoids in detail. In this paper we have revised the description and in many cases the assignment of previously described echinoderms (crinoids, blastoids and asterozoans) and have described substantial new collections from the South African Museum, the Geological Survey of South Africa (mainly collected by J.N.T. during stratigraphic field mapping during the 1970s to 1990s) and the private collection of Mr Roy Oosthuizen of Klaarstroom. STRATIGRAPHIC SETTING The clastic Cape Supergroup borders the southern and western parts of South Africa for 800km eastwards and almost 300km northwards from Cape Town (Fig. 1). The Supergroup 1s divided into a lower, predominantly arenitic Table Mountain Group, conformably overlain by the markedly argillaceous Bokkeveld Group, which in turn is overlain by the more arenitic Witteberg Group (Fig. 2). The Bokkeveld Group consists of a cyclical alternation of predominantly argillaceous and arenaceous units. Each of these extensive litho- stratigraphic units is given formation status (Fig. 2). The 6 lower formations which can be traced 116 Vanrhynsdorp a n S т. 2 © o [s] > z Cape Town MEMOIRS OF THE QUEENSLAND MUSEUM a BOKKEVELD GROUP ~ W(Port Elizabeth INDIAN OCEAN 24 FIG. 1. Sketch map of South Africa showing outcrop areas of the Bokkeveld Group in solid black. throughout the outcrop area are collectively referred to as the Ceres Subgroup. The upper part of the sequence in the west, with 5 formations, is designated the Bidouw Subgroup, whilst eastwards the laterally equivalent Taka Subgroup consists of 3 formations (Theron, 1972; Theron & Johnson, 1991). Thickness of the Bokkeveld Group is much greater in the east than in the west. These alternating lithostratigraphic units represent 5 major, sheet-like, superimposed, coarsening-upward cycles which feather out southwards into a relatively homogeneous mudstone-siltstone sequence. This southward decrease of coarser clastics is linked to a progressive thickening of the argillaceous units and of the Bokkeveld Group. The arenaceous units vary from fine-grained quartz arenites to immature arkosic arenites, either horizontally laminated or planar to trough cross-bedded. Hummocky cross stratification becomes a prominent structure northwards (Theron et al., 1995). The argillaceous units consist of dark grey shale, mudstone and siltstone with thin intercalated lenses of fine to medium grained lithic sandstone. The latter reveal swaley cross-stratification and often marked ripple cross laminated zones towards the north. Weathering of the sequence gives rise to hogback topography, the more resistant arenites creating ridges whereas the intervening argillites generally weather predominantly recessively. This distinctive weathering minimises good exposures of the argillites. Although numerous fossils have been collected and described, little attention has been paid to their stratigraphic occurrence so that for many years there was no clarity as to whether any zonal scheme could be developed. The faunas vary at a gross level with geography (Schwarz, 1906; Theron, 1972; Oosthuizen, 1984) in that brachiopods and echinoderms are more common in the west, with conulariids, corals and hyoliths more prevalent towards the east. Usually, marine invertebrate faunas are most common in the Ceres Subgroup, but are found up to the level of the Karoopoort Formation in the west and the Karies Formation in the east (Fig. 2). Although present throughout the Bokkeveld Group, plant and especially trace fossils, become abundant in the northwestern outcrop areas. A wide variety of typically shallow marine ichno- genera occur with large numbers of a variety of the uncommon pentameral stellate trace fossil Asteriacites (Fig. 3) at various horizons within these proximal arenites (Theron, 1972). Examination of sediment/phyla associations reveals distinct affinities of certain species for a particular lithology (Reed, 1907; Theron, 1972; Oosthuizen, 1984). Since a large percentage of the Bokkeveld fauna is benthic organisms, they are likely to be facies controlled. Trilobites and cephalopods tend to occur mainly in argillaceous horizons in contrast to the gastropods, bivalves, and brachiopods which are found more evenly in arenites and argillites. Among echinoderms, the crinoids are found in a variety of sediment types from mudstones to lithic arenites. Although stem fragments are prolific, well-preserved crowns are largely confined to finer grained sediments. Although scarce, ophiuroids, carpoids and EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA KAROQPOORT FORMATION OSBERG FORMATION KLIPBOKKOP FORMATION WABOOMBERG FORMATION BOPLAAS FORMATION TRA-TRA FORMATION HEXRIVER FORMATION VOORSTEHOEK FORMATIO GAMKA FORMATION GYDO FORMATION Vterbated some plant 117 Quartzose sandatono, siftetone, minor mudetone, medium- to fine-grained, thin- te thich-bedded, planar and trough crose-bedded, SNistone and sandstone, latter predominates i ertiwards, fine- te modium-grained, thin boddi horizoatal and ripple laminated, ripple marks common letrac! p e Sandstone and siltstene, fine- to vilem агн) thin- te thick-bedded, horizontal and ripple laminated, planar and trough crose-bedded, channeling occurs, intra ciast lenses, bloterbated, piant fragments. last lenses, bloturbated, rare plant fragments. I Sitetono, siity mudetone and minor sandstone beds, tine- 10 modium-gralned, horizoatal and ripple laminated, planer cross-bedding predominates, trowgh crose-bedding occurs, abundant ripple marke, intraciaet lences, bleturbated, seme plant fragmenta. Sandstone with thia histone Jan, lenses fine- to modium stained, | Frei to thich-bedded, horizontal, and PDT ad trough crose-bedded, а lant ripple Biftstone and mudstone, rare thia bedded sandstone, massive or horizontal and ripple laminated, intraciast lenses bloturbated, loeaifilerous lenses. Sandstone, Iino- to medlum-grained, thin- te thick-bedded, horizontal and ripple laminated, planar and trough crose-bedded, thia intraciaat lenses, as wel aa numerous thin bones of plaat fragments. marke, intraciast lenses, Silistone and silty mudstone and minor thin-bedded, sandstone, ripple and micro cross laminated, w abuadant ripple marke, bloterbation and trace fosalia, thia intraciact lenses, plant fragments. Sandstone, tino- to medium-grained, thin- to thickh-bedded, medium scale planar & trough crese-bedded, thin Intraciaet lenses. asive, horizental-, гірріо- and miore cresa seliiferouo tenses. i-lamina , bloturb Я Sandstone, Mne- to modium-grained, thin- te thick-bedded, paralel laminated, planar crece-Bedéed, thin Intraciaot lenses, 'ocolfereue lenses. 4 чә © соб. мобооте a eme A minor thin-bedded sandstone, scattered lossiferove nodules and lenses, iontiouiar bedding, abundant blotorbation Quartzees sandstone medium- te fhe-grained, thin- to thich-bedded, planar & shallow trough arece-beds FIG. 2. Stratigraphic section of South African Devonian. The coarse sandstones at top and bottom of section are contiguous parts of the Witteberg (C3) and Table Mountain (C1) Groups, respectively and indicate upper and lower margins ofthe Bokkeveld Group (C2). All sediments are siliciclastics; solid black = mudstone, horizontal dashes — down symbol = shells; convex up symbol = leaf shaped symbol — siltstone and close dots = fine sandstone. Fossil content is indicated by: star = echinoderms; convex trilobites; two curved lines crossing towards righthand end = fish; plant fragments. A shorthand nomenclature to reflect cyclicity within the Bokkeveld Group has emerged so that the shales (S) alternate with the sandstones (Q)and are numbered from the base up; thus the Gydo Formation is C2S1, the Gamka Formation is C2Q1, the Voorstehoek Formation is C2S2, etc. blastoids are similarly found mainly in mud- stones, shales or silty shales. Faunal community structures are recognised (Boucot, 1971; Hiller & Theron, 1988). The overall decrease of Bokkeveld invertebrates and increase in plant fragments and ichnofossils to the north as the argillaceous units become sandier, suggests a shallowing ofthe basin in that direction. There is a corresponding decrease in fossil content in a southerly direction approach- ing the deep basin (Theron, 1970, 1972; Theron & Loock, 1988). PALAEOGEOGRAPHICAL SETTING The conspicuous change from a few thousand metres of supermature sand (Table Mountain Group) to the predominantly muddy sediments of the Bokkeveld Group throughout the Cape Basin in the Early Devonian, is interpreted as an overall northward advance of the shoreline and progression of shelf and delta slope sediments (Gydo Formation) across the sand-shoal Rietvlei Formation. The latter constituting the uppermost unit of the Table Mountain Group, was deposited in a wide shallow embayment open to the southeast and flanked by a mature low gradient coastal plain (Rust, 1973). The Bokkeveld sequence reflects the most dynamic phase of the Cape Basin development, when, at the Pragian-Emsian transition, tectonic activity and accelerated downwarping evolved. Pulsatory cyclicity in the vertical stacking of the upward coarsening sequences implies tecton- ically controlled regressions and transgressions (Theron, 1972; Tankard & Barwis, 1982). These major cycles represent the progradation of lobate wave-dominated deltas along a coastline of moderately high marine energy (Tankard & 118 MEMOIRS OF THE QUEENSLAND MUSEUM FIC i 3. 11451 Barwis. 1982: Theron & Loock, 1988). Nearshore deposits grade southward into thick shelf mudstones, with the greater thickness towards the eastern Cape reflecting increased downwarping in that direction, An idealised Bokkeveld genetic sequence consists of sediments laid down successively in the shelf, delta slope and delta platform environments during the constructional phase of delta growth. This is in turn overlain by nearshore marine reworked deltaic deposits. which represent the sediments that evolved during the destructional phase of delta development (Tankard & Barwis. 1982). Storm activity is well-documented in the delta platform sediments. which represent the distributary mouth bar. interdistributary bay and tidal flat deposits, especially in the northern Bokkeveld facies (Theron et al., 1995). Fossils are generally sparse in these sand-, silt-, and mudstones, but occasionally occur as relatively thick lenticular Asteriacites sp. A, assemblage of 10 or more complete or partial individual traces in close proximity. from Vanrhynsdorp (C282), x12 . (Photo courtesy of John Almond). coquinites. Fossils are also generally sparse in the reworked sands of the delta platform. Wave and tidal activity created interspersed barrier washover sheets and tidal inlet and channel fill sequences that were not very conducive to the preservation of fossils. On the other hand the dark grey mud and siltstones of the Bokkeveld Group contain a rich invertebrate fauna. especially brachiopods and bivalves. which are preserved as scattered internal and external moulds. Coquinites, where present. are relatively thin but laterally persistent; as water depth increased coquinites became rarer. Ebbing storm surge currents entrained shells and sediment from the seafloor and carried them seawards, to where hollows and storm-generated channels acted as traps in which shells and disarticulated crinoidal material accumulated (Hiller & Theron. 1988), Generally. post-mortem transport of fossils was limited as indicated by the minimal EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA Vanrhynsdorp У дт Clanwilliam VA 26 Citrusdal V Аб 119 Prince Albert Ceres Worcester V Capetown А 24 Willowmore 7 Grahamstown iut ate iss wah | 5 Ladis ху Oudtshoorn i FIG. 4. Sketch map of South Africa indicating the localities from which Lower Devonian echinoderms are known. The 2 frames around the Ceres and Ladismith-Prince Albert districts are enlarged in Figs 5 and 6, respectively. Key to numbered localities outside the frames: 5 = Bucklands; 6 = Grootrivierhoogte; 11 = Platfontein; 14 = Kaaba/Alexandria; 15 = Nouga/Vredefort; 23 = Keurboomstrand; 24 = Warmwaterberg; 26 = Clanwilliam. mechanical damage to shells. Disarticulated shells are common but rarely display evidence of abrasion or breakage. Furthermore the vulnerability of the multiplated echinoderm skeletons to post-mortem disaggregation generally make preservation of whole crowns relatively rare. In the Gydo and Waboomberg Formations preservation of echinoderms still in life position, indicates predominantly gentle currents and sudden burial, perhaps by smothering mud clouds. Sudden influxes of fine sediment may have come either from the rivers feeding the deltas, or as a result of a storm generating a blanket of wave-stirred mud (Theron, 1972; Hiller & Theron, 1988). This environmental scenario is supported by high concentrations of brittle stars (adult as well as immature individuals), constituting lenticular ‘starfish beds’ in the Waboomberg mudstones in association with large numbers of infaunal bivalves preserved at a high angle to bedding with their umbones pointing upwards. Co-occurring fenestellid bryozoans, all with the apices of their cones directed upwards suggest overturning by gentle currents. In locally overlying beds well-preserved carpoids are associated with an ostracod fauna and with exquisitely preserved Lingula, which are not in life position (Ruta & Theron, 1997; Becker et al., 1994). The frequency of asterozoan trace fossils in the northern proximal shallow marine beds substantiate the original abundance of starfish and brittle stars in the marginal Bokkeveld seas. The relative rarity of asterozoan body fossils in the Bokkeveld sequence is therefore a reflection of the disintegration of their complex skeletons after death. Both ophiuroids and asteroids probably constituted quite an important component of the normal prevalent benthic marine biota of the Bokkeveld Group. Avallable data allow correlation (Boucot, 1971; Hiller & Theron, 1988) of several benthic invertebrate fossil communities with various depositional subenvironments in a delta complex: 1, a tidal flat community dominated by inarticulate brachiopods and infaunal bivalves inhabited the sheltered, back-barrier environ- ment; 2, the distributary mouth bar community dominated by brachiopods, occupied the relatively turbulent shallow water setting at the seaward edge of the delta platform; 3, the delta slope community was of intermediate aspect, dominated by brachiopods but more diverse with infaunal bivalves, gastropods, crinoids, cricoconariids and especially trilobites; 4, the shelf community, which although still dominated by brachiopods, is the most diverse community; brachiopods constitute «1/2 the diversity of the assemblage, with trilobites, bivalves and gastropods well represented and echinoderms, 120 MEMOIRS OF THE QUEENSLAND MUSEUM S £ 2 © о о | a FIG. 5. More detailed locality map of Ceres district as outlined by lefthand frame in Fig. 4. 1=Hottentotskloof; 2=Theronsberg Pass; 3=Tafelberg/Boplaas; 4=Matroosberg/Vredelus; 8=Hex River Pass; 9=Gydo Pass; 10=Die Vlakte; 12-Klipfontein/Lakenvlei; 13=Swaarmoed; 21=Eselfontein; 25=Ceres. Thick black lines are roads; thin crossed line (lower right) is a railway; thin black lines are streams. hyolithids, corals, bryozoans, conulariids and In the Gydo and Gamka Formations, which cephalopods comprising a significant proportion. constitute the oldest deltaic cycle, these communities are well-represented. Shelf and EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA 121 Prince Albert © o Et [ra с x E о о FIG. 6. More detailed locality map of Prince Albert-Ladismith districts as outlined by righthand frame in Fig. 4. 7 = Gamkapoort Dam; 16 = Vleiland; 17 =Koudeveld; 18 = Gamkaskloof; 19 = Bosluiskloof; 20 = Damascus; 22 = Vrischgewaagd. Thick black lines are roads; thin black lines are streams. slope communities are found in the Voorstehoek Formation and in western outcrops of the Waboomberg Formation, shelf community |cRINOIDS — — [ЕШ КЕ? assemblages have been identified as well. In — |opniocrinus stangeri Salter, 1856 a NH many of the other formations, available | Ophiocrinus sp. cf mariae (Kier, 1952) os m collections are generally too small for com- Corocrinus imbecillus Schmidt, 1941 S |a | munity analysis (Hiller & Theron, 1988). Mandelacrinus nelsoni gen. et sp. nov cfG|cf G FAUNAL AFFINITIES Monocyclic camerate indet = сај — | All Bokkeveld echinoderm specimens known |27022"? sp. = rae to the authors were included in the present study. | 2299075 interbrachiatus gen. et sp. nov | Affinities of the taxa identified (Table 1) are |Xopgferinus halbichi sp. nov. G T highly equivocal. One bias in this assessment is — | Monaldicrinus johni gen. et sp. nov cfG|efG| | the variation in levels of knowledge from other | Sacrinus gamkaensis gen. et sp. nov сга m parts ofthe world. The fauna of North Americas — |sacrinus hexensis gen. et sp. nov ча | | the best known and in general, most recently | Cradeocrinus plenarius sp. nov ala studied; in Europe comparably large faunas are Thalantocrinus arenacene sp. nov G known but many have not been given a modern eR PURIS On gr taxonomic treatment; South America and ae АКБ © E Australia have faunas of this age but they are DE ERASAN, — — LEGS poorly known (e.g. Australian crinoids (Jell, Aulacolatiaster breviramus gen. et sp. nov. | cf G [et G 1999) as demonstrated in this volume). Early Ulrichaster macrodentus gen. et sp. nov GG Devonian echinoderms from other parts of the Haughtonaster reedi Rilett, 197] ега сга | Е world are virtually unknown and not able to be | Hexuraster weitzi (Spencer, 1950а) ELDER D compared. At species level the South African Encrinaster tischbeinianus (Roemer, 1862)| S ] fauna has a crinoid and an asterozoan in common | Marginura hilleri sp. nov G Eugasterella africana sp. nov. |G | TABLE 1. Faunal affinities of the South African |57185 ohioensis Kesling & LeVasseur. 5 echinoderms dealt with herein compared with Europe РРР, 3 " | (1), North America (2), South Amietiea (3) А зене слее (йе, 171 S) Г Australia (4). С = ће genus occurs in that continent; | BLASTOIDS —- -— cf G =a closely related genus occurs in common; S= | Pachyblastus dicki Breimer & Macurda, 1972 dus Ze бреше» also occurs in that continent; апа Е = the сонна ootheizeni Breimer & G amily 1s ın common. dac Bkini = i 122 with Europe, a crinoid and an asterozoan with North America and a blastoid in common with South America. At the generic level most matches are with North America, then Europe, then a very few with South America and Australia. The 25 South African echinoderms dealt with herein do not provide a statistically large enough sample to make any compelling arguments and thus the affinities must be considered unknown at this stage. Brachiopod affinities (Boucot et al., 1969) place South Africa in the cool water Malvinokaffric Realm suggesting that further study of South American echinoderms may reveal closer affinities. SYSTEMATIC PALAEONTOLOGY The following abbreviations indicate repositories for the material discussed in the text:- British Museum of Natural History (BMNH), Geological Survey of South Africa, Pretoria (PRV), Geological Survey of South Africa, Bellville in Capetown (B), Roy Oosthuizen Collection, Zwartskraal, Prince Albert (RO), South African Museum, Capetown (SAM), Geological Collections, Stellenbosch University, South Africa (SU), Rhodes University, Grahamstown (RUGDNH), Natal Museum, Durban (NM) and Sedgwick Museum, Cambridge University, England (SM A). АП illustrations are of latex casts taken from external moulds unless otherwise stated; they are whitened with a sublimate of ammonium chloride for photography. Class CRINOIDEA Miller, 1821 Terminology follows Moore & Teichert (1978). Measurements are given as: length, parallel to the central axis; width, transverse to, but never cutting or joining the central axis; and depth, normal to, and may join central axis. MEMOIRS OF THE QUEENSLAND MUSEUM Subclass CAMERATA Wachsmuth & Springer, 1881 Order DIPLOBATHRIDA Moore & Laudon, 1943 Superfamily RHODOCRINITOIDEA Roemer, 1855 Family OPSIOCRINIDAE Kier,1952 Although Kier (1958) advocated elimination of the Opsiocrinidae after he recognised that Opsiocrinus was dicyclic, Ausich (1986) found the family useful in his classification of the Rhodocrinitoidea. Frest & Strimple (1981) and Ausich (1986) recognised that several of his familial characters are of generic standing in some cases in the same group. With this in mind the classification may be considered preliminary. Frest & Strimple (1981) and Ausich (1986) recognised the cofamilial relationship of Opsiocrinus and Ophiocrinus; we consider these genera synonyms (see below). We use Ausich's (1986) classification herein but following ICZN Article 40 the family name based on the junior synonym remains valid. Ophiocrinus Salter, 1856 TYPE SPECIES. Ophiocrinus stangeri Salter, 1856 from the Lower Devonian Bokkeveld Series, South Africa; by monotypy. DIAGNOSIS. Infrabasals 5, forming a pentagon completely or almost completely concealed by stem; interbrachials numerous, depressed, small, regular, especially in proximal part of interrays; CD interray conspicuous, with anitaxis of subquadrate anal plates variously developed in field of small irregular plates on either side. Arms 10 or 20, free and becoming cuneate then biserial distal to last arm division, unbranched distally. Stem circular or pentagonal in section, with narrow marginal crenularium, heteromorphic. REMARKS. The only significant differences between Opsiocrinus Kier, 1952 and Ophiocrinus appear to be the number of arms, length of anitaxis and cross section of the stem. Frest & Strimple (1981, table 1) showed that the2 genera FIG. 7. Ophiocrinus stangeri Salter, 1856. A, deformed crown with pinnulate biserial arms, RO39, х1. B, C-D interray view of crown showing 3 vertical columns of anal plates B4603, x2. C, small specimen showing uniserial arms well away from theca and biserial arm on left high up, B4544, x3. D, holotype crown with C-D interray on left, with Ist interprimibrach supporting 4 anal plates, with uniserial brachials proximally gradually becoming cuneate then biserial distally, SM A3441, *4. E, basal view of'small cup showing subpentagonal stem facet, from the Gydo Formation in a quarry on E side of road N from Prince Alfred’s Hamlet, Ikm S of Gydo, B4670, «5. Е, small deformed crown SAM13479, x2. G, two crowns compressed in the same direction, that on left with diminished crown length and that on right compressed laterally and showing the infrabasal circlet on top of the stem viewed from inside the cup. SAMK972, x1.25. EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA are distinguished only by the number of arms and in the nature of arm brachials. Although there is a reversal (clearly a typographical error) between their text and table 1 with Ophiocrinus being credited with cuneate brachials in the text but biserial ones in table |. Regardless of this confusion the larger collection of O. stangeri, now available, allows confirmation that the arms in both genera progress distally from rectangular to cuneate to biserial. Length of the anitaxis, with 4 plates in one genus and 2 in the other could well be considered a specific discriminator if consistent (1 CD interray is available for Ophiocrinus and 2 for Opsiocrinus). Some camcerate genera contain species with different numbers of arms. One specimen of О, stangeri (Fig. 2C) hasaray withonly 2 arms as opposed to the usual 4 in adjacent rays: a 2nd specimen (Fig. 1B) shows the beginnings of the radial interplate ridge patterns so prominent in some Opsiocrinus mariae Kier, 1952 (Kesling & Chilman, 1975, frontispiece, pl.40, figs 11-14): several specimens (Fig. 1A, 2A, ЗВ) have slight, barely visible re-entrants in the proximal margin of the basal plates which probably accomodated the corners of the infrabasal pentagon as in O. mariae. This indicates that orientation of the infrabasal pentagon to the basal circlet changed from the exterior to the interior as described by Kier (1958, fig. 1) in O. mariae because the interior of the cup in O. stangeri (Fig. 3E) shows the infrabasal pentagon with the angles at the sutures between basals and the sutures between infrabasals at the centre of the basals. The stem of the North American species is inferred to be pentagonal because of the shape of the attachment facet on the cup. In O. stangeri the stem is round but in at least 1 specimen (Fig. 1G) the attachment facet or the Ist columnal is pentagonal. Thus we synonymise these 2 genera despite their geographic and stratigraphic (Early Devonian vs Middle Devonian) separations. Ausich (1986:87) inferred 2 lineages within the Opsiocrinidae from the Llandovery into the Devonianwiththe 2 Devonian genera on separate lineages based on cup shape. prominence of ridges and arm numbers. However, cup shape and ridge pattern are identical in the 2 Devonian genera and taking the rudimentary ridge pattern in a specimen of Ophiocrinus even the interbrachial ornament may be allied. The relation of the infrabasal circlet to the basal circlet is another feature which seems to join the 2 Devonian genera. Other monophyletic camerate genera are known to have 10 and 20 armed MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 8. Ophiocrinus stangeri Salter, 1856, plate diagram showing stem facet as dashed circle, radials black and only 2 incomplete arms from half of one ray; posterior interray at 12 o'clock. members and we suggest that Ophiocrinus and Opsiocrinus constitute 1 genus and belong to | lineage rather than 2 that had been separate since the Early Silurian (Jell, 1999, fig. 2). Ophiocrinus stangeri Salter, 1856 (Figs 7-10) Ophiocrinus stangeri Salter, 1856:223, pl.25, lig.20; Ubaghs. 1978: 1428. fig. 238.2. MATERIAL. HOLOTYPE: SMA3441 from De Dooms, Hex Rivier Poort (donated to the Sedgwick Museum, Cambridge in 1932 from the collection of Dr W. Stanger. B4523 from Bucklands, eastern Cape Province(C281), B4526-4530 from Gamkapoort Dam (C281), B4544, B4553, B4603 trom Gydo Pass (C281), B4551, B4552 {rom Gamkapoort Dam (C281), RUGDNHI from Klein Kaaba, Alexandria district, eastern Cape (Voorstehoek Formation), B4579; RO S17, RO S20 from Grootrivier, EARL л Y DEVONIAN ECHINODERMS OF SOUTH AFRICA 12 FIG.9. Ophioerinusstaugeri Salter, 1856. A. crown showing B and C rays, RO S20. х5. B. small crown with arms on anterior (or A ray) side missing to expose anal tube, K 4532, х4. C, large crown showing some normal arms (onright) but mostly arms that have been broken off during life and regenerated producing major change in arm diameter, B4553, 42.5. D. crown on long stem suggesting high level feeder, SAMI3459. х1, 126 MEMOIRS OF THE QUEENSLAND MUSEUM MBA etenim у tmn | reser н fepe qa Mr AEA AVA He TES Е = = z А " 27-4 чанач Pas VIG. 10, Ophiocrinus stangeri Salter. 1856. A, small crown showing changing brachial type B4544A, х5. В. small crown SAMK973, х2. C, poorly preserved crown from most northerly outcrops in Calvinia district, B4559, x5. D. crown RO S17, *2.5. E, interior of crown showing 5 infrabasals. RUGDNHI. *2.5. EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA Ceres at 32?38'S, 19?24'15"E (C2S1), RO39 from Gamkapoort at 33° 18'S, 21? 38'E (С251), SAMK967, 969, 970, 972, 973, 975 from Wolfaardt’s Farm, Riet Valley, Ceres, SAM9703 from Clanwilliam, Cape Province, SAM13459 from Boschluis Kloof, SAM 13462, 13464, 13479 from Koudeveld Berg, above 2nd sandstone. DESCRIPTION. Crown up to 60mm long, subcylindrical, with arms about 3 times as long as cup. Cup bowl-shaped, up to 25mm maximum diameter, of moderate length, with smooth plates. Infrabasals 5, small, equal, diamond-shaped, concealed by stem. Basals 5, 7- sided, longer than wide, with greatest width in proximal 1/2, rarely with depressed and weakly isolated lateral tips beneath tips of radial (e.g., in holotype), often with slight re-entrant in proximal side presumably accomodating angle ofthe infrabasal pentagon. Radials 5, pentagonal, not in contact with other radials, penetrating deeply into basal circlet; radial facet plenary. First primibrach hex- agonal, wider than long; 2nd primibrach axillary, pentagonal; secundibrachs 2 or rarely 3, fixed, subquadrate; tertibrachs free, becoming triangu- lar distally. Arms 20 (rarely 18), uniserial proximally, becoming almost biserial distally, pinnulate with pinnules on long side of triangular tertibrachs alternating from side to side on suc- cessive plates, with well-developed oral groove. CD interray with 2 large hexagonal plates forming central anal column flanked by smaller polygonal anal plates; primanal hexagonal, in radial circlet, similar in size to radials. Inter- primibrachs small, numerous in each ray; proximal 1 septagonal, resting on basals; 2 in second row, 3 in next row, increasing in number and decreasing in size distally; interbrachial field decreasing in width adjacent to secundibrachs as arms spread laterally. Intersecundibrachs in larger specimens, | in first row, 2-3 in more distal rows. Tegmen unknown. Stem circular, hetero- morphic, noditaxis N212 proximally with nodals longer and of greater diameter than internodals, distally with noditaxis N3231323, with inter- columnal sutures strongly crenulate, greatest length among available specimens 15cm (uniform diameter of 2mm), greatest diameter 4mm. REMARKS. Among available specimens only the holotype shows the CD interray so we have no data on possible variation in this area of the cup, nor any data on the tegmen which is concealed by the arms. One specimen (Fig. 9B), here tentative- ly assigned to O. stangeri, has a long anal tube, about 1/2 length of arms and covered by small polygonal plates; the tegmen of this common species should be sought in future collecting to confirm this feature. The unbranched, free, pin- nulate arms in which the brachials become more and more cuneate distally together with a frag- ment of stem identical with that of O. stangeri lying beside the cup suggest assignment to O. stangeri; negative evidence is provided by the lack of any other species with this type of arm in the rest of the South African Bokkeveld crinoids available at present. We make the assignment tentative in the absence of detail of the cup. Some intraspecific variation is recorded in the description above with other variation: 1, in a few specimens (Fig. 7C, F) the distal noditaxis extends much further proximally i.e. proximal stem resembles distal stem of most specimens; 2, in some specimens, particularly smaller ones (c.7mm cup diameter) proximal tertibrachs are more quadrate than rectangular in lateral view, becoming cuneate distally; 3, intersecundibrachs are not present in these smaller or even some larger specimens; 4, in one specimen (Fig. 7B) the threeway intersections of sutures in the interprimibrach and intersecundibrach areas are depressed, suggesting a subtle interplate ridge system as in Opsiocrinus. At species level this taxon is unmistakeable and distinct from the most similar O. mariae Kier,1952 in number of arms, depression and ornament in interbrachial areas and CD interray plating. O. stangeri is the most common crinoid in available collections from the Bokkeveld Series but is still incompletely known. Ophiocrinus sp. cf. mariae Kier, 1952 (Fig. 11) MATERIAL. A cup RO179 from Cockscomb Mountains, Steytlerville, near Bucklands at 33?31'S, 24?44'30"E in the Gydo Formation. DESCRIPTION. Infrabasal circlet just visible laterally, with sutures between infrabasals directed at midline of basals, angles of infrabasals directed at sutures between basals; strong ray ridges elevated, occupying most of plate width from basals up to at least 2nd secundibrach; 10 arms; interprimibrach series 1-2-3 decreasing in size at level of arm branch, with strong radial ornament of central tubercle and radial lines sometimes consisting of a series of tubercles, with 7 radial ridges on the Ist interprimibrach, with 6 ridges on plates of next row; 1-2 intersecundibrachs with 5-rayed ornament. REMARKS, This specimen may represent an undescribed species but it resembles O. mariae Kier,1952 more than it does the South African O. 128 FIG. 11. Ophiocrinus sp. cf. О. mariae (Kier.1952) partial cup showing depressed interbrachials with stellate ornament, RO179, х5. stangeri. Xs 10 arms, 2nd primibrach axillary, strongly depressed interbrachial plates with stellate ornament, and strongly depressed 3-way sutural junctions in the cup are features of O. mariae but the laterally visible infrabasals, ray ridges standing high on the arm plates and 7-rayed ornament on the Ist interprimibrach differ from that species. It is simply distinguished from O. stangeri by its 10 rather than 20 arms and the ornament on the interprimibrachs. Without knowledge of the posterior. stem or higher arms we prefer to make tentative assignment only. Order MONOBATHRIDA Moore & Laudon. 1943 Suborder COMPSOCRININA Ubaghs.1978 Superfamily PERIECHOCRINOIDEA Втопп, 1849 Family PERIECHOCRINIDAE Bronn, 1849 Corocrinus Goldring.1923. TYPE SPECIES. Corocrinus ornatus Goldring,1923 from the Middle Devonian Ludlowville Shale, New York; by original designation. MEMOIRS OF THE QUEENSLAND MUSEUM Corocrinus imbecillus Schmidt, 1941 (Figs 12) Corocrinus imbecillus Schmidt, 1941: 97, pl. 14, figs 3,4. MATERIAL. HOLOTYPE: E3142 in the Museum für Naturkunde, Berlin, from the Lower Devonian, Upper Koblenz Shale, western Germany. South African material ROC25, ROT17 and B4551 all from Gamkapoort Dam in the Gydo Formation. DIAGNOSIS. Like type species but with 20 arms. DESCRIPTION. Cup high conical. up to 20mm maximum diameter. of moderate height: plates with strong radial ornament, with high ray ridges; ray ridges wide on basals, becoming higher relative to the plates and more narrowly arched in section above the axillary primibrach. Basals 3, sutures in B and E rays and CD interray, equal, pentagonal. visible in lateral view, with 3 prominent ridges radiating distally. Radials 5. hexagonal (A.C.D) or heptagonal (B.E). in contact with other radials except in CD interray. slightly longer than wide, with 6 strong radial ridges including ray ridge. First primibrach hexagonal, longer than wide, with 6 radial ridges: 2nd primibrach axillary, pentagonal, with 5 radiating ridges including the Y-shaped ray ridge. Secundibrachs elongate, fixed, with 2nd or 3rd axillary, (most often both axillary in 1 arm but any pattern not available). Tertibrachs free, uniserial, with interbrachial sutures becoming oblique distal to 2nd or 3rd brachial but brachials never triangular. Arms 20, uniserial, strongly pinnulate; pinnules long (up to lcm long in available specimens), of 8 or more pinnulars, 1 per brachial, alternating from side to side up arm. CD interray wide, with ridge up median line of plates (Fig. 12A); primanal large, heptagonal, in radialcirclet, similarin size to radials. supporting 3 hexagonal anals distally. Interprimibrach series elongate, with proximal one hexagonal. resting on 2 radials. supporting 2 smaller hexagonal interprimibrachs in next row, all with 6 radial ridges: more distal rows of smaller plates with central tubercules and radial ridges. Inter- secundibrachs numerous, | in proximal row. small, centrally tuberculate. Anal sac (Fig. 12C) of many small polygonal abutting plates. about as long as cup, possibly flexible. Stem circular, heteromorphic, with noditaxis of N212 pattern: intercolumnal sutures strongly crenulate: greatest length among available specimens 90mm. diameter 3mm proximally. 2mm distally. EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA FIG. 12. Corocrinus imbecillus Schmidt, 1941. A, crown, RO T17, x3. B, internal mould of crown, B4523, x3. C, crown showing arm branching and part of the anal tube curving to right, B4551, х4. D, crown showing pinnulate arms, RO C25, х2. REMARKS. Breimer (1962) noted the close similarity of this species to the type species and we can only make species distinction on the number of arms: none of Goldring’s (1923) material of the type species has any free arms attached so that if a further bifurcation occurred in the free arms it would not be available on her material. We therefore, reserve the possibility that C. ornatus and C. imbecillus could be synonymous. Breimer (1962) placed weight on the axillary primibrach having 7 sides: externally 129 the axillary primibrach of C. imbecillus is not clear on Schmidt's (1941) material or on the South African specimens but the internal mould (Fig. 12B) shows the shape of cup plates very well and the axillary primibrach 15 7-sided, agreeing with Ubaghs diagnosis. Mandelacrinus gen. nov. TYPE SPECIES. Mandelacrinus nelsoni sp. nov. ETYMOLOGY. For Nelson Mandela, President of South Africa. DIAGNOSIS. Cup low conical; plates smooth, with margins depressed in interadii, with strong wide ray ridges leaving narrow depressed marginal zone on fixed brachials. Basals 3, sutures in B ray, CD interray (and presumably E ray). Radials hexagonal, laterally in contact except in CD interray. First primibrach hexagonal; 2nd primibrach axillary, pentagonal. CD interrary wider than other interrays, primanal interrupting radial circlet, supporting 3 hexagonal anals in 2nd row, 5 in 3rd row, 6 in 4th row at level of 1st secundibrach then diminishing on to tegmen. Interprimibrachs small with central tubercle distal from 3rd row. Intersecundibrachs small, tuberculate. Arms 20, biserial distal to 2nd tertibrach, pinnulate; pinnules long, slender, a row along either side of arm, | per brachial. Stem circular, of alternating nodals and internodals proximally. REMARKS. Three basals, separation of C and D radials by the primanal, other radials in contact and hexagonal Ist primibrach suggest the Perie- chocrinidae but the single fixed secundibrach is not a feature of that family. However, co-familial Corocrinus has only a few fixed secundibrachs and makes a reasonable ancestor for the new genus. The shorter cup, wider interradii, less ornamented cup, biserial arms and single fixed secundibrach may all be considered derived character states relative to Corocrinus. Mandelacrinus nelsoni sp. nov. (Fig. 13, 14) MATERIAL, HOLOTYPE: RO740 from the Cockscomb Mountains, Steytlerville, near Bucklands at 33?3l'S, 24°44°30"Е in the Gydo Formation. PARATYPE: SAMK976 from Wolfaardt's Farm, Riet Valley, Ceres.. DIAGNOSIS. As for genus. DESCRIPTION. Cup 6-13mm long, low conical, up to 15mm diameter; plates smooth, with depressed margins, with well-developed ray ridges standing well above lateral parts of plates particularly about the secundibrach level where arms begin to stand away from cup. Basals 3, sutures in B and E rays and CD interray, equal, pentagonal, visible in lateral view. Radials 5, hexagonal, in contact with other radials except in CD interray, wider than long in holotype, longer than wide in small specimen. First primibrach hexagonal, wider than long; 2nd primibrach axillary, pentagonal; Ist secundibrach fixed, hexagonal, with flat lateral areas rising up strongly to high ray ridge; 2nd secundibrach axillary, pentagonal; tertibrachs free, biserial MEMOIRS OF THE QUEENSLAND MUSEUM \ Ñ N [| f H i j a NN We, AX E N /] S NN: My ON // f Ж) FIG. 13. Mandelacrinus nelsoni gen. et sp. nov., sketch of plate arrangement with radials darkened; anterior - posterior axis in NW-SE line; most arms shown as far distally as 1st tertibrachs with distal portions (not including tips) sketched for 2 arms on left of B ray. from 2nd. Arms 20, biserial from 2nd tertibrach, strongly pinnulate; pinnules long (up to Іст long in smaller specimen), of 7-8 pinnulars, 1 per 1/2 brachial in 2 rows along arm. CD interray wide, with ridge up median line of plates (Fig. 6B); primanal large, heptagonal, in radial circlet, similar in size to radials, supporting 3 hexagonal anals distally, followed by rows of 5 then 6 anal plates, then decreasing in size and number between maximum width of fixed arms. Interprimibrach series narrow, elongate; lst interprimibrach hexagonal, resting on 2 radials, supporting 2 large hexagonal interprimibrachs distally; 3rd row of 3 small centrally tuberculate plates and 4th row same but with 5 plates. Intersecundibrachs 1 or 3, 1 in proximal row, centrally tuberculate, 2 in distal row. Tegmen unknown. Stem circular, up to 4mm in diameter, heteromorphic proximally becoming uniform distally, noditaxis uncertain but probably N212; intercolumnal sutures strongly crenulate; greatest length available 3cm. EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA REMARKS. Of the 2 specimens 1 is twice the size of the other and both show the posterior of the cup to confirm that they belong to the same species. There do not seem to be any other genera closely comparable as mentioned above. Monocyclic camerate indet. (Fig. 15) MATERIAL. РКУ HR-5 trom the Hex River area. 131 FIG. 14, Madelacrinus nelsoni gen. et sp. nov. A, holotypecrown in C ray view with C-D interray to left. RO 740, x3. B, smaller crown in C-D interray view, B976, x3. DESCRIPTION. Cup probably conical. about 8mm long; plates smooth or with very subtle vermiform ornament. Basals apparently 5. pentagonal. Radials 5, largest plates in cup. 7-sided: 151 primibrach hexagonal; 2nd primibrach axillary: interprimibrachs and inter- secundibrachs normal polygonal plates decreasing in size distally. Arms 10, uniserial. of cuneate brachials, highly pinnulate: pinnules MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 15. Monocyclic camerate indet. Crown showing long pinnules and triangular brachials, PRV HR-5, х4. alternating from side to side on long side of brachials. REMARKS. Without details of the posterior it is not possible to classify this specimen, but the large radials suggest an advanced compsocrinine group suchas the Carpocrinidae and comparison, with for example Acacocrinus Wachsmuth & Springer1897 (see Ubaghs, 1978, fig. 272.2). show complete correspondence in all features available from the South African specimen, Ausich (1987) transferred Acacocrinus to the Periechocrinidae but considered it a likely ancestor of the Carpocrinidae. Identification of the South African species must await further material. Superfamily HEXACRINITOIDEA Wachsmuth & Ѕргіпрет,1885 Family HEXACRINITIDAE Wachsmuth & Springer.1885 Arthroacantha Williams, 1883 TYPE SPECIES. Arthroacantha ithacensis Williams, 1883 from the Upper Devonian of New York; by original designation. Arthroacantha? sp. (Fig. 16) MATERIAL. One set of 5 incomplete arms RO128 (part and counterpart) from Gamkapoort, Prince Albert (33°18°S, 21?38 E) in the Gydo Formation. DESCRIPTION. Arms biserial, pinnulate. occasionally with pair of adjacent 1/2 brachials extending into strong lateral spines. occasionally with other pairs of brachials in between spinose ones bearing circular facets; pinnules long, slender, with wide groove on inner surface: adoral groove on arm moderately deep, U-shaped in section; each brachial with concave pinnular facet. REMARKS. Identity of thes arms is very doubtful but among known spinose genera Arthroacantha appears to have the most similar organisation (Kesling & Chilman, 1975. pl. 135) with periodic pairs of strong spines of the same dimension and arrangement as in our specimen. However. our arms do not appearto branch at the spinose brachials which is usual in Arthroacantha. Nevertheless. Stewart (1940:56) noted that in a specimen from the Silica Shale at Silica, Ohio large strong tubercles are irregularly developed between the bifurcations. Goldring (1923: 290) noted a subspinose tubercle on each EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA 1/2 brachial above the axillary secundibrach in А. carpenteri Hinde, 1885. There is considerable variation in the arms of Arthroacantha so our specimen fits within the range. However. the assignment must be extremely tentative without knowledge of the cup: an alternative is that these arms may belong to a new crinoid genus. Subclass CLADIDA Moore & Laudon. 1943 Order DENDROCRINIDA Bather, 1899 McIntosh (1983. 1986) discussed phylogeny and classification of dicyclic cladid inadunates and concluded that the 2 suborders of Moore & Laudon (1943) and 3 suborders of Moore et al. (1978) need major review. Until that review is complete we retain all the cladids in this paper in one broad group. Family DENDROCRINIDAE Wachsmuth & Springer.1886 Eckidocrinus gen. nov. ТҮРЕ SPECIES. Eckidocrinus interbrachiatus sp. nov. ETYMOLOGY. Ananagram from the surname of Mr Roy Dick, who contributed much material to this study. DIAGNOSIS. Basals hexagonal, with broad low radial ridge ornament. Radial facet 71/2 radial width, gently declivate. concave. Inter- primibrach depressed. filling interradial space between primibrachs. Arms with 2 main rami per ray, with bilateral heterotomous branching; 3rd primibrach axillary; strong ramules alternating side to side along each arm from every 3rd or 4th brachial, cach ramule dividing at least twice. Stem circular, large diameter, heteromorphic. REMARKS. Among early Palaeozoic cladids bilaterally heterotomous arm branching is not widespread. Although not restricted to that family, the arm branching pattern resembles fairly closely that of the Ordovician dendrocrinid Grenprisia Moore, 1962 whichalso has brachials shaped the same as the new genus and small interprimibrachs. Family assignment is made on these tentative grounds and a specimen exhibiting the posterior of the cup will be necessary to be more definite. With available morphology it is established as a separate genus. If the similarities to Grenprisia are indicative of relationship it must be very distant as they are well separated in time but no other known cladid has the combination of features mentioned above. FIG. 16. Arthroacantha? sp., set of biserial arms with long pinnulesand paired spines, RO 128a andb, * 1.5. Eckidocrinus interbrachiatus sp. nov. (Fig. 17) MATERIAL. HOLOTYPE: B4534. PARATYPE: B4554 from the Voorstehoek Formation, at Matroosberg, Stinkfontein, Hex River Pass. DIAGNOSIS. As for genus. DESCRIPTION. Cup broadly conical (holotype 16mm max. diameter), short (holotype 9mm long): plates smooth, with broad low radial ridges and depressed corners in most parts of cup. Infrabasals 5, visible laterally, pentagonal, short. wider than long. Basals 5. largest plate in cup. hexagonal, with broad low ridges from infrabasals to radials and similar transverse ridge to adjoining basals. distal tip of basals depressed as are lateral parts, ridges less obvious at infra- basal to basal sutures. Radials 5. pentagonal, with scallop of radial facet in upper margin, with ridges running from basals towards facet and another similar ridge running laterally around cup near distal edge of radial circlet; radial facet >1/2 width of radials, a little less than semicircular, declivate, concave. Primibrachs 3. 2 subquadrate, wider than long; 3rd primibrach axillary. pentagonal. Interprimibrachs filling interradial notch to approximately base of 3rd VIG. 17. Eckidocrinus interbrachiatus gen. et sp. nov. A, set of arms viewed from inside, B4554, «1. B, holotype crown showing uniserial repeatedly branching arms, B4534, х2. MEMOIRS OF THE QUEENSLAND MUSEUM primibrach plate, 3 larger plates resting on radials, becoming smaller distally. Arms bilaterally heterotomous, with 2 main arms per гау. with strong ramules (at least 6 per ramus) alternating from side to side along each arm at about every 3rd or 4th brachial. with ramuli dividing at least twice; full extent of arms not clear: ventral groove deep. broadly V-shaped in section, with large cover plate series, about 3 or 4 per brachial. Anal area of cup not available: anal sac longer than arms. straight, of small strongly ornamented plates. apparently perforated. Stem circular, heteromorphic. with noditaxis 3231323. REMARKS. Part of the anal sac is preserved at the right of the arms in one specimen (Fig. 17A). The anal sac appears to have been long and composed of a few columns unornamented plates. Family CYATHOCRINITIDAE Bassler, 1938 Kopficrinus Goldring, 1954 TYPE SPECIES. Kopficrinus pustuliferus Goldring, 1954 from the Lower Devonian of New York; by original designation. REMARKS. Goldring's genus, for which the anal structure has been unknown except for the anal X supporting 3 small anals, was assigned to the Gasterocomidae by Moore et al. (1978) which family lacks an anal tube and has the anal opening through the side of the cup below the posterior radials. Thus the anal opening somewhere above the cup in the type species does not fit the family concept. The second species of the genus. described herein, with a long anal tube also argues against assignment to Gasterocomidae. We assign the genus to the Cyathocrinitidae based on the symmetrical anal plating, anal tube. the small pentagonal infrabasal circlet and the enlarged posterior basal. Kopficrinus halbichi sp. nov. (Fig. 18) Ophiurites sp. Rossouw, 1933: 75, fig.2. ETYMOLOGY. For Prof. LW. Halbich, Stellenbosch University, who kindly made this material available. MATERIAL. HOLOTYPE: SUK466. PARATYPE: SUK464 (2 specimens on one piece of rock (1 drawn by Rossouw, 1933), from the first shale band on Vredenhot, Prince Albert (locality quoted by Rossouw (1933) in the Gydo Formation. DIAGNOSIS. Infrabasal circlet apparently fused: arms atomous (or with at least 14 EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA 135 primibrachs): anal sac of 4 columns of hexagonal sutured plates. DESCRIPTION. Cup low. bowl-shaped, smooth. Infrabasal circlet pentagonal, apparently fused into a single plate, most of circlet visible in side view. Basals 5, pentagonal except for hexagonal posterior basal. Radials 5, with narrow angustary and declivate horseshoe-shaped facets, without axial canal separate from ventral groove. Anal X in radial circlet, slightly proximal to adjacent radials, resting symmetrically on posterior basal, with horizontal distal margin supporting 3 small anal plates at base of tall anal tube. Anal tube of 4 columns of close-packed hexagonal plates (distal end and anal opening not clear). Arms atomous (longest section available 14 brachials long). Stem circular in section. with vaguely pentalobate lumen. REMARKS. Rossouw (1933. fig.2) gave a line drawing only of the specimen he identified as an ophiuroid and which is here designated a paratype of this new crinoid species. The line drawing was inaccurate in several details which are corrected here with a photograph (Fig. 18C.D): for example the specimen shows 3 of the 4 columns of plates in the anal tube whereas Rossouw's figure shows only 2. The latex cast from the counterpart of Rossouw's specimen gives the oral aspect, with the mouth area unclear: oral plates are not clear but may be assumed to have been present from the edges of the radials. This species is assigned to Kopficrinus based on the matching posterior plating of the cup, the radial facets, the large round brachials and in particular the apparently atomous arms. Distal tip of any arm is not available in either the type or the new species but in both, arm sections are known with 13 or 14 brachials. Thus it is uncertain if the arms are atomous or not but it seems unlikely, from comparison with other crinoids. that these arms would divide for the first time so far from the cup; we consider the arms were most likely atomous. FIG. 18. Kopficrinus halbichi sp. nov. A, cup showing infrabasal circlet and anal tube (12 o'clock), 506464. x4. B, cup SUG464-1, х4. C-D, oral and basal views of holotype cup SUG466a and b, «3.5. E, sketch of plate arrangement with infrabasals shown as circlet with size of stem facet and section of central canal indicated. IB = infrabasal; B = basal; R = radial; X = anal X; A-E above radials indicate rays. Family EUSPIROCRINIDAE Bather, 1890 Monaldicrinus gen. nov. TYPE SPECIES. Monaldicrinus joluii sp. nov. ETYMOLOGY. An anagram for John Almond, Geological Survey of South Africa, Capetown, who greatly assisted us during this project. DIAGNOSIS. Cup low. strongly flared laterally. Infrabasals 5. Basals 5. Posterior basal heptagonal. supporting 2 almost equally sized anal plates distally. C radial virtually symmetrical distal to BC basal. Anal sac large and long, with opening at tip. with main column of plates both externally and adaxially. Arms 10. uniserial, stout, dividing isotomously once, ramulate; ramules alternating from side to side on every 5th brachial, dividing isotomously on 6th terlibrach. Stem circular in section. small diameter for the family, with moderately large lumen vaguely pentalobate in section. REMARKS. This genus is assigned to the Euspirocrinidae on the arrangement of anal plates in the cup which arrangement compares closely with Parisocrinus Wachsmuth & Springer.1880 and 4mpheristocrinus Hall.1879, on the nonpinnulate arms, on the 5 infrabasals and basals, onthe ambulacral groove cover plates and on the circular stem. It is distinctive in its arms branching only once isotomously. in being ramulate and their arrangement, in the nature of the large anal sac, in the low flared shape of the cup which does approach lasocrinus Lyon, 1857 and in the relative diameter of arm to stem. The anal plate arrangement is also found in Poteriocrinites (see Moore et al.,1978, fig.394.7) but that genus of the Poteriocrinina has pinnulate arms which is used by Moore ef al. (1978) as a subordinal discriminator The arm structure of Monaldicrinus occurs in the Barycrinidae but that family has a distinctly different anal plate arrangement. However, in discussing the Barycrinidae, Moore & Laudon (1943:40) considered the single isotomous branching followed by development of alternating ramules (their branchlets) to be a primitive stage of development leading to pinnulation. They also considered the transverse ridge on the articular facet, seen іп Afonaldicrinus, an advanced feature. It is tempting to suggest that perhaps Monaldicrinus from the Malvinokaffric Province was an advanced cyathocrininid ancestral to the Poteriocrinina. MEMOIRS OF THE QUEENSLAND MUSEUM Monaldicrinus johni sp. nov. (Figs 19-21) MATERIAL. HOLOTYPE: SAMK977, an extemal mould from 100m N of the house at Wolfaardt’s Farm, Riet Valley, near Ceres, PARATYPES: B4579 from Hex River Pass, 1.3 miles from Great Swaarmoed Farm; SAM3947 from Laken Vlei, Ceres (1st Shale), RO202 and 810 from Gamkapoort, Prince Albert, 33°18°S, 21?38 E (C281). DIAGNOSIS. As for genus. DESCRIPTION. Cup smooth, small relative to size of crown, 15mm long. conical. Arms apparently elevated only 10-20? above horizontal. Infrabasals 5, pentagonal, uniform in size. almost completely visible and as long as wide in side view. Basals 5. hexagonal except for heptagonal BC and CD basals. base against infrabasals with obtuse angle (7150?) at central junction with suture between infrabasals. Radials 5. large, wider than long, heptagonal: radial facet only slightly declined outwards, peneplenary. Facet on Ist primibrach with strong transverse ridge broken by central gap, with large circular canal just beneath broadly V-shaped ambulacral canal. Circularcanal migrating through the gap in transverse ridge by 5th primibrach to be close to outer edge of facet and well removed from sharply V-shaped ambulacral groove. Ambulacral groove covered by thick cover plates; cover plates more numerous than brachials (c. 8 to 5) but exact correspondence unclear. CD basal supporting hexagonal anal X and pentagonal radianal; radianal resting symmetrically on BC and CD basals, supporting the right tube plate (3rd anal plate in cup) adjacent to the C ray. Anal sac large, with terminal aperture, with numerous (710) vertical columns of poly gonal plates; some columns with plates having scalloped lateral margins (scallops never on proximal or distal margins) with perforations through wall at each scallop: perforations most prominent near base disappearing before (1р; main column of larger plates on outer side distal to anal X and on inner side rising from oral area; main column on inner side with short stubby spine usually on every 2nd plate but irregular and sometimes on consecutive plates; such spines less frequent and more irregularon some other columns on innerside but not on outer side of sac. Arms 10, each ray with 1 isotomous branching at axillary primibrach 8 (B4579)-12(RO810). Above this 1 specimen (RO202) with ramules developed alternately on each 5th secundibrach: 1st ramule on outside of ray; ramule branching isotomously at least twice 37 EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA 1. FIG 19.Monaldicrinusjohni gen. et sp. nov. A.C.E; RO $10. A, cup х2. C, cup showing stout uniserial arm, х 1.2. E, distal face of first primibrach *6. B, part of an arm showing ambulacral cover plates B4549. x6. D, holotype SAM K977, x2, each time at 5th brachial, circular in section. with moderately large lumen vaguely shorter than 4 secundibrachs. standing up pentalobate in section, vertically off the arm. Stem circular in section. А | small diameter for the family and relative to cup, REMARKS. The 4 specimens of the cup have the ` arms splayed out horizontally or almost so, mostly in the bedding plane. Also in each is à degree of dislocation and dislodgement along sutures and fracture across plates within the cup. These taphonomic features make it difficult to interpret original cup shape and arm attitude. In so far as the plates arc in contact around the cup at the radial circlet we could interpret minor flattening of the cup. However, inFig. 19D where the anal sac is assumed to have been vertical in life but is now horizontal. the dislocation of plates near its base is minimal and certainly does not suggest such a change in attitude after death. We therefore, have no reason to believe the horizontal arm position reflects the original attitude. While it is more likely, by comparison with other cladids, that the arms were erect. the much smaller stem diameter, relative to crown size in Monaldicrinus which is a fairly large crown among crinoids in general may suggest either 1, it lived in quiet water and so did not need strong attachment in currents (unlikely given the enclosing clastic sediments but nevertheless the animal is found in the finer shale layers and may have migrated with thc environment) or 2, that the arms were less erect and thus less of a baffle to currents. What ever the answer, we consider that the arms were probably not completely erect but that the ramules coming off them were probably vertical. In the 3 specimens with 4 rays preserved the A ray is missing and the plating near its base is unclear. In Fig. 19D there is the suggestion that it may lie on the basals but with only point contact to the other radials rather than a sutured junction of some length. We cannot be certain of this point due to preservation but if so this would provide a strong accentuation of the A ray - CD interray line of symmetry. The large specimen of the oral side of the arms and anal sac (Fig. 21C) is assigned to this species on the nature ofthe anal sac. size and nature of the arms and their branching pattern. Although very difficult to see on the latexes. ramules are identified on the mould of Fig. 21B (because of the penetration of limonite) at every 5th MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 20. Monaldicrinus johni gen. et sp. nov. sketch of plate arrangement (infrabasals shown as circlet with size of stem facet and section of central canal indicated) and arm branching (on right). IBB = infrabasal circlet, В = basal; R = radial; RA = radianal; X = anal X; rt = right tube plate; A-E above radials indicate rays. secundibrach. The prominent tubercles on that specimen which are lacking on other specimens are interpreted as only occurring on the oral side of the anal sac and thus not evident on the other specimens of the outer side. Without moulds of the 2 sides from the | individual this cannot be confirmed. Until it is determined objectively we remain confident of our identification. Family THALAMOCRINIDAE Miller & Gurley. 1895 Following McIntosh & Brett (1988) this family name replaces Bactrocrinitidae Jackel, 1918 as used by McIntosh (1979; 1983). Sacrinus is assigned to this family based on its affinity to Follicrinus which was assigned to the Bactrocrinitidae by McIntosh (1979. 1983) and althoughexcluded by McIntosh & Brett (1988) is retained in the family herein. Family level subdivision of the cladids remains in flux so we adopt a broad family concept here. We acknowledge that quadrangular vs. pentagonal radianal plate could indicate entirely separate lineagesbut this has not been demonstrated and is not applied herein. Sacrinus gen. nov. TYPE SPECIES. Sacrinus gamkaensis sp. nov. ETYMOLOGY, Sa- for South Africa. DIAGNOSIS. Cup conical: infrabasals approximately same length (or only slightly shorter) as basals; radials wider than long, with angustary radial facets projecting laterally. Anal plates in cup 3, with broad low radial ridge VIG. 21, A-C, Monaldicrinus johni gen. et sp. nov. A, crown splayed on bedding plane in proximal view, SAM3947, x1. B, crown showing anal tube on right B4579, x2, C, set of arms showing cover plates and widely spaced ramules and large anal tube RO 202, x1. D, stem indet.. section of distinctive indeterminate stem in lateral view RO 295a, х4. 139 EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA MEMOIRS OF THE QUEENSLAND MUSEUM 140 EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA FIG. 23. Sacrinus gamkaensis gen. et sp. nov., sketch of plate arrangement and arm branching (on right) with distal branching shown on only one branch but symmetrical on each other branch. IB = infrabasal; В = basal; R = radial; RA = radianal; X = anal X;rt = right tube plate; A-E above radials indicate rays. ornament; radianal pentagonal, proximal and left of C radial, directly supporting right tube plate: anal X not much larger than radianal; anal tube large. inflated, irregularly shaped. of many very small polvgonal plates irregularly arranged. Arms robust, branching isotomously 3 or4 times. Stem pentagonal to subrounded in section, heteromorphic. REMARKS. The major distinguishing feature of this new genus and of Fo/licrinus Schmidt, 1934 (type Zaxocrinus? grebei Follmann, 1887) from the Lower Devonian of Germany is the irregularly shaped, inflated anal sac of small irregularly arranged stellate plates. McIntosh (1983) queried whether this feature alone should carry generic status, particularly in the absence of knowledge of the anal sac in Bactrocrinites fusiformis Roemer, 1844, the type of Bactrocrinites. In Follicrinus the radianal is quadrate. there are only 2 anal plates in the cup. the radial facets may be interpreted as plenary (as by Moore et al. (1978) who clearly thought so by placing it in the Mastigocrinidae) and the stem is circular in section and composed of long columnals. In Sacrinus the radianal is pentagonal, there are 3 anal plates in the cup, radial facets are clearly peneplenary and protruding and the stem is heteromorphic and pentagonal in section. Despite these differences cup shape including shape and proportions of most individual plates, robustness and branching of the arms, size and plate ornament and arrangement of the anal sac and tuberculate ambulacral cover plates indicate a fairly close relationship between the 2 genera. We note some 141 comparison with Ordovician Grenprisia Moore, 1962 inthe very large cylindrical to inflated anal sac of small stellate or smooth plates, the 5- or 6-sided radianal, short wide radials and pentagonal siem. However, these comparisons could very well reflect parallel evolution and without intermediate forms no suggestion of affinity is made. The distinctive anal structure makes comparison with other cladids rather unnecessary. Sacrinus gamkaensis sp. nov. (Figs 22-25) ETYMOLOGY. From Gamkapoort. MATERIAL. HOLOTYPE: ROC25. PARATYPES: RO127,.255. 731.L7. C25, B4522, B4523, B4526, B4527 and B4530 all from Gamkapoort, all from the Gydo Formation. OTHER MATERIAL: B4574 from the Hex River Pass area on Montagu Road, 14km from the N9 хитой: RO283 from Swaarmoed Pass, Ceres, 33°21°30"S, 19°30°30E; and ROTIS from Matroosberg, Worcester (Hex River Pass) 33?30'S, 19?49* ED all in the Voorstehoek Formation. B4669 from the quarry on E of road N from Prince Alfred's Hamlet, about 1km S of Gydo in the Gydo Formation. DIAGNOSIS. Radials and basals with broad low ray ridges. Anal plates in cup 3; anal X only just larger than pentagonal radianal: anal plates and proximal anal sac plates with distinct ray ridges: anal sac long, cylindrical to moderately inflated, composed of many tiny stellate polygonal abutting plates. Arms isotomous, Ist branching on 4th. 5th. or 6th primibrach. Stem subpentagonal in section, heteromorphic. DESCRIPTION. Cup conical, up to 5mm long. with broad low radial ridges and (in some specimens) ornament of fine lines on top of ridges. Infrabasals 5, pentagonal, slightly wider than long (max). Basals 5, hexagonal except for heptagonal BC basal, with radial ridges forming a broad cross. Radials wider than long; articulating facet angustary, about 0.5 of plate width, only slightly declined. almost circular, projecting FIG. 22 Sacrinus gamkaensis gen. et sp. nov. A-B, crown in C-D interray view showing base of anal tube and arm branching RO 255, x6 and х4, respectively. C, ‘anal tube and interior of some arms RO L7a, х2. D-F, crown showing arm branching and ornament on cup ROC25 5, х2, x] and x3, respectively. G, crown showing anal tube, RO T21, x3. H, interior view of crown RO 127, x4. I, crown viewed from side opposite anal interray showing anal tube RO TI 8.x3. 142 MEMOIRS OF THE QUEENSLAND MUSEUM VIG. 24. Sacrinus gamkaensis gen. et sp. nov. A, crown in ambulacral view of arms and internal mould of cup RO 731, х2. В, ambulacral view of ann showing small cover plates RO 255, х5. C, partial crown showing ray ridges on radials RO T25, x3. D, crown showing ornament on anal tube RO L7, x3. laterally: radial ridges radiating from arm base in 4 directions towards centre of 4 lower sides (i.e. 2 cross to adjoining basals and other 2 laterally onto adjoining radials or radianal in case of C radial). Anal plates in cup 3. with radiating ridges directed to the middle of each side, with all 3 way sutural junctions depressed: anal X hexagonal, directly distal to CD basal, contacting D radial and other 2 anal plates towards C ray; radianal pentagonal, proximal and left of C radial, contacting BC and CD basals; right tube plate directly distal to radianal. Anal sac subcylindrical. inflated irregularly, reaching distally to about 3rd arm division, of many tiny irregularly polygonal abutting plates, aperture not observed: each plate with radial ridge ornament continuing and becoming finer distally. Arms slender, branching isotomously 7 times, laterally compressed in cross section, with deep adoral groove. with column of tiny cover plates on either side of groove; primibrachs 4, 5 or 6 axillary; in 2nd and more distal brachial series 5th brachial usually axillary (variable between or within arms even in | specimen), Stem subcircular to subpentagonal in section, noditaxis N3231323, with latus projecting laterally on nodals. REMARKS. Sacrinus gamkaensis is distinguish- ed from 5. hexensis by the radial ornament on its anal cup plates and anal sac, the latter being smooth. A few specimens of the type species also show a very fine multiple ridge ornament (Fig. 22D. F) ontop of the broad radial ridge ornament that is the specific distinguisher. Some specimens EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA 143 FIG. 25. Sacrinus gamkaensis gen. et sp. nov. A-B, part and counterpartof'incomplete cup on long stem RO TI15a & b, x3. C, crown showing arm branching on 6th primibrach RO 283, x1. D-E, adjacent views (different angles) of anal area of incomplete crown and stem B4574, x3. Е, posterior view showing stellate ornament on anal tube plates and large ambulacral cover plates proximally on arm, B4669, x3. SEUM MEMOIRS OF THE QUEENSLAND MU 44 | EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA (Figs 22А, 25C) suggest a pustulose surface on radial plates and in others (Fig. 24C) the radial ridge ornament is well developed on radials as well as anal cup plates. Variation noted here is considered intraspecific. Sacrinus hexensis sp. nov. (Fig. 26) ETYMOLOGY. From the Hex River Pass. MATERIAL. HOLOTYPE: B4571a (lower specimen of 2 close together). PARATYPES: B4571b (other 2 specimens), B4572, РКУ HR5 all trom Hex River Pass on Montagu Road, 14km from № tumoff (C282). DIAGNOSIS. Cup plates including anal plates and anal sac plates smooth; anal sac long, subcyl- indrical. of many small polygonal abutting plates: 151 arm branching variable at 3rd-6th primibrach. DESCRIPTION. Cup short. conical, with smooth plates, up to 5mm long. Infrabasals 5, pentagonal, as long as wide (max). Basals 5, hexagonal except for heptagonal CD basal. Radials wider than long in lateral view, with 6 sides plus radial facet: radial facet angustary. about 0.5 of plate width, only slightly declined, almost circular. Anal plates in cup 3; anal X hexagonal. distal to CD basal. contacting D radial and other 2 anal plates towards C ray; radianal pentagonal. proximal and left of C radial, contacting BC and CD basals; right anal tube plate distal to radianal. Anal sac subcylindrical, reaching distally to about 3rd arm division, composed of many tiny smooth polygonal abutting plates, opening terminal, with circlet of elongate plates around aperture. Arms slender, branching isotomously 4 times. laterally compressed in cross section, with deep adoral groove, with columns of tiny cover plates on either side of groove: primibrachs 3, 5, 6 or 7 axillary; similar variation in subsequent brachial series, number of brachials between axillaries not consistent even within one specimen. Stem subpentagonal to subrounded, heteromorphic, noditaxis N3231323; nodals slightly longer and wider, with rounded latus. REMARKS. This species is distinguished from K. gamkaensis above. FIG. 27. Cradeocrinus plenarius sp. nov. А-В. holotype crown B207A, x5. C, sketch of plate arrangement. IB = infrabasal; B = basal; R = radial; RA = radianal; X = anal X; rt = right tube plate. Cradeocrinus Goldring, 1923 TYPE SPECIES. Cradeocrinus elongatus Goldring, 1923 from the Upper Devonian West Falls Group of western New York; by original designation. FIG. 26. Sacrinus hexensis gen. et sp. nov. A, crown showing anal tube from side of cup opposite C-D interray B4571c, х5. B- C, holotype crown in A ray and C-D interray views B4571a & b, х4, respectively. D, crown in A-E interray view B4571D, x3. E, crown with D radial central and anal cup plates to right HR-5, х5. 146 Cradeocrinus plenarius sp. nov. (Fig. 27) ETYMOLOGY. For the plenary radial facets. MATERIAL. HOLOTYPE: B0207 from the Waboom- berg Formation at Boplaas Farm, N of Ceres. DIAGNOSIS. Cup long, cylindrical, very small; plates smooth. Radial facets plenary. Three anal plates in cup; radianal pentagonal, contacting Ist right tube plate; anal sac long slender, of 8-10 columns of polygonal plates, perforated in sutures. Arms slender, of long brachials with shallow ventral groove, branching isotomously at 4th primibrach. DESCRIPTION. Cup small, 2.2mm long, 1.2mm max diameter, high conical; plates smooth. Infrabasals 5, equal, long, pentagonal in lateral view. Basals 5, hexagonal, longest plates in cup. Radials 5, pentagonal, with plenary radial facets. CD interray wide, with 3 anal plates in cup, and proximal 1/2 of 2 others just in cup, anal X and radianal each supporting column of plates rising into anal sac; radianal pentagonal, proximal and left of C radial, contacting right tube plate; anal sac at least 10mm long, of 8-10 columns of regular hexagonal plates, with depressions lead- ing to perforations in the sutures. Arms very slender, tapering strongly over | st 2 primibrachs, long (longest available incomplete at 19mm), with 4th primibrach axillary, branching at least twice distally but type of subsequent branching uncertain (probably heterotomous with ramules from shapes of axillaries available), non- pinnulate; primibrachs more than twice as long as wide, with shallow ventral groove. Stem circular in section, heteromorphic, with alternating long and short columnals of uniform diameter, tapering for proximal 10mm then uniform in diameter, full length unknown. REMARKS. This species is distinguished in the genus by its plenary radial facets (other species have peneplenary facets), small size, smooth plates, and very slender arms. It has the cup shape of C. elongatus and the anal sac structure of C. nanus (Roemer,1863) (Schmidt,1934, fig.24) but in combination its features are separate. Thalamocrinus Miller & Gurley, 1895 TYPE SPECIES. Thalamocrinus ovatus Miller & Gurley, 1895 from the Ludlovian Brownsport Formation in Tennessee; by origianal designation. REMARKS. McIntosh & Brett (1988) reviewed the genus in detail providing a clear basis for MEMOIRS OF THE QUEENSLAND MUSEUM comparison. The South African material is not as well preserved as most of the North American species and on only one South African specimen is the posterior interray available, and then not clearly. However, the radianal, although pentagonal is similarly placed and the anal X is in the radial circlet, the cup is barrel-shaped to only very slightly flaring, the plates are of similar thickness and the proximal part of the stem is very similar with nodals much larger than internodals and well rounded laterally. T. arenaceus is distinguished within the genus by its very short infrabasal circlet. Thalamocrinus arenaceus sp. nov. (Fig. 28) ETYMOLOGY. For the sandstone matrix of all specimens. MATERIAL. B4538 and B4556 (numerous cups and arm and stem fragments on each slab) from the Gamka Formation at Grootrivier Hoogte. DIAGNOSIS. Infrabasals short in lateral view. DESCRIPTION. Cup barrel-shaped to slightly flaring at radials, about as long as wide (6-8mm in available specimens); plates smooth, convex, thick. Infrabasals 5, very short and pentagonal in lateral view. Basals 5, hexagonal, as wide as long; posterior basal heptagonal. Radials 5, pentagonal; radial facet angustary. Three anal plates in cup; radianal pentagonal, proximal and left of C radial, isolated from anal sac; anal X pentagonal; anal sac not available. Arms of thick brachials, with U-sectioned ventral groove. Stem circular in section, strongly heteromorphic with alternating nodals and internodals of markedly different diameters, with rounded latus. REMARKS. This material is preserved in medium to coarse sandstone, indicative of a higher energy environment than most Bokkeveld echinoderms which occur in the intervening shales. The two available slabs suggest consider- able postmortem movement with most arms and stems disarticulated. However, no disarticulated cups are evident suggesting strong interplate sutures and/or cup shape less susceptible to disaggregation. Family LECYTHOCRINIDAE Kirk,1934 Othozecrinus gen. nov. TYPE SPECIES. Othozecrinus royi sp. nov. EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA 147 FIG. 28. Thalamocrinus arenaceus sp. nov. A, group of 6 cups, 3 of them looking into the cup. 2 looking at the base and | on ils side B4538, <3. В. cup in lateral view and dissarticulated macro and microcolumnals B4538. «4. C, twocupsin lateral view B4556D, х3. D, cup B4556B, х4. E, surface with 4 cups including that from D. all inlateral view B45564A. х4. F, holotypecrown showing posterioranal plates and part ofanarm D4556C. х5. 148 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 29, Othozecrinus royi gen. et sp. nov. A-B, B4545, х2 and х1, respectively. C, internal mould showing large anal tube 34545. >], D, internal mould of cup SAMII69, x3. E, stem columnal B4539A. х4, EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA ETYMOLOGY. For Roy Oosthuizen, who collected much of the material described herein; an anagram to which is added only the ‘cr’. DIAGNOSIS. Cup wide bowl-shaped, with strongly convex plates; 5 infrabasals; radianal and anal X in cup in radial circlet. Anal sac of moderate length, inflated, of numerous large polygonal plates. Arms uniserial, branching isotomously; primibrach 7 axillary. Stem circular, relatively small diameter, with greatly expanded epifacet into 6-9 blunt spines. REMARKS. While the details of this species are not entirely available, in particular the oral area and the branching pattern of the arms, it is sufficiently known to be confident it does not fit an existing genus. The two similarly sized (but differently shaped anal plates in the cup, convex infrabasals, 7 primibrachs and very distinctive stem combine to isolate this species. The Lecythocrinidae has members with 2 anal plates in the radial circlet. The anal plate arrangement of Othozecrinus is most similar (though not closely) to that of Lecythocrinus (cf Moore et al.,1978, fig.375.7) but the concealed infrabasals and the subquadrangular stem with large axial and 4 peripheral canals make close relationship unlikely. Other members of the family, Cestocrinus and Corynecrinus, have 0 or 5 anal plates in the cup and Corynecrinus has infrabasals concealed by the stem. The Euspirocrinidae is distinguished by its 3 anal plates in the cup and the Barycrinidae has a small radianal proximal to C radial. If forced to place this genus, with such distinctive stem, into a family it has to be the Lecytho- crinidae but that is a tentative assignment. Othozecrinus royi gen. et sp. nov. (Figs 29-31) MATERIAL, HOLOTYPE: B4545 from Hex River Pass (C281, Суйо Formation). PARATYPES: SAM1169 from Boschluis Kloof (Gydo Formation), SAMK965 from Wolfaardt's Farm, Riet Valley, Ceres, B4524, B4539 from Grootrivier Hoogte (C2Q1, Gamka Formation), ROC20 Gamkapoort, Prince Albert at 33?18'S, 21?38'E (lst shale), ROL70 Warmwatersberg, Barrydale at 33?46'30"S, 20°55°30"E (Ist shale). SUGD300 from De Dooms.B4600 from Platfontein in the Gydo Formation. DIAGNOSIS. As for genus. 149 FIG. 30. Othozecrinus royi gen. et sp. nov., sketch of plate arrangement with arm branching as far as known on right and dotted lines on infrabasals indicating extent of stem facet. IB — infrabasal; B = basal; R = radial; RA = radianal; X = anal X; A-E above radials indicate rays. DESCRIPTION. Cup short, wide bowl-shaped, of thick, strongly convex plates. Infrabasals 5, forming pentagonal circlet, strongly convex, with short strong almost spinose projections on A, B and C infrabasals in larger specimens; stem facet central, hemispherical concavity, depressed distal to projecting convexity of infrabasals, with distinct rim. Basals 5, smooth, convex with sutures depressed, hexagonal (but with 2 proximal sides at very high obtuse angle) except heptagonal CD and BC basals. Radials 5, convex, with Ist primibrach narrower than radial; articulating facet slightly declined outwards, with strong transverse ridge and distinct muscle and ligament areas. CD basal supporting 2 anal plates, with right just proximal to left; distal edges of these 2 plates level with distal margin of radial circlet. Anal sac short, squat; polygonal plating apparently irregularly arranged, with finely scalloped edge to each plate indicating perforate wall, perforations circular along sutures. Arms uniserial, thick, of long brachials, with sharp deep ambulacral groove; primibrach 7 axillary; rest of arms unknown. Stem of large columnals (c. 15mm across not including blunt lateral spines), up to 4mm long, with small (up to 4mm diameter) central circular articulum, with up to 10 large blunt lateral spines. Articulum with small central lumen showing 5 rounded pits, one at each corner of the pentagon and circular jugulum centrally; rest of articulum with radial ridges and intervening grooves forming triangular segments, with fine peripheral rim to articulum. Latus (epifacet) with smooth gently convex (in radial direction) surface, extended into stout blunt spines; 6-9 blunt spines per 150 MEMOIRS OF THE QUEENSLAND MUSEUM VIG 31. Otliozecrinus royi gen et sp. nov. A. base of cup geol surv B4600, х3. В. base of cup B4524. х3, С. base of cup RO L70, #3. D, lateral view of different cup RO L70, «3. E. oral view of interior of cup SUGD300C, x2, F. basal circlet with stem attached RO C20, х3. G, columnal B4539B. х5. Н. columnals SUGD30t ЈА. х3. EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA columnal. Blunt spines on every visible columnal in lateral view. REMARKS. //yperexochus immodicus Moore & Jeffords,1968 from the Early Devonian of Tennessee has many lateral spines on its columnals and a small articulum but the surface of the articulum is differently ornamented and the spines are not in a single whorl, shaped differently апа more numerous. Sumarievstis radiatus Stukalina, 1978 from the Middle Ordovician of the southern Tien Shan in central Asia is the most similar columnal but it has many more lateral spines and a narrow peripheral crenularium on the articulum. Floricolumnus Donovan & Clark, 1992 is similar in its greatly expanded epifacets which are almost spinose but in that genus they are well-separated nodals with smaller columnals between whereas in Othozecrinus they occur on each columnal evident (Fig. 31F). However, the central depressionand narrow crenularium on both sides of these columnals indicate that at least one unexpanded columnal occurs between every pair of expanded ones. It seems unlikely that any of these genera is closely related to Othozecrinus and that the columnals with expanded epifacets have evolved independently in each case. Class ASTEROIDEA de Blainville. 1830 Terminology and suprageneric classification largely follow Spencer & Wright (1966) except that: 1) their ‘Mouth angle plate’ (MAP) (—'oral ossicles’ of Blake & Guensburg. 1989) is here termed Ist ambulacral plate (AMB1) following Smith & Jell (1990); 2) their Amb | is here termed ambulacral 2 following Bjork et al. (1968) and Smith & Jell (1990): 3) ‘dorsal’ and ‘ventral’ are used both for body and ossicle surfaces; ‘adradial’ and "abradial for directions toward and away from the arm axis, respectively: and "proximal and ‘distal’ fordirections toward and away from the mouth, respectively. Dimensions of plates in the arms are indicated by: (rad.) = in radial direction or length of arm: (ad.-ab.) = in adradial-abradial direction across arm. In general, dimensions are referred to as ‘long’ and ‘short’ in the radial direction and “wide or ‘narrow’ across the arm. Family PROMOPALAEASTERIDAE Schuchert, 1914 Aulacolatiaster gen. nov. TYPE SPECIES. Aulacolatiaster breviramus sp. nov. ETYMOLOGY, Latin aulax, furrow, latus, wide and aster; a star, referring to the wide ambulacral groove. DIAGNOSIS, Arms 5. short (no longer than diameter of disc), wide. Dorsal surface with primary ossicular ring of large ossicles centrally. Ventral surface dominated by wide elliptical ambulacral grooves; ambulacrals wide, opposite, with extremely shallow ambulacral channel, with fine parallel ambulacral ridges running abradially parallel to plate margins and separating podial basins; Ist ambulacrals large. oriented vertically. paired across interradii: inferomarginals with pair of short lateral spines. becoming elongate proximally, isolating from the margin of the frame at least 1 narrow axillary (odontophore) in each interradius: adambulacrals with transverse row of extremely large prominent tubercles. REMARKS. This genus is distinctive in its combination of meshwork dorsal plating in distinct radial columns, wide ambulacral groove with wide ambulacrals bearing fine parallel ridges separating podial basins, adambulacrals with row (ad.-ab.) of very large prominent tubercles and inferomarginals becoming wider proximally and each with 2 lateral spines. We consider it most closely allied to Promopalaeaster Schuchert. 1914 with which it shares the same type of dorsal plating. wide ambulacral groove, large Ist ambulacral, and interradial structure but from which it may be distinguished by its parallel ambulacral ridges indicating a single column of tube feet along each side of the arm. extreme size of the tubercules ina transverse row on the adambulacrals and short arms which could prove to be growth related when more material becomes available. Ambulacral ridge structure proximally distinguishes North American species, including the type species of Promopalaeaster but Promo- palaeaster elizae Spencer, 1930 from the Upper Ordovician of Scotland and Koactis simplex Spencer, 1914 from the Lower Silurian both exhibit (Spencer & Wright, 1966, figs 50.1 and 50.2c) ambulacral ridges parallel to each other and to the interplate sutures throughout as in Alulacolatiaster. Aulacolatiaster bears some resemblance to Palasterina McCoy. 1851 inthe wide ambulacral groove. parallel ambulacral ridges, and large Ist ambulacrals but is distinguished by its interradial structure and dorsal plating. Its placement will remain problematic until better preserved A һә MEMOIRS OF THE QUEENSLAND MUSEUM tA VIG. 32. Aulacolatiasterbreviramus gen. et sp. nov. A, ventral view of mouth region of large incomplete paratype with mouth wide open SAMK 1063, 4. B-C, dorsal and ventral views, respectively. of holotype SAMK 018a & b, х4 and х6, respectively. EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA l Un чө FIG. 33. Ulrichaster macrodentatus sp. nov. A-B, enlarged ventral view of oral region and ventral view of incomplete paratype with short section of crinoid stem in upper right КО 44, x6 and х3, respectively. C-D, enlarged dorsal view of arm and dorsal view of incomplete holotype PRV725, «6 and x4, respectively. 154 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 34. Ulrichaster macrodentatus sp. nov. A-B, ventral view of slightly disarticulated paratype and enlargementof its oral area РКУ737. x5 and x8, respectively. C, juvenile paratype in dorsal view PRV738, x10. EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA 15 material becomes available, particularly a dorsal surface. Aulacolatiaster breviramus sp. nov. (Fig. 32) ETYMOLOGY. Latin brevis, short and ramus. arm; for the relatively short arms. MATERIAL. HOLOTYPE: SAMKIOIS, dorsal and ventral external moulds from Riet River, N of Ceres Division on Wupperthal Road; S side of farm which is also called Groot Rivier in the neighbourhood; Gydo Fmn, 33m below top of unit, SAMK 1063 incomplete ventral extemal mould from Wolfaardt’s Farm near Ceres, about 100m NNE of the house. DIAGNOSIS. As for genus. DESCRIPTION. Stellate. Arms 5. short (10mm from oral axis to arm tip on holotype), wide (4mm at widest point). Dorsal surface with primary ossicular ring of large ossicles in radial positions and with central pit or perforation on each, outlines unclear but apparently sub- quadrate: a further circlet of large ossicles probably present in interradial positions and resting on the circlet of radially disposed ossicles (single dislocated subrectangular plate lying in interradial position (Fig. 32B); dorsal surface of arms with broad median groove (probably preservational), with midline bordered by columns of weakly tuberculate plates that abut along the midline of the arm. with at least 3 columns of tuberculate plates abradially separated by deep pits (suggesting some sort of meshwork stucture but this remains unclear from the available specimen), with columns of plates laterally aligned into rows across the arm, without clearly defined superomarginals. Ventral surface with small interradii and wide elliptical arms; ambulacral grooves elliptical, occupying about 2/3 arm width for most of arm length (more than 80% distally), extending to arm tip: ambulacrals wide, opposite, with extremely shallow median ambulacral channel: ambulacral ridges fine. parallel, with very small expansion at adradial end, running abradially parallel to plate sutures (this attitude not absolutely certain but where sutures between successive ambulacrals are evident they are parallel to the ridge), weakly expanded at abradial end to about same length as adambulacral; Ist ambulacrals large. plough shear-shaped, oriented vertically, paired with convex sides adjacent across interradii:; inferomarginals widerthanlong.becoming much wider proximally with at least 1 axillary (and possibly 3 or 4) separating adambulacrals from n inferomarginals in each interradius, with pair of short lateral spines directed abradially on each inferomarginal: adambulacrals with transverse row of prominent tubercles (tubercles may be irregularly arranged in smaller individual but well aligned in transverse rows from expanded abradial end of ambulacrals in large individual (Fig. 32A)) REMARKS. The holotype is known from external moulds of both surfaces but whereas the ventral mould is very well preserved the dorsal mould seems to have been abraded (probably since collection) and does not preserve much detail. The fragmentary external mould of the ventral oral area of a larger individual is well preserved but only a small fragment. Both casts of the ventral surface have a convex mound of matrix in the oral pole suggesting that the central dorsal structure was weak and collapsed easily upon death. Well-developed podial basins between the ambulacral ridges abradially are shared by contiguous ambulacrals but preservation is not good enough to determine whether or not a pore passes through the basin floor. Family URASTERELLIDAE Schuchert, 1915 Ulrichaster Spencer. 