-V Bulletin of the British Museum (Natural History s c * , LIBRARY Zoology series Vol 45 1983 British Museum (Natural History) London 1983 Dates of publication of the parts No 1 28 July 1983 No 2 28 July 1983 No 3 25 August 1983 No 4 25 August 1983 No 5 29 September 1983 No 6 27 October 1983 No 7 .24 November 1983 ISSN 0007-1498 Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset Contents Zoology Volume 45 Page No 1 Problems in catfish anatomy and phylogeny exemplified by the Neotropical Hypophthalmide (Teleostei: Siluroidei). By G. J. Howes 1 No 2 Miscellanea A revision of the genus Epiclintes (Ciliophora: Hypotrichida) includ- ing a redescription of Epiclintes felis comb. n. By P. G. Carey & E. C. Tatchell 41 Notes on the Family Lekythoporidae (Bryozoa, Cheilostomata). By P. L. Cook & P. J. Hayward 55 A new species of Arthrolepis (Anura: Ranidae) from the West Usambara Mountains, Tanzania. By A. G. C. Grandison 77 The distribution behavioural ecology and breeding strategy of the Pygmy Toad, Mertensophyrne micranotis (Lov.). By A. G. C. Grandison & S. Ashe 85 Additional notes on bariliine cyprinid fishes. By G. Howes 95 No 3 Bats (Mammalia: Chiroptera) from Indo- Australia. ByJ. E. Hill . 103 No 4 On Macropleurodus, Chilotilapia (Teleostei: Cichlidae) and the interrelationships of African cichlid species flocks. By Peter Humphry Greenwood 209 No 5 Miscellanea D enter ammina (Lepidodeuterammind) subgen. nov., and a re- description of Rotalina ochracea Williamson (Protozoa: Forami- niferida). By P. Bronnimann & J. E. Whittaker . . . ... .233 The freeliving marine nematode genus Sabatieria (Nematoda: Comesomatidae). I. Two new species from Stonington Island, Antarctica. By H. M. Platt 239 New species of marine nematodes from Qingdao, China. By Z. N. Zhang & H. M. Platt .253 Echinoderms of the Rockall Trough and adjacent areas. I. Crinoi- dea, Asteroidea and Ophiuroidea. By J. D. Gage, Margaret Pearson, Ailsa M. Clark, G. L. J. Paterson & P. A. Tyler 263 No 6 The cranial muscles of loricarioid catfishes, their homologies and value as taxonomic characters (Teleostei: Siluroidei). By Gordon J. Howes . . 309 No 7 Miscellanea A lectotype for Deuterammina (Deuterammind) rotaliformis (Heron-Allen & Earland) and new trochamminids from E. Ireland (Protozoa: Foraminiferida). By P. Bronnimann & J. E. Whittaker . . . . . .347 Notes on Atlantic and other Asteroidea. 3. The families Ganeriidae and Asterinidae, with description of new asterinid genus. By Ailsa M. Clark .... .... 359 Peniculus haemuloni, a new species of copepod (Siphonostoma- toida: Pennellidae) parasitic on Haemulon steindachneri from Ubatuba, Brazil. By P. D. Alexander .381 Bulletin of the British Museum (Natural History) Problems in catfish anatomy and phylogeny exemplified by the Neotropical Hypophthalmidae (Teleostei: Siluroidei) Gordon J. Howes Zoology series Vol 45 No 1 28 July 1983 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in for scientific series, Botany. Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself, available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History), Cromwell Road, London SW7 5BD, England. World List abbreviation: Bull. Br. Mm. nat. Hist. (Zool.) Trustees of the British Museum (Natural History), 1983 The Zoology Series is edited in the Museum's Department of Zoology Keeper of Zoology : Dr J. G. Sheals Editor of Bulletin : Dr C. R. Curds Assistant Editor : Mr C. G. Ogden ISSN 0007-1498 Zoology Series VoUSNo 1 pp 1-39 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 28 July 1983 j WUbTWH Problems in catfish anatomy and phylogeny exemplified by the Neotropical Hypophthalmidae (Teleostei: Siluroidei) Gordon J. Howes Department of Zoology, British Museum (Natural History), Cromwell Road, London SW75BD Contents Introduction 1 List of genera examined 3 Form and function in Hypophthalmus and comparison with other siluroids . 4 Body shape, sensory canals and lateral line system 4 The eye and infraorbital bones 6 Cranial musculature . . . . 9 The snout and jaws 17 The cranium 21 The complex vertebrae, swimbladder and posterior lateral line nerve . . 25 The suspensorium 39 The hyoid arch 32 The opercular series 34 The gill arches 34 Postcranial elements 35 Relationships of the Hypophthalmidae 35 Acknowledgements 33 References 38 Introduction Of the subgroups of otophysan fishes, the siluroids or catfishes are the most morpho- logically diverse. None of the other subgroups, gymnotoids, characoids and cyprinoids, has such a range of body shape, peculiarity of dermal covering when present, and cranial morphology. Neither do these groups display such a range of postcranial architecture, complex fusion patterns of the caudal fin skeleton or such diverse forms of swimbladder. This morphological diversity is reflected in the most recent familial classification of the siluroids (Greenwood et al., 1966), which recognises thirty-one families, a number exceeding the combined familial total of other otophysan groups. As presently recognised, most siluroid families are morphologically well differentiated, suggesting that each is probably a monophyletic assemblage. Indeed, the large morphological 'gaps' separating families are emphasised by the fact that seven families are monotypic, cf one among characoids and two among gymnotoids. That the interrelationships of siluroid families have scarcely been studied is not surprising when one considers the paucity of anatomical data available. Even the largest families, i.e. the Pimelodidae, Ariidae and Bagridae are represented in the anatomical literature by only meagre information, the anatomy of some neotropical families, Auchenipteridae, Ageneiosidae, and Aspredinidae, has scarcely been described. The siluroids offer a wealth of character complexes involving the bony elements of the cranium, vertebral column and fin supports, and their associated muscles. These character Bull. Br. Mus. nat. Hist. (Zool.) 45( 1 ) : 1-39 Issued 28 July 1983 2 G. J. HOWES complexes lend themselves readily to the application of cladistic methodology. The task of analysis is, however, a daunting one. Not only is the number of taxa considerable, over 2000 species, but, by their very nature, siluroids are difficult to study. Most adult specimens are large and highly ossified, and sutures between bones are often obscured in dry skeletal material. It is therefore necessary to work, for the most part, with cleared and stained prep- arations of small, and frequently juvenile specimens. Thus problems can arise in the interpretation of features which undergo considerable ontogenetic change. One of the generalisations made about siluroids is that they are benthic fishes. In fact, the majority of species in seven families, Siluridae, Schilbeidae, Pangasiidae, Ariidae, Ageneiosidae, Cetopsidae, Hypophthalmidae, are mid- water to surface dwellers, and only in one third of families, Loricariidae, Amphiliidae, Callichthyidae, Astroblepidae, Doradidae, Chacidae, Sisoridae, Cranoglanididae, can the majority of included taxa be classed as entirely benthic. In the past more attention has been given to siluroid inter- than to intra- relationships. The most recent analysis (Fink & Fink, 1981) recognises the siluroids as a suborder, the Siluroidei, coordinate with their supposed sister group, the Gymnotoidei which together form the order Siluriformes. The Siluriformes are, in turn, thought to be the sister group of the characoids, order Characiformes, with the cyprinoids, order Cypriniformes, representing the plesiomorph lineage and sister group of the Silurifomes + the Characiformes. Only the most rudimentary analyses have been attempted on the functional anatomy of siluroids (Eaton, 1948; Alexander, 1965; Gosline, 1973 & 1975). According to Gosline (1973) siluroids employ a feeding mechanism functionally different from that of other otophysans. But again, lack of comparative anatomical data prohibits any worthwhile discussion of this issue. One attempt at cladistic analysis involving several siluroid families has been published by Lundberg & Baskin (1969). In order to obtain an understanding of siluroid relationships these authors chose to focus efforts '. . . on comparative studies of a single structural complex in obtaining an understanding of siluroid relationships'. This approach, whilst accumulating valuable comparative data, can however, produce a misleading phylogeny. The weakness inherent in an approach that places reliance on single character complexes is discussed below (p. 27) with reference to patterns of vertebral fusion and swimbladder encapsulation. Roberts (1973) notes that the '...cosmopolitan distribution of siluroids and their pseudarchaic characters have led ichthyologists to believe catfishes to be older than other ostariophysans'. I presume 'pseudarchaic' as used by Roberts means 'neomorph' resulting in convergences with features found in ancient groups. But, neomorphs are no indication of the group's phyletic age. On the other hand, the cosmopolitan distribution of siluroids may suggest an early and extensive distribution whereas the more restricted distribution of other otophysans could indicate their later cladogenesis from the ancestral otophysan lineage. The subject of this paper, the family Hypophthalmidae, is chosen for anatomical investi- gation because it represents several aspects of the problems discussed above. The family is monotypic, apparently 'isolated' phyletically, is 'pelagic' in habits, and poses intriguing problems of functional anatomy. The object of the study is to describe the osteological and myological features, with particular regard to the cranium, to analyse these features in terms of their apo-plesiomorphic status, to account for their functional relevance, and to use these data as indicators of phyletic relationship. Giinther (1864:3:66) first recognised the distinctiveness of Hypophthalmus by placing it in a separate subfamily, Siluroidae Anomalopterae (Group Hypophthalmina). Helogenes was also included on the basis of its elongate anal fin, and its short dorsal and adipose fins. Cope (1871) established the Hypophthalmidae as one of three families of his Nematognathi. In Cope's view Hypophthalmus was so aberrant that he doubted its relationships with other catfishes. Eigenmann & Eigenmann (1890) continued to recognise Gunther's concept of the family but Regan (1911) placed Helogenes in a separate family thereby establishing the Hypophthalmidae as monotypic. Chardon (1968) chose to recognise the uniqueness of the CATFISH ANATOMY & PHYLOGENY 3 Hypophthalmidae by placing it in its own suborder, the Hypophthalmoidei. He was, however, unable to relate the taxon closely with another of equal rank. As yet, the taxonomy of the genus Hypophthalmus has received no thorough treatment. Eigenmann & Eigenmann (1890) synonymised the six nominate species with H. edentatus, the type of the genus, stating There is little doubt but that the species have been based on the different stages of one species'. More recently, however, Roberts (1972:139) noted there are '. . . three very distinct species of Hypophthalmus in the Amazon'. In the most recent literature authors have recognised a second species named as H. perporosus Cope, 1878 (see Goulding, 1980;Carvalho, 1980a). Although Hypophthalmus is a common commercial food fish in Amazonia (see Roberts, 1972: 140), its ecology is poorly studied (for accounts see Roberts, 1972; Meschkat, 1960; Goulding, 1980; Carvalho, 19800 & b). Hypophthalmus species are shoalers and filter-feed, chiefly on zooplankton. According to Carvalho (19806), the larger proportion of ingested food of H. edentatus comprises cladocerans and copepods. Gut contents of specimens personally examined contain insect (Hymenoptera) leg and antennal fragments, and fragmented plant material (portions of leaves, stems, pollen cuticles and seed spikes). Carvalho (19806) classifies Hypophthalmus as a pelagic zooplanktivore but perhaps, in view of the above catalogue of gut contents, it should more rightly be classed as a detrital filter- feeder. As in other detritivores, Hypophthalmus has a long, convoluted gut. Only two papers deal with any detailed aspect of hypophthalmine anatomy. Wright (1885) described the skull and swimbladder connections. This paper, however, omits some important features and misinterprets others, omissions obviously due to limitations imposed by the use of dry skeletal material in which many delicate structures are lost or obliterated. Chardon (1968) also described the swimbladder and Weberian structure. His account and illustrations supplement those of Wright's. Roberts (1972) gives a more general account of Hypophthalmus, and includes a fairly detailed description of the gill arch structure and its likely mode of function. List of genera examined Where availability of material has permitted, the cranial anatomy of at least two species in each polytypic genus has been examined. A = alizarin stained specimen, D = dry skeleton, S = alcohol preserved specimens (dissected). Aelurichthys D Ageniosus A, D, S Amphilius A Arius A, D, S Aspredo S Auchenipterus A, D, S Bagrus A, D, S Brachyplatystoma S Branchioica A Bunocephalus A CallophvsusS Chaca D Chaetostoma D Chrysichthys A, D, S Clarias A, D, S Clarotes D, S Cochliodon A Cryptopterus D, S Diplomystes D, S Doras D Eutropius D, S Gagata D Galeichthys D, S Geneidens D Glyptosternon D, S Goldiella S Helogenes A, D, S Hemisorubim D, S Heptapterus S Hoplosternum D, S Hypophthalmus A, D, S Ictalurus A, D, S Iheringichthys A, S Leiocassis D Liobagrus D Loricaria D, S Luciopimelodus D, S Megalonema S Mystus D Nematogenys D, S Notarius D Osteogeniosus A Ompok D, S Oxydoras D Pangasius S Pangasianodon D Parapimelodus S Perugia S Phyllonemus A Pimelodina S Pimelodus A, D, S Platysto malic hthys D Plotosus D, S Pseudauchenipterus S Pseudobagrus A, D Pseudocetopsis S Pseudopimelodus D, S 4 G. J. HOWES Pterodoras D Silurus D, S Tandanus D, S Rhamdia D, S Sorubim A, D, S Trichomycterus A, S Rhinodoras D Soruhimichthys D, S Wallago D, S Schilbe A, D, S Synodontis S Abbreviations used in the figures are given either in the relevant sections of the text, or in the figure captions. Form and function in Hypophthalmus and comparison with other siluroids The principal derived features of Hypophthalmus are listed below. The reasons for considering their derived nature are discussed under each character complex. 1 . Body shape and pattern of the sensory canals and lateral line system. 2. The position of the eyes and arrangement of their musculature. 3. The hypertrophy of the facial, and the complexity of the hyoid musculature. 4. The particular morphology of the mouth and the articulation of the mandibular barbels with cartilaginous plates. 5. The reduction of the neurocranium and its complete ankylosis with the anterior vertebral complex. 6. The nature of the complex vertebrae, anterior placement of the Weberian ossicles, reduction plus encapsulation of the swimbladder, and displacement of the vagus nerve. 7. The extent of the gill openings and elongation of the branchial arch elements. 8. The elongation and modification of the hyoid arch. Body shape, sensory canals and lateral line system (Figs 1 & 2) Hypophthalmus has a deep and moderately compressed body with a sloped dorsal profile; the head is broad (30% of its length) with a subtriangular snout, the eyes are set at the same horizontal level as the mouth. There is a single pair of long maxillary barbels and two pairs of similarly long mandibular barbels. The gill opening extends almost as far forward as the mandibular symphysis. The pectoral fin is narrowly triangular, its tip reaching beyond that of the small pelvic fin. The first ray of the pectoral fin is spinous and finely serrated along its outer margin. The anal fin is long with c. 62 branched rays, extending from the tip of the pelvic to below the adipose fin. The dorsal fin is short-based with 1 + 6 rays; the small adipose fin lies midway between the dorsal fin and the origin of the forked caudal fin. In life, Hypophthalmus edentatus has a silvery blue-green colour, the barbels are dark, the fins hyaline but with dusky tips to the pectorals, and a dark submarginal band on the anal fin. The skin above the orbit is permeated with a network of canals which are denser and more ramified below the frontal margin than elsewhere (Fig. 2). The entry points to this network are through the pterotic and frontal sensory canal openings. Similar ramified patterns occur over the antero-dorsal and ventral portions of the operculum. Their respective connections are with the pterotic and preopercular canals. The lateral line has dorsal and ventral branches. The longest dorsal branches are angled caudally at 45 to the main lateral line, some extending almost to the dorsal midline; shorter branches run cranially at a similar angle, thus forming a network (Fig. 1 ). The longest ventral branches extend to the base of the anal fin where they curve caudally almost at a right angle (one or two branches curve anteriorly). Shorter, angled branches cross the upper part of the vertical lines and form the ventral counterpart of the dorsal network. The lateral line is bifurcated caudally, the longer arm, itself often sub-branched, extends to the 6th or 7th ray of the upper caudal lobe. The lower branch hardly reaches the base of the caudal fin and is absent in some specimens. CATFISH ANATOMY & PHYLOGENY Fig. 1 Hypophthalmus edentatus\ (upper) lateral view showing ramifying lateral line system; (lower) ventral view of head to show inferior position of the eye. Scale = 20 mm. Drawn from specimen BMNH 1925. 10.28:327. The body shape of Hypophthalmus is like that described by Alexander (1965) for the African siluroid Schilbe. The salient features noted by Alexander of the schilbeid morpho- type are a compressed trunk; long anal fin; short body cavity; reduced dorsal and adipose fins; deep epaxial musculature reaching well-forward on the cranium and the supraoccipital bearing a tall median crest onto which most of the dorsal musculature attaches. All these features apply to Hypophthalmus and occur also in Old-World siluroids (Siluridae e.g. Ompok, Cryptopterus; Schilbeidae, Pangasiidae) and the Neotropical Auchenipteridae (Auchenipterus) and Ageniosidae. As such it seems to be a morphotype independently derived in several lineages. Nonetheless, such a body shape is assumed to be a derived feature for each group in which occurs as it is always accompanied by some marked, but different modification of the swimbladder and, or, lateralis system. Carvalho (19800) found a high proportion of fat in Hypophthalmus (up to 26% body weight) which varied inversely to the total water content. This high fat content doubtless contributes to the attainment of neutral buoyancy and appears to be a correlate of a reduced swimbladder (see p. 28). A highly ramified lateral line system similar to that in Hypophthalmus is also found in siluroids and pangasiids, but in no other taxon investigated do the lower branches extend so far ventrally or curve along the anal fin base. An orbital and opercular canal system like that of Hypophthalmus occurs similarly only in Luciopimelodus. A ramifying of canals are present on the cranial and postcranial regions of many siluroids, but generally as separate twiglets over the head and upper opercular region only. In the Ageneiosidae and Pimelodidae alone is there a lattice-like pattern aproaching that of Hypophthalmus. Further- more, only in Brachyplatystoma and Luciopimelodus does such a system of canals cross the interoperculum. Other siluroids with a similar body shape to Hypophthalmus (e.g. Pangasius, Schilbe, Ompok} lack an intricate epidermal network. Alexander (1965) observed that catfishes possessing the schilbeid morphotype (see above) were more pelagic than others, and accounted for the compressed, tapered body shape in terms of a specialised locomotion. A principal component of schilbeid locomotion is the ability to hover by inclining the body at a 45 angle to the surface. It has not been reported whether Hypophthalmus adopts a similar locomotory attitude. Neither is it clear whether Hypophthalmus should be classed as 'pelagic' for it seems that these fishes may make vertical migrations (Meschkat, 1960). G. J. HOWES io4-5 io1-3 Fig. 2 Hypophthalmus edentatus lateral view of head showing sensory canal system and infra- orbitals. Composite drawing. The eye and infraorbital bones (Figs 2-4) Two of the more noticeable features of Hypophthalmus are the position of the eye relative to other parts of the cranium, and the arrangement of the orbital muscles (Fig. 3). The eye is of moderate size (12% of head length) and is situated on the ventral margin of the head at the same horizontal level as the mouth. When viewed ventrally the eyes appear as almost entirely inferior (Fig. 1 ). The eye muscles, in their origins and position lateral to the adductor muscle complex, are unlike those of any other siluroid. The anterior muscles are long and strap-like. Both the superior obliquus (sob) and inferior obliquus (iob) have a common origin from a broad tendinous sheet that extends dorsally across the adductor mandibulae musculature to attach to the lateral margin of the frontal and to the lateral ethmoid. The optic nerve and those supplying the eye muscles all run parallel to one another within an almost vertical band of connective tissue. As in other siluroids the superior obliquus is innervated by the trochlear nerve (niv) and the inferior obliquus by a branch of the oculomotor (niii). The rectus internus (ri) arises from a thin tendinous strand that crosses the face of the adductor complex and runs between the superior obliquus and inferior obliquus muscles to attach to the anteromedial surface of the eyeball. The muscle is unnervated by a sub-branch of that branch of the oculomotor that feeds the inferior obliquus. The posterior set of muscles, rectus superior, r. inferior and r. externus (Fig. 4, rs; rin; re) originate from a tendinous band which is strongly attached to the dermis. The former two muscles are innervated by branches of the oculomotor and the latter by the abducens (nvi). The optic nerve (nii) is thick and exits from a slit-like foramen to loop over the adductor muscle complex and pass across its face which is grooved to accommodate it. The infraorbital bones (io Fig. 2) are, with the exception of the 3rd, reduced to ossifications of the sensory canal. The 1st infraorbital is axe-shaped, the common morphology of this element amongst siluroids, the 2nd a slightly curved, cylindrical bone, the 3rd with an CATFISH ANATOMY & PHYLOGENY aap et ao lo nso am mn Fig. 3 Hypophthalmus edentatus; (upper) superficial dissection of the head showing position of the eye muscles in relation to the cheek musculature; (lower) dorsal view of orbital region (eye muscles removed) with frontal cut away to display the dilatator operculi muscle (do) and olfactory nerve (ni). Drawn from specimen BMNH 1976.6.18: 123-129. Scales = 5mm. am = adductor mandibulae; nso = rami of supraorbital nerve trunk. antero-ventral lamellae, its posterior, tubular portion lying beneath the orbit; the 4th, the largest of the series, curves dorso-posteriorly to beyond the border of the eye, then, at the upper level of the orbit, curves dorso-anteriorly to extend across the interspace between the orbit and the cranium; the 5th a small tubular bone connected to the pterotic. Although unusual, the ventro-lateral position of the eye in Hypophthalmus is not unique amongst siluroids. It occurs similarly in Ageniosus, some Auchenipterus and Pangasius species. However, in all these taxa, the arrangement of the eye muscles is of the more normal teleost condition. In both Auchenipterus and Pangasius the eye ball is cupped by the surrounding muscles, the posterior set passing into the optic foramen. Ageniosus is markedly similar to Hypophthalmus in the displacement of the optic nerve tract across the adductor muscle face. However, the musculature is extended laterally and there is a consequent orientation of the nerve tract, unlike the vertical course followed in Hypophthalmus. Furthermore, the eye muscle arrangement in Ageniosus is less modified in that the posterior muscles, although highly tendinous, join in a single cylindrical tract across the adductor mandibulae muscle. niv Fig. 4 Hypophthalmus edentatus; the eyeball, associated muscles and nerves (the upper skin reverted). Drawn from specimen BMNH 1976.6.18: 123-129. Scale = 5 mm. Only in the pimelodid Sorubimichthys is there a muscle arrangement approaching that of Hypophthalmus. Although in the former genus the eye is superior in position and the posterior set of muscles are horizontally aligned, the anterior muscles originate from a tendinous band above the extensor tentaculi muscle (et Fig. 10). The inferior orientation of the eyes in Hypophthalmus may provide a partially stereo- scopic vision which would serve to alert the fish to an approaching predator. The degree of eye mobility in Hypophthalmus is unknown. The eye muscles are thinner, and appear 'weaker' than in the other taxa investigated, but their complete innervation suggests some degree of eye-ball movement. According to Dullemeijer & Barel (1977) the eye in teleosts is a functionally dominant feature and in their opinion one with little positional plasticity. This is obviously not the case in Hypophthalmus where the eye has shifted ventrally from the usual siluroid supero- lateral position. In Hypophthalmus the position of the eye has apparently been 'dictated' by the form of the adductor muscle complex. This is in direct contrast to phytoplankton feeding percomorphs (Dullemeijer & Barel, 1977) where the eye has retained a superior position and consequently only a small area is available in which the adductor muscle can develop and operate. In general, the eyes of siluroids appear to have a less dominant role than in other otophysans or indeed other teleosts. In many siluroids the trend has been for the eyes to have shifted into a superior position, taken to the extreme in some loricarioids (e.g. Chaetostoma and Astroblepus] where the orbits are separated by a minimal space. CATFISH ANATOMY & PHYLOGENY 9 Fink & Fink (1981) claim that the infraorbital series in siluroids is reduced to the canal bearing portion of the bone. Whilst this is true for the majority of taxa it does not hold for all; Hypophthalmus is one exception (see above). In the clariids, the posterior infraorbitals (includes 'supraorbital' aucct.} are greatly expanded. The recognition of reduced infraorbitals and the absence of a supraorbital bone as synapomorphies linking siluroids with gymnotoids (Fink & Fink, 1981) is dubious. Such reduction and loss also occurs in cyprinoids (Cobitidae). An even more pertinent point is the differing arrangement of the infraorbitals in gymnotoids and siluroids clearly shown by Fink & Fink (1981, fig. 7D-E). The anterior infraorbital bones of siluroids are always elongate and the 3rd extends someway past the orbit so that the posterior series curve dorso- anteriorly. In gymnotoids the 3rd infraorbital is a short bone and the posterior elements are directed diagonally in a dorso-posterior direction. Cranial musculature (Figs 5-6) Jaw and suspensorial muscles When the eye, together with its muscles and the optic tract have been removed, the entire face of the fish is seen to be covered by an almost undifferentiated adductor muscle (Fig. 5). The orbital socket is indicated on the face of the muscle by a circular tendinous area, the optic tract by a vertical groove (otf, Fig. 5). Dorso-posteriorly, lateral fibres of the levator arcus palatini muscle (lap) adhere to the adductor's surface, and a more medial section of the levator is indicated by a longitudinal division of the adductor. The posterior origin of the adductor mandibulae is from the lateral and antero-medial faces of the hyomandibula. Anteriorly, at the articulation of the lower jaw, there is a myocomma and the adductor can be differentiated into a number of segments. The upper, outer portion curves inward, its anterior border being tightly attached to the thick mandibular nerve tract (nvmd) which follows the muscles contour as it passes medially to lap otf etb etc eta Fig. 5 Hvpophthalmm edentatm\ superficial head musculature. Drawn from specimen BMNH 1976.6.18: 123-9. Scale = 5 mm. 10 G. J. HOWES aap etb nvmd atm pal amm bmx A 2 +A, Fig. 6 Hypophthalmus edentatus; lower jaw musculature. The palatine and extensor muscle have been raised. In relaxed position the tendon (atm) passes between the rami of the maxillaris nerve (nvmx). Drawn from specimen BMNH 1976.6.18: 123. Scale = 3 mm. attach to the inner face of the lower jaw. From its insertion this part of the muscle is regarded as section A 2 (Fig. 6). There is no recognisable A w , A 2 inserting via a broad tendon stretching across the medial face of the adductor mandibulae mass. The antero-ventral part of the muscle lying lateral to the mandibularis nerve trunk is divisible into an upper bundle of fibres inserting into the lip tissue covering the coronoid process, and a ventral section inserting partially into labial tissue and partially on the lateral face of the dentary (A,, Fig. 5). When the superficial layer of the dorsal part of A, is removed, a medial layer is found to be continuous with A 2 (amm, Fig. 6); this division appears to be homologous with the retractor tentaculi (see below). The posterior border of the ventral part of A, is marked by a myocomma. A short, pinnate bundle of fibres stems from the antero- medial face of the adductor complex and inserts into a long tendon (atm, Fig. 6). The tendon passes between the maxillary and mandibular branches of the trigeminal nerve to insert on the maxilla at the base of the barbel. There is no discrete medial division of the main body of the adductor mandibulae, only posteriorly is there a separation by a medial sheet of the levator arcus palatini muscle (see above), but the fibres of both muscles intermesh. The central portion of the adductor arcus palatini extends between the parasphenoid and the dorsal border of the pterygoid bones (aap, Fig. 6). A separate, posterior portion runs from the parasphenoid to the medial face of the hyomandibula, and is identified as the adductor hyomandibulae. The anterior part of the adductor arcus palatini is divisible into three sections which together comprise the extensor tentaculi. The antero-dorsal part of this muscle covers the lateral ethmoid and attaches to the posterior half of the palatine (eta, Fig. 5). The postero-medial part originates as two separate sections from the central region of the adductor arcus palatini. The longer, postero-medial section runs forward at an oblique angle to insert on the postero-medial portion of the palatine (etb, Fig. 5). The shorter, medial portion runs antero-dorsally to insert on the antero-medial part of the palatine (etc, Fig. 5). Innervation Two thick nerve tracts stem from the trigeminofacial ganglion and exit from a single prootic foramen. The supraorbital trunk diverges immediately. The infraorbital trunk is of great CATFISH ANATOMY & PHYLOGENY nvmd , rs aap ct 11 Imb am Fig. 7 Nematogenys inermis; dorso-lateral view of cheek musculature. Drawn from specimen BMNH 1883.li.27: 45-48. Scale=10mm. co = coronoid process; ct = connective tissue; 1mb = ligament connecting maxillary barbel with mandible. thickness, and above the jaw articulation it divides into the buccal and maxillo-mandibular rami which pass, respectively into the root of the maxillary barbel and the lower jaw. (Fig. 6). The buccal ramas gives off a branch to the adductor arcus palatini before bifurcating into an outer branch serving the area of the extensor tentaculi muscle around the palatine, and an inner branch which enters the base of the maxillary barbel. The mandibular ramus (nvmd) curves round the anterior border of the adductor muscle complex, sending off short branches into the musculature (Fig. 6). The trunk passes across the inner face of the dentary to run inside the length of the bone before terminating in a bifurcation, each branch of which enters its respective barbel (Figs 7 & 1 4). Jaw and suspensorial muscles in other siluroids It is generally accepted (see Winterbottom, 1974) that in teleosts the section of the adductor mandibulae termed A, is that which forms the outer, or dorsal muscle segment inserting on the maxilla. Section A 2 is that which lies medially and, or, ventrally to A,, and inserts on the lower jaw; the part of it which covers the medial face of the dentary is termed A w . Further divisions (A 2 a, A 3 etc) are those successively medial to A, and A 2 and which insert onto the inner face of the lower jaw. In siluroids, this seemingly straightforward nomenclature has become somewhat confused through the presence of a muscle, serving the base of the maxillary barbel, known as the retractor tentaculi, often referred to as the adductor tentaculi or protractor maxillae; see Winterbottom, 1974 for synonymy. The fact that this muscle inserts on the maxilla but stems from the medial aspect of the adductor mandibulae has led to its being homologised with various sections of the adductor complex. Takahasi (1925) thought that because of its maxillary insertion it was synonymous with division A,, a hypothesis followed by Edgeworth (1935). McMurrich (18846) and Lubosch (1938) considered the retractor tentaculi to be an inner division of the adductor. Eaton (1948) agreed, in referring to Ictalurus thus . . .'From the fact that a stout ligament runs from the base of the adductor tendon back and downward to join the ligamentous sheath of the 12 G. J. HOWES Imx Fig. 8 Callophysus macropterus; lateral view of muscular and ligamentous connections to the upper and lower jaws. Drawn from specimen BMNH 1913.7.30: 20-22. Scale = 5 mm. Imx = maxillary-mandibular 'ligament'; pet = palato-pterygoid connective tissue. quadrate-articular joint, I would infer that the muscle has simply taken hold, as it were, of the primitive ligament which runs, in such fishes as Amia calva, from the articular to the maxillary. Thus, the ligament has now become in large part a muscle tendon. . .' Alexander (1965) concurs with Eaton, believing that the retractor tentaculi (the adductor tentaculi of Alexander) is derived from '. . .A 3 rather than A,'. From the arrangement in siluroid taxa I have investigated, it appears that these latter authors are probably correct in assuming that the retractor tentaculi is part of the inner adductor series and that its attachment with the upper jaw is a de novo development. The possible evolutionary development of the retractor tentaculi as suggested by Eaton (1948) and Alexander (1 965) can be amplified by the following observations. The presumed plesiomorph condition of the adductor mandibulae muscle in siluroids is where divisions of the muscle are not well-differentiated and insertion is entirely, or almost entirely, on the lower jaw. Diplomystes and Nematogenys (Fig. 7) illustrate this condition where the lower, outer part inserts musculously on the lower jaw (anguloarticular). An upper, inner division can be distinguished by its attachment, anteriorly, to the lower part by a broad vertical tendon which covers the postero-dorsal rim of the lower jaw. Stemming from the antero-medial surface of this muscle, and extending across the dorsal surface of the mandible is a thick sheet of connective tissue (ct). This sheet bifurcates, the upper strand attaching to the posterior face of the maxilla, the lower to the distal portion of the maxilla where it forms a sheath around the maxillary barbel. In Callophysus, Pimelodidae, a continuous sheet of tissue extends medially from the dentary to the anterior pterygoid process (pet, Fig. 8). That area of tissue covering the coronoid process is thickened and, anteriorly, is separated from the main sheet so forming a distinct 'ligament' running to the base of the maxillary barbel (Imx, Fig. 8). A similar arrangement of the 'ligament' is present in Tandanus and Plotosus. Alexander (1965), following Takahasi (1925), states that the deepest part of the adductor mandibulae inserts on the articular-maxillary ligament. This is, however, incorrect. A strong ligament does indeed stretch from the posterior aspect of the maxilla to the coronoid process of the dentary, and appears to be homologous with the partially differentiated 'ligament' described above in Callophysus, but no muscle inserts on it (Imx, Fig. 9). The outer portion of the adductor mandibulae in Tandanus and Plotosus inserts on the lower jaw medial to the ligament. The inner sections of the adductor, be they interpreted as A 2 or A 3 , insert on the medial face of the anguloarticular (Fig. 9). CATFISH ANATOMY & PHYLOGENY 13 et mx Imx bmx Fig. 9 Tandanus tandanus; dorso-lateral view of jaw muscle attachments. Drawn from specimen BMNH 1871.2.10: 4-6. Scale= 10 mm. Kesteven (1943) also erroneously identified a retractor tentaculi, his protractor maxillae, in Tandanus. He states that the muscle '. . .arises from the side of the skull above the anterior attachment of the palatal arch. . .' to insert on the maxilla '. . .enswathing the posterior end of the bone.' I find no such muscle in Tandanus tandanus (Fig. 9) nor in Plotosus plotosus and must presume Kesteven misidentified part of the extensor tenatculi or else did not entirely dissect the connective tissue enswathing the maxilla. In the pimelodids Pimelodus and Iheringichthys and the bagrid Clarotes a distinct ligament connects the base of the maxillary barbel with the coronoid process of the dentary. However, the ligament then courses posteriorly from the coronoid process to the fascia of the medial section of the adductor mandibulae. A further modification of this condition occurs in one group of pimelodids, Parapimelodus and Pimelodus maculatus, whereby there are two distinct ligaments extending from the base of the barbel, the outer attaching to the coronoid process and the medial one to the pterygoid process. In none of these taxa does the musculature attach to the ligaments. In Megalonema, Pimelodidae, the maxillo-articular ligament bifrucates posteriorly, the inner branch becoming continuous with the tendon of the medial segment of the adductor muscle complex. The outer ligament is, however, still connected with the lower jaw. In Sorubimichthys, Pimelodidae, a distinct ligament connects the maxilla with the coronoid process of the dentary. However, the ligament is tightly attached along the length of the upper jaw (premaxilla) by connective tissue. A branch of the mandibularis nerve runs into the tissue joining the upper and lower jaws (Fig. 10). These examples serve to illustrate the probable pattern involved in the evolution of the retractor tentaculi. The hypothesised stages of this process are; (1) the palato-mandibular connective tissue becomes differentiated to form a stout, free ligament connecting the maxilla with the posterior part of the lower jaw, (2) medial fibres of the adductor mandibulae become associated with the maxillo-palatine part of the palato-mandibular tissues, (3) the connective tissue becomes further differentiated and tendinously links a discrete medial section of the adductor mandibulae with the root of the maxillary barbel. The basic conditions for the development of the retractor tentaculi are present in several members of the Pimelodidae, Bagridae and Plotosidae and it seems very likely that the muscle has been derived independently in several lineages. Various apomorphic conditions of the muscle can be recognised as for example in some pimelodids such as Rhamdia where the anterior segment is attached to the orbitosphenoid via a bifurcated tendon. 14 do am nvmd pmx Imb Fig. 10 Sorubimichthys planiceps; dorso-lateral view of superficial jaw musculature. Drawn from specimen BMNH 1977.5.24: 12. Scale = 5 mm. em = eye musculature, 1mb = maxillary barbel- mandibular ligament. If the adductor mandibulae muscles of siluroids are compared with those of other otophysans, their arrangement is seen to correspond most closely to that of characoids, where section A, also inserts on the dentary. Vari (1979: 317) considered that a lower jaw insertion for the outer part of the adductor represented the plesiomorph condition. In characoids a strong, well-differentiated maxillary-mandibular ligament is present and often the dorso-lateral fibres of A, attach to this ligament. Vari (1979) notes that incorporation of the ligament into the tendon of A, is a derived condition in distichodontid characoids. In cyprinoids the section A, never inserts on the lower jaw and there is no maxillary- mandibular ligament. In gymnotids there is a maxillary-mandibular ligament and in Gymnotus a few fibres of the outer part of the adductor muscle insert on this ligament; the remainder of the muscle inserts onto the outer face of the lower jaw, as in siluroids. However, this is not the case in Sternopygus, Eigenmannia and Rhamphichthys the only other gymnotid taxa examined, where the outer segment of the adductor attaches to the 1st infraorbital in Sternopygus and Eigenmannia, and to both the 1st infraorbital and the maxilla in Rhamphichthys. Two hypotheses are available to account for the arrangement of the outer adductor muscles in siluroids. 1. That the insertion on the lower jaw is plesiomorphic and the ligamentous strand connecting the maxilla with the dentary is homologous with that similar ligament in characoids. 