Jo, BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 29 1976 BRITISH MUSEUM (NATURAL HISTORY) LONDON: 1977 DATES OF PUBLICATION OF THE PARTS No. i ..... 22 January 1976 No. 2 . . . . .13 April 1976 No. 3 . . . . 13 April 1976 No. 4 . . . .23 April 1976 No. 5 . . . . . -27 May 1976 No. 6 . . . . -27 May 1976 ISSN 0007-1498 Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU CONTENTS ZOOLOGY VOLUME 29 PAGE No. i. A review of the family Centropomidae. By P. H. GREENWOOD . i No. 2. Fossil reptiles from Aldabra Atoll, Indian Ocean. By E. N. ARNOLD ........... 83 No. 3. A review of the family Canidae, with a classification by numerical methods. By J. CLUTTON-BROCK, G. B. CORBETT and M. HILLS . 117 No. 4. The cranial musculature and taxonomy of characoid fishes of the tribes Cynodontini and Characini. By G. J. HOWES . . . 201 No. 5. Designation of lectotypes of some ostrocods from the Challenger Expedition. By H. S. PURI and N. C. HULINGS (Pis 1-27) . . 249 No. 6. Some tertiary and recent Conescharelliniform Bryozoa. By P. L. COOK and R. LAGAAIJ (Pis 1-8) 317 A REVIEW OF THE FAMILY CENTROPOMIDAE (PISCES, PERCIFORMES) P. H. GREENWOOD BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 29 No. i LONDON: 1976 A REVIEW OF THE FAMILY CENTROPOMIDAE (PISCES, PERCIFORMES) BY PETER HUMPHRY GREENWOOD Pp. 1-81 ; 37 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 29 No. i LONDON: 1976 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Scientific Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 29, No. i of the Zoology series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation : Bull. Br. Mus. not. Hist. (Zool.) ISSN 0007-1498 Trustees of the British Museum (Natural History), 1976 BRITISH MUSEUM (NATURAL HISTORY) Issued 22 January 1976 Price £5.85 By P. H. GREENWOOD CONTENTS Page INTRODUCTION ........... 4 MATERIALS ........... 5 ABBREVIATIONS USED IN TEXT-FIGURES ...... 8 THE FAMILY CENTROPOMIDAE ........ 10 AN ANATOMICAL AND TAXONOMIC REVIEW OF THE Ldtes AND LutioldteS SPECIES ........... 12 The neurocranium . . . . . . . . . 14 Hyopalatine arch and the preoperculum . . . . . 27 Circumorbital bones ......... 31 Opercular bones .......... 32 Jaws 34 Branchial skeleton ......... 35 Hyoid arch .......... 38 Pectoral girdle and associated bones ...... 39 Vertebral column ......... 42 Caudal fin skeleton ......... 44 Dorsal and anal fins ......... 45 Swimbladder .......... 47 Baudelot's ligament ......... 47 Lateral line .......... 48 THE INTERRELATIONSHIPS OF SPECIES WITHIN THE GENUS Ldtes, AND THE TAXONOMIC STATUS OF Luciolates Blgr. ...... 49 A REVIEW OF THE GENUS Psammoperca Richardson . . . . 51 Introduction .......... 51 Osteology and anatomy of Psammoperca waigiensis . . . 52 Neurocranium .......... 52 Hyopalatine arch and the preoperculum . . . . . 53 Circumorbital bones ......... 54 Opercular series .......... 55 Jaws ....... 55 Branchial skeleton ........ 55 Hyoid arch skeleton ........ 56 Pectoral girdle and associated bones . . . . . . 56 Vertebral column ......... 58 Caudal fin skeleton ......... 58 Dorsal, anal and pelvic fins ...... 59 Swimbladder ....... 60 Baudelot's ligament ......... 60 Lateral line ........ 60 THE RELATIONSHIPS OF Psammoperca .... 60 THE RELATIONSHIPS OF Centropomus WITH THE LATINAE . . 62 Neurocranium ........ 62 Hyopalatine arch ...... 63 Preoperculum ....... 64 4 P. H. GREENWOOD Operculum .... . . . . . . 64 Circumorbital bones ......... 64 Jaws ........... 64 Gill arches ........... 64 Vertebrae ........... 65 Predorsal bones . . . . . . . . . - . 65 Dorsal fins . . . . . " . ' . . 65 Anal pterygiophores ......... 66 Caudal fin skeleton . . . . . . . . 66 Pectoral girdle and associated bones ...... 66 Swimbladder .......... 66 Baudelot's ligament ......... 67 Lateral line .......... 67 FOSSIL CENTROPOMIDAE ......... 68 Lates species .......... 69 Eolates species .......... 70 BlOGEOGRAPHY .......... 71 DIAGNOSES FOR THE CENTROPOMIDAE, ITS SUBFAMILIES, GENERA AND SUBGENERA ........... 75 Centropomidae .......... 75 Centropominae .......... 76 Latinae ........... 76 ACKNOWLEDGEMENTS ......... 78 REFERENCES ........... 78 INTRODUCTION THE Centropomidae, a family of tropical estuarine, marine and freshwater percoid fishes, is represented in the New World, Africa and Asia by a total of at least 18 species (the number varying mainly with the taxonomic limits set for the family by different authors). Of the 18 species recognized, 9 occur in the New World, 7 - all freshwater species - occur in Africa and 2 in Asia. Temporally the family has a good fossil record extending from the Eocene to prehistoric times. Geographically, however, the fossil record is restricted to Africa and Europe, and only one taxonomic division of the family, the closely related genera Lates and Eolates, is represented ; see Sorbini (1973) and Greenwood (1974). It was, in fact, a new fossil centropomid from the neogene of Africa (Greenwood & Howes, 1975) that led to this revision. Our attempts to identify the new fossil soon made it clear that the African centropomids are more varied anatomically than had been realized previously. Also, it became obvious that the current taxonomic arrangement of the family does not reflect the probable phyletic relationships of its constituent taxa. Indeed, except for Eraser's (1968) analysis of the New World Centropomus species, no fully reasoned attempt has been made to interpret intra- familial relationships along phyletic lines. Also, the presumed relationships of the Centropomidae with the Serranidae (Regan, 1913 ; Berg, 1947 ; Gosline, 1966 ; Greenwood, Rosen, Weitzman & Myers, 1966) appear to be based more on intuitive than on critical taxonomic reasoning, and need reappraisal. Although it was for these reasons mainly that the present review was undertaken, I also hope that it may provide a step towards the clearing of that taxonomic rag-bag, the 'lower percoid fishes' (see Greenwood et al., 1966 ; Gosline, 1966). GLAUCOSOMIDAE Osteological material : Glaucosoma burgeri CENTROPOMIDAE Osteological material : Lates calcarifer Lates niloticus Lates niloticus Lates niloticus Lates niloticus Lates niloticus Lates niloticus Lates niloticus Lates macrophthalmus Lates longispinis Lates microlepis Lates angustifrons Lates mariae Luciolates stappersi Luciolates stappersi Luciolates stappersi Luciolates stappersi REVIEW OF CENTROPOMIDAE MATERIALS BMNH 1884.2.26:60 China BMNH 1873.1.21:2 Fitzroy R. Unregistered No locality BMNH 1971.2.8:186 No locality Unregistered 'Red Sea' BMNH 1864.6.29:5 'West Africa' Unregistered Lake Rudolf Unregistered No locality Unregistered Lake No Unregistered Lake Albert Unregistered Lake Rudolf BMNH 1900.12.13:37 Albertville BMNH 1955.12.20:1722 Lake Tanganyika BMNH 1955.12.20:1667 BMNH 1955.12.20:1672 BMNH 1936.6.15: 1705-6 BMNH 1971.6.23:76-8 BMNH 1975.4.23:2 Lake Tanganyika Lake Tanganyika 293 mm S.L. 360 mm S.L. (skeleton) (alizarin preparation) (disarticulated skeleton) (disarticulated skeleton) (skull and pectoral girdle) (alizarin preparation) (alizarin preparations) (alizarin preparation) (skeleton, 102 mm S.L.) (skull) (disarticulated skeleton) (skull and pectoral girdle) (skull and pectoral girdle) (disarticulated skeleton) Psammoperca waigiensis BMNH 1892.9.2:10-11 Borneo Psammoperca waigiensis Centropomus undecimalis Centropomus ensiferus Centropomus pectinatus Dissected specimens : Lates niloticus Lates niloticus Lates niloticus Lates macrophthalmus Lates macrophthalmus Lates longispinis Lates angustifrons Lates mariae Luciolates stappersi BMNH 1872.10.18:90 BMNH 1883.12.16:1-2 BMNH 1861.12.12:13 BMNH 1894.12.1:5 BMNH 1907.12.2: 2915-6 BMNH 1907.12.2: 2952-3 BMNH 1931.11.20:1-2 BMNH 1929.1.24:341-4 BMNH 1975.1.18:1 BMNH 1932.6.13: 102-106 BMNH 1906.9.8:87-88 BMNH 1955.12.20: 1628-29 BMNH 1955.12.20: 1669-71 Lake Tanganyika (skull) Lake Tanganyika (alizarin preparation) Lake Tanganyika (disarticulated skeleton from a fish 390 mm S.L.) (skull and pectoral girdle) (circumorbital series) (skull) (skeleton, disarticulated) (skull and pectoral girdle) Cebu Jamaica No locality Jamaica Nile Nile Merowe (Paratype) Lake Albert (Syntype) Lake Tanganyika 227 mm S.L. 165 mm S.L. 218 mm S.L. 275 mm S.L. 315 mm S.L. 250 mm S.L. 300 mm S.L. Lake Tanganyika 220 mm S.L. Lake Tanganyika 215 mm S.L. P. H. GREENWOOD Psammoperca waigiensis Psammoperca waigiensis Centropomus undecimalis Centropomus undecimalis Centropomus pectinatus Specimens .examined : Lates calcarifer Lates calcarifer Lates niloticus Lates macrophthalmus Lates longispinis Lates angustifrons Lates mariae Lates microlepis Luciolates stappersi Luciolates stappersi Psammoperca waigiensis Psammoperca waigiensis Psammoperca waigiensis Psammoperca waigiensis Psammoperca waigiensis Centropomus undecimalis Centropomus undecimalis Centropomus pectinatus Centropomus ensiferus Specimens radiographed : Lates calcarifer Lates niloticus Lates macrophthalmus Lates macrophthalmus Lates macrophthalmus Lates longispinis Lates angustifrons Lates angustifrons Lates mariae Lates mariae Lates microlepis Lates microlepis Luciolates stappersi BMNH 1892.9.2:10-11 Borneo 150 mm S.L. BMNH 1872.10.13:90 Cebu 240 mm S.L. BMNH 1883.12.16:1-2 Jamaica 280 mm S.L. BMNH 1923.7.30:114 Rio de Janeiro 175 mm S.L. BMNH 1895-5-27:3-5 Mazatlan 245 mm S.L. BMNH 1863.2.23:29 Amoy 190 mm S L BMNH 1936.8.6:43 Queensland 260 mm S.L. Major part of the entire collection BMNH 1929.1.24: 340-344 (Paratypes) 145-283 mm S.L. BMNH 1932.6.13: 102-106 (Syntypes) 115-270 mm S.L. Entire collection Entire collection Entire collection Entire collection RGMARC 129887-889 (Tervuren Museum specimen) 407 mm S.L. BMNH 1933.3.11:312 Culion, no mm S.L. Philippines BMNH 1883.11.28:14 Singapore 165 mm S.L. BMNH 1891.10.29:66 Ceylon 132 mm S.L. BMNH 1939.1.17:11 Hong Kong 215 mm S.L. BMNH 1888.11.6:5 Madras 175 mm S.L. BMNH 1936.1.31:8 Trinidad 290 mm S.L. BMNH 1906.6.23:82 Trinidad 180 mm S.L. BMNH 1920.12.22: Trinidad 230 & 240 mm S.L. 57-58 BMNH 1903.5.15:3-5 Panama 183-230 mm S. L. BMNH 1891.11.30:1-8 BMNH 1900.12.2: 2329-38 BMNH 1929.1.24: 340-344 BMNH 1929.4.16:39-41 BMNH 1929.1.24: 341-344 BMNH 1932.6.13:102-6 BMNH 1936.6.15: 1687-97 BMNH 1906.9.8:87-88 BMNH 1955.12.20:1636; 1628-9; 1655-6 ; 1672-86 BMNH 1906.9.6:7 BMNH 1906.9.8:89 BMNH 1955.12.20: 1753-85 BMNH 1955.12.20: 1669-71 ; 1680 REVIEW OF CENTROPOMIDAE Luciolates stappersi Luciolates stappersi Psammoperca waigiensis Psammoperca waigiensis Psammoperca waigiensis Psammoperca waigiensis Psammoperca waigiensis Centropomus unionensis Centropomus robalito Centropomus nigrescens Centropomus ensiferus Centropomus undecimalis Centropomus undecimalis Centropomus undecimalis Fossil material : Eolates gracilis Eolates gracilis Eolates gracilis Eolates gracilis Eolates gracilis Eolates gracilis Eolates gracilis SERRANIDAE Osteological material : Epinephelus tauvina Epinephelus areolatus Epinephelus afer Epinephelus itajara Dissected specimens : Epinephalus alexandrinus BMNH 1964.7.14:1 Malta Specimens examined : Serranus radialis BMNH 1923.7.30:77-79 Rio de Janeiro BMNH 1936.6.15: 1705-6 BMNH 1971.6.23:76-78 BMNH 1872.10.18:90 BMNH 1939.1.17:11 BMNH 1892.9.2:10-11 BMNH 1870.12.27:17 BMNH 1888.11.6:5 BMNH 1903.5.15:8 BMNH 1895.5.27:6 BMNH 1883.7.28:14 BMNH 1903-5-15:3-5 BMNH 1895.5.27:2 BMNH 1936.1.31:8 BMNH 1924.2.29:16 BMNH P23803 Monte Bolca 130 mm S.L. BMNH Pi6i37 Monte Bolca 121 mm S.L. BMNH P39i8 Monte Bolca 139 mm S.L. BMNH P23798 Monte Bolca c. 135 mm S.L. BMNH Pi6374 Monte Bolca 41 mm S.L. BMNH Pi 6756 Monte Bolca 29 mm S.L. BMNH P37225 Monte Bolca 124 mm S.L. Unregistered Java (skeleton, 225 mm S.L.) BMNH 1888.12.29:24 Muscat (skeleton, 280 mm S.L.) Unregistered St Croix (skeleton, 140 mm S.L.) BMNH 1883.12.16:9 Jamaica (skeleton, 355 mm S.L.) 243 mm S.L. 115-140 mm S.L. AMBASSIDAE Osteological material : Ambassis urotaenia Ambassis commersonii Ambassis wolffii BMNH 1928.1.17:8-15 BMNH 1855.9.19:359 BMNH 1898.4.2:67 Specimens examined: The major part of the collections of Ambassis and Chanda (alizarin preparation) (disarticulated skeleton) (skeleton, 85 mm S.L.) GERRETDAE Osteological material : Gerres oyena Gerres oyena BMNH 1965.4.4:125-38 BMNH 1960.3.15:670-5 (alizarin preparations) {alizarin preparations) P. H. GREENWOOD ABBREVIATIONS USED IN THE TEXT-FIGURES AA Anguloarticular ART P Articular process of the pre- maxilla ASC P Ascending process of the pre- maxilla AHYF Anterior facet for hyomandi- bula Bb 1-3 Basibranchial 'BcF' 'Berycoid foramen' BOC Basioccipital BrR Branchiostegal ray BSP Basisphenoid Cb 1-5 Ceratobranchial of first to fifth arch Ch Ceratohyal CL Cleithrum COR Coracoid D Dentary Dhh Upper hypohyal D PROC Dorsal process of the maxilla E Mesethmoid Ei-E4 Epibranchials of the first to fourth arches EaTP, Tooth-plates associated with the E3TP second and third epibranchials ECT Ectopterygoid Eh Epihyal ENT Entopterygoid EP Epural EPI Epioccipital (= epiotic auct.} EXO Exoccipital FR Frontal FRC Frontal crest FRR Frontal ridge GF Gill filament Gh Glossohyal Gr Gill raker H1( H5 First and fifth hypurals Hbi Hypobranchial of first gill arch HsPU2 Haemal spine of second preural vertebra Hyomandibula Intercalar Interoperculum Lachrymal (first circumorbital bone) LATE Lateral ethmoid LAT SP Latero-sensory canal openings LC Lateral commissure LIG Ligament MET Metapterygoid NaPU2 Neural arch and spine of second preural vertebra OCS Occipito-spinal nerve foramen OP Operculum PAL Palatine PAR Parietal PARC Parietal crest Pbi-Pb4 Pharyngobranchials of the first to fourth gill arches Pb2 TP- Pb4 TP Tooth plates associated with pharyngobranchials of the sec- ond to fourth gill arches PCj, PC2 Upper and lower postcleithra PFr Pectoral fin ray PH Parhypural PHYF Posterior facet for hyomandi- bula PMAXP Posterior maxillary process PMXP Premaxillary process of the maxilla POP Preoperculum PRO Prootic PS Parasphenoid PTF Posttemporal fossa PTO Pterotic PTS Pterosphenoid PTSP Pterosphenoid pedicle PTSS Pterosphenoid spur PU1 + U1 Fused first ural and preural centra Q Quadrate R Radial for pectoral ray RA Retroarticular SC Supracleithrum Sc Scapula SLP Supralamellar tooth plate SOC Supraoccipital SOC S Supraoccipital shelf SOP Suboperculum SOS Subocular shelf SPO Autosphenotic SY Symplectic TP Tooth plate UR Uroneural URi, URa Upper and lower uroneurals V Vomer VHh Lower hypohyal II -V Second to fifth circumorbital bones IX Foramen for glossopharyngeal nerve X Foramen for vagus nerve ist APTY First anal pterygiophore and Asp Second anal fin spine REVIEW OF CENTROPOMIDAE FIG. i. Outlines of : (a) Psammoperca waigiensis, (b) Lates calcarifer, (c) L. niloticus, (d) L. longispinis, (e) L. macrophthalmus , (f) L. angustifrons , (g\ L. mariae, (h) L. wicro- /e^>is, (i) L. stappersi, (j) Centropomus undecimalis. io P. H. GREENWOOD THE FAMILY CENTROPOMIDAE Although in 1955 Matsubara classified several of the genera considered below in the family Serranidae, there is still a consensus of opinion among ichthyologists that these fishes constitute a natural taxonomic group, albeit one closely related to the Serranidae. (See Katayama (1956) for a short taxonomic history of the group.) There has, however, been rather less agreement on the definition and delimitation of the family Centropomidae in which these various genera are classified, or with which they are thought to be most closely related. In particular there is uncertainty about the affinities of Glaucosoma Temm. & Schl., of Chanda Ham. Buch. (= Ambassis of authors), and of genera related to Chanda. Regan (1913), for example, included both Glaucosoma and Chanda (as Ambassis) in the Centropomidae, as did Norman (1966) who, however, gave Chanda and related genera subfamilial rank (Chandinae) and placed Glaucosoma with Lates Cuv., and Psammoperca Richard- son in the subfamily Latinae. Other views were expressed by Jordan (1923) who gave familial rank both to Chanda and its related taxa (Ambassidae), and to Glauco- soma (Glaucosomidae). Berg's (1947) classification returned Chanda to the Centro- pomidae, but kept Glaucosoma as a monotypic family. Greenwood et al. (1966) followed Berg, as did Lindberg (1971). There have, of course, been several definitions of the Centropomidae, both sensu latu and stricto (see especially Gill, 1883, and Meek & Hildebrand, 1925, for the family as restricted to species of Centropomus ; Regan, 1913, and Norman, 1966, for the family sensu latu ; Munroe, 1961, for the Chandidae and Centropomidae, and Katayama, 1954, for the only comprehensive definition of the Glaucosomidae). Yet, from none of these definitions is it possible to determine the synapomorph features that could establish the phyletic relationships of the taxa involved, either as a holophyletic assemblage or as two or even three lineages. With the aim of establishing such relationships I have examined all the characters listed in these various definitions ; as is inevitable in such revisionary work I have discovered other characters which were not taken into account by earlier authors. Most of the characters used by Regan (1913), Norman (1966) and Katayama (1954) are either primitive features widely distributed amongst the lower percomorphs and percoids (i.e. symplesiomorphies), or, if derived ones, are characteristics also shared with several percoid families. In the symplesiomorphic category are the vertebral number, presence of frontoparietal crests, and the dentition and other jaw characters. The derived characters include the presence of an axillary pelvic scale, and the extension of lateral line pore scales onto the caudal fin. This latter character is of interest because, although the lateral line extends some way onto the caudal in several percoid taxa, rarely does it reach or almost reach to the margin of that fin, as it does in Centropomus, Lates and Psammoperca. (Only in the Sciaenidae does the lateral line extend as far posteriorly as in these genera.) This distinction in the degree to which the lateral line extends posteriorly has not been drawn by other workers. One character not used by previous authors (but mentioned with reference to Centropomus and Lates by Gosline, 1966) is the presence of an anteroposteriorly expanded neural spine on the second vertebra. Indeed, this feature, combined with REVIEW OF CENTROPOMIDAE n the extension of the lateral line far onto the caudal fin, provide the only synapo- morph characters at all widely distributed amongst taxa currently classified with the Centropomidae. Because the caudal lateral line character also occurs in the Sciae- nidae, the neural spine character is the sole truly synapomorph feature of the centropomids. Currently recognized genera with such an expanded neural spine are Lates, Luciolates, Psammoperca and Centropomus (Fig. i). Except for Luciolates, these genera also have the caudal lateral line scale row extending or almost extending to the fin margin. (The lateral line in Luciolates is discussed on p. 48.) Neither Glaucosoma nor Chanda (and its related genera) has either of these features. The lateral line extends only onto the basal third, or less, of the caudal fin, and the second neural spine is no wider than that of the first vertebra (in other words, the usual percomorph condition) . Thus, on the basis of these characters, and the lack of any other unifying features, it would seem that Glaucosoma and Chanda cannot belong to the same lineage as Centropomus, Lates, Luciolates and Psammoperca. These latter taxa alone are therefore retained in the family Centropomidae. Questions now arise as to the relationships and status of Glaucosoma and the Chanda-\ike genera, of their relationship to the Centropomidae as here defined, and of the interrelationships of the Centropomidae within the Percoidei. Nothing I have yet discovered suggests that Glaucosoma is a close relative of Chanda (and its immediate relatives). Both taxa are readily defined by various autapomorphies, but I cannot find any synapomorph characters uniting them. Unfortunately, the sort of detailed information needed for phyletic studies amongst percoid fishes is not yet available for many taxa, and I cannot suggest where the relationships of Chanda and Glaucosoma may lie. For the moment the only course available is to recognize two families, the Glaucosomidae and the Chandidae, and to consider both as of uncertain affinity amongst the Percoidei. The dorsal gill arch skeleton in the Chandidae I have examined (several species of Chanda) is certainly more derived than are those of the Centropomidae and Serranidae (see Rosen, 1973, for a discussion of the gill arches in percoid fishes). In the morphology of the pharyngobranchials, especially the second, Chanda is very similar to Eucinostomus argenteus (Gerridae) as figured by Rosen (op. cit., text-figs 98 & 99). Glaucosoma also shows more derived characters in its gill arch skeleton than does any member of the Centropomidae. I suspect that it will be from the gill arch skeleton that the relationships of these two families will ultimately be determined. Similar problems and lack of data limit the formulation of hypotheses regarding the phylogeny of the Centropomidae. It is generally thought, or implied, that the centropomids are closely related to the Serrandiae (see Regan, 1913 ; Katayama, 1954 ; Gosline, 1966 ; Greenwood et al., 1966 ; Norman, 1966). Again it has so far proved impossible to demonstrate within these families any but symplesiomorph or autapomorph features, none of which provides acceptable information for confirming or refuting this relationship. Thus, for the time being the Centropomidae too must remain as a family incertae sedis amongst the lower percoids. However, with the limits of the Centropomidae defined (see above) it is now possible to turn to problems of infrafamilial relationship and taxonomy. 12 P. H. GREENWOOD AN ANATOMICAL AND TAXONOMIC REVIEW OF THE LATES AND LUCIOLATES SPECIES The present taxonomic status of several Lates species must be reviewed before considering their anatomy and phyletic relationships. The probably monotypic genus Luciolates Blgr. is also included in this review, although a discussion of its ultimate status is deferred until p. 49. With one exception, namely Lates calcarifer (Bloch), all extant Lates species are confined to Africa but fossil remains of this genus are known from southern Europe as well as from several areas in Africa (Sorbini, 1973 ; Greenwood, 1974). The extinct taxon Eolates gracilis (Agassiz) from Monte Bolca will be considered later (p. 70), together with the extinct 'species' of Lates. Lates calcarifer, a coastal and estuarine species, is widely distributed in the Indo-Pacific region (India, Bangladesh, Burma, Malay Peninsula, Java, Sumatra, Borneo, Celebes, Sarawak, Philippines, Papua-New Guinea, northern and western Australia, southern China, and Japan). According to Weber & de Beaufort (1929), this species also occurs in the Persian Gulf ; their reference to L. calcarifer entering the mouths of the Nile, Niger and Senegal, and ascending these rivers, is clearly an error stemming from a confusion of this species with L. niloticus. Although essentially a marine fish, L. calcarifer freely enters and remains in rivers but always returns to estuarine or marine environments for spawning (Dunstan, 1959 ; Lake, 1971). Lates niloticus (L.) is widely distributed in the rivers and lakes of tropical Africa (Nile, Niger, Senegal, Volta and Zaire [= Congo] rivers ; Lakes Chad, Albert, Rudolf and some of the Ethiopian lakes). Not surprisingly in such a widespread taxon there are indications of some geographically limited morphotypes. As yet there has been insufficient study of these populations to determine the significance of their morphological differences, and none of the fluviatile populations has been given the formal status of a subspecies (see Daget (1954) on Pellegrin's (1922) L. niloticus var. macrolepidotus from Zaire). Worthington (1932), however, has described two subspecies, L. niloticus rudolfianus and L. n. longispinis from Lake Rudolf. Lates niloticus rudolfianus, a form attaining a large size (up to 148 cm total length) and apparently confined to inshore regions of Lake Rudolf (Worthington, 1932), is acknowledged by Worthington to be morphologically intermediate between L. niloticus of the Nile and populations of that species inhabiting Lake Albert (named L. albertianus by Worthington [1929], but shown by Holden [1967] to be indis- tinguishable from L. niloticus). I have re-examined the type material of L. n. rudolfianus and can find no reason for maintaining the subspecific status of this population. In all morphometric, meristic and gross morphological characters the type specimens lie within the range of variability determined for L. niloticus over its entire range. Thus, at least until larger samples are available from numerous localities in Lake Rudolf, I would consider L. niloticus rudolfianus to be a synonym of the nominate species. The second subspecies from Lake Rudolf, L. n. longispinis, presents a somewhat different problem. Apparently it is separated ecologically from the other Lates REVIEW OF CENTROPOMIDAE 13 species in the lake, being a fish of the deeper waters (Worthington, 1929 ; Hopson, unpublished report). Furthermore, it is morphologically differentiate from L. niloticus, and does not attain such a large adult size. The principal morphometric differences distinguishing the taxon 'longispinis' from L. niloticus are its larger eye (diameter 22-6-39-9 Per cen^ of head in fishes 118-273 mm standard length, cf. 18-3-22-9 per cent in L. niloticus of a comparable size ; in both taxa eye size is negatively correlated with standard length) and longer third spine in the dorsal fin (78-0-84-0 per cent of head, cf. 55-0-70-0 per cent) . The larger eye in 'longispinis' is most clearly manifest when small specimens of both species are compared ; for example the eye is 21-8 per cent of the head in a 107 mm S.L. L. niloticus but is 32-9 per cent in a 118 mm specimen of 'longispinis' '. Another difference, but one correlated with relative eye size, lies in the less marked posterior extension of the maxilla in 'longispinis'. In specimens of L. niloticus more than 125 mm S.L. the posterior tip of the maxilla lies at a point clearly behind a vertical through the posterior orbital margin ; in 'longispinis' above 125 mm long the maxillary tip lies in or a little anterior to that vertical. In fishes less than 120 mm S.L., the distinction is much less obvious (or even non-existent) because of the relatively larger eye in L. niloticus of that size. Since, in Lake Rudolf, 'longispinis' and L. niloticus are sympatric (albeit allotopic), and because the two taxa show various and consistent morphological differences, I can find no grounds for considering 'longispinis' to be a subspecies of L. niloticus. The obvious expedient of raising Worthington's (1932) L. n. longispinis to full specific rank, however, requires further consideration when the taxon is compared with L. macrophthalmus Worthington, 1929 (see above ; also Holden, 1967). Lates macrophthalmus is the endemic ecological counterpart in Lake Albert of 'longispinis' in Lake Rudolf (see Holden, 1967), and closely resembles that species as well, sharing with it the presumably derived features of enlarged eyes and elongate third spine in the dorsal fin. The only differential feature I can find is the relatively longer spine of 'longispinis' (78-0-85-0, mean 82-0 per cent of head, cf. 65-0-84-0, m = 74'4 per cent, in L. macrophthalmus). There also appear to be slight differences in the relative proportions of certain head parts, e.g. the vertical limb of the preoperculum lies slightly further forward in 'longispinis'. Detailed comparisons are hampered by the paucity of study material, there being only the five syntypes of L. n. longispinis* and the eleven syntypes of L. macrophthalmus available. Basically, the problem raised by 'longispinis' in Lake Rudolf and L. macroph- thalmus in Lake Albert is whether each should be considered a distinct and endemic species evolved locally from a population of L. niloticus (the generally accepted hypothesis, see Worthington, 1932, and Holden, 1967) or whether they should be looked upon as sister taxa derived from a common ancestor distinct from L. niloticus. This hypothetical species presumably invaded the developing Lakes Rudolf and Albert alongside L. niloticus. If this latter relationship could be determined it would, on the morphological evidence available, be more realistic to * The sixth syntype of L. n. longispinis mentioned by Worthington (1932) cannot be located, and neither is it recorded in the Museum's register. This suggests that the word 'six' in the original description is a lapsus for 'five'. I4 P. H. GREENWOOD treat ' longispinis' as a subspecies of L. macrophthalmus rather than as a distinct species. Unfortunately I do not have enough material at my disposal to test the two hypotheses, even assuming that anatomical criteria alone would be suitable for this purpose. For the moment then, and without prejudice to an ultimate solution of the taxon's true phyletic position, I propose treating Worthington's subspecies as a full species, namely L. longispinis Worthington (1932). The three other Lates species, L. angustifrons Blgr., L. microlepis Blgr. and L. mariae Steindachner (see Poll, 1953), require no further comment at this stage. All are morphologically distinct from the other species and from one another. A fourth Lates-like taxon from Lake Tanganyika is currently placed in the genus Luciolates* Blgr., principally because of the wide separation of the two dorsal fins (Boulenger, 1914 ; Poll, 1953, 1957). Luciolates is closely related to Lates, in particular to L. mariae. As I hope to demonstrate in the next section of this paper I believe that Luciolates should be included in Lates if the principles of phyletic classification are not to be violated. The anatomy of Lates and Luciolates The anatomy, and especially the osteology, of Lates and Luciolates has never been subject to a general review encompassing all known species. Gregory (1933) has given a rather superficial account of the syncranial osteology in Lates niloticus,^ and Katayama (1956) a more detailed description of Lates calcarifer which included some details of its soft anatomy. The account which follows is based on the examination of at least two skeletons of each species, and in the case of L. niloticus on several specimens over a wide size range. Radiographs of several specimens of every species were also examined. In all intrageneric comparisons made below the conditions found in L. calcarifer and L. niloticus are, with few exceptions, taken to be those primitive for the genus. This conclusion regarding the status of the two species was reached after all the species had been examined and a comparison made with members of other percoid groups apparently related to the Centropomidae (Gosline, 1966 ; Greenwood et al., 1966). Within the Centropomidae as a whole, L. calcarifer and L. niloticus-type cranial osteology should also be taken to represent the primitive condition. The neurocranium The overall morphology of the neurocranium in Lates and Luciolates can be judged from Figs 2-8. Basically, the neurocranium in Lates differs little from that of most serranids (sensu Greenwood et al., 1966). It has, however, well-developed and continuous frontoparietal crests with a sensory canal pore located at or near the junction of the crests, and the exoccipital facets are contiguous (separated in most serranids, * A second species, Luciolates brevior, has been described (Boulenger, 1914), but is known only from the holotype and has never been recorded again. In all probability L. brevior should be treated as a synonym of Luciolates stappersi Blgr., 1914, and is treated as such in this paper. f The neurocranium supposedly of Luciolates stappersi, figured by Gregory (1933), is wrongly identified; as far as I can judge it is from a specimen of Lates angustifrons. REVIEW OF CENTROPOMIDAE soc PHYF SPO PRO PAR PTF FR OCS 10mm SPO PTO FIG. 2. Lates niloticus, neurocranium. (a) Left lateral view, (b) Dorsal view. (From Greenwood & Howes, 1975.) personal observations ; see also Gosline, 1966). Since continuous frontoparietal crests (usually incorporating a sensory pore) occur in berycoid fishes (see Patterson, 1964), this condition must be considered a primitive one. Likewise, the medially contiguous exoccipital facets are also a primitive feature found in berycoids. The extensive interfrontal penetration of the supraoccipital, however, must be ranked as a derived feature. The dorsicranium shows some slight interspecific differences in detail but not in basic layout. The supraoccipital extends forward to the level of the median sensory pore of the supraorbital lateral line cross-commissure, and clearly separates the frontals posteriorly. The bone's relative anterior extension appears to be least marked in L. angustifrons, L. mariae and L. microlepis ; this is attributable to the i6 P. H. GREENWOOD EPI SOC FR LATE PRO PS BOC 10mm FIG. 3. Lates calcarifer, neurocranium, left lateral view. anteriorly more elongate frontals, a lengthening associated with the elongation of the ethmoid region in these species. In Luciolates stappersi, despite the attenuation of its snout, the supraoccipital extends forward to a point level with the anterior orbital margin ; the median sensory pore has a corresponding anterior displacement (Fig. 8). These four Lake Tanganyika species also have deeper grooves lying between the median supraoccipital crest and the fronto-parietal ridges on each side of the skull, a consequence, perhaps, of their narrower skulls (see below). All Lates species have a well-demarcated ledge on either side of the supraoccipital crest, the ledge being confluent anteriorly with the supraocciptal bone itself, and extending backwards almost to the posterior margin of the crest. The ledge is narrower and less conspicuous in Luciolates, and is confined to the anterior part of the crest. The posttemporal fossa is deep in all species except Luciolates stappersi, and in none do its constituent bones meet at the centre of the fossa ; even in the largest specimen examined the fossa is still open, its aperture closed off from the cranial cavity by a tough membrane. Amongst members of the Serranidae the Lates - Luciolates condition is characteristic of small and apparently juvenile fishes ; in larger individuals (many of which are, nevertheless, considerably smaller than adult Lates} the fossa has a completely bony floor. This interfamilial difference would suggest that the Lates condition is the primitive one. The wide cephalic lateral line canals of the dorsicranium are completely bone enclosed in all species (including Luciolates} . On each side of the skull the continuous supraorbital- temporal canal opens to the exterior through several pores. Dorsal and lateral skull outlines are essentially similar in L. calcarifer, L. niloticus, L. macrophthalmus and L. longispinis except for a marked narrowing of the inter- orbital region in the two latter species, and a more forward position of the orbit in L. calcarifer. The ethmoid region is relatively short, and the parasphenoid runs REVIEW OF CENTROPOMIDAE 17 forward in the same line as the base of the braincase. The preotic skull proportions of the largest L. niloticus examined (neurocranial length 228 mm) are very similar to those in a much smaller L. calcarifer skull (103 mm long, from a fish of c. 40 cm S.L.), and differ from those in smaller L. niloticus skulls. The most noticeable differences apparent when these smaller L. niloticus skulls are compared with the skull of an equal-sized L. calcarifer are the relatively more anterior position of the orbit, and the much longer precommissural skull region in the latter species (see below, p. 20) . The skull proportions of large L. niloticus (i.e. skulls > 150 mm long), however, PTO PARC FR LATE LC PRO 10mm PAR PTO SOC SPO LATE EXO BOC PRO 5mm FIG. 4. (a) Lates longispinis. (b) L. macrophthalmus. Neurocranium in left lateral view. For nomenclature of L. longispinis see p. 12 et seq. 18 P. H. GREENWOOD come to resemble those of L. calcarifer more closely, the resemblance increasing with the size of the skull. Katayama (1956) figures the neurocranium from a L. calcarifer of 28-6 cm S.L. ; judging from this figure there is little difference between a skull of that size and one from a L. niloticus of comparable length. Seemingly the orbital and precommissural skull proportions change much more rapidly in L. calcarifer ; compare, for example, the 103 mm skull of L. calcarifer (S.L. c. 40 cm) with the 123 mm skull of L. niloticus (S.L. c. 48 cm) in Figs 3 and 2. Compared with the four species from outside Lake Tanganyika, three endemic Tanganyikan species, L. microlepis, L. mariae and Luciolates stappersi (Figs 5-8) show a distinct narrowing of the skull (particularly the braincase), an elongation of the ethmoid region, and an angling of the parasphenoid relative to the basi- occipital. The slope of the parasphenoid is steepest in L. mariae and least in L. microlepis, with Luciolates occupying an intermediate position in the series. The fourth Lake Tanganyika endemic, L. angustifrons (Fig. 5), is, in most features of its neurocranial profiles, intermediate between the other endemic species and those from outside the lake. Nevertheless, it is clearly differentiated from the latter by FRC PARC LATE RO 20mm SPO PARC PTO EPI SOC PAR FIG. 5. Lates angustifrons. Neurocranium in : (a) left lateral view, (b) dorsal view. Subgenus Lates (see p. 77) REVIEW OF CENTROPOMIDAE 19 the elongation of its ethmoid region and by the shape of its ethmoid bones, charac- teristics that unite it with the other endemic species from Lake Tanganyika (see below) . This elongation of the ethmovomerine skull region in all species of Lates (and Luciolates) from Lake Tanganyika immediately distinguishes the group (see Table i), TABLE i Relative length of ethmovomerine region in various Lates spp., and in Psammoperca waigiensis Ethmovomerine length as % of Neurocranial1 Ethmovomerine2 neurocranial Species length (mm) length (mm) length Lates calcarifer 103-0 25-0 24-2% L. niloticus 16-0 5-0 3I-3% 76-0 21-5 28-3% 124-0 30-5 24-8% 228-0 60-5 26-7% L. macrophthalmus 32-0 8-5 26-5% L. longispinis 59-5 16-5 28-7% L. angustifrons 120-0 43-3 36-8% L. mariae 26-0 10-0 38-5% 77'5 32-0 4I-3% L. microlepis 44-0 18-0 4i-o% L. stappersi 71-0 31-0 43'5% 7*-o 32-5 45-8% 103-5 49-o 46-9% Psammoperca waigiensis 43-0 14-5 33'7% 1 Neurocranial length : measured directly from the anterior tip of the vomer to the posterior point on the lower margin of the basioccipital facet for the first vertebra. 2 Ethmovomerine length : measured directly from the anterior point of the vomer to that point on the dorsicranium where the lateral ethmoid-prefrontal passes under lateral margin of the frontal. and argues strongly for their monophyletic origin. The concave posterior face of the lateral ethmoid in L. angustifrons, L. microlepis, L. mariae and Luciolates stap- persi, as compared with members of the L. calcarifer-L. niloticus complex, has a distinct posterior slope (cf. Figs 2-4 with 5-8). Furthermore, and most strikingly, the lateral margins of the bone are much wider (or, as it appears in lateral view, much deeper) and have a pronounced downward slope. In the L. calcarifer- L. niloticus group the posterior margin of the lateral ethmoid is almost vertically aligned, and its lateral margins are narrow and horizontally aligned (cf. Figs 2-4 with 5-8). Once again it is Luciolates that shows the most profound modifications with, in this instance, L. microlepis showing the least modified condition and L. angustifrons and L. mariae (in that order) occupying the intermediate places in the series. There is little interspecific variation in the morphology of the lateral ethmoids of L. calcarifer, L. niloticus, L. macrophthalmus and L. longispinis (see Figs 2-4). Subgenus Luciolates (see p. 71) P. H. GREENWOOD FRC PARC socs LATE FR V PS PRO 10mm FIG. 6. Lates microlepis. Neurocranium in left lateral view. All Lates species, and Luciolates, have three facets on each lateral ethmoid ; two, ventrally placed, are for articulation with the palatine, and the third (situated dorsolaterally above the posterior palatine facet) for articulation with the first circumorbital bone (lachrymal). The facets are less well defined in the Tanganyika species, and are most poorly differentiated in Luciolates. A noticeable feature of the skull in L. calcarifer, L. niloticus and, to a slightly lesser degree, in L. angustifrons is the way in which the anterior wall of the neuro- cranium (i.e. the prootic, pterosphenoid and ascending arm of the parasphenoid) are extended forward beyond the level of the lateral commissure (Figs 2, 3 & 5) ; in L. calcarifer and L. niloticus the tunnel-like ventral part of this extension surrounds all but the anterior half or more of the basisphenoid. This feature is emphasized when the skulls of these species are compared with those of L. mariae, L. microlepis and Luciolates stappersi, species in which there is only a slight prolongation of the neurocranial wall beyond the level of the lateral commissure (cf. Figs 2, 3 & 5 and 6-8). The situation in L. macrophthalmus and L. longispinis is virtually inter- mediate between those in the other two groups. (See Table 2 and Fig. 4.) Closer examination of the precommissural extension in specimens of L. calcarifer (neurocranial length, ncl., 103 mm), L. niloticus (ncl., 75 mm and above) and L. angustifrons (ncl., 120 mm) reveals the existence of a pterosphenoid pedicle which, through its contact with the parasphenoid anteriorly and the outer lip of a horizontal groove in the prootic, forms a semi-tubular bridge over the oculomotor and profundus nerves and the internal jugular vein (Figs 9 & 10). Rognes (1973) has called a similar stucture in labrids an internal jugular bridge, and that name will be used here. REVIEW OF CENTROPOMIDAE 21 TABLE 2 Precommissural skull proportion in Lates and Psammoperca Precommissural length as % of Neurocranial1 Precommissural2 neurocranial Species length (mm) skull length (mm) length Lates calcarifer 103-0 25-0 24'2% L. niloticus 76-0 10-0 I3'I% 124-0 22-0 *7'7% L. longispinis 59-5 6-0 10-1% L. macrophthalmus 32-0 3-0 9'4% 74-0 10-0 13-5% L. angustifrons 120-0 14-0 H'7% L. stappersi 71-0 3-5 4-9% 103-5 6-0 5-8% Psammoperca waigiensis 43-0 2-5 5*7% 1 See Table i (p. 19). 2 Precommissural skull length: measured directly from the anterior margin of the lateral commissure to the anterior margin of the ascending limb of the parasphenoid. The relative contributions of the prootic and parasphenoid bones to the internal jugular bridge show marked intraspecific variability, and usually differ on either side (see Fig. 90 -d). The parasphenoid contribution is always the least important, the major part of the ventral wall (and the entire groove) coming from the prootic, and the dorsal and lateral walls from the pterosphenoid pedicle. Except in the small L. niloticus skulls examined (see below) there is always some contact between the three bones at the orbital (i.e. front) margin of the bridge. Since the smallest available skulls of L. calcarifer and L. angustifrons measure 103 mm and 120 mm long respectively, no comment can be made on the interrelationship of these bones in small individuals of those species. Apparently correlated with the degree of pterosphenoid development and the development of the precommissural braincase is the extent to which the auto- sphenotic is prolonged anteriorly. The correlation is a positive one in species with an extensive precommissural braincase and a well-developed pedicle (i.e. L. calcarifer and L. niloticus). The anterior extension of the autosphenotic is least marked in Luciolates stappersi, L. mariae and L. microlepis, and is of intermediate length in L. angustifrons, L. macrophthalmus and L. longispinis. Before describing and comparing the precommissural crania for all Lates and Luciolates species, it is necessary to consider the ontogenetic changes involved in the production of an adult L. niloticus-type pterosphenoid pedicle and internal jugular bridge. The smallest L. niloticus skulls examined (12 mm long) have no noticeable pre- commissural extension of the braincase ; the parasphenoid does not contact the pterosphenoid and there is no bony bridge over the nerves and blood vessel (Fig. ga). At the ontogenetic stages represented by those skulls there is also no obvious P. H. GREENWOOD PTO PARC PRO EXO 10mm PARC PTO SOC FIG. 7. Lates mariae. Neurocranium in : (a) left lateral view, (b) dorsal view. pterosphenoid pedicle, but a narrow ligament runs from the lower, anterior part of the pterosphenoid to the outer rim of the weakly developed prootic groove lying below the internal jugular vein (Fig. ga) ; in effect, the ligament occupies the position later taken by the pterosphenoid pedicle arm of the internal jugular bridge. At its dorsal base, the ligament is attached to a small spur of bone on the pterosphenoid, which I would interpret as an incipient pedicle. In progressively larger skulls (i.e. to a length of 76 mm), there is a gradual develop- ment and down-growth of the pterosphenoid pedicle, and of a dorsally directed a LATE REVIEW OF CENTROPOMIDAE PAR FR EXO PRO 20mm SPO LC PRO LATE BOG 1C PTO SPO PTO EPI FR PAR FIG. 8. Lates stappersi. Neurocranium in : (a) left lateral view, (b) ventral view, (c) dorsal view. For details on the altered generic placement of this species (previously Luciolates stappersi) see p. 50. P. H. GREENWOOD spur-like development from the prootic lateral to the internal jugular groove. As a consequence of these growth patterns (and a dorsal extension of the ascending parasphenoid arm) an at first narrow (Fig. gb), but gradually broadening, bony ridge is formed over the internal jugular vein and the associated oculomotor and profundus nerves. Concurrently, there is a gradual forward growth of the pre- commissural region of the skull. A L. niloticus skull 76 mm long has the pterosphenoid pedicle and precommissural skull developed to an extent comparable with that in a L. angustifrons skull 120 mm long. Growth of the precommissural skull wall in L. niloticus continues beyond this 2mm 1mm FR PTS 5 mm FIG. 9. Outline figures of internal jugular bridge, pterosphenoid pedicle and precommis- sural skull to show in : (a) & (b) growth changes in L. niloticus and in : (c) & (d) variability in the bridge of a single specimen of L. longispinis. (a) Lates niloticus ; left lateral view, neurocranial length 12 mm. Note ligamentous connection between spur of pterosphenoid and process on prootic. (b) L. niloticus ; left lateral view, neurocranial length 16 mm. Note downgrowth of pterosphenoid spur (= pedicle) to join prootic process, (c) & (d) L. longispinis. Left and right sides of skull showing variation in the interrelationships of bones contributing to the internal jugular bridge. Note direct pterosphenoid -para- sphenoid contact in (d). REVIEW OF CENTROPOMIDAE LC PTSP PRO 10mm FIG. 10. Lates calcarifer. Outline figure to show relative hyperdevelopment of pre- commissural skull (left lateral view), especially the pterosphenoid pedicle and internal jugular bridge. Compare with Figs g(c) & (d), 5(a), 2(a). From a skull of 10-3 cm neurocranial length. point ; in a skull 124 mm long it has attained, however, the overall morphology and proportions seen in the largest skull examined (230 mm long). Unfortunately no L. angustifrons skulls longer than 120 mm could be obtained so it has not been possible to determine the definitive form in that species. However, to judge from the totality of interspecific differences seen in skulls of about the same size it seems unlikely that this region of the neurocranium in L. angustifrons ever attains the proportions found in either L. niloticus or L. calcarifer (see above, p. 21). As was noted earlier (p. 20), the precommissural skull region in L. macrophthalmus and L. longispinis is less well developed than in adult L. calcarifer and L. niloticus. It must, however, be remembered that members of the two former species reach a much smaller adult size (Worthington, 1929, 1932). In a L. macrophthalmus skull 74 mm long (from a fish of 275 mm S.L.) the internal jugular bridge and the pterosphenoid pedicle have about the same degree of develop- ment as in a 76 mm long skull of L. niloticus (S.L. c. 290 mm) or a 120 mm long skull of L. angustifrons (S.L. c. 350 mm) ; the situation is similar in a slightly larger individual of L. macrophthalmus (320 mm S.L., neurocranial length no mm) Both specimens have a narrow parasphenoidal contribution to the bridge which is otherwise formed mainly from the pterosphenoid pedicle and the prootic spur. The smallest L. macrophthalmus skull examined (32-5 mm long, from a fish of no mm S.L.) shows a degree of development comparable with that in a L. niloticus skull only 16 mm long ; namely, a ligamentous bridge, and the pterosphenoid pedicle manifest only as a small spur of bone (Fig. ga) . Conditions in L. longispinis, as seen in a skull 59 mm long (from a fish c. 250 mm S.L.), are close to those in the 74 mm skull of L. macrophthalmus, but the bridge is a little narrower. In a larger skull (70 mm long from a fish 275 mm S.L.) the bridge and pedicle, and the precommissural skull proportions are similar to those in the 26 P. H. GREENWOOD 76 mm skull of L. niloticus described above (p. 24), with a distinct pedicle and, at least on one side of the skull, a parasphenoidal contribution to the internal jugular bridge (Fig. gd) ; on the left side of this specimen, the ascending parasphenoid limb fails to reach the level of the upper lateral margin of the prootic (Fig. gc). It would seem, then, that the internal jugular bridge and the precommissural skull in both L. longispinis and L. macrophthalmus are comparable with those in similar-sized skulls of L. niloticus, or are perhaps a little less advanced in some individuals. In other words, the adult skull of L. macrophthalmus and L. longispinis retains at least some of the pre-adult features of L. niloticus. A really marked reduction in the adult precommissural braincase and in the pterosphenoid pedicle and internal jugular bridge is seen in the skulls of three Lake Tanganyika taxa, namely L. mariae, L. microlepis and Luciolates stappersi. (This region of the skull is also relatively reduced, as compared with L. niloticus, in the fourth Tanganyika species, L. angustifrons , see pp. 20-25 above.) In none of these three species does the parasphenoid contact the pterosphenoid, always being separated from that bone by the prootic (Figs 6-8). No trace of a pterosphenoid pedicle, even as a low ridge, is detectable in the three Luciolates stappersi skulls I have examined (neurocranial lengths 71 (f. 2) and 113 mm), but a low ridge was found in the largest of the three L. mariae skulls (26-0, 77-5 and 104-0 mm long). A similar ridge is developed on the right but not the left pterosphenoid of a 44 mm long skull of L. microlepis. A larger skull (95 mm) of L. microlepis, however, has a well-developed, broad-based but distally narrowed pedicle which reaches almost to the level of the prootic spur (Fig. 6) . It is connected to the prootic spur by a short section of what appears to be ossified ligament. Thus, of these three species, L. microlepis is the only one in which the ptero- sphenoid pedicle makes a significant contribution to the internal jugular bridge. Even in the largest skulls of L. mariae and Luciolates stappersi there is only a liga- mentous bridge, a condition directly comparable with that in the smallest specimens of L. niloticus, except that in the Tanganyika fishes the ligament appears to be ossified. In other words, the precommissural braincase in large specimens of L. mariae and Luciolates stappersi (standard lengths 390 and 415 mm respectively) is like that in L. niloticus of about 60 mm standard length, while that of a L. micro- lepis 390 mm standard length is comparable with a L. niloticus of about 130 mm S.L. The pterosphenoid -prootic ligament found in juvenile L. niloticus and adults of Tanganyika taxa described above is readily separated from both its bones of attach- ment. Thus it seems unlikely that it is truly part of the pterosphenoid pedicle. Presumably the ligament is replaced by the pedicle as it grows down to meet the spur from the outer rim of the prootic groove. The large L. microlepis specimen noted above represents a late phase in this developmental sequence, the small L. macrophthalmus (ncl., 32-5 mm ; p. 25) an early phase, and the adult condition in L. niloticus, L. calcarifer and L. angustifrons the terminal state. An internal jugular bridge is of sporadic and phyletically widespread occurrence amongst living teleosts. Rognes (1973) gives detailed accounts of the bridge in labrine Labridae, and reviews records of its occurrence in other groups. I can REVIEW OF CENTROPOMIDAE 27 confirm its presence in certain ostariophysans (Alburnm ; see also Holmgren & Stensio, 1936, for Abramis), certain scorpaeniforms (Enophrys bison, Scorpaena scrofa, Trigla hirudo, see also Allis, 1909 ; Allen [1905] describes a bridge in Ophidian [Hexagrammidae]), and in several percoids (Epinephelus species [but not other serranids], Stizostedion volgensis, Perca fluviatilis [but not, apparently, in Gymno- cephalus}}, and in some sphyraenoids (Sphyraena sp.). In the majority of cases where a bridge is present, it is of the type found in juvenile L. niloticus, namely a ligament (generally ossified) joining a reduced pterosphenoid pedicle to a process developed on the prootic (see above, p. 21). Only in Enophrys bison is a bridge of the L. angustifrons type present. This list, based on samples taken from the families represented in the dry skeleton collection of the British Museum (Natural History), cannot by any means be con- sidered complete, especially since the bridge is not always preserved in dry skeletons. Nevertheless, it is interesting to find that in none of the beryciform skeletons at my disposal is there any indication of a bridge nor even of the pterosphenoid pedicle (which is usually obvious even if the ligamentous part of the bridge is missing). Neither a bridge nor a pterosphenoid pedicle was noted in any of the Mesozoic beryciforms described by Patterson (1964). Superficially, Salmo trutta has what appears to be an internal jugular bridge, but closer inspection shows that it is formed entirely within the prootic. Thus it would seem to be homologous with the 'prelateral commissure' described by Rognes (1973) in the labrid Ctenolabrus exoletus (see Rognes, op. cit., fig. 59). The pterosphenoid pedicle has a long history in actinopterygian fishes, being well developed in some leptolepids and pholidophorids, in Amia and its fossil relatives Sinamia and Ellenes, and at least partially developed in some palaeoniscids (Patter- son, 1975). As Patterson (op. cit., p. 409) observes : ' ... It is therefore likely that a pterosphenoid pedicle of some sort, or at least the potentiality to develop such a structure is a primitive actinopterygian feature.' Since the pterosphenoid pedicle is an integral part of the internal jugular bridge (see above) and because this bridge is of widespread occurrence among teleosts, one may conclude that the bridge too is a primitive feature. The absence or great reduction of the bridge and pedicle in certain Lates species can, therefore, be interpreted as an apomorphic feature, at least when individuals of these species attain a size at which the bridge would otherwise be present in related taxa. Lates macrophthalmus and L. longispinis (both species with reduced bridges) are examples of the situation where maximum adult size is about equal to that in preadult L. niloticus and L. calcarifer ; at that size, specimens of L. niloticus (and presumably L. calcarifer} have a poorly developed bridge. Thus, it is probably correct to consider L. macrophthalmus and L. longispinis as plesiomorphic with respect to the bridge character. Hyopalatine arch and the preoperculum (Figs n & 12) Apart from slight proportional changes in, particularly, the length of the palatine and ectopterygoid bones of the Tanganyika species, there is little interspecific variation in the hyopalatine arch of Lates species (see Figs n & 12). P. H. GREENWOOD PAL POP. FIG. ii. Hyopalatine arch, right (including preoperculum) in lateral view of (a) Lates mariae, (b) L. niloticus. The hyomandibula has two well-defined articulatory facets interconnected by a thin lamina of bone. The metapterygoid has a strong sutural union with the hyomandibula and with the posterior tip of the expansive endopterygoid. There is no true metapterygoidal lamina (sensu Katayama, 1956, and Gosline, 1966) but a slight ridge is detectable in the position where a lamina would occur ; also, in many species there is a small foramen (or fenestra) in the metapterygoid at the postero-dorsal end of the ridge. I would interpret these structures as the remnants of a greatly reduced metaptery- goidal lamina. Fine viliform teeth cover the entire ventral surface of the palatine. A similarly shaped (i.e. elongate ovoid) tooth patch occurs on the medial aspect of the anterior REVIEW OF CENTROPOMIDAE 29 arm of the ectopterygoid, sometimes extending a short way onto the vertical arm of that bone as well. The autopalatine is a fairly stout bone. Anteriorly, on its medial face are two well-defined articulatory surfaces for contact with the ethmoid ; dorsally there is a weakly demarcated facet for articulation with the lateral ethmoid. A panhandle- like, cartilage-tipped projection from the upper surface of the palatine provides articulation between this bone and the maxilla. In most details, including the presence of a reduced metapterygoidal lamina, the hyopalatine arch of Luciolates resembles that of Lates, particularly the Lake Tangan- yika species of the genus. However, all the bones (especially the endopterygoid) are thinner and the palatine is less robust, with poorly demarcated articulatory facets. The palatine tooth patch is much narrower in Luciolates, and there is a great 10mm PAL POP 10mm FIG. 12. Hyopalatine arch, left side (including preoperculum) in lateral view of (a) Lates stappersi, (b) L. angustifrons. 