1950b TYPE SPECIES. Urasterella ulrichi Schuchert, 1915 from the Middle Ordovician of central USA. REMARKS. Spencer (1950b) subdivided Urasterella McCoy into a group witha single row of ossicles in the dorsal midradius of the arms (further subdivided on features of dorsal arm ossicles between the radial and marginal columns) and a second group with 2 rows of ossicles in the dorsal midradius (1.е., on either side of the arm axis). He erected U/richaster for the latter group which accepts the new species described herein. Ulrichaster macrodentatus sp. nov. (Figs 33, 34) ETYMOLOGY. Greek macros, long and dentatus, tooth: for the long 1st ambulacrals. MATERIAL. RO44 from the Voorstehoek Shale (C282) at 33°30°S: 19°49°E near Matroosberg, Worcester (Нех River Pass). PRV738, 737 (and counterpart PRV725) trom Tafelberg in the Waboomberg Shale (C284). DIAGNOSIS. Dorsal surface with very open meshwork of ossicles in radial columns, columns symmetrical about arm axis. Ist ambulacrals large. elongate, reaching interradial margin or 156 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 35. Haughtonasterreedi Rilett, 1971. A-B, ventral and dorsal views of juvenile holotype SAM 3881, x5. C, dorsal view of juvenile paratype SAM3375, х5. D, ventral view of SAM K978, x6. close to it, in closely adjacent pairs across inter- radii. DESCRIPTION. Stellate. Arms 5, long (at least twice disc diameter). Dorsal surface an ordered meshwork (with large perforations) of small tuberculate elongate plates in 4 radial columns (2 columns on either side of arm axis smaller than next columns abradially) linked by less regular struts in transverse rows, without obviously differentiated primary ossicular ring or other plates (but disc plating not clear on available specimens). Ventral surface with long parallel sided ambulacral grooves; ambulacrals wide, opposite, with diagonal ambulacral ridges having curved ventral edges forming basins abradially. EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA Ist ambulacrals large. extending to interradial margin, straight but with strong lateral projection on adradial side, paired across interradii: adambulacrals narrow. subrectangular to sub- discoidal in ventral view, without spines: inferomarginals forming lateral margins of arms, small (same size as abradial dorsal plates). with 1 or 2 or more short stubby spines forming spinose margin. REMARKS. None of the available material is well preserved but enough is available to show the features of U/richaster in particular the short stubby spines of the inferomarginals and abradial dorsal plates and the ordered meshwork of radial columns of the dorsal plating with 2 columns either side of the arm axis. However. shape of the Istambulacralsand the larger pits between dorsal plates distinguish it from other members of the genus. The type species, U. ulrichi has more columns of ossicles covering the arm dorsally and does not appear to have the larger pits of the dorsal meshwork as іп U. macrodentatus. U. gutterfordensis Spencer. 1918 has small Ist ambulacral plates. Sa/teraster biradialis Withers & Keble. 1934 from the Ludlow of Victoria (included by Spencer (1950b) in U/richaster) is an ophiuroid with the 2 median rows of ossicles the dorsal side of the ambulacrals, any dorsal plating having been removed or not preserved. Class OPHIUROIDEA Gray. 1840 Order STENURIDA Spencer.1951 Suborder PAROPHIURINA Jaekel, 1923 Family EOPHIURIDAE Schóndorf. 1910 Haughtonaster Rilett, 1971 TYPE SPECIES. Haughtonaster reedi Rilett, 1971 from the Gydo Formation near Ceres. DIAGNOSIS. Disc pentagonal. dorsal surface of smooth oval plates surrounded by an irregular mass of very fine, almost spicular elements: ventral interradii triangular. with polygonal plates having some perforations on sutures (i.e. a few oppositely scalloped margins of plates). Ambulacrals offset across arm axis, rectangular in dorsal view: adambulacrals wide and gently oblique to arm axis in ventral view, forming lateral walls of arm; podial basins subcircular. very deep. Ist ambulacral with strong curved ridge forming distinctive elliptical basins interradially. REMARKS. Rilett (1971) assigned this genus to the Eophiuridae to which Spencer & Wright (1966) assigned only Fophiura from the Arenig of Czechoslovakia. The more mature specimens now available indicate that a close relationship is unlikely although the 2 genera should probably be retained in the same suborder. Hotchkiss (1976) established a subordinal level division of the Stenurida based on whether ambulacrals were opposite or offset across the arm axis, noting that pre-existing classification had been based on grades of development and inferring that the development of offset ambulacral columns indicates a monophyletic clade of Palacozoic ophiuroids. He erected the Scalarina for stenurids with opposite ambulacrals and the Parophiurina is available for those with offset ambulacrals. Phylogeny within the Paraophiurina is not clear and would require a review of known members, which is outside the scope of this paper. Haughtonaster is distinctive in that suborder in lacking spines on the arms, lacking a column of sublateral plates, in its long slender tapering arms, in its deep circular podial basins and in its wide adambulacrals enclosing the arm laterally. In the shape of ambulacrals along each arm, nature of adambulacrals and podial basins, and in particular the distinctive ridge on the Ist ambulacral. Haughtonaster very muchresembles Stenaster Billings. 1858. However. that genus belongs to the Scalarina and a close relationship would not be possible unless the offset arrangement of ambulacrals evolved more than once. We do not speculate on that possibility and retain Haughtonaster in the Parophiurina. Because we can see no workable family arrangement we retain Rilett’s (1971) family assignment. Haughtonaster reedi Rilett, 1971 (Figs 35-37) MATERIAL. HOLOTYPE: SAM3881 from Hottentots Kraal. PARATYPE: SAM3375 from Hottentots Kloof, Ceres, OTHER MATERIAL: ROC50, RO122, RO804 from Gamkapoort, Prince Albert at 33°18°S, 21?38 E in the Gydo Formation, B4567 from Swaarmoed Pass, 1.3 miles from Great Swaarmoed Farm, Ceres; PRV1484. SAMI3472 from Voetjies Kloof Suid; SAMI3470; SAMK 1016 from Riet River (also called Groote Rivier) N of Ceres Division on Wupperthal Road. S side of farm in Gydo Formation; SAMK978 from Wolfaardt's Farm near Ceres (100m NNE of house). DESCRIPTION. Overall form. Maximum radius 7-30mm, disc radius 5-6mm. Arms 5, short and blunt in small specimens, in large specimens weakly petaloid in proximal 1/2 and strongly tapered over distal 1/2, with elongate pointed tip. Disc pentagonal, extending to 5th or 6th 158 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 36. Haughtonaster reedi Rilett, 1971. A, dorsal view of RO 122, 2. B, ventral view of B4567, x3. C-D, dorsal views of RO C50, *4 and x2, respectively. EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA 159 FIG. 37. Haughtonasterreedi Rilett, 1971. A, ventral view of incomplete smaller specimen RO 804. х6. В, dorsal view of proximal part of arm RO C50 х4. С, ventral view of incomplete adult SAM13472 x3. D. dorsal view of incomplete adult RO C50 x2. E, ventral view of incomplete adult SAM13470 х4. F, ventral view of 2 adults PRV 1484 х4. 160 ambulacral; in small specimens only 3-4 tiny plates in interradius, in larger specimens 8-12 polygonal plates in a triangular interradius, plates irregularly polygonal, possibly disc plates with scalloped margins in larger specimens (no area of plating well preserved). Only small fragments of dorsal disc available. Arm plating. Ambulacrals in dorsal view subquadrate (in smaller specimens) to subrectangular (larger specimens), offset across arm axis distal from 2nd ambulacral, with sharp elliptical clefts between successive ambulacrals in each column, with weakly zigzag (obtuse projection at midlength of ambulacrals pointing at recessive interambulacral suture in opposite column of ambulacrals) line of suture between the 2 columns in arm axis. Ambulacrals in ventral view subtrapezoidal, separated from succeeding ambulacral in same column by shallow rounded basin shared by 2 ambulacrals (i.e. inter- ambulacral suture runs through middle of depression) and descending abradially into deeper part of podial basin, with straight radial axis between 2 ambulacral columns; ambulacral channel very shallow, barely discernible. Adambulacrals in dorsal view subtriangular, extending ventrally to form the abradial wall of the arm, with adradial tip directed at proximal end of ambulacral. Adambulacrals in ventral view wide, with parallel interadambulacral sutures, continuing abradially to form lateral wall of arm, without spines, with sharp adradial projection between podial basins. Podial basins subcircular with slight expansion on adradial side between adjacent ambulacrals, extremely deep adjacent to adambulacrals and shallower adjacent to radial axis of arm, apparently with some subtle ledges and dimples on the evenly curved walls (but preservation is not good enough to be certain of consistent structures). Mouth frame. Dorsal aspect not available except in small specimen; central circular dorsal depression but plating details not available. Mouth small; no buccal slit. Smallest specimen with Ist ambulacral unspecialised except for slight proximal projection. Ist ambulacral becoming more elongate (radially) with growth, developing strong curved ridge convex towards arm axis; curved ridges from adjacent arms forming distinctive elliptical basins interradially. REMARKS. The marked changes that take place in this species during growth involving increasing arm length relative to width and development of the pentagonal disc could be MEMOIRS OF THE QUEENSLAND MUSEUM considered indicative of separate species if it were not for the distinctive structure of the Ist ambulacrals and the broad adambulacrals forming the lateral walls of the arms that link the specimens over the whole size range. There is no discernible variation in the available material of the same size. As discussed under the generic remarks above the only species deserving of comparison is Stenaster obtusus, which has its ambulacrals opposite each other. Ophiuroid arm indet. A (Fig. 58A) MATERIAL. SAMK625 from Gamkapoort. DESCRIPTION. This specimen, preserved only as an external mould, is interpreted as the ventral aspect ofa part ofa free arm with the 2 alternating columns of vaguely hexagonal ambulacrals on the right, serving to identify the slightly zigzag arm axis; proximal is inferred as up the page from the gradually diminishing size of plates in the opposite direction (distal); a column of transverse plates abutting the ambulacrals laterally is interpreted as a column of sublaterals; the column of longitudinally elongate plates forming the lateral margin of the arm (and to which the abradial end of the sublaterals abut at the junction between successive plates) is interpreted as the lateral column and is not obviously spinose. The subquadrate depressed areas bordered by the ambulacral and lateral columns and separated by the sublaterals (podial basins?) are fully floored and have 2 distinct depressions on that floor; one is adjacent to the distal end of the ambulacral and the other is proximal and abradial against the lateral plate. REMARKS. Structure ofthe arm with 3 columns of plates (ambulacrals, sublaterals and laterals) suggests a primitive ophiuroid among the Stenurida (cf. Eophiura, Pradesura, Stuertzaster etc.). The alternating columns of ambulacrals exclude it from Hotchkiss's (1976) Scalarina and place it in the Parophiurina as used herein. However, there is no comparable form in this or any related group. Other stenurids with alternating ambulacrals may have the podial basin entirely on the ambulacrals (Eophiura, Pradesura) or have very wide podial basins (Stuertzaster) but none have the subquadrate podial basins of this South African form. This specimen represents a new genus but provides insufficient information for its definition forcing us to retain it in open nomenclature. The position ofthe sublaterals forming the proximal margin of EARLY DEVONÍAN ECHINODERMS OF SOUTH AFRICA 16! FIG. 38, Hexuraster weitzi (Spencer, 19504), lectoty pe. SAMI11055. A, ventral view (mould and thus latex cast incomplete) «1.5. B. dorsal vie «0.9. C-D, enlargements of parts of arms from A in ventral vie «2.5. 162 the podial basin suggests a possible affinity with Stuertzaster, which is as much as can be said at the moment. Order OEGOPHIURIDA Matsumoto, 1915 Suborder LYSOPHIURINA Gregory, 1896 Family CHEIROPTERASTERIDAE Spencer, 1934 Hexuraster gen. nov. not Hexura Simon, 1884: 314. Hexura Spencer, 1950a: 300. TYPE SPECIES. Hexura weitzi Spencer, 1950a from the Lower Devonian of South Africa. DIAGNOSIS. Body large (incomplete holotype 130x80mm); disc uncalcified, extending to arm tips. Arms slightly petaloid. Ambulacrals cylindrical with lateral projection (or toe of boot) shorter than radial length. Adambulacrals wide (ad-ab), expanded abradially to be T-shaped, with single very strong lateral spine on each adambulacral. Madreporite interradial, close to mouth frame on ventral surface. Mouth frame small for size of animal, plates not massive, with well-developed podial basin on 2nd ambulacral. Mouth large, with short buccal slit between Ist 2 ambulacrals. REMARKS. Spencer's (1950a) name was preoccupied by a spider genus so the replacement name is proposed herein. Spencer (19504, figs 1 - 3 - 4) included 3 specimens which are assigned herein to 3 different genera. Jell (1997) chose SAM11055 (Spencer, 1950a, figs 1,2) as lectotype of H. weitzi and placed it in the Cheiropterasteridae which he reinstated from synonymy with the Encrinasteridae (Spencer & Wright, 1966). Spencer (1950a) allied Hexuraster with his Euzonosomatidae (=Encrinasteridae of Spencer & Wright, 1966) and although he quoted several reasons that do not apply to the lectotype of the type species but rather to his figs 4 and 5 we consider the Cheiropterasteridae closely related to the Encrinasteridae. Following Jell (1997), the family is known from the Early Devonian of Germany and South Africa and the Early Carboniferous of Indiana. Hexuraster weitzi (Spencer, 1950a) (Figs 38, 39) Hexura weitzi Spencer, 1950a: 300, figs 1. 2 (not figs 3-5). MATERIAL. LECTOTYPE: (chosen Jell, 1997) SAM11055 (130x80mm) from the Lower Devonian Bokkeveld Series, at De Doorns; RO45 (about 30mm MEMOIRS OF THE QUEENSLAND MUSEUM diameter) from the Gydo Formation at Gamkapoort, Prince Albert, 33?18'S, 21?38' E. DIAGNOSIS. As for genus. DESCRIPTION. Overall form. 5-armed, preserved radius 15-65mm, arm radius just greater than disc radius. Arms slightly petaloid, with rounded only weakly tapered tips. Disc very large, uncalcified, extending to arm tips, with high obtuse angled re-entrant in some interradii but continuing almost circular in other interradii. Arm plating. Ambulacrals subquadrate in dorsal view, offset across arm axis distal from ambulacral 3, boot-shaped in ventral view, with narrow cylindrical leg, long lateral process (foot) having a slight concavity on proximal side (instep) to receive abradial tip of distal end of next proximal ambulacral. Ambulacral channel prominent, formed by half grooves along the adradial edge of each ambulacral, with straight line of suture between 2 ambulacral columns in bottom of amulacral channel. Adambulacrals with long lateral projection abutting against the toe of ambulacrals; in dorsal view abradial subtrapezoidal expanded part of adambulacral with sharp proximal-distal ridge aligned on all adambulacrals to form ridge along the length of arm; ridge sharper on adradial side than on abradial side, with furrow on abradial side; lateral face subcircular to oval, flat, with single stout lateral spine attached apparently by suture. In ventral view expanded head of adambulacral subcircular, with low proximal-distal ridge aligned along the arm into one long ridge. Madreporite. Subcircular to oval, large (4mm max. diameter in lectotype), possibly with vermiform ornament (not entirely clear in lectotype), situated only slightly asymmetrically interradially adjacent to ambulacral 2. Mouth frame. Mouth very large in available specimens, with Ist ambulacral situated interradially leaving large embayments in each arm axis. Ist ambulacral long and narrow with deep transverse groove near midlength in dorsal view, similarly shaped but without transverse furrow in ventral view, with sutured junction to 2nd ambulacral almost radial. Smaller specimen with smooth basin-like recess under distal part of Ist ambulacral. 2nd ambulacral little different from more distal ambulacrals, subquadrate, indistinct in dorsal view, subquadrate, with well- developed podial basin in ventral view. EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA FIG. 39. Hexuraster weitzi (Spencer. 19504). mconiplete juvenile m ventral view RO 45 «4. REMARKS. The lectotype has some iron oxide minerals deposited in the external mould obscuring details in some areas, particularly in the mouth [тате but details of the arms are available by reference to different parts of the mould. Family ENCRINASTERIDAE Schuchert. 1914 Encrinaster Haeckel, 1866 nol -lypidexamey Vitzinger, 1843. Aspiclosoma Goldluss. 1848: 145: Schöndori 1910: 4 Encrinaster Haeckel. 1866:67: Lehmann. 1937: 28; Spencer & Wright, 1900: DES, Juzonosoma Spencer, 1930; 411; Spencer & Wright. 1966: Lió: Lehmann, 1957: 24. TYPE SPECIES. рохо amoldi Goldtuss. 1838 from the Lower Devonian of Germans, 103% SPECIES ASSIGNED. arnoldi. tischheinianus. petaloides, eifelensis, goldfussi, pontis, roemeri, schinidti aud laevidiscus REMARKS. The several German species of this genus were reviewed by Schóndorf (1910) (as Aspidosoma) and Lehmann (1957), Spencer (1930) spread these German species and others from Britain and North America between Fnerinaster and Fuzonosoma. Generic placement of fiscltheinianus depends on the concepts of these 2 genera. Spencer (1930: 404) provided a key for genera of his Euzonosomaüdac. a junior synonym of Schuchert’s Encrinasteridae, in which he indicated the “Geno-holotype’ of /'uzonosonm as E. petaloides (Simonovitsch). In the same work (1930; 411) he appears to have tried to designate a different genotype in the sentence '"JKuzonosoma orhitoides. n. sp.. is chosen as the holotype of the species. Although his intention is not clear from this work. the first of these citations links the words genus (geno-) and type (holotype) and must be accepted as a valid designation of a tv pe species. The second citation does not link the words genus and type in any way and cannot be considered a valid designation of a genoty pe or type species. Therefore, the type species of Euzonosoma is Aspidosoma petaloides Simonovitsch, 1871 by original designation. No action by any subsequent author can change this designation, Even subsequent action by the original author (c... Spencer & Wright. 1966) cannot supplant the original designation, So regardless of Spencer s intentions. which may be inferred from his subsequent citation of £. orhitoides as the type species and despite subsequent workers acceptance of E, orbitoides as type. К. petaloides must be considered the type species. This recognition should not greatly change the concept of Fuzonosoma because Spencer (1930: 404) included that species in his generic concept. Spencer (1930) distinguished between these 2 genera in his key by only 1 feature. the degree of widening (ad-ab) of the adambulacrals in (he median regions (1,c.. midlength) of the arm: in Euzonosoma the adambulacrals are distinctly broader in the median region than ai the extremity whereas in &nerinasrer adambulacrals are only slightly differentiated (inferring some widening but only slight) compared to the extremity. The variation usually comes down to width of the adambulacrals within the disc as opposed to their width just outside the disc. Most authors and 164 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 40. Enerinaster tiselibeinianus (Roemer, 1862 ). ^. dorsal view of SAM K1018 x2. B-D, ventral view of whole specimen, of disc area and of madreporite. respectively. SUG299. x |.2 «3.5 an x 15, respectively. E. slab with 3 individuals in ventral vie «0.8 (figured by Spencer. 1950, figs 4, 5), EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA particularly Spencer accept that ambulacrals and adambulacrals could rotate in a transverse direction (ab.-ad.) and this is reflected in width of the ambulacral groove which varies from specimen to specimen. We suggest that such rotation 1s gretly restricted in the disc but not so restricted in the free arms. Therefore, we suggest that the specimens with wide adambulacrals in the proximal free part of the arm are ones where the adambulacrals have rotated laterally to expose their full width in dorsal view. Thus width of adambulacrals in direct dorsal or ventral view may often be influenced by the attitude in which a specimen is buried (1.е., whether adambulacrals are curled ventrally concealing much of the ambulacrals or flattened out on the sediment and often dislocated from the ambulacrals. In some instances, particularly where preservation is in fine mudstone, the degree of convexity of the arm is greater where the adambulacrals appear narrow and least where the adambulacrals appear widest; this suggests to us that rotation of the plates in the arm is a highly significant factor in determining the dorsal appearance of adambulacrals. Given the considerable width of the adambulacrals, small differences in attitude may make significant differences to perceived width. This feature must be considered unacceptable as a generic discriminator at least in the degree of widening. In the same key, Spencer (1930) distinguished a 3rd closely related genus, Mastigactis by its adambulacrals being of uniform width throughout the arm. Whether this uniformity is the result of the entire arm having been buried without any lateral rotation of the adambulacrals is not easy to determine without reference to the specimens and should remain an open question. Recognising that a key seeks to limit the number of features by which to make easy recognition of taxa a search of discussion of the 2 genera following the key reveals very few comparative statements. Spencer (1930: 418) stated Encrinaster may be distinguished by comparatively long thin arms containing many segments and that marginal disc plates of Encrinaster bear long spines. The number of segments in each arm determines the length ofthe arm and the number increases with growth so this is a very growth related difference; measure- ments of all illustrated specimens of the 2 genera fail to show any clear cut differentiation. These linked and growth related features do separate the forms that Spencer (1930) figured in the work in which he erected the second genus. Similarly the long spines on the marginal plates in Е. grayae are not evident in the type species or in most other species of the genus. Thus the features quoted by Spencer (1930) are parochial in their discriminatory application and are not suitable generic features on a broader scale. Thus we synonymise Encrinaster and Euzonosoma. Encrinaster tischbeinianus (Roemer, 1863) (Figs 40-43) Aspidosoma tischbeinianum Roemer, 1863: 144, pl. 23, fig. la, b (not pl. 25, fig. 11): Schóndorf. 1910; 23 and synonmy listed therin. Encrinaster tischbeinianus (Roemer): Schuchert. 1914: 244, Euzonosoma tischbeinianum (Roemer). Spencer, 1930: 404; Lehmann, 1957: 25, pl. 4, figs 1. 4-6. Hexura weitzi Spencer. 1950a: 300, figs 4. 5 (not figs 1-3). MATERIAL. South Africa. B4500-4502, B4505, B4506, B4510, B4511, B4548 from the Voorstehoek Formation at Hottentots Kloof. SAMK1018 from Riet River N of Ceres on Wupperthal road. S side of farm which is also called Groote Rivier in the neighbourhood, from Voorstehoek Formation. SAM11908 from Gamkapoort. SUG299 from De Doors. Spencer's specimen (19502, figs 4,5). B4555 from Boplaas Farm in the Waboomberg Formation. B4563 from Swaarmoed Pass, 2.1km from Great Swaarmoed Farm, near Ceres. RUGDNH2 from the Tra-Tra Formation. DIAGNOSIS. Reaching large size (130mm arm length); with arms distally whip-like; disc with concave margins (of more than 11) large plates between arms; rest of disc of thin tiny tuberculate plates; 151 ambulacrals short and stubby; ambulacral groove wide across disc (apparently unable to close?); ambulacrals and adamb- ulacrals as boot-shaped ossicles with toes pointing at each other; podial basins circular, similarly sized throughout arm; width of arm determined by width of ambulacrals or attitude of adambulacrals; madreporite ventral, adjacent to mouth frame, with strong ridge ornament. DESCRIPTION. Overall form. Arms up to 130mm long (95mm max. in South African material), petaloid, widest at disc margin (up to 5mm), tapering strongly in distal part, distally whip-like. Disc pentastellate, with concave disc margins between arms, diameter 7-40mm, 0.5 or more of arm length, with margin of large irregularly polygonal plates sometimes extending a 2nd or 3rd row away from margin, remainder of surface a tuberculate apparently lightly mineralised integument, with larger tubercles close to mouth frame ventrally, especially in larger specimens. Arm plating. Within the disc arms fixed into the plating, with adambulacrals and ambulacrals on 166 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 41. Fnerinaster tischbehianus(Roemer, 1862), Ventral view of juveniles in both ventral and dorsal views, B4501 3.5. adult holotype with numerous fragmentary sanie level suggesting ambulacral groove may be beyond ambulacrals and laterally enclosing a wide open and unable to be closed. Towards disc wide. deep ambulacral groove. Within disc. margin adambulacrals extending ventrally ambulacrals wide. with boot-shaped ridges well EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA 167 x3. B. C, ventral views un i FIG 42. Encrinaster tischbeinianus(Roemer, 1862). A, dorsal view of incomplete B45 ^ B4500 and SAM11908, respectively х2. D, incomplete ventral view B4567 х4. SEUM F THE QUEENSLAND MU SO MEMOIR 168 EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA away from the arm axis; near the disc margin, boot-shaped ridges closer to axis but arm of same width; distally, boot-shaped ridges contiguous along arm axis; ambulacrals narrowing abruptly to produce the whip-like termination. Ambulacrals offset across arm axis; in dorsal view subquadrate, with narrow (becoming wider in larger specimens) but deep elliptical clefts between successive plates formed by concave proximal and distal faces for muscle insertion; proximal and distal margins raised as low ridges, with posterior rim extending abradially, with dorsal rim of concavity raised as a transverse ridge proximally, becoming less distinct distally as the interambulacral cleft becomes smaller and dorsal surface becomes subcylindrical (1.е., flat radially but curved transversely). Laterally this subcylindrical surface changes at a sharp line into a gently convex adradial side to the podial basin. Podial basin with floor having circular gap in the plates between ambulacrals and adambulacrals. In ventral view, sutural junction between the 2 columns of ambulacrals obtusely zigzag (c. 160°). Proximal arm with thin, low, uniform, radial ridge along ambulacrals on each side of and close to («0.5mm) arm axis; these ridges running to the suture along arm axis at about the disc margin. Abradial to this line each ambulacral with concavity facing arm axis and inclined on the side of the leg of the boot-shaped ridge. Prominently raised boot-shaped ridge with a strong constriction above the ankle and a long toe with distinct arch of the sole. Narrowest point (above the ankle) at midlength of podial basin on abradial side; small concavity (mentioned above) on adradial side. Abradial side concave, descending rapidly into the podial basin. Distally in the whip-like portion of the arm ambulacrals much narrower, lacking podial basins. Adambulacrals. Shape and orientation of plates varying along column, with concommitant change in the ambulacral groove; ambulacrals 1 and 2 without associated adambulacrals. Adambulacrals proximally in dorsal view subtriangular, flat, in the same plane as the interradial surface, with articulation against ambulacrals a point separated from distal point by curved margin to aperture in floor of podial basin. Distinct groove parallel to axis along dorsal surface close to abradial margin, 169 continuing along 6 plates beyond the disc margin, petering out as dorsal view of adambulacrals becomes progressively narrower. Adambulacrals changing from horizontal to almost vertical approaching disc margin, developing en echelon rather than linear arrangement; on | arm of the holotype 3 adambulacrals with bases of small spines attached to distal face (no other spines known). Adambulacrals almost vertical and subquadrate in lateral view just beyond the disc, distally becoming thinner, closer to the radial axis and enclosing arm more completely. In ventral view, adambulacrals within disc as prominent L-shaped ridge with toe of base abutting against toe of boot-shaped ridge on ambulacral, with longer sections aligned and forming abradial margin of arm. Transverse ridge formed by toes of adjacent ambulacral and adambulacral almost knife-edged, of uniform height across both plates, giving very quadrate appearance to arms. Along abradial margins the continuity from plate to plate gives a sharp ridge along the whole arm to the disc margin. Gently sloping platform on adradial side of the L-shaped ridge descending into podial basins, formed by adradial face of adambulacral with ventral face directed laterally in this section of the arm. Approaching the disc margin podial basins smaller and deeper and adambulacrals arranged en echelon in column continuing so distally as adambulacrals become smaller but enclose the ambulacral groove more. Mouthframe. lst and 2nd ambulacrals conspicuous; tori and denticles rarely evident (dorsal margin of | torus (Fig. 42A) and a set of 3 denticles attached to a torus (Fig.41) are evident). Ist ambulacral with proximal ends vertical, bluntly pointed; groove for nerve ring extremely well impressed distal to where 1st Amb becomes higher and wider; groove for water ring wider and shallower, remaining on the mouthframe plates across the radial and interradial sutures so that the groove is evident as a full ring, with 2 pores in the groove on each 2nd ambulacral near the radial line of each arm. At the interradial junction between Ist ambulacra the water ring groove descends into the junction, creating a subcircular basin. 2nd ambulacral expanded only slightly distally, barely overlying any of 3rd ambulacral. FIG. 43. Encrinaster tischbeinianus (Roemer, 1862). A-B, large incomplete individual in ventral and dorsal views, respectively, RUGDNH2 x1.2. C,D, juvenile in ventral view, B4500 x2. E, juvenile in ventral view, B4548 x3. F, juvenile in dorsal view B4502 x7. G, group of juveniles, B4500 x2. H, group of juveniles, with adult at bottom, B4502 x3. 170 In ventral view, proximal faces of adjacent Ist ambulacrals forming high concave recess to accommodate tori. 1st ambulacral extending well ventrally at edge of the mouth, descending steeply distally; suture between Ist and 2nd ambulacrals about halfway down this slope at widest point. 2nd Amb with small shallow podial basin on distal adradial corner and beginning of sharp longitudinal ridge of adambulacrals on distal abradial corner. Distal face of 2nd ambulacral near arm axis with small concave facet opposing the same on 3rd ambulacral, both for insertion of longitudinal muscles. Madreporite. Interradially on ventral surface, just to the right of and contiguous with 2nd ambulacral, oval, outwardly convex, with irreg- ular straight and curved grooves peripherally. Disc. Dorsal surface a tessellated pavement of thin subquadrate to irregular plates each bearing a rounded tubercle, with margin of superior and inferior series of larger, thicker, differentiated plates without tubercles and with well defined sutural interplate boundaries. A few such plates are rarely seen in the second row from the margin. REMARKS. Dorsally, available specimens show the plating of the mouthframe and arms without any suggestion of integument covering them; in life there must have been some sort of integument over the central mouth area and this probably extended out over the arms as well. It is probably of some unmineralised tissue that does not fossilise. How such an unmineralised integument related to the interradii is not clear; the obvious arrangement would see a ‘skin’ enclose the whole disc but then the reason for tubercles on plates becomes unclear if they are internal. If the integument covered only the mouth frame and arms the arrangement for its connection to the body between adambulacrals and interradial plates and its method of growth are obscure. Adambulacrals within disc have sutural junctions with the interradii on dorsal and ventral surfaces so the height of the body could not have been more than 1-2mm at this point as that is the height of the abradial wall of adambulacrals; this would leave a very flat body cavity within the disc and adds support to the possibility of an outer integument covering the entire body except for the ambulacral grooves and mouth. MEMOIRS OF THE QUEENSLAND MUSEUM This South African material conforms closely to the German Е. tischbeinianus from the Lower Devonian Hunsrückschiefer in all features including relative disc size and shape, number of marginal disc plates, arm shape, relatively widely separated ambulacral column in each arm, and size and shape of the mouth frame. Considerably larger specimens are known from Germany but this disparity may be accounted for by the amount of collecting and by the German slates being a better matrix for yielding large whole specimens; in the coarser South African matrix the chances of obtaining large specimens is reduced. The finely tuberculate ornament on the disc is not evident on the German material but the slatey cleavage would be expected to obliterate such fine ornament. We consider all these discrepencies due to differences in preservation or intraspecific. A number of very small specimens among the South African material give some information on growth of the species. In general the small specimens are virtually identical to the larger ones except that the mouth frame is not so robust, particularly in dorsal view; the arms appear to be ofuniform width and the abradial aligned parts of the adambulacrals remain narrow and knife-edged throughout instead of thickening up and becoming en echelon arranged as in the larger specimens. Marginura Haude, 1999 not Marginaster Perrier, 1881; Marginaster Haude, 1995: 63. Marginura Haude, 1999: 1. TYPE SPECIES. Encrinaster yachalensis Ruedemann, 1916 from the Lower Devonian of western Argentina. DIAGNOSIS (from Haude, 1995). Encrinaster- inae with mosaic plated dorsal surface and interradii, concave margins to disc, concave outer margin of podial basin which has a round upper lamella. REMARKS. This genus is only known from South America (Haude, 1995) and now South Africa. FIG. 44. Marginura hilleri sp. nov. A, ventral view of disc including slightly disarticulated oral area B4566 x3. B, dorsal view of incomplete specimen RO P84B x3. C, ventral view of parts of 2 arms and an interradius with only marginal plating of latter remaining RO P84C x5, D, ventral view of 2 arms, an interradius and the oral area with plates adjacent to mouth mainly disociated and possible madreporite at lower right RO E11 x3. 171 EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA Y y © » Е Au A ates Es A | h 172 Marginura hilleri sp. nov. (Figs 44, 45) ETYMOLOGY. For Dr Norton Hiller who contributed material for this study. MATERIAL. HOLOTYPE: B4566 from Swaarmoed Pass, Ceres, 1.3 miles from Great Swaarmoed Farm. RO Ell from Damascus, Prince Albert at 33? 17'15"S; 21° 55'45"E. ROP84 from Swaarmoed Pass, Ceres at 33°21°30"S; 19°30°30"E. DIAGNOSIS. Disc surface a mosaic of poly- gonal plates; disc plates larger and subquadrate near the margin, much smaller, less regular and bearing strong circular tubercles proximally. Adambulacrals with club-shaped abradial expansion, with L-shaped ridge running along proximal and then abradial margins. Ambulacral 2 extending dorsally over 3rd and 4th and with pointed distal extremity (not truncated). Abradial spatulate spines on arms apparently continuing along interradial disc margin. DESCRIPTION. Overall form. Arms up to 40mm long, tapering distally, distally whip-like. Disc pentastellate, with concave disc margins between arms, with margin of large irregularly polygonal plates bearing marginal spines (2 per marginal), remainder of surface of small irregular tuberculate plates. Arm plating. Within the disc arms fixed into the ventral plating. Distally adambulacrals extending further ventrally to enclose a narrowing but deeper ambulacral groove. [Within disc ambulacrals wide, with boot-shaped ridges well away from the arm axis; near the disc margin boot-shaped ridges closer to axis but arm of same width; distally boot-shaped ridges contiguous along arm axis; ambulacrals narrowing dramat- ically to produce the whip-like termination. ] Ambulacrals offset across arm axis, with straight intercolumn suture along arm axis, proximally much wider than long in dorsal view but in 2 distinct sections: 1, a subquadrate raised adradial section with 1-2 tubercles on a fine granular background ornament and 2, an abradial section down a distinct abradial slope (abradial section becoming narrower distally along arm and disappearing at beginning of whip-like section), with wide deep elliptical clefts between successive plates formed by concave proximal and distal faces for muscle insertion; proximally dorsal proximal and distal margins with fine low ridges. In ventral view arm axis with straight sutural junction and distinct ambulacral channel MEMOIRS OF THE QUEENSLAND MUSEUM between the 2 columns of ambulacrals; each ambulacral concave towards arm axis (i.e. back of leg of boot shape). Prominently raised boot-shaped ridge with short leg, strong constriction above the ankle and a long toe with distinct arch of the sole; with narrowest point (above the ankle) at midlength of podial basin, with concave abradial side descending rapidly into the podial basin. Ambulacrals much narrower distally in the whip-like portion of the arm, apparently lacking podial basins. Adambulacrals. Shape and orientation of plates varying distally along column, shaped like the head of a large golf club (a *wood') with transverse projection (the shaft of the golf club) abutting the toe of the ambulacral ‘boot’; ambulacrals 1 and 2 without associated adambulacrals. Adambulacrals subquadrate in dorsal view, with articulation against ambulacrals at 2 points (anterior and posterior), with concave adradial margin in between combining with concave abradial margin of ambulacrals to define perforation through bottom of podial basin. Adambulacrals vertical throughout, forming lateral wall of arm. In oral view, adambulacrals very thickened abradially, with continuity from plate to plate of expanded abradial parts producing sharp ridge along the whole arm being base of lateral wall, with gently sloping platform on adradial side descending into podial basins, with short wide spatulate lateral spines. Mouthframe. lst and 2nd ambulacrals fused into large unit as typically forms mouth frame throughout family but suture between them not evident on available material, dorsally overriding 3rd and 4th ambulacrals; tori and denticles not evident but a few acicular plates in oral region of one specimen (Fig. 44B) may be disaggregated denticles, with small narrow proximal ends; groove for nerve ring shallow and crossing paired Ist ambulacrals just proximal to groove for water vascular ring; groove for water ring wide, well-impressed, remaining on the mouthframe plates across the radial and interradial sutures (so that the groove is evident as a full ring), with 2 pores in the groove on each 2nd ambulacral near the radial line of each arm. In ventral view, proximal faces of adjacent Ist ambulacrals forming relatively small concave recess to accommodate tori. 2nd Amb with small deep podial basin on distal adradial corner. Distal face of 2nd ambulacral near arm axis with small concave facet opposing the same on 3rd ambu- lacral, both for insertion oflongitudinal muscles. EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA 173 FIG. 45, Marginura hilleri sp. nov. RO P84a & b, prie pene holotype. 2 incomplete arms. an interradius and some oral plates. A, dorsal view «5. B, enlargement of lower right of C showing madreponte (left) and short spiny interradial plates disarticulated and lying on their side «7. C. ventral view x3. 174 Madreporite. Interradially on ventral surface, suboval, with ornament of vermiform groove defined by sharp ridges. REMARKS. This species is very close to the type and only congener, M. vachalensis but may be distinguished by its marginal spines, the pointed rather than truncated distal dorsal tips of the second ambulacrals, the course ofthe sharp ridge on the ventral side of the adambulacrals and the size ofthe tubercles or small spines on the ventral interradial disc plates, particularly proximally. The new species does not contradict any of the structural interpretation of Haude (1995). Family PROTASTERIDAE Miller, 1889 Eugasterella Schuchert, 1914 TYPE SPECIES. Eugasterella logani (Hall, 1858) from the Middle Devonian Hamilton Group of New York. DIAGNOSIS. Arms 5, long, slender, tapering distally, without dorsal arm plates or dorsal spines. Disc circular, inflated; dorsal surface of polygonal plates covered by finely granular integument. Madreporite ventral. Mouth frame large; Ist ambulacral long, narrow; 2nd ambulacral large, stout, with well-developed grooves, apophyses and pores dorsally for directing water vascular and nervous systems. Ambulacrals boot-shaped in ventral view; ambulacral groove slightly sinuous; dorsal surface with wide deep excavations for dorsal longitudinal muscles; podial basins large and well-defined. Adambulacrals roughly ear- shaped, narrow, wrapped around sides of ambulacrals, with large nodes for attachment to toe of boot on ambulacrals, with vertical spines. REMARKS. This diagnosis follows that of Harper (1985) with emendation as inferred by Hotchkiss (1993) who diagnosed Strataster, emphasising features of dorsal arm structure. Eugasterella africana sp. nov. (Figs 46-48) Hexura мей Spencer, 1950a: fig. 3 [not figs 1.2.4-6]. ETYMOLOGY. From A frica. MATERIAL. HOLOTYPE: B456la (a,b, dorsal and ventral external moulds). PARATYPES: B4561b and c all from Klipfontein near Swaarmoed Pass, Ceres, B4569 from Hex Rivier Pass on Montagu Road 14km from N9 turnoff and RO123 from Matroosberg, Worcester(Hex River Pass) (33°30°5, 19° 48°E); all from the Voorstehoek formation. SAMK1014, 1015 from Verstehoek Formation at Riet Rivier (i.e. Gydo Formation, N of Ceres Division on MEMOIRS OF THE QUEENSLAND MUSEUM Wupperthal Road; S side of farm which is also called Groot Rivier in the neighbourhood. DIAGNOSIS. Arms slender, tapering gently throughout. Disc subrounded to subpentangular, inflated dorsally; dorsal surface of thin plates covered by a finely tuberculate integument, without spines. Short spines in a central marginal triangle on ventral interradii. Ambulacrals boot-shaped, with leg and foot of about same length. Adambulacrals with row of spatulate spines along ventral edge and 1-2 vertical spines. Madreporite circular with marginal aperture around half circumference. DESCRIPTION. Overall form. 5-armed, disc radius 5-8mm, arm length up to 35mm. Arms slender, 2-3mm wide at disc margin, tapering distally throughout (no arm tip available). Disc circular to subpentagonal, gently inflated dorsally, with dorsal surface of small thin irregularly shaped plates covered by finely granular integument, without any spines on dorsal surface; ventral interradii triangular, with same finely granular integument as dorsal surface covering a lattice-like arrangement of plates increasing in aperture sizes towards margin of disc, with short stout spines in triangular zone involving entire disc margin and a third corner interradially about 1/3 disc radius away from margin. Arm plating. Dorsal surface of arms circular in section, with many small thin irregular plates possibly elongate across arm axis and irregularly with small nodes. Ambulacrals offset along arm axis; in dorsal view subtrapezoidal, with wide deep excavations proximally and distally for longitudinal muscles, with prominent (decreasing in prominence distally) transverse ridges proximally and distally above deep cleft for muscle attachment, slightly sinuous axial line formed by concave adradial margin on each ambulacral into which a proximal and distal tip of successive ambulacrals from the opposite column project, the sinuosity decreasing distally; ventrally boot- shaped, with leg and foot of about equal length; ambulacral groove shallow, slightly sinuous as in dorsal aspect; podial basin round, deep, almost entirely on 1 ambulacral with corresponding adambulacral forming outer side to basin. Adambulacral abradial to and corresponding 1 to 1 to ambulacrals, planar, roughly ear-shaped, oriented vertically to form the lateral walls of the arm; in dorsal view only slightly abradially convex dorsal edge around short abradial edge of ambulacrals; in ventral EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA 175 FIG. 46. Eugasterella africana sp. nov. A-B, ventral and dorsal views, respectively, of holotype B456 la & b #4, C-D, dorsal and ventral views. repectively. of specimen with dorsal plating removed thus exposing proximal ambulacrals, tori and oral denticles (not evident with dorsal surface in place as in B) RO123a & b x4. 176 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 47. Eugasterella africana sp. nov. A, dorsal view B4561E x5. B, dorsal view with some longer spines in marginal interradial areas B4569 x2.4. C, ventral view showing madreporite and marginal interradial spines 4561р, x6. D. dorsal view B4561C хб. EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA FIG. 48. Eugasterella africana sp. nov., ventral views. A, SAM K1015 x3. В, B4561B x4. C, SAM K1014 x3. view L-shaped. formed by strong lateral projection abutting toe of ambulacrals and narrow ventral edge in proximal- distal line, with line of 5 pits along narrow ventral edge each bearing short spatulate ventrally directed spine, with strong pointed spine directed distally arising from narrow platform formed by interruption to ventral edge towards distal end. Madreporite. Ventral, to left in CD interradius, circular, convex, with broadly U-shaped slit-like 177 opening close to and around 1/2 the circumference facing the D ray. Mouth frame. In dorsal view occupying about 1/2 disc area: Ist ambulacrals in contiguous pairs interradially, elongate radially, straight. with transverse groove for nerve ring, pointed distal tip. with junction to 2nd ambulacral at 45? on outer distal bevelled face. Torus biconvex, fitting into concave proximal end of each pair of Ist ambulacrals, with horizontal row of 5 flat blunt 178 spines projecting into mouth (central 3 widerthan others); 2nd ambulacral large, forming strong V distally along arm, extending over next 2 or 3 ambulacrals, with groove for water vascular system on proximal side, extending along proximal 1/2 across distal ends of Ist ambulacrals, with 2 pits in groove on each plate (leading to podial basins of 2nd and 3rd ambulacrals. In ventral view Ist ambulacrals paired, forming narrow Vs, not contiguous as in dorsal view, straight, with concave proximal face, abutting 2nd ambulacrals on abaxial side of podial basin; 2nd ambulacrals with well- developed podial basin beginning column of basins of each arm; reduced adambulacral with 5 spatulate ventral spines not visible in dorsal view where covered by expanded 2nd ambulacral. REMARKS. This species differs from the type, E. logani (Hall, 1858), from the Middle Devonian of New York in having marginal interradial spines on the disc, ambulacrals shorter and with deeper longer dorsal clefts between successive ambulacrals in each column and circular madreporite with slit-like aperture around most of its margin; it resembles more the only other assigned species, Е. devonicus (Kesling, 1972) from the Middle Devonian of Ohio, in structure of the disc and in spinosity of adambulacrals although it appears to have a maximum of 2 vertical spines on each adambu- lacral as opposed to 4 in the North American species and may be further distinguished by the structure of the disc dorsally, which in Е. devonicus was described by Kesling as having no discernible plates, thickly studded with small grains probably marking the position of papillae and bristly with short erect little spines. Harper (1985, fig. 5) described and figured the dorsal surface of the disc of E. logani as com- posed of overlapping polygonal plates bearing raised biaxial ridges. The South African species suggests that these ridged plates are part of the ventral interradial plating (Figs 46C, D) which are evident in both dorsal and ventral moulds of specimens where the dorsal surface of the disc has been removed. We would interpret Harper's figure (1985, fig. 2C) as representing a specimen in which the dorsal surface of the disc was well-preserved only in the interradial triangle on the right of the specimen as viewed; some dorsal surface is evident in the other interradial areas except that at the upper left where the ventral or inner surface of the ventral plating is evident. MEMOIRS OF THE QUEENSLAND MUSEUM Strataster Kesling & Le Vasseur, 1971 TYPE SPECIES. Strataster ohioensis Kesling & Le Vasseur, 1971 from the Lower Carboniferous of Ohio; by original designation. DIAGNOSIS. See Hotchkiss (1993:64). Strataster ohioensis Kesling & Le Vasseur, 1971 (Figs 49-53) MATERIAL. South African material - B4509, B4512, B4513, B0195 -0197, B0276 and PRVT82 all from Boplaas Farm, N of Ceres in the Waboomberg Formation. REMARKS. Some ofthese external moulds were coated with a thin layer of varnish after collection; this made them less absorbent and thus the latex casting has not been as successful as with other unvarnished material. The dorsal crest of each arm usually has a row of small air bubbles where the latex did not penetrate into the moulds of the carinal spines, but these spines are cast ina few places (Fig. 50C, 51B). The papillate dorsal disc integument is well preserved in several places draped over the strong mouth frame exactly as with Kesling and Le Vasseur's (1971) material; this indicates that the disc was distended with fluid or other soft tissue during life but had no solid material in it and no detritus in the gut. This is in contrast to Eugasterella africana where the disc has remained inflated indicating some solid disc filling such as sediment detritus or some structural integrity possibly conferred by the dorsal plating beneath the papillate surface. This material agrees with the species described by Kesling & LeVasseur (1971) in every respect and as those authors gave an extremely detailed description there is nothing further to add. In particular, the upper arm plates and row of carinal spines, considered by Hotchkiss (1993), to be generic features occur in the South African material. The dorsal clefts between ambulacrals appear sharper in some of the North American specimens but in the South African material the small dorsal arm plates are preserved in most places and the varnish applied to the external moulds has in most cases diminished the height of the ridges adjacent to the dorsal clefts. The South African occurrence extends the range to become Early Devonian to Early Carboniferous (Tournaisian) and the geographic extent to include Laurentia and Gondwanaland encompassing the Appalachian and Malvinokaffric EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA 179 FIG. 49, Strataster ohioensis Kesling & Le Vasseur, 1971, group of 3 individuals B4513 (C) x3. A. lower left specimenin dorsal view x5. В. upperspecimen in ventral view x5. D, lowerright specimen in ventral view «5. 180 MEMOIRS OF THE QUEENSLAND MUSEUM i T4 z s 7 FIG. 50. Strataster ohioensis Kesling & Le Vasseur. 1971. A-B, dorsal views В4509Д & B. х4. C. dorsal view РКУ T82A, #4. D, ventral view showing madreporite B0276, x4. EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA 181 FIG. 51. Strataster ohioensis Kesling & Le Vasseur, 1971. A, ventral view РКУ T82B, x6. B, enlargement of 2 arms from Fig, 37A to show the median row of dorsal spines on each B4513, x10. C, ventral view B0196, x4, D, ventral view B0276, x4. 182 MEMOIRS OF THE QUEENSLAND MUSEUM VIG. 52. Strataster ohioensis Kesling & Le Vasseur, 1971. A, ventral view B4509C, х5, B, dorsal view B0197, x4. C. dorsal view B4509D, х5. D, ventral view with some of ventral disc plating removed so exposing dorsal plating from the interior of B4513, х4. EARLY DEVONIAN ECHINODERMS OF SOUTH AFRICA FIG. 53. Strataster ohioensis Kesling & Le Vasseur, 1971, ventral view of large individual B4512 x3. Provinces of brachiopod palaeobiogeography (Boucot et al.. 1969). Some of the material referred to by Hotchkiss (1995) in his Appendix 1 at numbers 108-112 is included in this species. Strataster stuckenbergi (Rilett, 1971) (Figs 54-56) Taeniaster stuckenbergi Rilett, 1971: 32, figs 3-4. MATERIAL. HOLOTYPE: NM Type 1550 on NM831 from near Ceres. OTHER MATERIAL: B4504 from the Waboomberg Formation at Theronsberg Pass, B4533, B4536 from the Voorstehoek Formation at Matroosberg. DIAGNOSIS. Row of carinal spines on arms not extending onto disc or to arm tips. Disc integument dorsally and ventrally (interradii) with irregularly spaced spines on papillate 183 surface. Ambulacrals with short leg and longer foot to boot in ventral view. Adambulacrals with 6 or 7 curved, spatulate, ventral (or oral) spines, with 2 or 3 strong, straight, circular-sectioned lateral (vertical) spines. Madreporite large. circular. with slit-like aperture adjacent and parallel to margin for about 3/4 circumference. DESCRIPTION. Size and shape. Average disc diameter among available specimens 15-20mm. Disc apparently circular: a few specimens suggest the disc may be extended slightly down each arm to give a substellate shape but this is probably due to postmortem movement. All available arms are preserved in curved position but are estimated to be 40-50mm long: arms widest at edge of disc (up to 4mm), tapering strongly distally to be width) and depressus (width > length). We thus assign them to the broader species concept. When Teichert (1949) identified Storthingo- crinus there is a distinct possibility that he had material of this species because the plating arra- ngement is identical: however, the radial facets of 230 Storthingocrinus are quite different and it has been suggested that it is a camerate crinoid (Prokop & Petr, 1997). Subclass CLADIDA Moore & Laudon, 1943 Family CODIACRINIDAE Bather, 1890 Codiacrinus Schultze, 1867 TYPE SPECIES. Codiacrinus granulatus Schultze, 1867 from the Middle Devonian of Germany; by original designation. REMARKS. This genus was discussed by Jell (in Jell & Holloway, 1983:16); it contains 7 species, C. granulatus, C. schultzei Follmann, 1887, C. procerus (Prokop, 1973), C. ornatus (Prokop, 1973) (probably a junior synonym of C. granu- latus), C. rarus Jell in Jell & Holloway, 1983, C. nicolli sp. nov. and C. secundus Jell, 1999. Codiacrinus nicolli sp. nov. (Fig. 27) ETYMOLOGY. For Robert Nicoll who collected some of the material. MATERIAL. Holotype: WAM91.710 from QML1929. Paratypes: CPC34566-34577 from section 354 (9m level) on W side of McWhae Ridge (Nicoll & Playford, 1993). Other Material: QMF36204-36206 from QML1031, E side of Bugle Gap S of Wagon Pass. DIAGNOSIS. Cup small, with granular orna- ment but no ray ridges; basals small, pentagonal, almost equidimensional, with proximal margin shorter than others; radials long, with angustary radial facets, with strong distal projections both sides of facet. DESCRIPTION. Cup small, up to 11mm long and 8mm diameter, subglobose to subcylindrical, with broadly flared basal circlet, of very thick plates (body cavity less than 1/2 thecal diameter). Infrabasals 3 (in Fig. 22E there appear to be only 2, but it is a weathered base and the positions of the visible sutures suggest that the 3rd suture has been fused and thus the specimen aberrant), 2 large and equal and | small, separated by sutures in typical Y-shaped arrangement, with obtuse angle distally at base of sutures between basal plates, outflared away from stem. Basals pentagonal, with 4 equal sides and shorter prox- imal margin, up to 4mm across. Radials large, longer than wide, occupying most of theca; radial facet angustary, more than 1/2 radial width, sub- rectangular excavation into radials, with flat semicircular floor and convex butterfly-shaped inner surface, with lateral parts of radials of adjoining plates forming 5 projections distally. MEMOIRS OF THE QUEENSLAND MUSEUM No anal plates in theca. Stem circular in section, very small diameter, with fine central lumen. Arms unknown. REMARKS. Smaller size, type of radial facet, small infrabasal circlet, outflared infrabasal and basal circlets and stem diameter much less than that of cup distinguish this species within the genus. It is probably most similar to the type species particularly in comparison with Schultze's (1867, pl. 3, fig. 9C) second specimen which is more cylindrical than globose. It is quite distict from the other Australian species in the Pridoli and Lochkov of Victoria in its stem size, size of infrabasals, radial facets and ornament. Subclass FLEXIBILIA Zittel, 1895 Order TAXOCRINIDA Springer, 1913 Superfamily TAXOCRINOIDEA Angelin, 1878 Family TAXOCRINIDAE Angelin, 1878 ?Taxocrinus sp. (Fig. 28D) MATERIAL. QMF40360 from QML1031. DESCRIPTION. Cup plates smooth. Infrabasals 3, azygous in C ray, visible externally, forming narrow margin to stem facet. Basals 5, pent- agonal, equidimensional except posterior one; posterior basal much longer than others and also wider, hexagonal, with distal margin weathered and unclear but apparently with distal lateral corners curving towards axis of cup around a central semicircular part of margin that could be part ofan aperture that may lead into an anal tube. REMARKS. This basal cup fragment is too in- complete for species identification. The distal end of the posterior basal suggests the beginning ofa tubular structure as in an anal tube suggesting the Taxocrinidae. Within that family, Taxocrinus Phillips in Morris, 1843, which ranges from the Middle Devonian to Lower Carboniferous of Europe and North America, has a symmetrical posterior basal leading directly into an anal tube and also has the stem facet restricted to the infrabasal circlet. However, it is retained in open nomenclature because it is so incomplete. ECHINODERMS FROM THE CANNING BASIN 231 FIG, 27. Codiacrinus nicolli sp. nov. all from WCB354/9 (Nicoll & Playford, 1993) except M from QML 1029 and N, О from QML 1031. A-D, CPC34566. х3. A, lateral view. В. distal view. C, oblique lateral view. D. proximal view. E. proximal view of CPC34567, х3. F, distal view of broken theca showing thickness of shell. CPC34568, х3. G, lateral view of CPC34569, x3. Н, distal view of CPC34570, x3. LJ, lateral and proximal views of CPC34571, x3. K,L, distal and oblique views of C PC 34572, х3. М, lateral view of WAMO91.710. x7. N, О, proximal and lateral views of QMF36204, х5. VIG. 28. A-C, Forbesiocrinus sp. ^, proximal view of basal and radial circlets, NMVP100323 from NMVPL1930, x4. B, internal view of base of cup showing infrabasal, basal and radial circlets, NMVP100324 from NMVPL 1929, х4. C, weathered section through whole animal showing differentiated stem and strongly incurved and coiled arms, QMEF36207 trom the Virgin Hills Formation W of Hull Range, x2. D, ?Taxocrinus sp., basal view of'cup fragment with infrabasal circlet visible around stem facet and large posterior basal at upper left QMF40359 from QML 1031, х4. MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 29. Cyclocystoid indet., silicified marginal ossicle, QMF36209, x3, [rom Frasnian part of Sadler Limestone SW of Wade Knolls, Paddy's Valley. A, B, ventral views of marginal ossicle tilted slightly differently to show the cupule zone and eniries to radial ducts (A) and crest (B) more clearly. C, dorsal view showing entries to radial ducts. D, lateral view showing high rounded crest, cupule zone and circumferential canal. Order SAGENOCRINIDA Springer, 1913 Superfamily SAGENOCRINOIDEA Roemer, 1854 Family SAGENOCRINITIDAE Roemer, 1854 Forbesiocrinus sp. (Fig. 28A-C) MATERIAL. NMVP100323 from NMVPL1930 and NMVP 100324 [rom NMVPL1929 and probably QMF36207 from the Virgin Hills Formation W of the Hull Range. DESCRIPTION. Cup with flat base, plates with tuberculate ornament. Infrabasals 3, equal, pent- agonal, concealed within stem facet. Basals 5 pentagonal, with only distal triangular tips visible laterally forming margin to stem facet: posterior basal with an extra side distally, sup- porting 2 anal plates, separating C and D radials, Radials 5, hexagonal, in contact with each other except in posterior interray; radial facet plenary. Stem tapering slightly distally, of extremly short columnals proximally, becoming heteromorphic ECHINODERMS FROM THE CANNING BASIN 23 (alternating long and short columnals with angular latus) distally. REMARKS. Assignment to Forbesiocrinus is based on the posterior basal supporting 2 anal plates apparently symmetrically but the lack of arms prevents meaningful comparison with other species of the genus. The specimen in section from the Hull Range is doubtfully referred to this taxon but if correctly interpreted has the infra- basals completely concealed by the stem facet. In the other known flexible crinoid from the basin the infrabasals are evident laterally. Class CY CLOCYSTOIDEA Miller & Gurley, 1895 Family CYCLOCYSTOIDIDAE S.A. Miller, 1882 Cyclocystoid indet. (Fig. 29) MATERIAL. QMF36209 from the Frasnian part of the Sadler Limestone SW of Wade Knolls in Paddy's Valley. DESCRIPTION. Single marginal ossicle Imm wide, 2mm in radial length and 1mm high, with bevelled lateral margins indicating that the marginal ossicles were not in contact throughout their lateral margins. Crest high, with almost circular lateral profile, with ornament of rounded (in section) ridges aligned in parallel curves across crest. Cupule zone with 2 circular cupules each with strong circular central tubercle, with sharp ridge between cupules, with narrow deep circumferential channel, with 2 relatively large radial ducts from centre of each cupule. Dorsal surface smooth. REMARKS. The features of this ossicle are clearly in line with the Cyclocystoidea (Smith & Paul, 1982) but within the class it does not appear to fitany genus. The youngest described genus is Sievertsia last known from the Middle Devonian of Europe but that genus has flat or concave crests and dorsal surfaces and cannot accept the Australian ossicle. The ornament on the crest is unknown in any genus and this ossicle probably represents a new genus but it is retained in open nomenclature pending more complete material. Smith & Paul (1982: 677) reported an occurrence of the class in the Frasnian of Iowa, commun- icated to them by Terry Frest (pers. comm. 1980) but without illustration comparison is not pos- sible. However, the occurrences in Iowa and Western Australia are the youngest known occur- rences of the class and are valuable knowledge for that reason. Uu ACKNOWLEDGEMENTS For providing material we are grateful to Alex Cook, Queensland Museum, Phil Playford and Tony Cockbain, Geological Survey of Western Australia, Annette George, University of West- ern Australia, Scott Brownlaw, University of Queensland, Michael House, University of Southhampton and Thomas Becker, Humboldt University, Berlin. We thank Scott Brownlaw, University of Queensland, Alex Cook, Queens- land Museum and Gary Webster, Washington State University for their good company and collecting skills in the field. We thank Peter Murray, Museum and Art Gallery ofthe Northern Territory for hospitality and field support. LITERATURE CITED AUSICH, W.I. 1977. The functional morphology and evolution of Pisocrinus (Crinoidea: Silurian). Journal of Paleontology 51: 672-686. BECKER, R.T. & HOUSE, M.R. 1997. Sea-level changes in the Upper Devonian of the Canning Basin, Western Australia. Courier Forschungen- Institut Senckenberg 199: 129-146. BECKER, R.T., HOUSE, M.R. & KIRCHGASSER, W.T. 1993. Devonian goniatite biostratigraphy and timing of facies movements in the Frasnian of the Canning Basin, Western Australia. Geological Society Special Publication 70: 293-321. BREIMER, A. & DOP, J.A. 1975. An anatomical and taxonomic study of some Lower and Middle Dev- onian blastoids from Europe and North America. | and 2. Proceedings of the Koninklijke Nede- rlandse Akademie van Wetenschappen, Series B, 78: 39-61. BREIMER, A. & MACURDA, D.B. 1972. The phylo- geny of the fissiculate blastoids. Verhandelingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afdeling Naturkunde, Eerste Reeks 26(3): 1-390. BRIGGS, D.E.G. & ROLFE, W.D.I. 1983. New Con- cavicarida (New Order: ? Crustacea) from the Upper Devonian of Gogo, Western Australia and the palaeoecology and affinities of the group. Special Papers in Palaeontology 30: 249-276. BROWER, J. 1967. The actinocrinitid genera Abactino- crinus, Aacocrinus and. Blairocrinus. Journal of Paleontology 41: 675- 705. COCKBAIN, A.E. 1984. Stromatoporoids from the Devonian reef complexes, Canning Basin, Western Australia. Geological Survey of Western Australia Bulletin 129: 1-108. DRUCE, E.C. 1976. Conodont biostratigraphy of the Upper Devonian reef complexes of the Canning Basin, Western Australia. Bulletin of the Bureau of Mineral Resources, Geology and Geophysics 158: 1-303. DUBATOLOVA, Yu. A. 1964. Morskii lilii devona Kuzbassa [Devonian crinoids of the Kuznetz 234 Basin]. Akademiya Nauk SSSR, Sibirskoe Otdeleniye Trudy Instituta Geologii i Geofiziki 153p, 14pl. 1971. Morskie lilii Rannego i Crednego Devona Altay 1 Kuzbassa. [Crinoids of the Early and Middle Devonian of the Altay and Kuzbass.] Akademia Nauk SSSR, Sibirskoe Otdelenie Trudy Instituta Geologii i Geofiziki 124: 1-159. ETHERIDGE, R. Jr & CARPENTER, P.H. 1886. Catalogue of the Blastoidea in the Geological Department of the British Museum (Natural History), with an account of the morphology and systematics of the group, and a revision of the genera and species. (British Museum (Natural History): London). FAY, R.O. 1961. Blastoid studies. The University of Kansas Paleontological Contributions, Echino- dermata, Article 3: 1-147. FOLLMANN, O. 1887. Unterdevonische Crinoiden. Verhandlungen des naturhistorischen Vereines der preussischen Rhinelande, Westfalens und des Reg.-Bezirks Osnabruck, Vierundvierzigster Jahrgang 4: 113-138. GLENISTER, B.F. 1958. Upper Devonian ammonoids from the Manticoceras Zone, Fitzroy Basin, Western Australia. Journal of Paleontology 32: 58-96. GLENISTER, B.F. & KLAPPER, G. 1966. Upper Dev- onian conodonts from the Canning Basin, Western Australia. Journal of Paleontology 40: 777-842. GOLDFUSS, G.A. 1831. Petrefacta Germaniae. Vol 1, Part 2, Radiariorum reliquiae, pp. 115-221. (Arnz & Co: Dusseldorf). 1839. Bietrage zur Petrefactenkunde Acta Naturae. Curiosorum. Nova Acta Physico-medica Academie Caesar Leopoldino-Carolinae Naturae Curiosorum 19: 329-364, HILL, D. & JELL, J.S. 1970. Devonian corals from the Canning Basin, Western Australia. Geological Survey of Western Australia Bulletin 121: 1-158. HOROWITZ, A.S., MACURDA, J.B. & WATERS, J.A. 1986. Polyphyly in the Pentremitidae (Blastoidea, Echinodermata). Bulletin of the Geological Society of America 97; 156-161. JELL, P.A. 1999, Silurian and Devonian crinoids from central Victoria. Memoirs of the Queensland Museum 43: 1-114. JELL, Р.А. & HOLLOWAY, D.J. 1983. Devonian and ?Late Silurian palaeontology ofthe Winneke Res- ervoir site, Christmas Hills, Victoria. Proceedings of the Royal Society of Victoria 95: 1-21. JELL, P.A., JELL, J.S., JOHNSON, B.D., MAWSON, R. & TALENT, J.A. 1988. Crinoids from Dev- onian limestones of eastern Australia. Memoirs of the Queensland Museum 25: 355-402. KESLING, R.V. 1964. A new species of Melocrinites from the Middle Devonian Bell Shale of Michigan. Contributions from the Museum of Paleontology, University of Michigan 19: 89-103. LANE, N.G., WATERS, J.A. & MAPLES, С.С. 1997. Echinoderm faunas of the Hongguleleng MEMOIRS OF THE QUEENSLAND MUSEUM Formation, Late Devonian (Famennian), Xinjiang-Uygur Autonomous Region, People's Republic of China. Paleontological Society Memoir 47: 1-43 (Journal of Paleontology 71(2), supplement). LONG, J.A. 1991. The long history of Australian fossil fish. Pp. 337-428. In Vickers-Rich, P., Monaghan, J.M., Baird, R.F. & Rich, T.H. (eds) Vertebrate palaeontology of Australasia. (Pioneer Design Studio: Melbourne). MOORE, R.C. & TEICHERT, C. (eds) 1978. Treatise on invertebrate paleontology. Part T. Echino- dermata 2, 3 vols. (Geological Society of America & University of Kansas: Lawrence, Kansas). MORRIS, J. 1843. A catalogue of British fossils. Com- prising all the genera and species hitherto described; with reference to their geological dis- tribution and to the localities in which they have been found. (John van Voorst: London). MULLER, J. 1856. Über neue Crinoiden aus dem Eifeler Kalk. Koniglich Akademie des Wissenschaft Berlin, Monatsbericht 1856: 353-356. NICOLL, R.S. & PLAYFORD, Р.Е. 1993. Upper Dev- onian iridium anomalies, conodont zonation and the Frasnian-Famennian boundary in the Canning Basin, Western Australia. Palaeogeography, Palaeoclimatology, Palaeoecology 104: 105-113. PETERSEN, M.S. 1975. Upper Devonian (Famennian) ammonoids from the Canning Basin, Western Australia. Paleontological Society Memoir 8: 1-55. (Journal of Paleontology 49(5), Supplement). PHILLIPS, J. 1841. Figures and descriptions of the Palaeozoic fossils of Cornwall, Devon, and west Somerset. (Longman, Brown, Green & Longmans: London). PLAYFORD, P.E. 1981. Devonian reef complexes of the Canning Basin Western Australia. Field Excursion Guidebook, Fifth Australian Geol- ogical Convention, Perth. 44p. (Geological Society of Australia: Sydney). PLAYFORD, P.E. & LOWRY, D.C. 1966. Devonian reef complexes of the Canning Basin, Western Australia. Geological Survey of Western Aus- tralia Bulletin 188: 1-150. PROKOP, R.J. 1973. Elicrinus n. gen. from the Lower Devonian of Bohemia (Crinoidea). Vestnik Ustredniho Ustavu Geologickeho 48: 221-224. PROKOP, R.J. & PETR, V. 1997. The genus Pygmaeo- crinus Bouska, 1947 (Crinoidea, Inadunata) in the Devonian of the Barrandian area (Czech Repub- lic). Acta Musei Nationalis Pragae, Series B, Historia Naturalis 53: 1-10. REIMANN, I.G. 1945. New Devonian blastoids. Bulletin of the Buffalo Society of Natural Sciences 19(2): 22-43. RIGBY, J.K. 1986. Late Devonian sponges of Western Australia. Geological Survey of Western Aus- tralia Report 18: 1-59. ROEMER, С.Е. 1852. Bietrage zur geologischen Kenntnis des nordwestlichen Harzgebirges. Palaeontographica 3(2): 69-111. ECHINODERMS FROM THE CANNING BASIN 23 1855. Erste Periode, Kohlen-Gebirge. In Bronn, H.G. Lethaea Geognostica, vol. 2 (3rd ed.) (E. Schweizerbart: Stuttgart). ROZHNOV, S. 1981. Morskie lilii nadcemeistva Pisocrinacea [Sea lilies of the Superfamily Pisocrinacea]. Trudy Paleontological Institute 192: 1-127. SANDBERGER, G. & SANDBERGER, F. 1856. Die Versteinerungen des rheinischen Schichten- systems in Nassau. (Kreidel & Niedner: Wiesbaden). SCHULTZE, L. 1867. Monographie der Echinodermen des Eifler Kalkes. Denkschriften der kaiserlichen Akademie der Wissenschaft. Mathematisch- Naturwissenschafiliche Classe 26: 113-230. SEDDON, G. 1970, Frasnian conodonts from the Sadler Ridge-Bugle Gap area, Canning Basin, Western Australia. Journal of the Geological Society of Australia 16; 723-753. SMITH, A.B. & PAUL, C.R.C. 1982. Revision of the Class Cyclocystoidea (Echinodermata). Philo- sophical Transactions of the Royal Society of London B. Biological Sciences 296: 577-684. SPRINGER, Е. 1920. The Crinoidea Flexibilia, Smith- sonian Institution Publication 2501: 1-486. STRIMPLE, H.L. 1963. Crinoids ofthe Hunton Group. Oklahoma Geological Survey Bulletin 100: 1-169. STRIMPLE, H.L. & LEVORSON, C.O. 1973. Ad- ditional crinoid specimens from the Shellrock Formation (Upper Devonian) of Iowa. Proceed- ings ofthe Iowa Academy of Sciences 80: 182-184. TALENT, J.A., MAWSON, R., ANDREW, A.S., HAMILTON, PJ. & WHITFORD, D.J. 1993. Middle Palaeozoic extinction events: faunal and isotopic data. Palaeogeography, Palaeo- climatology, Palaeoecology 104: 139-152. TEICHERT, C. 1949. Observations on stratigraphy and palaeontology of Devonian, Western Portion of Kimberley Division, Western Australia. Report of the Bureau of Mineral Resources, Geology and Geophysics 2: 1-55. THOMAS, A.O. 1924. Echinoderms of the Iowa Dev- onian. lowa Geological Survey Annual Reports 1919 and 1920 29: 385-552. UBAGHS, С. 1978. Camerata. Pp. T408-T519. In Moore, R.C. & Teichert, C. (eds) Treatise on Invertebrate Paleontology. Part T. Echinodermata 3. (Geological Society of America & University of Kansas; Boulder & Lawrence, Kansas). VEEVERS, J.J. 1959. Devonian brachiopods from the Fitzroy Basin, Western Australia. Bureau of Mineral Resources, Geology and Geophysics Australia Bulletin 45: 1-220. WATERS, J.A. 1988. The evolutionary palaeoecology of the Blastoidea. Pp. 215-230. In Paul, C.R.C. & Smith, A.B. (eds) Echinoderm phylogeny and evolutionary biology (Clarendon Press: Oxford). WATERS, J.A. & HOROWITZ, A.S. 1993. Ordinal- level evolution in Blastoidea. Lethaia 26: 207-213. Un WHITEAVES, J.F. 1887. On some fossils from the Hamilton Formation of Ontario, with a list of the species at present known from that formation and province. Contributions to Canadian Palaeont- ology 1: 91-125. YAKOVLEV, N.N. 1949, Jaekelicrinus bashkivicus n. gen., n. sp. Journal of Paleontology 23: 435. APPENDIX Localities Register NMVPL1929. From more westerly of 2 stromatolitic limestone horizons mapped by Druce (1976, fig. 29) and Playford (1981, fig. 34) east of Millard Creek, 400-500m E of McWhae Ridge; collection made over 200m of strike 200-400m NNE of line of Section 4 of Druce (1976, fig. 29). Just NE of prominent westerly swing of creek this bed makes low ridge just above creek bank then after crossing creek outcrop area widens as its surface is exposed on low rising ground. GR4160 - 926262. Age: Although Druce (1976, fig. 29) mapped only 2 stromatolite horizons in his section 4 he mentioned 3 such beds in his text (Druce, 1976: 11). His first stromatolite bed is presumably west of Millard Creek and not mapped; his second stromatolite bed is the first stromatolite bed of Playford (1981: 42) based on the assigned ages ‘upper Palmatolepis triangularis’ zone (Druce, 1976: 11) and *im- mediately above the Frasnian-Famenian boundary' (Playford, 1981: 42). Fauna: Jaekelicrinus murrayi, Forbesiocrinus sp. NMVPL1930. From more easterly (i.e. younger) of two stromatolitic limestone horizons mentioned in siting NMVPL1929 and collecting from along a similar strike distance (200m) due E of that mentioned above for NMVPL1929. This horizon forms prominent line of ridges with E dip slope of 10-20 and W scarp over which Casey Falls pour. GR4160 - 926262. Age: Palmatolepis quadrantinodosa Conodont Zone of Druce, (1976: 11) or lower marginifera Conodont Biozone (Becker & House, 1997, fig. 9). Fauna: Jaekelicrinus murrayi, Forbesiocrinus sp. NMVPL1931 (=UQL3395 = GSWA21939 = OML1030). From red muddy carbonates of Virgin Hills Formation on left bank of creek above Casey Falls extending from near top of ridge of second stromatolite horizon E to sharp southerly bend in creek; collections from 30-40m of section almost immediately above stromatolite horizon. GR4160 - 926258. GPS location 18 Age: Palmatolepis quadrantinodosa Conodont Zone of Druce (1976: 12). Petersen (1975) assigned an age of do П within the Cheiloceras zone. This horizon equates to the “sponge garden facies’ of Becker & House (1997: 140, figs 7, 8) which they place in the Pernoceras delepenei Goniatite Biozone in the upper marginifera Conodont Biozone. Fauna: Wacrinus caseyensis, Wacrinus millardensis. NMVPL1938 (=K190). On GSWA track from Kelly’s Pass to Teichert Hills 200-300m N of90 turn from E to N 236 just NE of small prominent outlier of Permian Grant Formation (Playford, 1981, fig. 29; Playford & Lowry, 1966, plate 4); low lime knoll with some stromatolites. GR4160 - 933300. Age: Petersen (1975:53) assigned a probable age of do II and this equates to the late erepida or early rhomboidea Conodont Biozones which accords with the co-occurring gonialites. Fauna: Playfordierinus kellyensis. NMVPL1936 (=К 177). At first bend in creek downstream from spring due S of Teichert Hills (Playford & Lowry. 1966, plate 4). Rubbly outcrop above stromatolitic horizon. GR4160 - 942301, Age: Palmatolepis rhombotdea Conodont Biozone (do IT) (Glenister & Klapper, 1966: 838). Fauna: Jaekelicrinus murrayi, Playfordicrinus kellyensis, holdfasts, Catillocrinid indet. NMVPL/1939, 200-300m SSW of Millard Creek at S end (slightly W) of McWhae Ridge on ridge on left bank of minor left bank tributary marked by Druce (1976, fig. 29); 20m below base of Bugle Gap Limestone. GR4160 - 920256. Age: At level of Maenoceras Lsts (Becker & House, 1997, fig. 7) assigned to the lower marginifera Conodont Biozone. Fauna: Waerinus millardensis, Jaekelierinus murrayi. QML1031 (ZNMVPL 1940, = BC23-3 of Seddon (1970), = Тоб of Teichert (1949), = site of section 12 of Druce (1976)). On top of most southerly of 5 low hills stretching in a line (for about 1.5 km) SSW from Waggon Pass in Bugle Gap. GR4160 - 905355. GPS location 18 Age: Michael House (pers. comm.) assigns this locality to the Crickites lindneri Goniatite Biozone (Becker & House, 1997. fig. 8) which equates to the /inguiformis Conodont Biozone. Fauna; Hexacrinites brownlawi, Cudiacrinus nicolli, Hyperablastus. buglensis, Jaekelicrinus murrayi and Taxoerinus sp, NMVPL1942 (= BC4H-1 of Seddon (1970, р. 746)). From section at N end of Ngumban Cliff (Playford, 1981, fig. 29) MEMOIRS OF THE QUEENSLAND MUSEUM (i.e. E wall of S entrance to Bugle Gap, just N of Pinnacle Spring). Collection from some 40-50 m of section above lower stromatolite horizon. GR4160 - 891241. Age: Druce (1976, p. 16), in his Sechon 25, which is probably a parallel section, dated the lower stromatolite horizon in the Palmatolepis crepida Conodont Biozone (do IL) and the second stromatolite horizon in the basal Р, quadrantinodosa Biozone (do П). Very likely the rhomboidea Conodont Biozone. Fauna: Wacrinus millardensis, crinoid stems іп siromatolites. NMVPL1950-1956 (2T16 = WAPET Н = K495). Section between Margaret River and NeedleEye Rocks on first left bank tributary of first left bank creek from Margaret River N of Mount Pierre (Mount Pierre Creek); well exposed silty carbonates with cleaner limestone beds standing up above general outcrop near base, GR4061 - 042783 to 024776, 0-110m no fossils 18 А (= 1950) - 110-147т 18 B (= 1951) - 147-155m Jaekelicrinus murrayi, holdfasts. I8 C (= 1952) - 195-210m Jaekelicrinus murrayi, hold fasts. 