2. That insertion on the lower jaw is a derived feature brought about by regression of the maxilla and the loss of its intimate connection with the lower jaw. In effect, a regression of the cyprinoid condition. Fink & Fink (1981) favour the second hypothesis stating . . .'The conditions in some characiformes and in siluriformes are hypothesised to be secondary reductions from a primitive attachment to the maxilla.' Although Fink & Fink refer to some Characiformes, the outer element of the adductor mandibulae attaches to the lower jaw in the majority of characiform taxa. In those siluroids where the adductor mandibulae inserts on the upper jaw, Loricariidae, Astroblepidae, it is to the premaxilla and the muscles involved are derivatives of the medial and not the outer parts of the adductor complex (Howes, in press). Only in the Callichthyidae is there an insertion of A, on to the maxilla, and this appears to be a derived condition associated with the maxillary-ethmoid joint (Howes, in press). In view of these CATFISH ANATOMY & PHYLOGENY 15 observations I would regard a lower jaw insertion of the adductor madibulae A, in siluroids as the plesiomorph condition. The use of this character by Fink & Fink (1981) uniting the siluroids and the gymnotoids cannot be upheld, as in some (?the majority) of gymnotoids, the outer part of the adductor muscle inserts on the 1st infraorbital and not the lower jaw (see above). The opercular muscles (Figs 3 & 5) The dilatator operculi (do, Figs 3 & 5) is an exceptionally long muscle having its area of origin anterior to the orbit, the fibres adhering to, and covered by, the ventral margin of the frontal. Posteriorly, above the optic tract, the muscle is exposed before becoming tendinous and passing between the adductor mandibulae and the medial section of the levator arcus palatini. The long insertion tendon attaches to the antero-dorsal process of the operculum. The levator operculi (lo Fig. 3) comprises two thin segments, an upper one arising from the postero-dorsal face of the hyomandibula and inserting on the postero-dorsal margin of the operculum, and a medial one originating from the pterotic and inserting on the antero- medial face of the operculum. The adductor operculi (ao Fig. 3) originates from a fossa indenting the pterotic, exoccipital and 'posttemporaF. It inserts on the dorso-medial face of the operculum. The hyoid muscles (Figs 1 1 & 12) The ventral head muscles have a complex arrangement with much integration of the various segments. The ventral layer exhibits two groups of muscles; those in which the fibre direction is toward the midline and those in which it runs antero-laterally. The largest element (hma, Fig. 11 A), belonging to the latter group, occupies half the lateral aspect of the lower jaw. Its fibres run at an oblique angle from the anterohyal to the dentary; anteriorly they change direction to run almost longitudinally. The outermost fibres insert onto the forepart of the dentary whilst the remainder turn inward as a separate bundle and are incorporated into the medial section (hmb, Fig. 1 1 A). This medial part of the muscle also originates from the anterior part of the anterohyal. The muscle is formed from two sheets. The ventral, medial portion (hmbl, Fig. 1 1 A) curves antero-medially, its outer fibres inserting on the cartilaginous plate of the inner mandibular barbel, its inner fibres meeting those of its antimere at a midline raphe. The lateral portion (hmb2, Fig. 11 A) runs anteriorly to insert on a tendinous pad covering the cartilaginous base of the outer mandibular barbel (see below). A separate bundle of fibres insert into the base of the barbel. The medial fibres ofhmb2 overlie the lateral aspect of hmb 1 , but at the midline both muscle- layers meet at a raphe. Covering the branchiostegal rays and extending forward as a thin, longitudinally fibred segment is muscle hmc (Fig. 11 A). Anteriorly it joins an aponeurosis into which hmbl inserts; medially it joins its partner at a midline raphe. Extending forward from the aponeurosis is a discrete bundle of fibres that attaches to the base of the inner mandibular barbel (hmd, Fig. 11 A). The large muscle (hma) is considered, from its disposition and innervation by the ramus hyoideus VII to be the posterior protractor hyoidei. The inner divisions (hmbl and hmb2 are also innervated by branches of the r. hyoideus VII and appear to be anterior parts of the protractor hyoidei. The small muscle slip attaching to the outer mandibular barbel is merely a continuation of the dorsal portion of this muscle and can be identified with that element named by Singh (1967) as a retractor tentaculi, not to be confused with his use of this name also for the facial muscle. The longitudinal muscle (hmc) is the hyohyoidei adductores. It is unclear, however, if the small bundle of fibres (hmd) running from its anterior aponeurosis to the inner mandibular barbel is indeed part of this muscle or whether it is derived from the protractor hyoidei. This small barbel muscle should also be identified with Singh's (1967) retractor tentaculi, which Winterbottom (1974) considered to be homologous with the 16 G. J. HOWES im Fig. 11 Hypophthalmus edentatus; A, ventral view of hyoid musculature (right side); B, dorso- medial view of mandibular barbel supporting elements (left side). Thick dashed lines = nerve path, thin-dashes and dots = outline of barbel root; C, lateral view of mouth, upper lip cut sagittally. Drawn from specimens BMNH 1972.7.27: 675-678 & 1976.6.18: 123. Scales = 5 mm ul = upper lip. retractor hyoidei. Ghiot (1978) illustrated in Pimelodus five separate muscles controlling the mandibular barbels of which he ascribed origin to 'differentiation of the hyoid protractor'. The intermandibularis (im, Figs 1 1 A & C) forms the anterior transverse muscular floor of the mouth. It is divided medially and each segment is attached to its respective cartilaginous plate which serves to support the barbel (see below). In the midline, each half of the muscle joins in a raphe which, in turn, is attached by strong connective tissue to an overlying collar of fatty tissue (ft, Fig. 11C). This collar encircles the leading edge of the antero- and dorsohyals and covers the basihyal. Only in some pimelodid taxa is there such a complex arrangement of the hyoid musculature as in Hypophthalmus. In Pimelodus, Sorubim and Pseudoplatystoma the protractor hyoidei is complexly aponeurotic, and for its greater part is attached to the large CATFISH ANATOMY & PHYLOGENY 17 ib ha Fig. 12 Sorubim lima; ventral view of hyoid musculature (dashed lines = outlines of barbel supporting plates. Drawn from specimen BMNH 1969.7.15.26. Scale = 5 mm. ib, ob = inner and outer mandibular barbels. cartilaginous plates supporting the bases of the barbels (Fig. 12). Ghiot (1978) distinguished in Pimelodus clarias five separate sections of what he termed the superior hyoidian protractor muscle in addition to separate lateral and medial divisions. The complexity of the hyoid musculature with separate mandibular barbel muscles attached to basal cartilages appears to be derived for at least one group of pimelodids and the Hypophthalmidae. The usual condition in siluroids is for the mandibular barbels to be 'embedded' within the protractor hyoidei, with bundles of fibres inserting on the anterior and posterior bases of the barbels and performing the function of retractors and protractors. This arrangement of hyoid musculature is illustrated by Singh (1967) in Bagridae and Winterbottom (1974, fig. 12) in Ictaluridae. Unlike other otophysans, the siluroids exhibit considerable variability in the morphology of the hyoid musculature. A comparative study of these various arrangements appears to offer a fruitful field of potential synapomorphies. The snout and jaws (figs 11-15) Ramsay Wright's (1885) description of the snout region in Hypophthalmus can hardly be bettered and the following account is merely a supplement. The entire ethmoid region is of papyraceous bone interspersed with cartilaginous strips (ec). Such cartilaginous areas separate the ethmoid (eth) from the lateral ethmoids and are bridged by the spine-like nasal bones (na, Fig. 13 A). It is not possible to determine whether the ethmoid bone is a supra- or a mesethmoid. That it is so thin, and its lateral and posterior 18 pmx le mx Fig. 13 Hypophthalmus edentatus; (upper) dorsal view of ethmoid region, (lower) palatine- maxillary articulation. Scales = 5 mm. Drawn from specimen BMNH 1972.7.27: 675-678. margins grade into cartilage suggest that it is a single element and is entirely mesethmoidal in origin. The lateral ethmoid (le, Fig. 13) has a wide anterior surface with a convex antero-lateral margin; postero-laterally its border is produced into a long, ventrally directed spine which almost meets the third infraorbital. The medial border of each lateral ethmoid abuts the midline mesethmoidal cartilage. The vomer is needle-like, its head slightly broadened and resting below the ethmoid (v, Fig. 1 3). The nostril is bordered medially and laterally by the anterior arm of the bifurcated 1st infraorbital. The edentulous premaxillaries (pmx, Fig. 13) are barely identifiable as separate elements, being fused with the ethmoid and distinguishable only by their denser ossification. They are so firmly united at the symphysis that even under a magnification of 50 x a suture is not visible in alizarin preparations. The maxilla is a small, well-ossified, flattened and hook- shaped bone (mx, Fig. 13). The blade of the hook supports the barbel (bmx), its base is rounded and articulates withe the cartilaginous tip of the palatine. The rod-like palatine (pal) is ossified, apart from its anterior and posterior tips and a small medial disc at about its midpoint (Fig. 1 3). This disc articulates with the lateral border of the lateral ethmoid. Like the upper jaw, the lower jaw is also edentulous. The dentary (d) contributes to half the length of the mandible and is strongly sutured to the anguloarticular (aa). The coronoid process is formed largely by a dorsal extension of the Meckelian cartilage (com, Fig. 14). The horizontal segment of this cartilage extends anteriorly along the medial face of the dentary and also has a slight posterior extension. The mandibularis nerve passes across the medial face of the upright part of the cartilage and then between the horizontal segment and the dentary wall. The nerve trunk continues along the ventral rim of the dentary and then bifurcates, each ramus inserting into the base of a corresponding mandibular barbel. A separate rod-like cartilaginous element lies antero-dorsally to the coronoid cartilage (cr, Fig. 14). Its distal tip is forked, and attaches to the lateral fold of thick labial tissue. The cartilage is mobile. When the jaws are abducted, and the labial tissue made taught, the cartilaginous rod rises to an acute angle. CATFISH ANATOMY & PHYLOGENY 19 nvmd .com Fig. 14 Hypophthalmus edentatus, lower jaw (right) in medial view. Drawn from specimen BMNH 1972.7.27: 675-678. Scale = 5cm. com = coronoid process of Meckelian cartilage; lt = labial tissue. The barbels are supported on three cartilaginous plates (cp, Figs 1 IB & 14). The first, i.e. the element closest to the jaw symphysis, is the most complex. Its flat proximal tip attaches to the dentary rim and is curved where the mandibularis branch passes between it and the dentary. The cartilage extends dorso-posteriorly as an open cylinder, its distal tip providing the insertion site for the inter mandibularis muscle. The root of the inner barbel (ib) bifurcates, one part ending as a short hook on the outer part of the cartilage, the other extending along the outer side of the cylindrical section. The two inner plates are irregularly shaped. The 2nd, middle, plate contacts the symphysial plate dorsally; the 3rd, outer, is notched at its articulation with the dentary thus allowing transmission of the mandibular nerve branch to the outer barbel (ob). As with the inner barbel the outer one is also bifurcated, its thicker leg attaching to the 3rd plate, its inner, thinner leg becoming 'cartilaginous' and extending posteriorly to join the centre of the 2nd cartilaginous plate. Uniquely amongst siluroids, the anterior portions of the ceratobranchials enter the floor of the mouth and are curved upward, with the result that the basihyal is raised to the level of the roof of the mouth and the dorsohyals touch the underside of the ethmoid. The elevation of the floor of the mouth appears to be accomplished by the upward rotation of the medial cartilaginous plates supporting the intermandibularis muscle (see p. 16). The inter- mandibularis thus forms a hillock against the underside of the ethmoid and the thin upper lip tightly embraces the lower jaw when the mouth closes. The pressure created by the floor rising against the roof of the mouth serves to force water through the mesh-like gill-rakers which filter out particulate material. The length of the gill openings presumably allows a high volume of water to pass rapidly over the gill-arches. The skin covering the dentary contains 7-8 transverse tubes (mst, Fig. 14) filled with a jelly-like substance staining intensely with the mucus-specific stain alcian blue. Of the taxa examined Iheringichthys has a similar jelly-like material in the lower lip, but it is concentrated in ampullae in the tissue close to the jaw symphysis and not arranged in widely spaced rows. Gelinek (1978) has described ampullary organs in the epidermis of Sorubim. He found them to be densely grouped in the labial tissue of the upper jaw and noted their structural similarity to those organs in Plotosus. Gelinek observed the similarities between these organs to those of gymnotids and mormyrids, thus hypothesising a similar function, namely electroreception. The snout and jaws in other siluroids The presumed plesiomorph condition of the ethmoid in siluroids is that which occurs in 20 G. J. HOWES mx ethc Fig. 15 Upper, Iheringichthys labrosus, ventral view of mandibular barbel supporting elements. Specimen BMNH 1934.8.20: 70-72. Scale = 2 mm. Lower, Luciopimelodus pati, dorsal view of ethmoid region. Specimen BMNH 1 878.5. 1 6: 30 (dry skeleton). Scale = 3 mm. d = dentary. many taxa belonging to the Diplomystidae, Trichomycteridae, Bagridae, Ictaluridae, Pimelodidae, Mochokidae, Plotosidae and Loricariidae, here the ethmoid is elongate and T- shaped and often with a median anterior notch (see Fink & Fink, 1981). Howes (1980) mistakenly considered this type of ethmoid in Mochokidae as apomorphic. In its presumed plesiomorph condition, the siluroid lateral ethmoid is triangular, closely united with the parasphenoid ventromedially and with the orbitosphenoid posteriorly. The expanded and poorly ossified ethmoid region of Hypophthalmus is considered to be a derived condition. Other derived states of the ethmoid are those where the bone is flattened with divergent anterior forks (Ageniosidae) is curved ventrally (some Ariidae) or is cavitous with a medially grooved surface (some Auchenipteridae). Another derived form occurs in one group of Pime\oididae,Piramutana, Hemisorubim, Luciopimelodus, Brachyplatystoma, Pseudoplatystoma and Sorubimichthys. Here the mesethmoid is greatly depressed and CATFISH ANATOMY & PHYLOGENY na le 21 sbc Fig. 16 Hypophthalmus edentatus, dorsal view of cranium. Specimen BMNH 1972.7.27: 675-678 (alizarin). Scale = 10 mm. expanded laterally (Fig. 1 5). As in Hypophthalmus the lateral portion of the ethmoid forms a large cavity wherein lies the olfactory organ (etch, Fig. 1 5). Derived conditions of the lateral ethmoid are those where the bone has a posterior process contacting an anterior process of the frontal (some Ariidae and Pangasiidae) and where the bone is extensive, its antero-lateral portion lamellate, extensively cancellous and vacuolate. This latter condition occurs in Pangasiidae, Amphiliidae, Auchenipteridae and Pimelodidae, but among the latter family only one taxon, Parapimelodus, approaches Hypophthalmus in the width and shape of its mouth. The mandibular barbels of pimelodids are particularly well-developed and in some taxa, as in Hypophthalmus, they articulate with large cartilaginous plates, e.g. Sorubim, Pseudoplatystoma, Iheringichthys, Pinirampus, Pimelodus (part), Luciopimelodus. The two former genera differ from Hypophthalmus, however, in having the barbel origin remote from the jaw symphysis. Ghiot (1978) described the cartilaginous plates in Pimelodus 'clarias' and commented on the elasticity of the plates and their function of supporting and moving the barbels. In Pimelodus blochii, Iheringichthys, Pinirampus and Luciopimelodus the barbels lie close to the symphysis, the presumed plesiomorph condition, but the cartilaginous supporting plates have a complex form resembling that in Hypophthalmus (Fig. 15). These taxa also share with Hypophthalmus the peculiar mandibular sesamoid cartilage which supports the lip fold, and the coronoid extension of the Meckelian cartilage (see p. 1 8 above). The cranium (Figs 16-18) There is little purpose in presenting a detailed description of the cranium, already described 22 G. J. HOWES by Wright (1885). It is necessary only to comment on those features pertinent to the present discussion. The neurocranium is moderately broad, transversly convex, becoming high-vaulted in the occipital region, and has a deep supraoccipital crest (Fig. 16). All the cranial bones are of a honeycomb texture with the frontal papyraceous. Nearly all sutures are synchondral, and fatty tissue pervades bone-muscle interspaces. The otic region The prootic is pierced by two foramina (pro, Fig. 17). The hyomandibular and palatine branches of the VII cranial nerve emerge through the posterior one (fhvii) whilst the oculo- motor, trigeminal, abducens and facial nerves emerge through the anterior foramen in the border of the bone (ftf). In almost all siluroids investigated there is but a single foramen piercing or indenting the prootic. In some ariid taxa the prootic is imperforate and the foramen for the cranial nerves is contained within the pterosphenoid. The absence of a second, posterior, opening in the prootic of most siluroids is due to the peculiarity of the cranial blood vascular system in these fishes. In the majority of teleosts the posterior prootic foramen serves for the passage of the jugular vein (lateral head vein) as well as the hyomandibularis nerve trunk. In siluroids, the jugular lies lateral to the cranial wall and is joined by its various branches in the same aperture as that through which the trigeminal nerve complex passes. The absence of a complete trigemino-facialis chamber in siluroids has been commented upon by several authors as the following quotations indicate: Allis (1908:259) referring to Ictalurus ( = Amiurus) notes that, 'The artery (external carotid) does not apparently traverse a trigemino-facialis chamber, for although it would seem as if this chamber must be present in some form, there is no proper indication of but one cranial wall in this region, and that one wall would seem to be the inner wall of the chamber; for both the external carotid and the jugular vein lie external to it.' De Beer (1937:137) again in referring to Ictalurus comments, 'There is no lateral commissure or pila antotica. The side wall of the skull in the orbital region therefore presents a continuous fontanelle the sphenoid fontanelle between the hind edge of the preoptic root of the orbital cartilage and the anterior basicapsular commissure. . .' Bamford (1948:365, 367) referring to Galeichthys, 'The normal foramina for the exit of the various parts of V and VII are absent, and the exit of these is limited to the sphenoid fissures.' . . .'There is no trace of a pila lateralis or any similar structure.' Alexander (1965:95) finds that in siluroids, 'The chondrocranium lacks the lateral commissure. The trigemino-facialis chamber of the adult skull has consequently lost its walls.' All these authors are agreed that the siluroid condition is one where the trigemino-facialis chamber has been modified and the prefacial commissure suppressed. Bamford (1948) suggested that cartilaginous growth from the anterior edge of the otic capsule had crowded the nerves and vessels into the same foramen as the optic nerve (see below concerning the passage of the optic nerve). The absence or reduction of an internal carotid artery in certain siluroids has also been the subject of comment by Allis (1908), Goodrich (1930), De Beer (1937) and Bamford (1948). According to Bamford (1948) the internal carotid has all but disappeared, with a consequent modification of the arterial circuitry. As yet, too few siluroids have been examined to ascertain the extent of this specialization. The optic foramen and the suprasphenoid In Hypophthalmus passage of the optic nerve (II) is through a separate foramen between the pterosphenoid (pts) and the 'suprasphenoid' (see below). In many siluroids there is no separate optic foramen, the optic nerve sharing the same opening as the trigemino-facialis CATFISH ANATOMY & PHYLOGENY 23 obs fhvii pro ss pts fii Fig. 17 Neurocranium, otic region of: (upper) Hypophthalmus edentatus ventro-lateral view. Specimen BMNH 1972.7.27: 675 (alizarin). Scale = 3mm. (centre) Auchenipterus nigripinnis lateral view. Specimen BMNH 1910.5.26: 12 (skeleton). Scale = 3mm. (bottom) Pterodoras granulosus lateral view. Uncatalogued skeleton. Scale = 10 mm. ss = suprasphenoid. complex, e.g. Diplomystidae, Callichthyidae, Siluridae, Bagridae (part), Clariidae. It is more usual in siluroids for there to be a separate foramen for the optic nerve between the pterosphenoid and parasphenoid or the suprasphenoid. Wright (1885) referred to a paired element in Hypophthalmus as a basisphenoid. The bone in question is long and shallow and contacts the pterosphenoid, orbitosphenoid and prootic (ss, Fig. 17). Between it and the pterosphenoid is a slit-like foramen through which passes the optic nerve (fii, Fig. 17). In many taxa I have examined a similar element is present, and in the Auchenipteridae and Doradidae it is pierced by the optic foramen (Fig. 17), a feature recognised as synapomorphic. 24 v6 sbc v5 v4 Fig. 18 Hypophthalmus edentatus neurocranium, occipital region (upper) in lateral, and (lower) in ventral views. Composite drawing from alizarin and dry preparations, tsc = 'transcapular'. McMurrich (1884#) and Kindred (1919) have described the so-called basisphenoid in Ictalurus ( = Amiurus). Kindred (1919: 39-40) discussed the homology of this element and decided that the term 'suprasphenoid' rather than basisphenoid would be better applied to it. He defined the suprasphenoid as a '. . .connective tissue ossification above the parasphenoid and between the ventral ends of the alisphenoids ( = pterosphenoids).' Daget (1964) also advocates caution in applying the term 'basisphenoid' to this element in siluroids. Thus, Kindred's term, suprasphenoid is adopted here as one being free of any implied homology. The homology of the siluroid parasphenoid itself is in doubt. Allis (1919) considered the problem in Ictalurus, (=Amiurus) and argued that the siluroid parasphenoid was an ossification of the ventral myodome roof, the posterior myodome being absent in siluroids. Bamford (1948:375) remarks that in Ictalurus, '. . .there are so many exceptional features in this region of the skull . . . myodome basicranial relations, absence of trigemino-facialis chamber, blood systems, pilae etc that the matter cannot be regarded as settled.' Until more is known about the occurrence of the suprasphenoid amongst siluroids, and until more definite information on its ontogeny and relationships with other elements is available, the value of this bone as a taxonomic character cannot be determined. The occipital region There are several outstanding features in this region of the cranium in Hypophthalmus, not least of which is the ankylosis of the skull and the vertebral column (Fig. 18). CATFISH ANATOMY & PHYLOGENY 25 The supraoccipital (so) crest is high and posteriorly is firmly united with the succeeding neural spine (4th). Anteriorly the supraoccipital meets the frontal (f), and laterally it is bounded by the epioccipitals. Wright's (1885) description and figure give an incorrect shape to the supraoccipital, and those elements bounding it are misidentified. Posteriorly the epioccipital (epo) is separated from the parapophysis of the 4th vertebrae by an elliptical foramen through which passes the ramus lateralis (Fig. 16). This foramen is shown in Wright's (1885) figures (fe, figs 1 & 3), but in the wrong position, as lying between the pterotic and posttemporal. The 'posttemporaf is an extensive bone (ptt, Fig. 18), its anterior part meeting the pterotic (pte) and exoccipital (exo) and its medial margin bordering the parapophyses of the 4th, 5th and 6th vertebrae. In no other siluroid does the posttemporal extend so far posteriorly, nor articulate with the 6th vertebra. The ventral limb of the 'posttemporal' is broad and thick, and articulates with the basioccipital. A thick ligament runs between its upper medial border and the 4th centrum (Ipt, Fig. 18). There has been some considerable debate concerning the identity of the element connecting the cleithrum with the cranium in siluroids. The most recent discussions are by Lundberg (1975) and are reconsidered by Fink & Fink (1981). The latter conclude that the siluroid element is a composite formed from the supracleithrum, possibly the posttemporal and Baudelot's ligament. That part of the element contacting the basioccipital and identified by Fink & Fink (1981) as the ossified Baudelot's ligament was referred to by Kindred (1919) as the transcapular. I have little to add to this discussion except to confirm Lundberg's (1975) observation that the small plate-like bone (previously thought to be an extrascapular, but identified by Lundberg as the posttemporal) does not occur in the Hypophthalmidae. However, such an element is present in Luciopimelodus which I have labelled here as an extrascapular so as to conform with Fink & Fink's nomenclature (es, Fig. 20). Notwithstanding Lundberg's evidence (1975) from the sensory canal pattern that the posttemporal is lacking, it would seem that the supracleithrum would be the element of the clavicular series most 'easily lost' in phylogeny; see for example Rosen (1964) where the supracleithrum is lacking in certain atheriniforms. The complex vertebrae, swimbladder and posterior lateral line nerve The complex vertebrae and swimbladder (Figs 1 9-2 1 ) Both Wright (1 885) and Chardon (1968) have described the peculiar nature of the Weberian apparatus in Hypophthalmus. The most notable character is the complex encapsulation of the swimbladder formed of the parapophyses of the 4th centrum. The parapophyses are produced into dorsal and ventral laminae, the ventral lamina formed partly from superficial ossification; see below p. 28. The cavity so formed provides a capsule housing the swim- bladder (sbc, Fig. 18). Internally the swimbladder (sb, Fig. 19) is divided by a longitudinal septum. Externally it has a thick tunica externa but no tunica interna. The opening between the upper and lower parts of the parapophysis together with the lateral extension of the 'posttemporal' form a funnel-like aperture to the exposed outer wall of the swimbladder (Fig. 19). No transverse duct connects the swimbladder chambers. The tripus (tr) is connected to the tunica externa just anterior to the septum. A medial extension of the tripus articulates with the complex vertebrae (3rd centrum). Anteriorly the tripus contacts the scaphium (sc) via a minute intercalarium (ic). The claustrum is lacking. The scaphia provide the walls of the cavum sinus imparis. According to Wright (1885: 1 14), the roof of the cavum sinus imparis is an ossified plate of dura mater attached to the first centrum, but in the preparations examined it seems as if the roof is just a continuation of the exoccipital. The 1st centrum is so far forward that it lies between the exoccipitals and above the basioccipital (vl +2, Fig. 19). 26 G. J. HOWES Wright (1885) maintained that the Weberian apparatus in Hypophthalmus has been '. . .pushed (in the process of its reduction) into the foramen magnum, instead of being outside the skull . . .'. Although that author considered the Weberian apparatus and swimbladder to be functionless (a theory endorsed by Bamford, 1 948) this does not appear to be so. The swimbladder is constructed as in other siluroids and there is a mobile articulation of the Weberian ossicles, all of which suggests that the system is functional (see below). The complex vertebrae and swimbladder in other siluroids In siluroids, as in other otophysans, the anterior four vertebrae are modified to form the Weberian apparatus. However, unlike other otophysans, there is a greater degree of fusion between the centra involved, and the united elements are usually referred to as the complex vertebrae (Chardon, 1968). In addition to this fusion of the first four centra, sometimes the 5th, and more rarely the 6th, 7th and even 8th are also incorporated in this complex. The lateral processes of the various centra incorporated in the complex vertebrae are always well-developed, often forming sheets of bone which may contact one another to provide a dorsal shield to the swimbladder. The anterior ramus of the 4th vertebral para- pophysis (following Tavolga, 1962, referred to hereafter as the Miillerian ramus) may contact the inferior limb of the 'posttemporal' and, in some taxa where the swimbladder is encapsuled, it may form a substantial part of the capsule. Alternatively, the Miillerian ramus may remain free from the 'posttemporal' and curve ventrally to contact the tunica externa of the swimbladder. In some families (Doradidae, Auchenipteridae, Mochokidae, Malapteruridae, Pangasiidae, Ariidae) the Miillerian ramus is free from any attachment with the 'posttemporal', its distal tip is expanded into a plate which contacts the tunica externa of the swimbladder. Protractor muscles run from the bony expansion to the cranium and neural spine complex supporting the dorsal fin. Miiller (1842) named this mechanism the Spring- federapparat, now usually referred to as the elastic spring apparatus, and below as the ESA. The plesiomorph condition of the swimbladder in siluroids is assumed to be one in which v5 v6 Fig. 19 Hypophthalmus edentatus swimbladder and Weberian apparatus. Dorso-lateral view of transverse section through the anterior vertebrae and occipital region. Composite drawing from alizarin and dry preparations. fs2 = foramen of 2nd spinal nerve; ssc = semicircular canal. CATFISH ANATOMY & PHYLOGENY 27 V5 fx v6 Fig. 20 Luciopimelodus pati complex vertebrae in (left) dorsal, (right) ventral, and (bottom) lateral views. Specimen BMNH 1878.5.16: 30, dry skeleton. Scale = 5 mm. the organ is large and free from any intimate contact with the anterior vertebral parapophyses, the lumen partially divided by a T-shaped septum, and the pneumatic duct entering the anterior transverse chamber. In taxa with this type of swimbladder, the anterior vertebrae (i.e. those following the complex vertebrae) demonstrate a range of fusion patterns, from being completely unfused, Diplomystidae, to the fusion of centra 4-6 in some Pimelodidae. In those taxa with an ESA, the swimbladder is also large and not encapsuled, but it is notable that in all ESA taxa studied the complex centra display the maximal vertebral consolidation (4-6 + 4-7 in some ariids). In those taxa with encapsuled swim- bladders, a series of progressive anterior vertebral fusions is apparent. However, only three families have the maximal fusion of 6 centra, Loricariidae, Astroblepidae and Ageneiosidae. If progressive fusions, namely the 4th centra with the 5th, then fusion with 6th and 7th vertebrae, are taken as successively derived conditions the resulting cladogram indicates that swimbladder encapsulation has occurred independently in at least two lineages (Fig. 22). However, a more rigorous study of the individual taxa within those lineages shows incon- gruencies in this hypothesised phylogeny. For example, the nature of swimbladder encapsulation in the Callichthyidae, Astroblepidae and Loricariidae is virtually identical, the 'posttemporal' contributing to the lateral wall of the capsule (Alexander, 1964; 1965; Chardon, 1968). These taxa also share a derived rib structure and dermal ossification patterns (see Alexander, 1965). On the basis of vertebral fusion patterns alone, however, these families appear in different lineages. Likewise, the Ageneiosidae also appears with the Loricariidae and Astroblepidae, but the encapsulation of the swimbladder is quite different since it is derived from superficial ossification (see below). 28 G. J. HOWES Further incongruencies arise in this phylogeny when the detailed structure of the ESA and the associated swimbladder are taken into account. In the Doradidae and Auchenipteridae the organisation of this system is more elaborate than that of other ESA taxa. In both families the epioccipital has long posterior processes which contact the 5th and 6th parapophyses. Such features are lacking in other ESA groups. From these observations it seems very likely that the ESA has, as in the case of swimbladder encapsulation, been achieved independently in several lineages. Another incongruency arises if the superficial ventral ossification of the fused vertebral centra is taken into account. Superficial ossifications were described by Bridge & Haddon (1894) as a continuous sheet of bone uniting the paired parapophyses and enclosing the cardinal veins. According to Tilak (1965: 170-1 71) the superficial ossification is derived from the tunica externa of the swimbladder. There is much variability in development of superficial ossification. In some taxa it forms a channel or a complete tunnel enclosing the dorsal aorta. In others it contributes to the anterior part of the swimbladder capsule. There appears to be a trend for the more derived siluroids to have reduced and encapsuled swimbladders. The most extreme forms of encapsulation and swimbladder reduction are usually encountered among those taxa with accessory respiratory devices, Clariidae, Loricariidae and Callichthyidae. In this respect the Hypophthalmidae and Ageneiosidae are exceptional. Alexander (1964) makes the point that encapsulation of the swimbladder results '. . .when the anterior parapophyses have not been correspondingly reduced'. Undoubtedly this is so, the anterior (Miillerian) ramus of the 4th parapophysis progressively curving around the swimbladder and meeting the lamina of the 5th appears to be an ontogenetically obvious process of encapsulation. The swimbladder capsule may also incorporate part of the ventral superficial ossification (see above), in Loricariidae and Callichthyidae the posterior cranial bones (epioccipital and 'posttemporal') also contribute to its formation. Alexander (1964) advanced the hypothesis that a reduced swimbladder volume could have evolved to compensate for increased buoyancy resulting from gas held in the accessory respiratory organ. Qasin & Hasan (1961) suggested that such gas could even be used in buoyancy control. Gee (1976) does not agree, pointing out the unlikelihood of a simultaneous reduction of swimbladder volume (SBV) and development of an accessory air breathing organ (ARO) derived from some other part of the fish's anatomy. Gee also noted that loricariids although possessing the smallest SBV lack an ARO. Gee's thesis is that the overiding factor in swimbladder reduction is directed towards a demersal existence wherein a greater variety of environments may be exploited, e.g. substrate burrowing by tricho- mycterids; leaf litter and bank 'burrowing' by loricariids. Some of these environments, however, tend to be hypoxic and would have demanded compensatory air breathing devices. Reduction of SBV in Hypophthalmidae and Ageneiosidae is obviously due to different factors than those obtaining in loricariids etc, and ones related to attaining neutral, or near neutral buoyancy. Unlike Old World 'pelagic' siluroids, schilbeids, siluriids and pangasiids, whose neutral buoyancy depends on a large swimbladder, the Hypophthalmidae, and probably the Ageneiosidae, rely on a high body fat content (see p. 5). More positive asser- tions on the functional utility of SBV reduction can only be made in the light of detailed knowledge of swimbladder physics. Alexander (1965) states there is no known fish in which the Weberian apparatus is a functionless vestige. However, in loricariids, callichthyids and trichomycterids the ossicles are reduced, the claustrum and intercalarium lacking (see Chardon, 1968). To what degree the absence of these elements impairs or alters the function of the Weberian system is unknown. Bamford (1948) noted that little work had been done on the ontogeny of the siluroid Weberian apparatus as compared with that in cyprinids. There has been no redress of this situation. Bamford's implication was that the entire Weberian system could have been derived independently in siluroids. This hypothesis has still to be tested. CATFISH ANATOMY & PHYLOGENY 29 nviip nviip nviia cw Fig. 21 Pathway of vagus and posterior branch of the facial nerves in gymnotoids (upper), and siluroids (lower). Dorsal view, semi-diagrammatic drawn from dissections of Eigenmannia virescensand Pimelodus blochii. cw = cranial wall; op = operculum. The posterior lateral line nerve The anterior shift and gross re-shaping of the Miillerian ramus in Hypophthalmus has necessitated a diversion in the route of the vagal ramus lateralis, posterior lateral line nerve. As a result, after leaving the exoccipital (fX. Fig. 18) it takes an almost vertical course then passes through the foramen between the Miillerian ramus and epioccipital (see above, p. 25), finally, running posteriorly across the dorsal surface of the parapophysis. Nelson (1960) described the course of the vagus lateralis in Malapterurus as leaving the cranium and running laterally along the ossified Baudelot's ligament (inferior posttemporal limb and transcapular of authors) thence passing caudally along the length of the body; the ramus lateralis vagi of authors. Such is the usual course of the nerve in other siluroids investigated. Nelson (1960) also describes a dorsal or recurrent branch of the VII cranial nerve which, after leaving the cranium innervates the dorsal length of the body. This nerve appears to correspond with that described by Mithel (1964) as the ramus lateralis accessorius in Mystus and by Juge ( 1 899) as the 'nerf de Weber' in Silurus. The more direct course of the vagus lateralis in Hypophthalmus also occurs in Luciopimelodus (Fig. 20). Here too the posterior tip of the 'posttemporal' is elongated to extend as far as the 5th parapophysis, and the Miillerian ramus is elevated and strongly curved antero-ventrally to partially encapsulate the swimbladder. These features cause the nerve to follow a vertical route from its exit in the exoccipital and, as in Hypophthalmus to pass through a gap between the supraoccipital, epioccipital and Miillerian ramus (Fig. 20). Nelson (1960) noted that in gymnotoids the '. . .nerve (vagus) emerges from the postero- lateral aspects of the cranium associated with the opercular muscles and passes superficial to the pectoral girdle.'; after which it penetrates the body wall musculature and follows the horizontal myoseptum. Szabo (1974) following Nelson (1960) considers the dorsal nerve in siluroids '. . .to be of the type', presumably homologous, as the lateral nerve trunk in gymnotoids. Fink & Fink (1981) accept such a homology as a character in uniting siluroids and gymnotoids as sister groups. The very different pathways of the nerve trunk in these two groups suggests however the possibility of independent derivation. In siluroids the vagus lateralis passes laterally across the transcapular (tsc) (Baudelot's ligament fide Fink & Fink), over the Miillerian ramus and then ventrally, medial to the pectoral girdle. The posterior branch of the anterior lateral nerve (ramus recurvens nervi facialis; ramus dorsalis or ramus lateralis accessorium auctf) emerges through the supraoccipital to course along the dorsal aspect of the body (Fig. 2 1). 30 Diplomystidae Bagridae Chacidae Schilbfeidae Siluridae Ictaluridae Plotosidae Pimelodont.(pt) Cranoglanidae G. J. HOWES Clariidae Amphiliidae Sisoridae Callichthyidae Trichomycteridae Cetopsidae Aspredinidae Pimelodont.(pt) Hypophthalmidae Helogeneidae Ageniosidae Astroblepidae Loricariidae Sb En Ariidae(4748) Auchenipteridae Doradidae Mochokidae Malapteruridae Pangasiidae Sb .44546 ESA U Fig. 22 Cladogram of currently recognised siluroid families based on degree of fusion of the anterior vertebrae and nature of the swimbladder. UFU = vertebrae unfused; FU = vertebrae fused; 44-5+ = numbers of centra added to the complex vertebrae; SbF = swimbladder free; Sb En = swimbladder encapsuled; ESA = elastic spring apparatus. For the purposes of the diagram the Clariidae includes the Heteropneustidae. The Pimelodontidae is represented on two lineages, those taxa with an encapsuled swimbladder correspond with group 1 as denned on p. 37. Ariid taxa examined appear to have the 7th (? and 8th) centra added to the complex vertebrae. It is not certain that all ariid taxa have this many consolidated centra. Data mostly from Chardon (1968) and pers. obs. In gymnotoids the vagus lateralis runs postero-ventrally into the epaxialis and does not pass over the 4th parapophysis. The posterior branch of the anterior lateral nerve does not pass dorsally but laterally, to pierce the hyomandibula, then to run medial to the pectoral girdle, and into the body musculature where it joins the vagus lateralis (Fig. 21). The suspensorium (Figs 23 & 24) In Hypophthalmus a large pterygoid bone extends between the quadrate and hyomandibula to form a bipartite attachment with the lateral ethmoid. This articulation takes the form of a long triangular portion and an upright process (pty, Fig. 23). Between the two anterior forks a ligament runs anteriorly to attach to a minute bony element and thence to the lateral CATFISH ANATOMY & PHYLOGENY 31 syc op ih Fig. 23 Hvpophthalmus edentatus suspensorial and opercular bones, medial view (cartilage dotted). Specimen BMNH 1972.7.27: 675-678. Scale = 5 mm. ethmoid. The pterygoid (pty, Fig. 23) contacts the quadrate synchondrally, and the hyomandibular process via a dentate suture. The hyomandibular process (hy) extends anteriorly along the dorsal rim of the quadrate (q) which it meets synchondrally. The stem shank of the hyomandibula contacts the central dorsal rim of the quadrate via a synchondrosis. Posteriorly, it is separated by a wedge of cartilage (syc), which may represent the symplectic. The interhyal (ih) articulates between this cartilaginous element, the posterodorsal edge of the quadrate, and the upper rim of the interoperculum. The identity of the various bones comprising the suspensorium in siluroids is doubtful. Regan (1911) simply referred in pimelodids to 'pterygoids' lying between the hyomandibula and palatine. Compared with most other teleosts the pterygoid series is, for the most part, incomplete since it lacks one of the pterygoid elements usually present. Authors have been at variance as to whether it is the ecto- or endopterygoid which is absent (see comments of Kindred, 1919; Alexander, 1965 and Gosline, 1975). The problems of identifying siluroid suspensorial elements is illustrated by reference to the pimelodid Pinirampus (Fig. 24). The pterygoid is like that of Hypophthalmus in having a sharply demarcated ventral portion that articulates with the quadrate, whilst the dorsal portion articulates with the hyomandibular process. An area of cartilage lies between the branches of the pterygoid, the hyomandibular process and the quadrate. Lying close to the anterior border of the pterygoid is a scythe-shaped bone, its elongate portion directed ventrally. The bone is connected to the pterygoid and to the lateral ethmoid via ligaments. Between the scythe-like bone and the pterygoid border is a splint-like bone which is attached by connective tissue to both the scythe-like element and the palatine. Two identification schemes of the suspensorial elements are presented. In one (Fig. 24, left) the pterygoid is considered as the metapterygoid, the scythe-shaped bone as the endopterygoid and the splint-like bone as the ectopterygoid. In the other (Fig. 24, right) the hyomandibula plus metapterygoid are considered as forming a single element, or, alternatively the metapterygoid is considered lost, and the pterygoid as comprising the endo- plus, ectopterygoid. The two isolated bones are problematical. From its position in connective tissue the splinter-like element could be a sesamoid ossification. The scythe- shaped bone is characteristic of one group of pimelodids (see p. 37) and again, may be a 32 G. J. HOWES hy+mp ect Fig. 24 Pinirampus pirinampu suspensorial and opercular (part) bones. Specimen BMNH 1934.8.20: 104-5. Scale = 3 mm. The upper, labelled drawings represent two interpretations of the suspensorial elements. Shaded areas represent cartilage. sesamoid bone. A small bone occurs in Hypophthalmus in a similar position and having the same ligamentous attachments as the scythe-shaped bone in Pinirampus (see above). It seems unlikely that these 'isolated' elements represent an ectopterygoid as they occur in the wrong topographical positions. The problem of suspensorial bone homologies in siluroids will be dealt with elsewhere (Banister & Howes, in preparation). The hyoid arch (Fig. 25) In Hypophthalmus the anterohyal and posterohyal are exceptionally elongate and, unusually, are of equal length. The anterohyal (ahy) bears 8 branchiostegal rays (bsr) and the posterohyal (phy) 6. The dorsal and ventral hyals (dh, vh) are small and separated from the anterohyal by cartilage. The interhyal (ih) articulates with the posterior tip of the posterohyal. The urohyal(uh) is a short oval bone lacking a medial ridge. Gosline (1974) relates the number of branchiostegal rays to the length of the free sleeve of the branchiostegal membrane. This certainly appears to be a valid correlation in Hypophthalmus where the gill opening extends very far forward, and may, to a certain extent, be applied to some pimelodid taxa as well. Examples are Sorubim and Pseudoplatystoma which also possess long gill openings and a high number of branchiostegal rays (11-1 6). CATFISH ANATOMY & PHYLOGENY vh 33 ih Fig. 25 Hypophthalmus edentatus hyoid arch, lateral view. Specimen BMNH 1972.7.27: 675-678. Scale = 5 mm. According to McAllister (1968) a high number of branchiostegal rays is the primitive condition in teleosts. On this basis he considers that amongst siluroids the Old World taxa '. . .appear to be more primitive than the South American catfishes, having 4-20 branchiostegals as opposed to 3-17. . .'