30 P. H. GREENWOOD reduction in the area of the ectopterygoid teeth, the tooth-patch being either reduced to a small oval near the ectopterygoid-palatine articulation or it is completely absent. In one of the three skeletons examined the tooth-patch was present on one side only. Like the hyopalatine arch, the preoperculum in Lates shows little interspecific variation, although it does show some intraspecific variability. The entire posterior margin of the vertical limb, except for a short length near its ventral angle, is finely serrate, the individual serrae are slender, sharp-pointed and tall. In very large specimens of L. niloticus (> 150 cm S.L.), the serrations are considerably reduced in height, and consequently the posterior margin of the bone is merely irregular (see also Sorbini, 1973). At the posterior angle between the horizontal and vertical preopercular limbs there is a large, posteriorly directed and triangular spine (Fig. n) ; very rarely this spine is subdivided almost to its base, resulting in two narrower but still triangular spines. On the horizontal limb there are generally three triangular spines, each slightly shorter and narrower than the spine at the bone's posterior angle. In larger L. niloticus the spines become irregular in outline, relatively shorter, and may have rounded rather than acute points. Although three preopercular spines are modal for all species but L. macrophthalmus, a fourth spine is sometimes developed either on one or both sides. Usually the extra spine is a distinct entity, but sometimes it appears merely to be a subdivision of one of the other spines. Lates macrophthalmus is apparently exceptional in having a high proportion of individuals with four spines (seven of the eleven specimens examined). The proportion of four-spined fishes amongst samples of the other species is : L, calcarifer, none out of 18 ; L. niloticus, 7 out of 31 ; L. longispinis, 3 out of 6 (a high proportion, approaching that of L. macrophthalmus, which may be a related taxon, see p. 13) ; L. angustifrons, none out of 14 ; L. mariae, 4 out of 20 ; L, microlepis, 2 out of 27. The occurrence of four-spined individuals may be a population feature, hence my reservations about the seemingly unusual condition in L. macrophthalmus. All but one of the L. niloticus specimens with four spines came from a single sample (incidentally, the largest available for L. niloticus and one much larger than was available for any other species). Luciolates stappersi (Fig. I2a) also has a serrated posterior margin to the vertical preopercular limb, but here the serrations are lower and less well defined (in this respect resembling the condition in 16-20 mm standard length L. niloticus}. The spine at the preopercular angle is always present and prominent, although it is somewhat finer than in any Lates species. The horizontal limb may have three large and relatively short spines, but specimens with two or three groups of very small spines, or even what amounts to a crenellated border, are common. The incidence of bilateral asymmetry in the type of spination is also high. In both Lates and Luciolates the preopercular lateral line canal is completely bone enclosed, with its pores confined to the horizontal limb. Although a serrated or otherwise ornamented vertical preopercular limb is of common occurrence amongst the lower percoids (e.g. in the Serranidae), the presence REVIEW OF CENTROPOMIDAE of large and discrete spines on the horizontal limb and at its angle is extremely rare (Percalates and Siniperca [Serranidae] are, as far as I can determine, the only taxa having the same type of preopercular ornamentation as Lates). A similar generali- zation can be made for the lower percomorphs (sensu Rosen, 1973 ; for example, the 'Beryciforms'). Thus, it seems reasonable to conclude that the ventral pre- opercular ornamentation in Lates (and probably other centropomids as well, see below) is a derived condition (see also Rosen, 1973 : 469). Luciolates too can be included in this generalization, the condition here being interpreted as the secondary simplification of a derived condition mimicking a plesiomorphic one. Circumorbital bones (Figs I3a-d) The greater part of the ventral margin to the first circumorbital bone (the lachry- mal) is finely serrated in all Lates species ; only a short anterior part is smooth. In all species the entire margin of the second circumorbital is also serrated. The infraorbital lateral line canal in Lates and in Luciolates is enclosed throughout its length, communicating with the exterior through five pores in the lachrymal, one anteriorly on the third circumorbital bone, and through other pores found between successive bones in the series. LAC SOS LAC LAC L_LAC FIG. 13. Circumorbital bones (right side) in : (a) & (b) Lates niloticus, and in : (c) & (d) L. stappersi ; (a) and (c) lateral view, (b) and (d) viewed dorsally and somewhat anteriorly. 32 P. H. GREENWOOD All species (including Luciolates stappersi) have a well-defined facet developed at about the middle of the upper lachrymal margin ; it articulates with a similar facet on the lateral ethmoid. There is a general similarity in the shape of the first two circumorbital bones in all Lates species, although the three species from Lake Tanganyika (L. angustifrons, L. mariae and L. microlepis} have a slightly more elongate lachrymal. These species (except L. angustifrons) also differ from L. niloticus, L. calcarifer, L. macroph- thalmus and L. longispinis in having a relatively more elongate fifth circumorbital, and in having much narrower bony flanges developed from the ventral contours of the cylindrical canal-bearing portions of the third, fourth and fifth bones. Greatest departure from the L. niloticus -L. calcarifer situation is seen in the re- duced size of the subocular shelf in the Tanganyika species, again excepting L. angustifrons where the shelf is like that in L. niloticus and L. calcarifer, viz. a thin but broad bony plate that curves upwards from the third circumorbital to He along the entire length of the fourth bone. In L. microlepis the subocular shelf is reduced in width, and just reaches upwards to the level of the articulation between the third and fourth circumorbitals ; in L. mariae there is a further and marked reduction in width, and the shelf barely reaches to the level of the articulation between the bones. Both species have the ventral flange on the third and fourth circumorbitals reduced to a thin flange. These reductional trends are carried further in Luciolates, where the serrations on the lachrymal are very weak and are confined to about the posterior third of the bone ; serrations are completely absent from the second circumorbital. The facet for articulation with the lateral ethmoid is weakly developed, and its origin from the lachrymal is far less well defined than in the other species. The subocular shelf is, relatively, a little narrower than in L. mariae, but it does extend further up the fourth circumorbital (along about its lower third) ; see Fig. I3c-d. The depth of the ventral flange on the second to fourth circumorbitals is almost comparable with L. mariae, as is the flange on the fifth bone. In their gross morphology, the circum- orbital bones in Luciolates stappersi are noticeably more elongate than those in any Lates, including the other Lake Tanganyika species. Apart from differences in the overall proportions of the first, third and fourth bones, the morphology of the entire series in an adult Luciolates of 105 mm standard length closely resembles that in a juvenile Lates niloticus 32 mm long. Opercular bones (Figs I4a-b) There is little variation in the operculum, suboperculum and interoperculum of Lates and Luciolates, apart from a slight relative elongation of the interoperculum in the Tanganyika species, especially Luciolates. In all taxa there is a well-defined, curved ridge on the medial face of the interoperculum against the upper, concave surface of which the proximal end of the epihyal articulates. The operculum (Fig. I4a-b) is armed with a single stout spine formed from the posterior tip of the near-horizontal strut which runs backwards from the hyo- manibular facet of the bone. REVIEW OF CENTROPOMIDAE OP 33 SOP 10mm 10mm IOP OP. SOP IOP FIG. 14. Opercular series (medial aspect of bones from right side) in : (a) Lates angustifrons, (b) L. stappersi, (c) Psammoperca waigiensis. A characteristic feature in all taxa is the thinness of the sub- and interopercular bones. Jaws (Figs 15 & 1 6) Both the maxilla and the premaxilla show little interspecific variation amongst Lates species, and are of the generalized percoid type. There is also little difference between Lates and Luciolates in the morphology of these bones. However, in Lates species the ascending process of the premaxilla is from 30 to 60 per cent higher than the articular process (apparently being lowest in the Lake Tanganyika species) ; it is only a little higher than the ascending process in Luciolates stappersi. The premaxillary dentition in all Lates species is composed of numerous close-set rows of small conical to subconical teeth which form a villiform covering to the complete width of the bone over almost its entire length (Figs I5b-c). 34 ASCR P. H. GREENWOOD 10mm PMAXP FIG. 15. Lates niloticus. (a) Premaxilla (left) lateral view, (b) Dentary (left) lateral view, (c) Dentary (left) occlusal view. (All from Greenwood & Howes, 1975.) 5mm ASCP FIG. 1 6. Lates stappersi. (a) Premaxilla (right), occlusal view, (b) Premaxilla (right), lateral view, (c) Dentary (right), lateral view. REVIEW OF CENTROPOMIDAE 35 Although most premaxillary teeth in Luciolates stappersi are like those in Lates, the species is noteworthy for the presence of at least one greatly enlarged and two slightly smaller caniniform teeth adjacent to the symphysial surface of the pre- maxilla ; a few neighbouring teeth may also be somewhat enlarged. In general the larger teeth are linearly arranged, with the largest one situated lingually. The upper jaw elements in Lates and Luciolates show no derived characteristics and, of course, both genera retain the supramaxilla. The enlarged median teeth of Luciolates, however, would seem to be a derived feature. Like the upper jaw, the lower jaw elements (dentary, anguloarticular and retro- articular) show little interspecific variation. In Luciolates the anguloarticular is relatively shallow, but otherwise has a typical 'Lates' form. The dentition of the dentary mirrors that on the premaxilla, except that in Luciolates the outermost tooth row is composed of noticeably larger and more clearly caniniform teeth, and there are no enlarged symphysial teeth. Branchial skeleton (Figs 17-19) The branchial skeleton in both Lates and Luciolates is of a generalized percoid type (see Rosen, 1973), and it shows few interspecific differences, apart from a relative E2TP E2 E4 E3 E2 E3TP 1mm FIG. 17. Lates niloticus. Branchial skeleton, dorsal part (drawn from an alizarin pre- paration, 40 mm S.L.). (a) Dorsal aspect of left side, (b) Ventral aspect (left side) to show upper pharyngeal teeth and tooth plates. 34 ASCR P. H. GREENWOOD FIG. 15. Lates niloticus. (a) Premaxilla (left) lateral view, (b) Dentary (left) lateral view, (c) Dentary (left) occlusal view. (All from Greenwood & Howes, 1975.) ASCP 5mm FIG. 1 6. Lates stappersi. (a) Premaxilla (right), occlusal view, (b) Premaxilla (right), lateral view, (c) Dentary (right), lateral view. REVIEW OF CENTROPOMIDAE 35 Although most premaxillary teeth in Luciolates stappersi are like those in Lates, the species is noteworthy for the presence of at least one greatly enlarged and two slightly smaller caniniform teeth adjacent to the symphysial surface of the pre- maxilla ; a few neighbouring teeth may also be somewhat enlarged. In general the larger teeth are linearly arranged, with the largest one situated lingually. The upper jaw elements in Lates and Luciolates show no derived characteristics and, of course, both genera retain the supramaxilla. The enlarged median teeth of Luciolates, however, would seem to be a derived feature. Like the upper jaw, the lower jaw elements (dentary, anguloarticular and retro- articular) show little interspecific variation. In Luciolates the anguloarticular is relatively shallow, but otherwise has a typical 'Lates' form. The dentition of the dentary mirrors that on the premaxilla, except that in Luciolates the outermost tooth row is composed of noticeably larger and more clearly caniniform teeth, and there are no enlarged symphysial teeth. Branchial skeleton (Figs 17-19) The branchial skeleton in both Lates and Luciolates is of a generalized percoid type (see Rosen, 1973), and it shows few interspecific differences, apart from a relative E2TP E4 E3 E2 E3TP 1mm FIG. 17. Lates niloticus. Branchial skeleton, dorsal part (drawn from an alizarin pre- paration, 40 mm S.L.). (a) Dorsal aspect of left side, (b) Ventral aspect (left side) to show upper pharyngeal teeth and tooth plates. 38 P. H. GREENWOOD elongate so that it overlaps the joint between the two bones. The plates associated with the third arch are generally the largest of the series, approach one another medially and cover a great deal of the third basibranchial. In the one available branchial skeleton of L. angustifrons the individual plates appear to have fused together on the first two gill arches to form a long tooth plate on each side of the arch. A similar arrangement is seen in the alizarin preparation of a small (96 mm S.L.) Luciolates stappersi, but this specimen differs in other respects, especially in having a single, median plate on the third basibranchial and a small plate on each hypobranchial of that arch. The arches dissected from a much larger specimen (270 mm S.L.) have the plates of the third basibranchial narrowly separated medially, a long plate at the base of the second gill arch and a small plate intercalated between it and the basal plate of the first arch. Clearly, at least in this species, there can be quite considerable individual variability in the pattern of tooth plate distribution (see Nelson, 1969 : 500-501, for a description of variation in another percoid, Pomatomus saltatrix [Pomatomidae]). Hyoid arch (Figs 20 & 21) The hyoid arch in Lates and Luciolates is of a basal percoid type, with dorsal and ventral hypohyals, a large and complete 'berycoid' foramen and seven branchio- stegal rays. There is remarkably little interspecific variability in the shape of this Eh 'BcF' BrR 10mm 10mm FIG. 20. Hyoid arch and branchiostegal rays (right side), viewed laterally, in (a) Lates stappersi, (b) L. mariae, (c) L. niloticus. REVIEW OF CENTROPOMIDAE 39 10mm FIG. 21. Urohyal. (a) & (b) Lates stappersi (left lateral and ventral views respectively), (c) & (d) L. angustifrons (left lateral and ventral views respectively). arch, without even, as might be expected, clear-cut proportional differences in the arches from species with elongate skulls (i.e. the Tanganyika species). The first four branchiostegal rays articulate with the ceratohyal, the fifth with either the ceratohyal or at the cerato-epihyal suture, and the last two rays (the stoutest and broadest of the series) articulate with the epihyal. The first three rays contact the ventral face of the ceratohyal, the other four lie on the lateral aspect of the cerato- or epihyal. These latter rays have progressively broader heads, with the dorsal outline of the head on the last two, or occasionally three rays somewhat indented. The basihyal is an elongate bone, spatulate in dorsal outline, and does not carry a tooth plate. The urohyal (Fig. 21) is similar in all species, but is markedly more elongate in Luciolates, even when it is compared with the urohyal in the Lates species of Lake Tanganyika. Pectoral girdle and associated bones (Fig. 22) The pectoral girdle shows few interspecific or intergeneric differences, either in its overall proportions or in the shape of its individual bones. Judging from the only 4o P. H. GREENWOOD sc. CL CL COR COR FIG. 22. Pectoral girdle (right half) in : (a) Lates calcarifer, (b) L. stappersi. (The supracleithmm is removed from this specimen.) available skeleton of Luciolates stappersi the horizontal limb of the cleithrum is somewhat narrower than it is in Lates, and has less ventrolateral curvature ; the scapula and coracoid are also noticeably deeper in this species and the foramen enclosed between the coracoid and the medioventral margin of the cleithrum is larger (cf. Figs 22a & 22b). The posterior angle of the cleithrum in both genera is expanded and slightly protracted, and its hind margin is serrated. These serrations are most numerous in L. calcarifer, L. niloticus and L. angustifrons (6-10 serrae, the uppermost often ill-defined), are fewer in L. macrophthalmus (5-7) and fewest (3 or 4) in L. microlepis and L. mariae. Judged on the size range of available material for any one species, the number of serrae is not obviously correlated with the fish's size, and the number may differ on either side of an individual. In Luciolates the cleithral projection can have a smooth posterior border or be ornamented with from one to three weak serrations ; as in Lates there are lateral discrepancies in the number of serrae. The three upper radials articulate with the scapula, and the fourth either articu- lates with the coracoid or partly with the coracoid and partly with the scapula. The supracleithrum in both Lates and Luciolates is a slightly curved, dagger- shaped bone showing no interspecific variability in shape or size. The first postcleithrum is a flat, scale-like bone, the second is elongate and spini- form (Fig. 23). No obvious interspecific or intergeneric differences were detected in either element. The posttemporal is characterized, in both genera, by a deep and dorsally directed oval pocket formed in the body of the bone immediately lateral to the base of its REVIEW OF CENTROPOMIDAE PC2 FIG. 23. Lates calcarifer. Postcleithra (left). intercalar limb and a little anterior to the facet for articulation with the supra- cleithrum (Fig. 24). The pocket opens dorsally into the lateral line canal, and its lateral wall bulges slightly outwards ; in alizarin preparations of a young L. niloticus this wall has a pitted, 'strawberry-skin' appearance similar to that of the auditory bulla in many clupeomorph fishes. The pit is occupied by the distal end of the ligament which runs from the posttemporal to the tunica externa of the swimbladder (see p. 47 below). Posteriorly, the shield-like body of the posttemporal is serrated, the extent and size of the serrations apparently not differing between the various species. The extrascapula is a small Y-shaped bone, largely occupied by the lateral line sensory canals it carries (i.e. the supratemporal and temporal lines), and shows little FIG. 24. Posttemporal in : (a) & (d) Lates niloticus (right bone), (a) lateral, (d) medial aspect, (b) & (e) Psammoperca waigiensis (left bone), (b) lateral, (e) medial aspect, (c) & (f) Centropomus undecimalis (left bone), (c) lateral, (f) medial aspect. 42 P. H. GREENWOOD interspecific variability. It articulates closely with the posttemporal, the two bones together partially covering the posterior part of the posttemporal fossa. Vertebral column (Figs 25 & 26) The total count in all Lates species and in Luciolates stappersi is 25, viz., n ab- dominal, 13 caudal and the fused first ural and preural centra. There are nine pairs of pleural ribs, the first pair associated with the third vertebra. On those vertebrae with parapophyses, (the eighth and subsequent abdominals have obvious parapophyses but a small projection is visible on the seventh), the ribs articulate with the posterior face of the parapophysis ; at least in Lates the rib articulation on the preceding centra is through a shallow facet whose ventral lip is slightly produced laterally. The parapophyses in Luciolates differ from those in Lates in being almost vertically aligned, and by having, in all bar the first pair, a horizontal strut joining the para- pophyses of each centrum near their distal tips. Also, in this genus the articulatory pit on the first three rib-bearing centra has no ventral lip, but on the fourth rib- bearing centrum (i.e. the sixth abdominal vertebra) the lip is sufficiently produced to resemble a very short parapophysis. I have been able to check the dorsal ribs in only two species (L. niloticus and Luciolates stappersi). Lates niloticus has epineural ribs associated with the first FIG. 25. First three abdominal vertebrae in : (a) Lates angustifrons, (b) Psammoperca waigiensis, (c) L. stappersi, (d) Centropomus ensiferus. REVIEW OF CENTROPOMIDAE 43 two vertebrae, and epipleurals present on the first six pairs of pleural ribs ; Luciolates stappersi has epineurals as in Lates, but the epipleurals apparently are confined to the first three pairs of ribs only. The first three abdominal vertebrae are the most individually distinctive elements in the entire column (Fig. 25). The second vertebra is characterized by the great expansion anteroposteriorly of its neural spine, which is 2-3^ times broader than the spine of the first vertebra and about twice as broad as the spine of the third vertebra. Not only is the spine expanded but it has a characteristic outline. The anterior and posterior margins run almost parallel to one another for most of the spine's height (rather than converging with one another), and the spine narrows smoothly at a point about three-quarters of its height above the centrum. At this point, the front margin curves backwards to meet the posterior margin which may be almost vertical or, and more generally, it may have a slight posterior curvature. The hind margin of the first neural spine is closely applied to the front of the second spine, but the third neural spine slopes away from the second at a marked angle. Neither the first nor the third neural spine has parallel margins except basally ; the margins slope towards one another over most of their height, giving the spine a narrowly triangular outline. A variety of skeletons covering a wide size range of individuals (c. 16 to 1000 mm) is available only for L. niloticus. These skeletons indicate that the relative antero- posterior expansion of the second neural spine may at first show a positively cor- related increase with increasing standard length, but that in very large fishes the spine becomes relatively narrower. There are quite marked interspecific differences in the length -height proportions of certain centra, particularly those in the abdominal region of the column. In the descriptions that follow the first three abdominal vertebrae are excluded since those are not affected by proportional changes ; all measurements are maxima. In L. niloticus, L. macrophthalmus and L. longispinis, the abdominal and caudal centra are of approximately equal length and depth or are only a little longer than deep (the latter proportions applying especially to caudal vertebrae) . Lates calcarifer has caudal centra like those in the former species but its abdominal centra are slightly more elongate. The abdominal centra in L. angustifrons have proportions similar to those in L. calcarifer, as do the first five or six caudal centra. Beyond that point, however, the caudal centra are noticeably more elongate (i.e. they are about i| times longer than deep). Lates mariae shows slightly greater elongation of its abdominal centra (c. 1 1 times longer than deep), but the caudal centra are similar to those in L. angustifrons. This trend is accentuated in L. microlepis where, although the abdominal centra have proportions like those of L. mariae, the posterior caudal elements are from if to twice as long as deep ; the anterior caudals, however, are still about i \ times longer than deep. Finally, in Luciolates, all the centra are clearly elongate (c. if to twice as long as deep) and there is no difference in pro- portions between the caudal and abdominal elements of the column. There are three predorsal bones in all Lates and Luciolates (pace Fraser, 1968), the proximal end of the first lying just anterior to the first neural spine, the ends of 44 P. H. GREENWOOD the second and third bones lying, respectively, in front of and behind the second neural spine. Caudal fin skeleton (Fig. 26) There is but slight interspecific variation in the caudal skeleton of Lates (Fig. 26). All species have two epurals, two uroneurals and five hypurals ; the first, second and fifth hypurals are autogenous (as is the parhypural and the haemal arch and spine of the third preural centrum). The hypurapophysis is weakly to moderately developed. The neural spine on the second preural vertebra is reduced to a low crest in all species. The principal caudal fin ray formula for all species is 1,8 + 7,1. Although the caudal skeleton in Luciolates is basically similar to that in Lates, it differs in having the first to fourth hypurals fused into a single plate except for a narrow proximal gap between the fused first and second, and the fused third and fourth hypurals ; the fifth hypural is free and is autogenous basally. One small specimen (96 mm S.L.) of Luciolates stappersi, an alizarin preparation, has a small and free sixth hypural, the fifth hypural in this specimen being fused in with the third and fourth. Unlike the other Luciolates examined (by dissection and radiographically) the second and third hypurals in this fish are not apposed over their distal halves but are fused proximally instead. As in Lates, there are 1,8 + 7,1 principal caudal rays in Luciolates. The caudal fin margin in adult L. calcarifer, L. niloticus, L. macrophthalmus , L. longispinis and L. angustifrons is weakly truncate to markedly subtruncate (nearly rounded), in L. mariae it is truncate to weakly emarginate, but in L. micro- lepis it is so strongly emarginate as to be almost crescentic. (In juveniles, however, NaPU2 PU-i+U-i UR Caudal fin skeleton in : (a) Lates niloticus, (b) Eolates gracilis (After Sorbini, 1973). REVIEW OF CENTROPOMIDAE 45 the margin is distinctly truncate [see Poll, 1953] or weakly subtruncate [see Boul- enger, 1915].) A crescentic margin is also developed in Luciolates stappersi, and is deepest in fishes over 150 mm standard length. Dorsal and anal fins The number of pterygiophores supporting the rays of the dorsal fin (or fins) shows some slight interspecific variation ; viz : L. calcarifer 18, L. niloticus 18 or 19, L. microlepis 19 (rarely 20), L. macrophthalmus 18 (rarely 19), L. longispinis 18 or 19, L. angustifrons 19 and L. mariae 19. Each of the first eight or nine pterygiophores carries a single spine, and no medial radials are associated with these bones. An examination of the dorsal fin ray supports in alizarin preparations of small (16-20 mm S.L.) L. niloticus suggests that the medial radial fuses with the proximal one (the pterygiophore) to form the elongate head of that bone. Distinct medial radials are also absent from those pterygiophores carrying the branched dorsal fin rays. Luciolates stappersi has 19 dorsal pterygiophores, the first nine of which bear a single spine (again without the interposition of a medial radial). The seventh and eighth pterygiophores have markedly elongate heads, and each carries a short weak spine which is largely embedded in the epaxial body musculature. Superficially, these spines are well separated from each other and from the first and second dorsal fins. The ninth pterygiophore carries a longer and somewhat stouter spine which is the first ray of the second dorsal fin. Unlike Lates, the posterior branched rays of the dorsal fin in Luciolates do have distinct medial radials, even in the largest individuals examined. The wide gap between the dorsal fins of Luciolates was, and in published accounts of this taxon still is, the principal diagnostic feature for the genus. It is therefore of some importance to reconsider the relative positions of the dorsal fins (or of its two sections where the fin is apparently a single unit, as in L. niloticus}. As Poll has described (Poll, 1953) and I have been able to confirm, the Lake Tanganyika species of that genus show ontogenetically correlated changes in relative fin position. However, my observations also indicate that the definitive fin positions in these species are reached well before the cessation of obvious growth in body length. Lates calcarifer (as compared with L. niloticus} has a distinctly greater interval between the last and first spines of the two fins than that existing between the penultimate and last spines of the first fin ; the gap is bridged by a low membrane. In L. niloticus the spacing between these three rays is almost equal, and the inter- connecting membrane appears to be slightly deeper. The condition in L. macroph- thalmus and L. longispinis approaches that in L. niloticus but with a slightly greater distance between the spines in L. macrophthalmus. The condition in L. angustifrons (the seemingly most generalized of the Tanganyika species) is either comparable with that in L. calcarifer or, in some individuals, the inter-fin spacing may be a little greater. Some specimens I have examined (up to 345 mm S.L.) have no membrane connecting the two fins and in a few the 'last' spine of the first dorsal is not connected with the rest of the fin ; it is impossible to 46 P. H. GREENWOOD tell whether this latter condition is the result of damage. The smallest fish measured (90 mm S.L.) has a distinct but membrane-spanned gap between the fins. None of the L. microlepis examined has a membrane connecting the fins, and in several there is an isolated spine in the gap. The usual condition in L. mariae (except in fishes < 70 mm S.L.) is a distinct gap between the fins, with a single, isolated spine lying at about its midpoint. This species is unusual in having a modal dorsal spine count of nine (eight is the mode in other Tanganyika species, although occasional individuals with nine spines are recorded ; see Poll, 1953). Within the Lates species of Lake Tanganyika then, one finds a complete inter- gradation between a continuous, albeit deeply notched dorsal fin, and two separate fins with an isolated spine interposed. The condition in Luciolates differs from the latter state only in the greater width of the gap and the occurrence of two spines within it. Luciolates is, however, unusual in having only six spines in the first fin 2ndASp 2ndASo FIG. 27. First anal pterygiophore, and abdominal -caudal vertebral transition, in : (a) Lates niloticus, (b) Centropomus ensiferus (drawn from radiograph 1903.5.15:3-5 and dry skeleton 1861.12.12:13). REVIEW OF CENTROPOMIDAE 47 and one spine with only 9 or 10 branched rays in the second (compared with the usual seven spines and one spine plus 11-13, rarely 10, branched rays in the fins respectively). The two isolated spines in Luciolates may therefore represent the detached 'ultimate' and 'first' rays respectively of the ancestral type fin, with what we now consider to be the first spine of the second dorsal fin a neomorphic develop- ment from a branched ray. Alternatively, and as would seem more probable, the ancestral species could have had seven spines in the first dorsal fin, an isolated spine between it and the second dorsal, and the latter comprising one spine and 10 branched rays (a condition found in some specimens of L. mariae). The anal fin skeleton is similar in all Lates species. There are nine, rarely eight, pterygiophores, the first a large double structure carrying two spines (Fig. 2ya) ; it articulates with the cross-bar on the haemal arch of the first abdominal vertebra. All other anal pterygiophores, except the last, carry a single ray (that on the second a spinous one) ; the last pterygiophore carries two rays. Medial radials are absent except on the last three or four pterygiophores. Morphologically the anal fin skeleton of Luciolates is like that in Lates, although the first pterygiophore is less robust and there are nine others in the series (i.e. a total of 10). A medial radial is present in the last four pterygiophores. Swimbladder One outstanding feature of the swimbladder in Lates and Luciolates is the presence of a tough connective tissue strap running from a point anterodorsally on the tunica externa to the posttemporal, which has a well-defined ventrolateral recess for the reception and anchorage of the strap (see above, p. 41). Katayama (1956) does not describe this connection in L. calcarifer but I have been able to confirm its presence in that species. Apart from Psammoperca (see p. 60 below), I know of no other percoid species in which a similar swimbladder -posttemporal connection has been described, nor indeed of any connection between those two points. The functional significance of a swimbladder -posttemporal linkage is not readily apparent. The anterior end of the swimbladder in all Lates species and in Luciolates stappersi has a deep median indentation which gives that end of the swimbladder a distinctly bilobed appearance. Baudelot's ligament This ligament is well defined in Lates and Luciolates, and originates from a deep pit on the basioccipital. In L. niloticus and Luciolates stappersi (the two species dissected) little or no epaxial body musculature runs below the ligament medially ; laterally, however, there is a broad muscle band passing below and above it to insert partly on the anterolateral aspect of the basioccipital but mainly on the exoccipital. Thus at least the distal half of Baudelot's ligament is embedded in muscle. The relationship of the ligament to the epaxial musculature seems to combine certain features of both the percichthyid and serranid types described by Gosline (1966), but is more akin to the serranid type. 48 P. H. GREENWOOD Lateral line (Fig. 28) In all Lates species the pored lateral line scales of the body continue onto the caudal fin where they extend, or almost extend, to the posterior margin of the fin. Since the posterior margin is generally abraded or damaged it is difficult to tell in the latter cases whether the absence of scales from the immediate marginal area is artef actual or not. Two other rows of pore-bearing scales are present on this fin, one lying above and the other below the median row (from which they are separated by a space usually equal to that between two fin rays). These upper and lower scale rows generally do not quite extend to the posterior fin margin. Because of their small size the scales in these rows are difficult to see in fresh and spirit-preserved specimens unless the fin is allowed to dry out completely. Superficially there does not seem to be any linkage between the median and the other caudal lateral line scale rows ; scales in the latter rows cease to be pored at the base of the fin. Dissection of adult specimens does not reveal any deeper-lying connecting channels. The presence of a triple lateral line on the caudal fin in Lates has not been recorded before, and to the best of my knowledge has not been described in any other percoid species. Since it is clearly a derived condition it is a useful indicator of the mono- phyletic origin of these species. The posterior extremity of the lateral line in Luciolates is also triradiate, but here the three branches are interconnected by pore-bearing scales (Fig. 28b). The median row extends onto the caudal fin, but the line of pored scales is interrupted by the presence of poreless ones, and it never extends to the margin of the fin. The upper and lower lines do not extend for more than one or two scales beyond the limits of the body scales. However, in a few specimens an occasional pored scale is found some distance onto the fin membrane and in the same line as a basal branch. Although the condition of the caudal lateral line in Luciolates does differ from that in Lates it is still a triradiate one and the two taxa can reasonably be thought to FIG. 28. Caudal fin, showing : (a) lateral line pore scales in three rows (drawn from Lates niloticus, but typical for all species except L. stappersi), (b) L. stappersi showing 'trident' arrangement of lateral line pore scales at body -caudal fin junction ; note that pores do not continue onto membrane of fin. REVIEW OF CENTROPOMIDAE 49 share a derived character. It is difficult to tell from the evidence available whether the Luciolates condition should be considered a further derivative - albeit a reduc- tional one - of the Lates type, or whether it represents an early stage in the evolution of the Lates type. THE INTERRELATIONSHIPS OF SPECIES WITHIN THE GENUS LATES, AND THE TAXONOMIC STATUS OF LUCIOLATES BLGR. An analysis of the osteological and other anatomical features described in the previous sections shows that all seven Lates species share two derived characters, viz. (i) three rows of pored lateral line scales on the caudal fin and (ii) the ventral (i.e. horizontal) arm of the preoperculum has three or more large serrae. There are three other derived characters (the swimbladder-posttemporal ligament, the anterior extension of the supraoccipital, and the presence of two epurals in the caudal fin skeleton), but as these are shared with Psammoperca (see below, p. 61) they are of no value in establishing the monophyletic origin of the genus Lates on the basis of synapomorph characters occurring within its constituent species. Since, however, the first two apomorph features noted above do not occur in any other members of the Centropomidae except Lates species, they argue strongly for the monophyly of the genus. It is possible to subdivide the genus Lates by grouping together three species sharing one clear-cut apomorphy and at least four apomorph trends. Such a subdivision would bring together L. angustifrons , L, mariae and L. microlepis, species with an elongate ethmovomerine region in which the posterior face of the lateral ethmoid slopes backwards at a pronounced angle and the dorsolateral aspects of that bone slope sharply downwards ; this characteristic appearance of the snout region is clearly seen in Figs 5, 6 & 7. The apomorph trends shared by these species are an elongation of the caudal and posterior abdominal vertebrae (most marked in L. mariae and L. microlepis ; see p. 43), a division of the dorsal fin into two separate parts (reduced interconnecting membrane in L. microlepis, actual separation of the fins in L. mariae ; see p. 45), reduction of the pterosphenoid pedicle and internal jugular bridge (slight reduction in L. angustifrons, progressively greater reduction in L. microlepis and L. mariae ; see pp. 20-27) and, lastly, an elongation and narrowing of the entire skull (a trend not necessarily correlated with the former which is also manifest in species with broad skulls, e.g. L. macrophthalmus ; see pp. 17-19). Finally, and no doubt of significance, it may be noted that the three species are all endemic to Lake Tanganyika. In view of these characteristics, especially the changes in lateral ethmoid mor- phology, it would seem phyletically proper to recognize the species as more closely related to one another than to any other Lates species still extant. This topic will be taken up again later (p. 51). It is difficult to establish any well-founded scheme of interrelationships for the remaining species, L. calcarifer, L. niloticus, L. macrophthalmus and L. longispinis. Part of this difficulty stems from the problematical relationships of L. longispinis and L. macrophthalmus, as was discussed above, p. 13. These two species alone in the 50 P. H. GREENWOOD group show and share definite apomorph characters* (enlarged eyes and long dorsal fin spines ; for a discussion of the reduced pterosphenoid pedicle see p. 25). All four species otherwise exhibit only the synapomorph features of the genus, and are distinguished from each other by slight meristic and morphometric differences. The taxonomic status of Luciolates has never been reviewed critically since Boulenger (1914) first differentiated the genus from Lates on the grounds of its having ' . . . corps plus allonge, nageoires dorsales largement separees 1'une de 1'autre, et ventrales inserees en arriere de la base des pectorals'. It will be recalled (p. 47) that the condition of the dorsal fin in Luciolates represents a slight exaggeration of that existing in L. mariae. In turn, the L. mariae fin condition is a development of that in L. microlepis which is a further slight deviation from the condition found in the basic L. calcarifer-L. niloticus type. In other words, the apparently characteristic dorsal fin of Luciolates is in fact linked by intermediates with that of the most generalized Lates species. Amongst the various Lates species similar intermediate character states can be found for most of the features which, at first sight, might seem to distinguish Lucio- lates from a generalized Lates species. As examples of these 'distinguishing' features one can cite the relative elongation of the vertebral centra, the protraction of the snout (especially the ethmovomerine skull region) and the general elongation and narrowing of the neurocranium. But, all are features shared with the Lates species of Lake Tanganyika, especially the peculiarly shaped ethmoid (cf. Figs 8, with 5-7). Even the supposedly distinctive position of the pelvic fins in Luciolates is closely approached by L. mariae. There are, of course, certain characters in which Luciolates does differ trenchantly from all Lates species, and these features must be given particular attention. No Lates species has enlarged caniniform teeth such as occur, in small numbers, near the symphysis of the upper jaw in Luciolates (see p. 35), none shows such a high degree of hypural fusion (see p. 44), and Luciolates is unique in having the three caudal extensions of the lateral line restricted to the proximal part of the fin and visibly interconnected with each other. One may, I think, rate the dentition and fused hypural plates of Luciolates as derived characters. The condition of the lateral line may be primitive or it could be a secondary reduction of the Lates type (i.e. a derived character), although the interconnection of the lines might argue against such a conclusion. But, even if all these character states are derived ones, they are autapomorphies ; on the basis of synapomorphies Luciolates still has as its nearest relatives the three Lates species of Lake Tanganyika. Furthermore, Luciolates shares with these species one apomorph character (the morphology of the lateral ethmoid) which distinguishes the group as a whole from all other African species of Lates, as well as from the Indo-Pacific marine species L. calcarifer. For these reasons I propose that Luciolates should be united with its nearest rela- tives in the genus Lates. At the same time I propose placing the Lake Tanganyika * On the evidence currently available, L. macrophtkalmus (from Lake Albert) and L. longispinis (from Lake Rudolf) could either be sister taxa derived from a common ancestor (itself a sister species of L. niloticus) or each could have been derived locally, in late Pleistocene times, from the population of L. niloticus then inhabiting these lake basins. REVIEW OF CENTROPOMIDAE 51 Lates species, that is L. angustifrons , L. microlepis, L. mariae and now Lates stappersi in one subgenus (for which the name Luciolates Blgr. is available), separate from L. calcarifer, L. niloticus, L. macrophthalmus and L. longispinis which species comprise the subgenus Lates. Definitions and synonymies for these taxa are given on pp. 77-78. Interrelationships within the subgenus Luciolates may be delimited on the basis of vertebral morphometry, the division of the dorsal fin, the morphology of the lateral ethmoid, and on neurocranial anatomy and morphology (see relevant sections on pp. 14-45). Lates angustifrons is clearly the plesiomorph sister species of all others in the subgenus. Lates mariae and L. microlepis show generally similar degrees of speciali- zation in all the characters noted above, and can thus be considered sister species ; since in some features (e.g. the lateral ethmoid) L. microlepis is less specialized than L. mariae it can be considered the plesiomorph member of the pair. The greatest level of specialization is seen in Lates stappersi which is therefore ranked as the apomorph sister species of L. mariae and L. microlepis combined (see Fig. 37). The difficulties of ranking species within the nominate subgenus have been discussed above (see pp. 49-50). Indeed, it is not even possible to show that this subgenus is monophyletic since its 'diagnostic' features are those plesiomorphic for the genus as a whole. A REVIEW OF THE GENUS PSAMMOPERCA RICHARDSON Introduction There are no published accounts of the osteology and anatomy of Psammoperca. A brief outline of the osteology of P. waigiensis (Cuv.) is given here, together with some notes on various aspects of the soft anatomy in this species, particularly those features which have some bearing on the phyletic relationship of the taxon. Fishes of the genus Psammoperca (Richardson, 1844) occur in coastal waters from the Bay of Bengal, the Indo-Australian archipelago, northern Australia, the Philip- pines and the China Sea. To a considerable extent, this distribution overlaps that of Lates calcarifer (see above, p. 12 ; also Fig. 36, and Weber & de Beaufort, 1929). Two nominal species, P. waigiensis (Cuv.) and P. macroptera Gunth. are recognized, the latter restricted to Australia and known only from the holotype. The material I have examined is entirely of P. waigiensis, but the individual variability rep- resented in these samples certainly indicates that P. macroptera should be con- sidered a synonym. The question could be solved if large samples from the type locality and other regions of Australia were examined. Superficially, P. waigiensis is much like L. calcarifer (Fig. i), but is distinguished by its widely separated nostrils, smooth lower border to the preoperculum and to the lachrymal, and by the more extensive squamation of the dorsal and anal fins. 52 P. H. GREENWOOD Osteology and anatomy of Psammoperca waigiensis Neurocranium (Figs 2ga-b) The proportions and general appearance of the neurocranium closely resemble those of Lates macrophthalmus (cf. Figs 29 & 4b) ; that is to say, a member of the subgenus Lates in which there is a reduction in the length of the precommissural neurocranium without elongation of the ethmoid skull region. The ethmovomerine region is exactly like that in members of the subgenus Lates ; the posterior wall of the lateral ethmoid is slightly concave and rises steeply to meet LATE FR SPO BOC 10mm PAR PTF EPI SOCS SPO PTO FIG. 29. Psammoperca waigiensis. Neurocranium in : (a) left lateral view, (b) dorsal view. REVIEW OF CENTROPOMIDAE 53 the frontal, and there are two palatine and one lachrymal articulatory facets on this face. A slight intergeneric difference lies in the strongly concave posterior margin of the vomer, which gives the tooth patch in Psammoperca a distinct arrowhead outline in ventral view. The precommissural region (see p. 20) of the braincase in Psammoperca differs in certain details from that in L. macrophthalmus. The pterosphenoid is about the same relative size and the ascending limb of the parasphenoid meets the prootic to create a generally similar appearance for this region of the skull. However, in Psammoperca there is no trace of a pterosphenoid pedicle and there is no bridge, not even a ligamentous one, across the internal jugular vein and its associated nerves (see pp. 20-26 above). In this respect the skull of Psammoperca resembles, most closely, that of Lates (Luciolates) stappersi. The otic skull region in Psammoperca is like that in Lates (Lates} macrophthalmus as far as the relative sizes and relationships of the constituent bones are concerned, but the posterior half of the prootic is noticeably inflated and is thinner in Psammo- perca. The posttemporal fossa, like that in all extant Lates species, is large and deep, and does not have a complete bony floor. The exoccipital facets meet medially. The autosphenotic does not extend far into the orbit medially or dorsolaterally ; again the resemblance is more to L. macrophthalmus than to other members of the subgenus Lates, and there is some resemblance to species of the subgenus Luciolates. The dorsocranium is, in all respects save one, like that in L. macrophthalmus, with the supraoccipital extending forwards to separate the frontals, high frontoparietal crests, deep excavations between these crests and the supraoccipital, and a clearly demarcated lateral shelf on the supraoccipital where the crest extends posteriorly beyond the epioccipitals. The one difference I can detect is the absence of a bone- enclosed supraorbital transverse commissure in Psammoperca. The cephalic lateral line system in other respects, however, is like that in Lates. The parasphenoid resembles closely that in Lates but is more sharply angled upwards from the level of the ascending limb ; in this respect Psammoperca resembles species of the subgenus Luciolates. Hyopalatine arch and the preoperculum (Fig. 30) Again, it is only in certain details that the hyopalatine arch of Psammoperca differs from that arch in Lates. Psammoperca has no tooth patch on the ectopterygoid, and the dermopalatine tooth patch is very narrow. According to Weber & de Beaufort (1929), ectoptery- goid teeth are present in P. waigiensis but I have been unable to detect any on the specimens I have examined. Ectopterygoid teeth are absent in some specimens of Lates (Luciolates} stappersi, and it is interesting to recall that the dermopalatine tooth patch is narrowed to an extent comparable with that in Psammoperca. The palatine in P. waigiensis has a distinct dorsal ridge on the autopalatine immediately in front of the facet for articulation with the lateral ethmoid. This ridge is absent in all members of the subgenus Luciolates and is but weakly developed in species of the nominate subgenus. 