18 D (= 1953) - 210-232m 18 Е (= 1954) - 372m 31 F (= 1955) - 382m 31 G (71956) - last 10m of sechon beneath first prominent grey limestone bench on NE side of Needle Eye Rocks; in head of gully opening to SE. Age: Most of the WAPET H section belongs to the Pseudoclymenia australis Ammonoid Biozone (Thomas Becker pers. comm. 1997) which equates to the lower iracliytera Conodont Biozone (Becker & House, 1997, fig. 8). QML1029. In bank of Millard Creek slightly N of W from Casey Falls, on the line of section B-C on figure 33 of Playford (198 1:35). GPS location 18 44.07'S, 126 05.18 F. Age: Very late. Frasnian, late Pelmatolepis linguiformis Conodont Biozone equivalent to the Crickites lindneri Ammonoid Biozone (Becker & House, 1997, fig. 9). Fauna: Melocrinites solidus sp. nov., Codiacrinus nicolli 5р, nav, NEW CARBONIFEROUS CRINOIDS FROM EASTERN AUSTRALIA GARY D. WEBSTER AND PETER А. JELL Webster, G.D. & Jell, P.A. 1999 06 30: New Carboniferous crinoids from eastern Australia. Memoirs of the Queensland Museum 43(1): 237-277. Brisbane. ISSN 0079-8835. New crinods are described from the Carboniferous of Queensland and New South Wales. Early Carboniferous faunas are dominated by actinocrinitids and platycrinitids. The geo- graphic distribution of Aacocrinus, Dialutocrinus, Sampsonocrinus, Litocrinus, Prininocrinus and Holcocrinus is extended with the first report of these genera from Austra- lia. A fauna from the Neerkol Formation of Queensland containing acrocrinids, an euspirocrinid and a scytalocrinid is the first Late Carboniferous fauna recognised from the non-equatorial belt or higher latitudes. New genera and species introduced are Denarioacrocrinus neerkolensis, D.? ornatus, Neerkolocrinus typus and Kopriacrinus mckellari. New species described are Aacocrinus acylus, Manillacrinus acanthus, Sampsonocrinus cannindahensis, Prininocrinus namoiensis and Holcocrinus barrabaensis. A neotype is designated for Synbathocrinus ogivalis. Australian Early Carboniferous crinoid faunas are most closely allied to North American faunas, but developed geographically widely separated from them. O Crinoids, Carboniferous, Queensland, New South Wales. Gary D. Webster. Department of Geology, Washington State University, Pullman, Washington 99164-2812, USA; Peter A. Jell, Queensland Museum, P.O. Box 3300, South Brisbane 4101, Australia; 14 June, 1998. The few Carboniferous crinoids described from Australia have been reported from NSW and Queensland. De Koninck (1878, 1898) reported 5 species from 2 unknown horizons at Burragood and Glen William, NSW, Etheridge (1892) described 3 camerates from the Mirari Limestone at Greenhills and 1 from Chalky Gully, NSW. This was followed by description of camerates and cladids from the ‘Gympie Beds’ (incorrectly assigned to the Permian initially), Queensland (Etheridge in Jack & Etheridge, 1892). Identifications were based on the broad concepts oftaxa at the time and, with few exceptions, were tentative at best. More recent reports of crinoids are based on moderately to well-preserved calyces and crowns, allowing more detailed identifications based on modern concepts of taxa. These reports have been an actinocrinitid calyx from Swain's Gully (Pickett, 1960), an acrocrinid from the Late Carboniferous of Queensland (McKellar, 1966), a Visean Physetocrinus and two unidentified inadunates from Queensland (Campbell & McKellar, 1969), camerates and inadunates from the Goonoo Goonoo Mudstone and Namoi Formation, NSW (Campbell & Bein, 1971) and a glaphyrocrinid and eumorphocrinid from the New England Fold Belt (Lindley, 1979, 1988). Carboniferous crinoid specimens that have remained undescribed in survey, university, museum and private collections have been drawn together for detailed study. These specimens add significantly to the known diversity and stratigraphic distribution (Table 1) of the Australian faunas. Our purpose is to: 1, describe the available specimens; 2, provide new data or interpretations of some of the earlier described material; and 3, relate all this material to known faunas elsewhere in the world. FAUNAL ANALYSIS We recognise 35 Early and 5 Late Carbonif- erous crinoids (Table 1) in Queensland and NSW. This does not include taxa based on stem segments, disarticulated cup plates and fragmentary specimens described by de Koninck (1878, 1898), Etheridge (1892) and Etheridge (in Jack & Etheridge, 1892). Several of the taxa in these reports are based on more complete specimens; 3 are accepted and the others are reassigned herein. Early Carboniferous crinoids are recognised from 5 formations in Qld and 4 or 5 formations in NSW. Late Tournaisian faunas from the Namoi Formation, Goonoo Gonoo Mudstone and Dangarfield Formation of NSW and the Malchi Formation of Old are considered coeval and all contain | or 2 species common to 2 of the form- ations. No species occurs in all 4 formations. Camerate crinoids are the most diverse forms in each of the faunas and the only crinoids known from the Dangarfield Formation. The Malchi һә „ө oo Formation has the most diverse fauna and includes the only Tournaisian flexible crinoids recognised in Australia. Visean faunas are from 2 or 3 formations in Qld and | in New South Wales. Except for Aaco- crinus in the Tournaisian or Visean Tellebang Limestone and the Caswell Creek Group, these faunas contain no genera in common. The only non-camerate taxon in these faunas is the disparid Litocrinus in the Baywulla Form- ation. Camerate calyces and tegmens have also been found in reef talus ofthe type section ofthe Early Carboniferous Lion Creek Limestone west of Rockhampton. However, they are weathered, or so fragmentary, that it has been impossible to identify them below family level. At least 5 genera are present, based on cup shapes and plate structures. In situ crinoid holdfasts are present in the reef core, from where the calyces are thought to have been derived. Campbell & Bein (1971) noted that Australian Early Carboniferous crinoids have more affinity with North American faunas than do the co- occurring brachiopods. However, they also noted that when the interior of many ofthe brachiopods described from North America become known this difference may not be so great. Recognition of a rhodocrinitid, Aacocrinus, Sampsonocrinus, Cribanocrinus, Dichocrinus, Dialutocrinus, Litocrinus, Prininocrinus and Holcocrinus in the Early Carboniferous and an acrocrinid and scytalocrinid in the Late Carboniferous of Aust- ralia strengthens the crinoid affinities with North America and Europe. Aacocrinus, Cribanocrinus and Prininocrinus were restricted to North America and Dialutocrinus to Europe (Lane & Sevastopulo, 1987, 1992), Ranges for Lower Carboniferous crinoid genera were given in Lane & Sevastopulo (1987, 1992), and the differences in the ranges between North America and Europe were noted along with first and last occurrences. They also pointed out that, although some differences in ranges and origins and extinctions were noted, most were relatively minor and perhaps the result of better definition of the North American genera and sampling artifacts. All ofthe Australian crinoid genera that are known from North America and Europe are of Tournaisian age, although some of the genera range into the Visean or younger in North America or Europe (Lane & Sevastopulo, 1987, 1990). These genera strongly support a late Tournasian age for the Namoi Formation, Goonoo MEMOIRS OF THE QUEENSLAND MUSEUM Gonoo Mudstone, Dangarfield Formation and the Malchi Formation as had been suggested by other invertebrate fossils (Campbell & Bein, 1971; among others). Camerate crinoids, that evolved rapidly and that are diverse and most abundant in North America during the middle Tournaisian through Visean, are the Rhodocrinitidae, Actino- crinitidae, Batocrinidae, Coelocrinidae, Dichocrinidae and Platycrinitidae (Lane & Sevas- topulo, 1987). The Batocrinidae and Coelo- crinidae are known only in North America, the others are well represented in equivalent strata in Europe and Russia. Except for the Batocrinidae and Coelocrinidae these families are represented in equivalent strata of eastern Australia, but presently are known from fewer genera than in Europe or North America. Disparid (Allagecrinidae, Synbathocrinidae) and poteriocrinid (Poteriocrinitidae, Scytalo- crinidae, Graphiocrinidae) crinoids are represented in the Early Carboniferous Austral- ian faunas by 1 or 2 genera each. These families are represented in North America and Europe by several genera and underwent rapid divers- ification during the Early Carboniferous (Lane & Sevastopulo, 1990). Flexible crinoids are represented by 2 poorly preserved specimens assigned to taxocrinid and sagenocrinid species. Both of these groups are common in the Early Carboniferous of Europe and North America, and they are known in Russian and Chinese faunas (Lane & Sevastopulo, 1990). Late Carboniferous crinoids in a Westphalian horizon in the Neerkol Formation the Acro- crinidae, Euspirocrinidae and Scytalocrinidae. The Acrocrinidae range from Tournaisian into the Stephanian. Although known from the Early Carboniferous of North America, Europe and Russia, in the Late Carboniferous they are known only from North America where they underwent arapid diversification (Moore & Strimple, 1969). Euspirocrinids are most common in the Ordovician and Silurian, waning thereafter. They are represented in North America by Parisocrinus and Zygotocrinus in the Early Carboniferous. The discovery of euspirocrinids in Australia in the Late Carboniferous and Permian (Webster & Jell, this volume) extends their geographic and strati- graphic range. Scytalocrinids are common in Early and Late Carboniferous deposits world- wide. Ifall ofthe reported Early Carboniferous marine fossil occurrences are plotted on biogeographic NEW CARBONIFEROUS CRINOIDS reconstructions, such as Bambach (1990), they lie between 45" N and S latitudes, mostly within 30° of the palaeoequator (Campbell & McKellar, 1969). The faunas are equatorial belt organisms and not truly cosmopolitan (Bambach, 1990). On recent plate reconstructions of Early Carbonif- erous biogeographic regions (Bambach, 1990) Australia is located on the E edge of what was becoming Pangea. Thus, the Australian crinoid faunas evolving in basins along the W border of Panthalassa in the Tournaisian and Visean were well away from the European and North American faunas on the N and W sides of the continental masses, although still in the equator- ial belt. A developing Tethys lay to the N and W ofthe Australian plate. Migration routes and sites of origin are uncertain for many genera at this time (Lane & Sevastopulo, 1987, 1990). By Westphalian time the Yarrol Shelf of east- central Queensland was between 55° and 60°S latitude. The crinoid fauna of the Neerkol Form- ation on the Yarrol shelf is the only higher latitude, cooler water, non-equatorial belt fauna known from the Late Carboniferous. The 2 species of acrocrinids in this fauna show greatest affinity with correlative taxa in the midcontinent of North America described by Moore & Strimple (1969), The euspirocrinids (Kopriacrinus gen. nov., Neerkolocrinus gen. nov.) are an extension ofthe family into the Late Carboniferous, and the scytalocrinid (Prininocrinus) is an extension ofa genus previously known from the Early Carbonif- erous of NW Canada. SYSTEMATIC PALAEONTOLOGY Crinoid teminology follows Moore & Teichert (1978), with columnal patterns after Webster (1974). Measurements are given as: length, parallel to the central axis; width, transverse to, but never cutting or joining the central axis; and depth or thickness, normal to and may join the central axis, Curvature of the cup walls, plate circlets within the cup and fixed brachials are referred to as: incurved if distally bending toward, vertical if parallel to, weakly to strongly flaring if bending away from and horizontal if perpendicular to the central axis. Material collected by us came from localities entered in the Queenland Museum Locality Register (QML) and is curated in the Queensland Museum Palaeontological Collection (QMF). Other palaeontological collections referred to are indicated by the following prefixes: Geological Survey of Queensland, Brisbane (GSQ); Geol- ogical Survey of New South Wales, Lidcombe 239 (MM); Australian National University (ANU) and Australian Museum, Sydney (AMS). Local- ities are in Queensland unless otherwise noted. Subclass CAMERATA Wachsmuth & Springer, 1885 Order DIPLOBATHRIDA Moore & Laudon, 1943 Superfamily RHODOCRINITOIDEA Roemer, 1855 Family RHODOCRINITIDAE Roemer, 1855 Rhodocrinitid gen. nov. (Fig. 1F) MATERIAL. QMF38955, QMF38956, locality and horizon unknown, probably Tournaisian Namoi Formation, NSW. Collected by GM. Philip. DESCRIPTION. Crown small, 14.3mm long, 5.7mm wide, arms gently splayed. Cup bowl shaped, 5.1mm long, 5.0mm wide, moderately coarse stellate ray ornament, plates moderately inflated. Infrabasals 37, small, confined to shallow basal cavity. Basals 5?, hexagonal, 1.9mm long, 1.7mm wide, proximally forming base of cup, distally forming base of cup wall, strongly convex transversely and Jongitudinally. Radials 5?, heptagonal, 1.5mm long, 1.7mm wide, strongly convex transversely and longi- tudinally. Primibrach | hexagonal, 1.1mm long, 1.2mm wide. Primibrach 2 axillary, heptagonal, 1.1mm Jong, 1.2mm wide. Secundibrach | at- tached to calyx; secundibrach 2 free. All free brachials uniserial, very thick, proximally recti- linear, distally cuneate, narrow, strongly convex transversely, straight longitudinally, with slender pinnule on long side. Secundibrach 4 axillary, no further branching. Arms branching isotomously, 4 in exposed ray, 20 total if all rays branch uniformly. One pinnule per brachial. Pinnulars very slender, elongate, with longitudinal angular ridge, longer than brachials. Interradial series 1-2-2-2-tegmen plates. Tegmen and anals unknown. Stem circular in section, homeomorphic, prox- imal columnal 0.8mm in diameter. Lumen small, circular; crenularium narrow; latus gently convex. REMARKS. The crown (Fig. 1F) is flattened with one ray central and part ofa second ray along the right side of the specimen. The interray is well-developed, narrowing at the distal end but leaving an obvious gap between the rays at the summit of the fixed arms. The brachials resemble those of a 4-armed dichocrinid or a primitive poteriocrinitid such as Liparocrinus. Most rhodocrinitids have biserial 240 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 1. A-E, Actinocrinites sp. 1. A, A ray view of partial calyx QMF38927 x1.6. B, lateral view of distorted partial calyx QMF38932 x2.5. C, B ray view of partial calyx QMI'38930 x 1.6. D, exterior view of abraded calyx plates QMF38933 x2.2. E, lateral view of partial theca and tegmen with long anal tube QMF38928 x2.5. F, Rhodocrinitid gen. nov., lateral view of crown QMF38955 x4.8. arms or the uniserial brachials, a primitive condit- ion, are very wide. The stellate ornament, although known in rhodocrinitids, such as Diamenocrinus and Rhodocrinites, is not common, The arm branching pattern is slightly advanced as most rhodocrinitids branch on the 2nd secundibrach and then again on the 6th tertibrach or higher and commonly branch one or more times at a higher level. The unfigured specimen (QMF38956) is NEW CARBONIFEROUS CRINOIDS crushed, the orientation is uncertain, plate relationships are masked and the ornamentation is only partially preserved. The specimens may represent a new genus, judged to belong to the rhodocrinitids, but without exposure of the anals, neither specimen is adequate to serve as a holotype. They are associated on a small slab with a scytalocrinid? indeterminate and Dichocrinus cf. D. laudoni. Order MONOBATHRIDA Moore & Laudon, 1943 Superfamily PERIECHOCRINOIDEA Bronn, 1849 Family ACTINOCRINITIDAE Austin & Austin, 1842 REMARKS. Actinocrinitids are among the most common elements in most major Tournaisian or Visean faunas although Visean actocrinitids are relatively rare in Europe. Moore & Laudon (1943) recognised that the actinocrinitids were derived from the periechocrinids and Brower (1967) subdivided them into 4 sections, which were recognised as subfamilies by Ubaghs (in Moore & Teichert, 1978). Brower (1967) also consider- ed the possibility that Actinocrinites might be polyphyletic and Campbell & Bein (1971), noting the differences in the arm structure, considered the Eumorphocrininae to be polyphyletic. Webster & Lane (1987) expanded Ubaghs's (in Moore & Teichert, 1978) key to identification of the genera of the Actinocrinitidae to include taxa introduced between 1978 and 1987. We recognise that the present classification needs revision and does not reflect the phylogeny of a family that is considered polyphyletic. A systematic revision ofthe Actinocrinitidae would require inclusion of the Periechocrinidae and is beyond the scope of this study. We continue to use the present classification for convenience, noting the general morphologic relationships of the taxa described for future reference. Identification of genera ofthe Actinocrinitidae is difficult based on fragmentary thecae or calyces in which the tegmen or distal ends of the protruded arm lobes are absent. Fragmentary material commonly does not provide information about the shape ofthe calyx which is important in the present classification. The presence or absence of an anal tube is recognised as the major difference between Aacocrinus and Diatorocrinus. Without the distal ends of the protruded arms, it may be impossible to tell the number of arms per ray, which is the major 24] difference between Actinocrinites and Aacocrinus. In some instances the number and type of free arms may be needed, such as Cytidocrinus and Manillacrinus. Subfamily ACTINOCRINITINAE Austin & Austin, 1842 Actinocrinites Miller, 1821 TYPE SPECIES. Actinocrinites triacontadactylus Miller, 1821 from the Tournaisian Mountain Limestone, England; by subsequent designation of Wachsmuth & Springer, 1881. REMARKS. Actinocrinites is thought to have evolved from a periechocrinid in the Late Devonian or Early Carboniferous (Moore & Laudon, 1943) and was exceedingly abundant in the Tournaisian of North America and Europe declining in the Visean. Actinocrinites has also been reported in the Early Carboniferous of Japan (Minato, 1951; Minato etal., 1979), eastern Aust- ralia (de Koninck, 1878, 1898; Etheridge, 1892; Pickett, 1960) and northern Africa (Termier & Termier, 1950). Webster & Lane (1987) consider- ed Silurian, Devonian and Permian species (Bassler & Moodey, 1943) to be incorrectly assigned to the genus. They also believed many of the Carboniferous species to be synonyms; there are 61 Carboniferous species currently assigned to the genus (Webster, unpublished data). Ausich & Kammer (1991) placed 4 species in synonomy, while establishing one new species, in a morphometric and qualitative analysis of the late Osagean and Meramecian Actinocrinites of the Mississippian stratotype region. Additional studies of this type should help resolve the current taxonomic morass of the genus. We consider the Australian species of Actino- crinites described herein to be new species. They are left in open nomenclature, because some specimens are poorly preserved, unsuitable to serve as holotypes and the taxonomy of Act- inocrinites must be resolved to allow proper comparisons. Actinocrinites polydactylus Miller, 1821 Actinocrimus polydactylus de Koninck, 1878: 160, pl. 6, fig. 3. 1898; 122, pl. 6, fig. 3. REMARKS. De Koninck (1877, 1898) described an internal mould of a compressed calyx from Glen William as Actinocrinus polydactylus (sic). The basal view of the specimen has a pentagonal axillary second primibrach; it lacks the tegmen and the ornamentation is unknown. The generic assignment is questionable because the arms are not grouped and protruded as 1n Actinocrinites. Because the ornament is not preserved, the specific assignment is doubtful. It is possible that the specimen is immature and belongs with Actinocrinitid indet. described below. Actinocrinites sp. | (Figs 1A-E, 2; Table 1) MATERIAL. QMF38927-38935 from QMLS0S8, late ‘Tournaisian, Malchi Formation. All specimens crushed during burial and plates leached by weathering; description based on latex casts. QMF 38927, calyx oriented with basal circlet centred; Е and A rays show secundibrachs, B ray shows proximal tertibrachs. QMF38928, distal part of theca, tegmen and anal tube. QMF38929-38933, calyces with proximal tertibrachs. QMF38934, partial calyx, base up. QMF38935, partial calyx, on side. GSQF10866 and 13489 from GSQL K-21, Visean?, Caswell Creek Group. DESCRIPTION. Calyx medium sized; arms grouped: tegmen highly arched with long slender anal tube; all plates below tegmen inflated, with prominent hexagonal stellate ridge ornament. Basal circlet large, tripartite, horizontal prox- imally. widely flaring distally: base with large circular stem facet with narrow crenularium on outer margin: base of stellate ridges at horizontal to widely flaring flexure. Radials 5, hexagonal. large. subequal. moderately flared. Primanal large, in radial circlet, distal anals unknown. Primi- brachs 2: first primibrach hexagonal. adjoined on each side by 2 interprimibrachs; second primi- brachaxillary, heptagonal. adjoined laterally by 2 interprimibrachs on each side. Secundibrachs 2-4. normally 2. Tertibrachs biserial and free above 2nd tertibrach. Interbrachial series 1-2-3 or 4 -?. Minimum 6 arms per ray, where free. Intersecundi- brachs not common, rarely 1 orseries 1-1. Tegmen moderately high, many small to intermediate inflated ambulacral and interambulacral plates: ambulacral plates slightly larger and elevated above interambulacral plates, with rounded to irregular central nodes. Anal tube projecting above tegmen, with distal anal opening, slender, of alternating rectangular and larger hexagonal plates in tiers, with hexagonal plates of one tier interlock above and below with rectangular plates of adjacent tiers; considerable variability in plate size, extra plates inserted to compensate. Stellate ridges 4 (2 on either side of sutures to adjacent basal, inner 2 merging at centre of radial) or 5 (from centre of each basal plate) extend from basal circlet to radials and anal, 3 or 4 ridges continue onto primibrachs with ray ridge largest: 3 or 4 subhorizontal ridges from radials to adjacent radials or anal, central ridge largest: 3 subhorizontal cross ridges from Ist primibrach to MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 1. Aetinocrinites sp. | measurements (mm). * = incomplete or crushed. | 38927 | 38931 | 38928 Calyx diameter* | 34.5 Йй "Thecal length (estimated) 25.0 380 | Basal circlet diameter 99 11.1% Basal circlet length 3.6 40 | Radial length | Е 63 10.8 Radial width 8.6 11.7 | First primibrach length 5.9 8.5 First primibrach width + 66 | 84% Second primibrach length 33 6.0 Second primibrach width | 55 77 First secundibrach length me | First secundibrach width 1 6.6 Stem facet diameter | 37 | 33 Tegmen length* 15.5 Anal tube length* | Е 15.8 | Anal tube diameter | 63 Ist interprimibrach or 2nd anal. central ridge largest; 3 diagonal ridges from radials to Ist interprimibrachs, central ridge largest: subhoriz- ontal and diagonal ridges continue onto 2nd tier of interprimibrachs; lateral ridges merge across plates to form small triangles at apices of plate junctions; triple ridges continue on interprimi- brachs to tegmen; single ridge continues on secundibrachs, tertibrachs, and intersecundibrachs: distally ridges grading into discontinuousaligned nodes or decrease in number in some interrays. Stem circular transversely. heteromorphic; nodi- taxis N323 132313231323. Columnals much shorter than wide: latus convex: lumen large, circular, articulum with narrow crenularium, slightly narrower areola and narrower spatium; nodals may bear short spines between cirral facets. REMARKS. Actinocrinites sp. 1 differs from 4? sp. 3 by having more complex stellate ornament- ation. The ornament of 4.? sp. 3 consists of inflated ossicles with apical pits and single stel- late ridges that are confined to the impressed sutures, not expressed as ridges across the inflated surface. Actinacrinites sp. 2 lacks thecal plate ornament except for impressed sutures and apical pits. The 3 species suggest a progression from unridged to complex stellate ornament, This is not an lineage as А. sp. 1 and 4.? sp. 3 occur together in the Rockhampton Group, whereas 4. sp. 2 occurs inthe younger Caswell Creek Group. Both single and complex stellate ridge ornaments are developed in Actinocrinites and Wachsmuth & Springer (1897) included specimens of both FIG 2. Actinocrinites sp. 1. A, lateral view, ray uncertain. partial calyx QMF38935 “3.5. B, basal view, rays uncertain, of partial calyx QMF38934 x2, C, lateral view, ray uncertain, partial calyx GSQF13489 x2. D, lateral view of partial calyx QMF38929 x2. E, lateral view of partial calyx QMEF38931 «1.7. F, lateral view, ray uncertain, partial theca GSQF 10866 x2. NEW CARBONIFEROUS CRINOIDS 24: types in species such as 4. multiradiatus (Shumard, 1858) and A. verrucosus (Hall. 1858). Single and multiple stellate ridge ornaments are also present in other actinocrinitids and other camerate families. The widespread occurrence and possible repeated development of these features suggest that it is a functional ornament. Brower (1967) suggested that actinocrinitids may be polyphyletically derived from the periechocrinids, many of which have the single or multiple stellate ridge ornament. The dimero- crinitids are periechocrinoids with infrabasals and should also be considered as possibly ancestral to the actinocrinitids. However. origin of the actino- crinitids is beyond the scope of this study. A pluricolumnal and disarticulated columnal at the base of the calyx on the slab with QMF38929 are included in А. sp.1, as they have the identical articular facet as that of the calyx. A 12.3mm diameter encrusting holdfast with a round articular facet (3.7mm diameter) bearing a central depression and large circular lumen impressionis on this same slab and may belong to an immature Actinocrinites sp. 1. This description is based on QMF38927, 38928 and 38931. with variations noted from other specimens. Actinocrinites sp. 2 (Figs 3, 4D) MATERIAL. GSQF 13490-13494 from GSQL3006, early or middle Tournaisian, Neil’s Creek Clastics. DESCRIPTION. Calyx bowl shaped. 20mm long (estimated). 335mm wide (estimated), sutures impressed: calyx plates inflated, without ribbing: arms grouped, flaring with lst primibrach: tegmen probably moderately inflated with anal tube. Basals 3, equal, horizontal proximally. flared distally, visible in side view of сир; basal circlet diameter 8.4mm. Radials 5, heptagonal, large, 6.9mm long. 7.6mm wide, moderately flaring. Primanal hexagonal, in radial circlet: anal series: P-2-3-3 minimum, continuing onto tegmen. First primibrach hexagonal, 5.4mm long, 5.8mm wide, incurved slightly from radials. Axillary 2nd primibrach heptagonal. adjoined laterally by 2 interray plates, outflaring distally. First secundibrach hexagonal, wider than long, strongly outflaring. Axillary 2nd secundibrach pentagonal. wider than long, gently upflaring. Arms free with first or 2nd tertibrach. Four arms FIG 3 MEMOIRS OF THE QUEENSLAND MUSEUM per ray. Interray series: 1-2-3-2-?, extending onto tegmen. Tegmen of many interambulacral plates. 1.7 long. 1.1 wide to 2 long. 2 wide. Numerous ambulacral plates increase in size towards anal tube. 0.6 long, 0.6 wide to 1.8 long. 1.7 wide. Anal tube subcentral. Stem impression circular. 4mm diameter. REMARKS. GSQF13490 consists of the internal and partial external moulds of a theca. The internal mould of the calyx is distorted slightly along the D-AB axis, with the tegmen crushed down into the visceral cavity. Ambulacral track- ways are elevated above the interambulacral areas. and the proximal part of the slightly eccentric anal tube projects above the ambulacral areas. The tegmen would have been moderately inflated. The external mould preserves the non- stellate character of the inflated plates of the basal circlet. the B ray and adjacent parts of the interrays on either side. GSQF13493 is the external mould of a calyx crushed along the A-CD axis. retaining the basal circlet, radials, primibrachs and parts of the interrays; the anal series is lost. GSQF 13492 is an internal mould of a part of the tegmen, showing growth lines on some ossicles. The unnumbered specimen is an external impression of a fragment of 2 rays and the interray from the distal ends of the radials through the 151 secundibrachs. It would have been the largest individual of the 5 specimens as a primibrach is 10.3mm long and 10.9mm wide. nearly twice the dimensions of the primibrach of СОЕ 13490, the most complete specimen. GSQF 13491 is a crushed partial theca and GSQF- 13494 the external mould of a partial theca. Actinocrinites? sp. 3 (Fig. 4E.F) MATERIAL. QMF38936 and 38937, moulds of crushed partial thecae from QML508, late Tournaisian, Malchi Formation. DESCRIPTION. Calyx high, truncated, conical. arms grouped, plate structure like Actinocrinites sp. 1. lacking intersecundibrach plates: interprimi- brach series 1-2-3-?. Anal series and tegmen unknown. Minimum 4 arms per ray, inner facet on axillary secundibrach wider than outer. Stem facet circular; lumen circular; articulum narrow. areola more than twice width of articulum, spatium intermediate width. Actinocrinites sp. 2. A.B, B ray and D-E interray views of partial calyx GSQF13493. C-E., B ray, internal A- B interray and oblique internal tegmen views of GSQF 13490. F, lateral view of partial tegmen GSQF 13492. G, interray view of partial calyx GSQF 13494. All x2. [a] NEW CARBONIFEROUS CRINOIDS QMF38936: Calyx 25.5mm long (estimated); basals 5.2mm long (estimated); radials 6.8mm long, 7.4mm wide; 151 primibrach 5.1mm long, 6.0mm wide; 2nd primibrach 4.6mm long, 5.5mm wide. QMF38937: Basal circlet diameter 11.8mm; basals 4.4mm long; radials 8.0mm long, 10.0mm wide; stem facet 4.5mm in diameter. REMARKS. The lack of stellate ridge ornament across the inflated calyx plates of Actinocrinites? sp. 3 is not an artifact of weathering or abrasion, because the stem facets of the basal circlet and broken proximal columnal of QMF38937 show sharp detail of the culmina and crenellae of the articulum and surfaces of the areola and spatium. The generic assignment is questioned because the tegmen is unknown. Aacocrinus Bowsher, 1955 TYPE SPECIES. Aacocrinus nododorsatus Bowsher, 1955 from the Kinderhookian (Tournaisian) Chouteau Limestone of Missouri; by original designation. Aacocrinus acylus sp. nov. (Fig. 4A-C) ETYMOLOGY, Latin, acylus, acorn of the holm-oak; refers to the acorn shape of the calyx. MATERIAL. HOLOTYPE: QMF38953. PARATYPE: QMF- 38954, external moulds of calyces from QML1248, Tournaisian or possibly Visean Tellebang Limestone. DIAGNOSIS. Calyx small, equidimensional; calyx bowl shaped, with single stellate ridge ornament; arms grouped, protruded, flare with axillary 2nd primibrach; tegmen strongly arched, as long as calyx; slightly eccentric anal tube; arms free with tertibrachs, 4 arms per ray; stem facet circular. DECRIPTION. Calyx small, equidimensional. Calyx bowl shaped; coarse, single-ridge stellate ornament. Basal circlet large, short, proximally horizontal; large impressed circular stem facet, distally upflared, forming base of walls, visible in lateral view. Radials moderately large, strongly convex longitudinally, moderately convex transversely, forming majority of cup wall. Primanal large, in radial circlet, series unknown; Ist primibrach and Ist interprimibrach subvertical, forming distal part of cup wall. Axillary 2nd primibrach widely flared, lacking stellate ridge ornament; 2nd secundibrach axillary; 4 arms per ray; arms free with MEMOIRS OF THE QUEENSLAND MUSEUM tertibrachs. Tegmen strongly arched, as long as calyx, formed of orals, three series of ambulacrals and interambulacrals; all plates nodose, commonly with large blunt nodes. Anal tube narrow, slightly eccentric, formed of medium- sized strongly nodose to blunt spined plates. Free arms and stem unknown. REMARKS. Silicification of the siltstone to fine grained sandstone external moulds of Aacocrinus acylus obliterated cup plate sutures except those of the basal circlet. Sutures of the brachials commencing with the 2nd primibrach and tegmen plates are well preserved. The stellate ridge ornament is rounded, vague, but obvious. QMF38953 is slightly crushed from compaction and oriented on its side, whereas QMF38954 does not appear to be crushed and is oriented obliquely on its side with nearly all ofthe tegmen covered. Neither specimen shows the anal series. The generic assignment is based on the plate arrangement and shape of the calyx. Aacocrinus acylus belongs to the 20-armed group of the genus and has a bowl-shaped calyx, whereas all other species of the genus have a more conical shape. The tegmen has third-order ambulacrals that Brower (1967) noted as the difference between 10- and 20-armed species. Both A. tetradactylus Brower, 1967 and A. chouteauensis (Miller, 1892), the other 20-armed species, have longer thecae, shorter tegmens and strongly eccentric anal openings. Aacocrinus acylus is the first report of the genus outside the United States. Brower (1967) noted that all identified species are from the Kinderhookian of the midcontinent of the United States, but he also recognised an unnamed Osagean form from the same area. Webster & Lane (1987) reported an early Osagean specimen from the Anchor Limestone of southern Nevada. Aacocrinus sp. | (Figs 5A-G, 6B-H) MATERIAL. QMF38938-38944, latex moulds of crushed partial thecae from QMLSOS, late Tournaisian, Malchi Formation. GSQF10865, 10867, 10868 and 10871a and b, from GSQL K-21, Visean? Caswell Creek Group. DESCRIPTION. Calyx medium sized; arms grouped; tegmen highly arched with one row of plates forming anal tube; all plates below tegmen inflated with prominent hexagonal single ridge FIG. 4. A-C, Aacocrinus acylus sp. nov. A,B, basal and lateral views of paratype, QMF3 8954 x2.6. C, lateral view of holotype theca, QMF38953 x2.6. D, Actinocrinites sp. 2, posterior interray view of GSQF13491 x1.6. E,F, Actinocrinites? sp. 3. E, basal view of partial calyx QMF38936 x2. F, lateral view of partial theca QMF38937 x2. NEW CARBONIFEROUS CRINOIDS 247 248 stellate ornament. Basal circlet large, tripartite, horizontal proximally, upflared distally; base with large circular stem facet with narrow crenularium on outer margin; fluting at base stellate ridges at horizontal to upflared flexure. Radials 5, hexagonal, large, subequal dimensions variable, upflared. Primanal large, in radial circlet; distal anals unknown. Primibrachs 2; Ist primibrach hexagonal, adjoined on each side by 2 interprimibrachs; 2nd primibrach axillary, heptagonal, adjoined laterally by 2 interprimi- brachs on each side. Secundibrachs 2. Tertibrachs becoming biserial and free after 2nd tertibrach. Interbrachial series 1-2-3-4-tegmen. Tegmen high, formed of many small to intermediate sized inflated ambulacral and interambulacral plates; ambulacral plates slightly larger and with circular to irregular central nodes, elevated above interambulacral plates. Anal opening projecting above tegmen with single row of plates. Stellate ornamentation of elevated sharp ridges; double and single ridge from basals onto radials and primanal, single ridges on all calyx plates thereafter; double ridge along basal sutures merging at centre of radials to continue as ray ridge; ray ridges largest, subhorizontal and diagonal ridges smaller. Occasional additional accessory node or very short ridge on radials or Ist interprimibrach. Minimum 4 arms per ray, inner facet on axillary secundibrach wider than outer. Stem facet large, circular; lumen large, circular to subpentagonal; articulum narrow, areola more than twice width of articulum, spatium intermediate width. QMF38938: Calyx 26mm long (est.); basal circlet diameter 10mm, 5.3mm long; radials 7.6mm long, 8.3mm wide; Ist primibrach 6.4mm long, 6.4mm wide; 2nd primibrach 5.7mm long, 6.7mm wide; proximal columnal 5.2mm in diameter. REMARKS. This description is based on QMF- 38938, 38941 and 38942 and GSQF10867. Other specimens are included in this taxon because they have the same type of plate ornament. The single stellate ridge ornament results in a less intricate ridge pattern on the calyx. The wider facets on the inner axillary secundibrachs suggest additional branching of the arms, which would result in 6 MEMOIRS OF THE QUEENSLAND MUSEUM arms per ray. Known species of Aacocrinus have 2 - 4 arms per ray, whereas most species of Actinocrinites have 6 arms per ray and an anal tube of variable length. Aacocrinus sp. 1 differs from A. acylus by having a longer, more conical cup and calyx, and a longer tegmen. The tegmen is formed of more numerous plates than any of the American species ofthe genus and is more similar to that of Actinocrinites. Aacocrinus sp. 1 is considered a new species but all specimens are incomplete and unsuitable to serve as a holotype. Manillacrinus Campbell & Bein, 1971 TYPE SPECIES. Cactocrinus? brownei Dun & Benson, 1920 from the Namoi Formation, NSW; by original designation. REMARKS. Manillacrinus was incorrectly included in the Eumorphocrininae by Ubaghs (in Moore & Teichert, 1978) and Webster & Lane (1987) as they did not recognise the arm grouping of M. brownei (Campbell & Bein, 1971, pl. 50, figs 1, 2, 6, 7) and no mention was made in the original descriptions about the arms grouped and protruded, a character of the Actinocrinitinae. Manillacrinus is distinguished from other genera of the Actinocrinitinae by the biserial ramules given off the outer sides of the 2 uniserial arm trunks of each ray. Manillacrinus brownei (Dun & Benson, 1920) (Figs 5H, 6A) Cactocrinus? brownei Dun & Benson. 1920: 342. pl. 19. fig. 1. Manillacrinus brownei (Dun & Benson); Campbell & Bein, 1971: 427, pl. 50, figs 1-7; text-fig. 7. MATERIAL. QMF38945, an external mould of a partial theca, from QML508, late Tournaisian, Malchi Formation. GSQF13495 from GSQL3012, late Tournaisian Malchi Formation, external mould of partial theca. REMARKS. Manillacrinus brownei has pro- truded grouped arms; sharply elevated, centrally inflated, single ridge, stellate ornament; and 2 ramule-bearing arms per ray. The 2 ramule- bearing arms, combined with the protruded grouped arms, are the most diagnostic characters of the genus. The single ridge stellate ornament of M. brownei is not unique to the genus, but the stellate intersecundibrach plate is not common in actinocrinitids. Also, the stellate ridges continue FIG. 5. A-G, Aacocrinus sp. 1. A, lateral view of partial calyx QMF38939 x1.7. B, lateral view of partial calyx QMF38938 x1.6. C, lateral view of partial calyx QMF38943 x4. D, interray view of partly disarticulated calyx QMF38942 x2.6. E, view of crushed theca QMF38944 x2.6. F, view of basal circlet and stem facet QMF38941 x4.2. G, basal view of partial calyx QMF38940 x3.6. H, Manillacrinus brownei (Dun & Benson, 1920), basal view of partial calyx GSQF13495 х2. 249 NEW CARBONIFEROUS CRINOIDS tw Un © without significant decrease in development to the base of the tegmen. The partial calyx GSQF13495 is a basal circlet with one ray to the secundibrachs and proximal parts of 2 other rays. It is assigned to M. brownei because the stellate ornament is so sharp and the ridges are centrally inflated and highly elevated. It also has the double ridges from the basal circlet to 2 of the radials. The 2nd specimen QMF38945 is the distal.part of a calyx with the stellate ridged intersecundibrach in both rays. Manillacrinus acanthus sp. nov. (Figs 7, 8) ETYMOLOGY. Greek acanthos, spine; referring to the short thoms on some of the ambulacral plates of the tegmen. MATERIAL. HOLOTYPE: MMF33605. PARATYPES: MMF33431 and 33606; other specimens MMF33435 and AMSF65556, all from the late Tournaisian Namoi Form- ation, near Barraba, NSW. DIAGNOSIS. Ambulacral plates with 1 or 2 short blunt spines midway between free arms and anal tube; calyx high conical. DESCRIPTION. Calyx 62.3mm long; 30.2- 50.6mm wide, (40.4mm average), widest at base; tegmen high, conical; plates tumid; apical pits on calyx; multiple stellate ridge ornament to base tegmen; arms grouped, distinctively protruded; tegmen high arched; short central anal tube. Basal circlet tripartite, 9mm long, 12.7mm wide (average), proximally concave, with circular stem facet, distally strongly upflared. Radials 5, septagonal, 12.5mm long, 11.3mm wide, concave to convex longitudinally, moderately convex transversely. Primanal 10.1mm long, 8.6mm wide, gently convex longitudinally, moderately convex transversely, in radial circlet; anal series P-2-3-4-4-tegmen. First primibrach hexagonal, 7.9mm long, 9.0mm wide, moderately convex longitudinally and transversely, adjoined laterally by 2 ilBrr on each side. Axillary 2nd primibrach septagonal, 5.1mm long, 8.2mm wide, straight to slightly concave longitudinally, moderately convex transversely, adjoined on outer side by 2 interprimibrachs. First sec- undibrach pentagonal, 3.4mm long, 4.8mm wide, longitudinally concave, moderately flaring outward, convex transversely. Axillary 2nd MEMOIRS OF THE QUEENSLAND MUSEUM secundibrach pentagonal, 2.5mm long, 5.3mm wide, straight to gently convex longitudinally, convex transversely, widely flaring outward. First and 2nd tertibrachs fixed in calyx, arms free thereafter. Interprimibrach series 1-2-3-2-tegmen; plates convex longitudinally and transversely, decreasing in size distally, last row incurved. Single intersecundibrach octagonal, elongate, with short proximal and distal facets. Multiple stellate ridge ornament of 4-5 ridges from basals to radials, decreasing to single ridge thereafter, continuing to secundibrachs in rays and to tegmen on interbrachs. Tegmen plates large, tamid. Ambu- lacral plates elevated above interambulacrals, 1 or 2 midway between free arms and anal tube bearing short blunt spines. Anal tube subcentral, formed of 2 rows of polygonal plates. Stem circular; proximal facet 9.5mm diameter (average), with narrow crenularium. Free arms unknown. REMARKS. Manillacrinus acanthus is distin- guished from M. brownei by the spines on the tegmen and the much more conical theca. The adult specimen is much larger than adult M. brownei. Measurements were taken on the holotype, which is crushed normal to the BC-D plane. Paratype MMF33431 lacks the tegmen. Paratype MMF33606 lacks the basal circlet and the teg- men is crushed into the visceral cavity, but the spines on the ambulacral plates are well-preserved. The stellate ridge ornament is poorly preserved on the types but is recognised best on paratype MMF33431. MMF33435 is abnormal with both the A and B radials followed by 4 primibrachs. The distal parts of the calyx are missing and the distal relationship of the ray and interray plates is unknown. AMSF65556 lacks the basal circlet, but the two radials and next distal plates have well-preserved stellate ridge ornament. Sampsonocrinus Miller & Gurley, 1895 TYPE SPECIES. Sampsonocrinus hemisphericus Miller & Gurley, 1895 from the Chouteau Limestone, Missouri; by original designation. REMARKS. Sampsonocrinus is distinguished from Actinocrinites by having a lower, more bowl- shaped cup. relatively larger basals and radials and only 4 arms per ray. Sampsonocrinus is à FIG. 6. A, Manillacrinus brownei (Dun & Benson, 1920), lateral view of partial calyx QMF38945 x1.8. B-H, Aacocrinus sp. 1. B, lateral view of base of partial calyx GSQF1087 1b x2. C-G, B ray, oral, posterior views of tegmen, interior B ray and summit oblique of interior of tegmen of GSQF10867 x2. H, lateral view of partial theca GSQF 10868 x2, NEW CARBONIFEROUS CRINOIDS 2 un me" [e] [97 кә MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 7. Manillacrinus acanthus sp. nov. A,B, A-E interray and posterior views of paratype MMF33431.lacking tegmen x1.7. C,D, C ray and A ray views of holotype MMF33605 x12. Tournaisian genus, previously reported from the United States. United Kingdom records are now excluded from the genus (George Sevastopulo pers. comm.) This is the first report of the genus in Australia. Sampsonocrinus cannindahensis sp. nov. (Fig. 9) ETYMOLOGY. From Old Cannindah. MATERIAL. HOLOTYPE: GSQF10864, external and internal mould of calyx from GSQL K-21, Visean? Caswell Creek Group. PARATYPE: QMF 17784, internal mould of cup. Collected by C. W. De Vis (no. 649). DIAGNOSIS. Cup low, broad, with blunt spines or coarse nodes on each of the tegmen plates. DESCRIPTION. Calyx low bowl-shaped, 19.6mm long (incomplete), 39.8mm wide; plates inflated, bulbous: sharp simple stellate ridge ornament continuous across sutures on all adjacent plates below tegmen; tegmen moderately inflated; arms grouped. projecting subhorizontally. Basal circlet 4.0mm long (estimated), 8.2mm diameter NEW CARBONIFEROUS CRINOIDS FIG. 8. Manillacrinus acanthus sp. nov., A-C, A гау x1.2), oblique oral x1.5) and posterior x 1.5) views of paratype MMF33606, lacking basal circlet and tegmen crushed into the visceral cavity. (internally). Radials 5, hexagonal, 4.8mm long (internal), 7.3mm wide (internal), gently convex longitudinally and transversely. widely out- flared. Primanal heptagonal, large, in line with radials; anal series P-2-3-3-2-tegmen, wide. First interradial hexagonal, 5.7mm wide; series 1-2-3-2-tegmen. First primibrach pentagonal. 4.1mm long, 5.4mm wide. C ray 2nd primibrach axillary, pentagonal, 3.0mm long, 3.6mm wide, with distal tip curving outward toward sub- horizontal. Single secundibrach axillary, slightly upflared. Arms free with 2nd tertibrach, 4 arms No Un чө per ray, 20 arms if branching uniform. Tegmen plates relatively small, strongly inflated into short blunt spines. Orals 5, off centre toward anal interray, larger than other tegmen plates, sur- rounding anal opening at tegmen apex. No anal tube. Ambulacral plates inflated more than inter- ambulacrals with short blunt spines. Stem and free arms unknown. REMARKS. The holotype internal mould is slightly distorted along the D-AB axis; weather- ing destroyed most of the external mould of the cup. Ambulacral grooves were elevated above the interambulacral areas, distally merging with them toward the centre of the tegmen, and the eccentric anal tube projected above the tegmen surface. The short calyx of the paratype is crushed inward concentrically around the basal circlet and the tegmen is not exposed. Dialutocrinus Wright. 1955 TYPE SPECIES. Dialutocrinus milleri Wright, 1955 from the Tournaisian of England; by original designation. Dialutocrinus? sp. (Fig. 10A-C) MATERIAL. QMF33864, а partial calyx, from the Visean? Caswell Creek Group, 7.1 km NE of Monto, on hill on side of road; 24°51°6"S, 151°91°54"E. Collected by Paul Tiemey. DESCRIPTION. Calyx 19.1mm long. 36.3mm wide, conical, with concave sides. with single ridge stellate ornament. Arms very weakly grouped. slightly portruded. Basal circlet 2.5mm long, 1l.1mm wide, flat proximally, widely flaring distally, circular stem impression in basal impression. Radials 5, 7.2mm long and wide, heptagonal, moderately bulbous. Primanal small- er than radials, 6.5mm long, 5.5mm wide, moderately bulbous. Anal series P:2:3:4:?, con- nected with tegmen. First primibrach hexagonal, 3.9mm long, 6.0mm wide, bulbous, slightly con- cave longitudinally. Axillary 2nd primibrach heptagonal, 3.8mm long, 6.2mm wide, slightly convex longitudinally and transversely. Inter- primibrach series 1:2:3:4:3:?, plates decrease in size distally, forming moderately wide gap bc- tween arms at rim of calyx. First secundibrach hexagonal, 3.1mm long, 4.7mm wide, weakly flaring. Axillary 2nd secundibrach pentagonal, 2.8mm long. 4.4mm wide, weakly flaring. Inter- secundibrach series 1:1. Minimum 2 tertibrachs. Arms not free until 3rd tertibrach or | st quartibrach. Probably 8 arms per ray. Stem circular transverse- ly. 4.3mm diameter proximally. t3 [7 гм MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 9. Sampsonocrinus cannindahensis sp. nov., A-D, external oral and E, ray viewsand oral and basal views of internal mould of holotype, GSQF 10864 x2. E, B-C interray view of internal mould of paratype, ОМЕ 1 7784 x1.6. REMARKS. The specimenlacksthetegmenand inthe DE interray and adjacent parts of the D and free arms, and has been slightly distorted by Е rays. The specimen is questionably assigned to compaction of the cup into the fixed brachials. — Dialutocrinusbecause it has 2 secundibrachs and Plates are recrystallised and the stellate ornament the free arms are unknown. The type species, D. is nearly lost except on the lower 2 rows of plates milleri Wright, 1955, has a single secundibrach NEW CARBONIFEROUS CRINGIDS 255 and the inner arm branches repeatedly above ihe axillary seeundibraeh; the puter arm remains unbranched, The wide interprimibrach series (1-2-3-4-3-7), single iniersecundibrach. calyx shape, cup plate structure; and ornamentation of I? sp agrees with D. milleri. The specimen тау represent a geographical variant. Actinocrinitid indet. (Fig. JOD,E) Кшт sp. and. Татре, 1892: 77, p. 20. figs b. 7 ‘Perwchacrimy зейін епок, 1892. 78, pl. 22 fin. 3 “аресте? sp, md, Бетине, 1822; 79, vewrslig. Т MATPRIAL. AMSE27094 and 28189 (pari and counter- part). locality uncerizun within the Clarencetowr area. UOQFT3204, from the Toumaisia Wootton Beds. 2,25 miles N of Clurencetowa, on Glen William Rd, NSW DESCRIPTION. Calyx 85.5mm long, 56min wide (incomplete. crushed). Basal eirclet. tripartite. 23 31mm diameter (minimal), widely flaring. Radials large. 21.3mm long. 21.0mm wide, hexagonal, gently convex longitudinally and transversely, upflared. First primibrach hexagonal, 168mm long, 174mm wide, gently convex longitud- inally and transversely, upflared. Axillary 2nd primibrach pentagonal. 12.7mm long, 15 3mm wide (estimated), concave longitudinally and trans- versely, Second secundibrach hexagonal, 5.6tmim long, 10. тит wide. Iiterprimibrach series 1-2-3-7. Anal series. tegmen and stem unknown, REMARKS. Both specimens are crushed thecae. Measurements were made оп AMSF28389. The plates show 9 major growth rings and are preserved as very thin interior rinds. Edges of the plates are pteserved as elevated walls marking the sutures. No omamentation is preserved as the exteriors Were lost by weathering, Plates of UQF 13204, are weathered and leached. with the remaining calcite very sofi and chalky, shawing 8 major growth rings. Provenance of the calyx is given as Clarencetown, NSW. There are numerous lenses of limestone within the Wootton Beds between Clarencetown and Dungog ( Lishmund etal., 1985). Both specimens probably derive from this arca, Actinocrinus sp. ind. of Etheridge (1892) 15 probably from the same locality as the described material, or the same horizon from a nearby locality: preservation is similar. Etheridge (1892) reported specimens from the Carboniferous Mirari Limestone, at Greenhills, Paterson to Dungog Road. Periechocrinus indicator, from Carbomferous deposits at Chalky Gully, and Periechoerinus? sp. ind., from the Carboniferous Mirari Limestone at Greenhills, described by Etheridge (1892. p. 69-7]. nl. 22, fig. 4: text-fig. T) may also he conspecifie with the material describe. Neither of the Periechocrinys specimens are quite as large, but they have similar plate structure. Also, the former is preserved in the same manner us AMSF28389_ If all of these specimens are conspecific, the specifie name indicator will have рпогцу if a genus is specified. Family DICHOCRINIDAE 5. ^. Miller, 1889 Subfamily DICHOCRININAE S.A, Miller, 1889 Dichocrinus Munster. 1839 TYPE SPECIES. Dichocrimis radiatus Münster, |839 from the Barly Carbonilerous. of Germany: by monotypry, Dichocrinus ci. D. Тамо Broadhead, 1981 (Fig. 11) MATERIAL, QMF38957. locality and horizon in NSW unknown. probably late Toumaisian Матор Formation Collected by Li M. Philip. DESCRIPTION, Crown small, 12.6mm long (incomplete), 6.0mm wide, expanding gently upward. Cup sleeply conical, 7.2mm long. 5.1mm wide, smooth, unornamented. Ваза! circlet bipartite, steeply upflared, visible in side view, 4.2mm long, 2.6mm wide, Radials 5, longer (4.2mm) than wide (2.6mm). expanded gently distally. Radial facet angustarv, projecting slightly above shoulders of radial. Anal not exposed. Brachials strongly convex transversely, straight longitudinally, rectilimear. Second primi- brach axillarv, isotomous branching, no distal branching on preserved secundibrachs; 10 uni- serial arms if branching same in all rays. Stem heteromorphic proximally (noditaxis pattern N1). homeomorphie distally. Columnals circular transversely, diameter 1.5mm proximally; latus gently to moderately convex. REMARKS. This small crown ts flattened in the anterior-posterior plane of symmetry. Bracluals at the base of the arms are slightly disarticulated from the radials. The conical cup with the basals making up half the cup wall, 10 arms and uniserial proximal brachials are all primitive features in Dichocrinus. The specimen is similar to D, ladon, but the stem does not taper away trom the cup as il does on the holotype of D. laudani (Broadhead, 1981, pl. 2. fig. 8). Taper of the stem may be a variable feature as the paratype (Broadhead, 1981, pl. 2, fig. 6) and other specimens (Webster, 1997, pl. 2. figs 5, 10) do по! show as strong a taper. Distally, the brachials of 256 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 10, A-C. Dialutocrinus? sp.. basal, B ray. and D-E interray views of clayx QMF33864 «1.8. D.E, Actinocrinitid indet. D, lateral view of weathered partial calyx AMSF'28389 х1. E, lateral view of exterior of AMSF27094 x 1.1 NEW CARBONIFEROUS CRINOIDS FIG. 11. Dichocrinus cf. D. laudoni Broadhead, 1981, lateral view of upright crown QMF38957, with Scytalocrinid? indet., lateral view of horizontal crown QMF38958 x3. D. laudoni become cuneate, whereas the distal brachials of D. cf. D. laudoni are unknown. They probably become cuneate as this is a feature common to all uniserial species of the genus. If new material is found to show that the brachials do not become cuneate distally, this Australian specimen could represent a new species. In Dichocrinus only D. cintus Miller & Gurley, 1890 and D. fusiformis Austin & Austin, 1844 have 10 arms, but both have surface ornament on the cup plates. All 3 of the 10-armed species of Dichocrinus are of middle or late Tournaisian аре, suggesting a similar age for D. cf. D. laudoni. The specimenis ona small slab associated with anindeterminate scytalocrinid? and a rhodocrinitid. 257 Family ACROCRINIDAE Wachsmuth & Springer. 1885 Subfamily GLOBACROCRININAE Moore & Strimple, 1969 Denarioacrocrinus gen. nov. YPE SPECIES. Denarioacrocrinus neerkolensis gen. et sp. nov. from the late Namurian or early Westphalian part of the Neerkol Formation, near Stanwell, Queensland. ETYMOLOGY. Latin denarius, containing 10; refers to the 10 distal intercalaries. DIAGNOSIS. Calyx vase-shaped; basal circlet unknown; strongly rounded. angustary radial facets, sloping downward outward: minimum 5 rows of intercalaries: 10 plates in distal row, 1 in line with single anal. 3 in line with A, D and E radials, all others interradial; 4 arms per ray. REMARKS. Moore & Strimple (1969) noted the explosive evolution in the acrocrinids in the Middle to Late Carboniferous of the midcontinent region of the United States. The Globacrocrin- inae retains the narrow radial facets and advanced genera have a flat basal circlet not visible in lateral view. Genera are defined mainly on the number of distal intercalaries. Denario- acrocrinus, with 10 distal intercalaries, is intermediate between the Morrowan to Des- moinesian G/obacrocrinus,. with 8, and the Missourian Cauacrocrinus, with 11 distal intercalaries. Denarioacrocrinus is the first Late Carboniferous globacrocrinid with part of the arms preserved. The 4 arms per ray are an advanced condition compared to the Early Carboniferous Protacrocrinus with 2 and Springeracrocrinus with 3. Denarioacrocrinus neerkolensis sp. nov. (Fig. 12A-C) ETYMOLOGY. From the Neerkol Formation. MATERIAL. HOLOTYPE: GSQF10875a and b, internal and external thecal moulds from GSQLK106, late Namurian or early Westphalian, Neerkol Formation. DIAGNOSIS. As for genus. DESCRIPTION. Calyx vase-shaped, 22.9mm long (incomplete), 16.5mm wide (slightly crushed), widest at midlength. suture flush, fine vermiform ornament. Basal circlet not preserved. Intercalar- ies mostly hexagonal, moderately large, minimum 5 rows. Distal row of intercalaries 10 plates, 3 in line with radials (A. D and E). 1 under anal, all others interradial. Distal intercalary next to anal series below C radial, 5.1 mm long, 5.1mm wide (estimated). gently convex, subvertical. A and D radials hexagonal, proximally adjoining 3 inter- calaries; C and E radials pentagonal. proximally adjoining 2 intercalaries; E radial pentagonal with proximal tip adjoining distal tip of in line intercalary; all radials gently convex transversely, straight to slightly convex longitudinally, sub- vertical. C radial 4.7mm long, 6.3mm wide. Radial facet angustary, sloping downward outward, strongly convex externally, below radial summit, with transverse ridge. Anal plate large, hex- agonal, 5.5mm long, 4.5mm wide, distal half in line with radials. Tegmen formed of small plates, projecting above radial summit. Arms slender, isotomously branching on 2nd primibrachs and 2nd secundibrachs, 4 per ray, 20 arms if all rays branch as C ray. Proximal brachials cuneate, convex longitudinally, strongly rounded transversely, horseshoe-shaped, with pinnule on long side. Distal brachials unknown, probably biserial. REMARKS. This calyx is slightly distorted from compaction and lacks the basal circlet. The vase-shape, multiple rows of intercalaries and angustary radial facets are primitive features of the Globacrocrininae, whereas 4 arms per ray is an advanced feature. This was the first crinoid reported (McKellar, 1966) from the Late Carbon- iferous of Australia. Denarioacrocrinus? ornatus sp. nov. (Fig. 12D-F; Table 2) ETY MOLOGY. Latin ornamentus, ornament; referring to the stellate and granulate ornamention. MATERIAL. HOLOTYPE: GSQF10877a and b from GSQL334. PARATYPE: GSQF10876 from GSQLK 106, late Namurian or early Westphalian Neerkol Form- ation. DIAGNOSIS. Alternating 6 rectangular and 6 hexagonal first row of intercalaries above the basals and stellate ornament on cup plates; sutures impressed. DESCRIPTION. Calyx vase-shaped, widest at midlength. Bipartite basals horizontal, in basal impression, followed by 7 rows of intercalaries increasing in size distally. Sutures impressed. Plates mostly hexagonal after first row of alternating 6 rectangular and 6 hexagonal plates. Rows staggered with plates interlocking distally and proximally. Occasional pentagonal or hepta- gonal plate where interlocking adjusted for smaller plate. Plates with stellate ridge ornament radiating from centre of plate to а! adjacent plates. Longi- tudinal ridge developed most strongly on basal MEMOIRS OF THE QUEENSLAND MUSEUM row of intercalaries. Fine nodose to vermiform ornament, nodes mav be aligned to stellate ridges or sides of plates. First row of intercalaries horizontal, rectangular ossicles strongly convex longitudinally. hexagonal ossicles forming base of calyx; hex- agonal plates distal tips visible in lateral view. All following intercalaries subvertical. Stem circular transversely; columnals with wide aureola, narrow crenularium, small ?pentagonal lumen. REMARKS. GSQF10877a and b are the internal and external moulds of the base of one partial calyx up to the 3rd row of intercalaries. GSQF- 10876 is a partial calyx lacking the radials and tegmen. It is crushed along the enterior-posterior plane of symmetry. D.? ornatus is distinguished from D. neerkolen- sis by the stellate ornament and plate arrangement of the 1st row of intercalaries. This is the Ist acrocrinid known with stellate ornamentation. Lacking the radials and complete row of distal intercalaries generic assignment must be quest- ioned. Suborder GLYPTOCRININA Moore, 1952 Superfamily PLATYCRINITOIDEA Austin & Austin, 1842 Family PLATYCRINITIDAE Austin & Austin, 1842 Platycrinites J.S. Miller, 1821 TYPE SPECIES. Platycrinites laevis Miller, 1821 from the Early Carboniferous of England; by subsequent designation of Meek & Worthen, 1865. Platycrinites nux? (Etheridge, 1892) nomen correctum (Fig. 141,J) MATERIAL. GSQF10870, an internal mould from GSQLK-21, Visean? Caswell Creek Group. REMARKS. An internal mould of a cup, des- ignated Platycrinus? nux by Etheridge (in Jack & Etheridge, 1892) and mentioned by Etheridge TABLE 2. Denarioacrocrinus? ornatus sp. nov. measurements (mm), T == | - |. Fl0877. F10876 — Calyx length чш ‘pal 2 242 | Calyx width 156 | Diameter basal circlet (maximum) 69 Diameter basal circlet (minimum; _ 59 Intercalary | rectangular, length JL Intercalary | rectangular, width | intercalary 1 hexagonal, length Jotercalary | hexagonal, width | Diameter stem impression 24 | 56 | NEW CARBONIFEROUS CRINOIDS 2 Un D FIG. 12. A-C, Denarioacrocrinus neerkolensis gen. et sp. nov., holotype GSQF10875a and b, x2.5. A, posterior view of calyx. B,C, A ray and posterior views of cop mould. D-F, sie eae hans y Ser rode sp. nov. D. B- C interray view of calyx, paratype GSQF 10876, 3.3. E-F, basal and interior view of holotype GSQF 10877, (1892) has a high bowl-shaped basal circlet and Carboniferous Caswell Creek Group. near Old vertical elongate radials. An internal mould Cannindah is nearly the same size. 4 tiic (GSQF10870) of a cup from the Early same shape and plate proportions, Cooneraene 260 of these specimens in size and proportions, as well as the tvpe of preservation, suggests that they are from the same stratigraphic unit. Etheridge (in Jack & Etheridge. 1892) reported the specimen from the Middle or Marine Series of the Bowen river Coal Field, 21720'S, 148730 E; this is inthe Lizzie Creek Volcanics. We wonder if. for unknown reasons, the locality information given by Etheridge (in Jack & Etheridge.1892) could be incorrect or if the specimen was reworked from Carboniferous deposits. Disar- ticulated unornamented plates described below as Platvcrinites sp. 4 have radial ratios similar to those of this specimen. Neither the new specimen described or that of Etheridge (in Jack & Etheridge, 1892) resolves the problem of the lack of ornamentation and other external features of the plates. Also. the tegmen and arms are unknown. It could be argued that P. nux is based on an internal mould and the surface ornament is unknown. However. future collections in these areas should provide additional specimens to support the acceptance of P. nux. Platycrinites testudo Campbell & Bein. 1971 (Fig. 13) Platycrinus sp. Etheridge, 1892: 131, pl. 20, fig 8. Platycrinites testudo Campbell & Bein, 1971: 430, pl. 51, figs 10-21. Platycrinites? crokeri Campbell & Bein. 1971: 433, pl. 49, figs 9-15; pl. 51. figs 3-4. Platycrinites? sp. 1 Campbell & Bein, 1971: 434, pl. 51. figs 1-2. MATERIAL. MME33433, 33437, 33438, 33441, 33445- 334552, 33607. AMS F59512, 59513, 59516, 65664, 104701, 104702 from the late Tournaisian Namoi Formation, near Barraba, NSW; collected by J. Irving. REMARKS. The nodose ornament of previously illustrated specimens indicates the variation in P. testudo. New specimens from the Namoi Formation support this interpretation and show additional variation in the size and alignment of the nodes and ornamentation on the radials as well as the size and arrangement of the ornament- ation on the tegmen plates. The incomplete specimen illustrated by Eth- eridge (1892: 20. fig. 8) was described as 4 plates of a basal circlet. but it is 2 radials with distal tegmen plate and a proximal part of the basal circlet. The radial facet is visible on the radial to the right centre and the inverted V of the nodose ornament points to the facet on the radial to the left of centre. MEMOIRS OF THE QUEENSLAND MUSEUM Platycrinites sp. 1 (Fig. 14F-H) MATERIAL. MMFE33440, 33442, 33443 and 33453 from the late Tournaisian Namoi Formation near Barraba, NSW. DESCRIPTION. Cup bowl-shaped, 25mm long (crushed). 12-25.2mm wide (18.6 av.), fine granular or vermiform ornament. Basal circlet low conical. widely flaring: plates fused. Radials 5. 16.1mm long, 13.1mm wide, weakly convex longitudinally, gently convex transversely, shoulders flat to laterally sloping. Radial facet angustary. 5.4mm wide, crescent-shaped. moderate- ly convex aborally. Primibrach axillary, concave longitudinally. moderately convex transversely. Stem facet elliptical, relatively small, 3.4 x 4.4mm. Proximal columnal with convex latus. Arms. tegmen and stem unknown. REMARKS. P/aytcrinites sp. 1 differs from P testudo by lacking the coarse nodose ornament- ation, having a more upflaring basal circlet and relatively smaller stem facet. It differs from P? sp. 2 of Campbell & Bein (1971) by being more equidimensional, lacking the pustulose ornament and having shallower radial facets. The specimens are all distorted and cracked from compaction. Platycrinites sp. 2 (Fig. 14A) MATERIAL. QMF38946 from QMLS30S, late Tournais- ian, Malchi Formation. DESCRIPTION. Radial large, 12.8mm long. 13.6mm wide. gently convex longitudinally and transversely, thin, with fine granular ornament; radial facet angustary, projecting outward from radial surface, deep, horseshoe-shaped, strongly convex outer edge, transverse ridge wide, gently rounded. ends elevated slightly. REMARKS. The fine granular ornament of Platycrintes sp. 2 readily distinguishes it from P. testudo. The projected horseshoe-shaped facet is much deeper and more projected than that of P sp. 1. Lacking the rest of the cup and tegmen it is left in open nomenclature. Platycrinites sp. 3 (Fig. I4D.E) MATERIAL. Basal circlet and partial radials QMF'38947 and partial radial plate, QMF38948, from QML508, late Tournaisian, Malchi Formation. DESCRIPTION. Basal circlet bowl-shaped, thin. unornamented: plates fused. Radial large. NEW CARBONIFEROUS CRINOIDS 261 VIG. 13. Platycrinites testudo Campbell & Bein. 1971. A, lateral view of calyx. orientation uncertain. MMF 33433 х1.6. B, oblique posterior view of theca AMF59513. MMT3343 x1.4. С.П, A ray and posterior views of theca MMF33607 «1.7. EI basal and oral views of theca posterior view of theca MMF33438 «1.6. 15.5mm long (incomplete), 15.5mm wide, thin, smooth; angustary radial facet strongly convex outwardly, slightly elevated above radial surface, small inwardly pointed V-shaped ridges at outer ends of transverse ridge. Stem facet small, 2.0 х 2.4mm, elliptical outline, with narrow crenularium. REMARKS. Platycrinites sp. 3 differs from P. testudo, P. sp. | and P. sp. 2 by lacking ornament. The specimens are fractured and incomplete. They are illustrated to show the variation in the platycrinitids of eastern Australia. Platycrinites sp. 4 (Fig. 14B,C) MATERIAL. Partial basal circlet QMF38949 and partial radials QMF38950 from QML508, late Tournaisian, Malchi Formation. REMARKS. The basal circlet is an internal mould with the bare edges of parts of the exterior surface. It is uncrushed, high conical and thin. The thin partial radials are very elongate, unornamented and straight longitudinally. These specimens, if from one species, would have similar shape and proportions to P. nux, and the specimen would be tentatively referred to P. nux from the Caswell Creek Group. Camerate indet. Crinoid calyx Etheridge in Jack & Etheridge, 1892: 210, pl. 44. fig. 8. MATERIAL. QMF 1194, late Tournaisian ?Malchi Form- ation, Rockhampton District. Collected by C.W. De Vis. REMARKS. The weathered internal mould of a partial calyx reported by Etheridge (in Jack & Etheridge, 1892) is an indeterminate camerate. Plate sutures are indistinguishable but the long tegmen shows parts of the ambulacral trackways of two rays. Etheridge (in Jack & Etheridge, 1892) reported the specimen from the Gympie Beds, now considered to probably be from the Malchi Formation. Subclass DISPARIDA Moore & Laudon, 1943 Superfamily ALLAGECRINOIDEA Carpenter & Etheridge, 1881 Family ALLAGECRINIDAE Carpenter & Etheridge, 1881 Litocrinus Lane & Sevastopulo, 1982 TYPE SPECIES. Kallimorphocrinus punctatus Lane & Sevastopulo, ' 982 from the Visean New Providence Shale of Tennessee; by original designation, MEMOIRS OF THE QUEENSLAND MUSEUM Litocrinus sp. (Fig. I5D-F) MATERIAL. QMF39022, 39075 from QML878, Visean Baywulla Formation. DESCRIPTION. Cup conical, 0.65mm long. 0.8mm wide. Basal circlet fused, 0.1 5mm long, 0.5mm wide. Radials straight longitudinally, mod- erately convex transversely, moderately flaring: radial facets convex outward, subhorizontal. Small anal notch. Oral circlet moderately arched. Orals 5, concave; posterior oral larger, separating BC and DE orals. REMARKS. The cup of QMF39022 is coated with small secondary crystals masking plate sutures and giving the specimen a false ornament. The specimen has a higher basal circlet than L. scoticus and is not as elongate as L. extensus, both described from the Visean of Scotland (Wright, 1932, 1952). It is not as elongate as L. angulatus and L. tintinabuium, both from the Visean Nunn Shale of New Mexico (Strimple & Koenig, 1956). It also lacks the concave cup walls of L. protuberans and L. pansus, both reported from the Permian of Western Australia (Webster & Jell, 1992). It may represent a new species, but is an inappropriate specimen to serve as a type. The smaller immature specimen QMF39075, is 0.55mm long and 0.45mm wide, weathered and shows poor plate margins. This is the first report of Litocrinus from the Carboniferous of Australia. Superfamily BELEMNOCRINOIDEA S.A. Miller, 1883 Family SYNBATHOCRINIDAE S.A. Miller. 1889 Synbathocrinus Phillips, 1836 TYPE SPECIES. Syabathocrinus conicus Philips, 1836 from the Tournaisian of England: by monotypy. Synbathocrinus ogivalis de Koninck, 1878 (Fig. 15A-C) Sinbathocrinus ogivalis de Koninck. 1878: 158. pl. 6. figs 1-16: 1898: 121, pl. 6, figs 1-16, Bassler & Moodev. 1943: 696. Aynbuthocrinuis sp. Campbell & Bem. 1971: 424. pl. 49. figs 16-21, text-fig. 5. Webster. 1977-164. MATERIAL. MMF33432 from the late Tournaisian Namoi Formation near Barraba, NSW. DESCRIPTION. See de Koninck (1878, 1898) and Campbeil & Bein (1971). NEW CARBONIFEROUS CRINOIDS 263 HG 14. А. Plarverinites sp. 2. lateral view of radial QMF38946 «32 B.C. Platverinitessp. 4, B interior view of partial basal cirelet QMF38949 x2.5. C, lateral view of partial radial and associated columgal QME38950 «2.5. D.E, Platyeriuites sp. 3, D, view of disaruculated basal circlet and radials QMPE 38947 «2.9. E, lateral view of partial radial QME38948 »2 7. F-H, Platverinites sp. L. F, basal view of slightly distorted basal circlet MME33443 «1.7. GH, lateral views ol Weathered partial interior and exterior of opposite side of crushed calvx MME33240. X17, 1J. Platvermites nux? (Etheridge, 1892). lateral views of calyx interior, orientation uncertain. GSQF10870 «15 264 REMARKS. Synbathocrinus ogivalis was reported by de Koninck (1878) from an unknown horizon at Burragood, Paterson River, NSW. As noted by Campbell & Bein (1971), it probably came from the Lower Carboniferous. De Koninck's original description is minimal and his illustrat- ions are stylised. However, they are sufficient to define and recognise the species. De Koninck (1878) mistakenly reported the basal circlet to be a single plate for all species of Svabathocrinus including S. ogivalis, whereas Campbell & Bein (1971) showed 3 plates in their S. sp. Both de Koninck's (1873) and Campbell and Bein's (1971) descriptions are considered correct for the specimens investigated and show the variation in S. ogivalis. The basal circlet of Synbathocrinus is highly variable within most species known from multiple specimens. Webster & Lane (1987) reported the basal circlet of 2 undesignated species in the late Tournaisian Anchor Limestone of southern Nevada to vary from 1-4 plates. Two Artinskian species from Western Australia (Webster, 1987) varied from 1-4 plates іп $. campanulatus and from 1-3 in S. constrictus. The specimen illustrated herein has 3 plates in the basal circlet, with the EA basal azygous, but the sutures are difficult to define, because of partial fusing and recrystallisation. The type specimen of S. ogivalis is presumed lost in the Garden Palace fire of 1886. Therefore, ANU18893, (Campbell & Bein, 1971, pl. 49, figs 16-18) is designated the neotype. Subclass CLADIDA Moore & Laudon, 1943 Order CYATHOCRINIDA Bather, 1899 Superfamily CYATHOCRINITOIDEA Bassler, 1938 Family EUSPIROCRINIDAE Bather, 1890 Neerkolocrinus gen. nov. TYPE SPECIES. Neerkolocrinus typus from the West- phalian Neerkol Formation, Rockhampton. ETYMOLOGY. For the Neerkol Formation. DIAGNOSIS. Crown robust, expanding gently distally; cup medium to high bowl-shaped, with upflared basals; plate inflated, with stellate ridge ornament on basals and radials and fine node ornament on cup plates continuing on brachials; 5 anals in cup; prominent anal tube tapering distally above posterior interray; at least 10 arms, branching isotomously on 6th primibrach; at least one ramule on 3rd secundibrach in some rays; brachials rectilinear, strongly rounded trans- versely, lacking pinnules. MEMOIRS OF THE QUEENSLAND MUSEUM REMARKS. Neerkolocrinus shows some relation- ship to both euspirocrinids and botryocrinids. Genera assigned to both the euspirocrinids and botryocrinids have a conical or bowl-shaped cup, lacking stellate ornamentation and 1-4 anals. The presence of the proximal 1/2 of the first 2 tube plates below the radial summit does not alter the number of anals in the cup significantly, but is considered a primitive condition. First branching of the arms on the 6th primibrach is also con- sidered a primitive condition for such a young form of euspirocrinid or botryocrinid. The tapered anal tube above the posterior interray is present in some euspirocrinids (Euspirocrinus and Parisocrinus), whereas most botryocrinids have longer or recurved anal tubes. The radianal is beneath or below to the left of the C radial in most euspirocrinids and botryocrinids. Botryo- crinids typically have ramules and the brachials lack cover plates, which euspirocrinids and Neerkolocrinus have. Muscular articulation and an internal axial canal in the brachials of Neerkolocrinus also show relationships to the euspirocrinids rather than the botryocrinids. Euspirocrinids may be divided into 3 groups based on cup shape. Ampheristocrinus, Clostero- crinus, Parisocrinus and Zygotocrinus have narrow, long cone-shaped cups. Caelocrinus and Vasocrinus have short conical to bowl-shaped cups and Euspirocrinus and Neerkolocrinus have long wide bowl-shaped cups. Neerkolocrinus is distinguished from Euspirocrinus by the more bowl-shaped cup, more distal Ist branching of the arms, stellate ridge ornament and continu- ation of fine node ornament onto the brachials. Neerkolocrinus typus sp. nov. (Figs 16,17A,B) ETYMOLOGY. Latin typus, model or example; the type species. MATERIAL. HOLOTYPE: GSQF10872 from GSQLK 106, late Namurian or early Westphalian Neerkol Form- ation. DIAGNOSIS. As for genus. DESCRIPTION. Crown robust, elongate, flaring upward gently, 43.9mm long (incomplete), 19.4- 23mm wide (21.2mm av.). Cup medium high bowl-shaped, plates inflated, 17.6mm long, 13.6- 23mm wide (18.3mm av.); ornament of stellate ridges to adjoining plates on basals and radials, ray ridges more strongly developed than non-ray ridges; fine granular nodes with some alignment parallel to plate margins, nodes continuing on all brachials. Infrabasal circlet diameter 11.8mm NEW CARBONIFEROUS CRINOIDS 265 FIG. 15. A-C, Synbathocrinus ogivalis de Koninck, 1878, oral, posteriorand basal views of cup MMF33432 х4.2. D-F, Litocrinus sp., A ray, oral and posterior views of cup QMF39022 x50. internally. Infrabasals 5, slightly inflated. form- ing base of cup. upflared, visible in lateral view, distal tips forming base of cup walls. Basals 5. equidimensional, 9.5mm long and wide, mod- erately convex longitudinally and transversely, hexagonal: BC basal heptagonal supporting primanal on left shoulder; CD basal octagonal supporting first and second anals on right side and shoulder. Radials 5, slightly wider (10.2mm) than long (9.1mm), moderately convex longitudinally and transversely, subvertical to slightly incurved. Radial facet angustary, horseshoe- shaped. strongly declivate. Anals 5 in cup. Primanal pentagonal, below and left of C radial. also adjoining BC and CD basals, 2nd and 3rd anals. Second anal largest, hexagonal, adjoining primanal, CD basal, D radial. 2 anal tube plates and 3rd anal. Basal 1/2 of 4th and 5th anals below summit of radial facets. Anal tube extending well above radial summit over posterior interray, tapering distally. Arms at least 10, if isotomous branching in all rays on 6th primibrachs as in D and E rays. Brachials uniserial. rectilinear, : strongly convex transversely, wider than long proximally. distally subcircular transversely, nonpinnular. primibrachs 3.2mm long, 5.5mm wide. Third secundibrach giving off ramule in D ray. All brachial facets with transverse ridge across approximate middle of facet. deep transversely elongate ligament pit on outer margin, very small internal axial canal in middle of transverse ridge, elevated adoral area between transverse ridge and ambulacral groove, large muscle areas on inner margins, triarthrial articulation. Ambulacral groove deep V-shaped, notched on both sides; small cover plates not preserved. REMARKS. The type is preserved as external and internal moulds of a partial crown com- pressed along the A-CD plane of symmetry. Kopriacrinus gen. nov. TYPE SPECIES. Kopriacrinus mckellari gen. et sp. nov. from the Westphalian Neerkol Formation, W of Rock- hampton. ETYMOLOGY. Greek Kopria, dunghill, refers to the appearance of the cup as a cluster of pellets or a composite fecal pellet. DIAGNOSIS. Crown small; cup high cone-shaped; plates inflated; sutures impressed, apical pits; radial facet angustary, declivate; 3 anals; tegmen plates large, minimum 10; quadrate orals separated by large ambulacrals; tegmen sub- horizontal or projecting slightly above radial 266 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 16. Neerkolocrinus typus gen. et sp. nov., holotype, GSQF 10872. A, B-C interray view of internal mould x2.8. B, D-E interray view of external moul «2.8. C, basal view of internal mould x2.9. D, С ray view of external surface of cup x3.9. FIG. 17. (Opposite) A.B. Neerkolocrinus typus gen. et sp. nov., holotype, GSQF 10872. A, lateral view of external surface of partial arm x3.7. В, D-E interay view of internal mould x2.8. C-E, Kopriacrinus mckellari sp. nov C.D, internal x3.5) and external x4.7) views of paratype GSQF10874. Е. C ray view of cup with associated brachial fragments, holotype GSQF 10873 x3.7 NEW CARBONIFEROUS CRINOIDS 267 268 summit; at least 10 arms; rectilinear brachials strongly rounded transversely: stem transversely pentagonal, heteromorphic. REMARKS. Except for the pentagonal stem Kopriacrinus fits the Euspirocrinidae, which were defined by Lane & Moore (in Moore & Teichert, 1978) as having a round stem. Phylo- genetic significance of this feature is uncertain, but may imply a polyphyletic origin of the family. Kopriacrinus differs from all genera in the family by one or more other morphologic features. It is most similar to lasocrínus, which differs by having a low conical cup with large axial canals in the brachials and radials. Kopriacrinus mckellari sp. nov. (Fig. 17C-E; Table 3) ETYMOLOGY. For R.G McKellar in recognition of his contributions to the geology of Queensland. MATERIAL. HOLOTYPE: GSQF10873. PARATYPE: GSQF10874 from GSQL K106, late Namurian or early Westphalian Neerkol Formation. DESCRIPTION. Crown small, cylindrical to slightly expanding distally. Cup high cone shap- ed. with inflated plates, sutures impressed. apical pits. Infrabasal circlet with shallow invagination for pentagonal stem facet. Infrabasals 5, prox- imally horizontal, distally upflared forming base ofcup wall, visible inlateral view, bulbus, strongly convex longitudinally and transversely. Basals 5. hexagonal, wider than long. straight long- itudinally, moderately convex transversely, inflated, surface irregular, BD and CD basals heptagonal, with extra side adjoining anal plates. Radials 5, pentagonal, wider than long, strongly convex longitudinally and transversely, inflated, surface irregular. Radial facet angustary, declivate, projecting out from radial, horseshoe-shaped, with faint transverse ridge. Anals 3; radianal large. adjoining BC and CD basals, inflated. Anal X and right tube plate probably projecting above radial summit. Tegmen plates large. at least 10. Orals quadrate, filling radial notches, abutting large ambulacrals adorally. Ambulacrals large. expand- ing adorally, with longitudinal ridge. Tegmen subhorizontal or projecting slightly above radial summit. Anal tube unknown. Rectilinear brach- ials horseshoe- shaped transversely, straight to slightly concave longitudinally, with deep V- shaped ambulacral groove, with facets for cover plates. At least one isotomous branching of arms: probably 10 arms. Stem transversely pentagonal. heteromorphic. Noditaxis pattern N1. Columnals with large lumen. strongly convex latus. MEMOIRS OF THE QUEENSLAND MUSEUM REMARKS. The smaller holotype is an external mould of a slightly crushed cup with some associated brachials, whereas the larger paratype is an external and internal mould of a crown with disarticulated arm plates adjacent to the cup and 28.0mm of stem. The well-developed bulbous infrabasals are nearly equidimensional on the holotype, but longer than wide on the paratype. The holotype is compressed along the AB- posterior interray/D plane. The paratype is compressed along the A-CD plane of symmetry. Order POTERIOCRINIDA Jaekel. 1918 Superfamily POTERIOCRINITOIDEA Austin & Austin, 1842 Family POTERIOCRINITIDAE Austin & Austin, 1842 ‘Poteriocrinites’? smithii (Etheridge, 1892) (Fig. 18) Poteriocrinus? smithii Etheridge in Jack & Etheridge, 1892: 209, pl. 8, fig. 1. Poteriocrinites smithi Etheridge; Bassler & Moodey, 1943: 645. Branson, 1948: 209. MATERIAL. Plasticine cast BME15661 and plaster cast GSQF1590 stratigraphic unit uncertain, possibly from the Rockhampton Group, near Stanwell, Queensland. DESCRIPTION. Crown elongate, 33.0mm long. 21.2mm wide (arms flaring distally). Cup bowl- shaped, with stellate ridge ornament on basals and radials. Infrabasal circlet large, proximally horizontal, bearing stem facet, distally upflared. probably formed of 5 plates. Basals large. strongly convex longitudinally and transversely. forming base of cup walls; BC basal 3.9mm long. 2.9mm wide (estimated). Radials large, 2.1mm long, 4.0mm wide (estimated), moderately con- vex longitudinally and transversely, subvertical. with wide laterally sloping shoulders distally. Radial facet angustary. 1.7mm wide, with strong- ly convex outer rim, declivate, with radiating crenulae and culmina on outer margin. Radial notches wide. Anals 3, convex longitudinally and TABLE 3. Kopriacrinus тскеПағі measurements (mm). Holotype Paratype | F10873 F10874 | Cup length 84 —— 94 Cup width (maximum) 70 10,4 Infrabasal circlet diameter | 5.0 52 Basal length А Ao S8 Basal width 3.1 50 | Radial length 38 _ 44 || Radial width 37 6,7 Stem diameter Г 53 NEW CARBONIFEROUS CRINOIDS transversely. Radianal below C radial. pentag- onal, 2.8mm long, 1.7mm wide. adjoins C radial, BC basal, infrabasal circlet, CD basal and anal X. Anal X hexagonal. large, length 2.5mm, width 2.0mm, adjoins C radial. radianal, infrabasal circlet. CD basal, D radial, right tube plate. Right tube plate rectangular. in radial circlet above anal X. Stellate ridges sharply elevated; 4 ridges radiating from the B radial downward, 2 onto each subjacent basal; 3 ridges radiating from the C radial downward, | each onto BC basal, radianal anal X. One subhorizontal ridge ex- tending laterally from anal X onto radial to right and CD basal to left. Brachials slightly cuneate, narrow. straight longitudinally, strongly convex transversely, horseshoe-shaped transversely, with | very slender pinnule on long side. with medium nodes on exterior. First branching isotomous on 2nd primibrach and exotomous on 2nd secundibrach in 2 rays visible. Arms 3 per ray. 15 if all rays branch in same manner. Stem circular transverse- ly. heteromorphic, 1.5mm diameter, 37.0mm long. incomplete. Noditaxis N3231323 proximally, N; distally: nodals cirrate. multiple cirri per nodal. Columnals wide, with well rounded latus. Сїттї small, homeomorphic, transversely round. REMARKS. The description differs consider- ably from that of Etheridge (in Jack & Etheridge, 1892) in recognition of the infrabasal circlet and anals. Plate sutures are difficult to determine on the casts, but are visible with magnificationas the stellate ridges are slightly offset. The original interpretation is that the visible part of the cup consists of 2 partial radials, 1 on either side of a centrally complete radial; the underlying very wide basals are fractured and only the distal tips of 2 of the infrabasals are visible at the top of the stem. If this interpretation is correct, then the stellate ridge ornament varies considerably on the different radials and basals. The consistency in the stellate ridge ornament is accounted for in the new interpretation. The specimen is not a Poteriocrinites. It probably represents a new genus of the Poteriocrinitidac, based on the arm branching pattern and cup ornament, both of which differ significantly from all other genera assigned to the family. In hopes the original will be found or other specimens discovered we do not alter the original citation. Etheridge (in Jack & Etheridge, 1892) reported the specimen to be an impression in a hard sand- stone. Attempts to locate the original specimen failed but resulted in location of the materials listed above. The original specimen is apparently 269 FIG. 18. ;Poteriocrinites ? smithii (Etheridge, 1892), С ray view of plasticine cast of holotype, BME15661 x].7. lost. A note with the plaster cast, GSQF1590, reported the original specimen was loaned to F.A. Bather and returned to the Geological Survey of Queensland in 1915 with three plaster casts, only one of which remains. Etheridge's figure was drawn from ап earlier cast, perhaps one of the two plasticine casts in the British Museum (E15661). The stratigraphic position of ‘P’? smithii is uncertain. Etheridge (in Jack & Etheridge, 1892) reported the specimen from the Gympie Beds in the vicinity of Stanwell. As mapped by Dunstan 27) (1898) the Gympie Formation was the wide- spread basal umt of the Carboniferous in the Stanwell area. Currently, the Gympie Group is restricted to the Middle Permian of the Gympie Block (Day etal., 1982) and not recognised in the Stanwell area. There are several tossiliferous horizons in the Carboniferous and Permian in the vicinity of Stanwell. Without the original specimen the lithology cannot be matched and age of the specimen is in question. Supertamily SCYTALOCRINOIDEA Moore & Laudon, 1943 Family SCYTALOCRINIDAE Moore & Laudon, 1943 Prininocrinus Goldring, 1938 TYPE SPECIES, Prininocrinus robustus Goldring, 1938 from the Late Devonian of Canada; hy original designation, REMARKS. The A ray of the holotype of Pri- ninocrinus robustus Jacks the arm above the 1st primibrach and Goldring (1938) noted that 3 other rays branch on the 2nd primibrach, Furthermore, she described one of the paratypes as having 3 primibrachs in one ray, which could be the A ray. Onentation of the specimen did not allow unquestioned identification of the ray. Thus, it is unknown if Prininocrinus has ап atomous A ray or branched at some level above the 3rd primibrach. An alomous А ray is à primitive character of some Seytalocrinidac (Histocrinus, Hypselocrinus) among other primitive inadunates. However, it is not an invariable character. as the A ray branches (olten on primibrach 6 or higher) on some species assigned ta these genera. Although listed under the Seytaloerinidae Prininocrinus was incorrectly placed in synonomy with Sevta/ocrinus (Moore & Strimple, in Moore & Teichert, 19782640), as Ihe arms of Sevtalocrinus branch on the single primibrach in all rays. Prininocrinus is here aceepted and considered to have an atomous or branched A ray, Prininocrinus namoieusis sp. nov. Hiseceinis sp. Campbell & Bein, 197]. 423. pl 50. figs 8-9. їсхї-їг.. 4. ETYMOLOGY, From the Namoi Formation, MATERIAL. HOLOTYPE: ANU21344 from the late Tournaisian Namoi Formation, Crinoid Creek, near Burraba, NSW, DIAGNOSIS, Crown slender, elongate: cup low bawl-shaped. distal tips of basals visible in lateral view; 3 anals in cup; radial facets plenary: MEMOIRS OF THE QUEENSLAND MUSEUM brachials strongly convex transversely, rectilinear, isatomous branching on 2nd primibrach in all rays except atomous A ray; pinnules very slender, elongate. DESCRIPTION, Sce Campbell & Bein (1971: 423) REMARKS. The partial crown assigned to //isto- rinus by Campbell & Bein (1971) ts reassigned to Prininacriaus because the brachials are rectilinear not cuneate as in /Tisroerinus (Kammer & Ausich, 1992), Prininocrimus namotensis has a low howl-shaped cup with the distal tips ofthe infrabasals visible in lateral view, 2 anals in the cup and reculincarstrongly rounded brachials, P. nameirensis differs from P robustus by having a slightly shorter cup. less robust arms and more slender pinnules. The inlrabasals, essentially confined io the basal plane of the cup (except for the distal upflared tips), and the intermediate position of the 3 anals (primanal moved wholely or partly into the CD interray position) are slightly advanced conditions in P robustus and P. namoiensis, Seytalocrinid? indet, (Fig. 11) MATERIAL QME38958, lacality and horizon unknown in NSW, probably late Toumaisian Namor Formation. Collected by GM. Philip. DESCRIPTION, Cup small, truncated cone- shaped, no ornament. infrabasal circlet forming truncated base proximally, weakly flared distally forming lower 1/4 of cup wall. Basals subequal length and width, straight longitudinally, gently convex transversely, gently flaring distally; CD basal truncated distally for reception of anal X. Radials gently flaring longitudinally, moderately convex transversely, Radial facet plenary. projecting slightly, rimibraeh widest at base, lade-shaped proximally, constricted medially, strongly convex transversely shortly distal io base, concave longitudinally; distal end unknown. Proximal stem heteromorphic, with noditaxis Ni, becoming homeomorphic distally. Columnals circular transversely; latus strongly convex. REMARKS. This small partial erown is crushed and the cup plates are parily disarticulated with the ыча ам of the radial covered by sub- jacent basals, The truncated end of the CD basal 15 narrow, suggesting there was a radianal to the lower right of anal X and 3 anals in the cup. Cup and primibrach shapes, in conjunction with the NEW CARBONIFEROUS CRINOIDS plenary radial facet, suggest affinity with Scytalocrinidae such as ZHvdriocrinus and Hypselocrinus but without the arms and anal series no generic assignment can be made. The specimen is associated with an unnamed new genus of rhodocrinitid and Dichocrinus cf. D. laudoni on a small slab. Superfamily ERISOCRINOIDEA Wachsmuth & Springer, 1886 Family GRAPHIOCRINIDAE Wachsmuth & Springer. 1886 Holcocrinus Kirk, 1945 TYPE SPECIES. Graphiocrinus longicirrifer Wachsmuth & Springer, 1890 from the Tournaisian Hampton Formation, Iowa; by original designation. Holcocrinus barrabaensis sp. nov. (Fig. 19) ETYMOLOGY. From Barraba, NSW. MATERIAL. HOLOTYPE: MMF33608 (AMSF59510 and 59511 are casts of the type) from the late Tournaisian Namoi Formation, Crinoid Creek, near Barraba, NSW; found by J. Irving. DIAGNOSIS. Crown elongate; cup truncated medium cone, primibrachs axillary. intermediate length: brachials intermediate length, moderately cuneate; unornamented. DESCRIPTION. Crown clongate, 54.9mm long (incomplete), 21.5mm wide. Cup truncated medium cone, 7.0mm long, 7.5-11.5mm wide (9.5mm av.). lacking ornamentation. Infrabasals 5, horizontal proximally, upflared distally, visible in lateral view. Basals 5, 3.6mm long, 3.8mm wide, hexagonal, except CD basal heptagonal, gently convex transversely and longitudinally. Radials 5, 5.8mm long. 4. 1mm wide, moderately convex transversely, gently convex longitudin- ally. Radial facet plenary. Single anal, widest distally, 4.2mm long, 3.8mm wide, slightly con- vex transversely and longitudinally, projecting slightly above radial summit. Axillary single primibrachs equidimensional, 5.8mm, strongly convex transversely, concave longitudinally. hourglass-shaped. Secundibrachs cuneate, short, moderately convex transversely, concave longitudinally, faintly staggered, probably with pinnule on shoulder of longerside. Arms 10, 2 per ray, branching isotomous. Stem homeomorphic. 58.0mm preserved, proximally transversely pent- agonal for 14.5mm (5.0mm diameter), round distally (3.7mm diameter). Tegmen unknown. 271 FIG.19. Holcocrinus barrabaensis sp. nov.. A.B, posterior (AMSE 59511) and anterior (AMSF'59510) views, х1. REMARKS. The holotype was embedded in mudstone. A cast (AMF59510) shows 58.0mm of stem, Part of the stem is now missing on the original. Webster (1997) noted that Holcocrinus has cuneate brachials and suggested that it belongs to a clade of conservative inadunates retaining à conical cup. The infrabasals forming the trun- cated base of the cup with only the distal tips visible in lateral view and a single anal are more advanced features within this conservative clade. The crown of H. harrabaensis is crushed normal to the BC-DE plane. Holcocrinus barrabaensis is distinguished within the genus by having a truncated medium conical cup, brachials of intermediate length and lacking ornamentation. The cup of H. wachsmuthi (Meek & Worthen, 1861) is a much higher truncated cone. H. spinobrachiatus (Hall, 1861) has a broader medium bow! shape and all other species have low bowl-shaped cups. Brachials of Н. /ongi- cirrifer (Wachsmuth & Springer in Miller, 1889) are very short and strongly cuneate, whereas those of H. wachsmuthi are longer, triangular and nearly biserial. Brachials of H. nodobrachiatus (Hall, 1861) and H. spinobrachiatus have short nodes or spines on the distal ends of the mod- erately cuneate brachials. Brachials of =H. barrabaensis lack the spines, are longer than those of H. longicirrifer and shorter than those of H. spinobrachiatus. 