. The extent of these ranges and the degree of overlap hardly justify such a statement. From McAllister's counts for siluroid families, the pre- dominant number of branchiostegal rays would seem to be 9-1 1 and perhaps this, by virtue of its widespread occurrence, should be thought of as the plesiomorph number rather than the high number advocated by McAllister. Further comparative data, particularly on the morphology and the placement of the branchiostegal rays on the hyoid bar are necessary before any valid statement can be made concerning the apo- or plesiomorphy of their number. The relationships of the interhyal to the preoperculum shows some interesting modifi- cations in the Pimelodidae. The usual siluroid condition, as in other ostariophysans, is for the interhyal to articulate between the quadrate, or symplectic, and the hyomandibular limb. In most siluroids the interhyal attaches, in the absence of a symplectic, ligamentously to both quadrate and hyomandibula, abutting against the intervening cartilage (see description of Hypophthalmus above). In Pseudoplatystoma the interhyal has shifted posteriorly to such an extent that its connection with the hyomandibula and quadrate is lost, 34 G. J. HOWES Fig. 26 Hypophthalmus edentatus upper and lower (part) branchial arches, dorsal view, left side (cartilage stippled). Specimen BMNH 1972.7.27: 675-678. Scale = 3 mm. and instead it articulates entirely with a preopercular fossa. Intermediate stages in the development of this condition are to be found in Goldiella, Sorubim and Brachyplatystoma where the interhyal has shifted even further posteriorly and is ligamentously attached to the rim of a preopercular fossa alongside the hyomandibula. The opercular series (Fig. 23) The preoperculum (po) of Hypophthalmus is a thin, narrow bone covering the posterior margins of the hyomandibula and the quadrate, virtually obscuring the interoperculum. The interoperculum (iop) is roughly triangular but with a concave ventral border where it overlaps the pectoral fin base. The operculum (op) is almost equilaterally triangular, its posterior edge much fretted, and its entire structure cancellous. The opercular bones of Hypophthalmus appear to be of a 'generalised' siluroid type, i.e. a large operculum, almost vertical preoperculum and small interoperculum. There is considerable variability in size and shape of these elements throughout the Siluroidei (Howes, in press). Fink & Fink ( 1 98 1 ) maintain that the triangular shape of the operculum is synapomorphic for the siluroids and gymnotoids. However, whereas the ventro-posterior border of the operculum in siluroids is either straight or concave, that in gymnotoids is more often convex, as in other otophysans. The gill arches (Figs 26-27) Upper elements (Fig. 26). The epibranchials of Hypophthalmus edentatus are narrow- waisted, with expanded medial surfaces and bear 25-27 long, fine gill-rakers, As in all siluroids examined, epibranchial (eb) 3 has an uncinate process which in Hypophthalmus is rounded and overlies the central portion of eb4. Infrapharyngobranchial(if) 3 is well-ossified and elongate, its posterior margin separated from eb3 and eb4 by a cartilaginous wedge. Of the two alizarin preparations at my disposal, the larger shows two distinct chondrifications in CATFISH ANATOMY & PHYLOGENY uh 35 cm Fig. 27 Hypophthalmm edentatus lower branchial arch elements; (upper) dorsal view. Specimen BMNH 1972.7.27: 675-678. Scale = 3mm; (lower) cross section of left side gill-rakers and ceratobranchials. cm = ceratobranchial membrane. this wedge. These cartilages suspend a large oval toothplate bearing on its ventral surface many minute, conical teeth (tp, Fig. 26). Lower elements (Fig. 27). The basibranchials (bb) and hypobranchials (hb) are poorly ossified. Basibranchial 1 is a triangularly shaped cartilage lying in the plane of the antero- hyals; basibranchials 2 and 3 are rod-like ossified elements; basibranchial 4 is incorporated within a broad cartilaginous cornua, the ossified lateral margins of which represent hypo- branchials 4; basibranchial 5 is represented by a triangular cartilage whose apex is ossified. Hypobranchials 1-3 are ossified only along their leading edges. The ceratobranchials are the most elongate of any siluroid and carry at least 200 long gill-rakers. The first two ceratobranchials (cb) have only a single row of gill-rakers (gr), along the outer edge; the inner edge has a low wall of fibrous connective tissue (cm). The other ceratobranchials have a double row of gill-rakers; the 5th is dentigerous, its anterior stem is excessively elongate and meets its fellow along the midline. The gill-rakers on cb 5 occur only along the inner margin; anteriorly the rakers of each ceratobranchial intermesh. Roberts (1972) described the sieving mechanism of the gill-rakers. The arrangement of this mechanism is shown in Fig. 27 as a transverse section of the lower gill-arch. Such an arrange- ment does not occur in any other siluroid. 36 G. J. HOWES Postcranial elements As this study is concerned primarily with the cranial anatomy of Hypophthalmus, no detailed description of the postcranial anatomy is presented. The following features, however, are worthy of note. The pectoral girdle is of simple construction, with a greatly elongated horizontal cleithral limb and the coracoid reduced to a horizontal lamina. Such elongation of the cleithrum is unknown in any other siluroid. There are 62 vertebrae, excluding the complex vertebrae. This is an exceptionally high number amongst siluroids and is exceeded only by members of the Clariidae. Virtually no comparative data exist for vertebral numbers in siluroids. A superficial survey indicates that the mean siluroid count is 40-45. High counts are found among Ariidae 49-50, Ageneiosidae 49-50, Siluridae 50-51 and Clariidae 60 + . Among the Pimelodidae only Luciopimelodus has an exceptionally high count 47-48. The caudal fin skeleton of Hypophthalmus is of a plesiomorph pattern with no fusion of any hypural elements (PH;1;2;3;4;5). Lundberg & Baskin (1969) found hypural fusion and reduction of principal caudal fin rays to be widespread conditions. The most derived hypural fusion patterns occur in the Plotosidae, Chacidae and Loricariidae. In the latter family there is also an expansion of the last neural spine, which contacts the upper caudal element, i.e. the fused hypurals 3-5, uroneural and epural. Certainly the derived caudal fusion patterns shared by the Loricariidae, Callichthyidae and Astroblepidae are congruent with synapomorphies involving the encapsulation of the swimbladder (see p. 27). Other caudal fin patterns are not so easy to interpret and characters such as the absence of an hypurapophysis in the Clariidae and Plotosidae are interpreted as parallelisms by Lundberg & Baskin. Although both clariids and plotosiids trend toward anguilliform morphology, it is interesting to note that the hypural fusion pattern in clariids (excluding the Heteropneustidae) is, in contrast to plotosiids, plesiomorph. Even in such a seemingly derived taxon as Allabenchelys all the hypurals are free. Amongst the Pimelodidae, the predominant caudal fusion pattern is a free parhypural; 1 +2; 3 + 4; 5, which as Lundberg & Baskin (1969: 40) remark is significantly more advanced than that of any bagrid. The hypural fusion patterns yield little support for the proposed groupings of pimelodids presented below, except that Rhamdia and Pimelodella share a similar fusion of hypurals, 1 +2 and 3 + 4. Relationships of the Hypophthalmidae Anatomically, the Hypophthalmidae is a unique group. No other siluroid has such an arrangement of the eye musculature, hypertrophy of the jaw adductor muscles, reduction of ethmoidal ossification, complexly developed mandibular barbel supporting elements, elongated gill- and hyoid arch elements, and profoundly modified vertebral and swimbladder elements. Attention was drawn in the Introduction to the poverty of published data on siluroid comparative anatomy. In order to apply cladistic methods in resolving interrelationships extensive comparisons must be made thus, at this stage one cannot, with any degree of certainty identify plesiomorph or apomorph characters amongst siluroids. However, from this survey it is clear that the sister-group to the Hypophthalmidae lies within the Pimelodidae. Synapomorphies are manifest in the structure of the ethmoid region, the mandibular barbel articulation, the eye muscle arrangement, the complex vertebrae, and the sensory canal system. The following is a synopsis of these characters: 1 . Ethmoid region with lateral cavity (p. 20) shared with Luciopimelodus, Pinirampus and Pimelodus (part). 2. Articulation of the mandibular barbels with large cartilaginous plates (p. 21) shared with Luciopimelodus, Iheringichthys, Sorubim, Pinirampus, Pseudoplatystoma, Pimelodus (part). CATFISH ANATOMY & PHYLOGENY 37 3. Anterior eye musculature of tendinous origin shared with Sorubimichthys 4. Network of epidermal canals crossing the orbital and opercular regions shared with Luciopimelodus and Iheringichthys. 5. Posterior extension of the 'posttemporal' passing beyond the edge of the Miillerian ramus, elevation of that ramus, and vertical path of the anterior portion of the vaugus lateralis nerve shared with Luciopimelodus. In the course of this study only a limited number of pimelodid taxa was studied, but even this preliminary investigation reveals that there are at least three major groups contained within the Pimelodidae as presently recognised, viz: 1. Those species with the swimbladder partially encapsuled and usually with a caecal posterior border; the 5th and 6th fused within the vertebral complex; the ventral superficial ossification of the 4th and 5th centra forming a tunnel for the dorsal aorta; absence of a retractor tentaculi muscle. Included taxa: Iheringichthys, Pinirampus, Callophysus, Parapimelodus, Pimelodina, Perugia, Megalonema, Luciopimelodus, Pimelodus (part). In addition, the first four genera and Pimelodus maculatus all have a scythe-shaped pterygoid bone lying anterior to the main pterygoid series and attaching to the lateral ethmoid via a ligament (p. 31). 2. Those species with the swimbladder well-developed, not encapsuled, and with attached muscles (see below); 6 vertebrae forming the anterior complex centrum, the 4th and 5th centra elongate and forming an aortic tunnel. A retractor tentaculi muscle present. Included taxa: Pseudoplatystoma, Sorubim, Hemisorubim, Pimelodus (part). The distribution of swimbladder muscles throughout the Pimelodidae as a whole has still to be recorded. Bridge & Haddon (1894) described swimbladder compressor muscles in Pseudoplatystoma, (their Platystomd) as originating from the occiput and inserting on the antero-ventral wall of the swimbladder. These authors also described a tensor tripodis muscle in Pseudoplatystoma and some other pimelodontid taxa. This muscle runs from the exoccipital and inserts tendinously into the anterior wall of the swimbladder. Alexander (1965: 110) was unable to trace the tensor tripodis in Pimelodus blochii, Alexander's P. clarias. However, it appears in this species as a mere strip of tendinous fibres. The muscle is also present in Sorubim and Hemisorubim 3. Those species with the swimbladder well-developed, not encapsuled, and without compressor muscles; 5 fused centra, sometimes with superficial ossification but without an aortic tunnel. The lateral ethmoid is transversly convex. Included taxa: Rhamdia, Pimelodella, Heptapterus The above groups include taxa presently distributed amongst four subfamilies, the Pimelodinae, Luciopimelodinae, Callophysinae and Sorubiminae. None of the three groups defined here corresponds with any subfamily as presently recognised. In addition, there is another subfamily, the Pseudopimelodinae (Gomes, 1946). Representatives of this latter group have not been thoroughly investigated in the course of this study, but in those that have been examined the anterior part, at least, of the swimbladder is divided by an extension of the complex vertebrae. No meaningful phylogenetic classification is accomplished in recognising the above named pimelodid subfamilies, based as they are on a mixture of plesio- and apomorphies. Similarly, placing the Hypophthalmidae in a separate suborder (Chardon, 1968) simply to recognise its morphological distinctiveness, serves no purpose in elucidating its relationships. From the characters analysed above synapomorphies have been identified that indicate the sister group of the Hypophthalmidae to be in that assemblage which includes Luciopimelodus and Iheringichthys. A more refined definition of this group must await a thorough anatomical study of the Pimelodidae. 38 G. J. HOWES Acknowledgements This paper has benefited greatly from the critical and constructive comments of Drs Humphry Greenwood, Keith Banister and Paul Skelton, to all of whom I extend my sincere thanks. I particularly thank Bernice Brewster, Lynne Parenti, Chris Sanford and Rob Travers for so much helpful advice and technical assistance. I am deeply indebted to those many ichthyological colleagues and associates who, over the years, have provided specimens and given me so freely and patiently much information on catfishes. References Alexander, R. McN. 1964. The structure of the Weberian apparatus in the Siluri. Proc. zool. Soc. Lond. 142:419^40. 1965. Structure and function in catfish. J. Zool. Lond., 148: 88-152. Allis, E. P. 1908. The pseudobranchial and carotid arteries in Ameiurus. Anal. Anz. 33: 256-270. The myodome and trigemino-facialis chamber of fishes, and the corresponding cavities in higher vertebrates. J. Morph. 32: 207-322. Bamford, T. W. 1948. Cranial development of Galeichthys felis Proc. Zool. Soc. Lond. 118 (2): 364-391. Beer, G. R. de, 1937. The development of the vertebrate skull Oxford. 552pp. Bridge, T. W. & Haddon, A. C. 1894. Contributions to the anatomy of fishes II. The air-bladder and Weberian ossicles in the siluroid fishes. Phil. Trans. Roy. Soc. B, 184: 65-333. Carvalho, F. M. 1980a Composic.ao qufmica e reproduc.ao do mapara (Hypophthalmus edentatus Spix, 1 829) do lago do Castanho, Amazonas. Ada Amazonica 10 (2): 379-389. 19806 Alimentacao do mapara (Hypophthalmus edentatus Spix, 1829) do lago do Castanho, Amazonas (Siluriformes, Hypophthalmidae). Acta Amazonica 10 (3): 545-555. Chardon, M. 1968. Anatomic comparee de 1'appareil de Weber et des structures connexes chez les Siluriformes. Ann. Mus. roy. Afr. ser. 8 Zool. 169: 1-277. Cope, E. D. 1871. Observations on the systematic relations of the fishes. Proc. Amer. Ass. Adv. Sci. 20: 317-343. Daget, J. 1 964. Le crane des teleosteens. Mem. Mus. natn. Hist. nat. Paris (A) 31 : 1 63-34 1 . Dullemeijer, P. & Bare), C. D. N. 1977. Functional morphology and evolution. In: Major Patterns in Vertebrate Evolution Hecht, M. K., Goody, P. C. & Hecht, B. M. (Eds): 83-1 1 7. New York. Eaton, T. H. 1948. Form and function in the head of the channel catfish, Ictalurus lacustris punctatus. 7.A/0r/7/z.83:181-194. Edgeworth, F. H. 1935. The cranial muscles of the Vertebrates. Cambridge. Eigenmann, C. H. & Eigenmann, R. S. 1890. A revision of the South American nematognathi, or catfishes. Occ. Pap. Calif. Acad. Sci. 1: 1-508. Fink, S. V. & Fink, W. L. 1 98 1 . Interrelationships of the ostariophysan fishes (Teleostei). Zool. J. Linn. Soc. 72(4): 297-353. Gee, J. H. 1976. Buoyancy and aerial respiration: factors influencing the evolution of reduced swim- bladder volume of some Central American catfishes (Trichomycteridae, Callichthyidae, Loricariidae, Astrobelpidae). Can. J. Zool. 54 (7): 1030-1037. Gelinek, E. 1978. On the ampullary organs of the South- American paddle-fish Sorubim lima (Siluroidea, Pimelodidae). Cell Tiss. Res. 190: 357-369. Ghiot, F. 1978. The barbel movements of three South American pimelodid catfishes. Zool. Anz. 200 (5-6): 395-401. Gomes, A. L. 1946. A review ofMicroglanis, a genus of South American catfishes with notes on related genera. Occ. Pap. Mus. Zool. Univ. Mich. 494: 1-19. Goodrich, E. S. 1930. Studies on the structure and development of vertebrates. Reprint edit. New York (1958). Gosline, W. A. 1973. Considerations regarding the phylogeny of cypriniform fishes, with special reference to structures associated with feeding. Copeia 1973 (4): 761-776. 1975. The palatine-maxillary mechanism in catfishes, with comments on the evolution and zoo- geography of modern siluroids. Occ. Pap. Calif. Acad. Sci. (120): 1-3 1 . Goulding, M. 1 980. The Fishes and the Forest. Berkeley, Los Angeles and London. CATFISH ANATOMY & PHYLOGENY 39 Greenwood, P. H., Rosen, D. E., Weitzman, S. H. & Myers, G. S. 1966. Phyletic studies of teleostean fishes with a provisional classification of living forms. Bull. Am. Mus. nat. Hist. 131: 339-456. Gunther, A. 1 864. Catalogue of fishes in the British Museum 5. London. 455pp. Howes, G. J. 1980. A new catfish from Sierra Leone. Bull. Br. Mus. nat. Hist (Zool.) 38 (3): 165-170. in press. The cranial muscles of loricariid fishes, their homologies and use as phylogenetic characters. Bull. Br. Mus. nat. Hist. (Zool.) Juge, M. 1899. Recherches sur le nerfe cerebraux et la musculature cephalique de Silurus glanis. Rev. Suisse Zool. 6: 1-171. Kesteven, H. L. 1943. The evolution of the skull and cephalic muscles. The fishes. Pt. 1. Mem. Australian Mus. 8 (2): 63-1 32. Kindred, J. E. 1919. The skull of Amiurus. Illinois biol. Monogr. 5 (1): 1-120. Lubosch, W. 1938. Muskeln des Kopfes: Viscerale Muskulatur (Fortsetzung). B. Teleostier. In: Handbuch der vergleichenden Anatomie der Wirbeltiere. Bolk, L., Goppert, E., Kallius, E. & Lubosch, W. (Eds) 5: 1011-1024. Berlin & Vienna. Lundberg, J. G. 1975. Homologies of the upper shoulder girdle and temporal region bones in catfishes (order Siluriformes), with comments on the skull of the Helogeneidae. Copeia 1975 (1): 66-74. McAllister, D. E. 1968. The evolution of branchiostegals and associated opercular, gular and hyoid bones and the classification of teleostome fishes, living and fossil. Bull. natn. Mus. Can. (Biol.) 77 (221): 1-239. McMurrich, J. P. 1 8840. The osteology of Amiurus catus (L.) Gill. Proc. Can. Inst. 2: 270-3 10. 1 844/7 The myology of Amiurus catus (L.) Gill. Proc. Can. Inst. 2: 3 1 1-35 1 . Meschkat, A. 1960. Report to the Government of Brazil on the fisheries of Amazon region. BRA/TE/FI Rept. No. 1 305. FAO, Rome. Mithel, M. 1 964. The cranial nerves of Mystus seenghala (Sykes) Agra Univ. J. Res. 13: 67-78. Muller, J. 1842. Beobachtungen iiber die Schwimblase der Fische, mit Bezug auf einige neue Fischgattungen. Arch. Anal. Physiol.: 307-329. Nelson, E. M. 1960. The morphological relationships of the lateral-line nerve in certain 'Electric Fishes'. Copeia 1960 (2): 130-134. Qasin, S. Z. & Hasan, R. 1 96 1 . A hydrostatic function of the accessory respiratory organs in air breathing fishes. Nature 191, 396-7. Regan, C. T. 191 1. Classification of the teleostean fishes of the order Ostariophysi, Siluroidea. Ann. Mag. nat. Hist. 8, (8): 553-577. Roberts, T. R. 1972. Ecology of fishes in the Amazon and Congo basins. Bull. Mus. Comp. Zool. 143 (2): 117-147. 1973. Interrelationships of ostariophysans. In: Interrelationships of fishes. Greenwood, P. H., Miles, R. S. & Patterson, C. (Eds) 373-395 London & New York. Rosen, D. E. 1964. The relationships and taxonomic position of the halfbeaks, killifishes, silversides and their relatives. Bull. Am. Mus. nat. Hist. 127 (5): 2 19-267. Singh, B. R. 1 967. Movements of the barbels in some siluroid fishes. Zool. Anz. 178 (5-6): 402-4 1 2. Szabo, T. 1974. Anatomy of the specialized lateral line organs of electroreception. In: Electroreceptors and other specialized receptors in lower vertebrates. Fessard, A. (Ed.) 13-58. Berlin. Takahasi, N. 1925. On the homology of the cranial muscles of the cypriniform fishes. J. Morph. 40: 1-109. Tavolga, W. N. 1962. Mechanisms of sound production in the ariid catfishes Galeichthys and Bagre. Bull. Amer. Mus. nat. Hist. 124 (3): 5-30. Tilak, R. 1965. The comparative morphology of the osteocranium and Weberian apparatus of Tachysuridae (Pisces, Siluroidei)./. Zool. 146(2): 150-174. Vari, R. P. 1979. Anatomy, relationships and classification of the families Citharinidae and Distichodontidae (Pisces, Characoidea). Bull. Br. Mus. nat. Hist. (Zool.) 36(5): 26 1-344. Winterbottom, R. 1974. A descriptive synonymy of the striated muscles of the teleostei. Proc. Acad. nat. Sci. Philad. 125: 225-3 17. Wright, R. Ramsay 1885. On the skull and auditory organ of the siluroid Hypophthalmus. Trans Roy. Soc. Canada. Sect. IV: 107-1 1 7. Manuscript accepted for publication 18 October 1982 British Museum (Natural History) Associated publication on fishes The Cichlid fishes of Lake Victoria, East Africa. The biology and evolution of a species flock. P. H. Greenwood. Bull. British Museum (Natural History) Zool. Suppl. No. 6, 1974 vi + 1 34 pp, 1 coloured plate, 77 text figures, 4to, (paper) 6.00 (boards) 8. 2 5 Lists are available free on request to: Publications Sales British Museum (Natural History) Cromwell Road London SW7 5BD Standing orders placed by educational institutions earn a discount of 1 0% off our published price. Titles to be published in Volume 45 Problems in catfish anatomy and phylogeny exemplified by the Neotropica Hypophthalmidae (Teleostei: Siluroidei). By G. J. Howes Miscellanea Bats (Mammalia: Chiroptera) from Indo- Australia. By J. E. Hill. On Macropleurodus, Chilotilapia (Teleostei: Cichlidae) and the interrelationships of African cichlid species flocks. By P. H. Greenwood. Miscellanea The cranial muscles of loricarioid catfishes, their homologies and value as taxonomic characters (Teleostei: Siluroidei). By G. J. Howes. Miscellanea Printed by Henry Ling Ltd, Dorchester Bulletin of the British Museum (Natural History) Miscellanea Zoology series Vol 45 No 2 28 July 1983 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in for scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both the scientific staffof the Museum and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself, available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History), Cromwell Road, London SW7 5BD, England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Zool.) Trustees of the British Museum (Natural History), 1983 The Zoology Series is edited in the Museum's Department of Zoology Keeper of Zoology : DrJ. G. Sheals Editor of Bulletin : Dr C. R. Curds Assistant Editor : Mr C. G. Ogden ISSN 0007-1498 Zoology series Vol45 No 2 pp 41-101 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 28 July 1983 Miscellanea Contents < 1 GENERAL 29 JUL1983 A revision of the genus Epiclintes (Ciliophora: Hypotrichida) including a redescription of Epiclintes felis comb. n. By P. G. Carey & E. C. Tatchell . Notes on the Family Lekythoporidae (Bryozoa, Cheilostomata). By P. L. Cook & P. J. Hayward A new species of Arthroleptis (Anura: Ranidae) from the West Usambara Mountains, Tanzania. By A. G. C. Grandison The distribution behavioural ecology and breeding strategy of the Pygmy Toad, Mertensophyrne micranotis (Lov.). By A. G. C. Grandison & S. Ashe Additional notes on bariliine cyprinid fishes. By G. Howes 41 55 77 85 95 A revision of the genus Epidintes (Ciliophora: Hypotrichida) including a redescription of Epidintes felis comb. n. Philip G. Carey Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD. Eric C. Tatchell Anatomy Department, Guy's Hospital Medical School, London Bridge, London SE1 9RT. Introduction Several hundred species of ciliated protozoa have now been documented from marine interstitial sediments (Dragesco, 1974). However few of these organisms have been adequately described; indeed some forms although regularly encountered in the extensive literature on this group (Hartwig, 1980) have never been redescribed since the original accounts given by Muller (1786) and Ehrenberg (1830). In an early study, Faure-Fremiet (1950) put forward the hypothesis that interstitial ciliates had a cosmopolitan distribution. This has subsequently been proved correct (Hartwig, 1977), and it has been recognised that these organisms may play a dominant role in the benthic ecosystem (Fenchel, 1969). The opportunity to redescribe one such truly cosmopolitan species came with the discovery of large numbers of the highly contractile hypotrich ciliate Epidintes felis (Muller, 1 786) comb, n. in muddy creeks near the estuary of the river Thames. A brief glance through the literature concerning this species indicated that it had been poorly described. In addition there was some confusion concerning synonymies and it was apparent that a revision of the genus Epidintes was long overdue. Materials and methods Epidintes felis were collected from two locations on the south shore of the Thames estuary: Oare creek, National Grid reference TR0163 and Faversham creek, National Grid reference TR0262. Both locations are near Faversham, north Kent. The organisms were found in small shallow (5- 10 cm) lagoons just below the high tide line along the sides of the two creeks. They were found on the surface of the fine mud and detritus at the bottom of the lagoons or beneath the surface film of the water. The salinity of the water was 70% sea water (SW). Subsequently this species has been found at two sites on the north shore of the Thames, at Shoeburyness, Essex, National Grid reference TQ9485. At this locality the organisms were found firstly at the sand surface, 20 metres from shore, and secondly in association with the red alga Ceramium sp. collected 10 metres from shore. All sites were permanently covered with water at low tide. Cells were cultured in filtered 70% SW, pH 7-8 at 18C under constant illumination from an 8 W fluorescent lamp. Subcultures were made every 5 days. Some difficulty in handling the organisms was experienced due to their thigmotactic nature and inate fragility but when quickly transferred using a wide bore micropipette, this problem was solved. Silver stains of the cortex were prepared by the method of Tuffrau (1967). Silver stained preparations were supplemented by observations using Nomarski interference, phase contrast and brightfield Bull. Br. Mus. not. Hist. (Zool.) 45 (2): 41-54 Issued 28 July 1983 41 42 P. G. CAREY & E. C. TATCHELL illumination, photographs and video recordings. It was found that good structural preser- vation could be obtained with certain fixatives, consequently some specimens were prepared for transmission electron microscopy. Two methods of fixation were employed: (1) Nouzarede's Method (Nouzarede, 1977) for fragile ciliates. The specimens were first fixed in 2% osmium tetroxide in 70% SW neutralised with solid calcium carbonate, for 10 minutes. They were then rinsed in 70% SW and transferred to 10 drops of sea water to a dish and two drops of 25% glutaraldehyde (also neutralised with solid calcium carbonate) were added. The organisms were fixed in this solution for 30 minutes. The ciliates were then returned to the osmium solution for a further 20 minutes. (2) Specimens were fixed in a solution of equal parts of 3% glutaraldehyde in phosphate buffer and 1% osmium tetroxide in phosphate buffer for 20 minutes. These two fixatives were prepared in Millonig's buffer (Millonig, 1961). In both cases the ciliates were washed in 10% ethanol and then dehydrated in graded alcohol solutions before being embedded in TAAB resin. Sections were stained in a saturated solution of uranyl acetate in 50% ethanol for 10 minutes, followed by 0-4% lead citrate in 0-1 M NaOH (CO 3 2 ~ free) for 5 minutes. The first fixative preserved the pellicle most effectively and the second was employed to preserve the subpellicular structures. Nomenclature Epiclintes felis (Muller, 1 786) was first identified by Muller (1 786) under the name Trichoda felis. Although the description was terse and the accompanying diagram gave little information, it is still possible to recognise this very distinctive hypotrich. Muller (1786) described the form of the body as being slightly curved and transparent. It was rather rotund but posteriorly attenuated in the 'tail'. The ventral surface was seen to be covered by 'vibrating hairs' which covered the surface of the 'trunk' right up to the apex of the tail. Bory St Vincent (1824) added a little more information to Muller's description and transferred the organism to the genus Oxitricha as O. felis. He noted that the tail was frequently held curved. Once again the diagram of the species gave few details as to the placement of the cirri. The organism was also independently decribed by Claparede & Lachmann (1858) as a new species in the family Oxytrichina, Oxytricha auricularis. Without knowledge of Muller's T. felis, these authors gave a fairly detailed account of their 'new' species. They carefully described the characteristic shape of the body and the fact that the dorsal surface was seen to be ornamented by 'short batonnets' implanted in the tegument. Detailed infor- mation regarding arrangement of cirri was lacking. In his paper of 1862, Stein proposed the name Epiclintes auricularis to combine Muller's (1786) T. felis and O. auricularis of Claparede & Lachmann (1858). Stein (1864) gave the first detailed description of E. auricularis and supplied further details in his major work (Stein, 1 867). However Stein ( 1 862) was in error in attributing the specific name ''auricularis'' to his new genus Epiclintes. Since the name 'felis' was presented in 1786, 70 years before O. auricularis was published, the earlier name must take precedence. The correct name of the taxon is Epiclintes felis (Muller, 1786) comb. n. Diesing (1866) was the third author to independently identify E. felis and ascribe it to a new genus; Claparedia Diesing, 1866. Although an invalid name, his useful description noted that the peristome of the organism was situated asymmetrically and the body was extremely plastic or deformable. It was also noted that the tail region was highly contractile and caudal 'styli' were seen to be present. Wallengren (1900) in his description of the organism he called Epiclintes ambiguua made a taxonomic error which has subsequently led to great confusion. He stated that the species E. ambiguua was first named by Stein (1862) after Muller's (1786) original description of Trichoda ambiguua. This is incorrect, Stein (1862) described E. auricularis, based on the description of T. felis Muller (1786). However Muller (1786) did describe a T. ambiguua but it is doubtful if this organism is a hypotrich and most certainly does not belong in the genus REVISION OF EPICLINTES 43 Epiclintes. Kahl (1932) failed to note the error and adopted the name E. ambiguus; so that subsequently the name E. ambiguus appears regularly in the literature. Apart from this lapse in the nomenclature of the organism, Wallengren (1900) went on to give an excellent description of his species, which was undoubtedly E. felis. Since these first accounts, many workers on interstitial ciliates have included information regarding this hypotrich, but none has clarified the important details of cirral arrangement, division and behaviour. Morphological Description Epiclintes felis (Muller, 1786) comb. n. E. felis may be described as a genuinely psammophilic species, that is to say, it can be found on, or at least temporarily associated with, sand of the intertidal zone rather than between the interstices. From Figs 1 & 4 it can be seen that the body of this hypotrich is divided into three distinct regions; a dorsoventrally flattened 'head' region leading via a thin neck to a rotund 'trunk' which terminates in an attenuated 'tail'. Many authors have commented on the fact that the organism is highly contractile, with the ability to shorten to at least 25% of its initial length. It is quite clear that the diversity of form recorded for this species is the result of the organism having undergone contraction to a varying degree. All observations recorded here are based on extended cells unless otherwise stated. However in either con- tracted or relaxed state, the three regions of the body are a permanent feature. The true appearance of the body is best seen when the organisms have been left undisturbed for some time. In normal locomotion over the substrate, the body is held extended and frequently slightly curved. The head and tail regions are transparent but the central trunk region is translucent and is frequently seen to be full of food materials. In culture E. felis was seen to feed on bacteria but frequently, in freshly collected material, diatom frustules were seen within the body. The hypotrich is colourless but Biitschli (1887-1 889) noted a yellowish tint. Claparede & Lachmann (1858) were the first authors to quote an accurate size for Oxytricha auricularis. They recorded a length of 300 um. Other reports quote a range of sizes varying from 200-300 um, Stein (1864, 1867), Mereschkowsky (1879), Rees (1884), Kahl (1932). In this study the species was found to vary in size between 100-350 urn. 'Head' Region This region illustrated in Figs 2 & 5 is anteriorly rounded and dorsoventrally flattened. It is truly auriform or 'ear-like' hence the etymology of Claparede & Lachmann (1858). In extended, normally moving animals, the head is equal to or one and a half times as broad as the trunk region. Many earlier authors regarded the trunk as possessing greatest breadth but it is clear that their observations were based on at least partially contracted animals. From Fig. 5 the asymmetric adoral zone of membranelles (AZM) extends around the anterior head region on the ventral surface and along the left side of the oral area entering a distinctive tubular buccal cavity. The cytopharynx is clearly visible but the cytostome is not. The AZM itself comprises some 25-35 fine membranelles held erect. These are twice the length of the cirri on the body, and extend as far as the buccal oveture. Here the membranelles form a distinctive 'cage'. 'Trunk' Region This region is ovoid tapering anteriorly to the head region and posteriorly to the elongated tail. Its length varies with the degree of contraction; in the extended state it is at least twice the length of the anterior portion. When contracted it may equal the head region in length. In normal locomotion the surface of the cell adjacent to the substrate is somewhat flattened. This would account for the thigmotactic nature of this region. On contraction the dorsal surface of the organism becomes distinctly convex in shape, the ventral surface retaining its flattened appearance. This central region may become distended by food material. The 44 P. G. CAREY & E. C. TATCHELL contractile vacuole is single and as Kent (1880-1882) has correctly stated, it may be situated centrally or may be found on the left near the termination of the peristome field. The cytoproct is clearly visible at the dorsal surface, just at the junction of the trunk region and tail. The number and form of the macronuclei has been poorly described; early authors described this organelle as a single hoop-shaped body. The macronuclei are in fact small Figs. 1-3 Epiclintes felis photographed by Nomarski interference microscopy: (1) Entire organism, dorso-lateral view, bar represents lOjim; (2) 'Head' region, ventral view, bar represents 7 urn; (3) Tail' region, ventral view, partly contracted, bar represents 8 )im. REVISION OF EPICLINTES 45 and numerous and are, as Wallengren (1900) noted, spherical or ovoid. Due to difficulties in staining this hypotrich, the micronuclei have never been observed. 'Tail' Region The tail is relatively long, see Figs 1 & 4, at least the length of head and trunk combined, and tapers quickly from the hind regions of the trunk, then less so to its end which is gently rounded. As seen from Figs 1 & 3 this caudal region is very characteristic due to the arrange- ment of cirri which give it a plaited or braided appearance, especially when contracted. Kahl (1932), for example, noted that it resembled a fishbone. It is usually dorso ventral ly flattened and distinctly ribbon-like. This region is highly contractile, much more so than the head or trunk. Possibly longitudinal contractile fibres, akin to the myonemes of peritrich ciliates, are present in the tail as shortening is so rapid. These could contribute to the braided appearance seen so clearly on contraction. The tail serves to periodically jerk the body backwards during normal locomotion. Also on shortening it is frequently noticed that the tail may bend or flex at the junction with the trunk region. It has the ability to bend at least 90 from the longitudinal axis of the cell without sustaining damage. Many of the early workers noted correctly that this region contained no food materials. Cirri Dorsal Cirri It can be seen from Fig. 6 that the dorsal surface of the organism is ornamented by sensory bristles, which take the form of very short non-motile cilia each apparently arising from a shallow pit. There are three distinct longitudinal rows of single, equally spaced bristles in E. felis. The first row arises at the edge of the peristome at the extreme anterior edge of the cell, passes over the head in the midline thence to the trunk and terminates at the posterior extremity of the tail. This row is most easily observed from the lateral view. The two other rows lie on the dorsal surface, at the lateral edges of the organism. They pass round the anterior margin, down the sides of the body and meet the first row at the extremity of the tail. Claparede & Lachmann (1858) were the first authors to identify these organelles in this species. These were the 'batonnets of the tegument'. Ventral Cirri Unlike the dorsal surface, the ventral surface of E. felis is equipped with many rows of short cirri. These take the form of thick bristles that taper distally. They are used extensively in swimming. Biitschli (1887-1889) noted correctly that these cirri constantly oscillate. Due to the constant movement of the organism and the translucency of the trunk region, it was very difficult to observe the numbers and arrangement of cirri without silver impregnation methods, and this could account for the great discrepancies reported by earlier workers. The ventral surface of the head region is equipped with three diagonal rows of cirri, Fig. 5, running right to left from the extreme right margin of the cell to just over two thirds the width of the body. The first lies parallel to the long axis of the cell, the second and third becoming rather more diagonal. The third row terminates at the end of the buccal cavity. Again, almost impossible to observe without staining is the paroral apparatus which runs from the base of the buccal cavity and part way up its right side. The membranes are small and may be subdivided into two or three parts. The fourth ventral row, which may be termed the first row of the trunk region, starts in the head, passes close to the end of the buccal cavity but continues over the trunk region becoming less diagonally slanted as it does so. The cirri in these rows are fairly short and stumpy and are all separated to the same degree, thus cannot be sub-divided into frontoventrals or midventrals. Stein (1864, 1867) and Biitschli (1887-1889) both stated that three rows were present in this area. Again the number of rows of cirri on the trunk region has been poorly reported. Earlier authors did not state clearly whether or not these ventral rows extended into the tail. Muller's (1786) description gave four or five rows for this central portion while Stein (1864, 1867) claimed six or seven. 46 P. G. CAREY & E. C. TATCHELL -azm pa cc 6 Figs. 4-6 Epiclintes felis: (4) Entire organism, ventral view, (azm) adoral zone of membranelles, (be) buccal cavity, (cv) contractile vacuole, (tc) transverse cirri, (me) marginal cirri, (cc) caudal cirri, (vc) ventral cirri, (cp) cytoproct; (5) 'Head' region, ventral view, (db) dorsal bristles, (pa) paroral apparatus, (c) cytopharynx; (6) Lateral a, and dorsal b, views illustrating arrangement of dorsal sensory bristles. REVISION OF EPICLINTES 47 Claparede & Lachmann (1858) noted five short strong rows in the tail region but these probably included rows of marginal cirri which are known to be present, no definite number of trunk rows is mentioned in their description. Rees (1884) claimed a total of twelve rows over the entire body, his diagram shows eight on the trunk and one on the tail. Mereschkowsky (1879) gave a total of twenty for the entire cell, nine for the central portion and five upon the tail. The excellent diagram of Perejaslawzewa (1886) shows a total of seven, three on the anterior and four on the trunk, two of which continue into the tail. Wallengren's (1900) description of E. ambiguua depicted six oblique ventral rows on the trunk and six rows beginning in the mid-trunk region and entering the tail. Nevertheless it can be seen that there are eight rows running diagonally across the body from right to left (Figs, 4 & 7), the last two enter the tail region and are continued to the tip, passing near the mid-line. These cirri are of similar length and are equally separated, but all of the eight rows can only be visualised in stained preparations. Frequently it is seen that these rows of cirri become interlaced and form a distinctive crest down the mid-line of the tail. Marginal cirri are a characteristic feature of this hypotrich species, see Figs 1 & 4. There is a continuous peripheral row of short marginals beginning just behind the termination of the peristome field on both right and left sides, which pass down the lateral edges of the body and join at the apex of the tail. The cirri on the right side appear slightly longer than the left. Stein (1864) noted bristle-like marginals whilst Calkins (1902) showed that they projected from the lateral edges and that they were elongated at the posterior of the cell. The two marginal rows lying at the periphery of the tail and the two ventral rows lying adjacent to the mid-line have, in earlier reports, often been seen as four separate rows in this caudal prolongation. A band of transverse cirri, clearly distinguishable from the ventral cirri by their separate rhythm of oscillation, run down the left side from the first trunk row to the first beginnings of the tail (Fig. 7). They approximate in size and shape to other cirri of the ventral surface. A very short band of caudal cirri, numbering four or five emerge from the ventral surface of the tail. These are finer and slightly longer than the ventrals and marginals and are the caudal 'styli' seen by earlier workers. CC Fig. 7 Epiclintes felis, ventral view of silver stained preparation, (tc) transverse cirri, (me) marginal cirri, (cc) caudal cirri, (vc) ventral cirri. 