54 P. H. GREENWOOD ENT .PAL POP. FIG. 30. Psammoperca waigiensis. Hyopalatine arch, including preoperculum (right side), viewed laterally. As in Lates, the vertical limb of the preoperculum has a finely serrated posterior border, and the mandibular-preopercular sensory canal is bone enclosed (but opening through three ventrally directed and elongate pores on the horizontal arm) . The margin of the ventral limb, however, is entirely smooth except for a stout, posteriorly directed spine at the angle between the vertical and horizontal limbs (Fig. 30) . A similar spine is, of course, present in all Lates species but Psammoperca lacks the three or four stout and ventrally orientated spines on the horizontal limb. Lates (Luciolates} stappersi, it will be recalled (p. 30), often shows some reduction in the size of these spines, but in no individual are they entirely wanting. Circumorbital bones (Fig. 31) The five circumorbital bones are very similar to those in Lates ; the relative elongation of the lachrymal and of the fifth circumorbital in Psammoperca is more like that seen in members of the subgenus Luciolates. The lateral line canal is bone enclosed but opens to the exterior through five pores on the lachrymal, a pore between each articulation of the individual bones, and a ventral pore on the third circumorbital bone. The suborbital shelf (on the third bone) is well developed to an extent almost equalling that found in members of the subgenus Lates ; it extends dorsally to about the upper end of the fourth circumorbital bone. The most marked difference between the circumorbital series in Psammoperca and Lates lies in the completely smooth ventral margin to the lachrymal and second circumorbital bones. These bones are strongly serrated in all Lates species, except L. stappersi, but even in that species some definite trace of the serrations does remain on the posterior part of the lachrymal (see p. 32 and Fig. i3c-d). REVIEW OF CENTROPOMIDAE 55 Opercular series (Fig. 140) The operculum of Psammoperca, like that in Lates, is armed with a single stout spine developed at the posterior end of the stay running from the articular facet for the hyomandibular boss. Indeed, the entire opercular series is like that of Lates, the relatively elongate interoperculum having the proportions of that bone in L. (Lucio- lates) mariae, L. (Luciolates) microlepis and L. (Luciolates} stappersi rather than that in other species of the genus. As in Lates, the sub- and interopercular bones of Psammoperca are thin. Jaws (Figs 32a-c) The maxilla, supramaxilla and premaxilla are, except for the coarser teeth on the latter bone, identical with those elements in species of the subgenus Lates. The bones of the lower jaw (dentary, anguloarticular and retroarticular) are also like their counterparts in members of that subgenus ; again, the teeth are stouter than in Lates. Branchial skeleton In its basic morphology and in the details of its upper pharyngeal dentition the gill arch skeleton of Psammoperca is identical with that of Lates niloticus (see p. 35). The only difference I can detect from the one Psammoperca skeleton studied is that the regularly arranged, small, rectangular tooth plates lying laterally on the gill arch above the filaments (the supralamellar plates, see p. 37) are restricted to the outer side of the first four gill arches (in Lates plates are present on both aspects of an arch). This reduction in tooth plates should be considered as a derived condition since a marked reduction or even the complete loss of free dermal tooth plates is a feature of the more specialized percomorph groups. SOS LAC FIG. 31. Psammoperca waigiensis. Circumorbital bones (right) in : (a) lateral view, (b) viewed dorsally and somewhat anteriorly. P. H. GREENWOOD PMAXP ASCP SMX DPROC PMX P AA 10mm FIG. 32. Psammoperca waigiensis. (a) Premaxilla (right) in lateral view, (b) Maxilla (right) in a slightly oblique dorsal view to show supramaxilla. (c) Dentary (right), with anguloarticular and retroarticular, in lateral view. Hyoid arch skeleton (Fig. 33) The only marked difference between the hyoid skeletons of Psammoperca and Lates (especially members of the subgenus Lates} is the presence in the former of a moderately large, ovoid tooth-patch firmly attached to the broadly spatulate basihyal. Psammoperca has seven branchiostegals, the posterior two of which articulate laterally with the epihyal, the next two with the ventrolateral face of the ceratohyal, and the first three with the ventral margin of that bone. The presence of a basihyal tooth plate must be considered a plesiomorph character for the genus, the only living member of the Centropomidae in which it has persisted. Pectoral girdle and associated bones (Fig. 34) The one obvious difference between the pectoral girdles (i.e. supracleithrum, cleithrum, scapula and coracoid) of Psammoperca and Lates is the absence of serra- tions on the posterolateral angle of the cleithrum. In all other respects the girdles REVIEW OF CENTROPOMIDAE 57 2 mm FIG. 33. Psammoperca waigiensis. Dorsal view of basihyal, showing tooth plate. in the two genera are similar, but with a greater resemblance in overall proportions between the girdle of Psammoperca and that in the subgenus Lates. Although in Psammoperca there are no serrations at the posterior cleithral angle, the bone in that region is drawn out into a short but well-demarcated spine. As in Lates, the three upper fin radials articulate with the scapula, and the lowest with the coracoid. There are two postcleithra, but in Psammoperca the upper member of the pair is less expansive than in Lates. The posttemporal and extrascapula are similar in both genera, the posttemporal in Psammoperca even having the same kind of pit for the reception of the swimbladder ligament (see p. 41), but it does lack serrations on its hind margin. SC PFr COR 10mm FIG. 34. Psammoperca waigiensis. Pectoral girdle (right half) with supracleithrum in situ and post-cleithra displaced posterodorsally. Lateral view. 58 P. H. GREENWOOD Vertebral column The total vertebral count in P. waigiensis is 25, comprising n abdominal verte- brae, 13 caudal, and the fused first preural and ural elements of the caudal fin skeleton. There are nine pairs of pleural ribs, the first pair carried on the third vertebra, the last pair on the eleventh abdominal vertebra ; this latter pair of ribs, instead of sloping gently backwards parallel with the preceding pair (as in Lates), runs almost horizontally and generally overlaps the proximal tip of the first anal pterygiophore. The first definite parapophyses appear on the seventh vertebra, and are but a little shorter than those on the succeeding centra, although there is a slight and posteriorly progressive elongation of these processes. Anterior to the seventh vertebra, the ribs articulate with a shallow pit on the centrum. Where parapo- physes are present, the rib articulates with the posterior face of the process. In all these features, except for the better developed first and second parapophyses, and the angling of the last pair of ribs, Psammoperca is like Lates (see p. 42). Epineural ribs are present on the first three vertebrae, and epipleural ribs on at least the first four pleural ribs. (These data were obtained from radiographs.) The first three vertebrae are shown in Fig. 25b ; their close resemblance to those in Lates is obvious (cf. Fig. 25a). One slight difference is in the development of a low median ridge on the ventral face of the second centrum of Psammoperca. As in Lates, the neural spine of the second vertebra is much broader than the spine of the first and third centra, has its anterior and posterior margins parallel over much of their lengths, and tapers rather abruptly to form a slightly hooked tip. The angle between the posterior face of the second spine and the anterior face of the third spine is from 20° to 25 °. Except in the first four vertebrae, all centra are a little longer than deep, the relative length of the centrum increasing somewhat in the posterior abdominal vertebrae, which have about the same proportions as the caudal vertebrae. In this respect the centra in Psammoperca are rather more like those in Lates (Luciolates) angustifrons than in other species of that subgenus or in species of the nominate subgenus. There are three predorsal bones, the first lying immediately anterior to the first neural spine, the second and third situated immediately before and behind the tip of the second neural spine. Caudal fin skeleton (Fig. 35) The caudal skeleton in Psammoperca differs from that in Lates in one important respect, namely the presence of a single uroneural (see p. 44). Otherwise there is great intergeneric similarity in this structure (viz. 2 epurals, 5 hypurals, 1,8 + 7,1 principal fin rays and a low neural crest on the second preural vertebra [lower, in fact, than in Lates]). There is, as far as can be detected from radiographs, probably no fusion between any of the hypurals, although in one fish (240 mm S.L.) of the eight examined, hypurals 3 and 4 are so closely apposed as to appear fused. The first and fifth hypurals are autogenous, the others are fused to the underlying vertebral support. REVIEW OF CENTROPOMIDAE 59 UR NaPU2 PH HsPU2 5mm FIG. 35. Psammoperca waigiensis. Caudal fin skeleton (drawn from specimen 1872.9.2:10-11). The posterior margin of the caudal fin is rounded. A single uroneural must be considered a derived feature, and in this respect the caudal fin skeleton in Psammoperca is, relative to that in Lates, more specialized. Indeed, since there are in Psammoperca two and not three epurals the caudal fin skeleton is more specialized than that in any member of the Serranidae (where there are, invariably, three epurals - but one uroneural - in the fin skeleton ; see Gosline, 1966). Dorsal, anal and pelvic fins There are 19 pterygiophores (proximal radials) in the dorsal fin skeleton, each one except the last supporting a single fin ray. Although an occasional member of the subgenus Lates may have 19 pterygiophores (see p. 45), the usual number in that taxon is 18. Nineteen, however, is the modal number of pterygiophores (20 the unusual one) in species of the subgenus Luciolates. Unlike Lates, Psammoperca has some of the dorsal fin spines (the fifth through the eighth) associated with discrete medial radials ; a medial radial is also associated with the last branched ray in this fin. Lates (Luciolates} stappersi alone amongst the Lates species has medial radials (associated with the posterior four or five branched rays). 60 P. H. GREENWOOD The dorsal fin is deeply indented to form an anterior part with seven spines, and a posterior portion with one spine and 12 branched rays. A continuous but low membrane connects the two parts of the fin. The spacing between the spines of the two fin divisions is fairly even (cf. Lates species, p. 45). The anal fin skeleton comprises nine pterygiophores, of which the first is a double structure and carries two spines. All other pterygiophores, except the last, carry a single spine or ray. (Total fin ray count 111,8.) Medial radials are absent from all pterygiophores except the last. (In Lates species medial radials are present on the last three or four pterygiophores, see p. 47.) The first of the nine anal pterygiophores, like that in Lates, is a stout and elongate bone and is in contact proximally with the haemal spine of the first abdominal vertebra. The origin of the pelvic fins lies slightly behind that of the pectoral fins ; in other words the fins have the same positions as in Lates (Luciolates] stappersi. Swimbladder As in Lates, so in Psammoperca there is on each side of the swimbladder anteriorly a tough connective tissue strap extending from the tunica externa to the posttemporal. The position, shape and size of the strap are identical in both genera (as are the modifications to the posttemporal, see pp. 41 and 47). The gross morphology of the swimbladder resembles that in Lates. The tunica externa is thick, and a pair of short blunt processes extends forward on either side of a median invagination of the swimbladder. Psammoperca does differ, however, in having a narrow posterior diverticulum extending outside the visceral cavity. In a single specimen dissected, this caudal swimbladder prolongation lay on the left side of the first anal pterygiophore ; it is embedded in the body musculature of that region and does not penetrate into the haemal arches of any caudal vertebrae. Baudelot's ligament The ligament is well developed and its relationships with the body musculature in the cervical region are like those described for Lates niloticus on p. 47 ; that is, it closely approximates to the serranid type described by Gosline (1966). Lateral line Unlike Lates there is only one series of pore-bearing scales on the caudal fin of Psammoperca. These small scales are an uninterrupted continuation of the body lateral line scales ; they extend almost to the posterior margin of the caudal fin. In one of the nine specimens available, a few widely separated pore scales were found on the fin membrane between a pair of rays on the lower part of the fin. THE RELATIONSHIPS OF PSAMMOPERCA The close overall resemblance between Psammoperca and Lates has long been recognized (Regan, 1913), and has even resulted in a false record of Psammoperca for the Japanese fauna (see Katayama, 1956). REVIEW OF CENTROPOMIDAE 61 .A detailed examination of the characters shared by the two taxa shows that many must be ranked as primitive features (i.e. symplesiomorphies) and therefore of little value in estimating relationships. Included amongst the symplesiomorphies are the vertebral count, the presence of a single opercular spine, similarities in gill arch anatomy and dentition, the single spine carried on the first pterygiophore of the dorsal fin, and many details in syncranial morphology and anatomy. There are, however, four derived characters shared by Psammoperca and Lates which are not present in any other members of the Centropomidae. These syna- pomorphies are : (i) A swimbladder-posttemporal ligament (and correlated modifications to the posttemporal bone) ; see p. 60. (ii) A large spine at the posterior angle of the preoperculum. (iii) Two epurals in the caudal fin skeleton ; see p. 58. (iv) An anteriorly extended supraoccipital which separates the posterior parts of the f rentals. On the basis of these characters it is reasonable to conclude that Lates and Psammo- perca are members of the same lineage, a lineage distinct from that of Centropomus (see below, p. 62). To indicate this relationship I propose placing Psammoperca and Lates together in one subfamily, the Latinae. Members of the genus Centro- pomus would thus constitute a second subfamily, the Centropominae, which can be readily defined on the basis of several specialized characters (see p. 67 below). Psammoperca waigiensis (and, where these features can be checked, also P. macroptera ; see p. 51 above) differs from all or most species of the genus Lates in at least 14 features. In the list that follows, the condition of these features in Lates is given in parentheses. 1 . A single series of lateral line scales on the caudal fin. (Three series.) 2. A single uroneural. (Two uroneurals.) 3. Some spine-bearing dorsal fin pterygiophores with a median radial. (None.) 4. No tooth patch on the ectopterygoid. (Present, but reduced in L. stappersi.) 5. No spines on the ventral (horizontal) limb of the preoperculum. (Three or four spines.) 6. Ventral margin of the first infraorbital bone (lachrymal) smooth. (Serrated, strongly so in most species.) 7. No pterosphenoid pedicle or internal jugular bridge. (Pterosphenoid pedicle present in all species except L. stappersi and L. mariae ; internal jugular bridge present in all species, even if reduced to a ligament.) 8. Transverse commissure of supraorbital lateral line system absent or poorly developed. (Present and well developed.) 9. Dermal tooth patch fused with basihyal. (Absent.) 10. Supralamellar tooth plates (p. 37) present only on the outer side of each gill arch. (Pre- sent on both sides.) 11. Posterior margin of the posttemporal smooth. (Serrated.) 12. A single short spine at the posterior angle of the cleithrum. (One large and two smaller spines.) 13. Posterior extravisceral extension of the swimbladder. (None.) 14. Second dorsal and anal fin entirely covered by small but densely arranged scales. (Squamation restricted to about the proximal two-thirds of the fin.) In some of these characters (e.g. i, 3, 5 and 9) Psammoperca is more primitive than any Lates species ; in others (2, 4, 7, 10 and 13) it shows derived characters. The 62 P. H. GREENWOOD status of characters 6, n, 12 and 14 is at present indeterminable. (See discussion on pp. 30-32.) It is on the basis of the unique derived characters (i.e. autapomorphies) found in each of the two taxa that I would maintain them as distinct genera, the implication being that Psammoperca split off from the common latine lineage before the evolution of a serrated preoperculum and the tripartite lateral line extension onto the caudal fin. The derived characters seen in Psammoperca (especially the loss of a ptero- sphenoid pedicle, the presence of a single uroneural, the loss of certain branchial arch tooth plates, and the loss of ectopterygoid teeth) must have evolved after this split occurred. In these features Psammoperca is certainly more 'advanced' than is Lates. It is interesting to note that a reduction and ultimate loss of the pterosphenoid pedicle is seen in certain Lates species of the subgenus Luciolates (see pp. 20-27), and that Lates (Luciolates} stappersi also shows a considerable reduction in, and oc- casionally the loss of, ectopterygoid teeth. Furthermore, this species also shows a marked weakening of the serrations on the lachrymal. Similar parallel trends in all three characters are found amongst the species of Centropomus (see below) thus suggesting that this is the manifestation of a potentiality possessed by the common ancestor of all living centropomids. THE RELATIONSHIPS OF CENTROPOMUS WITH THE LATINAE Fraser (1968) has given a good account of the osteology of five species of Centro- pomus but he was unable, through lack of published information, to compare fully these species with members of the genera Lates and Psammoperca. He did, however, list a number of differences between Lates and Centropomus and these will be com- mented upon below. In my comparison of Centropomus and the Latinae I have drawn on Fraser's (op. cit.) information and supplemented it from dissection, radiographs and dry skeletons of C. parallelus Poey, C. pectinatus Poey, C. armatus* Gill, C. unionensis* Bocourt, C. robalito* Jordan & Gilbert, C. nigrescens* Giinther, C. ensiferus Poey and C. undecimalis (Bloch) ; species not described by Fraser (op. cit.) are marked with an asterisk. The neurocranium in all Centropomus species is narrow and elongate, with a pronounced relative lengthening of the ethmovomerine region. In these respects it resembles the neurocranium of Lates (Luciolates} mariae and L. (Luciolates} stappersi, but it does differ in having only a gently angled parasphenoid (or even a straight one ; cf. C. pectinatus, text-fig. 4, and C. undecimalis, text-fig. 5, in Fraser (1968), with Figs 7 & 8 above), and in having the postotic region of the skull relatively longer. Within the Centropomus species I have examined, there is a trend of neurocranial elongation which closely parallels that found in members of the Lates subgenus Luciolates. Another parallelism with Lates is seen in the reduction of the pterosphenoid pedicle and internal jugular bridge. The pedicle and bridge are best developed in C. ensiferus (see text-fig. 6 in Fraser, 1968) where the condition of the bridge is like that in L. (Luciolates} angustifrons REVIEW OF CENTROPOMIDAE 63 (see p. 24 and Fig. 5a). A noticeable difference, however, is that, in C. ensiferus, there is no ascending arm of the parasphenoid and the bridge is formed by contact between the pterosphenoid and prootic. (The prootic in all Centropomus contributes to the posteroventral margin of the orbit ; in Lates this rarely happens because part of the ascending parasphenoid arm usually rises in front of the prootic. This tongue of parasphenoid is, however, very narrow in the more specialized species of the subgenus Luciolates.) In all other Centropomus species I have examined or which are figured by Fraser (1968), excepting C. undecimalis, the pterosphenoid pedicle is either reduced (e.g. C. pectinatus) or is greatly reduced to a small bony knob (that is, to conditions com- parable with those in very small L. (Lates) niloticus or those in adult L. (Luciolates} microlepis ; see p. 26). In C. undecimalis there is no trace of a pterosphenoid pedicle ; in other words a situation directly comparable with that in L. (Luciolates) stappersi and in Psammoperca waigiensis. Correlated with this reduction in the pedicle, the internal jugular bridge is reduced from a narrow bony strut in C. ensiferus to a ligament in the other species (except C. undecimalis), again paralleling exactly the trend seen in Lates (pp. 21-26). In C. undecimalis even the ligament has disappeared (at least in the specimen of 175 mm S.L. I dissected) ; this, it will be recalled, is the condition also found in Psammoperca (p. 53). Probably as a correlate of the lengthening ethmoid-vomerine skull region, the shaft of the vomer in all Centropomus species is much broader anteriorly and has a closer sutural union with the lateral ethmoid than it does in any latine species. In other details, however, this region of the skull is generally similar in both Centro- pomus and the Latinae. The otic region in Centropomus is bullate, more markedly so in some species than in others, but always more inflated than in any Lates species and rather more so than in Psammoperca. An outstanding inter-subfamilial difference is found in the lateral line system of the dorsicranium. In Lates and in Psammoperca all three major canals are bone enclosed. In Centropomus the canals are in the form of laterally orientated open gutters, with only the posterior part of the supraorbital line completely tubular. The frontal cross-commissure is also open (with the gutter directed medially), as is the entire length of the frontoparietal branch (whose gutter is directed laterally). Fraser (1968) has corrected Regan's (1913) erroneous observation that parietal crests are absent in Centropomus, but as compared with Psammoperca and Lates, the parietal crests, and their counterparts on the frontals, are low and very poorly defined, and do not extend to the posterior margin of each parietal (often being confined to the anterior half of that bone) . The supraoccipital in Centropomus does not extend so far anteriorly as it does in Lates and Psammoperca, its tip barely separating the frontals and only reaching a level with a vertical through the middle or the posterior third of the prootic. There are few noteworthy differences in the hyopalatine arches of Centropomus and the Latinae. As Fraser (1968) noted (pace Regan, 1913), ectopterygoid teeth are present in Centropomus. From Fraser's drawing (op. cit., text-fig, n) one gains 64 P. H. GREENWOOD the impression that a metapterygoidal lamina is present in at least some Centropomus species, but I cannot confirm this from the dry skeletons I have examined. There are several inter-subfamilial differences in the morphology of the pre- operculum. First, the lateral line canal in Centropomus is an open gutter, the opening orientated posteriorly on the vertical limb of the bone and ventrally on its horizontal limb. It is the upper rim of this gutter that has been described as a 'ridge' on the preoperculum in Centropomus species (see Fraser, 1968). In the Latinae, where the canal is bone enclosed and tubular no 'ridge' is, of course, de- tectable. A second and pronounced difference is in the ornamentation of the preoperculum, a difference most obvious when Centropomus is compared with Lates. In Centro- pomus, as in Lates and Psammoperca, the posterior margin of the vertical limb is serrated (less regularly so in Centropomus), but the horizontal limb in that genus has a number of low, rather irregular serrations that are enlarged posteriorly at the angle of the bone. In no Centropomus species is there any indication of the three (or four) large triangular spines that characterize all Lates species ; likewise, no Centropomus has the completely smooth horizontal preopercular arm of Psammoperca. Also, unlike both Lates and Psammoperca, there is no single, stout spine at the pos- terior angle between the two preopercular arms ; instead, in Centropomus there are a variable number of spines, all of which are somewhat larger than those preceding and succeeding them on the arms of the bone, but none is as large nor as distinctive as the single spine of the latines. The operculum in Centropomus lacks a spine at its posterodorsal angle (see p. 55) but otherwise the opercular series shows no marked departure from its counterpart in the Latinae. The open lateral line gutters of the circumorbital bones, the reduced serration of the ventral lachrymal border, and the relatively short fourth and fifth circumorbitals are the most obvious inter-subfamilial differences noted in this region of the skull. Apparently the subocular shelf in Centropomus is like that in Psammoperca and members of the latine subgenus Lates, but I have been unable to check this point in all Centropomus species, and in particular those with narrower and more elongate heads. Apart from some slight proportional differences, the major feature differentiating jaw elements in the two subfamilies is the much shorter ascending process of the premaxilla in Centropomus. In the Latinae the ascending process is at least one and a half times the height of the maxillary process (see p. 34) but in Centropomus the two processes are of equal height (cf. text-fig. 12 in Fraser, 1968, with Figs 15, 16 & 32 above). The basic gill arch morphology and dentition are similar in Centropomus and the Latinae, although the tooth plates associated with the basibranchials are slightly more elongate in at least some members of the Centropominae. The supralamellar tooth plates in most Centropomus species which I have dissected (C. ensiferus, C. parallelus, C. pectinatus, C. undecimalis and C. armatus) show a unique arrangement not found in any member of the Latinae. The plates are present only on the outer aspect of the second to fourth gill arches, and are fused, in pairs, with the bases of REVIEW OF CENTROPOMIDAE 65 the gill rakers on that aspect of the arch ; occasionally a single plate may occur between a pair of gill rakers. An exception is provided by a small (160 mm S.L.) specimen of C. parallelus in which the plates are serially arranged like those in Psammoperca. A slightly larger fish (220 mm S.L.) shows a condition intermediate between that in the smaller specimen and that of the other species (and specimens of C. parallelus) I examined. Possibly this change in plate arrangement is a growth phenomenon. All Centropomus species have 24 vertebrae (including the fused first ural and preural centra), comprising 9 abdominal and 15 caudal elements. (Eraser, 1968, gives a count of 10 + 14, indicating that we differ in our interpretation of which vertebra constitutes the first caudal element ; I identify it as that vertebra with which the first anal pterygiophore articulates.) All members of the Latinae, in contrast, have 25 vertebrae (n abdominal and 14 caudal elements). There are seven or eight pairs of pleural ribs in centropomines (nine in the Latinae) , the number apparently showing some intraspecific variability. The first rib articu- lates directly with the third vertebra. Definite parapophyses are developed on the seventh abdominal centrum but a low process occurs on the sixth vertebra. Anterior to these centra the ribs articulate as in the Latinae, that is, with a pit in the centrum. Also as in the Latinae, the ribs articulate with the posterior face of the parapophysis when these are present. As far as I can tell from radiographs, the shape and proportions of the centra in all Centropomus species are like those in Lates calcarifer. That is, posterior to the third vertebra all the centra are slightly longer than deep, with little difference in pro- portion between abdominal and caudal elements. Apart from the neural spine on the second vertebra the first three vertebrae are like their counterparts in latine fishes. The second vertebra, however, has a very greatly expanded neural spine (see Fig. 25d) into the anterior face of which the entire posterior margin of the first neural spine is fitted. Fraser (1968) has shown that the proportions of the second neural spine change with age in at least some species of Centropomus ; the spine in young fishes resembles that in adult Lates and Psammoperca (see Fraser, op. cit., text-fig. 14, and pp. 454-5 ; and cf. Figs 25a-c above). All Centropomus species have three predorsal bones, the first situated above the tip of the first neural spine, the second at about the middle of the expanded second spine, and the third lying immediately behind that spine. Fraser (1968) states that there are only two predorsals in Lates, but this is not so (see p. 43 above) ; there are, in fact, no intergeneric differences in this feature. A distinct gap separates the two dorsal fins in all Centropomus species ; the size of the gap, however, shows some specific variation. Unlike those Latinae with separate dorsals (members of the subgenus Luciolates ; see p. 45), the centropomines have no isolated spines between the fins. The head of the seventh pterygiophore is drawn out posteriorly so that it effectively underlies the gap between the fins ; the spine which this pterygiophore carries thus becomes the first (and only) spine of the second dorsal fin. There are 16 or 17 dorsal pterygiophores in Centropomus (cf. 18 or 19, rarely 20 in the Latinae), none of which, as far as I can determine, has a separate medial 66 P. H. GREENWOOD radial (see p. 45). The first dorsal pterygiophore, unlike that of the latines, carries two spines ; except for the last dorsal and first anal pterygiophores, all the others carry a single spine or ray. The dorsal fin ray counts in Centropomus are VII or VIII and 1,8 or 9 (cf. VI-VIII and 1,10-13 in all Latinae except Lates (Luciolates} stappersi which has VI + I + I and 1,9 or 10). Thus in the centropomines there has been not only a trend towards separation of the dorsal fins (a trend also apparent in the latines, see p. 46 above) but also a reductional trend in the number of dorsal fin rays, particularly the bran- ched rays. Interestingly, if the two independent and much reduced fin spines are 'removed' from the fin formula of Lates (Luciolates) stappersi, the result - save for an extra branched ray -is the formula of a Centropomus (i.e. VI + I + I and I,io — > VI and I,io). A most characteristic feature of all Centropomus, and one not even approached by any member of the Latinae, is the very strong and long first anal pterygiophore (see Fig. 27b) ; in many species there is also hypertrophy of the second anal spine. Despite the length of this pterygiophore it extends only a little further distally (i.e. towards the vertebral column) than does its counterpart in the Latinae. The greater length of the bone in Centropomus is accommodated by the bone sloping obliquely backwards so that the articulation for the spines lies in a vertical below about the seventh rather than the second or third abdominal vertebra as is the case in Lates (Fig. 2ya) and Psammoperca. The caudal fin skeleton in Centropomus differs from that in the Latinae in either one (Lates) or two (Psammoperca) characters and is of a more primitive kind. Primi- tive features in Centropomus are the three epurals (two in Latinae) and the two uroneurals. Lates also has two uroneurals but only one is present in Psammoperca (the fin skeleton in that genus being the most evolved within the Centropomidae) . All Centropomus species have a deeply forked caudal fin whereas in the Latinae the fin is usually rounded or truncate, although it is weakly forked in some species of the Lates subgenus Luciolates. Like the Latinae, the caudal fin formula of the Centropominae is 1,8 + 7,!. The pectoral girdle and fin skeleton are basically alike in the Centropominae and Latinae except for slight differences in the postcleithral elements. The posttemporal in Centropomus lacks the cavity and associated bullation that characterize this bone in Lates and Psammoperca, a consequence of there being no swimbladder- posttemporal ligament in Centropomus (see below). Otherwise the posttemporal is similar in both subfamilies. The extrascapula in Centropomus is also basically like that in the Latinae, but it is characterized by having the lateral line canals situated in open gutters and not enclosed in bony tubes (see p. 41 above). In those Centropomus species which I have been able to dissect, the anterior end of the swimbladder has no medial invagination (see p. 60 above). However, in some species there are a pair of short horns arising from the dorsolateral aspect of the bladder and extending part way towards the skull ; in none could I find any direct connection between the skull and the horns, and neither could I find any trace of a swimbladder -posttemporal ligament such as occurs in all members of the Latinae. The development of the swimbladder horns seems to be restricted to certain species. REVIEW OF CENTROPOMIDAE 67 When present these appendages may be short, simple and anteriorly directed, or, as in C. imdecimalis, they may be elongate and curved backwards to lie laterally along the swimbladder (see Meek & Hildebrand, 1925). This latter condition is reminiscent of that found in certain species of Sciaenidae (a family in which there is also an extension of the lateral line onto and reaching the margin of the caudal fin ; the possibility of there being some phyletic relationship between sciaenids and centropomids is under review). Baudelot's ligament is present in Centropomus and is moderately well developed. The relationships between this ligament and the body muscles are like those in the Latinae (see p. 60 above), with little or no muscle passing medially below the liga- ment, but with a broad band passing underneath it laterally to insert on the basi- occipital and exoccipital. A single extension of the body lateral line scale series onto the caudal fin is found in all species of Centropomus (there are three extensions in Lates but only one in Psammoperca). In Centropomus, as in the Latinae, the caudal extension of the lateral line is continuous to the margin of the fin or almost so. When all these characters are taken into account, it is clear that the Centro- pominae (i.e. Centropomus species) differ from the Latinae (Lates and Psammoperca species) in a number of features. Some of these differences involve the retention of characters primitive for the family whilst others represent the development of unique specializations shared only by Centropomus species. In the former (i.e. plesiomorph) category may be listed the caudal fin skeleton, the short supraoccipital bone, the single lateral line extension onto the caudal fin, and the absence of a swimbladder -posttemporal ligament. The autapomorphic features of the Centro- pominae are more numerous and include the open cephalic lateral line canals, the separation of the dorsal fin, the hypertrophy of the first anal pterygiophore (and at least relative hypertrophy of the second anal spine), the absence of medial radials throughout the dorsal and anal fins, the development of a curved and posteriorly protracted head on the seventh pterygiophore of the dorsal fin (see p. 65), the development in most species of anterior horns on the swimbladder, the incorporation of the supralamellar tooth plates into the gill rakers, and the elongation of the skull, especially its ethmovomerine region (with which feature may be correlated changes in the shape of the ethmoid and vomer). There are other differences, like the absence of an opercular spine, the forked caudal fin, and the markedly reduced squamation of the dorsal, caudal and anal fins, whose apo- or plesiomorph status is uncertain. On the basis of those characters that are clearly synapomorphic the Centropomus species can be recognized as a monophyletic group and one which, although sharing a common ancestry with the Latinae, is clearly distinct from that lineage. It is for this reason that I propose giving the Centropomus species-group coordinate ranking (as the subfamily Centropominae) with the Latinae (see also above p. 61 ; and p. 75 below for diagnoses). When the mosaics of apo- and plesiomorph characters within the two subfamilies are compared it becomes impossible to decide which taxon should be considered the plesiomorph sister group of the other. However, it does seem that what we are now 68 P. H. GREENWOOD observing is the product of vicariant differentiation from a once widespread basic centropomid taxon, a differentiation that produced the Centropominae in America, and the Latinae in Asia and the Mediterranean region (including Africa), leaving each group with its own association of primitive and derived features. Fraser (1968) noted certain shared characters amongst the various species of Centropomus, and from their pattern of occurrence concluded that three phyletic lineages are represented amongst the living species. Unfortunately, Fraser does not give a really critical analysis of the characters on which his phylogeny is constructed and it is thus impossible to test the supposed interrelationships of the three lineages he hypothesizes. In particular it would seem that his monotypic lineage comprising C. pectinatus is more likely a member of the C. ensiferus - C. robalito lineage, and not, as is expressed in Fraser 's diagram, one distinct from the other two lineages and occupying an equal phyletic relationship with both of them (see Fraser, 1968, text-fig. 9). Although Fraser's analysis is not documented in terms of synapomorph and symplesiomorph characters it obviously shows that similar trends can be found within the Centropominae and the Latinae. This aspect is particularly well demon- strated in the neurocranial morphology and in the reduction of the pterosphenoid pedicle and internal jugular bridge. There is also inter-subfamilial similarity in the trend towards greater separation of the two parts of the dorsal fin. In this trend the Latinae appear not to have evolved much beyond the early phases, whereas the centropomines have carried the trend further and no longer preserve traces of its earlier stages within their numbers. FOSSIL CENTROPOMIDAE Apart from identifications based solely on otoliths,* all species of fossil centro- pomids so far discovered are currently referred either to Lates or to Eolates Sorbini (see Sorbini, 1973), that is, to the subfamily Latinae. The time range of these fossil taxa extends from the Eocene to the Holocene, and their geographical range from the Paris Basin, through Austria, Portugal, northern Italy and Croatia to Egypt, the Sahara and eastern Africa (Sorbini, 1973 ; Green- wood, 1974 ; Greenwood & Howes, 1975). With the exception of some material from Europe (Sorbini, 1973) the majority of fossils are from Africa and are in the form of disarticulated and damaged bones. The problems of specific (or, indeed, generic) identification when dealing with material of this nature need not be stressed. In most instances the fossil bones have been compared with their counterparts in Lates calcarifer and L. niloticus. If the fossils are from Africa, and the bones are not noticeably different from their counter- parts in L. niloticus the material was either referred to that species or, and probably more accurately since diagnostic features are rarely preserved, merely to Lates sp. * Psammoperca sheppeyensis Frost 1934, Centropomus superpendens Frost, 1934 an(l C. excavatus Stinton, 1966, all from the London Clay (Eocene), are species described from otoliths only. Since so little is known about otolith morphology in living centropomids and because no skeletal material is available for the species, these records can at present contribute little to our understanding of centro- pomid phylogeny and biogeography. REVIEW OF CENTROPOMIDAE 69 When obvious morphological differences could be detected the remains have been taken to represent different species (e.g. L. fajumensis Weiler, 1929 ; L. karungae Greenwood, 1951 ; L. rhachirhinchus Greenwood & Howes, 1975). Because these species are based on fragmentary, disarticulated bones it is impos- sible to determine their phyletic relationships with each other or with the extant species of Latinae (see discussions on L. rhachirhinchus in Greenwood & Howes, 1975). About all that can be said with any certainty is that latine centropomids had, by late Miocene times, a distribution that included Egypt, Tunisia and eastern Africa (Lakes Victoria and Albert regions) and that at least one species, L. rhachi- rhinchus, showed several derived characters even when compared with extant species of that genus (Greenwood & Howes, 1975). All these remarks are, of course, based on the assumption that the taxa are correctly placed in the genus Lates ; in no case is it possible to check on the autapomorph characters used here to define the genus (see p. 77), the generic identity being based on an overall similarity between the fossil bones and their counterparts in extant Lates species. The situation is little better for the three European species in which the entire skeleton is preserved, viz. L. partschii Heckel, 1855 (Miocene, Vienna Basin) ; L. croaticus Kramberger, 1902 (Miocene, Croatia) and L. macropterus Bassani, 1899 (Oligocene of Vicenza). I have not been able to examine any material of L. croaticus, and the only pub- lished description and figures of this species are inadequate for critical interpretation, although Kramberger (1902) does give a vertebral count of 27, that is two more vertebrae than in any other member of the Latinae for which the count is available. The status and relationships of this nominal species must therefore remain incertae sedis. Sorbini (1973) has re-examined L. macropterus, but was unable to draw any definite conclusions about its relationships. Again it is impossible to check on any diagnostic characters of phyletic importance. Lates partschii (Miocene of Vienna) has been thoroughly redescribed by Sorbini (1973), who also published a photograph of the holotype, and a close-up picture of its caudal skeleton. But once again certain critical details are either not preserved, are obscured, or are damaged. For instance, there seems to be only a single and median row of lateral line scales on the caudal fin but one cannot be certain that dorsal and ventral scale rows were not present. There are certainly only two epurals in the caudal fin skeleton, and there are, apparently, two uroneurals, both features which are characteristic of Lates (see pp. 44 and 77). From this and other circumstantial evidence given in Sorbini's account, it seems likely that partschii can be placed in Lates, but it is impossible to determine its relationships with any extant species of that genus. Fortunately, many important features are preserved in the extensive material of Eolates that is available for study (Sorbini, 1973 ; personal observations on specimens in the collections of the British Museum (Natural History))*. * Two nominal species are recognized, Eolates gracilis (Agassiz) 1833, and E. macrurus (Agassiz) 1833. According to Sorbini (1973), E. macrurus may yet prove to be a synonym of E. gracilis. For this reason, and because the osteology of E. gracilis is much better known, only that species is taken into account in the discussions that follow. 7o P. H. GREENWOOD Eolates gracilis is distinguished from all Lates species by at least one character complex (the caudal fin skeleton), and probably by two other characters as well (the absence of upper and lower lateral line scale rows on the caudal fin, and the disposition of the branchiostegal rays). I have examined eight specimens of E. gracilis (from the BMNH collections) in which the caudal fin is well preserved. In all, the median lateral line scale row is clearly developed and it is also possible to see other scales, often still in their rows, on other parts of the fin. None of these other scales is perforated and I am confident that only one lateral line scale row (the median one) is preserved. My colleague, Dr K. Banister, has recently examined E. gracilis holotype in the Paris Museum and reports that only a median row can be detected in this specimen as well. The caudal fin skeleton in E. gracilis (Fig. 26b) shows a well-developed neural spine on the second preural centrum (spine greatly reduced in Lates), three epurals (two in Lates) and two uroneurals (two uroneurals also present in Lates). In other words, the caudal fin skeleton is of a more primitive type than that in Lates. (It will be recalled that Centropomus also has three epurals, but the second preural arch and spine are reduced and resemble those in Lates.) A possible third intergeneric difference concerns the number and disposition of the branchiostegal rays, but this requires confirmation since it is based on data available from only one of the E. gracilis specimens examined by Sorbini (1973). In the sole specimen from which a branchiostegal ray count could be made Sorbini (op. cit.) records, with some uncertainty, a total of eight rays (seven in Lates and other centropomids). Judging from the photograph of this specimen (Sorbini, 1973, Plate IV, fig. i), I should doubt that the fragment at the anterior end of the ceratohyal is indeed part of a branchiostegal ray. There is, however, no doubt that in this specimen all the branchiostegal rays are associated with the ceratohyal. According to McAllister (1968) this condition is not found in any living percoid fish ; there is always at least a half articulation between a ray and the epihyal. The rays in the E. gracilis specimen are in no way disarranged, and the posterior one is well forward of the epi-ceratohyal junction. Clearly no decision can be made on the validity of this apparent intergeneric differ- ence (or its apparent uniqueness amongst percoids) until further specimens can be examined. Like the preoperculum in Lates this bone in Eolates has three large ventral spines on its horizontal limb, and an enlarged spine at the posterior angle of the bone. Also as in Lates, there is a single, large spine on the posterodorsal margin of the operculum in Eolates. Ornamentation of the cleithrum and on the first infraorbital bone (lachrymal) is similar in Eolates and Lates, but the phylogenetic importance of these latter characters is probably not great. Regrettably, little detailed information can be obtained about the morphology of the neurocranium in Eolates. Sorbini (1973) gives no description of the posterior orbital region of the skull, presumably because in his material, as in that of the BMNH, this area of the head is either crushed or obscured by other bones overlying it. Thus it is impossible to determine what type of pterosphenoid pedicle and inter- nal jugular bridge is present. REVIEW OF CENTROPOMIDAE 71 The ethmoid region is generally well preserved, and resembles that found in members of the subgenus Lates (see p. 19 above). In a few E. gracilis specimens the posttemporal is well preserved ; it seems to show the slightly bullate outer surface that, in living centropomids, is associated with the insertion point of a swimbladder- posttemporal ligament (see p. 41 above), a derived feature characterizing members of the subfamily Latinae (see above, p. 66). In brief, Eolates (as represented by E. gracilis) is clearly a member of the sub- family Latinae and shares at least one derived character (the ventral preopercular spines) with the genus Lates (see p. 31). Eolates differs from Lates in having only one series of lateral line scales on the caudal fin (presumed in this context to be a primitive feature, see p. 48 above), and in having a caudal fin skeleton that is primitive in relation to this skeleton in Lates (see above, p. 66). A third intergeneric difference is in the less deeply indented dorsal fin of Eolates, a feature with which may be cor- related the equal spacing between the 'last' (i.e. shortest) spine of the anterior part of the fin and the longer 'first' spine of the fin's posterior half. This character too should be considered a plesiomorph one because a deeply divided fin is a basal condition in the centropomid trend leading towards completely separate first and second dorsal fins (see above, pp. 46 and 65). All the features discussed so far indicate that Eolates should be considered more primitive than Lates. In phyletic terms it should be ranked as the plesiomorph sister group of that taxon. The relationship of Eolates within the subfamily Latinae is, therefore, best indicated by uniting Eolates with Lates in a single tribe (Latini, new tribe) which would then become the sister taxon of the tribe containing only the genus Psammoperca (tribe Psammopercini nov.). Sorbini (1973) also recognizes the affinity of Lates and Eolates, but he would regard the relationship as an ancestor -descendant one (op. cit. : 41) rather than that of recent shared common ancestry as is proposed here. Sorbini's claim that ' . . . The living marine species L. calcarifer presents the greatest relationship to fossil Tertiary species, which lived in a similar habitat' (Sorbini, 1973 : 41) certainly cannot be substantiated by the meristic and morpho- logical data available from these fossils. For example, as interspecific similarities between L. calcarifer and E. gracilis Sorbini lists (op. cit. : 36) ' . . . disposizione delle vertebre, n. raggi branchiostegi, habitat . . .'. The habitat is similar, but what importance can be attached to this feature in a family with several euryhaline species? The arrangement of the vertebrae in Eolates is like that in several Lates species, while the reference to the number of branchiostegal rays is, I presume, a lapsus for 'spine branchiali'. Eolates has either seven or eight branchiostegal rays (there are seven in all other centropomids ; see above, p. 70), but nine gill rakers (the same number as L. calcarifer}. However, a low gill raker count (8-12) is com- mon to several Lates species, and is apparently the primitive state for the family as a whole. BIOGEOGRAPHY The contemporary world distribution of the Centropomidae (Fig. 36) strongly suggests a Tethyan distribution for the common ancestor of its two subfamilies, the P. H. GREENWOOD FIG. 36. World distribution of extant species of Centropomidae. Stippled areas : Lates species (outside Africa = L. calcarifer). Black spots : Psammoperca species (probably only one, P. waigiensis). Black area ; Centropomus species. Centropominae (America) and the Latinae (Mediterranean, African freshwaters and Asia). The Centropominae, on this hypothesis, have evolved in the tropical New World, probably in estuarine and marine habitats, and the Latinae principally in African freshwater habitats. There is, however, a major dichotomy in the Latinae, between the tribes Latini and Psammopercini, which must have taken place before the Latini invaded Africa. As evidenced by various European fossil species (see above, pp. 68-71, and Sorbini, 1973) and by the wide dispersal of Lates calcarifer (see p. 12), the Latini were and still are successful coastal fishes. The greater diversification of the tribe in African freshwaters can probably be attributed to the greater opportunities for speciation provided by the developing tropical lakes and river systems of later Tertiary and Quaternary Africa. (For a summary of these historical factors see Beadle, 1974.) It will be recalled that there are seven extant and at least three extinct Lates species in Africa, compared with the single extant (L. calcarifer} and three extinct marine or estuarine species (see above, p. 69 ; also Sorbini, 1973 ; Greenwood, 1974 ; Greenwood & Howes, 1975). There are, of course, at least nine species of Centropomus (Centropominae) all of which are essentially marine species (although some freely enter freshwater ; Meek & Hildebrand, 1925). Trans-isthmian isolation could account for four of these species (Eraser, 1968) but there still remain the other five species to contrast with the single marine Lates species (L. calcarifer} of the Indo-Pacific region. The causal factors involved in this aspect of Centropomus speciation are not apparent. It is interesting to compare the morphological radiation undergone by the Centro- pominae and Latinae, and to notice the marked parallelism apparent in the two REVIEW OF CENTROPOMIDAE 73 groups. For example, in both subfamilies there are trends of specialization leading to the reduction and loss of the pterosphenoid pedicle and internal jugular bridge (p. 63), to elongation of the skull through differential lengthening of the ethmoid region (p. 62), to an increase in the number of gill rakers on the first gill arch, to a reduction in the number of supralamellar tooth plates (p. 64), and towards the division and then separation of a primitively continuous dorsal fin (see p. 66). So similar are all the features involved in any one of these trends that one can eliminate any possibility of convergence. The similarities must reflect shared genotypic factors stemming from common ancestry. The absence of Lates (or some related latine fish) from the present-day Medi- terranean Sea may, as Sorbini (1973 : 40) suggests, be due to climatic changes adversely affecting the one or more species that were present in the Mediterranean basin during parts of the Tertiary (Sorbini, op. cit., especially text-fig. 10). In- creasing salinity in the developing Mediterranean may also have had its effect on local populations. During the Eocene and Miocene, species of Lates were also present in Africa (Sorbini, 1973 ; Greenwood, 1974). The Eocene fishes from the Fayum in Egypt may have been estuarine and marine (Weiler, 1929), as may have been the Miocene species from Tunisia (Greenwood, 1973). However, Miocene records of Lates from the equatorial regions of Lake Victoria (Greenwood, 1951) and Lake Albert (Green- wood & Howes, 1975) show that some latine species had adjusted fully to freshwater environments, and that enough time had elapsed since the first invasion for latine species to have reached areas some 3750 km inland from the Mediterranean coast. Like all other fossil Latinae from Africa, the Miocene species are referred to Lates solely on the overall similarity between the preserved fossil bones and their counter- parts in extant Lates species. Such critical features as the nature of the lateral line scales on the caudal fin and the morphology of the posttemporal bone are unknown for any one of them. One Miocene species, L. karungae Greenwood, 1951, from Rusinga Island, Lake Victoria, is represented by only a few vertebrae ; the specific diagnostic features for this taxon relate to the morphology of the third vertebra (Greenwood, 1951). The other taxon, L. rhachirhinchns, from the Lake Albert-Lake Edward region of Zaire is better represented by numerous skeletal parts (Greenwood & Howes, 1975). It differs from all other Lates species in several features, many of which can be considered as derived, and one of which (vertebral proportions) is shared with certain members of the endemic subgenus Luciolates from Lake Tangan- yika (see p. 43 above, and Greenwood & Howes, op. cit.). Even though it is im- possible to identify specifically the Lates remains from the Miocene and Pliocene deposits in North Africa and Egypt, L. rhachirhinchus is morphologically quite distinct from those taxa. Thus, one may conclude from this situation either that more than one taxon invaded Africa or that, by Miocene times, the population of Lates in the Lake Albert -Lake Edward region had undergone marked morphological differentiation, presumably in isolation from its parental stock. The same arguments could be applied to L. karungae although in this instance there is less evidence for the extent to which the morphological differentiation had progressed. 