'The axillary primibrachs of H. smythi are much longer than those of H. barrabaensis which are longer than those of 77. longicirrifer. Poteriocrininid cup indet. Crinoid allied to Stemmatocrinus Etheridge in Jack & Etheridge, 1892: 206, pl. 44, fig 7. MATERIAL. QMF1196 probably from the late Tournaisian Malchi Formation, Rockhampton District. Collected by C.W. De Vis. REMARKS. An indeterminate poteriocrininid cup 10.6mm wide is a slightly distorted internal mould and parts of 3 weathered radial facets. The basal circlet is subhorizontal and formed of 5 plates. Basals were horizontal proximally, upturned on the distal tips. One basal is wider than the other 4 and presumably had | or 2 anals above it. Cup walls were largely formed by the subvertical radials. Etheridge (in Jack & Etheridge, 1892) apparently thought the speci- men lacked an anal plate when he considered it allied to Stemmatocrinus (=Erisocrinus). The cup could belong to any number of the poteriocrini- nids, depending on the number of anals and is considered indeterminate. Etheridge (in Jack & Etheridge, 1892) reported the specimen from the Gympie Beds, now considered to probably be from the Malchi Formation. Poteriocrininid arms indet. #1 (Fig. 20A) Arms of crinoid Etheridge in Jack & Etheridge. 1892: 210, pl. 7. fig. 8. Рагітеу, 1996: 242. MATERIAL. GSQF1588, external mould ofa partial set of arms from the Tournaisian Rockhampton Group, Stony Creek, Stanwell. Collected by J. Smith. DESCRIPTION. Partial set of 10 arms of at least 3 rays, 27.0mm long, 22.0mm wide. flaring distally. MEMOIRS OF THE QUEENSLAND MUSEUM FIG: 20). A, Poteriocrininid indet. arms #1, lateral view of arms GSQF 1588, «2.5. B, Poteriocrininid indet. arms #2, lateral view of arms GSQF1587, *2. Brachials smooth, no ornament, weakly cuneate, straight longitudinally. strongly convex NEW CARBONIFEROUS CRINOIDS transversely. deep. U-shaped in transverse section, with slender pinnule on longer side. Ambulacral groove small. V-shaped. First secundibrach much longer than following secundibrachs. formed by fusing of 2 brachials. Isotomous branching on only primibrach preserved and axillary secundibrach 6. REMARKS. The brachial shape. branching pattern and longer 1st secundibrach are common to aphele- crinids to which this specimen may be related. The aphelccrinids are most common in Tour- naisian and early Viscan. strata of the United States and Scotland (Bassler & Моойсу. 1943: Webster, 1973, 1977, 1986, 1988, 1993). The specimen is re-illustrated because the original does not show the distal part of the arms and is. in part. a reconstruction of part of the arms that are masked by matrix. No part of the cup is recognisable in the few fragmentary plates at the base of the arms, Poteriocrininid arms indet. #2 (Fig, 20B) Arms of cnmoid Etheridge in Jack & Etheridge, 1892: 210, pl. 7, fig, 7. Parfrey. 1996; 242, MATERIAL, 5011597. external mould of partial set of arms, from the Rockhampton Group, Rhytichonella Gully. Stanwell. Collected by J. Smith. DESCRIPTION. Partial set of 8 arms. 34.6mm long. 30.8mm wide, Brachials weakly cuneate, straight longitudinally. strongly convex trans- versely. nearly circularin transverse section. bear transverse row of 3-5 coarse nodes projecting as exceedingly short spines. Ambulacral groove small, V-shaped. Pinnules slender. elongate. REMARKS. Brachials of most poteriocrinids lack опе. When present. the ornamentation commonly consists of fine granules or longi- tudinal or transverse ridges. Thus, the projecting spine-like nodes on the brachials of Poleriocrinid arms indet. #2 are very distinctive. Ch/idenocrin- us echinatus, à Late Carboniferous poteriocrinid from the United States (Strimple & Watkins. 1969), has similar spine-like node ornamentation but only 1 - 2 per brachial and not in an aligned transverse row. The brachials of Neerkolocrinus ivpus have irregular nodose ornament. not aligned in а transverse row and the brachials are much longer. The original illustration From the extemal mould does not show pinnules, which were not mentioned in the description (Etheridge in Jack & Etheridge. 1892). Without the cup. the specimen is left in open nomenclature. FIG. 21. l'axoerinidindet., lateral view of partial crown ОМЕЗ8951. «3.3. Subclass FLEXIBILIA Zittel, 1895 Order TAXOCRINIDA Springer, 1913 Superfamily TAXOCRINOIDEA Angelin, 1878 Family TAXOCRINIDAE Angclin. 1578 Taxocrinid indet. (Fig. 21) MATERIAL. QME3895] from QML308_ laic. Tourn- aisan. Маі l'ormatior, DESCRIPTION, Crown slender. elongate. Cup high truncated cone. with unornamenied plates, with apical pits. Basals large. 7.3mm wide. strongly convex transversely with shallow longi- tudinal medial trough. distally upflared. distal tips incurved. Radials large. 14.0mm. long, 14.2mm wide (incomplete). slightly conves longitudinally. strongly convex. transversely, Radial facets angustary. with sharply convex outer rim. sloping inward. wiih short central transverse ridge 1/3 width of facet. with elongate ligament pit and large outer area aboral of ridge: inwardly with transverse pit adjacent to trans- verse ridge. Anal large. proximal 1/2 in line of radials, Brachials rectilinear, concave or straight longitudinally. strongly convex transversely. deep. not m contact laterally. with faint development of patelloid process. with deep wide V-shaped аиби lacral groove. Brachial facets concave. with fine 274 MEMOIRS OF THE QUEENSLAND MUSEUM VIG. 22. A.B, Sagenocrinitoid indet., lateral views of opposite sides of disarticulated crown QMI' 389522 and b x 2.6. crenellae and culmina radiating from apex of ambulacral groove to aboral edge of facet cover- ing an arc of 195°. Arms branching at least once and probably several times. Interprimibrachs unknown, but probably present. Anal tube plates large. with crenulate edges. Stem unknown. REMARKS. QMF38951 is the external mould of both sides of a crushed. disarticulated crown. The specimenis assigned to the Taxocrinidae because the radials are large. upflaring and have angustary radial facets, and the arms were not abutting. It is uncertain whether the cup includes more than 1 anal. Articular facets of the brachials of Forbesio- crinus nobilis de Koninck & Le Hon, 1854 (Springer, 1920. pl. 24, figs 13, 15, 18. 19. 23), Svnerocrinus incurvus (Trautschold, 1867) (Springer, 1920, pl. 42, figs 8i, 8k) and Parichthvocrinus nobilis (Wachsmuth & Springer, 1880) (Springer. 1920, pl. 61, figs 5-8) have narrow crenularia only along the outer edges of the facet. The crenularium of this taxon is quite different, radiating from the growth centre to the outer edges of the ossicle. The specimen probably represents a new genus, but is too incomplete to serve as a holotype. Order SAGENOCRINIDA Springer, 1913 Superfamily SAGENOCRINITOIDEA Roemer, 1854 Sagenocrinitoid indet. (Fig. 22) MATERIAL. QMF38952 from QML508, late Toum- aisian, Malchi Formation. DESCRIPTION. Partial set of arms, small. Brachials weakly convex longitudinally. moder- ately convex transversely; (smooth fine granulate surface reflects grains in matrix). Patelloid process small, weakly developed. Arms branch isotomously twice; Ist branching on 3rd brachial preserved. 2nd branching on 4th brachial of adjacent parts of first branching. Distal part of interbrachial. interprimibrach. or anal series 2 NEW CARBONIFEROUS CRINOIDS 275 large plates followed by numerous small irreg- ular plates. REMARKS. This fragmentary specimen may represent parts of 2 rays or part of ] ray. It is assigned to the Sagenocrinitoidea based on the interprimibrach or interbrachial series and branch- ing pattern of the brachials. ACKNOWLEDGEMENTS Andrew Rozefelds helped collect the Rock- hampton Group. Loan of specimens by thc curators in charge of collections at the Queens- land Geological Survey, Geological Survey of New South Wales, Australian Museum, Aust- ralian National University and British Museum is greatly appreciated. Paul Avern processed photographs. GDW extends his appreciation to Washington State University for granting professional leave and to the Director of the Queensland Museum for use of facilites and office space during prosecution of this project. The reviews of Bill Ausich and George Sevasto- pulo are kindly acknowledged. LITERATURE CITED AUSICH, МІ. & KAMMER, T.W. 1991. Late Osagean and Meramecian Actinocrinites (Echinodermata: Crinoidea) from the Mississippian stratotype region. Journal of Paleontology 65: 485-499. AUSTIN, T. & AUSTIN, T. 1844. A monograph on Recent and fossil Crinoidea, with figures and descriptions of some Recent and fossil allied genera. 3: 33-48, pls 5-6 (London). ВАМВАСН, R.K. 1990. Late Palaeozoic provinciality in the marine realm. Geological Society Memoir 12: 307-323. BASSLER, R.S. & MOODEY, M.W. 1943. Biblio- graphic and faunal index of Paleozoic pelmatozoan echinoderms. Geological Society of America Special Paper 45: 1-734. BRANSON, C.C. 1948. Bibliographic index of Permian invertebrates. Geological Society of America Memoir 26: 1-1049. BROADHEAD, T.W. 1981. Carboniferous camerate crinoid subfamily, Dichocrininae. Palaeontographica, Abteilungen A 176: 81-157. BROWER, J.C. 1967. The actinocrinitid genera Abactinocrinus. Aacocrinus and Blairocrinus. Joumal of Paleontology 41: 675-705. CAMPBELL, K.S.W. & BEIN, J. 1971. Some Lower Carboniferous crinoids from New South Wales. Journal of Paleontology 45: 419-436. CAMPBELL, K.S.W. & McKELLAR, R.G. 1969, Eastern Australian Carboniferous invertebrates: sequence and affinities. Pp. 77-119. In Campbell, K.S.W. (ed.), Stratigraphy апа palaeontology. (Australian National University Press: Canberra). DAY, R.W, WHITAKER, W.G. MURRAY, C.G. WILSON, І.Н. & GRIMES, K.G. 1982. Queens- land geology. Geological Survey of Queensland Publication 383: 1-194. DUN, W.S. & BENSON. W.N. 1920. The geology and petrology of the Great Serpentine Belt of New South Wales. Part 9 — The geology, palaeontology and peirography of the Currabubula district, with notes on adjacent regions. Section B — Pal- aeontology. Proceedings of the Linnean Society of New South Wales 45: 337-374, pls 18-24. DUNSTAN, B. 1898. The Mesozoic Coal Measures ol Stanwell and associated formations. Geological Survey of Queensland Publication 131: 1-21. ETHERIDGE, R. Jr 1892. A monograph of the Carbon- iferous and Permo-Carboniferous invertebrata of New South Wales. Part П. Echinodermata. Annelida, and Crustacea, Memoirs of the Geological Survey of New South Wales, Palae- ontology 5: 65-131, pls 12-22. GOL DRING. W. 1938. Devonian crinoids from the Mackenzie River Basin (NWT) Canada. Bulletins of American Paleontology 24(81): 1-23. HALL, J. 1858. Palaeontology of Iowa. Report of the Geological Survey of the State of lowa 1(2): 473-724. 1861. Descriptions of new species of Crinoidea from the Carboniferous rocks of the Mississippi Valley. Journal of the Boston Society of Natural History 7: 261-328. JACK, R. L. & ETHERIDGE, R. Jr 1892. The geology and palaeontology of Queensland and New Guinea. (J.C. Beal, Govt Printer: Brisbane). KAMMER, ТА. & AUSICH W.I. 1992. Advanced cladid crinoids from the Middle Mississippian of the east-central United States; primitive-grade calyces. Journal of Paleontology 66: 461-480. KONINCK, L.G de 1877-1878. Recherches sur les Fossiles Paléozoiques de la Nouvelle Galles du Sud (Austrálie). Mémoires de la Société Royale Liege, (1877), ser. 2, Part 1, Texte: 1-373: Part 2. atlas, 24 pl. F. Hayez. Bruxelles. Crinoiden: Troisiéne Partie, Fossiles Carboniléres, pp. 158-166, pl. 6 (1878). 1898. Descriptions of the Palaeozoic fossils of New South Wales (Australia): Memoirs of the Geological Survey of New South Wales. Palae- ontology 6: 1-298, pls 1-24. KONINCK. d: .G. de & Le HON, Н. 1854, Recherches sur les crinoides du terrain carbonifere de la Belgique. Academie Royal de Belgique Memoir 28(3): 1-215. LANE, N.G. & SEVASTOPULO, G.D. 1987. Strati- graphic distribution of Mississippian camerate crinoid genera from North America and western Europe. Courier Forschunginstitut Senckenberg 98: 199-206. 1990. Biogeography of Lower Carboniferous crin- oids. Geological Society Memoir 12: 333-338, LINDLEY, LD. 1979. An occurrence of the camerate crinoid genus Eumorphocrinus in the Early 276 Carboniferous of New South Wales. Journal and Proceedings of the Royal Society of New South Wales 112:121-124. 1988. Glaphvrocrinus, a new camerate crinoid genus from the Lower Carboniferous of New South Wales. Alcheringa 12:129-136. LISHMUND, S.R., DAWOOD, A.D. & LANGLEY. W.V. 1985. The limestone deposits of New South Wales. Geological Survey of New South Wales Mineral Resources 25: 1-373. McKELLAR, R.G. 1966. A revision of the blastoids "Mesoblastus? australis’, “Granatocrinus? wachsmuthii and “Tricoelocrinus? carpenteri^ described by Etheridge (1892) from the Carboniferous of Queensland. Memoirs of the Queensland Museum 14: 191-198, pl. 24. MILLER, S.A. 1889. North American geology and paleontology. (Western Methodist Book Concern: Cincinnati ). 1892. North American geology and paleontology, first appendix. ( Western Methodist Book Concern: Cincinnati) pp. 665-718. MILLER, S.A. & GURLEY, W.F.E. 1890. Description of some new genera and species of Echinodermata from the Coal Measures and Subcarboniferous rocks of Indiana, Missouri, and Iowa. Journal of the Cincinnati Society of Natural History 13: 1-25, pls 1-4. MINATO, M. 1951. On the Lower Carboniferous fossils of the Kitakami Massif, NE Honsyu, Japan. Journal of the Faculty of Science, Hokkaido University ser. 4, 7: 355-382. MINATO, M.. HUNAHASHI, M., WATANABE, J., KATO, M. (eds) 1979. The Abean Orogeny. (Tokai University Press: Tokyo). MOORE, R.C. & LAUDON, L.R. 1943. Evolution and classification of Paleozoic crinoids. Geological Society of America Special Paper 46: 1-151. MOORE, R.C. & STRIMPLE, H.L. 1969. Explosive evolutionary differentiation of unique group of Mississippian-Pennsylvanian camerate crinoids (Acrocrinidae). University of Kansas Paleontological Contributions Paper 39: 1-44. MOORE, К.С. & TEICHERT. К. (eds) 1978. Treatise on invertebrate paleontology. Part T. Echino- dermata 2. 3 vols. (Geological Society of America and University of Kansas: Lawrence, Kansas). PICKETT, J.W. 1960. Note on a Carboniferous crinoid from Swain's Gully, Babbinboon, NSW. The Australian Journal of Science 23: 88. SHUMARD, В.Е. 1858. Description of new fossil Crinoidea from the Palaeozoic rocks of the western and southern portions of the United States. ‘Transactions of the St Louis Academy of Science (1857) 1: 71-80. SPRINGER, F. 1920. The Crinoidea Flexibilia (2 vols). Smithsonian Institution Publication 2501: 1-486, pls A-C, 1-76. STRIMPLE, H.L. & KOENIG, J.W. 1956. Missis- sippian microcrinoids from Oklahoma and New Mexico. Journal of Paleontology 30: 1225-1247. MEMOIRS OF THE QUEENSLAND MUSEUM TERMIER, G. & TERMIER, H. 1950. Paléontologie Marocaine II. Invertébres de l'Erde Primaire. 4. Annélides, Arthropodes, Echinodermes, Conularides et Graptolithes. Service Carte Géologique Morocco, Notes et Mémoires 79(4): 1-279, pls 184-241. TRAUTSCHOLD, H. 1867. Einige crioideen und andere Thierrests des Jungeren Bergkalks im Bouvemment Moskau. Bulletin de la Societie Imperial Naturalistes de Moscou 15(3-4): 1-49. WACHSMUTH, C. & SPRINGER, F. 1880. Revision of the Palaeocrinoidea. Part 1. Proceedings of the Academy of Natural Sciences of Philadelphia 1880: 226-378, pls 15-17. 1897. The North American Crinoidea Camerata. Harvard College Museum of Comparative Zoology Memoir 20 (3 vols), 21: 1-897. WEBSTER. GD. 1973. Bibliography and index of Paleozoic crinoids, 1942-1968. Geological Society of America Memoir 137; 1-341. 1974. Crinoid pluricolumnal noditaxis pattern. Journal of Paleontology 48: 1283-1288. 1977. Bibliography and index of Paleozoic crinoids, 1969-1973. Geological Society of America Microform Publication 8; 1-235. (3 cards) 1986. Bibliography and index of Paleozoic crinoids, 1974-1980. Geological Society of America Microform Publication 16: 1-405. (5 cards) 1987. Permian crinoids from the type-section of the Callytharra Formation, Callytharra Springs, Western Australia. Alcheringa 11: 95-135. 1988. Bibliography and index of Paleozoic crinoids and coronate echinoderms, 1981-1985. Geol- ogical Society of America Microform Publication 18; 1-235. (3 cards) 1993. Bibliography and index of Paleozoic crinoids, 1986-1990. Geological Society of America Microform Publication 25: 1-204. (3 cards) 1997. Lower Carboniferous echinoderms from northern Utah and western Wyoming. Utah Geological Survey Bulletin 128, Paleontology Series 1: 1-65. WEBSTER, G.D. & JELL, P.A. 1992. Permian echinoderms from Western Australia. Memoirs of the Queensland Museum 32(1): 311-373. 1998. New Permian crinoids from Australia. Mem- oirs of the Queensland Museum 43(1): 279-339. WEBSTER, GD. & LANE, N.G. 1987. Crinoids from the Anchor Limestone (Lower Mississippian) of the Monte Cristo Group southern Nevada. PEST of Kansas Paleontological Contributions Paper 119: 1-55. WRIGHT, J. 1932. The Scottish species of Allage- crinus. Geological Magazine 69: 337-366, pls 23-25. 1952. The British Carboniferous Crinoidea. Pal- aeontographical Society Monograph 1(5): 149-190. 1955. The British Carboniferous Crinoidea. Palaeontographical Society Monograph 2(1): 191-254, pls 48-63. NEW CARBONIFEROUS CRINOIDS XRF APPENDIX 1 Locality Register. GSQ K-21 - Hill behind Pearson's Homestead. 0.8mile N of Old Cannindah road intersection along Cannindah Road, Caswell Creek Group. Collected by R. McKellar, GSQ K-106 - Along side road off Stanwell-Dalma road. Rockhampton Sheet 1:250,000 yd grid:31760852; Ridge- lands 1:100,000 Sheet GR 22€ 34101: Late Carboniferous, Neerkol Formation. Collected by R. McKellar. GSQL334 - Neerkol Formation, Late Carboniferous, W of Rockhampton. GSQL3006 - Crow's Nest, NW of Mt Morgan, Mount Morgan 1:100,000 Sheet GR. 284854; Gympie Beds of Jack & Etheridge (1892). Collected by J. Smith, 1888. GSQL3012 - Malchi Creek, probably Malchi Formation, W of Rockhampton, Queensland. ОМІ.508 - Mrs Harding’s property, low hills Ikm SE of homestead, above limestone hardground. K V252152; 8915 Ridgelands 1:100,000 Sheet, Rockhampton Group. Collected by Р.А. Jell & A. Rozefelds. QMLS878 - 2000" W of Old Cannindah Homestead, Queensland. Oolitic limestone in Baywulla Formation, Early Carboniferous. QML1248 - North side of hill behind Old Cannindah homestead, Early Carboniferous Caswell Creek Group, Tournaisian or possibly Visean Tellebang Limestone. 18.0 43.5 Monto 1:100,000 Sheet. APPENDIX 2 Listof described Carboniferouscrinoid taxa from stratig preceded by an asterick (*) are not treated systematically Neerkol Formation, Westphalian, Old. Denarioacrocrinus neerkolensis gen. et sp. nov. Denarioacrocrinus? ornatus sp. nov. Neerkolocrinus typus gen. et sp. nov Kopriacrinus mckellari sp. nov. Prininocrinus namoiensis sp. nov. Baywulla Formation, Visean, Old. Litocrinus sp. Caswell Creek Group, Visean?. Old. Aacocrinus sp. 1 Sampsonocrinus cannindahensis sp. nov. Dialutocrinus? sp. Platycriniies nux? (Etheridge, 1892) Rockhampton Group, Malchi Formation, late Tournaisian, Old. Actinocrinites sp. 1 Actinocrinites? sp. 3 Aacocrinus sp. 1 Manillacrinus brownei (Dun & Benson. 1920) Platycrinites sp. 2 Platycrinites sp. 3 Platycrinites sp. 4 Camerate indet. *Poteriocrinites '? smithii (Etheridge, 1892) Poteriocrininid cup indet. Poteriocrininid arms indet. #1 Poteriocrininid arms indet. #2 Taxocrinid indet. Sagenocrinitoid indet. Мез Creek Clastics, early or middle Tournaisian, Old. Actinocrinites sp. 2 raphi units of Queensland and New South Wales. Taxa y erein. Tellebang Limestone, Tournaisian or Visean, Old. Aacocrinus acylus sp. nov. Unknown horizon, possibly Namoi Formation, late Tournaisian, NSW. Rhodocrinitid gen. nov. Dichocrinus cf. D. laudoni Broadhead, 1981 Scytalocrinid? indet. Namoi Formation, late Toumaisian, NSW. *Cribanocrinus biseriatus Campbell & Bein, 1971 *Glyphyrocrinus expansus Lindley, 1988 Manillacrinus acanthus sp. nov. Manillacrinus brownei (Dun & Benson, 1920) Platycrinites testudo Campbell & Bein, 1971 Platycrinites sp. 1 Holcocrinus barrabaensis sp. nov. Goonoo Goonoo Mudstone, late Tournaisian, NSW. *Manillacrinus sp. Campbell & Bein, 1971 Platycrinites testudo Campbell & Bein, 1971 *Platycrinites sp. 2 Campbell & Bein, 1971 Synbathocrinus ogivalis de Koninck, 1878 *Cyathocrinites sp. Campbell & Bein, 1971 Dangarfield Formation, late Tournaisian, NSW. D umorphocrinus elongatus Lindley, 1979 *Glaphyrocrinus expansus Lindley, 1988 *Glaphyrocrinus minutus Lindley. 1988 Flagstaff Formation?, Visean?, NSW. Actinocrinitid indet. NEW PERMIAN CRINOIDS FROM AUSTRALIA GARY D. WEBSTER AND PETER A. JELL Webster, G.D. & Jell, P.A. 1999 06 30: New Permian crinoids from Australia. Memoirs of the Queensland Museum A3( 1). 279-339. Brisbane. ISSN 0079-8835. New crinoids are described from the Permian of Queensland, New South Wales, ‘Tasmania. and Western Australia. They 1. strengthenatfinities with Timorand North America: 2, add to knowledge of biodiversity: 3, improve knowledge of some earlier described taxa: and 4. extend the stratigraphic value of Хеосатріосгіпих. Range of the Isocrinidae is extended down to the Artinskian, based on Archaeoisocrinus occiduaustralis gen. et sp. nov. The new Order Ampelocrinida which is recognised by syzvgial brachial pairs in which muscular articulation alternates with ervptosyzygialarticu- lation 1s assigned to the Articulata and includes the Ampelocrmidae, Corythocrinidae. Calceolispongiidae and Tribrachvocrinidae. Euspirocrinids are recognised in the Artinskian and possibly Roadian of eastern Australia. extending their range from the Visean. Three flexible crinoids are recognised їп the Artinskian of WA. Cymbiocrinus cherrabunensis is designated the type species of the A/etacalceolispongia gen. nov. Other new genera and species described are Anaglyptocrinus willinki, Necopino- crinus tvcherus and Eidosocrinus condaminensis, New species described are Platverinites halos, Auliskocrinus? bananaensis, Neocamptocrinus catherinensis. Spaniocrinus geniculatus, Glaukosocrinus middalyaensis, Pedinocrinus? nodosus, Moapacrinus cuneatus and Sundacrinus medius. O Crinoids, Permian, Queensland, New South Wales, Tasmania, Western Australia. Gary D. Webster, Department of Geology, Washington State University, Pullman, Washington 99164-2812, USA; Peter A. Дей, Oueensland Museum, P.O. Box 3300, South Brisbane 4101, Australia; 23 Мау, 1998. Permian crinoids of Western Australia werc reviewed by Webster (1987) and Webster & Jell (1992). Permian crinoids of eastern Australia were reviewed in Webster (1990). Those reviews may be summarised here by noting that the earliest descriptions of Australian crinoids were in 1847, most species have been described since 1949, and that crinoids are widespread in the marginal Permian basins of both eastern and Western Australia. Paleobiogeography of Australian Permian echinoderms (Webster et al.. in press) may be summarised by noting that: I. Permian echino- derms are common clements in deposits of eastern Australia from Sakmarian into Wordian time and in WA from Sakmarian into Wuchiaping- ian time: 2. Australian echinoderm faunas are dominated by taxa endemic to Australia and the Tethys. but contain a few taxa found throughout the equatorial belt; 3. Australian faunas show greatest affinity to the faunas of Timor. but con- tain some North American Carboniferous taxa that are holdovers in Australia or the Tethys: and 4. Australian echinoderms lived in a cooler water. clastic-richenvironment of deposition S of 35 S The purpose of this paper is to: 1. describe new Permian crinoids from Australia: 2. provide new information on some previously described taxa: 3. relate the new material to previously described faunas: and 4. denote the significance of an Early Permian articulate crinoid in WA. FAUNAL ANALYSIS Western Australian crinoids are reported from 5 horizons. This includes previously unreported taxa in 3 horizons. and new information on taxa from the other 2 horizons (Table 2). The Permian crinoid fauna of the Cally tharra Formation is the most diverse in Australia and second largest in the world. Webster & Jeli (1992) reported 16 camerates. 37 inadunates. and | flexible. increased here to 42 inadunates (6 disparids. 3 cvathocrinids. and 33 poteriocrin- inids) and 7 flexibles. Most of the new material consists of disarticulated radial plates and frag- mentary sets of arms. The 3 species each of Prophvllocrinus and Loxocrinus. and the arms of a timorechinid show increased affinity of Cal- lytharra and Timor (especially Basleo) faunas. This supports proposed correlation of the Cal- Iytharra fauna with part of the Basleo fauna and an Artinskian age for that part of the Basleo faunas (Webster. 1987: Webster & Jell. 1992). Glaukosocrinus in the Callytharra fauna is its first report outside North America, extending the range from the Late Carboniferous into the Permian. A crown of Archaeoisocrinus occiduaustralis gen. et sp. nov., found in the arms of a crown of Jimbacrinus bostocki, from the Cundlego Form- ation on Jimba Jimba Station is the earliest known articulate crinoid. The articulate crinoids were previously known from the Early Triassic to the Recent, thus their range is extended into the Palaeozoic approximately 28 m.y. The Wandagee Sandstone yielded an abund- ance of loose columnals of several species of Calceolispongia, whereas articulated cups and crowns are rare. No new taxa are described from the Wandagee, but details of C. abundans and C. rubra suggest that all calceolispongiids: 1, lived either resting on or partly buried within the substrate, with the stem serving as a runner or vestigial tether in the adult stages; 2, had syzygial articulation. facilitating differential movement between laterally adjacent Ist and 2nd brachials of each ray; and 3, that together, the Ist and 2nd brachials of all rays formed a facultative tegmen capable of gentle expansion and contraction, as needed for capacity adjustment of the gut tract. Discovery of an in situ nest and its partly weathered components consisting of several crowns, partial crowns, and fragments of Neocamptocrinus millyitensis in the Cherrabun Member of the Hardman Formation provides new information on the arms, and proximal and medial stem of this Wuchiapingian camerate. Crinoids of eastern Australia are described from 7 stratigraphic units from early Artinskian to Wordian. This includes new taxa from 5 of the units and new information on taxa from 2. A reconstructed cup and proximal brachials of Calceolispongia gerthi from the Sakmarian Berridale Limestone of SE Tasmania has a cylindrical shape that rested on the substrate probably attached by a runner type stem. This supports the interpretation as discussed for C. abundans and C. rubra. Pentastellate columnals and disarticulated cup plates of Nowracrinus ornatus from the early Artinskian Kansas Beds of NW Tasmania are recognised in the lineage of the early articulate crinoids. Columnals have a crenularium that parallels the stellate outline and nodals have 5 elliptical cirral facets. Neocamptocrinus catherinensis sp. nov. is the first report of the genus from the late Artinskian Catherine Sandstone. Gissocrinus? sp. MEMOIRS OF THE QUEENSLAND MUSEUM (=Anaglyptocrinus sp. herein) was the only crinoid previously reported from the Catherine Sandstone (Willink, 1979b). These two genera are Tethyan and Australian endemics, respectively. The discovery of Anaglyptocrinus willinki in the Wandrawandian Siltstone enlarges that fauna to 12 species in 7 genera. The Wandrawandian Siltstone, with the second largest Permian crinoid fauna in E Australia, contains Calceolispongia (3 spp), Neocamptocrinus (2 spp), Notio- catillocrinus (2 spp) and Tribrachyocrinus (2 spp). The Wandrawandian fauna could be referred to as an Australian fauna with Australian endemics Notiocatillocrinus, Nowracrinus, Tribrachyocrinus and Anaglyptocrinus and Tethyan endemics Neocamptocrinus and Calceolispongia; only Dichocrinus is found throughout the equatorial belt in the Carbon- iferous and appears to be a Permian holdover in Australia. Notiocatillocrinus, Neocamptocrinus and Calceolispongia show affinity with the Callytharra Formation and suggest an Artinskian age. The Wandrawandian fauna is in situ, as many specimens retain stem and arms attached and associated brachiopods and corals are in living positions suggesting they lived below storm wave base. Shi & McLoughlin (1997) considered the Wandrawandian Siltstone to represent an off- shore environment on an unstable palaeoslope. For the first time crinoids are reported from the latest Artinskian Condamine Beds of SE Queens- land. The Condamine fauna is the most diverse known from E Australia, contains several very large specimens, and is not typical of other Permian faunas of E Australia. It shows affinity with the Basleo faunas of Timor containing Neo- camptocrinus, Platycrinites, Spaniocrinus, and Sundacrinus in common. It also shows affinity with the Wandagee Sandstone, with Eoindo- crinus praecontignatus in common, which supports a late Artinskian age for the Condamine Beds. Occurrence of Calceolispongia sp. shows affinity with E and W Australian faunas. Other identified elements in the fauna are Necopino- crinus tycherus gen. et sp. nov., the youngest known euspirocrinid, and Moapacrinus cuneatus sp. nov., perhaps the youngest known cromyo- crinid and showing affinity with Artinskian faunas of North America. Two interesting elements of the Condamine fauna are sets of arms questionably assigned to а stellarocrinid and an indeterminate poteriocrinid. Both have brachials considerably larger than in NEW PERMIAN CRINOIDS most Palacozoic crinoids. The stellarocrinid(?) arms are unusual. with the pinnules attached to the shorter end and a large protruded node on the longer end of each brachial. Large nodes on the mner side of the brachial are unknown m the crinoids and the pinnules are normally attached to the longer end of the brachial. The nodes would have served as protection for food part- icles moving along the ambulacral trackway in the wide ambulacral groove in the open feeding posture. Development of the рише on the shorter endofthe hrachial may be a result of enlargement of the longer end to accomodate the prominent node. Brachials of Poteriocrinid indet.. arm frag- iment 1, are the first pseudobisenal reported. They are also Inpinnulate and apparently represent a Ler- minal late Palacozoicevolutionary development, The Condamine Fauna is judged to be near ut siti, with several specimens retaining arms and proximal stem. Specimens probably represent storm kills transported a short distance and buried ш a silty mud. Auliskacrinus? bananaensis and а tribrachyo- crinid(?) arm fragment from the Wordian Flat Top Formation of central Е Queensland are associated with starfish and plant. fragments. These are (he first crinoids reported from the Flat Top Formation, merease the number of crinoids found ina sandstone matrix апа arc interpreted as living in a clastic environment. There are 120 species of Permian crinoids recognised in WA (Teichen. 1949. 1954: Web- ster, 1987, 1990: Webster & Jell, 1992, herein). However. only 55 (45.8%) are identified species. or referred (cf. ) to a named species. All others are referred to generic or higher taxonomic calegorics and are represented by partial cups or crowns. sets of arms. arm fragments, loose radials. and rarely columnals. In E Australia 60 species of Permian crinoids are recognised (Willink, 1978, 19798, 1979b, [980a, 1980b: Webster. 1990; herein). Of these. 49 (81.7%) are identified to. or referred (cf.) to species level, All others are identified like the WA (аха and represented by similar types of incomplete speennens, Among the E and W Australian taxa there are several that merit special comment The Dichocrinidae were common in the Earls Carboniferous and rare in Late Carboniferous of North America and Europe (Broadhead, 1981; Webster. unpublished compilation), In the Permian they are unknown ontside the Tethys. where they are moderately common, especially in Australia, Veocanploerinys 1s of stratigraphic utility in both E and W Australia although after introduction to Australia lineages may have evolved independemly in the two regions Neocampracrinmis catherinensts sp, nov. in rhe Catherine Sandstone and A. sp. in the Condamine Beds inerease the stratigraphic utility of the genus. Reports of the stems of Camplocrinus (= Veocamptocrinis) in Russia (Yakovlev. 1950) and Timor (Wanner, 1924) show the widespread distribution of the genus in thc Tethys and suggest its ппу for correlation Шетел. Calceolispongia and Jimbacrinus are wide- spread in the Permian of Australia and the former is know n (as Dinefeerinus) from Timor (Wanner. 1916, 1924, 1937) and India (Reed, 1928, 1933) Their straligraphic utilits in Australia has been i alley hy Teichen (1949), Willink ( 1979h), and ebster & Jell (1992), Caleeofispongie is considered to have evolved as 2 separate lineuges in E апа W Australa in the Early Permian (Willink, 1979b: Webster & Jell, 1992), Neocainplocrinus commonly. occurs wiih Calceolispongia. These 2 genera had simular. widelv tolerant, ecological requirements as reflected in theirability to live in either clastic or carbonate deposilional environments. Both taxa were lower level feeders. living attached to runner stems, or in some species of Ca/ceafi- spongia tethered by a dysfunctional stem in the adult stages. When in association, Calceali- spongia is typically more abundant, Veg- canplocrinus has a greater stratigraphic range. extending into the Wuchiapingian. Platverinites (Late Devonian то Late Permian) is one of the few long ranging crinoid genera of the Paleozoic, I also had widely tolerant ccologi- cal requirements as it is found in the claystones, тай. and arenaceous limestones of the Cal- Ivtharra Formation and the mudstones of the Condamine Formation. It is one of the few equatorial genera in the Permian and a higher level trophic feeder. Euspirocrinids were reported to have a range of Middle Ordovician, Mohawkian, lo Early Carboniferous, Tournaisian, by Lane & Moore (in Moore & Teichert, 1978), The discovery of Anaglyproerinuis willinki gen, et sp. nov, in the Wandrawandian Siltstone at Warden Head. NSW. and Necopinocrinmus tveherus gen. et sp. nov. 1m the Condamine Beds. extends the range of the family into the late Artinskian and possibly early Roadian. Webster el al. (1n press) recognised 37 Permian crinoid genera in Australia. Of these: 34 were 282 MEMOIRS OF THE QUEENSLAND MUSEUM based on cups and crowns and identified to genus; 2 genera were based on stems; and | was referred to as Rhenocrinidae n. gen. In this study, 11 genera are reported for the first time from Australia. These are: Auliskocrinus?, Ana- glyptocrinus, Necopinocrinus, Eidosocrinus, Archaeoisocrinus, Spaniocrinus, Glaukosocrinus, Pedinocrinus?, Sundacrinus, Moapacrinus, Loxocrinus and Prophyllocrinus. In addition, 9 indeterminate genera, that are probably new for Australia, are based on poorly preserved crowns, cups, sets of arms, arm fragments and loose cup ossicles. Together these bring the total number of Permian crinoid genera for Australia to 56, with Anaglvptocrinus replacing Gissocrinus? of Willink (1979a). Of the 46 named genera 14 (30.4%) are endemic to Australia, 13 (28.3%) are endemic to the Tethys, 7 (15.2%) are found throughout the equatorial belt, and the other 12 (26.1%) are Permian holdovers in Australia or known from Australia and one other locality outside the Tethys, but at this time, not considered to be cosmopolitan. Bambach (1990) pointed out that there are no true cosmopolitan crinoid taxa in the Permian, since no taxon is found in all four of the biogeographical regions he recognised. Genera referred to as cosmopolitan by Webster et al. (in press) are found throughout the equatorial latitudes and the cooler water higher latitude Australian localities. Eastern Australian crinoid faunas contain 8 endemic genera whereas WA faunas contain 3. In addition there are 3 Australian endemics common to both, Undoubtedly, there will be ad- ditional endemics recognised as more complete specimens of indeterminate taxa are found. These taxa represent evolution in cooler water, clastic- rich environments, not the equatorial belt carbonate rich enviroments typical of most Pal- aeozolc crinoid faunas. We suggest that the E Australian endemic genera will continue to be a greater number than those of WA. Western Aust- ralian taxa currently identified as genus, family, and order indeterminate will probably contain a good percentage of taxa described from Timor. Eastern Australian faunas have less affinity with Timor and evolved in latitudes farther S than those of WA (Webster et al., in press). AGES AND CORRELATION The Permian stratigraphy of Western Australia was correlated internationally on moderately frequent occurrences of ammonoids (Glenister et al., 1993). Few Permian ammonoids have been reported from E Australia so there the regional biostratigraphy is based on brachiopods (Dickins et al., 1964, among others). International cor- relations of some Permian units of E Australia are not clear (Day et al., 1975). Eastern and Western Australian crinoid faunas have in common Dichocrinus. Neocamptocrinus, Platycrinites, Notiocatillocrinus, Eoindocrinus, Calceolispongia, and possibly Jimbacrinus. Only E. praecontignatus is common at species level. The Wandagee Sandstone of WA is late Artin- skian (Glenister et al., 1993) and is correlated using E. praecontignatus with the Condamine Beds of SE Queensland. An Artinskian age for the Condamine Beds is supported by Moapacrinus which is Artinskian in North America (Webster & Lane, 1967). Occurrence of crinoid genera and species in E and W Australia implies that both regions were connected by seaways in the late Sakmarian to allow the incursion of Ca/ceolispongia. In the latest Sakmarian or early Artinskian Veocampto- crinus and Notiocatillocrinus invaded both areas. In the middle Artinskian Jimbacrinus and Dicho- crinus are common to both regions. Platycrinites and Eoindocrinus praecontignatus are found in both areas in the late Artinskian. This implies that E and W Australia were interconnected repeated- ly in the Early Permian, Because the lineages of Calceolispongia and Neocamptocrinus are apparently separate in E and W Australia, it would appear that there was a common source for both areas, but not an interconnection for two-way exchange between them. At the generic and specific level WA faunas correlate most closely with the Basleo faunas of Timor. Loxocrinus booni, 2 other species of Loxocrinus, and 3 species of Prophyllocrinus in the Callytharra Formation support the suggestion (Webster & Jell, 1992) that part of the faunas of the Basleo Beds are of Artinskian age. Spanio- crinus and Sundacrinus in the Condamine Beds show affinity with the Basleo Beds and support an Artinskian age. All other faunas of E Australia show little affinity with the Basleo Beds, except for the longer ranging, Tethyan endemics Calceolispongia and Neocamptocrinus. STEM ARTICULATA Discovery of Archaeoisocrinus gen.nov. in the Artinskian of Western Australia requires a review of characteristics defining the subclass Articulata. Simms (1988) defined the Articulata on a combination of morphological characters, but most significantly on the absence of the anal NEW PERMIAN CRINOIDS plate in ihe cup. He pointed out that all of the morphological characters recognised in the Articulata were individually or in various in- complete combinations known in various Palaeozoic taxa, but not in the total combination as found in the post-Palaeozoic articulates. Simms & Sevastopulo (1993) reviewed the origin of the articulate crinoids, noting that, as defined by Miller (1821), a number of late Palaeozoic crinoids could be included in the Articulata. Furthermore, applying a cladistic study of 9 primitive and derived morphologic characters (Simms & Sevastopulo, 1993, text-fig 2), they compared 3 Middle Triassic articulate genera to 11 Palaeozoic taxa which have been proposed as ancestral, or have close morpho- logical affinities, to the articulates. They also suggested a revised classification of the Palaeo- zoic crinoids that included major revisions of, as well as supression of the term, Inadunata. We agree with most of the proposed clades of Simms & Sevastopulo (1993), but do not agree, with excluding their middle to late Palaeozoic 'stem-group articulates'from the Articulata. They referred to the post-Palaeozoic Articulata as the ‘crown-group articulates’. This makes the Articulata a horizontally defined (Simpson, 1961) taxon. Simms & Sevastopulo (1993) justified the new definition of the Articulata by adding the recognition of the entoneural system enclosed within the thecal plates. Simms & Sevastopulo (1993) noted several late Palaeozoic crinoids with the entoneural system enclosed within thecal plates. However, all of the Palaeozoic taxa that they discussed had one or more anal plates within the cup, and thus could be excluded from the crown-group articulates or Articulata, following the definition of Simms (1988) in combination with the entoneural system enclosed within thecal plates. We assert that the synapomorphic feature that defines the Articulata is the development in the arms of syzygial brachial pairs in which muscular articulation alternates with cryptosyzygial ligamentary articulation as illustrated by Willink (1979b, text-fig. 16) for Meganotocrinus. We also conclude that brachial morphology described as rectilinear, weakly cuneate, moderately cuneate, strongly cuneate, cuneate biserial and rectilinear biserial may be considered an evo- lutionary lineage. However, such evolution could and did stop anywhere along this sequence within different crinoid lineages. Thus the 2 states of uniserial (= primitive) and biserial (= derived) arms, as given by Simms & Sevastopulo (1993, text-fig. 2), are insufficient for defining the complex brachial morphology. We agree that biserial arms evolved more than once in the Palaeozoic, once in the Mesozoic, and that they provided greater flexibility of the feeding arm. We also assume that biseriality became a dead- end. We suggest that the reason biseriality was a deadend may be that the interior axial canals could not function efficiently in the short zigzag relationship between adjacent brachials in the cuneate and rectilinear biserial conditions, if it ever developed in those forms. Development of the interior axial canals in the cuneate brachials provided greater protection from injury to the canals in the development of muscles at articular facets, at points of arm branching and on the facets of syzygial paired brachials. They were not restricted by short spaces between zigzag facets. Removal of the anal from the cup was an evolutionary trend that was repeated many times throughout the Palaeozoic. The loss of the anals from the cup in the stem-group articulates is considered unrelated to the development of the entoneural system being enclosed in thecal plates, as some genera (Calceolispongia, among others) developed an entoneural system enclosed in thecal plates while retaining an anal within the cup. We agree with Simms & Sevastopulo (1993) that: 1) the Ampelocrinidae and Cymbiocrinidae should be combined and revised; 2) they contain genera that are not stem-group articulates and should not be included within the family; 3) several taxa of stem-group articulates are insufficiently defined to fully evaluate their position in the lineage; 4) Articulata, as here defined, is a monophyletic clade. We propose that the primitive Articulata possessed the following morphologic features: 1, dicyclic or cryptodicyclic cup; 2, cirri with multi- radiate articula distally and transverse ridge articula proximally or cirri with transverse ridge articula throughout; 3, pinnulate arms; 4, brachial articula with ligamentary and clearly defined muscular fossae: 5, first arm division on primi- brachs 2-4; 6, entoneural system enclosed in paired canal; 7, syzygial brachial pairs in arms; 8, anals in cup, | to 3; and 9, uniserial arms, with cuneate brachials. These morphologic features are found in Chlidonocrinus, Ampelocrinus, and Nowracrinus as shown by Simms & Sevastopulo (1993, text-fig 2), but they included these taxa in their stem-group articulates. It should also be noted that each has a pentagonal stem proximally. 284 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 1. Comparison of major morphologic characters of genera assigned to the Order Ampelocrinida and isocrinid Archaeoisocrinus. Stem X-sect Family and Genus | Cup Shape | No. Anals | No. IBr. | No. Arms | Сук аї сир ч at | Anal Sac | Еасеї Туре Corythocrinidae n i Corythocrinus conical 1 3 20-30 pinnulate round no ? _plenary Araeocrinus conical 3 45 20+ pinnulate round 2 _ long plenary Campbellicrinus conical 1 3 12 pinnulate | — round _ ? short plenary Ampelocrinidae E Е g i E р Ampelocrinus | med bowl 3 2 10 _ pinnulate | pentag | yes recurved plenary Chlidonocrinus med bow 1 2 20 min pinnulate pentag yes ? plenary | Cymbiocrinus | low bowl 1 m | 10 | pinnulate | pentag or rd yes ? | plenary Halogetocrinus low bow 1 3-4 10+ pinnulate round yes ? plenary Moundocrinus med bowl 1 2 10 pinnulate subpentag ? short plenary Oklahomacrinus discoid 1! 2 10 pinnulate subpentag ? ? plenary Calceolispongiidae Е Allosocrinus med bow 1 - b pinnulate | subpen to rd ? ? plenary Calceolispongia high bowl 1 _- 5 pinnulate | subpen to rd yes ? plenary Jimbacrinus high bow 1 - 5 pinnulate | subpen to rd no ? plenary Metacalceolispongia | med bowl 1 2 11+? pinnulate pentag ? ? plenary Tribrachyocrinidae Tribrachyocrinus . globe 3-4 | 2 12 min ramules round no short plenary Meganotocrinus globe 1 2 20 ramules round no short plenary Nowracrinus globe |. 1 2 20 ramules round yes short plenary Insertae sedis | | i Tasmanocrinus conical O or 1? 2 6min pinnulate pentag yes short | peneplenary | Isocrinidae " Archaeoisocrinus discoid 0 2 10 pinnulate pentag ? ? | plenary We consider Corythocrinus, from the late Tournaisian of Indiana (Tables 1, 2). the oldest Articulata. This is followed, in order of earliest occurrence, by Chlidonocrinus, Cymbiocrinus and Ampelocrinus, from the Visean of North America. We consider the report of Ampelocrinus from the Visean of England (Wright. 1951) a questionable identification. Offshoots of the Ampelocrinidae include the Calceolispongiidae, a Late Carboniferous— Permian lineage, and Tribrachyocrinidae, a Permian lineage. Thus, our higher level classification is: Subclass Articulata Order Ampelocrinida ord. nov. Order Millericrinida Order Cyrtocrinida Order Bourgueticrinida Order Isocrinida- with Archaeoisocrinus gen. nov. Order Comatulida Order Uintacrinida Order Roveacrinida Origin of the Ampelocrinida is uncertain. Strimple & Watkins (1969) suggested that Corvthocrinus was derived from a rhenocrinid because the plicate plates of the anal tube indicated affinities between these two forms: however, plicate tube plates are also known in poteriocrinitids and are common among dendrocrinids. Moore & Teichert (1978) considered the Ampelocrinidae derived from the Decadocrinidae, but gave no explicit reasons for supporting this relationship. Because so many stem articulates have | anal, we suggest that the Ampelocrinida might be derived from a cyathocrininid, such as Lecythocrinus, where the radianal had already been eliminated from the cup, or Corynecrinus, where the radianal and anal X are above the posterior basal. However, the 3 anals in Ampelocrinus and Araeocrinus suggest that the presence of a single anal may not be a primitive character of the group. Carboniferous evolution of the Ampelocrinida occurred in North America and Europe, whereas the Permian record is within the Tethys, especially E Australia, except for North American species of /7alogetocrinus and Alloso- crinus (Table 2). Moore & Jeffords (1968) described several taxa with pentagonal and pentastellate columnals from Devonian and Carboniferous strata of the United States. The cups are either unknown or not recognised in association with the column- als. The geographic distribution of such NEW PERMIAN CRINOIDS 28 Un TABLE2. Rangechart of genera assigned to the Ampelocrinida. (x) indicatesage of type species. (—) indicates age of species assigned to the genus. United States series names used for Carboniferous because known record is restricted to North America. Carboniferous (part) Family & Genus Wuch. Corythocrinidae Corythocrinus x Araeocrinus x Campbellicrinus Ampelocrinidae Ampelocrinus x Chlidonocrinus — x Cymbiocrinus Halogetocrinus - Moundocrinus Oklahomacrinus - Calceolispongiidae Allosocrinus Calceolispongia Jimbacrinus Metacalceolispongia Tribrachyocrinidae Tribrachyocrinus Meganotocrinus Nowracrinus Incertae sedis Tasmanocrinus columnals is unknown. We suggest that such forms should be investigated as possible stem articulates. SYSTEMATIC PALAEONTOLOGY Crinoid teminology follows Ubaghs et al. (in Moore & Teichert, 1978), with columnal patterns after Webster (1974). Measurements are given as: length, parallel to the central axis; width, transverse to, but never cutting or joining the central axis; and depth, normal to, and may join the central axis. Curvature of the cup walls, plate circlets within the cup and fixed brachials are referred as: incurved if distally bending toward. vertical if parallel to, weakly to strongly flaring if bending away from and horizontal if perpen- dicular to the central axis. Material collected by us came from localities entered in the Queenland Museum Locality Register (QML), and is curated in the Queens- land Museum Palacontological Collection (QMF). Other palaeontological collections referred to are indicated by the following prefixes: Geological Survey of Queensland. Brisbane (GSQ): Geological Survey of Western Australia, Perth (GSWA); Geological Survey of New South Wales, Lidcombe (MM): Department of Geology. University of Queensland, (ПО); The Natural History Museum London (BME); and Tasmanian Museum (TM). Osag. Mrmc. Chst. Morw. Atok. Dsmn.Mssr. Vrgl. | Permian (part) Assl. Skmr. Artk. Road. Word. Capt. Subclass CAMERATA Wachsmuth & Springer, 1885 Order MONOBATHRIDA Moore & Laudon. 1943 Superfamily HEXACRINITOIDEA Wachsmuth & Springer, 1885 Family DICHOCRINIDAE Miller, 1889 Subfamily DICHOCRININAE Miller, 1889 Dichocrinus Münster, 1839 TYPE SPECIES. Dichocrinus radiatus Münster, 1839 from the Early Carboniferous of Belgium; by monotypy. Dichocrinus? sp. (Fig.1B) MATERIAL. GSWAF50172, from GSWAL119377, Billidee Formation, Artinskian. DESCRIPTION. Crown small, 29.4mm long (incomplete), 25.8mm wide (arms flared). Cup elongate, cylindrical, unornamented. Basal circlet unknown. D radial 7.7mm long. 3.5mm wide (incomplete), gently convex longitudinally and transversely, with narrow shoulders sloping abmedial. Radial facet angustary, rounded aborally. Anal large, 7.4mm long, 4.6mm wide, widest at base. tapering distally. in line with radials. Arms 2 per ray. isotomous branching on 2nd primibrach, biserial above secundibrach 4-7. Brachials cuneate, moderately convex longitudinally, strongly convex transversely. One 286 slender pinnule per brachial on long side. Stem and tegmen unknown. REMARKS. This is the first unornamented Dichocrinus reported from the Permian of Australia. Several species of Dichocrinus from the Early Carboniferous of the United States have a distally tapering anal plate and most have 10 arms (Broadhead, 1981). The 10 arms are a prim- itive condition in the genus. Lacking the tegmen, the generic assignment is questioned. The Billi- dee Formation specimen probably represents a new species, but lacking the basal circlet and tegmen, it is left in open nomenclature. Auliskocrinus Broadhead, 1981 TYPE SPECIES. Dichocrinus crassitestus White, 1862 from the late Tournaisian upper part of the Burlington Limestone, lowa; by original designation. Auliskocrinus? bananaensis sp. nov. (Fig.l A) ETYMOLOGY. From Banana in central Queensland. MATERIAL. HOLOTYPE: QMF38897 from QML806. DIAGNOSIS. Anal tube large, conical, distally tapering above the posterior interradius; cup truncated cone-shaped; basal circlet very short; brachials rectilinear. DESCRIPTION. Specimen small, 26.2mm long. Crown small, cylindrical, 16.1mm long, 8.2mm wide at tip ofanal tube. Calyx robust, 12mm long to tip of anal tube. Cup truncated conical, 7.4mm long, 6.9mm wide at radial summit; plates smooth; sutures flush. Basal circlet shallow distally flared bowl, 2mm long, 4.8mm wide. Radials large, 5.9mm long, 5mm wide at base of radial notch, longitudinally moderately convex proximally becoming gently convex distally, moderately convex transversely, forming most of cup wall, subvertical distally. Radial facet angustary, 3mm wide, moderately convex pro- jecting from radial aborally. Single anal large, 6mm long, in radial circlet. Tegmen formed of numerous small plates. Anal tube conical, formed of small irregular polygonal plates, tapering distally, distal opening above posterior inter- radius. Arms relatively short, 11mm long, slender, 4 per ray, isotomously branching on axillary 2nd primibrach. Brachials rectilinear, MEMOIRS OF THE QUEENSLAND MUSEUM wider than long, slightly convex longitudinally. strongly convex transversely, with single pinnule on opposite sides of arm. Primibrach 1 much wider (2.6mm) than long (0.7mm). Axillary primibrach 2 3mm wide, 0.8mm long. Stem round, 1.2mm diameter at cup, 10.1 mm preserv- ed, heteromorphic. Noditaxis N1; nodals longer and wider than internodals; latus strongly rounded. REMARKS. Auliskocrinus? bananaensis is preserved as an external mould with the C ray centred. Arguments could be made for erecting a new genus for 4.? bananaensis or assigning it to Dichocrinus. The cup shape. position and plate structure of the anal tube, and rectilinear brachials of 4.? bananaensis are atypical of the closely related Auliskocrinus and Dichocrinus. Cup shape of Auliskocrinus is relatively high, subconical or slightly globose whereas Dic/io- crinus is relatively high conical. In both genera the basal circlet forms a significant part ofthe cup wall. Only D. dichotomus, an early Visean species with biserial arms, has a bowl-shaped cup with a low upflaring basal circlet, similar to 4.? bananaensis. No member of the Dichocrininae (Broadhead, 1981, fig. 2) has a conical, distally tapering anal tube projecting above the tegmen over the posterior interradius as in 4.? Рапапа- ensis. The small more centrally located vertical anal tube of Auliskocrinus is formed of laterally interlocking columns of hexagonal rather than irregular plates, and the genus has slightly cuneate brachials. The tegmen of Dichocrinus is typically low, but may be moderately elevated with the anal opening flush or only slightly projected above the tegmen (Broadhead, 1981). Brachials of Dichocrinus are either cuneate or biserial, most commonly rectilinear proximally becoming moderately to strongly cuneate distally. Most advanced species have biserial brachials with 20 arms (Broadhead, 1981). With the exception of the truncated conical cup, 20 arms, and projected anal tube, 4.? bananaensis retains primitive features of Dichocrinus. Variation in cup shape and tegmen structure of monotypic Auliskocrinus is unknown (Broadhead, 1981). We assign this specimen to Auliskocrinus because the tegmen forms a high cone with a terminal anal opening and the FIG. 1. A, Auliskacrinus? bananaensis sp. nov., C ray view of crown with tegmen, holotype QMF38897, «4.3. B, Dichocrinus? sp., lateral view of partial crown GSWAFS50172, x2.5. C,D, Neocamptocrinus catherinensis sp. nov. C, D-E interray view of partial crown, paratype GSQF 13487, 2.5. D, С ray view of crown, holotype GSOF 13486, «2.5, 287 NEW PERMIAN CRINOIDS rectilinear brachials are more similar to the slightly cuneate brachials of Auliskocrinus. Subfamily CAMPTOCRININAE Broadhead, 1981 Neocamptocrinus Willink, 1980 TYPE SPECIES. Neocamptocrinus bundanoonensis Willink, 1980a from the Wordian Berry Formation, NSW; by original designation. REMARKS. Willink (1980a) defined Neo- camptocrinus primarily on the distinctive inflated tegmen formed of 5 large orals, and numerous small interambulacral and anal plates. He noted Neocamptocrinus as acommon element in E Australia, recognised 3 species on the basis of cups, another 7 species on columnals, and considered the genus of potential stratigraphic value. In WA there are 3 species (Webster, 1990; Webster & Jell,1992), a pluricolumnal from the Callytharra Formation (Webster, 1987) and very large pluricolumnals in the Wandagee Sandstone (Webster & Jell, 1992).Willink (1980a) considered the coiled elliptical stem typical of Neocamptocrinus. Broadhead (1981) noted that the elliptical stem distinguishes the Campto- crininae, whereas Dichocrininae have a round stem. Webster (1987) reported stems of Campto- crinus cf. C. indoaustralicus (considered Neocamptocrinus by Webster & Jell, 1992) to vary from slightly elliptical proximally to elliptical to subrectangular distally. Webster & Jell (1992) noted that the proximal stem of М. millyitensis is nearly circular in section, becoming elliptical distally. New material of N. millyitensis shows the curvature of the enrolled proximal nearly circular part and the transition from that into the strongly elliptical part (Figs 2A-C; ЗА,В; 4C,D). The range of Neocamptocrinus in E Australia is from the Sakmarian, Billop Formation of Tasmania, into the Wordian, Condamine Beds of Queensland. In stratigraphic sequence, species recognised are: N. sp. nov., Condamine Beds N. bundanooensis Willink, 1980a, Berry Formation N. catherinensis sp. nov. Catherine Sandstone N. wardenensis Willink, 1980a, Wandrawandian Siltstone ӨМ. tasmaniensis (Sieverts-Doreck, 1942), Crin- oidal Zone N. millerensis Willink, 1980a, Billop Formation MEMOIRS OF THE QUEENSLAND MUSEUM The following columnal (о) species are con- sidered junior synonyns of øN. tasmaniensis: ӨМ? sievertsae Willink, 1980a, oN.? doreckae Willink, 1980a, oN. bernacchiensis Willink, 1980a, oN.? banksi Willink, 1980a, and oN.? sp. cf. N.? tasmaniensis. These are all from the Crinoidal Zone on Maria Island and represent different parts of the stem of one species. In general, species of Neocamptocrinus are distinguished on cup plate ornamentation, cup plate shape, ornament of tegmen plates, and number of arms. The oldest form known, N. millerensis, has a slender high cup with a vermiform ornament, whereas N. wardenensis has a lower, more bulbous cup and coarse node and irregular ridge ornament. The plates of №. catherinensis sp. nov. are smooth, but the tegmen is a prominent conical projection above the posterior interray. Ornament on N. bundanoonensis consists ofpits on the cup plates. Cup plates of N.? sp. indet. (Willink, 19802, pl. 4, figs 17-26) probably belong with oW. tas- maniensis from the same stratigraphic unit (Crinoidal Zone). This form would have had a lower cup with a longitudinal trending vermi- form ornament. In WA Neocamptocrinus ranges from the late Sakmarian, basal Callytharra Formation, into the Wuchiapingian, Cherrabun Member of the Hardman Formation: N. millyitensis Webster & Jell, 1992, Cherrabun Member oN. sp. Webster & Jell, 1992, Wandagee Sandstone N. occidentalis Webster, 1990, Cundlego Sandstone N. barrabiddyensis Webster & Jell, 1992, Bul- gadoo Shale oN. sp. Webster, 1987 (as Camptocrinus cf. C. indoaustralicus), Callytharra Formation N. barrabiddyensis lacks ornamentation, N. occidentalis has fine granular ornament on cup plates, and №. millyitensis is smooth but has nodes on some oral plates. All have more globose cup shapes than E Australian species. With 8 arms per ray N. millyitensis has the greatest number known forthe genus. Most species have 4-7 arms per ray, but some rays of N. millerensis may have 2. The general trends in evolution of Neocamptocrinus in Australia were to: 1, lower the cup by flat- tening the basal circlet and shortening the length of the radials and primanal; 2, increase the number of arms per ray; and 3, increase in size. Cup and tegmen are smooth or have ornament of NEW PERMIAN CRINOIDS 289 FIG. 2. Neocamptocrinus millvitensis Webster & Jell, 1992. A-C. distal facet, lateral stem and proximal facet views of proximal pluricolumnal showing enlargement below theca, QMF38031. х5, D-G. oral, A ray, posterior and basal views of theca QME 37981. «2.6. 290 variable type. Neocamptocrinus has the longest stratigraphic range of any Australian Permian crinoid and may have the greatest geographic distribution (columnals referred to as Campto- crinus in Timor and Russia are herein considered Neocamptocrinus), Only Calceolispongia has comparable stratigraphic and palaeogeographic ranges among Australian Permian crinoids. Neocamptocrinus catherinensis sp. nov. (Fig. 1C,D; Table 3) ETYMOLOGY. From the Catherine Sandstone. MATERIAL. HOLOTYPE: GSQF 13486 trom the Guad- alupian, Catherine Sandstone, in the upper part of Sandy Creek, Springwood Homestead, Queensland. PARATYPE: GSQF'13487, same. DIAGNOSIS. Cup small to medium sized, high bowl-shaped: tegmen conical projected toward posterior interray; 4 arms per ray. DESCRIPTION. Crown small to medium sized, cylindrical. Cup high bowl-shaped, plates un- ornamented. Basal circlet upflared, 2 equal plates. suture in A-CD plane of symmetry, forming basal one third of cup. Radials 5, hept- agonal, gently convex longitudinally and transversely, gently flaring proximally, sub- vertical distally; distal facets with tegmen plates sloping downward gently. Radial facet angustary, nearly 1/2 radial width, gently rounded below distal tips of radial. Anal plate in radial circlet, gently convex longitudinally and transversely. Arms slender, elongate. 4 per ray, branching isotomously on axillary 151 primibrach and 2nd secundibrach. Brachials rectilinear, uniserial proximally, cuneate, becoming biserial distally. Tegmen with rounded conical projection on posterior. Stem nearly circular proximally becoming strongly elliptical distally, hetero- morphic in strongly elliptical part; noditaxis №. Nodals formed by fused columnals, with incipient cirral scars on outer side. REMARKS. The holotype is preserved in the enrolled position as original calcite embedded in fine grained sandstone. The cup is crushed with 2 rays and the anal or edge of a 3rd radial exposed. Proximal parts of the arms show the branching pattern and the distal part of the stem is cirrate. Iron oxide replacement of the plates of the paratype is very soft and partly lost on the enrolled proximal part of the stem and partial MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 3. Neocamptocrinus catherinensis sp. nov. measurements (mm). *crushed, estimated. holotype paratype | GSQFi3486 | GSQFI3487. Crown length, incomplete 265 | E = Calyx length { 8.5 Cup length | 99 65 Cup width 10.1* 5:0. Basal circlet diameter |. 48 3.1 Basal circlet length | 3.2 1.2 Radial length 5.4 4.2 Radial width 44+ 24 First primibrachial length _ | 1 | First primibrachial width 2.1 | Proximal columnal diameter 2.1 11 Stem length 65 63.7 crown. The uncrushed cup is smaller than the holotype, the D and E rays are centred, the short anal tube projects on the right and the distal part of the preserved stem is cirrate. The tegmen of other species of Neocampto- crinus is inflated and may be slightly elevated towards the posterior side, but is not elevated into a conical projection as sharp or prominent as that of N. catherinensis, Only N. wardenensis with a rounded posteriorly elevated tegmen is compar- able. Also the cup of N. wardenensis is a lower bowl shape and the basal circlet is shorter and more outflared. The cup of М. catherinensis is most similar to that of №. millerensis, which has a very low tegmen and relatively longer radials. Neocamptocrinus millyitensis Webster & Jell, 1992 (Figs 2-4) Neocamptocrinus sp. nov. Webster,1990: 57, pl. 1, figs 7-11. Neocamptocrinus millyitensis Webster & Jell, 1992: 320, figs 3A-L. MATERIAL. Crowns QMF37980-F37985, partial calyces (QMF37920, F38986-F38024), partial sets of arms (QMF38025, F38026), radials (QMF37921-F37928), columnals and pluricolumnals (QMF37929-F37971, F38027-F38864), and cirri (QMF37972-F37979. F38865-F38873) from QML772 and 1146). DESCRIPTION. This description only adds to that of Webster & Jell (1992). Radial facet an- gustary, approximately half maximum width of radial, sloping outward gently. Brachials cuneate, strongly convex transversely, straight to FIG. 3. Neocamptocrinus millyitensis Webster & Jell, 1992. A,B, lateral and proximal facet views of proximal pluricolumnal and part of basal circlet, QMF'38027, «3.8. C, lateral view of distal end of radial and proximal brachials showing branching pattern, QMF38029, х4. D-F. A ray, posterior and basal views of theca QMF37980, x2.6. NEW PERMIAN CRINOIDS 291 292 slightly convex longitudinally, deep, uniserial proximally, biserial distally. Arms slender, delicate; endotomous branching heterotomous on single primibrach and secundibrach and isotomous on single tertibrach; arms 40, 8 per ray. Brachials with single slender pinnule (up to 9mm long) on long side. Anal series 3-2, with the 2 plates to the left of the anal opening. Stem elliptical in transverse section; short, more equidimensional (4.6 х 2.4mm) adjacent to cup; becoming longer, flattened and extended elliptical (10.1 х 3.4mm) within 7-8mm, 2-3cm from the cup; becoming less extended elliptical (9.4 х 6.8mm) an unknown distance distally. Noditaxis pattern heteromorphic, with nodals separated by 2 internodals normally, but varying from 1-3. Cirri attach at ends of ellipse, 2 per nodal; cirral facet extending laterally onto 2 adjacent columnals with growth. Cirri not developed in proximal more equidimensional part of stem. REMARKS. The revised description is based upon new material listed above. Five of the calyces were in situ in a nest with the broken stem segments in the surrounding matrix. Many of the pluricolumnals are coiled, indicating specimens were enrolled prior to fracturing and disag- gregation from compaction and weathering. The calyces and pluricolumnals were encased in a clay to silt and fine sand matrix, rather than the typical fine to medium sand of the Cherrabun Member. This suggests that when enrolled, the cirri formed a protective screen around the crown shielding it from the coarser grained sediments. As burial proceeded, finer grained sediments infiltrated the cirri entombing the crowns. Compaction after burial distorted and fractured some of the calyces and broke the stem into pluricolumnals. Modern weathering left a lag gravel of columnals, pluricolumnals, partial and complete calyces, and arm fragments over 3 m? on a very gently sloping surface. Five complete and 7 partial calyces and numerous stem segments were recovered in situ by excavation to 20cm beneath the lag gravel in the weathered zone. The arms are delicate, quite slender and, based on a partial set of arms lacking all parts of the calyx, extended a minimum of 25cm above the tegmen. Although uniserial proximally, they become biserial in the middle and distal parts of the arm. MEMOIRS OF THE QUEENSLAND MUSEUM Neocamptocrinus sp. nov. (Fig. 5A-C) MATERIAL. QMF38900, part of exterior side of 2 rays of partial set of arms, QMF39006, part of interior side of 3 rays of partial set of arms, and QMF39007, pluricolumnal, from QMLS18. DESCRIPTION. Axillary primibrach triangular, lateral ends nearly overlap Ist primibrach. Second secundibrach axillary. Brachials strongly convex transversely, straight longitudinally, very deep, cuneate, becoming biserial on the 8th tertibrach, rectilinear biserial on 11th tertibrach. Fine granular ornament on primibrachs and secundibrachs, smooth thereafter. Ambulacral groove narrow, deep V-shaped. Arms 20, 4 per ray, slender, very elongate, 65.8mm (incomplete). Pinnules slender, narrow, one per brachial. REMARKS. Preservation of both specimens is moderately good, with some parts poorly preserved through oxidation by weathering. The arm branching is typical of Dichocrinus or Neo- camptocrinus with 4 arms per ray. Arms are very delicate and larger than most dichocrinids, They are assigned to Neocamptocrinus based on the shape of the brachials, uniserial to biserial arm development, arm branching pattern, and pluri- columnals and columnals of Neocamptocrinus in the same interval of the Condamine Beds. Neocamptocrinus? sp. (Fig. 5D) MATERIAL, QMF38880 from QML1237. REMARKS. The partial set of arms is 20.1mm long, 9.4mm wide, and consists of parts of 14 arms. They are assigned to Neocamptocrinus because they closely resemble the arms of N. millyitensis, as the cuneate brachials are small, biserial, strongly rounded transversely, and bear small delicate pinnules. Webster (1987) reported pluricolumnals of Camptocrinus cf. indoaustralicus from the type section of the Callythara Formation. Although no cup or calyx has been recovered from the Callytharra Form- ation, these columnals are now considered to belong to Neocamptocrinus, because they are similar to those reported from several strati- graphic units in WA. NEW PERMIAN CRINOIDS 293 i | : | і li e ai a mm FIG 4. Neocamptocrinusmillyitensis Webster & Jell, 1992. AB, lateral and inner views of coiled pluricolumnal QMF38028. х2.5. C.D, proximal facet and outer views of expanding part of proximal pluricolumnal QMF38030. x6.9. Г.Г. lateral views of partial set of arms QMF 38025, «3.3. 294 MEMOIRS OF THE QUEENSLAND MUSEUM 44 FIG. 5. A-C, Neocamptocrinus sp. nov. A, view of interior side of 3 rays of partial set of arms QMF39006, х1.6. В. lateral view of exterior side of 2 rays of partial set of arms QMF 38900. * 1.6. C, lateral view of pluricolumnal QMFE39007, х2, D, Neocamptocrinus? sp.. lateral view of partial set of arms QMF38880. «5.1. Alte" ИЙИП Тү 117271 \ЙДШҮ UU LL d TOU VIT ae ave "FT ^ L MAT ш itin Ил 4 ЛАЛ, ПАЛАТИ ny | Чу NEW PERMIAN CRINOIDS 295 Superfamily PLATYCRINITOIDEA Austin & Austin, 1842 Family PLATYCRINITIDAE Austin & Austin, 1842 Platycrinites Miller, 1821 TYPE SPECIES. Platycrinites laevis Miller, 1821 from the early Carboniferous of England, by subsequent desig- nation of Meek & Worthen, 1865. Platycrinites halos sp. nov. (Fig. 6C,D) ETYMOLOGY. Greek halos, a circle around the sun: refers to the elevated platform around the radial facet. MATERIAL. Holotype, internal mould with part of proximal tegmen and external mould of basal circlet, 2 radials, and 1 interambulacral, QMF39008 from QMLS18. DIAGNOSIS. Cup very large, bowl-shaped; radial facets concave, elliptical, subvertical; arms pro- jecting horizontally away from cup; radial facets large, elliptical, on slightly projecting platforms. DESCRIPTION. Cup large bowl-shaped, c. 25mm long, estimated 51mm wide, granular texture. Basal circlet large, 14,5mm long, 25mm wide. Basals 3, azygous half size 2 zygous, down widely flaring proximally beneath stem facet, upward widely flaring distally, forming proximal part of cup wall. Radials 5, large, 27mm long, estimated 27mm wide, moderately convex longi- tudinally, strongly convex transversely, distally shoulders incurved partly around angustary radial facet. Radial facet large, estimated 12.5mm long and wide, concave, elliptical outline, surrounded by narrow platform with sloping rim. Tegmen arched, unknown length. First interambulacral plates very large, estimated 12mm long, 15.7mm wide, laterally flanked by series of small plates covering ambulacral trackways. Stem facet large, estimated 14 x 10mm, separated from cup wall by narrow groove. REMARKS. This is one of the very large calyx type Platycrinites. The arms proximally project horizontally away from the cup. The basal circlet forms a small part of the cup walls as it flares outward much more than upward. The radials are the main part of the cup wall, subvertical proximally and incurved distally. A disarticulated associated columnal beside the base of the cup is 3.2mm long and 18.6mm by an estimated 8mm in transverse section; latus moderately concave; fulcral ridge elevated well above adjacent pits. It is one of the straight columnals of the segmented twist type of Webster (1997) and probably from the same specimen as the cup. The size of the radials is close to that of Platycrinites sp. of Webster & Lane (1967) from the Artinskian part of the Bird Spring Formation of southern Nevada. However, no other species of Platy- crinites has the elevated or rimmed radial facets like P. halos. Platycrinitid indet. (columnals) (Fig. 6A,B) MATERIAL. QMF38899, 39009, 39010 from QMLS518. REMARKS. Elliptical columnals belonging to a platycrinitid, such as Platycrinites, Neoplaty- crinus, or Stomiocrinus are of the segmented twist type (Webster, 1997). The facets bear a dual transverse ridge divided by a shallow groove along the long axis. They have an axial canal and 2 or 3 coarse crenulae and culmina on the distal ends of the long axis. Straight and twist columnals are present. They are mentioned to show crinoid diversity in the Condamine Beds. Subclass CLADIDA Moore & Laudon, 1943 Superfamily CYATHOCRINITOIDEA Bassler, 1938 Family EUSPIROCRINIDAE Bather, 1890 Anaglyptocrinus gen. nov. TYPE SPECIES. Anaglyptocrinus willinki, late Artinskian Wandrawandian Siltstone at Warden Head, NSW. ETYMOLOGY. Greek anaglyptos, wrought in relief, and krinon, lily; refers to the low relief, weathered out condition of the holotype. DIAGNOSIS. Cup medium bowl, with nodose ornament, with shallow apical impressions; infrabasal circlet flat to gently upflared; radial facet angustary, wide radial notches; single large anal above posterior basal; brachials rectilinear, strongly convex transversely, with 4 rows of cover plates above V-shaped ambulacral groove; arms branching isotomously on 4th primibrach and once or twice more; brachials with very small internal axial canal, brachial facets trifacial; anal tube narrow, elongate; stem round, with round lumen. REMARKS. Anaglyptocrinus is distinguished from all other euspirocrinids by the flat to very low basal circlet, medium bowl-shaped cup, and the single anal plate. Cup shape is most similar to but shorter than Euspirocrinus. Other taxa assigned to Anaglyptocrinus are Gissocrinus? voiseyi Willink, 1979 and Gissocrinus? sp. Willink, 1979, 296 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 6. A,B, Platycrinitidindet. columnals. A, facetal view of twist columnal QMF 38899, x3.6. B, facetal view of weathered straight columnal QMI'39009, 3.8. C.D. Platverinites halos sp. nov., external («1.5)and internal (x1.3) views of slightly disarticulated and distorted partial theca, holotype QMF39008. Anaglyptocrinus willinki sp. nov. (Fig.7) ETYMOLOGY. For R. Willink in recognition of his studies of the Permian crinoids of eastern Australia, MATERIAL. HOLOTYPE: QMF38913 from QML859. DIAGNOSIS. As for genus, cup ornament nodose. DESCRIPTION. Crown slender, elongate. 20.8mm preserved. Cup medium bowl-shaped. nodose ornament on all cup plates, double row of nodes on proximal edge of radial parallel to basal-radial sutures, shallow impressions at junction of basals and radials. Infrabasal circlet flat to widely flaring, barely visible in lateral view. Basals 5, large. 4mm long (incomplete), 5.6mm wide, moderately convex longitudinally and trans- versely, incurved proximally, subvertical distally. forming lower half of cup wall. Radials 5. 4.4mm long, 5.7mm wide, straight longitudinally below facet, convex longitudinally adjacent to facet. moderately convex transversely. Radial facet narrow, strongly convex outer edge, flaring in- wardly to merge with radial shoulders. inneredge smooth with wide concave ambulacral notch. declivate: transverse ridge prominent, divided by gap in middle, dividing facet into inner and outer halves: outer half with small 4-lobed elevation off centre, slightly aboral to gap in transverse ridge: outer fossa divided by low rise from 4-lobed elevation into 2 shallow transversely elongate parts, deepest central aboral; inner fossa transversely elongate, shallow. Radial notches wide. Single anal large. pentagonal, projecting slightly above radial summit, proximally abut- ting terminated end of CD basal, distally adjoining 2 proximal tube plates. Arms slender, branching isotomously on 4th primibrachand 4th or 5th secundibrach: more distal branching NEW PERMIAN CRINOIDS 297 VIG. 7. Auaglvptocrinus. willinki gen. et sp. nov. А. camera lucida sketch of posterior (х6). B- A гау (42.8) Mg view, OMF 38913, t= tegmen plates. a = anal X. r 7 C f гау radial and b= CD Helt) and BC (right) basals. unknown. Brachials straight longitudinally. strongly convex. transversely. deep. with wide V-shaped ambulacral groove. Brachial facets irifacial: transverse ridge wide V. apex pointed adoral. with small single axial canal in slightly elevated centre. Ambulacral cover plates small, 0,05mm long and wide. poly ропа], interlocking with adjacent plates longitudinally and laterally: 4 across ambulacral groove. merging with small polygonal plates of tegmen proximally. Anal tube slender. elongate, of 10-11 vertical rows of smooth. laterally interlocking hexagonal plates. 2mm wide, 1.8mm long. Column heteromorphic: noditaxis pattern N3231323. Columnals. round transversely; latus moderately to strongly convex on nodals and internodals: nodals cirrate throughout 21mm of preserved column. Axial canal round. REMARKS. The crown 15 crushed with distal paris of the arms and tegmen lost to weathering and the infrabasals and anals are not exposed, Excavation of the under side exposed the C radial. BC and CD basals. parts of the infrabasal circlet, primanal and proximal parts of 2 tube plates (Fig.7A). Ornamentation of the cup was Jost or very faintly preserved on parts of the exposed cup. but well-developed on the ex- cavated cup plates. Willink (1979a) described A. voisevi andl, sp. from the Cataract River Formation and Catherine Sandstone, respectively. These forms show similarity of the cup shape and arm branching pattern to the Wandrawandian specimen. They differ from the nodose ornamented cup of 21. willinki by A. voisevi being ornamented with nodes and sharp ridges on all cup plates as well as the brachials and l. sp. bearing prominent inter- connected plate ridges on the cup and proximal and distal expanded rims on the brachials giving them an hourglass shape. Thus. the 3 forms make a series from simple nodose ornament to highly ornate, with the simple nodose form the oldest (late Arunskian) and the 2 younger more orna- mented forms of approximately the same age (Roadian). Ray ridge and interconnected ray ornament inthe crinoidsis common in the actino- criniuds. primitive poteriocrinitoids, and а few flexibles. It is not as соттоп in the late Palaco- zoic as in the carly and middle Palaeozoic. Thus. the ray ridge ornament of the younger forms is considered heterochronous homeomorphy. Willink (19793) considered Gssocrinus in need of revision. noting that the diagnosis provided by Moore & Teichert (1978) was narrower than that of Angelin (1878) or Bather (1893). Furthermore, he suggesied that the Australian forms probably represented the end members of a conservative stock of the cyathocrinitids that was most closely allied to the Silurian Gissocrinus. However, he also suggest- ed the possibility, that the Australian specimens represented heterochronous homeomorphy. The first branching of the arms of Gissocrinus is on the single primibrach and the cup is a low bowl, both advanced evolutionary features for this Silurian taxon. By comparison, the type species of Cyathocrinites, an Early Carboniferous taxon, has a medium bowl cup and the first branching of the arms is on the 3rd primibrach, both more primitive features. A. willinki has even more prim- itive arm branching than Cyathocrinites, but a slightly more advanced cup form. The primitive arm branching with ambulacral cover plates sug- gests evolution from an unknown conservative stock of the cyathocrininids. The trifacial artic- ulation facets of the brachials and internal dual axial canals are advanced features found in some of the stem articulate crinoids (Simms & Sev- astopulo, 1993). Necopinocrinus gen. nov. TYPE SPECIES. Necopinocrinus tycherus sp. nov. from the Condamine Beds, Elbow Valley area, near Warwick, SE Queensland. ETYMOLOGY. Latin necopinus, unexpected, and crinon, lily; refers to a euspirocrinid not being expected to occur in the Permian. DIAGNOSIS. Cup expanded low bowl, with con- stricted base, with incurved radial, with coarse nodose ornament on all cup plates; 3 infrabasals, small infrabasal in C ray; radial facets angustary, 1/3 radial width, horseshoe-shaped; 3 small anals above posterior basal; single primibrach axillary. Arms widely spread, branching isotomously. Brachials cuneate; stem round transversely. REMARKS. The expanded low bowl-shaped cup and axillary 151 primibrach are the 2 most dis- tinctive charaters of Necopinocrinus. The cup is most similar to, but more bowl-shaped, than the low cone-shaped cup of Vasocrinus. Necopinocrinus tycherus sp. nov. (Fig. 8) ETYMOLOGY. Greek tyche, luck or chance, and refers to the Lucky Valley Creek wherein the specimen was found. MATERIAL. HOLOTYPE: QMF38901, from QML518. DIAGNOSIS. As for genus. DESCRIPTION. Cup expanded, low bowl, with constricted flat base, 15mm long, 39mm wide (crushed, compacted in part); all cup plates with MEMOIRS OF THE QUEENSLAND MUSEUM coarse nodes, some grading into very short ridges. Infrabasal circlet large, with large circular stem facet, constricted subvertical above stem facet, expanding upflaring distally, divided into 3 plates, 2 large equal plates and | smaller plate in C ray, visible in lateral view. Basals large, upflaring, gently convex longitudinally and transversely, forming major part of cup; D-E basal 12mm long (estimated), 20mm wide. Posterior basal 14.8mm long, 10.4mm wide, truncated distally by 3 small facets for anals. Radials large, 12mm long, 18.4mm wide, strongly convex longitudinally, moderately convex transversely, strongly incurved distally to near subhorizontal. Radial facet angustary, horseshoe- shaped, 7.4mm wide, deep, subhorizontal. Three anals small, in line of radials, probably projected slightly above radial facet. Anal tube not pre- served. Single primibrach axillary, 5.2mm long, 5.2mm wide, straight longitudinally, strongly convex transversely; distal facets wide spread, separated by narrow concave trough. Secundi- brachs wider than long, weakly cuneate, straight longitudinally, strongly convex transversely. Primibrachs and proximal 2 secundibrachs with coarse nodes, with line of coarser nodes along lateral edges. Stem large, circular in transverse section, 9mm diameter. REMARKS. The crown of Necopinocrinus tycherus is crushed along the A-CD plane of symmetry. Radials are cracked and impacted downward, overlapping the distal tips of the basals, the E-A basal is inset and edges are over- lapped by adjacent plates, the infrabasal circlet is compressed, brachials are slightly offset from the cup and one another, and the stem and distal parts of the crown are lost. This specimen represents a conservative stock of the Euspirocrinidae showing an advanced condition of: 1, the anals restricted to the area above the extended posterior basal; 2, the infra- basal circlet of 3 plates; and 3, the arms branching on the single primibrach. Anaglyptocrinus and Necopinocrinus are the first post Carboniferous euspirocrinids reported, extending the range of the family into the Late Permian. Superfamily SCYTALOCRINOIDEA Moore & Laudon, 1943 Family SPANIOCRINIDAE Moore & Laudon, 1943 Spaniocrinus Wanner, 1924 TYPE SPECIES. Spaniocrinus validus Wanner, 1924 from the Permian Basleo Beds of Timor; by original designation. NEW PERMIAN CRINOIDS Spaniocrinus geniculatus Sp. nov. (Fig. 9) ETYMOLOGY. Latin geniculatus, like the bent knee; refers to the knee-shaped brachials. MATERIAL. HOLOTYPE: QMF38987 from QML518. PARATYPE: QMF39011, same. DIAGNOSIS. Crown slender, elongate; cup medium bowl; ornament of coarse nodose to short irregular ridges cont- inuing onto brachials, with prominent longitudinal ridge or keel along middle of brachials: brachials recti- linear to slightly cuneate, interlocking laterally. Arms 5. Stem round, hetero- morphic. DESCRIPTION. Crown slender, elongate, 57.3mm long (incomplete), 21.2mm wide, widest at first brachial, tapering distally. Cup medium bowl, 6mm long, 18mm wide, with coarse nodose to irregular ridge ornament, continuing onto brachials; sutures impressed. Infrabasal circlet small, not exposed, may be in shallow impression. Basals 5, convex longi- tudinally and transversely, proximally forming base of cup. distal part forming base of cup walls, widely outflaring. Radials 5, large. wider (9.5mm) than long (6mm), strongly convex longitudinally, moderately convex transversely, tumid, outflared. Radial facet plenary. Brachials much wider than long (first brachial 4.5mm long, 10.5mm wide), rectilinear to slightly cuneate, moderately convex transversely, straight longi- tudinally, prominant central longitudinal ridge or keel. with coarse nodose ornament, interlocking laterally, transverse ridges and grooves on lateral ends exterior to pinnular facets; 2 small pinnules on each side, transverse outline angular. Arms 5. tapering distally. Anals not exposed. Stem round, 5mm diameter, heteromorphic: noditaxis pattern N212. cirriferous on second nodal below cup: 53.6mm preserved. Columnals moderately long, (nodals 3.5mm long. internodals 2.6mm long 31mm below cup); latus convex, with coarse nodose ornament proximal to cup. smooth distally. Cirri round, 2.5mm diameter. 299 FIG. 8. Necopinocrinus tycherus gen. et sp. nov., D-E interray view of partial crown QMF38901, x1.7. REMARKS. The external mould of the crown and proximal stem of the holotype has the distal part of the arms partly disarticulated and central parts of the arms missing. Parts of 4 rays are preserved. The paratype is a set of arms, lacking the cup. The medial ridge on the brachials is well developed on both the holotype and paratype. Measurements taken from latex cast of holotype. Comparisons are made with species of Span- iocrinus and Parspaniocrinus because the two genera are closely related. The medium bowl- shaped cup of S. geniculatus is lower than that of either S. validus Wanner, 1924, S. transcaucas- icus Yakovlev, 1933 or Parspaniocrinus beinerti Strimple. 1971. all of which have truncated medium cones. and S. trinodus Weller, 1909 has a much narrower turbinate cup. The coarse nodose ornament of S. validus and S. transcaucasicus, the triple nodes on the radials of <. trinodus. and the fine granular ornament of P. beinerti, lack the irregular longitudinal ridges of 5. geniculatus. Brachials of P. beinerti have a rounded exterior in transverse section, whereas brachials of 5. geniculatusare like the angulartransverse outline of S. validus. The cup and shorter brachials of 5. geniculatus are advanced features, probably derived from S. validus. 300 MEMOIRS OF THE QUEENSLAND MUSEUM Fig. 9. Spaniocrinus geniculatus sp. nov. А,В. enlarged (1.7) view of ray to left in figure B and lateral view of crown * 1,1), holotype QMIE 38987. C. D. exteriorand interior views of partial set of arms, paratype QMI'39011. x].8. Superfamily DECADOCRINOIDEA cone or bow! shaped with small basal concavity: Bather, 1890 five infrabasals with only distal tips at most Family DECADOCRINIDAE Bather. 1890 Visible in side view; five medium-sized basals: DIAGNOSIS. Moore & Strimple (in Moore & five radials with articular facets as wide as plates: Teichert, 1978: 685) gave the diagnosis as: One to three anals in cup; anal sac tall, slender. "Crown slender. Cup widely expanded, truncate Arms ten, formed of cuneate uniserial brachials. NEW PERMIAN CRINOIDS branching isotomously on primibrachs 2 in geologically older forms, and on primibrachs | in later ones, no further branching, arms sinuous or zigzag in appearance. pinnules stout, tending to resemble ramules. Stem preponderantly round transversely and noncirriferous (except Aulo- crinus).