48 P. G. CAREY & E. C. TATCHELL Division Division in E.felis was observed on a few occasions during this study but precise details are lacking, therefore a description of stomatogenesis and associated development is not given. Wicklow (1979; pers. comm.) clarified the events accompanying stomatogenesis in this unusual hypotrich. In a detailed study of its morphogenesis he found that E.felis possessed a ventral primordium from which many of its ventral rows develop. The row of transverse cirri develop from the posteriormost ends of proter and opisthe cirral streaks. Fission is isotomic, the opisthe retaining the elongate tail and the proter swimming away from division with a simple cylindrical body. The new AZM was seen to develop from the anteriormost of the transverse cirri, in the mid-trunk region. infrastructure When sectioned, E.felis is seen to be unique with the dorsal surface of the cell covered by a multilamellate pellicle of about sixteen layers thick (Fig. 8). This thick pellicle extends down either side of the organism but is modified on the ventral surface. In the vicinity of the ventral cirri the pellicle reverts to the more normal ciliate structure (Pitelka, 1969), but in other regions the multilamellate appearance is seen but the number of layers is reduced to five or six. This unusual pellicle does not extend up onto the dorsal sensory bristles but terminates in a funnel-shaped structure (Fig. 9) rather than a simple pit as seen for example in Euplotes (Ruffolo, 1976), which surrounds the base of the organelle. The repeat distance in this thick pellicle is about 6 nm, thus giving a total width for the pellicle of about 100 nm. The lamellae have the appearance of a series of unit membranes in a collapsed condition. The existence of this multilayered pellicle and its function in this highly contractile hypotrich is certainly worthy of further investigation. Diagnosis of the Genus EPICLINTES Stein, 1862 Trichoda Muller, 1 786 (In part) Oxytricha Claparede & Lachmann, 1858 (In part) Claparedia Diesing, 1 866 A free swimming genus of hypotrich, with a flexible and elastic body. The organism exhibits characteristic cycles of contraction and relaxation. The body is divided into three regions; anteriorly the head is dorsoventrally flattened, bounded at the apex of the cell by the peristome. It tapers gradually to the central portion which is cylindrical. The trunk is twice the length of the head and is usually filled with food materials and granular inclusions. The posterior portion takes the form of an attenuate tail, the length of which may roughly equal or extend to at least three times the length of both head and trunk combined. The trunk and tail are flattened ventrally contributing to the organisms thigmotactic abilities. The peristome is asymmetric, commencing right and enlarging as it traverses the anterior of the cell. The AZM terminates on the left, within the head region, in a distinct buccal cavity. Paroral membranes are present on the right side of this buccal cavity. The ventral surface of this organism is equipped with several rows of short stumpy cirri passing longitudinally or diagonally across the body. These cannot be separated into frontoventrals or midventrals. Marginal cirri are placed in one or two rows along the sides of the trunk and tail. A row of transverse cirri is present, running from just behind the peristome to where the trunk narrows at the start of the tail. A short band of caudal cirri is present also at the tip of this tail. Marine. Species Descriptions From the literature, nine species have been attributed to the genus Epiclintes Stein, 1862. Clearly some of these species are not valid and some have been described on more than one REVISION OF EPICL1NTES 49 Figs. 8-9 (8) Electron micrograph of Epiclintes felis illustrating the multilamellate pellicle of the dorsal surface, fixed by method 1 . The pellicle is 16 layers thick with a repeat distance of 6-2 nm, bar represents 40 nm; (9) Electron micrograph of Epiclintes felis illustrating a section through a dorsal sensory bristle, fixed by method 2, bar represents 0-1 urn. 50 P. G. CAREY & E. C. TATCHELL occasion. Thus study reduces the number of nominal species to three, E. felis, E. caudatus Bullington, 1940 and E. radiosa Calkins, 1902. Corliss (1979) noted that one species of Epiclintes resembled the genus Uncinata. However no member of the genus Epiclintes recorded in the literature resembles this giant species with its characteristic beak-like uncinus. Two species that have been assigned to the genus, but differ fundamentally from the others are Epiclintes vermis Gruber, 1884 and Epiclintes pluvialis Smith, 1900. E. vermis with its vermiform body closely resembles other hypotrich genera such as Holosticha. E. pluvialis superficially resembles Epiclintes but clearly is in possession of a symmetrical peristome. It lacks ventral cirri and the dorsal cirri were said to be long and 'hispid'. These two species differ sufficiently from the diagnosis of the genus Epiclintes to warrant exclusion. Key to species of Epiclintes 1 Frontal 'styles' absent Frontal 'styles' well developed, numbering four or five radiosa 2 Ventral cirri in two longitudinal rows caudatus Ventral cirri in eleven diagonal rows felis Epiclintes felis (Muller, 1 786) comb. n. Trichoda felis Muller, 1 786 Oxitricha felis Bory St Vincent, 1 824 Oxytricha auricularis Claparede & Lachmann, 1858 Epiclintes auricularis Stein, 1 862 Epiclintes ambiguua Wallengren, 1900 Epiclintes ambiguus Kahl, 1932 DIAGNOSIS A psammophilic species commonly found on or adjacent to the sand surface. The body is characterised by its extreme flexibility and contractility. When extended, three regions of the body are apparent. There is a typically auriform anterior, a cylindrical 'trunk' region and an elongate 'tail'. The peristome is asymmetric running around the apex of the cell and into a buccal cavity on the left. The paroral apparatus is small and is situated on the right of the buccal cavity. The cytopharynx is clearly visible. The width of the anterior is equal to, or one and a half times as broad as the trunk. This central region tapers posteriorly to the tail. It is translucent and full of food materials. The contractile vacuole is single and placed centrally or just behind the peristome. A cytoproct lies adjacent to the dorsal surface at the junction of trunk and tail. Macronuclei are small, oval and numerous. The tail is elongate, at least the length of the anterior and central portions combined. It is rounded at the tip and often displays a plaited appearance. The dorsal surface is equipped with three longitudinal rows of sensory bristles, running from the extreme anterior to the tip of the tail, one in the mid-line, the other two at the lateral edges. Ventrally the head is equipped with three rows of short cirri running diagonally from right to left. The trunk has eight rows similarly arranged, the last two of which enter and run down the tail. These two progress right to the tip. Two lateral rows of marginal cirri are present, beginning near the peristome and extending to the point of the tail. A longitudinal row of transverse cirri runs down the left side of the trunk, beginning at the first trunk row and terminating where the tail region begins. In addition, a short row of four or five caudal cirri are present at the extremity of the tail. A prepared slide of Epiclintes felis stained by the silver impregnation technique of Tuffrau (1967) has been deposited in the collection of the British Museum (Natural History), accession number 1982 : 2 : 12. Average length 100-300 um but may shorten to at least 25% of this initial length. Epiclintes felis has been recorded from many locations. Claparede & REVISION OF EPICLINTES 51 Lachmann (1858) found the organism on the coast of Norway and noted that it had also been collected in the Baltic Sea. Stein (1864) similarly found this species in the Baltic but more recent researches have shown it to be truly cosmopolitan. Hartwig (1977) gave an extensive list of its distribution and locations will only be listed briefly here; North Sea, Baltic Sea, White Sea, Gulf of Naples, French Adriatic and Mediterranean Coasts, Black Sea, Caspian Sea, Sea of Japan, Bay of Bengal, American Atlantic Coast, Coast of Mauretania, Brazilian Coast and the Coast of Great Britain. 10 Fig. 10 Epiclintes caudatus After Bullington, 1 940, ventral view. Epiclintes caudatus Bullington, 1940 DIAGNOSIS (Fig. 10) A very long and slender hypotrich, widest centrally and dorsoventrally flattened. It is extremely extensile and contractile. The body is divided into three regions, the anterior portion is almost but not quite equal to the central portion in width. The 'tail' is long and narrow, slightly enlarged at the tip. The peristome is asymmetric and traverses around the head region terminating in a buccal cavity on the left side. Sensory bristles emerge from the dorsal surface, at the lateral edges of the cell, beginning in the mid- peristome region and extending to the tip of the tail. Two longitudinal rows of ventral cirri are present, situated on either side of the midline. Two rows of marginal cirri are also present. In the original description the colour was said to be yellowish and the length at rest was 354 um and the width 28-6 um. The three regions of the body, an asymmetric AZM, the presence of dorsal sensory bristles, marginal cirri and a high degree of contractility con- tribute to the inclusion of this species in the genus Epiclintes. Only two rows of ventral cirri were noted, running longitudinally. This, together with a very elongate head region and a very long tail process serve to distinguish it from E.felis. A marine species isolated from the 52 P. G. CAREY & E. C. TATCHELL 11 12 13 15 Figs. 11-15 (1 1) Oxytricha retractilis After Clapareide & Lachmann, 1858, ventral view of re- laxed and contracted organisms; (12) Oxytricha longicaudata After Strethill- Wright, 1862, ven- tral view of relaxed and contracted organisms; (13) Mitra radiosa After Quennerstedt, 1867, lateral and ventral views of relaxed organism; (14) Mitra retractilis After Kahl, 1932, ventral view of relaxed organism; (15) Epiclintes radiosa After Calkins, 1902, lateral and ventral views of relaxed organism. REVISION OF EPICLINTES 53 West Indies. These organisms usually swim in left spirals but when locomoting over a sub- strate or the underside of the surface film they were seen to move without spiralling, in circles, to the left and right. Epiclintes radiosa Calkins, 1 902 Oxytricha retractilis Claparede & Lachmann, 1858 Oxytricha longicaudata Strethill-Wright, 1862 Mitra radiosa Quennerstedt, 1 867 Epiclintes retractilis Kent, 1 88 1-1 882 Mitra rectractilis Kahl, 1932 DIAGNOSIS A marine species which like E. felis has been described by many authors in various states of contraction. O. retractilis Claparede & Lachmann (1858) and O. longicaudata Strethill-Wright (1862) were both described in the extended state. (Figs 1 1 & 12). They identified three distinct regions of the body, a narrow anterior with asymmetric peristome, a central region that was distinctly ovoid and an elongate highly contractile 'tail' which nearly disappeared on shortening. Strethill-Wright (1862) noted that this tail had a plaited appearance. Five long anterior cirri were observed by both authors. Quennerstedt (1867), Kahl (1932) and Calkins (1902) all described the organism in the contracted state whereupon the tail shortens considerably and the head and central portions tend to be seen as a single unit. Quennerstedt (1867) (Fig. 13) noted the organism was five times as long as broad and that five long straight frontal cirri, he termed 'styles', were present. This author also noted that a single row of marginals were present upon each side of the anterior and central portions, and a double row bordered the tail. Kahl (1932) noted that M. retractilis had a short row of five transverse cirri by which the organism attaches to the substrate. Again four or five long frontal bristles were noted (Fig. 14). Calkins (1902) described the peristome as wide on the right and narrowing as it approached the left side of the body, mention was also made of attachment to the substrate by posterior cirri (transverse cirri) of the tail (Fig. 15). Kahl (1932) was the first to group these independently described organisms under the name Mitra retractilis. However as Kent (1881-1882) had already pointed out, the name Mitra had previously been assigned to a genus of mollusc and was therefore preoccupied. Thus Kahl's designation is invalid. The additional description by Calkins (1902) erected the name Epiclintes radiosa for the organism described by Quennerstedt (1867). The sizes quoted for these organisms are 80 urn for O. retractilis, 75 urn for M. retractilis and 45 um for E. retractilis. This marine species has been recorded from Norway, Scotland, Heligoland, Kiel Sea, Finland, White Sea and the Atlantic Coast of the USA. Vital information on numbers and arrangement of cirri are unfortunately lacking, but they have been grouped due to the characteristic frontal cirri and the conspicious highly contractile tail. Further information is required on this interesting species before a definite decision can be taken regarding its validity. References Bory de St. Vincent, J. B. 1824. Encyclopedic Methodique II, Histoire Natur. des Zoophytes, fais suite a I'hist. nat. des vers de Bruguiere. Paris. Bullington, W. E. 1940. Some Ciliates from the Tortugas. Papers Carnegie Inst. Wash. Tortugas Lab. 32: 179-221. Butschli, O. 1887-1889. Protozoa, Abt. Ill, Infusoria und System der Radiolaria. In Bronn, H. G. (Ed.) Klassen undOrdnungdes Thiers-Reichs 1: 1098-2035. C. F. Winter, Leipzig. Calkins, G. N. 1902. Marine Protozoa from Woods Hole. Bull. U.S. Fish. Commn. 21: 413-468. Claparede, E. & Lachmann, J. 1858. Etudes sur les infusoires et les rhyzopodes. Mem. inst. Genevoise. 5: 1-482. Corliss, J. O. 1979. The Ciliated Protozoa: Characterisation, Classification and Guide to the Literature. 2nd ed. Oxford: Pergamon. 54 P. G. CAREY & E. C. TATCHELL Diesing, K. M. 1866. Revision der Prothelminthen. Sber. haver. Akad. Wiss., Math. Natur. Kl. Munch. 52: 505-580. Dragesco, J. 1974. Ecologie des Protistes Marins. In De Puytorac, P. & Grain, J. (Eds) Actualites Protozoologiques 1: 219-228. Ehrenberg, C. G. 1830 (1832). Beitrage zur Kenntnis der Organisation der Infusorien und ihrer Geographischen Verbreitung, besonders in Siberien. Abh. Akad. Wiss. DDR, Year 1832, 1-88. Faure-Fremiet, E. 1950. Ecologie des Cilies Psammophiles Littoraux. Bull. Biol. France. Beige. 84: 35-75. Fenchel, T. 1969. The Ecology of Marine Microbenthos. IV. Structure and Function of the Benthic Ecosystem, its Chemical and Physical Factors and the Microfauna Communities with Special Reference to the Ciliated Protozoa. Ophelia 6: 1-1 82. Gruber, A. 1884. Weitere Beobachtungen an Vielkernigen Infusorien. Ber. Naturf. Gesell. Freiburg, IB. 3: 57-69. Hartwig, E. 1977. On the Interstitial Ciliate Fauna of Bermuda. Cah. Biol. Mar. 18: 1 13-126. 1980. A Bibliography of the Interstitial Ciliates (Protozoa): 1926-1979. Arch. Protistenk. 123: 422-438. Kahl, A. 1932. Urtiere oder Protozoa I. Wimpertiere oder Ciliata (Infusoria) III. Spirotricha. In Dahl, F. (Ed.) Die Tierwelt Deutschlands 25: 399-650. G. Fischer, Jena Pt. Kent, W. S. 1880-1882. A Manual of the Infusoria, 1-3. David Bogue, London. Mereschkowsky, C. 1879. Studien iiber die Protzoen des Russlands nordlichen. Archiv. fur Mikros Anat.16: 153-248. Millonig, G. 1961. Advantages of a phosphate buffer for OsO 4 solutions in fixation. J. appl. Phys. 32: 1637. Muller, O. F. 1 786. Animacula Infusoria Fluviatilia et Marina. Havniae et Lipsiae. Nouzarede, M. 1977. Cytologie fonctionelle et morphologic experimental de quelques protozoaires cilies mesopsammiques geant de la famille des Geleiidae (Kahl). Bull. Stat. Arcachon (N.S.) 28, (Suppl.) (year 1976), 1-315. Perejaslawzewa, S. 1886. Protozoen des Schwarzen Meers. Zapiski Novorossiiskago Obs. Estestvoispytatelei 10: 79-1 14 (formerly: Memoiren der Neurus. Ges. derNaturh. zu Odessa.) Pitelka, D. R. 1969. Fibrillar Systems in Protozoa. In T. T. Chen (Ed.) Research in Protozoology 3: 437-469. Pergamon Press N.Y. Quennerstedt, A. 1867. Bidrag till Sveriges Infusorie fauna. Ada Univ. Lund. Vol. II, Tomes 4-5 pg.41. Rees, E. 1884. Protozoaires de 1'escault de Test. Tijschr. d. Nederl. Dierk. Vereenig. Supp. Dl: 593-673. Ruffulo, J. J. 1976. Fine Structure of the Dorsal Bristle Complex and Pellicle of Euplotes. J. Morphol. 148:469-487. Smith, J. C. 1900. Notices of some undescribed infusoria, from the infusorial fauna of Louisiana. Proc. Am. Mic. Soc. 21: 87-94. Stein, F. R. 1862. Neue oder nicht genugend bekannten Infusorienformen Tiere. Amtl. Ber. Dl. Naturf. u. Aerzte. 37: 161-162. 1 864. Ueber die neue gattung Epiclintes Stein. Sber K. bohm. Ges Wiss Math. Nat. Kl. (1): 44-46. 1867. Der Organismus der Infusionsthiere nach eigenen Forschungen. In Systematischer Reihenfolge Bearbeitet, II, Leipzig. Strethill-Wright, T. 1862. Observations on British Protozoa. Quart. J. Mic. Sci. New Ser. 2: 2 1 7-22 1 . Tuffrau, M. 1967. Perfectionnements et practique de la technique d'impregnation au protargol des infusoires cilies. Protistologica 3: 9 1-98. Wallengren, H. 1900. Studierofver Ciliata Infusorien Ada Univ. Lund. 36: 1-52. Wicklow, B. J. 1979. Epiclintes ambiguus and Kerona polyporum: comparative ultrastructure, cortical morphogenesis and systematics of two hypotrich ciliates. J. Protozooi 26: 1 6A. Manuscript accepted for publication 22 March 1 982 Notes on the Family Lekythoporidae (Bryozoa, Cheilostomata) P. L. Cook Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD P. J. Hayward Department of Zoology, University College of Swansea, SA2 8PP Introduction (a) Cheilostome Ovicells In recent years, several kinds of cheilostome ovicell have been investigated, and the origins and relationships of the various body wall layers involved have been traced through ontogeny. As a result, it has become obvious that the ovicell, which may be regarded as all the walls defining and protecting the brood chamber and the contained embryo, has several very different origins. Ovicells may be derived wholly, or in part, from diverticula of terminal walls or from extensions of frontal walls and, or, frontal shields. They may be derived wholly, or in part, from one or more zooids. They may be modified, interzooidal, frontally budded kenozooids, or may be formed from one or more kenozooids budded from the maternal zooid, which produces the ovum, or from a sequentially distal zooid or zooids (see Cook, 1979 for brief review). The general term 'ovicell' may therefore be defined functionally, but not morphologically, and it appears that the protective coverings for brood chambers have often been evolved convergently and show only superficial similarities. All ovicells which have been investigated in detail, and all those which may be inferred by inspection to have analogous ontogeny, are 'distal' in position to the opening edge of the operculum of the maternal zooid. Generally, this 'distal' orientation is also the same as the direction of budding of zooids 'away from' the ancestrula as the colony develops. Even in colonies with 'reversed frontal budding' (where the distal part of the orifice is directed towards the ancestrular region, see below), the opening of the ovicell is placed close to the orifice of the maternal zooid, on that side of the operculum which opens to allow protrusion of the lophophore (see Cook & Lagaaij, 1976). This position is correlated with the position of the coelomopore through which the ovum passes to the exterior. In all cheilostomes which have been investigated, the coelomopore is placed at the base of the 'distal' pair of tentacles. Passage of ova into ovicells is achieved by protrusion of the lophophore and apposition of the coelomopore to the ovicell orifice (see Silen, 1 945). Proximally placed ovisacs, which are principally uncalcified have been reported in the anascan genus Aetea, but their ontogeny may be intussusceptive and quite unlike that of other known ovicells (Cook, 1977). Proximal calcified ovicells have also been described in several cheilostome ascophoran genera, which were grouped together in the family Lekythoporidae by Levinsen (1909), and further reviewed by Canu & Bassler (1929). The reported occurrence of 'proximal' ovicells infers that the position of the coelomopore, or at least the behaviour of the maternal lophophore, is radically different from all other cheilostomes in only one group, the Lekythoporidae. Such a difference would be fundamental, although it could be postulated for the genus Inversiula (Microporellidae), which is apparently equally aberrant, and where the operculum is hinged distally (see Bull. Br. Mus. nat. Hist. (Zool.) 45 (2): 55-76 Issued 28 July 1 983 55 56 P. L. COOK & P. J. HAYWARD Moyano, 1972). The brood sac in Inversiula is an interior diverticulum of part of the cuticular outer body wall (tentacle sheath) housed in specially enlarged brooding zooids. The operculum is orientated almost vertically, which would apparently allow passage of ova from a partially protruded lophophore into the ovisac in the 'normal' manner. This has never been observed in living specimens, but Nielsen (1981) has described passage of ova into ovicells closed by the operculum, in which the lophophore was not protruded, but was dimorphic, with very short tentacles. The hingeing of the operculum in the Lekythoporidae is not like that of Inversiula and investigation of specimens of the five genera attributed to the family shows that the ovicell is placed in the 'normal' distal position with respect to the orifice, but that the pattern of astogeny is one of 'reversed frontal budding'. The modifications in zooidal morphology resulting from this type of growth are described below. (b) Frontal budding and colonial and zooidal morphology Frontal budding is a common phenomenon only in cryptocystidean ascophorans, although it occurs sporadically, in a modified form, initiating overgrowths in umbonuloid ascophorans and some anascans. In some cryptocystidean ascophorans, columns of successive, frontally budded zooids may occur (see Banta, 1972, fig. 1 (1), 3). Each zooidal element arises as an intussusceptive expansion of frontal cuticle, with underlying epidermis, of the primary zooid of the series. These together form the boundary wall of an expansion of hypostegal coelom, which is presumed to be provided with nutrients from the visceral coelom by means of the marginal frontal septulae in the frontal calcified shield. As the cuticle-covered bud reaches its full size, a secondary cryptocystidean frontal shield, also with marginal septulae, grows into, and partitions, its coelom into visceral and hypostegal elements. Cryptocystidean frontal shields are interior walls (see Boardman & Cheetham, 1973), and because there is communication between hypostegal and visceral coeloms, these zooids have the potential to form successive series of frontally budded members vertically to the horizontal plane of the primary zooids. Functionally, frontally budded zooids may be regarded as being identical to primary zooids budded through septulae in distal or lateral walls; but morphologically, they may also be regarded as extensions only of the originating primary zooid, because they have no basal walls. An even more integrated kind of frontal budding is found where more than one primary zooid contributes to the development of more than one frontal bud. These interzooidal frontal buds may form nodular masses or erect branches, and are sometimes composed of groups of integrated polymorphic zooids with differing functions (e.g. Hippoporidra, see Cook, 1983). Frontally budded zooids frequently have morphologies and orientations different from those of the primary series (see Boardman et al, 1969, fig. 5), and apart from not possessing basal walls, often have no recognisable lateral walls. In some minute, globular colonies, all budding is frontal, and the position and orientation of the zooids is rigidly patterned and 'reversed' with respect to the direction of growth. In these forms, the distal side of the orifice, the side on which the operculum opens, is directed towards, not away from the ancestrular area, as it is in 'normal' colony growth. In addition, all the calcified walls may be regarded as frontal shields, which consist of a 'concealed' portion, from which new frontal buds arise, and an 'exposed' portion which surrounds the orifice (see Cook & Lagaaij, 1976). Combinations of 'normal' and 'reversed' orientations of frontal buds occur in the genus Sphaeropora (see Cook & Chimonides, 198 1 ). In the Lekythoporidae all colonies are known or inferred to have minute, encrusting bases. In Lekythopora and Poecilopora zooids are budded distally from the ancestrula for only one or two astogenetic generations. The rest of the colony is composed of erect branches of frontally budded zooids, each of which is orientated with the distal edge of the orifice towards the outer face of the branch. Each zooid bud arises interzooidal ly from more than one zooid of the earlier astogenetic generation and develops a cryptocystidean frontal shield with numerous frontal septulae. Fully formed zooids are vase-shaped or columnar and all the vertical calcified walls (whether axial or peripheral) form one continuous 'frontal' shield. FAMILY LEKYTHOPORIDAE 57 m m Fig. 1 Diagrammatic, idealised longitudinal section through axis of an erect branch of Lekythoporidae (based on Lekythopora and Poecilopord). Direction of colony growth (astogenetically distal), arrow at right. External cuticles only shown (thin lines); Primary calcification (thick lines); secondary and extrazooidal calcification stippled; as ascus; ax axial frontal septulae; b topographically distal, frontal bud; ec extrazooidal coelom (derived from amalgamation of hypostegal coelom of several zooids); he hypostegal coelom; ia interzooidal, frontally budded avicularium; m mandible; o brood chamber within peristomial ovicell; op operculum (direction of opening arrowed); p peristome; pa peristomial avicularium; ps peristomial frontal septula; t tube connecting peristomial avicularium with frontal septula; ts tentacle sheath. 58 P. L. COOK & P. J. HAYWARD FAMILY LEKYTHOPORIDAE 59 Septulae are present in the axially directed 'concealed' part of the shield, and in the out- wardly directed part. This, together with the circumoral region forms the equivalent of the 'exposed' part of the frontal shield in conescharelliniform colonies with 'reversed frontal' budding (see Cook & Lagaaij, 1976, and Fig. 1). One or more circles of septulae surround the primary orifice. The septulae allow the expansion of hypostegal coelom axially to form new zooid buds, and peripherally to allow thickening of the exposed frontal shield calcification and, or, the budding of peristomial or interzooidal avicularia and kenozooids. The primary orifice is terminal, and is orientated with the proximal edge of the operculum directed axially and in a more 'distal' position, with respect to the direction of growth of the branch, than the outwardly directed (morphologically) distal edge (see Fig. 1). This is because the longitudinal axes of the zooids are curved towards the outside of the branch. The peristome arises as a ridge between the circum-oral septulae and other frontal septulae. It becomes tubular and often very elongated, continuing the outward curve of the zooid axis. The ovicell is an expanded diverticulum on the peripheral, morphologically distal, side of the peristome, but the curvature of the zooid and peristome is such that it appears to be topographically 'proximal' in position (see Fig. 1). This illusion is completed in several species when the peristome develops a small secondary 'sinus' on its astogenetically distal, but topographically proximal side (Fig. 18). The problem of interpreting the relationships of the primary orifice and the ovicell is exacerbated by the length of the peristome and by the rapid development of secondary calcification and frontally budded kenozooids in most species. The hypostegal coeloms may even amalgamate, with the breakdown of interzooidal cuticle, and form extrazooidal coelom with massive extrazooidal calcification. The presence of septulae and extension of hypostegal coelom on the outer sides of the peristome also assists this calcification. The exposed frontal shields, peristome and even ovicells sometimes become completely obscured within only two astogenetic generations of the growing edge or tip of the branch (Fig. 2). Longitudinal and transverse sections reveal details of the relationships of zooid budding and communication, but because most zooidal axes are curved in more than one plane, com- plete axial sections cannot be obtained. In colonies with regularly cylindrical branches, Catadysis, or irregular cylindrical extensions, Lekythopora and Orthoporidra, all zooid orifices are orientated with their proximal edges pointing axially and nearly vertically, i.e. in the direction of growth. In colonies which appear superficially to be composed of two 'laminae' of zooids, Poecilopora, the proximal sides of orifices oif one 'lamina' are directed towards those of the opposing 'lamina', in the same manner as those of the erect, but rooted genus Flabellopora (see Harmer, 1957). In colonies with rounded branches with one 'frontal' surface, Turritigera, the zooidal axes are twisted as well as curved, but the proximal edges of the orifices are directed towards the axis of the branch. The 'basal' side of these colonies is entirely composed of extrazooidal calcification (Fig. 8). Avicularia occur on the edge of the peristome in all species. They are derived from circum-oral frontal septulae, and one proximo-lateral avicularium at least is always present, although several may occur. A long, tubular connection between the avicularian chamber and the originating septula passes through the peristomial calcification. This often traces a curved path, which perhaps reflects the distortions in zooidal axes which occur during calci- fication of the bud. Sometimes the avicularia are raised on long hollow processes, which may bear small secondary avicularia laterally as well as terminally. The long processes become obscured during later thickening, leaving the avicularia at the surface of the branch, where they may appear to be frontally budded interzooids. In addition, large frontally budded, interzooidal avicularia also develop later in astogeny, especially at the bases of branches. The apparently 'confused' budding pattern (especially in Lekythopora and Orthoporidra), and the effects of extrazooidal calcification therefore make it difficult to distinguish the origins of avicularian types except near the growing tips. Similar difficulties in the genus Turbicellepora have been described by Hayward (1978), who also discussed the descriptive terminology applicable to the large 'spatulate' avicularian rostra which often occur in 'celleporine' forms. Measurements of zooids are also difficult to define and obtain, because 60 P. L. COOK & P. J. HAYWARD L Figs 6-9 Scanning electron micrographs of orifices of Lekythoporidae; arrows indicate direction of colony growth: (6) Catadysis immersum (Busk) BMNH 1963.2.12.247, Montevideo, part of branch showing secondary orifices and frontal pores x26; (7) C. immersum, secondary orifice with avicularia. Note astogenetically distal (topographically proximal) edge of primary orifice at lower level x!30; (8) Turritigera reticulata sp. nov. BMNH 1890.4. 16. 2 A, Marion Island, broken edge of branch from distal side showing one primary orifice (centre), zooid cavities, and thick 'basal' extrazooidal calcification of branch x57; (9) T. fenestella sp. nov. BMNH 1890.4. 16. 2B, Marion Island, detail of zooid at branch tip, showing sinuate primary orifice, secondary orifice and large avicularium x 1 12. the shape of zooids is often irregular, the primary orifice is completely hidden and secondary calcification obscures all limits of structures. One measure of the relative length of zooids may be the distance between the centres of secondary orifices in distal series, but this is very susceptible to error if peristomes are worn or broken. Opercula and primary orifices may often be visible in broken branches, and examination from the topographically 'proximal' (zooid interior) side may reveal differences in shape between them. FAMILY LEKYTHOPORIDAE 61 Systematic Section LEKYTHOPORIDAE Levinsen Lekythoporidae Levinsen, 1909 : 89, 383. TYPE GENUS. Lekythopora MacGillivray. DESCRIPTION. The characters of the family are redefined as follows. Colonies erect or semi-erect, arising from a small encrusting base which becomes strengthened by frontally budded kenozooids, avicularia and, or, extrazooidal calcification later in astogeny. All zooids of the secondary zone of change and repetition (see Boardman et ai, 1969) are frontally and interzooidally budded. Zooids columnar, with a centrally placed, terminal primary orifice. Proximal sides of orifices and asci orientated towards the central axis of the branch. All vertical calcified walls formed as an interior, cryptocystidean shield; axial part concealed, peripheral part and circum-oral region exposed. Shield with a hypostegal coelom and frontal septulae, some of which form a circum-oral ring. Primary calcified orifice straight, curved or sinuate proximally, surrounded by a raised, tubular peristome. Avicularia arising from circum-oral septulae, occasionally numerous, always one proximo-laterally, which appears to be topographically distal. Subrostral chambers terminal, on the edge of the peristome or a mucronate process arising from the peristome. Rostra acute or rounded, mandible slung on a complete bar. Subrostral chamber connected to the originating septula by a long tube, which passes through the calcified wall of the peristome. Interzooidal, frontally budded avicularia sporadic, variously orientated, mandible slung on a complete bar, which may bear a ligula. Brood chamber protected by a large, globular peristomial ovicell, sometimes with an exposed, frontal cuticular area, or with pores and small avicularia on its surface. REMARKS. The family has been defined (e.g. by Bassler, 1953) to include the genera Lekythopora MacGillivray, Poecilopora MacGillivray, Turritigera Busk, Catadysis Canu & Bassler, Orthoporidra Canu & Bassler and Actisecos Canu & Bassler. Of these, all but Actisecos have been described as having 'proximal' ovicells. Actisecos, a rooted, lunulitiform genus, has little in common with the other genera and was assigned to the family Actisecidae by Harmer (1957 : 854). The remaining genera differ from each other principally in details of budding pattern, the shape of the primary orifice and operculum, and the characters of avicularia and ovicells. Most of the specimens examined are small, rarely more than 10-15 mm in height, and type material is often worn and fragmentary. A few colonies, C. immersum, O. compacta, T. reticulata and T. fenestella, are relatively large and exceed 50 mm in height or width. Some species show distinct substratum preferences. Lekythopora is often found on erect hornerid and adeonid bryozoans, Poecilopora on flexible cellariiform and cellulariiform bryozoans or on hydroids, Turritigera is often associated with other bryozoans or with polychaete tubes, and colonies of Orthoporidra may originate on small stones. With one exception, Poecilopora cribritheca, all species are from the Southern hemisphere, and perhaps apart from Lekythopora, are from fairly deep, sometimes very deep, cold waters. Generally, bottom temperatures have been given only for Antarctic localities in published data, and those for low latitude, 'tropical' and 'subtropical' specimens are not known (see Table 2). Several Antarctic species have been reported as abundant, but material examined here, from the Zoological Museum Amsterdam (ZMAC), the National Museum of Victoria (NMV), the British Museum (BMNH), and the Manchester Museum (MM) collections, is not generally plentiful. In view of the enormous, but unworked collections which have been made from the Antarctic shelf, and the increasing number of species becoming known from deep waters, it should be possible, theoretically at least, to obtain larger quantities of some of the species described here, and to analyse their population variation in colony form and zooidal morphology. Several species appear to belong to com- plexes, composed of widely distributed populations, each with different character corre- lations. Additional information on early astogeny, and internal features such as tentacle 62 P. L. COOK & P. J. HAYWARD Table 1 Average measurements of zooids of Lekythoporidae (in mm) Lz Iz Lpo Ipo Lov lov L. hystrix 0-70 0-40 0-12 0-11 0-20 0-40 P.anomala 0-40 0-28 0-08 0-07 0-15 0-20 P. cribritheca 0-50 0-30 0-12 0-11 0-18 0-20 T.stellata 0-50 0-30 0-13 0-16 0-20 0-25 Stn 320 T.stellata 0-60 0-35 0-13 0-18 0-25 0-35 Stn 142 T. reticulata 1-20 0-50 0-13 0-11 0-40 0-50 T. fenestella 0-75 0-35 0-17 0-12 0-25 0-30 C. immersum 0-65 0-50 0-12 0-17 0-25 0-35 O.compacta 0-80 0-50 0-19 0-23 0-40 0-50 O.solida 1-20 0-50 0-20 0-25 0-40 0-50 O.petiolatus 0-60 0-33 0-12 0-13 0-24 0-30 Table 2 Nominal Recent records of Lekythoporidae and Orthoporidroides. Additional temperature data from Murray, 1895 and Livingstone, 1928; estimated depths and temperatures in parentheses Depth Temperature Species Locality data Latitude Longitude (metres) (C) Reference Lekythoporidae L. hystrix Port Phillip Heads 3830'S 14430'E (60) (+17) L.hvstrix Port Western 3830'S 14530'E L. hystrix Port Jackson 34S 151E (60) (+17) BMNH BMNH BMNH L.avicularis Port Jackson 34S 151E 146 (+17) Maplestone 1909 P.anomala Port Phillip Heads 3830'S 14430'E (60) (+17) P.anomala Twofold Bay 3659'S 15020'S 275 (+12-8) P.anomala Maria Island 4237'S 148E 73 BMNH BMNH Thornely 1924 P. cribritheca Sulu Archipelago 68'N 12119'E 275 (+12) Harmerl957 T.stellata Montevideo 3717'S 5352'W 1100 +2-8 T.stellata Cape of Good Hope 3504'S 1837'E 275 +8-3 T.stellata Cape Horn 53S 68W T.stellata Is. Topar 508-5'S 7441'W 360 T.stellata Caleta Hale 4757'S 7441'W 40-50 T.stellata BellinghausenSea 70S 8048'W 7500 +0-9 T.stellata BellinghausenSea 7020'S 8323'W 459 +0-8 T.stellata BellinghausenSea 7015'S 846'W 659 +0-8 T.stellata BellinghausenSea 7023'S 8247'W 480 +0-8 T.stellata BellinghausenSea 7114'S 8914'W 460 -0-3 T.stellata BellinghausenSea 7118'S 882'W 435 -0-3 Busk 1884 Busk 1884 Waters 1905 Moyano 1974 Moyano 1974 Waters 1904 Waters 1904 Waters 1904 Waters 1904 Waters 1904 Waters 1904 T. reticulata Marion Island 38S 4640'W BMNH T. fenestella Marion Island 38S 4640'W BMNH Continues FAMILY LEKYTHOPORIDAE 63 Species Locality data Latitude Depth Longitude (metres) Temperature (C) Reference T. spectabilis Uruguay 3649'S 5315-4'W 1661-1679 d'Hondt!981 3816-9'S 5156-1'W 4382-4402 d'Hondt 1981 3655-7'S 5301-4'W 2707 d'Hondt!981 C. immersum Montevideo 3717'S 5352'W 1100 + 2-8 Busk 18 84 C. immersum Burdwood Bank 5425'S 5732'W 103 Hay ward 1980 O. compacta BellinghausenSea 70S 8048'W 7500 + 0-9 Waters 1904 O. compacta Bellinghausen Sea 7118'S 8802'W 435 -0-3 Waters 1904 O. compacta Bellinghausen Sea 7119'S 8737'W 436 -0-2 Waters 1904 O. compacta Halley Bay 7531'S 2636'W BMNH O. compacta Scotia Bay 61S 45W BMNH O. compacta McMurdo Sound 7705'S 16417'E 256 BMNH O. compacta? South Chile 6228-5'S 5941-5'W 119 BMNH O. compacta? Queen Maude Land 7019-4'S 2412-6'E Redier 1965 to to 7020-9'S 2413-4'E O. setosa Commonwealth Bay 6432'S 9720'E 201 Thornely 1924 O. setosa Commonwealth Bay 6608'S 9417'E 220 Thornely 1924 O. setosa Adelie Land 6655'S 14521'E 782 + 1-8 Thornely 1924 O. setosa Adelie Land 6632'S 14139'E 287 + 1-62 Thornely 1924 O. setosa Graham Land 65S 64W 750 Vigeland 1952 O. setosa Graham Land 65S 64W 90 Vigeland 1952 O. setosa Graham Land 65S 64W 140 Vigeland 1952 O. petiolata Cape Horn 5313'S 6831'W 97 + 6-6 Waters 1905 O. petiolata Burdwood Bank 5425'S 5732'W 103 Hayward 1980 O. petiolata Houtjes Bay 3410'S 1810'E BMNH O. solida S. W. Australia 4242'S 13410'E 4758 + 1-1 Busk 1884 Orthoporidroides O. erectus Magellan Straits 55S 72W 320 ( + 7-8) BMNH O. erectus Montevideo 3717'S 5354'W 1100 + 2-8 BMNH O. erectus Is. Innocentes 5033'S 7453'W 150 Moyano 1974 O. erectus Is. Topar 508-5'S 7441'W 360 Moyano 1974 O. erectus Canal Zenteno 5249'W 7340'S 30-40 Moyano 1974 O. aff. erectus E. Falkland Is. 5209'S 5814'W 79 + 8-3 BMNH O. aff. erectus E. Falkland Is. 5029'S 5852'W 140 + 4-78 BMNH O. aff. erectus E. Falkland Is. 5030'S 5819'W 141 + 5-4 BMNH (9. aff. erectus E. Falkland Is. 5231'S 5829'W 146 + 5-45 BMNH O. aff. erectus E. Falkland Is. 5050'S 5713'W 144 + 5-61 BMNH O. robusla Chile 3737'S 7340'W 600 Moyano 1981 number, opercular variation etc., should result in a better understanding of the systematic relationships within the Lekythoporidae, and the relationships of the family with other ascophoran groups. LEKYTHOPORA MacGillivray Lekythopora MacGillivray, 1883 : 194. TYPE SPECIES. L. hystrix MacGillivray. 64 P. L. COOK & P. J. HAYWARD DESCRIPTION. Colonies forming short, irregularly cylindrical branches. Zooids with several series of frontal septulae. Primary orifice sinuate, operculum extended proximally but not distinctly sinuate. Ovicells with a frontal, marginally porous area covered by cuticle. REMARKS. L. hystrix appears to be the only species referable to the genus. Harmer (1957 : 884) referred Phylactella lucida Hincks to Lekythopora', this species has been discussed by Cook (1968 : 220) and referred to Celleporina. L. perplexa Harmer (1957 : 884, pi. 59, figs. 5-7, 10) is also a 'celleporine' species with 'normal' frontal budding and tabulate ovicells and may be provisionally referred to Celleporina. Harmer (1957 : 885) also mentioned an unnamed species of Lekythopora from Japan. Examination of his specimens shows that these colonies, too, are referable to Celleporina. L. laciniosa Calvet (1907 : 445, pi. 29, figs. 13, 14) is attributable to Celleporina. Lekythopora hystrix MacGillivray Lekythopora hystrix MacGillivray, 1883 : 194, pi. 2, figs. 6, 6a-d; 1885 : 113, pi. 2, fig. 6: 1888 : 201, pi. 156, figs. 4-10; 1895: 106, pi. 14, figs. 1-2. Waters, 1885 : 308. Brown, 1958 : 83. Lekythopora avicularis Maplestone, 1909 : 273, pi. 78, fig. 12. SPECIMENS EXAMINED. BMNH, Port Phillip Heads, Victoria, 1897.5.1.934,937, 1899.5.1.1328, 1934.10.20.103. Port Western, S. Australia, 1934.2.20.23. Australia, 1883. 10.15.125-145. Port Jackson, Sydney, 1981.4.1.1. NMV, Port Phillip Heads 646 II, 64612 (fig'd MacGillivray, 1888) and 64602. DESCRIPTION (Figs. 14a, 15). Primary zooids with short peristomes, frontally budded zooids with long, curved peristomes, directed peripherally, sometimes with a secondary, topo- graphically proximal, sinus. Peristomal avicularia with curved, acute rostra, placed on the edge of the peristome. Interzooidal avicularia elongated, only slightly expanded and rounded terminally, orientated proximally to branch growth. Ovicell with frontal area surrounded by minute pores. REMARKS. MacGillivray (1888 : 21 1) noted that the primary orifice was difficult to see, and described it variously as 'rounded' (1883) or 'with a notch in the lower lip' (1888). In one illustration (1 885, pi. 2, fig. 6), he actually figured the relationship of the proximal sinus with a distally placed ovicell, but apparently did not notice that this orientation was in contra- diction to his description of the ovicell as 'proximal'. However, he was aware of the reversal of orifice orientation with respect to direction of growth in P. anomala, and compared it with that of L. hystrix. The relationships of L. mooraboolensis Maplestone (1902 : 25, pi. 2, fig. 18) and L. kitsoni Maplestone (1902 : 25, pi. 2, fig. 19), from the Australian Tertiary, are difficult to evaluate without specimens. L. mooraboolensis was described from a single, small, globular specimen of 3-4 zooids and may have been the young astogenetic stage of a colony of Conescharellina. L. kitsoni was described as ligulate and bilaminar and appears to have had 'proximal' ovicells with a frontal area. It may be assignable to either L. hystrix or P. anomala (see below). Apart from L. kitsoni, L. hystrix is apparently the only species of the family which has a fossil record. Both Waters (1885) and MacGillivray (1895) noted its occurrence from several Tertiary Australian localities (see also Brown, 1958). POECILOPORA MacGillivray Poecilopora MacGillivray, 1886 : 136. TYPE SPECIES. P. anomala MacGillivray. DESCRIPTION. Colonies with compressed, cylindrical, or 'bilaminar' branches, bifurcating in one plane. Zooids in opposing interdigitating series. Primary orifice sinuate, operculum extended proximally but not sinuate. Ovicells with a frontal area covered by cuticle, visible late in ontogeny. FAMILY LEKYTHOPORIDAE 65 66 P. L. COOK & P. J. HAYWARD Fig. 14 Silhouettes of opercula of Lekythoporidae and Orthoporidroides, Scale bar = 0-50 mm; (a) Lekythopora hystrix; (b) Poecilopora cribritheca; (c) Turritigera stellata; (d) T. reticulata; (e) T. fenestella; (f) Catadysis immersum; (g) Orthoporidra compacta; (h) 0. petiolatus; (i) 0. solida; (j) Othoporidroides erectus. Poecilopora anomala MacGillivray Poecilopora anomala MacGillivray, 1886 : 136, p. 1, fig. 9; 1888 : 21 1 (as P. anomola), pi. 156, figs. 11-13. ^Turritigera stellata Thornely (not Busk), 1924 : 18. SPECIMENS EXAMINED. BMNH, Port Phillip Heads, 1887.12.10.67, 1888.11.14.103,134,291, 1897.5.1.947.948,949, 1889.7.1.3613. Port Western, South Australia, 1899.7.1.5129. ChallengerStn. 163 A, Twofold Bay, New South Wales, 220 m, 1899.7.1.4081, 1963.2.12.15. DESCRIPTION (Figs. 3, 12, 13) The small colonies rarely exceed a height of 10 mm, and are branched in one plane. The 'bilaminar' growth resembles that of Flabellopora. The primary orifice has an indistinct sinus proximally. Zooids small, with porcellanous frontal shields with scattered septulae. Peristomes long at lateral margins of colony, flaring terminally; short elsewhere, with one acute avicularium. Interzooidal avicularia short, slightly expanded, and rounded terminally, randomly orientated. Ovicells prominent even late in ontogeny, protruding beyond the secondary thickening, frontal area surrounded by minute pores. REMARKS. The early stages of the colony may consist of the ancestrula and one encrusting zooid generation only ( 1 899.7. 1 .361 3). MacGillivray (1886 : 137) noted that the reversal of orifice orientation was the explanation for the apparently 'proximal' ovicells, and postulated that L. hystrix might have a similar structure. Thornely's (1924) record of 'Turritigera stellata' was not from an Antarctic locality. Although most of the stations mentioned in her paper were from Wilkes Land, Antarctica, FAMILY LEKYTHOPORIDAE 67 the locality 'Maria Island' (4237'S, 148E, 73 m) is Tasmanian (see Livingstone, 1928 : 8). Thornely noted that her colonies were 'bilaminar', that the zooids had tubular peristomes with only one marginal avicularium, and that the ovicells had a perforated frontal area. The locality of her specimens indicates that they were either L. hystrix or P. anomala, and the colony form suggests strongly that they were the latter. The specimens from Challenger Stn 163 A were not described by Busk (1884). The other bryozoans from this Station include a large number of cellariiform and cellulariiform species, which provide substrata for the colonies of P. anomala. Poecilopora cribritheca (Harmer) Catadysis cribritheca Harmer, 1957 : 886, pi. 59, figs 9, 12-15, 22, 23. SPECIMENS EXAMINED. ZMA, LECTOTYPE, Siboga Stn 105, Sulu Archipelago, 275 m. BMNH, Paralectotype, as above, 1 98 1 .5.6. 