74 P- H. GREENWOOD Sorbini (1973) postulated certain time sequences and migration routes to explain the present-day distribution of Lates species in Africa. Basically the problem Sorbini sets out to explain is the widespread occurrence of one species, L. niloticus, in the Nile, Niger, Zaire, and Senegal river systems, and in Lakes Rudolf, Albert and Chad, in contrast to the occurrence of four endemic species (one supposedly a distinct genus) in Lake Tanganyika. He notes the former occurrence of Lates in other lakes (Edward and Victoria) but is not concerned with the factors that led to these local extinctions, and neither does he take into account the endemic species that coexist with L. niloticus in Lakes Rudolf and Albert. There are two basic tenets in Sorbini's hypothesis, first that the various invasions he postulates originated in Egypt, and second that fossils identified as L. niloticus are indeed representatives of that species. As I have discussed above the latter assumption is not necessarily acceptable, and neither can I find any a priori grounds for postulating repeated and temporally extended invasions from a single area (in this argument, Egypt). That a species of Lates had reached the regions of Lake Victoria and Lake Albert - Lake Edward by Miocene times is not disputed (see above), and Sorbini's argument for the contemporaneous presence of a Lates species in the Lake Tanganyika basin is also acceptable. Why, then, should Lates not have occurred in other Miocene rivers and water bodies, environmental conditions, of course, permitting such colonization? To the best of my knowledge there is no evidence to show that suitable conditions were confined to the regions from which Miocene fossils have been re- covered. Thus I find it difficult to understand why, in order to explain the present distribution of L. niloticus, Sorbini should postulate two invasions, each following different routes, but both originating from Egypt during the Pliocene and continuing through the Pleistocene. Presumably a major reason for putting forward this hypothesis is the fact the fossils identified as L. niloticus are first recorded from the Pliocene of Egypt, thereby implying the origin of that species in Egypt at a later date than the one at which another species (L. karungae] was already present in the Lake Victoria area (and, had he known it, a second species L. rhachirhinchus was present in the area of Lake Albert-Lake Edward ; Greenwood & Howes, 1975). In view of the known distribution for Miocene Lates and because of the uncertain- ties associated with the specific identification of most fossil Lates remains, a simpler hypothesis can be made, viz. : At some stage prior to the late Eocene a species of Lates invaded Africa, possibly through more than one entry point, but almost certainly from the north. In the course of time this species gradually dispersed through the various river systems with some isolated populations evolving into distinct species now extinct (e.g. L. rhachirhinchus and L. karungae, possibly also L. fajumensis), and others or their descendants (like the endemic species of Lake Tanganyika) still surviving. A little modified descendant of the original invader, the species now recognized as L. nilo- ticus, continued to spread (by such means as river capture or lake extension) until it came to have its present distribution. The L. niloticus-like fossils of Pleistocene times (Greenwood, 1959, 1974 ; Sorbini, 1973) stand witness to a much wider area REVIEW OF CENTROPOMIDAE 75 for the distribution of Lates and even probably for the species L. niloticus (but of that point we must remain uncertain). It is unnecessary here to discuss the history of L. niloticus in lakes such as Rudolf and Albert which may, at some time in their histories, have dried out completely, and which have had complex relationships with the River Nile and other lakes (see discussions in Greenwood, 1959, 1974 ; also Beadle, 1974). There is still, however, the problem of the endemic Lates species in the two lakes, L. longispinis in Lake Rudolf and L. macrophthalmus in Lake Albert. In brief, on morphological criteria (p. 12) these species are apparently more closely related to one another than either is to L. niloticus, the species from which each was supposed to have been derived at some time during the Pleistocene (Worthington, 1932 ; Holden, 1967). On the evidence currently available it is impossible to determine whether L. longis- pinis and L. macrophthalmus do in fact represent survivors of a distinct lineage or if, as Worthington (1932) postulated, they are offshoots of earlier L. niloticus popula- tions that once inhabited the two lakes (see discussions on pp. 13 and 14). DIAGNOSES FOR THE CENTROPOMIDAE, ITS SUBFAMILIES, GENERA AND SUBGENERA CENTROPOMIDAE Poey, 1868 Poey, F., 1868, Repertorio Fisico-Natural de Cuba, 5, no. 2 : 280. (See also Gill, T., 1883, Proc. U.S. natn. Mus., 5 : 484-485). TYPE GENUS : Centropomus Lacepede, 1802. DIAGNOSIS. Percoid fishes, some attaining a large size (up to 2m), with the neural spine of the second vertebra markedly expanded in an anteroposterior CENTROPOMINAE LATINAE Eolatest Lates(Luciolates) Lates( Lates) FIG. 37. Cladogram to illustrate phyletic relationships within the Centropomidae. 76 P. H. GREENWOOD direction, and the pored scales of the body lateral line continued onto the caudal fin, reaching the posterior margin of that fin in all but one species. Twenty-four or 25 vertebrae (including the fused first ural and preural centra of the caudal skeleton) ; pleural ribs associated with parapophyses except on the first three to five rib-bearing vertebrae (the first two vertebrae of the column are without ribs) ; 3 predorsal bones. Dorsal fin either deeply divided, the first part entirely spinous (7 or 8 spines), the second of one spine and 8-n branched rays, or the two parts of the fin separated by a distinct gap ; anal fin with 3 spinous and 6-9 branched rays ; caudal fin with 17 principal rays, its posterior margin rounded, truncate or forked. Scales ctenoid, small to moderate in size, dorsally not extending forward on to the head beyond the level of the midpoint of the eye (usually only to the level of the posterior margin of the orbit) but present on the cheek and operculum ; scaly sheath at the base of the anal and soft dorsal fins, but squamation extending onto all fin membranes (including that of the caudal). No scales on the maxilla ; a small supramaxilla present. Teeth on the premaxilla, dentary, vomer, palatine and, in most species, the ectopterygoid ; teeth absent, except in Psammoperca, from the glossohyal. Jaw teeth generally small, viliform or conical, and arranged in several rows. Pterosphenoid pedicle and internal jugular bridge present in all but three or four species, although variously developed ; frontoparietal crests present. Seven branchiostegal rays. Pseudobranch present. About 20 extant species from marine, estuarine and freshwater habitats in the tropical New World (Atlantic and Pacific coasts), tropical Africa (predominantly fresh- or brackish water species), and from Indo-Pacific coastal waters. Six extinct species (some from Europe), the earliest being from the Eocene of Monte Bolca. Subfamily GENTROPOMINAE Centropomid fishes with 24 vertebrae ; the cephalic lateral line canals not enclosed in bony tubes but carried in skin-covered bony gutters ; the supraoccipital barely separating the frontals ; the first anal pterygiophore hypertrophied and inclined backwards at an oblique angle ; no opercular spine but three or four enlarged spines at the posterior angle of the operculum ; no swimbladder-posttemporal ligament developed ; no isolated spine or spines situated between the first and second dorsal fins (these fins always separated by a distinct gap) ; pseudobranch superficial. TYPE GENUS : Centropomus Lacepede, 1802. A single genus Centropomus (type species Sciaena undecimalis Bloch, 1792), generic diagnosis as for the subfamily with, additionally, caudal fin skeleton having three epural and two uroneural bones. The genus is confined to the tropical waters of North, Central and South America, and occurs on both the Pacific and Atlantic coasts. A key to the species of Centropomus is provided by Meek & Hildebrand (1925), and supplementary information by Chavez (1961) and Rivas (1962). Subfamily LATINAE Jordan (1923) Centropomid fishes with 25 vertebrae ; the cephalic lateral line enclosed in bony tubes ; the supraoccipital extending far forward between the frontals ; the first REVIEW OF CENTROPOMIDAE 77 anal pterygiophore not hypertrophied, and inclined backwards at only a slight angle ; a single well-developed opercular spine and a single, enlarged spine at the posterior angle of the preoperculum ; a stout ligament connecting the swimbladder with the posttemporal (which is itself modified to receive the ligament) ; dorsal fin deeply indented or separated into two fins (between which there are one or two isolated spines) ; pseudobranch covered. TYPE GENUS : Lates Cuvier & Valenciennes, 1828. Three genera, two extant and one extinct. The two extant genera are : PSAMMOPERCA Richardson, 1844 TYPE SPECIES : Ldbrax waigiensis C. & V., 1828. Latine fishes with : a smooth horizontal limb to the preoperculum, a basihyoid tooth plate, supralamellar tooth plates present on the outer face of the first four gill arches only ; with a single series of lateral line scales on the caudal fin, with the nostrils of each side widely separated, and a caudal fin skeleton in which there are two epural bones and a single uroneural. Probably only one species, P. waigiensis (a second nominal species P. macroptera Giinther, 1859, is almost certainly a synonym), widely distributed in the coastal waters of the Indo-Pacific. LATES C. & V., 1828 TYPE SPECIES : Perca nilotica, L., 1758. Latine fishes with the horizontal limb of the preoperculum produced into three or four (rarely more) large, flattened and triangular spines, no basihyoidal tooth plate but supralamellar tooth plates present on both the anterior and posterior faces of the first four gill arches, with three series of lateral line scales on the caudal fin, with the nostrils of each side close together, and a caudal fin skeleton with two epurals and two uroneurals. Eight extant species (seven of which are African and confined to freshwaters, and one marine or estuarine and widely distributed in Indo-Pacific coastal waters) arranged in two subgenera : LATES (LATES) TYPE SPECIES : L. niloticus (L). Species of the genus in which the posterior face of the lateral ethmoid has only a slight slope posteriorly, the dorsolateral parts of that bone are almost horizontally aligned, and the entire ethmovomerine region of the skull is not noticeably elongate. Four species : L. calcarifer (Indo-Pacific), L. niloticus (rivers of northern and western tropical Africa, and also in Lakes Chad, Albert and Rudolf [introduced into Lakes Victoria and Kioga]), L. macrophthalmus (Lake Albert only) and L. longispinis (Lake Rudolf only). 78 P. H. GREENWOOD LATES (LUCIOLATES) TYPE SPECIES : Luciolates stappersi Boulenger, 1914. Species of Lates having a characteristically shaped and elongate ethmovomerine skull region in which the posterior face of the lateral ethmoid slopes backwards at a pronounced angle, and the dorsolateral aspects of that bone are directed ventrally at a steep angle. Four species, all endemic to Lake Tanganyika : L. angustifrons, L. microlepis, L. mariae and L. stappersi. The single extinct genus is : EOLATES Sorbini, 1970 TYPE SPECIES : Lates gracilis Agassiz, 1883. See Sorbini, 1973, for full description, synonymies, etc. Eolates, with one species (E. gracilis} and possibly a second, E. macrurus (Ag.), 1833, is known only from the lower Eocene deposits of Monte Bolca, northern Italy. Eolates differs from Lates in the structure of its caudal fin skeleton (three epurals ; a well-developed neural spine on the second preural vertebrae), in having only a single series of lateral line scales (the median one) on the caudal fin, and in having a less deeply indented dorsal fin (see p. 70 above). The phyletic relationships of Eolates within the Latinae are discussed on p. 71, where it is suggested that Lates and Eolates are sister taxa and should be placed in the Tribe Latini nov., the sister group of the Tribe Psammopercini nov. (a taxon containing only the genus Psammoperca) . ACKNOWLEDGEMENTS Again, it is a pleasure for me to thank Drs Colin Patterson, Donn Rosen and Gareth Nelson for the numerous and illuminating discussions we have had on the centropomid fishes and on aspects of their anatomy in a broader context. To my colleague, Gordon Howes, goes my gratitude for the great amount of work and skill he has put into producing the figures illustrating this paper, and, as always, for his invaluable assistance in innumerable other ways. To Dr Thys van den Audenaerde go my thanks for lending me specimens of Luciolates stappersi from the Musee Royal de 1'Afrique Centrale, Tervuren, to Dr Mary Burgis (the City of London Poly- technic) for her considerable efforts in getting material from Lake Tanganyika, and to my colleague, Dr Keith Banister, for examining the type of Eolates gracilis on my behalf. REFERENCES ALLEN, W. F. 1905. The blood vascular system of the Loricati, the mail-cheeked fishes. Proc. Wash. Acad. Sci. 7 : 27-157. ALLIS, E. P. 1909. 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Neue Beitrage zur Kenntniss der fossilen Fische Osterreichs. Denkschr. Akad. Wiss. Wien, 11 : 187-274. HOLDEN, M. J. 1967. The systematics of the genus Lates (Teleostei: Centropomidae) in Lake Albert, East Africa. /. Zool., Lond. 151 : 329-342. HOLMGREN, N. & STENSIO, E. A. 1936. Kranium und Visceralskelett der Akranier, Cyclosto- men und Fische. In : Bolk, L., Goppert, E., Kallius, E. and Lubbock, W. (Eds), Handbuch der vergleichenden Anatomie der Wirbeltiere, 4 : 233 — 500. Berlin and Vienna. JORDAN, D. S. 1923. A classification of fishes including families and genera as far as known. Stanford University Publications, University Series, Biological Sciences, 3, no. 2 : i-x + 77-243- 8o P. H. GREENWOOD KATAYAMA, M. 1954. Systematic position of the genus Glaucosoma. Bull. Fac. Ed. Univ. Yamaguchi, 4, no. i : 23-29. 1956. On the external and internal characters of Lates calcarifer (Bloch), with its systema- tic position. Bull. Fac. Ed. Univ. Yamaguchi, 6, no. i : 133 — 140. KRAMBERGER, K. G. 1902. Palaeichthyologische Beitrage. Mitt. Jb. K. ung. geol. Anst. 14 : 1-21. LAKE, J. S. 1971. The freshwater fishes and rivers of Australia. 61 pp. Melbourne. LINDBERG, G. U. 1971. Opredelitel' i kharakteristika semelstv ryb mirovolfauny. Izdatel'stvo 'Nauka' Leningrad. (Translated as : Fishes of the world. Halstead Press, New York, 1974- 545 PP-) 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. no. 221 : xiv-239pp. MATSUBARA, K. 1955. Fish morphology and hierarchy. Pts I-III. 1605 pp. Tokyo, Ishizaki-Shoten . MEEK, S. E. & HILDEBRAND, S. F. 1925. The marine fishes of Panama. Pt II. Publs Field Mus. nat. Hist. no. 226, Zool. Ser., 15 : 331-707. MUNROE, I. S. R. 1961. The marine and freshwater fishes of Ceylon. 351 pp. Dept. of External Affairs, Canberra. NELSON, G. J. 1969. Gill arches and the phylogeny of fishes, with notes on the classification of vertebrates. Bull. Am. Mus. nat. Hist. 141 : 475-552. NORMAN, J. R. 1966. A draft synopsis of the orders, families and genera of recent fishes and fish-like vertebrates. 649 pp. British Museum (Nat. Hist.), London. PATTERSON, C. 1964. A review of Mesozoic acanthopterygian fishes, with special reference to those of the English Chalk. Phil. Trans. R. Soc. B, 247 : 213-482. 1975. The braincase of pholidophorid and leptolepid fishes, with a review of the actino- pterygian braincase. Phil. Trans. R. Soc. B, 269 : 275-579. PELLEGRIN, J. 1922. Poissons de 1'Oubanghi-Chari recueillis par M. Baudon. Description d'un genre de cinq especes et d'une variete. Bull. Soc. zool. Fr. 47 : 64-76. POLL, M. 1953. Poissons non Cichlidae. Result, scient. Explor. hydrobiol. Lac Tanganyika (1946-1947), 3 (sA) : 1-251. I957- LCS genres des poissons d'eau douce de 1'Afrique. Annls Mus. r. Congo Beige Ser. 8°, Sci. Zool. 54: 1-191. REGAN, C. T. 1913. The classification of the percoid fishes. Ann. Mag. nat. Hist. (8), 12 : 111-145. RICHARDSON, J. 1844. Ichthyology. In : Richardson, J. & Gray, J. E. (Eds), The zoology of the voyage of H. M.S. Erebus and Terror, under the command of C apt. Sir J. C. Ross during 1839-43. 139 pp. London. RIVAS, L. R. 1962. The Florida fishes of the genus Centropomus, commonly known as snook. Q. Jl Fla Acad. Sci. 25 : 53-64. ROGNES, K. 1973. Head skeleton and jaw mechanism in Labrinae (Teleostei: Labridae) from Norwegian waters. Arbok Univ. Bergen Mat.-Naturv. Ser. no. 4 : 1-149. ROSEN, D. E. 1973. Interrelationships of higher euteleostean fishes. Zool. J. Linn. Soc. 53 (Suppl. i) : 397-5I3- SORBINI, L. 1973. Evoluzione e distribuzione del genere fossile Eolates e suoi rapporti con il genere attuale Lates ( Pisces -Centropomidae). In : Studi e ricerche sui giacimenti terziari di Bolca. Memorie Mus. civ. Stor. nat. Verona (fuori Ser. 2), 2 (i) : 1-43. STINTON, F. C. 1966. Fish otoliths from the London Clay. In : Casier, E., Faune ich- thyologique du London Clay : 404-496. British Museum (Nat. Hist.), London. WEBER, M., & DE BEAUFORT, L. F. 1929. The fishes of the Indo-Australian Archipelago, 5:xiv + 458. Leiden. WEILER, W. 1929. Die mittel- und obereocane Fischfauna Agyptens mit besonderer Beriick- sichtigung der Teleostomi. Abh. bayer. Akad. Wiss., Munich, n.f. 1 : 1-57. REVIEW OF CENTROPOMIDAE 81 WORTHINGTON, E. B. 1929. New species of fish from the Albert Nyanza and Lake Kioga. Proc. zool. Soc. Lond. 1929 : 429-440. 1932. Scientific results of the Cambridge Expedition to the East African Lakes, 1930-1. -2. Fishes other than Cichlidae. /. Linn. Soc. (Zool.), 38 : 121-134. P. H. GREENWOOD, D.Sc. Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON, SWy 5BD A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) 1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96 ; 14 Plates. 1965. (Out of Print.) 2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125 ; 29 Plates, 77 Text-figures. 1969. £4'5o. 4. HAYNES, J. R. Cardigan Bay Recent Foraminifera (Cruises of the R.V. Antur) 1962-1964. Pp. 245 ; 33 Plates, 47 Text-figures. 1973. £10-80. 5. WHITEHEAD, P. J. P. The Clupeoid Fishes of the Guianas. Pp. 227; 72 Text-figures. 1973. £9-70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : the Biology and Evolution of a Species Flock. Pp. 134 ; i Plate, 77 Text-figures. 1974. £3-75. Hardback edition £6. Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU fSAPRf > UHUURY FOSSIL REPTILES FROM ALDABRA ATOLL, INDIAN OCEAN E. N. ARNOLD BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 29 No. 2 LONDON: 1976 15 APR FOSSIL REPTILES FROM ALDABRA ATOLL, \^AJ INDIAN OCEAN BY EDWIN NICHOLAS ARNOLD Pp. 83-116 ; 3 Text-figures, 4 Tables BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 29 No. 2 LONDON: 1976 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Scientific Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 29 No. 2 of the Zoology series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). ISSN 0007-1498 Trustees of the British Museum (Natural History), 1976 BRITISH MUSEUM (NATURAL HISTORY) Issued 13 April, 1976 Price £2-25 FOSSIL REPTILES FROM ALDABRA ATOLL, INDIAN OCEAN By E. N. ARNOLD CONTENTS Page SYNOPSIS ........... 85 INTRODUCTION .......... 86 Family TESTUDINIDAE - LAND TORTOISES ...... 88 Geochelone ........... 88 Family CROCODYLIDAE - CROCODILES ....... 89 Crocodylus ........... 89 Family GEKKONIDAE - GECKOES ....... 90 Paroedura ........... 90 Geckolepis ........... 91 ? Phelsuma ........... 94 Family IGUANIDAE - IGUANID LIZARDS ...... 95 Oplurus ........... 95 Family SCINCIDAE - SKINKS ........ 98 'Scelotes' ........... 98 Mabuya ........... 100 ORIGIN OF THE POINT HODOUL LIZARD DEPOSIT ..... 101 COMPARISON OF ALDABRA REPTILES WITH THOSE OF NEIGHBOURING AREAS 104 HABITAT REQUIREMENTS OF EXTINCT ALDABRA REPTILES . . . 106 POSSIBLE RESOURCE PARTITION BY EXTINCT ALDABRA REPTILES . . 107 BODY SIZE AND 'ISLAND GIGANTISM' ....... 109 POSSIBLE CAUSES OF EXTINCTION . . . . . . . in COMPETITIVE EXCLUSION . . . . . . . . in EXTERMINATION BY AN INVADING PREDATOR . . . . . 112 LOSS OF ECOLOGICAL RESOURCES . . . . . . . 113 COLONIZING ABILITY OF ALDABRA REPTILES . . . . . 113 ACKNOWLEDGEMENTS . . . . . . . . . 114 REFERENCES ........... 114 SYNOPSIS The present reptiles of Aldabra comprise only the giant tortoise, Geochelone gigantea, two geckoes, Phelsuma abbotti and Hemidactylus mercatorius, and the skink, Cryptoblepharus boutonii, but a richer and quite different fossil fauna has recently been discovered. Giant tortoises occur in most terrestrial deposits on the atoll and remains of crocodiles and lizards have been found in two : the Bassin Cabri Calcarenites (undated, but considerably older than 125 ooo years B.P.) and cavity fillings in the Aldabra limestone at Point Hodoul formed since 100 ooo years B.P. The Bassin Cabri Calcarenites contain remains of a crocodile similar to Crocodylus niloticus and of an iguanid lizard of the genus Oplurus. These also occur in the Point Hodoul deposits together with five other kinds of lizards, two or three of which are geckoes (a Paroedura similar to P. sanctijohannis and P. stumpffi, a Geckolepis close to G. maculata and what is possibly a Phelsuma) and two skinks (a 'Scelotes' similar to 'S.' johannae and a Mabuya very like M. maculi- labris). The Point Hodoul lizard remains may be the food residue of a predator, perhaps an owl. The reptile fauna of Aldabra is much more similar to that of Madagascar and the Comores 6* 86 E. N. ARNOLD Islands than to that of East Africa or of the Seychelles. Composition of the Point Hodoul fauna and the presumed requirements of some of its members suggest that conditions on Aldabra at that time may have been rather similar to those now occurring on the Comores. It is likely that the ecological requirements of the fossil forms were sufficiently different for them all to be able to coexist. Some of the fossils clearly differ in size from their closest modern relatives, both the Geckolepis and the Opiums being very large ; possible reasons for this are discussed. Aldabra was completely submerged after the laying down of the Bassin Cabri Calcarenites, but not since the Point Hodoul deposits were formed. Extinction of the reptiles found in the latter may have been largely caused by transient or permanent loss of ecological resources, although competition from the species existing on the atoll today could have been a minor factor. The possible importance of invading predators is difficult to assess. All the reptiles known from Aldabra seem to have been well adapted to the problems of transmarine colonization. There is evidence that the giant tortoises reached the island three times and the crocodile and Oplurus at least twice. INTRODUCTION THE study of island faunas has had a long history and is again fashionable, one of the most influential events in bringing this about being the appearance of The theory of island biogeography by MacArthur and Wilson in 1967. Among the topics that have recently received attention is the problem of what factors limit the number of species on islands and the importance of extinction rates in this process. A restriction on such investigations is the paucity of direct evidence of natural faunal change on islands. This is especially true in the case of reptiles. Instances are known where late Pleistocene or more recent island reptiles have become extinct (see, for instance, Etheridge, 1964, 1965, 1966 for the West Indies, Bravo, 1953 for the Canary Islands and Vinson & Vinson, 1969 for the Mascarenes), but often only part of the previous fauna has disappeared and there is frequently circumstantial evidence that human influences have been important in bringing such changes about. Situations where there has been an extensive faunal turnover in probably more natural conditions would consequently be of interest. Aldabra appears to be a case in point. Aldabra is situated about 640 km east of the African mainland, about 380 km northeast of the Comores Islands and some 420 km northwest of Madagascar (see Fig. i). It is a low atoll, being only about 10 m above sea level at its highest point, and is some 34 km long by 14-5 km wide. There is a large central lagoon and the total land area is 155 km2. Much of the present surface consists of coral limestone, which is often covered by scrub. There are few natural large trees and the lagoon is fringed by mangrove. The present reptile fauna consists of giant tortoises (Geochelone gigantea Schweigger, 1812), two geckoes (Hemidactylus mercatorius Gray, 1842 and Phelsuma dbbotti Stejneger, 1893) and the skink, Cryptoblepharus boutonii (Desjardins, 1831). None of these species is confined to the atoll and all of them have relatively wide distributions in the West Indian Ocean. The island has been the subject of considerable scientific research by a series of expeditions organized by the Royal Society since 1966. Some of the results of this work have been published and a number of papers on the atoll form volume 260 of Philosophical Transactions of the Royal Society of London, series B (1971). Braith- waite, Taylor & Kennedy (1973) have given an account of the depositional and FOSSIL REPTILES FROM ALDABRA 87 SEYCHELLES . '• AMIRANTES -.- AFRICA ALDABRA COMORES MADAGASCAR FIG. I. West Indian Ocean, showing position of Aldabra and other nearby islands mentioned in text. erosional history of Aldabra. During investigations on the island, Dr J. D. Taylor of the British Museum (Natural History) collected substantial amounts of fossil material including the reptiles that form the basis of this paper. Large numbers of tortoise remains were also observed in situ but could not be extracted for detailed examination. Tortoises occur in many of the terrestrial deposits of Aldabra (Braithwaite et aL, 1973). Lizards and crocodiles have also been found, but only in two deposits, the Bassin Cabri Calcarenites and cavity fillings in the Aldabra Limestone at Point Hodoul (see Fig. 2). The Bassin Cabri Calcarenites are of unknown age, but they are older than the Aldabra Limestone which has yielded dates by the 23 nj ,Q n5 II r % CU 12 ^ 1Z2 £13 co T; (-] 'O r3 T3 r^, •— < •s 5 c rt « c x rt PsaJridcJ^ ffl feffi^g^S H ' be more or h 'cn >«% 0 PH + a> ' H M 'O ^ 1 1 O 2 T3 "3 O o 8 ^ "d ° ^ .5 o tuO O M ID N N O OH 1 MM J CD M fl pS °° 1 5 .s CO ,g i-i n ^ «» o* , ^%t c^« ^5 1 1-1 i— i M '5? H W 1 S "O "S m (2 •S o ... vO N H m W 1 H HI a s~~* M O "ti .•2 o ^3 S 'I S ° o to rt G i If? H "2 cn" ^ o M g -H 60 .2 d t r5n MH "o O (U CO n CD O H H -M CO ** OJ ^ 4J s cu T5 fH a 60 3 "Tfl 0 w w " f 1 | 43 J 1 o 5 11 a s ^ P .3 _N _cn O « ««H to c 2 -53 5 ° rd JOT S^S^"^« Jf o ^g 'i^^s^^,^^ &» a ^5^^^s 3 . 5 "o 1 T f\ ? •*; »*i «* "S •4-> i Ilil^ll5 104 E. N. ARNOLD (93-0% nocturnal) and even if it is regarded as diurnal, nocturnal forms still pre- dominate 34 : 23 (597% nocturnal). Limited size distribution and a high proportion of nocturnal animals would fit predation by an owl. In fact, apart from the single large Opiums and the one juvenile Paroedura, both of which may not necessarily have been part of a predator food residue concentration, the size distribution of the Point Hodoul lizards is well within the limits recorded for West Indian owl deposits. While no owls are known from the fossil fauna of Aldabra and none are present now, the cosmopolitan barn owl, Tyto alba (Scopoli), was in residence there in 1906 (Benson & Penny, 1971). This species occurs on many quite isolated islands throughout much of the world and is thought to have been one of the agents that produced the West Indian deposits, so it might have produced the Aldabra concentration although some other species could have been responsible. COMPARISON OF ALDABRA REPTILES WITH THOSE OF NEIGHBOURING AREAS It is not possible to compare the extinct Aldabra reptiles with even roughly contemporaneous faunas in nearby areas for, with the exception of relatively late material from the Mascarenes, very few Quaternary reptile remains, especially those of lizards, are known from these regions. Table 2 shows the known Aldabra reptile fauna, past and present, and its distribution in neighbouring areas. It is apparent from this that Aldabra has more similarity to Madagascar and the Comores than to mainland East Africa and the Seychelles. The Comores are thus the island group that shows most resemblance in its present fauna to the Point Hodoul fossil assemblage, which is not unexpected as this archi- pelago lies less than 400 km from Aldabra. However, the Comores have a number of forms that are unrepresented in the fossil material or indeed in the extant fauna of the atoll. Thus, there are up to three species of Phelsuma on each island and on one or more of them occurs the small gecko, Ebanavia inunguis Boettger, 1878, chameleons, the skink, Mabuya striata, and up to three species of snake. It cannot be ruled out that some of these forms were present on Aldabra when the Point Hodoul deposits were being laid down, for the sample of fossil lizards is very likely not to be fully representative. It is small, consisting of a minimum of 57 individuals, and of the six species present three could be represented by only one, one and two individuals respectively. Even assuming that the remains were randomly accumulated, sampling error alone might have excluded some species. In fact, random accumulation is most improbable, for even a wide-ranging predator is unlikely to have hunted evenly over all habitats of the atoll and, as has been shown, small forms and diurnal ones are under-represented. So, if minute lizards like Ebanavia and diurnal forms like chameleons and additional species of Phelsuma were present, they might well have been excluded from the sample. For the same reasons, the possibility cannot be discounted that the tiny diurnal skink, Crypto- blepharus boutoni, now on the atoll may have been present when the Point Hodoul deposit was formed. The extant gecko, Hemidactylus mercatorius Gray, 1842, on FOSSIL REPTILES FROM ALDABRA 105 a b«c M I •9 ******** * ^ £ § 5 «s '-S ^* S3 1 3,8 1 II 2 Ifl •§ § S« to s '2 o IN « • a •§ 9 •2 |-§ ^> §-^ « % ~ •^ s 'il-il ^a^^l • bx) O* -i-> "o |' 0 £ 0 0 o o p , cio FT.. a fS c o CD 5 2 3 T3 0 i >l It It It CD oj > > •gi o o bx> bX) a X3 a 2 fe x> 13 o rH t +> CD in CD +J m 0 +j CO CO 0) CD I 1 _0 '-3 2 s >-, ctf o3 it it 4J ^ X) X) Xi ^Q p^ O r™, 1 CD CD CD -i-J CD «O •H 2 L B l-i D It It CD CD »H ^J (!_) ^j S CO s fi ^ > jjj _c > > •S .S li T) 9 CQ 1^ I iG a -g- -o p _a ^ 13 rt £j , .^ be o f^ in co o o 1—1 a M w M H CN vO X CD rt 0 C 3 03 St 1-1 ft ft "CJ CTJ 0 N a 'rd 3 a 0 -i-> J3 O ^ s 4j 4-> 4-> It ft it fli O ffi d X) 0 'I -M S 'S ' — be a g ^ 3 'O rt cd .« £ - -S a, ^ s £••§ FOSSIL REPTILES FROM ALDABRA 109 nocturnal forms. Habitat differences are often based on variations in insolation, humidity and substrate (e.g. ground type, whether more or less horizontal or vertical surfaces are preferred and, if the latter, whether they are trees, rock faces etc.). As most lizards are general predators, food size is often a more important parameter in reducing competition than food type, although a minority of species are specialists taking high proportions of, for instance, vegetation, ants or small vertebrates. As stated, not much information is available about the habits of the modern relatives of the members of the Point Hodoul fauna. But what little there is can be augmented by various means, including examination of stomach contents (for dietary differences), speculation based on their morphology and analogies drawn from related forms. Thus, lizard head length often correlates with the size of prey usually eaten, well developed toe-pads in geckoes typically indicate good climbing ability, and large corneas and vertically slit-shaped pupils suggest at least partial nocturnality. Potentially important differences between the Point Hodoul lizards are given in Table 3. From this it can be seen that members of any pair of species may well have differed in at least one parameter of probable ecological significance and usually in more. So, it is unlikely that any of them would have been precise ecological equivalents and there is no reason to think that they could not have coexisted, assuming of course that the atoll supported the minimum microhabitat diversity to allow this. BODY SIZE AND 'ISLAND GIGANTISM' The most striking differences between the Point Hodoul lizards and their modern relatives are in apparent maximum size. Thus the linear dimensions of the Opiums may have been almost twice those of the largest living member of the genus and the Geckolepis was about 30% longer than the largest species alive today. The Paroedura and 'Scelotes', on the other hand, may have been somewhat smaller than their extant relatives. A general tendency to large body size in a number of lizard and other groups on small islands is often given formal recognition in the literature as 'island gigantism'. Certainly, in many of the more important lizard families some of the largest forms (although not all) occur on small islands. This is true for geckoes, iguanids, lacertids, skinks and varanids and a similar trend is apparent in land tortoises. One probable reason for this is that many reptiles are much better transmarine colonizers than most mammals and are better at surviving transient periods of adverse conditions. Consequently they often get to, and persist on, small islands where the relatively large mammals that would normally occupy predator and vegetarian niches in mainland areas are entirely absent. The reptiles can therefore expand into this vacant ecological space with consequent increase in body size (or if they were large to start with, without reduction). Very large lizards occur, or have quite recently occurred, on at least five islands or island groups in the African region (see Table 4). Significantly, four out of the five giant forms involved are, or were probably, sub- stantially vegetarian (viz. Gallotia, Macroscincus, Phelsuma gigas and, by analogy with its living relatives, the Aldabra Opiums ; the diet of Didosaums is uncertain). no E. N. ARNOLD aximum size, CD a o-^ i L o g ^J t-H 0 210 Geckoes (Tarentola) Skinks (Chalcides) 150 Geckoes (Tarentola) Skinks (Mabuya) 132 Geckoes (see text) Skinks (see text) 80* Geckoes (e.g. Phelsuma) o I cn ^ |1 1 bo CJ CD O 0 N 2 cn 4 in the African region Largest related forms in possible source £ CO CD c d 0 IH O nf z CO mainland or large island Lacerta lepida (S.W. Europe, N.W. Africa) Mabuya perrotetii (West Africa) Oplurus cuvieri (Madagascar and Comores) Members of Group II of the Leiolopisma assemblage (Greer, 1974) (East Indies) P. madagascariensis (Madagascar etc.) cn H 'O •k £ 1 fj N d CD ^i * I §s "\ w 1 •2 J cn SH •« -i Jo co S IH S •*•* V § B "S co O if* « * 0-2 u -^ « CO B I Td auritius N CD 3 bo •E 1 3"S ,0 CQ # -1— I-H > h- 1 5j X K FOSSIL REPTILES FROM ALDABRA in Yet, although this habit is not uncommon in iguanids, it is not, or only weakly, developed in mainland geckoes, skinks and lacertids. It is also noteworthy that none of the five giants in Table 4 are closely related to each other, even when they occur on neighbouring islands that were probably colonized from the same general source area as with Mauritius and Rodriguez. This indicates that the ability to produce giants is widespread amongst lizards, but possibly once a large form has developed on a small island, its presence inhibits the development of further giants. This is supported by the fact that, in the listed examples, other species coexist with the giants but have not become large, even when they are members of groups that have produced giants elsewhere (the Mabuya stock including Macroscincus, Phelsuma). Evolution and maintenance of size differences seem to be common phenomena in island lizards, for instance Schoener (1970) found that Anolis (Iguanidae) species in the West Indies often diverge in body size when sympatric. Such differences may be particularly marked on small islands because restricted habitat diversity limits the extent to which species can partition resources by the selection of different microhabitats and therefore increases the need to evolve differences in prey size and hence body size. The large apparent size difference between the two probably nocturnal geckoes in the Point Hodoul cavity filling may be a case in point. Here, Paroedum seems to have grown to about 60 mm snout to vent, while Geckolepis attained 90 to 100 mm. On the Comores and on Madagascar, members of these two genera do occur together with wide overlap of size, but it is certain that these localities provide a wider range of environments than Aldabra did, so evolution of clear size differences has not been necessary. As might be expected from experience with domestic animals, size in vertebrates is often very labile and easily modified by selection. Certainly in lizards the body size of populations may vary quite extensively through time. For instance Etheridge (1964, 1965, 1966) found that in the West Indies the body size of several species was considerably less than that of presumed ancestral fossil populations on the same island. POSSIBLE CAUSES OF EXTINCTION Although one cannot be sure that none of the present lizards of Aldabra was on the atoll when the Bassin Cabri Calcarenites and the Point Hodoul deposits were formed, it is certain that the crocodile and the six lizards represented in them are now extinct on the island. Among possible reasons for their disappearance are : (i) com- petitive exclusion by forms now present on the island ; (2) extermination by an invading predator ; and (3) transient or permanent loss of essential ecological resources. Competitive exclusion If the three present Aldabra lizards arrived after the Point Hodoul deposit was formed, it is possible that they could have displaced some of the species represented 112 E. N. ARNOLD in it. However, in a stable situation, with no accompanying changes in ecological resources, this would only be likely to happen if the ecological space required by the invader more or less completely overlapped that of one or more of the species already there. Also, in a stable situation, an island lizard is quite likely to be better adapted to its immediate environment than one coming from elsewhere. This, together with the fact that an invader would be initially greatly outnumbered and would just have completed a debilitating transmarine journey, should give the original inhabitant great advantages over potential competitors arriving from outside. If, on the other hand, the island environment was undergoing change at the time of invasion in a way that favoured the incursor, then replacement would be more likely. It is possible that Hemidactylus mercatorius has roughly the same requirements as the Aldabra Paroedura (see p. 104) and Phelsuma abbotti might be the ecological equivalent of the putative Phelsuma in the Point Hodoul deposits. The third species now on Aldabra, the skink Cryptoblepharus boutonii, is too small to have been a competitor of either of the fossil skinks, unless of course it competed with their young. But this is not very likely, especially as it is known that Cryptoblepharus can coexist on small islands with forms identical with, or very similar to, the fossils : it occurs with Mabuya maculilabris on Europa and with 'Scelotes' valhallae on Glorioso. So, the present species would only have been likely to displace at most two of the six Point Hodoul lizards. Extermination by an invading predator On small islands, introduced predators often destroy a high proportion of indigen- ous reptiles. Mongooses and rats were probably responsible for the extermination of many endemic West Indian populations and a similar process seems to have taken place on Rodriguez and Mauritius, the former island having lost its two native geckoes. Ten endemic Mascarene reptile species probably occurred on Mauritius of which six are no longer found on the main island and a seventh is apparently restricted to a single locality there (for the lizards, see Vinson & Vinson, 1969, and Vinson, 1973). Habitat destruction may have contributed, but it does not seem to have been the main factor, for five of the forms now extinct on Mauritius itself still survive on offshore islands that are free of mammalian predators even though the natural vegetation is greatly reduced. In such cases, the predators concerned are ones with a broad dietary tolerance, so that a fall in numbers of a particular prey-type does not produce a corresponding drop in predator numbers through starvation. Consequently, predator pressure on a declining species is not reduced. Presumably, island endemics are exterminated because they lack the requisite antipredator strategies to resist an invader and, as the numbers of individuals and their range is restricted, they are hunted out swiftly, before these can evolve. One can envisage a natural situation like this with perhaps an avian predator arriving on an isolated small island like Aldabra. As has been suggested, owls may have been active when the Point Hodoul deposit was formed and rats have reached the island since (presumably by human agency). Both of these might have been capable of reducing the fauna, but it is very difficult to assess their actual importance. FOSSIL REPTILES FROM ALDABRA 113 Loss of ecological resources It is likely that the number of forms an island can support is partly dependent on habitat diversity. Reduction in diversity, even transiently, may mean that some species would not survive. Such reduction may have occurred on Aldabra. For instance, the presence of Geckolepis and Mabuya nr. maculilabris in the Point Hodoul deposit suggests that large trees may have been more abundant at this time ; reduction or temporary complete loss of these could have hazarded the lizards. However, it is not easy to see why so many of the fossil forms disappeared. One possibility is that the island suffered a transient period of very adverse conditions, for instance an extended drought. Such an event seems quite possible in the varying climatic conditions of the late Quaternary. A low atoll like Aldabra would be more prone to extreme drought than high islands like the Comores where orographic in- fluences probably produced a more stable rainfall. These islands still possess a fauna similar to that represented in the Point Hodoul deposits. In the West Indian Ocean, low islands certainly seem more prone to extinction of their faunas, for their level of endemicity is much lower than that on high islands (Peake, 1971). Braith- waite et al. (1973) suggest that the breaching of the Aldabra land rim and the flooding of the lagoon, which took place perhaps 4000 to 5000 years ago and reduced the area of the island by 60%, may have been responsible for widespread destruction of habitat and consequent extermination. Whether one or both these factors were responsible, permanent or transient loss of ecological resources seems a likely primary cause of the extinction of the Point Hodoul reptile fauna. Even if com- petitive replacement was also involved it would have been most likely to take place in such a changing environment. COLONIZING ABILITY OF ALDABRA REPTILES It is virtually certain that Aldabra received its reptiles by transmarine migration (although one or more of the modern species might possibly have been transported by man), so one would expect the forms that reached the atoll to have been well adapted to the hazards of sea crossings and the perilous early stages of colonization. There is some circumstantial evidence that this is so. One indication is that all the reptile groups known from Aldabra have got to at least one other oceanic island that would have required a transmarine journey. Paroedura, Geckolepis and Oplurus are all on the Comores, Mabuya and Crocodylus reached this archipelago and the Sey- chelles as well, and the other groups (if 'Scelotes' is taken to include the endemic scincines of Mauritius and the Seychelles) got to a relatively large number of islands in the West Indian Ocean. Aldabra has probably been completely submerged twice during its occupation by giant tortoises, which would imply that they colonized the atoll not once but three times (Braithwaite et al., 1973). Similarly, remains of Crocodylus and Oplurus occur in both the Bassin Cabri Calcarenites and the Point Hodoul cavity fillings. These deposits are separated by the marine Aldabra limestone, so the reptiles common to them must have colonized at least twice. n4 E, N. ARNOLD In the West Indies, the species of the iguanid genus Anolis that are successful colonizers are all typical of savannah or open forest situations, none of them being mainly rain forest or montane forms (Williams, 1969). Wilson (1959, 1961) has found an analogous situation in Melanesian ants. With the exception of the crocodile, the reptiles that have reached Aldabra fit this pattern. All those in which habits are known or can be guessed would be expected to occur at least some- times in fairly dry habitats, such as littoral situations (in the case of Cryptoblepharus) or savannah. This is true even of the probably tree-associated forms, Geckolepis and Mabuya nr. maculilabris. Williams thinks that this general trend results from the greater ability of dry-adapted animals to survive the almost inevitable desic- cation of a sea passage. Another characteristic to be expected in successful colonizers is the ability to swim, or float, or cling to rafting vegetation. Crocodylus niloticus sometimes lives perma- nently in coastal areas and is obviously well fitted for survival at sea, and giant tortoises are known to float well (see e.g. Grubb, 1971). Virtually all the Aldabra lizards have, or had, good climbing ability and consequently might be expected to maintain their position well on floating objects. The only exception is the very short-legged, probably ground-dwelling 'Scelotes'. Parthenogenesis is obviously an initial advantage in a colonizing species since any individual of an all-female species, irrespective of age, can form a propagule. Some of the most widespread species of oceanic reptiles seem to be parthenogenetic including the worm snake, Ramphotyphlops braminus (Daudin, 1803) - McDowell, 1974, and the gecko Lepidodactylus lugubris (Dumeril & Bibron, 1836) - Gorman, 1973, but all four reptile species on Aldabra at the present time are bisexual and none of the fossil forms belong to genera in which parthenogenesis has been reported. ACKNOWLEDGEMENTS Special thanks are due to the Royal Society who enabled the material on which this paper is based to be collected and made it available to me. I should also like to acknowledge the considerable help given to me by various individuals, in particular that of Dr J. D. Taylor who collected the specimens. Mr C. J. McCarthy made skeletons and radiographs of comparative material and the fossils were prepared by Mr F. M. P. Howie and Mr N. Hughes. Fig. 3 was drawn by Dr A. W. H. Bartram. Professor J. Guibe (Museum Nationale d'Histoire Naturelle, Paris) arranged the loan of some critical specimens from Madagascar and the Comores in his care. The manuscript was read by Miss A. G. C. Grandison, Mr J. F. Peake and Dr Taylor, all of whom provided helpful criticism. REFERENCES ANGEL, F. 1936. Deux gekkos nouveaux de Madagascar appartenant au genre Phyllo- dactylus. Bull. Soc. zool. Fv. 61 : 508—511. — 1942. Les lezards de Madagascar. Mem. Acad. malagache, 36 : 1-193. ANTHONY, H. E. 1919. Mammals collected in eastern Cuba in 1917, with descriptions of two new species. Bull. Am. Mus. nat. Hist. 41 : 625—643. BELLAIRS, A. D'A. 1969. The life of reptiles, vol. II. London. FOSSIL REPTILES FROM ALDABRA 115 BENSON, C. W. & PENNY, M. J. 1971. The land birds of Aldabra. Phil. Trans. R. Soc. B. 260:417-527. BLANC, C. P. 1966. Notes sur les Gekkonidae de Madagascar : I - Observations relatives a deux especes de Geckolepis : G. maculata Peters, 1880 et G. typica Grandidier, 1867. Vie Milieu, 17 : 453-460. 1972. Les reptiles de Madagascar et des iles voisines. In Battistini, R. & Richard- Vindard, G. (eds), Biogeography and ecology in Madagascar : 501-614. The Hague. & BLANC, C. F. 1967. Observations ecologiques sur les Sauriens du Mont Bity. Ann. Univ. Madag. (Serie Sci. Nat. Mat.}, 5 : 67-74. BRAITHWAITE, C. J. R., TAYLOR, J. D. & KENNEDY, W. J. 1973. The evolution of an atoll : the depositional and erosional history of Aldabra. Phil. Trans. R. Soc. B, 266 : 307-340. BRAVO, T. 1953. Lacerta maxima n. sp. de la fauna extinguida en el Pleistocene de las Islas Canarias. Estudios geol. Inst. Invest, geol. Lucas Mallada, 9 : 7-34. BRYGOO, E. R. 1966. Note sur les reptiles terrestres recoltes a Europa en Avril 1964. Mem. Mus. natn. Hist. nat. Paris (Serie A], 41 : 29-31. COTT, H. B. 1961. Scientific results of an enquiry into the ecology and economic status of the Nile crocodile (Crocodilus niloticus) in Uganda and Northern Rhodesia. Trans, zool. Soc. Land. 29 (4) : 211-350. DIXON, J. R. & KROLL, J. C. 1974. Resurrection of the generic name Paroedura for the phyllodactyline geckoes of Madagascar, and a description of a new species. Copeia, 1974 (i) : 24-30. ETHERIDGE, R. 1964. Late Pleistocene lizards from Barbuda, British West Indies. Bull. Fla St. Mus. biol. Sci. 9 (2) : 43-75. 1965. Fossil lizards from the Dominican Republic. Q. Jl Fla Acad. Sci. 28 (i) : 83-105. 1966. Pleistocene lizards from New Providence. Q. Jl Fla Acad. Sci. 28 (4) : 349-358. GORMAN, G. C. 1973. The chromosomes of the Reptilia, a cytotaxonomic interpretation. In Chiarelli, A. B. & Capanna, E. (eds), Cytotaxonomy and vertebrate evolution : 349—424. GREER, A. E. i97oa. A subfamilial classification of scincid lizards. Bull. Mus. comp. Zool. Harv. 139 (3) : 151-183. i97ob. The systematics and evolution of the subsaharan Africa, Seychelles and Mauritius scincine scincid lizards. Bull. Mus. comp. Zool. Harv. 140 (i) : 1-23. 1974- The generic relationships of the scincid lizard genus Leiolopisma and its relatives. A ust. J. Zool. Supplementary series 31 : 1-67. — (in press). On the evolution of the giant Cape Verdes scincid lizard Macroscincus coctei. J. nat. Hist. GRUBB, P. 1971. The growth, ecology and population structure of giant tortoises on Aldabra. Phil. Trans. R. Soc. B. 260 : 327-372. GUNTHER, A. 1879. On mammals and reptiles from Johanna, Comoro Islands. Ann. Mag. nat. Hist. (5) 3 : 215-219. — 1880. Description of new species of reptiles from Eastern Africa. Ann. Mag. nat. Hist. (5) 6:234-238. HECHT, M. K. 1951. Fossil lizards of the West Indian genus Aristelliger (Gekkonidae). Am. Mus. Novit. 1538 : 1-33. LOVERIDGE, A. 1957. Check list of the reptiles and amphibians of East Africa (Uganda; Kenya ; Tanganyika ; Zanzibar). Bull. Mus. comp. Zool. Harv. 117 (2) : 153-362. MACARTHUR, R. H. & WILSON, E. O. 1967. The theory of island biogeography. Princeton, New Jersey. MCDOWELL, S. B. 1974. A catalogue of the snakes of New Guinea and the Solomons, with special reference to those in the Bernice P. Bishop Museum, part I. Scolecophidia. /. Herpetol. 8 (i) : 1-57. MERTENS, R. 1955. Studien iiber die Reptilienfauna Madagaskars, I. Beobachtungen an einigen madagassischen Reptilien im Leben. Zool. Gart. Lpz. 22 : 57-73. MILLER, G. S. 1929. A second collection of mammals from caves near St. Michel, Haiti. Smithson. misc. Collns, 74 (3) : 1-8. Ii6 E. N. ARNOLD MILLOT, J. 1951. Un lizard d'eau & Madagascar (Scelotes astrolabi Dum. et Bib.). Naturaliste malagache, 3 : 87-90. OELRICH, T. M. 1956. The anatomy of the head of Ctenosaura pectinata (Iguanidae). Misc. Publs Mus. Zool. Univ. Mich. 94 : 1-122. PAULIAN, R. 1953. A propos des lezards aquatiques. Naturaliste malagache, 5 : 108. PEAKE, J. F. 1971. The evolution of terrestrial faunas in the Western Indian Ocean. Phil. Trans. R. Soc. B, 260 : 581-610. SAVAGE, J. 1952. The correct generic names for the iguanid lizards of Madagascar and the Fiji islands. Copeia, 1952 (3) : 182. SCHOENER, T. W. 1970. Size patterns in West Indian Anolis lizards. II. Correlations with the sizes of particular sympatric species - displacement and convergence. Am. Nat. 104:155-173. THOMPSON, J. & WALTON, A. 1972. Redetermination in the chronology of Aldabra Atoll by 230Th/234U dating. Nature, Lond. 240 : 145-146. VINSON, J. & VINSON, J.-M. 1969. The saurian fauna of the Mascarene Islands. Bull. Mauritius Inst. 6 (4) : 203-320. VINSON, J.-M. 1973. A new skink of the genus Gongylomorphus from the Macabe Forest (Mauritius). Revue agric. sucr. lie Maurice, 52 : 39-40. WETMORE, A. 1922. Remains of birds from caves in the Republic of Haiti. Smithson. misc. Collns, 74 (4) : 1-4. & SWALES, B. H. 1931. The birds of Haiti and the Dominican Republic. Bull. U.S. natn. Mus. 155 : 234-236. WILLIAMS, E. E. 1969. The ecology of colonisation as seen in the zoogeography of anoline lizards on small islands. Q. Rev. Biol. 44 (4) : 345-389. WILSON, E. O. 1959. Adaptive shift and dispersal in a tropical ant fauna. Evolution, Lancaster, Pa, 13 : 122-144. 1961. The nature of the taxon cycle in the Melanesian ant fauna. Am. Nat. 95 : 169-193. E. N. ARNOLD D. Phil. Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) 1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96 ; 14 Plates. 1965. (Out of Print.) 2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125 ; 29 Plates 77 Text-figures. 1969. £4.50. 4. HAYNES, J. R. Cardigan Bay Recent Foraminifera (Cruises of the R.V. Antur] 1962-1964. Pp. 245 ; 33 Plates, 47 Text-figures. 1973. £10.80. 5. WHITEHEAD, P. J. P. The Clupeoid Fishes of the Guianas. Pp. 227 ; 72 Text-figures. 1973. £9.70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : the Biology and Evolution of a Species Flock. Pp. 134 ; I Plate, 77 Text-figures. 1974- £375. Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol 884 A REVIEW OF THE FAMILY CANIDAE, WITH A CLASSIFICATION BY NUMERICAL METHODS J. GLUTTON -BROCK, G. B. CORBET & M. HILLS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 29 No. 3 LONDON : 1976 A REVIEW OF THE FAMILY CANIDAE, WITH A CLASSIFICATION BY NUMERICAL METHODS BY JULIET GLUTTON-BROCK, GORDON B. CORBET & MICHAEL HILLS Pp 117-199 ; ii Text-figures, 9 Tables BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 29 No. 3 LONDON: 1976 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Scientific Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 29 No. 3 of the Zoology series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). ISSN 0007-1498 Trustees of the British Museum (Natural History), 1976 BRITISH MUSEUM (NATURAL HISTORY) Issued 13 April, 1976 Price £5.95 A REVIEW OF THE FAMILY CANIDAE, WITH A CLASSIFICATION BY NUMERICAL METHODS By JULIET CLUTTON-BROCK, G. B. CORBET & M. HILLS CONTENTS Page SYNOPSIS ........... 119 INTRODUCTION .......... 120 CHARACTERS OF THE CANIDAE . . . . . . . . 121 SOURCES OF DATA .......... 