^ REMARKS. The Decadocrinidae were recog- nised primarily on the zigzag nature of the 10 arms and Moore & Strimple (in Moore & Teichert. 1978) considered them intermediate in evolutionary development between some of the genera with rectilinear uniserial arms and some withbiserial arms. Taxa with more than 10 zigzag arms were assigned to one of several families based on arm branching patterns and other cup features (i.e. Plummericrinus in the Pachylocrinidae; Sp/ieniscocrinus in the Ampelo- crinidae). Using the zigzag nature ofthe 10 arms, Holcocrinus should have been assigned to the Decadocrinidae instead of the Graphiocrinidae. Without the zigzag appearance of the arms, genera assigned to the Decadocrinidae could have been assigned to the Scytalocrinidae or Graphiocrinidae on the basis of cup shape and number of anals within the cup. However, except for Parascytalocrinus all scytalocrinids have a truncated cone-shaped cup. Parascytalocrinus was established by Kammer & Ausich (1993) for species with a low bowl-shaped cup with a flat or shallow basal invagination and an atomous A ray previously assigned to Scytalocrinus. Inthe same paper, they erected Lanecrinus for species with 10 zigzag arms previously assigned to Scytalo- crinus. This restricted Scvtalocrinus to species with conical cups and non zigzag arms. If the zigzag pattern of the brachials is looked at closely. most genera show that it is dominantly the result of a slight to moderate extension on the long side of the cuneate brachial into a distal shoulder where the pinnule attaches. A node or blunt spine, which accentuates the zigzag appearance when present. may be positioned on the distal shoulder adjacent to the pinnule facet on the outer side of the brachial. 7rautscholdi- crinus lacks the zigzag appearance of the arms, but shows a faint zigzag pattern on the medial keel of the cuneate brachials. There is considerable difference in the length of the brachials in the Decadocrinidae. The brachials of 7rautscholdicrinus, Zostocrinus and Fireocrinus are the longest, G/aukosocrinus has intermediate length brachials and all other genera have very short brachials. With the exception of 301 FIG. 10. Glaukosocrinus middalvaensis sp. nov., posterior view of holotype QMF 38881, x2.2. Decadocrinus and Zostocrinus they branch on the single primibrachials. The genera of the Decadocrinidae do not fit into an evolutionary lineage and the family is herein considered polyphyletic. They probably represent advanced taxa evolved from several conservative genera within the cuneate brachial clade recognised by Webster (1997) or other rectilinear brachial genera. Revision of the Decadocrinidae is beyond the scope of this study and should be incorporated in a revision of the Poteriocrinina. Until such a study is completed the Decadocrinidae is retained for convenience. Glaukosocrinus Strimple, 1951 TYPE SPECIES. Malaiocrinus parviusculus Moore & Plummer, 1940 from the Desmoinesian Millsap Lake Formation, Parker County, Texas; by original designation. Glaukosocrinus middalyaensis sp. nov. (Fig.10) ETMOLOGY. From Middalya Station, WA. MATERIAL. QMF38881 from QML1240. DIAGNOSIS. Crown cylindrical, with very fine nodose to vermiform ornamentation; cup with basal invagination; radial facets peneplenary; 3 anals in cup; radianal and anal X large; single axillary primibrach elongate; brachials cuneate; large pinnules relatively short; 10? arms distinctly zigzag; stem round, with narrow crenularium, with wide areola, with round small lumen. DESCRIPTION. Crown cylindrical, medium size, incomplete length 43.1 mm, crushed width 26.4mm, very fine nodose to vermiform orna- mentation extending onto arms. Cup medium bowl, shallow basal invagination, crushed length 10mm, crushed width 20mm maximum, 8.2mm minimum. Infrabasals 5, small, horizontal, in basal invagination, not visible in lateral view. Basals 5, medium size, strongly convex longi- tudinally, moderately convex transversely, forming walls of basal invagination, basal plane, and base of cup walls. Radials 5, large, length 8.2mm, width 9.1mm, gently convex longitud- inally and transversely. Radial facet peneplenary, deep, sloping outward strongly. Anals 3; radianal large, 8mm long, 6mm wide, adjoining C radial, CD and DE basals, anal X, and right tube plate; anal X pentagonal, large, 7.8mm long, 6mm wide, widest near distal end. Right tube plate elongate, 5.6mm long, 4.1mm wide, narrowest on proximal end, proximal 1/3 below radial summit. Single primibrach axillary, constricted medially, length 8.1mm, width 7.7mm, widest on proximal end, strongly convex transversely, concave longitudinally. Brachials cuneate, ap- proximately equidimensional, strongly convex transversely, straight longitudinally, with wide pinnule facet on alternating distal ends giving arms distinct zigzag appearance. Pinnules wide, stout, relatively short. Ambulacral groove deep V-shaped. Arms 10? Stem round; facet with narrow crenularium, wide areola, narrow round lumen, REMARKS. The crown is crushed and cup plates are dislocated in part. The infrabasal circlet was partly exposed by cleaning and barely extends MEMOIRS OF THE QUEENSLAND MUSEUM beyond the stem facet. Only 3 basals are preserved and all are distorted by compaction. Solution weathering has destroyed most surface ornament except along part of the D radial, anal X and first tube plate. Glaukosocrinus middalyaensis is distinguished from G parviusculus (Moore & Plummer, 1940) and G planus Strimple & Moore, 1971 by the very fine anastomosing ornament. In addition, the primibrach is longer than that of G planus. This is the first report of Glaukosocrinus outside North America and the first in the Permian. The peneplenary radial facets make relatively narrow radial notches as on the cup of G. parviusculus. The arm branching on the single primibrach and shallow basal invagination are advanced features, while the 3 anals in the cup is a primitive feature. These features did not change significantly in the Late Carboniferous or Early Permian. Eidosocrinus gen. nov. TYPE SPECIES. Zidosocrinus condaminensis sp. nov. from the Condamine Beds, Elbow Valley arca, near Warwick, SE Queensland. ETYMOLOGY. Greek eidos, form or likeness, and krinon, lily; refers to the types based on latex casts. DIAGNOSIS. Crown cylindrical; cup low bowl-shaped, base invaginated, one anal, with single axillary primibrachs of differing lengths in different arms, cuneate brachials, 10 arms zigzag, coarse horn like nodes or blunt spines on the basals, radials, and distal tips of all brachials, fine granulate ornament on basals and radials. REMARKS. Mild to moderate tumidity of cup and arm plates in the Poteriocrinina is known in Spheniscocrinus and Cromyocrinus, among others. Likewise, coarse nodes or blunt spines on the axillary brachials are developed on the Pir- asocrinidae (Pirasocrinus, Sciadocrinus) and Zeacrinitidae (Tholocrinus), among others. Tri- ceracrinus (assigned to the Pirasocrinidae) has coarse horn-like nodes on the basals, radials and primibrachs (similar to those of Eidosocrinus), but lacks the nodes on the very short, weakly cuneate secundibrachs that have a medial transverse ridge. Thus, the ornamentation of Eidosocrinus is a distinguishing character. The differing length from ray to ray of the FIG. 11. Eidosocrinus condaminensis sp. nov. A, lateral view of disarticulated partial crown, paratype QMF 38904, x2.6. B, basal view of slightly disarticulated crown, paratype QMF38903, х2.7. C,D, basal (x2.2) and B ray (x1.8) views of holotype QMF38902. E, internal view of posterior interray, paratype QMF38905, x4.3. NEW PERMIAN CRINOIDS 303 104 axillary primibrachs of &idosocrinus is found in several Scvtalocrinidae, Aphelecrimdae. Graph- iocrinidac, among other poteriocrininids. This feature is considered intermediate between branching above the first primibrach and on the single primibrach. Combined with features of the cup il may гећесі а closer evolutionary relation- ship of the Scvtalocrinidae, Aphelecrinidae and Decadocrinidae than with the rectilinear brach- jals of the Stachyocrinidae. Eidosocrinus condaminensis sp. nov. (Fig. 11) ETYMOLOGY. From the Condamine Beds. MATERIAL. HOLOTYPE: QMF38902 from. ОМІ 518 PARATYPES: QMF389(3-38905, ишим. DIAGNOSIS. As for genus. DESCRIPTION. Crown incomplete. 34. 1mm preserved, cylindrical, ornament of single coarse hom-like node or blunt spine on all basal. radial, and brachial plates. Cup low bowl-shaped. 17.8mm wide, 8.8mm long, base invaginated. fine granular ornament. Infrabasal circlet small. 5 Imm diameter. subhorizontal. not visible in lateral view. Basals 5. 4,5mm long. 5.3mm wide, strongly tumid, outflared. forming base of cup and walls, Radials large. 5mm long. 7.5mm wide, strongly tumid. slightly flaring. forming most of cup wall. Radial facet plenary. subhorizontal. slightly concave transversely, deep, withelevated transverse ridge on central 1/3. deep elongate ligament pit, wide outer margin. Primanal large. 4inm long. 4.7mm wide, very tumid, abutting distally terminated posterior basal, proximal 1/2 in line of radials. distal 1/2 projecting above radial summit, distally adjoined by 2 anal tube plates. Single primibrachs axillary in all rays, A ray longest (8. этот). C ray intermediate (greater than 4.7mm), В and E rays shortest (4.9mm), strongly comvex transversely, concavo-coivex longitudinally, hourglass-shape in exterior view, Secundibrachs cuneate. approximately as wide as long. deep. strongly convex transversely. straight proximally becoming convex distally, node adjacent to pinnule facet on long side. Branching isotomously, 10 arms. Stem round. heteromorphic: noditaxis pattern N1. Columnals moderately long: crenularium narrow: lumen small. circular?: latus roundly convex on nodals, gently convex on internodals. REMARKS. Descriptionof Lidosocrinus conda- minensis is based on the casts of all types because no specimenis complete. Measurements made on MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 12. A.B, Eoindocrinus praecontignatus Arendt, 1981. oblique lateral and lateral views of cup ОМЁЗ38910. х5.2, C, Pedinocrinus? nodosus sp. noy., C-D iaterray view of holotype QMF 38906. with Е. praecontignatiscup on arins m uppernght, “17. the holotype. The holotype has part of the D. A, B. and C rays; some cup and ann plates are dislocated slightly. The 1nfrabasal circlet, D-E basal plate. E ray. and anal are lost through weathering. A poorly preserved pluricolumnalin alignment with. directly below. and 6mm from the cup is probably part of the stem. It is round, 2.2mm diameter. heteromorphic (noditaxis NEW PERMIAN CRINOIDS pattern N1) with moderately elongate columnals. Paratype QMF38903 is crushed, all cup plates are slightly dislocated, the proximal columnal nearly covers the infrabasal circlet, and only the prox- imal part of the E and A rays are preserved. Paratype QMF38904 is also crushed and retains part of the cup, proximal columnals, and primi- brach of one ray. Paratype QMF38905 is an internal and external mould of a cup showing the basals, radials, anal, and C ray primibrach. The fine granular ornament is preserved on some cup plates of all 4 specimens. Tentative assignment of Eidosocrinus conda- minensis to the Decadocrinidae is for convenience and based primarily on the zigzag nature of the arms. Superfamily LOPHOCRINOIDEA Bather, 1899 Family STELLAROCRINIDAE Strimple, 1961 Pedinocrinus Wright, 1951 TYPE SPECIES. Pachylocrinus clavatus Wright, 1937 from the Early Carboniferous, Tournaisian, Lower Limestone Group, Scotland; by original designation. Pedinocrinus? nodosus sp. nov. (Fig. 12C) ETYMOLOGY. Latin nodus, nodes; refers to the nodose omament of the cup and proximal brachials. MATERIAL. HOLOTYPE: QMF38906, from ОМІ5 18. DIAGNOSIS. Crown flaring distally, cup low bowl-shaped, 3 anals, coarse nodose ornament continuing onto proximal brachials, Ist primi- brach, 8th secundibrach and 9th tertibrach axillary; bulbous tegmen; stem round, hetero- morphic. DESCRIPTION. Crown moderately large, 47.7mm long (incomplete), 42.3mm wide (still expanding), pear-shaped, arms flaring. Cup low bowl, 20mm wide (incomplete?), 6.5mm long (estimate), base invaginated. Coarse nodose ornament on all cup plates, primibrachs and proximal 3-4 secundibrachs; all axillary brachials above primibrachs nodose or bearing short blunt spines. Infrabasal circlet not exposed, within impressed basal cavity. Basals relatively small, 4mm long, 4.8mm wide, tumid, moder- ately convex longitudinally and transversely, subhorizontal to gently upflared. Radials largest cup plates, 3.6mm long, 8.8mm wide, moder- ately convex transversely, strongly convex longitudinally. Radial facets plenary. Anals 3, moderately large; radianal in CD interray, 305 supporting both rectangular anal X and right tube plate directly above in radial circlet. Brachials cuneate, uniserial to biserial, strongly convex longitudinally and transversely, sutures impress- ed, one pinnule on widest side. In C ray Ist primibrach, 8th secundibrachs, and 9th terti- brachs axillary, probably | or 2 additional branchings distally. All branching isotomous, minimum of 40 arms if all rays branch as in C ray. Arms flare moderately laterally. Anal tube large, probably bulbous, formed of many small poly- gonal plates. Stem round, heteromorphic; noditaxis of N212 or NI in proximal 20mm preserved. Columnals with strongly convex latus. REMARKS. The external mould of Pedino- crinus? nodosus preserves part of the posterior side of the cup, the proximal parts ofthe C and D rays including some quartibrachs and tegmen plates. Cup plates are partly dislocated with the C radial nearly covering the right tube plate. The cup is covered by a cladoporid coral in part. All measurements are approximate, from latex casts. At first glance P.? nodosus appears to resemble Plaxocrinus, Tholocrinus and Hydreionocrinus, all of which have moderately large flaring crowns with a low bowl-shaped or discoidal cup, axillary brachials bearing short spines or blunt nodes and large inflated tegmens. However, these taxa are placed in different familes based on the number of anals in the cup, type of brachials, and arm branching patterns. These taxa range in age from Early Carboniferous into the Late Permian and represent heteromorphic evolution within different lineages of the poteriocrininids in the late Palaeozoic. Lacking ornamentation this specimen would be placed in Pedinacrinus without question. Arguments could be made for erecting a new genus for P.? nodosus. However, we do not believe that ornament alone is sufficient for establishing a new genus. The significant time gap between the Tournaisian P. clavatus (Wright, 1937) and the Artinskian P.? nodosus suggests Pedinocrinus may be a holdover in Australia. The coarse nodose ornament on the cup plates and proximal brachials of P.? nodosus should assist future recognition. Stellarocrinid? gen. et sp. nov. (Fig. 13C) MATERIAL. UQF12211A, from QMLS18. DESCRIPTION. Cup unknown. Arms broad, widespread. Brachials uniserial, 4.6mm long, 11.5mm wide, mildly cuneate, deep, with coarse 306 nodose ornament; with very large blunt node elongated parallel to arm length on inner side of long end, with notch for pinnule facet on outer side of short end. Ambulacral groove large, 2mm wide, V-shaped, joined by side grooves from short end of brachials. C and D ray arms branching isotomously on single primibrach, may branch again distally. Branches widely flared laterally. Tegmen formed of several in- flated small (3.6mm diameter) to medium sized (7mm diameter) polygonal plates adjacent to 2 inflated very large orals (11mm diameter) at base of large anal tube (19mm diameter). Base of anal tube formed of 5 columns of vertically stacked plicate hexagonal plates (5.6mm long, 6.4mm wide); anal tube length unknown. Additional columns of tegmen plates not preserved, estimated minimum of 4 or 5 present. Smaller (3.6mm long and wide) anal tube plates projecting outward from the 3rd row of tube plates. REMARKS. The specimen is an external mould of the oral surface of a large partial crown con- sisting of parts of 7 arms probably belonging to 5 rays. The 2 arms ofthe C and D rays branch close to the tegmen and the ambulacral groove of each of the 2 adjacent arms (B and E rays) join the ambulacral groove of the C and D rays before passing into the interior ofthe tegmen. It could be proposed that each of these sets of 3 arms are part of 1 ray, which would require 2 more rays behind the tegmen and not preserved. This is not likely with the excellent preservation and spacing ofthe arms. The 7th arm, the A ray, is undivided on the preserved part. The distal end of the E ray is regenerated, as distal brachials are much smaller than proximal brachials. The first brachial of the regenerated section is axillary. All arms probably branch again distally. All measurements are ap- proximate, taken from a latex cast. Although pinnules are not preserved their presence 1s presumed because notches for their attachment are present on the outer side of the brachials and a large U-shaped ambulacral groove along the short end of the brachial adjoins the main ambulacral groove. Attachment of the pinnules to the short end of the brachials is an exception to the normal attachment on the long end. Both the development of the large nodes on the inside of the long end of the brachial and the pinnule attachment on the short end are MEMOIRS OF THE QUEENSLAND MUSEUM considered evolutionary developments of the specimen, not known in other poteriocrininids. The small anal tube plates projecting laterally from the anal tube probably represent the distal parts of a recurved anal tube. The plates are slightly disarticulated and adjoined more distal plates that are not preserved. The specimen represents a new genus but is considered inadequate to serve as a holotype, lacking the cup. It is assigned to the stellaro- crinids because the laterally projecting arms spread widely, structure of the large elongate anal tube, presumed branching pattern of the arms, and cuneate brachials bear coarse ornamentation. These are all features of the stellarocrinids. Family SUNDACRINIDAE Moore & Laudon, 1943 Sundacrinus Wanner, 1916 TYPE SPECIES. Sundacrinus granulatus Wanner, 1916 from the Permian Basleo Beds, Timor; by original designation. Sundacrinus medius sp. nov. (Fig. 13A,B) ETYMOLOGY. Latin medius, middle; refers to the cup shape intermediate between that of 2 previously described species. MATERIAL. QMF38908 from QMLS18. DESCRIPTION. Crown medium size, pear- shaped, 32.4mm long (incomplete), 19.6mm wide. Cup medium to high bowl-shaped, 13mm long, 18.6mm wide at radial summit, base gently upflared, all plates very thick with coarse nodose ornament grading into irregular anastamosing ridges. Infrabasal circlet large, 9.5mm diameter, gently upflared, visible in lateral view. Basals largest plates in cup. gently convex longitudinally and transversely, widely flaring, of variable size and shape; posterior basal hexagonal, 7.5mm long, 8.4mm wide, adjoining radianal, BC basal, infrabasals, CD basal, D radial and anal X. Radials large, 6.5mm long, 7.5mm wide, subvertical to slightly incurved distally, weakly convex longi- tudinally and transversely. Radial facet plenary, strong outward-downward slope. Two pentagon- al anals in cup; radianal largest, adjoining C radial, BC and CD Basals, anal X, and first tube plate, distal tip projecting slightly above radial summit; anal X adjoining radianal, CD basal, D FIG. 13. A,B, Sundacrinus medius sp. nov., posterior and D ray views of holotype, QMF38908, х2.6. C, Stellarocrinid? gen. et sp. nov., oral view of tegmen and arms, UQF12211A, x1. D, Moapacrinus cuneatus sp. nov., posterior view of slightly disarticulated crown, holotype, QMF38909, x 1.9. 307 NEW PERMIAN CRINOIDS 308 radial, D primibrach, and overlying tube plate, distal 1/3 above radial summit. D ray primibrach elongate, straight longitudinally, strongly convex transversely, may be axillary. Secundibrachs cuneate uniserial, elongate, deep, straight longi- tudinally, strongly convex transversely, with wide V-shaped ambulacral groove. Anal tube stout, projecting above cup, formed of thick hex- agonal plates of uncertain structural pattern, probably laterally interlocked stacked columns, length unknown. Stem round transversely, 4.5mm diameter, heteromorphic, of variable noditaxis pattern in 24mm length preserved, at least 4 distinct sizes of columnals; latus mod- erately to strongly convex. REMARKS. This partial crown consists of a slightly crushed cup below a jumbled pile of dislocated brachials and anal tube plates. It is assigned to Sundacrinus based on cup shape, thick plates, plenary radial facet sloping outward- downward and irregular shape of cup plates. Sundacrinus medius has a cup shape inter- mediate between the conical cup of S. triangulus Wanner, 1924 and the bowl-shaped cups of S. granulatus Wanner, 1916 and S. vastus Wanner, 1924. The elongate cup of S. elongatus is much more slender than that of S. medius. Moore et al. (in Moore & Teichert, 1978) recognised that the number of anals in Sundacrinus varied, reporting 1, rarely 2. However, there are 2 in S. cf. vastus (Wanner, 1937, pl. 10, fig. 25) and 3 in S. tri- angulus (Wanner, 1937, pl. 10, fig. 21). Thus 2 anals in S. medius is intermediate. This is the first report of the anal tube of Sun- dacrinus and the first report of the genus in Australia. It provides additional support for inte- connections of E Australia and Timor. Superfamily CROMYOCRINOIDEA Bather, 1890 Family CROMYOCRINIDAE Bather, 1890 Moapacrinus Lane & Webster, 1966 TYPE SPECIES. Moapacrinus rotundatus Lane & Webster, 1966 from the Artinskian part of the Bird Spring Formation, Nevada; by original designation. Moapacrinus cuneatus sp. nov. (Fig. 13D) ETYMOLOGY. Latin cuneatus, wedge-shaped. MATERIAL. HOLOTYPE: a crushed, partly dis- articulated, partial crown, QMF38909 from QML518. DIAGNOSIS. Crown elongate, cup medium bowl-shaped, shallow basal invagination, sutures MEMOIRS OF THE QUEENSLAND MUSEUM impressed, coarse nodose ornament, single large anal, axillary 15 primibrach, brachials strongly cuneate. DESCRIPTION. Crown elongate, 46.2mm long, incomplete. Cup medium bowl-shaped, 7.6mm long, 16mm wide, shallow basal invagination, sutures impressed, coarse nodose ornament, slightly incurved at radial summit. Infrabasal circlet not visible in lateral view. Basals large, 5.3mm long, 6.7mm wide, strongly convex longitudinally and transversely, forming base of cup and lower part of cup wall. Radials of intermediate size, 4.8mm long, 8.1mm wide, moderately convex longitudinally and trans- versely, subvertical. Radial facet plenary, deep; transverse ridge slightly concave externally; liga- ment pit elongate, deep; narrow outer margin; muscle fields large, intermuscular furrow shallow. Single anal large, 4.5mm long, 5.2mm wide, directly above posterior basal, distal 2/3 above radial summit. Single primibrach axillary. Brach- ials uniserial, strongly cuneate, gently convex longitudinally, strongly convex transversely; 10 arms. REMARKS. Moapacrinus cuneatus has wedge- shaped brachials and is ornamented with coarse nodes, whereas other species of the genus have rectilinear brachials and lack coarse nodose ornament. Pabian & Strimple (1993) reported fine granular ornament on M. elexensis Pabian & Strimple, 1993 known only from a cup. Only the posterior 1/2 of the cup of M. cuneatus is exposed; the C and D ray arms are dislocated and brachials partly disarticulated. This is the first report of Moapacrinus outside North America, the first record of a cromyocrinid in E Australia, and the youngest cromyocrinid known. Cromyocrinids are common in late Pal- aeozoic faunas of the Midcontinent and Rocky Mountain regions of the USA. Pabian et al. (1989) reported the cromyocrinids in their ‘Terrigenous Facies Belt’, implying some clastic sediment entering the living environment. Webster & Houck (1998) noted that cromyocrinids dom- inate Late Carboniferous faunas in intermontane basin settings of the Rocky Mountain region. Although carbonates dominated the environment, some sand size clastic sediment was deposited wherein the cromyocrinids were living. Thus, a cromyocrinid was probably well adapted for living in the mudstone environment of the Con- damine Beds. NEW PERMIAN CRINOIDS FIG. 14. A, B, Parabursacrinus granulatus Wanner, 1949, D ray and basal views of crushed crown QMF38882, x32. Family INDOCRINIDAE Strimple, 1966 Eoindocrinus Arendt, 1981 TYPE SPECIES. Eoindocrinus praerimosus Arendt, 1981 from the late Artinskian Sarginsk Horizon, Ural Mts; by original designation. Eoindocrinus praecontignatus Arendt. 1981 (Fig. 12A,B) MATERIAL. External mould of cup, QMF38910 from OMLS518. REMARKS. This small cup (5.4mm long. 5.8mm wide), on the arms of Pedinocrinus? nodosus, is oriented on its side with the C-D basal centred, the basal circlet upturned and the oral rim crushed downward (not visible). The large stellate ridge ornament converges in the centre of the basals and forms triangles across adjacent plates. Smaller inflated triangles are formed within these at the apices of triple plate junctions. A ridge junction also occurs on the radianal which 309 FIG. 15. A.B, Timorechinid gen. indet., lateral views of partial set of arms QMF38883, x3.3. supports the right tube plate distally and is adjacent to anal X. A specimen of К. praecontignatus from the Wandagee Sandstone of Western Australia has partly developed secondary ridges forming a secondary triangle within the primary ridge triangle (Webster, 1990). The Condamine and Wandagee forms are considered conspecific with variation in ornament comparable to that in Æ. praecontignatus from the Urals (Arendt. 1981). Superfamily ZEACRINITOIDEA Bassler & Moodey, 1943 Family ZEACRINITIDAE Bassler & Moodey, 1943 Parabursacrinus Wanner, 1924 TYPE SPECIES. Bursacrinus procerus Wanner, 1916 from the Basleo Beds, Timor. by original designation. MEMOIRS OF THE QUEENSLAND MUSEUM 310 jw me^ а! DL М "Juli wp» n g Fa um wes № 38884, C.D, Poteriocrinitid indet., arms 3, interior (х3) and exterior (х4) lateral views of QMF38887, х4. 6. A.B, Poteriocrinitid indet., arms 1. A, lateral view of QMF38885, 3.2. B. lateral view of FIG. 1 3.1. NEW PERMIAN CRINOIDS 31] Parabursacrinus granulatus Wanner, 1949 (Fig. 14) MATERIAL. QMF38882 from QML 1232. REMARKS. This small crown is probably an immature or young adult, It is crushed against the C-EA axis, infrabasals and basals are not visible and distal parts of the arms are lost. The single anal projects 1/2 above the radial summit. Arms all bifurcate on the Ist primibrach. Granulose ornament continues onto the rectilinear brach- ials. Heteromorphic proximal stem columnals are round in section. Family TIMORECHINIDAE Jaekel, 1918 Timorechinid gen. indet. (Fig. 15) MATERIAL. QMF38883 from QML 1237. DESCRIPTION. Arm fragment incomplete. slender. 32mm long, 13.5mm wide, parts of , or possibly 4 rays present. Arms 3. slender. Brachials uniserial, rectilinear, gently convex longitudinally. moderately convex transversely. Isotomous branching on 4th and 5th brachials, againon4th and 5thbrachialsonouter 1/2 of arm, probably endotomous. Pinnules and ambulacral groove not visible. REMARKS. This specimen has the arms enclosed. is crushed, and probably represents parts of 3 rays. The main part visible is judged to represent | ray which had bifurcated isotomously below the preserved part. As interpreted there are 6 arms in the ray. total of 30 arms if all rays bifurcate uniformly. Brachials and arm branching pattern of this type occur in Notiocrinus and Parabursacrinus of the Timorechinidae to which the specimen is referred. Poteriocrinitid indet., arms 1 (Fig. 16A.B) MATERIAL. QMF38884 and 38885 from QML 1237. DESCRIPTION. Fragment 1. Arms slender, elongate: fragment 29.2mm long. 19.4mm wide, incom- plete, including medial portions of a minimum of 16 arms with one additional distal branching on most arms. Brachials rectilinear to moderately cuneate, gently convex longitudinally, strongly convex transversely. Axillary brachials strongly protruded. One slender pinnule per brachial on alternate sides of arm. All branchings isotomous. but only branch on one half of arm distally, probably endotomous. FIG. 17. Poteriocrinitid indet., arms 2, lateral view of QMFS388806. x1.8. Fragment 2. 32mm long. 14mm wide, incomplete, medial portions of a minimum of 9 arms. Description as for fragment 1. REMARKS, These 2 fragments may belong to a single specimen as they were found within 15 cm of one another. They are the medial and distal parts of the arms and, if from 1 specimen, there were a minimum of 40 arms. In the enclosed position the arms have a jointed or knotted ap- pearance at the branchings, similar to those of several poteriocrinitids, such as Abrotocrinus and Anchicrinus. Poteriocrinitid indet.. arms 2 (Fig. 17) MATERIAL. QMF38886 from QML 1240. DESCRIPTION. Partial set of arms 53.8mm long. 39.5mm wide. incomplete. arms unbranched, loosely parallel. Brachials medium size. mod- erately cuncate, straight to weakly convex longitudinally. roundly convex transversely, with pinnule on long end; ambulacral groove shallow. rounded V shape. 2 — N FIG. 18. A-C, Poteriocrinitid indet.. arm fragment 1, oral. lateral and exterior views, QMFE3891 1, «2.5. REMARKS. The arms have a slight zigzag appearance as a result of weathering, especially MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 19. Poteriocrinitid indet., arm fragment 1, camera lucida sketch of exterior surface of 4 brachials shown in Fig. 18C. *3. Similar parallel ruled sections are external parts of a single brachial. sides of the specimen. It is not known if the A ray was unbranched and there were only 9 arms or a 10th arm was lost with weathering. These arms are similar to a number of poteriocrinitids. especially some scytalo- crinids and decadocrinids. Poteriocrinitid indet.. arms 3 (Fig. 16C.D) MATERIAL. QMF38887 from QML 1237. DESCRIPTION. Arm frag- ment 30mm long, 17.9mm wide, with parts of 10 unbranched arms. Arms slender. Brachials uniserial. strongly cuncate, gently convex longitudinally, strongly convex trans- versely, Ambulacral groove wide, open rounded V- shaped, One slender pinnule per brachial on long end. REMARKS. The specimen represents the distal the long pinnule bearing end of the brachials with part of a minimum of 10 arms, which may rep- greater relief. Where there is little weathering the resent only 1/2 the arms of the specimen as the zigzag is slight. Nine arms are present on the 2 pinnules and interior of the arms are visible on 1 NEW PERMIAN CRINOIDS side ofthe small slab, the exterior of 4 arms on the other side and some arms between these are visible in end view. Poteriocrinitid indet., arm fragment | (Figs 18, 19) MATERIAL. OMF38911 from QML518. DESCRIPTION. Fragment, very large 29.2mm long, 25.2mm wide, 10.5mm deep, consisting of 8 brachials. Single brachial 7.8mm long, 25.2mm wide, 10.5mm deep. Brachials uniserial but ap- pearing biserial (=pseudobiserial). In exterior view each brachial divided into 2 parts; larger pentagonal section borders 2 adjacent brachials in middle of arm in apparent biserial interlocking fashion, sides adjoining 2 pentagonal sections of alternate brachials, end forming arm margin with pinnule facet; smaller section triangular, longer isosceles sides tapering toward centre of arm, ad- joining adjacent brachials on either side, shorter 3rd side forming arm margin with pinnule facet. Both sections convex longitudinally and transversely, continuous beneath the 2 adjacent brachial sections. Interior convex with central small V-shaped ambulacral groove. REMARKS. These brachials are the largest known for Palaeozoic crinoids, the only known pseudobiserial form, and one of the few bipin- nular forms, bearing one pinnule on each end ofa compound brachial. The bipinnular condition of Zeacrinus is a minimal form of hyperpinnulation. Hyperpinnulation, in which multiple pinnules are present on both sides of the arms, developed in a few camerates (Briarocrinus, 4 per brachial, 2 on each side) and poteriocrinitids (Cupressocrinites, 6 or 8 per brachial, 3 or 4 on each side; Neo- zeacrinus, 4 per brachial, 2 on each side), Hyperpinnulation is thought to have developed by fusion of adjacent brachials (Ubaghs in Moore & Teichert, 1978). Development of the pseudobiserial form requires overlapping of the 2 adjacent brachials, as well as the bipinnular condition. The following possible origins are suggested. First, that they evolved by fusion of 2 cuneate uniserial brach- ials, one becoming the shorter end and the other the longer end, each bearing a pinnule, with simul- taneous overgrowth of the 2 adjacent brachials. Second, that they evolved from biserial brachials in which 2 pinnule bearing brachials fuse at the midline with the simultaneous overgrowth of the 2 adjacent brachials. Third, that they evolved from cuneate brachials with development of a pinnule on the short non pinnule bearing end of a brachial, concurrent with overgrowth of the 2 adjacent brachials. The Ist or 2nd origin is most likely as the 3rd requires redevelopment of a pinnule on a non pinnule bearing end ofa brachial. Although no pinnules are attached to the brach- ials, 2 ossicles adjacent to the arm fragment are pinnulars, both with the wide V-shaped ambu- lacral goove exposed. The largest is 5mm long, 5mm wide and 3.5mm deep; the ambulacral groove is 1.5mm wide and 0.6mm deep. Their size and association with the arm fragment suggest that they belong to 1 species. Poteriocrinitid indet., arm fragment 2 (Fig. 20B,C) MATERIAL. QMF38912 from QMLS18. DESCRIPTION. Arm large, 59mm long (incom- plete), unbranched. Brachials large, (proximal brachial 4mm long on wide end, 7mm deep, estimated 8mm wide) strongly cuneate, biserial proximally, uniserial distallly, straight to gently convex longitudinally, strongly convex trans- versely, with large pinnule on wide end, ambulacral groove large, V-shaped. Proximal pinnule 3mm long, 4mm deep, concave longi- tudinally, strongly convex transversely. More distal pinnules slender, elongate, concave longi- tudinally, strongly convex transversely. REMARKS. This arm fragment is curved back- wards in a feeding or death posture. Pinnules are larger than brachials of many crinoids. It probably belongs to an unknown poteriocrininid. Poteriocrinitid indet., arm fragment 3 (Fig. 20A) MATERIAL. QMF39013 from QML518. DESCRIPTION. Brachials large, 3.3mm long, 4.8mm wide, moderately cuneate, slightly convex longitudinally, strongly rounded transversely, coarse nodose ornament; single pinnule on long end, pinnulars slender, elongate. Anal tube slender, formed of irregularly arranged polygonal plates with coarse nodose ornament; basal plate of tube with sharply pointed centrally expanded spine. REMARKS. The nodose ornament is randomly distributed externally with up to 20 on a single brachial. Nodose brachials occur in several Condamine fauna species. Nodes are more num- erous and smaller on Poteriocrinitid indet., arm fragment 3 than on Pedinocrinus? nodosus and coarser than on Poteriocrinites? smithii Etheridge, 1892. 314 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 20. A, Poteriocrinitid indet., arm fragment 3, lateral view, QMF 39013, x2. B,C, Poteriocrinitid indet., arm fragment 2, counterpart lateral views, QMF38912, x1.7. Subclass FLEXIBILIA Zittel, 1895 Order SAGENOCRINIDA Springer, 1913 Superfamily LECANOCRINOIDEA Springer, 1913 Family MESPILOCRINIDAE Jaekel, 1918 Loxocrinus Wanner, 1916 TYPE SPECIES. Loxocrinus globulus Wanner, 1916 from the Basleo Beds, Timor; by original designation. Loxocrinus booni Marez Oyens, 1940 (Fig. 21E.F) MATERIAL. QMF38888 from QML759. REMARKS. This partial cup consists of the D and E radials and distal tip of the DE basal. if the A radial is symmetrical. The cup is low bowl- shaped with the infrabasals probably not visible in lateral view. It is slightly abraded and NEW PERMIAN CRINOIDS 3l encrusted with bryozoans. Radial facets of the short, thick radial plates are shifted to the right of centre with the left shoulder wider than the right. The facets are concave, deep with a narrow rim of crenulae and culmina along the rounded outer edge; other details of the surface are lost by solution or covered by encrusting organisms. Only L. booni has a low bowl-shaped cup with the infrabasals not visible in lateral view, as the other 2 species, L. globulus Wanner, 1916 and L. dilatatus Wanner, 1916, are globose with the infrabasals visible in lateral view. Loxocrinus sp. | (Fig. 22A) MATERIAL. QMF38889 from QML757. DESCRIPTION. Radial small, 5mm long, 5.1mm wide, thick, proximally straight longitudinally, distally incurved, moderately convex trans- versely, fine granular to vermiform ornament; shoulders extended longitudinally, left shoulder wider than right, both wrapping around radial facet to ambulacral groove. Radial facet angust- ary, skewed right of centre, elliptical in outline, concave with marginal rim; transverse ridge aboral of centre, low; ligament pit in centre of radial, small, transversely elongate; outer margin- al area crescent-shaped; muscle areas large, with irregular surface; ambulacral groove moderately wide V shaped, confined to adoral edge of facet. REMARKS. If the radial facet of Loxocrinus sp. 2 were horizontal, the radial sloped inward for its entire length. Most likely the radial facet sloped downward, outward and the proximal part of the radial was vertical with the distal part curving inward. The shoulders left small obvious notches between arm bases. The specimen is the D or E radial if the A radial is symmetrical. Loxocrinus sp. 2 (Fig. 22B,C) MATERIAL. QMF38890, 38891 from QML758. DESCRIPTION. Radial small, thick, proximally straight longitudinally, distally incurved, mod- erately convex transversely, unornamented; shoulders extended longitudinally, left shoulder wider than right, wrapping around radial facet to ambulacral groove, right shoulder terminating against radial facet adoral of transverse ridge. Radial facet angustary, skewed right and left of centre, nearly circular in outline with extended right side muscle area, concave with marginal un rim; elevated transverse ridge 3/4 of distance aboral of facet; ligament pit aboral of centre of and culmina on aboral 1/2; muscle areas large, irregular surface; ambulacral groove moderately wide V-shaped, notched into adoral 1/4 of facet. QMF38891 8.1mm long, 7.6mm wide; QMF38890 8.2mm long, 10.1mm wide. REMARKS. Orientation of Loxocrinus sp. 2 would have been very similar to L. sp. 1. The two forms are distinguished by the lack of ornamentation on L. sp. 2. They both differ from L. booni by being much longer. QMF38891 is the D or E radial, if the A radial is symmetrical, as the facet is skewed to the right and the facet shows evidence of solution weathering. QMF38890 is the B or C radial with the facet skewed to the left. Family PROPHYLLOCRINIDAE Moore & Strimple, 1973 Prophyllocrinus Wanner, 1916 TYPE SPECIES. Prophyllocrinus dentatus Wanner, 1916 from the Basleo Beds, Timor; by original designation. Prophyllocrinus sp. 1 (Fig. 21A) MATERIAL. QMF38892 from QML758. DESCRIPTION, Radial medium size, 10.8mm long (incomplete), 10.4mm wide, medium depth, proximally straight longitudinally, distally incurved, moderately convex transversely, fine granular ornament; shoulders greatly extended longitudinally, left shoulder wider than right, ex- tending slightly beyond radial facet, right shoulder broken off, probably terminating against distal end of radial facet. Radial facet angustary, skewed right of centre, elongate U-shaped, con- cave with marginal rim; elevated transverse ridge 3/4 aboral length of facet; ligament pit small, centre of radial; outer marginal area crescent- shaped; muscle areas large, shallowly concave; ambulacral groove deep, wide V-shaped, notched into adoral 1/2 of facet. REMARKS. Solution etching has destroyed the proximal edge of the radial and some surface features of the transverse ridge and ligament pit of the radial facet. The right shoulder and a small piece of the left edge of the outer margin of the radial facet were broken off. The long U-shaped radial facet readily distinguishes the specimen from those of Loxocrinus and allies it with Pro- phyllocrinus. Breakage suggests a very short right shoulder, a character of Proapsiocrinus and Ancistrocrinus. MEMOIRS OF THE QUEENSLAND MUSEUM 316 FIG. 21. A, Prophyllocrinus sp. 1, lateral view of radial QMF 38892, x5.7. B.C, Prophyllocrinus sp. 2. B, lateral vieof radial QMF38893, «4.4, C, lateral view of radial QMF 38894, x4.1. D, Prophyllocrinussp. 3, lateral view of radial QMF38895, x9.2. EF, Loxocrinus booni Marez Oyens, 1940, oral and lateral views, QMF38888, x5.5, Prophyllocrinus sp. 2 DESCRIPTION. Radial medium size and (Fig. 21B.C) thickness, proximally straight longitudinally, MATERIAL. Radials QMF38893 from QML758 апа distally incurved, moderately convex transversely, QMF38894 from QML737. no ornament; shoulders greatly extended NEW PERMIAN CRINOIDS longitudinally, left shoulder longer than right, both terminate against distal end of radial facet. Radial facet angustary, skewed right of centre, elongate U-shaped, concave with marginal rim; elevated transverse ridge 3/4 aboral length of facet; ligament pit very small, centre of radial; outer marginal area crescent-shaped; muscle areas large, shallowly concave; ambulacral groove deep, wide V-shaped, notched into adoral 1/2 of facet. QMF38893 10.3mm long (incomplete), OMF38894 8mm long, 7.9mm wide (both incomplete). REMARKS. QMF38893 lacks the distal end of the right shoulder, the proximal edge ofthe radial and adoral edge of the ambulacral groove from solution etching. Solution etching has destroyed the proximal edge of the radial, some surface features of the transverse ridge and ligament pit of the radial facet and distal parts of the muscle area and ambulacral groove of QMF38894. Prophyllocrinus sp. 2 differs from P. sp. 1 by lacking ornament. Prophyllocrinus sp. 3 (Fig. 21D) MATERIAL. QMF38895 from QML758. DESCRIPTION. Radial small, 4.9mm long, 4.6mm wide, medium depth, proximally straight long- itudinally, distally incurved, moderately convex transversely, with coarse granular ornament; shoulders greatly extended longitudinally, left shoulder longer than right. Radial facet angust- ary, skewed right of centre, elongate U-shaped, concave with marginal rim; elevated wide trans- verse ridge located close to ambulacral groove at 1/3 adoral length of facet; ligament pit very small; outer marginal area crescent-shaped; muscle areas small, shallowly concave; ambulacral groove narrow V-shaped, notched into adoral 1/3 of facet. REMARKS. Solution etching has destroyed the distal end of the right shoulder and some surface features of the transverse ridge. Prophyllocrinus sp. 3 differs from Р. sp. 1 and P. sp. 2 by the coarse granular ornament. The specimen is probably a juvenile and with growth would have a facet much like that of P. sp. 1 or P. sp. 2. Sagenocrinitid indet. (Fig. 22D) MATERIAL. QMF38896 from ОМІ 1233. REMARKS. A weathered, poorly preserved, partial crown of an indeterminate sagenocrinitid 317 shows part of the cup plates, part of one ray including an interbrachial series of 2 plates, and distal brachials of 2 or 3 rays. Critical parts ofthe cup, if preserved are not exposed to identify the genus. This is the first crown of a sagenocrinitid reported from the Callytharra Formation. It is illustrated to show the faunal diversity. Loose flexible ossicles perhaps belonging to this taxon, are uncommon in bulk samples of the Callytharra Formation from the type section. Both cup and arm ossicles are present and the brachials show well-developed patelloid processes. Subclass ARTICULATA Zittel, 1879 The Articulata is revised to include 8 orders, 7 as in the Treatise (Moore & Teichert, 1978) plus Ampelocrinida below. Articulata are characterised by brachial pairs with alternating muscular and cryptosyzygial articulation. Order AMPELOCRINIDA ord. nov. REMARKS. The Ampelocrinida (Table 2) includes Corythocrinidae, Tribrachyocrinidae, Calceoli- spongiidae, Ampelocrinidae (as constituted below) and the unassigned Zasmanocrinus. It may be defined as Palaeozoic genera not previously in- cluded in the Articulata but with brachial pairs with alternating muscular and cryptosyzygial articulation and lacking perfect pentameral symmetry. Family AMPELOCRINIDAE Kirk, 1942 DIAGNOSIS. Cup bowl-shaped to discoidal, small; infrabasals small, subhorizontal to down- flaring, commonly not visible in lateral view; 1 anal (exception, 3 in Ampelocrinus); radial facets plenary; arms commonly 10, rarely more, 2 primibrachs (exception, 3-4 in Halogetocrinus); isotomous branching; cuneate pinnulate brach- ials ; brachial pairs with alternating muscular and cryptosyzygial articulation; short anal tube where known (exception, recurved in Ampelocrinus); proximal stem commonly pentagonal or sub- pentagonal, rarely circular, in transverse section; very cirriferous close to cup where known. GENERA INCLUDED. Ampelocrinus, Chlidonocrinus, Cymbiocrinus, Halogetocrinus, Moundocrinus, Oklahomacrinus. REMARKS. Some species assigned to these genera may not belong to the Ampelocrinidae but a review of them is beyond the scope of this study. Genera assigned to the Ampelocrinidae by Moore et al. (in Moore & Teichert 1978), here excluded are Arroyocrinus, Polusocrinus, MEMOIRS OF THE QUEENSLAND MUSEUM VIG. 22. A, Loxocrinus sp. 1, lateral view of radial QMF38889, «9.4. B.C, Loxocrinus sp. 2. B, lateral view of radial QMF38890, «5.8. C, lateral view of radial QMF38891, «5.8. D, Sagenocrinitid indet., lateral view of weathered crown QMF 38896, x1.7. Proampelocrinus, Spheniscocrinus, because they lack the brachial pairs with alternating muscular and cryptosyzygial articulation. Inclusion of some cymbiocrinids within and exclusion of some ampelocrinids from the Ampelocrinidae requires revision of both families. a revision beyond the scope of this paper. Ampelocrinidae are intermediate to upper tier feeders. adapted to carbonate or clastic sub- strates in equatorial latitudes. Family CALCEOLISPONGIIDAE Teichert, 1954 DIAGNOSIS. Bowl-shaped to cvlindrical cup: thick plates; basals often extended as prongs or spines: | anal; radial facets plenary; arms 5 or more; 2 primibrachs if arms not atomous; iso- tomous branching; brachials cuneate: brachial pairs with alternating muscular and cryptosy- турла! articulation; pinnulate; arms incurling distally when enclosed: no tegmen; stem sub- pentagonal or pentagonal proximally, commonly becoming circular distally; cirriferous near cup where known. GENERA INCLUDED. Calceolispongia, Alloso- crinus, Jimbacrinus. Metacalceolispongia gen. nov. REMARKS. The calceolispongiids are best characterised by their thick plates, bowl-shaped orcylindrical cup, often with extended prongs on the basals and distally incurled arms. They are a bottom or very low tier feeding animal adapted to a carbonate or clastic environment in equatorial and higher latitudes. NEW PERMIAN CRINOIDS Calceolispongia Etheridge, 1915 TYPE SPECIES. Calceolispongia hindei Etheridge, 1915 from the late Artinskian upper Noonkanbah Formation, Canning Basin, WA; by monotypy. REMARKS. Two major papers by Teichert (1949) and Willink (1979b) described most of the 22 species of Calceolispongia from Australia. Teichert (1949) described 12 species from Western Australia and demonstrated their strati- graphic value in the Carnarvon and Canning Basins. He reported that one of the major ev- olutionary trends of the genus was towards enlargement of the basals which were used for resting on, or anchoring within, the substrate in adult stages. The stem ofthese forms was a tether in the immature stages and so small as to be of no or little functional value in the adult stage (Webster,1990). Willink (1979b) described 7 species, reassigned 2 species questionably assigned to Phialocrinus by Etheridge (1892) to Calceolispongia, and proposed an evolutionary lineage for the E Australian taxa. He based his lineage largely on the basals and 2nd brachials. Although E Australian species include forms with nodose ornamented or thick, slightly to moderately enlarged bulbous basals, they never developed the extremely enlarged bulbous basals typical of the youngest WA species. The oldest WA species are quite small and have slightly to moderately enlarged basals (Teichert, 1949). Willink (1979b) considered that E Australian species evolved separately from the WA species. The earliest forms of both E Australian and WA lineages began in the Sakmarian. It is of more than passing interest that the 8 non Australian taxa of Calceolispongia, known from the Basleo deposits of Timor (Wanner, 1916, 1924, 1937; Marez-Oyens, 1940), and 1 from the Artinskian deposits of peninsular India (Reed, 1928) are all the greatly enlarged basal type. This suggests that the WA, Indian and Timor forms all developed in an interconnected area of the Tethys and were geographically isolated from the E Australian taxa. Willink (1979b) also described in detail the brachial muscle and ligament structure and proposed a 2 dimensional rheophilic feeding fan for species of Calceolispongia with only slightly to moderately enlarged bulbous basals. This was based on a stemmed form supposedly elevated off the substrate. He compared the arms to some modern crinoids noting their similarity of brachial and muscle structure. This same basic arm structure is present, where known, on all 319 species of Calceolispongia, whether the cup is a greatly enlarged or slightly to moderately en- larged basal form. This implies that the feeding strategy of the greatly enlarged basal cup form, living on or partially buried within the substrate, and the slightly to moderately enlarged basal cup form, elevated off the substrate by the stem, was the same. We suggest that the slightly to mod- erately enlarged basal form was not elevated significantly above the substrate. Instead we interpret it to have had a runner-like stem, with the proximal end upturned so that the cup was ina position very similar to, but perhaps slightly higher than, the cup with a vestigal tether stem. The slightly to moderately enlarged and nodose basals of the stemmed form helped hold the cup in an upright or inclined position, preventing over- turning and fouling of the arms in the substrate. The 2nd brachial, where known, of all species of Calceolispongia has protruded nodes to short blunt spines or coarse irregular ridges. This includes the extremely enlarged or slightly to moderately enlarged basal forms of the taxon. Willink (1979b) suggested that these enlarged brachials: 1, added weight to the crown enabling the cup to orient on its side in slower currents; 2, acted as stabilisers if the crown was suddenly bent in strong currents or detached and oriented on it side; and 3, would have produced con- siderable enhancing eddying particulate feeding when the cup was oriented horizontally. We agree with Willink’s analysis of the plate, muscle, and ligament structure of the arms of the calceolispongiids, but, in part, question the functional significance of the nodes, blunt spines, or irregular ridges on the 2nd brachial and feeding model. The added plate material of the enlarged 2nd brachial to the overall weight of the crown would be minimal, as they account for a small percentage of the overall bulk of the crown. In a detached crown the nodes or blunt spines could serve to stabilise the crown by projecting into the substrate, but this would normally require the 2nd brachials of 2 rays. With the nodes protruded into the sediment it would severely limit, if not completely negate, the function of the rest of those 2 arms, limiting the filtration fan to 3 rays. In soft sediment the base of the ambulacral trackways would probably be fouled by burial in sediment. It is very doubtful that the crown commonly survived if detached from the stem. Autotomy of the stalk of Palaeozoic crinoids has been discus- sed by Donovan (1993) and Baumiller (1997). Although the ability to autotomise the stalk developed in late Palaeozoic cladids (Porugassizocrinus, Webster & Lane, 1970; among others), it has not been demonstrated that other taxa could regenerate (he distal stem and reattach, If they autotomised the distal parts of the stem and then grew additional stem proximally, living by grasping or burial attachment of the cirri, it should be recognised hy finding а stem preserved with the cirri їп the grasping or burial position and the distal end autotomised. The only regenerated stems reported from the late Palacozoic are those of a flexible crinoid (Strimple & Frest, 1979) and Lichenaerinus (Ausich & Baumiller, 1993), Use of the nodes or spines to produce eddy currents useful for feeding is considered minimal. Because the arms projecLoulwards away from the cup inte the current, only the proximal part of the arms could have benefitted if the calceoli- spongnds fed in u 2 dimensional or normal parabolic filtration fan posture. The projections on the 2nd brachial of both greatly enlarged and slightly to moderately en- larged basal cups suggest that they served a vital function necessary to either a stemmed existence with the cup barely elevated above the substrate (slightly to moderately enlarged basals, runner type stem) or the cup resting on, or partly buried within, the substrate (greatly enlarged basals, stem vestigal in mature forms). We suggest that the nodes or blunt spines served a dual purpose, as stops and protection, and that the feeding posture was similar in both types of calceoli- spongtid cup. As stops, the nodes and spines prevented the continued rotational movement of the arms. as they opened to feed. On forms with the basals buried within the sediment the nodes or spines could have stopped against the sediment or abut against another organism living in close proximity. The first brachial on C. abundans had to rotate outward 76" from a vertical position for the nodes on the brachial above the outer lig- ament pit to stop against the distal tip of the radial external to the ligament pit on the radial facet. The nodes or spines of the 2nd brachial would have to rotate 135° to abut against the proximal parts of the radials and distal parts of the brachials. With the immovable ligamentary articulation between the Ist and 2nd brachials (Willink, 1979b) it is impossible for the nodes or spines to abut the cup plates. However, they could abut against adjacent organisms and deter MEMOIRS OF THE QUEENSLAND MUSEUM predators mpping at the base of the arms m cups with the basals resting on the substrate or cups barely elevated above the substrate on à runner stem. In either of these positions the proximal brachials would be elevated above (he substrate and the arms spread in a narrow to widespread filtration Гай or they could have spread into a subhorizontal feeding position with pinnules elevated into the current, Teichert (1949, pl. 8. fig. 2) illustrated the oral view af a crown of C. abundans with the proximal part of the enrolled portion of the arms unrolled along the bedding surface. The first 2 brachials are buried in the sediment at a high angle to the more distal brachials, with only the distal part of the 2nd brachial partly visible at the base of the arms. The cup is completely huried. We interpret this specimen to have been in the feeding posture at the time of rapid burial followed by death. If the specimen died before burial, any current (such as that necessary to bury the specimen in sand) would have moved the arms 1nto a semi-aligned postion, if not broken them off. This suggests that the enlarged basal calceolispongiid species, with the cup resting on or buried within the sediment, Fed with the arms extended along the substrate rather than above the substrate up in the current in à 2 dimensional or normal parabolic filtration fan posture. The enrolled arms in the nonfeeding position prevent- ed sediment from fouling the inactive ambulacral trackways and removed the arms from the paths of predators and benthic scavengers. This. also explains the relatively large space in the muscle area between the radial and Ist brachial facets (Willink, 1979b, fig 11) in the relaxed and flexed positions. Most cladid erinoids have relatively small spaces between the radial and Ist brachial facets when closed (musele contracted) and larger spaces when open (muscle relaxed), because the muscle area of the radial facet is subhorizontal. The facet of the radial is steeply downflared into the cup of Calceoli- spongia, while the brachial facet 1s subhorizontal with the arm in the enrolled contracted position, not the relaxed position as shown by Willink. In the relaxed position the arm is unrolled and extended out into the feeding position with the ligament in the outer ligament pits of the radial and Ist brachial contracted. In both conditions the cross-sectional area between the muscle areas of the facets is relatively larger than in most poteriocrininids. When closed the arms ofthe calecolispongiids NEW PERMIAN CRINOIDS 321 FIG. 23. Calceolispongia abundans Teichert, 1949, oblique C ray (A), distal (B) and B-C interray (C) views of partial crown QMF38874, x2.3. formed an open pentagonal petaloid structure in distal view, not the tightly enclosed crown of most poteriocrininids. However, the 1st and 2nd brachials were more tightly enclosed, surround- ing the visceral mass protruded above the radial summit in the tegmen. The base of the enrolled arms formed a constriction, the arm girdle (Lane & Webster, 1966), above the 2nd brachial. Another use of the node or spines on the 2nd brachial could have been to deter settlers from using the shelf developed atop the 2nd brachial and deny predators access to the visceral mass. Calceolispongia abundans Teichert, 1949 (Figs 23, 24A-C) MATERIAL. QMF38874 from QML 1217. REMARKS. The partial crown, an immature specimen (24mm long, 33mm maximum width), lacks the stem, tips of the protruded part of 4 basals and all distal parts of the arms beyond the m L3 H3 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 24. A-C, Calceolispongia abundans Teichert. 1949. basal, posterior and A ray views of partial crown QMF38874, х2.3. D,E, Calceolispongia rubra Teichert, 1949. proximal and distal facet views of 2nd brachial QMF38876, x4. 1. NEW PERMIAN CRINOIDS 323 4th brachial. It was transported to the site of burial with loss of the stem and perhaps some of the distal parts of the arms. At the site of depos- ition some of the Ist to 4th brachials were disarticulated and moved into the visceral cavity as the specimen was buried. The silt and sand enclosing the partial crown and the associated disarticulated plates are compacted and slightly cemented. Most of the enclosing matrix is easily FIG. 25. A-D, Calceolispongia rubra Teichert, 1949, distal facet (х4), exterior (х4), proximal facet (x4) and lateral (х3.2) views of 2nd brachial QMF38878. removed by wetting, brushing with a moderately stiff tooth brush and light scraping with a needle under the microscope. Features not previously described on C. abundans are the ridges and grooves of the lateral ends of the Ist and 2nd brachials. The Ist ridge and groove are 1.7mm above the proximal end of the Ist brachial and the last is at the distal end of the 2nd brachial. Initial ridges and grooves are vey to A short and of low amplitude, whereas distal ones have higher amplitude. With the brachial in a living or near vertical position, the ridges and grooves are oriented nearly horizontally with a very gentle curvature, centrally convex upward. There are 7 ridges on the Ist brachial in 2.8mm, and 6 ridges in 2.7mm on the 2nd brachial. In lateral view, with the arms enclosed, they appear to interlock with false symplectic articulation. Development of the ridges and grooves on the ends of the proximal brachials is known in other species of Calceolispongia, such as C. lizziensis (Willink, 1979b, pl. 9, fig. 18) and C. spectabilis (Teichert, 1949, pl. 19, fig. 40), as well as poterio- crinitoids such as Spaniocrinus, as described above. Most commonly the apparent interlocking occurs between apposing brachials and may extend along most ofthe arms of adjacent rays, such as in Parastachyocrinus. In these forms alternate brachials form the groove and ridges of | arm and the brachials of the adjacent arm form the counterparts to produce the interlocking fit. Webster & Lane (1967) referred to this as an ‘interlocking structure’ and interpreted it as adding strength to the enclosed crown. They also considered that this limited the movement of the proximal brachials below the arm girdle or constriction in the arms occurring immediately above the interlocking on the cromyocrinid Moapa- crinus. The interlocking of adjacent brachials is structurally quite different from the ridge and groove development on individual brachials of Calceolispongia, although they may have served a similar purpose. We suggest the structures in the calceoli- spongiids served as sliding guides for differential rotational movement of adjacent proximal brachials as the arms were opening and closing as well as guides for forming a close fit when the arms were entirely extended and were being tightly enclosed. A rotational movement of the arms above the transverse ridge ofthe radial facet occurred as the arms opened and closed. Thus gently curved ridges and grooves allowed a relatively smooth differential movement between laterally adjacent plates. The crinoid could flex | arm to a greater or lesser degree than the 2 adjacent arms. When the arms are flexed to a position where the Ist and 2nd brachials were not in lateral contact with the brachials of adjacent rays (Teichert, 1949, pl. 9, figs 1, 2), the ridges and grooves of one arm were not in contact with those of adjacent rays, but would have served as guides for proper positioning of the brachials as the arms enclosed. MEMOIRS OF THE QUEENSLAND MUSEUM In the enclosed position the ridges and grooves would have also restricted translational move- ment of the 2 proximal brachials parallel to the long axis of the arms. This would allow tensional forces (such as predators nipping the tips of the arms or accidentally hitting the arms while chasing prey) applied to the distal parts of the arms to be mitigated by the added strength of the 2 adjacent arms on the proximal 2 largest brach- ials. Depending upon the amount of tension, this could have resulted in retention of the 2 proximal brachials, the largest in the arm, when the distal brachials were lost. Retention of the proximal part would require regeneration of fewer brachials and thus less time and energy to replace the lost part of the food gathering network. The enlarged proximal 2 brachials surrounded an expanded visceral cavity. We suggest that they served as a plated structure surrounding the tegmen in the enclosed position, while also serving as moveable arm bases. To accomodate expansion ofa full gut tract, the 2 proximal brachials would have rotated outward. The ridge and groove structure would have allowed smooth expansion and contraction with a slight rotational move- ment of the proximal brachials. The nodes or blunt spines on the 2nd brachial would have served for protection analogous to tegmen spines. In the feeding position, the opened arms would have allowed some exposure of the visceral mass above the radials, covered only by the tissue of the tegmen. Teichert (1949) described the nervous system of Calceolispongia as a series of fine canals on the inner side of the cup plates. He did not men- tion the dual entoneural canals of the brachials. From the radials there are 4 canals that pass into the first brachial on the oral side ofthe transverse ridge of the arm facet. On the distal crypto- syzygial facet with the 2nd brachial the 4 canals continue into the inner side of the 2nd brachial and as they come out on the distal side they merge into 2 canals continuing throughout the more distal brachials. The 2 very small entoneural canals are on the oral side of the centre of the transverse ridge of facets with muscular articulation and at the growth centre of brachials with crypto- syzygial articulation. First and 2nd brachials (QMF38875-38878) of C. rubra Teichert, 1949 from the Wandagee Sandstone (QML 1222) show the canals (Fig. 24D,E, 25, 26A-D). The dual internal canals also occur in the brachials of Jimbacrinus bostocki. In J. bostocki they pass across the radial facet into the 1st brachial on the oral side of the transverse ridge. In the distal NEW PERMIAN CRINOIDS 325 FIG. 26. A-D. Calceolispongia rubra Teichert, 1949. A.B, distal and proximal facet views of first brachial OMF38875. 4.4. C.D, proximal and distal facet views of second brachial QMF38877, x4.]. E-G, Calceo- lispongia gerthi Willink. 1979b, basal, posterior and D ray v iews of reconstructed crown GSQF 13488, x1.5. 326 brachials thereafter, there are dual canals as in Calceolispongia. Calceolispongia gerthi Willink, 1979b (Fig. 26E-G) MATERIAL. GSQF13488a-r, 5 basals (13488a-e), 5 radials (13488fj), 4 first brachials (13488k-n), and 4 second brachials (134880-r) from the early Artinskian Berridale Formation, Rathbone's Quarry, Granton, Tasmania. Collected by S. Parfrey. REMARKS. Willink (1979b) based the descript- ion of Calceolispongia gerthi on disarticulated second brachials. He recognised that the basals were not distinguishable from those of C. diemen- ensis and described and illustrated basals referred to as C. gerthi-diemenensis. No cups or crowns were reported for either species. All ofthe material came from the Crinoidal Zone on Maria Island. The discovery of disarticulated plates from a single specimen in a thin shale between 2 lime- stone layers in the Berridale Limestone allowed the reconstructon ofa partial crown including the first 2 brachials. Among the 5 basals, only the CD basal has a truncated distal end for reception of the anal plate, which was not recovered. Under the microscope the sutures of the basal and radial plates along the anal interseries were shorter than those between comparable plates in other inter- rays. Likewise, facets between the radials and Ist brachials of the C and D rays were shorter than those in other rays, and had a short gap on the end adjacent to the anal interray, allowing recognition of their positions within the cup. Facets of the Ist and 2nd brachials of the C and D rays were shorter and slightly twisted compared to those in other rays allowing easy recognition. The rest of the specimen was reconstructed around the C and D rays. The BC and DE basals are broken, but the pieces were glued together. Unfortunately the BC basal was also distorted by compaction and the DE basal was solution weathered. It is uncertain if they are in the correct positions or interchanged on the reconstruction. The AB and EA basals would not fit the sutures for the BC and DE, and are thus presumed to be in their correct positions. The third 1st brachial could fit on the E, A, or B radials, and thus its position, though correct with- in a ray, cannot be more precise. Likewise the 2 second brachials have been solution weathered and it is uncertain which, if either, fit the Ist brachial. The infrabasal circlet was not recovered. The reconstructed partial crown is cylindrical and probably rested on the enlarged basals on the MEMOIRS OF THE QUEENSLAND MUSEUM substrate with a runner or tether stem. The proximal side of the blunt spine projections is essentially horizontal, and the infrabasal circlet is not visible in side view, as it is downflaring or subhorizontal and confined to a shallow basal concavity. The radials are subvertical and the protruded knobs of the 2nd brachials are sub- horizontal to slightly upflared in the enclosed position. When the arms were fully extended laterally the knobs would have rested against the apex area of the subjacent radial and 2 basals. If C. gerthi lived elevated slightly above the substrate, the basal projections could have served as bumpers to keep the crown in a near upright position when tilted by currents or scavengers. The blunt spines on the basals along with the knobs of the 2nd brachials would help to prevent the arms getting into the sediment and ambulacral trackways from getting fouled when hit by scavengers or strong current surges. From the shape ofthe reconstructed cup itis most likely that C. gerthi rested on the substrate with a runner stem. Calceolispongia sp. (Fig. 27) MATERIAL. One slab with 3 basals, anal, and numerous brachials, all external moulds, QMF39012 from QMLS18. DESCRIPTION. Plates thickened, basals pro- truded into short double spine projecting out and upward, surface ornament coarse nodes and an- astomosing ridges aligned towards spine tips. Anal pentagonal, with central rounded node. Brachials cuneate, concave longitudinally, strongly rounded transversely; ambulacral groove wide, deep V- shaped. REMARKS. The plates of Calceolispongia sp. are part of a single specimen disarticulated by currents or scavengers leaving them in close association. Other plates are unknown, either not exposed or lost by weathering. The basals are similar in shape to C. gerthi and C. diemenensis (Willink, 1979b), except they have surface ornament in addition to the double spine. If these plates are from a fully grown individual, the specimen would have been small, cylindrical, and lived on the substrate with a runner stem. It probably represents a new species, based on the nodose to anastamosing ridge ornament, that evolved from C. gerthi or C. diemenensis. Lacking radials and proximal brachials it is left in open nomenclature and mentioned for complete- ness of the Condamine fauna. NEW PERMIAN CRINOIDS 327 FIG. 27. Caleeolispongia sp., overlapping views of dislocated cup and arm plates, QMF 39012, 2.5. Jimbacrinus Teichert, 1954 TYPE SPECIES. Jimbacrinus bostocki Teichert, 1954. from the Artinskian Cundlego Sandstone of WA; by original designation. Jimbacrinus donnellyensis Webster & Jell. 1992 Jimbacrinus donnellyensis Webster & Jell, 1992: 333, fig. 21. REMARKS. The locality for J. donnellvensis was givenas from the upper part ofthe Artinskian Bulgadoo Formation in the type section near Donnelly's Well (Webster & Jell, 1992), Because this is one of the few limestone environments above the Callytharra Formation in the Permian of WA the locality was deemed worthy of reinvestigation for additional information and more precise location. The yellow weathering limestone is a small lense, forming a weak bench at the base of the slope. QML1141. Collection yielded 45 crowns of J. donnellvensis (OMF39151-39195), 3 crowns of Stomiocrinus merlinleighensis (QMF39196-39198). 7 basals of Calceolispongia sp. (QMF39210-39216), 8 columnals of Neocamptocrinus sp. (QMF39202- 39209), 2 radials of Thaumatoblastus sp. (QMF- 39200, 39201) and 1 spine of Archaeocidaris? sp. (QMF39199), All specimens are in loose blocks or occur as free elements. Metacalceolispongia gen. nov. TYPE SPECIES. Cymbiocrinus cherrabunensis Webster & Jell, 1992 from the Wuchiapingian Cherrabun Member, Hardman Formation, WA. ETYMOLOGY. Greek meta, between or change, and Calceolispongia, implying a derived form. DIAGNOSIS. Bowl-shaped cup, thick inflated plates, apical pits, 1 anal, plenary radial facets, 2 primibrachs, 11 arms minimal incurl distally, brachials cuneate, muscular and cryptosyzygial paired brachials, pentagonal proximal columnals. REMARKS. Metacalceolispongia differs from all other calceolispongiids by having more than 5 arms. It is similar to Cymbiocrinus, differing by having more than 10 arms, thick plates and a pentagonal stem proximally. It could have evolved from 4//osocrinus by further branching of the arms or from Calceolispongia by branch- ing ofthe arms and less protrusionof the basals. Metacalceolispongia cherrabunensis (Webster & Jell. 1992) Cymbiocrinus cherrabunensis Webster & Jell, 1992:351, fig. 20. DESCRIPTION. See description of Cymbiocrinus cherrabunensis Webster & Jell (1992: 351). REMARKS. Webster & Jell (1992) commented that the 3rd arm on the C ray of C. cherrabunensis is atypical of Cymbiocrinus and noted the affinities to Jimbacrinus. Evaluation of Cymbio- crinus and the Calceolispongiidae within the Articulata lineage resulted in reassignment. Family TRIBRACHYOCRINIDAE Arendt & Willink, 1981 DIAGNOSIS. Cup globose. relatively large: infra- basals upflared, distal tips visible in lateral view: radial facets plenary; 1-4 anals; 12 or 20 arms: isotomously branching: cuneate brachials ramu- late: brachial pairs with alternating muscular and MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 28. Tribrachyocrinus corrugatus Ratte, 1885, B ray view of crown Z3256, x2.3. cryptosyzy gial articulation; stem relatively large, round in transverse section, with round lumen; may be very cirriferous close to cup. GENERA INCLUDED. Tribrachyocrinus, Mega- notocrinus and Nowracrinus. REMARKS. T7ribrachyocrinus was assigned to the Sundacrinidae on the basis of arms developed in 3 rays (Moore & Laudon, 1943) and made the monotypic type of the Tribrachyocrinidae by Arendt & Willink (in Arendt, 1981). Meganotocrinus was questionably assigned to the Ampelo- crinidae and Nowracrinus was considered incertae sedis (Willink, 1979b). Except for the non-development of arms in 2 rays and a larger number of anals there is little difference between Tribrachyocrinus and Meganotocrinus or Nowracrinus. Nowracrinus differs from Mega- nolocrinus by stellate crenulations extending across plate boundaries. The tribrachyocrinids differ from all other Ampelocrinida by the develop- ment of ramules instead of pinnules and the large transversely circular stem. They were an NEW PERMIAN CRINOIDS 329 intermediate or upper tier feeder. adapted to carbonate and clastic substrates in higher latitudes. Tribrachyocrinus M Coy. 1847 TYPE SPECIES. Tribrachyacrinus clarkii M'Cov. 1847 from Roadian or Wordian sediments m ihe Maitland district. NSW: by original designation. Tribrachyocrinus corrugatus Ratte; 1885 (Fig. 28) MATERIAL. TMZ3256, from the Malbina Formation. late Artinskian, Storm. Bay Sheet 8311, 1: 100.000, end relurence 773 348, Tasmania. Collected by Andrew Rozefelds, Max Bunks, and Noel Kemp. DESCRIPTION. Crown small, 42mm long. 26,6mm wide, all cup plates with coarse corrugate ornament, Part of infrabasals. basals, A-C radials. radianal. 3rd anal. parts of A and C rav arms preserved in flattened plane. Radial facet plenary, First primibrach wedge-shaped. tapering distally, Axillary 2nd primibrach triangular. distal tip separating facets for widely diverging Ist secundibrachs. Axillary 2nd or 3rd secundi- brach, shaped like axillary primibrachs. Both branchings isotomous. All distal brachials cuneate. uniserial straight to weakly concave longitudinally. strongly convex transversely. with smooth external surface. Large ramule or small arm on every 2nd tertibrach, alternating sides of main arm for next 4-6 brachials, un- known thereafter. In A ray first ramule on 3rd tertibrach, 2nd ramule on 5thtertibrachon 1/2 ray adjacent to C ray. In 1/2 ray adjacent to D ray Ist ramule on 2nd tertibrach. 2nd ramule on Sth tertibrach. Thereafter ramules every other terti- brach. Brachials paired. musculature articulation оп branching facets and after ramules alternating with cry ptosyzy gial. REMARKS. The small crown is crushed by com- paction, It is. the first crown of this species known with well-preserved arms of 2 rays. The descript- ion of the arms supplements the excellent cup analysis and description of 7: corrugatus by Willink (1979b). Tribrachyocrinus granulatus Etheridge. 1892 (Fig. 29) MATERIAL. TMZ3258. from the Мала Formation, late Artinskian. Storm Bay Sheet 8411. 1: 100.000, grid veftrence 773 348, Tasmania. Collected by A. Rozefelds. M. Banks and М Kemp REMARKS. The cup has a 1.8mm diameter hole drilled through the left central part of the large FIG. 29. 1892. C ray view of cup with drill hole in таШапаї. 221258. *2,6 Tribrachvocrinus granulatus Fiheridge, A-C basal plate in the centre of the flattened specimen, Non-predatory drill holes inthe tegmen plates of Early Carboniferous camerate crinoids were described by Bauiniller (1990) and epizoan pits on cup. tegmen and arm plates of early and middle Palacozoic crinoids were reported by Brett (1978, 1985), To our knowledge. no predatory drill holes through cup plates of Palaeozoic crinoids have been reported. The hole was there prior to deposition. is filled with the surrounding fine silty clay matrix and has a compaction fracture through the plate crossing the right edge ofthe hole. Burial was rapid as no overgrowths or abrasion occurred. Solution weathering has destroyed part of the surface calcite of most exposed plates. Scavengers have not dis- articulated the cup plates and the degree of articulation of the parts of the arms present can- not be determined because of matrix cover. A small. gastropod. Peruvispira sp.. 3.7cm away from the cup. is not known to be carnivorous (А. Cook. pers. comm., 1998) and is not thought to have been associated in a coprophagous relationship with the specimen. Willink (1979b) considered T. granu/atus uninterpretable and recommended suppression MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 30. Tribrachyocrinus? sp. arm fragment, A-C, lateral view of whole specimen (* 1.7), enlarged proximal (x2.4) and distal (x3.1) views of QMF38898. of the species because no additional material had been found. He also considered 7. granulatus as possibly representative of T. rattei Willink, 1979. If such were proven 7. granulatus would have precedence. 7. rattei has aligned nodes coalesc- ing into ridges. 7. granulatus has only nodes, which may show alignment, therefore we accept both species. Tribrachyocrinus? sp., arm fragment (Fig. 30) MATERIAL. QMF38898 from QML 806. DESCRIPTION. Arm large, 60mm long (incom- plete), branching of indeterminate type on 6th brachial, may branch again distally. Brachials large, (proximal brachial 3.2mm long, 6.7mm wide, 7mm deep), uniserial, moderately cuneate, concave longitudinally. strongly convex trans- versely, large ramule given off on every other brachial on opposite sides of arm (2 ramules per 4 brachials). Articular facet of proximal brachial with large rounded transverse ridge; narrow crescent-shaped outer margin with deep ligament pit adjacent to transverse ridge; muscle areas large, narrowing distally. Ramulars moderately cuneate, straight to weakly convex longitud- inally, strongly convex transversely. Ramules branching on ramular4 or 5, may branch again on secundiramular 2, and tertiramular 2; may give off small unbranched ramules of pinnular size. Brachials paired with muscular articulation where branching or a ramule is given off alter- nating with cryptosyzy gial articulation where no branching occurs. REMARKS. The branching pattern of the ramules is similar to that of 7ribrachyocrinus corrugatus above the branching on the second secundibrach. The branching of the ramules may represent a specific difference. The branching also resembles that of the Silurian flexible Cholocrinus obesus (Angelin, 1878), the Early NEW PERMIAN CRINOIDS 331 FIG. 31. Meganotocrinus princeps (Etheridge, 1892), lateral view of abnormal crown BME68151, х1.7. Carboniferous cyathocrinitid Barverinus asteris- cus Van Sant, 1964 and flexible Onyvchocrinus exsculptus Lyon & Casseday. 1860. There is no indication of patelloid processes on either brach- ials or ramulars and the articular facet of the Ist brachial is of a form common to many poterio- crinitids. Meganotocrinus Willink, 1979 TYPE SPECIES. Phialocrinus princeps Etheridge, 1892 from Artinskian Muree Sandstone Member, Branxton Formation, NSW; by original designation. Meganotocrinus princeps (Etheridge, 1892) (Fig. 31) MATERIAL. BME68151, Middle Permian, from an m- known locality in Queensland or NSW. REMARKS. A request for loan of the type specimen of Poteriocrinites smithi resulted in not only the plasticine type of P. smithi, but, the external mould of a partial crown of Meganoto- crinus princeps with an identification label of Р. smithi, from the Gympie Beds, Stanwell, near Rockhampton. The specimen is part of the Dunstan Collection, purchased by the British Museum, July 1935. There is obviously a mixup in the locality and identification of the specimen. This specimen of M. princeps is an external mould of parts of 3 rays of an abnormal cup and proximal brachials, with associated, but not attached, distal parts of 3 or 4 arms. It is em- bedded in a volcaniclastic matrix. Permian strata in the Stanwell area are the Early Permian Youlambie Conglomerate and early Late Permian Dinner Creek Conglomerate. Permian volcaniclastic deposits are present in and SE of the Rockhampton area, about 25-45km E of Stanwell. Other Permian volcaniclastics and sedimentary deposits are present 20-30km W and SW of Stanwell. Thus the specimen could be from the Stanwell or Rockhampton area. All reported localities of M. princeps are in NSW (Willink, 1979b). The abnormality occurs on one of the radials. which lacks the development of the arm and terminates in a distally projected wide V-shaped extension. The Ist primibrachs of the 2 adjacent arms partly overlap the radial. the one onthe right more than that on the left. There is no indication of a radial facet, nor of any injury. This appears similar to B and E radials in 7ribrachyocrinus where no arm is developed. If the 3 basals were not exposed the specimen would have been assigned to 7ribrachyocrinus. It is interpreted as a genetic defect and is illustrated to show the abnormality, but may also indicate the close FIG. 32. Nowracrinus ornatus Willink, 1979, A, distal facet view of primibrachial QMF39020, «2.8. B, internal view of radial QMF39015, x2.4. C, external view of basal QMIE39014, x3.4. relationship between 7ribrachyocrinus and Meganotocrinus. Nowracrinus ornatus (Etheridge, 1892) (Figs 32. 33) Tribrachyocrinus ornatus Etheridge, 1892: 94, pl. 19. Nowracrinus ornatus (Etheridge); Willink, 1979a: 124, figs 3-6f. MEMOIRS OF THE QUEENSLAND MUSEUM MATERIAL. Basal, QMF39014, radial QMF39015 primi- brachs, QMFE39019, 39020 and columnals QMF39016- 39018, 39021, 39076, 39077 trom QML 1247. DESCRIPTION. This description only adds or alters that of Willink (1979a). Pluricolumnal heteromorphic, pentagonal in transverse section. Noditaxis N3231323 minimal, may be more complex. Columnals large, pentagonal nodal 10. Imm diameter. c. Imm thick: internodals sub- round to pentagonal, weakly pentastellate, 8.5mm diameter, «0.7mm thick. Nodal latus narrow, strongly protruded. relatively sharp, rounded: internodal latus similar. Facets with narrow crenularium parallel to pentagonal sides of columnal, crenulae and culmina coarse, straight sided, slightly longerat angles of columnal, other- wise equal length. Areola narrower to slightly wider than crenularium. Lumen large, subcirc- ularto pentagonal, parallel to outline of columnal. Symplexy articulation. Nodals with 5 cirri. Cirral facet elliptical, long axis parallel to columnal facets, with small central axial canal. REMARKS. Combination of the intraplate cren- ulations and nodose to vermiform ornamentation are the distinctive features of cup plates of N. ornatus. Columnals lack surface ornament but the pentagonal and pentastellate outline com- bined with distinctive facets could be used for correlation in absence of the cup plates. All columnals are slightly to moderately distorted from compaction. Family INCERTAE SEDIS Tasmanocrinus Willink, 1979 TYPE SPECIES. Tasmanocrinus mariensis Willink, 1979, from Sakmarian strata on Maria Island, Tasmania; by original designation, Tasmanocrinus sp. (Fig. 34) MATERIAL. TMZ3259, from the Malbina Formation, late Artinskian, Storm Bay Sheet 8411, 1:100,000, grid reference 773 348, Tasmania. Collected by Andrew Rozefelds, Max Banks, and Noel Kemp. DESCRIPTION. Crown cylindrical, 24.5mm long, 9mm wide (incomplete, plates slightly dis- associated). Cup conical, 3.7mm long, crushed. Radials 4 or 5. 3.7mm long, 3.2mm wide. subvertial longitudinally, gently convex trans- versely, proximal end weakly convex, distal end with peneplenary radial facets. Arms 10? Brachials cuneate, strongly convex transversely, with open V-shaped ambulacral groove. NEW PERMIAN CRINOIDS 333 FIG. 33. Nowracrinus ornatus Willink, 1979. A,B, overlapping views of crushed pluricolumnal, QMF39018, x2.8. C, facetal view of columnal QMF39077, x4.2. D, facetal view of columnal QMF39021, «4.2. E, facetal view of slightly disarticulated pluricolumnal QMF39016, х3.3. Е, facetal view of slightly disarticulated pluricolumnal QMF39017, x3.4. G, facetal view of columnal QMF39076, х4.4. Cryptosyzygial and muscular articulation on noded on petals, with strongly convex latus. Ситі alternating pairs of brachials distally. Pinnules close to cup, probably 5 per nodal. Cirrals short. stout. Stem pentagonal, 4mm attached to cup, ound transversely, with convex latus. 36mm unattached, heteromorphic; noditaxis REMARKS. This is the second specimen of Tas- N3231323. Columnals pentalobate, lobed to manocrinus and probably represents a new FIG 34. A.B, Tasmanocrinus sp. lateral. views of partial proximal stem and crown. 73259. »3.3. species, И 15 crushed with plates slightly to moderately disassociated. Weathering and recrystallisation have destroyed facets on most exposed surfaces and ornamentation. A weak line of nodes or granules parallel to the 1niraradial sutures may be the remnant of coarser nodes simular to the aligned nodes on 7. mariensis Willink. 1979. The elongate conical cup was possibly eryptodicyclic with the basals and MEMOIRS OF THE QUEENSLAND MUSEUM infrabasals not visible in lateral view whereas the basal circlet formed a visible part of the cup of 7. mariensis. In the radial circlet 3 plates are ex- posed and 2 are discernible through the thin layer of silt and clay covering the opposite side of the specimen. Willink (19792) described Tasmano- crinus as having 3 radials bearing arms. a 4th radial lacking the distal end and a Sth plate as a radial-like anal. This specimen does not provide additional information concerning the un- certainty of the 4th radial, Only 2 of the Ist primibrachs are visible among the somewhat dis- articulated arm plates. Family affinities of Tasmanocrinus are un- certain. Cup shape has affinity with the Corythocrinidae. If there are only 3 arms it could be related to Tribrachvocrinus, but the arms are pinnulate not ramulate. The pentagonal stem, cirriferous close to the cup, is similar to some Ampelocrinidac. Brachial and columnal artic- ulation show affinity with the Articulata and the peneplenary facets are unique within the Ampelocrinida, Order ISOCRINIDA Sicveris-Doreck, 1952 Family ISOCRINIDAE Gislén, 1924 Archaeoisocrinus gen. nov. TYPE SPECIES. Arcliaeoisocrinus occiduaustralis from the middle Artinskian Cundlego Sandstone. Jimba Jinba Station, WA ETYMOLOGY. Greek алле, beginnmg, old. or primitive: and /secrimis. relers to the beginning of the tsocrimds. DIAGNOSIS. Crown small. cylindrical, with arms enclosed. Cup discoidal. cryptodicyclic. infra- basals and basals in deep basal cavity. covered by proximal columnals; radials form base and cup wall: no anal or anal notch in cup. Radial facets plenary: wide gape between radials and Ist primibrach. Pentameral symmetry. Arms 10, branching isotomously on 2nd primibrach: brachials cuncate uniserial. with small dual internal entoneural canals. with | pinmule on long side of brachial above primibrachs. Brachial articulation alternating between oblique muscular and cryptosyzygial, Stem pentalobate: columnals thin. with strongly rounded convex latus. REMARKS. .Irclhaeoisocrinus dilfers from all other isocrinids. based on cups. in that the basals are within the basal cavity, and not visible in lateral view. Archaeoisocrinus isthe oldest genus ofthe Isocrinidae.cvolved from an ampelocrinid. possibly /7alegetocrinus.in the Early Permianby NEW PERMIAN CRINOIDS 33 B FIG. 35, A-D. Archaeoisocrinusocciduaustralissp. nov. 3 lateral (A,B,D)basal (C) views of crown QMF 38879, «5, removal of the anal plate from the cup and re- striction of the basals to the basal invagination. Previously. isocrinids were reported to range from Early Triassic to the Recent. with the Early Triassic specimens consisting of poorly ргеѕегу- ed columnals lacking details of the articular facet (Rasmussen in Moore & Teichert. 