1 . DESCRIPTION (Fig. 14b). Colonies with compressed cylindrical branches. Primary orifice with a wide sinus. Zooids with porcellanous calcification and scattered frontal septulae. Peristomial avicularia not protuberant, rounded or subtriangular. Ovicell immersed, but the frontal area, which has minute, slit-like pores, remains visible. REMARKS. Harmer (1957) assigned P. cribritheca to Catadysis, which has cylindrical branches of zooids without long peristomes but with numerous frontal pores. The porcellanous calcification, compressed branches, long peristomes and exposed frontal area of the ovicells of P. cribritheca are all characters typical of Poecilopora. P. cribritheca differs from P. anomala in its slightly larger zooids, and the shape and distribution of the avicularia. Interzooidal avicularia seem to be absent, but the material is fragmentary, and incomplete. P. cribritheca is the only species of Lekythoporidae known to occur in the tropical waters of the Northern hemisphere, but is from fairly deep water (275m) near the equator. TURRITIGERA Busk Turritigera Busk, 1884 : 129. TYPE SPECIES. T. stellata Busk. DESCRIPTION. Colonies with cylindrical branches, sometimes becoming reticulate with anastomoses. Most zooids curved so that their peristomes open on one side of the branch. Branches dichotomous or formed by frontal ly budded zooids arising in groups at right angles to the primary branch, in more than one plane. Primary orifice sinuate, but operculum curved or sinuate proximal ly. Peristomial avicularia sometimes numerous, occurring on the edge of peristomes, often on raised processes. Ovicell rapidly occluded by secondary calcification, but often remaining prominent. Turritigera stellata Busk Turritigera stellata Busk, 1884 : 130, pi. 24, fig. 1. Waters, 1888 : 22, pi. 1, figs. 22, 25; 1904 : 76, pi. 5, fig. 3, pi. 8, fig. 13; 1905 : 242, pi. 29, figs. 19, 20. Moyano, 1974 : 1 8, figs. 4, 8, 3 1-34. not Turritigera stellata Thornely 1924, see P. anomala. SPECIMENS EXAMINED. BMNH, Challenger Stn. 320, 1100m, 1887.12.9.517.520B, 1899.7.1.3166,3167, 1944.1.8.240,241. Challenger Stn. 142, 275m, 1887.12.9.518,519, 1899.7.1.498,499,500, 1934.2.16.13, 1944.1.8.239,242. Locality? New Zealand, 1936.12.30.155. MM, Challenger Stn. 320, T38-40; Expedition Antarctique Beige, T 17-37. DESCRIPTION (Figs. 4, 10, 14c). Colonies reaching a height of 20-30 mm, branched irregularly. Primary orifice with a small, rounded proximal sinus, operculum with a distal flange, curved proximally. Zooids with numerous frontal septulae, which become slit-like as 68 P. L- COOK & P. J. HAYWARD calcification increases. Peristomes long, flaring terminally, with 6ne to seven terminal avicularia, which are sometimes raised on short processes. Mandibles and rostra acute. Interzooidal avicularia elongated, slightly expanded and rounded terminally, randomly orientated, sometimes present on non-zooidal 'basal' side of colony. Ovicell with a small frontal area which is rapidly obscured by frontal thickening. 18 tentacles (Waters). 'Basal' thickening extrazooidal, with slit-like pores and minute avicularia. REMARKS. The South American (Stn. 320), South African (Stn. 142) and Antarctic popu- lations show some differences in characters. Colonies from Stn. 320 are rather delicate, with numerous small peristomial avicularia, and interzooidal avicularia on the 'frontal' and 'basal' sides. There is relatively little extrazooidal thickening (see also Moyano, 1974). Colonies from Stn. 142 are at a later stage of ontogenetic development and consist of astogenetically earlier fragments. They appear more robust, with considerable 'basal' and 'frontal' thickening. The axially directed, proximal, peristomial avicularium is large and the remainder are reduced in number, usually to two or three. Interzooidal avicularia are absent. Colonies from Waters's (1904) Antarctic Stations are very robust, and the zooids tend to be arranged biserially (c.f. T. reticulatd). Waters noted that some colonies possessed anastomoses (1904, pi. 5, fig. 3a). The zooids have numerous peristomial avicularia, but interzooidal avicularia are very rare. In contrast, specimens from ?New Zealand (locality doubtful) are delicate but have one enlarged peristomial avicularium, like those from South Africa. T. stellata would therefore appear to be a very variable species. The extent and significance of this variation can only be assessed when complete colonies, showing all astogenetic and ontogenetic stages can be compared from a wide range of populations. Recently, for example, d'Hondt( 1981 : 41, pi. 5, figs. 1-3) has described a new, abyssal form, T. spectabilis, from a depth range of 1661-4402 m, off Uruguay. T. spectabilis resembles some forms of T. stellata in having only one, large peristomial avicularium. Ovicells were absent. The operculum of T. stellata figured by Waters (1904, pi. 5, fig. 3b) was distinctly sinuate, like the primary orifice. Waters's preparation of opercula (MM T33) is badly preserved and no sinus is visible. None of the numerous opercula in the Busk preparations from South African material (BMNH 1 899.7. 1 .498,499) shows any sinus. Turritigem reticulata sp. nov. SPECIMENS EXAMINED. BMNH, HOLOTYPE and Paratypes, all parts of one colony, Marion Island, 1890.4.16.2A. ETYMOLOGY. Reticulatus (L) net-like, referring to the colony form. DESCRIPTION (Figs. 8, 11, 14d, 17, 18). Colonies large, 50mm in height, reticulate, with anastomosing branches. Zooids very large, alternating and biserial, peristomes short. Primary orifice very small, with a fairly wide sinus, operculum distinctly sinuate. Secondary orifice often forming a topographically proximal sinus. Frontal pores numerous, becoming slit-like. Peristomial avicularia small, one to three are bourne on a stout suboral, topo- graphically distal process, mandibles short, triangular. Interzooidal avicularia absent. Ovicells very large, with marginal pores; prominent at first, becoming partially obscured by extrazooidal thickening, often orientated medially. REMARKS. The reticulate growth form of T. reticulata is shared by T. fenestella (see below) and, according to Waters (1904), by some colonies of T. stellata. The large zooidal dimensions of T. reticulata, together with its very small primary orifice distinguish it from T. fenestella. Turritigera fenestella sp. nov. SPECIMENS EXAMINED. BMNH, HOLOTYPE and Paratypes, all parts of one colony, Marion Island, 1890.4. 16.2B. FAMILY LEKYTHOPORIDAE 69 70 P. L. COOK & P. J. HAYWARD ETYMOLOGY. Fenestella (L) a little window, referring to the colony form. DESCRIPTION (Figs. 9, 14e, 16). Colony large, 50mm in height, reticulate, with anastomosing branches. Zooids irregularly arranged, fairly small, with a primary orifice with a proximal plate, in which there is a distinct round sinus; operculum also with a distinct sinus. Frontal pores fairly numerous, rounded, becoming obscured. Peristomial avicularia one to three, one bourne on a very long, curved suboral process. Mandibles triangular, the suboral mandible large, rostrum hooked terminally. Ovicells small, smooth, remaining prominent, although obscured by extrazooidal thickening, often orientated medially. REMARKS. The similarities in colony form of T.fenestella and T. reticulata, which were from the same locality, tend to obscure the distinct differences in budding pattern, zooidal size and orificial characters which exist between them. The zooids of T. fenestella are similar in size to those of T. stellata, but differ completely in the form of the avicularian processes and in the shape of the operculum. The three species described above form a complex, which requires more plentiful material for elucidation. CATADYSISCanu & Bassler Catadysis Canu & Bassler, 1927 : 12, 23, 25. TYPE SPECIES. Myriozoum immersum Busk ( = Schizoporella challengeria Waters). DESCRIPTION. Colonies with regularly cylindrical branches, zooids opening on all sides. Zooids with numerous frontal septulae, peristomes not prominent. Primary orifice with a rounded sinus, operculum curved proximally. Peristomial avicularia small, interzooidal avicularia absent. Ovicells imperforate. Catadysis immersum (Busk) Myriozoum immersum Busk, 1884 : 170, p. 25, fig. 4. Schizoporella challengeria Waters, 1888 : 30, pi. 2, figs. 25-28. Catadysis challengeria (Waters) Canu & Bassler, 1927 : 11, Bassler, 1953 : G233. Catadysis immersum (Busk) Harmer, 1957 : 885. Hayward, 1980 : 712, figs. 5A-D. SPECIMENS EXAMINED. BMNH, Challenger Stn 320, 1100m, 1887.12.9.683,684,6856, 1899.12.12.18, 1899.7.1.2295, 1934.11.12.57, 1944.1.8.333,334, 1963.2.12.221,247. 'Bruce' Stn 346 (Scottish National Antarctic Expedition) Burdwood Bank, 1936.12.30.297. DESCRIPTION (Figs. 2, 6, 7, 140- Colonies large, 50x30 mm, branching in all planes. Zooids with numerous frontal septulae which become immersed by extrazooidal calcification as a series of slit-like pores. Branches cylindrical, smooth, composed of four series of zooids. Peristomes never prominent, orifices rapidly obscured by secondary calcification, leaving rounded pits on the branch surface. Peristomial avicularia small, proximal and distal, mandibles triangular. Ovicells obscured very early in ontogeny. REMARKS. Waters (1888) introduced the name 'Schizoporella challengeria' for Busk's species because he considered that it belonged to the genus Schizoporella and that the combination was therefore preoccupied by Onchopora immersa Haswell (1 880), which he also assigned to Schizoporella. O. immersa is referable to the genus Tetraplaria (see Harmer, 1957 : 1055), and as Catadysis has little in common with Schizoporella, Waters's name 'challengeria' is unnecessary (see also Harmer, 1957 : 885, and Hayward, 1980). Most specimens are small and fragmentary, but one from Challenger Stn 320 exceeds 50 mm in width, with numerous branches. ORTHOPORIDRA Canu & Bassler Orthopora Waters, 1 904 (preoccupied). Onhoporidra Canu & Bassler, 1927 : 12, 23, 34. TYPE SPECIES. Orthopora compacta Waters. FAMILY LEKYTHOPORIDAE 71 DESCRIPTION. Colony with irregularly cylindrical branches. Primary orifice usually straight proximally, but opercula may be curved. Zooids with numerous frontal septulae, calcifi- cation smooth and very thick. Peristomial avicularia often raised on long, hollow mucronate prominences. Ovicells large, usually imperforate, but with small frontal avicularia. REMARKS. A large number of specimens has been recorded from many Antarctic and sub- Antarctic localities. These appear to have the general character of O. compacta as described below. Small differences in the primary orifice and operculum, however, indicate that population studies are necessary, and it is possible that several distinct species will be found to exist, once plentiful material has been examined. Orthoporidra compacta (Waters) Orthopora compacta Waters, 1904 : 75, pi. 5, figs. 4a-i. ?Redier, 1965 : 4, 32. Orthoporidra compacta: Canu & Bassler, 1927 : 12; 1929 : 515, fig. 214. Rogick, 1965 : 406. Moyano, 1978:44. ICellepora setosa Thornely, 1924 : 17, fig. 5. Livingstone, 1928 : 76, pi. 3, fig. 8, Figs. 18-20. Moyano, 1978:44. ?not Cellepora setosa: Redier, 1965 : 30; 1966 : 2. SPECIMENS EXAMINED. MM, T10-15, 343, Waters's( 1904) preparations including operculum (T14) and tentacles (T12), Expedition Antarctique Beige. BMNH, Halley Bay, 1966.3.4.4. Bahia, Chile, 1971.3.26.29. Scotia Bay, S. Orkneys, 1920.12.1 1.3. McMurdo Sound, 1981. 3.1.1. DESCRIPTION (Figs. 14g, 21, 22). Colonies large, branched in several planes. Primary orifice and operculum slightly curved proximally. Zooids large, frontal septulae distinct, marginal and circum-oral, in two rows across exposed frontal shield. Suboral, topographically distal, avicularium raised on a long tubular process, mandible rounded or triangular, rostrum bar with a ligula. Interzooidal avicularia oval or expanded terminally, rostrum bar with a ligula, usually orientated proximally. Ovicell large with small avicularia on the surface. 24 tentacles (Waters). REMARKS. The only type material consists of decalcified sections and preparations etc. The operculum (T14) differs from that figured by Waters (1904) and resembles that of 0. solida in being slightly curved proximally, with lateral proximal lacunae which may be muscle insertions (cf. Waters, 1904). Thornely (1924) introduced C. setosa for specimens from several Antarctic localities. Her specimens were redescribed by Livingstone (1928), who noted that neither the primary orifice nor operculum were sinuate. Both authors figured large ovicells with small pores, or avicularia?, on their surfaces. The only available specimen named as O. setosa, which is part of the material described or listed by Redier (1965, 1966), is a bifurcated branch from Antarctica identified by Dr A. B. Hastings (Stn 136, Expedition Antarctique Beige, BMNH 1964.8.2.5.). Although similar to O. compacta in most characters including those of the large, wide, ligulate interzooidal avicularia, this specimen differs in the dimensions of the orifice and in possessing a sinuate operculum. The primary orifice is straight proximally but has a small central notch, is longer than wide and generally far smaller than that of O. compacta. 'C. setosa Redier' is therefore, in part at least, a separate taxon from C. setosa Thornely and O. compacta. Additional material is required before all the Antarctic records of these forms can be reassessed. Redier's (1965) material of O. compacta also requires re-examination. His record of the Philippine Islands as a locality for the species is a mistaken reference to the illustrations included by Canu & Bassler (1929) in their paper on Philippine bryozoans. Orthoporidra solida (Busk) Cellepora solida Busk, 1884 : 200, pi. 29, fig. 12. Waters, 1904 : 76, pi. 5, fig. 5. 72 P. L. COOK & P. J. HAYWARD FAMILY LEKYTHOPORIDAE 73 SPECIMENS EXAMINED. BMNH, LECTOTYPE (chosen here), Challenger Stn 160, 4758m. 1899.7.1.3525, the specimen figured by Busk, 1884. Paralectotypes 1887.12.9.775-777; 1899.7.1.3526. DESCRIPTION (Figs. 5, 14i, 19, 20). Orthoporidra with very large zooids, with numerous, scattered frontal septulae. Primary orifice and operculum curved proximally. Peristome not very prominent. Peristomial avicularia proximal and paired, lateral, raised on blunt mucros, mandibles rounded, rostra subtriangular. Interzooidal avicularia oval, large. Ovicells large, with small avicularia and scattered pores. REMARKS. The colonies form branched cylindrical masses 5-10 mm in height. Secondary thickening is less developed than in O. compacta, and consists of numerous kenozooids as well as extrazooidal tissue. The avicularian processes are short and blunt, quite unlike those of O. compacta. Busk (1884: 200, pi. 29, fig. 12a) described the large interzooidal avicularia (as 'operculum' in the explanation of Plate 29), and Waters (1904, pi. 5, Fig. 5) also illustrated an operculum. The preparations in the BMNH show that the opercula are far larger than the dimensions indicated by Waters's figure. Orthoporidra petiolata (Waters) Celleporapetiolata Waters 1904 : 76 (nom. nud.); 1905 : 241, pi. 29, figs. 19, 20. Catadysis petiolata: Hayward, 1980 : 714, Figs 5 E-H. SPECIMENS EXAMINED. MM, T2-T6, Expedition Antarctique Beige. BMNH, Houtjes Bay, S. Africa, 1936.12.30.300c. Stn 346, Scottish National Antarctic Expedition, Burdwood Bank, 1981.3.1.12. DESCRIPTION (Fig. 14h). Colony arising from a minute thickened base of kenozooids, encrusting erect bryozoa, 10-15 mm in height. Zooids with marginal and circum-oral septulae. Primary orifice and operculum with a small sinus. Suboral peristomial avicularium raised on long processes, mandible small, rounded or elongated and triangular. Ovicells small, prominent at first, with frontal pores and small avicularia. 14 tentacles (Waters). REMARKS. Waters (1904) mentioned that a species which he proposed to name 'C. petiolata' was similar in character to O. compacta. In his formal introduction (1905) he gave no further details, and figured only the operculum and mandibles. Waters's preparation (T5) shows an operculum with a small rounded sinus. The generic assignment of O. petiolata is arbitrary. The orificial characters are similar to those of some species assigned here to Turritigera (e.g. T. fenestella), and to Catadysis immersum (see Hayward, 1980). The budding pattern of the zooids, and the distribution of the frontal septulae resemble those of Orthoporidra compacta. Comparison of the Lekythoporidae with the genus Orthoporidroides Moyano Frontally budded zooids with cryptocystidean frontal shields, which form massive or cylindrical, erect branches are typical of the Family Celleporinidae Harmer. Genera such as Turbicellepora (see Hayward, 1978) and Celleporina (see Harmer, 1957) also have oral avicularia derived from circum-oral frontal septulae, and often have elongated peristomes with terminal avicularia. Large, interzooidal (vicarious) frontally budded avicularia, and ovicells with porous frontal areas (tabulae) are also typical of these genera. The genera included in the Lekythoporidae are therefore distinguished from those in the Celleporinidae principally by their 'reversed' frontal budding pattern. The apparent close similarity between the Families is illustrated by Orthoporidroides erectus (Waters). CELLEPORINIDAE Harmer Celleporinidae Harmer, 1957 : 899. 74 P. L. COOK & P. J. HAYWARD ORTHOPORIDROIDES Moyano Orthoporidroides Moyano, 1 974 : 20 1 . 1 98 1 . TYPE SPECIES. Cellepora armata var. erecta Waters. Orthoporidroides erectus (Waters) Cellepora armata var. erecta Waters, 1888: 36. pi. 3, figs 4, 4 1 , 43. Orthoporidroides erectus : Moyano, 1 974 : 2 1 , Figs. 6, 35^40. SPECIMENS EXAMINED. BMNH, Challenger Stn 308, 320m, 1889.12.12.7. Challenger Stn 320. 1100m. 1888.3.14.3, 1899.7.1.3584, 3587, 3592, 3602, 3606; 1934.11.12.5. Shackleton-Rowett Exped., ?S. Orkney, 1923.12.1.39,48. DESCRIPTION (Fig. 14j). Colonies erect, branching, arising from a small, encrusting base. Zooids budded frontally and distally, primary orifice with a shallow sinus directed towards the outside of the branch, operculum produced into a triangular sinus. Avicularia suboral, terminal on a long, proximally produced peristome, mandible acute. Ovicell hyperstomial, with a small, frontal slit. REMARKS. Moyano (1974) has illustrated the morphology of O. erectus fully and compared it with that of Turritigera stellata. Moyano also noted similarities between O. erectus and Cellepora petiolata Waters (1905), see above. The opercula of O. erectus figured by Moyano resemble those illustrated by Waters, but differ slightly from those of the specimens from Challenger Stn 320, which are very variable (Fig. 14j). These specimens were not described by Busk ( 1 884) or Waters (1888). Discussion Study of the relationships of the ovicell in the Lekythoporidae illustrates the importance of analysis of the astogenetic structure of colonies, as well as the ontogeny of their member zooids. It is interesting that since the ovicells of these genera were first described as 'proximal' in position, this monothetic 'character' has been accepted almost without question, and has even been modified (e.g. Harmer, 1957 : 884) in an attempt to include other taxa in the family, a frequent occurrence in bryozoan taxonomy (see Boardman et al, 1969). The explanation of the unusual relationships of the ovicell has awaited first, the demonstration of frontal budding by Banta (1972) and second, the realisation that in both erect and rooted colonies, this type of astogeny may include reversal of zooidal orientation with respect to the direction of colony growth (see Cook & Lagaaij, 1976, and Cook & Chimonides, 1981). The similarities in several characters among the genera assigned to the Lekythoporidae and Celleporinidae suggest a close relationship between the families. It is possible that parallel complexes exist, each containing groups of species with totally different budding patterns, but with similar zooidal characters and even with similar resultant colony forms. Orthoporidroides robusta Moyano (1981 : 182, Figs 1-7), for example, resembles some other, unnamed Recent specimens from the Falkland Islands (BMNH, 1981.3.1.6,9,10) in illustrating how the type of budding pattern found in O. erectus may have been transformed into the type found in the Lekythoporidae. Colonies of both species are large, maximum height 50-60 mm, and profusely branched, with a range of zooid orientations. In Orthoporidroides sp. there are several sequential series of outwardly directed interzooidal, frontally budded zooids. The more centrally placed zooids of branches, which are those budded earlier in the astogenetic sequence, tend to have 'reversed' orientation of orifices, while the more numerous peripheral zooids have a 'normal' orientation. Some intermediate orientations, and ovicells in a 'proximal', lateral and distal topographical position occur in both O. robusta and the unnamed species. This differs from O. robusta in FAMILY LEKYTHOPORIDAE 75 its orifice shape, which has a very long, narrow sinus, and the number of peripheral zooids in its branches. At present it is difficult to determine whether or not astogeny and structure reflect closer genetic links than those of zooidal morphology. In order to trace possible changes in, for example, zooidal orientation and astogeny with time, a range of fossil specimens is necessary. Colonies of Lekythoporidae are robust enough to be preserved in fossil sediments; unfortunately only L. hystrix is unequivocally known to have a fossil record. Lack of records of other species, both in Recent and fossil deposits, is probably caused by difficulties of recognition, due to the small size of the colonies, and rapid thickening of extrazooidal calcification, which obscures the zooidal features. Recent species tend to inhabit the deeper shelf waters, and until more collections from these regions, particularly from high latitudes in the Southern hemisphere become available, detailed analysis of colonies is not possible. Although the characters of the type-species of the five genera of Lekythoporidae are distinct, several of the other species described here show intermediate sets of correlations which make generic assignment somewhat arbitrary. The 'easily recognised' character of the 'proximal ovicell' has to a great extent obscured analysis of other structures in the past. The fragmentary nature of most of the type material requires amplification by well preserved, complete colonies before the range of astogenetic, ontogenetic, polymorphic and microenvironmental variation of the taxa can be analysed and described. Acknowledgements We are grateful to Dr M. V. Hounsome (Manchester Museum), Dr B. Smith (National Museum of Victoria) and Dr S. van der Spoel (Zoological Museum, Amsterdam) for the opportunity to examine specimens, and to Mr P. J. Chimonides (British Museum, Natural History) for discussions and scanning electron microscopy. References Banta, W. C. 1972. The body wall of cheilostome Bryozoa V. Frontal budding in Schizoporella unicornis Jloridana. Mar. Biol. Berlin 14(1): 63-7 1 . Bassler, R. S. 1953. In R. C. Moore. Treatise on Invertebrate Paleontology. Part G. Bryozoa. Lawrence, Kansas. Boardmun, R. S. & Cheetham, A. H. 1973. Degrees of colony dominance in stenolaemate and gymnolaemate Bryozoa. In R. S. Boardman, A. H. Cheetham & W. A. Oliver. Animal Colonies. Stroudsburg. Boardman, R. S., Cheetham, A. H. & Cook, P. L. 1969. Intracolony variation and the genus concept in Bryozoa. Proc. N. Am. Paleont. Conv. 1969, Part C: 294-320. Brown, D. A. 1958. Fossil cheilostomatous Polyzoa from South-West Victoria. Mem. geol. Surv. Viet. 20: 1-90. Busk, G. 1884. 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Morphology and systematics of some rooted cheilostome Bryozoa. J. nat. Hist. 15: 97-134. 76 P. L. COOK & P. J. HAYWARD Cook, P. L. & Lagaaij, R. 1976. Some Tertiary and Recent conescharelliniform Bryozoa. Bull. Br. Mm. not. Hist. (Zool.) 29 (6): 3 1 7-376. Harmer, S. F. 1957. The Polyzoa of the Siboga Expedition. Pt. 4. Cheilostomata Ascophora II. Siboga Exped. 28d: 641-1 147. Haswell, W. A. 1881 (1880). On some Polyzoa from the Queensland coast. Proc. Linn. Soc. N.S.W. 5: 33^4. Hay ward, P. J. 1978. Systematic and morphological studies on some European species of Turbicellepora (Bryozoa, Cheilostomata). J. nat. hist. 12: 55 1-590. 1980. Cheilostomata (Bryozoa) from the South Atlantic, J. nat. Hist. 14: 701-722. d'Hondt, J.-L. 1981. Bryozoaires Cheilostomes bathyaux et abyssaux provenant des campagnes oceanographique americaines (1969-1972) de 1' "Atlantis II", du "Chain" et du "Knorr" (Woods Hole Oceanographic Institution). Bull. Mus. natn. Hist. nat. Paris ser. 4, 3 (Al): 5-71. 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Manuscript accepted for publication 10 October 1982 A new species of Arthroleptis (Anura: Ranidae) from the West Usambara Mountains, Tanzania Alice G. C. Grandison Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD The herpetology of the Usambara Mountains, Tanga Province, Tanzania is primarily based on the extensive collections made by Loveridge (Barbour & Loveridge, 1928). He concen- trated his efforts on the more accessible eastern half of the Usambaras, especially around Amani (506'S: 3837'E) at an elevation of around 923 m and appears to have been the only person who collected amphibians in the Mt. Lutindi outlier (Hindu of Moreau, 1935), the only part of East Usambaras that rises above 1219 m and has a patch of highland and cedar forest on its peak (1 506 m) similar to that which, until comparatively recently, covered a vast area of West Usambaras (Moreau, 1935). Both the East and West Usambaras consist of high mountains with dramatically steep cliffs and ridges but the western peaks and plateaux are all above 1219m, their highest ridges reach 2286 m and the rainfall is considerably less. The eastern and western divisions of the mountains are divided by the hot, dry trench-like Lwengera Valley which is nearly 914m deep. The forests of the Highland Zone of the West Usambaras, as defined by Moreau (1935) in his excellent account of the topography, ecology and climate of these ancient mountains, remained virtually untouched by herpetologists until the last decade when collecting has been carried out by the writer and other zoologists, notably at ca 1 530 m, 37-8 km NE Soni above Mombo at the Mazumbai Forest Reserve and tea plantation, 448'S: 3829'E. This Forest Reserve consists of over 600 acres given to the University of Dar es Salaam by the owner of the tea plantation, Mr John Tanner. The most profitable collecting by the writer was done at 1530 m where an irrigation channel about 1-25 m wide follows the contour from a waterfall and river in forest passing through the forest to emerge in part of the plantation. The stream of approximately 140 mm depth had a slack to moderate current with a fine gravel and mud base. On its right bank it was overhung by forest trees while its left bank had a muddy track of sodden leaves plus debris backed by balsam and wild banana. The forest trees included Ocotea usambarensis and Newtonia. Average annual rainfall measured over a seven year period (1954/60), was 1041 mm. From a stone and debris in the stream and from the stream margin five Arthroleptis were obtained. These together with specimens found later by other collectors, at Mazumbai and additional localities in the West Usambaras between 1450m and 2134m, surpass in adult size all known species of Arthroleptis. In colour pattern and some proportions they resemble A. affinis Ahl, and to a much lesser extent A. adolfifriederici Nieden. A comparison of them with long series of affinis obtained mainly at the type locality, Amani in the East Usambaras, and with the type and paratype of Ruandan adolfifriederici suggests that an undescribed species of Arthroleptis exists in the West Usambara Mts. The new species is named after Mr John Tanner in recognition of the generosity and hospitality afforded the collectors and other zoologists and his keen interest in the natural history of Mazumbai. Abbreviations BM(NH)British Museum (Natural History), London FMNH Field Museum of Natural History, Chicago R Universitetets Zoologiske Museum, Copenhagen ZMB Zoologisches Museum der Humboldt-Universitat Berlin Bull. Br. Mm. nat. Hist. (Zool.) 45 (2): 77-84 Issued 28 July 1 983 77 78 A. G. C. GRANDISON Arthroleptis tanneri sp. nov. Arthroleptis adolftfriederici: Barbour& Loveridge 1928 (part) HOLOTYPE. Adult female BM(NH) 1974.59, collected by A. G. C. Grandison and F. V. Slade at Mazumbai 448'S 3829'E West Usambara Mountains, Tanzania, ca. 1530m elevation, 1000 hrs, 8 October, 1973. PARATYPES. Same locality data as for holotype: BM(NH) 1974.60 (stained and cleared preparation), BM(NH) 1974.61-63, 2245-2300 hrs., 8th October, 1973, BM(NH) 1974.200 16.30 hrs. 1 1th October 1973,m BM(NH) 1982.525-36, R 773 1-35, R 7736-68, R 7797-98. Phillipshof 445'S 3818'E (4 mis N of Lushoto), W. Usambara Mts: MCZ 13166. Shume- Magamba Forest Reserve, 440'S 3815'E, W. Usambara Mts, 2134m elevation, 11-16 February 1981: BM(NH) 1982.537-40. Balangai Reserve, 456'S 3837'E, W. Usambara Mts, 1450 m elevation, May 1981: BM(NH) 1982.541. DIAGNOSIS. A large Arthroleptis related to A. adolfifriederici, separable from that species and all other species of the genus by exceptional size (males 33-7 3-3: females 50-7 mm 4-9) and stout build. Further distinguished from the Usambara A. affinis by its shorter tibia, more widely set nares, slimmer, less pointed terminal phalanges, absence of supernumerary tubercles under the metatarsals, chin spinules but no denticulations along the second and third fingers in the adult male and by minor differences in the temporal colour pattern. DESCRIPTION OF HOLOTYPE. Gravid female 54-0 mm SVL. Head slightly broader than long, its width 44% SVL. Length of snout from anterior border of eye to snout tip 35% of head width. Distance between the nares 11% of SVL. Canthus rostralis sharp, loreal region oblique, lips moderately flared. Interorbital region wider than upper eyelid, subequal to internarial distance. Tympanum clearly visible, the diameter of the tympanic annulus slightly greater than half the internarial distance. Snout rounded in dorsal view. Body stout. Terminal phalanges of fingers bluntly rounded, not or barely wider than distal joint, without circummarginal grooves. Toes long and slender, their terminal phalanges bluntly rounded with circummarginal grooves, the maximum width of the terminal phalanx of the 3rd toe about half the tympanic diameter and only 1/3 wider than the midpoint of the penultimate phalanx. No outer metatarsal tubercle or tarsal fold. Inner metatarsal tubercle broadly oval, its length 1/2 the distance from the tip of the 1st toe to the distal margin of the tubercle. Tibial length less than length of foot (measured from tip of 4th toe to proximal margin of inner metatarsal tubercle) 49% of SVL. No supernumerary tubercles proximal to basal subarticular tubercles of the toes. No toe web. Skin on the head and back smooth with a slight ridge extending along the canthus and lateral edge of the upper eyelid to form a curved dorsolateral line on the back. COLOUR IN ALCOHOL. Dorsum mid brown with a darker rather obscure cruciform area extending from the interorbital area at the level of the middle of the eyelids to the middle of the shoulders, and bordered posteriorly by a broad pale chevron-shaped zone that is of the same intensity of brown as the top of the snout and upper eyelids. Behind the somewhat indistinct chevron is a darker area that reaches to the vent. The tip of the snout has a short light vertical streak. The lores are darker than the top of the snout and a prominent dark brown band passes from the posterior corner of the upper eyelid through the upper third of the tympanum and curves round the posterior rim of the tympanum ending at a point above the arm insertion. The under surfaces of the tarsus and metatarsus are dark brown, except for their outer edges which have a pale line from the heel along the external margin of the 5th toe to its tip. The legs and forelimbs are cross banded dark and lighter brown. The back of the thighs are mottled with pale grey. The middle of the belly is immaculate dirty white, the gular skin and chest pale fawn with creamy white mottling. The lower lip is medium brown, spotted or barred with white. Undersurfaces of the hind limbs dirty white with a faint greyish network. Undersurfaces of toes rather pale, the subarticular and metatarsal tubercles cream. Undersurfaces of hand uniform dirty cream. NEW SPECIES OF ARTHROLEPTIS 79 HABITAT. The holotype and some of the British Museum paratypes were associated with Rana angolensis Bocage and Phrynobatrachus kreffti Boulenger. One adult female was on a stone jutting out of the stream, others in holes in rotten logs or under logs or on leaf litter, while the holotype and two other paratypes were among floating plant debris and twigs at the stream margin. The Copenhagen paratypes were reported as being on the forest floor about 1-6 km from water near to where the forest is replaced by heather on Mt. Sagara above Mazumbai. COLOUR IN LIFE. The general dorsal colouration of the British Museum series was claret-brown with the chain of vertebral markings from the interorbital region to the lumbar region of a darker shade. The hands and feet were pinkish, especially their undersurfaces. Upper half of iris was pale gold. VARIATION IN THE PARATYPES. There is a limited amount of variation in the distinctness of the dorsal pattern, the dark area over the occiput and the pale chevron immediately posterior to it being more clearly defined in younger individuals. Furthermore in the long series of smaller specimens (R 7736-68) some tend to have dark specks at the posterior margin of the chevron and small dark brown spots on flanks and temporal region. The cruciform area is replaced in some of the paratypes by a dark inverted triangular zone from the posterior limit of which two dark oblique bands extend towards the flanks. The undersurface of the head varies from grey mottled white to an almost uniform brown which in a sexually mature adult male has an even darker band laterally. The dorsal skin of the juvenile BM(NH) 1974.200 has small scattered warts. SECONDARY SEXUAL CHARACTERS. As already indicated in the species diagnosis, male tanneri are smaller than females. Only four males are available and all have unpigmented testes. The smallest R7735 (30-0 mm SVL), is the only one that has spinules on the chin, and a dark brown almost black area along the lower jaw medial to the barred lip. No other secondary sexual differences are apparent. All the gravid females have large unpigmented eggs with a maximum diameter of 3-5 mm. COMPARISON. Although A. tanneri attains a larger size than A. affinis Ahl, the taxon described from Amani, East Usambaras and reinstated by Skelton-Bougeois (1961), there is a strong superficial resemblance. However, statistically significant differences at the level P< 0-001 using two sample t-test exist in the following percentage ratios of samples of the two species. A. tanneri A. affinis gravid 9 9 ^99, subadults gravid 9 9 cf^99, subadults Head width n6 n!9 nlO n32 SVL x 44-17 x 43-34 x 42-0 x 42-01 S 1-17 S 1-86 S 1-56 S 1-29 Head width n 6 n 19 n 11 n 32 Tibia length x 88-5 x 84-31 x 77-2 x 75-36 S 5-0 S 6-09 S 2-40 S 3-38 Tibia n 6 n 19 n 13 n 32 SVL x 49-8 x 51-83 x 54-8 x 55-78 S 3-06 S 3-36 S 3-10 S 2-60 Internarial distance n 6 nil SVL x 11-15 x 12-62 S 0-52 S 0-51 80 A. G. C. GRANDISON No statistically significant differences in the percentage ratios of the internarial distance to SVL were found in samples of tanneri and affinis that included halfgrown individuals as well as adults of both sexes. Breeding males of tanneri and affinis may be distinguished by their different secondary sexual characters. None of the sexually mature male affinis has chin spinules but all have a row of denticulations along the inner border of the 2nd and 3rd fingers and the 3rd finger is elongated. Of the two males obtained by Loveridge at Phillipshof West Usambaras and identified by Barbour & Loveridge (1928) at adolfifriederici Nieden, the larger individual (37-4 mm SVL) MCZ 13166 has neither chin spinules nor denticulated fingers but agrees in proportions and in other characteristics with tanneri while the smaller specimen (29- 1 mm SVL ) MCZ 1 3 1 67 is a typical affinis and has finger denticulations. Like all the males and females of affinis from the East Usambaras, both those from the Highland Zone (Mt. Lutindi) and from the Intermediate Forest of Amani, the Phillipshof example MCZ 13167 bears a prominent supernumerary tubercle under the distal portion of the 2nd and 3rd metatarsals and additional but smaller tubercles scattered over the sole of the foot. In some individuals of affinis a small tubercle is also present at the base of the first toe between the inner metatarsal tubercle and the subarticular tubercle (Fig. 2). Ahl (1939) in his type descriptions of affinis and of the junior synonym A. schoenbecki, both of which were obtained at Amani, mentions the supernumerary tubercles which bedeck the soles of the feet in the type specimens but makes no reference to the more distal tubercles under the 2nd and 3rd metatarsals. Examination of the type material in the present study has confirmed their presence. Supernumerary tubercles under the metatarsals are absent from every individual of tanneri, as well as from the holotype and paratype of A. adolfifriederici from Rugege and Bugoya Forests Ruanda (Fig. 2). A third example also collected at Rugege and referred to adolfifriederici by Nieden (1912) also lacks these tubercles. However adolfifriederici differs from A. tanneri in its smaller size, the adult female holotype and paratype being 41-1 mm Fig. 1 The Usambara Mountains, NE Tanzania. Land over 1000m is indicated by cross hatching. NEW SPECIES OF ARTHROLEPTIS 81 Dorsal view and undersurface of the foot of: Arthroleptis tanneri sp. nov. (A & C); Arthroleptis af finis (B & D). and 43-8 mm SVL, as well as in its marbled dorsal pattern (Nieden, 1912) and further differs from both tanneri and affinis in its much narrower head and shorter internarial distance and tibia. Percentage ratios in the holotype and paratype of adolfifriederici of head width/SVL are 35-0 and 33-1, of internarial distance/SVL 10-70 and 10-27, of head width/tibia length 61-0 and 55-34. A. tanneri seems to be further distinguished from either 82 A. G. C. GRANDISON adolfifriederici or affinis by its terminal phalanges being bluntly rounded and barely wider than the distal joint. In the Ruanda adolfifriederici and in the East Usambara series of affinis they tend to be sharply pointed and are decidedly wider than the proximal phalanges and joints. In colour pattern tanneri and affinis are rather similar but whereas in tanneri the ventral surfaces of the toes are cream, in affinis the phalanges and the soles of the feet are dark brown and the cream subarticular tubercles are in sharp contrast. Also in affinis the supratympanic stripe generally curves round the top of the tympanic annulus and does not pass across the tympanic membrane. Osteology The stained and cleared adult female paratype of tanneri was compared with a similarly prepared Amani adult of affinis. Osteological differences that are interpreted as being of taxonomic significance are limited to the skull, in particular the shape, extent of the sphenethmoid including the degree of its dorsal exposure, the medial separation of the nasals and the shape of the anterior borders of the frontoparietals. In tanneri the sphenethmoid is a short squat bone, sharply concave posteriorly and with a straight anterior border that fails to reach the palatines except for their medial posterior tips. It has no dorsal exposure, being completely covered by the frontoparietals. The sphenethmoid in affinis not only projects much farther forward to extend well beyond the palatines and to the level of the anterior edges of the choanae, but its anterior and posterior borders are of a different shape from those of tanneri, being deeply convex and straight respectively. Furthermore the sphenethmoid has a crescentic dorsal exposure. In both taxa the palatines are robust and contact the maxillae broadly but fail to reach the pterygoids. Not only are the medial borders of the nasals more convex and wider apart in affinis but there is a greater separation between the nasals and the frontoparietals; moreover the anterior borders of the frontoparietals project forwards instead of as in tanneri being straight. It is considered unlikely that these differences are attributable to individual variation in view of the author having discovered in an extensive analysis of bufonid skulls that there is virtually no intraspecific variation in adults in the shape and size of the sphenethmoid, palatines, nasals and frontoparietals. Distribution of A. tanneri and its relationship with /. affinis In the eastern half of the Usambara Mts A. affinis is known from Amani, 923 m, and from 1219 m on Mt. Lutindi (Hundu). In the neighbourhood of Amani during the dry season the writer found affinis to be abundant among leaf litter, in cracks in the impacted earth of road cuttings and in mounds of earth, sometimes in association with Callulina kreffti Nieden and Nectophrynoides tornieri Roux. Loveridge obtained 192 examples from leaf strewn paths at Amani (Barbour & Loveridge, 1928). At comparable elevations in the western half of the Usambaras no herpetological fieldwork has been undertaken below 1219m and only one Arthroleptis has been taken at 1219m, on the Ambangulu Estate, Korogwe District, 505'S, 3826'E. The single individual, a juvenile of 20-0 mm displays features of both tanneri and affinis. It resembles affinis in having supernumerary tubercles under its metatarsals but its proportions correspond more closely to those of tanneri, its percentage ratios being Head width/SVL 45-0, Head width/Tibia 90-0 and Tibia/SVL 50-0. The existence of this aberrant individual raises the question of whether tanneri and affinis coexist at Ambangulu and whether the 'mixed' characters of this juvenile can be attributed to hybridisation. Further material from this and other localities at lower elevations in the West Usambara Mts is awaited with interest. Evidence of sympatry of A. tanneri and A. affinis rests on the two adult males collected by NEW SPECIES OF ARTHROLEPTIS 83 Loveridge in 1926 during ten days spent at Phillipshof in the West Usambaras. Barbour & Loveridge (1928) quote the elevation of Phillipshof as approximately 5500ft (1676m), which accords with information given to the author by the Royal Geographical Society based on the 1916 War Office maps of Tanganyika Territory and the Lushoto sheet 1 : 250 000 which place Phillipshof four miles north of Lushoto, 445'S 3818'E, elevation 1524-1829 m. Barbour & Loveridge (1928) describe Phillipshof as 'rolling downs of grazing land, marshy swamps and slow-flowing streams in the bottoms, scattered patches of rain-forest on the uplands, with vast stretches of forest nearby on either side of the Malindi road'. According to Moreau's (1935) zonation of the Usambaras Phillipshof lies in the same Highland Zone as Mazumbai. It is not known whether the Phillipshof examples of tanneri and affinis were found in similar niches and closely associated with each other or whether they occupied quite different biotopes; no precise field data accompany the specimens. Except for the Phillipshof record of sympatry and the unique characters of the Ambangulu juvenile the available material points to tanneri and affinis being separated geographically and by altitude with A. tanneri confined to the West Usambara Mts in the Highland forest zone of Moreau (1935) and A. affinis occurring at the lower elevations of the Intermediate forest zone of the East Usambara Mts. Additional material used in study Arthroleptis adolfifriederici Nieden Bugoya Forest, Ruanda: ZMB 2 1 787 (Paratype). Rugege Forest, Ruanda: ZMB 25287, FMNH 73836 (ZMB 2 1 789) (Holotype). Arthroleptis affinis Ahl Amani 506 ' S 3837 ' E, E. Usambara Mts, Tanzania 923 m: BM(NH) 1 974. 1 77 (stained and cleared preparation), BM(NH) 1974.170-176, BM(NH) 1974.178-185, BM(NH) 1974.186-195, MCZ 88131-278, MCZ 13153-60, FMNH 73836, ZMB 25289 (holotype of A. schoenebecki Ahl), ZMB 23093 (holotype of A. affinis), R 7797-98, R 77 1 00. Mt. Lutindi, E. Usambara Mts, Tanzania: MCZ 13161-65. Phillipshof, W. Usambara Mts, Tanzania: MCZ 13167. Acknowledgements Most cordial thanks are extended to Dr K. M. Howell, University of Dares Salaam and to Dr A. Schiotz, Director, Danmarks Akvarium, Copenhagen for making available collections from the Usambaras and generously allowing me to describe the new species. Mr H. Marx (Field Museum of Natural History, Chicago), Dr G. Peters (Zoologisches Museum Humboldt-Univ. Berlin) and Dr E. E. Williams (Museum of Comparative Zoology, Harvard) also kindly lent material in their care. Dr I. Walker (Imperial College of Science, London) and Dr N. Jago (Centre for Overseas Pest Research, London) helped in formulating plans for my Tanzanian visit. Dr R. Laurent (Institute Miguel Lillo, Tucuman) and my colleagues Dr G. Boxshall and Dr M. Hills gave freely of their time in discussing morphometric analyses, Dr Hills being especially helpful with the statistical interpretation of data. I wish also to record my thanks to members of the British Council, Staff of the Zoology Department, University of Dar es Salaam, and the Tanzanian Government for facilitating my research. It is difficult to express adequately my appreciation of Mr John Tanner's kindness, generosity and hospitality towards members of the team while staying at his home in 84 A. G. C. GRANDISON Mazumbai. He ensured that our visit was scientifically worthwhile and enjoyable, and I thank him most sincerely. Dr S. N. Stuart's help with field data and literature is also gratefully acknowledged. Field work was supported financially by the Trustees of the British Museum (Natural History). References Ahl, E. 1939. Beschreibung neuer afrikanischer Frosche der Gattung Arthroleptis. Sber. Ges. naturf. FreundeBeri 303-310. Barbour, T. & Loveridge, A. 1928. A comparative study of the herpetological faunae of the Uluguru and Usambara Mountains, Tanganyika Territory with descriptions of new species. Mem. Mus. comp. Zool. Harv.. 50: 87-265. Moreau, R. E. 1935. A synecological study of Usambara, Tanganyika Territory, with particular reference to birds. J. Ecol. 23: 1-43. Nieden, F. 1910. Verzeichnis der bei Amani in Deutschostafrika vorkommen den Reptilien und Amphibien. Sber. Ges. naturf. Freunde Be rl: 44 1-4 52. Nieden, F. 1912. Amphibia. Wiss. Ergebn. dt. Zent Afr. Exped. 