123 SELECTION OF SPECIES ......... 123 DERIVATION OF DATA ......... 124 NATURE OF THE DATA ......... 125 KINDS OF CHARACTERS . . . . . ... 125 USE OF THE DATA MATRIX . . . . . . . . 125 MEASUREMENT OF SIMILARITY ........ 126 NUMERICAL RESULTS ......... 126 DISTRIBUTION OF SIMILARITY ....... 126 NEAR NEIGHBOURS ......... 128 TWO-DIMENSIONAL PLOTS . . . . . . . . 128 SIMILARITY VALUES FOR THE EXISTING CLASSIFICATION . . . 133 HOMOGENEITY OF THE THREE MAIN GENERA ..... 136 AUTOMATIC CLASSIFICATION ........ 137 GENERAL TAXONOMIC CONCLUSIONS ....... 139 SYSTEMATIC ACCOUNT ......... 141 GENUS Cam's .......... 141 GENUS Vulpes .......... 150 GENUS Alopex .......... 161 GENUS Otocyon .......... 162 GENUS Nyctereutes ......... 164 GENUS Dusicyon .......... 165 GENUS Chrysocyon ......... 176 GENUS Speothos .......... 177 GENUS Cuon .......... 179 GENUS Lycaon .......... 180 ACKNOWLEDGEMENTS . . . . . . . . . 181 APPENDIX I: DATA MATRICES ........ 182 APPENDIX II: LICE (PHTHIRAPTERA) OF THE CANIDAE .... 194 REFERENCES ........... 195 SYNOPSIS Within the accepted classification of the Canidae it is usual to recognize three subfamilies, fourteen genera and thirty-seven species, excluding the domestic dog. Using numerical methods and a total of ninety characters an analysis has been carried out of the overall similarity between thirty-five of these species plus two breeds of domestic dog. The specimens used for this analysis are in the collections of the British Museum (Natural History). Classification above 120 J. CLUTTON-BROCK ET AL. the level of species has been critically examined on phenetic characters. The results demon- strate the isolated positions of the monospecific genera Lycaon, Speothos and Cuon and do not strongly support their grouping as a discrete subfamily. The status of Otocyon, Nyctereutes, Chrysocyon and Alopex as monospecific genera is also upheld although Alopex is clearly related to the other foxes of the genus Vulpes. Urocyon and Fennecus are included in Vulpes, and Cerdocyon and Atelocynus in Dusicyon. The three larger genera, Canis, Vulpes and Dusicyon, are retained although they are closely interrelated. Vulpes vulpes is shown to be a distinctly atypical member of its genus. The position of the extinct Falkland Island wolf was found to be enigmatic but it is provisionally retained with other species of Dusicyon. A systematic description is given of each species and the data are presented as a series of tables that may be used for reference. INTRODUCTION THE Canidae, comprising the dogs and foxes, is one of the most clear-cut families of mammals and its content has rarely been seriously disputed since Gray (1821, 1825) first used the name in its present form. Exceptions to this unanimity amongst taxonomists have been the proposal to include the bears and the dogs in a single family (Winge, 1924) ; and at the other extreme the splitting of the Canidae by giving family rank to some of its more aberrant members, e.g. the Otocyonidae of Trouessart (1885) for the single species Otocyon megalotis, the bat-eared fox. Neither of these courses has received general acceptance. The relation of the Canidae to other families of carnivores has been rather more controversial. This was discussed in some detail by Simpson (1945) who accepted the grouping of the Canidae with the Ursidae (bears), Procyonidae (racoons etc.) and Mustelidae (weasels etc.) in a superfamily Canoidea (frequently called the Arctoidea in the older literature), contrasting with the superfamily Feloidea (frequently Ailuroidea in the past) containing the Felidae (cats), Viverridae (civets etc.) and Hyaenidae (hyaenas). In contrast to the stable concept of the family Canidae, classification within the family has been distinctly unstable. At the species level re visionary work has progressively clarified the limits of the separate species until at present the only real uncertainty concerns the South American forms usually placed in the genus Dusicyon s.l. Classification at the generic level has been particularly unstable and even those generic allocations that have stood intact for the past century have done so by neglect rather than by the soundness of their foundations. The genus Cuon for example, containing the single species C. alpinus, the red dog of S.E. Asia, is usually diagnosed in terms of a single character, namely the absence of third lower molars. The Simien jackal of Ethiopia has been variously considered to be the sole member of a genus Simenia or has been placed with the other jackals in Canis. Likewise the Arctic fox has been given generic rank as Alopex lagopus or has been included in Vulpes. Although these are to some extent questions of taxonomic rank there has been a tendency to attempt to settle them in isolation without taking into account other members of the genera concerned. THE FAMILY CANIDAE 121 Grouping of the genera into subfamilies has been equally controversial. Simpson (1945) probably followed some degree of consensus in recognizing three subfamilies : Otocyoninae (Otocyori), Simocyoninae (Lycaon, Cuon, Speothos) and Caninae (all other genera). But these members of the Simocyoninae are so diverse that many doubts have been expressed about the validity of such a grouping. The last comprehensive revision of the Canidae was that of Mivart (1890) since when an enormous amount of relevant data has accumulated and many piecemeal taxonomic changes have been made. The main objective of the present study was therefore to revise the classification within the family, taking account of all species and all available characters. No attempt was made to resolve outstanding prob- lems at the species level. The principle followed in determining which characters to use was to include all characters that showed clear-cut interspecific differences anywhere within the family. Most previous studies of classification in mammals using numerical methods have been confined to limited sets of characters, e.g. that of parts of the Felidae by Imaizumi (1967) using only the skull and that of New Guinea rodents by Lidicker (1973) using only the penis. These seem to have the weakness of including many characters that show very little variation within the group concerned and of ignoring major differences in other parts of the body. By using all those characters showing major, clear-cut differences between species, it was hoped that a sufficiently large and representative sample of characters would be obtained without having to include the more subtle differences that can only be detected in terms of differences in mean value between one species and another. As is the case with most groups of mammals, literature on the Canidae is exces- sively fragmented. It was therefore considered that the data matrices would in themselves be a useful source of information and they are presented here (Tables 4-9) in a form that we hope will be of value for reference. Recently a comprehensive popular account of the family has been produced by Bueler (1974), and a more technical review, within the framework of behaviour patterns, by Fox (1975). CHARACTERS OF THE CANIDAE The Canidae are cursorial, terrestrial carnivores that have their young in burrows or dens in the ground. The family is distributed over the greater part of the habit- able world with the exception of some oceanic islands. The species may be solitary, hunting on their own or in restricted family groups and living off small prey, or they may be social pack animals like the wolves that hunt large prey. All canids will feed on some vegetable matter and carrion, especially when the preferred diet is scarce. In all species the jaws are well developed, the head is longer than in the cat family, and the ears are prominent. The body is lightly built and the limbs are long. Pelage characters are variable but the dominant colours are black, white, grey and ochreous or tawny brown. There is usually a dense underfur mixed with longer dark or 'agouti' guard hairs (Little, 1957). The tail is usually bushy, often with a contrasting black or white tip and a patch of dark hairs on the dorsal part covering a glandular area (called the 'tail gland' throughout this work, see 122 J. CLUTTON-BROCK ET AL. Hildebrand, 1952!)). The fur is thicker and lighter coloured in the winter and in low temperature zones. The underparts of the body, inner sides of the limbs and insides of the ears are usually paler in colour than the rest of the body. The sense organs are well developed and most species are predominantly nocturnal. Individuals communicate with each other by facial expression, body and tail posture and by howling, yelping or barking. The limbs are long and slender and adapted for swift running. There are four functional digits on each limb. On the fore-limb in all species except Lycaon pictus the first digit is vestigial and is represented by a claw and small pad higher up on the foot than the four functional pads. In L. pictus this digit is totally absent. In the domestic dog and dingo a vestigial first digit may also be present as a 'dew claw' on the hind limb as well. This may be occasionally observed in wild canids (see, for example, Lonnberg, 1916, for a report of 'dew claws' on a wild fox). The facial part of the skull in the Canidae is elongated, the zygomatic arches are wide and the bony orbits never form a complete ring. The temporal ridges may be either wide and enclose a lyriform sagittal area or they may be fused to form an interparietal crest. The 'brows', that is, the part of the frontal bones above and between the eyes, may be slightly dished, as in the foxes, flat as in most of the species of Dusicyon or convex as in the dog and wolf. The development of the 'brows' is a reflection of the size of the frontal sinuses. The postorbital process of the frontal bone usually ends in a small point. The auditory bulla is relatively large, rounded and divided internally by an incomplete septum. 3142 The dental formula for all species is, I -, C -, P -, M -, with the following excep- 3143 3 J 4 3 3I4I tions : Otocyon megalotis -I-, C-, P -, M-; Speothos venaticus - I -, C -, P -, M - ; 3144 3142 Cuon alpinus - I -, C -, P -, M -. In every species except 0. megalotis P4 and Mj (the carnassials) are larger than all the other teeth ; they are trenchant and bite together with a shearing action. In Otocyon, however, these teeth are molariform and no larger than the rest. In all canids the canine teeth are long and more or less sharply pointed ; the premolars are also pointed and have one main cusp. The molars, with the exception of the lower carnassial (Mj), are bunodont. In the majority of species the talonid or heel of the lower carnassial has two cusps, but in 5. venaticus, C. alpinus and L. pictus it is crested and has only one cusp. The homologies and development of the cusp patterns in the Canidae were discussed by Marett Tims (1896) but it is a subject that has since received little attention. In general, the canids being highly cursorial, the family is not found in dense forest areas; three exceptions are C. alpinus (Oriental region), 5. venaticus and Dusicyon microtis (both of the Brazilian subregion of South America) . All the races of wolf and almost all the species of true fox (genus Vulpes] are found in the northern hemisphere. The coyote, Canis latrans, of North America is replaced in southern Europe, Africa and the Orient by the several species of jackal. In Africa the ecological niche of the wolf is taken by L. pictus, the hunting dog. THE FAMILY CANIDAE SOURCES OF DATA 123 Selection of species The 37 species of wild canids are listed below, named and arranged according to what can be considered a consensus of current views on their classification. The , arrangement of subfamilies and genera follows Simpson (1945) except that Cerdocyon and Atelocynus are given generic rank following Cabrera (1958). The species are delimited according to the most recent major regional works, namely those of Ellerman & Morrison-Scott (1966) for the Palaearctic and Oriental regions, Ellerman et al. (1953) for southern Africa, Hall & Kelson (1959) for North America and Cabrera (1958) for South America. Cam's simensis, from Ethiopia, is the only species not included in these and its specific distinctness has never been in doubt. All of these species were used in this study with two exceptions, marked *, of which no specimens were available in the British Museum (Natural History). These are Urocyon littoralis, the grey fox from the Santa Barbara Islands, California, treated as speci- fically distinct from the continental Urocyon cinereoargenteus by Hall & Kelson, and Canis rufus (= C. niger), the red wolf of southeastern U.S.A. Both of these appear to be sufficiently similar to their better known relatives, U. cinereoargenteus and Canis lupus respectively, that their generic allocations can be presumed to follow those of the larger species. The generic name Oreocyon, replaced by Dasycyon (sic), has been proposed for a new species of canid, Dasycyon hagenbecki, based on a single skin from the Andes of South America (see Krumbiegel, 1953). The existence of this species has not been corroborated by further finds and we have not included it in this study. In addition to these wild species two forms of domestic dog were included, the dingo of Australia to represent a primitive breed and the bloodhound as an example of a highly differentiated breed. Present classification of the family Canidae : Subfamily CANINAE Canis lupus Wolf Canis rufus Red wolf* Canis latrans Coyote Canis aureus Golden jackal Canis mesomelas Black-backed jackal Canis adustus Side-striped jackal Canis simensis Ethiopian jackal Alopex lagopus Arctic fox Vulpes vulpes Common or red fox Vulpes corsac Corsac fox Vulpes ferrilata Tibetan sand fox Vulpes bengalensis Bengal fox Vulpes cana Blanford's fox Vulpes rueppelli Sand fox Vulpes pallida Pale fox Vulpes chama Cape fox Vulpes velox Kit fox Europe, Asia, N. America, Arctic Central N. America N. America S.E. Europe, N. Africa, S. Asia Africa south of the Sahara Africa south of the Sahara Mountains of Ethiopia Arctic Europe, N. Africa, Asia, N. America Central Asia Tibetan plateau India S.W. Asia N. Africa, S.W. Asia Southern edge of Sahara S. Africa N. America 124 J- CLUTTON-BROCK ET AL. Fennecus zerda Fennec fox Urocyon cinereoargenteus Grey fox Urocyon littoralis* Island grey fox Nyctereutes procyonoides Raccoon dog Dusicyon australis Falkland Island wolf Dusicyon culpaeus Colpeo fox Dusicyon culpaeolus Dusicyon gymnocercus Azara's fox Dusicyon inca Dusicyon griseus Argentine grey fox Dusicyon fulvipes Chiloe fox Dusicyon sechurae Sechura desert fox Dusicyon vetulus Hoary fox Cerdocyon thous Common zorro Atelocynus microtis Small-eared zorro Chrysocyon brachyurus Maned wolf Subfamily SIMOCYONINAE Speothos venaticus Bush dog Cuon alpinus Dhole Lycaon pictus Hunting dog Subfamily OTOCYONINAE Otocyon megalotis Bat-eared fox N. Africa, Arabia N. America, northern S. America Islands off California E. Asia Falkland Is., extinct since c. 1880 S. America - Patagonian subregion Uruguay Eastern Patagonian subregion Mountains of Peru S.W. Patagonian subregion Island of Chiloe N.W. Peru, Ecuador Brazil S. America - Brazilian subregion Central S. America - Brazil Southern Brazilian subregion S. America - Brazilian subregion E. and Central Asia Africa south of the Sahara Africa south of the Sahara Derivation of data The data relating to each species were derived primarily from specimens in the collections of the British Museum (Natural History), supplemented by information from the literature. Whenever possible, three skulls and skeletons of each species were selected and included one male and one female. All measurements were taken with dial calipers. The skeletal measurements are defined in the figures accom- panying Tables 4, 5, and 8. A character that has been used in diagnosing the genera of canids is the relative development of the frontal sinuses in the skull. In order that this character could be assessed correctly at least one skull from each genus was sectioned diagonally through the cranium to expose the sinuses. For characters of the pelage all the skins of most of the species were examined and scoring was done on between 3 and 10 skins. In defining the pelage characters account was taken of the genetics of coat colour as described by Little (1957). Assessments of hair and skin colours were made by eye and estimates of the thickness of the hair were made by rubbing one hair at a time between the thumb and fore- finger. Characters concerning internal anatomy, body proportions of the live animal, and comparative behaviour were extracted from published works and were scored in the THE FAMILY CANIDAE 125 same way as the directly measured variables. The sources of these data are given with the descriptions of the characters in Tables 4-9. NATURE OF THE DATA Kinds of characters The list of characters given in Tables 4-9 (p. 182) includes qualitative, quantita- tive and derived characters. Qualitative. These are characters whose values are simply alternatives from a list. Comparisons of magnitude between the different values are meaningless. Quantitative. There are two main types of quantitative character. The first takes values on an ordinal scale for which comparisons of order are possible. For example character 7, Table 7, dark patch on dorsal surface of tail, takes the values absent /short/ long and short is intermediate between absent and long. The second takes values on a scale for which differences and ratios of values may be compared. All the linear measurements on the skull fall into this category. Derived. These are characters whose values are derived from the actual observations on the specimen. The condylo-basal length of the skull was used as the best avail- able measure of overall size. Since the range of size in the family is very considerable, all other linear measurements were used in the form of ratios, frequently to condylo- basal length. The attempt to eliminate size-dependence by using characters derived as ratios can only be partially successful, but at least the derived characters are measuring aspects of shape which are far less size-related than the original characters. All the character values were obtained separately for each specimen. For quantitative characters these values usually varied within a species and a value for the species was obtained by averaging the values for the specimens. For characters taking values on an ordinal scale the coded values (such as I, 2, 3) were averaged. This is not ideal because it presumes that 2 is exactly halfway between i and 3 whereas it is only known that 2 is intermediate between i and 3. Because the values of these characters did not vary much in this study the difficulty was ignored. The values of qualitative characters showed no variation so the common value was used for the species. Use of the data matrix The characters employed have been presented in Tables 4-9 in a form suitable for general-purpose reference. There are, however, limitations on their use that must be stressed. In particular the mean values given for the quantitative charac- ters should not be used for further statistical studies without taking full account of the very small sample size (usually three). In the context of the analysis presented here, using 90 characters, it is thought that the errors inherent in these mean values are not important, especially where the range of values for a given character is 126 J. CLUTTON-BROCK ET AL. great, e.g. condylo-basal length of the skull for which the mean value for a species varies from 82 to 226 mm. It would, however, be quite inappropriate to use these figures as a precise measure of the difference in value between closely similar species, especially in the case of those species such as Canis lupus and Vulpes vulpes that have enormous ranges and considerable geographical variation. For the same reason elaborate and more accurate methods of measuring characters of the pelage were considered to be inappropriate. The main value of the data matrix is, we believe, in showing how the characters of a particular species relate to the variation found in the family as a whole, rather than as a basis for the detailed comparison of closely related species. MEASUREMENT OF SIMILARITY A measure of similarity between each pair of species was obtained by first allowing both qualitative and quantitative characters to contribute amounts between o and 100, and then averaging the contributions over characters. The rules were : (i) a qualitative character contributed 100 if the two species had the same value and o otherwise, regardless of whether the value represented the 'presence' or the 'absence' of something ; (ii) a quantitative character contributed an amount proportional to the differ- ence in the character value for the two species ; the proportion was chosen so that the largest difference between any pair of species in the study contributed 100 and a zero difference contributed o ; (iii) if a character was recorded as missing on a species because its value was not known then that character was ignored when assessing similarity of all other species with that species. This way of measuring similarity is the method used in the CLASP package of programs for numerical taxonomy (Rothamstead Experimental Station) and has been discussed in detail by Gower (1971). A number of minor variations in the method are included in the package. All analyses of similarity values were carried out twice : once using all the charac- ters listed in Tables 4-9 (referred to as 'All characters') and again using only charac- ters of the skull and teeth (Tables 4 and 5), these being the characters that traditionally have been given greatest weight in mammalian classification. NUMERICAL RESULTS Distribution of similarity The result of the process of selecting characters, observing their values and measuring similarity is a set of similarities on a scale o-ioo consisting of one for each pair of species. In this study there were 37 species and 666 similarities. Frequency distributions of similarities are shown in Fig. I and indicate the range of THE FAMILY CANIDAE 127 10 8 3 CJ 0) flj O OC 0 36 44 52 60 68 76 Similarity 84 92 100 10r ~ 8 o c o> 3 CJ (1) 0) - ro 2 0) OC 0L 36 44 52 60 68 76 84 92 S imilarity FIG. i. Relative frequency distribution of the similarity values : a. (above) all characters ; b. (below) skull and teeth only. values observed. Such distributions are useful for assessing which values of simi- larity correspond to 'high' and 'low'. The distribution based on the cranial and dental characters is similar to that based on the full list but has a longer left tail with some similarity values as low as 36. 128 J. CLUTTON-BROCK ET AL. Near neighbours A set of 666 similarities is far too large to scan by eye. Some method of arranging them is necessary and the simplest possible method is to list, for each species, the near neighbours of that species. The nearest neighbour to a species A is the species with highest similarity to A and the near-neighbour list which was used consisted of the five closest species to each species in turn, in order of similarity. Thus the set of 666 similarities is replaced by a subset of 37 x 5 = 185 similarities and the subset is far more readily scanned than the full set. This is due both to the reduction in the number of similarities and to the ordering in terms of similarity with each species separately. These near-neighbour lists are extremely useful when considering the taxonomic status of each individual species and for this reason they have been included in the systematic account of each species rather than given as a separate table. Two-dimensional plots A good overall view of the similarities is obtained from a plot in which the points represent species and the distances between points represent taxonomic distance. This distance must be defined in terms of similarity and a mathematically convenient definition is to set the squared taxonomic distance equal to 2 (100 — similarity) so that taxonomic distance itself is the square root of this quantity. It follows that a similarity of 100 corresponds to zero taxonomic distance and a similarity of o corresponds to a distance of V200- The total set of such taxonomic distances may not be exactly reproducible in a plane, for three points must obey the law that the distance round two sides of a triangle is greater than the distance along the third side. If they do not, a plot is found in which the geometric distances are as close as possible to the taxonomic distances. Major groupings are usually faithfully reproduced but some taxonomic distances can be rather distorted. In this study all conclusions from plots were checked against the original list of similarities. Fig. 2a, b shows two-dimensional plots which were prepared using the principal co-ordinates algorithm (Gower, 1966) . These figures demonstrate the remote position of some of the monospecific genera (Speothos, Lycaon, Cuon, Otocyon). Within the main group the species of Canis are well separated from those of Vulpes with Dusicyon occupying an intermediate position. Fig. 2b, based on skull and teeth, suggests a close relationship between Lycaon, Cuon and Speothos (currently forming the subfamily Simocyoninae) but when all characters are considered (Fig. 2a) Cuon is less closely associated with this group. The close association of Speothos and Lycaon in this figure is however spurious and provides a good example of the kind of distortion that can arise in this kind of plot. The taxonomic distance between them is V(2 x 32) = 8-0 whereas the distance on the plot is only 0-8. On the other hand, the taxonomic distance between Speothos and Vulpes bengalensis is \/(z x 41) = 9-0 and the distance on the plot is 7-5. The relationship between Speothos (and Lycaon) and the Caninae is generally well represented but not the relationship between Speothos and Lycaon. The distortion could be slightly reduced by adding a third dimension, but the improvement is bought at the cost of a far more cumbersome diagram (the so-called 'plumber's diagram'). *Sp ven ' L. p ictus 9 Ny. proc • Al. lag 9V. vu/pes F. zerda * 9Cu. alp V. ruep ' Udlld V. pall 9 V.chama • Ch. brach V. corsac* 9 9V V. ve/ox bang 9 D. vet 9 D. gris . D aust V ferr • D. culp 9 A. D.sech9 9 • D.futv hound* Ce.thous9 9 9D.gym _. .-. . C. simen • D culpaeo 9 C. lupUS ^ 9 rj /nca C. latrans A „ • C. adust 9 C. aureus z „ • C. meso -5 -A -3 -2 -1 0 1 2 3 A First dimension • Of. meg 9Sp. ven 9 Cu. alp 9 (Jr. ciner • Lpictus 9Ny.proc 9 • At. lag 9Vpa// F. zerda 9D.vet V.rueP9 .Vcana • • V. chama V. corsac 90. aust D.sech . 9V. ve/ox • hound 9C aureus v vu/pes C. lupus 9 •dm9° • Ce. thous C. latrans 9 At. mic C. meso D. culpaeo V. ferr D. fulv • Q. gris nic 9 D. gym • Ch. brach 9 D. inca C. adust 9 1 1 ' • D. culp • C. simen < j 5 -A -3 -2 -1 0 1 i 3 4 First dimension FIG. 2. Two-dimensional plot of all 37 species using the principal co-ordinates algorithm : a. (above) all characters ; b. (below) skull and teeth only. 30 £20 '•o -10 -20 • D.aust V vulpes 1 V cana • V. ferr V. corsac 9V. ruep , V. ve/ox • V. beng • V. chama • D. cu/paeo , V. pa/1 • D. gris D.vet* D.sech» D* 9D.cu/p gym • D. inca 40r | 30 5 s LO 10 -10 -20 -30L • D.vet D. sech • D.aust J L -20 -10 0 10 20 30 First dimension • V. cana • V. corsac V. ferr • V. pall 1 V. chama • V.beng •* ruep • V. ve/ox tD.fu/v V. vulpes • D gris •D. gym • D. cu/paeo D. inca • D. culp -30 -20 -10 0 10 20 30 First dimension 40 FIG. 3. Two-dimensional plot of members of Vulpes and Dusicyon using the principal co-ordinates algorithm : a. (above) all characters ; b. (below) skull and teeth only. 30 c - 20 tn ^u c o -10 -20 -30 dingo hound • D. aust ' C. lupus i C. aureus • C. latrans D. sech I D.vet • D. fulv D. inca 1 D. gym • D. gris C. simen •D. culpaeo • D. culp C. adust C. meso 30 20 10 10 20 30 40 First dimension 40r o 30 20 10 -10 -201- L • D. ausf • C. aureus hound • •dingo • C. lupus • D.vet D. sec/? • C. meso . fulv • D. gym C. adust C. latrans D. culpaeo • C. simen J_ _L • D. />?ca X • D. culp _L . pr/s -50 -40 -30 -20 -10 0 10 20 30 40 First dimension FIG. 4. Two-dimensional plot of members of Cam's and Dusicyon using the principal co-ordinates algorithm : a. (above) all characters ; b. (below) skull and teeth only. 30 20 T3 E O O 10 -10 -20 -30 • Ce. thous i At. mic • D. inca C. aureus • C. simen • C. adust • C. meso • C. latrans • D. aust • D. fulv D. gym *»D.cu/p • D. gris • D. vet • D. culpaeo • V. ferr v. chama hound • C. lupus din9° • (Jr. ciner V. pa/1 • F. zerda V. beng V. corsac • V. velox V. cana V. ruep • Al. lag • V. vu/pes -40 -30 -20 -10 0 10 20 First dimension 30 | 30 20 10 -10 -20 • D. culp D. inca • • c. simen • C. adust D. culpaeo » • C. meso ' D. gym D. gris • At. mic • D. fulv • Ce. thous • C. latrans • V. ferr i D. sec/? • V. vulpes _ V. velox C. aureus D. aust • V. beng • V. ruep (Jr. C/'ner • _ p •V. chama • V. cana *h • dingo • C. lupus hound •Al.lag v corsac •D.vet V. pall 40 30 20 10 10 20 40 First dimension 50 FIG. 5. Two-dimensional plot of members of the subfamily Caninae using the principal co-ordinates algorithm : a. (above) all characters ; b. (below) skull and teeth only. THE FAMILY CANIDAE 133 A better approach is to concentrate on different parts of the main plot and prepare separate plots for these parts. In this study we were interested in looking more closely at the overlap between the genera Vulpes and Dusicyon (Fig. 3a, b) and between Canis and Dusicyon (Fig. 4a, b). In Fig. 3a, using all characters, there is not much overlap between Vulpes and Dusicyon ; V. pallida is the most Dusicyon- like of the Vulpes (confirmed by its nearest neighbours according to similarity) and V. vulpes and V. ferrilata are rather atypical foxes ; the position of D. australis suggests a low similarity with both Vulpes and Dusicyon. For cranial and dental characters only (Fig. 3b) the picture does not change much although D. sechurae and D. vetulus move up closer to D. australis and V. vulpes appears more fox-like. In Fig. 4 the distinction between the genera Canis and Dusicyon is less clear than between Vulpes and Dusicyon ; the position of D. australis suggests a higher simi- larity with members of Canis than with Dusicyon, and C, simensis, C. adustus and C. mesolomas are all closer to the Dusicyon group than to other members of Canis. The same situation occurs in a more acute form using cranial and dental characters only. The other aspect of Fig. 2 that is worth studying more carefully is the position of the less distinctive monotypic genera in relation to the large genera. These are shown in Fig. 5a, b. Using all characters there is a strong case for including Urocyon and Fennecus in Vulpes, and Atelocynus and Cerdocyon in Dusicyon. An additional point of interest from this figure (and Fig. 4b) is the grouping of the bloodhound, the dingo and C. lupus, with D. australis not far away. These points will be discussed in more detail in the systematic section of the paper. Similarity values for the existing classification The existing classification is displayed graphically in Fig. 6. The ranks of the taxa are species, genus, subfamily and family. A species such as Otocyon megalotis simply changes its rank as it becomes a monotypic genus and then a monotypic subfamily. If a numerical value is associated with each rank the figure becomes a dendrogram and a useful way of studying the existing classification is to construct the dendrogram based on mean similarity between species. The ranks are given numerical values as follows (similarities based on all characters). Family - mean similarity between pairs of species, each member of the pair being from a different subfamily. There are 33x3 + 33x1 + 3x1 = 135 such com- binations and the mean of the 135 similarities is 65-0. Subfamily - mean similarity between pairs of species, each member of the pair being from a different genus, but the same subfamily. The number of such combinations is 431 and the mean similarity is 79-8. Genus - mean similarity between pairs of species, each member of the pair being from a different species but the same genus. The number of such combinations is 100 (monotypic genera contribute no pairs) and the mean similarity is 87-3. Species - this is given the value 100 which would be consistent with the way values have been given to genus if the specimens within a species were identical. In fact they were not, so the correct level for species should be rather less than 100. 134 J. GLUTTON-BROCK ET AL. 60 70 80 90 100L CANIDAE Caninae Simocyoninae Family Otocyoninae Subfamily Genus Species of Vulpes Species of Canis Species of Dusicyon Species of monotypic genera Species FIG. 6. Dendrogram of the existing classification : rank level equal to mean similarity between species. To see how well this dendrogram fits the data it is necessary to examine the distribution of similarity values that go to make up each of the means described above. These are shown in Fig. 7 in the form of cumulative frequency rather than frequency distribution because the former are easier to compare. Ideally there should be little or no overlap between the ranges of values at different ranks ; such a situation would indicate a very strong hierarchic structure in the similarities. In this case the overlap is rather large, particularly between subfamily and genus, which is to say that there are too many high similarities between species from different genera within the same subfamily. It is clear from Figs 3 and 4 that there is not much one can do about this. Even if D. australis were to be placed in Canis and if C. simensis, C. mesomelas and C. adustus were to be placed in Dusicyon the hierarchic structure would still be rather weak. The overlap between the range of similarity at family and subfamily level is not so high because of the low similarity of Otocyon megalotis and the three members of the Simocyoninae with all other canids. To enable impressions from the two-dimensional plots to be checked against actual similarity values a table of mean similarities between and within genera was pre- pared (Table i). The mean similarities of Fennecus and Urocyon with members of Vulpes are in bold print, as are those of Atelocynus and Cerdocyon with members of Dusicyon. Apart from some distortion in Fig. 2 the plots are in good agreement with the table of mean similarities. (a) All characters THE FAMILY CANIDAE 100 r Crf(%) 135 Similarity (b) Skull and teeth Similarity FIG. 7. Cumulative relative frequencies (Crf) of similarity values corresponding to each rank for the existing classification : a. all characters ; b. skull and teeth only. TABLE i Mean similarities between and within genera of the existing classification (a) All characters 13 3 Vulpes Canis Dusicyon Alopex Fennecus Urocyon i i 86-9 2 78-0 3 86-0 4 79-2 5 85-1 6 85-0 2 74-2 72-0 74-8 3 90-5 79-0 83-6 4 78-2 74'4 5 82-5 6 * 7 8 9 IO II 12 Nyctereutes 7 70-6 70-9 73'3 72-2 68-4 73'3 * Atelocynus 8 75-7 79'3 82-2 69-6 72-4 79-3 78-1 * Cerdocyon 9 78-0 79'4 84-8 72-9 76-2 82-1 76-8 86-1 * ' Chrysocyon 10 67-1 69-4 71-4 68-9 597 64-9 673 73'4 68-6 * Speothos ii 58-6 61-9 63-3 61-1 53'1 61-5 65-9 68-2 60-0 53-8 * Cuon 12 7O-O 75'5 75-6 74'7 69-3 71-0 71-9 70-4 69-9 65-3 73-5 * Lycaon 13 57-6 62-9 61-4 61-1 5i'9 55-6 57'1 56-6 S3'2 50-0 67-9 69-7 Otocyon I4 70-7 63-2 68-3 69-8 71-6 72-6 70-8 67-0 64-1 62-6 51-5 59'5 (b) Skull and teeth only Vulpes Canis Dusicyon Alopex Fennecus Urocyon Nyctereutes i i 87-6 2 76-6 3 82-9 4 84-1 5 83-0 6 81-4 7 78-0 2 8i-5 79'2 70-8 80-0 3 80-8 74'9 76-2 78-7 4 * 77-0 80-1 82-9 5 * 77'2 68-4 6 82-2 7 8 9 IO II 12 Atelocynus 8 81-5 82-2 85-6 78-9 7I-3 81-0 82-0 * Cerdocyon 9 79'6 77-8 826 76-0 75-3 79-2 80-5 85-6 * Chrysocyon 10 75-1 83-2 79-7 76-0 637 73'3 74-8 82-7 75-2 * Speothos ii 61-7 66-1 65-1 71-3 50-0 59'2 74-1 62-5 58-6 57-3 * Cuon 12 62-O 70-6 65-3 71-9 50-5 58-5 72-2 63-3 57'6 63-9 87-0 * Lycaon 13 58-3 73'2 62-8 68-5 45-5 51-5 68-1 62-9 56-5 63-3 73'7 80-9 Otocyon 14 62-7 52-2 56-9 63-8 59'6 75'9 66-0 61-5 61-4 58-4 43-0 41-7 13 14 36-6 136 J. CLUTTON-BROCK ET AL. Homogeneity of the three main genera To study the effects of the marginal species on the homogeneity of the three main genera the members were listed in order of 'typicality', defined as the mean similarity of a species with all other members of the same genus. This was done both for the existing classification and for a revised classification in which the marginal fox genera, Alopex, Fennecus and Urocyon, are included in Vulpes, and Atelocynus and Cerdocyon in Dusicyon (see Tables 2 and 3). There are several interesting features of these lists. For the existing classification, the typicalities of V. vulpes and D. australis are relatively low. There is a high degree of concordance between the lists based on all characters and those based on cranial and dental characters only. For the revised classification the new arrivals mingle with the others in a gradual way, i.e. there is no sudden drop in typicality, except for the low typicality of Alopex, suggesting that it is best left out of Vulpes. The homogeneity is worst for the genus Dusicyon where similarity is based on all characters, but the new range of similarity is more in line with that for Vulpes and Canis. TABLE 2 List of members of Vulpes, Canis and Dusicyon (existing classification) in order of typicality. The measure of typicality is shown next to each species (a) All characters Vulpes V. bengalensis V. velox V. chama V. corsac V. rueppelli V. pallida V. ferrilata V. vulpes V. cana (b) Skull and teeth only Vulpes 90-5 V. chama 90-5 89-7 V. velox 90-1 89-2 V. bengalensis 89-9 88-6 V. corsac 88-1 88-2 V. pallida 88-0 86-0 V. rueppelli 87-4 84-6 V. vulpes 87-2 83-4 V. cana 84-8 82-4 V. fewilata 82-6 Canis Canis C. aureus C. latrans C. mesomelas C. adustus C. lupus Dingo C. simensis Hound Dusicyon D. gymnocercus D. culpaeolus D. fulvipes D. griseus D. sechurae D. inca D. culpaeus D. vetulus D. australis 86-8 C. latrans 85-6 Dingo 84-8 C. aureus 84-2 C. lupus 83-0 C. adustus 82-5 C. mesomelas 82-3 Hound 82-2 C. simensis Dusicyon 93'2 D. gymnocercus 91-8 D. culpaeolus 9i-3 D. fulvipes 91-1 D. sechurae 90-8 D. griseus 90-4 D. inca 90-0 D. australis 89-3 D. culpaeus 86-2 D. vetulus 88-7 87-8 87-3 86-0 85-8 85-7 83-6 81-9 87-5 86-4 85-3 85-1 83-3 82-7 82-5 80-4 THE FAMILY CANIDAE TABLE 3 137 List of members of Vulpes plus Alopex lagopus, Fennecus zerda and Urocyon cinereoargenteus; also members of Dusicyon plus Atelocynus microtis and Cerdocyon thous. Both lists in order of typicality with the measure of typicality shown next to each species (a) All characters Vulpes V. bengalensis V. chama V. velox V. corsac V. rueppelli V. pallida F. zerda U. cinereoargenteus V. ferrilata V. vulpes V. cana A. lagopus Dusicyon D. gymnocercus D. fulvipes D. culpaeolus D. griseus D. sechurae D. inca D. culpaeus D. vetulus D. australis C. thous A. microtis (b) Skull and teeth only Vulpes 89-2 V. chama 89-1 88-3 V. bengalensis 88-7 88-3 V. velox 88-3 87-4 V. pallida 87-1 87-1 V. corsac 87-0 86-9 V. rueppelli 86-8 84-2 V. vulpes 86-2 83-8 V. cana 83-8 83-1 A. lagopus 83-1 82-0 F. zerda 81-9 81-4 U. cinereoargenteus 80-9 78-7 V. ferrilata 79-8 91-5 90-6 90-2 89-6 89-4 89-1 88-3 87-9 85-0 85-0 82-6 Dusicyon D. gymnocercus D. culpaeolus D. fulvipes A. microtis D. sechurae D. griseus D. inca C. thous D. australis D. culpaeus D. vetulus 87-4 86-7 85-6 85-3 84-9 83-0 82-9 82-7 81-9 80- 1 Automatic classification A natural question to ask at this stage is what happens if the 37 species are grouped using the 666 similarities with some standard linkage algorithm? The choice of linkage algorithm is never easy (see Sneath & Sokal, 1973, for a full dis- cussion) ; several were tried and an average linkage method, equivalent to the way the dendrogram of Fig. 6 was constructed, was chosen : two groups merge at level S if the mean similarity between pairs of species, one from each group, is greater than or equal to S. A sensible name for this algorithm would be 'weighted average linkage' because the mean similarity between groups takes account of the sizes of the groups. Unfortunately 'weighted' has other connotations in taxonomy and so the term 'centroid linkage' is used instead. The reader should be warned that the terminology in Sneath & Sokal is different (1973 : 235). The results of the algorithm are shown in Fig. 8a, b. For both 'all characters' and 'skull and teeth only' the dendrograms retain the more homogeneous parts of the main groups. At a higher level some odd combinations occur, e.g. D. australis with 138 J. CLUTTON-BROCK ET AL. 100 90 80 70 - t J 1 t C. lupus C. latrans — | _l C. meso C. adust nJ V. ve/ox V. beng 1 V. corsac V. ruep V. pall V. chama O. sech [ D. vet 1 D. culpaeo — [ D. gym -J ^ > — ] D. fulv D. gris 1 V ferr - D aust - — | Cf (hot/1; hound dingo Al lag 100 90 80 70 60 i 1 — i — i — r— T i V. vulpes V. ve/ox — — — — V. beng 1 V. pall -, \— V. chama — ' V. ruep ' Al. lag V. corsac V. cana F. zerd h H h h- i D. sech 1 h D. vet ' C meso 1 C. adust 1 D. culpaeo — i D. gym — ' h h b] /I- C. simen Ch. brach D. fulv 1 D. gris ' U- ~n C. lupus — i dingo — ' C. aureus D. aust H — Sp. ven „ , ~>- FIG. 8. Centroid linkage dendrogram : a. (left) all characters ; b. (right) skull and teeth only. V. ferrilata which then join with the Vulpes and Dusicyon groups. Such oddities are common in average linkage dendrograms because small changes in similarity can cause large changes in the dendrogram. Another disadvantage is that the dendrogram as such gives no indication of how well a hierarchy fits the data ; further analysis along the lines followed for the existing classification would be necessary. In general, the two-dimensional plots provided a much better way of looking at the relationships between species than the dendrograms from automatic classification algorithms. THE FAMILY CANIDAE 139 GENERAL TAXONOMIC CONCLUSIONS Before drawing any general taxonomic conclusions from this analysis of phenetic relationships it is necessary to consider any other sources of relevant data that were not taken into account in the numerical analysis. Additional evidence might, for example, be forthcoming from the fossil record of the family, from studies of kary- ology, from immunology, from parasitology or from the results of attempted inter- breeding, although it was believed that none of these categories of data could usefully be incorporated into the numerical analysis. In practice the data from these additional sources proved to be so scanty that they contribute almost nothing to the problems of classification above the species level. The fossil history of the Canidae has been discussed by Matthew (1930), Romer (1955), Radinsky (1973) and Todd (1970). Data on chromosome numbers are summarized by Chiarelli (1975) who also tabulated the authorities for intergeneric hybrids. Further information on interspecific interbreeding was listed by Gray (1972). The species of lice that have been found on members of the canid family are presented in the Appendix (p. 194). The general conclusion that emerges from the various forms of numerical analysis is that the majority of species, including most species of Canis, Dusicyon and Vulpes, form a large cluster with complex interrelationships and no major discontinuities, surrounded by a number of more or less distinctive and isolated species. To consider these isolated species first, the most distinctive is undoubtedly Lycaon pictus whose similarity to its 'nearest neighbour' is only 71. Next in order of separation come Speothos venaticus (73), Otocyon megalotis (76), Nyctereutes procyonoides (78), Cuon alpinus (79), Chrysocyon brachyurus (So) and Alopex lagopus (82). These are currently considered to represent monotypic genera except that the last (A . lagopus) is sometimes included in Vulpes. All the other species that have at one time or another been considered distinctive enough to warrant generic separation have 'nearest-neighbour' values of over 85 and are therefore no more distinctive than many species that are consistently classified within the large genera. We can therefore conclude that the species detailed above should continue to rank as monospecific genera, namely Lycaon, Speothos, Otocyon, Nyctereutes, Cuon, Chrysocyon and Alopex. The last of these is the least distinctive, and the skull and teeth show a very high degree of similarity to certain species of Vulpes, especially V. corsac. On the basis of all characters, however, it is clearly the most distinctive of the foxes, as shown for example by the low 'typicality' in Table 3, and there are therefore reasonable grounds for retaining it as a monospecific genus. We can now proceed to consider whether there is justification for grouping any of these distinctive genera at subfamily level. The only such grouping with any claim to consideration seems to be the one that is currently employed, namely the grouping of Lycaon, Cuon and Speothos as a subfamily, contrasting with the re- mainder. Although many of the similarities between these are higher than between them and other species, they are nevertheless very low, ranging from 73-5 between Cuon and Speothos to 68 between Lycaon and Speothos. In contrast Cuon has a mean similarity of 76 with species of Dusicyon. In addition it must be remembered that I4o J. CLUTTON-BROCK ET AL. many of the individual similarities are simply due to the shared absence of a special- ized character. The higher similarity of skull and teeth between Cuon and Speothos (87) might support such a grouping but in general it seems that the similarities between these three genera (or any two of them) are so tenuous that no useful purpose is served by uniting them within a subfamily. The case for subfamily rank of any other genus is equally tenuous, e.g. Otocyon has an overall similarity of 73 with Urocyon cinereoargenteus which is normal for intergeneric similarities (Table i). All the remaining species that have been given generic rank emerge from this analysis as an integral part of the main cluster of species and there appear to be no grounds for continuing to place any of them in monospecific genera. These are Fennecus and Urocyon which fall clearly within Vulpes ; and Cerdocyon and Atelo- cynus which fall so close to Dusicyon that it seems reasonable to include them (Figs i and 5, Table 3). The status of all these is considered in more detail in the systematic section. The question of the recognition of generic limits within the C anis/ Vulpes / Dusicyon complex is more difficult. No objective analysis of the results of this study would produce these three genera as presently composed but nor would it produce any other clear-cut grouping of species. On the other hand, the retention of these genera does not produce any serious anomalies and they are capable of definition. They are therefore retained here. Arising from a detailed study of the South Ameri- can species, Langguth (1975) proposes to recognize Cerdocyon thous, Atelocynus microtis and Lycalopex vetulus as additional, monotypic, 'differentiated' genera and to include the remaining species of Dusicyon in Canis. These proposals are not greatly at variance with our conclusions although the distinctiveness of these 'differentiated' species seems marginal, and if Dusicyon were merged with Canis it would be difficult to argue that Vulpes should not be treated likewise. This examination of the family Canidae as an integrated whole has enabled some anomalies in the taxonomy to be straightened out. It is hoped that it has also enabled some misconceptions to be erased. Perhaps the most notable of these is the belief that the common fox, Vulpes vulpes, is a typical representative of its genus. This belief has led to the classification of those species of fox that do not conform with V. vulpes in separate genera. In fact the typical fox is Vulpes bengal- ensis and V. vulpes should be considered almost as an aberrant species. When this fact is recognized the classification of the genus becomes more straight- forward. The extinct Falkland island wolf, Dusicyon australis, is seen to be a very anomalous species but lack of adequate specimens precludes any very clear assessment of its affinities. The revised classification proposed is presented below, and the relationships of each genus and species are considered in more detail in the systematic account that follows. In this list the 'species' of Dusicyon that are indented may be conspecific with the species listed above them, but this is a question that can only be answered by more detailed study of distribution and variation in South America. THE FAMILY CANIDAE 141 Proposed classification of Canis lupus (Canis rufus) Canis (domestic) Canis latrans Canis aureus Canis mesomelas Canis adustus Canis simensis Vulpes vulpes Vulpes corsac Vulpes ferrilata Vulpes bengalensis Vulpes cana Vulpes rueppelli Vulpes pallida Vulpes zerda Vulpes chama Vulpes velox Vulpes cinereoargenteus (Vulpes littoralis) Alopex lagopus Otocyon megalotis Nyctereutes procyonoides Dusicyon australis Dusicyon culpaeus Dusicyon culpaeolus Dusicyon gymnocercus Dusicyon inca Dusicyon griseus Dusicyon fulvipes Dusicyon sechurae Dusicyon vetulus Dusicyon thous Dusicyon microtis Chrysocyon brachyurus Speothos venaticus Cuon alpinus Lycaon pictus the family Canidae : Wolf (Red wolf) Dingo and domestic dogs Coyote Golden jackal Black-backed jackal Side-striped jackal Ethiopian jackal Common or red fox Corsac fox Tibetan sand fox Bengal fox Blanford's fox Sand fox Pale fox Fennec fox Cape fox Kit fox Grey fox (Island grey fox) Arctic fox Bat-eared fox Raccoon dog Falkland Island wolf - extinct since c. 1880 Colpeo fox Azara's fox Argentine grey fox Chiloe fox Sechura desert fox Hoary fox Common zorro Small-eared zorro Maned wolf Bush dog Dhole Hunting dog SYSTEMATIC ACCOUNT Genus CANIS L. Type species Canis familiaris L., the domestic dog. For the purposes of this analysis the genus has been taken as comprising six species plus two domesticated forms, the feral dingo as an example of a primitive domestic breed and the bloodhound which exemplifies advanced domestication without gross abnormalities. These two dogs have been treated as separate 'species' I42 J. GLUTTON-BROCK ET AL. on an equal level with the wild species. Although Canis familiaris is the type species for the genus the name has not been used in this work because we believe that formal zoological nomenclature should be avoided in naming domestic animals (see Groves, 1971). Recent classifications of the Canidae usually include the three groups, wolves, coyote and jackals, within the genus Canis and the results of this numerical taxonomy show that this is consistent with the phenetic relationships of the species. Heller (1914) separated the jackals from the wolves and coyote in the genus Thos Oken, and this classification was followed by Allen (1939). Heller defined the genus Thos as a group of Canidae having long slender Vulpes-like canines, small outer incisors, small carnassials, upper molar teeth with well-marked cingula and the fourth lower premolar with a minute extra cusp on its hinder border. He distinguished the genus Canis by the much thicker and shorter canines, greatly enlarged outer incisors, large carnassials, molars without a definite cingulum, and the fourth lower premolar without a third cusp on its posterior border. None of these characters is definitive and they are all very variable in their development. It is therefore more appropriate to include the jackals within the genus Canis. Allen (1939) also afforded separate generic status to the Ethiopian jackal, Canis simensis, placing it in the genus Simenia Gray. This somewhat aberrant canid appears from the results of the numerical taxonomy to be phenetically close to the genus Dusicyon but the skull has a high similarity with that of Canis adustus and we therefore include it with the jackal group in the genus Canis. There is no evidence to support the suggestion of Brink (1973) that C. adustus should be placed in a separate genus. The wolf is the largest species within the family and it is separated from the coyote and jackals by its more highly evolved social behaviour patterns that are closely reflected in its descendant, the domestic dog. The skull in all members of the genus has well-defined diagnostic features. It is always rather heavy and has large frontal sinuses. The temporal ridges are close together and are often fused into a strong interparietal crest. With the exception of Canis simensis, the facial region is relatively shorter than in the genera Vulpes or Dusicyon. The canine teeth are robust and the carnassials are relatively large. There is no subangular lobe to the mandible. The genus is very flexible in its habitat preferences, again with the exception of C. simensis which is confined to montane areas of Ethiopia, where it is nearly extinct. Most wild Canis species have wide distributions but they are not found in areas of dense tropical rain forest. The domestic dog has been taken to all parts of the world that are inhabited by man and it is presumed that the dingo was taken to Australia by man during prehistoric times. The wild members of the genus Canis are distributed throughout Europe, Asia, North America and Africa. Canis lupus L., 1758 Wolf DISTRIBUTION. Widespread in the northern hemisphere and as a species without particular habitat preferences. May be found in tundra regions, woodland, open THE FAMILY CANIDAE 143 plains or the edges of desert areas, from sea level to more than 3000 m. As their habitat has been restricted the wolf populations have been moved into progressively more inhospitable regions. Formerly widespread throughout the temperate areas of the northern hemisphere, but now extinct in western Europe except for small dwindling populations in Portugal, Spain, Italy, Sicily and Scandinavia. Still widely distributed in the U.S.S.R. and extending eastward into Mongolia, western China, Korea, Tibet and southwards into India. There are still small numbers in southwestern Asia. Widespread in Canada and Alaska but extinct in the U.S.A. except in wildlife parks. Classified as a vulnerable species by the Red data book (Goodwin & Holloway, 1972). DESCRIPTION. The largest member of the family. Body heavy with large head and long legs. Tail long and bushy. Ears erect. Fur very thick and with long guard hairs, especially in the arctic regions of the range. Pelage characters very variable - may be white, tawny, grey or black, but grey agouti with some tawny is the predominant colouring. Legs, ears, muzzle and underparts are usually reddish or pale. There is often a dark saddle and a dark patch over the tail gland. Highly social. Hunt large prey in packs of between two and eight individuals, although larger groups may be found in exceptional circumstances (Mech, 1970). Skull large with very well-developed interparietal crest. Massive jaws with powerful canine and carnassial teeth. Facial region long. Bullae large. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Cam's latrans 90-0 Cam's aureus 87-7 Bloodhound 84-0 Dusicyon culpaeolus 82-0 Cam's mesomelas 81-8 Skull and teeth only Dingo Bloodhound Cam's aureus Cam's latrans Dusicyon australis 94'7 92-4 89-2 89-1 87-5 It is of great interest that the numerical results show such a close phenetic simi- larity between Canis lupus, the dingo and the bloodhound, especially as this breed of dog bears little superficial resemblance to the wolf. This is discussed in the following sections on the dogs. The wolf is also closely related to the coyote, Canis latrans, and it may be mentioned here that there is evidence to suggest that Canis rufus Audubon & Bachman, 1851 (formerly known as Canis niger Bartram, 1791 - see Paradiso & Nowak, 1972), which has not been included in this analysis, is a composite species resulting from wolf -coyote hybrids (see Lawrence & Bossert, 1967, 1975 ; Mech, 1970 ; Paradiso, 1971 ; Gipson et al., 1974). REMARKS. Canis lupus (when not persecuted by man) is one of the most wide- spread and successful of large mammals. It is exceedingly variable in size, pelage and body proportions, but probably not in its behaviour patterns, over its wide range. For the purposes of this analysis four specimens were chosen for measure- ment and scoring of characters, the localities being selected to cover as much as possible of the range. The localities were Sweden, British Columbia, Spain and India. Indian wolves are considerably smaller than northern animals and have a shorter coat. I44 J. CLUTTON-BROCK ET AL. In order that a predator may kill its prey efficiently it must be either larger than or approximately the same size as the prey or it must hunt in a group and use a concerted effort to obtain its food. The wolf is adapted to feed on animals that are much larger than itself and the features that distinguish it from other canids are all integrated with this predator -prey relationship which has resulted in a highly evolved social system. The wolf pack is held together by strong social bonds and the suppression of aggression between individual members. So much work has been carried out in recent years on the social behaviour and signals of communication in the wolf and it is now so well known that it will not be repeated here (amongst others, see Crisler, 1959 ; Fox, 1970, 1971, 1975 ; Mech, 1970 ; Pulliainen, 1967 ; Schenkel, 1967 ; Scott, 1967 ; Woolpy & Ginsburg, 1967). It may be said, however, that the basic difference between the wolf and the other highly social canid, Lycaon pictus, is that the wolf pack is based on a hierarchical system (as in man) whereas in L. pictus the pack is held together by individual dominant and submissive relationships, with no established hierarchy and no highly evolved system of communication by facial and body signals (Fox, 1970 ; see p. 181). Canis (domestic) Dingo DISTRIBUTION. Open country throughout most of the continent of Australia except Tasmania. Absent from central New South Wales and the northern agri- cultural districts of western Australia (Ride, 1970). DESCRIPTION. A medium to large-sized dog. Usually a tawny-yellow colour but may show other colour variations including black. There is often a white tip to the tail and white feet. Of 15 skins examined in the British Museum only one had the first digit on the hind feet represented by a claw ('dew claw'). As noted by Lonnberg (1916), a vestigial first digit may be very occasionally present in wild canids but it is certainly exceptional. Mivart (1890, p. iv) stated that no wild species of canid ever has this first digit and we have not noticed any example of it, but it is relatively common in domestic dogs. The same dingo skin that had 'dew claws' (no. 25.8.1.9) had a slight dark patch on the dorsal side of the tail in the position of the tail gland. Hildebrand (i952b) quoted the belief that the tail gland is not found in the domestic dog, but further observation might well show that it can be present. Skull like that of a small wolf. Teeth large and evenly spaced, carnassials strongly developed. Bullae large but rather flatter than in the wolf. Frontal sinuses well developed. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Bloodhound 88-7 Canis lupus 94-7 Dusicyon australis 87-1 Canis latrans 91-7 Canis aureus 84-0 Canis aureus 91-5 Dusicyon inca 83-5 Dusicyon australis 91-1 Dusicyon sechurae 83-3 Bloodhound 90-9 THE FAMILY CANIDAE 145 The dingo is a fascinating relic of the primitive domestic dogs that must have been widespread in Asia during the early Holocene. It is not a biological species but a feral dog that is closely related to the New Guinea singing dogs and Indian pariah dogs. It is probable that these dogs are all directly descended from the Indian wolf, Canis lupus pallipes Sykes, 1831. This supposition is supported by the 'near neighbours' table for skull characters which shows a similarity of nearly 95 for the dingo and the wolf. The only other taxa in this analysis that are linked at this high level of similarity are Dusicyon gymnocercus with Dusicyon culpaeolus (which may well not be separate species) and Vulpes chama with Vulpes pallida. Corbett & Newsome (1975) have made a preliminary analysis of the social behaviour of the dingo in the wild. On the two-dimensional plots the dingo, wolf and bloodhound can be seen to be closely linked, and on the 'near neighbours' table for cranial and dental characters the dingo is linked with the 'typical' species, Canis aureus and Canis latrans, at a similarity of just under 92. The enigmatic position of Dusicyon australis, the extinct Falkland Island wolf, in association with the dingo and bloodhound is discussed in the section on that species (p. 166). Canis (domestic) Bloodhound ORIGIN. The bloodhound is probably descended from the French black and tan hounds that were bred for several hundred years at the St Hubert Monastery in the Ardennes. It has been established as a British breed since the Medieval period. DESCRIPTION. A pure-bred hound of ancient descent. Large, massively built, short-coated with long pendulous ears, a wrinkled face and long tail. May be black and tan, all tan or red and tan ; the skin that was used for this project was a mottled grey and tawny. The first digit on the hind foot (hallux) is sometimes developed as a 'dew claw' as is common in all domestic dogs. No black patch on the dorsal part of the tail in the position of the tail gland. Like all domestic dogs the bloodhound is a highly social animal. Skull typically 'hound-like'. Facial region long with heavy jaws. Teeth large and well-spaced but carnassial teeth relatively short. Postorbital processes of the frontal bones much swollen so that the profile of the skull has a marked 'stop'. This is a characteristic feature of advanced domestication in the dog and is associated with enlarged frontal sinuses. The reduced carnassial teeth and rather small flat bullae are also features of domestication. The interparietal crest is usually well- developed. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Dingo 88-7 Canis lupus 92-4 Dusicyon australis 85-9 Dingo 90-9 Canis lupus 84-0 Canis aureus 86-3 Canis aureus 83-5 Canis latrans 85-4 Canis latrans 81-1 Dusicyon australis 85-3 I46 J. CLUTTON-BROCK ET AL. It has been known for some years, from studies of social and agonistic behaviour, that the domestic dog is more likely to be descended from the wolf than from the coyote or the jackal (see, for example, Scott, 1967). It is most interesting that our numerical results, which included only a few behavioural characters out of the total of 90, so closely corroborate this deduction. Previous work on the osteological differences between the skulls of dogs and the wild Canis species has often failed to show clear distinction between the different groups. The present analysis shows that the skull of the bloodhound is phenetically closer to that of the wolf and the dingo than it is to the skulls of coyote or jackal. The two-dimensional plots and centroid linkage dendrogram also show the same relationships. It is surely rather remarkable that the dingo and the bloodhound, which bear so little superficial resemblance to each other and have such widely separated origins, should be so phenetically similar. The inference must be that they share a common ancestor in the wolf. Canis latrans Say, 1823 Coyote DISTRIBUTION. Widespread in North America. Formerly confined to areas west of the Mississippi river from southern Canada to central Mexico ; now ex- tending to Alaska and Costa Rica but still not very common in the eastern regions. The preferred habitat is open plains and desert and the coyote is not found in damp tropical areas (Hershkovitz, 1972 : 359 ; Miller & Kellogg, 1955 ; Van Wormer, 1964) . DESCRIPTION. 'Wolf-like' but smaller. An adult North American male wolf weighs an average of 45 kg, whereas the average weight of a male coyote is only 12 kg (Mech, 1970 ; Van Wormer, 1964). The coyote stands nearly as high at the shoulder as the wolf but it is much more lightly built with long slender legs, large ears and a bushy tail. Pelage characters are variable as with all canids that cover a wide geographical range. The coat is usually long and has an overall buffy-grey colour with long dark-banded guard hairs. Legs and sides may be fulvous. Under- parts and chin pale or nearly white. Tip of tail usually black. Not highly social but may live in family groups and take part in communal howling. Feeds on rodents, small game animals and birds. Size of skull between that of a small wolf and a jackal. The teeth, especially the canines and carnassials, well developed. Interparietal crest present but not so pronounced as in the wolf. Bullae rounded. Differences between the skulls of wolves, coyotes and domestic dogs have been analysed by Lawrence & Bossert (1967). SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Canis lupus 90-0 Canis aureus 89-7 Dusicyon culpaeolus 87-5 Canis mesomelas 87-4 Dusicyon culpaeus 87-2 Skull and teeth only Dingo 91-7 Canis mesomelas 90-8 Canis lupus 89-1 Canis aureus 88-9 Dusicyon australis 88-9 THE FAMILY CANIDAE 147 On phenetic characters the coyote lies between the wolf and the jackals. It is unlikely that the coyote has played any great part in the origins of the domestic dog. The American Indians may have crossed their dogs with coyotes from time to time but it is probable that this had only a local effect. Mengel (1971) has shown that gene flow from dogs to wild populations of Canis latrans (and also to Canis lupus] is unlikely to occur because of a phase shift in the breeding time of the hybrids. This prevents further reproduction after the first generation. An interesting aspect of Mengel's work was his demonstration that wild male coyotes are only fertile for about two months in the year whereas male domestic dogs are always in breeding condition. Canis aureus L., 1758 Golden jackal, Asiatic jackal DISTRIBUTION. Wooded and open country in the Balkan states, Romania, countries of the eastern Mediterranean, including Greece, Libya, Egypt and west- wards into Morocco. South to Senegal, the Sudan, Somalia, Ethiopia and Kenya. Eastwards through western Asia, the Middle East, Baluchistan and Sind. Through- out the peninsula of India to Ceylon and east to Assam, Burma and Thailand (Ellerman & Morrison-Scott, 1966). DESCRIPTION. Like the coyote, this jackal covers a very wide geographical area and it is very variable in size and pelage characters. The skins in the British Museum from localities in S.W. Asia and S.E. Europe were described in detail by Pocock (1938). In general the fur is rather coarse and not very long. The dorsal area is mottled black and grey. The head, ears, sides and limbs may be tawny or rufous, the underparts pale ginger or nearly white. Tail tip black. Feeds on small animals, carrion, insects and some fruit and vegetable matter. Not highly social but will hunt in packs. An exceptionally large subspecies, C.a. lupaster Hemprich & Ehrenberg, 1833, occurs in Egypt and Libya ; a specimen of this subspecies was included in the analysis. Skull like that of a very small wolf, with well-developed, high-crowned teeth. Interparietal crest present. Facial region somewhat short. There is often a well-marked cingulum on the labial side of M1. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Canis mesomelas 89-9 Canis latrans 89-7 Dusicyon australis 88-8 Canis adustus 88-7 Canis lupus 87-7 Skull and teeth only Dingo 91*5 Dusicyon australis 91-4 Canis lupus 89-1 Canis latrans 88-9 Canis mesomelas 88-3 For the all-characters similarity Canis aureus is the most typical of the Canis species (Table 2) and this is in fact obvious from one look at this jackal, for it is of I48 J. CLUTTON-BROCK ET AL. medium size and has no outstanding features. The pelage is typical of the family and the wide range that it covers precludes specialization. The skull of C. aureus is not as similar to the African jackals, Canis mesomelas, Canis adustus and Canis simensis, as it is to the dingo, wolf or coyote which is somewhat surprising. The position and relationships of Dusicyon australis are anomalous and are discussed in the section on that species. The behaviour of the golden jackals of the Ngorongoro Crater in Tanzania has been studied by H. & J. van Lawick-Goodall (1970), and detailed observations were made on a pair of jackals in Israel by Golani & Keller (1975). Canis mesomelas Schreber, 1778 Black-backed jackal DISTRIBUTION. Widespread in open savannah throughout eastern and southern Africa, northwards to the Sudan and west to Cameroun (Allen, 1939 ; Ellerman et al, 1953). DESCRIPTION. Distinguished by the dark saddle which extends the length of the back to the black tip of the tail. This saddle, which is mottled black and cream, contrasts strongly with the rufous sides of the body. The head and ears are also rufous flecked with white and dark hairs. The limbs are tawny or rufous, the underparts pale ginger. Underfur, except on the abdomen, consistently rufous, the colour of the saddle being due to banded black and white guard hairs. Lives alone or in small family groups. As with all species of Canis this jackal may par- ticipate in communal howling. Feeds on small prey and carrion. Skull smaller than that of Canis aureus. Parietal crest may be poorly developed and there may be a narrow lyriform sagittal area enclosed by weak temporal ridges. Dentition, especially the canines which are rather pointed, may resemble that in the genus Dusicyon. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Dusicyon culpaeolus 92-9 Dusicyon gymnocercus 91-5 Canis adustus 90-3 Canis aureus 89-9 Dusicyon inca 88-7 Skull and teeth only Canis adustus 92*7 Dusicyon australis 92-5 Dusicyon culpaeolus 91-9 Canis latrans 90-8 Dusicyon gymnocercus 90-7 The two-dimensional plots and centroid linkage dendrograms show that Canis mesomelas and Canis adustus are closely related and they have a similarity of 92 on cranial and dental characters. REMARKS. The black-backed jackal is looked upon as vermin in South Africa and it is persecuted by farmers because it kills sheep. THE FAMILY CANIDAE 149 Canis adustus Sundevall, 1846 Side-striped jackal DISTRIBUTION. Covers the same regions as Canis mesomelas, but prefers a heavier density of vegetation and wooded areas. Widespread in southern and eastern Africa. Northwards into the Sudan and Cameroun (Ellerman et al., DESCRIPTION. The pelage of this jackal differs considerably from that of C. mesomelas and it is a larger, heavier animal. The coat is long and soft-haired. There is no marked saddle but a line of white guard hairs, followed below by a line of dark hairs, runs along each side of the body, giving the jackal its name. The under- fur is ochreous, the guard hairs banded dark and white, giving a generally mottled grey appearance to the pelt. Head buffy-grey with darker grey ears. Underparts pale grey. Tail tip white. Feeds on carrion, rodents, insects and vegetable matter. Skull slightly longer and narrower than that of C. mesomelas, but the teeth are smaller and less high-crowned, especially the carnassials. The bullae are smaller and flatter. Interparietal crest slightly developed. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Canis mesomelas 89-9 Canis latrans 89-7 Dusicyon australis 88-8 Canis aureus 88-7 Dusicyon inca 88-5 Skull and teeth only Canis mesomelas 92-7 Dusicyon culpaeolus 92-0 Dusicyon gymnocercus 91-5 Canis simensis 91-2 Dusicyon microtis 90-4 As stated above, it is clear from the numerical results that Canis adustus has a high similarity with C. mesomelas. According to Van der Merwe (1953) this jackal is mainly nocturnal and feeds on smaller prey than the black-backed jackal. Cer- tainly the relative conformation of its skull and teeth suggest that this is likely. The side-striped jackal has no reputation as a killer of sheep and consequently it is not exterminated by farmers in the same way as C. mesomelas. These two jackals are a good example of closely related sympatric species. Canis simensis Riippell, 1835 Ethiopian jackal, Simien jackal DISTRIBUTION. Montane ; inhabits grassland plateau areas associated with giant lobelia at an altitude of 2900 to 3900 m on the Simien and other mountains in central Ethiopia. Probably nearly extinct and classified as an endangered species in the Red data book (Goodwin & Holloway, 1972). DESCRIPTION. Very little is known about this rare canid. The overall colour is a tawny rufous with pale ginger underfur. The chin, insides of ears, chest and 150 J. CLUTTON-BROCK ET AL. underparts are white. There is a distinctive white band around the ventral part of the neck and the inner sides of the limbs are also white. The tail is rather short ; the posterior end is dark with black ends to the guard hairs ; the anterior part is white underneath and around the anus. There is no dark patch to mark the tail gland. Skull 'jackal-like' but with an elongated facial region. Teeth, especially the upper carnassials, rather small. Canines long and sharply pointed. Interparietal crest slightly developed. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Dusicyon gymnocercus 88-5 Canis adustus 91-2 Dusicyon inca 87-6 Dusicyon gymnocercus 89-7 Dusicyon culpaeolus 87-3 Chrysocyon brachyurus 88-7 Dusicyon fulvipes 86- 1 Dusicyon culpaeolus 88-5 Dusicyon culpaeus 86- 1 Dusicyon culpaeus 86-5 Canis simensis is sometimes called the Simien fox. It is not, however, at all closely linked to the Vulpes group and the postorbital processes of the frontal bones do not have the little depressions that signify the lack of frontal sinuses ; a character that within the Canidae is only found in the genera Vulpes, Alopex and Otocyon. The near-neighbours tables and two-dimensional plots show a seemingly close similarity with the genus Dusicyon but it can be seen that both Canis adustus and Canis mesomelas are also close to Dusicyon, and it is possible that the numerical results for C. simensis are biased by lack of data on the postcranial skeleton and on behaviour. Gray (1868) placed this species in a separate genus, Simenia, and this classification was followed by Allen (1939). However, the 91-2 similarity that the cranial and dental characters have with C. adustus shows that separate generic status is not justified and the species is therefore retained within the genus Canis, In general appearance the skull of C. simensis looks like an elongated skull of C. adustus, in the same way as, in the foxes, the skull of Vulpes ferrilata looks like an elongated skull of Vulpes corsac. It may be worth comment that both C. simensis and V . ferrilata are adapted to a montane environment. Genus Vulpes Frisch, 1775 Type species Canis vulpes L. 1758. The work of Frisch (1775) has been declared unavailable by the International Commission on Zoological Nomenclature (Anon., 1950). This author was accepted by Simpson (1945) and is used here because the next available uses of Vulpes (Bow- dich, 1821 ; Fleming, 1822) postdate the generic name Fennecus Desmarest, 1804 (Oken, 1816 also being unavailable, see Opinion 417, 1956). As we propose in this classification to include Fennecus zerda with the foxes this would mean changing the generic name for the entire group of foxes from Vulpes Fleming, 1822 to Fennecus THE FAMILY CANIDAE 151 Desmarest, 1804 if Fleming were accepted. This change would clearly be most impractical, as Vulpes is in such general use. A proposal has therefore been submitted to the Commission to place Vulpes Frisch, 1775 on the Official List of Generic Names in Zoology (Clutton-Brock & Corbet, 1975). The genus Vulpes covers nearly the same geographical range as Canis except that there is no species of fox in central Africa. Twelve species have been included in this classification. As well as all those that are generally recognized as true foxes it has been found necessary to include the fennec fox (Fennecus zerda}, and the American grey fox (Urocyon cinereoargenteus). Justification for the changes are given on p. 134 and in the sections on these species. All the species of fox are solitary carnivores and they mostly live in burrows that they dig themselves. They prey on small mammals, birds, reptiles, insects and eggs, whilst some species feed on a considerable amount of fruit and vegetable matter. All foxes have a pointed muzzle, large erect ears and a long bushy tail. They tend to be rather low-bodied and have long, thick fur, but the wide distribution of the genus is reflected in modifications to these characters, as for example in adaptations to desert and montane conditions. The skull of all members of the genus Vulpes is distinctive in that the frontal sinuses are only slightly developed, if present at all, and there are small depressions that can be seen and felt on the frontal bones just medially from the postorbital processes. It may be noted that the skulls of Vulpes zerda, Vulpes cinereoargenteus and Alopex lagopus show these depressions and so, incidentally, does the skull of the South African bat-eared fox, Otocyon megalotis. In all foxes the skull is slender and flattened compared to that of Canis. The temporal ridges may be nearly fused as in Vulpes vulpes or they may be indistinct and wide apart as in the desert foxes. The raised temporal ridges and rugose parietal bones of V. cinereoargenteus can be seen as an exaggerated form of a common character when the genus is looked at as a whole. The sagittal and parietal bones of Vulpes pallida, in fact, closely resemble those of V. cinereoargenteus but are, in comparison, only feebly developed. Huxley (1880) made a comprehensive comparative study of the skulls and den- tition of V. vulpes and Dusicyon culpaeus as a basis for his wider study of the whole family Canidae. He concluded that although the skulls of the two species were very alike there were outstanding differences in the absence or slight development of the frontal sinuses in the fox and in the relative shapes of the cranial cavities, reflecting the shape of the brain. On these differences Huxley divided the Canidae into two groups, the alopecoids which included all the true foxes, and the thooids which included Canis, Dusicyon and Lycaon. This division of the genera into two groups on the basis of brain morphology has been repeated recently by Radinsky (1973) who found distinctions in the relative size and shape of the prorean gyrus and hence in the profiles of the frontal lobes of the brain between the species of Canis and Vulpes. Radinsky, however, found that species of Dusicyon were intermediate between these two genera in the shape and size of the proreari gyrus (defined as the dorsal part of approximately the anterior two-thirds of the frontal lobe : Radinsky, pers. comm., 1974). 152 J. CLUTTON-BROCK ET AL. Vulpes vulpes (L., 1758) Common fox, red fox DISTRIBUTION. The common fox is the most widespread of all wild canids and even exceeds the wolf in its distribution. It is found in wooded and open country throughout the Palaearctic region, including North Africa, and in southeastern Asia, northern Indo-China and much of North America. The natural range has been extended by human agency, perhaps most notably into Australia. DESCRIPTION. The largest member of the genus but very variable in size, as in many other characters, throughout its wide range. Typically the pelage is a rich rufous colour. The backs of the ears are black or dark brown and contrast strongly with the head and neck. There may be a black patch or mask between the nose and eyes. The insides of the ears are light in colour as are the chin and underparts. The tail, or 'brush', is very long and bushy and has a white tip. There is a dark stripe down the front of the foreleg, and the hindleg is black from the hock down- wards. There are, however, many variations to this coat colour pattern and a melanistic form is quite common. A black and silver-grey variety (the silver fox) is bred in captivity for its fur. The skull is one of the largest in the genus Vulpes, with a long narrow palatine region. An interparietal crest may be present but is not normally strongly de- veloped. The temporal ridges lie close together. The cheek-teeth are sharp but rather small. P2 and P3 have no posterior secondary cusps. The canines are long and finely pointed. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Vulpes bengalensis 87-4 Vulpes velox 86-5 Vulpes rueppelli 86-3 Vulpes chama 84-9 Vulpes cor sac 84-5 Skull and teeth only Vulpes velox 91-8 Vulpes bengalensis 91-4 Vulpes corsac 90-2 Alopex lagopus 89-6 Vulpes chama 89-2 Although Vulpes vulpes is the type species of the genus it bears a rather low similarity to the rest of the foxes on all characters (discussed in the section on typicalities, see p. 136), and on the two-dimensional plots it can be seen to be hardly less peripheral than Alopex lagopus, Vulpes cinereoargenteus and Vulpes zerda (Figs 2a, 3a, 5 a). On cranial and dental characters, however, the common fox does lie well within the genus. REMARKS. The common fox would undoubtedly have been domesticated by man if its solitary nature and pungent smell had not made it so intractable, for the species has had an almost symbiotic relationship with man since the prehistoric period. Fox bones are commonly found amongst Neolithic animal remains, especi- ally in Western Asia (Glutton-Brock, 1969) where foxes appear to have been an important source of meat. Their pelts remain of economic value at the present day in many parts of the world. On the other hand, since the beginnings of livestock THE FAMILY CANIDAE 153 husbandry the fox has preyed on domestic animals and scavenged for food around homesteads. Attempts to control the fox's depredations on livestock have de- veloped into one of the most highly ritualized of sports. Paradoxically, hunting of the fox has been the means, not only of its preservation, but also of an increase in its distribution. Before the emigration of Europeans to North America, the common fox may have had a much more restricted distribution over the whole continent, for it is known that foxes were imported into the eastern regions in the seventeenth century (Gilmore, 1946). British foxes were introduced by a hunt club into Australia in 1868 (Troughton, 1957). Vulpes cor sac (L., 1768) Corsac fox DISTRIBUTION. Steppe-lands of southeastern Russia, Volgo-Ural steppes, Russian Turkestan and Kirghizia, to Chinese Turkestan, Mongolia and Trans- baikalia. Possibly also Manchuria and Northern Afghanistan (Ellerman & Morrison-Scott, 1966). DESCRIPTION. A small fox, similar in size to Alopex lagopus, but with relatively longer legs. The fur is thick, soft and pale straw-coloured with darker, slightly tawny markings along the back. Except for a small black patch over the tail gland and a slightly black tip to the tail there is no dark colouring on the head, body or limbs. The underparts are pale. The three specimens in the British Museum (Natural History) have closely similar markings and Ognev (1962) described the same pelage characters. According to Ognev, this fox is less solitary than most species and may hunt in small packs. It feeds on rodents, birds, small reptiles and insects. Like A, lagopus the corsac fox may inhabit communal breeding dens. Skull similar to that of A . lagopus but the teeth may be relatively smaller. The temporal ridges are flat and may enclose a lyriform sagittal area. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Vulpes bengalensis 93-7 Alopex lagopus 90-8 Vulpes velox 93'3 Vulpes bengalensis 90-7 Dusicyon gymnocercus 90-9 Vulpes vulpes 90-2 Vulpes ferrilata 89-8 Vulpes velox 90-0 Dusicyon vetulus 89-8 Vulpes chama 88-8 As shown by the 'near neighbours' tables the corsac fox lies closest to A. lagopus on cranial and dental characters and closest to Vulpes bengalensis on all characters. Phenetically the species is a typical fox, despite its small size and on the two- dimensional plots it lies within the Vulpes group. 154 J. GLUTTON-BROCK ET AL. Vulpes ferrilata Hodgson, 1842 Tibetan sand fox DISTRIBUTION. High plateau country of Tibet and Nepal between 4500 and 4800 m (Ellerman & Morrison-Scott, 1966; Pocock, 1941). DESCRIPTION. The skins of this apparently rare fox have been described in some detail by Pocock (1936). The body colour is pale grey agouti or sandy with a tawny band along the dorsal region. The fronts of the legs are also tawny ; the underparts pale. Insides of the ears white, the outsides similar in colour to the rest of the body. The fur is soft and thick and the tail bushy. The end of the tail is white, whilst the anterior part has a wide band of dark guard hairs. There may be a dark streak over the tail gland. Very little is known of the habits of this fox or of the functions of its extraordinarily long head. Skull peculiarly elongated and with a very narrow maxillary region. Canine teeth also remarkably elongated and pointed. Cheek teeth well developed but widely spaced in the long jaws. Mandible correspondingly elongated and with relatively little depth. Temporal ridges of the cranium flat and enclosing a narrow, lyriform sagittal area. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Vulpes corsac 89-8 Dusicyon gymnocercus 89-2 Dusicyon culpaeolus 89-1 Dusicyon australis 88-9 Vulpes velox 87-9 Skull and teeth only Vulpes corsac 88-4 Vulpes velox 88-0 Dusicyon gymnocercus 84-9 Vulpes chama 84-8 Dusicyon microtis 82-7 Vulpes ferrilata is phenetically closer to the species of Dusicyon than is any other member of the genus Vulpes. Despite the unique appearance of the skull, however, it is of interest that in both near-neighbours tables this fox is slightly closer to Vulpes corsac than it is to any other species. The distribution of V. corsac lies to the north of that of V. ferrilata and it may be that the Tibetan fox has evolved from the more typical V. corsac in response to a specialized environment. An analogous situation may have occurred with Canis simensis which is phenetically close to Canis adustus and has a somewhat similar elongated muzzle. Vulpes bengalensis (Shaw, 1800) Bengal fox DISTRIBUTION. Open country, thorny scrub or semi-desert areas in southern peninsular India, Travancore, northwards to Sind, Bihar and Orissa, Kangra in Punjab, Haldibari and Nepal up to 1350 m (Ellerman & Morrison-Scott, 1966 ; Pocock 1941). DESCRIPTION. This Indian fox is medium-sized and sandy-coloured with soft fur that is not as thick or long as it is in Alopex lagopus or Vulpes corsac. The THE FAMILY CANIDAE 155 dorsal region of the pelt may be darker or more tawny than the rest which is either pale agouti or fawn, with tawny legs. The insides of the ears are white, the outsides grey, and the underparts are light-coloured or pale ginger. The black tip to the tail is the only dark colouring in the pelage except that in a few specimens there is a small dark patch over the tail gland. Feeds on small animals, including insects, and eggs as well as a fairly high proportion of fruit and berries. Skull typically 'fox-like', with long sharply pointed canines and well-developed molar teeth. The temporal ridges are flat and may enclose a lyriform sagittal area. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Vulpes velox 94-5 Vulpes chama 93-8 Vulpes cor sac 93-7 Vulpes rueppelli 93-3 Dusicyon gymnocercus 91-9 Skull and teeth only Vulpes chama 94-9 Vulpes rueppelli 92-3 Vulpes velox 91-7 Vulpes pallida 91-4 Vulpes vulpes 91-4 The Bengal fox has a similarity of over 90 with other species of fox shown in the table above and as can be seen from Table 2, it is the 'most typical' member of the genus for the 'all-character' results and is only three below Vulpes chama for the skull characters. It is reasonable therefore to assume that Vulpes bengalensis typifies the 'basic fox'. Vulpes cana Blanford, 1877 Blanford's fox DISTRIBUTION. Not well known but probably the mountain areas of Kopet Dag, southwestern Russian Turkestan, Afghanistan, northeastern Iran and Baluchi- stan (Ellerman & Morrison-Scott, 1966 ; Ognev, 1962). DESCRIPTION. A small fox with extremely soft fur and a long very bushy tail. The colouring is a blotchy black, grey and white with a dark tip to the tail and a dark patch over the tail gland. There is an almost black mid-dorsal line and the hind legs may be dark. Blanford (1888) described the pelt as having a 'rufescent tinge' but the skins examined in the British Museum appear to have no red pigment in the hair (see Table 6). The underparts are almost white ; the ears are grey, and there is a small dark patch between the eyes and nose. The condylo-basal length of the skull exceeds that of Vulpes zerda (the smallest species of canid) by only a few millimetres. Despite its small size the skull and dentition are typically vulpine with small sharply pointed teeth, flat temporal ridges and a narrow maxillary region. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Vulpes rueppelli 85-5 Vulpes bengalensis 84-6 Vulpes velox 84-3 Vulpes zerda 83-4 Vulpes chama 83-0 Skull and teeth only Vulpes zerda 89-5 Vulpes velox 88- 1 Vulpes chama 87-6 Vulpes rueppelli 87-0 Vulpes pallida 87-0 156 J. CLUTTON-BROCK ET AL. The numerical results show a rather low similarity for this species with the rest of the genus Vulpes. On the two-dimensional plots, however, Vulpes cana lies close to the sand foxes and its phenetic relationships must be with this group. Vulpes rueppelli (Schinz, 1825) Sand fox DISTRIBUTION. Arid areas of North Africa, southern Arabia, Persian Baluchistan and Afghanistan (Ellerman & Morrison-Scott, 1966 ; Harrison, 1968). DESCRIPTION. A large-eared, desert, sand fox. The pelage is reddish-grey agouti with dark guard hairs on the tail and a dark patch between the eyes and nose. Light underparts. Ears not distinct in colour from the rest of the body. As with the other desert foxes, Vulpes pallida and Vulpes zerda, the facial vibrissae are particularly long and black. Skull small but typically vulpine with a straight profile, narrow maxillae and small sharp canine teeth. Bullae large but not so expanded as in V. zerda. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Vulpes bengalensis 93-3 Vulpes bengalensis 92-3 Vulpes chama 91-1 Vulpes chama 90-9 Vulpes velox 91-1 Vulpes velox 89-8 Dusicyon culpaeolus 88-9 Vulpes pallida 89-6 Vulpes corsac 88-7 Dusicyon gymnocer cus 87-1 Vulpes rueppelli is a small fox that is well adapted to life in dry sandy environ- ments but it does not have the extreme desert-characters that are seen in V. zerda. The 'near-neighbours' tables and two-dimensional plots show that the species is phenetically close to Vulpes bengalensis and to the desert foxes (described in the following sections). Vulpes pallida (Cretzschmar, 1826) Pale fox DISTRIBUTION. Dry sandy areas in a line running across Africa from Senegal through Nigeria and Cameroun to the Sudan and Somalia (Allen, 1939 ; Eller- man & Morrison-Scott, 1966). DESCRIPTION. A small ginger-coloured fox with large ears that are the same colour as the body. The tail is dark and has a black tip and a dark patch over the tail gland. The underparts are pale and may be a pinkish ginger. Legs rufous. No dark patch between the eyes and nose. Skull small with a wide lyriform sagittal area and a relatively short maxillary region. The upper molars are well developed in relation to the carnassial teeth (P4) which are weak. THE FAMILY CANIDAE 157 SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Vulpes chama 92-4 Vulpes chama 95-2 Dusicyon sechurae 91-0 Vulpes bengalensis 91-4 Dusicyon fulvipes 90-9 Vulpes rueppelli 89-6 Vulpes bengalensis 90-5 Vulpes velox 89-0 Dusicyon gymnocercus 90-0 Vulpes cana 87-0 The systematic position of Vulpes pallida was discussed by Thomas (1918) in a short note on the sand foxes of North Africa. Thomas associated V. pallida with Vulpes rueppelli and Vulpes zerda and this grouping has been generally followed since then. Our numerical analysis of the phenetic characters suggests that these desert foxes are more closely related to the Indian fox Vulpes bengalensis and to the South African Vulpes chama than had been previously realized. The situation can be seen best as a series of species ranging in an arc from V. chama through V. pallida, V. zerda and V. rueppelli to V. bengalensis, with V. zerda as the most highly special- ized desert form. Vulpes cana (Blanford's fox) falls geographically within this arc, but it is adapted to a montane rather than a desert environment and its unusual pelage characters set it apart from the rest of the series. Vulpes zerda (Zimmermann, 1780) Fennec fox DISTRIBUTION. Desert areas of Morocco, Algeria, Libya, Egypt and east to Sinai and Arabia. Also south to the Sudan (Ellerman & Morrison-Scott, 1966 ; Harrison, 1968). DESCRIPTION. The smallest species of canid, with extraordinarily large ears. The pelage has no agouti hairs but is an evenly pale fawn colour with almost white underparts. The tail tip is dark and there is a dark patch over the tail gland ; these being the only parts of the body that are not pale in colour. There may be a ginger line along the back. The facial vibrissae are very long. Skull very small with exceedingly large swollen bullae. Otherwise typically vulpine, except that the cranium is rather rounded and the dentition is weak. The sagittal area, enclosed by barely perceptible temporal ridges, is very wide. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Vulpes chama 89-7 Vulpes cana 89-5 Vulpes pallida 88-9 Vulpes pallida 86-9 Vulpes bengalensis 88-5 Vulpes bengalensis 85-9 Dusicyon fulvipes 88-0 Vulpes chama 85-2 Vulpes velox 87-7 Vulpes rueppelli 83-7 158 J. CLUTTON-BROCK ET AL. V. zerda has been traditionally placed in a separate genus, Fennecus, and this classification is generally followed on account of this fox's huge ears, pale colouring and rounded skull. These characters should be seen, however, in their true context as adaptations to a most specialized environment. Like Alopex lagopus the fennec lies on the periphery of the fox group because it is adapted to extreme conditions where the biotic abundance is very low. Table 3 shows that the fennec fox lies above Vulpes ferrilata, Vulpes vulpes and Vulpes cana in order of typicality and in the 'near neighbours' table it is seen to have similarity values of nearly 90 with the other small species of fox. The two-dimen- sional plots and centroid linkage dendrograms also show that the fennec lies well within the genus, more so than V. vulpes. It would therefore be irrational for us to exclude this species and although it makes for nomenclatural difficulties (see p. 150) we are constrained to transfer it to the genus Vulpes. Vulpes chatna (A. Smith, 1833) Cape fox DISTRIBUTION. Dry areas of southwestern Africa, Transvaal and possibly western Rhodesia. Probably extinct in the Capetown area (Allen, 1939 ; Ellerman etal, 1953). DESCRIPTION. A relatively large fox, similar in size to Vulpes bengalensis. The fur is soft and short and the colouring of the body is rufous agouti. There may be long black guard hairs on the bushy tail and on the posterior dorsal region. The tail has a distinct black tip and there is a diffuse dark patch over the tail gland. The underparts are a pale rufous with a lighter chin. The legs are more tawny than the general body colour. There is no dark mark between the eyes and nose. Skull very similar to that of F. bengalensis, but the cranium is slightly wider and the maxillary region slightly shorter (see Table 4). SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Vulpes bengalensis 93-8 Vulpes bengalensis 95-2 Vulpes pallida 92-4 Vulpes pallida 94-9 Dusicyon culpaeolus 92-2 Vulpes velox 92-6 Vulpes velox 92-0 Vulpes rueppelli 90-9 Dusicyon gymnocercus 91-3 Dusicyon sechurae 89-8 The numerical results show that Vulpes chama is surprisingly closely related in its phenetic characters to the Bengal fox, F. bengalensis, and to the more northerly African fox, Vulpes pallida. On skull characters F. chama heads the list as the most typical member of the genus (Tables 2, 3) and as described under the description of F. pallida, it seems clear that this South African fox should be considered as one end of an arc of related species that have evolved in response to varying degrees of desert conditions. THE FAMILY CANIDAE 159 Vulpes velox (Say, 1823) Kit fox For the purposes of this analysis Vulpes macrotis Merriam, 1888 was included within the species Vulpes velox. DISTRIBUTION. Prairies of western North America. Distribution not well known but certainly decreasing. The northern subspecies is classed as endangered by the Red data book (Goodwin & Holloway, 1972) ; it is extinct in Canada. DESCRIPTION. A medium-sized fox with very thick soft underfur and long agouti guard hairs. The body colouring may be tawny or light ochreous and grey. Tail relatively short and very bushy with a black tip and a slight black patch over the tail gland. There is a dark patch between the eyes and the nose. Large ears white inside and grey or ochreous outside. Almost white underparts. Limbs tawny. Skull typically 'fox-like' and very similar to that of Vulpes chama and Vulpes bengalensis. Upper molar teeth slightly less well developed than in the latter species. Temporal ridges flat and enclosing a narrow lyriform sagittal area. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Vulpes bengalensis 94-5 Vulpes cor sac 93-3 Vulpes chama 92-0 Vulpes rueppelli 91-1 Dusicyon culpaeolus 90-5 Skull and teeth only Vulpes chama 92-6 Vulpes vulpes 91-8 Vulpes bengalensis 91-7 Vulpes corsac 90-0 Vulpes rueppelli 89-8 It is clear from the results of this analysis that the phenetic affinities of V. velox lie with the 'most typical' members of the genus, these being V. bengalensis from India and V. chama from South Africa. On the other hand, the similarity of 91-8 that the skull of V. velox bears to that of V. vulpes may provide a link between this widespread but somewhat discrepant species and the more typical group. Support for this may be seen in the work done by Creel et al. (1971, 1974) on hybridization between the kit fox and the common fox. Vulpes cinereoargenteus (Schreber, 1775) Grey fox DISTRIBUTION. Widespread in wooded country and along river valleys throughout Central and North America and the northern part of South America but not in the high plains (Cabrera, 1958 ; Hall & Kelson, 1959 ; Miller & Kellogg, 1955). A versatile carnivore that will easily adapt from a wooded to a pastoral environment (Hershkovitz, 1972 : 372). DESCRIPTION. A medium-sized, typically 'fox-like' canid. Body colour grey agouti with white jaws and throat. Ears and sides of neck ochreous or tawny. 160 J. CLUTTO-NBROCK ET AL. Chin grey or brown ; underparts pale ; legs and feet tawny. Long bushy tail. A dorsal black stripe extends from the mid-line of the back along the whole of the tail to the tip which is black. Hildebrand (i952b) stated that the tail gland is longer in this species than in any other canid. The gland is covered by a ridge of stiff guard hairs. (This character was used by Gray (1868) to support his classification of the grey fox in the separate genus Urocyon Baird, 1857.) Feeds on the usual small animals and birds that all foxes prey upon. The grey fox is said readily to climb trees to escape from hunters or other enemies - a most unusual habit for a canid. Teeth well developed. Canines not as long as is typical for the fox group as a whole, and the premolars high-crowned in relation to their length. Carnassial and molar teeth 'fox-like'. The cranium is distinctive and similar to that of Otocyon megalotis in having the temporal ridges well developed but separated by a wide sagittal area. The surface of the parietal bones is rugose, whilst that of the sagittal area is smooth. The frontal sinuses are present only as barely visible pockets (in a bisected skull) below the postorbital processes. There is a subangular lobe on the mandible but it is not so well developed as in 0. megalotis. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Vulpes bengalensis 89-5 Vulpes pallida 85-5 Vulpes velox 87-8 Vulpes chama 85-1 Vulpes corsac 87-8 Vulpes rueppelli 84-9 Vulpes fueppelli 87-6 Vulpes velox 83-5 Vulpes pallida 87-5 Vulpes bengalensis 83-3 A second species of grey fox, usually described as Urocyon littoralis Baird, 1858, is an island form that is probably closely linked to the mainland species. It has not been included in this analysis. The systematic position of the grey fox has been one of the most interesting problems to emerge from the present analysis. There has been little work done on the affinities of the species in the past although both Huxley (1880) and Guilday (1962) have observed the similarity in skull conformation between the grey fox and 0. megalotis. The separation of the grey fox in the genus Urocyon is generally accepted, and it has been asserted since the time of Mivart (1890) that this fox has more in common with the South American canids (genus Dusicyon) than with the common fox (Vulpes vulpes), the only member of the fox group with which it has been compared by mammalogists. The misconception has arisen, in part, because of the atypical appearance of the common fox in comparison with the rest of the genus, and it may also stem from the work of Osgood (1934) who described the grey fox in his paper on the South American canids and, by implication, clearly thought of it as belonging to the South American group. He made no mention of any possible relationship of the species with Vulpes, and inferred that Vulpes cinereoargenteus was more closely related to Dusicyon than was Chrysocyon brachyurus, the maned wolf, a theory for which this analysis gives no support at all. It can, in fact, be asserted THE FAMILY CANIDAE 161 that the grey fox bears less phenetic resemblance to the South American genera of canids than do most other members of the family, and it can be shown, for the first time, that there is a considerable similarity (almost 90 per cent for all characters) between the grey fox and the typical Vulpes bengalensis. Furthermore V. cinereo- argenteus has no similarity with any genus other than Vulpes in the 'near neighbours' tables. The development of the temporal ridges and subangular lobe of the mandible do place the skull apart from all other members of Vulpes but within the terms of this numerical taxonomy it would not be consistent to keep the grey fox as a separate genus. It lies on the periphery of the Vulpes group, as can be seen from the two- dimensional plots, but less so than V. vulpes or V. ferrilata. The southerly distribution of V. cinereoargenteus in relation to its phylogenetic origin has been much discussed. Most authors (including Mivart, 1890) have agreed that it must be a latecomer to South America. Hershkovitz (1972 : 312) described the species as a 'varicant', straddling the Nearctic and Neotropical regions and not clearly derived from either. He postulated further (p. 359) that the grey fox may have originated in Middle America and spread during the Quaternary into Canada and South America. Now that the species has been critically examined in relation to all the other members of the canid family it may be said that derivation from Dusicyon or an autochthonous origin in Middle America seems unlikely. It appears that it has closer phenetic links in the Asiatic species of Vulpes than was previously suspected and perhaps it has been pushed south as a result of competition with the other canid species in North America, and in particular the highly successful V. vulpes which was probably aided in the extension of its range by the activities of man (see p. 153). Genus ALOPEX Kaup, 1829 One species. Alopex lagopus (L., 1758) Arctic fox DISTRIBUTION. Arctic tundra of Europe, Asia and North America and areas of montane tundra in Scandinavia. In Asia southwards to Kamchatka (Ellerman & Morrison-Scott, 1966 ; Macpherson, 1969). DESCRIPTION. A small compact fox. The pelage is distinct in that it has two colour phases. One phase is pure white in winter, whilst in summer the back, legs, tail and head are dark brown and the underparts are light. The other phase is described as 'blue' and is more variable, being grey, brown or black in summer and winter. The fur is thick and very soft with guard hairs as fine and long as the underfur. Muzzle and ears relatively short, tail very thick and bushy. The Arctic fox is solitary in its hunting habits but the breeding dens are often found in colonies. 162 J. CLUTTON-BROCK ET AL. Skull rather shorter in the palatine region than in Vulpes vulpes and the frontal bones are slightly swollen at their junction with the nasals, but this does not affect the depressions on the postorbital processes which are characteristic of the genus Vulpes. Dentition as in V. vulpes but the canines may be relatively shorter. The Arctic fox feeds on small mammals, especially lemmings (Dicrostonyx and Lemmus spp.) and carrion. When lemmings are scarce the fox will feed more heavily on birds' eggs, insects, berries and other fruits and seeds. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Vulpes pallida 81-6 Vulpes cor sac 90-8 Vulpes chama 81-5 Vulpes vulpes 89-6 Vulpes cor sac 81-2 Vulpes rueppelli 86-7 Dusicyon gymnocercus 81-0 Vulpes bengalensis 86-3 Dusicyon culpaeolus 80-7 Dusicyon culpaeolus 85-2 Up to the present time the Arctic fox has been commonly classified in a separate genus, Alopex Kaup, 1829, although some authors, including Bobrinskii (1965), have preferred to make Alopex a subgenus of Vulpes. Miller (1912, p. 318) listed the following distinctive characters in support of separate generic status : 'Skull intermediate in general form between that of Canis and Vulpes ; occipital depth about one third condylo-basal length ; interorbital region more elevated than in Vulpes ; postorbital processes thin, flat, or slightly concave above, with bead-like overhanging edges ; dorsal profile of forehead rising abruptly above rostrum as in Canis ; teeth moderately heavy and large ; external form fox-like, but ear short and rounded, not conspicuously overtopping the surrounding fur.' Miller went on to state that although in most respects intermediate between Canis and Vulpes the Arctic foxes form such a natural group that they should be in a distinct genus. When the skull of the Arctic fox is compared with that of Vulpes vulpes most of the above distinctions can be seen to hold, although we have not noticed any difference in the degree of depression in the postorbital processes in Alopex lagopus. When the skull is compared with that of Vulpes corsac, however, there are fewer differences and as can be seen from the 'near neighbours' table, A. lagopus, has a similarity of 90 with the skull of this species. Alopex lagopus is a species of fox that has special adaptations to life in an arctic environment where there is low biotic abundance, and although the skull is similar to that of V. corsac, on the numerical results for all characters the species is separated at a similarity of only just over 80. It is clear from the two-dimensional plots that the Arctic fox lies close to the genus Vulpes but its inclusion amongst the foxes in the table of 'typicality' (Table 3) shows that it is the most aberrant of the foxes and there are therefore grounds for retaining it in a separate genus. Genus OTOCYON Muller, 1836 One species. THE FAMILY CANIDAE 163 Otocyon megalotis (Desmarest, 1822) Bat-eared fox DISTRIBUTION. Arid areas in South Africa, southern Angola, Botswana, perhaps western Rhodesia, East Africa and northwards to the Sudan, Ethiopia and Somalia (Ellerman et al., 1953). DESCRIPTION. A long-legged, medium-sized fox with very large, wide ears, long fur and a very bushy tail. The general colour is brownish or ochreous with grey agouti guard hairs. Throat, underparts and insides of ears pale. Limbs nearly black as are the outsides of the ears and the muzzle. Black tip to the tail. Omnivorous, social animals, living in groups and feeding on insects, small rodents, fruit and berries. Adapted to life in a desert environment. Apart from the dentition the skull bears a singular resemblance to that of Vulpes cinereoargenteus, with well-developed temporal ridges enclosing a wide sagittal area and rugose parietal bones that contrast with the smooth surface of the sagittal area. Bullae large. No frontal sinuses. Dentition unique in that there are always at least three upper and four lower molar teeth. Carnassials much reduced in length. Canines large and 'fox-like'. Premolar teeth high-crowned in relation to their length as in V. cinereoargenteus. Subangular lobe of the mandible unusually large. Basal line of the horizontal ramus very straight. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Vulpes chama 75-6 Vulpes cinereoargenteus 75-9 Vulpes vulpes 73-4 Vulpes pallida 67-1 Vulpes pallida 72'7 Nyctereutes procyonoides 66-0 Vulpes cinereoargenteus 72-6 Vulpes rueppelli 65-6 Vulpes velox 72-0 Vulpes vulpes 65-6 Huxley (1880) suggested that Otocyon megalotis was the most primitive member of the canid family and that its extra molar teeth represented the basic mammalian dentition. Matthew (1930 : 123) believed that 'an extra upper and lower molar had appeared', but he gave no further explanation of this appearance. Guilday (1962) put forward the theory that the extra teeth were the result of a mutation that duplicated the upper first and lower second molars at the expense of the carnassial teeth which were correspondingly shortened in length. This theory seems sound, for the molars in question are most similar to each other. That this mutation is of considerable age is shown by the finding of a primitive Otocyon in the Villafranchian of Olduvai, Tanzania. This specimen was named Protocyon reckii by Fetter (1964) who considered it to be more primitive than the Recent form and ancestral to it. Simpson (1945 : 224) tentatively allowed the subfamily rank of Otocyoninae Troues- sart, 1885 to stand for this monotypic genus but it is clear that he did not really approve of it. Our numerical results support Simpson's hesitation and there seems little doubt that 0. megalotis should be considered as an aberrant fox with affinities to Vulpes cinereoargenteus. There is therefore no justification for the recognition of a 164 J. CLUTTON-BROCK ET AL. subfamily Otocyoninae, but the generic status of Otocyon is clearly established by the low level of similarity that it bears to all other species. The similarities in behaviour between 0. megalotis and Nyctereutes procyonoides are discussed in the next section. Genus NYCTEREUTES Temminck, 1839 One species. Nyctereutes procyonoides (Gray, 1834) Raccoon dog DISTRIBUTION. River valleys and the edges of forests in the Amur and Ussuri region of eastern Siberia, Japan, Manchuria, China and Indo-China (Ellerman & Morrison-Scott, 1966 ; Ognev, 1962). Introduced and now widespread in European Russia and eastern Europe. DESCRIPTION. A rather slow-moving, heavy-bodied canid with a small head and short limbs. The pelage characters give it a superficial resemblance to the raccoon, Procyon lotor (L.). The back is a mottled tawny and black, the guard hairs being long, banded, rather coarse and shiny ; the underfur is abundant, soft and fawn in colour. The tail is rather short and dark at the end but without a distinct black tip. Limbs fawn or dark brown. The facial region is short. The raccoon dog is the only species of canid that has a distinct dark mask around the eyes and between the eyes and ears. Beneath ea.ch eye there is a diffuse white band stretching back to the ears and emphasizing the dark mask. Underparts brown, beige or fawn. Nocturnal, fossorial, hunting in pairs or family groups. Preferred habitat, small forested areas near water, and river valleys. Diet very varied : often eats fish and feeds on small rodents, amphibians, eggs, shellfish, berries and acorns (Ognev, 1962). Hibernates. Skull small with short nasals and maxillary region. Distinct subangular lobe to the mandible but not so highly developed as in Otocyon megalotis. Teeth small and weakly developed. Molars somewhat bunodont. Palatine bones extend backwards beyond M2. Surface of parietal bones rugose. Temporal ridges fused to form a slight interparietal crest. Orbits relatively small. Frontal sinuses moderately large. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Dusicyon microtis 78-1 Dusicyon sechurae 86-2 Dusicyon australis 77-4 Canis aureus 83-5 Dusicyon thous 76-8 Dusicyon australis 83-2 Dusicyon vetulus 76-4 Alopex lagopus 82-9 Dusicyon sechurae 75-7 Vulpes cinereoargenteus 82-2 THE FAMILY CANIDAE 165 It is difficult to assess the systematic position of Nyctereutes procyonoides as it has no close affinities with any of the other canids. The 'near neighbours' tables place the genus at a low level of similarity with the Dusicyon group and in this context it may be mentioned that in 1880 Huxley wrote, in his study of the Canidae : 'Nyctereutes is essentially a low Thooid of the South American type.' Radinsky (1973) has suggested on the basis of the shape of the prorean gyrus of the brain that the raccoon dog should be linked with the foxes. The presence of fairly well-developed frontal sinuses in the skull does not, however, lend support to this view. Kleiman (1967) in her study of some aspects of the behaviour of the Canidae asserted that Nyctereutes is unique in that a submissive animal does not wag its tail. She noted several striking similarities between the raccoon dog and the bat-eared fox (0. megalotis). Both engage in communal sleeping and social grooming which she says may be related to the black facial mask that is present in both species although much more extensive in Nyctereutes where, as previously stated, it surrounds the eyes and goes back to the ears. Both species share a peculiar tail posture ; in dominant animals or in a sexually aroused male the tail is carried in an inverted U-shape, and the black hair on the tail is erected and stands up prominently. There can be little doubt about the generic status of Nyctereutes. On the two- dimensional plots and on the centroid linkage dendrograms the raccoon dog is always an outsider and it bears a similarity value of less than 75 with the genera Canis, Vulpes and Dusicyon (Table i). Genus DUSICYON Hamilton Smith, 1839 Type species Dusicyon attstralis (Kerr, 1792) All the species within the genus Dusicyon are restricted to the continent of South America and its neighbouring southern islands. The number of taxa that we suggest should be included in the genus differs from that of the accepted check list of Cabrera (1958) which excludes Dusicyon thous and Dusicyon microtis. Of the eleven species that were examined for this analysis eight form a phenetically closely linked group and it is suggested that further work at the specific level might lead to the elimination of four of these. Three species, D. australis, D. thous, and D. microtis lie on the periphery of the group and their taxonomic position is discussed. It is difficult to give a diagnosis for the genus as in many characters it lies between Canis and Vulpes, with D. australis presenting the most 'dog-like' features and D. vetulus the most 'fox-like'. An indication of this intermediate state is apparent in the descriptions of the early authors who wrote of the animals as 'foxes', 'wild dogs' or 'wolves'. Gray (1868) described them as 'fox-tailed wolves'. The pelage is usually grey agouti with some ochreous or tawny colouring, with the exception of D. microtis which is dark all over. The ears are fairly large and erect ; the head is rather narrow, and the tail is very long, bushy and has a contrasting dark tip (white in D. australis}. The underparts are usually pale and the legs ochreous or tan. The skull is rather long and narrow with temporal ridges either apart and enclosing a lyriform sagittal area or nearly fused. There is no well-marked inter- parietal crest. Dentition is more 'fox-like' than 'dog-like'. The canines are long 166 J. GLUTTON-BROCK ET AL. and finely pointed ; the premolars and carnassials are high-crowned, and the molars are well developed. The carnassial teeth are short relative to the lengths of the molars and to the condylo-basal length. The palatine bones may extend backwards beyond M2. Early writers on this group of South American canids usually placed them all within the genus Cams until the work of Thomas (1914) which brought the following generic names into common use : Dusicyon Hamilton Smith, 1839 ; Cerdocyon Hamilton Smith, 1839 > Pseudalopex Burmeister, 1856 ; Lycalopex Burmeister, 1856. Thomas designated the Falkland Island 'dog' (formerly known as Canis ant- arcticus Shaw, 1800) as the type of Dusicyon. With the general acceptance of Pocock's paper (1913) on the affinities of the Falkland Island 'wolf, in which he allied this species closely with the other South American canids, Thomas's classification was followed, with minor alterations, by Kraglievich (1930) and Cabrera (1931). Osgood (1934) reduced Cerdocyon, Pseuda- lopex and Lycalopex to subgenera of Dusicyon and retained D. australis as D. (Dusi- cyon) australis. Simpson (1945) accepted this classification, but most recent authors have followed Cabrera's further modifications, in which he placed D. thous in the genus Cerdocyon and separated D. microtis into a new genus Atelocynus Cabrera, 1940, leaving the remainder as Dusicyon. Langguth (1970, 1975) went further and separated Lycalopex vetulus as an additional monospecific genus, while including the remaining species of Dusicyon in Canis. The subgenus has been avoided throughout this work because of the absence of sufficiently discrete groups at the appropriate level. This being so, and after a careful examination of the numerical results, it has been decided to include all the genera and subgenera mentioned above within the genus Dusicyon. Dusicyon australis (Kerr, 1792) Falkland Island wolf Although well known from the descriptions of Darwin and others, no detailed examination has been made of the available material of this extinct species since the account of Pocock (1913). It was therefore considered appropriate to give here a fresh description of the material that is held in the British Museum (Natural History) especially as there are many interesting features about the skulls and skins that make the systematic position of the species hard to define. MATERIAL. No. 37.3.15.47. The holotype of Dusicyon darwini Thomas, 1914. Skull, mandible and skin from East Falkland Island. Collected by Charles Darwin and presented by Burnett and Fitz Roy. There are shot holes in the frontal bones behind the orbits and the occipital region of the skull is missing. Young adult male. Data on this specimen were used in the analysis. No. 37.3.15.48. Skull, mandible and skin from West Falkland Island. Collected by Charles Darwin and presented by Burnett and Fitz Roy. Skull complete except for the left zygomatic arch which is missing. Young adult female. Data used in the analysis. THE FAMILY CANIDAE 167 No. 69.2.24.3 (i692a). Skull without mandible ; complete except for the right canine tooth which is missing. No history except that the skull was purchased from E. Gerrard, jun. It is, however, likely that this skull came from the live animal that was brought to the Zoological Gardens, London, by Mr A. A. Lecombe in 1868 (Newton, 1868). Young adult. This skull was not available at the time the numerical analysis was carried out but it agrees in every important respect with those specimens that were used. No. 85.10.12.1 (ig62b). Skull without mandible. North coast, West Falkland Island. Dentition and zygomatic arches incomplete. Presented by E. A. Holmsted. The skull had obviously lain in the sea for some time as it is covered with tiny barnacles. This could raise the possibility that it actually came from a domestic dog rather than from Dusicyon australis. The skull, however, has all the characters that are typical of the Falkland Island 'wolf rather than of a domestic dog ; these being the raised sagittal area and lack of interparietal crest, the extension of the palatine bones backwards from M2, and the development of the malar bone (see Pocock, 1913). Data on this skull were included in the analysis. No. 1974.483 (i692b). Left mandibular ramus with P2, P3, P4, Mx. Although this mandible has the same number as the old registered number of the skull above (no. 85.10.12.1) it cannot be from the same animal as it is too large ; nor does it fit the other skull with no mandible (no. 69.2.24.3). There is no history for this specimen and it is not recorded in the British Museum catalogues. It can be identified as D. australis on the unique character of the lower carnassial in which the little cusp (metaconid) at the base of the main cusp, on the lingual side of the tooth, lies only slightly above the inner cusp of the talonid. Data from this mandible have not been used in the analysis. A further two skulls from the collection of the Royal College of Surgeons (nos 635 & 636) were described by Pocock (1913) and Thomas (1914) but unfortunately these have been missing since the 1939-45 war. These two skulls were catalogued as follows by Flower (1884) : No. 635. Skull O.C. 4363. Presented by Admiral Sir Francis Beaufort. No. 636. Skull. Found by the donor on West Falkland Island. Presented by E. A. Holmsted, Esq., 1878. DISTRIBUTION. Inhabited East and West Falkland Islands until about 1880 when the species became extinct. DESCRIPTION. A 'large wolf-like fox' (Darwin, 1860) with a short face, wide muzzle and short ears. The tail short with a white tip. Coat thick and soft, mainly brown in colour with some rufous and speckled with white from pale guard hairs. Underparts pale, becoming cream at the posterior ventral surface. As remarked by Mivart (1890) and Pocock (1913) there is a dark reddish patch above the hock of the hind leg. The middle part of the tail has long dark guard hairs that contrast with the white tip. They fed mainly on birds, especially the upland goose, Chloephaga picta. Skull large, with relatively wide palatine region. Temporal ridges well developed, enclosing a lyriform sagittal area which is flattened and only formed into an inter- parietal crest at the posterior end of the cranium. Enlarged frontal sinuses. No 10* 168 J. CLUTTON-BROCK ET AL. subangular lobe to the mandible. Palatine bones extend backwards beyond the posterior edge of M2. Teeth large and somewhat compacted in the premolar region. Canines 'fox-like', premolars simple and high-crowned. P4 with the protocone drawn backwards and reduced. Inner tubercle (metaconid) of Mx placed very low, on a level with the posterior cusps (talonid), as figured by Pocock (1913 : 390). The animals that inhabited the East Falkland Island were said to be larger and less red than those on the West Island. Hamilton Smith (1839) recorded a legend that the eastern group was descended from dogs left on the island by the Spanish, whereas the western group consisted of truly wild indigenous 'foxes'. Thomas (1914) examined the material in the British Museum and in the Royal College of Surgeons and decided that two species were represented. These he named Dusicyon darwini (eastern) and Dusicyon antarcticus (western). Apart from there being some doubt about which island two of the skulls originated from, with Thomas altering 'west' to 'east' to suit his case, an examination of the four skulls and two skins now available shows a remarkable similarity between them. Aside from the variation in size that could be due to sex there are few differences in the skull that could be even ascribed to individuality. When Darwin visited the Falkland Islands in 1834 it was apparent to him that the wild 'foxes' that he saw there were doomed to extinction (Darwin & Waterhouse, 1840) and fifty years later they had indeed been exterminated. There remain, however, several contemporary accounts of the species, as well as the few specimens preserved in museums. The extinction of this species was due to indiscriminate slaughter and to the value of its fur to traders as far away as New York. The generic name of Dusicyon was given to this species by Hamilton Smith in 1839 after he had seen a large collection of pelts in a fur store in New York, owned by a Mr Astor. Hamilton Smith described these pelts as indistinguishable from those of Lyciscus cagottis (the Mexican coyote, Canislatrans}. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Vulpes ferrilata 88-9 Canis aureus 88-8 Dusicyon sechurae 88-6 Dusicyon culpaeolus 88-6 Dusicyon gymnocercus 87-7 Skull and teeth only Canis mesomelas 92-5 Canis aureus 91-4 Dingo 91-1 Canis latrans 88-9 Dusicyon sechurae 88-6 Pocock (1913) examined the skulls and skins of D. australis and decided that they bore a close affinity with Dusicyon culpaeus and that the species could be in no way a near relation of Canis latrans. Pocock was incited to pay attention to the Falkland Island 'wolf by a quotation of Lydekker's from Huxley's work on the cranial and dental characters of the Canidae (1880). In this work Huxley concluded that D. australis was in some skull characters close to C. latrans. The results of the present analysis support Huxley's observations in showing that the skull and teeth are closer to Canis than to Dusicyon. There are definite charac- ters, however, like the length and shape of the canines that more closely resemble THE FAMILY CANIDAE 169 Dusicyon, and the shape of the lower carnassial tooth is unique. So it will not be proposed here that the Falkland Island 'wolf be returned to the genus Canis ; the results are too uncertain and there is not enough material to make a thorough investigation possible. It may be remarked, however, that Pocock was somewhat hasty in his total rejection of Huxley's observations which in this, as in other parts of the work, are found to agree very well with the results obtained from our numerical analysis. In the two-dimensional plots as well as in the dendrogram for cranial and dental characters D. australis lies as close to, or closer to, Canis than to Dusicyon (Figs 2b and 8b). The Falkland Islands lie within the continental shelf, approximately 400 km east of Patagonia (51-53° S, 57-61° W). It is possible that at some stages of the Pleistocene the islands were connected with the mainland and may have supported a mammalian fauna. If so, it could be argued that the canid became isolated on the islands when they became finally detached from the continent. It would be most surprising, however, for the only relic of a Pleistocene fauna to be one large car- nivorous species. It seems much more likely that D. australis was taken to the Falkland Islands as a domestic animal by early man. This could have happened thousands of years ago, allowing the population to evolve into an autochthonous race, similar to the dingo. Support for this view is seen in the white tip of the tail (all other Dusicyon species have a black tip), the enlarged frontal sinuses and the wide muzzle when compared with other species of Dusicyon. These characters can signify domestication and frequently occur in the dingo. If the Falkland Island 'wolf was descended from domesticated animals it is perhaps possible that a species of Dusicyon was the progenitor rather than a species of Canis. Hamilton Smith (1839), amongst other early writers, described a domesti- cated form of D. culpaeus but he stated that the Indians preferred imported European dogs and that these were superseding the indigenous varieties. Unless further evidence from fossil or archaeological sources comes to light, the origin of D. australis must remain speculative and although the results of this analysis show that the species was quite distinct from the mainland canids, the evidence does not justify giving it separate generic status. Dusicyon culpaeus (Molina, 1782) Colpeo fox DISTRIBUTION. Widespread throughout the Andes mountains and hilly regions of the western and southern countries of South America up to 4000 m (the 'Pata- gonian subregion' of Hershkovitz, 1957, 1972 ; see also Cabrera, 1931, 1958 ; Lang- guth, 1970). DESCRIPTION. Variable in size - can be large and 'wolf-like'. Head, neck, ears and legs tawny or rufous. Underparts pale. Back and shoulders grey with agouti (banded) guard hairs. Underfur fawn. Tail bushy with black tip ; length over half that of the head and body combined. 170 J. CLUTTON-BROCK ET AL. Skull longer and narrower in the facial region than in Dusicyon australis. Frontal bones flat. Interparietal crest poorly developed. Palatine bones do not extend backwards beyond the posterior edge of M2. Canines and premolars simple and 'fox-like' as in D. australis. The metaconid of Mj higher than the level of the talonid as is usual in the Canidae. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Dusicyon gymnocercus 93-9 Dusicyon culpaeolus 93-3 Dusicyon inca 92-7 Dusicyon griseus 92-7 Dusicyon fulvipes 90-0 Skull and teeth only Dusicyon inca 88-5 Dusicyon gymnocercus 88 -i Dusicyon culpaeolus 87-5 Canis simensis 86-5 Canis latrans 86- 1 The three species Dusicyon culpaeus, Dusicyon gymnocercus and Dusicyon culpae- olus are phenetically very close to each other. The pelage characters are so similar that it would be hard to define differences between them and perhaps the distinctions that have been found in the skulls may be attributable to individual variation. It is not possible, however, within the scope of this work to discuss problems of speciation and although it may appear from the numerical taxonomy that these three should be placed in one species they could be valid biological entities whose ecological distributions do not overlap. Dusicyon culpaeolus (Thomas, 1914) DISTRIBUTION. Uruguay (Cabrera, 1958). DESCRIPTION. Very similar to Dusicyon culpaeus, but smaller. Considered by Kraglievich (1930) to be a subspecies of Dusicyon gymnocercus and it certainly bears the closest phenetic resemblance to this species. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Dusicyon gymnocercus 96-2 Dusicyon inca 93-6 Dusicyon culpaeus 93 '3 Canis mesomelas 92-9 Dusicyon griseus 92-7 Skull and teeth only Dusicyon gymnocercus 95-3 Canis adustus 92-0 Dusicyon inca 92-0 Canis mesomelas 9i-9 Dusicyon fulvipes 89-1 D. culpaeolus was not known as a separate species until the description of Thomas (1914) which was made from one skull and skin in the British Museum. Further examination might show that it should be included, with D. gymnocercus, in D. culpaeus. THE FAMILY CANIDAE 171 Dusicyon gymnocercus (Fischer, 1814) Azara's fox DISTRIBUTION. Paraguay, northern Uruguay, southeastern Brazil and eastern Argentina (Cabrera, 1931, 1958). DESCRIPTION. Like Dusicyon culpaeolus the phenetic characters of this species show close similarity to Dusicyon culpaeus, from which it differs only in the shorter, wider rostrum and more uniform pelage. SYSTEMATIC DISTRIBUTION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Dusicyon culpaeolus 96-2 Dusicyon culpaeolus 95-3 Dusicyon griseus 95-5 Dusicyon griseus 91-8 Dusicyon fulvipes 94-3 Dusicyon fulvipes 91-8 Dusicyon culpaeus 93*9 Canis adustus 91 -5 Dusicyon inca 93-8 Canis mesomelas 90-7 Like D. culpaeolus this form may prove to be conspecific with D. culpaeus. Dusicyon inca (Thomas, 1914) DISTRIBUTION. Peru at 4000 m (Cabrera, 1958). DESCRIPTION. A fairly large canid similar in size to Dusicyon culpaeus but distinguishable from it by a more evenly grizzled pelage. Tail with a distinct black tip. Chin black, ears and outer sides of legs a dull tawny brown. Underparts brownish white. Skull similar to that of D. culpaeus. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Dusicyon gymnocercus 93-8 Dusicyon culpaeolus 92-0 Dusicyon culpaeolus 93-6 Dusicyon gymnocercus 88-8 Dusicyon culpaeus 92-7 Dusicyon culpaeus 88-5 Dusicyon fulvipes 91-3 Canis adustus 88-3 Dusicyon griseus 90-3 Canis mesomelas 88-0 This is another of Thomas's species that was described from a single skull and skin (the type is in the British Museum), and like Dusicyon culpaeolus it is possible that a study of further material might show that it should be included with D. culpaeus. It should be pointed out, however, that the pelage of the one skin of D. inca in the British Museum is distinguishable from that species and in fact more closely resembles that of Dusicyon griseus. I72 J. CLUTTON-BROCK ET AL. Dusicyon griseus (Gray, 1836) Argentine grey fox DISTRIBUTION. The plains and low mountains of Patagonia, western Argentina and Chile (Cabrera, 1931, 1958). DESCRIPTION. A small species. Ears large, head rust-coloured flecked with white. Agouti guard hairs with pale underfur giving a generally pale appearance to the back. Underparts pale grey. Feet tawny. Tail long and moderately bushy. The pelage of this species looks very like that of Dusicyon fulvipes, Dusicyon inca, Dusicyon sechurae and Dusicyon vetulus but (from the skins in the British Museum) it is less red than D. fulvipes and more red than the remaining species. Skull small and 'fox-like' with faintly marked temporal ridges enclosing a wide lyriform sagittal area. Teeth widely spaced and 'fox-like'. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Dusicyon gymnocercus 95-5 Dusicyon fulvipes 92-8 Dusicyon fulvipes 94-4 Dusicyon gymnocercus 91-9 Dusicyon culpaeolus 92-7 Vulpes velox 89-0 Dusicyon culpaeus 92-7 Dusicyon culpaeolus 87-9 Vulpes bengalensis 91-1 Vulpes bengalensis 86-9 The skull of Dusicyon griseus has little to distinguish it from that of Dusicyon culpaeus except for its small size and lack of interparietal crest (absence of a crest appears to be associated with small size in the Canidae). Dusicyon fulvipes (Martin, 1837) Darwin's fox, Chiloe fox DISTRIBUTION. The southern part of the Island of Chiloe. This is one of the very many islands that lie off the coast of Chile between latitudes 40-45°, separated from the mainland by a narrow channel, the Gulf of Corcovado. DESCRIPTION. Smaller than Dusicyon griseus with a uniformly dark and rufous pelage. The ears, head and legs are tawny, the back dark grey with agouti guard hairs. The tail is neither long nor bushy but has a black tip. Skull as in D. griseus but smaller. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Dusicyon griseus 94-4 Dusicyon griseus 92-8 Dusicyon gymnocercus 94-3 Dusicyon gymnocercus 91-8 Dusicyon sechurae 93-1 Cerdocyon thous 90-0 Dusicyon culpaeolus 92-7 Dusicyon culpaeolus 89-1 Dusicyon inca 91-3 Vulpes chama 89-0 THE FAMILY CANIDAE 173 It appears that this canid has always been somewhat rare or otherwise very shy and not many specimens have been collected. Osgood (1943 : 72) described how he trapped a pair of adults on the beach in 1922. These two were very similar in pelage characters and skull conformation to Darwin's specimen. Osgood stated that there is a close agreement in characters between Dusicyon fulvipes and D. griseus and suggested that Darwin's fox is merely an island form of D. griseus rather than a separate species. The results of this analysis support Osgood's suggestion. REMARKS. The type specimen of this species was collected by Darwin and the skull and skin are now in the British Museum (no. 55.12.24.431) together with one other skull and skeleton (no. 51.11.8.4 (99&a) purchased from Mr Brandt). The identification of this second specimen is not certain, however, because the original entry in the catalogue has the word 'Chili', and there is no indication that the animal came from the Island of Chiloe. The following account of the 'fox' that Darwin collected may be quoted from his Voyage of the Beagle (1860 : 280) : 'December 6th. 1834. I*1 the evening we reached the island of San Pedro, where we found the Beagle at anchor. In doubling the point, two of the officers landed to take a round of angles with the theodolite. A fox (Canis fulvipes), of a kind said to be peculiar to the island, and very rare in it, and which is a new species, was sitting on the rocks. He was so intently absorbed in watching the work of the officers, that I was able, by quietly walking up behind, to knock him on the head with my geological hammer. This fox, more curious or more scientific, but less wise, than the generality of his brethren, is now mounted in the museum of the Zoological Society.' Dusicyon sechurae (Thomas, 1900) Sechura desert fox DISTRIBUTION. The arid coastal zone of northwestern Peru and southwestern Ecuador, including the Sechura desert (Cabrera, 1931, 1958). DESCRIPTION. A small light species with pale agouti guard hairs and fawn underfur. Cream to fawn underparts. Little or no rufous colouring on the body. Tail with distinct black tip. Skull small with lyriform sagittal area and no interparietal crest. Palatine bones extend backwards beyond the posterior edge of M2. Teeth small with 'fox-like' canines. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Dusicyon vetulus 94-5 Dusicyon vetulus 92-1 Dusicyon fulvipes 93-1 Vulpes chama 89-8 Dusicyon gymnocercus 92-6 Dusicyon australis 88-6 Vulpes pallida 91-0 Dusicyon fulvipes 88-5 Dusicyon culpaeolus 90-4 Vulpes velox 87-6 In pelage characters this species lies close to Dusicyon griseus and Dusicyon vetulus. Its small size may be an adaptation to desert conditions. 174 J- GLUTTON-BROCK ET AL. Dusicyon vetulus (Lund, 1839) Hoary fox DISTRIBUTION. The most northeastern of the species of Dusicyon that have been described so far. Found in south-central Brazil, Minas Gerais and Mato Grosso. DESCRIPTION. The smallest species of Dusicyon, similar in size to the smallest true foxes, for example Vulpes pallida. Pelage as for Dusicyon sechurae but with a marked dark stripe along the dorsal line of the tail. Skull small with faintly marked temporal ridges, a very narrow lyriform sagittal area and a slight interparietal crest. Teeth small with widely spaced premolars and reduced upper carnassial (P4). Canines sharply pointed and 'fox-like'. Anterior part of the frontal bones slightly swollen. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Dusicyon sechurae 94 '5 Dusicyon sechurae 92-1 Dusicyon gymnocercus 91-2 Vulpes bengalensis 86-0 Vulpes bengalensis 90-1 Dusicyon australis 85-4 Dusicyon fulvipes 90-0 Vulpes chama 85-2 Dusicyon griseus 89-9 Canis mesomelas 84-6 This species is noted for its small teeth and reduced carnassials which, combined with its somewhat isolated distribution in the central and eastern parts of the continent, have inclined previous authors to place it in a separate genus. The first description of the species was by Burmeister (1854 : 99) wno created the genus Lycalopex for it. This was followed by Gray (1868) and by all subsequent authors until Osgood (1934) reduced Lycalopex to a subgenus of Dusicyon. Cabrera (1958) and Simpson (1945) accepted this change and this nomenclature has been in general use up to the present. Langguth (1970, 1975) has, however, reverted to classifying the species in a separate genus, that is, Lycalopex vetulus. Although the two-dimensional plots show that Dusicyon vetulus lies somewhat on the edge of the Dusicyon group the analysis provides no evidence that the species should be separated at the generic level and for all phenetic characters it is clear that it lies very close to Dusicyon sechurae. The reduction in the size of the teeth may be more apparent than real for they are in proportion to the small size of the skull. Dusicyon thous (L.) Common zorro, crab-eating fox DISTRIBUTION. Savannah and woodland areas of northeastern South America, Columbia, Guiana, Brazil and south into northern Argentina (Cabrera, 1931, 1958 ; Hershkovitz, 1957 ; Langguth, 1970). THE FAMILY CANIDAE 175 DESCRIPTION. A fairly small, dark canid with a grizzled-brown or grey pelage. The legs may be tawny, underparts brownish-white and ears ochreous or rufous. The tail is fairly long, bushy and either totally dark or with a black tip. Ears short. The caecum was said by Garrod (1873) to be nearly straight rather than convoluted as in most canids. Temporal ridges faintly marked and enclosing a lyriform sagittal area. Frontal sinuses well developed and nasal bones slightly swollen in the facial region. Teeth large but canines not particularly long. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Dusicyon fulvipes 91-1 Dusicyon fulvipes 90-0 Dusicyon griseus 86-7 Dusicyon gymnocercus 88-4 Dusicyon microtis 86- 1 Dusicyon culpaeolus 86-2 Dusicyon gymnocercus 85-3 Dusicyon microtis 85-6 Dusicyon culpaeolus 85-3 Canis mesomelas 84-3 Following Thomas (1914) many authors have separated the zorro from Dusicyon and placed it in either the subgenus or genus Cerdocyon Hamilton Smith, 1839. Cabrera (1931) distinguished the species from Dusicyon at the generic level on the long dark tail, large feet and characters of the molar teeth and mandibular condyle. The present analysis shows that, although the species lies somewhat apart from the main Dusicyon group for some characters, for example the somewhat enlarged frontal sinuses and dark pelage, the numerical results provide no evidence that would justify separate generic status. Ducisyon microtis (Sclater, 1882) Small-eared zorro DISTRIBUTION. Tropical forests of the Amazonian basin in Brazil, Peru, Ecuador and Colombia. From sea level to 1000 m (Hershkovitz, 1957, 1961). Classified as rare by the Red data book (Goodwin & Holloway, 1972). DESCRIPTION. Larger than the common zorro with a large head, very short, rounded ears, short legs and a long bushy tail. Distinctive, dark, grizzled brown pelage with dark underparts except in the pelvic region where the hair is lighter in colour. The behaviour of this species in captivity has been described by Hersh- kovitz (1961). Temporal ridges strongly developed forming a raised, narrow, slightly lyriform sagittal area (as in Dusicyon australis). Frontal sinuses quite large as in Dusicyon thous, and nasal bones slightly swollen in the facial region. Canines long and 'fox-like'. Cheek-teeth robust. 176 J. CLUTTON-BROCK ET AL. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Dusicyon thous 86- 1 Cam's adustus 90-4 Cam's adustus 84-4 Dusicyon gymnocercus 89-7 Dusicyon fulvipes 84-3 Canis mesomelas 87-9 Dusicyon gymnocercus 82-9 Dusicyon culpaeolus 87-8 Dusicyon sechurae 82-9 Dusicyon sechurae 86-6 As with most other members of the South American Canidae there has been a fair amount of vacillation in the classification of the small-eared zorro. Thomas (1914) placed it with the common zorro in the genus Cerdocyon Hamilton Smith, 1839. Osgood (1934), on the other hand, believed it to be a true Dusicyon within the subgenus Dusicyon, whilst he placed only D. thous in the subgenus Cerdocyon. Cabrera (1940 : 14) considered the small-eared zorro to be quite distinct from Dusicyon and he placed it in a new genus Atelocynus Cabrera, 1940. Simpson (1945 : 109) noted the new genus but did not use it in his classification. Hersh- kovitz (1961), however, fully supported Cabrera and believed that the new genus was valid. His reasons were based on the combination of characters that appear to distinguish the small-eared zorro from the rest of the Dusicyon species ; these being the distinctive pelage, large size, small ears, large heavy teeth and development of the mandibular condyle as in D. thous. These characters were observed by Osgood who, nevertheless, retained the species within the genus Dusicyon. The results of this analysis show that Dusicyon microtis is phenetically fairly close to D. thous and that it lies on the periphery of the main Dusicyon group. It could only be argued that it should be given separate generic status if this was also done for D. australis. Hershkovitz (1972 : 390) believes that D. microtis is a specialized canid adapted to living in tropical rain forest areas. Genus CHRYSOCYON Hamilton Smith, 1839 One species. Chrysocyon brachyurus (Illiger, 1811) Maned wolf DISTRIBUTION. Tall grasslands and the outskirts of forests in eastern and southern Brazil, Paraguay, eastern Bolivia and northern Argentina (Cabrera, 1958 ; Hershkovitz, 1972 : 390). Classified as vulnerable by the Red data book (Goodwin & Holloway, 1972). DESCRIPTION. The largest of the South American canids with a very striking appearance, 'like a fox on stilts'. Shy and solitary, feeding on small prey and some vegetable matter. It is believed that it never digs, and indeed this might be difficult with its long legs. The pelage is distinctive and different from that of any other canid. The hair is long and reddish in colour over the whole body. Muzzle and THE FAMILY CANIDAE 177 chin dark, anterior part of throat white and inside of ears white. Feet black from the hocks, which are elongated, downwards. White tuft to rather short bushy tail. The hair along the nape of the neck and back is longer than the rest and dark coloured. Ears large and erect. Flower (1879) recorded that the caecum of a specimen that died in the Zoological Gardens was quite straight. Skull large and elongated. Frontal bones flat. Temporal ridges close and fused into a well-developed interparietal crest. Palatine bones extend slightly further back than the posterior edge of M2. Auditory bullae relatively very small. Teeth simple, widely spaced and 'fox-like'. Premolars simple and high-crowned. P4 short. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Canis simensis 79-9 Dusicyon gymnocercus 74-7 Dusicyon culpaeolus 74 '5 Dusicyon inca 73-4 Dusicyon microtis 73-4 Skull and teeth only Canis simensis 88-7 Canis adustus 88-7 Dusicyon culpaeolus 84-3 Dusicyon gymnocercus 83-6 Canis lupus 82-8 C. brachyurus clearly stands apart on its own. It is not a fox, as is often main- tained ; neither does it lie close to the Canis group, for although the 'near neighbours' tables do show a fairly high level of similarity with Canis simensis and Canis adustus these are the two species of Canis that are closest to the Dusicyon group. A rather low level of similarity with the genus Dusicyon is therefore probably the best inter- pretation of the affinities of the maned wolf and its position on the two-dimensional plots supports this view. Genus SPEOTHOS Lund, 1839 One species. Speothos venaticus (Lund, 1842) Bush dog DISTRIBUTION. Common throughout tropical rainforests and savannah areas in the Brazilian subregion of South America. Also found in one locality in southeastern Panama where Hershkovitz (1972 : 359) suggests that it may have been introduced by man (Cabrera, 1958). DESCRIPTION. Small, rather 'otter-like' with short legs and tail. Head heavy with a wide muzzle and small ears. Head and neck ochreous fawn or tawny merging into dark brown or black along the back and tail. Chin and underparts as dark as the back. There may be a light patch behind the chin on the throat. Skin of body yellow or tan in colour. The caecum is said to be straight as in Chrysocyon brachy- urus (Flower, 1880 : 73). The brain has relatively high and massive frontal lobes 178 J. CLUTTON-BROCK ET AL. (reflected in the swollen frontal lobes of the skull) and a relatively untwisted cere- bellar vermis (Radinsky, 1973). A social carnivore that hunts in packs of up to ten animals and swims well. As observed by Huxley (1880), the occiput is unique amongst canids in being drawn out into a short tube (unfortunately this character was missed and has not been taken into account in the numerical analysis). Facial region short with swollen frontal bones producing a slightly convex skull profile. Dentition reduced with M2 nearly always missing and M3 always absent, as in Cuon alpinus. Canine teeth 'dog-like', that is short and robust. Upper premolars 1-3 unusually thick in cross-section and with no posterior secondary cusps. The talonid or heel of the lower carnassial (Ma) has only one cusp as in C. alpinus and Lycaon pictus (Table 5). Symphysis of the mandible very long and strongly ankylosed. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Skull and teeth only Cuon alpinus 73-5 Cuon alpinus 87-0 Dusicyon microtis 68-2 Nyctereutes procyonoides 74-1 Lycaon pictus 67-9 Lycaon pictus 73-7 Dusicyon australis 67-8 Dusicyon australis 72-6 Bloodhound 67-5 Dingo 72-3 Speothos venaticus, L. pictus and C. alpinus have been placed in the subfamily Simocyoninae by Simpson (1945 : 109, 223) on palaeontological evidence. Accord- ing to Matthew (1930 : 128) there were two branches of primitive canids during the Miocene. One led to the present-day true canids (subfamily Caninae) whilst the second (the Simocyoninae), which was equally widespread and abundant, later became extinct except for these three representatives. The only diagnostic charac- ter that distinguishes the two groups is the development of the talonid of the lower carnassial as a single cusp or ridge in the Simocyoninae. In all other canids the talonid has two cusps and was described by Matthew as 'basined'. One of the objects of the present work was to test the validity of this grouping on phenetic grounds. While it does appear that the three species are closer to each other than to any other groups on the basis of cranial and dental characters, the overall similarities are very low and it seems best to refrain from emphasizing their very few points of resemblance. Whatever the validity of their common origin they have clearly diverged very greatly and their recognition as isolated monospecific genera seems appropriate. The bush dog is a highly social animal. Unfortunately there are no detailed studies of the behaviour of C. alpinus and the few observations that have been made on S. venaticus show that its behaviour patterns are markedly different from those of L. pictus. Some habits are shared, however. Both species practise communal sleeping and hunting, neither uses the gape or teeth-baring threats and neither has very highly developed tail-wagging behaviour (Kleiman, 1967). The behaviour of the bush dog is clearly interrelated with its body-shape and pelage characters. The ventral surface is seldom exposed because the animal has THE FAMILY CANIDAE 179 short legs and it therefore has no need for a colour contrasting with the back. The short legs may also be related to the lordosis-like posture held by the female during courtship. The female is said to lower her front legs and raise her hindquarters and tail, as cats do (Kleiman, 1967 : 368). Similarly the lack of facial markings is probably related to the exaggerated submissive grin which exposes the molar teeth rather than a paler cheek region as in the other social canids. Kleiman maintained (1967 : 371) that S. venations and L. pictus cannot be allied on their behaviour patterns. It would certainly be remarkable if they could be, as the bush dog is highly specialized for hunting in the tropical rainforests of Brazil and the Cape hunting dog (L. pictus) for following the migrating herds of large mammals in the African savannah. S Genus CUON Hodgson, 1838 One species. Cuon alpinus (Pallas, 1811) Dhole, red dog, Indian wild dog DISTRIBUTION. Montane forest areas of the Indian peninsula, Malaysia, Java, Sumatra, Burma and northwards into Korea, China and eastern U.S.S.R. Not found in Ceylon (Ellerman & Morrison-Scott, 1966). Formerly fairly common but now the distribution of the dhole is much reduced and it is rare. Classified as a vulnerable species by the Red data book (Goodwin & Holloway, 1972). DESCRIPTION. A fairly large 'dog-like' canid with rounded ears and a long, moderately bushy tail. The legs are rather short, the pelage an evenly tawny or dark red colour with slightly darker tail and lighter underparts. The winter coat may be yellowish-grey in cold regions. A social carnivore that lives and hunts in packs. As observed by Huxley (1880 : 276), there is a notable similarity between the skulls of Cuon alpinus and Lycaon pictus. In both species the facial region is short and wide, although more so in Lycaon than in Cuon, and the frontal and maxillary bones are swollen so that the skulls have a convex profile (as in Speothos venaticus) . The palatine foramina are long in both species and the nasal bones widen at the point where they meet the suture between the frontal and maxillary bones (the nasals are often described as having a sigmoid shape) . In both species the dentition is 'dog-like' and strongly developed except that M2 is reduced in size and, in Cuon, M3 is absent. Secondary posterior cusps are present on P2, P3, and on the lower premolars in both species. The talonid of the lower carnassial (Mx) has only one cusp in Cuon as it has in Speothos and Lycaon. This character was first observed by Major in 1872 (1900 : 834). i8o J. CLUTTON-BROCK ET AL. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Dusicyon australis 79-2 Dusicyon gymnocercus 78-2 Cam's latrans 77'9 Dingo 77-3 Canis aureus 77-0 Skull and teeth only Speothos venaticus 87-0 Lycaon pictus 80-9 Dingo 78-6 Dusicyon australis 76-0 Canis lupus 75-2 Pocock (1941 : 146) was not impressed with the assumed similarities between C. alpinus and L. pictus that had been described by previous authors and he was even less impressed by the similarities between Cuon and Speothos. Our numerical results, however, show that there are certain phenetic resemblances in the skulls and teeth of the three genera but the pelage and postcranial characters are widely different and although all three are social species without highly developed facial expressions it is not known whether there are any inherent behaviour patterns that link the three genera. Kleiman's comparative study (1967) did not include Cuon, and Lycaon is the only one of the three on which serious ethological studies have been carried out (van Lawick-Goodall, 1970 ; Kuhme, I965a, b). Although the dhole may resemble the dingo and the Indian pariah dog in colouring and superficial appearance, the skull and teeth are so distinctive that it is most unlikely that this species has contributed to the ancestry of the domestic dog. Genus LYCAON Brookes, 1827 One species. Lycaon pictus (Temminck, 1820) Hunting dog DISTRIBUTION. Formerly widespread throughout the African savannah south of the Sahara wherever game was abundant, up to 2700 m. Now becoming increasingly restricted to game reserves (Allen, 1939 ; Ellerman et al., 1953). Classified as a vulnerable species by the Red data book (Goodwin & Holloway, 1972). DESCRIPTION. A large canid with long legs and a heavy, rather 'hyaena-like' head. Ears large, rounded and nearly naked. Body hair may be scant. Mottled pelage which is variable in pattern and colouring. The irregular spots may be black, brown, grey or white, on a basic colour of yellowish-grey or black. The muzzle is dark and may have a dark stripe leading along the side of the head. Tail moderately bushy with a white tip. This species is the only member of the Canidae in which the first digit is absent or vestigial in the fore feet as well as the hind feet. Highly social but has never been domesticated. Short wide facial region with swollen frontal maxillary bones that give a convex shape to the skull profile, as in Cuon alpinus. The anterior palatine foramina are large and the nasal bones are wide. Frontal sinuses well developed. Interparietal THE FAMILY CANIDAE 181 crest may be pronounced. Dentition complete with strong, 'dog-like' canines and carnassials. Posterior secondary cusps are present on P2, P3 and on the lower premolars. The talonid of the lower carnassial, Mj, has only one cusp, as in Speothos and Cuon. SYSTEMATIC POSITION. Percentage similarity to 'near neighbours' : All characters Dusicyon australis 71-0 Cuon alpinus 69-7 Bloodhound 68-8 Dingo 67*9 Speothos venaticus 67-9 Skull and teeth only Dingo 81-8 Cuon alpinus 80-9 Canis lupus 79-4 Canis aureus 77 -6 Bloodhound 77-4 The numerical results show that Lycaon pictus is a most aberrant canid and there can be no dispute about its generic status. The phenetic relationships of this species with Speothos and Cuon have already been discussed in the sections on these genera. REMARKS. The hunting dog occupies the 'wolf niche' in Africa. The species has evolved a system of ritualized communal feeding whereby a whole pack can be sustained on the hunting efforts of a few individuals (Kuhme, I965a, b). This system is based on the regurgitation of food by the hunters for the juveniles and all members of the pack that have not joined in the killing of prey (usually antelope or gazelle). Many species of canid will regurgitate food for their young but in the hunting dog this habit is extended and has evolved into a basis for highly organized social behaviour. Although it is perhaps the most social of all canids the hunting dog has not evolved the elaborate facial expressions and signals of communication that are now so well known from studies of behaviour in the wolf. Fox (1970) suggested that the reasons for this are that the social organization of the hunting dog is based on individual dominant and subordinate relationships, mutual submission and strong group-orientated activities, rather than on a hierarchy of relationships as occurs within the wolf pack. He further suggests that communication by facial expression is important to groups of wolves that frequently undergo separation and congregation, whereas the hunting dog packs remain together as cohesive units for longer periods. Perhaps the strikingly individual markings of the hunting dog also assist in communication and identification of conspecifics. ACKNOWLEDGEMENTS Computer time for this project was kindly supplied by Rothamsted Experimental Station and we are most grateful to Miss Linda Jackson for her help in running the programs. Our thanks are also due to Miss Jenny Paul who assembled much of the data during tenure of a vacation studentship at the British Museum (Natural History). 182 J. CLUTTON-BROCK ET AL. APPENDIX I: DATA MATRICES TABLE 4 Characters of the skull i 2 3 4 5 6 7 8 9 10 II 12 13 Cam's lupus 226 65 33 5i 88 57 M 3 3 2 I 2 O Dingo 1 80 64 32 50 80 57 M 2 3 2 I 2 O Bloodhound 221 63 35 56 83 49 12 3 3 2 I 2 O Canis latrans 180 59 28 48 93 52 14 2 3 2 I 2 o C. aureus 1 60 65 34 52 85 57 16 2 3 2 I 2 o C. mesomelas 144 64 32 5i 79 59 16 2 3 2 I I o C. adustus 149 60 30 50 89 5i M 2 3 2 I I o C. simensis 183 52 25 49 94 53 13 I 2 2 I I o Vulpes vulpes 130 55 29 54 88 54 15 I 2 2 I O o V. corsac 112 58 3i 53 81 59 17 I I 2 O O o V. ferrilata 145 49 24 5i H5 57 15 I I 2 O O o V. bengalensis 112 58 29 50 89 56 18 I I 2 I o o V. cana 90 58 26 45 87 56 19 O O O O o 0 V. rueppelli I O2 57 27 47 94 55 20 I I 2 I o o V. pallida 98 59 30 5i 83 55 18 I O I I o o V. zerda 82 60 29 49 89 54 26 O O O o o o V. chama III 60 30 50 83 55 18 I I 2 I o o V. velox II4 56 26 46 89 54 17 I I I I o o V. cinereoargenteus 119 55 28 5i 85 56 16 2 O 2 2 o I Alopex lagopus I24 62 35 56 81 56 16 I 2 2 O o o Otocyon megalotis II4 54 32 60 81 54 *7 2 0 I 2 o 2 Nyctereutes procyonoides 110 59 35 59 83 57 17 I 2 2 2 2 I Dusicyon australis 157 64 34 53 75 15 2 I 2 I 2 0 D. culpaeus 165 53 24 45 124 55 14 2 3 2 I I o D. culpaeolus 139 57 29 50 9i 56 15 I 2 2 I I o D. gymnocercus 139 55 28 5i 9i 53 16 I I 2 I I 0 D. inca 147 55 28 5i 98 56 13 I 3 2 2 I o D. griseus 120 55 27 48 92 5i 15 O i I O I o D. fulvipes H5 59 30 50 93 54 16 O o I I I 0 D. sechurae 117 58 32 54 82 57 15 I i 2 I 2 o D. vetulus 104 65 36 55 74 59 19 I i 2 I 2 o D. thous 126 62 3i 50 90 53 15 O i I I I I D. microtis 149 60 32 54 94 58 15 2 i 2 I I I Chrysocyon brachyurus 213 54 28 52 95 56 12 3 3 2 I I o Speothos venaticus I24 67 4i 62 75 58 15 i 2 2 2 2 o Cuon alpinus I78 70 40 57 72 61 15 i 3 2 I 2 o Lycaon pictus 188 76 47 62 61 68 15 2 3 2 O 2 o THE FAMILY CANIDAE 183 TABLE 4 cont. Key to characters i. Condylobasal length 2. Palate - greatest width as % of length of palate (a : b). 3. Rostrum - width as % of length of palate (c : b). 4. Rostrum. - width as % of width of palate (c : a). 5. Premaxillae - anterior palatine length as % of width of rostrum (d : c). 6. Zygomatic width as % of condylobasal length (e : i). 7. Bullae - maximum length as % of condylobasal length (f : i). 8. Temporal ridges - size : o = absent ; 3 = highly developed. 9. Temporal ridges - proximity : o = wide apart ; 3 = fused. 10. Interparietal crest : o = absent ; 2 = well developed. 11. Parietal bones - rugosity : o = smooth ; 2 = distinctly rugose. 12. Post-orbital processes - convexity : o = concave; i = flat; 2 = strongly convex. 13. Mandible - size of subangular lobe : 0-2. i84 J. CLUTTON-BROCK ET AL. TABLE 5 Characters of the teeth upper lower deciduous i 2 3 4 5 6 7 8 1 — 9 10 II / — 12 — > 13 Canis lupus ii 50 2 ii 0 I 6 o I I o o I Dingo ii 48 2 ii o I 6 o I I 0 0 I Bloodhound ii 54 2 9 o I 6 0 I I o 0 I Canis latrans 12 42 2 ii 0 I 7 o I I 0 C. aureus II 50 2 IO o I 6 o I I 0 I I C. mesomelas II 41 I n o I 7 o I I o I I C. adustus II 40 I 9 o I 7 o I I 0 I I C. simensis 12 39 I 9 o I 6 o I I o I I Vulpes vulpes 12 37 I 10 o I 6 0 I I o I o V. corsac 12 41 I 10 o I 6 o I I o V. ferrilata 14 35 I 9 0 I 6 o I I 0 o I V. bengalensis 12 36 I 9 o I 8 0 I I o I 0 V. cana 9 38 0 ii o I 7 o I I o V. rueppelli 9 47 I IO o I 7 o I I 0 I o V. pallida IO 41 I 8 o I 8 0 I I o V. zerda 10 35 I 9 o I 8 o I I o I o V. chama 10 37 I 9 0 I 7 o I 1 0 V. velox II 37 I IO o I 6 0 1 I o V. cinereoargenteus 8 5i I 8 o I 7 0 I I o I o Alopex lagopus ii 49 A *7 2 10 o I 6 0 I I o I o Nyctereutes procyonoides 10 47 46 I . 9 o I 6 o I I 0 o Dusicyon australis 12 43 I ii o I 6 0 I I 0 D. culpaeus 13 36 I IO 0 I 7 o I I 0 I o D. culpaeolus II 41 2 IO o I 7 o I I o D. gymnocercus II 40 I 10 o I 7 0 I I o D. inca 13 39 2 II o I 7 o I 1 0 D. griseus II 37 I 10 0 I 7 o I I o I 0 D. fulvipes II 35 I IO o I 7 0 I I o D. sechurae 12 38 I 9 o I 7 o I I 0 I I D. vetulus 12 37 0 7 0 I 8 o I I o D. thous 9 46 I 10 o I 8 o I I 0 I I D. microtis 13 39 I 9 o I 7 o I I o Chrysocyon brachyurus II 44 O 8 o I 6 o I I 0 Speothos venaticus II 50 2 10 o 0 0 0 0 0 o o Cuon alpinus 10 53 2 ii 0 I 4 0 0 0 o 0 o Lycaon pictus 12 50 2 ii 0 I 5 o o I 0 o I THE FAMILY CANIDAE 185 TABLE 5 cont. Key to characters 1. C1 - height as % of condylobasal length (a : b). 2. C1 - alveolar length as % of height (c : a). 3. P3 - posterior cusps present : 0-2. 4. P4 (carnassial) - length as % of condylobasal length (d : b). 5. P4 - shape : o = carnassial; i = molariform. 6. M2 present : o-i. 7. M2 - greatest width as % of condylobasal length (e : b). 8. M3 present : o-i. 9. M! (carnassial) - two cusps on heel : o-i. 10. M 3 present : o-i. 11. M4 present : o-i. 12. DP3 - protocone developed as a cusp : o-i. 13. DP4 - posterior border concave, so that metacone appears as a separate lobe : o-i. d v P right M2 right MI left M1 left Lycaon pictus 186 J. CLUTTON-BROCK ET AL. TABLE 6 Pelage of head and body i 2 3 4 5 6 7 8 9 10 II 12 13 14 15 16 17 Canis lupus 2 C 0 o o o o o I o 2 o 2 I I 2 o Dingo 3 O 0 o o o I o o o o o I I o I o Bloodhound 2 I o o o o 0 o o 0 o o 2 I I o o Canis latrans I I o o 0 o o o I o I o 2 I I 2 o C. aureus I 2 o o o o 2 o I o I o 2 I I 2 o C. mesomelas I I o o o o 2 o I o 3 0 2 I I 2 o C. adustus I I o o o o 2 o o I o o 2 I I 2 o C. simensis O 3 o o o o I o I o o o I I I I o Vulpes vulpes I 3 o o I o 2 o o o o o I 2 I 2 o V. corsac I i 0 o I 0 2 o o o 0 o I I I I o V. ferrilata I i o o o o 2 o o o o o I I I 2 o V. bengalensis I i o o I o 2 o o o o o I I I 2 0 V. cana I o o o I o 2 o o o I o 0 I I 2 o V. rueppelli I i o o I o 2 o o o o o I I I 2 o V. pallida I i o o o o 2 o o o o o I I o I o V. zerda O i o o o o 2 o o o o o o I I 2 o V. chama I i o o o o 2 o 0 o o o I 2 I 2 o V. velox O i o o I o 2 o o o o o I I I 2 o V. cinereoargenteus I i o o I o 2 o o o o o I I I I o Alopex lagopus 2 o o o o o I o o o o o o 2 o 2 I Otocyon megalotis I I o I o o 2 0 0 o o o I 2 I 2 o Nyctereutes procyonoides I 2 0 o o I I o o o o o 2 2 I 2 o Dusicyon australis I 2 o o 0 o o o o 0 o I I I 2 o D. culpaeus I I o o o o 2 o o o I o I I I I o D. culpaeolus I I o o 0 o 2 o o o I o I I I 2 o D. gymnocercus I I o o o o 2 o o o o 0 I I I I o D. inca I I o o o o 2 o o o o o I I I I o D. griseus I I o o o o I o o o o o I I I I o D. fulvipes 2 I o o 0 o 2 o o o o o I I I I o D. sechurae I O o o o o 2 o 0 o o o I I I I o D. vetulus I I o o o o 2 o o o o o I I I I o D. thous 2 I o o o o I 0 I o I o 2 I I I o D. microtis 3 I o o o o I o o o o 0 2 o I I o Chrysocyon brachyurus i 3 o I o o I o 2 o 2 o I 2 o I 0 Speothos venaticus 2 i o o o o O o o o 0 I I O o O o Cuon alpinus O 3 o o o o O o o o o o 2 o o I o Lycaon pictus 2 2 I I 0 o O I o o o o I o 0 O o THE FAMILY CANIDAE 187 TABLE 6 cont. Key to characters 1 . Overall colour - intensity of black pigment : o = absent ; i = grey or banded hairs ; 2 = general appearance dark ; 3 = very dark. 2. Overall colour - intensity of red pigment: o = absent; i = present as yellow or red underfur ; 2 = general appearance reddish or tan ; 3 = extensive red colour. 3. Pelage boldly spotted : o-i. 4. Muzzle dark : o-i. 5. Facial mask between nose and eye : o-i. 6. Facial mask behind and below eye : o-i. 7. Mystacial vibrissae - length and thickness : 0-2 (Hildebrand, I952b). 