1978). The middle Artinskian occurrence of 4. occidu- australis extends the range of the isocrinids back approximately 30 m.y. Archaeoisocrinus occiduaustralis sp. nov. (Figs 35. 36) ETYMOLOGY. Latin occidens, westem, and australis, southem. MATERIAL. HOLOTYPE: QMF38879 from a nest of Jimbacrinus bostocki Teichert, 1954, from the Cundlego Sandstone on Jimba Jimba Station, WA. Slab found by Kevin Davy. Chns Johnston, and ‘Tom Witherspoon and specimen found in preparation by Scott Vergiels. Crown retains the proximal 3 columnals, proximal part of all 10 arms, and medial parts of 2 arms. DIAGNOSIS. As for genus. DESCRIPTION. Crown small, cylindrical, with arms enclosed, 11.5mm long (incomplete), 9.6- 16.7mm wide (13.1mm av. with arms partly opened). Cup discoidal. 0.8mm long, 4.2-4.5mm wide, with deep basal cavity, unornamented, with pentameral symmetry, without anal series in cup. FIG. 36. Archaeoisocrinus occiduaustralis sp. nov., camera lucida drawing of distal facet of 9th secundi- brach showing dual entoneural canals, QMF38879, x20. without anal notch on tip of indeterminate pos- terior radials, cryptodicyclic. Infrabasals? and basals within basal cavity, not visible in lateral or aboral views, covered by proximal columnals. Radials large, 1.4mm long (minimum), 2.6mm wide, strongly convex longitudinally, gently con- vex transversely, with proximal end in basal cavity, with medial part forming base of cup, distally gently upflared. Radial facet plenary, sloping down outward, 68° from horizontal, with large deep outer ligament pit, narrow outer marginal area. Large gape between radial and Ist primibrach. First primibrach 0.8mm long, 2.6mm wide on proximal end, 3.1mm wide on distal end, slightly convex longitudinally, moderately con- vex transversely. Axillary 2nd primibrach large, 1.3mm long, 3.2mm wide, slightly convex longi- tudinally, moderately convex transversely. AII secundibrachs except Ist and 2nd moderately cuneate proximally, less cuneate distally, gently convex longitudinally, strongly convex trans- versely, nearly circular in transverse section, with single pinnule on long side; small dual entoneural canals circular in transverse section, centrally located; ambulacral groove small, V-shaped. First secundibrach rectilinear, 0.9mm long, 2.1mm wide. Second secundibrach small, nearly resorbed?, externally lens-shaped, restricted to middle of arms, 0.2mm long, Imm wide. Third secundibrach cuneate, 1.6mm long on long side. 0.7mm long on short side, 2.1mm wide. Branching isotomous on 2nd primibrach in all rays, 10 arms, no distal branching on preserved arms. Straight muscular articulation between radials and primibrachs. Oblique muscular articulation between axillary 2nd primibrach and Ist secundibrach and between 3rd and 4th secundibrachs. Syzygial articulation between Ist and 2nd primibrachs and between 1st and 2nd, and 2nd and 3rd secundibrachs. Cryptosyzygial and muscular articulation on alternating pairs of brachials distally. Cryptosyzygial facet with faint culmina and crenellae on outer half of facet, MEMOIRS OF THE QUEENSLAND MUSEUM radiating from entoneural canal. Stem pentalobate, facet not preserved. Columnals with strongly convex latus. REMARKS. This delicate specimen is preserved with the arms slightly splayed around a sandstone matrix. The proximalmost columnals are dis- torted, masking the articular facet. Distal parts of the stem and arms are lacking. ACKNOWLEDGEMENTS We sincerely appreciate the support of Don Mackenzie, Alex Cook and Bev Webster in field investigations that recovered many of the spec- imens described herein. Loan of specimens by the Queensland Geological Survey (through Sue Parfrey), Geological Survey of Western Australia, and Tasmanian Museum and Art Gallery are gratefully acknowledged. We are grateful to the managers of Williambury, Wandagee, Cherrabun, Carey Downs and Middalya Stations for access to collecting localities on their properties. Parks and Wildlife Service, granted access for collection of some Tasmanian specimens. Larry Davis pro- vided some references. Paul Avern processed photographs. This project was supported by the National Geographic Society Grant 5982-97. GDW extends his appreciation to Washington State University for granting professional leave and to the Director of the Queensland Museum for use of facilites and office space during prosecut- ion of this project. The reviews of Tom Baumiller and Gary Lane are kindly acknowledged. LITERATURE CITED ANGELIN, N.P. 1878. Iconographia Crioideorum: in stratis Sueciae Siluricis fossilium. (Samson & Wallin: Holmiae). ARENDT, YU. A. 1981. Trekhrukie morskie lilii (Three armed crinoids). Trudy Palaeontological Institute, 189:1-195, (In Russian) AUSICH, W.L & BAUMILLER, Т.К. 1993. Column regeneration in an Ordovician crinoid (Echino- dermata): paleobiologic implications. Journal of Paleontology 67: 1068-1070. BAMBACH, R.K. 1990. Late Palaeozoic provinciality in the marine realm. Geological Society Memoir 12: 307-323. BATHER, F.A. 1893. The Crinoidea of Gotland: Pt. 1, The Crinoidea Inadunata. Kongliga Svenska Vet- enskaps-Akademiens Handlingar 25(2): 1-201. BAUMILLER, T.K. 1990. Non-predatory drilling of Mississippian crinoids by platyceratid gastro- pods. Palaeontology 33: 743-748. 1997. Crinoid functional morphology. In Waters, J. A, & Maples, C.G. (eds), Geobiology of Echino- derms. The Paleontological Society Papers 3: 45-68, NEW PERMIAN CRINOIDS BRETT, C.E. 1978. Host-specific pit-forming epizoans on Silurian crinoids. Lethaia 11: 217-232. 1985. Tremichnus: a new ichnogenus of circular- parabolic pits in fossil echinoderms. Journal of Paleontology 59; 625-635. BROADHEAD, T.W. 1981. Carboniferous camerate crinoid Subfamily Dichocrininae. Palaeonto- graphica Abteilung A 176(4-6): 81-157. DAY, R.W., WHITAKER, W.G, MURRAY, C.G, WILSON, І.Н. & GRIMES, К.С. 1975. Queens- land geology. Geological Survey of Queensland Publication 383: 1-194, DICKINS, J.M., MALONE, E.J. & JENSEN, A.R. 1964. Subdivision and correlation of the Permian middle Bowen Beds, Queensland. Bureau of Mineral Resources Geology and Geophysics Report 70: 1-12. DONOVAN, S. 1993. Contractile tissues in the cirri of ancient crinoids: criteria for recognition. Lethaia 26: 163-169. ETHERIDGE, R. Jr 1892. A monograph of the Carboniferous and Permo-Carboniferous invertebrata of New South Wales. Part II. Echinodermata, Annelida, and Crustacea. Memoirs of the Geological Survey of New South Wales, Palaeontology 5: 65-131, pls 12-22. 1915. Western Australian Carboniferous fossils, chiefly from Mount Marmion, Lennard River, West Kimberley. Western Australia Geological Survey, Bulletin 58: 7-49. GLENISTER, B.E, ROGERS, ES. & SKWARKO, S.K. 1993. Ammonoids. Geological Survey of Western Australia Bulletin 136: 54-63. KAMMER, T. & AUSICH, W.I. 1993, Advanced cladid crinoids from the Middle Mississippian of the east-central United States: intermediate-grade calyces. Journal of Paleontology 67: 614-639. LANE, N.G. & WEBSTER, G.D., 1966. New Permian crinoid fauna from southern Nevada. University of California Publications in Geological Sciences 63: 1-60. LYON, S.S. & CASSEDAY, S.A. 1860. Description of nine new species of Crinoidea from the sub- Carboniferous rocks of Indiana and Kentucky. American Journal of Science and Arts, series 2, 29: 68-79. MAREZ OYENS, F.A.H.W. De 1940. Neue Permische Krinoiden von Timor. Geological Expedition Lesser Sunda Island 1: 285-348. MILLER, J. S. 1821. A natural history ofthe Crinoidea, or lily-shaped animals; with observations on the genera, Asteria, Euryale, Comatula and Marsupites. (Bryon & Co.: Bristol). MOORE, R.C. & JEFFORDS, R.M. 1968. Classific- ation and nomenclature of fossil crinoids based on studies of dissociated parts of their columns. The University of Kansas Paleontological Contrib- utions, Echinodermata Article 9: 1-86, 28 pls. MOORE, R.C. & PLUMMER, F.B. 1940. Crinoids from the Upper Carboniferous and Permian strata 337 in Texas. University of Texas Publication 3945: 1-468. MOORE, R.C. & TEICHERT, C. (eds) 1978. Treatise on invertebrate paleontology. Part T. Echino- dermata 2. 3 vols. (Geological Society of America & University of Kansas: Lawrence, Kansas). PABIAN, R.K., BOARDMAN, D.R., II & HOLTER- HOFF, P.F. 1989. Paleoecology of Late Penn- sylvanian and Early Permian crinoids from north-central Texas. Texas Tech University Studies in Geology 2: 291-303. PABIAN, R.K. & STRIMPLE, H.L. 1993. Taxonomy, paleoecology and biostratigraphy of the crinoids of the South Bend Limestone (Late Pennsylvan- ian-Missourian, ?Virgilian) in southeastern Nebraska and southeastern Kansas. Conservation and Survey Division, University of Nebraska- Lincoln, Professional Paper 1: 1-55. REED, F.R.C. 1928. A Permo-Carboniferous marine fauna from the Umaria Coal-field. Records Geological Survey India 60: 367-398, pls 31-34. 1933. Notes on some lower Palaeozoic fossils from the southern Shan States. Records Geological Survey of India 66: 188-211. SHI, GR. & McLOUGHLIN, S. 1997, Permian strati- graphy, sedimentology and palaeontology of the southern Sydney Basin, eastern Australia. School of Aquatic Science and Natural Resources Man- agement, Deakin University Technical Paper 1997/2: 1-59. SIEVERTS-DORECK, H. 1942. Crinoioden aus dem Perm Tasmaniens. Zentralblatt für Mineralogie, Geologie und Paläontologie, Monatschefte 3: 80-87. SIMMS, M.J. 1988. The phylogeny of post-Palaeozoic crinoids. Pp. 269-284. In Paul, C.R.C. & Smith, A.B. (eds), Echinoderm phylogeny and evol- utionary biology. (Clarendon Press: Oxford). SIMMS, M.J. & SEVASTOPULO, G.D. 1993. The origin of articulate crinoids. Palaeontology 36: 91-109. SIMPSON, С.С. 1961. Principles of animal taxonomy. (Columbia University Press:New York). STRIMPLE, H.L. 1971. A Permian crinoid from Coahuila, Mexico. Journal of Paleontology 45: 1040-1042. STRIMPLE, H.L. & FREST, Т. 1979. Points of gener- ation and partial regeneration of the column of Euonychocrinus simplex (Crinoidea: Flexibilia). Journal of Paleontology 53: 216-220. STRIMPLE, H.L. & MOORE, R.C. 1971. Crinoids of the Francis Shale (Missourian) of Oklahoma. University of Kansas Paleontological Contrib- utions Paper 55: 1-20. STRIMPLE, H.L. & WATKINS, W.T. 1969. Carbonifer- ous crinoids of Texas with stratigraphic implications. Palaeontographica Americana 6(40): 139-275, pl. 30-56. TEICHERT, K. 1949. Permian crinoid Calceoli- spongia. Memoir of the Geological Society of America 34: 1-132. 1954, A new Permian crinoid from Western Australia. Journal of Paleontology 28: 70-75. VAN SANT, J. 1964. Crawfordsville (Indiana) crinoid studies. University of Kansas Paleontological Contributions, Echinodermata Article 7: 1-136. WANNER, J. 1916. Die permischen Echinoderm von Timor, I Teil. Paláontologie von Timor 6: 1-329, pls. 94-115. 1924. Die permischen Echinoderm von Timor, II Teil. Jaarboek van het Mijnwezen in Neder- landsch Oost-Indié 5. 1921, 3: 1-328, 22 pls. 1937. Neue Beiträge zue Kenntnis der permischen Echinodermen von Timor, VII-XII. Palaeon- tographica, Supplement 4(2): 59-212. WEBSTER, G.D. 1974. Crinoid pluricolumnal noditaxis patterns. Journal of Paleontology 48: 1283-1288. 1987. Permian crinoids from the type-section of the Callytharra Formation, Callytharra Springs, Western Australia. Alcheringa 11: 95-135. 1990. New Permian crinoids from Australia. Palaeontology 33: 49-74. 1997, Lower Carboniferous echinoderms [rom northern Utah and wx Wyoming. Paleontology Series. v. 1, Utah Geological Survey Bulletin 128: 1 5. WEBSTER, GD. & HOUCK, K.J. 1998. Middle Pennsylvanian, late Atokan-early Desmoinesian, echinoderms from an intermontane basin, the Central Colorado Paleontology.72: 1054-1072. WEBSTER, GD. & JELL, PA. 1992. Permian echino- derms from Western Australia. Memoirs of the Queensland Museum 32: 311-373. WEBSTER, GD., JELL, РА. & DEREWETZKY, A.N. in press. Palaeobiogeography of Permian echino- derms of Australia. Permian of Eastern Tethys Symposium Volume. Trough. Journal of MEMOIRS OF THE QUEENSLAND MUSEUM WEBSTER, GD., & LANE, N.G. 1967. Additional Permian crinoids from southern Nevada. University of Kansas Paleontological Contrib- utions, Paper 27: 1-32. 1970. Carboniferous echinoderms from the south- western United States. Journal of Paleontology 44: 276-296, pls 55-58. WELLER, S. 1909. Description of a Permian crinoid fauna from Texas. Journal of Geology 17: 623-635. WILLINK, R. 1978. Catillocrinids from the Permian of eastern Australia. Alcheringa 2: 83-102. 1979a. Some conservative and highly-evolved Permian crinoids from eastern Australia. Alcheringa 3: 117-134. 1979b. The crinoid genera Tribrachyocrinus M Coy, Calceolispongia Etheridge, Jimbacrinus Teichert and Meganotocrinus n. gen. in the Permian of eastem Australia. Palaeontographica Abteilungen A 165: 137-194. 1980a. A new coiled-stemmed camerate crinoid from the Permian of eastern Australia. Journal of Paleontology 54: 15-34. 1980b. Two new camerate crinoid species from the Permian of eastern Australia. Alcheringa 4: 227-232. WRIGHT, J. 1937. Scottish Carboniferous crinoids. Geological Magazine 74: 385-411, pls 13-16. 1951. The British Carboniferous Crinoidea. Pal- aeontographical Society Monograph, 1(4): 103-148, pls 32-40. YAKOVLEV, N.N. 1933. Dve verkhnepermskhe mor- skie lilii iz zakavkaziya [Two Upper Permian crinoids from the Transcaucasus]. Izvestiya Akad- emii Nauk SSSR, Leningrad 7: 975-879. (In Russian) 1956. Organism i sreda. Stati po paleoeckologii besnozvonochnykh 1913-1956. [Organisms and surroundings. Writings on paleoecology of invertebrates, 1913-1956]. Akademii Nauk SSSR. 139p. (In Russian) APPENDIX 1 Queensland Museum localities referred to in text. QML518 - Late Artinskian Condamine Beds, S side Lucky Valley Creek, Elbow Valley area, SW of Warwick, SE Qld. Magellan GPS coordinates 28722731"S, 152708 19"E, QML 737 - Late Sakmarian or early Artinskian Callytharra Formation, N side of bladed track from Callytharra Homestead to Byro Homestead in W most exposures 1.5km W of type section S of Wooramel River, WA. 25°52°30"S, 115°29°E. Wooramel Sheet SG50-5, 1966. QML758 - Late Sakmarian or early Artinskian Callytharra Formation, N side of bladed track from Callytharra Homestead to Byro Homestead as track enters dry wash tributary to Wooramel River; lowermost fossiliferous shale and marl capped by limestone in lower fossiliferous unit. Type section of Callytharra Formation. 25°52730"S, 115?30705"E, Glenburgh Sheet SG50-6, 1963. QML759 - Late Sakmarian or early Artinskian Callytharra F ormation, second fossiliferous shale and marl capped by limestone in type section of Callytharra Formation. South side of track and stratigraphically higher than QML758. QML772 - Wuchiapingian Cherrabun Member of Hardman Formation; bench in lower slope below cliff 0.5-1.0km NNW of type section, Millyit Range: GR767877 Crossland Sheet SE51-16, 1977, WA. Magel- lan GPS coordinates 19°10°45"S, 125°32735"E. OML S06 - Late Permian, Flat Top Formation, Back Creek Group; halfway up small rise, 1.3 km E on Uncle Tom road from Leichardt Highway, S of Banana, Queensland. Coll. A. Cook & M. Wade. QML859 - Wandrawandian Siltstone; Point Upright, wavecut platform below lighthouse, Uladulla, NSW. NEW PERMIAN CRINOIDS QML1141 - Middle Artinskian Bulgadoo Shale, upper part of type section SE of Donnelly's Well, Williambury Stn, WA. Magellan GPS coordinates 24°05’45"S, 115"05'40"E. QML1145 - same as QML772. QML1146 - Wuchiapingian Cherrabun Member of Hardman formation; bench in lower slope below cliff, Millyit Range, WA. Magellan GPS coordinates 19°10°28"S, 125°32°26"E. QML1217 - Late Artinskian, basal massive sandstone of the Wandagee Sandstone, exposed in type section along the Minilya River, Wandagee Station, WA. Magellan GPS coordinates 23°44’20"S, 114?25'02"E, QML1232 - Early Artinskian, upper part of Callytharra Formation, E limb of Gooch Range. W side of valley, E facing slope, S of Minilya-Lyndon road, up small drainage just S of where bladed road turns E to cross valley, WA. Magellan GPS coordinates 23°54’22"S, 114^56'54"E. QML1233 - Float in limestone below QML 1232. 339 QML1237 - Early Artinskian, upper part of Callytharra Formation, lower crinoidal rich zone, N side of Minilya- Lyndon Road, flat below slope, 50m SE of ridge, E limb of Gooch Range, WA. Magellan GPS coordinates 23°53’48"S, 114756'48"E, QML1240 - Early Artinskian, upper part of the Callytharra Formation, top of first bench nearest road, N side of Minilya-Lyndon road, E limb of Gooch Range, WA. Magellan GPS coordinates 23°53°41"S, 114°56°35"Е. OMLI247 - Kansas Beds, Early Permian, Artinskian; roadcut and quarry on W side of cut at top of first hill on Blackwells Road, 0.8km W of junction with Highway А10. Blackwells Road is 14km S of junction A10 and B26, Yolla, Tasmania. Coordinates 83.25 35.80 Burnie Sk55-3, 1973. GSWAL119377 - Artinskian, Billidee Formation, 4th lime- stone, Australian Map Grid coordinates Zone 50, 326340E, 7296860N, S of Mt Sandiman sheds. Coll. A.J. Mory. APPENDIX 2 List of described Permian crinoid taxa from stratigraphic units of Western Australia, Queensland, New South Wales and Tasmania. Cherabun Member, Hardman Formation, Wuchiapingian, Neocamptocrinus millyitensis Webster & Jell, 1992 Metacalceolispongia cherrabunensis (Webster & Jell, 1992) WA. Wandagee Sandstone, late Artinskian, WA. Calceolispongia abundans Teichert, 1949 Cundlego Sandstone, middle Artinskian, WA. Archaeoisocrinus occiduaustralis gen. et sp. nov. Billidee Formation, early Artinskian, WA. Dichocrinus? sp. Callytharra Formation, late Sakmarian to early Artinskian, WA. Neocamptocrinus? sp. Glaukosocrinus middalyaensis sp. nov. Parabursacrinus granulatus Wanner, 1949 Timorechinid gen. indet, Poteriocrinitid indet., arms 1 Poterioerinitid indet.. arms 2 Poteriocrinitid indet., arms 3 Loxocrinus booni Marez Oyens, 1940 Loxocrinus sp. 1 Loxocrinus sp. 2 Prophyllocrinus sp. 1 Prophyllocrinus sp. 2 Prophyllocrinus sp. 3 Sagenocrinit id indet. Flat Top Formation, Wordian, Qld. Auliskocrinus? bananaensis sp. nov. Tribrachyocrinus? sp.. arm fragment Condamine Beds, latest Artinskian or early Roadian, Qld. Neocamptocrinus sp. nov. Platycrinites halos sp. nov. Platycrinitid indet., columnals Necopinocrinus tycherus gen, et sp. nov. Eidosocrinus condaminensis gen. et sp. nov. Pedinocrinus? nodosus sp. nov. Stellarocrinid? gen. et sp. nov. Spaniocrinus geniculatus sp. nov. Sundacrinus medius sp. nov. Moapacrinus cuneatus sp. nov. Eoindocrinus praecontignatus Arendt, 1981 Calceolispongia sp. Poteriocrinitid indet., arm fragment 1 Poteriocrinitid indet., arm fragment 2 Poteriocrinitid indet., arm fragment 3 Catherine Sandstone, late Artinskian, Qld. Neocamptocrinus catherinensis sp. nov. Unknown stratigraphic unit, late Artinskian or early Roadian. Old.? Meganotocrinus princeps (Etheridge, 1892) Berridale Limestone, early Artinskian, Tas. Calceolispongia gerthi Willink, 1979 Kansas Beds, early Artinskian, Tas. Order indet., basal and radial plates Nowracrinus ornatus (Etheridge. 1892) Malbina Formation, late Artinskian, Tas. Tribrachyocrinus corrugatus Ratte, 1885 Tribrachyocrinus granulatus Etheridge, 1892 Tasmanocrinus sp. Wandrawandian Siltstone, late Artinskian, NSW. Anaglyptocrinus willinki gen. et sp. nov. 340 A MONASTERID STARFISH FROM THE PERMIAN OF TIMOR. Memoirs of the Queensland Museum 43(1): 340. 1999:— Among a large collection of Permian fossils made by Brad Macurda from the Indonesian island of Timor and deposited in the Museum of Paleontology at the University of Michigan is a fragment (2 arms) of a small starfish, the first record of the group from that island which has yielded the world's most diverse Permian echinoderm fauna. The locality label reads ‘No. 51702, 1 specimen, Permian, Tonino I, Timor’. This locality refers almost certainly to the known Permian locality Tonino (= Toeninoe of Macurda, 1983) Noil (=stream) 1-2km SE of Basleo, SE Timor. Order PUSTULOSIDA Spencer, 1931 Suborder MONOMARGINALINA Kesling, 1969 Family MONASTERIDAE Schuchert, 1915 Genus nov. (Fig. 1) DESCRIPTION. Arms 8mm long, 4mm wide proximally and 2mm deep, upturned strongly at distal tip. Dorsally all plates strongly convex to bulbous: median column of radials or carinals of 4 plates, with distal one greatly inflated and terminating arm; inferomarginal columns enclosing arms laterally and visible in both dorsal and ventral views, each column of 3 convex plates: with 3 interradial plates not inflated, with small axil visible dorsally and ventrally. Ventral surface mainly 2 columns of large adambulacrals; adambulacrals short and wide, becoming narrower distally, transversely convex, in contact along axis so concealing ambulacrals. Mouth frame disarticulated, unclear, REMARKS. Following Kesling’s (1969) review, this specimen fits the concept of the Monasteridae because it has short wide adambulacrals ventrally and dorsally the column of strong dorsal carinals contiguous with the inferomarginals that enclose the arms laterally. The family is known from Australia (Kesling, 1969) and South West Africa (Lane & Frakes, 1970) but all known Australian species are represented by specimens very much larger (at least 5 times) than this Timorese specimen so its morphology is probably that of a juvenile. It is, therefore, difficult to make meaningful comparisons with confidence and makes any formal taxonomic decisions inadvisable. The most distinctive feature of the new specimen is the extremely large terminal radial plate which is highly suggestive of a new genus. It is difficult to imagine such a distinctive feature disappearing with growth and none ofthe known species ofthe family have such an arm termination. However, with only 2 arms of a juvenile and no disc details available I find myself in the same position as Lane & Frakes (1970), unable to name a new taxon based MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 1. Monasteridae gen. nov. UMMP51702, 7.5. A, dorsal view. B, ventral view. on it. The occurrence of monasterids in Timor further strengthens the faunal similarities between that island and Western Australia so evident among crinoids and blastoids (Webster & Jell, 1992). Literature cited KESLING, R. V. 1969, Three Permian starfish from Western Australia and their bearing on revision of the Asteroidea. Contributions from the Museum of Paleontology, at the University of Michigan 22: 361-376 LANE, N.G. & FRAKES, L.A. 1970. A Permian starfish from South West Africa. Journal of Paleontology 44: 1135-1136. MACURDA, D.B. 1983. Systematics of the fissiculate Blastoidea Papers in Paleontology from the Museum of Paleontology at the University of Michigan 22: 1-291 WEBSTER, G.D. & JELL, P.A. 1992. Permian echinoderms from Western Australia, Memoirs of the Queensland Museum 32 311-373. Peter А. Jell, Queensland Museum, Р.О. Box 3300, South Brisbane 4101, Australia; 17 May 1999. A NEW CORNUTE CARPOID FROM THE UPPER CAMBRIAN (IDAMEAN) OF QUEENSLAND ANDREW B. SMITH AND PETER A. JELL Smith, A.B. & Jell, P.A. 1999 06 30: A new cornute carpoid from the Upper Cambrian (Idamean) of Queensland. Memoirs of the Queensland Museum 43(1): 341-350. Brisbane. ISSN 0079-8835. The first Cambrian carpoid from Australia, Drepanocarpos australis gen. et sp. nov. is described from the Chatsworth Limestone in the Lily Creek section at Chatsworth 100km north of Boulia, western Queensland. Its age is the Peichiashania secunda - Prochuangia glabella Zone, latest Idamean Stage (=mid Franconian Stage), in the medial Late Cambrian. It belongs to the cornute Family Phyllocystidae, having cothurnopores, rigidly plated dorsal surface and flexible plated ventral surface, a well-defined marginal frame with ventral vertical strut and dorsal transverse strut and 6 well-defined rings in the proximal part of tail. O Carpoid, Phyllocystidae, Upper Cambrian, Queensland. Andrew B. Smith, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom; Peter A. Jell, Queensland Museum, P.O. Box 3300, South Brisbane 4101, Australia; received 31 August 1998. Carpoids are rare in Upper Cambrian rocks. Ubaghs (1963) described one from a unique specimen and two others from fragmentary material of uncertain generic placement; all 3 came from a single locality and horizon in the early Trempeleauan Whipple Cave Formation in Nevada. Sumrall et al. (1997) featured 6 carpoids from the Upper Cambrian of Wyoming and Nevada with none of them represented by a complete specimen and all known from 1 or 2 specimens or a single slab; the 3 cornutes are assigned specific names in 2 new genera, Acuticarpus and Archaeocothurnus, the other 3 taxa are left in open nomenclature. Ubaghs (1999) described a new genus, Lobocarpus, from the Upper Cambrian of Montagne Noire, southern France. Known Australian carpoids are reported elsewhere in this Memoir (Ruta & Jell, 1999a-e) and derive from the latest Ordovician to Early Devonian clastic sequences of Victoria and Tasmania. However, the Cambrian and great majority of the Ordovician in Australia have yielded no carpoids, and cornutes have never been recorded from Australia. At Museum of Victoria Locality NMVPL1597 (= Bureau of Mineral Resources Locality K204 of Shergold, 1982) in a 4m thick grey micaceous limestone forming a bench on the low limestone rise 3.5km S of Chatsworth Homestead 60km SW of Duchess, W Queensland is a rich silicified fauna of trilobites (Connagnostus sp. undet., lveria iverensis, Lorretina depressa, Peichia- shania secunda, Prochuangia glabella, Pseudagnostus parvus, Pseudagnostus sp. undet.), gastropods, monoplacophorans, brachiopods, hyoliths, sponges and echinoderms. The last mentioned group was partly described by Jell et al. (1985) who treated the ‘eocrinoid’ Riddersia watsonae and noted an isorophid edrioasteroid and by Smith & Jell (1990) who described the isorophids Hadrodiscus parma and Chatsworthia spinosa (=isorophid of Jell et al. 1985) and the edrioblastoid Cambroblastus enubilatus. Details ofthe location are available in Shergold (1982, figs 3,4) who also dated the bed, based on trilobites, as within his Peichiashania secunda - Prochuangia glabella Zone, the youngest within the Idamean Stage. The fossils are coarse silica replacements of extremely fine structures and in no specimen is replacement complete. Unravelling the structure of the species has been achieved by gathering some information from each of 12 available specimens and piecing it together into a picture of the whole animal. This approach necessitates more illustrations and camera lucida drawings than is normal to understand the species. Illustrated material is deposited in the Queensland Museum, Brisbane (QMF) and the Natural History Museum, London (BMNH). Plate lettering, orientation and terminology follow Jeffries et al. (1987) without necessarily entering the debate on the affinities and possible biology of carpoids. 342 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 1. Drepanocarposaustralis gen.et sp. nov., ventral surface uppermost, x9. A, QMF 17862. B, QMF 17860. SYSTEMATIC PALAEONTOLOGY Order CORNUTA Jaekel. 1901 Family ?2PHYLLOCYSTIDAE Derstler, 1979 Drepanocarpos gen. nov. TYPE SPECIES. Drepanocarpos australis sp. nov. ETYMOLOGY. Greek drepanon, a sickle or blade, alluding to the curved marginal frame, and carpos, a fruit - the common name applied to this group. RANGE AND DISTRIBUTION. Upper Cambrian (Idamean = Franconian) of Queensland. DIAGNOSIS. Cornute with body longer than wide, with strong marginal frame and ventral strut; ventral surface composed of a few large plates; dorsal tegmen of many small platelets. Gill slits 4-5, as cothurnopores in the posterior left-hand side in an embay ment of marginal plate k, framed by skeletal elements. A transverse bar on the dorsal surface separates the body into proximal and distal portions. Proximal part of appendage with 6 well-defined rings, distal part narrower. No strongly differentiated stylocone. REMARKS. This genus belongs to the Cornuta on account of its marginal frame and distinctive bipartite appendage. Of the 3 families currently recognised, Scotiaecystidae has a very dis- tinctive gill slit morphology quite different from that in Drepanocarpus and thus need not be considered further. The elongate body form, tesselated dorsal plating and finer platelets of the ventral surface are typical of phyllocystids such as Lobocarpus Ubaghs, 1999. the only named phyllocystid NEW CAMBRIAN CORNUTE FROM QUEENSLAND ?anterior U-shaped plates of U) ь „ә d? cothurnopores e— marginal plate ?posterior U- p shaped plate of cothurnopores FIG. 2. Drepanocarpos australis gen. et sp. nov., camera lucida drawings of ventral surface with plating interpretation (plate lettering follows the system of Jefferies et al., 1987). A, QMF 17862. B, QMF 17860. C, posterior view of the left hand side of QMF 1 7860, dorsal surface uppermost. Cross hatching = sediment. Scale bar = Imm. knownfrom the Cambrian. Note that Ubaghs was equivocal in his family assignment. Lobocarpus differs from Drepanocarpos in having a much more heart-shaped body, with broad, flange-like marginals and a less well-developed central strut. Drepanocarpos differs from the type species of Phyllocystis in having a more asymmetric body, deeper left marginal indentation and in having larger and fewer dorsal plates. With the uncertainty of classification among the early Palaeozoic cornutes some comparisons with ceratocystid and cothurnocystid Cambrian forms are warranted. Drepanocarpos resembles Middle Cambrian Ceratocystis Jackel, 1901 from Europe in its overall shape, although the body is more rounded proximally. The ventral surface is composed of a small number of large sutured plates. these probably being extensions of the marginal plates. MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 3. Drepanocarpos australis gen. et sp. nov., dorsal surface uppermost, х9. A, QMF17861. B. BMNH EE6344. Drepanocarpos differs from Ceratocystis, however. in a number of important details. Most importantly its gill slits are not in the form of sutural pores but rather cothurnopores. Protocystites meneviensis from the Middle Cambrian of Wales (Jefferies et a/., 1987) closely resembles Ceratocystis in shape but has reduced ventral plating and a better defined proximal tail, like Drepanocarpos. It differs from Drepanocarpos in the absence of a sagittal strut on the interior of plates a and g and the very different shape and arrangement of the distal spines and gill slits, which resemble those of Ceratocystis. "Phyllocystis sp.’ and Nevadaecystis americana Ubaghs from the latest Cambrian of Nevada (Ubaghs, 1963), Cothurnocystis ? bifida Ubaghs & Robison from the Middle Cambrian of Utah (Ubaghs & Robison. 1988) and an unnamed cothurnocystid from the Middle Cambrian Spence Shale of Utah, U.S.A. (Sprinkle, 1976) have cothurnopores (oval spout-like openings within the dorsal tegmen) and well-defined rings in the proximal part of the tail. Cothurnocvstis ? bifida is more L-shaped in outline than Drepanocarpos and plate 1 forms the distal left-hand angle of the head carrying a very large process. It also has a ventral tegmen of retiform NEW CAMBRIAN CORNUTE FROM QUEENSLAND c/d Ee cothurnopores: strut on interior of ventral plates B traces of plating FIG. 4. Drepanocarpos australis gen. et sp. nov., camera lucida drawings of dorsal surface with plating interpretation (plate lettering follows the system of Jefferies et al., 1987). A, ОМЕ17861. B, BMNH EE7344. Scale bar = 1mm. stereom, possibly in the form of a continuous sheet of calcified integument. Unlike Drepanocarpos plates a and c are not united by a transverse bar, although this may be a result of poor preservation. Nevadaecystis is laterally elongate and shaped like Cothurnocystis, with similar highly developed lateral blade-like processes on plate 1. Also, like Cothurnocystis but unlike Drepano- carpos, plates a and c in Nevadaecystis are unconnected and the distal border of the buccal cavity lacks plate 4. Like Drepanocarpos it has a ventral surface of large plates, extensions of the marginals, and a dorsal tegmen of much smaller platelets. There is also a saggital strut formed presumably from the internal thickening of two of these plates. Jefferies et al. (1987, fig. 12) interpreted Nevadaecystis as having an additional plate x, like Cothurnocystis fellensis. However, in the reconstruction of Ubaghs & Robison(1988, fig. 11.1) no such plate is shown. The unnamed cornute from the Middle Cambrian Spence Shale in Utah (Sprinkle, 1976) resembles Drepanocarpos in shape and plating arrangement, in lacking obvious appendages to the marginal frame, in having a dorsal tegmen of small platelets, in having a smoothly rounded right margin and in having cothurnopores across 346 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 5, Drepanocarpos australis gen. et sp. QME]17803, x8. C, BMNH EE6345, «8. FIG. б. Drepanocarpos australis gen, et sp. nov.. camera lucida drawings of specimens in dorsal view showing cothumopores. A. QMF 17866. B, QMT 17863. C. BMNH EE6345. Scale bar = Imm. NEW CAMBRIAN CORNUTE FROM QUEENSLAND 347 FIG. 7. Drepanocarpos australis gen. et sp. nov. A, QMF 17867 in dorsal aspect, x 10. B, QMF17865 in dorsal aspect showing fully plated ventral surface, x10. C, BMNH EE6346, in ventral view, x12. the posterior edge running between plates | and k. Unfortunately the ventral plating is unknown and we do not know whether plates a and c are united to form a transverse bar behind the buccal area as in Drepanocarpos. Archaeocothurnus Sumrall et al., 1997 from the Middle and Upper Cambrian of Utah and Nevada, and including 'PAyllocystis" (Ubaghs, 1963). differs from Drepanocarpos in that plate gis reduced to a narrow strut and there is a plated dorsal tegmen. This genus is thus more closely comparable to Cothurnocystis. Acuticarpus Sumrall et al. (1997) from the Upper Cambrian of Wyoming is distinguished by its triangular shape. the more slender marginal plates, presence of a stylocone and lack of spines along the thecal margin. Poor understanding of the anterior of the theca makes comparison more difficult. There are some similarities, indicating affinity. between Drepanocarpos and Hanusia Cripps, 1989 from the Ordovician of Czechoslovakia. particularly in possession of the spike on plate 1. However, that genus has an l-spike and an e-spike, very limited extension of marginals onto the ventral surface, virtually straight right thecal margin, elongate thecal shape, and probable cothurnopores on dorsal surface. In summary Drepanocarpos appears to be distinct from Ceratocystidae and Cothurno- cystidae and bears most resemblance to the Phyllocystidae in particular Lobocarpus and Sprinkle's (1976) Spence Shale stylophoran. We thus make tentative assignment to the Phyllocystidae in line with Ubaghs (1999) caution in assigning Lobocarpus. Drepanocarpos australis sp. nov. (Figs 1-9) ETYMOLOGY. Latin australis, southern, the only comute so far known from Australia. MATERIAL. Holotype, QMF17860, paratypes QMFT17861-17871 and BMNH EE6344-6346. DIAGNOSIS. As for genus. DESCRIPTION. Body up to 10mm long and 7mm wide. Appendage more than 10mm long in an individual with a 5.5mm long body (Figs 8С, 10A) generally shorter, abruptly truncated. Ventral surface flat, with left laterodistal margin deeply embayed at plate a, with right margin uniformly convex. Proximal margin slightly embayed close to the appendage. The appearance of individuals varies with the extent of weathering. In better preserved specimens distal plates b and c appear as curved processes resembling appendages (Figs 1A, 2A, 3, 4), while in the holotype these 2 processes are connected distally, forming a continuous margin- al frame (Figs 1B, 2B). Marginal frame of 7 plates possibly, with 2 dorsal and 2 ventral plates above and below the tail. Plate k embayed dorsally for cothurnopores; cothurnopores as narrow slits surrounded by plating (Figs 4, 5) or as deep notches (Fig. 2), partially underlain by a narrow ledge. Plate 1 with a short pointed spike not much longer than the thickness of the marginal frame. Plate boundaries on the marginal frame are very difficult to make out, especially anteriorly because of the coarseness of silicification and because the anterior is available in only 2 specimens. Plates a and c/d apparently abutting to form a transverse strut across the body. Plate c/d (these 2 plates are inferred but no separating suture can be discerned with certainty) long and curved, tapering distally, with short lateral bar distally; one specimen (Fig. 1B) suggests that a short lateral bar connects to plate b across the distal extremity of the body via a small plate. We do stress that this interpretation of a single specimen requires verification. Distal processes not present. Plates c/d, e and f forming a continuous smooth curved right-hand margin. Plate f with an internal notch proximally. Dorsal plating best seen from the interior. Suture separating plates a and g evident; suture defining a V-shaped region towards the right. Other sutures not discernible; dorsal surface of a few large plates sutured together but the precise pattern of plate sutures is unknown. No large calcite plates are preserved distal to the trans- verse bar formed by plates a/c. Ventral surface largely lost; small plates occasionally seen inside marginal frame may be the remains of a plated tegmen. Cothurnopores 4 or 5, in the proximal angle of the head close to the marginal frame in the embayment of plate k (Figs 5, 6). Fine structure masked by preservation, apparently a row of closely spaced oval mounds, each with an irregular surface sometimes showing a median depression. MEMOIRS OF THE QUEENSLAND MUSEUM plate lost during acid etching ventral plates tegmen plates ? FIG. 8. Drepanocarpos australis gen. et sp. nov., camera lucida drawings of QMF17865 with plating interpretation. A, dorsal view. B, proximal view. Scale bar = 1mm. Appendage of a thicker proximal part of 6 well-defined rings and a thinner distal part (whose structure is not well seen in any specimen). No specimen well enough preserved to show the stylocone. REMARKS. Although 12 specimens of this species are known, none is sufficiently well-preserved to show all of the characteristic features. The reconstruction (Fig. 9) has therefore had to be composite. In particular plate sutures are often difficult to identify due to the coarse silica replacement, and sutures are only indicated if they appear consistenly in a number of specimens. The different extent of weathering also poses a problem of interpretation. However, enough is known about this species to show that it merits separation at generic level from other cornutes as discussed above under the generic heading. The most closely related cothurnocystid is the unnamed solute from the Spence Shale, Utah (Sprinkle, 1976, pl. 1, fig. 1) which differs from D. australis in having a slightly broader head, no spike on plate l, a longer and better developed plate i and no apparent connection between plate a and c (though this may be a preservational artefact; the connection is not seen either in QMF17860). Furthermore, the Spence Shale cothurnocystid has a narrower, less blade-like right-hand margin. NEW CAMBRIAN CORNUTE FROM QUEENSLAND VENTRAL plating unknown — possible tegmen internal strut formed by plate thickening 349 tegmen of small plates- organization! unknown tegmen of small plates assumed — organization unknown tegmen plates ? VIG.9. Possible reconstruction of Drepanecarpas australis ger. et sp. nov. We stress thal inlerplate sutures are hot always clearand their representation with dashed lines indicates this uncertainty. The sketch is provided as the best estimate, though by по means certain, of the skeleton from available material. ACKNOWLEDGEMENTS We are grateful to Dick Jefferies. Natural History Museum. London and Jim Sprinkle, University of Texas. Austin for strong reviews, acknowledging that neither referee necessarily agrees with all or any of the conclusions. LITERATURE CITED CASTER, KE., 1907. Homoiostelea, Pp, 581-623, In Moore, R.C, fed.) Treatise on invertebrate paleontology. Part S, Echinodermata 1. (Geological Society of America & University of Kansas: New York). CRIPPS, A.P, 1989, A new genus of stem chordate (Comuut)from the Lower and Middle Ordovician of Czechoslovakia and the origin of bilateral symmetry in the chordates. Geobios 22: 2 13-245, DERSTLER, К. 1979. Biogeography of the stvlophoran carpoids (Echinodermata), Pp. 91-104. In Gray. J. & Houcot. A. (eds) Historical biogeography. plate tectonics and the changing environment (Oregon State University Press: Corvallis). JAEKEL, O.. 1901. Ueber Carpoiden, eine neue Klasse von Pehnatozoen. Zeitschrift der Deutschen geologischen Gesellschaft 52: 661-677. JEFFERIES, RPS., 1986. The ancestry of the vertebrates. (British Museum (Natural History): London). JEFFERIES, RPS., LEWIS. М, & DONOVAN. S.K.. 1987. Protacystites menevensis — a stem-group chordate (Comuta) trom the Middle Cambrian of south Wales. Palacontology 30: 429-484, JELL, PA. BURRETT. C.F & BANKS, M.R.. 1985. Cambrian and Ordovigian echinodenns from eastern Australia, Alcheringa 9: 183-208, RUTA, M. & JELL. P.A.1999a. Protoeytidium gen nov. a new anouralocystitid: mitrate from the Victorian latest Ordovician and evolution of the Allanievüdiidae. Memoirs of the Queensland Museum 43: 353-376. 1999b. Jdokeracarpusgen. nov..a new mitrate [rom the Ludiovian Kilmore Silistone and Lochkovian Humeyale Formation of central Victoria Memoirs of the Queensland Museum 43: 377-398. 199090. Two new anomalocystiitid milrales trom the Lower Devonian Humevale Formation ol central Victoria. Memoirs of the Queensland Museum 43: 390-422, [999d. A nole on Fictoriacystis wilkinsi (Anomalovystitida, Mitratay from the Upper Silurian of Victoria. Memoirs of the Queensland Museum 43: 423-130, 19996, Revision of Silurian and Devonian Allanicytidudae. (Anomalocystitida, Mitrata) from southeastern Australia. Tasmania and New Zealand. Memoirs ol the Queensland Museum 43; 431-451. SHERGOLD. LIL. 1982. Late Cambnan trilobites from the Chatsworth Limestone, western Queensland 350 Bulletin of the Bureau of Mineral Resources Geology and Geophysics Australia 186: 1-111. SMITH, A.B. & JELL, P.A. 1990. Cambrian edrioasteroids from Australia and the origin ofthe starfishes. Memoirs of the Queensland Museum 28: 715-778. SPRINKLE, J. 1976. Biostratigraphy and paleoecology of Cambrian echinoderms from the Rocky Mountains. Brigham Young University Geology Studies 23(2): 61-73. SUMRALL, C.D., SPRINKLE, J. & GUENSBURG T.E. 1997. Systematics and paleoecology of Late Cambrian echinoderms from the western United States. Journal of Paleontology 71: 1091-1109. MEMOIRS OF THE QUEENSLAND MUSEUM UBAGHS, G. 1963. Cothurnocystis Bather, Phyllocystis Thoral and an undetermined member of the Order Soluta (Echinodermata, Carpoidea) in the uppermost Cambrian of Nevada. Journal of Paleontology 37: 1133-1142. 1969. Les échinodermes carpoides de l'Ordovicien inférieur de la Montagne Noire (France). (Cahiers de Paléontologie, éditions du Centre National de la Recherche Scientifique: Paris). 1999, Echinodermes nouveaux du Cambrien supérieur de la Montagne Noire (France Meridionale). Geobios 31: 809-829. UBAGHS, G. & ROBISON, R.A. 1988. Homalozoan echinoderms of the Wheeler Formation (Middle Cambrian) of western Utah. University of Kansas Paleontological Contributions Paper 120: 1-17. ABERRANT PANDANOCRINUS ABERRANT PANDANOCRINUS, EARLY DEVONIAN CRINOID FROM NORTH QUEENSLAND. Memoirs of the Queensland Museum 43(1): 351. 1999:- Pandanocrinus martinswellensis Jell et al., 1988 was described from the Early Devonian (Pragian; sulcatus Biozone) Martins Well Limestone Member of the Shiels Creek Formation at Martins Well on Pandanus Creek Station, north Queensland. Collections from the type locality in the University of Queensland (UQL3579) and the Queensland Museum (QMLS50) include several hundred individuals with uniform plate arrangement except for QMF25736, which has 7 plates in the radial circlet and a small triangular plate at the D-E interray between basal and radial circlets. BB 3, equal, 2 with 3 sides against RR, 3rd with 4 or 5. BB circlet 7-sided; sides are not even either in length or in angle of meeting. Orientation is difficult because free arms and most of the upper theca are missing, only a few fixed brachials and interbrachials preserved. 7-sided anal plate bisected by diameter with one of the interplate sutures in the basal circlet indicating A ray-CD interray line of symmetry. Other interplate basal circlet sutures in C & D rays; in normal specimens the sutures are in the A, C and E rays (Jell et al., 1988). With this orientation (Fig. 1B) A, B and C rays appear normal although the upper margin of the B radial is not symmetrical. D & E rays are irregular in plate arrangement and shape. D & E radials appear to be divided. each into 2 plates of comparable size; thus the 7 radials. Between adjacent D and E radials (i.e. 1 from each pair) and resting on the basal circlet is a small triangular plate which has no homologue in other crinoids. If the D radial is correctly interpreted as being 2 plates then the anal is not in contact with the C radial as in normal specimens. Moreover, there appear to be 2 plates in the CD interray in the Ist brachial circlet; thus producing a circlet of 11 rather than the normal 10 plates. Apart from the more numerous plates the shapes of most posterior plates from the C to E rays are irregular. This is clearly a unique specimen. The most obvious explanation involves traumatic injury as a juvenile: growth centres of several plates were fragmented and 2 plates grew where formerly there had been 1. In radials, division was fairly equal whereas the small triangular plate must have had only a tiny piece of growth centre to start it off. Alternative explanations could involve a growth response to a constricted growth position (e.g., in a crevice or, given the number of fossils at the site, in a tightly packed meadow of large crinoids) or a single mutation. The chance of 1 mutation dividing 4 or more growth centres is remote and dismissed here. The influence of the surroundings could affect overall shape and symmetry but is unlikely to have induced growth of entirely separate plates. С FIG. 1. Pandanocrinus martinswellensis Jell et al., 1988. А. basal view of aberrant theca QMF25736, «1.5. B, plate diagram of QMF25736 Literature Cited JELL, P.A., JELL, LS, JOHNSON, B.D.. MAWSON, К. & TALENT, J.A. 1988. Crinoids from Devonian limestones of eastern Australia, Memoirs of the Queensland Museum 25(2) 355-402 Peter A. Jell, Queensland Museum, P.O. Box 3300, South Brisbane 4101, Australia; 17 May 1999. ws) un кә THE CRINOID MELOCRINITES TEMPESTUS IN THE DEVONIAN CAMPWYN BEDS ON THE SHORE OF REPULSE BAY, SOUTH OF PROSERPINE. Memoirs of the Queensland Museum 43: 352. 1999:- Along the western shores of Repulse Bay, just south of Proserpine, the Campwyn Voleanics (Fergusson et al., 1994) extend from around Seaforth in the south to near Lethebrook in the north (Paine, 1972). About midway between Midgeton and Lethebrook. in the vicinity of the Laguna Quays Resort (termed Aqua del Rey in Fergusson et al, 1994), the Campwyn Volcanics outcrop in a broad intertidal band more than 100m wide. Strike is virtually parallel to the waterline and dip is to the west at very low angles. Muddy volcanoclastic limestones predominate with the percentage bioclasts highly variable between beds. Ina few of these beds crinoidal and other shelly (including coral and brachiopod) debris is common; sections of stem up to 100mm long and 10mm in diameter are common. Cup and arm plates are rare. In 1991 Mr Bob Spencer forwarded the crinoid calyx figured herein to the Queensland Museum after having collected it in the intertidal area now disturbed by excavations for the Laguna Quays Resort; he also sent a badly weathered specimen of a large pleurotomariacean gastropod and a solitary rugose coral. Subsequent collection by the author has confirmed the provenance but failed to discover any further identifiable crinoids. This crinoid (Queensland Museum Fossil 40967). with a high conical, smooth cup, 52mm long, is identified as a very large specimen of Melocrinites tempestus Jell et al., 1988. This assignment is based on the shape, arrangement and ornament of cup plates and cup shape. The specimens figured by Jell et al. (1988) are mostly weathered smooth but one (Jell et al, 1988. fig. 13G) shows the corners of cup plates depressed indicating that the unweathered cup had strongly convex plates as in the Campwyn Volcanics specimen. The age of the Campwyn Volcanics has been given as Upper Devonian based on corals determined by Hill from near Seaforth (Jensen et al., 1966; McKellar in Roberts et al., 1971) and Lower Carboniferous based on brachiopods from the Mackay sheet, determined by McKellar (Jensen et al., 1966). In the Proserpine sheet area the age has been given as Upper Devonian based on the Seaforth corals (Paine, 1972) or on conodonts in samples from the Laguna Quays locality (Fergusson et al.. 1994, fig. 3b). However. the crinoid species determined herein has previouslv only been found in the Middle Devonian (Givetian) part of the Papilio Formation of the Broken River Province. Melocrinites is widely recorded from the Silurian to Frasnian in the Northern Hemisphere and Australia but is only recorded from the Famennian in New York. The only Australian Frasnian record is in the Canning Basin reef complex (Jell & Jell, 1999). Since the conodont age suggested for this locality is Famennian (R.A. Henderson pers. comm. 1999) there appears to be some conflict with the crinoidal indication. This single crinoid could be a long ranging species providing a second Famennian occurrence for the genus or far less likely the section at Laguna Quays could represent a longer period of time than previously thought. MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 1. Melocrinites tempestus Jell et al.. 1988, lateral view of cup QMF40967, «1.5 Literature cited FERGUSSON, C.L., HENDERSON, R.A. & WRIGHT, J.V. 1994 Facies in a Devonian-Carboniferous volcanic fore-arc succession, Campwyn Volcanics, Mackay district, central Queensland, Australian Journal of Earth Sciences 41: 287-300 JELL, Р.А. & JELL, I.S. 1999. Crinoids, a blastoid and a cyclocystoid from the Upper Devonian reef complex of the Canning Basin, Western Australia. Memoirs of the Queensland Museum 43: 201-236 JELL, P.A., JELL, J.S., JOHNSON, B.D.. MAWSON, R. & TALENT, J.A. 1988. Crinoids from Devonian Limestones of eastern Australia. Memoirs of the Queensland Museum 25: 355-402. TENSEN, A.R., GREGORY, C.M. & FORBES, V.R. 1966. Geology of the Mackay 1:250000 sheet area, Queensland. Bureau of Mineral Resources, Geology and Geophysics Report 104:1-58 PAINE, A.G.L. 1972. Proserpine Qld. Sheet SF/55-4. 1.250000 Geological Series - Explanatory Notes (including map) 24p. ROBERTS, J., JONES, P.L, JELL, J.S., JENKINS, T.B.H., MARSDEN, M.A.H., McKELLAR, R.G., McKEL VEY, B.C & SEDDON, G. 1971. Correlation of the Upper Devonian rocks of Australia. Journal of the Geological Society of Australia 18(4): 467-490 Peter A. Jell, Queensland Museum, Р.О, Box 3300, South Brisbane 4101, Australia; 26 May 1999. PROTOCYTIDIUM GEN. NOV., A NEW ANOMALOCYSTITID MITRATE FROM THE VICTORIAN LATEST ORDOVICIAN AND EVOLUTION OF THE ALLANICYTIDIIDAE MARCELLO КОТА AND PETER A. JELL Ruta, M. & Jell, Р.А, 1999 06 30: Protocytidium gen. nova a new anomalocystitid mitrate from the Victorian latest Ordovician and evolution of the Allanicytidiidae. Memoirs uf the Queensland Museum 43(1): 353-376. Brisbane. ISSN 0079-8835. Theanomalocystitid mitate Prorocyridium elliottae gen. et sp. nov. is described from the up- permost Ordovician Darraweit Guim Mudstone in central Victoria, Protacytidium has: 1) [2 plates on the convex surface, with only 3 in distal row and the remaining 9 as in Ёпор{онги; remarkably asymmetrical lateral and median orifice plates of plano-concave surface divided into a ‘thecal’ portion framing the body orifice and a ‘lip’ projected distally beyond the ori- fice: 3) much reduced plate B failing to contact right LOP; 4) narrow, elongate plate A sutured with proximal 2/3-3/4 of medial margin of left intermediate lateral marginal plates; 3) short, slender, almost straight left spine and longer. more robust, convex, sickle-shaped right spine; 6) cancellate to honeycomb-like stereom, often replaced by radiating trabeculae near periphery of plates; 7) styloid trapezoidal, bearing semicircular styloid blades with radi- aling trabeculae; 8) proximal blade about half as large as distal blade. Protocytidium is intermediate between Enoploura popei and the basal allanicytidiid Occu/tocystis koeneni. The Allanicytidiidae are reviewe ed i in the light of this new find. O Protocytidium, Anomaloeystitida, Allanicytidiidae, Bolindian, Victoria. Marcella Ruta, Department of Palaeontology, The Natural Flistory Museum, Cromwell Road, London SW7 SBD, United Kingdom; Peter A, Jell, Queensland Museum, P.O, Box 3300, South Brishane-4101, Australia; 10 June 1998. During field mapping of the Kilmore 1:50,000 sheet (Vandenberg, 1992), which included a study of Ordovician/Silurian boundary sections in the Deep Creek and Ben Dhui Creek areas ( Vandenberg et al., 1984), numerous specimens of a small mitrate were discovered at NMVPL660 in Ben Dhui Creek (Vandenberg et ul. 1984, figs 1. 2A). This species has a combination of skeletal features making it transitional between Enoplaura Wetherby, 1879 from the Middle ta Late Ordovician of North America (Caster, 1952: Parsley, 1991) and the allanicytidiid Occultocystis koeneni Haude, 1995 from the Lower Devonian of Argentina. The new taxon is the oldest and only Ordovician member of the Allanicytidiidae. In this paper we describe and reconstruct the taxon, assess its phylogenetic position and discuss ils bearing on the origin and evolution of the Allanicytidiidae. GEOLOGICAL SETTING The material described herein comes trom NMVPL660 (Vandenberg et al., 1984, fig. 2А. appendix 2). It is in the bed of Ben Dhui Creek, 750m N of the Wallan to Woodend road. about LOkm W of Wallan at AMG311740-5860360 on the Kilmore 1:50,000 sheet 7823-11 Series R 754, Edition |-AAS (1979). It is in the type section of the Darraweit Guim Mudstone which consists of medium to thick hedded massive calcareous mudstone. The mudstone is black when fresh but surface exposures are usually decalcified, weathered and grey or green as at NMVPL660. The associated fauna. which is preserved in situ, includes the trilobite Songxites darraweitensis (Campbell). the graptolite Climacograptus angustus Perner and nautiloids. This is a thin unit (20m in the type section) precisely dated by the widespread graptolite Climacograptus? exiraordinirius (Sobolevskaya) which occurs in the youngest Ordovician Zone at the lop of the Bolindian Stage. SYSTEMATIC PALAEONTOLOGY Anatomical terminology and plate nomenclature ( Appendix) follow Ruta (in press). Ruta & Bartels (1998, fig. 5А,В) and Ruta & Jell (19992,b.c). Specimens are housed in the Museum of Victoria, Melbourne (NMVP) wherein the locality is also registered (NMVPL). All photographic illustrations are of latex casts taken from decalcified moulds and whitened with ammonium chloride sublimate. 354 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. L Protoevtidium elliontae gen. et sp. nov. A, inside of proximal part of convex surface and distal part of plano-concavesurface of NMVP100439. «5. В, С, partial plano-concave surface and detail of adjacent isolated spine (probably right spine) of NMVP100424. х5 and +7, respectively. C, fragment of marginal plate from plano-concave surface showing stereom fabric. NMVP100405, ¥11. D, partial plano-concave surface and appendage. NMVP100401, «5.5, I. partial plano-concave surface of NMVP100415, х9, F, fragment of marginal plate from plano-concave surface showing stereom fabric. NM VP 100400. x11. A NEW VICTORIAN ANOMALOCYSTITID MITRATE 35 Class STYLOPHORA Gill & Caster, 1960 Order MITRATA Jaekel, 1918 Suborder ANOMALOCYSTITIDA Caster, 1952 Family ALLANICYTIDIIDAE Caster & Gill, 1967 DIAGNOSIS (modified from Caster & Gill, 1967; Caster, 1983; Haude, 1995; Ruta, in press). Median orifice plate (MOP) longer than each lateral orifice plate (LOP). Spines longer than distal margin of plano-concave surface. Distal part of convex surface of 3 or, more frequently, 2 plates with transverse thickening along inside of their distal margins. Distal styloid blade inclined proximally, sometimes with lateral ear-like projections. Sharp, longitudinal keel on external surface of styloid, failing to reach free margin of proximal blade (only in basal allanicytidiids). Proximal blade semicircular. REMARKS. Ruta (in press) amended and expanded the diagnosis of the Allanicytidiidae, formalising Haude's (1995) proposal to include Occultocystis koeneni. As discussed below, some characters supporting the sister-group relationship between the new anomalocystitid and the Allanicytidiidae sensu Haude (1995) are diagnostic of the latter (Ruta, in press). Our diagnosis is, therefore, more generalised than those of Haude (1995) and Ruta (in press) and aims to avoid the problem of defining the Allanicytidiidae mainly on the basis of the simplified plating of the convex surface in the most derived representatives of the group (Caster, 1956, 1983; Caster & Gill, 1967; Philip, 1981; Ruta & Theron, 1997; Ruta & Jell, 1999c). However, reversal of some of the characters listed above occurs to some extent within the Allanicytidiidae. Protocytidium gen. nov. TYPE SPECIES. Protocytidium elliottae sp. nov. ETYMOLOGY. Greek proto, the first and cytidium, a small box. Neuter. DIAGNOSIS. Body subelliptical to pyriform, rarely subrectangular. Convex surface of 12 plates, with 3 in distal row. Medial and lateral orifice plates of plano-concave surface asymmetrical, divided into a 'thecal' portion framing the body orifice and a ‘lip’ projected distally beyond the orifice. Plate B reduced, not in contact with right lateral orifice plate. Plate A narrow, elongate. Left spine short, slender, almost straight; right spine longer, more robust, Un convex, sickle-shaped. Stereom cancellate to honeycomb-like, often replaced by radiating trabeculae near periphery of plates. Styloid trapezoidal, with semicircular blades with radiating trabeculae; proximal blade c.1/2 as large as distal blade. REMARKS. The phylogenetic position of this new genus is discussed after the specific treatment below but simple distinguishing features are as follows: it differs from all anomalocystitids in the arrangement of the median and lateral orifice plates of the plano-concave surface, shape of the spines and elongate plate A with tiny plate B. Enoploura has 5 plates in the distal row on the convex surface, larger C21, different ornament and less expanded styloid. Occultocystis has C21 isolated from the proximal body margin, very few plates in the convex surface and C11 and C13 as marginal plates. Most of the advanced genera of the Allanicytidiidae are distinguished by having only Cl and C5 in the distal row on the convex surface, by having straight spines and more elaborate surface ornament. Protocytidium elliottae sp. nov. (Figs 1-14) ETYMOLOGY. For Tracey Elliott of the Palaeontology Department at the Natural History Museum, London. MATERIAL. Holotype: NMVP100401. Paratypes: NMVP100390-100400, 100402-100403, 100405- 100406, 100408-100419, 100421-100436, 100438- 100439, 100487-100488 all from NMVPL660. DIAGNOSIS. As for genus. DESCRIPTION. EXTERNAL. Measurements. Holotype (Fig. ID): 11mm long, 6mm wide. Largest specimen (Fig. 7C): 14mm long, 7mm wide. Plano-concave surface (Figs 1A-B,D-E, 2B-D, 4A-B, 6D, 7A,D, 8A, 9B, 100, 13A). Mostly flat, except for slightly raised lateral margins, with lateral marginal plates divided more or less equally into subhorizontal and vertical parts. Lateral body walls of uniform depth, except for rapid tapering near proximal and distal ends (Fig. 13C-E). PM usually 1.5 times as wide proximally as distally, with straight to concave lateral margins on either side of slight lateral projection just proximal to midlength, with proximal margin strongly embayed for insertion of appendage. PLM with subhorizontal part subtriangular, with convex lateral margin and straight or broadly concave distal margin. ILM as long as PLM, 2 Un on MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 2. Protocytidium elliottae gen. et sp. nov. A, distal view of distolateral corner of body NMVP100391, x12. B, partial plano-concave surface of NMVP100397, x10. C, plano-concave surface incomplete distally. NMVP100410, x10. D, partial plano-concave surface of NMVP100403, х7. A NEW VICTORIAN ANOMALOCYSTITID MITRATE 337 FIG, 3, Proteevtidium elliottae gen. et sp. nov. A, convex surface of NMVP 100488, +7. B. distolateral corner of mostly disarticulated plano-concave surface of NM VP100391. «11.5, C. partial proximal convex surface with inside of plano-concave surface in background, NM VP100390, х6. D, inside of plano-concave surface with C21 still in position and with abapical surface of styloid in lower right, NMVP100417. «11.5. m un о MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 4. Protocytidium elliattae gen. et sp. nov. A. proximal appendage and partial plano-concave surface of NMVP100396, <8. В, proximal plano-coneave surface of NMVP100408, *10. C. inside of plano-concave surface of NMVP100413, «12. A NEW VICTORIAN ANOMALOCYSTITID MITRATE subrectangular, with medial margin of right ILM sigmoidal and medial margin of left ILM similarly sigmoidal distal to its junction with the A-C suture and proximally straight, with lateral and distal margins straight. DLM as long as ILM, subrectangular to subtrapezoidal, with shallow distal surface deepening laterally, with subcentral tubercle for spine articulation. LOP and MOP remarkably asymmetrical, at least twice as long as wide, divided into proximal 2/3 articulated with rest of plano-concave surface, and distal 1/3 projecting beyond distal margin of transverse orifice, usually giving rise to irregular process. Irregular gaps between distal parts of adjacent margins of orifice plates. Right LOP generally slightly shorter and narrower than left LOP or MOP. Plate B subtrapezoidal (based on shape of available plate margins), sutured to left LOP, MOP, A and C. Plate А at least 5 times longer than wide, oblique to longitudinal body axis, sometimes remarkably shortened, with medial margin either gently convex or strongly convex in distal 1/2 and gently concave to straight in proximal 1/2, with pointed proximal wedged between C and left ILM. Plate C widening proximally. in contact with MOP. Convex surface. Al plates except C21 and C3 arranged in pairs (indicated by plate margins and by mirror image arrangement of medial and lateral plates sutured with C20-C22); distal 1/3 of convex surface invariably disrupted and poorly preserved. C21 with width 40% of length, shield-shaped, with slightly convex longitudinal and transverse profiles; margins very gently convex except straight to weakly concave proximal margin, extreme proximolateral section also with short concave section, producing proximolateral projection. C20 and C22 subtrapezoidal, with maximum width proximally; transverse profile of plates strongly arcuate, especially near their proximo-lateral angle, with thickening immediately distal to proximal margin, with blunt proximo-lateral angles, with medio-distal angles proximal to latero-distal angles. C15 and C19 irregularly pentagonal, with greatest dimension at 45° to body axis, with proximo-medial margins in contact with latero-distal margins of C21, with sigmoidal medio-distal margins. C11 and C13 longer than wide, irregularly hexagonal, sutured along straight median suture. C6 and C9 subtrapezoidal, smallest plates of convex surface, lateral to C11 and C13, respectively. Distal 1/3 of convex surface poorly preserved in available material, but apparently consisting of 3 large plates in a transverse row (7 C1, C3 and C5 based on comparison with Enoploura and Occultocystis). C5 (Fig. 3A) subrectangular, slightly longer than wide. C3 slightly longer than C5, subpentagonal. Body stereom and sculpture. External surface of coarse stereomic fabric forming radial pattern on each plate. Terrace-like ridges and riblets absent. Stereom of thick, generally straight, sometimes bifurcating trabeculae often connected by irregularly spaced, short transverse bar-like connections; trabeculae and connections delimit deep pits and become more irregular and randomly branching near plate margins, where the pits are smaller, more numerous and polygonal. Trabeculae of proximal 1/2 of convex surface generally thicker than elsewhere. C20-C22 (Figs 3A,C-D, 5C) with trabeculae almost parallel to each other, rarely bifurcating, gently arcuate or straight near lateral and medial margins of C20 and C22, becoming confluent near centre of proximal 1/2 of external surface of both plates, where they become slightly thicker and irregularly sinuous, with trabeculae on distal 1/2 thinner, subparallel to body axis, converging to centre of plates proximally. C21 with broadly arcuate trabeculae separated by granular fabric on proximal 2/3, with distal trabeculae straight, bifurcating, in fan-like arrangement. Remaining body plates with gradual changes in stereom structure from centre to periphery (Figs | A-B,D-E, 2C-D, 4B, 7A-B,D, 8A-C, 9B, 10D); centre compact or honeycomb-like, with short, anastomosing trabeculae; more peripheral stereom of elongate trabeculae with transverse connections; surface with cancellate or honeycomb-like fabric proportional to plate size. Orifice plates with shallow pits and poorly defined, flat-topped trabeculae or coarse and without obvious surface pattern. Proximo-distally elongate trabeculae near proximal and distal margins of PLM, ILM and DLM (Fig. 1F-G). External surface of A апа B (Fig. 1D) with large, widely spaced polygonal pits delimited by thin trabeculae. PM with honeycomb fabric, with radiating peripheral trabeculae, especially along distal margins. Spines. Left spine straight or gently convex externally, round in cross-section, with clavate proximal end, with hemispherical articular surface, tapering distally, with blunt terminus. Right spine slightly longer and twice as wide as left spine, sickle-shaped, flattened except for 360 MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 5, Protocytidium elliottae gen. et sp. nov. A. styloid and proximal part of plano-concave surface of NMVP100424, х8. В, inside of plano-concave surface of NMVP 100423, x12. C, partial convex sulace with distal row removed to show inside of orifice plates of NMVP100398, #12. D, proximal appendage and proximal plano-concave surface of NMVP100396. «12. A NEW VICTORIAN ANOMALOCYSTITID MITRATE 361 NG. 6. Protecytidium elliettae een, et sp. nov. А. inside of plano-concave surface of disarticulated NMVP100398, «8. B, styloid of NMVP 100430, * 12. C, inside of plano-concave surlace of NM VP 100487, «7, D. plano-concave surface of NMVP 100394. «9 P D һә MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 7, Protocytidium elliottae gen. et sp. nov. A. plano-concavesurface showing orifice platesof NMVP 100402. “10. B, proximal parts of plano-concave surface and appendage of NMVP100414, «10. C. interior of plano-concave surface but with proximal part concealed by C20-C22 still in place of convex surface. NMVP100403. +7. D, plano-concave surface with orifice plates intact, NMVP100395. x9, A NEW VICTORIAN ANOMALOCYSTITID MITRATE being conical distally, with sharp lateral and blunt medial margins, tapering distally to point, with subconical articular projection with 2 facets separated from short subcylindrical region by poorly pronounced ridge, with stereom of minute, irregular, shallow pits separated by coarsely granular to compact texture. INTERNAL. Plano-concave surface. Internal surface of orifice plates divided into 2 unequal parts and different depths by transverse, distally concave thickening (Figs 7C, 8D, 9D, 11D, 12D). Internal surface of left latero-distal angle of body occupied by subtriangular area straddling suture between left DLM and left LOP and medial to tubercle for spine insertion on left DLM. Proximo-lateral 1/2 of subtriangular area slightly deeper and more raised than medio-distal 1/2 and delimited laterally and proximally by 2 ridges, continuing distally on internal surface ofleft LOP where the proximal, thicker ridge (pr in Fig. 12A) almost parallels the body axis, with the thinner ridge (lar in Fig. 12A) widening rapidly in its distal 1/2 before giving rise to thickened lateral margin of internal surface of left LOP, with central part of subtriangular area occupied by pit (p in Fig. 12A). Proximal and lateral ridges merging and continuing as a low sinuous ridge (lr in Fig. 12A) coinciding with junction between subhorizontal and vertical parts of plate. Distal 1/4 of low ridge with triangular lateral process (Ip in Fig. 12A) partially delimiting shallow subelliptical area (sa in Fig. 12A). Low ridge of right side apparently fading gradually distally near medio-distal angle of internal surface of right ILM, immedialely lateral to point where the oblique septum (see below) straddles the suture between C and right DLM and abruptly changes direction. Oblique septum (se in Fig. 12C) with asymmetrical transverse section running from distal right angle to proximal left angle of inside of plano-concave surface (Figs 3C-D, 4C, 5B, 6A,C, 7C, 8D, 9A,C-D, 10C, 11A, 12C), with proximal 1/3 straight or gently convex to left, with proximal end merging into left scutula (sc in Fig. 12C). Left scutula transversely elongate (Figs 3D, 6A,C, 9C). Straight, transverse ridge (tr in Fig. 12C) (sensu Ruta & Jell, 1999a) delimited proximally by transverse furrow (tf in Fig. 12C). Second ridge, probably homologous with the proximal ridge of Ruta & Jell (19992) (dr in Fig. 12C), running from left scutula to lateral margin of left PM, bending rapidly latero-distally and continuing on internal surface of left PLM as the most proximal part of the low ridge described above (lr in Fig. 12C). Apophyseal horns (ah in 363 Fig. 12C) gently convex in transverse section, with straight proximal margin and gently convex distal margin in plan view, with medial ends separated by small gap (Fig. 9C). Septum (se) on plate C divided into a straight proximal 1/2 and narrower, gently convex to the right or sinuous distal half. Distal 1/3 of septum gently convex to the right, proximo-distal on inside of right DLM. Distally, septum has T-shaped birfucation near latero-distal angle of right DLM. Convex surface. Inside of convex surface poorly preserved (Figs ІА, 11E). Co-operculum cup-like, just distal to midpoint of proximal margin of C20 and C22, with irregular rim, thicker in its distal 1/2 than in its proximal 1/2, with narrow slit at proximo-medial angle. Low oblique crest running medio-distally from medio-distal angle of co-opercular rim (Fig. 1A). C21 with weak struts radiating from point proximal to midlength (Fig. 1A) and reaching its proximal margin, with proximo-distally elongate, low, poorly defined ridge centrally (Fig. 11E), gradually disappearing distally. Stereom. Inside of plano-concave surface with minute pits or, more frequently, honeycomb-like. Plate peripheries usually with minute pits sometimes very shallow and surrounded by poorly defined trabeculae. Proximal part of MOP and LOP compact or coarsely fibrillar, rarely with shallow, subelliptical pits and small lumps; distal part of orifice plates coarsely granular or with radiating and often bifurcating trabeculae separated by deep furrows (Figs 7C, 9D, 11D). Stereom of inside of convex surface cancellate to coarsely granular, of low anastomosing trabeculae separated by shallow elongate pits (Fig. 11E). APPENDAGE. Proximal part of 6-7 tetramerous overlapping rings; ring plates of uniform width, in sutured contact mid-longitudinally and laterally. Ring plates on plano-concave side of body subrectangular, convex in transverse section, with straight proximal margins, with distal margins gently convex or sinuous and slightly thickened. Ring plates on convex side narrower, with straight proximal and distal margins. Styloid 3 times as wide as long, maximum width at distal blade, with low median keel. Proximal blade with semicircular, blunt free margin and flat to gently convex distal surface. Distal blade proximally recumbent, with semicircular sharp free margin, with flat surfaces, with proximal surface sometimes divided into 2 slightly convex halves (Figs 2B, 7B). Abapical surface of styloid (Figs 3D, 12E) gently concave 364 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 8. Protocytidium elliottae gen. et sp. nov. A. proximal plano-concave surface and appendage plates of NMVP100413. +7. B. proximal plates of plano-concave surface and parts of appet idage of NMVP100391_ «6. C. PM plates over inside of convex surface plates of NMVP100419. «12. D, inside of plano-concave surface of NMVPI00401, +12. A NEW VICTORIAN ANOMALOCYSTITID MITRATE 365 FIG 9. Protocytidium elliottae gen. et sp. nov. A, inside of plano-concave surface and orifice plates of NMVP100392, «7. B, partial plano-concave surface of NMVP 100409. х1 2. C, inside of plano-concave surface of NMVP100418, x12. D, detail of distal part of Fig. SD. NMVP100401. х 15. 366 MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 10. Protocytidium elliottae gen. et sp. nov. A. partial convex surface. NM VP10043 5. 77. В. lateral view of proximal appendage and plano-concave surface of NM VP 100430, «8 C, inside of plano-concave surface of NMVP]100399, 12. D, partial convex surface of NMVP 100416. «12. A NEW VICTORIAN ANOMALOCYSTITID MITRATE in transverse section, with slightly raised proximal and distal margins, with straight central longitudinal furrow (sf in Fig. 12E) along proximal 2/3 of styloid, of uniform depth and width, shallowing to rounded proximal termination and distally in subelliptical pit (sp in Fig. 12E), with proximal margin slightly concave medially. Articular surface (Fig. 12E) with 4 irregular pits laterally on left and right. Distal part. Largest observed number of ossicles 11 (Fig. 1D), articulated with paired plates. Ossicles only slightly higher than wide, triangular in cross-section (Fig. 10C), diminishing in size distally; lateral surfaces flat or gently convex in abapical 1/2, slightly depressed in apical 1/2; apical margin blunt, sloping distally, with blunt bulbous apex. Apex developed mainly on 3 or 4 most proximal ossicles, decreasing rapidly distally, absent on distal ossicles. Articular margins of ossicles along zig-zag line, divided into smaller, distal portion in contact with plate of next distal segment and larger, proximal part sutured with plate of corresponding segment. Plates subrectangular, partly overlapping each other proximo-distally. Articular margin of plates thicker proximally, gently tapering distally. Stereom. Appendage externally with shallow irregular pits. Tetramerous rings and free margins of styloid blades sometimes coarsely granular. Radiating trabeculae on both styloid blades. REMARKS. Material is often considerably disrupted and partly deformed. Very few specimens approach completeness, and the distal part of the convex surface is invariably damaged or missing. In some cases, the body plates are disarticulated but lie close to each other so that their mutual spatial relationships are almost unchanged. Preservation of the plano-concave surface is generally better than that of the convex surface, but the precise arrangement ofthe orifice plates can be deduced only by combining information from different individuals. In some specimens, both spines are visible and in the holotype, these are found in close proximity to the body, although only the left spine is preserved articulated with the toroidal projection on the distal surface of left DLM. The appendage is always incompletely preserved, although isolated ring plates of the proximal part, the abapical surface of the styloid and external ossicle morphology can be reconstructed. Despite tectonic deformation, it is possible to distinguish 2 morphological variants. 367 Homologous plates in individuals belonging to these variants show slightly different length/width ratios (e.g. PLM, ILM and DLM) and, sometimes, remarkably different shapes (e.g. A and C). The possibility that the 2 variants represent the effects of compression of the body along different directions cannot be entirely ruled out. Intraspecific and ontogenetic variation or sexual dimorphism could also be responsible. Similar problems were discussed by Ruta & Bartels (1998) in their analysis of tectonic deformation in Rhenocystis latipedunculata Dehm, 1932. Unlike Rhenocystis, specimens of P. elliottae have not been found in proximity to one another on the same slab surface and, therefore, retrodeformation techniques could not be applied to the Australian taxon. The first variant is more frequently represented with elongate, subelliptical to pyriform body outline and usually has markedly elongate plates A and C. In the second, rarer variant the body is only slightly longer than wide, subrectangular; both marginal and central plates of plano-concave surface are shorter; in particular, A and C are wider than long, subrhomboidal in outline and of equal width. The 2 variants do not seem to be related to body size, although some specimens belonging to the first category are among the largest known. ORIGIN AND EVOLUTION OF THE ALLANICYTIDIIDAE Established by Caster & Gill (1967) to accommodate Early Devonian Allanicytidium flemingi from the Reefton Group of New Zealand, the Allanicytidiidae has expanded to now include 7 Southern Hemisphere anomalocystitids (Allanicytidium, Notocarpos, Tasmanicytidium and Protocytidium from Australia, Placocystella from South Africa and Occultocystis and Australocystis from South America). Philip (1981) recognised the almost identical plating pattern in Allanicytidium and Notocarpos garratti (Ludlow), although he did not note the lateral orifice plates in the latter taxon and misinterpreted the arrangement of C20 and C22 (Caster, 1983). Caster (1983) provided a diagnosis and revision of the allanicytidiids with his description of Tasmanicytidium burretti (Llandovery). Haude (1995) modified the diagnosis to include Occultocystis koeneni (Early Devonian). Prior to discovery of Protocytidium, Occultocystis provided the only link between Late Ordovician mitrate faunas from Laurentia and mid-Palaeozoic mitrates from Gondwana. 368 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. П. Protocytidium elliottae gen. et sp. nov. Inside of plano-concave surface, convex surface, orifice plates and stereom, A. inside of plano-concave surface of NMVP100421. «12. B, partial convex surface of NMYP100400, *12. C, E, partial convex surface of NMVP100428. «7. D. detail of distal part of Fig, 5C showing inside of orifice plates of NMVP 100398, х2(), A NEW VICTORIAN ANOMALOCYSTITID MITRATE 169 VIG. 12. Protocytidium elliottae gen. et sp. nov. Reconstruction of internal features of body. A, lett DLM and left LÖP. based on NMVP100401. B. right DLM based on NMVPI00418. C, left PLM and left PM, based on NMVP100393. D, inside of MOP and LOP, reconstructed from NMVP100398, 100401, 100403 and 100488. Г. ubapical surface of styloid. based on NMVP100417. Abbreviations as in text. Drawings not to scale, 370 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 13, Reconstruction of Prorocytidium elliottae gen. et sp. nov, A, plano-coneave surface, B. convex surface. С, right lateral view. D, proximal view (spines omitted). E, distal view (appendage omitted), A NEW VICTORIAN ANOMALOCYSTITID MITRATE Character analysis of the allanicytidiids and relatives was undertaken by Ruta & Theron (1997). Enoploura is the sister-group to monophyletic Allanicytidiidae (Ruta in press). Ruta & Theron (1997) assigned Placocystella africana (Reed, 1925) and Australocystis langei Caster, 1956 (Early Devonian) to the Allanicytidiidae and recognised that the plano-concave surface of Tasmanicytidium is similar to that of more derived allanicytidiids (Caster, 1983; Ruta, in press). Mongolocarpos minzhini Rozhnov, 1990 (Ludlow; Mongolia) is removed from the Allanicytidiidae despite the proximo-distal elongation of plate A. It is here considered a close relative of Placocystites, Rhenocystis and Victoriacystis following Ruta (in press). ENOPLOURA AND THE ORIGIN OF THE ALLANICYTIDIIDS. Enoploura popei Caster, 1952 (Fig. 14A) has been described in great detail (Caster, 1952; Parsley, 1991). Its distalmost transverse row of plates on the convex surface has 5 elements as in Barrandeocarpus norvegicus Craske & Jefferies, 1989 and the anomalocystitids Anomalocystites cornutus Hall, 1858, Bokkeveldia oosthuizeni Ruta & Theron, 1997, Mongolocarpos minzhini Rozhnov, 1990, Placocystites forbesianus de Koninck, 1869, Rhenocystis latipedunculata Dehm, 1932 and Victoriacystis wilkinsi Gill & Caster, 1960. The lateral plates of the distalmost transverse row in Enoploura overlap the admedian plates as well as the marginal plates just proximal to them (Parsley, 1991). These marginal plates and the 2 large central plates lying medial to them may correspond to C6-C9 in Bokkeveldia, in which the admedian plates ofrow II, C7 and C8, are relatively large in comparison with other plates ofthe convex surface, as well as in other anomalocystitids. As an alternative hypothesis, the marginal plates sutured with C20 and C22 and the two large central plates of the convex surface of Enoploura may be homologous with the lateral (C15 and C19) and admedian (C16 and C18) plates of row IV in Bokkeveldia. Enoploura and Ateleocystites guttenbergensis Kolata & Jollie, 1982 are similar in configuration of rows П and Ш in the latter resembling that of the 2 distalmost rows of Enoploura. Thus, Enoploura differs from Ateleocystites in possessing a shortened convex side (Parsley, 1991) in which the distal, 5- plated row would correspond to row П of Ateleocystites; the row 371 proximal to it would be homologous to row III of Ateleocystites and the 2 marginal elements in contact with C20 and C21 on the right side and with C21 and C22 on the left side would correspond to the lateral elements of row IV (C15 and C19) in Ateleocystites. Furthermore, the proximo-distal imbrication pattern of the 2 large central plates of Enoploura is similar to that of the admedian elements of row Ш (C11 and C13) in Ateleocystites. According to Parsley (1991), the body of Enoploura is progenetically shortened in comparison with that of other mitrates. This is an intriguing hypothesis, but it needs to be corroborated by additional fossil evidence. A derivation of Enoploura from taxa with a polyplated convex surface is likely, but not certain. Despite its specialised features (e.g. morphology of the styloid; plate arrangement of the convex surface), Enoploura retains a primitive skeletal configuration on the plano-concave surface, as indicated by plate B. Such a config- uration suggests that this genus probably evolved from an ancestor resembling such Laurentian genera as Afeleocystites (Ruta, in press). PROTOCYTIDIUM: BASAL ALLANI- CYTIDIID. Skeletal configuration of Proto- cytidium elliottae (Fig. 14B) invites comparisons with Enoploura and with such basal allanicytidiids as Occultocystis. Affinities of Protocytidium with the allanicytidiids are suggested by the central plates of the plano- concave surface, especially the proximo-distally elongate plate A, and by the longitudinal styloid keel, a semicircular, proximal styloid blade and a proximally recumbent distal blade. The distal 2/3 of C21 is similar in shape and proportions to its homologue in Zasmanicytidium. However, although plate A is more elongate and narrower than in Notocarpos and is shaped like its homologue in such allanicytidiids as Placocystella, it does not contact left PLM as in all allanicytidiids more derived than Notocarpos (Caster, 1956, 1983; Caster & Gill, 1967; Ruta & Theron, 1997; Ruta & Jell, 1999c; Ruta, in press). Although MOP is longer than LOP, it is not as expanded transversely as in Notocarpos, Tasmanicytidium, Placocystella, Allanicytidium and Australocystis. Furthermore, both LOP plates are wider proximally than distally and not wedged obliquely between MOP and DLM (MOP and LOP not known in detail in Occultocystis). FIG. 14. Plano-concave (left) and convex (right) surfaces of A, Enoploura popei (redrawn and modified from Parsley, 1991). B, the basal allanicytidiid Protocytidium elliottae. C, convex surface of Occultocystis koeneni (redrawn and modified from Haude, 1995). Drawings not to scale. Except for the occurrence of 3 (as in Occultocystis) rather than 5 distal plates, skeletal configuration of the convex surface of Protocytidium differs from that of Enoploura mainly in the relative size and proportion of various plates. The 3 distalmost plates of the MEMOIRS OF THE QUEENSLAND MUSEUM convex surface (especially C3) are larger in Protocytidium than in Enoploura but smaller than in Occultocystis. Both in Enoploura and in Protocytidium LOP and MOP project distal to the body orifice and are divided into proximal and distal parts ('thecal’ and 'lip’ of Parsley, 1991). Gaps are present distally between MOP and LOP but they are smaller and more irregular than in Enoploura. The transverse thickening on the internal surface of LOP and MOP is another similarity between Protocytidium and Enoploura. More striking resemblances are on the internal side of the plano-concave surface, especially in the asymmetrical development of internal ridges near the latero- distal angles ofthe left and right DLM. Plate B, generally regarded as a primitive character for the anomalocystitids (Craske & Jefferies, 1989; Ruta & Theron, 1997; Ruta, in press), is much smaller in Protocytidium than in Enoploura and other mitrates and appears to be displaced slightly to the left of the longitudinal body axis. Furthermore, B does not contact right LOP and is strongly asymmetrical in outline. C6, C9, C15 and C19 of Enoploura differ from their homologues in Protocytidium in being much longer than wide. Relative size and proportions of C11, C13 and C21 are similar in both taxa. The coarsely pitted to labyrinthine stereom in Enoploura popei (Caster, 1952; Parsley, 1991) is only vaguely reminiscent of the external skeletal texture of Protocytidium. The latter appears to be less coarse, presumably as a result of the much smaller body size of Protocytidium, and more variable both on the surface of single body plates and on different plates. Resemblances between the body stereom of Enoploura and that of Protocytidium nevertheless occur in the vermicular to ridge-like pattern of C20 and C22 and the lateral body walls. TRANSITIONAL OCCULTOCYSTIS. In Occultocystis (Fig. 14C), C21 is notas large as in other allanicytidiids and is not interposed between C20 and C22. On both right and left of its convex surface, a plate may occur which is perhaps incompletely fused with C20 and C22 respectively (Haude, 1995). The distal margin of the convex surface consists of 3 plates, of which the median one is narrow and elongate with concave lateral margins as in Enoploura. It is likely that the marginal plates sutured along the midline of the convex surface of Occultocystis are homologous with the 2 large central elements of Enoploura. A NEW VICTORIAN ANOMALOCYSTITID MITRATE 373 Allanicytidium flemingi Australocystis langei Placocystella africana Tasmanicytidium burretti 1 Notocarpos garratti 2 Occultocystis koeneni 3 Protocytidium elliottae 7 Enoploura popei 5 Rhenocystis latipedunculata | Victoriacystis wilkinsi 8 Placocystites forbesianus 9 Mongolocarpos minzhini 1 Bokkeveldia oosthuizeni Anomalocystites cornutus 7 Willmanocystis denticulata 7 Kopficystis kirkfieldi 4 Barrandeocarpus jaekeli 8 Barrandeocarpus norvegicus 10 Aleleocystites guttenbergensis 2 Diamphidiocystis drepanon 8 Kierocystis inserta | Mitrocystella incipiens 2 Mitrocystella barrandei 4 Eumitrocystella savilli here to establish the phylogenetic position of Protocytidium elliottae. Morphological characters are those discussed by Ruta (in press) with coding for Kierocystis inserta Parsley, 1991 accounting for the reconstruction of the proximal 1/3 of the convex surface proposed by Parsley (1991: pers. comm., 1997). Some characters of the spines are entered as polymorphic for Protocytidium, The matrix includes 24 taxa and 106 binary characters and was analysed with PAUP Version 3.1.1 on a Power Macintosh 7500/100 using the same heuristic search settings as detailed by Ruta (in press). The analysis yielded a single tree (length = 230 steps: consistency index excluding uninformative characters = 0.456: retention index = 0.68; rescaled consistency index = 0.322) (Fig. 15). Major differences between this tree and the 3 equally parsimonious solutions found by Ruta (in press) are: 1) Barrandeocarpus jaekeli Ubaghs, 1979 and B. norvegicus Craske & Jefferies, 1989 are sister taxa and, together, form the sister-group of FIG. 15. Most parsimonius tree resulting from cladistic analysis. Numbers indicate branch lengths. Plating of the convex side of Occultocystis differs from that of other allanicytidiids in several details. The transition from this genus to more derived allanicytidiids was probably charac- terised by the disappearance of the median element of the distalmost transverse row and by elongation of C21 both distally and proximally. The evolutionary history of more derived allanicytidiids is characterised by remarkably few character changes, most of which pertain to general proportions and relative size of plates of convex surface, skeletal sculpture and appendage morphology (Ruta & Theron, 1997; Ruta & Jell, 1999c; Ruta, in press). PHYLOGENETIC ANALYSIS Data from a study of interrelationships of the anomalocystitid mitrates (Ruta, in press) are used Ateleocystites guttenbergensis Kolata & Jollie, 1982; 2) Kierocystis inserta Parsley. 