4: 165-195. Skelton-Bourgeois, M. 1961. Reptiles et batraciens d'Afrique orientale. Rev. Zool. Bot. afr. 63: 309-338. Manuscript accepted for publication 15 July 1982 The distribution, behavioural ecology and breeding strategy of the Pygmy Toad, Mertensophryne micranotis (Lov.) Alice G. C. Grandison Zoology Department, British Museum (Natural History), Cromwell Road, London SW7 5BD Sanda Ashe P.O. Box 3, Watamu, Kenya Introduction Sexually mature males of the East African pygmy toad, Mertensophryne micranotis (Loveridge) have been reported as having series of small spines around the rim of the cloaca and at the entrance to the cloacal tube (Grandison, 1980). Development of these spines correlates with development of the cluster of heavily keratinised thumb spines and their function was interpreted as an adaptation for holding the vents of the male and female securely together during mating to ensure direct transfer of sperm to the small clutch of large eggs produced by the female. Spines in the area of an anuran's vent were first reported by Anderson (1871). He described a large zone of curved cornified papillae in Rana gammi Anderson, a species currently synonymised with Rana (Paa) sikkimensis Jerdon. Dubois (1976) figured the spines in sikkimensis and suggested that cornified spinules occur also in Rana (Paa) delacouri Angel. The life histories of neither of these Nepalese species is known. In order to test the theory that had been postulated of internal fertilisation in Mertensophryne micranotis fieldwork and a captive breeding programme were recently undertaken in Kenya. New information on the species' occurrence in Kenya, its habits and breeding behaviour is presented here. Range and ecological preferences M. micranotis is known to occur in Tanzania, Zanzibar and Kenya. Few examples have been collected, perhaps because such a small-sized toad (sexual maturity is attained at 16 mm in males) is difficult to detect in leaf litter. Although the type and paratype found at Kilosa, Tanzania and a single individual from the Uluguru Mts. were said by Loveridge (1925) and Barbour & Loveridge (1928) respectively not to have been found in forest, else- where the species seems to be restricted to forest or recently cut down forest. In Kenya the species occurs both along the coast from Gede in the north (31 1 'S: 40 1 'E) to Shimoni in the south (438'S: 3923'E), in the Shimba Hills National Park in the Coast Range (416'S: 3922'E) and Mrima Hill (429'S: 3916'E). Until comparatively recently the Coast Province of Kenya was a near continuous forested belt overlying the approximately three kilometre wide Pleistocene coral reef and the Pliocene and Pleistocene sandy soils adjacent to it farther inland. Today little true forest remains in the Coast Province and very few remnants extend to more than 50 acres. Agriculture, notably plantations of sugar, coconut, cashew nut, maize and sisal combined with the rapid growth of Kenya's tourist industry have taken their toll of the coastal forests. Two of the small remaining pockets are contained in the Bull. Br. Mus. nat. Hist. (Zool.) 45 (2): 85-93 Issued 28 July 1 983 85 86 A. G. C. GRANDISON & S. ASHE 3 9 Fig. 1 Known distribution of Mertensophryne micranotis (Loveridge) in Kenya. Records indicated by an inverted triangle. MER TENSOPHR YNE MICRA NO TIS 87 National Monuments of Gede and Mtwana, in which are preserved the ruins of thirteenth century Arab-African towns. Diani forest (sometimes also referred to as Jadini forest) con- sists of 50 acres of largely untouched forest similar in composition to Gede forest; it is privately owned. Other small forest relics encircle cleared areas that until the nineteenth century were settlements, each housing up to 2000 of the Mijikenda peoples but which today contain only a few houses occupied by elders of the clans and graves of their ancestors. Most of these kayas, as these sacred villages are called, were situated along the ridge of the Coast Range from the Shimba Hills northwards to Kilifi and at vantage points to resist incursions from the north by the Galla farmers. By the seventeenth century the most southerly of the Mijikenda, the Digo, vacated their Shimba kaya and established minor kayas along the coastal plain (Spear, 1978). At one such sub-kaya, Kinondo, and at the Kombeni kaya M. micranotis has been found. On account of their religious significance and present day use for rituals and processions the kaya forests are provided with a measure of protection which may safeguard their populations of micranotis. In the southern extremity of the Coast Province two densely forested hills, Jombo and Mrima, jut up spectacularly from the surrounding flat plains (Britton et al., 1980). No herpetological survey has apparently ever been made of Jombo (476 m) but the senior author made two short visits to nearby Mrima which rises to 299 m. The very deeply weathered volcanic plug of Mrima has been prospected since the early 1950s for its rich deposits of manganese and niobium. Its rocks weather extremely rapidly and up to 236 m of terra rosa cover the carbonatite plug. It is the weathered material that has been mined and the abandoned and completely unprotected mineshafts are a serious hazard to contend with when scouring the upper reaches of the forest for the anuran denizens of the leaf litter. A circular, flat-topped pile of moist leaves which contained duiker droppings and may have been a communal duiker latrine on to which some animals, perhaps ants, had piled leaves and other debris yielded juvenile M. micranotis. The pile was on a trail and was the only damp spot found; no rain appeared to have fallen for some considerable time and no water- filled treeholes were evident. On the bank of a path and among small pieces of weathered carbonatite and terra rosa and harmonising perfectly in colouration another juvenile micranotis was obtained. In these forests there are no surface ponds or streams and even after heavy and prolonged rain the water probably rapidly disappears through the red earth and litter that overly and form pockets in the labyrinth of holes in the coral rag of the narrow coastal strip. On the sandy soils farther inland the rains likewise drain rapidly. The only standing water that becomes available with the highly seasonal rains are ephemeral reservoirs in treeholes formed by broken off limbs, in fissures in tree buttresses and in live and fallen tree trunks, as well as in abandoned shells of the large land snail, Achatina, and in the occasional discarded can on the forest floor. Consequently the only anurans that can survive in such forests need to have a highly specialised breeding strategy, such as direct development that completely omits the larval stage or one that produces a terrestrial larva, or need to be opportunistic breeders capable of taking advantage of the seasonal water pockets for egg laying sites. As the water holes are small and subject to rapid desiccation small clutches capable of an accelerated rate of development would be to the species' advantage. The searches made of the forest litter, of trees and shrubs in these Kenyan forests have revealed only three species of Anura, two of them arthroleptines a group known to develop directly from eggs laid in underground chambers to fully transformed miniatures of the adult frogs. The third, Mertensophryne micranotis, has been reported as breeding in water-filled treeholes and land snail shells and has an unusual-shaped tadpole with an angled head that is surrounded by a raised ring of tissue (Grandison, 1980). The anuran composition of a water- less forest in SE Tanzania is recorded by Loveridge (1942). The area is Nchingida 108'S, 3912'E on the Rondo Plateau which is the type locality of Mertensophryne micranotis rondoensis (Loveridge), a form that Loveridge distinguished from the nominate on its less heavily pigmented throat. Loveridge (1944) described the Rondo Plateau as waterless where even the heaviest rainstorms drain rapidly through the sandy soils. In addition to the M. m. 88 A. G. C. GRANDISON & S. ASHE rondoensis he recorded (1942, 1944) four other species of Anura two arthroleptines and two microhylids, Breviceps mossambicus Peters and Spelaeophryne methneri Ahl. While the life history of Breviceps is known to be entirely independent of water (Wager, 1965), very large eggs being laid in burrows often far from water and the tadpole stage passed within the egg capsule, no aspect of the breeding cycle of Spelaeophryne has been recorded. The right ovary removed from an adult female S. methneri in the BM collections contains 1 8 large ova, average diameter 2 mm, and a few much smaller ova: all are unpigmented. Such a small clutch of unpigmented eggs suggests a specialised reproductive mode that has freed the species from an aquatic environment, although since the species lacks any obvious means of burrowing into the litter, unlike Arthroleptis stenodactylus Pfeffer and Breviceps with their well developed shovel-shaped metatarsal tubercles, it seems unlikely that Spelaeophryne is a burrowing breeder. Perhaps like Nectophrynoides malcolmi Grandison (1978) it has a terrestrial tadpole with a vascularised tail that serves as a respiratory organ, but perhaps like Mertensophryne micranotis it depends on waterfilled holes in which to lay its small complement of eggs. The senior author's field study was conducted during the month of May. Mean monthly rainfall records had suggested that this period would be the most likely to coincide with the long rains and consequently produce water-filled breeding sites and the likelihood of witness- ing breeding behaviour in the species. However the rains were capricious in May 198 1 and at Diani Beach in the southern coastal belt only 19 cm fell during the entire month and for half the period there was no rain or only a few drops (C. Harcourt, pers. comm.); as a result the number of water-filled treeholes and land snail shells was exceedingly small and no mating pairs were found. The standing water in all treeholes found was siphoned and checked for the presence of anuran eggs and tadpoles but only one sample contained micranotis tadpoles; it was in one of three holes in the buttress formation of a tree that branched near its base into three trunks and was the only one obscured by a green leafy liana which partly shaded it from the sun. The diameter of the tree trunk at the level of the hole was 54 cm. The other two waterfilled holes in the same tree were at a similar height from the forest floor and although their sizes fall within the range of variation of other oviposition sites for micranotis the holes contained only insect larvae (Table 1). All the tadpoles, in excess of 45, had developed limbs but still had larval mouthparts. The pH of the water was not taken and the identity of the tree is not known. The mammalogist, Dr G. Rathbun, reports (pers. comm.) that while studying elephant shrews and cutting forest trails at Gede he was attracted by a faint squeak to a hole in a tree Table 1 Records of water-filled treeholes occupied by Mertensophryne micranotis. From data supplied by G. Rathbun (t) and L. P. Lounibos (*) Height from Volume of Larvae (L) forest floor Diameter Depth water or Location Date (cm) (cm) (cm) (cc) Toads (T) t Gede 18.V.71 150-0 10-16 Full T(cf and 9) * Kombeni 16.V.75 700 L 16.V.75 230 L 24.iv.76 90 L 24.iv.76 30-48 3-8x5-1 5-08 55 L 27.iv.76 101-60 3-8x7-6 7-62 20 L 7.V.77 60 T (in amplexus) * Makadara ll.iv.76 17-78 2-5x2-5 6-35 70 L ll.iv.76 96-52 2-5x2-5 6-35 25 L 19.iv.76 121-92 3-8x3-8 7-37 45 L Shimoni 12.V.81 20-32 2-5x3-8 10-16 Full L MER TENSOPHR YNE MICRANO TIS 89 7-6 cm diameter which he tentatively identified as Lecaniodiscus fraxinifolius. The hole was in a stump formed by a fallen limb 1-5 m from the forest floor. It contained a pair of sexually mature M. micranotis which suggests that the hole had been selected as a potential breeding site. Dr L. P. Lounibos (pers. comm.) provides the only other known records of the occurrence of micranotis in water-filled treeholes. His records and details of the Gede and Shimoni treeholes are given in Table 1 . Dr Lounibos' observations were made in the course of his research into mosquito habitat segregation in Makadara Forest Shimba Hills and Kombeni Forest, Rabai Location (355'S: 3934'E). Samples of his Makadara Forest tadpoles and adults were identified by A. McKay, National Museums, Kenya and the identity of a sexually mature male obtained at Kinondo Forest was confirmed by the senior author. It is assumed that his Kombeni tadpoles and toads were also correctly identified. Lounibos (1981) should be consulted for rainfall records and seasonality of water in treeholes in his study areas. Movements, colour change and territorial behaviour Skulking movements and exceptionally effective camouflage are striking features of M. micranotis. Adults and subadults when disturbed on the forest floor tend to remain motion- less or to very slowly back under a dead leaf or twig or disappear down a hole. A gravid female unearthed from loose soil at the base of a tree flattened its body and remained inert for some time after capture and to the extent that it resembled a dead leaf. The disruptive dorsal colour pattern of shades of brown closely matches the leaf litter in which micranotis is usually found and makes the animal exceedingly difficult to detect. The darkest areas of the body are invariably the lateral band and the interocular bar. A pale hair- like vertebral line of variable length, but usually extending from behind the eyes to the sacrum is present in most specimens. It is sometimes flanked by an irregular dark brown band that divides in the sacral region to form a large ring enclosing a pale zone which extends posteriorly to a brilliant white supra-anal triangle. This white triangle is characteristic of the species. The dark vertebral band and ring are separated from the even darker lateral band by a broad lighter area which is particularly subject to variation in colour according to the animal's background. For instance, in an adult male transported on white plastic foam this dorsolateral area was a very pale grey but after a day spent on dead leaves, twigs and bark the entire dorsum changed to dark brown, while two days later the pale grey areas changed to khaki. Both in its dark and khaki phases the toad was barely discernible among the litter. Recently transformed juveniles and halfgrown individuals are usually black when they emerge from their hiding places but may acquire a greenish-brown tinge when more active. Ventrally, both adults and young have prominent dark blotches on a white background. Males tend to have a brighter, more contrasting colour pattern than females. Observations on captive individuals suggest that the species is diurnal and particularly active in the morning, also that it is territorial, with individuals having their own burrows in the leaf litter. In the wild it is likely that members of the species occurring in forest overlying coral rag will use as retreats the abundant holes in the coral as well as burrows in soil and leaf litter. After active foraging the toads usually back down into their burrows. Adults are more secretive than juveniles and were not seen to make their presence obvious by hopping or other rapid move- ments; their movements were exceedingly slow, laborious and consisted of usually no more than five steps at a time followed by a long pause. On the other hand, juveniles tended to scramble and hop but such activity was interspersed with long periods of immobility. The junior author noticed the species' propensity while in captivity to climb. Day old toadlets climbed with apparent ease the glass sides of a casserole in which they were reared. Generally they moved hand over hand with the body raised but when climbing clean glass they lowered their bodies. Adults not only readily climbed the plants in the terrarium, occasionally roosting on leaves and crawling along narrow stems but they persistently climbed the glass sides of the rectangular tank usually by straddling a corner of the terrarium 90 A. G. C. GRANDISON & S. ASHE and bracing their limbs. As they climbed the glass it was noticed that they left behind a trail of liquid and that the posterior part of the toad's abdomen was flattened on the substrate. It is believed that the liquid is expelled from the toad's cloaca and that the surface tension set up facilitates progression over a smooth surface. By developing a technique that allows a short and spindly legged species to climb smooth vertical surfaces the availability and variety of treehole oviposition sites may also be increased thus enhancing the chances of survival. Captive breeding programme One subadult and eleven juveniles collected between 29 June and 1 1 July 1981 were reared in captivity to sexual maturity. They were collected from an area of less than an acre in the vicinity of the junior author's house at Watamu. The area once covered by primary forest and an extension of the Gede Forest was cleared of all but the large trees for house building in 1980. The toads were found on and among the leaf litter overlying coral rag in or at the edge of a surveyor's cut line as well as in a large termite-ridden rotten log in the undergrowth. The terrarium consisted of a glass tank 40x28x22-5 cm furnished with a 2-5 cm layer of leaf litter and sand, growing plants and logs. A coconut shell with a little rain water and a plastic jar 12 cm deep and a 6 cm diameter, filled with rainwater and tilted at an angle of 45 against a twisted root provided potential oviposition sites. The root acted as a climbing frame and provided numerous hiding places. A wooden ramp in the jar of water formed an exit. By the 22 October, five days after the rains had started the day temperature exceeded 30C and night time temperature 25C and there was high humidity, the first batch of eggs was laid but neither mating nor egg laying was witnessed. Several toads appeared gravid. The following accounts are condensed from the detailed notes made by the junior author on her observations on the mating behaviour of her captive specimens. Courtship It remains uncertain whether females approach individual males and what signals are used to initiate courtship because pairs were already in amplexus on each occasion when observations were begun. However it was noticed that the soft chirp of a male in amplexus prompted three other males that had been chasing each other around in the water container to call and mount each other indiscriminately and their calls appeared to induce two females to move in the direction of their calls but stop and wander off when the calling ceased. The behaviour of the three males in the jar suggests that the normal calling-station may be a water-filled container and that the receptive female moves in that direction when stimulated by the advertisement call. However the mating behaviour of only six pairs has been observed and while one pair mated in water the other pairs mated mostly out of water. Visual as well as auditory cues may contribute to mate selection in M. micranotis for it was noted that the three calling males in the water jar were of a brighter colour and more con- trasting pattern than the amplectic male which was very dark coloured. Amplexus and fertilisation The mating of one pair, which spanned a period of over eight hours, is described in detail. When the male first mounted the female, their eyes were in the same vertical plane while his hind legs trailed on the ground. The amplectic position was axillary. As he drew up his legs he stimulated the female's sacroiliacal region by drumming it with his long fourth toe then placed his feet over her tibiotarsal joints. Half an hour later he uttered an almost inaudible rapid ticking sound which the female answered with a soft chirp. The pair were clamped closely to each other from snout to vent but although their white supraanal patches were vertically in line a distance of about 2 mm separated them. After an interval of about a few MER TENSOPHR YNE MICRA NO TIS 9 1 minutes the female alternately inflated and deflated her body, as if sighing, while his feet were placed on her flanks. As her body deflated at 30 second intervals the male exerted a downward squeeze with his cloacal region. Despite the female moving away and trying to dislodge the male by scratching him first with her hind leg then with a front leg, the male remained firmly attached, although at one point he was shifted sideways as she forced her way under a piece of bark. Mating movements were discontinued while the female fed on white ants but were resumed approximately two hours after observations on the amplectic pair were begun. The male resumed tactile stimulation of the female by drumming his fourth toe. His heels which were then placed on either side of and slightly above her white supraanal patch seemed to channel the drop of clear liquid that trickled from his cloaca down to hers where it appeared to be absorbed. As the female adopted a more upright stance the male moved farther back and lower down and his mating movements became more forceful. As he engaged in a series of thrusts his white supraanal patch was seen to curl inwards towards that of the female. During the ensuing two hours mating movements continued intermittently, with a rest period while the female, still with male attached, moved to a hiding place in the litter but when the amplectic male produced a rapid, very faint ticking sound another male called from 25 cm away in the water jar; the vocalisation elicited a marked increase in activity by all the other males in the terrarium which chased and mounted each other indiscriminately while their calls became louder until each call sounded like a ten note chirp which could be heard 2-5 m away. Several females were active on the floor of the terrarium but although one of the calling males attempted to mount a female it did not persist when the female moved away. When the amplectic pair emerged from the hiding place more vigorous mating movements occurred, despite the female's attempts on two occasions to dislodge the male, and he thrust twice each time the female deflated her body. Although both sexes were visibly vibrating no call from the female was audible but the male emitted a ticking sound. During the next hour sporadic mating movements occurred but his vent was above hers and although the pair entered the coconut shell partly filled with water and moved round the slope in a clockwise direction only her foot entered the water and the pair climbed out of the shell. The female again tried to dislodge the male while she ate insects. Observation was discontinued for ten minutes during which time the pair disappeared. In five other pairs in which mating was witnessed the duration of amplexus varied from five and a half hours to ten hours. It was noticed that the male's grip on the female's vent is so tight that his vent drags upwards the skin surrounding her vent. One pair found in the water jar in the 'normal' amplectic position were joined by a second male which clung to the female's axillae in an inverted, belly to belly position. Both males engaged in mating move- ments but two hours later when the dorsal male had disappeared the protruding cloacal region of the underslung male was seen to curl upwards to contact that of the female and the moisture he expelled from his vent as he thrust appeared to immediately be absorbed by the female. Belly to belly amplexus is known to occur also in the internally fertilised Ethiopian bufonid, Nectophrynoides malcolmi Grandison (Grandison, 1978 and Wake, 1980). Although clutches of eggs were later found in the jar of water in the terrarium, egg laying by mated females had not been witnessed and because males were present the possibility of the eggs having been externally fertilised could not be ruled out. So on the next occasion when a pair began to mate they were removed to another terrarium containing a water bowl of similar capacity and quantity of water. When mating was completed and the male had dismounted he was removed and returned to his original terrarium. Mating had taken place in water. Six hours after mating the female, still in the water, began laying eggs, two strings emerging together. Three and a half hours from the start of egg laying and at 2300 hours oviposition had not been completed, but the following morning two eggs strings containing a total of 22 eggs were found attached to the rim of the jar. The female had returned to the floor of the terrarium. The eggs proved to be fertile and they developed at a similar rate to those in previous clutches. Each of the six eggs in one string that was severed and preserved within forty minutes of being laid was found to be at the late cleavage stage of development, but eggs 92 A. G. C. GRANDISON & S. ASHE from another clutch where the time of laying was unknown were at early to late gastrula. The number of eggs in five clutches were 17, 19,22,32, 32. The aforegoing description emphasises the close contact of the cloacae during mating in Mertensophryne micranotis and the tenacious grip that the male exerts on the female's vent while he engages in downward thrusts and his feet are placed on each side of the female's vent. While it can be assumed that the male's cloacal spines play a significant part in the coupling and that internal fertilisation of the eggs takes place it is still not known whether the spines interlock in the furrows of the female's vent. Function of the tadpole's head 'crown' Broadley (in Channing, 1978) suggested that the function of the head 'crown' in tadpoles of Stephopaedes anotis (Boulenger) might be to exclude from the eyes and nostrils scum accumulating on the water surface. The angled head and ring of raised tissue ('crown') of the M. micranotis tadpole is closely similar to that of S. anotis (Grandison, 1980) but observations on tadpoles of M. micranotis in the wild and in captivity suggest that the angled head with the raised ring of tissue surrounding a saucer-like depression in which lie the eyes and nostrils is a simple adaptation for suspending the tadpole at the meniscus, where the highest concentration of oxygenated water is available in the small pocket of stagnant water that is selected as a breeding site. With the 'crown' breaking the water surface the tadpole has access to oxygen, both during its gill breathing stages and later when it has acquired lungs, at the same time maintaining itself in a tail-down position with the minimum expenditure of energy. In the field study it was noticed that each time the sun struck the water surface of the treehole the tadpoles tended to rise quickly and suspend themselves at the meniscus, particularly around the rim of the hole so that their angled heads were parallel to the contour of the meniscus whilst the body and tail of the tadpole hung down vertically. In such a position the ventral mouthparts had access to the algal growth lining the rim of the cavity. When the head 'crown', which is at an angle of 45 to the body and tail, breaks the surface tension at or near the centre of the hole where the meniscus is virtually horizontal the body and tail of the tadpole are suspended parallel to the angle of the head. Diet The contents of the stomachs of three subadults that were known to have been preserved immediately after capture were analysed. Ants constituted the major food element but mites were found to have been almost equally favoured, particularly Linopodes (Eupodidae). Although nine other genera of mites were present only one or two examples of each were identified (Eupelops, Trachygalumna, Pilizetes, Liodes, Eremaezetes, Scapheremaeus, Bdella and Spinibdelld). Beetles and thrips of kinds typical of the litter/subcortical layer were also represented in the analysis, as well as Collembola (Symphypleona) and the macerated remains of spiders and fly larvae. The only trace of termites was a crumpled set of wings of Termitidae. In captivity the toads thrived on a diet of small white ants, termites and aphids. Acknowledgements This study was supported by the Fauna and Flora Preservation Society and the British Museum (Natural History). Field work was made possible through the assistance of the Office of the President of Kenya and the cooperation of Mr A. D. McKay, National Museums of Kenya. The senior author thanks Drs L. P. Lounibos and G. Rathbun for making available their observations on treehole occupancy by M. micranotis; her colleagues Dr W. Sands and Miss A. Baker for analyses of stomach contents; Miss C. Harcourt for field MER TENSOPHR YNE M1CRA NO TfS 93 assistance and rainfall records; and Mr and Mrs M. McKay and Mrs D. Webb for kind hospitality. The assistance given by Dr R. C. Drewes, Mr S. Reilly and Mr E. Wederkinch in making material and field data available is also gratefully acknowledged, as is Professor Ruth Bellairs' cooperation. References Anderson, J. 1871. A list of the reptilian accession to the Indian Museum, from 1865 to 1870, with a description of some new species. J. Asiat. Soc. Beng. 40: 12-39. Barbour, T. & Loveridge, A. 1928. A comparative study of the herpetological faunae of the Uluguru and Usambara Mountains, Tanganyika Territory with descriptions of new species. Mem. Mm. comp. Zool. Harv. 50: 87-265. Britton, P. L., Britton, H. A. & Coverdale, M. A. C. 1980. The avifauna of Mrima Hill, South Kenya Coast. Scopus 4: 73-78. Channing, A. 1978. A new bufonid genus (Amphibia: Anura) from Rhodesia. Herpetologica 34: 394-397. Dubois, A. 1976. Les grenouilles du sous-genre Paa du Nepal (famille Ranidae genre Rand). Cahiers nepalais. Documents no 6. C.N.R.S. vi + 275 pp. Paris. Grandison, A. G. C. 1978. The occurrence of Nectophrynoides (Anura Bufonidae) in Ethiopia. A new concept of the genus with a description of a new species. Monit. Zool. ital. N.S. Suppl. XI: 119-172. 1980. Aspects of breeding morphology in Mertensophryne micranotis (Anura: Bufonidae): secondary sexual characters, eggs and tadpole. Bull. Br. Mus. nat. Hist. (Zool.) 39: 299-304. Lounibos, L. P. 1981. Habitat segregation among African treehole mosquitoes. Ecol. Entom. 6: 129-154. Loveridge, A. 1925. Notes on East African Batrachians, collected 1920-1923, with the description of four new species. Proc. zool. Soc. Land.: 763-79 1 . 1942. Scientific results of a fourth expedition to forested areas in East and Central Africa. V. Amphibians. Bull. Mus. comp. Zool. Harv. 91: 377-436. 1944. Scientific results of a fourth expedition to forested areas in East and Central Africa. VI. Itinerary and comments. Bull. Mus. Comp. Zool. Harv. 94: 191-214. Spear, T. T. 1978. The Kava Complex. A history of the Mijikenda peoples of the Kenya Coast to 1900. xxiv+ 172 pp. Nairobi. Wager, V. A. 1965. The frogs of South Africa. 242 pp. Cape Town and Johannesburg. Wake, M. 1980. The reproductive biology of Nectophrynoides malcolmi (Amphibia: Bufonidae), with comments on the evolution of reproductive modes in the genus Nectophrynoides. Copeia no. 2: 193-200. Manuscript accepted for publication 3 August 1982 Additional notes on bariliine cyprinid fishes Gordon Howes Department of Zoology, British Museum (Natural History), Cromwell Road, London SW75BD This addendum serves to correct and amplify statements and observations published in an earlier study on the anatomy, phylogeny and classification of bariliine cyprinid fishes (Howes, 1980). The generic status of Barilius lujae Boulenger, 1909 In the synoptic account of Barilius species given in Howes (1980), Barilius lujae Blgr, 1909 was omitted. Examination of syntypes (BMNH 1908.11.26:8; 1909.4.26: 18), together with a series of more recently collected specimens (BMNH 1975.6.20:429^438; 1976.6.20 : 407^413, including alizarin preparations), makes it clear that the species lujae should be assigned to the genus Leptocypris. Synapomorphies characterising Leptocypris (modified from Howes, 1980) are: shallow lower jaw; truncated lateral ethmoid; absence of, or reduced intermandibularis muscle; absence of, or few gill-rakers; elongated pectoral and pelvic axial scales, and overlap of the antero-dorsal portion of the 2nd infraorbital by the posterior border of the 1st. The species lujae has all these characters and, in addition, a type of ethmoid architecture which supports the hypothesis that Leptocypris and Engraulicypris are closely related genera (Howes, 1980: 182). In Leptocypris lujae the medial ethmoid notch is horseshoe-shaped as in Engraulicypris sardella (cf. Fig. 1 here with fig. 9 in Howes, 1980). In Engraulicypris the ethmoid indentation coincides with a foramen in the underlying vomer, whereas in L. lujae the indentation is floored partly by the ethmoid cartilage and partly by the vomer. Compared with other species of Leptocypris, L. lujae is longer-jawed, the posterior tip of the maxilla extending to, or beyond, the posterior rim of the orbit; there is also a lower jaw symphysial process. As in L. niloticus and L. weynsii, L. lujae has a non-papillate maxillary valve. There is some variability in the length of the pelvic axial scale, it varies between 42%-50% of the pelvic fin length. However, this appears to be positively correlated with the length of the fish and it is noted that in L. niloticus the pelvic scale is not developed to its maximum length (25% of the pelvic fin length) until the fish is over 35 mm SL. In the length of the jaws and the morphology of the ethmoid region, L. lujae appears to be the most derived species. The species of Leptocypris may be identified by the following key: 1. Gill-rakers on first ceratobranchial absent; branched anal fin rays 8-9 . . . L.modestus Gillrakers on first ceratobranchial 2-3; branched anal fin rays 1 1-15 .... 2. Branched anal fin rays 1 1-12 L. niloticus Branched anal fin rays 14-15 3 3 Posterior tip of maxilla extending to centre of eye; lateral line scales 44-45 . . . L. weynsii Posterior tip of maxilla extending to, or beyond posterior border of the eye; lateral line scales 38-40 L. lujae Attention is drawn to 'Barilius' guineensis Daget, 1962, which appears, from its gross morphology, to belong to Leptocypris (see Howes, 1980 : 191). It differs from other species, however, in (according to Daget, 1962) having a total of 8 gill-rakers on the first arch. Bull. Br. Mm. not. Hist. (Zool.)45 (2): 95-101 Issued 28 July 1983 95 96 G. J. HOWES Mec Fig. 1mm 1 Leptocypris lujae\ dorsal view of ethmovomerine region of the neurocranium. Mec = mesethmoid cartilage, Pe = preethmoid, Se = supraethmoid, Vo = vomer. Comments on south-east Asian Barilius species When offering a revised concept of Barilius (Howes, 1980), based on the type species of the genus B. barila, few of the south-east Asian species had been examined and their generic attribution remained doubtful. Of those species in this category (Howes, 1980 : 190) the following have now been examined: B. bernatziki Koumans, 1937 (Holotype, Basle Museum, NHMB 5155, 77 mm SL); B. huahinensis Fowler, 1934 (USNM 103104, 52 & 66 mm SL); B. infrafasciatus Fowler, 1934 (USNM 107910, 65-75 mm SL); B. koratensis Smith, 1931 (Holotype, USMN 90298 47-3 mm SL); B. nanensis Smith, 1945 (Paratypes USNM 107939, 107940, 119474-119476, 53-64 mm SL); B. pulchellus Smith, 1931 (Paratypes USNM 90299, 51 & 53 mm SL; USNM 107967,39-5-69-5 mm SL). Osteologically and meristically, these species are intermediate between the generic subgroups (i) and (ii) previously postulated (Howes, 1980: 180), possessing characters of both groups. This suggests that those characters used in defining the two groups are a mixture of plesio- and apomorphies. In the south-east Asian species, and noticeably in B. pulchellus, the ethmoid bloc is somewhat flattened with the vomer extending anteriorly to the mesethmoid. The lateral edges of the supraethmoid are slightly raised to produce a shallow channel. The maxilla has a tall mid-lateral (palatine) ascending process and the palatine is broad anteriorly, slightly overlapping the maxilla and supporting the base of the anterior barbel. Rows of tubercles appear on the lower jaw and infraorbitals. These characters are shared with the Indian species, B. gatensis and B. bendelisis, and those species referred to group (ii) by Howes (1980: 190). The morphology of the ethmo-vomerine region, lateral extension of the palatine, and the pattern of tubercle development on the lower jaw appear synapomorphic for this group of species. As such, the concept of Barilius presented previously (Howes, 1980) is shown to be a paraphyletic one; see below, p. 99. The taxonomy of the south-east Asian species is in an unsatisfactory state. Smith (1945) separates B. nanensis and B. huahinensis on the differences in lateral line scale counts, dorsal fin position, and length of the anterior (rostral) barbel. However, there appears to be BARILIINECYPRINID FISHES 97 variability and overlap in all these characters, although the differences in body markings between the two species quoted by Smith (1945 : 156-157) are consistent. In B. nanensis there are 7-8 gill-rakers on the 1st ceratobranchial cf. 9-10 in B. huahinensis. Barilius pulchellus Smith, 1931 is a distinctive species characterised by a large pectoral axial fin lobe, 3-6 minute, spiny gill-rakers on the 1st ceratobranchial, and a black dorsal fin membrane. Specimens identified as B. infrafasciatus Fowler, 1934 (type not seen) are similar in virtually every respect to B. pulchellus apart from the length of the pelvic axial scale which is 50% of the pelvic fin length cf. 25% in B. pulchellus. Smith (1931) described Barilius koratensis as without barbels. My examination of the unique holotype reveals both anterior and posterior barbels. Smith described only two vertical bars, above the pectoral and below the dorsal fin, but there is also a dark patch above the last ray of the anal fin. In other meristic and morphometric characters B. koratensis resembles B. nanensis; these taxa may well be only representatives of different populations of one species. Barilius bernatziki Koumans, 1937 has 30-31 lateral line scales, the least number of all the south-east Asian species. In its markings (7 dark vertical bands and basal caudal spot), large pectoral axial lobe, and narrow cranium, B. bernatziki most closely resembles the Indian species, B. gatensis. The phyletic position of Zacco Zacco was excluded from the bariliine group by Howes (1980) on the grounds that it lacked a posttemporal-subtemporal connection. Fink & Fink (1981) claim this feature is present, and an examination of additional material leads me to agree with them. If Zacco is included within the bariliine assemblage, characters which were previously regarded as examples of homoplasy (peculiar morphology of the anal fin, ventral prolongation of the caudal peduncle and colour pattern), must now be viewed as synapomorphies. Zacco must be regarded as the sister taxon to Opsariichthys and because of its less derived cranial architecture and relatively unmodified lower jaw, it is the plesiomorph partner. Inclusion of Zacco within the bariliine group does not alter the group's distribution mapped in Howes (1980, fig. 47). Monophyly of the bariliine group According to Fink & Fink (198 1) a number of characters used by Howes (1980) to define the bariliine group are plesiomorphic. Of these they list the ventrally open posterior myodome; trigeminal foramen entirely within the prootic; lateral hyomandibular flange; frontal fossa; lateral temporal foramen, and the metapterygoid postero-dorsal process. They also believe that fusion of the 2nd and 3rd centra, only present in some bariliines, may be synapomorphic for those taxa and other Cyprinidae. Posterior myodome. Whilst it is recognised that a ventrally open posterior myodome often occurs in what are regarded as plesiomorphic teleosts (Patterson, 1975) it does not follow that the feature is itself plesiomorphic. Amongst otophysans, the posterior myodome in characoids is developed to various degrees. In some taxa currently recognised as mono- phyletic groups (Alestinae) there are some species with an open and others with a closed myodome. Also, in those taxa usually regarded as plesiomorphic (e.g. Hepsetus) the myodome is closed, whereas in those considered derived (Serrasalmidae) it is open. Ontogenetic evidence for polarity is contradictory; in a cyprinid investigated the myodome closes during ontogeny (Opsariichthys; see Howes, 1980), whereas in salmonids it opens (Verraes, 1974). Functionally, an open myodome serves to extend the length of the posterior eye muscles which then originate either from the rim of the basioccipital or from the anterior vertebrae. From its mosaic distribution and varying morphology, it is more parsimonious to 98 G. J. HOWES suppose that an open posterior myodome has developed independently in several lineages, and as such is an unreliable indicator of relationship. Hyomandibular flange, frontal fossa and temporal foramen. For the most part the same argument may be applied to these features as to the posterior myodome, namely, mosaic distribution through independent derivation, and thus I agree with Fink & Fink (1981) in disregarding them. However, some mention must be made of the lateral temporal foramen. Wu, Chen, Chen & Chen (1981) considered that the lateral temporal fossa in the Cobitidae and Gyrinocheilidae, and the supratemporal fossa in Catostomidae are homologues of the post-temporal fossa in other cyprinoids. This certainly appears to be so, as in all cases the fossa is bounded by the epioccipital, pterotic and, usually, the parietal. The lateral temporal foramen described in Opsaridium (Howes, 1980) is not to be confused with the above as it occurs between the pterotic and sphenotic and is thus homologous with that aperture in some characoids (e.g. Salminus). Metapterygoid process. Although a metapterygoid dorso-posterior process occurs in other otophysans, it differs markedly from the condition considered derived for some bariliine taxa (Howes, 1980). In Raiamas the process is directed anterodorsally at an angle of 45 and is the same length as the upper part of the hyomandibula. Furthermore, the process has a gutter along its posterior border into which the levator arcus palatini muscle inserts. In other bariliines the process is not so well developed, but in Engraulicypris and Leptocypris it is tall and lamellate, occupying a mid-dorsal position on the metapterygoid. In these taxa it also serves to support the LAP muscle. Whilst it is accepted that a metapterygoid process is a common otophysan feature, in none does it possess the morphology of the process in bariliines, nor does it provide the site of attachment of almost the entire LAP muscle. Vertebral fusion. Fink & Fink's (1981) claim that all other members of the bariliine group are '. . . more specialised than Opsariichthys in having the second and third centra fused' must be treated with caution. Fusion of centra 2 and 3 is a variable character throughout the Cyprinidae. There may be only partial (dorsal) fusion, as in Zacco and Engraulicypris (Howes, 1980) and Pseudo- laubuca (Howes, 1978). Monophyletic groups contain taxa exhibiting both fused and unfused centra (e.g. aspinines and squaliobarbines) and in such cases it is not always the more derived members of the groups which display the fusion pattern. Thus, vertebral fusion must be regarded as having been derived independently in several lineages. Supporting Fink & Fink's argument for the derived nature of vertebral fusion simply leads to an unresolved reticulate pattern of relationships (Fig. 2). Interrelationships of bariliines Disregarding the posterior open myodome and those characters considered by Fink & Fink (1981) as plesiomorphic, the bariliine group can be defined on only a single synapomorphy, ie a connection between the posttemporal and subtemporal fossae. That this character has now been found in Salmostoma (placed in the cheline group by Howes, 1979) revises the concept of the bariliine group. Accepting the fossae connection as synapomorphic, then two lineages can be recognised, (1) those species with elongate pectoral and pelvic axial scales, long and expanded processes of the 1st vertebrae, with development of a condylar joint with the basioccipital, and a modified ethmo-vomerine region (see caption to Fig. 3), and (2) those with the pectoral and pelvic axial scales lobate or fleshy, the rostral barbel attached to the lateral border of the palatine, parallel rows of tubercles on the lower jaw, and distinctive body and fin patterning. Lineage (1) includes Barilius barila, B. evezardi, B. modestus and B. vagra, Salmostoma, Engraulicypris, Raiamas and Leptocypris. Salmostoma was regarded by Howes (1979) as the plesiomorph sister group of 'cheline' genera. The 'chelines' share with Barilius and Salmostoma the derived form of anterior vertebrae and ethmovomerine region, but lack the BARILIINECYPRINID FISHES 99 Other Some Zacco.Opsariichthys 'bariliines' cyprinids & some cyprinids Centra 2 3-^ - Centra fused; \ / unfused 'derived' Fig. 2 A cladogram of cyprinid relationships based on the pattern of vertebral fusion proposed by Fink & Fink (1981); see text. posttemporal-subtemporal connection. Salmostoma, however, shares with all other chelines a deep, posteriorly sloped neural complex. Only one 'cheline' genus, Securicula shares the elongate pectoral and pelvic axial scales with Salmostoma and Barilius. In the absence of synapomorphies linking the 'cheline' genera with any other cyprinid taxon, the absence of these features is considered as a phylogenetic loss. Current observations indicate that the genera Rastrineobola, Neobola and Chelaethiops also belong to this lineage, and work in progress attempts to define more precisely the relationships of these three genera. Lineage (2) contains the remaining Barilius species recognised as group (ii) by Howes (1980: 1 80) together with Opsaridium, Opsariichthys and Zacco, plus the luciosomine genera (sensu Howes, 1980). In summary, it would seem that the bariliine and cheline groups (sensu Howes, 1979; 1980) are paraphyletic assemblages; a revised hypothesis of the interrelationships of the relevant taxa is presented in Fig. 3. The dates of Opsaridium Peters and Rastrineobola Fowler Jordan (1919) mistakenly gives the publication date of Opsaridium Peters as 1855. Howes (1980) followed this error. The correct date is 1853 (see citation in references). The specific name is correctly zambezensis. The publication date of Rastrineobola Fowler is 1936, not 1934 as misprinted in Howes (1980: 195). Bengala Gray, 1832, a senior synonym of Megarasbora (, tint her, 1868 Giinther (1868) established the genus Megarasbora to contain Cyprinus elanga Hamilton, 1822. Megarasbora was recognised by Brittan (1954) and Howes (1980) in their respective revisions and reviews of rasborine cyprinids. However, all three authors had overlooked the fact that Gray (1832) had already established the genus Bengala for Hamilton's species. Jordan (1919) recorded Megarasbora as a junior synonym of Bengala. 