1991 and Diamphidiocystis drepanon Kolata & Guensburg. 1979 are successively more closely related to the remaining ingroup taxa. Protocytidium occupies an intermediate position between Hnoploura popei Caster, 1952 and the Allanicytidiidae as defined by Haude (1995) and Ruta (in press), thus confirming the transitional nature of several of its features. The following characters, all showing state change 0-1 numbered in the same order as they appear in the data matrix (Ruta in press) and accompanied by theirconsistency index (ci) values, support the sister-group relationship between Protocytidium and the Allanicytidiidae under the accelerated character-state transformation (character changes are placed as close to the root of the tree as possible. thus emphasising reversals): 21 (сі= 1). plate MOP longer than each of the 2 plates ГОР; 30 (ci=0.333), spine length greater than 174 length of distal margin of plano-concave surface; 87 (ct=0.25), 3 plates along distalmost margin of convex surface; 90 (ci=1), interior of distal margin of convex surface with transverse thickening with asymmetrical cross-section; 97 (сі=1). distal styloid blade inclined proximally; 99 (ci=0.25). a sharp, longitudinal keel on external surface of styloid; 102 (ci-1). proximal styloid blade semicircular in outline. With the exception of characters 30, 87 and 99, the other characters are uniquely derived features of the clade (Protocytiditim elliottae + remaining Allanieytidiidae). When the delayed character- state transformation 1s used (character changes are placed as far from the root of the tree as possible, thus emphasising parallelisms), the clade comprising Prorocytidium elliottae and the remaining Allanicytidiidae is supported by state changes relative to characters 21. 30, 97, 102 as well as by character 53 (ci=0.25), absence of overlapping elements on convex surface, Prom this pattern of character distribution, it is possible to highlight the major changes during allanicytidiid evolution. Modifications of the plano-concave surface include: reduction in size and subsequent loss of B; proximo-distal elongation of A and subsequent interposition of it between C and left lateral marginal DLM, ILM and PLM; narrowing and acquisition of oblique orientation of right and left LOP, which became wedged between MOP and right and left DLM, respectively; widening of MOP; acquisition. of flexible articulation between orifice plates and adjacent plates of plano-concaye surface. Modifications of the convex surface include: reduction of distal row of plates from 5 to 3 to 2 elements; great expansion of C71; arrangement of marginal plates in 4 sets of paired elements surrounding C21: projection of most distal pair of plates beyond distal margins of LOP and MOP. Modifications of the appendage include: reduction in number of tetramerous rings; acquisition of semicircular outline of free margin of proximal styloid blade: proximally recumbent position of distal styloid blade: lateral ear-like projections of distal blade. CONCLUSIONS The diverse mitrate fauna of Australasia is entiched by addition of the latest Ordovician anomalocystitid Prorocytidium ellivitae. Spine morphology, external stereum texture and skeletal configuration of body plates distinguish MEMOIRS OF THE QUEENSLAND MUSEUM this mitrate from other anomalocystitids. Character analysis indicates that Protacyriditum is the most primitive member of the Allani- cytidiidae. Character distribution patterns and comparison of this genus with Enoploura and Oceultocystis suggest several skeletal modifications in evolution of the allanicytidiids including: |) simplification of the convex surface plating through loss of plates: 2) modification of the orifice plutes to form a flexibly articulated siructure; 3) inerease in the degree of bilateral symmetry of the body: 4) loss of plate B; 5) proximo-distal elongation of plate A; and 6) lateral expansions of distal styloid blade. ACKNOWLEDGEMENTS lan Stewart and Fons Vandenberg helped collect the material described herein. А.К. Milner (Birkbeck College, University of London), A.C. Milner. 5.1, Culver and L.R.M. Cocks (Natural History Museum, London) read the manuscript. P. Crabb (Natural History Museum, London) photographed the specimens. B. Lefebvre and Ron Parsley provided useful information. We arc grateful to the reviewers, Ron Parsley and Jim Sprinkle for their helpful suggestions and vouch that the authors alone are responsible for the above. A European Community grant (Training and Mobility of Researchers) enabled MR to visil the Queensland Museum (Brisbane) and the Museum of Vicioria (Melbourne), whose stall are thanked for their hospitality. LITERATURE CITED CASTER, К.Е. 1952. Concerning Enoplonra of the Upper Ordovician and its relation to other carpoid Echinodermata. Bulletins of American Paleontology 34: 1-47. 1956, A Devonian placocystoid echinodermi front Paraná, Brazil. Paleontologia do Paraná (Centennial Volume): 137-148. 1983, A new Silurian carpoid echinoderm. from Tasmania and а revision of the Allanicytidudae, Alcheringa 7: 321-335. CASTER, K.E. & GILL, E.D. 1967. Family Allanteytidtidae, new family. Pp. 5561-5504. In: Moore, К.С. (ed.) Treatise on invertebrate paleontology. Part 5. Echinodermata 1(2). (Geological Suciety af America & University of Kansas: New York). CRASKE, A.l. & JEFFERIES. R.P.S. 1989. А new milrate from the Upper Ordovician of Norway, and a new approach to subdividing a plesjon. Palaeontology 32; 69-99, DEHM, R. 1932. Cystoideen aus dem rheinischen Unterdevons. Neues Jahrbuch fiir Mineralogie, Geologie und Paläontologie. Beilage-Band, Abteilung A 9; 63-93. A NEW VICTORIAN ANOMALOCYSTITID MITRATE GILL, E.D. & CASTER, K.E. 1960. Carpoid echmoderms from the Silurian and Devonian of Pete Bulletins of Americun Paleontology 41: 5-71. HALL, J. 1858. Scientific intelligence, IT, geology, 4. Crinoids of New York. American Journal of Science and Arts 25: 277-279. HAUDE, R. 1995, Echinodermen aus dem Unter-Devor der argentinischen Prükordillere. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 197: 37-86. JAEKEL. О. 1918. Phylogenie und System der Tienen. Paliontologische Zeitschrift 3: 128, KOLATA, D.R. & GUENSBURG, T.E. 1979. Diamphidiocystis, a new mitrale carpoid from the Cincinnatian (Upper Ordovician) Maquoketa Group in southern Ilinois. Journal of Paleontology 53: 1121-1135. KOLATA, D.R. & JOLLIE, M. 1982. Anomalocystitid mitrates (Stylophora, Echinodermata) from the Champluinian (Middle Ordovician) Guttenberg Formation of the Upper Mississippi Valley Region. Journal of Paleontology 56; 531-565. KONINCK, M.L. de. 1869. Sur quelques Echinodermes remarquables des terrains paléozoiques. Bulletin de l'Academie Royale des Sciences Belgique 28: 544-552. PARSLEY. R.L. 1991. Review of selected North American mitrate stylophorans (Homalozoa: Echinodermata), Bulletins of American Paleontology 100: 5-57. PHILIP, GM. 1981. Notocarpos garratli gen. et sp. nov., anew Silurian mitrate carpoid from Victoria, Alcheringa 5: 29-38. REED, ER.C. 1925, Revision of the fauna of the Bokkeveld beds. Annals of the South African Museum 22: 27-226. KOZHNOV, S,V. 1990, New representatives of the class Stylophora (Echinodermata). Paleont- ological Journal 24: 34-45, 375 RUTA, M. in press. À cladistic analysis of the anomalocystitid mitrates. The Zoological Journal of the Linnean Society. КОТА, M, & BARTELS, С. 1998, A redescription of the anomalocystitid mitrate Rhenoevstis latipedunculata from the Lower Devonian of Germany. Palaeontology 41: 771-806. RUTA, M, & JELL, P. A. 1999a, Adoketocarpus gen. nov. а mitrate from the Ludlovian Kilmore Silistone and Lochkovian Humevale Formation of central Victoria. Memoirs of the Quecnsland Museum 43: 377-398. 1999b. Two new anomalocystitid mitrates from the Lower Devonian Humevale Formation of central Victoria, Memoirs of the Queensland Museum 43: 399-422, 1999c. Revision of Silurian and Devonian Allanicytidiidae (Anomalocystitida, Mitrata) from southeastern Australia, Tasmania and New Zealand. Memoirs of the Queensland Museum 43: 431-451. RUTA, M. & THERON, JN. 1997. Two Devonian mitrates from South Africa. Palaeontology 40: 201-243, UBAGHS, G. 1967. Stylophora, Pp. 5496-5565. In: Moore, R.C. (ed.) Treatise on invertebrate paleontology. Part S, Echinodermata 1(2). (Geological Socicty of America & University of Kansas: New York). 1979, Trois Mitrata (Echinodermata: Stylophora) nouveaux de l'Ordovieien de Tchécoslovaquie. Paldontologische Zeitschrift 53: 98-119. VANDENBERG, A.H.M. 1992, Kilmore 1:50,000 map and geological report. Geological Survey of Victoria Report 91: 1-86, + map. VANDENBERG, A.H.M., RICKARDS, К.В. & HOLLOWAY, D.J. 1984. The Ordovician- Silurian boundary at Darraweit Guim, central Victoria. Alcheringa 8: 1-22. WETHERBY, A.G. 1879. Description of a new family and genus of Lower Silurian Crustacea. Journal of the Cincinnati Society of Natural History |: 162-166. 376 MEMOIRS OF THE QUEENSLAND MUSEUM APPENDIX Through this and the following 4 papers by the same authors reference is made to a system of plate nomenclature proposed in a paper by the senior author that remains in press with The Zoological Journal of the Linnean Society of London (Ruta, in press). To facilitate the use of this nomenclature in the papers published in this volume a key to that plate notation is provided below. Plating of the convex surface 1s shown on the lefthand diagram and of the plano-concave surface on the right. The convex surface is based on the maximum regular plating known which occurs in the South African Bokkeveldia oosthuizeni Ruta & Theron, 1997. This terminology has been developed to avoid entirely any implied interpretation of orientation or function and although no thanks may be forthcoming for introducing another terminology in an already contentious area we believe use of terminology which removes all interpretation is desirable and should be of benefit to the enduring arguments surrounding these animals. The following abbreviations are employed on the figure:- On the convex surface ‘c’ prefix is for convex and equates to the ‘v’ for ventral used by Ruta & Theron (1997). On the plano-concave surface PM - proximal marginal plates PLM- proximal lateral marginal plates ILM- intermediate lateral marginal plates DLM - distal lateral marginal plates LOP - lateral orifice plates MOP - median orifice plate Central plates:- A — anomalocystid plate B = second asymmetrical plate of some genera C - largest central plate SE А0" ADOKETOCARPUS GEN. NOV., A MITRATE FROM THE LUDLOVIAN KILMORE SILTSTONE AND LOCHKOVIAN HUMEVALE FORMATION OF CENTRAL VICTORIA MARCELLO RUTA AND PETER A. JELL Ruta, M. & Jell, P.A. 1999 06 30: Adoketocarpus gen. nov., a mitrate from the Ludlovian Kilmore Siltstone and Lochkovian Humevale Formation of central Victoria. Memoirs of the Queensland Museum 43(1): 377-398. Brisbane. ISSN 0079-8835. The mitrate Adoketocarpus gen nov. is the first representative of the Paranacystidae from Australia. It occurs in central Victoria with А. acheronticus sp. nov. in the Ludlovian Kil- more Siltstone and А. janeae sp. nov. in the Lochkovian part of the Humevale Formation. Adoketocarpus has a simplified plate arrangement on the convex surface and a strongly twisted orifice; plating of the plano-concave surface resembles that of Middle Ordovician Eumitrocystella savilli from Morocco suggesting that the paranacystids may derive from boreal mitrocystitids. Fewer lateral marginal plates of the plano-concave surface, loss and size reduction of distal plates and enlargement of proximalmost plates of the convex surface would attend evolutionary transition from boreal mitrocystitids to paranacystids. Parana- cystids formed a clade whose origin and evolution were apparently in Gondwanaland, where they probably dispersed in the Ordovician/Silurian. O Paranacystidae, Adoketocarpus, Silurian, Devonian, Victoria. Marcello Ruta, Department of Palaeontology, Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom; Peter A. Jell, Queensland Museum, P.O. Box 3300, South Brisbane 4101, Australia; received 3 December 1997. The known Australian carpoid fauna of 4 en- demic genera with 6 species is more abundant than faunas from other parts of Gondwana. Some Australian carpoids have affinities with species from Europe and North America (Gill & Caster, 1960; Ruta, 1997a), whereas others have affinities with taxa from South America, South Africa and New Zealand (Caster, 1956, 1983; Caster & Gill, 1967; Philip, 1981; Haude, 1995; Ruta & Theron, 1997). The new genus described herein has its nearest relatives in the Lower Devonian of South America and South Africa. Among Australian carpoids none are closely related to the new genus. Rutroclypeus is the only solute known from Australia; it occurs in the Lower Devonian of Kinglake, Victoria, with А. junori (Withers, 1933), R. victoriae Gill & Caster, 1960 and R. ? withersi Gill & Caster, 1960. The other 3 monotypic genera are mitrate stylophorans of the Anomalocystitida: Tasmanicytidium burretti Caster, 1983 from the upper Llandovery Richea Shale of SW Tasmania, Notocarpos garratti Philip, 1981 from the upper Ludlovian part of the Humevale Formation near Whittlesea, central Victoria and Victoriacystis wilkinsi Gill & Caster, 1960 (Ruta, 1997a) from the lower Ludlovian in the Dargile Formation near Heathcote and the Melbourne Formation at Hawthorn. Extensive collections of echinoderms from the mid-Palaeozoic of central Victoria made over many years, but mainly during the early 1980s, are housed in the Museum of Victoria. This paper deals with anew paranacystid mitrate genus from the Upper Silurian and Lower Devonian of central Victoria. It is important because: 1, it is the first Australian record for the Paranacystidae Caster, 1954; 2, its occurrence in the Upper Silurian is the earliest for the family, predating the Lower Devonian record in South America (Caster, 1954; Haude, 1995); 3, its anatomy can be reconstructed in detail, thus throwing light on several poorly known aspects of the parana- cystids and prompting a new interpretation of their skeleton (Caster, 1954; Ruta, 1997c). 4, it provides additional evidence of the affinities between Siluro-Devonian mitrate faunas of Australia and the Malvinokaffric Realm (Caster, 1954, 1956, 1983; Gill & Caster, 1960; Caster & Gill, 1967; Philip, 1981; Parsley, 1991; Haude, 1995; Ruta & Theron, 1997; Ruta, 1997c). LOCALITIES, The mitrates described in this work are from 4 different localities in the Museum of Victoria locality register (NMVPL) representing 3 horizons. NMVPL252 Middendorp’s Quarry at Kinglake West is marked on the map of Williams (1964) and discussed elsewhere (Ruta & Jell, 1999); it 378 occurs in the Humevale Formation and the brachiopod fauna indicates the Boucotia janeae Brachiopod Zone (Garratt, 1983) of mid Lochkovian age. NMVPL1924 (=T95 of Williams (1964)) is in fine sandstones in the bed of Mathieson Creek, 2km S ofthe Kinglake West to Flowerdale Road. Carpoids are a minor component of an extensive fauna dominated by diverse crinoids and stel- leroids and including brachiopods, bryozoans, corals (Pleurodictyum only), bivalves and several other minor groups. Williams (1964) showed this site to be in the same Sandstone Member (probably the Flowerdale Sandstone Member) as Collins Quarry near Kinglake West which is Lochkovian (Vandenberg, 1988) based on the brachiopods (Garratt, 1983). NMVPL1927 is on the eastern bank of Broad- hurst Creek, where it crosses the Kilmore to Wandong Road, SSW of Kilmore in central Victoria. The locality occurs at 37°20°30"S, 144°59°35"E (Grid Reference 220652) on the Kilmore 1:50,000 Geological Map (Vanden- berg,1992). The fossils come from the Kilmore Siltstone (Vandenberg,1992), a sequence of predominantly thin (5-10cm), horizontally banded siltstones and very thin (less than 1cm), cross-bedded sandstones with rare, irregularly interbedded turbidite sandstones. The sequence is more than 3,500m thick. This huge thickness of siltstone indicates very rapid deposition on a tectonically stable shelf. Graptolites indicate an early Ludlow age ( Van- denberg, 1992) for the upper Kilmore Siltstone in the Kilmore district, from which most of the shelly faunas have been collected. Trilobites pre- dominate, with rare brachiopods, bryozoans, ostracodes and rugose corals, all of which are concentrated mainly in coarse sandstones. Echinoderms are locally abundant in thick sandstone beds (Vandenberg, 1992). NMVPL 1927 occurs lower in the sequence and is most probably early Ludlow in age, although the possibility that it belongs to the late Wenlock cannot be ruled out. NMVPL1960 refers to material excavated from a pipeline trench about 2km towards Kilmore along the Kilmore to Wandong Road from NMVPL1927 in similar lithology and horizon. SYSTEMATIC PALAEONTOLOGY A standard anatomical nomenclature for mitrates does not exist (Caster, 1952; Caster & Gill, 1967; Ubaghs, 1967; Kolata & Jollie, 1982: MEMOIRS OF THE QUEENSLAND MUSEUM Jefferies, 1986; Cripps, 1990; Kolata et al., 1991; Parsley, 1991, Beisswenger, 1994; Ruta, 1997a, b; Ruta & Theron, 1997), The terminology of Ruta (in press) is followed for the external skeleton and plating pattern except that, for brevity, ‘left’ and ‘right’ replace ‘anomalocystid’ and ‘abanomalocystid’ respectively. For the internal anatomy, we use Ubaghs’ (1967, 1969) nomenclature, although reference is necessary to some other sources (Jefferies, 1986; Jefferies, 1973; Jefferies & Lewis, 1978; Cripps, 1990; Beisswenger, 1994; Ruta & Theron, 1997). Unless otherwise stated illustrations are of latex casts from decalcified siltstone, whitened with ammonium chloride sublimate. Class STYLOPHORA Gill & Caster, 1960 Order MITRATA Jaekel, 1918 Suborder incertae sedis Family PARANACYSTIDAE Caster, 1954 DIAGNOSIS (modified from Caster, 1954 and Ruta, 1997c). Plates A and C proximo-distally elongate, markedly different in shape and size; C separated from left DLM and left PLM by A. Left PM slightly larger than right PM, with chevron- shaped distal margin. Lateral and proximal marginal plates with well-developed subvertical projections. Skeletal sculpture of small elliptical knobs or tooth-like serrations along lateral mar- gins of PLM plates, sometimes extending on proximal parts of lateral margins of left DLM and right EXM/ILM. Proximal 1/2-2/3 of convex body surface formed by proximo-distally elongate, shield-like C20 and C22. Distal part of convex surface of 2 smaller, subtrapezoidal plates with sinuous to gently concave lateral and distal margins and pronounced latero-distal angles. Small, subquadrangular to lozenge-shaped plate, possibly homologous with C21, between C20 and C22, proximally and subtrapezoidal plates, distally. Proximal part of appendage shorter than maximum width of each PM plate. Styloid with slightly expanded to flared, non-recumbent blades and proximal stout artic- ulation process. Proximal ossicles of distal part of appendage expanded transversely and much more robust than successive ossicles. Adoketocarpus gen. nov. TYPE SPECIES. Adoketocarpus acheronticus sp. nov. ETYMOLOGY. Greek adoketos, unexpected and carpos, a fruit; refers to body shape and unusual skeletal features. Masculine. A LUDLOVIAN MITRATE FROM CENTRAL VICTORIA left proximal A Q 1 larger than right DLM, contributing to distal 1/3 of left lateral body margin. Subvertical projections of left and right PM straight to gently convex externally; suture between left and right PM bending slightly rightward distally. Plate C about as wide proximally as distally, with chevron-shaped proximal margin. Suture beiween A and C sinuous, gently concave rightward proximally, more deeply concave leftward distally, Suture between distal subtrapezoidal plates of convex surface shorter than their proximal and proximo- medial margins. FIG. 1. Orientation of the skeleton of Adoketocarpus acheranticus gen. et sp. nov, A, plano-coneave surface. B, convex surface. DIAGNOSIS. C at least twice as wide as A. Length of left and right PLM exceeding that of. other lateral marginal plates. Lateral serrations on PLM plates decreasing in size proximo- distally, sometimes extending on proximal part of lateral margins of left DLM and right EXM/ILM. Maximum width of proxiimal part of appendage less than maximum width of proximal margin of each PM plate. Adoketocarpus acheronticus sp. nov, (Figs 1-2, 3D, 4-10, ПА,р,Е, I2A-D) ETYMOLOGY. Greek Acheron. а river of the underworld in Greek mythology: most specimens were collected on the bank of a stream. MATERIAL, Holotype: NMVP100330. Paratypes: NMVP100331-100348 from NMVPL1927. Other material: NMVP100349-100356, QMF37202, 37208, 37212. 37213, 37214 from NMVPLI927; NMVP149357-149358 from NMVPL 1960. DIAGNOSIS. Lateral body margins gently convex, Plate A subpentagonal to wedge-shaped, slightly to much wider proximally than distally and c. 1/2 as large as C. Right LOP slightly smaller than C, roofing over the body orifice, with almost straight proximo-lateral margins and sinuous latero-distal margins; distal process pro- nounced, with blunt, rounded end. Left DLM much sharterthan right EXM/TLM, about as long as and slightly narrower than left LOP. Left LOP DESCRIPTION. EXTERNAL. Body longer than wide, ovato-lanceolate to pyriform, slightly (Figs 1-2, 3D, 4A-E, 5C, 1L A,B,D). Maximum width about twice as fat away from distal process of right LOP as from proximal excavation for appendage insertion. Lateral margins more strongly convex proxim- ally than distally; right margin slightly longer than left margin. Subvertical projections of lat- eral marginal plates visible when the convex surface is oriented towards the observer (Figs 4F, 6A.C.D. 7A,B, 11E, 12D). Distal body orifice twisted leftward and framed by radially arranged platelets of different shape and size (Figs 4F, 6A-D). Plano-concave surface slightly raised along lateral margins and near proximo-lateral angles and shallower centrally (Figs 4A-E, 5A-D). Maximum curvature of convex surface at the level of its proximal 1/3 (Figs 6A, C, 7B, 9B). Measurements. Holotype (Fig. 4A): c.4.7mm wide and 6.7mm long. Smallest specimen (Fig. 4C,F): c.2.7mm wide and 3.7mm long. Largest specimen (Fig. 4E): estimated body width and length c.5.7mm and 7.7mm, respectively. Plano-concave surface. Weakly concave surface ol L1 medium to large plates. Marginal plates in 2 groups of 3, a distal element roofing over the body orifice and 2 proximal elements contribut- ing to excavation for appendage insertion (Figs LA, 2A. 3D. 4A-E, 5A.C. 11A,B,D). Height of subvertical projections of lateral marginal plates approximately constant over most oftheir length, 380 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 2. Reconstruction of Adoketocarpus acheronticus gen. et sp. nov. (appendage omitted). A, plano-concave surface; B, left lateral view. C, convex surface. D, proximal view. E, distal view (hatched area shows space possibly occupied i in life by polyg gonal plates). only slightly decreasing near latero-distal angles of plano-concave surface and at the level of proximal 1/3 of both left and right PLM (Figs 2B, 4D-E, 5A.C, 6A.C, 7А, 11E); subvertical pro- jections of PM plates higher than those of lateral marginal plates, rectangularto trapezoidal in prox- imal view and flat to gently convex externally (Figs 2A-D. 4D. 5A-D). Subpentagonal left LOP and trapezoidal right DLM forming latero-distal angles of plano-concave surface; left LOP slightly wider and longer than right DLM. Left DLM and right EXM/ILM subrectangular and proximo-distally elongate; length of left DLM c.2/3 that of right EXM/ILM. Left and right PLM subequal in shape and size. triangular and contributing to proximo-lateral angles of body: lateral margins of both plates PLM with 4-5 serrations rapidly decreasing in size proximo- distally and either gradually merging into each other or neatly separated, 2-3 proximalmost serrations with blunt, rounded apex, steep distal margin and gentle proximal margin; 1-2 very shallow serrations near proximal 1/3 of lateral margins of left DLM and right EXM/ILM (Figs 1A. 2A, 3D, 4A-D, 5C, 7A, 11A). Distal margin of left PM longer than distal margin of right PM and chevron-shaped. the right arm of the chevron being as long as or longer than its left агт; median 1/2 of proximal margins of both plates PM excavated for insertion of appendage: suture between such plates generally bending slightly rightward distally; transverse thickening sometimes observed along their proximal margins (Figs 4D-E, 5A-D). Irregularly pentagonal right LOP slightly smaller than C, with prominent plectrum-shaped to semicircular distal process, gently sinuous latero-distal mar- gins and almost straight proximo-lateral margins (Figs 4A-E, 5A.C, 11A,B,D); distal process im- mediately left of longitudinal body axis. Plate A A LUDLOVIAN MITRATE FROM CENTRAL VICTORIA LT т чапта ащ FIG. 3. Plate nomenclature of the plano-concave (left) and convex (right) surfaces in 3 mitrocystitids and in Adoketocarpus acheronticus gen. et sp. nov. Specimens not to the same scale. A, Mitrocystella barrandei (redrawn from Ubaghs, 1968). B, Mitrocystella incipiens (simplified from Jefferies, 1986). C, Eumitrocystella savilli (modified from Beisswenger, 1994). D, Adoketocarpus acheronticus gen. et sp. nov. subpentagonal to wedge-shaped, longer than wide, slightly to much wider proximally than distally, comparable in size with left and right DLM (Figs 4A-D, 5C, 11A,B,D) and in contact with left LOP latero-distally, left DLM laterally, left PLM proximo-laterally and left PM proximo-medially. C twice as long and wide as A and in contact with right LOP distally, right DLM latero-distally, right EXM/ILM laterally, right PLM proximo- laterally and right and left PM proximally (Figs 2A, 3D). Suture between A and C with deep leftward concavity distally and less pronounced rightward concavity proximally (Figs 4A-D). Convex surface. Convex surface of 5 small to large plates symmetrically arranged in centre, plus 8 plates extending up lateral and proximal margins from plano-concave surface. Proximal 2/3 of convex surface of large, proximo-distally elongate, shield-like C20 and C22 plates, 1.5-2 times longer than wide (Figs 1B, 2C, 3D, 4F, 6A,C, 7B, 9A-C, 12B,D). Lateral 1/2 of proximal margins of such plates straight to strongly convex; median 1/2 deeply excavated for appendage insertion; lateral margins of both plates gently to moderately convex, sometimes showing abrupt curvature about half-way along their length; lateral and proximal margins merging into each other smoothly or at c.120° (Figs 4F, 6A,C-D, 7B,8B-D, 9A-C, 11C,E). Proximo-lateral part of external surface of both C20 and C22 distinctly sloping, almost vertical with respect to the rest of the plate in its distal 1/2 and delimited distally by straight, slightly pro- nounced keel with steeper distal and gentler proximal slope (Figs 2B-D, 6C, 7B, 9B-C). Keels running latero-medially and proximo-distally from proximo-lateral angles of both C20 and C22, their length being less than 1/2 the width of these plates. External margin of small facet for articulation with plates PLM visible on both C20 and C22 just latero-distal to lateral end of each keel (Figs 1B, 2B-D, 4F, 6A, C, 7B, 9B). Sub- elliptical to teardrop-shaped pit proximal to 382 MEMOIRS OF THE QUEENSLAND MUSEUM VIG. 4. Adoketocarpusacheronticus gen. et sp.nov., all (тот NMVPL 1927.x10, A-E, plano-concave surface. F, convex surface, B, D, Е. showing part of the appendage. A. NMYP 100330. holotype. В, NMVPIQUA31. C. F: NMVP100332. D. NMVP100333. E, NMVP100334. median 1/3 of keel on C20 (Figs 1B, 2B-D. 6С. 7B.9B). Distal 1/3 of convex surface occupied by 2 subtrapezoidal plates. slightly wider than long and in contact with each other along very short, straight suture and with C20 and C22 along gently sinuous sutures: latero-distal angles of subtrapezoidal plates with distally directed. blunt-ended processes (Figs 1B, 2B-C.E. 3D, 4F. 6A-D. 9B). Subquadraugular to lozenge-shaped C21 with straight to slightly convex margins in contact with medio-distal margins of C20 and C22 and with medio-proximal margins of sub- trapezoidal plates (Figs 2B-C.E. 6A-D. 7B. 9B). Distal orifice. Body orifice strongly twisted leftward, roofed over by right LOP and floored bv 6-7 platelets arranged radially (Figs 1B, 2B-C, E. AF. 6A-D). Right platelet subelliptical. with major axis almost perpendicular to longitudinal body axis; remaining platelets subrectangular to spike-shaped and decreasing progressively in A LUDLOVIAN MITRATE FROM CENTRAL VICTORIA 383 PIG. 5. Adoketocarpus acheronticus gen. et sp. nov., all from NMVPL 1927. A.C. plano- concave surface. B.D. proximal part of appendage and styloid. A.B. NMVP100335, x10 and «20, respectively. С.р, NMVP100336, ~10 and x20, respectively. size from right to left; left platelet subtriangular. between proximal margins of orifice platelets and with majoraxis parallel to longitudinal body axis. distal margins of subirapezoidal plates. more Polygonal elements of irregular shape inserted numerous on the right than on the left, smaller 384 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 6. Adoketocarpus acheronticus gen. et sp. nov., all from NMVPL 1927. A.C. convex surface and proximal part of appendage. B,D, close-ups of orifice platelets. A,B, NMVP100337, 10 and x30, respectively, C,D, NMVP 100338, x10 and x25, respectively. and transversely clongate near distal margins of subtrapezoidal plates, larger near proximal margins of orifice platelets: polygonal elements probably also covering proximal part of right latero-distal margin of right LOP (Fig. 6C-D). Stereom. Plates of plano-concave surface with irregular, highly variable external texture, ma consisting of subcircular to subelliptical pores separated by trabeculae (Figs 4A-E, 5A-C). Thinner trabeculae and smaller subcircular pores visible near lateral body margins. Trabeculae often forming a fringe near plate sutures. where the pores are narrow and elongate and arranged radially. Irregularly radiating pattern of sinuous, elongate, bifurcating trabeculae on marginal and subcentral plates, especially near their centres: adjacent trabeculae often sending at irregular intervals shorter, transverse trabeculae de- limiting subrectangular to subelliptical pores. A LUDLOVIAN MITRATE FROM CENTRAL VICTORIA 385 FIG. 7. Adoketocarpus acheronticus gen et sp. nov., all from NMVPLI927. A, NMVP100339, inside of plano-concave surface and partially preserved appendage. «10. B, NM VP100340, partially preserved convex surface, distal 1/3 of inside of plano-concave surface and disrupted appendage, *15. Stereom of subvertical projections of lateral mar- ginal plates generally compact or consisting of minute subcircular pores. Stereom of convex surface almost uniformly composed of circular pores and short trabeculae, the latter arranged radially near the plate sutures (Figs 4F, 6A-D, 7B. 9B-C). Stereom of orifice platelets coarsely granular or with very shallow pits surrounded by weak ridges (Figs 4F. 6B.D). INTERNAL. Plate C. right LOP (in part) and left and right PM reveal most of the inside of the plano-concave surface, but little is known about the interior of the lateral marginal elements (Figs 7A. 9B-C). So far as the convex surface is concerned. only the internal aspect of C20 and C22 is known in detail, whereas the inside of the distal 1/3 of this surface is only partially preserved. Plano-concave surface. Right latero-distal angle of internal surface of right LOP occupied by median 1/3 of subcircular depression straddling suture with right DLM and continuing on adjacent part of internal surface of latero-distal angle of this plate (Figs 1B, 2C, E. 6C-D). In- ternal side of plano- concave surface divided into 2 fields by oblique septum (Ubaghs. 1967) (= oblique ridge of Jefferies, 1986) (Figs 7A-B, 8A; sc in Fig. 10B): distal 1/3 of septum with marked 386 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 8. Adoketocarpus acheronticus gen. et sp. nov., all. from NMVPL1927. Internals of skeletal plates. A, NMVP100340, disrupted distal 1/3 of plano- concave surface: the element near top centre is right LOP, x25, B, NMVP100342. inside of C20, x20. C, NMVP100343. plates C20, C21, C22, left and right PM, left PLM, x16. D, NMVP100344, inside of C20, x12. A LUDLOVIAN MITRATE FROM CENTRAL VICTORIA 387 е АХ 160. uT ‘ 3 x. d > А + t LI r й "el x FIG. 9. Adoketocarpus acheronticus gen. et sp. nov., all from NMVPL1927. A, NMVP100345, inside of C20, x12. B, NMVP100346. convex surface, partially disrupted distal 1/3, x10. С, NMVP100347, slightly damaged C20 in external view, x12. rightward convexity occupying distal 2/3 of internal surface of C (Figs 6C-D. 7A-B. 8A); distalmost end rapidly weakening where it straddles suture between C and right LOP and vanishing distally on the internal surface of the latter; proximal 2/3 of septum straight, thicker than convex part and bending slightly to the right in cross-section; convex part passing gradually into straight part: boundary between these 2 parts delimited by elongate and poorly defined thickening (= diminutive spur of Ubaghs, 1967 and base of middorsal process of Jefferies, 1986) (Fig. 7A-B); straight part of septum running obliquely from internal side of proximal 1/3 of C to internal side of left PM, straddling suture just lateral to medio-distal angle of left PM (Fig. 10A-B). Proximalmost end of septum merging into cup- like left scutula (Ubaghs, 1967) (= dorsal calcitic cup of left pyriform body of Jefferies, 1986) (sc in Fig. 10B) delimited by slightly raised, thick, subcircular scutular rim. Deep transverse furrow MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 10. Adoketocarpus acheronticus gen. et sp. nov., from NMVPL1927, internal surface of left PM plate. A, NMVP100348, x25, latex cast coated with ammonium chloride. B, camera lucida drawing of the same specimen: abbreviationsas in text. C, NMVP100339, natural internal mould of proximal 1/3 of the2 PM plates, x25. D, NMVP100344, natural internal mould of proximal 1/3 of C20, «25. (7 transverse anterior groove of Ubaghs, 1967 and anterior boundary of posterior coclom of Jefferies, 1986) (tf in Fig. 10B) running from medio-distal part of scutular rim to median margin of both left and right PM and delimited distally by prominent transverse ridge. straight to slightly convex distally (tr in Fig. 10B). On the left PM, the median end of such a ridge merges into the proximalmost end of the septum, where the latter becomes confluent with the medio- distal part of the scutular rim. Two slightly curved ridges on internal side of the left PM plate. A LUDLOVIAN MITRATE FROM CENTRAL VICTORIA poorly developed or absent on the right PM, running almost parallel to each other or strongly diverging medio-laterally, and less robust than proximal part of septum; distal ridge (dr in Fig. 10B) projecting proximo-distally or transversely along gently curved direction from lateral wall of scutular rim to lateral margin of both PM plates where it widens and becomes less pronounced; distal ridge of left PM probably corresponding to accessory septum of Ubaghs (1967) (= posterior boundary of left pharynx of Jefferies, 1986); proximal ridge (pr in Fig. 10B) slightly shorter than distal ridge, almost straight and parallel to proximal margin of each of the two PM plates. Right PM differing from left PM in the small pit (7 infundibulum of Ubaghs, 1967 and pit for dorsal end of lateral line ganglion of Jefferies, 1986) lying proximo-lateral to right scutula (Fig. 8C). Shallow groove (sg in Fig. 10B) for artic- ulation with C20 and C22 running parallel to lateral part of proximal margins of left and right PM, and becoming progressively shallower near its lateral end; median end of groove interrupted by small ridge projecting latero-medially from medio-proximal part of scutular rim to lateral angle of external margin of excavation for appendage articulation (= cerebral basin of Jefferies, 1986). Proximalmost part of internal side of lateral margin of left and right PM show- ing ‘step’ (^s! in Fig. 10B) delimiting sudden change in the curvature of this margin and pre- sumably representing interlocking articulation with left and right PLM respectively. Median part of both left and right scutular rim merging medial- ly into transversely expanded area occupying median 1/3 of internal margin of excavation for appendage articulation, and presumably repres- enting basal portion of each of the 2 apophyseal horns (Ubaghs, 1967) (=hypocerebral processes of Jefferies, 1986) (ah in Fig. 10B), not preserved in available material. Excavation on both PM plates consisting of shallower distal and deeper proximal parts; distal part (=prosencephalar part of cerebral basin of Jefferies, 1986) delimited by internal margin of excavation and by thin ridge running along slightly oblique direction from bases of apophyseal horns to proximo-medial angle of both plates; distal part (=deuteren- cephalar part of cerebral basin of Jefferies, 1986) delimited by above mentioned thin ridge and by external margin of excavation. Convex surface. Cup-like co-operculum (Ubaghs, 1967) (= dorsal calcitic cup of pyriform body of Jefferies, 1986) near proximo-lateral angles of C20 and C22 (Figs 8B-D, 9A), 389 delimited by rim with thicker proximo-lateral and thinner medio-distal margins; rim interrupted medially by short, deep, transverse sulcus extending from excavation of co-operculum to portion of inner surface between co-operculum and proximal margins of C20 and C22; at the level of sulcus, both ends of co-opercular rim continue medially into slightly raised ridges parallel to median excavation of proximal margin of C20 and C22, gently convex distally and delimiting very shallow groove. Lateral wall of left and right co-opercula merging gradually into proximalmost part of lateral margin of both C20 and C22. Lateral 1/2 of proximal 1/3 of internal surface of both plates much deeper than the rest of the internal surface and showing 2 ridges (=? nl and n3 branches of palmar complex of Jefferies, 1986) projecting from distal wall of co-operculum, running close to each other distally for a short distance, then bending slightly medially and diverging from each other before disappearing abruptly medially (Figs 8C-D, 9A). Proximo-lateral part of co-opercular rim of C20 (Figs 8B, D, 9A) housing small subcircular pit (= pit of lateral line ganglion of Jefferies, 1986), c. 1/3 size of co-operculum and corresponding in position to infundibulum on inner surface of right PM. Straight, poorly pronounced ridge (= ? ventral anterior boundary of posterior coelom of Jefferies, 1986) running obliquely from a point just distal to each co-operculum to median margin of C20 and C22, at c.45? to longitudinal body axis (Fig. 8D). Transversely elongate facet for articulation with PM plate visible along lateral 1/2 of distal margins of C20 and C22, immediately proximo-lateral to co-opercula. Stereom. Internal texture of subcentral and lateral marginal elements of plano-concave surface resembling external texture in the radial arrange- ment of trabeculae and pores (Figs 7A-B, 8A). Stereom of inside of left and right PM (Figs 8C, 10A) consisting of irregular meshwork of stout trabeculae and extremely irregular pores of different shapes and sizes, replaced by more compact texture on septum, scutular rim and bases of apophyseal horns as well as along proximal edge of both PM plates. Elongate, sometimes confluent pores along proximal and distal ridge on lateral half of PM plates. Stereom of excavation for appendage insertion with generally small, rounded pores and thin trab- eculae (Figs 8B, D, 9A, 10A). Texture of inside of C20 and C22 consisting of regularly arranged circular pores, slightly smaller peripherally than 390 centrally. Stereom of co-opercula compact (Figs SB-D, 9A). APPENDAGE. Proximal part. 5-6 tetramerous rings overlapping each other proximo-distally; degree of overlap greater proximally than distally; distalmost ring slightly smaller than others and tightly wrapped around proximal styloid process; latero-distal angles of ring plates gently rounded (Figs 2A-B, 4B, D, F, 5A-D, 6A, C). Intermediate part. Styloid with robust, stout proximal blade, triangular in section, less expanded transversely than distal blade and carrying a blunt apex; distal blade slightly bent towards the proximal blade in lateral view and with small, flat, triangular distal bearing surface. Sharp longitudinal keel running between the two blades, with parabolic outline in lateral view. Lateral styloid surfaces gently concave (Figs 5A-D, 12C). Distal part. Few proximal ossicles preserved (mostly disarticulated), subrectangular in lateral view and apparently poorly developed, blunt apexes, except in the first two ossicles, possibly carrying a small distal bearing surface (Figs 7A,B, 12A-D). Stereom. Minute pores and thin trabeculae centrally on proximal ring plates and styloid; more compact stereom near margins of ring plates and on ossicles. REMARKS. Adoketocarpus acheronticus displays little morphological or ontogenetic variation, as evidenced by small changes in the shape and proportions of several isolated skeletal elements of different sizes, which presumably belong to individuals of different ages, and by the similar proportions of complete specimens. The largest known individuals are either pyriform or ovato-lanceolate, whereas small to medium-sized individuals are only slightly longer than wide and their lateral margins are convex. Such differences are only partly caused by deformation. Deformation is, however, negligible in most cases, as the specimens are often found slightly disrupted but without signs of breakage. Likewise, disarticulated plates lying in proximity to each other are rarely broken; often, they are turned upside down or slightly rotated. The right LOP, A, left and right PLM, left and right PM and C20 and C22 plates are most variable. The right LOP is sutured to the rest of the skeleton in most specimens, slightly displaced in a few specimens and rarely isolated. In the smallest specimen (Fig. 4C,F), the right MEMOIRS OF THE QUEENSLAND MUSEUM LOP is more asymmetrical and elongate proximo-distally than in larger specimens (Figs 4A-B, D-E, 7B, 8A); its distal process is distinctly triangular with a blunt, latero-distal angle, and its latero-distal margins are deeply embayed. Together with the left PM, this is the largest element of the plano-concave surface. The distal process and bilateral symmetry of the right LOP vary in larger specimens. Thus, in some individuals the right LOP becomes almost bilaterally symmetrical (Figs 4D-E, 7B, 8A), whereas in others left and right latero-distal margins of this plate differ in length and orient- ation with respect to the longitudinal body axis, so that its distal process is displaced with respect to this axis (Figs 4A,B, 11A). The outline of plate A varies from subrectangular in small and medium specimens to elongate pent- agonal or subtriangular in large individuals (Fig. 4A-E). The PLM plates have 3-4 sharp, proximal lateral serrations with steep distal margins in small to medium specimens; in large specimens, the serrations are more numerous, blunt-ended and with less steep distal margins; size of the serrations does not seem to change at different ontogenetic stages (Figs 4A-E, 5C). In the smallest individual (Fig. 4C,F), distal margins of the PM plates are much shorter than proximal margins, and the proximal excavation for insertion of the articulated appendage is shallower than in larger individuals. Plates C20 and C22 are 2.5-3 times as long as wide and differ mainly in degree of curvature of their lateral margins and in length and orientation of their medio-distal margin. One isolated C20 (Fig. 8B) has a more elongate and concave medio-distal margin and broad median emargin- ation in its proximal margin. In the same plate, a ridge runs for c. 2/3 plate width from the prox- imal end of its medio-distal margin to a thickened triangular area latero-distal to co-operculum; median 2/3 of ridge almost straight; lateral 1/3 bent strongly towards co-operculum. Shallow areas present just proximal to the ridge, and delimited proximally by a second, fainter transverse ridge running from median margin of C20 to a point c.1/2 along maximum plate width before dis- appearing abruptly laterally. Adoketocarpus janeae sp. nov. (Figs В,С, I2E, 13) ETYMOLOGY. For Jane Jell, who helped collect the material described herein. A LUDLOVIAN MITRATE FROM CENTRAL VICTORIA 391 FIG. 11. A,D.E, Adoketocarpus acheronticus gen. et sp. nov., from NMVPL 1927. A, plano-concave surface of QMF37208, x9. D,E, plano-concaveand convex surfaces of QMF 37214. x12. D.C, Adoketocarpus janeae gen. et sp. nov., from NMVPL 232, plano- concave and convex surfaces of NMVP149391, x12. MATERIAL. Holotype: NMVP100359. Paratype: NMVP100360 from NMVPL 1924 on Mathieson's Creek north of Kinglake West (=T95 of Williams 1964). NMVP149391, 149392 from NMVPL232. DIAGNOSIS. Lateral margins almost straight. Left DLM and right EXM/ILM subequal in length. Left LOP and right DLM smaller than other marginal plates and approximately equal in size. Left LOP median to left latero-distal angle of plano-concave surface. Plate A narrow and elongate. Plate C slightly wider proximally than distally. DESCRIPTION. General aspect. Only those skeletal features which distinguish 4. janeae from A. acheronticus are discussed in detail here. Lateral margins of body straight for most of their length and only slightly converging distally. Left DLM and right EXM/ILM much longer than wide and of similar shape and size: distal margin of left DLM at 45? with longitudinal body axis. Free margins of left LOP (distal in position) and of right DLM gently convex; free margin of right DLM about 3 times as long as medio-proximal margin. Left LOP lying median to left latero- distal angle of plano-concave surface and contributing only to a small extent to left lateral 392 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 12. A-D. Adoketocarpus acheronticus gen. et sp. nov., [rom NMVPL1927. «10. A.B. disrupted body and appendage QMF37202. C, isolated styloid and ossicles QMF37213. D. partial convex surface and ossicles QMF37212. Е, Adoketocarpus janeae gen. et sp, nov., disrupted convex surface of NMVP149392 from NMVPL232. x10. A LUDLOVIAN MITRATE FROM CENTRAL VICTORIA 393 FIG. 13. Adoketocarpus janeae gen. et sp. nov., from NMVPL 1924. A, NMVP100359, holotype, almost complete, slightly disrupted, plano-concave surface, x10. B, NMVP100360, partially disrupted, incomplete, plano-concave surface, x10. body margin. Right LOP with semi-elliptical distal process and markedly sinuous proximo-lateral margins, and c.1/2 as large as С; left latero-distal margin of right LOP at с.120° with free margin of left LOP. Lateral 1/2 of proximal margin of each of the 2 PM plates passing gradually into median 1/2: the latter only slightly concave and meeting its antimere at c.120*; suture between left and right PM straight. C slightly wider proximally than distally. Suture between A and C gently concave leftward throughout most of its length. Plate A at least 3 times as long as wide and in contact with left arm of distal, chevron-shaped margin of left PM along very short suture. Robust. sharp serrations partly visible along disrupted lateral margin of right PLM inthe less complete ofthe 2 specimens Observed. Stereom structure of coarse pores and irregular, sinuous trabeculae: pores decreasing in size towards plate sutures. Measurements. NMVP100359 (Fig. 13A) is 5.7mm wide and 7.8mm long. REMARKS. Attribution of the Lower Devonian species to Adoketocarpus is based on the plating pattern of the plano- concave surface, but only 4 individuals permit limited comparisons with the type species. The differences between the Lower Devonian and the Upper Silurian taxa warrant a distinct specific assignment for the former. Such features of A. janeae as shape and relative proportions of left LOP and left DLM and high length/width ratio of A are similar to those in Æ. savilli Beisswenger, 1994 from the Llandeilo of Morocco (Fig. ЗС). In E. savilli and 4. acheronticus, the left LOP builds the left latero-distal angle of the plano-concave surface and the most proximal part of its lateral margin contributes to the left lateral body wall. In both species, the left latero-distal margin of the right LOP and the free margin of the left LOP meet at an obtuse angle. In А. janeae, the left LOP lies almost completely median to the left latero-distal angle of the plano-concave surface and its free margin is at c.120? with the left latero-distal margin of the right LOP. Adoketocarpus janeae shows a more bilaterally symmetrical body out- line than either.4. acheronticus or E. savilli (Figs 3C.D. 13A). although elements of asymmetry are evident in the arrangement of the distal marginal 394 plates of the plano-concave surface and in the different length of the left and right lateral margins of the body. The higher degree of bilateral symmetry of A. janeae is also observed in the shape of the right LOP, the distal process of which lies only slightly to the left of the longitudinal body axis. Because the only convex surface of A. janeae is not well preserved (Fig. 12E) its degree of asymmetry is uncertain. COMPARISONS BETWEEN ADOKETOCARPUS AND OTHER PARANACYSTIDS Members of the Paranacystidae Caster, 1954 (Ruta, 1997c) are readily identified by several features such as the plate configuration of the convex surface and the 2 unequal subcentral plates on the plano-concave surface. As defined by Caster (1954) the family includes Parana- cystis petrii Caster, 1954, P. simoneae Ruta, 1997c, Adoketocarpus acheronticus and A. janeae. Haude (1995) placed Yachalicystis triangularis from the Lower Devonian of Argentina in the Paranacystidae, based on the 2 large, proximo-distally elongate plates on the convex surface and on details of the stereom. Ruta (1997c) tentatively assigned the Middle Devonian Dalejocystis casteri Prokop, 1963 from Bohemia to the Paranacystidae, based on its overall resemblance to Y. triangularis. However, both Y. triangularis and D. casteri are too poorly preserved for their affinities to be fully evaluated, and will not be considered further here. Because the material of A. janeae is incomplete, the following discussion focusses mainly on А. acheronticus, except where a direct comparison between the Lower Devonian form and other paranacystids is necessary. The body outline is more asymmetrical in A. acheronticus than in either Paranacystis species. With the exception of the left and right PLM, the remaining lateral marginal plates of the plano- concave surface of A.acheronticus do not form pairs of elements of similar shape and size, and are smaller than either left or right PLM; conversely, in both Paranacystis species, the lat- eral marginal elements immediately distal to PLM are almost mirror images of each other and comparable in size with these plates. In A. janeae, the left DLM and the right EXM/ILM are similar in shape and size. A. acheronticus differs from both Paranacystis species in that plate A is only slightly longer than MEMOIRS OF THE QUEENSLAND MUSEUM wide and much smaller than C; in addition, C is much wider in A. acheronticus than in either Paranacystis species; in P. simoneae, C has a robust, distal triangular process (Ruta, 1997c). As in P. simoneae, the 2 PLM plates of А. acheronticus have a lateral longitudinal row of denticulations. In P. simoneae, these are smaller than in A. acheronticus and shaped like sub- elliptical, proximo-distally elongate knobs confined to the proximal 2/3 of left and right PLM. In both Paranacystis species, the proximal excavation for appendage insertion is wider and deeper than in A. acheronticus. The distal, subtrapezoidal plates of the convex surface are much larger with respect to C20 and C22 in Р petrii (convex surface not preserved in P. simoneae) than in A. acheronticus; in addition, the suture formed by the 2 subtrapezoidal plates is shorter in the latter than in P. petrii. For Caster (1954), the distal plates of P. petrii were imbricate in life and acted as an ostial cover; he considered this feature diagnostic of the family. A. acheronticus suggests that ostial elements identified by Caster (1954: figs la-e, 2b) in P. petrii are homologous with the left LOP, right DLM and/orright LOP plates. In A. acheronticus, the right LOP is tightly sutured with the rest of the plano-concave surface and does not seem to have formed a flexible articulation in life. In a disrupted paratype of P. petrii (Caster, 1954, fig. 3b), the distal 1/2 of the internal mould of plano-concave surface shows a subpentagonal plate sutured on the right with a small trapezoidal element. These 2 plates correspond to the right LOP and right DLM of 4. acheronticus respectively, based on shape and position (see Caster, 1954, fig. le). In A. acheronticus the distal margins of the left LOP and right DLM are partly visible when the convex surface is oriented towards the observer, suggesting that the so-called ostial elements of P. petrii are likely to be displaced and/or disrupted plates of the distal margin of the plano-concave surface. Distal plate configuration ofthe plano-concave surface in A. acheronticus differs from that of P. simoneae, in which C carries a distal triangular process. The shape and position of this process resemble those of the right LOP in 4. acher- onticus; homology of this C process is uncertain. In P. simoneae, a small pit near the right proximal angle ofthe C process occupies the same relative position with respect to the body orifice as the subcircular depression on the underside of the right latero-distal angle of the right LOP of A. A LUDLOVIAN MITRATE FROM CENTRAL VICTORIA acheronticus. However, ibe distal part of the plano-concave surface of P. simoneae js not well-known; Ruta (1997c) mentioned the pos- sibility that some skeletal elements (possibly the left LOP and right DLM) in the only known specimen of this mitrate are either displaced or not preserved. А. ucheroiticus permits a more accurate diag- nosis of the Paranacystidae. The distal elements of the plano-concave surface did not act as an оза! cover, as surmized by Caster (1954) in Р petrii. Rather, this part of the skeleton was rigid апа roofed over the body orifice. The distalmost region of the convex surface, on the other hand, was probably flexible, as indicated by the small, irregular plates surrounding the orifice platelets. This flexible integument is documented in sey- eral mitrates (Ubaghs, 1967, 1979: Jefferies & Lewis, 1978; Kolata & Jollie, 1982; Jetfenes, 1986; Cripps. 1990: Parsley, 1991; Beisswenger, 1994; Ruta, 1997b). DISCUSSION. In 4. ucheranticus the simplified plating pattern of the proximal 2/3 of the convex surface is probably a derived condition compared with that of the mitrocystitids and anomalocystitids. On the other hand, the pit an C20 is shared with several mitrocystitids (Chauvel, 1941; Ubaghs, 1961, 1967, 1979, 1994; Cripps. 1990; Ruta. 1997b) which are con- sidered by many workers to be more primitive than the anomalocystitids (Jefferies. 1957. 1968. 1973, 1986. 199]; Craske & Jefferies, 1989; Cripps. 1990: Beisswenger. 1994: Ruta, 1997b. in press: Ruta & Theron, 1997; but see Ubaghs, 1979 and Parsley, 1991). A twisted orifice at a high angle to the axis of the body is found in Mecuinocarpus dentiger (Ruta, 1997b), in young individuals of Mitra- cvstires mitra (Jefferies, 1968; Ubaghs, 1967) and in Eumitrocystella savilli (Beisswenger, 1994) among the mitrocystitids, although less pronounced asymmetry of this structure is ob- served in other mitrates (Thoral, 1935; Ubaghs. 1967, 1969, 1979; Jefferies, 1986). There are fewer tightly sutured polygonal elements on the plano-concave surface of 4. acheronticus than on other mitrocystitids ar anomalocystitids, The 2 subcentral elements on the plano-concave surface also occur in £. savilli and mosi anomalocystitids, although this con- dition can be shown to have evolved in parallel in the 2 latter groups (Beisswenger, 1994; Ruta & Theron, 1997; Ruta, in press). In particular, the pasition of plate A. between C and the left lateral marginal elements of the plano-concave surface. recalls the skeletal configuration of the allani- evtidiid anomalacystitids (Ruta & Theron, 1997), The fewer plates of the convex surface and large C20 and C22 also occur in peltocystid mitrates (Thoral, 1935; Ubaghs, 1967. 1969; Jefferies, 1986: Parsley. 1991); however, (hese differ from 4, «cherónticus in several aspects of the plating pattem of the plano-concave surface A. ucheronticus and E. savilli have 2 subcentral, unequal plates on the plano-concave surface and à distal plate roofing over the orifice. In mast mitrocystilids and all anomalocysititids, the orifice is delimited by a row of distal marginal plates on the plano-cancave surface, consisting of a median (MOP) and two lateral (LOP) elements, Based on topological similarity, Beiss- wenger ( 1994) hypothesized that MOP (=platen) was lost in E. savilli, and that the distalmost clement of the plano-concaye surface 1s an en- larged and medially displaced right LOP (= plate c of Beisswenger. 1994). This hypothesis was supported hy three arguments, two of which also apply to 4. acheronticus. His first argument js based on the reconstructed internal features of the distalmost plate of the plano-concave surface of E. savilli, which '... carries the beginning of the oblique ridge and builds part of the lateral [body] wall as in other nitrates’ (Beisswenger, 1994: 448). Part of the internal side of the plano-concaye surface cun be observed in 4. aeheronticus (Figs 7A-B. 8A) where the distal end of the oblique ridge is partly visible on the proximal 1/3 of the internal surtace of right LOP. The right LOP is larger than the elements lying immediately proximal tà и, shows the same spatial relationships with these as in £, suvilli and closely resembles the like-named plate of the latter in its general shape and in the dev- elopment of a blunt distal projection covering the orifice platelets. The right LOP of 4. acheron- licus does not contribute to the right lateral body wall io the same extent as its proposed homo- logue in E. sevilli, in that itis slightly displaced to ihe left with respect ta the main body axis, presumably as a result of increased torsion of the orifice caused by the loss of a left lateral marginal clement. 1n А. acheronticus, almost the complete course of the oblique ridge is visible on the convex surface as a result of compaction (Figs 4h, бА, C-D, 7B. 9B). showing that the distalmost end af the ridge lies on the proximal part of the internal surface of the right LOP. Ori the plano-concave surface, the suture between the righr LOP and C (= plate 12 of Beisswenger, 1994) straddles the distalmost portion of the oblique ridge. The distalmost part of the ridge appears as an impression on the median hulf of the right distal subtrapezoidal plate of the convex surface. The second of Beisswenger's (1994) ar- guments is an hypothesis of morphological transformation inferred from the first argument: loss of MOP and increase in size and leftward displacement of the right LOP would explain leftward twisting of the orifice. If correctly in- terpreted, this transformation is more evident in A. acheranticus, where the orifice opening forms an angle of about 30° with respect to the main body axis. than in Æ savélli, where this angle is about 45°. Beisswenger's (1994) third argument is not directly applicable to 4. acheronticus, as it in- volves comparison between plate organization on the plano-concave surface of E. зам (Fig. 3С) and Mitrocystella. incipiens (Fig. 3B) which closely resemble each other in proportions and relative positions of the subcentral and of lett and right lateral marginal plates. Comparing these similar arrangements to the more primitive arrangement in M. burrandei (Fig. ЗА), major differences in the distal 1/3 of the plano-concave surface ure interpreted as resulting from loss of plate В (= plate 11 of Beisswenger, 1994) in £, savilli. Plate configuration of the distal 1/3 of the plano-concave surface in 4. aclheraniicus matches that of E. savili, and suggests that B may be secondarily absent in the Australian taxon, The number of lateral marginal plates in 4. acheronticus 15 reduced by one on the right and on one the left with respect 10 E, savilli. and the relative size and spatial relationships of the elements lying to the left of C are very similar to those of the left LOP, left DLM and A plates (= plates b, 3 and 10 of Beisswenger (1994). res- pectively). Based on their shape and position, proximal left and right lateral marginal plates correspond ta PLM (= plates | and f of Betss- wenger (1994), respectively). Assuming the correct identification of the other plates, it is reasonable to hypothesize that the left ILM (= plate 2 of Beisswenger. 1994) was last in A. uchernticus, and that such a loss brought A in contact with the lefi PLM and left PM. The right lateral marginal plates are more dil- ficult to interpret. Comparison with E. savilli indicates that a plate of the right side of the body was lost in 4. acheronticus. This plate might MEMOIRS OF THE QUEENSLAND MUSEUM correspond to the right EXM or ILM (= plates б and 7 of Beisswenger (1994). respectively) in £. savilli (Ruta, in press), but it cannot be identified unambiguously: hence, our right EXM/ILM not- ation for the marginal plate lying to the right of C. Finally, the left and right PM correspond to plates гапа h of Beisswenger ( 1994), respectively, based on their shape, position and internal surface; The resemblance between E. savilli and Ado- ketocarpus has implications for the origin and diversification of the Paranacystidae. We propose that the. Paranacystidae derive from an ancestor resembling E. sevilli through a simplification of the plate configuration. Further steps in this lineage may have been the acquisition of bilateral symmetry and a greater elongation of the body. Compensation for the asymmetrical body ouilinc may have been achieved through modification of the relative proportions of the marginal plates of the plano-concave surface. In A. aeheronticus, plate asymmetries are evident on this surface, but in 4. janeae and in other paranacystids (Caster. 1954; Ruts, 1997c), such asymmetries disappear or are drastically reduced. Bilateral symmetry probably occurred proximo-distally, affecting the marginal elements first (as in A. janeae) followed by the subcentral plates (as in Parunacyytis). CONCLUSIONS Adaketocarpus: |) is the first representative of the Paranacystidae Caster, 1954 recorded trom Australia; 2) from the Upper Silurian predates the next earliest family record wn the Lower Dev- onian of South America (Caster, 1954; Caster & Eaton. |956; Haude, 1995); 3) provides great detail of its extemal and internal anatomy, throw- ing light on several poorly known aspects of the paranacystids and prompting re-interpretation of skeletal organization of the family (Caster, 1954. Haude. 1995; Ruta, 1997c), 4) provides ad- ditional evidence of the affinities between Siluro-Devonian mitrates from Australia und those from the Malvinokaflric Realm (Caster, 1954, 1956, 1983; Gill & Caster, 1960; Caster & Gill, 1967; Philip. 1981; Parsley, 1991; Haude, 1995; Ruta & Theron, 1997: Ruta. 19970): 5) shows that the paranacystids probably evolved from (he paraphyletic mitrocystitids; Æ. savilli closely resembles .4dokerocarpus. especially in the plano-concave surface and in the sirongly twisted orifice. Evolution of the paranacystids was characterized by progressive increase m the bilateral symmetry of the body and of the lateral marginal plates. A LUDLOVIAN MITRATE FROM CENTRAL VICTORIA ACKNOWLEDGEMENTS M.R. thanks the Queensland Museum and its staff for use of facilities and for hospitality, A.R. Milner (Birkbeck College, University of Lon- don) for comments and suggestions, B. Lefebvre (Université Claude Bernard, Lyon) and R.L. Parsley (Tulane University, New Orleans) for stimulating discussions and Julia Day and Simone Wells (Natural History Museum, Lon- don) for encouragement. R.P.S. Jefferies, A.C. Milner, S.J. Culver and L.R.M. Cocks (Natural History Museum, London) read the manuscript. P. Crabb (Natural History Museum, London) took most of the photographs. We thank Ron Parsley and Reimund Haude for their useful reviews but the views set out above are those of the authors alone. M.R. is in receipt of a European Community grant (Training and Mobility of Researchers). The Museum of Vic- toria is thanked for loan ofthe material and David Holloway and Fons Vandenberg are thanked for help with field collecting. LITERATURE CITED BEISSWENGER, M. 1994. A calcichordate interp- retation of the new mitrate Eumitrocystella savilli from the Ordovician of Morocco. Palàontol- ogische Zeitschrift 68: 443-462. CASTER, K.E. 1952. Concerning Enoploura of the Upper Ordovician and its relation to other carpoid Echinodermata. Bulletins of American Paleontology 34: 1-47. 1954. A new carpoid echinoderm from the Paraná Devonian. Anais da Academia Brasileira de Ciéncias 26: 123-147. 1956. A Devonian placocystoid echinoderm from Paraná, Brazil. Paleontologia do Paraná (Centennial Volume): 137-148. 1983. A new Silurian carpoid echinoderm from Tasmania and a revision of the Allanicytidiidae. Alcheringa 7: 321-335. CASTER, K.E. & EATON, T.H. Jr 1956. Microstructure of the plates in the carpoid echinoderm Parana- cystis. Journal of Paleontology 30: 611-614. CASTER, K.E. & GILL, E.D. 1967. Family Allanicytidiidae, new family. Pp. 561-564. In Moore, К.С. (ed.) Treatise on invertebrate paleontology. Part S. Echinodermata 1. (Geological Society of America & University of Kansas: New York). CHAUVEL, J. 1941. Recherches sur les cystoides et les carpoides armoricains. Mémoires de la Société Géologique et Minéralogique de Bretagne 5: 1-286. CRASKE, A.J. & JEFFERIES, R.P.S. 1989. A new mitrate from the Upper Ordovician of Norway, and a new approach to subdividing a plesion. Palaeontology 32: 69-99. 397 CRIPPS, A.P. 1990. A new stem craniate from the Ordovician of Morocco and the search for the sister group of the craniata. Zoological Journal of the Linnean Society 100: 27-71, GARRATT, M.J. 1983. Silurian and Devonian stratigraphy of the Melbourne Trough, Victoria, Proceedings of the Royal Society of Victoria 95:77-98. GILL, E.D. & CASTER, К.Е. 1960. Carpoid echino- derms from the Silurian and Devonian of Australia. Bulletins of American Paleontology 41: 5-71. HAUDE, R. 1995. Echinodermen aus dem Unter- Devon der argentinischen Prákordillere. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 197: 37-86. JAEKEL, O. 1918. Phylogenie und System der Pelmatozoen. Paláontologische Zeitschrift 3: 1-128. JEFFERIES, R.P.S. 1967. Some fossil chordates with echinoderm affinities. Pp. 163-208. In Millot, N. (ed.) Echinoderm Biology. (Academic Press: London). 1968. The Subphylum Calcichordata (Jefferies 1967) — primitive fossil chordates with echinoderm affinities. Bulletin of the British Museum (Natural History), Geology Series 16: 243-339. 1973. The Ordovician fossil Lagynocystis pyramidalis (Barrande) and the ancestry of Amphioxus. Philosophical Transactions of the Royal Society of London, Series B 265: 409-469. 1986. The ancestry of the vertebrates. (British Museum (Natural History): London). 1991. Two types of bilateral symmetry in the Metazoa: chordate and bilaterian. Pp. 94-127. In Bock, G. R. & Marsh, J. (eds) Biological asymmetry and handedness. (John Wiley & Sons: Chichester). JEFFERIES, R.P.S. & LEWIS, D.N. 1978. The English Silurian fossil Placocystites forbesianus and the ancestry of the vertebrates. Philosophical Transactions of the Royal Society of London, Series B 282: 205-323. KOLATA, D.R.. FREST, T.J. & MAPES, R.H. 1991. The youngest carpoid: occurrence, affinities and life mode of a Pennsylvanian (Morrowan) mitrate from Oklahoma. Journal of Paleontology 65: 844-855. KOLATA, D.R. & JOLLIE, M. 1982. Anomalocystitid mitrates (Stylophora, Echinodermata) from the Champlainian (Middle Ordovician) Guttenberg Formation of the Upper Mississippi Valley Region. Journal of Paleontology 56: 531-565. PARSLEY, R.L. 1991. Review of selected North American mitrate stylophorans (Homalozoa: Echinodermata). Bulletins of American Paleontology 100: 5-57. PHILIP, GM. 1981. Notocarpos garratti gen. et sp. noy., anew Silurian mitrate carpoid from Victoria. Alcheringa 5: 29-38. 398 RUTA, M. 1997a. Redescription of the Australian mitrate Victoriacystis with comments on its functional morphology. Alcheringa 21: 81-101. 1997b. A new mitrate from the lower Ordovician of southern France. Palaeontology 40: 363-383. 1997c. First record of a paranacystid mitrate from the Bokkeveld Group of South Africa. Palaeont- ologia Africana 34: 15-25. In press. A cladistic analysis of the anomalocystitid mitrates. The Zoological Journal of the Linnean Society. RUTA, M. & JELL, P.A. 1999, Two new anomalo- cystitid mitrates from the Lower Devonian Humevale Formation of central Victoria. Memoirs of the Queensland Museum 43: 399-422. RUTA, M. & THERON, J.N. 1997. Two Devonian mitrates from South Africa. Palaeontology 40: 201-243. TALENT, J.A. 1967. Silurian sedimentary petrology and palaeontology. Bulletin of the Geological Survey of Victoria 59: 24-29, THORAL, M. 1935. Contribution à l'étude paléont- ologique de l'Ordovicien inférieur de la Montagne Noire et révision sommaire de la faune cambrienne de la Montagne Noire. (Imprimerie de la Charité: Montpellier). 363p. UBAGHS, G. 1961. Un échinoderme nouveau de la classe des Carpoides dans l'Ordovicien inférieur du département de l'Herault (France). Compte MEMOIRS OF THE QUEENSLAND MUSEUM Rendu Hebdomadaire des Séances de l'Academie des Sciences, Paris 253: 2565-2567. 1967. Stylophora. Pp. 496-565. In Moore, R.C. (ed.) Treatise on invertebrate paleontology. Part S. Echinodermata 1(2). (Geological Society of America & University of Kansas: New York). 1969. Les échinodermes carpoides de l'Ordovicien inférieur de la Montagne Noire (France). Cahiers de Paléontologie. (éditions du Centre National de la Recherche Scientifique: Paris). 112p. 1979. Trois Mitrata (Echinodermata, Stylophora) nouveaux de l'Ordovicien de Tchécoslovaquie. Paláontologische Zeitschrift 53: 98-119. 1994, Echinodermes nouveaux (Stylophora, Eocrinoidea) de l'Ordovicien inférieur de la Montagne Noire (France). Annales de Paléontologie. Invertébrés 80: 107-141. VANDENBERG, A.H.M. 1988. Silurian-Middle Devonian, Pp. 103-146. In Douglas, J.G. & Ferguson, J.A. (eds) Geology of Victoria. (Victorian Division of the Geological Society of Australia: Melbourne). 1992, Kilmore 1:50,000 map and geological report. Geological Survey of Victoria Report 91: 1-86, + map. WILLIAMS, GE. 1964. The geology of the Kinglake District, central Victoria. Proceedings of the Royal Society of Victoria 77: 273-328. WITHERS, R.B. 1933. A new genus of fossil king crabs. Proceedings of the Royal Society of Victoria 45: 18-22. TWO NEW ANOMALOCYSTITID MITRATES FROM THE LOWER DEVONIAN HUMEVALE FORMATION OF CENTRAL VICTORIA MARCELLO RUTA AND PETER A. JELL Ruta, M. & Jell, P.A. 1999 06 30: Two new anomalocystitid mitrates from the Lower Devonian Humevale Formation of central Victoria. Memoirs of the Queensland Museum 43(1): 399-422. Brisbane. ISSN 0079-8835. The anomalocystitid mitrates Victoriacystis holmesorum sp. nov. and Pseudovictoriacystis problematica gen. et sp. nov. are described from the Lower Devonian Humevale Formation of central Victoria. V. holmesorum varies consistently from the type species, V. wilkinsi, in the size (larger), shape and proportions of some body plates, the larger more robust spines and the shape of ossicles of distal part of appendage. Some specimens have a sinuous to crook-shaped right spine; others have a proximally geniculate right spine; the left spine is more robust than the right and cigar-shaped. Pseudovictoriacystis problematica has an unusual plate configuration on convex surface, which consists of 14 plates, without interven- ing row II, apparently without C16 and C18, and with a greatly elongate C17. Otherwise it is very similar to V. holmesorum, especially in distribution of terrace- like ridges and shape and proportions of plates on plano-concave surface. O Anomalocystitida, Victoriacystis, Pseudovictoriacystis, Devonian, Australia. Marcello Ruta, Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom; Peter A. Jell, Queensland Museum, P.O. Box 3300, South Brisbane 4101, Australia; received 2 March 1998. Victoriacystis was the first anomalocystitid described from Australia (Gill & Caster, 1960). The type species, И wilkinsi Gill & Caster, 1960, redescribed by Ruta (1997), occurs in the lower Ludlovian of the Dargile Formation near Heathcote and the Melbourne Formation at Hawthorn, Melbourne. Although the material from the Melbourne suburb of Hawthorn has been considered Lower Silurian (Gill & Caster, 1960; Talent, 1967; Ruta, 1997), it occurs in the Melbourne Formation of Vandenberg (1988) and is thus Ludlovian. Its age is, therefore, similar to that of the specimens from Heathcote. An incomplete and poorly preserved mitrate (NMVP16880, 16881) from the Lower Devonian part of the Humevale Formation near Kinglake West was attributed by Gill & Caster (1960) to V. aff. wilkinsi. Ruta (1997) considered it insepar- able from the type species. However, additional, more complete specimens from the same locality show that the Lower Devonian Victoriacystis is specifically distinct. Herein, we revise the diagnosis of Victoria- cystis (Gill & Caster, 1960; Ruta, in press), taking account ofthe Lower Devonian species and newly available specimens of Upper Silurian И wilkinsi which are treated elsewhere (Ruta & Jell, 1999b). A second Lower Devonian anomalocystitid mitrate genus from the Humevale Formation, known from a single, partially disrupted individual, is similar to the new species of Victoriacystis in the plano-concave surface, but has an unusual plating pattern on the convex surface. GEOLOGICAL SETTING Material described, about 30 partially to fully articulated specimens, comes from NMVPL252 (=Davies Quarry (Gill, 1948); =Middendorp’s Quarry (Williams, 1964)) on the western branch of Stony Creek, about 1.6km N of Kinglake West State School and 40km NNE of Melbourne; the site is Lochkovian (Gill & Caster, 1960; Strusz, 1972; Vandenberg, 1988; Vandenberg et al., 1976; Holloway & Jell, 1983; Jell, 1983). Fossils are found in a steely grey pyritic siltstone and are concentrated in a few thin bands interspersed through about 30m exposed in the quarry wall. The diverse fossil faunas are considered to be *... pockets of organic debris ... that do not represent natural assemblages’ (Jell, 1983: 210) based on disrupted bedding, attitude of various fossilised individuals and the great concentration of animals in a few thin beds. Analysis of Gillocystis runcinata (ophio- cistioid), Hillocystis atracta (rhombiferan) and Sphagoblastus adectus (blastoid) (Jell, 1983) suggests that the animals were probably buried when they were still alive and were flattened or 400 slightly crushed by pressure from overlying sediment, depending upon the degree of rigidity of their thecae. Plate dislocation is minimal or does not occur at all. Almost complete absence of skeletal disruption, fractured individual plates, preservation of oral and aboral sides as internally contiguous surfaces (e.g. jaw apparatus of Gillo- cystis against inner aboral surface) and collapse of periproctal plates onto internal side of dorsal thecal surface (e.g. Hillocystis) indicate that sediment did not usually enter the body cavity and that geostatic compression of skeletons occurred soon after burial and before soft tissue decay. Mode of preservation of several individuals of V. holmesorum is similar to that of Sphagoblastus but may differ from that of Gillocystis and Hillo- cystis. Most mitrate specimens are preserved as external moulds, often covered with thin layers of iron oxides. As in Sphagoblastus, the theca of Victoriacystis is a rigid structure composed of tightly sutured polygonal plates. In most cases, both its convex surface and its plano-concave surface are found almost completely articulated. Disruption is minimal and affects mainly LOP, MOP and C1-C9. Such plates generally lie in close proximity to each other and to the rest ofthe skeleton and their mutual spatial relationships are often almost unchanged. Fractures occur usually along lateral margins ofthe plano- concave surface and on plates PLM, C and C20-C22 and are more numerous in the proximal 1/3 of the body, where, as in the case of other mitrates, the skeleton reaches maximum thickness and greatest curvature (Parsley, 1991). Fractures are sometimes visible at junctions between horizontal and subvertical projections of plates DLM, ILM and PLM. In these cases, subvertical projections often lie flush with con- vex surface plates while retaining their mutual contacts with them. In some specimens, the con- vex surface is collapsed onto the plano-concave surface and disruption occurs mainly at level of sutures between lateral plates of convex surface and subvertical projections of DLM, ILM and PLM. C17 is often found sutured with C16 and C18 in contrast with the situation observed in other anomalocystitids in which, when present, this plate is rarely in place (Dehm, 1932; Jefferies & Lewis, 1978; Kolata & Jollie, 1982; Parsley, 1991; Ruta, 1997; Ruta & Bartels, 1998). In a number of specimens, one or, exceptionally, both distal spines are found articulated to DLM, or ata short distance from the body. More frequently, both spines are missing. MEMOIRS OF THE QUEENSLAND MUSEUM Few specimens retain intact appendages, and even in those cases, only proximal and inter- mediate parts are still in place; ossicles and paired plates of distal part are not preserved. Usually, the tetramerous rings of the proximal part are complete but collapsed while retaining their telescopic arrangement. Separation of ring elements is rare as is their preservation as fully undeformed structures. In some specimens, paired plates and ossicles of the distal part are preserved intact and undeformed, albeit rarely articulated with each other. Frequently, plates are disarticulated, collapsed onto the abapical surface of ossicles or missing altogether. Sometimes, paired plates (especially proximal) are found detached from ossicles while still overlapping each other proximo-distally. The ossicles frequently maintain their alignment. Proximal and distal articular surfaces are observed in at least one individual. Although rare and largely incomplete in their proximal 1/2, internal moulds are sometimes as- sociated with partially disrupted external moulds. Unlike other Lower Devonian echinoderms from the Humevale Formation, Victoriacystis has a relatively large, transversely elongate distal orifice through which fine sediment could easily enter the body cavity during burial. Similar pres- ervation is known in other carpoids, both solutes (Jefferies, 1990) and cornute stylophorans (Jefferies, 1968; Woods & Jefferies, 1992). SYSTEMATIC PALAEONTOLOGY External skeletal terminology and plate nomenclature follow Ruta (in press) with modifications as in Ruta & Jell (19992). Description of internal body anatomy is based on Ubaghs (1968, 1969). Morphological term- inology of ossicles is that of Jefferies & Lewis (1978) and Ruta & Theron (1997). The terms ‘apical’ and ‘abapical’ indicate the position of structures close to or away from the ossicular process (or apex) respectively. Specimens are deposited in the Palaeontological Collections of the National Museum of Victoria, Melbourne (NMVP) and the locality is registered in the locality register at the same Museum (NMVPL). Study and illustration of skeletal details was made on latex casts whitened with ammonium chloride. NEW ANOMALOCYSTITID MITRATES FROM VICTORIA 401 FIG. 1. Vietoriacystis holmesorum sp. nov. All plano-concave surfaces showing terrace-like ridges, spines, tetramerous rings, styloid and proximal ossicles. A, NMVP100361. B. NMVP100387b, C, NMVP100385. D. NMVP100369. E, NMVP100373. All *3. 402 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 2. Victoriacystisholmesorumsp. nov. A,E,lateraland plano-concavesurface views of NMVP100382, x5 and x3, respectively. B, appendage in lateral view of NMVP100371, x7. C, plano-concave surface of NMVP 108627, x2. D, plano-concave surface of NMVP100378b, х2. NEW ANOMALOCYSTITID MITRATES FROM VICTORIA 403 FIG. 3. Metoriacystis holmesorum sp. nov. A, left spine of NMVPI 00365. х5. B. right spine and articulation of NMVP100371, *7. C. right spine and articulation of NMYP100367, x5, D. plano-concave surface of NMVP100381. х3. Е. plano-concave surface of NMVP 100384, х3. 404 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 4. Fictoriacystis holmesorum sp. nov. All convex surfaces, showing terrace-like ridges, spines, tetramerous rings, styloid and ossicles. A, NMVP100362, x3. B, NMVP100363 (holotype), x2. C. (distally damaged) NMVP100378b. х2. D, (partial) NMVP 100385, х3. E, NMVP100376, «3. Е NMVTP100374, х2. NEW ANOMALOCYSTITID MITRATES FROM VICTORIA Superorder STYLOPHORA Gill & Caster, 1960 Order MITRATA Jaekel. 1918 Suborder ANOMALOCYSTITIDA Caster, 1952 Family PLACOCYSTITIDAE Caster, 1952 REMARKS. Family groupings in the Anomalo- cystitida are not satisfactory (Parsley, 1991; Ruta & Theron, 1997; Ruta, in press; Ruta & Bartels, 1998). Analysis of character distribution (Ruta. in press) suggests that, with the exception of the Allanicytidiidae Caster & Gill, 1968, all families, including Placocystitidae as defined by Parsley (1991). arenot monophyletic. Parsley (1991) and Ruta (in press) concur in recognising Pic- foriacystis as sister taxon to (Placocystites + Rhenacystis). However, the allanicytidiids are nut closely related to these 3 genera. As Parsley (1991:16) pomted out, their '... assignment to the Placocystitidae is admittedly speculative’. In the light of the revised family concept of Parsley (1991), we restrict this family to Placocvstites, Rhenocystis and Vietoriacvstis only. Victoriacystis Gill & Caster, 1960 TYPE SPECIES, Victoriucystis wilkinsi Gill & Caster, 1960 from the early Ludlow Graptotite Beds, Dargile Formation, Victoria; by original designation. DIAGNOSIS. Rows I-V with 5,4,3,5.3 plates, respectively. C1 and C5 smaller than C2-C4. C17 elliptical to rounded, c.1/2 as long as adjaceat C16 and CIB. Sutures between C15 and Cl6 and between C18 and C19 medially convex. C21 shield-shaped to rhomboidal. deeply but not completely inserted between C20 and C22. B absent. A-C suture oblique. Robust, transverse terrace-like ridges mainly confined to C20-C22 and PLM. Lateral margins of PM convex laterally. Tetramerous rings wider proximally than distally, with fold of palyplated integument between rings. Styloid with median longitudinal keel, proximal blade elliptical in section, distal blade spine-like. Successive ossicles decreasing rapidly in size. Victoriacystis holmesorum sp. nov. (Figs 1-13) Vietoriacystis alf, нама Gill & Caster. 1960:54. pl, 10, figs L3 Vieterideysts wilkinsi Gill & Caster: Ruta. 1997,85. fig ETYMOLOGY. For Frank and Entd Holmes. for their assistance in collecting the material. MATERIAL. HOLOTYPE: NMVP100863. PARATY PES. NMVP100361-100362. 