100 G. J. HOWKS Fig. 3 Revised cladogram of 'bariliine' and 'cheline' relationships (dashed lines indicate uncertain affinities). 1, posttemporal-subtemporal fossa connection; 2, pectoral and pelvic axial scales elongate (lost in 'chelines' and Engraulicypris); 2a, 1st vertebra with expanded lateral processes; 2b, omega-shaped ethmoid notch, vomerine arms extending straight forward; 2c, reduction of dilatator fossa, derived condition of jaw adductor muscles; 2d, loss of intermandibularis muscle, jaw elongation, hypertrophy of metapterygoid spine; 3, pectoral and pelvic axial scales lobate or fleshy; 3a, palatine extended laterally, supports rostral barbel (when present); 3b, parallel rows of tubercles on dentary; 3c, dentary with anterior notch, sexual dimorphism exhibited by extension and expansion of anal fin rays in males; 3d, bowl-shaped depression in supraethmoid. Characters described in Howes (1980), except 2b and 2c, taken from work in preparation. Acknowledgements My sincere thanks are due to Dr Richard Vari for facilitating my study of Smith's specimens in the United States National Museum and for his warm hospitality. I am most grateful to Maurice Kottelat for providing information on south-east Asian Barilius and arranging loans of specimens, and to Margaret Clarke for her assistance with nomenclatural problems. Drs Humphry Greenwood and Keith Banister have kindly read, and critically commented on the manuscript. References Brit tan, M. R. 1954. A revision of the Indo-Malayan fresh-water fish genus Rasbora. Institute of Science & Technology, Manila. 224 pp. Daget, J. 1962. Les poissons du Fouta Dialon et de las basse Guinee. Mem. Inst. fr. Afr. noire 65: 1-210. BARILIINECYPRINID FISHES 101 Fink, S. V. & Fink, W. L. 1981. Interrelationships of the ostariophysan fishes (Teleostei). Zool. J. Linn. Soc. 72 (4): 297-353. Gray, J. E. 1 832. Illustrations of Indian Zoology. Part 2, pi. 96. London Giinther, A. 1 868. Catalogue of fishes in the British Museum 7: 1-5 1 2. Howes, G. J. 1978. The anatomy and relationships of the cyprinid fish Luciobrama macrocephalus (Lacepede). Bull. Br. Mus. nat. Hist. (Zool.) 34: 1-64. 1979. Notes on the anatomy of the cyprinid fish Macrochirichthvs macrochirus with a review of the subfamily Cultrinae. Bull. Br. Mus. nat. Hist. (Zool.) 36 (3): 147-200. 1980. The anatomy, phylogeny and classification of bariliine cyprinid fishes. Bull. Br. Mus. nat. Hist. (Zool.) 37 (3): 129-198. Jordan, D. S. 1919. The genera of fishes Part 3. Leland Stanford Junior Univ. Pubs. University Series:285^MO + i-xv. Peters, W. C. H. 1853. Uebersicht der in Mossambique beobachteten Seefische. Monats. Ber. Acad. Berlin: 783. Smith, H. W. 1945. The fresh-water fishes of Siam, or Thailand. Bull. U.S. natn. Mus. 188: 1-622. Verraes, W. 1974. Discussion on some functional-morphological relations between some parts of the chondrocramium and the osteocranium in the skull base and the skull roof, and of some soft head parts during postembryonic development of Salmo gairdneri Richardson 1836 (Teleostei: Salmonidae). Forma Functio 7: 28 1-292. Wu, X., Chen, Y., Chen, X. & Chen, J. 1981. A taxonomical system and phylogenetic relationship of the families of the suborder Cyprinoidei (Pisces). Scientia Sinica 24 (4): 563-572. Manuscript accepted for publication 1 7 June 1982 British Museum (Natural History) An Atlas of Freshwater Testate Amoebae C. G. Ogden & R. H. Hedley 1980, Hardcovers, 222pp, 1 7.50 (1 8.00 by post). Co-published by British Museum (Natural History) and Oxford University Press. This book illustrates, using scanning electron micrographs, most of the common species of testate amoebae that are found in freshwater habitats. Information on the biology, ecology, geographical distribution and a classification are followed by descriptions of ninety-five species. Each of these is illustrated by several views of the shell. The text is designed not only to enable biologists to identify species of testate amoebae, but to serve as an introduction to students interested in the taxonomy and biology of these fresh- water protozoa. It will be of special interest to prptozoologists, ecologists, limnologists, water treatment specialists and micropalaeontologists interested in recent sediments. British Museum (Natural History) Publication Sales, Cromwell Road, London SW7 5 BD Titles to be published in Volume 45 Problems in catfish anatomy and phylogeny exemplified by the Neotropical Hypophthalmidae (Teleostei: Siluroidei). By G. J. Howes Miscellanea Bats (Mammalia: Chiroptera) from Indo- Australia. By J. E. Hill. On Macropleurodus, Chilotilapia (Teleostei: Cichlidae) and the interrelationships of African cichlid species flocks. By P. H. Greenwood. Miscellanea The cranial muscles of loricarioid catfishes, their homologies and value as taxonomic characters (Teleostei: Siluroidei). By G. J. Howes. Miscellanea Printed by Henry Ling Ltd, Dorchester Bulletin of the British Museum (Natural History) Bats (Mammalia: Chiroptera) from Indo- Australia J. E. Hill Zoology series Vol 45 No 3 25 August 1983 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself, available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the series on either an Annual or Per Volume basis. Prices vary according to the contents of the individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History), Cromwell Road, London SW7 5BD, England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Zool.) Trustees of the British Museum (Natural History), 1983 The Zoology Series is edited in the Museum's Department of Zoology Keeper of Zoology : Dr J. G. Sheals Editor of Bulletin : Dr C. R. Curds Assistant Editor : Mr C. G. Ogden ISSN 0007- 1 498 Zoology series Vol 45 No. 3 pp 103-208 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 25 August 1983 Bats (Mammalia: Chiroptera) from J.E.HH1, Department of Zoology, British Museum (Natural History), Cromwell Road, SW7 5BD Contents Synopsis . . . . Introduction Systematic section . Megachiroptera . Pteropodidae . Pteropodinae. Harpyionycterinae Nyctimeninae Macroglossinae . Microchiroptera . Emballonuridae Megadermatidae Rhinolophidae . Hipposideridae . Vespertilionidae Vespertilioninae . Miniopterinae Murininae Kerivoulinae . Molossidae Summary .... Acknowledgements . References. Addendum 103 103 104 104 104 104 127 130 132 140 140 142 143 148 153 153 171 190 194 195 198 199 199 208 Synopsis Accessions of Indo-Australian bats to the collections of the British Museum (Natural History) during the past fifteen years are reviewed in detail, with a particular reference to the collections made in Papua New Guinea and Sulawesi by 'Operation Drake'. Numerous species and species groups from the region are examined and in some cases revised; a new species of Hesperoptenus from Sulawesi and a new subspecies of Myotis adversus from the New Hebrides are described. The more important taxonomic studies and notes made in this study and the new range records that it reports are listed in a terminal summary. Introduction A variety of collections of bats from the region bounded by India in the west and the New Hebrides, New Caledonia and Fiji to the east has been received at the British Museum (Natural History) during the years 1967-1982. Some have come from individuals who have obtained small numbers of bats while visiting the region, others from large, organised expeditions that have carried out biological studies, or by donation from workers connected with organisations and institutions in the area. Individually much of this newly accessed material scarcely justified separate publication and record, although often unusual or of taxonomic or faunal interest. Towards the end of this period, however, extensive collections of bats were made in Papua New Guinea and in Sulawesi by Mr Ben Gaskell, an ecologist and collector with 'Operation Bull. Br. A/MS. nat. Hist. (Zool.) 45(3): 103-208 Issued 25 August 1983 104 J. E. HILL Drake', the commemorative round-the-world voyage of the brigantine Eye of the Wind in 1978-1980 that marked the 400th anniversary of the circumnavigation of the world by Sir Francis Drake in the Golden Hind. The voyage was interrupted from time to time so that the scientists, technicians and young people aboard could undertake adventurous, scientific or voluntary aid projects in various parts of the world. Bats were collected in Indo-Australia during two of these periods, first at and around Buso and Wau in Morobe Province, Papua New Guinea, and later in central Sulawesi, chiefly around Morowali where the objective was ultimately to prepare a Management Plan for the Morowali Nature Reserve. The study of the specimens obtained by 'Operation Drake', especially of those from Sulawesi, has entailed a further and more wide-ranging examination of much of the Indo-Australian bat collection already in London, and has prompted a further examination of the more recently accessed material that has not before been reported in the literature. In particular it has led to further study of a number of long standing taxonomic problems in the area and to some taxonomic changes. Although basically this paper is concerned with specimens obtained through the explorations initiated and carried out through 'Operation Drake' it has thus been possible to add a variety of other studies and notes drawn from material from other localities and sources, much of it directly relevant to the 'Drake' collections. Place names in many parts of the Indo-Australian region can present difficulties, there being sometimes a choice of as many as three variants of any one designation. In general the traditional and conventional European spelling or usage (i.e. Amboina, Ceram) has been adopted but Sulawesi has been used throughout for Celebes, this name having come into general use: however, the New Hebrides remain so called, although recently renamed Vanuatu. The Indonesian part of New Guinea appears as West Irian, while Papua New Guinea is used for the rest of the island: here, 'Province' has been omitted from locational data so that localities appear as 'Wau, Morobe' i.e. Wau, Morobe Province. Further west, Borneo is used as a general term for the entire island, divided into Sarawak, Sabah, Brunei and Kalimantan. Measurements of specimens are in millimetres and with the exception of those of individual teeth have been made with a dial-reading micrometer. Teeth have been measured with a mechanical stage fitted to a stereoscopic microscope. Although no standard suite of external and cranial measurements has been used, conventional measurements are given where required, amplified in particular cases to conform with those employed by the describer or by previous workers on the species concerned. To avoid repetition, a notation has been adopted for the wing elements whereby III m for example indicates the metacarpal of the third digit, III 1 its first phalange, IIP its second: when length measurements of these elements are given in the text these designations appear without preamble. Systematic Section MEGACHIROPTERA PTEROPODIDAE PTEROPODINAE Rousettus amplexicaudatus stresemanni Stein, 1 933 RousettusstresemanniSlein, 1935: 91 . Japen ( = Jobi) I, Geelvinck Bay, NE West Irian. SPECIMENS EXAMINED. Papua New Guinea: cf BM(NH) 69.1416 Madang, 5 14'S, 14545'E (skin, skull; coll. J. I. Menzies); <*BM(NH)73.1967 Rauit, West Sepik, 525 m,336'S, 142 1 5 'E (in alcohol; INDO-AUSTRALIAN BATS 105 coll. Aberdeen University Exploration Society Expedition to New Guinea, 1973); 3dd BM(NH) 78.850-852 Baku Forest Station, Gogol Valley, Madang (78.850, 851 in alcohol, skull of 78.850 extracted, 78.852 skin only; coll. P. A. Morris); 9 BM(NH) 78.853 Baiyer R, c. 50 km NW of Mt. Hagen, Western Highlands (in alcohol, skull extracted; coll. P. A. Morris). REMARKS. Until recently there have been few records of stresemanni and for many years (Laurie & Hill, 1954) it has been considered a distinct species co-existing in New Guinea with R. amplexicaudatus brachyotis (Dobson, 1877). Latterly, however, a number of reports from Papua New Guinea has appeared, McKean (1972) recording a series of stresemanni from Ihu, Greig-Smith (1975) a specimen (listed above) from Rauit, with a record of an example (also listed above) from Madang first noted (in litt.) by Menzies, and with Koopman (1979) reporting material from Bagabag I, off the northeastern coast near Madang and later (1982) from Dabora, Tapio and Mornuna, Milne Bay, east Papua New Guinea. Menzies (1977) tentatively referred sub-fossil remains from Kiowa in Ghimbu Province to stresemanni. Finally, Rookmaaker & Bergmans (1981) have listed numerous records from localities in much of New Guinea, some evidently based on specimens previously identified as R. a. brachyotis. Specimens in the American Museum of Natural History suggested to Koopman (1979) that stresemanni might be properly regarded as a subspecies of R. amplexicaudatus while Rookmaaker & Bergmans (1981) in their comprehensive review of this species have synonymised stresemanni with R. a. amplexicaudatus (Geoffrey, 1 8 100). Specimens recorded here from Papua New Guinea are large, corresponding closely with the original description of stresemanni by Stein. In size they also agree with specimens from Ihu reported by McKean (1972) and with those from New Guinea measured by Rookmaaker & Bergmans (1981). They are generally a little larger than six from Timor examined by Goodwin (1979) and are similarly near or exceed the upper size limits of Timorese speci- mens (including some of those seen by Goodwin) studied by Rookmaaker & Bergmans. Specimens from Timor may be assumed to be topotypical or nearly so of R. a. amplexicaudatus, described originally from that island. The dimensions given by Rookmaaker & Bergmans for examples from New Guinea are likewise generally a little greater than those for specimens from Timor and its associated islands. External measurements of 4dd and 1 9, with cranial measurements of 2dd and 1 9: length of forearm 85-8-904, 85-5; greatest length of skull 38-6, 394, 38-8; condylobasal length 36-8, 37-0, 37-5; condylocanine length 35-6, 35-8, 36-0; length front of orbit to tip of nasals 13-1,13-0, 13-1; length palation to incisive foramina , 17-5, ; least interorbital width 8-6, 8-3, 8-3; least postorbital width 7-5, 7-6, 8-3; zygomatic width 24-7, 24-0, 22-3; width of braincase 154, 15-6, 154; mastoid width 14-6, 14-1, 144; greatest width c'-c 1 7-1, 74, 7-6, c'-c 1 (alveoli) 6-7, 6-9, 7-0; m^m 2 (alveoli) 10-2, 10-3, iO-8; c-m 2 13-5, 13-7, 14-0; length complete mandible from condyles 28-6, 28-3, ; length right ramus from condyle 30-1, 29-9, 30-0; c-m 3 15-1, 14-7, 15-6. DISCUSSION. Rookmaaker & Bergmans (1981) have reviewed R. amplexicaudatus in detail, with a summary of earlier classification, synonymies, and many measurements derived from a relatively large number of specimens. These authors considered size to be the only major subspecific character in the species and recognised three size groups from the Solomon Islands, New Guinea, the Philippine Islands, Timor and Java. Of these, the smallest examples were found to occur on the Solomon Islands, the largest in New Guinea and the Philippines (these latter being mutually indistinguishable); specimens from Java, like those from the Solomons, proved significantly smaller than specimens from New Guinea and the Philippines. Examples from Timor were found to be generally a little larger than those from Java, but not significantly so, and a little smaller on the whole than those from New Guinea and the Philippines, although again the differences lacked significance. However, such speci- mens were thought by Rookmaaker & Bergmans to have a greater similarity to examples from New Guinea and the Philippines than to those from Java. On this basis they recognised three subspecies, R. a. brachyotis (Dobson, 18770) from the Solomon and Bismarck Islands, R. a. infumatus (Gray, 1 870) from Sumatra and Java east to Flores and possibly Alor Island, 106 J. E. HILL and R. a. amplexicaudatus from Sumba Island, Timor and some smaller associated islands, New Guinea, possibly from the Molucca Islands, and from the Philippines. Specimens from Sulawesi were not allocated to subspecies: others from Borneo, Mentawei and Engano Islands, Malaya, Thailand and Burma were referred to R. a. amplexicaudatus, but few examples are available from the western part of the range. At the eastern limit, however, Smith & Hood (1981) have suggested that the population on the Bismarck Islands (R. a. brachyotis) is subspecifically separable from that on the Solomon Islands (R. a. hedigeri Pohle, 1953), although the distinction is less marked than the divisions between these island subspecies and R. a. stresemanni from New Guinea. Considerations of relative size led Rookmaaker & Bergmans (198 1 ) to associate specimens from Timor with those from New Guinea and thus to synonymise stresemanni with R. a. amplexicaudatus. However, their account of specimens from Sulawesi and some of its associated islands indicates that in some ways these unallocated examples are intermediate between R. a. infumatus and Timorese specimens ofR. a. amplexicaudatus. A study of their detailed tabulated measurements for these populations confirms this view: also of two specimens (BM(NH) 98.11.3.20-21) from Alor Island provisionally allocated to R. a. infumatus by Rookmaaker & Bergmans the male is similar in size to males from Timor, but the female to females from Java. Rookmaaker & Bergmans also made a detailed examination of the relation between condylobasal length and zygomatic width in some of the populations of R. amplexicaudatus. Their diagrams show that while in these dimensions Timorese specimens lie in the lower part of the range of variation of those from New Guinea and the Philippines and are separated more distinctly from the Javanese population the limited Sulawesian sample tends to bridge this interval. It is also evident that the majority of specimens from New Guinea and the Philippines exceed Timorese and Sulawesian specimens in one or both of these dimensions. There seems, therefore, at least as much to justify the association of Timorese specimens (R. a. amplexicaudatus) with those from Sumatra, Java and the Lesser Sunda Islands (R. a. infumatus) as with those from New Guinea. For these reasons stresemanni has been retained as a distinct subspecies in New Guinea and possibly also in the Philippine Islands. Measurements by Rookmaaker & Bergmans (1981) indicate that only small overall differences exist between these populations. Philippine females have longer wing elements than the very limited New Guinea sample examined and on the basis of a larger representation of females from New Guinea are cranially rather smaller on the whole but with longer toothrows. Until more specimens are available from the Molucca Islands, Sulawesi and the western part of the range of R. amplexicaudatus from Borneo and Sumatra to Burma it seems appropriate to regard R. a. amplexicaudatus as a valid link between the smaller R. a. infumatus and the larger R. a. stresemanni. Rousettus celebensis Andersen, 1 907 Rousettus celebensis Andersen, 1907a: 509. Mount Masarang, N Sulawesi, 3500 ft. SPECIMENS EXAMINED. N Sulawesi: 99BM(NH) 78.964-967 Tangkopo, Batuangus, near Bitung (in alcohol, BM(NH) 78.965 head, skin, others heads only; coll. A. M. Jones). C Sulawesi: rfrf BM(NH) 81.1066-1068 Ganda Ganda, 1 57' S, 121 21' E; dd 106-125 117 99 93-128 108 IP-3 dd 116-142 133 (c.u.) 99 110-142 128 IIP dd 651-669 658 99 636-694 660 III 1 dd 455^69 462 99 438^74 455 IIP dd 548-621 585 99 554-615 583 IV m dd 631-652 637 99 617-659 635 IV dd 330-356 344 99 321-359 340 IV 2 dd 368-401 378 99 343-385 367 ym dd 623-651 639 99 615-645 637 V dd 291-306 299 99 275-304 292 V 2 dd 333-355 348 99 325-352 321 ally long and narrow, and, as in lanosus, the molars have little cuspidation. Andersen (1912) considered that celebensis probably represented a modification of the R. amplexicaudatus type, but it seems more appropriate to include this Sulawesian species with R. lanosus in the subgenus Stenonycteris if this is to be recognised rather than in the nominate subgenus where Andersen originally placed it. Pteropus hypomelanus macassaricus Heude, 1 896 Pteropus macassaricus Heude, 1 896 : 1 77, footnote, pi. 5, fig. 4. Makassar, S Sulawesi. SPECIMENS EXAMINED. C. Sulawesi: d BM(NH) 81.1092 Ganda Ganda, 1 57' S, 121 21' E- 99 BM(NH) 81.1093-1096 1 km NE of Tandiondo, 1 45' S, 121 17' E (all in alcohol, all except BM(NH) 81.1092 heads only; coll. B. H. Gaskell, 'Operation Drake'). Pteropus alecto alecto Temminck, 1837 Pteropus alecto Temminck, 1837, 2 : 75. Menado, N Sulawesi. SPECIMENS EXAMINED. C Sulawesi: rf, 4 99 BM(NH) 81.1097-1 101 1 km NE of Tandiondo, 145' S, 121 17' E (in alcohol, heads only; coll. B. H. Gaskell, 'Operation Drake'). INDO-AUSTRALIAN BATS 109 REMARKS. Musser et #/.(1982) have demonstrated that among the several forms of Pteropus described or reported from Sulawesi P. arquatus Miller & Hollister, 192 1 is in fact Acerodon celebensis, as are the specimens from the northern part of the island recorded originally by Tate (1942c) as P. argentatus. Others known from Sulawesi besides P. hypomelanus macassaricus and P. alecto alecto are P. griseus mimus Andersen, 1908, P. caniceps dobsoni Andersen, 1908 and P. personatusTemminck, 1825. Pteropus conspicillatus Gould, 1 850 Pteropus conspicillatus Gould, 1 850 : 109. Fitzroy I, Queensland, Australia. SPECIMENS EXAMINED. Papua New Guinea: rf, 9, 9 juv. BM(NH) 80.525-527 Lababia Cave, Lababia I, Morobe, 7 15 ' S, 147 09 ' E (in alcohol, coll; B. H. Gaskell, 'Operation Drake'). REMARKS. Laurie & Hill (1954) listed P. c. conspicillatus from Papua New Guinea after Matschie ( 1 899) who reported this subspecies from Bongu (5 30 ' S, 1 45 50 ' E) and Madang (5 13' S, 145 48 'E): a second subspecies, P. c. chrysauchen Peters, 1862 was listed from northwestern New Guinea after Tate (1942c) who recorded it from Geelvinck Bay. These examples from Lababia Cave cannot be allocated positively to either of these subspecies. The adult c? BM(NH) 80.525 has a uniformly dark head, the forehead, crown and sides of the muzzle blackish, mixed especially on the forehead and on the muzzle anterior to the eyes with buffy hairs. Beyond a slight band of buffy hairs there is no indication of the paler eye rings characteristic of P. c. conspicillatus and the sides of the muzzle are not pale as in this subspecies. The adult 9 BM(NH) 80.526, however, although predominantly black on the forehead, crown and cheeks has relatively distinct paler buffy eye rings and the sides of the muzzle above the mouth and the corresponding areas along the lower jaw are distinctly ochraceous buff. Moreover, the black of the crown extends forward as an acutely triangular patch whose apex intrudes between the pale superciliaries to the base of the rostrum. The adult male agrees therefore with P. c. chrysauchen, the adult female with P. c. conspicillatus. The collections of the British Museum (Natural History) include two further specimens (c?, 9 22.1.22.1-2) from Simbang, on the east coast of Papua New Guinea (probably at 3 35 S, 147 43' E) that are referable to P. c. conspicillatus although even in these the facial markings of the male are a little less pronounced than in the female example. Length of fore- arm in specimens from Lababia Cave (c?, 9) 1 8 1 , 1 79. Styloctenium wallacei (Gray ', 1866) Pteropus wallacei Gray, 1 866a : 65, fig. 1 . Makassar, S Sulawesi. SPECIMENS EXAMINED. C Sulawesi: 9 (yg. ad.), cf BM(NH)81.1 102-1103 R Ranu, 151 ' S, 121 30' E (in alcohol): cf (yg. ad.) BM(NH) 81.1 104 (in alcohol), 9 BM(NH) 81.1 105 (skin, skull, skeleton), 9 (yg.) BM(NH) 81.1 106 (in alcohol) Tambusisi Damar, Mt. Tambusisi, c. 4000 ft, 1 39 ' S, 121 22 ' E (all coll. B. H. Gaskell, 'Operation Drake'). REMARKS. Although this distinctive bat is by no means common in collections there is a number of records (Jentink, 1883; Matschie, 1899; Andersen, 1912) from N Sulawesi and Tate (1942c) has reported a long series from Malenge in the Togian Islands in the Gulf of Gorantalo. There is also a hitherto unreported specimen (cf MZB 12671) from Titaeli, Minahassa, in the collections of the Museum Zoologicum Bogoriense, Bogor. HO J. E. HILL Adults obtained by Operation Drake display very clearly the badger-like white facial markings characteristic of the species: the white shoulder patches although always present are rather small in the two male examples, one not quite fully adult. External measurements of an adult d and 9 (BM(NH) 81.1103, 81.1105): length of forearm 96-6, 95-5; thumb (c. u.) 43-2, 42-3; I m 12-2, 12-0; I 1 23-7, 234; II m 50-7, 51-5; II 1 12-4, 12-3; II 2 - 3 (c. u.) 12-0, 11-3; III m 69-0, 67-2; III 1 51-3, 51-4; III 2 69-0, 63-7; IV m 69-4, 68-6; IV 39-7, 38-5; IV 2 40-6, 40-4; V m 7 1 -7, 7 1 -8; V 3 1 -4, 3 1 -5; V 2 33-8, 33-9; tibia 42-4,. Cranial measurements of an adult 9 (BM(NH) 81.1105): total length of skull to gnathion 51-8; condylobasal length 48-5; condylocanine length 444; length front of orbit-tip of nasals 16-8; palatal length 28-0; length palation-incisive foramina 23-8; length palation-basion 18-3; lachrymal width 10-5; least interorbital width 6-8; least postorbital width 5-9; zygomatic width 28-3; width of braincase 19-7; mastoid width 17-9; orbital diameter 114; c'-c 1 (crowns) 9-5, (alveoli) 8-8; m'-m 1 (crowns) 13-9, (alveoli) 12-7; c'-c 1 (internally, cingula) 5- 1 ; pm 4 -pm 4 (internally) 6-9; width of mesopterygoid fossa 74; c-m 2 19- 1 ; length of complete mandible from condyles 364; length right ramus from condyle 38-1; coronoid height 19-8; c-m 2 19-9. Dobsonia viridis (?) viridis (Heude, 1896) Cephalotes viridis Heude, 1 896 : 1 76, footnote, pi. 5, fig. 1 . Kei Is. Dobsonia viridis umbrosa Thomas, 1910a : 384. Ceram I. SPECIMENS EXAMINED. C Sulawesi: 9, cT BM(NH) 81.1107-1108 R Ranu, 1 51' S, 121 30' E (in alcohol, skulls extracted; coll. B. H. Gaskell, 'Operation Drake'). REMARKS. These are apparently the first of viridis to be reported from Sulawesi: it occurs otherwise on the Kei Islands and on the islands of Amboina, Buru, Ceram and Banda. Thomas (1910#) separated specimens from Ceram as D. v. umbrosa to which Andersen (1912) subsequently referred others from Buru and Amboina: this author (p. 825) remarked that the reference by Thomas to a number of specimens from 'Aru' is a misprint for Buru but in his personal copy of his paper, now in the Library of the British Museum (Natural History), Thomas has corrected Aru to Kei [Islands]. The collections in London include several specimens collected by W. Stalker on the Kei Islands but only a single immature from Buru, obtained by this collector. A small average difference in colour separates umbrosa from viridis of the Kei Islands and the two are synonymised in the listing by Laurie & Hill (1954). A closely related form, D. crenulata Andersen, 1909 occurs on the Halmahera group of islands to the northwest of New Guinea and has been reported recently from the Sanghir ( = Sangihe) Islands and on the Togian Islands. It is separated from D. viridis chiefly on account of its greater size and larger teeth. Specimens from Sulawesi have the characteristic dentition of the viridis group of Andersen (1909, 1912) in which m 1 has a well-developed antero-internal ledge and pm| and m} have cuspidate labial and lingual ridges. The surface cusps or ridges of mj and m 2 are also strongly developed and prominent, of pm 4 much less so. The collector, B. H. Gaskell remarks of the coloration of the adult that when freshly obtained the dorsal pelage was a light dull green in hue, yellower towards the rear and flanks, with the upper surface of the head yellowy green grey, sharply divided on the neck from the colour of the body. The ventral surface was generally similar in colour to the lower back but medially light orange. The indefinite greenish tinge in the pelage appears characteristic of D. viridis: Goodwin (1979) also commented upon the unusual olive green colour of adults of D. peronii peronii (Geoffroy, 1810#) when living. The colour faded rapidly in specimens preserved as dry skins, in alcohol, or in formalin. INDO-AUSTRALIAN BATS 1 1 1 In size both Sulawesian examples are similar to or exceed the largest of Moluccan speci- mens of viridis and their cheekteeth, especially of the male, are generally longer and slightly larger, particularly pm 4 , mj and m 2 . To some extent, therefore, they approach crenulata but their canines are smaller and their cheekteeth generally narrower, like those of viridis from the Moluccas. Measurements of an adult 9 and c? (BM(NH) 81.1107-1108): length of forearm 113-5, 125-4; III m 71-6, 77-1; IV m 65-4, 70-0; V m 66-2, 71-0; total length of skull to gnathion 48-2, 50-1; condylobasal length 46-2, 47-8; condylocanine length 45-7, 47-6; rostral length 15-6, 15-3; length front of orbit-tip of nasals 12-2, 1 1-9; palatal length 24-1, 25-0; length palation- incisive foramina 21-1, 22-1; length palation-basion 19-1, 19-6; lachrymal width 11-8, 11-6; least interorbital width 8-4, 8-2; least postorbital width 7-5, 6-4; zygomatic width 29-4, 31-1; width of braincase 19-7, 19-7; mastoid width 18-4, 18-3; orbital diameter 9-9, 9-9; c'-c 1 (crowns) 9-3, 9-5, (alveoli) 8-7, 8-7; m'-m 1 (crowns) 14-6, 15-1, (alveoli) 13-8, 14-4; m 2 -m 2 (crowns) 12-8, 12-9, (alveoli) 12-5, 12-6; c'-c 1 (internally, cingula) 3-4, 3-7; pm 4 -pm 4 (internally, 7-6. 7-7; width of mesopterygoid fossa 6-0, 5-8; c-m 2 (crowns) 18-8, 19-8, (cingula) 18-4, 19-4; length complete mandible from condyles 36-5, 37-4; length right ramus from condyle 37-7, 38-9; coronoid height 20-3, 21-8; c-m 3 (crowns) 20-0, 20-7, (cingula) 19-8, 20-4. Length/width of: c 1 3-86/2-48, 3-82/2-54; pm 3 3-94/2-88, 4-02/3-22; pm 4 4-02/2-72, 4-32/3-14; m 1 5-15/2-44, 5-17/2-83; m 2 2-34/1-56, 2-37/1-61; c, 2-64/2-17, 2-65/2-22; pm 2 1-40/1-34, 1-30/1-45; pm 3 3-72/2-31, 3-90/2-52; pm 4 4-02/2-53, 4-22/2-62; m 1 3-99/2-15, 4-21/2-35; m 2 3-30/2-08, 3-31/2-16; m 3 1-90/1-31, 1-85/1-43. DISCUSSION. The viridis group of Dobsonia as proposed by Andersen (1909, 1912) includes besides D. viridis and D. crenulata the further species D. praedatrix Andersen, 1909 from the Bismarck Archipelago and D. inermis Andersen, 1909 and D. nesea Andersen, 1909 from the Solomon Islands. Since Andersen wrote a number of changes have been made to this classification. Troughton (1936) regarded nesea as a subspecies of D. inermis while Rabor (1952) in describing D. viridis chapmani from Negros Island in the Philippine Islands treated crenulata as a further subspecies of viridis. Pohle (1953) considered crenulata and praedatrix to be subspecies of D. viridis and inermis (including nesea) probably so. Laurie & Hill (1954) retained the arrangement of Andersen but united nesea with inermis as a subspecies. More recently, Bergmans (1975) discussed the viridis group in some detail, describing a further species, D. beauforti from Waigeo Island in the northern Moluccas, and rejecting the views of Rabor and Pohle. Since then he has (1978) examined the group yet further and has established that chapmani does not fulfil the appropriate diagnostic criteria but seems more likely to belong to the moluccensis group. Moreover, Bergmans is now of the opinion that praedatrix and inermis (including nesea} differ sufficiently from the two (sic) other species that they should form one or two species groups by themselves. Dobsonia beauforti is said to be morphologically allied to D. viridis but is appreciably smaller in forearm and skull measurements, in size much like D. inermis from the Solomon Islands. No specimens of D. beauforti have been examined. Only the holotype (BM(NH) 60.8.26.2) of crenulata is available for study in London. However, De Jong & Bergmans (1981) have reviewed this taxon, recording it for the first time from the Sanghir ( = Sangihe) Islands and from the Togian Islands, and have provided measurements of a number of specimens. These authors refer to earlier views (Rabor, 1952; Pohle, 1953) that crenulata is a large subspecies of D. viridis, but regard the union of the two into the same species as premature, since specimens of each sex from any one population are as yet insufficient to establish its range of size variation. The specimens from Sulawesi approach and in some respects equal crenulata in external and cranial dimensions, but on the whole have slightly narrower cheek teeth similar in width to those of viridis. They have smaller canines than the subadult holotype of crenulata, corresponding closely to those of viridis. This limited sample therefore suggests strongly that viridis and crenulata are likely to prove conspecific, a view provisionally adopted here. 112 J. E. HILL Dobsonia praedatrix Andersen, 1 909 Dobsonia praedatrix Andersen, 1909 : 532. Duke of York I, Bismarck Archipelago. SPECIMENS EXAMINED. Bismarck Archipelago: 9, rf, 9 BM(NH) 69.304-306 Kareeba Plantation, Keravat, New Britain, 4 1 8 ' S, 1 52 01' E (skins, skulls; coll. J. I. Menzies). REMARKS. These relatively recently collected specimens confirm the diagnostic features used by Andersen ( 1 909, 1 9 1 2) in separating praedatrix from viridis and crenulata. As pointed out by Andersen, the rostrum is heavily built and is broader and more massive, with a consider- ably wider interorbital region. The teeth of the male example are similar in size to those of the holotype but in the female specimens are generally slightly smaller, overall much as the teeth of viridis. Andersen (1912) observed correctly that pm| in the holotype of praedatrix, however, are practically as large as in crenulata but in each of the specimens reported here these teeth are smaller than those of crenulata and are nearer in size to pm| of viridis. The labial and lingual longitudinal ridges of pmf and m{ are faintly cuspidate in BM(NH) 69.305 but scarcely if at all cuspidate in BM(NH) 69.304 and 69.306. The longitudinal ridges of these teeth in the holotype are slightly more cuspidate, but the teeth are little worn. As in viridis, crenulata and in the holotype of praedatrix mj and m 2 have well developed surfacial cusps or ridges: the surface cusp of pm 4 is low and very undeveloped. The three specimens closely resemble the holotype in colour. Dorsally, the shoulders and neck are brownish, strongly tinged with black in the male, the head blackish brown, this darker colour extending medially down the nape and neck in a narrow line; the ventral surface of the body is brownish drab, medially with a faint wash of brighter tawny olive, the underside of the neck and chin with a much sparser covering of longer, brownish hairs. The male example is larger cranially than the two female specimens, adding support to Bergmans (1975) who suggested that sexual dimorphism in size might well occur in some at least of Dobsonia. Measurements of d 1 BM(NH) 69.305, 9969.304, 69.306 in that order: length of forearm 116-3, 117-3, 112-2; total length of skull to gnathion 50-6, 48-3, 48-5; condylobasal length 47-6, 45-6, 45-5; condylocanine length 47-3, 45-2, 45-0; length front of orbit-tip of nasals 14-5, 13-3, 13-1; palatal length 25-5, 23-7, 24-2; length palation-incisive foramina 22-6, 21-7, 21-7; length palation-basion 19-1, 18-8, 18-5; lachrymal width 13-8, 12-8, 12-9; least interorbital width 10-1, 9-7, 10-0; least postorbital width 8-2, 7-7, 7-9; zygomatic width 31-4, 30-0, 30-4; width of braincase 20-4, 19-4, 19-6; mastoid width 19-0, 17-8, 17-8; orbital diameter 10-8, 10-8, 10-6; c'-c 1 (crowns) 9-7, 9-4, 9-4, (alveoli) 9-2, 8-8, 8-8; m'-m 1 (crowns) 15-5, 14-7, 14-1, (alveoli) 14-7, 14-0, 1 3-5; c'-c 1 (internally, cingula) 4-0, 4-1, 3-9; pm 4 -pm 4 (internally) 8-0, 7-8, 7-5; width of mesopterygoid fossa 5-8, 5-9, 5-5; c-m 2 20-0, 18-4, 18-5; length complete mandible from condyles 37-8, 36-0, 36-0; length right ramus from condyle 39-2, 37-2, 37-5; coronoid height 21-6,21-7,21 -6; c-m 3 21-1,19-2,19-3. Length/width of cheekteeth: pm 3 4-41/3-08, 4-04/2-81, 4-06/2-84; pm 4 4-22/3-14, 3-97/2-82, 3-95/2-83; m 1 5-31/2-74, 4-73/2-61, 4-70/2-72; m 2 2-45/1-64, 2-06/1-49, 2-15/1-42; pm 2 1-39/1-55, 1-13/1-43, 1-24/1-44; pm 3 4-09/2-61, 3-87/2-39, 3-84/2-32; pm 4 4-42/2-78, 3-89/2-48, 4-01/2-46; m, 4-14/2-46, 3-60/2-21, 3-91/2-20; m 2 3-31/2-37, 2-98/2-02, 2-96/2-01 ;m 3 1-95/1-55, 1-79/1-46, 1-60/1-33. DISCUSSION. Examination of these adult examples confirms that Bergmans (1975) correctly rejected the suggestion by Pohle (1953) that praedatrix should be allied subspecifically to D. viridis, but I am less convinced that it should be removed from the viridis group as the former author (1978) has since suggested. If dental characters are to remain the chief criteria by which the species of Dobsonia are classified, then praedatrix must be included in the viridis group as Andersen (1909, 1912) envisaged it, except that the degree of cuspidation of the longitudinal ridges of pm| and m{ is less than is general in the group or is sometimes virtually absent. In this respect praedatrix approaches the peronii group of Andersen (loc. cit.) in which these ridges are simple, but m 1 reputedly lacks the well marked antero-internal basal ledge characteristic of the viridis group. The limited material of D. peronii in London INDO-AUSTRALIAN BATS 1 1 3 indicates that in this species m 1 has at least a small antero-internal ledge and that the character may not be as emphatic as Andersen implied, although the ledge is much more developed in D. viridis and its allies. As Bergmans (1978) suggested, D. peronii, by current classification the sole member of the peronii group, may be more closely related to the viridis group than to any other but as this author pointed out, D. peronii differs from D. viridis and from its allies in the outline of the rostrum, which is lower, relatively longer and curves downward less abruptly from the braincase. Bergmans also remarked that within the viridis group peronii comes closest to the viridis-crenulata-beauforti (sub)group (sic) but dentally it seems to approach more closely to praedatrix. The greenish coloration of D. viridis when alive also appears in D. peronii peronii according to Goodwin (1979). Bergmans (1978) advanced the opinion that the inclusion of praedatrix from the Bismarck Archipelago and inermis (including nesea) from the Solomon Islands in the viridis group indicated a disregard of zoogeographical considerations. I find no conviction in this assertion since the group as presently constituted occupies the arc of islands to the west, northwest, north and northeast of New Guinea, perhaps without interruption since some remain zoologically poorly known. A biogeographic model of this nature was suggested by Smith & Hood (1981). Indeed, if D. peronii is regarded as a member of the viridis group it then includes a succession of island species and subspecies extending from Nusa Penida (near Bali) in the west to the Solomon Islands in the east, but apparently excluding New Guinea. These considerations suggest that the peronii and viridis groups of Andersen (1909, 1912) might be merged to form a peronii group that includes D. peronii, D. viridis and its close allies, D. praedatrix and D. inermis. It can be summarized: D. peronii grandis Bergmans, 1978 Nusa Penida I, Sumbawa I. D. peronii sumbana Andersen, 1 909 Sumba I. D. peronii subsp. W. Flores I, Alor I, (see Bergmans, 1 978) Wetar I, Babar I. D. peronii peronii (Geoffrey, 1810#) Timor I. D. viridis (?) viridis (Heude, 1 896) Sulawesi, Amboina I, Buru I, Ceram I, Banda Is, Kei Is. D. viridis (?) crenulata Rau I, Morotai I, Andersen, 1909 Halmahera I, Ternate I, Batchian I, Togian Is, Sanghir ( = Sangihe) Is. D. beauforti Bergmans, 1 975 Waigeo I. D. praedatrix Andersen, 1 909 New Britain, New Ireland, Duke of York I. D. inermis nesea Andersen, 1 909 N and WC Solomon Is: Shortland, Alu, Ghizo, Rubiana, Bougainville, New Georgia. D. inermis inermis Andersen, 1 909 S and EC Solomon Is: San Christoval, Ugi, Ysabel, Rennell. Phillips (1968) considered nesea a synonym of inermis, but McKean (1972) retained it as a valid subspecies on the basis of Troughton's (1936) assertion that inermis is darker in colour. The occurrence of D. viridis in Sulawesi and (as crenulata) on the Togian and Sanghir Islands also has some bearing on Bergman's (1978) view of possible former distribution routes for Dobsonia towards the Lesser Sunda Islands. He suggested that D. peronii and D. moluccensis or their ancestors in these islands (there is only one local record of the latter, from Semau ( = Samoa) Island, near Timor) might have originated from New Guinea (moluccensis} or from the Aru, Kei or Timorlaut Islands (peronii), possibly moving along 114 J. E. HILL former land bridges, or alternatively along a more northerly route through the southern Moluccas and Sulawesi. A species of the moluccensis group, D. exoleta, has been known from Sulawesi for many years and Bergmans pointed out that the concept of the more northerly route implied that it is the closest living relative of D. peronii, although he considered exoleta to be less closely related to peronii than to the members of the viridis group. If Bergman's views are accepted the presence of D. viridis in Sulawesi resolves this apparent paradox. Dobsonia moluccensis moluccensis (Quoy & Gaimard, 1830) Hypoderma moluccensis Quoy & Gaimard. 1830.1 : 86, Atlas, pi. 11. Amboina. SPECIMENS EXAMINED. Molucca Is: BM(NH) 75.2140 Lihura limestone caves, near Ruhuwa, SC Ceram I (crania, mandible, fragments, teeth; coll. R. F. Ellen). REMARKS. This cave material compares favourably with D. m. moluccensis from Buru, Amboina and Ceram. Length of c-m 2 (alveoli) in two examples 23- 1 , 23-5. Dobsonia moluccensis magna Thomas, 1905 Dobsonia magna Thomas, 1905# : 423. Tamata, Mambare R, Papua New Guinea, 100 ft. SPECIMENS EXAMINED. Papua New Guinea: 9 BM(NH) 69.307 Brown R, near Port Moresby, c. 9 27 ' S, 147 08 ' E; d BM(NH) 69.308 Tupuselaia ( = Tupuselei), 9 33 ' S, 147 19 ' E (both skins, skulls; coll. J. I. Menzies): d 1 BM(NH) 73.1969 Kairiru Ridge, centre of Kairiru I, near Wewak, East Sepik, c. 2000ft; <5 BM(NH) 73.1970 Victoria Bay, NW end of Kairiru I (both in alcohol; coll. Aberdeen University Exploration Society Expedition to New Guinea, 1973); BM(NH) 78.202-209 ? Near Mt. Hagen, Western Highlands (crania, damaged); BM(NH) 78.210-211 Baiyer R, Western Highlands (mandibles, damaged); BM(NH) 78.212-249 Tuman, Kubor Range, Western Highlands (crania, damaged); BM(NH) 78.250-255 Upper Lai Valley, Southern Highlands (crania, mandibles, damaged); 9 BM(NH) 78.854 About 10 km S of Madang(in alcohol); d BM(NH) 78.855 About 10 km S of Madang, c. 40 m (skin); rf BM(NH) 78.856 Baiyer R, c. 50 km NW of Mt. Hagen, Western Highlands, 1300 m (in alcohol) (all coll. P. A. Morris); BM(NH) 79.2015 (skin), BM(NH) 78.3004 (skull) Haelaelinga Settlement, on Was ( = Wage) R, Nipa, Southern Highlands (coll. P. Sillitoe); rf, 9 BM(NH) 80.528-529 Buso, Morobe, 7 17 ' S, 147 08 ' E (in alcohol; coll. B. H. Gaskell, 'Operation Drake'). REMARKS. The cranial and mandibular material BM(NH) 79.202-255 was obtained from kitchen middens: most show signs of heating or burning. The rear of the cranium in such specimens has been broken open to allow the extraction of the brain, a circumstance noted by Menzies (1977) of material from similar accumulations at the Kiowa and Yaku rock shelters. Dobsonia exoleta Andersen, 1909 Dobsonia exoleta Andersen, 1909 : 531, 533. Tomohon, Minahassa, Sulawesi. SPECIMENS EXAMINED. C Sulawesi: cf BM(NH) 8 1 . 