100364-100382, 100384- 100386 all from NMVPL252, 405 DIAGNOSIS. C10. C12 and C14 larger һап adjacent proximal and distal plates, Proximal half of C16 and C18 narrower than distal half. C17 not in contact with either proximal angle of C12 or distal angle of C21. A larger than LOP or MOP. Strongly arcuate or geniculate suture between A and C. Distal portion of appendage not differentiated. Styloid large and robust. Proximal ossicular blades strongly recurved. Paired plates distally in appendage without tubercles. Left spine robust, cigar-shaped, >1/2 as long as body, with lateral cutting edge; right spine more slender. sigmoid to geniculate, without cutting edges, DESCRIPTION. EXTERNAL. Measurements. Holotype: body c.28mm long, 15mm wide. Smallest specimen (Figs 1C, 4D): body c.21mm long. 15пип wide. Largest specimen (Fig. 1E); body c.30min long, 17mm wide. Plano-concave surface. Plano-concave surface subrectangular, slightly wider than convex surface. with sharp (metal кайтпа of I | marginal and 2 subcentral plates (Figs LA-E, 2C-E, 3D-E, 12A), with maximum width at or slightly prox- imal to latero-distal angles of PLM. PM flat, almost às wide proximally as distally. with convex lateral margins except for abrupt curve before joining proximal margin, Curvature of distal margin of left PM usually higher than that of right PM (Figs I A-E, 2A,C-E. 3D-E. 5D, 6E. 12А). PLM, ILM and DLM with flat horizontal projections: well-developed, subvertical projections with gently convex cross-section and meeting plano-concave surface at 60° (Fig, 12D-E). PLM more than twice as wide distally as proximally, with subvertical projection subtrapezoidal, of uniform depth in distal 1/3, decreasing in depth proximally (Figs LA-B, 2A, C-E, 3D. 4B, Е, 5А-С. 12B); ILM trapezoidal. slightly shorter than PLM, with distally diverging, gently concave medial margins: subyertical projection subrectangular, comparable in size with those of DLM and PLM, with sutural margin in 2 shallow embayments of about same size, DLM irregularly pentagonal, with convex medial margins. with longer less convex lateral margins, with medial 2/3 of distal margin thickened and shghtly domed. Sub- vertical projections subpentagonal, with sutural margin in 2 embayments, Distal surface of DLM vertical or sloping distally towards convex surface. subtriangular, shallower peripherally. with subcentral vertically elongate narrow toroidal process for spine insertion sitting on 406 MEMOIRS OF THE QUEENSLAND MUSEUM VIG. 5. Fictoriaevstis holmesorum sp. nov. Convex surfaces showing terrace-like ridges. spines. tetramerous rings. A. NMVP100387a. B. (proximal part only) NMVP100382.C_(2 individualsside by side) NM VP108625. D. extemal ol'plano-concavesurface proximally and inside of convex surfacedistally of NMVP 100376. All «3. л NEW ANOMALOCYSTITID MITRATES FROM VICTORIA 407 FIG. 6. Metoriacystis holmesorum sp. nov. Spines (A-D), plano-concave surface (C.D) and proximal body excavation (E). A, left spine NMVP100367. х5. D. right spine NMVP100374. «5. C, rightspine NMVP 100384. x5. D, right spine NMVP100361. #5. E. NMVP100370, x7. 408 slightly raised, lens-shaped projection, Small gap between lateral margin of each LOP and medio-distal margin of each DLM (Figs 1A-B, 2A,C, 3D, 4A-B,F, 5A, ПА, 12А). MOP and LOP forming almost transverse row along distal margin. Vertical projections about 1/3 as large as horizontal projections, MOP about twice as wide as each LOP, rectangular, with straight distal margin. LOP c.3/5 size of DLM, subpentagonal, with blunt round latero-distal angles (Figs 1 A-D, 2C-D, 6A-B,D, 7A, 12A). Plate A generally sub- pentagonal to shield-shaped, c.1.5 times as large as each DLM, wedged between left ILM and C, with medial margin arcuate or geniculate, with lateral margin twice as long as latero-distal margin. Plate C largest plate in body, c.1/2 as long and wide as plano-concave surface, with medio-distal margin almost 1.5 times as long as latero-distal margin. Small, round tubercles subcentrally on A and on distal 1/3 of C (Figs I A,D, 2C, 3D, 6B,D, 7A). Convex surface. 20 plates in 5 transverse rows. Rows П-У gently concave distally. Rows Land IV with 5 plates each. Rows III and V with 3 plates each. Row П with 4 plates (Figs 4A-F, 5A-C, 12C). Maximum convexity c.2/3 of way proximally along C20 and C22 (Fig. 12B). Plates generally decreasing in size distally, except C15 and C19, which are smaller than C10 and C14. C2-C4 subpentagonal, subequal, larger than subrectangular Cl and C5 and just smaller than hexagonal C7 and C8. Distal margins of C1-C5 broadly convex. Lateral margins of C3 usually concave, sometimes straight (Figs 4A-F, 5A,C, 12C). Proximo-medial and proximo-lateral margins of C7 and C8 and proximal margins of C6 and C9 convex. C6 and C9 subpentagonal, often more expanded transversely than proximo-distally (unlike C2-C4), with sub- parallel laterally diverging (rarely converging) proximal and distal margins (Figs 4A-F, 5A,C, 12C). CIO and CI4 wider than long, subhexagonal, with very short medio-distal and medio-proximal margins. C12 hexagonal, slightly wider proximally than distally (Figs 4A-F, 5A-C, 12C). C15 and C19 subtrapezoidal, with convex margins, much wider distally than proximally (Figs 4A-C, D-E, 5A-C, 12C). C16 and C18 with lateral margins strongly diverging proximally, with latero-distal margins 1/2-1/3 length of medio-distal margins. Suture between C16 and C18 extremely short distally, slightly longer in proximal section. C17 subrounded to subelliptical, <1/2 as long as C16 and C18, never in contact with C12 or C21, flat or slightly raised MEMOIRS OF THE QUEENSLAND MUSEUM above convex surface (Figs 4A-C,E-F, 5A- C, 8D, 12C). C21 shield-shaped, subhexagonal, with gently convex or rarely straight latero-distal margins meeting at obtuse angle, with proximo-lateral margins straight in distal 1/2 and convex in proximal 1/2 (Figs 4A-F, 5A-C, 8D, 10C, 12C). C20 and C22 largest plates on convex surface, almost twice as long as wide, with oblique distal margins, with convexity of lateral margins increasing slightly proximally (Figs 4A-C,E-F, 5A-C, 7B, 10C, 12C). Sculpture. Terrace-like ridges on PLM (Figs 1 A-E, 2A, C-E, 3D-E, 5D, 11C, 12A) and C20- C22 (Figs 4A-F, 5A-C, 8D, 10C, 12A-C), transverse, mostly uniform distance apart, never anastomosing, interrupted abruptly at interplate sutures. Ridges near proximo-lateral angles of PLM, C20 and C22 and near proximal margins of C20 and C22 usually short, interrupted medially, sometimes bifurcating, more crowded together than elsewhere. Bifurcations of ridges rare, in either direction. Ridges on PLM more robust than on C20-C22, with slightly deeper and steeper proximal slope, with distal slope almost flat, slightly convex proximally and gently concave distally. Ridges on C21 variable, with 5-6 most proximal ridges rarely transverse (Fig. 5B), more often convex distally (Figs 4A-C, 8D), with convexity decreasing in straighter more distal ridges, with subcentral ridges straight or chevron-shaped with apex of chevron pointing proximally, with lateral 1/2 of chevron arms transverse or slightly diverging distally, with most distal ridges transverse. Body stereom. Uniformly compact (Fig. 6E) or of minute perforations with irregular outline and no evident distribution pattern (Fig. 8D). Perforations absent or very small along plate margins, frequently replaced by narrow band of short close spaced straight striations. Stereom of centre of plates rarely coarser than peripheral stereom, with more widely spaced larger pores and thicker trabeculae especially on MOP, LOP, A and C (Fig. ТА,р). Stereom of terrace-like ridges on convex surface granular, more compact at free margins. Stereom of ridges on plano-concave surface compact along free margins, coarse and irregularly perforated by small pores on distal 1/2, giving rise to faint striations on proximal 1/2. Striations at c.45? in medio-lateral direction. Spines. Left spine straight, massive, 1/2-5/8 body length, cigar-shaped, tapering to a blunt end in distal 1/3, with cross-section of proximal 2/3 asymmetrical, with semicircular medial margin, NEW ANOMALOCYSTITID MITRATES FROM VICTORIA 409 FIG 7. Vietoriacystisholmesorum sp. nov. A. plano-concavesurface partly disarticulatedand with spine: C. detail of proximal appendage of NMVP100374, х2 and #5, respectively, В. inside of plano-concave surface of NMVP100371.*3. D.F, proximal and distal articularsurfaces of ossicles NMVP108627. » 10. E, inside of distal part of convex surface of NMVP100376, x7. +10 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 8, Vietoriacystis holmesorum sp. nov. A, tetramerous rings and styloid of NMVP100373. 10. B, styloid. ossicles and paired plates of appendage of NMVP100363( holotype), *5. C. inside of plano-concave surface of NMVP100369, х3. D, proximal surface ornament of NMVP100370. х7. NEW ANOMALOCYSTITID MITRATES FROM VICTORIA 411 FIG. 9. Victoriacystis holmesorum sp. nov. Tetramerous rings, styloid, ossicles and paired plates. A. NMVP100378b. x5. B.C, NMVP100366, х3. D, NMVP100364, x5. E, NMVP100387b. х7. 412 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 10, FHetoriacvstis holmesorum sp. nov. Convex surface (C). terrace-like ridges (C), ossicles and plates (A.B.D). A.C. NMVP100389, х6 В.р, NMVP100366, «15 NEW ANOMALOCYSTITID MITRATES FROM VICTORIA 413 FIG. 11. Victoriaeystis holmesorum sp. nov. Internal side of central part of convex surface (A.B) and transverse section of terrace-like ridges of plano-concave surface (С). A.B, NMVP100367, «3 and «12, respectively. C. NMVP108627, «10. 414 with sharp lateral margin becoming more indistinct distally, with distal 1/3 circular or broadly elliptical in section (Figs 3A, 4A, 6A, 11A), with oblique gently convex, surrounded medially by slightly thickened margin and carrying low conical subcentral projection, giving insertion to toroidal process (Figs 2E, 3A, 5D, 6A, 7E). Right spines described individually: some (Fig. 3B-C) more slender, slightly shorter than left spine, uniformly decreasing in diameter proximo-distally, slightly flattened in proximal 1/2, with weak lateral keel, with proximal 1/4 separated from rest of the spine by abrupt geniculation; with distal 3/4 of spine straight or very gently curved medially; most others gently (Figs 2D-E, 3D-E, 5A-D, 6B-C, 7A-B) to strong- ly convex laterally (Figs 1A, 5C, 6D) proximally, laterally concave distally, forming a distinct distal hook in one specimen, with almost uniform cross-section proximo-distally and with blunt distal end, with articular surface similar to that of left spine, but perpendicular to main axis and without thickened medial margin. INTERNAL. Plano-concave surface. Slightly oblique septum along left 1/2 ofinside of C and of left PM, slightly sinuous in its distal 1/2, ending in poorly defined spur-like process subcentrally on distal 1/3 of C, with proximal part of septum convex laterally. Proximo-distally elongate, trough-like area of inside of left PM between septum and lateral margin of left PM. Faint ridge diverging laterally from the spur-like process, running on inside of C and right PM, flanking the lateral margin of the latter. Transverse ridge near distal margins of A and C, slightly concave distally, more robust on A, occupying almost the entire width of A, interrupted laterally on C. Proximal and distal surfaces of ridge sloping gently. Inside of other plates poorly preserved but apparently smooth. Convex surface. Internal surface of C1-C5 margins slightly thickened (Fig. 5D), surrounded by very shallow peripheral groove, with distal 1/3 (Figs 5D, 7E) occupied by proximally recumbent walls at c.20° to internal surface of plates, extending almost completely across their width. Distal surfaces of transverse walls merging gradually into distal 1/3 of internal side of plates, also slightly raised with respect to their proximal 2/3. Walls on C1 and C5 c.1/2 as wide as walls on C2-CA, slightly thicker than these, torus-shaped with strongly convex, blunt free margin. Walls on C2-C4 rectangular, with straight, thinner free margin and with clear-cut separation of lateral MEMOIRS OF THE QUEENSLAND MUSEUM margins from inside of plates. Internal surface of C6-C9 smooth and featureless. Subcentral, proximo-distally elongate thickening on internal surface of C10, C12 and C14. Lateral surfaces of thickenings almost vertical. Thickenings on C10 and C14 roughly elliptical, c. twice as long as wide. Thickening on C12 almost 3 times longer than wide, with subparallel lateral margins. Similar thickening, spindle-shaped in outline and with greater axis at c.60? to body axis, visible in the middle of plate C15 in NMVP100367 (Fig. 11A-B). Internal surface of C16 (Fig. 11A-B). Distal 1/2 not clear. Proximal 1/2 with a shallow cruciform area with irregular arms, outlined by thin ridges, Proximal arm short and stout, with gently concave margins, reaching proximo-lateral margin of C16. Distal margin of lateral arm curving distally as sinuous ridge joining lateral margin of distal arm of cross. Distal arm about twice as long as lateral and proximal arms. APPENDAGE. 1-1.25 times body length; proximal part 3 times wider at proximal ring than at distal ring, with segments of proximal 1/3 of distal part decreasing in size and changing shape distally. Segments of central 1/3 of distal part decreasing uniformly in size and displaying less remarkable changes in shape. Distally appendage whip-like, with segments of approximately constant proportions. Proximal part. Tetramerous rings with proximal one largest, with remainder of subequal length but decreasing width. Paired ring plates on plano-concave side, slightly smaller than those lying on opposite side, less convex in cross-section, with narrower thickening along distal margins (Figs 1 A-B,E, 2A,D-E, 3D, 4A-B,F, 5B, 7B, 8A, 9A-C,E, 12A-C). Distal margin of proximal ring straight, without tubercles. Distal margins of other rings concave, with regularly spaced, subconical to hemispherical tubercles (Figs 1E, 2C-D, 3D, 4A-C, F, 5A-B, 8A, 9A-C,E). Articular surfaces on lateral ends of ring plates on same side as convex surface, proximo-distally elongate, triangular, with a subcentral, roughly circular shallow area (Figs 1B,E, 2C-D, 8A, 9A). Intermediate part. Styloid (Figs 1B,E, 2D, 4B,F, 7A,C, 8A,B, 9A,E, 12A-B, 13B) robust. Proximal articular process just beneath proximal blade, subhorizontal, flattened, subtriangular. Proximal blade expanded transversely, recumbent, with free margin blunt, semicircular to broadly semielliptical. Narrow, sharp, median 415 NEW ANOMALOCYSTITID MITRATES FROM VICTORIA ALII D m zz EE = Е БЕ SS da plano-concave surface. D, le > edspinesomitted). E, proximal surface (appen „distal surface (distally articulat: FIG. 12. Reconstruction of Victoriacystis holmesorum sp. nov. A convex surface. D 416 longitudinal keel ending abruptly before reaching free margin of proximal blade and widening slightly in its distal 1/2. Region of blade facing proximally between free margin and insertion of proximal articular process vaulted. Distal blade 3 times as high as wide, spike-like. Proximal margin of distal blade narrowly acute but not sharp, slightly concave in lateral view in its lower 2/3 and almost straight in its upper 1/3 before merging into blunt blade apex. Lateral surfaces of distal blade flat to very gently concave in upper 2/3. Distal part. (Figs 2B,D, 4B,F, 7A-C, 8B, 9A-E, 10A-B,D, 12A-C, 13A,C-D). Ossicles of first 5-6 segments slightly more than 3 times as high as long and divided into massive body and blade- like process, or apex. Processes of first 3 or 4 ossicles (Fig. 13A) almost as high as their bodies, gradually decreasing in height and length distally, wedge-shaped in transverse section, with strongly curved proximal margin. Proximal margins of processes merging into sharp median longitudinal keel with asymmetrical parabolic profile and occupying proximal 1/3 (in first 3 ossicles) to 1/2 (in subsequent 2 or 3 ossicles) of ossicular length. Articular margins for insertion of paired plates bearing smaller, shallower proximal facet and larger, deeper distal facet. Ossicles 7-9 (Fig. 10D) 3 times as high as long; processes narrower than on more proximal ossicles, decreasing in size distally, confined to distal 1/3-1/4 of ossicular length, subtriangular in lateral view. Keel slightly concave to straight in lateral view. Remaining ossicles (Figs 9B-D, 10A-B) subrectangular in lateral view, 3 times as high as long, with almost straight proximal and distal margins; processes diminishing distally, cuspate, shaped like an equilateral triangle in cross-section. Terminal ossicles without processes. Proximal (Fig. 13C) and distal (Fig. 13D) articular surfaces parabolic, slightly taller than wide, comprising an outer interossicular depres- sion, an inner interossicular depression and a median interossicular groove. Outer interossic- ular depression delimited laterally by slightly thickened ossicular margin of uniform width and medially by thinner, ascending ridges forming external subelliptical boundary of inner inter- ossicular depression, becoming thinner and narrower apically. Inner interossicular depression elliptical, occupying most of apical 2/3 of articular surface. Internal boundary of depression delimited by faint vertical ridges flanking median interossicular groove. Abapical MEMOIRS OF THE QUEENSLAND MUSEUM ends of ascending ridges turned abruptly laterally, merging into free, semicircular rims of 2 articulation bosses on proximal articular surface of ossicle. Ridges marking proximal margins of subtriangular facets for articulation with paired plates of preceding segment of distal part of appendage. Distal articular surface of ossicles (Fig. 13D) differing from proximal articular surface in apically tapering median interossicular groove, comparatively deeper lateral regions of outer ossicular depression and trough-like, elongate articular pit delimited by 2 thick ridges diverging apically and laterally, occupying same position as articular bosses on outer ossicular depression, Paired plates of distal part of appendage chang- ing shape and size throughout appendage length (Figs 2D, 4B,F, 8B, 9A-C, 10A-B,D). Each plate articulating with distal facet of overlying ossicle and with proximal facet of next distal ossicle. First 4 pairs of plates at least twice as high as long, with gently convex distal margin, with sinuous to straight proximal margin. Subsequent 4 pairs of plates with decreasing height/length ratio almost as long as high. Remaining plates subsemicircular in lateral profile. Appendage stereom. Stereom of proximal rings generally compact, especially along free margins of ring plates, or with small subcircular pores, resembling that of central surface area of body plates, often with small granulations distributed randomly, rarely with coarse trabeculae and irregular pores. Styloid stereom usually compact, rarely microperforate, sometimes with irregular, elongate trabeculae near margins of lateral surfaces of distal blade. External surface of plates and ossicles of distal part of appendage usually covered with regular, subcircular shallow pits separated by short trabeculae. Stereom near free margins of ossicles and plates generally more compact, with smaller pits and more irregular trabeculae, the latter sometimes elongate and giving rise to faint striations, especially near apex of ossicular processes. DISCUSSION. Victoriacystis holmesorum closely resembles V. wilkinsi and Rhenocystis latipedunculata Dehm, 1932 especially in the number and arrangement of plates on the convex surface (Ruta, 1997, in press; Ruta & Bartels, 1998). Rhenocystis is distinguished from Victoriacystis by the plate arrangement of row Il, in which C7 and C8 are separated by interposition of C3 and C12. И holmesorum is larger and more robust than V. wilkinsi and R. latipedunculata. It NEW ANOMALOCYSTITID MITRATES FROM VICTORIA FIG. 13. Reconstruction of Victoriacystis holmesorum sp. nov. A, most proximal segments of distal part of appendage. B, styloid in left laterodistal view. C,D, proximal and distal articular surfaces of ossicle of intermediate region of distal part of appendage. differs from V. wilkinsi in its differentiated distal part of the appendage, larger plates of row Ш, C10 C12 and C14, with respect to more proximal and distal elements, narrower proximal 1/2 of C16 and C18 with respect to distal 1/2, lack of contact between C17 and either proximal angle of C12 or distal angle of C21, larger A with respect to LOP and MOP, usually strongly arcuate or geniculate suture between A and C, more robust and larger styloid, more strongly recurved proximal ossicular blades, absence of tubercles from the paired plates of the distal part of the appendage and longer and more robust spines, sometimes with remarkably different morphology. Pseudovictoriacystis gen. nov. TYPE SPECIES. Pseudovictoriacystis problematica sp. nov. 417 ETYMOLOGY, Greek pseudo, false, plus Victoriacystis; alluding to resemblance between new genus and Victoriacystis. Feminine. DIAGNOSIS. Distal 1/2 of convex surface of rows Land III, with 5 and 3 elements respectively. Distal margin of convex surface slightly convex centrally, concave laterally. C12 in contact with C2-C4. Lateral margins of C12 diverging slightly distally. C10 and C14 larger than C12, sutured with СІ-С2 and with C4-C5, respectively. Proximal 1/2 of convex surface ofrows IV and V, the former including C15, C17 and C19. C15 and C19 subtrapezoidal, their lateral margins 1.5 times as long as medial margins. C17 shield-shaped, at least 3 times as wide distally as proximally, with 3-lobed distal margin. C20 and C22 almost 1/2 as long as body, with strongly sinuous medial margins and very gently convex lateral margins. C21 narrowly inserted between C20 and C22, at least twice as long as wide, reaching maximum width in proximal 1/3. Pseudovictoriacystis problematica sp. nov. (Figs 14-16) ETYMOLOGY. Greek problema, question; alluding to the puzzling plate configuration of the convex surface. MATERIAL. HOLOTYPE: NMVP100383 from NMVPL232. DIAGNOSIS. As for genus. DESCRIPTION. Length 24mm, width 16mm, broadly rectangular. DLM and LOP of left side, C15, spines and articulated appendage not preserved. Plano-concave surface. Plating pattern and gen- eral plate proportions similar to Victoriacystis holmesorum. Differences as follows (Figs 14A, 15A): maximum body width about halfway along PLM; lateral margins of ILM and DLM straight; shallower lateral body walls; much greater PLM/ILM length ratio; length of distal margin of PLM/length of proximal margin of PLM slightly lower; proximal 1/3 of lateral margins of PLM more strongly curved medially; distal margins of PLM sinuous, with medial 1/2 proximally convex and lateral 1/2 proximally concave; PM about as long as C; lateral margins of PM more gently convex and with most proximal part forming deep semicircular notch; proximo- lateral angles of PM extended as narrow lateral elongate processes; proximal margins of PLM and PM at obtuse angle rather than merging gradually; medial margins of ILM slightly more concave; DLM about twice as long as wide, with 418 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 14. Pseudovictoriacystis problematica gen. et sp. nov. Holotype, NM VP100383. A.B, plano-concave and convex surfaces, respectively, х3, C, close-up of C20-C22, x10. NEW ANOMALOCYSTITID MITRATES FROM VICTORIA 419 ү! 1 "i ne A Sorin bavi an FIG. 15. Pseudovictoriacystis problematica gen. et sp. nov. Holotype, NMVP 100383, A. close-up of night LOP MOP and articular surface of right DLM. > 15. В. close-up of distal part of convex surface. *10. 420 distally protruding articular region for spine insertion and shallow depressed area medial to articular region; distal surface of articular region teardrop-shaped, with lateral end forming an acute angle and divided into 2 distinct parts; lateral part broadly triangular and shallowly concave; medial part subcircular and markedly convex distally, bearing small, subcentral toroidal process giving rise to a short. faint medial and lateral ridge; suture between A and C mostly straight, at c.60° to longitudinal body axis, gently curved and meeting proximal margin of MOP in distal 1/4. Convex surface (Figs 14B-C. 15B, 16). Slightly disrupted: proximal 1/2 reconstructed tentatively. Transverse rows of plates 4: distal row with 5 plates, homologous with Cl-C5 of other anomalocystitids (e.g. Rhenocystis and Victoriacystis). C2 and C4 subtrapezoidal. with distally converging lateral and medial margins and with lateral 2/3 of proximal margins broadly convex and lying proximal to proximal margin of C3. Cl and C5 subrectangular about twice as long as wide, with broadly concave distal margins, Row II of 3 plates (7C10. C12 and C14 based on shape and proportions). C10 and C14 longer than wide. subpentagonal. with medial angles strongly developed into points. C12 in contact with C3 medially and C2 and C4 laterally. Lateral margins of C12 mostly straight, slightly convex at proximal and distal ends, C15 and C19 smaller. subtrapezoidal, with convex margins. C17 with L/W ratio of 1.4, with lateral margins divided into sinuous distal 1/2 and distally diverging concave proximal 1/2. with distal margin broadly convex, with expanded central section. C20 and C22 with distal to proximal margins; distal margins «1/3 plate length. oriented slightly oblique to longitudinal axis. slightly concave; lateral margins very gently convex in distal 1/2, more strongly convex in proximal 1/2. Proximal margins merging into lateral margins. forming proximally protruding blunt-ended angles, with lateral 2/3 of proximal margins slightly convex. with medial 1/3 gently sinuous, forming central excavation into proximal margin. Medial margins convex but with smooth invagination distal to rounded medial projections at proximal end. C21 (Fig. 14B-C) trapezoidal. inserted between C20 and C22, with gently convex proximal marpin, with 4-5 irregular ridges. with one bifurcated laterally. Sculpture. PLM with robust terrace-like ridges. with thickened free margin; ridges irregularly sinuous, unevenly spaced, sometimes interrupted MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 16. Reconstruction of convex surface of Pseudo- victoriacystis problematica gen. et sp. nov. medially or giving rise to irregular lateral bi- furcations, with most proximal ridges short and at c.45° to body axis, with smooth broadly trapezoidal area of external surface (Fig. 14A). Terrace-like ridges on C20 and C22 (Figs 14B-C) less steep than on PLM, without thickened free margin, oriented mainly transverse to long- itudinal body axis but irregularly sinuous. Single, sigmoidal ridge at 45° to longitudinal axis on proximolateral 1/4 of external surface of C19. Body stereom. Stereom of plano-concave surface similar to that of lictoriacystis holmesorum. but consisting of slightly coarser meshwork, especially in the centres of the plates, often form- ing a reticulate pattern of irregular pores and trabeculae. Peripheral surface. especially on A. C. MOP and ILM, with elongate pores and thin. straight trabeculae arranged radially (Figs 14A. NEW ANOMALOCYSTITID MITRATES FROM VICTORIA 15A). Stereom of convex surface retiform except on C20-C22 (Fig. 15B-C). All other plates, but especially those of the 2 most distal transverse rows, with distinct surface pattern with pores increasing in size, becoming more elongate and arranged more regularly from centre to margins of plates, separated by radiating trabeculae, often dichotomously branching towards the periphery. Pores near plate margins polygonal, separated by short trabeculae, giving rise to coarse, cancellate surface pattern of stereom. Stereom of periphery of C17 and of medial 1/2 of C19 of densely spaced, circular to subelliptical pores, without obvious radiating arrangement of trabeculae. REMARKS. The possibility that NMVP100383 is a teratological individual of Victoriacystis holmesorum cannot be entirely ruled out. It closely resembles the latter in plating of the plano-concave surface. Although rare, pathological mitrate specimens are known (Ruta, 1998). However, in all known cases, abnormal individuals can be assigned to known species, based on their possession of most of the characters shared with normal individuals. Pending discovery of additional specimens to confirm diagnostic features, NMVP100383 is here placed in a new taxon because the config- uration of its convex surface reveals a unique combination of attributes not observed in any other species. Plate arrangement of distal 1/2 of convex surface resembles that of Placocystites forbesianus de Koninck, 1869 (Jefferies & Lewis, 1978) from the English Wenlock. In Pseudovictoriacystis and in Placocystites, C10, C12 and C14 are sutured with row I, there is no intervening row II and C12 is slightly smaller than both C10 and C14. Pseudovictoriacystis differs from Placocystites in being much longer than wide, in displaying a more limited distribution of terrace-like ridges, in possessing 5 rather than 3 plates in row I and in showing a much simpler plate configuration on proximal 1/2 of convex surface, apparently without plates C16 and C18 and with enlarged C17. Further- more, C3 is separated from C10 and С14 by interposition of C2 and C4 and appears to be comparable in size with C1 and C5 and slightly smaller than C2 and C4. The much older Kopficystis kirkfieldi Parsley, 199] from the Trentonian of Ontario, vaguely resembles Pseudovictoriacystis in 1ts 3-plated transverse row on distal 1/2 of convex surface. However, identification of the skeletal plates in that taxon is problematic (Ruta, in press). Unlike Pseudovictoriacystis, Kopficystis has 5 rather than 4 transverse rows of plates. Assuming the correctness of plate homologies discussed by Parsley (1991) and Ruta (in press), the most distal skeletal elements in this anomalocystitid correspond to C1, C6, C9, C10, C12 and C14 and to СМІ, o, j, gl, i and e respectively in the terminology of Parsley (1991). Although Pseudovictoriacystis displays a unique set of skeletal features, it is impossible to ascertain its affinities; we regard this form as deriving from a Victoriacystis-like ancestor. ACKNOWLEDGEMENTS We thank David Holloway (Museum of Victoria), Steve Eckardt and Frank and Enid Holmes for help in the field. M.R. is grateful to the Queensland Museum (Brisbane) for generous hospitality and for the use of facilities. A.R. Milner (Birkbeck College, University of Lon- don), R.P.S. Jefferies, A.C. Milner, S.J. Culver and L.R.M. Cocks (Natural History Museum, London) read a draft of the manuscript. P. Crabb (Natural History Museum, London) photo- graphed the specimens. We thank Dennis Kolata and Colin Sumrall for useful reviews but the views expressed herein are those of the authors alone. This work was carried out while M.R. was in receipt of a European Community grant (Training and Mobility of Researchers). LITERATURE CITED CASTER, K.E. 1952. Concerning Enoploura of the Upper Ordovician and its relation to other carpoid Echinodermata, Bulletins of American Paleont- ology 34: 1-47. CASTER, K.E. & GILL, E.D. 1967. Family Allanicytidiidae, new family. Рр. $561-S564. In Moore, R.C. (ed.) Treatise on invertebrate paleontology, Part S. Echinodermata 1(2). (Geol- ogical Society of America & University of Kansas : New York). DEHM, R. 1932. Cystoideen aus dem rheinischen Unterdevons. Neues Jahrbuch fiir Mineralogie, Geologie und Paléontologie, Beilage-Band, Abteilung A 9: 63-93. GILL, E.D. 1948. A new trilobite from the Yeringian (Lower Devonian) rocks of Kinglake, Victoria. Proceedings of the Royal Society of Victoria 59: 8-19, GILL, E.D. & CASTER, K.E. 1960. Carpoid echinoderms from the Silurian and Devonian of Australia. Bulletins of American Paleontology 41: 5-71. HOLLOWAY, D.J. & JELL, P.A. 1983. Silurian and Devonian edrioasteroids from Australia. Journal of Paleontology 57: 1001-1016. JAEKEL, O. 1918. Phylogenie und System der Pelmatozoen. Paláontologische Zeitschrift 3: 1-128. JEFFERIES, R.P.S. 1968. The Subphylum Calcichor- data (Jefferies 1967) — primitive fossil chordates with echinoderm affinities. Bulletin of the British Museum (Natural History), Geology Series 16: 243-339. 1990. The solute Dendrocystoides scoticus from the Upper Ordovician of Scotland and the ancestry of chordates and echinoderms. Palaeontology 33: 631-679. JEFFERIES, R.P.S. & LEWIS, D.N. 1978. The English Silurian fossil Placocystites forbesianus and the ancestry of the vertebrates. Philosophical Transactions of the Royal Society of London, Series B 282: 205-323. JELL, P.A. 1983. Early Devonian echinoderms from Victoria (Rhombifera, Blastoidea and Ophi- ocistioidea). Memoirs of the Association of Australasian Palaeontologists 1: 209-235. KOLATA, D.R. & JOLLIE, M. 1982. Anomalocystitid mitrates (Stylophora, Echinodermata) from the Champlainian (Middle Ordovician) Guttenberg Formation of the Upper Mississippi Valley Region. Journal of Paleontology 56: 531-565. KONINCK, M.L. de 1869. Sur quelques échinodermes remarquables des terrains paléozoiques. Bulletin de l'Académie Royale des Sciences Belgique 28: 544-552. PARSLEY, R.L. 1991. Review of selected North American mitrate stylophorans (Homalozoa: Echinodermata). Bulletins of American Paleontology 100: 5-57. RUTA, M. 1997. Redescription of the Australian mitrate Victoriacystis with comments on its functional morphology. Alcheringa 21: 81-101. 1998. An abnormal specimen of the Silurian anomalocystitid mitrate Placocystites forbes- ianus. Palaeontology 41: 173-182. In press. A cladistic analysis of the anomalocystitid mitrates. The Zoological Journal of the Linnean Society. RUTA, M. & BARTELS, C. 1998. A redescription of the anomalocystitid mitrate Rhenocystis MEMOIRS OF THE QUEENSLAND MUSEUM latipedunculata from the Lower Devonian of Germany. Palaeontology 41: 771-806. КОТА, M. & JELL, P. A. 1999а. Adoketocarpus gen. nov., a mitrate from the Ludlovian Kilmore Siltstone and Lochkovian Humevale Formation of central Victoria. Memoirs of the Queensland Museum 43: 377-398. 1999b. A note on Victoriacystis wilkinsi (Anomalocystitida: Mitrata) from the Upper Silurian of Victoria. Memoirs of the Queensland Museum 43: 423-430. RUTA, M. & THERON, J.N. 1997. Two Devonian mitrates from South Africa. Palaeontology 40: 201-243. STRUSZ, D.L. 1972. Correlation of the Lower Devonian rocks of Australasia. Journal of the Geological Society of Australia 18: 427-455. TALENT, J.A. 1967. Silurian sedimentary petrology and palaeontology. Bulletin of the Geological Survey of Victoria 59: 24-29, UBAGHS, G. 1967. Stylophora. Pp. 496-565. In Moore, R.C. (ed.) Treatise on invertebrate paleontology. Part S. Echinodermata 1(2). (Geological Society of America & University of Kansas: New York). 1969. Les échinodermes carpoides de l'Ordovicien inférieur de la Montagne Noire (France). Cahiers de Paléontologie. (Editions du Centre National de la Recherche Scientifique: Paris). VANDENBERG, A.H.M. 1988. Silurian-Middle Devonian. Pp. 103-146. In Douglas, J.G. & Ferguson, J.A. (eds) Geology of Victoria. (Victorian Division of the Geological Society of Australia: Melbourne). VANDENBERG, A.H.M, GARRATT, M.J. & SPENCER-JONES, D. 1976. Silurian-Middle Devonian. Special Publications of the Geological Society of Australia 5: 45-76. WILLIAMS, С.Е. 1964. The geology of the Kinglake district, central Victoria. Proceedings of the Royal Society of Victoria 77: 273-328. WOODS, LS. & JEFFERIES, R.P.S. 1992. A new stem-group chordate from the Lower Ordovician of South Wales, and the problem of locomotion in boot-shaped cornutes. Palaeontology 35: 1-25. A NOTE ON VICTORIACYSTIS WILKINSI (ANOMALOCYSTITIDA: MITRATA) FROM THE UPPER SILURIAN OF VICTORIA MARCELLO КОТА AND PETER A, JELL Ruta, M. & Jell, P. A. 1999 06 30: A note on l'jetoriacystis wilkinsi (Anomalocysttida: Mitrala) from the Upper Silurian of Victoria. Memoiry of lhe Queensland Museum 43(1): 423-430. Brisbane. ISSN 0079-8835. New material of l'fetoriac stis wilkinsi from the Upper Silurian of Victoria reveals new features of the convex surface and permits a more detailed comparison with the congeneric V. holmesorum from the Lower Devonian. A revised diagnosis is provided for P. wilkinsi, together with a description of the best preserved among the new specimens. © Anomalocystitida, Vietortucystis, Silurian, Victoria. Marcello Ruia, Department of Palavontology, The Natural History Museum, Cromwell Road, London SW7 SBD, United Kingdom, Peter А. Jell, Queensland Museum, Р.О. Box 3300, South Brisbane 4101, Australia; received 2 July 1998. Subsequent to the redeseription of the Ludlow anomalocystitid Fictoriacystis wilkinsi Gill & Caster, 1960 from the Dargile Formation, Heathcote and Melbourne Formation, Hawthorn (Ruta, 1997), we examined additional material in the Museum of Victoria, Melbourne, part of which extends. the known distribution of this species, New specimens are generally more complete and fully articulated than those figured by Gill & Caster (1960) and Ruta (1997) and clarify details of the external and internal anatomy of this mitrate. Major differences between K wilkinsi and K holmesorum Ruta & Jell, 1999 from the Lower Devonian Humevale Formation of central Victoria were highlighted by Ruta & Jell (1999h), SYSTEMATIC PALAEONTOLOGY Terminology and plate nomenclature are as used elsewhere in this volume (Ruta & Jeli, 19992). All material is housed in the Museum of Victoria Palaeontological Collections (NM VP) and localities entered in the fossil locality register ol the same Museum (NMYPL). All illustrations are of latex casts from internal and external moulds whitened with ammonium chloride sublimate unless otherwise stated, Class STYLOPHORA Gill & Caster, 1960 Order MITRATA Jaekel, 1918 Suborder ANOMALOCYSTITIDA Caster, 1952 Family PLACOCYSTITIDAE Caster, 1952 DIAGNOSIS. Lateral margins of PM convex for most of their length. C3 and C12 in contact with each other. Proximo-lateral angles of СЗ trun- cated (condition of second and third characters reversed in Victoriacystis) (Caster, 1952; Parsley, 1991; Ruta & Jell. 1999b: Ruta, in press). Victoriacystis Gill & Caster, 1960 TYPE SPECIES. Metoriaeystis wilkinsi Gill & Caster, 1960 from the Ludlow Dargile Formation, Victoria. DIAGNOSIS. бее Ruta & Jell (1999b) and Ruta (in press). OTHER SPECIES, Vicroriacystis halmesorun Ruta & Jell; 1999b from the Lochkovian of the Humevale Formation, central Victoria. Victoriacystis wilkinsi Gill & Caster, 1960 (Figs 1-5) MATERIAL. NMVP23086, 109203 trom F41-42 (type locality of Gill & Caster, 1960) (Dargile Fm: Ludlow). NMVP18313-18317 from City Brick Co. pit, Camberwell Rd, Hawthom (Gill & Caster, 1960) (Melbourne Fm: Ludlow). NMVP22160-22161, 22348. 24111. 100457-100458, 100461-100462, 100464-100468 from NMVPL299 (= F31 of Williams, 1964) road cutting S of Bald Hills, 3.2km Е of Kilmore (Dargile Fm: Ludlow). NMYP100446-100448 from NMVPL300 (= X64 of Williams, 1964) vicinity of disused mine on Comet Creek, 4.6km SE of Clonbinane (Humevale Fm: Ludlow). NMVP149332 from NMVPL1927 on Broardliurst Creek at the crossing of the Kilmore to Wandong Rd (sce Vandenberg, 1992) (Kilmore Siltstone: Ludlow). DIAGNOSIS (see also Ruta & Jell (1999b) and Ruta (1997. in press)). Lateral body walls slightly diverging ventrally. C-ILM sutures straight, convex or, rarely, sinuous. A-C suture straight or slightly geniculate, at <40° to body axis. PLM much wider distally than proximally, Medial 424 MEMOIRS OF THE QUEENSLAND MUSEUM VIG 1. Fictoriaevstis wilkinsi Gill & Caster from NMVPL300. A-C. convex surface, detail of proximal part of appendage and detail of distal part of inside of A, respectively, of NMVP100447., х3, x6 and х8, respectivelv: D, detail of right spine from Fig. 2C, NMVP100448. «8, Un NEW MATERIAL ОЕ V7CTORIACYSTIS WILKINSI 42 FIG. 2, Vietoriacvstis wilkinsi Gill & Caster. A-B, D, inside of convex surface, detail of left spine and convex surface, respectively. of NMVP100462, from NMVPL229. «5, «$ and *5. respectively; С. plano-coticave surface of NMVP100448 from NMVPL300, #4. 426 margins of DLM straight. Proximal margins of PM occupying 71/2 proximal body excavation. Rows II-IV perpendicular to longitudinal axis or only gently concave distally. C10, C12 and C14 only slightly larger than C16 and C18. Sutures between C15 and C16 and between C18 and C19 strongly diverging proximally. C16 and C18 longer and wider than C15 and C19. C3 much larger than C2 or C4. Few, widely spaced ridges on proximal 1/2-1/3 of C16 and C18 and near lateral margins of PM. Spout-shaped thickenings internally on C2-C4. Tetramerous rings with weak distal thickenings lacking knobs. Styloid with slightly recurved. poorly developed distal blade only slightly higher than proximal blade. Distal part of appendage not differentiated. Distal ossicles with straight to gently concave distal margins, poorly developed ossicular apices and markedly sloping apical margins. DESCRIPTION. EXTERNAL. We describe those features not reported by Gill & Caster (1960) or Ruta (1997). Body outline sometimes vase-shaped, with maximum width halfway along length of PLM. LOP subtrapezoidal to subpentagonal. MOP subrectangular. markedly asymmetrical, Distal margins of C2-C5 slightly distal to distal margins of LOP and MOP. Proximal margin of C17 sometimes accommod- ated by shallow notch on distal angle of C21 (Figs 1A, 2D, 3E). C21 otherwise shield-shaped (Fig. 1A), with geniculate lateral margins. Spines c. 1/3 as long as body. uniformly tapering distally, with blunt medial and lateral margins, with proximal 1/4 shaped like a truncated cone. with central shaft straight and slightly depressed, with distal 1/3 gently curved medially (Figs ID. 2A-D. 3D). Terrace-like ridges often irregular and branched near proximolateral angles of plano- concave and convex surfaces (Figs 3E, 4E), irregularly sinuous and widely spaced on proximal 1/2 of C16 and C18. INTERNAL. Septum on inside of plano-concave surface (Figs ЗЕ, 4A); distal 1/2 of septum on interior of C straight and almost parallel to longitudinal axis. widening slightly halfway along its proximal part immediately lateral to the medio-distal angle of left PM. Septum becoming much shallower and broader beyond this point. giving rise to vaguely L-shaped structure straddling the triple junction formed by C with the 2 PM plates. deepening again and gently convex laterally on the lateral 1/2 of left PM. Cross-section of septum more asymmetrical (steeper to right) at PM than at C. Most distal part MEMOIRS OF THE QUEENSLAND MUSEUM of C portion. corresponding to the diminutive spur of Ubaghs (1967), thickened and bent abruptly to the right at c. 30° to longitudinal axis. Most proximal part of septum on left PM not visible. Distal part of diminutive spur termin- ating abruptly. Rest of plano-concave surface almost featureless except for poorly pronounced ridge parallel to medio-distal margin of A, fainter medially than in its lateral 1/2, continuing on C along straight course, almost perpendicular to longitudinal axis and stopping abruptly before reaching lateral margin of C (Figs 1A,C, 4A). Inside of convex surface (Fig. 2A) with button- like projections on C12, C14, C18 and C19, spout-shaped thickenings on C2, C3 and C4 and tortuous ridges on C16 and C18. Button-like projections only slightly raised with respect to surrounding plate surface. gently merging into the latter and slightly longer than wide: greater axis of projections on C 14 and C19 at acute angle with longitudinal body axis. C12 projection slightly larger than remaining projections, c. 1/2 as large as C19 projection. Spout-shaped projections on C2, C3 and C4 (Figs 2A, 4B) of approximately equal size. occupying distal 1/3 of inside of plates, с. 1/3 as wide as these. delimiting a central and 2 lateral depressions, continuing distally into flat, sloping, rectangular area, and proximally into flat or gently convex trapezoidal area. Free margins of spout-shaped projections blunt, continuing laterally into narrow, transversely arched. vertical septa with sharp free margins. Ridges on C16 and C18 poorly preserved (Fig. 2A), apparently interrupted in places, with irregularly sinuous course. APPENDAGE. Tetramerous rings 7-8, telescopic, with poorly developed distal thicken- ings without tubercles and blunt parasagittal section, with ring plates on the same side as convex surface of body showing gently convex distal margins (Figs ТА-В. ЗА, D-E, 4A, D-E, 5A-E). Two distalmost ring plates on the same side as plano-concave surface of body slightly bent proximally, wrapped around proximal styloid process. Most distal ring plates lying on the same side as convex surface much smaller than more proximal plates, flanking the most proximal part of the styloid body immediately underneath free margin of proximal blade. Free margins of distal blade slightly concave proximally in lateral view, especially in abapical 1/2, Keel between proximal and distal blade sharp. with wedge-shaped lateral profile (Fig. 5A, C-D). Ossicles scarcely overlapping each other proximo-distally, with gently concave to NEW MATERIAL OF VICTORLIACYSTIS WILKINSI 427 51 V 5 ач FIG 3. Victoriacystis wilkinsi Gill & Caster, all from ММУ PL299. A. plano-concave surface and appendage of NMVP22161. 5. В. poorly preserved convex surface and appendage of NMVP22348. «6, С, plano-concave surface of NMVP24111, *8. D. plano-concave surface of NMVP18314, «5, J, convex surface partially disarticulated and revealing distal half of inside of plano-concave surface of N MVP18316. х2 428 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 4. Vietoriacystis wilkinsi Gill & Caster. A, inside of plano-concaveand part of exterior of convex surface of NMVP100446 from NMVPL300, х3. B. interior of distal part of convex surface of NMVP100462. х10. C. detail of distal part of appendage of NMVP 100457. «10, D, tetramerous rings of proximal appendage and mierior of distal appendage of NMVP23086, <8. B-D from NMVPL299. E. plano-concave surface of NMVP149352. from ММУРІ.1927, +5. NEW MATERIAL OF VICTORIACYSTIS WILKINSI 429 FIG. 5. Fictoriacvstis wilkinsi Gill & Caster. A-D from NMVPL299. Tetramerous rings, styloid, ossicles and paired plates. A, NMVP100457. «10. B-D. NMV P 100466, all «8. E, NMVP 100446 from NMVPL300, x6. 430 almost straight distal margins, poorly pro- nounced apices and blunt, markedly sloping apical margins. REMARKS. New specimens of E wilkinsi show that the range of variation in several anatomical features (e.g. body outline; shape of LOP and МОР; proportions of C21) is broader than previously reported (c.g. Ruta, 1997), None of the new specimens shows the knobbly sculpture reported by Ruta (1997) on plate MOP. It is, therefore, impossible to establish whether such a sculpture is a genuine feature of this anom- alocystitid. NMVP100462 (Figs 2A-B,D, 4B) provides the most complete information available on the inside of the convex surface. Internal characters of this surface are readily comparable with those of the congeneric K holmesorum (Ruta & Jell. | 999b), although the 2 species differ in the shape of the internal thickenings on C2-C4. Several skeletal features show consistent variations in the 2 species of Victoriacystis (Ruta & Jell, 1999b) and new specimens of | wilkinsi confirm this. Important differences are observed at the level of spines, and position, shape and proportions of several plates of the body and appendage. ACKNOWLEDGEMENTS We thank David Holloway and Andrew Sandford (Museum of Victoria, Melbourne) for loan of specimens; Andrew Milner (Birkbeck College, University of London) for comments on the manuscript; Phil Crabb (Natural History Museum, London) for the photographs. M.R. is grateful to the Museum of Victoria and to the Queensland Museum, Brisbane, for providing access to facilities. MEMOIRS OF THE QUEENSLAND MUSEUM LITERATURE CITED CASTER, К.Е. 1952. Concerning Enaploura of the Upper Ordovician and its relation to other carpoid Echinodermata. Bulletins of American Paleont- ology 34: 1-47. GILL, E.D. & CASTER. K.E. 1960. Carpoid echino- derms from the Silurian and Devonian of Australia, Bulletins of American Paleontology 41: 5-71. JAEKEL, O. 1918. Phylogenie und System der Pelmatozoen. Palüontologische Zeitschrift 3: 1-128. PARSLEY, R.L. 1991. Review of selected North American mitrate stylophorans (Homalozoa: Echinodermata). Bulletins of American Paleont- ology 100: 5-57. RUTA, M. 1997, Redeseription of the Australian mitrate Vietoriaeystis with comments on its functional morphology. Alcheringa 21: 81-101. In press. A cladistic analysis of the anomalocystitid mitrates, Zoological Journal of the Linnean Society. КОТА, M. & JELL, Р.А. 1999a. Adoketocarpus gen. nov, a mitrate from the Ludlovian Kilmore Siltstone and Lochkovian Humevule Formation of central Victoria. Memoirs of the Queensland Museum 43; 377-398, 1999b. Two new anomalocystitid mitrates from the Lower Devonian Humevale Formation of central Victoria. Memoirs of the Queensland Museum 43; 399-422. UBAGHS, G. 1967. Stylophora. Pp. 496-565. In Moore, R.C. (Ed.) Treatise on invertebrate paleontology. Part S. Echinodermata 1(2). (Geological Society of America & University of Kansas: New York). VANDENBERG, A.H.M. 1992, Kilmore 1:50,000 map and geological report. Geological Survey of Victoria Report 91: 1-86, = map. WILLIAMS, GE. 1964. The geology of the Kinglake district, central Victoria. Proceedings of the Royal Society of Victoria 77: 273-328. REVISION OF SILURIAN AND DEVONIAN ALLANICYTIDIIDAE (ANOMALOCYSTITIDA: MITRATA) FROM SOUTHEASTERN AUSTRALIA, TASMANIA AND NEW ZEALAND MARCELLO RUTA AND PETER A. JELL Ruta, M. & Jell, P. A. 1999 06 30: Revision of Silurian and Devonian Allanicytidiidae (Anomalocystitida: Mitrata) from southeastern Australia, Tasmania and New Zealand. Memoirs of the Queensland Museum 43(1): 431-451. Brisbane. ISSN 0079-8835. Additional specimens of the allanicvtidiid anomalocvstitids Notocarpos garratti (Upper Silurian, Victoria), Tasmanicytidium burretti (Lower Silurian, Tasmania) and Allanicytidium flemingi (Lower Devonian, New Zealand) yield new information, allowing revised diagnoses. №. garrattihasa system of ridges on the internal surface of C20-C22 anda row of orifice platelets along distal transverse thickening on inside of C1 and C5. Internal ridgeson C21 are homologouswith similar structures in A//anievtidium and Placocystella. T. burretti has orifice platelets, faint ridges internally on C21, tetramerous rings proximally in the appendage and sculpture on the distal styloid blade. New material of 4. flemingi shows external sculpture and stereom texture of convex surface, shape of C21 and proximal body excavation. O Allanicytidiidae, Silurian, Devonian, Victoria, Tasmania, New Zealand. Marcello Ruta, Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom; Peter A. Jell, Queensland Museum, P.O. Box 3300, South Brisbane 4101, Australia; received 20 August 1998. Despite extensive discussion on the interrelationships of allanicytidiid anomalo- cystitids (Caster & Gill, 1967; Philip. 1981: Caster, 1983; Haude, 1995; Ruta & Theron. 1997; Ruta & Jell, 1999a; Ruta. in press). numerous features of several species remain obscure or misinterpreted. All species except for the 2 most basal allanicytidiids, Protocytidium elliottae Ruta & Jell, 1999a from the Upper Ordovician of Victoria and Occultocystis koeneni Haude, 1995 from the Lower Devonian of Argentina, display very similar plate configurations. especially on the convex surface. making specific recognition difficult. Differ- ences in external sculpture, in body proportions and in shape of individual plates provide diagnostic features (Ruta & Theron, 1997: Ruta, in press). Newly available material of the allanicytidiids Notocarpos garratti Philip, 1981. 7asmanicy- tidium burretti Caster, 1983 and Allanicytidium flemingi Caster & Gill, 1967 clarifies poorly understood aspects of their anatomy, permitting revised diagnoses. SYSTEMATIC PALAEONTOLOGY Specimens are housed in the Museum of Victoria, Melbourne (prefix NMVP), and Department of Geological Sciences, University of Canterbury, Christchurch (UCM). Most localities are entered on the Museum of Victoria locality register (NMVPL). Terminology, orientation and plate nomenclature follow Ubaghs (1967, 1969) and Ruta (in press), with modifications as in Ruta & Jell (1999a-c). All illustrations are of latex casts from decalcified moulds and whitened with ammonium chloride. Class STYLOPHORA Gill & Caster, 1960 Order MITRATA Jackel, 1918 Suborder ANOMALOCY STITIDA Caster, 1952 Family ALLANICYTIDIIDAE Caster & Gill, 1967 DIAGNOSIS. See Ruta & Jell (1999a). Notocarpos Philip, 1981 TYPE SPECIES. Notocarpos garratti Philip, 1981 from the Ludlow Clonbinane Sandstone Member of the Humevale Formation, central Victoria, by original designation. DIAGNOSIS (modified from Philip, 1981 and based on the largest available, undeformed specimens). Plate A about as wide as long, with long axis at about 45? to body axis. not in contact withleft PLM. Plate B lacking. Plate C as wide as long. with subparallel lateral margins. LOP narrowly wedge-shaped. DLM with poorly dev- eloped truncatoconical projection. with process for spine insertion. C1 and C5 about 1.5 times as wide as long. C20 and C22 subrhomboidal, wider than long. with gently sinuous distal margins. PS 122 io MEMOIRS OF THE QUEENSLAND MUSEUM FIG 1. Notocarpos garratti Philip from NMVPL300. A, plano-concave surface of NMVP 100452. «5. B, distal half of plano- concave surface of NMVP 100453, х8. С.Р, plano-concaveand convex surfaces, respectively. of NMVP100441, «5 REVISION OF SILURIAN AND DEVONIAN ALLANICYTIDIIDAE 43 C21 twice as long as wide, occupying about 40% of width of convex surface, narrowly inserted between C20 and C22, with proximo-lateral margins smoothly concave, with proximal mar- gin about 1/10 plate width, with straight lateral margins parallel to body axis. Spines straight, robust. cigar-shaped, without cutting lateral edges. Transverse ridge ornament on at least some part of A, C. DLM, ILM and PLM. C20 and C22 with same ornament laterally. Riblets and irregularly confluent. short transverse ridges laterally on C11, C13, C15 and C19. C21 without sculpture. Proximal part of appendage of 4 tetramerous rings overlapping each other to a small extent. Proximal styloid blade 1/3 width of distal blade, approximately semicircular, smooth. Distal blade massive. fan-shaped. with straight radiating ridges. Distal part of appendage narrow, truncated abruptly (probably incomplete in all available specimens). presumably not longer than body. Notocarpos garratti Philip. 1981 (Figs 1-11. 16A) Notocarpos garratti Philip, 1981: 36, figs 3-6: Caster, 1983: fig. 2C. MATERIAL. NMVP100459-100460, 22350-22351 from NMVPL299 (= F31 of Williams (1964)) in road cutting south of Bald Hills, c. 3.2km E of Kilmore, Victoria (Ludlow, Dargile Formation), NMVP100440-100445, 100449-100456, 21939-21942, 22349, 22353-22354 from NMVPL300 (= X64 of Williams (1964)) near disused mine on Comet Creek, c. 4.6km SE Clonbinane, Victoria (Ludlow, Clonbinane Sandstone Member, Humevale Formation); NMVP65008-65010, 65022-65028, 65030-65031. 65035-65038. 65040-65042, 65044, 65052-65053 from the type locality. in a cutting on Dry Creek Road, Clonbinane; mid Ludlow, Clonbinane Sandstone Member, Humevale Formation. DIAGNOSIS. As for genus. DESCRIPTION. EXTERNAL. (including only new or additional data not in Philip, 1981). MOP broadly trapezoidal. 1.5-2 times as wide as long, with straight distal margin. LOP wedge-shaped, with major axis at c.45° to body axis, narrowly inserted between MOP and DLM, with straight or convex lateral margins, with straight or concave medial margins. C21 much longer than wide. with concave proximo- lateral margins. C11 and C13 shorter than C15 and C19. C15 and C19 same size or larger than C20 and C22. Terrace- like ridges rarely extending across width of marginal plates of plano-concave surface, frequently replaced laterally by short ridges or riblets. Similar pattern on convex suface, except Uu for more numerous riblets and confluent short ridges on lateral 2/3 of C11. C13, C15 and C19. External stereom texture usually compact to coarsely granular, rarely microporous or with vermicular surface pattern. INTERNAL. C21 and C20 with system of ridges proximally (Figs 9A. 10C, 11). Ridges variable in width, meandering on C21 to form almost bilaterally symmetrical pattern, proximally consisting of straight left and irregularly sinuous right sections normal to proximo-lateral margins of plate running medially to points directly distal to left and right ends of proximal plate margin, distally in sinuous bilaterally symmetrical pattern with 2 lateral and 2 medial lobes. Distally the ridges recurve proximally in thin parallel left and right ridges: proximal ends separated by wide, medial. proximally tapering septum: septum slightly higher than ridges, with almost vertical lateral walls. Ridge on C20 with first lobe after crossing from C21 V-shaped and apex pointing at co- operculum of C20; second lobe large. club- shaped. with medial arm crossing back to C21. curving latero-distally to surround right proximo-lateral angle of C21, continuing as thinner. straight ridge back on C20; third lobe subtrapezoidal. pointing towards co-operculum, with straight diverging arms; fourth and fifth lobes very weak ridges. finger-like: fifth lobe lateral to co-operculum. Co-operculum of C20 fan-shaped. raised. with slightly thickened margin, with proximal part continuing to proximo-lateral corner of plate as subtriangular. raised structure with almost straight medial margin and broadly concave lateral margin, distally with thick subquadrate body on margin centro-distally: faint, distally concave ridge projecting laterally from co-operculum, almost parallel to proximal arm of fifth lobe. widening and fading laterally. Orifice platelets overlying transverse thicken- ing on distal interior of Cl and C5. 6-8 in transverse row, rectangular, Imm wide, 0.75mm long (Figs 5C. 8A). APPENDAGE. Proximal part of appendage about 1/5 body length. «1/3 body width. Tetramerous rings 4, overlapping each other narrowly, with thickening along distal margins. gently arcuate transversely. Proximal styloid blade smooth, as long as wide. 1/3 distal blade width, slightly broader than and hardly separated from central part of styloid, with roughly semi- circular thick free margin. with convex distal 434 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 2. Notacarpos garratti Philip from NMVPI 300. A.D, detail of proximal appendage and plano-concave surface, respectively of NMVP100453. «10 and х5. respectively. B.E, detail of proximal appendage and plano-concave surface of. NMVP100443_ x8 and х5. respectively, C. detail of proximal appendage of NMVP100441. «10. F, convex surface of NMVP 100440. +5. REVISION OF SILURIAN AND DEVONIAN ALLANICYTIDIIDAE 43 surface, sometimes exceptionally well- developed and flared (Fig. 10B,D). Distal styloid blade transversely expanded, fan-shaped, slightly recumbent proximally, with sharp free margin, with weak straight radial ridges on proximal and distal surfaces (Fig. 9A,C-D); ridges of proximal surface slightly more robust and wider medially than laterally; ridges of distal surface slightly wider and flatter than on proximal surface; lateral ear-like projections with flat to gently depressed proximal surfaces and semicircular to parabolic margins. Appendage incomplete in most specimens. Ossicles and plates similar to those of Placocystella africana in lateral view (Ruta & Theron, 1997); ossicles with slightly more sinuous proximal and distal margins, with apical margin at 45° to horizontal plane, with apex projecting distally, with knobbly sculpture on lateral surfaces. External stereom texture of appendage similar to that of body, sometimes irregular due to coarse, irregular ridges, pustules and shallow pits, especially on tetramerous rings and styloid. MORPHOGENY. In smallest specimens: length/ width ratio of body slightly greater; lateral body margins more convex in proximal 1/3; plate A at least twice as long as wide; MOP as long as wide; proximal and distal margins of DLM and ILM more strongly converging medially; DLM wider than long; angle between lateral and medial margins of Cl and C5 >120°; lateral and medial margins of C1 and C5 sometimes merging into each other along smooth curve; lateral and distal margins of C20 and C22 merging gradually into one another along gently convex curve in plan view, without forming latero-distal angle; C11 and C13 as long as or longer than C15 and C19; C15 and C19 smaller than C20 and C22; proximolateral margins of C21 gently sinuous, diverging at greater angle from longitudinal axis; C21 width/body width slightly higher; C21 length/C1 (C5) length ratio slightly lower; spines more robust, shorter than distal body margin, club-shaped; proximal part of appendage «1/5 as long as body, at least 1/2 as wide distally as proximally; distal styloid blade 71/2 body width; ridges on both surfaces of the body frequently interrupted and replaced by riblets or transversely aligned short ridges; most distal ridges often pustule-like, especially on convex surface. REMARKS. Although preservation is sometimes poor due to relatively coarse matrix Un and despite little deformation in some specimens, new material of Notocarpos supplements information on external ornament of body plates and styloid. Previously unrecorded features include the rectangular orifice platelets on the inside of C1 and C5 and the system of sinuous ridges on interior proximal 1/2 of convex surface. The latter feature has been documented only in Allanicytidium and Placocystella among allanicytidiids (Caster & Gill, 1967; Ruta & Theron, 1997). Orifice platelets are known in some Northern Hemisphere anomalocystitids (Jefferies & Lewis, 1978; Kolata & Guensburg, 1979; Kolata & Jollie, 1982; Parsley, 1991). Tasmanicytidium Caster, 1983 TYPE SPECIES. Tüsmanicytidium burretti Caster, 1983 from the Lower Silurian Richea Siltstone, Tasmania; by original designation. DIAGNOSIS. (modified from Caster, 1983). Plate A as long as and c. 1/3 as wide as C; A-C suture straight. LOP-DLM suture strongly concave distally. DLM with latero-distal angles subcylindrical for spine insertion. C20 and C22 with weak transverse keels occupying more than 1/2 of plate width. C21 shield-like, with most proximal part narrowly inserted between C20 and C22, with gently concave proximal margin very narrow. Spines round in cross-section, straight, needle- shaped. Scale-like riblets on lateral 1/2 of PLM and on C11, C13, C15, C19, C20, and C22. C20 and C22 with few, short, transverse terrace-like ridges proximo-laterally. Tasmanicytidium burretti Caster, 1983 (Figs 12, 16B) Tasmanicytidium burretti Caster, 1983: 334, figs 2-4. MATERIAL. NMVP148541 from NMVPL296 near Terry Walshe Road, Tiger Range, on 1:100,000 Wedge Sheet 8112 DN512850, Tasmania; Llandovery, Richea Siltstone (Baillie, 1979). DIAGNOSIS. As for genus. DESCRIPTION. EXTERNAL. Largest riblets on convex surface scale-like, closely spaced, with straight to irregular distal margin, confined to C20-C22, about 0.2mm wide; smallest riblets subcircular, sparse, conferring pustulose aspect to distal part of convex surface, about 0.1mm wide (Fig. 12B). Transverse ridges on C20 and C22 faint. Riblets on CI5 and C19 never confluent, irregularly spaced, scale-like or sub- rectangular proximally, subcircular to pustulose distally. Riblets on C11 and C13 sparse, small, 436 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 3. Norocarpos garratti Philip trom NMVPL300. A.C. convex and plano-concave surfaces of NMVP100443, x3. B. proximal convex surface and appendage of NMVP100440, «10. D. convex surface of NMVP100442, «4, REVISION OF SILURIAN AND DEVONIAN ALLANICYTIDIIDAE 437 FIG. 4. Notocarpos garratti Philip from NM VPL300. A. detail of distal part of'convex surface of NMVP100442, x8. B-C, plano-concave and convex surfaces, respectively of NMVP21939, x5. pustulose, Riblets on C21 varying in shape and C21 with narrow, marginal zone of short, size proximo-distally, uniformly distributed. transverse striations, Stereom fabric apparently 438 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 5. Netocarpos garraiti Philip trom NMVPL300. A, proximal part of convex surface and appendage of NMYVP 100451, х8, B, convex surface of NMVP 100450, х5. C. distal part of plano-concave surface [rom interior, with spines and orifice platelets on NMVP 100442, ©8. REVISION OF SILURIAN AND DEVONIAN ALLANICYTIDIIDAE 439 FIG. 6. Notocarpos атташ Philip. A, convex surface of NMVP100459. В, partial convex surface of NMVP100460.C, partial (distal) convex surface of NM VP65040. D, convex surface of NMVP65027.E, partial convex surface of NMVP65037. F, plano-concave surface of NMVP65031. A-B from NMVPL299, х5. C-F from G23, х3. LAND MUSEUM 5 MEMOIRS OF THE QUEEN 440 REVISION OF SILURIAN AND DEVONIAN ALLANICYTIDIIDAE microporous to compact. Stereom centrally on C1 and C5 of irregular, subcircular shallow pits separated by weak trabeculae, proximally compact or minutely porous, distally vermicular, with irregularly sinuous, thick bifurcating trabeculae separated by shallow, narrow furrows and often flanked by short thickenings of variable shape and size. INTERNAL. C5 and C13 with laterally displaced, subhemispherical thickenings. C21 with subcentral straight ridge on proximal half surrounded by poorly defined, sinuous ridges as in Allanicytidium and Notocarpos. Distal margin of Cl and C5 with transverse row of 5-6 central subquadrate orifice plates, 0.3-0.5mm wide, bordered proximally by smaller, rectangular platelets. Orifice plates and inside of convex surface with compact to microporous stereom. APPENDAGE, Tetramerous ring plates of convex side arcuate in cross-section, with thin ridge running close to the thickened distal margins. Styloid with large distal blade; blade with concentric, externally convex, closely spaced ridges on proximal (Fig. 12A) and distal surfaces (Fig. 12B), arranged in sectors delimited by radial furrows running on distal surface from insertion of blade on styloid to blade margin, and corresponding to radial ridges with zig-zag course on proximal surface; concentric ridges alternating on both sides of zig-zag ridges and radial furrows, with degree of curvature increasing in more apical sectors (i.e. sectors which are closer to midpoint of blade margin). REMARKS. The new specimen adds inform- ation on orifice plates, tetramerous rings of appendage, shape and ornament of styloid blade, external body sculpture and inside of convex surface. The specimen 15 slightly deformed, but the convex surface is fully articulated. Compar- ison with the holotype permits an approximate estimate of body outline and general proportions. The elongate-ovoid or vasiform body (Caster, 1983) is almost certainly a genuine feature, although it may be slightly exaggerated in Caster’s reconstruction. 44] Allanicytidium Caster & Gill, 1967 TYPE SPECIES. Allanicytidium flemingi Caster & Gill, 1967 from the Lower Devonian Reefton Group, New Zealand; by original designation. DIAGNOSIS. (modified from Caster & Gill, 1967). Body subquadrate to ovoid, with broad margined re-entrant for insertion of appendage. A and C longer than wide, almost mirror images of each other. DLM with latero-distal angles projected into subconical processes for spine insertion. СІ and C5 thickened internally along distal margin, with sinuous proximo-lateral margins. Cll and C13 approximately equal in size to Cl and C5, with broadly concave medial margins and irregular proximal margins. C15 and C19 2/3 size of C11 and C13, each with sinuous proximal margin. C20 and C22 larger than C15 and C19, as wide as long. C21 with concave proximo-lateral margins interrupted by marked expansion in proximal half, with straight subparallel lateral margins. Closely spaced, polygonal to scale-like riblets on lateral marginal plates of plano-concave surface, near lateral margins of A and C and along most of the lateral margins of PLM. Proximal styloid blade semicircular, with thick, radial ridges on its distal surface. Distal styloid blade much wider than tall. Ossicles with lateral vertical grooves and poorly developed, nodose apices. Allanicytidium flemingi Caster & Gill, 1967 (Figs 13-15, 16C) Allanicytidium flemingi Caster & Gill. 1967: S564. figs 360. 361; Caster, 1983: figs 1. 2А. MATERIAL. NMVP27474 (plaster replica of holotype NZGS38/370203) and UCM440 from Rainy Creek near Reefton, Westland, South Island, New Zealand in the Emsian, Reefton Group. DIAGNOSIS. As for genus. DESCRIPTION. EXTERNAL (Figs 13, 14A). Riblet size (0.1-0.5mm wide), shape and distribution variable on different plates and on the same plate of convex surface. Riblets closer together on lateral 1/2 of marginal plates. Largest riblets on lateral 1/2 of C15, C19, C20 and C22, crescent-shaped, rectangular or irregularly polygonal, in irregular transverse rows, often resulting in vermicular surface, as opposed to FIG. 7. Notocarpos garratti Philip. A, convex surface of NMVP65028. B, convex surface of NMVP22354. С, plano-concave surface of NMVP21941. D, proximal appendage of NMVP65053. E, convex surface of NMVP65035. Е, convex surface of NMVP65037. A, D-F from G23, all x3. B-C, from NMVPL300, x8 and х6, respectively. 442 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 8. Nofocarpos garratti Philip, А. exterior view of plano-concave surface of NMVP100442 from NMVPLS300. х4. В, convex surface of NMVP65042. C, partial convex surface of NMVP63025. D, partial plano-concave surface of NMVP65044. B-D. from (23. *3. tuberculate or scaly elsewhere. Smallest riblets poorly defined margins, sometimes reduced to distally and proximo-medially on C20 and C22 small pustules without distal slope. Riblets on and on proximal 1/2 of C21, generally scale-like C21 less pronounced than (hose on lateral 1/2 of to broadly rounded. slightly wider than long. with marginal plates. with more regular, convex distal REVISION OF SILURIAN AND DEVONIAN ALLANICYTIDIIDAE 443 FIG. 9. Natocarpos garratti Philip, NM VP100444 from NM VPL300. A. inside of'convex surface with system of ridges on C20 and C21 and disrupted co-operculum on C20, х4. B, detail of C20 from D, x8. С, detail of styloid from A, х8. D, partial convex surface, х4, 444 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 10. Notocarposgarratti Philip. A-B, D, inside of plano-concavesurface ( B.) with detail of laterodistal comer (A)showing articulation of'spine and proximal appendage(D) of NMVP 100463 from NMVPL299. «10. х5 and * 10, respectively, C, inside of C20 and C21 of NMVP100444 from NMVPL300, «8 REVISION OF SILURIAN AND DEVONIAN ALLANICYTIDIIDAE margins, 1.5 times wider than long. Riblets on marginal plates 1.5-3 times as wide as long, with sharp, straight to crescentic, rarely polygonal distal margins. INTERNAL (Figs 14В-С, 15A). Internal surface of lateral marginal plates finely granular, of small irregular pores and furrows. Deeper portions of stereom sponge- like to coarsely perforate. especially on C21. Peripheral bands, corresponding to external bands, of fibrillar stereom composed of parallel striations perpendicular to plate margins and intercalated with transversely elongate to subcircular pores. Bands divided into lateral part and medial part by thin ridge reaching maximum thickness along distal margins of marginal plates апа latero- distal margins of C21; medial part 2-3 times as wide as lateral part. Portion of band running along distal angle of C21 1.5 times wider than elsewhere, with coarse stereom, without radial pattern of trabeculae in proximal 1/2, with fine, closely spaced, short trabeculae forming a fringe in distal 1/2 (Fig. 14B-C). Subelliptical. proximo-distally elongate, subcentral thickenings on lateral marginal plates of convex surface, with slightly raised margins. Thickening stereom compact peripherally, porose to sponge-like centrally. Stereom (Figs 13-14). External skeleton of microporous stereom without surface pattern. Surface of largest riblets coarsely granular to compact. with minute pores in smallest ones. Stereom near plate margins of larger pores of more irregular shape than those on central part of plates. Plate margins bordered by band 0.2-1mm wide, generally without riblets and with stereom of fine, closely spaced, straight parallel trabeculae and subcircular irregularly distributed pores. Changes of stereom structure through plate thickness visible if skeletal surface is eroded or broken. Stereom of deeper parts of skeleton generally coarser than surface stereom, labyrinthine, of short irregular thin branching trabeculae delimiting irregular pores. Deeper 445 FIG. 11. Camera lucida drawing of system of ridges on internal surface of C20 and C21 in Notocarpos garratti Philip, based on NMVP100444. stercom at level of peripheral bands composed of sinuous, branching trabeculae delimited by irregular furrows. replaced by granular fabric immediately adjacent to margins. Proximal body excavation. Proximal margins of PM crescent-shaped, with medial 1/2 slightly raised above subhorizontal projections (Fig. 15A). Apophyses subvertical, c. 1/2 as wide as each PM, subtriangular. with 1/3 of external surface closer to proximal margins of PM almost flat. remainder gently convex, with lateral ends turning abruptly towards convex surface and continuing into bases of apophyseal horns. Left horn missing. Right horn of uniform width. slightly distal to apophysis, subhorizontal, flat in cross-section, with free margins gently convex externally and only slightly diverging latero-medially (Fig. 15B). Medial end of right horn truncated abruptly without signs of breakage, therefore presumably complete. straight, oblique to longitudinal axis, reaching point at c. 2/3 of apophysis width in latero- medial direction. Space between internal margin of right horn and apophysis transversely 446 MEMOIRS OF THE QUEENSLAND MUSEUM VIG. 12. Tasmanicvtidium burretti Caster, incomplete plano-concavesurface, external (B) and partially disrupted internal side(A )of'convex surface, broken left spine, body sculpture, tetramerousrings and styloid with external ornament on distal blade. NMVP148541 from NMVPL296, х6. elongate, tapering slightly latero-medially and bent gently towards plano-concave surface. REMARKS. Caster & Gill (1967) gave a succinct but comprehensive description of Allanicytidium based on an almost complete external mould of the plano-concave surface and internal mould of the convex surface of a single individual (see also Caster. 1983). Replicas of type material in the Museum of Victoria include the partial external mould of the convex surface of the holotype, illustrated here for the first time together with a second specimen. Both individuals provide data on external sculpture and changes in stereom fabric of the interior of the convex surface, the shape of C21 (especially its proximal 1/4) and the morphology of the proximal body excavation. Knowledge of other skeletal features (e.g. MOP: internal side of plano-concave surface; ornament of distal styloid blade and distal ossicles) is still not available. SUMMARY AND CONCLUSIONS Configuration of the convex surface of the 3 allanicytidiids discussed herein (Fig. 16A-C) are compared to that of P/acocvstella africana (Fig. 16D). Although similar to 4//anicytidium flemingi, the Lower Devonian Australocystis langei Caster, 1956 from Brazil is omitted from the following discussion because of the incomplete preservation of its convex surface REVISION OF SILURIAN AND DEVONIAN ALLANICYTIDIIDAE 447 VIG. 13. Allanicytidiumflemingi Caster & Gill, from Rainy Creek near Reefton, NZ. A, partial convex surface and body sculpture of UCM440, x5. B, partial convex surface of NMVP27474 (replica of holotype NZGS 38/370203), x5. (Ruta & Theron, 1997; Ruta, in press). The prox- imal neck-like region in C21 in A//ani- cytidium and Placocystella suggests that these taxa are closer to each other than either is to Tasmani- cytidium or Notocarpos (Ruta & Theron, 1997: Ruta, in press). In Tasmanicytidium, the neck is not developed, but a short subtrapezoidal region between the proximo-medial angles of C20 and C22 indicates where the neck developed in more derived allanicytidiids. In passing from Tasmanicytidium through Placocystella to Allanicytidium, the C20/C21 and C21/C22 sutures became progressively more tortuous, resulting in development of lateral angular projections, poorly developed and close to the proximal margin of C21 in Placocystella, prominent and displaced halfway along the length of the neck in A//anicytidium. External sculpture in Notocarpos has transitional features between those of certain anomalocystitids from the Northern Hemisphere (e.g. Placocystites forbesianus de Koninck. 448 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 14. Alanieytidium flemingi Caster & Gill. UCM440. from Rainy Creek near Reefton. NZ A. detail of sculpture and stereom fabric of convex surface, x10. B,C, inside of distal part of C21, show ing peripheral band, * 10 and х8. respectively. 1869; Jefferies & Lewis, 1978) and thoseof more — Placocystites, Ruta & Theron (1997) and Ruta (in derived allanicytidiids. In the light of Jefferies’ press) invoked paedomorphosis to explain the (1984) reconstruction of ontogenetic changes in sculpture pattern of the allanicytidiids. REVISION OF SILURIAN AND DEVONIAN ALLANICYTIDIIDAE 449 FIG. 15. Allanievtidium flemingi Caster & Gill, UCM440, from Rainy Creek near Reefton. NZ. A. inside of convex surface showing stereom fabric and peripheral bands, х5. B, proximal body excavation with apophy seal horns. «10. "od Paedomorphic changes in allanicytidiid evolution are suggested by the smallest specimens of Notocarpos with sculpture of short ridges or riblets. The strongly diverging proximo-lateral margins of C21 in young Nofocarpos are reminiscent of those of more derived allanicytidiids (e.g. Tasmanteytidium and Placo- cystella). It is possible that paedomorphie changes occurred repeatedly in the allanicytidiid clade and affected various structures (ornament; plates) differently. Few changes affected the general plating pattern of the body (Ruta & Jell, 19993), These involve relative proportions of MOP, LOP, A and C plates, on the plano-concave surface, and outline of C21, on the convex surface. A detailed analysis of character changes is provided by Ruta (1n press), ACKNOWLEDGEMENTS David Holloway, Museum of Victoria, Melbourne and David Mackinnon, University of Canterbury, Christchurch lent material in their care. Steve Eckard! provided access to his collection, Andrew Milner, Birkbeck College, University of London offered suggestions to improve the text. We thank referees Chris Paul and Reimund Haude for useful suggestions but assume responsibility for the above ourselves. M. R. thanks the Museum of Victoria, Melbourne and the Queensland Museum, Brisbane for hospitality and for providing access to facilities. LITERATURE CITED BAILLIE, PW. 1979. Stratigraphic relationships of Late Ordovician to Early Devonian rocks in the Huntley Quadrangle, south-western Tasmania. Papers and Proceedings of lhe Royal Society of Tasmania 113: 5-13. CASTER, K.E. 1952. Concerning Enoploura of the Upper Ordovician and its relation to other carpoid Echinodermata. Bulletins of American Paleontology 34: 1-47. MEMOIRS OF THE QUEENSLAND MUSEUM D FIG. 16. Reconstruction of plating pattern of convex surface in Notocarpos garratti Philip (A), Tasmanicytidium burretti Caster (B), Allanicytidium flemingi Caster & Gill (C) and Placocystella africana (Reed) (D). Drawings not to scale. 1956. A Devonian placocystoid echinoderm from Paraná, Brazil. Paleontologia do Paraná (Centennial Volume): 137-148. 1983. A new Silurian carpoid echinoderm from Tasmania and a revision of the Allanicytidiidae. Alcheringa 7: 321-335 CASTER, K.E. & GILL, E.D. 1967. Family Allanicytidiidae, new family. Pp. 561-564. In Moore, К.С. (ed.) Treatise on invertebrate paleontology. Part S. Echinodermata 1(2). (Geological Society of America & University of Kansas Press: New York). GILL. E.D. & CASTER, K.E. 1960. Carpoid echinoderms from the Silurian and Devonian of Australia. Bulletins of American Paleontology 41: 5-71. HAUDE, R. 1995, Echinodermen aus dem Unter- Devon der argentinischen Prakordillere. Neues Jahrbuch fiir Geologie und Paläontologie, Abhandlungen 197: 37-86. REVISION OF SILURIAN AND DEVONIAN ALLANICYTIDIIDAE JAEKEL, O. 1918. Phylogenie und System der Pelmatozoen. Paláontologische Zeitschrift 3: 1-128. JEFFERIES, R.P.S. 1984. Locomotion, shape, ornament and external ontogeny in some mitrate calcichordates. Journal of Vertebrate Paleont- ology 4: 292-319. JEFFERIES, R.P.S. & LEWIS, D.N. 1978. The English Silurian fossil Placocystites forbesianus and the ancestry of the vertebrates. Philosophical Transactions of the Royal Society of London, Series B 282: 205-323. KOLATA, D.R. & GUENSBURQG, T.E. 1979. Diamphidiocystis, a new mitrate carpoid from the Cincinnatian (Upper Ordovician) Maquoketa Group in southern Illinois. Journal of Paleontology 53: 1121-1135. KOLATA, D.R. & JOLLIE, M. 1982. Anomalocystitid mitrates (Stylophora, Echinodermata) from the Champlainian (Middle Ordovician) Guttenberg Formation ofthe Upper Mississippi Valley region. Journal of Paleontology 56: 531-565. KONINCK, M.L. De 1869. Sur quelques échinodermes remarquables des terrains paléozoiques. Bulletin de |’ Academie Royale des Sciences Belgique 28: 544-552. PARSLEY, R.L. 1991. Review of selected North American mitrate stylophorans (Homalozoa: Echinodermata). Bulletins of American Paleontology 100: 5-57. 451 PHILIP, GM. 1981. Notocarpos garratti gen. et sp. nov., a new Silurian mitrate carpoid from Victoria. Alcheringa 5: 29-38. RUTA, M. (in press). A cladistic analysis of the anomalocystitid mitrates. The Zoological Journal of the Linnean Society. RUTA, M. & JELL, P.A. 1999a. Protocytidium gen. nov, a new anomalocystitid mitrate from the Victorian latest Ordovician and evolution of the Allanicytidiidae. Memoirs of the Queensland Museum 43: 353-376. 1999b. Adoketocarpus gen. nov., a mitrate from the Ludlovian Kilmore Siltstone and Lochkovian Humevale Formation of central Victoria. Memoirs of the Queensland Museum 43: 377-398. 1999c. Two new anomalocystitid mitrates from the Lower Devonian Humevale Formation of central Victoria. Memoirs of the Queensland Museum 43: 399-422. RUTA, M. & THERON, J.N. 1997. Two Devonian mitrates from South Africa. Palaeontology 40: 201-243. UBAGHS, G. 1967. Stylophora. Pp. 496-565. In Moore, R.C. (ed.) Treatise on invertebrate paleontology. Part S. Echinodermata 1(2). (Geological Society of America & University of Kansas: New York ). 1969. Les échinodermes carpoides de l'Ordovicien inférieur de la Montagne Noire (France). Cahiers de Paléontologie (Editions du Centre National de la Recherche Scientifique: Paris) 1-112. WILLIAMS, G.E. 1964. The geology of the Kinglake district, central Victoria. Proceedings ofthe Royal Society of Victoria 77: 273-328. + un Nm MEMOIRS OF THE QUEENSLAND MUSEUM DITHYROCARIS PRAECOX IS A CARPOID. Memoirs of the Queensland Museum 43/1): 452. 1999:- Chapman (1904) erected Dithvrocaniy praecox. for a specimen 15mm long from fine siltstone of the Upper Silurian Melbourne Formation in Merri Creek, al Craigieburn just E of the Hume Highway. about 25km N of Melbourne (Sections 2 & 3, Parish of Kalkallo, Geological Survey of Victoria Sheet Bb3). Dithvrocaris, based on its type species, (D. festudinens (Scouler, 1835) from the Lower Carboniferous of Scotland. belongs to the primitive crustacean group Phyllocarida (Rolfe. 1969: 321, 118.147). Chapman s species does nol belong to the Phyllocarida because the surface exposed ts multiplated and thus the animal could not have been enclosed by a single piece carapace as in the Phyllocarida, Rather. praecox 1% a mitrate сагро of the Family Allanicytidiidae. The plating arrangement (Fig. 1B) shows that the specimen represents the plano-concave surface of Notocarpos and because the anomalocystid plate C is on the right instead of the left (as normal in external view) it is an internal mould. Identifying Nofocarpos are the wedge-shaped lateral orifice plates (not recognised by Philip (1981) but clear [rom his figs 4B. 5D. 6D). shape of the proximal lateral plates. proximal median plates and large central plate. While this specimen has no major features separating it from Notecarpes garratti Philip. 1981 and seems likely to belong to that species, seniority of its name would destabilise a widely used species name and provide a wholly unsuitable holotype as basis for the type species. I advocate isolation of the name praecox оп the type specimen separating it from garratt by its smaller plate C and subquadrate body outline, thus preserving garratti as the type of Notecarpos and the widely used concept of the genus based on Philip's material and that presented by Ruta & Jell (1999). Literature cited CHAPMAN. F 1904. New or little known Victorian fossils in the National Museum, Melbourne, Part 4, -Some Silurjan Ostracoda and Phyllocarida. Proceedings of the Royal Society of Victoria 17: 298-319 PHILIP, G.M 198). Netocarpox gerratit gen. et sp nov, a new Silurian mitrate carpoid from Victoria. Alcheringa 5. 29-38 КОТ, W D.I. 1969, Phyllocarida. Рр R296-R331. In Moore. RC (ed. ) Treatise on invertebrate paleontology. Part R Arthropoda + vol 1 (Geological Society of America & University of Kansas: Boulder. Colorado & Lawrence, Kansas) RUTA, M. & JELL, РА. 1999 Revision of Silurian and Devonian Allanieviidiidae (Anemalocystitida’ Mitrata) from southeastern Australia, Tasmania and New Zealand. Memoirs at the Queensland Museum 43, 431-451 PA, Jell, Queensland Museum, P0, Box 3300, South Brisbane 4101. Anstralia; 20 May 1999. FIG, |, Notacarpos praecox (Chapman. 1904) A latex cust of the plano-concave surface of the internal mould of Museum af Victoria specimen. NMVP4662. 4. В. plating arrangement of plano-coneave surface drawn from A CONTENTS JELL, P.A. Silurian and Devonian crinoids from central Victoria ‚...............‚..... oe Bele tips 1 JELL, Р.А. & THERON, J.N. Early Devonian echinoderms from South Africa ...................-......+..у..%8.. 33 115 JELL, Р.А. & JELL, J.S. Crinoids, a blastoid and a cyclocystoid from the Upper Devonian reef complex of the Canning Basin, Western Australia... 2.02 ce cee ee ee Daraar tyres SAFE rohea] 201 WEBSTER, G.D. & JELL, P.A. New Carboniferous crinoids from eastern АишвїгаНа...............................+... 237 WEBSTER, G.D. & JELL, P.A. New Permian crinoids from Айшзїгайа................................. diac ped actin hac 279 SMITH, A.B, & IELL, P.A. A new cornute carpoid from the Upper Cambrian (Idamean) of Queensland. ............... 341 КОТА, M. & JELL, Р.А. Protocytidium gen. nov., a new anomalocystitid mitrate from the Victorian Latest Ordovician and evolution of the Allanicytidiidae ........ aL. ake See ass mas dena 02234 RUTA, M. & JELL, P.A. Adoketocarpus gen. nov., a mitrate from the Ludlovian Kilmore Siltstane and Lochkovian Humevale Formation of central Уісіопа. ............ ны аа а feats ыр е СЭД КОТА, M. & JELL, Р.А. Two new anomalocystitid mitrates from the Lower Devonian Humevale Formation of central Victoris отла E pad ue V eS а Р аА nettle а. og lcs 2399 RUTA, M. & JELL, P.A. A note on Victoriacystis wilkinsi (Anomalocystitida: Mitrata) from the Upper Silurian Of Wittotia e. cesi оту hed Peta АРТ А A ePi EI EpPrbeIPIIAWITOTIpept 423 RUTA, M. & JELL, P. A. Revision of Silurian and Devonian Allanicytidiidae (Anomalocystitida: Mitrata) from southeastern Australia, Tasmania and New Zealand... ....... sisse e 431 (continued inside cover) CONTENTS JELL, P.A. Silurian and Devonian crinoids from central Victoria ................................... 1 JELL, Р.А. & THERON, J.N. Early Devonian echinoderms from South Africa .................................+... 115 JELL, P.A. & JELL, J.S. Crinoids, a blastoid and a cyclocystoid from the Upper Devonian reef complex of the Canning Eram Vent RIS AME, узу DL Mi hy duo CIR oath octet OP FS E os Kore Xon ed 201 WEBSTER, G.D. & JELL, P.A. New Carboniferous crinoids from eastern АпвїгаНа.................................. 237 WEBSTER, G.D. & JELL, P.A. New Permian crinoids from Australia... ec eee erem ehh hh rh hr 279 SMITH, A.B. & JELL, P.A. A new cornute carpoid from the Upper Cambrian (Idamean) of Queensland................ 341 RUTA, M. & JELL, P.A. Protocytidium gen. nov., a new anomalocystitid mitrate from the Victorian Latest Ordovician and evolution of the Allanicytidiidae ......................................... 353 RUTA, M. & JELL, P.A. Adoketocarpus gen. nov., a mitrate from the Ludlovian Kilmore Siltstone and Lochkovian Humevale Formation of central Victoria... 2.2.2... eee ne 377 RUTA, M. & JELL, P.A. Two new anomalocystitid mitrates from the Lower Devonian Humevale Formation of БШШ И CC vate eg Wis pie Buel CUS SK aie «Stale sta IEEE a бы кл 399 RUTA, M. & JELL, P.A. A note on Victoriacystis wilkinsi (Anomalocystitida: Mitrata) from the Upper Silurian ТЕЛЕ рез ЖМА oer Op as HA EET RONDE ERE RR CH ee alee egg Ug me RR A 423 RUTA, M. & JELL, P. À. Revision of Silurian and Devonian Allanicytidiidae (Anomalocystitida: Mitrata) from southeastern Australia, Tasmania and New Zealand. ............................. 431 (continued inside cover)