1 1 09 Ganda Ganda, 1 57 ' S, 1 2 1 2 1' E (in alcohol, skull extracted); <5 BM(NH) 81.1238 Tapu Waru, 1 51' S, 121 22' E (in alcohol) (both coll. B. H. Gaskell, 'Operation Drake'). REMARKS. The relatively unworn dentition of BM(NH) 81.1109 agrees closely with the holotype (BM(NH) 99. 1 0. 1 .4) and with the account by Andersen (1912). The antero-internal corner of pm 4 is developed into a conspicuous ledge with elevated rim; centrally the ledge is raised into a small cusp; pm 3 has a similar, narrower ledge; the antero-internal corner of m 1 is INDO-AUSTRALIAN BATS 1 1 5 low and platform-like but little differentiated except by a shallow antero-internal notch in the longitudinal ridge; a slight antero-internal basal ledge in pm 3 , rather more developed in pm 4 ; m, with simple lingual ridge, no trace of an antero-internal ledge or cusp. There is a well developed posterior basal ledge on pm 3 , a similar but slightly narrower ledge on pm 4 ; pm 3 has likewise a slightly wider posterior basal ledge than pm 4 . Median surface cusps or ridges are well developed in mj and m 2 , in the latter the ridge extending uninterruptedly through the length of the tooth; there is a low surface cusp on the posterior face of pm 4 that is absent from the more eroded dentition of BM(NH) 81.1238. Measurements (rf BM(NH) 81.1 109, rf 81.1238, skull of 81.1 109): length of forearm 1 16-8, 1 16-7; total length of skull to gnathion 5 1 -7; condylobasal length 49-9; condylocanine length 49-8; length front of orbit-tip of nasals 14-4; palatal length 25-8; length palation-incisive foramina 23-4; length palation-basion 20-8; lachrymal width 12-4; least interorbital width 8-4; least postorbital width 7-0; zygomatic width 31-8; width of braincase 20-3; mastoid width 19-5; orbital diameter 11-1; c'-c 1 (crowns) 10-5, (alveoli) 9-7; m'-m 1 (crowns) 16-9, (alveoli) 15-9; c^c 1 (internally, cingula) 4-1; pm 4 -pm 4 (internally) 8-2; width of mesopterygoid fossa 6-1; c-m 2 22-0; length complete mandible from condyles 39-7; length right ramus from condyle 4 1 -0; coronoid height 21-2; c-m 3 23-3. DISCUSSION. Dentally D. exoleta agrees with D. moluccensis from the Molucca Islands, New Guinea and from some of its associated islands rather than with D. peronii from the Lesser Sunda Islands. The Sulawesian species differs most conspicuously from peronii in its lack of differentiation of the antero-internal corner of m, into a distinct cusp or small ledge, the inner or lingual ridge of the tooth being perfectly simple and lacking any notch or division. Andersen (1912) drew attention to a number of similarities between exoleta and D. pannietensis (De Vis, 1905) from the Trobriand Islands, which he thought differed from exoleta only in smaller size and in the lack of surface ridging on m p so considering the two taxa closely related. Later, Thomas (1914^) in describing D. anderseni remarked that this form from the islands of Manus and Ruk in the Bismarck Archipelago was intermediate in size between exoleta and moluccensis, an opinion confirmed to some extent by Bergmans (1979) who has examined specimens from a number of other islands in the Archipelago. As in pannietensis the dentition of anderseni is like that of moluccensis. Laurie & Hill (1954) recognised exoleta as a distinct species but listed both pannietensis and anderseni as sub- species of D. moluccensis. More recently, De Jong & Bergmans (1981) reviewed known specimens of exoleta in some detail and considered it to be a distinct species related in dental morphology to chapmani Rabor, 1952 from the Philippines, moluccensis, anderseni and pannietensis. Bergmans (1975, 1978, 1979) has discussed and re-examined pannietensis and anderseni. He concluded (1975, 1979) that both should be considered specifically distinct on account of their smaller size when compared with D. moluccensis moluccensis and D. m. magna but it is not clear to what extent they differ from each other. This author (1975) suggested that to consider all of these conspecific involved the acceptance of an exceptional size range within one species, although (1979) he has himself accepted a considerable range of size among specimens from the islands (Louisiades, D'Entrecasteaux, Trobriand, Woodlark) immedi- ately to the east of New Guinea that he considers all referable to pannietensis. Examination of the measurements provided by Bergmans (1979) suggests that some at least of the speci- mens that he refers to pannietensis are similar in size to anderseni: indeed, they overlap the measurements given by this author (1975, 1979) for the latter, although admittedly sexual differences in size may be involved. Moreover, Koopman (1979) in recording specimens from some of the small islands (Karkar, Bagabag, Umboi) off northeastern New Guinea and from the Bismarck Archipelago suggested that populations intermediate between magna and anderseni are to be found on Karkar and Umboi. This author has since (1982) discussed the question in some detail and while retaining anderseni as a subspecies of D. moluccensis considered pannietensis specifically valid, at least until the genus is fully revised. Insufficient specimens are available in London to determine the matter definitively, but for the present I 116 J. E. HILL am unconvinced that anderseni and pannietensis should be considered species distinct from moluccensis: Bergmans (1978) himself admits that both are more closely related to moluccensis than to other species of the genus. Bergmans (1979) provided a brief discussion of exoleta and compared it with D. moluccensis pannietensis, drawing attention to a number of differences. Of these, the relatively narrower interorbital and postorbital regions of exoleta also separate it (in the limited sample available) from D. m. anderseni, D. m. magna and D. m. moluccensis. For the present, therefore, I retain it as a distinct species and agree with Andersen (1912) in regarding it as the Sulawesian representative of the moluccensis group. The species is known to occur (De Jong & Bergmans, 1981) on Sulawesi, on the Togian Islands (Malenge) in the Gulf of Gorantalo, and on Muna Island. Dobsonia minor (Dobson, 1879) Cephalotes minor Dobson (1878), 1879 : 875. Amberbaki, NW New Guinea. SPECIMENS EXAMINED. Papua New Guinea: 9 9 BM(NH) 73.1971-1973 Rauit, 3 36' S, 142 15' E (in alcohol); BM(NH) 74.337-338 Near Rauit (skulls) (all coll. Aberdeen University Exploration Society Expedition to New Guinea, 1973). Cynopterus brachyotis brachyotis (Muller, 1838) Pachysoma brachyotis Muller, 1838 : 146. R Dewei, Borneo. SPECIMENS EXAMINED. Burma: 3 cTrf, 4 99 BM(NH) 78.64-70, 99 BM(NH) 78.151-153 British Embassy Residence Compound, Rangoon (in alcohol; coll. D. W. & G. Walton). N Sumatra: dd, 9 BM(NH) 81.675-677 Bohorok R, near Bukit Lawang, Langkat Reserve, Gunung Leuser Reserve complex (in alcohol; coll. R. Aveling). S Sumatra: dd BM(NH) 78.1114-1115 Banda hurip, Pulas District, S Lampong; (?), 9 BM(NH) 78.1116-1117 Asahan, Jabung District, Lampong; 9 BM(NH) 78.1118 Lebung Dadup, Asahan, Jabung District, Lampong; 9 BM(NH) 78.1 1 19 Sukaraja tiga, Sukadana District, Lampong; d BM(NH) 78.1120 Pring Kumpul, Pring Sewn District, Lampong; 3d BM(NH) 78.1122-1123 Air Nanigan, Pulau Penggung District (all skins, skulls; presented by Museum Zoologicum Bogoriense). N Sulawesi; d (?), (?) BM(NH) 78.968-970 Tangkopo Batuangus, near Bitung (in alcohol; coll. A. M. Jones). C Sulawesi: (?) BM(NH) 79.2333-2337 Shore of Lake Matano, c. 5 km W of Soroako (in alcohol, very bad condition; coll. P. Holmes); 21 dd (2 yg.), 1 9 9 , neonate BM(NH) 81.1 006- 1 036, R Ranu, 1 51' S, 121 30' E:2 dd, 5 99 (2 yg.) BM(NH) 8 1.1037-1044 GandaGanda, 1 57' S, 121 21' E (all in alcohol; coll. B. H. Gaskell, 'Operation Drake'). Banggui Archipelago: cTBM(NH) 81.1045 Potil Besar 1; rfdBM(NH) 81.1046-1048 Kelara, Besar 1 (all in alcohol; coll. B. H. Gaskell, 'Operation Drake'). REMARKS. Although C. brachyotis is a relatively commonly collected bat in southeastern Asia these specimens are of especial interest since those from Rangoon appear to be the first recorded from Burma and the furthest west the species has been reported on the Asian mainland while there have been few previous records from Sumatra and Sulawesi. Specimens of C. brachyotis from southern Sumatra are from an area that proves to be of particular taxonomic interest when the status of its congeners sphinx and titthaecheileus is considered. In size all of these specimens agree well with the extensive representation of C. b. brachyotis in the British Museum (Natural History) which suggests a range of forearm length of 57-66 for this subspecies over its range from Burma and Indochina to Sumatra and Sulawesi. Specimens from southern Sumatra with length of forearm 61-0-65-7 and m'-m 1 (crowns) 8-2-9-0 approach the weakly defined subspecies C. b. javanicus Andersen, 1910 from Java but fall within the limits given by this author for C. b. brachyotis. INDO-AUSTRALIAN BATS 1 1 7 DISCUSSION. Cynopterus minor Revilliod, 1911. The status of this nominal taxon must remain uncertain for the present. It is known so far apparently only from the holotype from Lambuja, SE Sulawesi, a small individual with a forearm length of 53 that may be a young adult or unusually small example of C. brachyotis. It seems unlikely, however, that Revilliod should fail to recognise it as such. In addition the name minor poses an involved nomenclatorial problem. It was proposed by Lyon in 1 908 as Niadius minor for the taxon subsequently recognised as the Sumatran representative of Cynopterus horsfieldi by Andersen (1912) who considered Niadius Miller, 1906 a synonym of Cynopterus. As a result, Andersen (1912 : 827) regarded Cynopterus horsfieldi minor (Lyon, 1908) to be preoccupied by the combination Cynopterus (Cynonycteris) minor used by Trouessart (1878) for the species otherwise known as Rousettus minor (Dobson, 1873<2,6), as it is today. In rejecting minor of Lyon, 1908 on this account Andersen proposed Cynopterus horsfieldi lyoni in its place, a substitute that subsequently (Robinson & Kloss, 1918, 1919; Chasen, 1940) came into use. Hill (196 la) reviewed the circumstances surrounding this situation and on the basis of verbal advice in 1960 from the International Commission for Zoological Nomenclature that secondary homonyms are not to be permanently rejected concluded that since Cynopterus (Cynonycteris) minor as used by Trouessart referred to a Rousettus the name minor as proposed by Lyon in 1908 should become again the valid subspecific epithet for the Horsfield fruit bat of Sumatra and Malaya. However, the current International Code of Zoological Nomenclature provides (Art. 59, (b), (i)) that a junior secondary homonym rejected before 1961 is permanently rejected and cannot be restored unless the employment of the replacement name is contrary to existing usage. Thus Cynopterus horsfieldi lyoni Andersen, 1912 must become the valid name for the Sumatran and Malayan representatives of C. horsfieldi. At the same time the Code (Art. 59, (b), (ii)) states that if the secondary homonymy has been overlooked or the junior name not replaced, and the taxa in question are no longer congeneric, the junior name is not to be rejected, even though one name was originally proposed in the current genus of the other. Therefore Cynopterus minor Revilliod, 1911, having by these rulings no competitor within Cynopterus (minor of Lyon, 1908 having been rejected and replaced by lyoni of Andersen, 1912) remains a valid name in this genus, and is unaffected by the combination Cynopterus (Cynonycteris) minor used by Trouessart (1 878). Cynopterus sphinx angulatus Miller, 1 898 Cynopterus angulatus Miller, 1 898 : 3 1 6. Trang, S Thailand. SPECIMENS EXAMINED. S Sumatra: 99 BM(NH) 78.1112-1113 Wai Miring, Kota Agung District, Lampong; rf BM(NH) 78.1121 Pring Kumpul, Pring Sewn District, Lampong; 5o o op^ I I I I I I I I I O (NsOsOONOOfN J^ I I Tt SO so OsOsOsoOsfNsOOOONONO o so t~- OO oo O o^oo ON I I I I *>O OO O*^ OO O* 1 * ^^ ^"^ ^^ i fsj ^N) ^^ o" O~ o" O" O" O" O' C2 O~ O~ oo Os" oT O O ON 1 i/->t--OOOOON ON OO CM o I 120 J. E. HILL the other two species. Dammerman (1938) while reporting C. sphinx angulatus from Krakatoa and Verlaten I also recorded a specimen of C. titthaecheileus from the nearby island of Sebesi. DISCUSSION. Andersen (1912) associated titthaecheileus with C. sphinx as a subspecies, notwithstanding that his arrangement of this part of the genus involved a wide geographical hiatus between C. s. sphinx in India, Burma and northern Thailand and 'C. s. titthaecheileus' in Sumatra and Java. This author considered angulatus, which fills this gap, to be a subspecies of C. brachyotis, a circumstance leading him to the conclusion that sympatric subspecies of C. brachyotis of different geographic origins, namely C. b. brachyotis and C. b. angulatus were to be found in the Malay Peninsula and in Sumatra. Although initially challenged by other authors, notably by Kloss and by his colleague Robinson (Kloss, 1911, 1916, 1917, 1919, Andersen & Kloss, 1915, Robinson & Kloss, 1915a, 19156, 1918) who thought that angulatus might more properly be considered a subspecies of C. sphinx, this concept prevailed until quite recently. Chasen (1940) attempted to resolve the distributional dilemma created by Andersen by limiting C. b. brachyotis to Malaya, Sumatra, Borneo and some of the associated islands, C. b. angulatus to southern Thailand and its coastal islands, and to the Natuna and Anamba Islands. Thus this author envisaged a large form in the north of the Malay Peninsula, a smaller form occupying the rest of the area from northern Malaya to Borneo, Sumatra, and, as C. b.javanicus Andersen, 1910, to Java. Hill (1961> Q fN O T3 cS | 55 oo u 3 1 1 _ ^t r- oo O m O oo ON r- O 3 _^ O O so ^t r- so o "CJs ^, r-- >n m so fl r*^ r m co r ^c Q . ; | ^< 1 CO fN OO so ^ m r- T^- ^- OO fN O m oo ZN S V) c ^ r^- >n so oo oo m so ^t Tf 1 1 1 r- r- m t-j- CO CO ON oo ON Sao CQ rn 5^ r- oo oo m ON in fN co iTso Si g so ^3" m so r^ m so co co ON'""' >, s: g*; s^< o ^^ Cd """ S ^ ^ 3 o ^r - ^- _L oo r- oo ON m O fN r~ o r- oo co so so c?N 05 &^ so ^~ in SO CO Tf SO CO CO -LT "U 5j = 1 c O CO C ^, CO , OO , ^H Q so i n ^D so f") ^" ^D CO CO t fNJ O 12 SDO _' OO b: *"" 00 ~". -2 5 | oo r~ _ oo o O Tf ON O so <"*"> ^t" OO OO SO so in m SO CO CO II s: ^2 s CQ CQ 6 fN O i'"' in ON fN r<^ ON fN O ON co m in m m og CM SO ^J" SO SO ^ ^ so co co ^^ ^ ^ O ^ II 1 so 'T fN in 311 00 O *A *^ CO ro t r O so oo fN fN CO oo O o \\ so ^J* in SO CO CO 21 CO CO ON fN Z, m oo fN OO 00 fN c^ r-~ fN fN fN Tf fN fN - SO ^ SO so f"l "^t* SO CO CO ^~" o gcQ C/3 O MX '5 c2 .S C si 00 c 8 ^ '5 ** t~^ "^ u - 'so a ff t t/3 D.^C CtM HrZ 2 E _ >> i. CQ 126 J. E. HILL 4-4-^-5, (alveoli) (3) 4-1-4-2, (cingula, internally) (3) 1-9-2-2; pm 4 -pm 4 (crowns) (3) 6-4-6-8, (alveoli) 6-1-6-6, (internally) (3) 3-8-4-1; m'-m' (crowns) 6-1,6-5, (alveoli) 6-1-6-4; width mesopterygoid fossa 3-1-3-3; c-m 1 (crowns) (3) 6-6-7-3; length complete mandible from condyles 15-1-16-3; length right ramus from condyle 16-1-17-2; coronoid height 7-6-8-7; c-m 2 (crowns) 7-5-7-9. Thoopterus nigrescens (Gray, 1870) Cynopterus marginatus var. nigrescens Gray, 1 870 : 123. Morty ( = Morotai) Island. Cynopterus latidens Dobson, 1878 : 86. Morty ( = Morotai) Island. SPECIMENS EXAMINED. C Sulawesi: 9, 2 dd BM(NH) 81.1049-1051 R Ranu, 1 51 ' S, 122 22' E; 8 rfrf, 6 99 BM(NH)81.1052-1065 Tambusisi Damar, Mt. 4000 ft. 1 39' S, 121 22 ' E (in alcohol. BM(NH) 8 1.1054, 81.1056, 81.1058-1061, 81.1063-1064 heads only; coll. B. H. Gaskell, 'Operation Drake'). REMARKS. There are few records of this long-haired cynopterine fruit bat which is known so far with certainty from Morotai Island and from Menado and Minahassa in northern Sulawesi. Its reported occurrence on Luzon Island in the Philippines is thought doubtful by Taylor (1934) although it may extend to the southernmost of these islands. Length of forearm in four examples 76-5-78-9. DISCUSSION. This large bat is similar in many respects to the relatively recently described genus Latidens Thonglongya, 1972 from southern India, which is almost identical in external form and colour and has a similarly long, strong rostrum. Dentally, however, Latidens approaches the Malaysian genus Penthetor, differing in the outline of m 1 , which is more or less square and not wedge-shaped, and in the wider, rather more square outline of pm 4 and m,. The latter teeth, however, also have a low surface cusp and in this respect approach Thoopterus. These and other features suggest that Thoopterus is represented in Malaysia by Penthetor as was thought by Andersen (1912) and in southern India by Latidens. The three genera can be readily distinguished from each other by the incisive dentition, Thoopterus having incisors \ \, Penthetor \ \, and Latidens \ \. Aethalops alecto (Thomas, 1923) Aethalodes alecto Thomas, 1923a : 251. Indrapura Peak, Sumatra, 7300 ft. SPECIMENS EXAMINED. W Java: _d f ^ ' 1^ '3p " .^ __ _- Q\ oo OO ON *S~t f^ ~" "M" ^ r^ f*^ ro r**) ro ro r^j P ^^ U _c c o. 2 O sp /^m_ sO . I "^>ro ^S E E c!> 148 J. E. HILL external features they considered tatar to be nearest to R. arcuatus. However, Bergmans & Rozendaal (1982) appear to have made no direct comparison with any representative of R. arcuatus, and of R. euryotis only with examples of the nominate subspecies from Amboina. Comparison with all but angustifolius among R. arcuatus, and with all of the described form of R. euryotis leads me to the conclusion that tatar is best considered a subspecies of this latter, a finding that confirms the allocation of similar specimens from Sulawesi to R. euryotis by Tate & Archbold (1939). In view of the relatively small size of R. e. tatar it is of some interest to remark that Koopman (1982) has reported three specimens of R. euryotis from Kiriwina island, off eastern Papua New Guinea that are of similar size (Table 4), and one that is little larger (approaching R. e. timidus) from New Britain, but Smith & Hood (198 1 ) have also recorded a much larger example from the same island. Koopman (1982) was inclined to doubt the validity of some at least of the subspecies ofR. euryotis then recognised. HIPPOSIDERIDAE Hipposideros bicolor bicolor (Temminck, 1834) Rhinolophus bicolor Temminck, 1834: 19, pi. 1, fig. 3; 1835: 18 (further description). Lectotype designated and type locality restricted to Anjer coast, northwestern Java by Tate ( 1 94 1 a). Hipposideros javanicus Sody, 1937 : 215. Babakan, Kroja, Tjilatjap, central Java. SPECIMEN EXAMINED. W Java: 9 Rijksmuseum van Natuurlijke Historic 29304 Tjilatjap (skin, skull; coll. 25 October 1929, apparently by H. J. V. Sody). REMARKS. This specimen appears to be one of those recorded by Sody (1930) as H. galeritus longicauda (Peters, 1861). However, the noseleaf lacks the lateral supplementary leaflets characteristic of//, galeritus and its allies and the specimen proves on further examination to represent H. bicolor bicolor. Dorsally, it is pale brown, the hairs with creamy or whitish bases that show through the darker tipping: the throat and chest are whitish, the remainder of the ventral surface brownish buff. The skull is elongate in outline, the rostrum and palate sharply tapered anteriorly; the narial swellings are only slightly inflated and there is a low sagittal crest; the zygomatic width is less than the mastoid width, the zygoma robust with low jugal eminence; the interparietal region is swollen and rather pronounced; the palation is shallowly V-shaped with wide mesopterygoid fossa; the sphenoidal bridge is wide, partially concealing elongate lateral apertures and there is a shallow oval sphenoidal depression; the cochlea are a little wider than their distance apart. The anterior upper premolar (pm 2 ) is very small, slightly extruded into a recess between the canine and the second upper premolar (pm 4 ); the posterior ridge of m 3 is about one half the length of the anterior ridge; the crown area of the outer lower incisors is very slightly greater than the crown area of the inner pair; anterior lower premolar (pm 2 ) threequarters the length and two thirds the height of the second lower premolar (pm 4 ) and two thirds its crown area. Measurements: length of forearm 45-8; ear not measurable; length of tail c. 39; length of tibia c. 21; length of foot (c. u.) c. 8; greatest length of skull 19-0; condylobasal length 17-0; condylocanine length 16-8; basal length 14-6; palatal length 6-5; width across rostral swellings 4-8; least interorbital width 2-9; zygomatic width 9-3; width of braincase 8-7; mastoid width 9-6; c'-c 1 (alveoli) 4-1; m 3 -m 3 6-1; c-m 3 6-5; m 1 " 3 4-0; length complete mandible from condyles 1 1 -7; length right ramus from condyle 12-0; c-m 3 7-0. DISCUSSION. This example agrees closely with the account by Tate (194 la) of the lectotype of H. bicolor (Temminck, 1834) in the Rijksmuseum van Natuurlijke Historic and with the description of H. javanicus by Sody (1937), thought by Tate (loc. cit.) and by Hill (1963) to be a synonym of bicolor. Sody almost concurrently (1937&) recorded javanicus from the island of Banka: the specimen that he reports is slightly smaller in some respects either than the holotype of javanicus or the specimen of bicolor from Java reported here. There seems little doubt \hatjavanicus must be considered a synonym of//, b. bicolor. INDO-AUSTRALIAN BATS 149 Hipposideros ater aruensis Gray, 1858 HipposiderosaruensisGray, 1858 : 107. Aru Islands. SPECIMENS EXAMINED. Papua New Guinea: 99 BM(NH) 73.2036-2037 (both yg.) Kairiru Cave, near St Xavier's Mission, Kairiru I, near Wewak, East Sepik, c. 3 2 1 ' S, 1 43 36 ' E (in alcohol; coll. Aberdeen University Exploration Society Expedition to New Guinea, 1973); d BM(NH) 80.612 S Tunnel, Bulolu Gorge, Wau, Morobe (in alcohol; coll. B. H. Gaskell, 'Operation Drake'). REMARKS. Little material is available from the more eastern part of the range of//, ater and for the present, following Hill (1963), specimens from New Guinea are referred to H. a. aruensis. McKean (1972) records specimens from Ambunti, Papua New Guinea. Length of forearm (BM(NH) 80.6 1 2) 40-9. Hipposideros maggietaylome erroris Smith & Hill, 198 1 Hipposideros maggietaylorae erroris Smith & Hill, 1981 : 14. Yaguam Sulfur Cave, 5 miles S and 3 miles W of Madang, Madang Province, Papua New Guinea, 5 1 7 ' S, 145 45 ' E. SPECIMENS EXAMINED. Papua New Guinea: 4 dd (2 yg.), 499 BM(NH) 73.2038-2045 Kairiru Cave, near St Xavier's Mission, Kairiru I, near Wewak, East Sepik, c. 3 21' S, 143 36' E (in alcohol; coll. Aberdeen University Exploration Society Expedition to New Guinea, 1973); 99 (1 yg.) BM(NH) 75.1863-1865 St Xaviers, Kairiru I (in alcohol, skull of BM(NH) 75.1864 extracted; coll. A. M. Jones, Aberdeen University Exploration Society); d, 9 BM(NH) 78.875-876 About 10 km S of Madang, 5 1 5 ' S, 145 45 ' E (in alcohol; coll. P. A. Morris); d 1 , 9 9 (1 yg.) BM(NH) 80.638-643 War Tunnel, Salamana, Morobe, 7 03' S, 147 03' E (in alcohol; coll. B. H. Gaskell, 'Operation Drake'). REMARKS. These specimens are listed by Smith & Hill (198 1) in their account of//, maggie- taylorae: those obtained by 'Operation Drake' are recorded erroneously as BM(NH) 80.516-522 (the suffixes in fact their collection numbers). Hipposideros ridleyi Robinson & Kloss, 1911 Hipposideros ridleyi Robinson & Kloss, 1911 : 241 Botanic Gardens, Singapore. SPECIMEN EXAMINED. Borneo: d BM(NH) 82.160. Sepilok Forest Reserve, Sabah (in alcohol; coll. P. Zborowski). REMARKS. This specimen is the first of H. ridleyi to be recorded from Borneo, the species being otherwise known from the type locality and from Selangor on the Malayan mainland (Medway, 1978). It may be easily recognised by its very large sub-triangular ears (length from meatus c. 21-23) broad noseleaf that lacks lateral supplementary leaflets, the expansion of the internarial septum to form a concave, saucer-like disc between and in front of the nostrils and by its high posterior leaf, the upper part supported by three septa defining four deep pockets. Length of forearm in the Bornean example 48-1, in the holotype (BM(NH) 61.329) 47-3 and in a specimen (BM(NH) 75.2000) from a locality between Kuala Kangsar and Rawang, Selangor, 48-5. Hipposideros cervinus cervinus (Gould, 1863) Rhinolophus cervinus Gould, 1863 : pi. 34, letterpress. 'Caves on Albany Island' (label on skin of holotype). Cape York, Queensland, Australia. 150 J. E. HILL SPECIMENS EXAMINED. C Sulawesi: 8 dd, 5 99 BM(NH) 82.99-1 1 1 Taronggo, 1 44' S, 121 40' E; 99 BM(NH) 82.112-113 R Ranu, 1 51' S, 121 30' E (all in alcohol, skulls of BM(NH) 82.100, 82.109-1 10, 82. 1 12 extracted; coll. B. H. Gaskell, 'Operation Drake'). S Sulawesi: 99 BM(NH) 82.1 14-1 15 Lalonggasu Meeto (Tomba Watu Cave), 18 km from Kendari Central (in alcohol, skull of BM(NH) 82. 1 1 5 extracted; coll. B. H. Gaskell, 'Operation Drake'). Papua New Guinea: dd BM(NH) 75.1866-1867 Wageo I, Schouten Is, East Sepik; 9 BM(NH) 75. 1 868 Kadovar I, Schouten Is; d , 3 9 9 BM(NH) 75. 1 869-1 872 East Sepik (all in alcohol; coll. A. M. Jones, Aberdeen University Exploration Society); d BM(NH) 76.386 Karkar I, Madang (in alcohol; coll. H. King); 9 BM(NH) 78.877 About 10 km N of Baku, Gogol Valley, Madang, c 40 m (flat skin); 4 dd, 1 99 BM(NH) 78.878-888 8 km W of Baku, c. 50m; (in alcohol) (all coll. P. A. Morris); 99 BM(NH) 80.607-6 1 1 N Tunnel, Bulolu Gorge, Wau, Morobe, 7 1 9 ' S, 1 46 44 ' E; d BM(NH) 80.6 1 2 S Tunnel, Bulolu Gorge; d BM(NH) Lae, Morobe, 6 49 ' S, 1 47 03 ' E; 9 9 BM(NH) 80.6 14-617 Buso, Morobe, 7 1 7 ' S, 1 47 08 ' E; 6 dd, 8 9 9 BM(NH) 80.6 1 8-63 1 Siboma, Morobe; 2 dd , 4 9 9 BM(NH) 80.632-637 Coastal cave W of Kui Village, Morobe, 7 22' S, 147 1 1' E (all in alcohol; coll. B. H. Gaskell, 'Operation Drake'). New Hebrides: 2 dd, 2 99 BM(NH) 73.1334-1337 Grotte Montmartre, Port Vila, Efate I, 20 m; 6 dd, 3 99 BM(NH) 73.1338-1346 Mission Montmatre, Port Vila; 4 dd, 2 99 BM(NH) 73.1347-1352 'Pig Cave', Harris Plantation, N coast Efate I, 40 m; c?, 9 BM(NH) 73.1353-1354 'Pig Cave', Narabut Camp, Efate I; 3 dd, 3 99 BM(NH) 73.1355-1360 Lomboh Cave, Litzlitz, Port Stanley Bay, Malekula I, 5m; cf, 3 99 BM(NH) 73.1361-1364 Lipelip Cave, Amok, Malekula I, 440m; 2 dd, 9 BM(NH) 73.1365-1367 Aouta Plantation, Aore I; dd BM(NH) 73.1368-1369 New Hebrides (all in alcohol, obtained by Earl of Cranbrook, Royal Society Expedition to the New Hebrides, 1973). REMARKS. Specimens from Sulawesi appear to be the first of//, c. cervinus to be recorded as such from that island, although earlier records of//, galeritus galeritus from Gimpoe or Bada and Peleng Island by Shamel (1940) and of E. galeritus celebensis from Talassa and Banti-moerang by Tate ( 1 94 1 a) may refer in fact to cervinus. A recent study at the British Museum (Natural History) by Jenkins & Hill (1981) has indicated that two species, namely //. galeritus Cantor, 1846 (including insolens Lyon, 1911 as a subspecies) and H. cervinus (including labuanensis Tomes, 18590 as a subspecies) occur in Borneo. Hipposideros celebensis Sody, 1936 based on specimens initially referred by this author (1930) to H. galeritus galeritus, has from the measurements given by Sody a broader braincase and shorter toothrow than the Sulawesian specimens here referred to H. cervinus, but Jenkins & Hill found two paratypes of celebensis to have longer toothrows and narrower zygomatic and braincase widths than Sody's measurements indicated. The paratypes unquestionably represent H. cervinus and these authors suggested from the available evidence that only H. c. cervinus occurs in Sulawesi, specimens from that island approaching the more easterly nominate subspecies rather than the geographically nearer Bornean subspecies //. c. labuanensis. Measurements of specimens from Sulawesi: length of forearm (17) 45-0-49-1; condy- locanine length (5) 14-3-14-9; width of rostrum (5) 4-9-5-1; least interorbital width (5) 2-5-2-8; zygomatic width (5) 8-8-9-1; width of braincase (5) 7-7-8-0; mastoid width (5) 8-6-8-9; c'-c 1 (alveoli) (5) 3-7-3-9; m 3 -m 3 (5) 6-0-6-1; c-m 3 (5) 5-8-6-0; length complete mandible from condyles (5) 10-3-10-8; length right ramus from condyle (5) 10-7-1 1-1; c-m 3 (5)6-3-6-4. Hipposideros diadema pullatus Andersen, 1 905 Hipposideros diadema pullatus Andersen, 1905c: 498. Haveri, Papua New Guinea, 700 m. SPECIMENS EXAMINED. Papua New Guinea: d BM(NH) 78.889 (flat skin), 9 ,rf 78.890-891 (in alcohol) About 10 km S of Madang (coll. P. A. Morris). Hipposideros dinops pelingensis Shamel, 1940 Hipposideros pelingensis Shamel, 1 940 : 353. Peling ( = Peleng) I. E of Sulawesi. INDO-AUSTRALIAN BATS 1 5 1 SPECIMENS OBTAINED. S Sulawesi: d, 2 99 BM(NH) 82.116-118 Lalonggasu Meeto (Tomba Watu Cave), 18 km from Kendari Central (in alcohol, skulls of BM(NH) 82.1 17-1 18 extracted; coll. B. H. Gaskell, 'Operation Drake'). REMARKS. Shamel (1940) diagnosed pelingensis solely on the basis of its shorter tibia (length 38-2-41-0) when compared with H. dinops Andersen, 1905c from the Solomon Islands. Tate (1941<2 : 376) recorded pelingensis from Talassa, (Maros), south Sulawesi, listing (p. 391) the specimens as H. diadema pelingensis although (p. 376) comparing and associating pelingensis with dinops. Hill (1963) considered pelingensis a subspecies of//, dinops but had seen no examples of the Sulawesian form. These specimens agree favourably in size and structure with the nominate subspecies from the Solomon Islands but the rostrum is very slightly narrower, the upper canines a little smaller at the base and the toothrows a little shorter. However, in most of these features they closely resemble the smallest of the available specimens from the Solomons, an example (BM(NH) 67.21 18) from Malaita Island reported by Hill (197 la) who gave measurements of H. d. dinops and discussed variation in this subspecies. External measurements: length of forearm (3) 93-4-96-9; length of tibia (3) 39-9-40-9. Cranial measurements (d 1 BM(NH) 82.117, 9 82.118): greatest length of skull 36-5, 36-0; condylobasal length 32-4, 32-4; condylocanine length 31-5, 31-4; palatal length 13-2, 13-0; rostral width 10-2, 9-9; anteorbital width 10-0, 9-0; length of anterorbital foramen 2-8, 2-7; width of anteorbital foramen 0-7, 0-8; least interorbital width 3-6, 3-9; zygomatic width 20-8, 20-2; width of braincase 13-9, 13-7; mastoid width 16-7, 16-1; d-c 1 (alveoli) 9-3, 8-9; m 3 -m 3 12-7, 12-7; c-m 3 13-8, 13-4; length c 1 4-12, 3-97; width c 1 2-83, 2-85; length complete mandible from condyles 24-7, 24-5; length right ramus from condyle 25-5, 25-4; top of condyle-tip of angular process 5-8, 6-1; bottom of condyle-tip of coronoid process 7-2, 7-6; tip of angular process-tip of coronoid process 10-9, Il-7;c-m 3 15-6, 14-9. DISCUSSION. Specimens of//, d. pelingensis are of particular interest in connection with the very large //. inexpectatus Laurie & Hill, 1954, no material of pelingensis being available in London when this was described. Direct comparison shows pelingensis to be considerably smaller, with lower, narrower rostrum, much less developed sagittal and lambdoidal crests, less expanded zygomata and much smaller teeth. There is especially a great contrast in the large and heavy mandible of inexpectatus and the relatively lighter and by comparison apparently delicate mandible of pelingensis. Until recently H. inexpectatus was known in the literature solely from its holotype (BM(NH) 25.6.5.19) from Poso, north Sulawesi but Fieler (1981) has reported two further examples, from Gorontalo and Minahassa. They are specimens that have been for many years in the collections of the Staatliches Museum fur Tierkunde in Dresden, formerly identified as //. diadema (Geoffroy, 1813). Aselliscus tricuspidatus novehebridensis Sanborn & Nicholson, 1950 Aselliscus tricuspidatus novehebridensis Sanborn & Nicholson, 1950 : 461. Cave on Segond Channel, Espiritu Santo I, New Hebrides. SPECIMENS EXAMINED. New Hebrides: 15 -s: ~s5 *z TT rNt^o r^TfNoofNONi^ r~- ' C ^e f"" */^ ^^ C^ CO ^v f^ 1 ^^ *^ -i . ; T r ^r t G ^ oo i/**j fsj ro ^- - oo % -S 7 r *? ^ ro TT ro so /> I r- oo s< m ON O o 3 10 s \ 1 /"> sO *= .'| so v .^rn mONt-t-rr^^ -s: !5 e^ C oo oooooo oomoooooooooo oo ooa 3' 03 ^ o "3 ^ Z~~ ^^^v^^v. (N oo r- oo "- r*^ t**^ t r* C/5 ob w "1 T t r D m ON m - - o T C/5 _ _ _ p^j r~~ I ^ sD SO s/ \ \ \ \ \ \ - \ /^ >/^^ >/^fsoNOO oo r c . VCt so u "* ^ f*i rnr---r i --'^->owo o si 2 -5! >) en M ^ O ^^v /- v ^ -^ x -*s ^ -V ^-~. ^-v x-v s^ * a 5- 03 ON c ^j r ^ ^ ^j r -- ^- O r ^ N o v' C2- 2- >3- Q> r v c u f^t f^ p^- ^S O "O _c O ^O ^ f n r- i^ rn ^^ m - OO 03 _X* 1= rn ON^ r^ 00 TJ- s CN V O CN ^D rs ^ O^ */^ ON ^* ?? 7 7^ o Tt ^t oo c > ^^ c o c o c o c O oo r~ oo c OO v C c < -s: -^ C ^ Jo" sb i O ("*"} ro r^i o^ ^"O t**" ^if */^ */^ ^^ ^^ ^ 3 311 rn -v . * ' ' -v ^ sT ' -^ ^ ^ ^- S 1^*^ rf -^ SS: g --*. f * ' ^ * 1 T^ m ^^ so m ON 5 c ^o * o t ' ^ n <~n r~- rN c r- D :D ' -s: Q <3 2* ~ m < n r~ i ^ v 2 ' Pk *pM -Sf rt rn " q^" o wC " " t~-^ r^T cT t~-^ s$ ON' o wj 2 "^ sb i O n ( n < n r~- r~~ ^ v o m 5 v S, ^^ "^S i* m ^ '- o _4J in (~l 'c '5 u c c C ab 3 ao c -r : 1 i 1 M j= o .'ti ^ ^ ^ ^J w _y aj *- S - &0 c 15 y. | B C, ; 1 -"2 'i I " c' v> o ^ B^Jl^'i^S-o^ ^ ^^ > '~ 43 6 js'S 5- o JC o 'C : 8 E z C^ S *o C C ^ '^ S) 1 ^ g "o j" 'g c 1 ^ ^^2 H jc5 ^ ^ aj UU -J N ^ S 1> i J V INDO-AUSTRALIAN BATS 159 ro so /"> r~- N< 3 ^^ ^o ^/^ ^* fN NO t~- NO I O _ - fNl *^ 03 s> ^, c ^J- _ M TT O OO ^J < . v^^ i/-} ^^ NtJ 1 ^ _ ^^ ^) ifi xj r*^ ON I ^ OO ^^ */^ */^ C 5 O m 'o "^t 'ni CO a .- x: ON t/5 < H > , ^, 'o UM | c 5 o t-3 ^J Cy a ~s; 10 fN ro rN 10 NO w-> "- OO Tt -s: ^ ffl OO ^O ^O '^ - Tt obob^j- ^> vb M SO o 'o jU C/5 X) C ') U J PJ urem 00 g |>g- j (U g x< cd 4^o c/2 3 C/5 J IU ^ 2^ "Sb 15 "6 ,o c % c o <-^- m ca ca u. t*. - x> o _a> ^1.^5 1^ a'8 || - K3 >% 1- >> ' x: " 3 .P o SO x; u ^ "^"^-E '"" 15 r^ (-1 r x; P "Si ca ^ "O "O -^ Sd'S-S' ' "anf 5 'Sc o ^ Z C ^o f" f" CO '> oo"U i "o | P c * H j- u. C C9 Cj ^* O O flj ^* ^^ 1 r *^ 1 flj H di ^~ ' i JO U U J N > S "o E c!> J J 160 J. E. HILL Hill (1976#) thought peshwa Thomas, 1915a from India and dryas Andersen, \9Qlb closely related to M. horsfieldii of which both are here considered provisional subspecies. It is also possible that M. jeannei Taylor, 1934 from Zamboanga Island, Philippine Islands may represent M. horsfieldii: there is reasonable agreement in size and the wing membrane in jeannei is attached to the foot about halfway between the heel and the base of the toe, a characteristic feature of M. horsfieldii when compared with M. adversus or M. hasseltii in which it attached at the end of the tibia or at the ankle. Myotis hasseltii continents Shamel, 1 942 Myotis adversus continentis Shamel, 1942 : 323. Bangkok, Thailand. SPECIMEN EXAMINED. Burma: cf BM(NH) 78.154 British Embassy Residence compound, Rangoon (in alcohol, skull extracted; coll. D. W. & G. Walton). REMARKS. This specimen is the first of M hasseltii to be reported from Burma, although the species is known to occur farther north in Thailand, at Chiangmai (BM(NH) 9.10.1 1.6-9). It agrees closely with examples of M. h. continentis from Thailand but is slightly larger in some respects (Table 6), approaching M. h. hasseltii from the Malay Peninsula and Java, or the specimens from Sri Lanka discussed by Hill ( 1 91 6a). The second upper premolar (pm 3 ) of the Burmese specimen is very small, totally intruded from the row, with pm 2 and pm 4 in contact, while pm 3 is small, slightly intruded but nevertheless separates pm 2 and pm 4 . No examples of M. hasseltii were obtained by 'Operation Drake' in Sulawesi, whence its reported occurrence apparently rests on specimens recorded by Jentink (1887, 1888) from Gorontalo in the northern part of the island. Myotis adversus moluccarum (Thomas, 1915) Leuconoe moluccarum Thomas, \9\5d : 1 70. Ara, Kei Islands. Anamygdon solomonis Troughton, 1929 : 89. Rovianna I, New Georgia group, Solomon Is. SPECIMENS EXAMINED. C Sulawesi: 2 rfrf, 9 BM(NH) 82.131-133 R Ranu, 1 51' S, 121 30' E (in alcohol, skulls extracted; coll. B. H. Gaskell, 'Operation Drake'). REMARKS. Tate (\94\d) referred specimens of M. adversus from Sulawesi, Peleng Island and West Irian to M. a. moluccarum, with which these agree closely. They are rather smaller than M. a. adversus (Horsfield, 1 824) from Java: the Bornean and Sumatran subspecies M. a. carimatae Miller, \9Q6d is very similar to moluccarum but only limited material is available for comparison. Myotis adversus orientis subsp. nov. HOLOTYPE. New Hebrides: 9 BM(NH) 73.1412 Eastern cave at cliff base behind Aouta Plantation, Aore I. Collected 24 August 1971 by the Earl of Cranbrook, presented by the Royal Society Expedition to the New Hebrides, 1971. Original number 104-02. Skin and skull. OTHER MATERIAL. New Hebrides: 3 cfrf, 7 99 BM(NH) 73.1404-141 1, 73.1413 (in alcohol, skulls extracted), 73.1414 (skin, skull). All from type locality (obtained Earl of Cranbrook, Royal Society Expedition to the New Hebrides, 1971). DIAGNOSIS. A member of the subgenus Leuconoe, generally slightly larger (Table 7) in most respects than the geographically adjacent M. a. moluccarum from the Solomon Islands, New ^^ 'C ajr CM >? 1-2 75 S 1 (NONOm-^ -Or^ivDONO Tf r- Tf ^& < JH 03 r- 05 to (U 'S "O r^uouo-^-TtOoboOTi-vovb 'O ON m 10 t SO O r^ vo n T^- *p rn rp op O r^l ^ .11 V T ! *____ T J.__ OOOOT i. r ^ vO ^..^^^Q 1 1 II 1 II III sooo rnoNooO^or^^ fNr~-Tt oi <3 -~"~>''i' T ^'TJ-ONf^oo - ^t-^oio \o ^z OOOOmOOOO ^OO OJ S 1 2 si ^ m vo O O m ir> _!, _!. in JL ,t l^ftTTt ^O Q^ rfmoo ON (N a & , 3 ^o r^, TJ-vbino ^b m ^< oo m m r^ r<^ ro m ^ s ' rn m rsi m oo f^vbinTJ-^oO-^vOOOtN _L.J.n % (L> J= =- .JL ^ 3 35 5 ^___, T i._ OOOOT ;.^,^ ) ^-.^.^ '^^^Tj-oooN'^tONmrNir~~ ' o P Q -^ co o t "/~> T 3-mmQsr~-r~-r;|->sb>/~> so CO r~* S 13*0 t boONONONr^bNONONONON ^OOVON xamined in j < s: on V5 ^ g (N >O 7 fl ? ?t7i ) 771 5 r^OfNTt Tj"ONi/^inr~-fN SO ^O s 3 a a co ir^'nTJ-^-ori.oo^.^o^c, rsvb inso^o\ovorr ) \ovo'sOvoso invovo Table 7 Measurements ofMyotis adversus < ^ Length of forearm (1 Greatest length of skull Condylobasal length Condylocanine length Least interorbital width Zygomatic width Width of braincase Mastoid width c'-c 1 (alveoli) m 3 -m 3 c-m 3 Length of complete mandible from condyles i Length of right ramus i c-m, ( 162 J. E. HILL Guinea, the Moluccas and Sulawesi, approaching M. a. adversus from Java in size; differing from these, from M. a. carimatae from Borneo and from M. a. macropus (Gould, 1855) from Australia in a slightly greater degree of frontal elevation of the rostrum, the frontal profile less concave, in wider, more substantial interorbital region, and in greater inflation of the braincase, which is more globose, especially anteriorly, slightly higher, and more elevated posteriorly. DESCRIPTION. Externally very like M. a. adversus or M. a. moluccarum with relatively large, rounded ears reaching almost to tip of muzzle when laid forward; tragus in length a little less than half length of ear, slender, pointed, its anterior margin straight, its posterior margin gently convex; wing inserted at ankle. Pelage dense and woolly, dorsally blackish brown, the hairs dark at the base and for most of their length, tipped lightly with greyish or buffy white, the tipping forming no more than a slight overlay, the ventral surface greyish white, the hairs blackish at the base, heavily and densely tipped with grey white, posteriorly with little or no black at the base. Dorsal pelage darker and blacker than brownish examples of moluccarum from Sulawesi and the Molucca Islands but these are old specimens that have been preserved for many years: a similarly old specimen from Choiseul Island in the Solomon Islands approaches examples from the New Hebrides in dorsal colour. Skull strongly constructed, with broad, high rostrum, its upper surface flattened posteriorly with a broad, shallow median longitudinal depression; frontal profile only slightly depressed; interorbital region broad, relatively massive, least interorbital width 28-30% of condylobasal length, 54-57% of width of braincase; 26-28% of condylobasal length, 49-54% of width of braincase in adversus, carimatae and moluccarum; braincase inflated, globose, anteriorly relatively full, elevated posteriorly; palate short and wide, with strong post-palatal spine, supported by thin lateral laminae; second upper premolar (pm 3 ) small, its crown area about one third crown area of anterior tooth (pm 2 ), almost in row, slightly intruded, or intruded from row to lie in a recess between pm 2 and last upper premolar (pm 4 ); second lower premolar (pm 3 ) in crown area about one third crown area of anterior tooth (pm 2 ), almost in row but compressed between pm 2 and last lower premolar (pm 4 ), or sometimes slightly intruded. Measurements of M. a. orientis appear in Table 7. ETYMOLOGY. An eastern representative of M. adversus. DISCUSSION. Excluding very large species such as M. macrotarsus (Waterhouse, 1845) from the Philippine Islands and northern Borneo and M. stalkeri Thomas, 19100 from the Kei Islands leaves in Indo-Australia a group of large-footed bats of the subgenus Leuconoe with forearms ranging in length from 36-44 and with pm 2 and pm 3 variably reduced and variably situated in the toothrows. Their current classification has been established by views and opinions scattered through several publications. These include Tate (194 Id), Medway (1965, 1977), Findlay (1972), Hill &Thonglongya (1972) and Hill, (1972, 1974, 1976a).The arrangement adopted here has been developed from these and may be summarized: 1 . Wing inserted on metatarsus M - horsfieldii Wing inserted at end of tibia or at ankle 2. Fur short, velvety; post-palatal extension short, its median spicule lacking thin bony supporting laminae; pm 3 minute, intruded, pm 2 and pm 4 in contact or nearly so, pm 3 small, only partially in row or completely intruded M. hasseltii Fur, dense, woolly; post-palatal extension long, its median spicule supported by thin bony laminae; pm 3 , not greatly reduced, pm 3 not usually much intruded, pm, usually in row or only slightly intruded M. adversus Myotis horsfieldii deignani Shamel, 1 942 Thailand, Hainan I, Hong Kong (Findley, 1972). INDO-AUSTRALIAN BATS Myotis horsfieldii horsfieldii (Temminck, 1 840) (incl. lepidus Thomas, \9\5d) Myotis horsfieldii (!)peshwa Thomas, 19 15a (see Hill, 1976a) Myotis horsfieldii (?) dryas Andersen, 19076 (see Hill, 1976a) Myotis horsfieldii (l)jeannei Taylor, 1934 Myotis hasseltii continentis Shamel, 1942 Myotis hasseltii hasseltii Temminck, 1840 Myotis hasseltii macellus Temminck, 1840 Myotis hasseltii (?) abbotti Lyon, 1916 Myotis adversus adversus Horsfield, 1824 Myotis adversus carimatae Miller, 1 906 d Myotis adversus moluccarum Thomas, \9\5d Myotis adversus orientis Hill, 1983 Myotis adversus macropus Gould, 1855 163 Malaya, Borneo, Java, Bali, Sulawesi. India. South Andaman I. Zamboanga I, Philippine Is. Burma, Thailand, Kampuchea, (?) Sri Lanka. Malaya, Rhio Archipelago, Sumatra, Java. Borneo. North Pagi I, Mentawei Is, offW Sumatra. Java, (?) Malaya. Borneo, Karimata I. Sulawesi, Molucca Is, New Guinea, Solomon Is. New Hebrides. Australia. Pipistrellus javanicus (Gray, 1838) Scotophilus javanicus Gray, 1838 : 498 (renaming of Vespertilion javanais F. Cuvier, 1832). Java. SPECIMENS EXAMINED. C Sulawesi: 9 BM(NH) 82.134 R Ranu, 1 51' S, 121 30' E (in alcohol, skull extracted; coll. B. H. Gaskell, 'Operation Drake'). REMARKS. This specimen from Sulawesi agrees in most respects with examples of P. javanicus from Java and from the island of Madura. It differs from these only in the slightly greater concavity of its rostral profile, in having the frontal part of the braincase a little more inflated and in a slightly higher posterior upper canine cusp. Measurements: length of fore- arm 32-1; greatest length of skull 13-6; condylobasal length 12-7; condylocanine length 12-3; least interorbital width 3-5; zygomatic width 8-6; width of braincase 6-7; mastoid width 7-3; c'-c 1 (alveoli) 4-3; m 3 -m 3 6-2; c-m 3 4-9; length complete mandible from condyles 9-4; length right ramus from condyle 9-8; c-m 3 5-2. DISCUSSION. Several species of Pipistrellus have been reported or described from Sulawesi. These include P. javanicus (Gray, 1838) by Dobson (1878) (as abramus, a subspecies of P. javani