8. Crown - dark median stripe : o-i. 9. Neck and back with crest or mane : 0-2. 10. Side - dark and light longitudinal bands : o = absent ; i = present. 11. Back -dark longitudinal band: o = absent; i = narrow stripe; 2 = wide stripe; 3 = saddle. 12. Ventral pelage dark : o = paler than rest of body ; i = dark. 13. Guard hairs - coarseness : 0-2. 14. Dorsal guard hairs - length in relation to body size : 0-2. 15. Dorsal guard hairs banded (agouti) : o-i. 16. Underfur - density : 0-2. 17. Seasonal colour change : o-i. i88 J. CLUTTON-BROCK ET AL. TABLE 7 Pelage of extremities ; other external characters ears tail fore legs hind legs r "\ f ^ I •\ 1 •\ I 2 3 4 5 6 7 8 9 IO II 12 13 14 15 16 17 Canis lupus II O o I 38 2 i 2 o I I o o 22 o o 10 Dingo II O o o 35 O o I 0 o I o o 20 i o Bloodhound 0 o o O o I o o I o o i o IO Canis latrans 13 O o o 33 2 I I o I I o o 21 o o 8 Canis aureus 12 O o o 35 I 2 o I I o o 17 o o C. mesomelas 15 O o o 44 2 2 o o I o o 22 o o C. adustus 12 O o o 52 I 2 o I I o o 22 o o C. simensis II O o o 25 I o 2 o o o o 20 o Vulpes vulpes 16 O I 59 2 o O o I I 0 o 25 o o 8 V. corsac 9 O o o 35 2 I 2 o 0 o o o V. ferrilata 8 0 o o 43 I I I o o o o 19 o V. bengalensis 15 o o o 58 2 I 2 o o o o 24 o 6 V. cana 18 o I o 71 2 0 I o o o I 21 o V. rueppelli 21 o o o 76 2 I O o 0 0 o 25 o V. pallida J7 o o o 60 I I 2 o o I o o 24 o o V. zerda 25 o o o 56 I I 2 o o I o 0 25 o o V. chama 20 o o o 69 2 I 2 o o o o 26 o V. velox 17 o o o 35 2 I 2 o o o o 26 o V. cinereo- argenteus 14 o o o 69 I 2 2 0 o I o o 24 o o 6 Alopex lagopus 9 I o o 59 2 O I o o I o o 24 o o 12 Otocyon megalotis 23 o I o 55 2 O 2 I I I 0 25 0 0 Nyctereutes procyonoides 9 I o I 37 I O I I I I 20 0 0 Dusicyon australis 7 o o o 29 I I 0 0 0 o o 19 o D. culpaeus 13 o o o 58 2 I 2 o o I o o 23 0 0 D. culpaeolus o o o 2 I 2 o o o o o D. gymnocercus 13 0 o o 53 2 I 2 o o o o 22 o D. inca 15 o o o 50 I I 2 o o o 22 o D. griseus 15 o o o 62 2 I 2 o o I o 24 o o D. fulvipes 14 o o o 41 I I 2 o o o 22 o D. sechurae 14 o o o 57 - I I 2 o o o o 22 o D. vetulus 12 o o o 54 2 I 2 o o o o 21 o D. thous IO o o o 45 I 2 2 o o I o I 21 o o D. microtis 10 o o o 43 I 2 2 I I 19 0 Chrysocyon brachyurus M o o o 22 2 O O I I I 22 o o Speothos venaticus 8 I o 0 24 O O I I I I 18 o I 8 Cuon alpinus 12 o o o 41 I O 2 o o I o o 24 o o 14 Lycaon pictus 14 I o I 36 I O O o o o o o 24 o 2 10 THE FAMILY CANIDAE 189 TABLE 7 cont. Key to characters 1. Ears -length as % of length of head and body (from skin labels and collector's notes, therefore only approximate). 2. Ears rounded : o-i. 3. Ears dark : o-i. 4. Ears - dark rim : o-i. 5. Tail - length as % of length of head and body (as for i). 6. Tail - bushiness : 0-2. 7. Tail - dark patch on dorsal surface (see Hildebrand, i952b) : o = absent ; 2 = long. 8. Tail - tip dark : o = white ; i = same as rest of tail ; 2 = black. 9. Fore legs entirely dark : o— i. 10. Fore legs with black line on front : o-i. 11. Fore feet — claws on digit i : o-i. 12. Hind legs dark : o— i. 13. Hind feet - dark plantar surface : o-i. 14. Hind feet - length as % of length of head and body (as for i). 15. Hind feet — claw on digit i : o-i. 1 6. Skin - darkly pigmented : 0-2. 17. Mammae -total number (from Hildebrand, i952b). J. GLUTTON-BROCK ET AL. TABLE 8 Body proportions ; post-cranial skeleton ; internal anatomy baculum [ ^ i 2 3 4 5 6 7 8 9 10 ii 12 13 14 Canis lupus 7i 79 39 i I 73 97 8 42 53 o o o Dingo 67 73 43 I I 70 1 02 6 40 Bloodhound 69 77 43 i I 99 8 39 Canis latrans 70 78 4i i o 45 o o 0 C. aureus 62 73 44 i I 67 97 7 44 34 o o 0 2 C. mesomelas 66 75 46 0 I 67 IOO 6 43 35 0 o o C. adustus 7i 77 43 o I 56 93 7 46 C. simensis Vulpes vulpes 70 81 38 0 0 7i 96 7 41 36 I o 0 2 V. corsac V. ferrilata V. bengalensis V. cana V. rueppelli 2 V. pallida 63 78 40 I o 76 85 8 52 4i o 0 O V. zerda 68 83 39 2 0 75 80 8 53 0 o O 2 V. chama 2 V. velox 66 78 39 V. cinereoargenteus 59 73 39 O o 63 96 8 50 43 o o O Alopex lagopus 70 79 34 I o 76 89 7 49 49 o o 0 2 Otocyon megalotis 70 82 42 I o 77 94 7 46 58 I o 0 2 Nyctereutes procyonoides 59 69 42 2 o 53 95 8 43 50 0 0 0 I Dusicyon australis 2 D. culpaeus 65 76 38 I o 63 93 7 4i 0 o O 2 D. culpaeolus D. gymnocercus 2 D. inca D. griseus 64 75 38 I 0 95 6 43 43 0 o O D. fulvipes 43 o o 0 D. sechurae D. vetulus 57 67 40 47 o o O D. thous 57 67 4i I o 54 48 43 o o 0 I D. microtis Chrysocyon brachyurus 92 103 47 2 o 60 95 6 49 34 o o I 0 Speothos venaticus 52 58 37 O o 7i 1 08 8 37 0 Cuon alpinus 60 70 35 2 o 60 104 8 42 o o O 2 Lycaon pictus 68 74 38 O I 68 97 7 41 52 I I O 2 THE FAMILY CANIDAE 191 TABLE 8 cont. Key to characters 1 . Fore legs - length as % of length of body spine (cervical to lumbar vertebrae) (from Hildebrand, iQ52a : fig. 6). 2. Hind legs - length as % of length of body spine (as above). 3. Neck -length of cervical vertebrae as % of combined length of thoracic and lumbar vertebrae (from Hildebrand, I952a : fig. 14). 4. Scapula - shape of teres major muscle scar on posterior angle : o = on posterior border only, with plane at right angles to lateral face ; i = intermediate ; 2 = whole scar on lateral face. 5. Scapula - extent of scar of serratus magnus muscle on medial side : o-i. 6. Pelvis - width as % of length (a : b). 7. Femur - length as % of length of tibia (c : d). 8. Femur - minimum width of shaft as % of length (f : c). 9. Third metatarsal - length as % of length of femur (e : c). 10. Baculum - length as % of condylobasal length. 11. Baculum - anterior end bifurcate : o-i. 12. Baculum - well-defined protuberance on dorsal keel : o-i (see Hildebrand, 1954, fig. 15). 13. Baculum - well-defined dorsal protuberance but no keel : o-i (as for 12). 14. Caecum - shape : o = straight ; i = nearly straight ; 2 = convoluted (from Flower, 1879, 1880 ; Garrod, 1873, 1878). Scapula Scapula (med.) Pelvis Femur Tibia Metatarsal IQ2 J. CLUTTON-BROCK ET AL. TABLE 9 Behaviour I Cam's lupus 2 Dingo i Bloodhound 2 3 4 5 6 7 8 9 10 ii 12 13 14 15 16 230002001001 121 O2OOOIOOIOOOI2I IIOIOOI 2 Canis latrans i 2IOOO2OOIOOII2I C. aureus O OOO2OOIOOII2I C. mesomelas o OIOOO OOIOOI 21 C. adustus o OIOOO OOIOOI 21 C. simensis Vulpes vulpes o V. corsac OOOOOOOOIOOOOI I I V. ferrilata V. bengalensis V. cana I V. rueppelli V. pallida V. zerda I V. chama V. velox I V. cinereoargenteus Alopex lagopus o IIOOO OOIO OOII Otocyon megalotis o I2OII OOIO 3O2I Nyctereutes procyonoides O2OII OOIO 3OOI Dusicyon australis D. culpaeus D. culpaeolus D. gymnocercus D. inca I IIOOO OOIO IO2 D. griseus D. fulvipes D. sechurae - D. vetulus D. thous o D. microtis Chrysocyon brachyurus o OOOOOOOOIOI2OIO Speothos venaticus 2 23001211110 02 Cuon alpinus 2 22OOO OOIO O2I Lycaon pictus 2 23IOI I 00 021 THE FAMILY CANIDAE 193 TABLE 9 cont. Key to characters (from Kleiman, 1966, 1967) 1. Diet - size of prey relative to body size : 0-2. 2. Diet - proportion of meat: o = varied - insects, vegetable, small vertebrates, carrion; 1 = varied - insects and small vertebrates ; 2 = mainly vertebrates. 3. Hunt socially : o = singly ; i = singly or in pairs ; 2 = pairs or family groups ; 3 = packs. 4. Ritual feeding : o-i. 5. Social grooming : o = rare and only between pairs ; i = well developed. 6. Communal sleeping : o-i. 7. Howling : o = absent or only as long-distance contact call ; i = present but no physical contact ; 2 = close-contact call, social howling in unison. 8. Frequency of oestrus phases for year : o = once ; i = twice. 9. Female courtship posture : o = normal standing position as in domestic dog ; i = crouch- ing position (lordosis) as in the cat. 10. Copulatory tie present : o-i. 11. Urination in a spray : o-i. 12. Defecation at specific sites : o-i. 13. Tail posture in dominant animals : o = no distinct posture ; I = straight and horizontal ; 2 = raised in a J-shape ; 3 = inverted U-shape. 14. Extent of teeth-baring in dominant threat posture : o-i. 15. Frequency of tail- wagging in submissive posture : 0-2. 16. Regularly occupies an underground den : o-i. 194 J- GLUTTON-BROCK ET AL. APPENDIX II: LICE (PHTHIRAPTERA) OF THE CANIDAE The identification of ectoparasites, particularly lice which are often rigidly host-specific, can sometimes expose interesting relationships between different groups of animals. The following species of lice are listed by Hopkins (1949) as known to parasitise members of the Canidae : Mallophaga, biting lice Trichodectes (Trichodectes) canis de Geer Cam's lupus f Domestic dog, including dingo * | Canis latrans f Canis aureus, one record from a captive host Dusicyon culpaeus, one record Dusicyon fulvipes, one record, apparently from a wild host Dusicyon thous f Felicola (Suricatoecus) vulpis Denny Vulpes vulpes * f Vulpes cinereoargenteus f Felicola (Suricatoecus) guinlei Werneck Otocyon megalotis * Felicola (Suricatoecus) fahrenholzi Werneck Dusicyon fulvipes * f Dusicyon sechurae, one record from a museum skin Heterodoxus spiniger Enderlein Domestic dog * f Canis latrans f Canis aureus f Canis adustus and Cuon alpinus (see Keler, 1971) Anoplura, sucking lice Linognathus setosus von Olfers Canis lupus, one record, apparently on a wild host Domestic dog * f Canis latrans, one record, no details Canis aureus, one record, apparently on a wild host Canis mesomelas f Vulpes vulpes, one record on a captive host Alopex lagopus f Linognathus taeniotrichus Werneck Dusicyon fulvipes, one record on a captive host Dusicyon thous * f * Nominal host for the species of louse listed. t Natural occurrence of the species of louse established on this canid. THE FAMILY CANIDAE 195 The genus Heterodoxus is particularly interesting for, with the single exception of Heterodoxus spiniger, its hosts are confined to Australian marsupials. H. spiniger has the domestic dog for its nominal host and has been recorded frequently from the coyote and jackals. Until recently this species of Heterodoxus was not known to occur on marsupials but there is now a record (Keler, 1971) for its presence on the wallaby, WallaUa agilis. Before this confirmed report Hopkins (1949) sug- gested that the species had evolved after transference to the dingo from a marsupial, perhaps shortly after dogs were introduced by man to Australia. This could have been in the early Holocene. The louse would then have spread to domestic dogs and thence to other wild canids as human populations moved about the world. At the present day H. spiniger is widespread on canids in many parts of Africa, Australia, America and Asia, but not apparently in Europe, Antarctica nor the northern regions of North America. Now that it is known, however, that H. spiniger does occur on a marsupial host it is possible that the transference to the dingo occurred at a later period ; on the other hand, its presence on the wallaby could be a secondary transference back to a marsupial host. The relationship between H. spiniger and its canid and marsupial hosts is obviously complicated, but it is possible that further work could throw light on the movements of human populations and the origins of their domestic dogs during the prehistoric period. Support for the inclusion of the American grey fox (Urocyon cinereoargenteus) within the genus Vulpes, as proposed in this classification, is given by the louse, Felicola vulpis, which has been identified from both the common fox and the grey fox. It was hoped there might be evidence for louse infestation on the skins of the extinct Falkland Island wolf, Dusicyon australis, and that this would lead to infor- mation on the relationships of this enigmatic canid. An examination (by Mr C. Moreby, British Museum (Natural History) ) of the two skins that are in the Museum collections failed to produce any signs of lice ; as incidentally did the mummified skin of an Ancient Egyptian dog, also in the collections. REFERENCES ALLEN, G. M. 1939. A checklist of African mammals. Bull. Mus. comp. Zool. Harv. 83 : 1-763- ANON. 1950. Bull. zool. Nomencl. 4 : 549. BLANFORD, W. T. 1888. The fauna of British India including Ceylon and Burmah. Mammalia. London. BOBRINSKII, N. A. 1965. Order Carnivora. In Bobrinskii, N. A., Kuznetsov, B. A. and Kuzyakin, A. P. [Key to the Mammals of the USSR]. Moscow (in Russian). BOWDICH, T. E. 1821. An analysis of the natural classifications of Mammalia. Paris : 40. BRINK, F. H. VAN DEN. 1973. 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Native mammals of Australia. Oxford. ROMER, A. S. 1955. Vertebrate palaeontology. Chicago. SCHENKEL, R. 1967. Submission : its features and functions in the wolf and dog. Am. Zool. 7 : 3I9-331- SCOTT, J. P. 1967. The evolution of social behaviour in dogs and wolves. Am. zool. 7 : 373-381. SIMPSON, G. G. 1945. The principles of classification and a classification of mammals. Bull. Am. Mus. nat. Hist. 85 : i-xvi, 1-350. SNEATH, P. H. A. & SOKAL, R. R. 1973. Numerical taxonomy. San Francisco. THOMAS, O. 1914. On various South American mammals. Ann. Mag. nat. Hist. ser. 8, 13 : 345-365- 1918. Some notes on the small sand-foxes of North Africa. Ann. Mag. nat. Hist. ser. 9, 1 : 243-245. TODD, N. B. 1970. Karyotypic fissioning and canid phylogeny. /. theor. Biol. 26 : 445-480. THE FAMILY CANIDAE 199 TROUESSART, E. L. 1885. Catalogue des mammiferes vivants et fossiles. Carnivora. Bull. Soc. Etud. Scient. Angers, 14 : 1-108. TROUGHTON, E. 1957. Furred animals of Australia. Sydney. VAN DER MERWE, N. J. 1953. The jackal. Fauna Flora Pretoria 4 : 4-77. VAN WORMER, J. 1964. The world of the coyote. Philadelphia & New York. WINGE, H. 1924. Pattedyr-Slaegter, vol. 2 : Rodentia, Carnivora, Primates. Copenhagen. WOOLPY, J. H. & GINSBURG, B. E. 1967. Wolf socialization : a study of temperament in a wild social species. Am. Zool. 7 : 357-363. JULIET CLUTTON-BROCK, Ph.D. G. B. CORBET, Ph.D. Department of Zoology M. HILLS, Ph.D. Department of Central Services BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) 1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96 ; 14 Plates. 1965. (Out of Print.) 2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125 ; 29 Plates, 77 Text-figures. 1969. £4.50. 4. HAYNES, J. R. Cardigan Bay Recent Foraminifera (Cruises of the R.V. Antur) 1962-1964. Pp. 245 ; 33 Plates, 47 Text-figures. 1973. £10.80. 5. WHITEHEAD, P. J. P. The Clupeoid Fishes of the Guianas. Pp. 227 ; 72 Text-figures. 1973. £9-70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : the Biology and Evolution of a Species Flock. Pp. 134 ; i Plate, 77 Text-figures. 1974- £3.75. Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol 884 sNU THE CRANIAL MUSCULATURE AND TAXONOMY OF CHARACOID FISHES OF THE TRIBES CYNODONTINI AND CHARACINI G. J. HOWES BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 29 No. 4 LONDON: 1976 THE CRANIAL MUSCULATURE AND TAXONOMY OF CHARACOID FISHES OF THE TRIBES CYNODONTINI AND CHARACINI ^£S% 2 designation by Benson (1972, p. 50, pi. n, fig. 15 ; nomen dubium.} Length 0-81 mm ; height 0-43 mm. Type locality: Stat. 146, deep-sea, east of Prince Edward's Island, 1375 fathoms. (46°46'o"S, 45°3i'o"E, trawled, Globigerina ooze, bottom temp. 35'6°F, surface temp. 43°F, December 29, 1873.) DESCRIPTION. Shape and ornamentation as described by Brady (1880). Eyespots well developed. OSTRACODS FROM THE CHALLENGER EXPEDITION 295 Cythere wyvillethomsoni Brady (PI. 13, figs 10 -18 ; PL 14, figs 1-3) Cythere wyvillethomsoni Brady, 1880 : 82, pi. 20, figs 4a-f. LECTOTYPE. Disarticulated left and right valves, BM 80.38. 123. Left valve: length 0-83 mm ; height 0-38 mm ; right valve : length 0-83 mm ; height 0-38 mm. Type locality: Christmas Harbour, Kerguelen Island, 20-50 fathoms (on slide) ; January 29, 1874. (Stat. 149 ; depth 120 fathoms on p. 17.) DESCRIPTION. Shape and ornamentation as described by Brady (1880). See PL 13, figs 10 and n. Inner lamella: both the anterior and the posterior vestibula are well developed with the anterior vestibule slightly larger. Marginal pore canals numerous on antero-ventral corner reducing in number towards dorsal corner. Mostly straight, a few branched and a few false. Posterior canals mostly straight and few in number, see PL 13, figs 12 and 15. Hinge holamphidont. Normal pores of open type. Central muscle scars : muscle scar pattern present not easily discerned. Eyespots present, but not well developed. Second antenna: see PL 13, fig. 18. Ob- tained from the lectotype. Penis: see PL 13, fig. 18. Obtained from the lectotype. REMARKS. Brady (1880, p. 82) reported this species from Stat. 149, Balfour Bay, Stat. 150, 151 and questionably from Stat. 185. He figured a female carapace (pi. 20, figs 4a-d) and a male (pi. 20, figs 46, f). Topotypic material: four specimens, BM 1974.349-52, of this species were recovered from sediment sample M-i85 (Stat. 151). Genus KRITHE Krithe hyalina Brady (PL 18, figs i, 2) Krithe hyalina Brady, 1880 : 115, pi. 27, figs 3a-d. LECTOTYPE. Whole carapace, BM 81.5.34. Length 0-59 mm ; height 0-31 mm. Type locality: Stat. 233b, Inland Sea, Japan, 15 fathoms. (34°i8'o"N, i33°35'o"E, trawled, mud, surface temp. 66-3°F, May 26, 1875.) DESCRIPTION. Shape and ornamentation as described by Brady (1880) and see PL 18, fig. i. Hinge adont. Krithe producta Brady (PL 17, figs 16-18) Krithe producta Brady, 1880 : 114, 115, pi. 27, figs xa-j. LECTOTYPE. Whole carapace, BM 80.38.127. Length 0-84 mm ; height 0-47 mm. Type locality: Stat. 146, 1375 fathoms. (46°46'o"S, 45°3i'o"E, trawled, Globigerina ooze, bottom temp. 35'6°F, surface temp. 43-o°F, December 29, 1873.) DESCRIPTION. Shape and ornamentation as described by Brady (1880). See PL 17, fig. 18. Hinge adont. Central muscle scars: see PL 17, figs 16 and 17. 2g6 H. S. PURI AND N. C. HULINGS REMARKS. Brady (1880, pp. 114, 115) lists this species from seventeen stations ranging in depth from 50 to 1675 fathoms and considered it to be cosmopolitan. The lectotype is from Stat. 146, depth 1375 fathoms. Krithe proditcta is common in sediment sample M-i69 (off Prince Edward's Island, 50-150 fathoms) and as a species should be split into several subspecies. Topotypic material: two left valves, BM 1974.353 & 355, and a right valve, BM 1974.354. Krithe tumida Brady (PI. 18, figs 3-5) Krithe tumida Brady, 1880 : 115, 116, pi. 27, figs 4a-d. LECTOTYPE. Whole carapace, BM 81.5.36 (valves separated after photography). Length 0-62 mm ; height 0-37 mm. Type locality : Stat. 323. (35°39'o"S, 50°47'o"W, trawled, grey ooze, 1900 fathoms, bottom temp. 33-i°F, surface temp. 73'5°F, February 28, 1876.) DESCRIPTION. Shape and ornamentation as described by Brady (1800). See PI. 18, fig. 3. Hinge adont. REMARKS. Brady (1880, pp. 115, 116) reported this species from Stat. 323 although he also lists (p. 13) Krithe tumida from Stat. 64 (35°35'o"N, 50°27'o"W, 2750 fathoms, grey ooze, June 20, 1873.) A left valve was recovered from sediment sample M-86 (Stat. 64) and is registered BM 1974.356. Genus LOXOCONCHA Loxoconcha africana Brady (PI. 18, figs 13, 14) Loxoconcha africana Brady, 1880 : 118, pi. 28, figs sa-d. LECTOTYPE. Disarticulated right and left valves, portion of anteroventral margin missing, BM 80.28.130. Right valve : length 0-59 mm ; height 0-40 mm ; left valve : length 0-59 mm ; height 0-37 mm. Type locality: St Vincent, Cape Verde, 1070-1150 fathoms, April 26, 1876. DESCRIPTION. Shape and ornamentation as given by Brady (1880), except for absence of papillae on lectotype. See PL 18, fig. 13. Inner lamella narrow anterior-h- and posteriorly with anterior and posteroventral vestibula. Marginal pore canals : about 17 simple canals. Hinge: aberrant amphidont type. In the right valve there is an anterior crescent-shaped socket within which there is a small tooth. The median element is crenulated. The posterior element consists of a divided tooth with a socket between. Normal pores large, scattered, round and elongate sieve-type. Central muscle scars: row of four adductor scars, single V-shaped frontal scar. Eyespot present. OSTRACODS FROM THE CHALLENGER EXPEDITION 297 Loxoconcha anomala Brady (PI. 18, figs 6-9) Loxoconcha anomala Brady, 1880 : 123, pi. 27, figs 5a-d. LECTOTYPE. Disarticulated right and left valves, BM 80.38.132. Right valve : length 0-65 mm ; height 0-43 mm ; left valve : length 0-62 mm ; height 0-43 mm. Type locality: from reefs at Honolulu, 40 fathoms, July 1875. DESCRIPTION. Shape and ornamentation as given by Brady (1880). See PI. 18, figs 6-9. Inner lamella wide throughout, line of concrescence and inner margin coincide ; no vestibula. Marginal pore canals about 22 branching, mostly bifurcate, a few trifurcate and single. Hinge adont. Central muscle scars: adductors consist of a vertical row of four elongate scars and a frontal scar. Eyespot present. REMARKS. Topotypic material: one specimen, a left valve (lost) and a right valve, BM 1974.360, was found in sediment sample M-324 (reefs at Honolulu). Loxoconcha australis Brady (PI. 18, figs 17, 18 ; PI. 19, figs 1-4) Loxoconcha australis Brady, 1880 : 119, 120, pi. 28, figs 5a-f ; pi. 29, figs 3a-d. LECTOTYPE. Left valve, BM 80.38.133. Length 0-78 mm ; height 0-37 mm. Type locality: Port Jackson, Australia, 2-10 fathoms, April 20, 1874. DESCRIPTION. Shape and ornamentation as given by Brady (1880). See PI. 19, figs 2 and 4. Inner lamella: see PI. 18, fig. 18 ; PI. 19, figs 2 and 3, the anterior vestibule is present but postero ventral vestibule absent. Inner lamella of about uniform width, selvage strong. Marginal pore canals : nine anterior and six ventral and posterior simple, straight canals. See PI. 19, fig. 3. Hinge: see PI. 19, fig. 3, aberrant amphidont, anteriorly a strong tooth followed by a crenulate bar. Pos- teriorly a large crescent-shaped socket with weak tooth. Normal pores scattered, large, rounded, sieve-type pore canals. Central muscle scars: vertical rows of four adductors, frontal scar kidney-shaped. Eyespot present and located low on carapace. REMARKS. Sediment sample M-242 (Stat. 187, off Booby Island, lat. io°36'o"S, long. i4i°5'o"E, 6-8 fathoms) yielded six complete carapaces, one of which was disarticulated. Topotypic material: disarticulated carapace, BM 1974.363. Loxoconcha honoluliensis Brady (PI. 19, figs 5, 6) Loxoconcha honoluliensis Brady, 1880 : 117, 118, pi. 28, figs 6a-f. LECTOTYPE. Disarticulated right and left valves, BM 80.38.136. Right valve : length 0-62 mm ; height 0-43 mm ; left valve : length 0-59 mm ; height 0-43 mm. Type locality: reefs off Honolulu, 40 fathoms, July 1875. 2g8 H. S. PURI AND N. C. RULINGS DESCRIPTION. Shape and ornamentation as given by Brady (1880). See PI. 18, figs 5 and 6. Lectotype and topotype pitted. Opaque areas present over most of the valves. Inner lamella widest anteriorly and posteroventrally. Line of con- crescence irregular and indistinct. Marginal pore canals: most of the branching type terminating in two, mostly three branches near the outer margin. Less than 30 canals somewhat evenly spaced throughout, a few false canals at the anterior end. Hinge adont. Right valve with a medium groove between two bars. Left valve with a median bar. Normal pores : sieve type, large and numerous. Central muscle scars: a vertical row of four elongate scars with the dorsal-most the longest and concave dorsally. Frontal scar V-shaped. Eyespots low and somewhat indistinct. REMARKS. Type locality for the lectotype, sediment sample M-324, yielded only two specimens, a right and a left valve. Loxoconcha honoluliensis was selected as type species of Loxoconchella by Triebel, 1954. Topotypic material: a right valve, BM 1974.361 ; a left valve lost. Loxoconcha pumicosa Brady (PL 18, figs 10-12) Loxoconcha pumicosa Brady, 1880 : 118, 119, pi. 28, figs 2a-d. LECTOTYPE. Whole carapace, BM 81.5.37 (left valve lost after photography). Length 0-50 mm ; height 0-34 mm. Type locality: Nares' Harbour, Admiralty Islands, 16 fathoms, March 2, 1875. DESCRIPTION. Shape and ornamentation as described by Brady (1880). See PI. 18, figs 10 and n. Inner lamella widest anteriorly, anterior and posteroventral vestibula present. Selvage strong. Marginal pore canals simple and straight. Hinge : aberrant amphidont hinge. Anteriorly, a crescent-shaped socket with a weak tooth followed by a crenulate bar. Posteriorly, a large crescent-shaped and incompletely divided tooth with a small socket inside the crescent. Eyespot present but inconspicuous. REMARKS. Two complete carapaces of this species were recovered from sediment sample M-279 (Nares' Harbour). Brady (1880, pp. 118, 119) also reported this species from off Booby Island (Stat. 187). Topotypic material: a complete carapace, BM 1974-359- Loxoconcha subrhomboidea Brady (PL 18, figs 15, 16) Loxoconcha subrhomboidea Brady, 1880 : 121, pi. 28, figs 4a-d. LECTOTYPE. Whole carapace, Hancock Museum (left valve missing after photography). Length 0-37 mm ; height about 0-24 mm. Type locality : Stat. 140, Simon's Bay, South Africa, 15-20 fathoms, October, 1873. DESCRIPTION. Shape as given by Brady (1880) except that the posterior is more angular than rounded and the caudal process is present but weak (see PL 18, fig. 15). OSTRACODS FROM THE CHALLENGER EXPEDITION 299 Ornamentation as given by Brady (1880) plus a heavy ventral ridge and deep furrow (see PI. 18, fig. 15). Inner lamella: anterior and posteroventral vestibula present. Well-developed posteroventral flange. Marginal pore canals simple and straight. Hinge: median element smooth. Posterior element comprises a large crescentic tooth which is incompletely divided, and a small socket inside the crescent. Normal pores few, large rounded sieve-type pores. REMARKS. Topotypic material: a left valve, BM 1974.362, was recovered from sediment sample M-i64 (Stat. 140, Simon's Bay), this is the only locality where Brady (1880, p. 121) found this species. Genus XESTOLEBERIS Xestoleberis africana Brady (PI. 19, figs 15, 16) Xestoleberis africana Brady, 1880 : 126, pi. 30, figs 4a-c. LECTOTYPE. Disarticulated right and left valves. Portion of anteroventral region missing on left valve, BM 81.5.40 (valves separated after photography). Right valve : length 0-40 mm ; height 0-31 mm ; left valve : length 0-40 mm ; height 0-28 mm. Type locality: Stat. 140, Simon's Bay, South Africa, 15-20 fathoms, October 1873. DESCRIPTION. Shape and ornamentation as given by Brady (1880) except for absence of papillae. Inner lamella widest anteriorly narrowing posteroventrally ; anterior and posteroventral vestibula present. Marginal pore canals short, simple canals throughout. About 20 canals, anteriorly. Hinge merodont. Right valve with terminal crenulated teeth. Crenulations continue into groove for median bar of left valve. Median bar smooth. Normal pores large, of sieve type. Central muscle scars: four elongate adductor scars, others obscure. Eyespot: eyepit large and 'Xestoleberis spot' small. The spot, located low, is short and semicircular in shape. REMARKS. This species was reported by Brady (1880, p. 126) from only one station (Simon's Bay). Topotypic material: sediment sample M-i64 (Stat. 140) yielded a single right valve, BM 1974.388. Xestoleberis expansa Brady (PL 19, figs 13, 14) Xestoleberis expansa Brady, 1880 : 129, 130, pi. 30, figs 3a-d. LECTOTYPE. Left valve, BM 81.5.41. Length 0-31 mm ; height 0-19 mm. Type locality: Stat. 323, off Uruguay, 1900 fathoms. (35°39'o"S, 50°47'o"W, trawled, grey ooze, bottom temp. 33'i°F, surface temp. 73'5°F, February 28, 1876.) DESCRIPTION. Shape and ornamentation as given by Brady (1880). The surface is extremely smooth and polished. Hinge merodont. Central muscle scars consist of four scars and a frontal scar. Eyespot: eyepit and 'Xestoleberis spot' present. 300 H. S. PURI AND N. C. RULINGS REMARKS. Brady (1880, p. 129) found one specimen, a complete carapace, which he figured (pi. 30, figs 3a-d). The British Museum slide catalogued as BM 81.5.41 contained two specimens, a complete carapace figured here on PI. 19, figs 13 and 14, which, unfortunately, is not a Xestoleberis. Consequently the other specimen, a left valve, is designated lectotype. Xestoleberis foveolata Brady (PL 19, figs n, 12) Xestoleberis foveolata Brady, 1880 : 130, pi. 30, figs ia-g. LECTOTYPE. Disarticulated right and left valves, BM 80.38.141 (valves separated after photography). Right valve : length 0-54 mm ; height 0-37 mm ; left valve : length 0-54 mm ; height 0-40 mm. Type locality: Stat. 187, Booby Island, north of Australia, 6-8 fathoms. (io°36'o"S, i4i°55'o"E, dredged, coral and sand, surface, temp. 77'7°F, September 9, 1874.) DESCRIPTION. Shape and ornamentation as given by Brady (1880). See PL 19, fig. ii. Inner lamella widest anteriorly ; anterior and postero ventral vestibula present, the former is wider. Marginal pore canals simple, straight and short. Most numerous anteroventrally. Hinge merodont. Right valve with anterior and posterior crenulate teeth connected by a smooth ridge with a groove above. Left valve with smooth median bar and crenulate sockets joined by a narrow groove and above the latter, an accommodation groove. Normal pores large and open. Central muscle scars: four elongate adductor scars arranged vertically and a frontal scar. Eyespots: eyepit small and not visible externally. 'Xestoleberis spot' long, slender and uniform in width. Overlap: left valve larger than right valve. REMARKS. The type locality is Stat. 187 and topotypes have been recovered from sediment sample M-242 (Stat. 187). Topotypic material: two carapaces, a left and a right valve, BM 1974.364 and 366-8. Xestoleberis granulosa Brady (PL 19, figs 17, 18) Xestoleberis granulosa Brady, 1880 : 125, 126, pi. 30, figs 5a-d. LECTOTYPE. Disarticulated right and left valves, Hancock Museum (valves separated after photography). Right valve : length 0-61 mm ; height 0-33 mm ; left valve: length 0-63 mm ; height 0-35 mm. Type locality: Port Jackson, Australia, 2-10 fathoms, April 20, 1874. DESCRIPTION. Shape and ornamentation as given by Brady (1880), except for the absence of papillae. Inner lamella wide anteriorly, narrowing towards posterior. Lamella widest posteriorly in region of posteroventral vestibule. Anterior vestibule wider than posterior. Line of concrescence more irregular in anteroventral region, less irregular elsewhere. Marginal pore canals most numerous anteriorly, with OSTRACODS FROM THE CHALLENGER EXPEDITION 301 about 35 simple canals and a few false canals. Concentrated anteroventrally, and less numerous ventrally and posteriorly. Some branching canals occur ventrally. Hinge merodont. Right valve with anterior and posterior crenulate teeth. Left valve with median bar. Bar as prominent as the teeth. Normal pores large and of sieve type. Central muscle scars: a vertical row of four elongate scars and a frontal scar. Eyespot: eyepit obscure. 'Xestoleberis spot' large and about twice as long as wide. Overlap: left valve larger than right. REMARKS. Brady (1880, pp. 125, 126) reported this species from only two localities (Stat. 162, East Moncceur Island, Bass' Strait, 38-40 fathoms, and Port Jackson). The lectotype is from Port Jackson and 10 complete carapaces and detached valves were recovered from sediment sample M-ig8 (Port Jackson). Topotypic material: a complete carapace, BM 1974.323, and a right valve, BM 1974.324. Xestoleberis nana Brady (PI. 20, figs 14, 15) Xestoleberis nana Brady, 1880 : 126, pi. 31, figs 5a-c. LECTOTYPE. Right valve, BM 80.38.143. Length 0-43 mm ; height 0-28 mm. Type locality: Stat. 172, off Nukualofa, Tongatabu, 18 fathoms. (20°58'o"S, i75°09'o"W, dredged, coral, surface temp. 75-o°F, July 22, 1874.) DESCRIPTION. Shape and ornamentation as given by Brady (1880), see PI. 20, figs 14 and 15. Inner lamella: see PI. 20, fig. 15. Widest anteriorly, vestibula present anteriorly and postero ventrally. Marginal pore canals short, simple and straight ; about 30 anteriorly with most of them concentrated anteroventrally. Hinge merodont. Right valve with anterior and posterior crenulate teeth and median groove. Normal pores large and open. Many with a 'halo' around the pore on the surface. Central muscle scars: a vertical row of four adductor scars and additional scars. Eyespot: eyepit indistinct, 'Xestoleberis spot' of two distinctly separate slender filaments, one below the other. Both slightly convex anteriorly. Xestoleberis setigera Brady (PL 20, figs 9-11) Xestoleberis setigera Brady, 1880 : 125, pi. 31, figs 2a-d and figs 3a-c. LECTOTYPE. Disarticulated right and left valves. Portion of ventral surface of right valve missing, BM 80.38.145 (valves separated after photography). Right valve : length 0-59 mm ; height 0-31 mm ; left valve : length 0-59 mm ; height 0-25 mm. Type locality: off Prince Edward's Island, 50-150 fathoms. (46°48'o"S, 37°49'3°"E, dredged, grey sand, surface temp. 4i-o°F, December 26, 1873.) DESCRIPTION. Shape and ornamentation as given by Brady (1880), except for the papillae which are absent in the lectotype but are present in the topotype. Inner 16 302 H. S. PURI AND N. C. RULINGS lamella widest anteriorly ; anterior and ventral vestibula present with the former the larger. The ventral vestibule is very narrow and terminates at the postero- ventral corner. Marginal pore canals most abundant anteriorly with about 20 simple straight and short canals plus a few false canals anteroventrally. Ventral canals simple and straight. Hinge merodont. Right valve with anterior and posterior crenulate teeth, left valve with smooth median bar. Normal pores large and open. Central muscle scars consist of four scars, the uppermost in the shape of a wide and shallow U. Frontal scar heart-shaped. Eyespot : eyepit present, 'Xestoleberis spot' club shaped. Overlap: left valve larger than right. REMARKS. Brady (1880, p. 125) reported this species from three places (off Christmas Harbour, Kerguelen Island, 120 fathoms ; the station from which he figured this species ; Stat. 151 and off Prince Edward's Island). The lectotype is from Prince Edward's Island. Topotypic material: a complete carapace, BM 1974.369, from sediment sample M-i69, Prince Edward's Island. Xestoleberis tumefacta Brady (PI. 20, figs 12, 13 ; PI. 26, figs 1-3) Xestoleberis tumefacta Brady, 1880 : 128, 129, pi. 31, figs 4a-d. NEOTYPE. Complete carapace, BM 1974.370. Length 0-54 mm ; height 0-33 mm. Type locality: Nares' Harbour, Admiralty Islands, 16 fathoms, March 2, 1875- DESCRIPTION. Shape as described by Brady (1880). See PI. 26, figs 1-3. Ornamentation, smooth, see PI. 26, figs I and 2. Inner lamella wide throughout. Hinge merodont. Eyespot: external eyespot absent, but internal 'Xestoleberis spot' is present. REMARKS. Brady (1880, pp. 128, 129) apparently was dealing with two different forms ; the form figured by him (pi. 31, figs 4a-d) is a true Xestoleberis but in his description he says, 'This has very much the general aspect of Loxoconcha' '. The only specimen in the BM collection (BM 81.5.44) (see PI- 2O> figs I2 and I3) *s a Loxoconcha, the specimen of Xestoleberis being lost. Brady found Loxoconcha pumicosa and Xestoleberis tumefacta only at Nares' Harbour (see p. 24) and several specimens of the only Xestoleberis present at this station were found in sediment sample M-279. One of these is here made neotype and figured (see PI. 26, figs 1-3). Topotypic material: two right valves, BM 1974.371-2. Xestoleberis variegata Brady (PI. 20, figs 16-18) Xestoleberis variegata Brady, 1880 : 129, pi. 31, figs 8a-g. LECTOTYPE. Disarticulated right and left valves, BM 80.38.146. Right valve : length 0-52 mm ; height 0-37 mm ; left valve : length 0-50 mm ; height 0-34 mm. OSTRACODS FROM THE CHALLENGER EXPEDITION 303 Type locality: Stats 93, 94, off St Vincent, Cape Verde, 1070-1150 fathoms. (i6°42'o"N, 25°i2'o"W, mud, surface temp. 78-o°F, August 5, 1873.) DESCRIPTION. Shape as given by Brady (1880) for the left valve, except for the absence of a ventral sinus ; the right valve is characterised by a flat dorsal surface in the region of the median bar/groove and a truncated posterior end, especially the dorsal half. See PI. 20, figs 16-18. Ornamentation: the lectotype has only two small opaque spots, both posteriorly rather than variegated all over as indicated by Brady (1880). Inner lamella: anterior and posteroventral vestibula present with the former the widest. Marginal pore canals simple, straight canals, in excess of 20 anteroventrally. Also, numerous ventrally, but decreasing in number posteriorly. Some false canals present. Hinge merodont. Right valve with anterior and pos- terior crenulate teeth. Left valve with median smooth bar. Normal pores large and open. Central muscle scars: a vertical row of four adductor scars; frontal scar kidney shaped. Eyespot: ' Xestoleberis spot' long and of uniform width except at the dorsal tip where it is narrow. The spot has a coarse granular appearance. Overlap: left valve larger than right. Genus CYTHERURA Cytherura clavata Brady (PL 19, figs 7-10) Cytherura clavata Brady, 1880 : 133, pi. 29, figs ya-d. LECTOTYPE. Left valve, BM 80.38.148. Length 0-60 mm ; height 0-29 mm. Type locality: Stat. 316, Stanley Harbour, Falkland Islands, 6 fathoms. (5i°32'o"N, 58°o6'o"W, dredged, 4 fathoms, mud, surface temp. 5i-2°F, February 3, 1876.) DESCRIPTION. Shape and ornamentation: see Brady (1880). Cytherura clausi Brady (PL 21, figs 9, 10) Cytherura clausi Brady, 1880 : 134, pi. 32, figs 8a-d. LECTOTYPE. Left valve, BM 81.5.47. Length 0-49 mm ; height 0-26 mm. Type locality: Stat. 140, Simon's Bay, South Africa, 15-20 fathoms, October 1873. DESCRIPTION. Shape and ornamentation as given by Brady. Surface strongly reticulate. Inner lamella very wide anteriorly where the inner margin runs a normal course ; very wide posteriorly where it forms a strong curve. Normal pores moderately small, numerous, open. REMARKS. Brady's syntypes (BM 81.5.17 and BM 81.5.45) labelled 'Cytherura clausi' really represent C. mucronata Brady. Several specimens of C. clausi Brady were found in sediment sample M-i64. Hornibrook (1952, p. 51, pi. 15, figs 242- 244) reported and figured Cytherura clausi Brady from New Zealand, conspecific with the form described here. 304 H. S. PURI AND N. C. HULINGS Cytherura costellata Brady (PL 21, figs 7, 8) Cytherura costellata Brady, 1880 : 134, pi. 32, figs ya-d. LECTOTYPE. Right valve, BM 80.38.149. Length 0-47 mm ; height 0-23 mm. Type locality: Stat. 149, Balfour Bay, Kerguelen Island, 20-50 fathoms, mud, January 1874. DESCRIPTION. Shape and ornamentation as described by Brady (1880). Cytherura cribrosa Brady (PI. 21, fig. 2) Cytherura cribrosa Brady, 1880 : 132, pi. 32, figs 5a-d. LECTOTYPE. Whole carapace, BM 80.38.150. Length 0-59 mm ; height 0-37 mm. Type locality: Stat. 305, 160 fathoms. (47°48'o"S, 74°46'o"W, trawled, mud, surface temp. 55-o°F, January i, 1876.) DESCRIPTION. Shape and ornamentation as described by Brady (1880). Cytherura cryptifera Brady (PL 21, fig. i) Cytherura cryptifera Brady, 1880 : 134, 135, pi. 32, figs 4a-c. LECTOTYPE. Left valve (damaged) Hancock Museum. Length 0-40 mm ; height 0-22 mm. Type locality: Stat. 162, off East Moncceur Island, Bass Strait, 38-40 fathoms. (39°io'3o"S, i46°37'o"E, dredged, sand, surface temp. 63-2°F, April 2, 1874.) DESCRIPTION. Shape and ornamentation as described by Brady (1880). Cytherura curvistriata Brady (PL 21, fig. 13) Cytherura curvistriata Brady, 1880 : 131, pi. 32, figs loa-d. LECTOTYPE. Whole carapace, BM 81.5.46. Length 0-37 mm ; height 0-21 mm. Type locality: Port Jackson, Australia, 2-10 fathoms, April 20, 1874. DESCRIPTION. Shape and ornamentation as described by Brady (1880). Cytherura lilljeborgi Brady (PL 21, figs 3-6) Cytherura lilljeborgi Brady, 1880 : 132, 133, pi. 32, figs 6a-d. LECTOTYPE. Whole carapace, BM 80.38.151. Length 0-43 mm ; height 0-25 mm. Type locality: Stat. 149, Balfour Bay, Kerguelen Island, 20-50 fathoms, mud, January 1874. DESCRIPTION. Shape and ornamentation as described by Brady (1880). OSTRACODS FROM THE CHALLENGER EXPEDITION 305 Cytherura mucronata Brady (PI. 21, figS II, 12) Cytherura mucronata Brady, 1880 : 133, 134, pi. 32, figs ga-d. LECTOTYPE. Whole carapace, BM 81.5.45. Length 0-50 mm ; height 0-28 mm. Type locality: Stat. 140, Simon's Bay, South Africa, 15-20 fathoms, October 18, DESCRIPTION. Shape and ornamentation as described by Brady (1880). See PL 21, fig. n. REMARKS. Brady (1880, pp. 133, 134) described both Cytherura mucronata and Cytherura clausi from Stat. 140 (Simon's Bay). The lectotype of C. mucronata (BM 81.5.45) and syntype specimens labelled by Brady as 'C. clausi' really represent C. mucronata. Several specimens of Brady's C. clausi were found in sediment sample M-i64 (Stat. 140). Genus CYTHEROPTERON Cytheropteron abyssorum Brady (PI. 23, fig. 8) Cytheropteron abyssorum Brady, 1880 : 138, pi. 34, figs 3a-d. LECTOTYPE. Whole carapace, BM 81.5.49. Length 0-37 mm ; width (from ala of right to ala of left valve) 0-50 mm. Type locality: Stat. 160, Southern Australian Basin, 2600 fathoms. (42°42'o"S, i34°io'o"E, trawled, red clay, bottom temp. 33'9°F, surface temp. 55'0°F, March 13, 1874.) DESCRIPTION. Shape: a large portion of each valve is missing and no attempt was made to study the specimen in detail. The description given by Brady (1880) fits the lectotype. Cytheropteron (?) angustatum Brady (PI. 23, figs 15-17) Cytheropteron (?) angustatum Brady, 1880 : 137, pi. 34, figs $&, b ('angustum' on explanation to Pi- 34)- LECTOTYPE. Left valve, BM 80.38.152. Length 0-46 mm ; height 0-28 mm. Type locality: Stat. 149, Balfour Bay, Kerguelen Island, 20-50 fathoms, mud, January 1874. DESCRIPTION. Shape as given by Brady (1880), except that the entire dorsal margin slopes towards the posterior. See PI. 23, figs 15 and 16. Ornamentation as given by Brady (1880). Inner lamella widest at anterior end. Line of concrescence and inner margin coincide throughout, no vestibule at either end. Hinge of Infra- cytheropteron type (holoperatodont) . Central muscle scars: see PI. 23, fig. 17. REMARKS. Brady (1880, p. 137) reported this species from Stat. 149 (Balfour Bay) and Stat. 185 (Torres' Straits, 155 fathoms) and he figured a left valve (pi. 34, 306 H. S. PURI AND N. C. HULINGS figs 5a, b). The lectotype is also a left valve from Stat. 149. Topotypic material: sediment sample M-237 (Stat. 185) yielded a single right valve, BM 1974.375. Cytheropteron assimile Brady (PI. 23, figs 1-7) Cytheropteron assimile Brady, 1880 : 138, 139, pi. 34, figs 2a-d. LECTOTYPE. Whole carapace, BM 80.38.153. Length 0-70 mm ; height 0-42 mm. Type locality: Stat. 151, off Heard Island, 75 fathoms. (52°59'3o"S, 73033'3°"E, dredged, mud, surface temp. 36-2°F, February 7, 1874.) DESCRIPTION. Shape as described by Brady (1880), except for posterior portion of dorsal margin. See PL 23, fig. i. Ornamentation as given by Brady (1880). Inner lamella: vestibule present at anterior end. Otherwise line of concrescence and inner margin coincide. Marginal pore canals few in number and most are simple. Hinge with only the terminal portions of the median element crenulate, see PL 23, figs 3-7. Normal pores open and scattered. Central muscle scars: four undivided adductors. Frontal scar single and V-shaped. REMARKS. Brady (1880, pp. 138, 139) described this species from off Christmas Harbour (Stat. 149) and off Heard Island (Stat. 151). The lectotype is from Stat. 151. Topotypic material: this species is common in sediment sample M-i83 (Stat. 149) and two right valves and a left valve are registered, BM 1974.376-8. Cytheropteron fenestratum Brady (PL 23, fig. 18 ; PL 24, figs 1-6) Cytheropteron fenestratum Brady, 1880 : 139, 140, pi. 34, figs 6a-d. LECTOTYPE. Right valve, BM 80.38.154-155. Length 0-93 mm ; height 0-56 mm. Type locality: Stat. 149, off Christmas Harbour, Kerguelen Island, 120 fathoms. (48°43'45"S, 69°6'i5"E, dredged, mud, surface temp. 38-8-39-0^, January 29, 1874.) DESCRIPTION. Shape and ornamentation as described by Brady (1880), except that the ventral surface is not 'irregularly nodulated'. Inner lamella: see PL 23, fig. 1 8. Line of concrescence and inner margin coincide except anteriorly where a vestibule is present. Marginal pore canals numerous and mostly simple. Hinge: see PL 24, figs 5 and 6. Terminal elements lobed, groove of median element smooth. Normal pores, numerous, scattered and open. Central muscle scars: five adductor scars arranged in a vertical row ; middle three more elongate than the dorsal- or ventralmost scars. Single frontal scar. REMARKS. Brady (1880, pp. 139, 140) reported this species from two stations (149 and 335). He figured a complete carapace (pi. 34, figs 6a-d). The lectotype is a right valve from Stat. 149. Topotypic material: sediment sample M-i83 (Stat. 149, off Christmas Harbour) yielded a right valve, BM 1974.379. OSTRACODS FROM THE CHALLENGER EXPEDITION 307 Cytheropteron mucronalatum Brady (PI. 22, figS 14-18) Cytheropteron mucronalatum Brady, 1880 : 140, 141, pi. 33, figs 8a-d. LECTOTYPE. Left valve, BM 80.38.157. Length 1-30 mm ; height 0-84 mm. Type locality: Stat. 296, near the Chile Rise in the eastern Pacific, 1825 fathoms. (38°6'o"S, 88°2'o"W, trawled, Globigerina ooze, bottom temp. 35'3°F, surface temp. 59'8°F, November 9, 1875.) DESCRIPTION. Shape: the lectotype differs from description of C. mucronalatum by Brady (1880) in the following ways : height is not equal to more than two-thirds of the length, anterior end scarcely dentate, posterior end devoid of spines, dorsum not broadly arched, see PI. 22, figs 15 and 18. Ornamentation: the elevated ridge mentioned by Brady (1880) is practically absent anteriorly and does not terminate in a strong short spine on the lectotype. Right valve does show the spine. Inner lamella: line of concrescence and inner margin coincide, vestibula absent. Marginal pore canals few, mostly simple, few false. Hinge: median element smooth. Normal pores, scattered and open. Central muscle scars: see PI. 22, figs 15 and 17. REMARKS. Brady (1880, pp. 140, 141) described this species from Stats 70, 224, 246, 296, 300 and 302 and he figured a complete carapace (pi. 33, figs 8a-d). The lectotype is a right valve from Stat. 296. Topotypic material: sediment sample M-92 (Stat. 70, 38°25'o"N, 35°5o'o"W, 1675 fathoms) yielded a single left valve, BM 1974.380. Cytheropteron patagoniense Brady (PI. 22, figS 12, 13) Cytheropteron patagoniense Brady, 1880 : 139, pi. 33, figs ya-d. LECTOTYPE. Left valve, eroded, BM 80.38.158. Length 0-56 mm ; height 0-37 mm. Type locality: Stat. 305, 160 fathoms. (47°48'o"S, 74°46'o"W, trawled, mud, surface temp. 55-o°F, January i, 1876.) DESCRIPTION. Shape: see PI. 22, fig. 12. Exceptions to Brady's (1880) descrip- tion include broadly rounded anterior, anterior half of dorsal margin regular and gently sloping, posterior half of dorsal margin steeply sloping. Ornamentation: the ridge mentioned by Brady is absent on the lectotype. A posterior dorsal knob is present at about the position where the ridge terminates on Brady's specimen. Hinge: median element smooth. Central muscle scars: four adductor scars with single kidney-shaped frontal scar. Cytheropteron scaphoides Brady (PI. 21, figS I4-l8) Cytheropteron scaphoides Brady, 1880 : 136, pi. 33, figs la-d. LECTOTYPE. Disarticulated right and left valves, BM 80.38.159. Right valve : length 0-37 mm ; height 0-19 mm ; left valve : length 0-37 mm ; height 0-15 mm. 3o8 H. S. PURI AND N. C. HULINGS Type locality: Stat. 149, Balfour Bay, Kerguelen Island, 20-50 fathoms, mud, January 1874. DESCRIPTION. Shape and ornamentation as given by Brady (1880). Inner lamella: anterior and posterior vestibula present with the former the larger of the two. Marginal pore canals few and simple. Hinge as follows : in the larger right valve, the sequence of elements anterior to posterior is socket, wedge-shaped (in dorsal view) tooth, crenulate bar, rounded (in dorsal view) tooth, socket. In smaller left valve, crenulate groove, socket, wedge-shaped tooth. In both valves, the high point of the wedge is towards the anterior end. See PI. 21, figs 17 and 18. Central muscle scars: adductor scars consist of four adjacent but distinct scars arranged in a vertical row. Frontal scar single somewhat circular and large. Overlap : right valve larger than left. Cytheropteron wellingtoniense Brady (PI. 23, figs 9-14) Cytheropteron wellingtoniense Brady, 1880 : 136, 137, pi. 34, figs 4a-d. LECTOTYPE. Right and left valves of a once articulated specimen, BM 80.38.160. Right valve: length 0-56 mm ; height 0-34 mm. Type locality: Wellington Harbour, New Zealand, depth unknown. DESCRIPTION. Shape essentially as given by Brady (1880). Notable exception is prominent indentation in dorso-anterior region. See PI. 23, fig. 10. Ornamenta- tion as given by Brady (1880). Inner lamella: anterior vestibule present. Hinge: see PI. 23, figs 11-13 ; terminal and median element crenulate. Central muscle scars: see PI. 23, fig. 14, frontal scar V-shaped. Genus BYTHOCYTHERE Bythocythere arenacea Brady (PI. 22, figS 1-5) Bythocythere arenacea Brady, 1880 : 142, pi. 33, figs 3a-g (arenosa on pi. 19). LECTOTYPE. Left valve, BM 81.5.50. Length 0-75 mm ; height 0-37 mm. Tip of posterior caudal process broken off and missing. Type locality: Stat. 185, Torres' Straits, 155 fathoms. (n°35'o"S, i44°3'o"E, dredged, sand and shells, surface temp. 77'0°F, August 31, 1874.) DESCRIPTION. Shape and ornamentation as described by Brady (1880). See PI. 22, fig. i. Inner lamella: see PI. 22, figs 2 and 3. Well defined anterior vestibule present. Hinge: see PI. 22, figs 4 and 5. Central muscle scars: five, arcuate, ver- tically arranged, elongate adductor scars and two frontal scars. REMARKS. Brady (1880, p. 142) found 'several examples' from Stat. 185 and figured (see pi. 33, fig. 3) complete carapaces of both a male and a female. Topo- typic material: two specimens, both left valves, were found in sediment sample M-237 ; one left valve is registered BM 1974.382. OSTRACODS FROM THE CHALLENGER EXPEDITION 309 By thocy there (?) exigua Brady (PI. 3, figs 6- 10) Bythocythere (?) exigua Brady, 1880 : 143, 144, pi. 6, figs ya-d (exigna on pi. 6). LECTOTYPE. Disarticulated right and left valves, BM 81.5.51. Right valve : length 0-50 mm ; height 0-28 mm ; left valve : length 0-50 mm ; height 0-28 mm. Type locality: Stat. 313, Straits of Magellan, 55 fathoms. (52°2i'o"S, 68°o'o"W, trawled, sand, bottom temp. 47'8°F, surface temp. 48-2°F, January 20, 1876.) DESCRIPTION. Shape and ornamentation as described by Brady (1880). Bythocythere pumilio Brady (PI. 22, figs 6-8) Bythocythere pumilio Brady, 1880 : 142, 143, pi. 33, figs 4a-d. LECTOTYPE. Whole carapace, BM 81.5.52. Length 0-45 mm ; height 0-21 mm. Type locality: Stat. 149, Balfour Bay, Kerguelen Island, 20-50 fathoms, mud, January 1874. DESCRIPTION. Shape and ornamentation as described by Brady (1880). Bythocythere velifera Brady (PI. 22, figs 9- ii ; PI. 27, fig. 3) Bythocythere velifera Brady, 1880 : 143, pi. 33, figs 5a-c. NEOTYPE. Left valve, BM 1974.381. Length 0-59 mm ; height 0-47 mm. Type locality: Stat. 185, Torres' Straits, 155 fathoms. (ii°35'o"S, i44°3'o"E, dredged, sand and shells, surface temp. 77-o°F, August 31, 1874.) DESCRIPTION. Shape and ornamentation as described by Brady (1880). REMARKS. The only specimen in the British Museum labelled Bythocythere velifera (BM 81.5.53) is a Cytheropteron (see PI. 22, figs 9-11), one of the two velate Bythocythere specimens found at the type locality is designated neotype. Genus PSEUDOCYTHERE Pseudocythere fuegiensis Brady (PI. i, figs 9, 10) Pseudocythere fuegiensis Brady, 1880 : 145, pi. i, figs ya-d. HOLOTYPE. Right valve, BM 81.5.54. (This is the only specimen found by Brady and figured as 7a-c.) Length i-i8mm ; height 0-50 mm. Type locality: Stat. 311, 245 fathoms. (52°5i'o"S, 73°53'o"W, trawled, mud, bottom temp. 46-o°F, surface temp. 5o-o°F, January n, 1876.) 3io H. S. PURI AND N. C. HULINGS DESCRIPTION. Shape and ornamentation as given by Brady (1880), except that the longitudinal striae are present over the entire surface but stronger in the posterior half. Inner lamella: anterior and posteroventral vestibula present, see PI. i, fig. 9. Marginal pore canals straight and simple. Cluster of three near centre of caudal process. Hinge adont. Central muscle scars: see PI. i, fig. 10. Genus CYTHERIDEIS Cytherideis laevata Brady (PI. 2, fig. 18; PI. 3, figs i -5) Cytherideis laevata Brady, 1880 : 146, 147, pi. 6, figs 5a-d ; pi. 35, figs 6a-d. LECTOTYPE. Disarticulated right and left valves, BM 80.38.164. Right valve : length 0-88 mm ; height 0-34 mm ; left valve : length 0-90 mm ; height 0-34 mm. Type locality: Stat. 151, off Heard Island, 75 fathoms. (52°59'3o"S, 73°33'3o"E, dredged, mud, surface temp. 36-2°F, February 7, 1874.) DESCRIPTION. Shape and ornamentation as described by Brady (1880), see PI. 2, fig. 18. Hinge adont, see PI. 3, fig. 3. Central muscle scars: see PL 3, fig. 2. REMARKS. Brady (1880, pp. 146, 147) did not mention the frequency of this species and he figures two complete carapaces (pi. 6, fig. 5 ; pi. 35, fig. 6). Topotypic material: this species is a common form in the sediment sample M-i85 and a right and a left valve are registered BM 1974.385-6. Genus XIPHICHILUS Xiphichilus (?) arcuatus Brady (PI. 24, figs 7, 8) Xiphichilus (?) arcuatus Brady, 1880 : 148, 149, pi. 35, figs 2a-d. LECTOTYPE. Disarticulated right and left valves, BM 81.5.55. Right valve : length 0-53 mm ; height 0-19 mm ; left valve : length 0-53 mm ; height 0-19 mm. Type locality: Stat. I74C, 610 fathoms. (i9°o7'5o"S, I78°I9'35"E, trawled, Globi- gerina ooze, bottom temp. 39-o°F, surface temp. 78-o°F, August 3, 1874.) DESCRIPTION. Shape: the lectotype differs from the description by Brady (1880) in that the anterior end is more rounded, ventral margin slightly concave anteriorly and convex posteriorly. Ornamentation: entire surface with many small longi- tudinal ridges that follow the general curvature of the dorsum. Ridges can be seen with transmitted light. Inner lamella widest anteriorly where a vestibule is present. Elsewhere the line of concrescence and inner margin coincide. Marginal pore canals few, straight and mostly single. Few false canals ventrally. Hinge merodont- lophodont. Right valve with small, terminal teeth and a long groove between. Left valve with a long smooth median bar between terminal sockets, the posterior OSTRACODS FROM THE CHALLENGER EXPEDITION 311 tooth is the largest. Central muscle scars: adductor scars consist of four closely adjacent scars. Xiphichilus complanatus Brady Xiphichilus complanatus Brady, 1880 : 148, pi. 35, figs 4a-d. LECTOTYPE. Left valve, BM 81.5.56 [specimen lost]. Type locality: Stat. 149, off Christmas Harbour, Kerguelen Island, 120 fathoms, January 29, 1874. DESCRIPTION. Shape as given by Brady (1880). Inner lamella, very wide anteriorly and posteriorly. Hinge adont. Normal pores frequent, open. REMARKS. Brady described this species only from Stat. 149. Three specimens (one adult and two juveniles) were found in sediment sample M-i.83. Topotypic material: a complete carapace, BM 1974.253. Genus POLYCOPE (?) Poly cope cingulata Brady (PI. 26, fig. 4) Polycope cingulata Brady, 1880 : 170, pi. 35, figs ya-d. MATERIAL. Complete carapace, BM 1974.384. Length 0-49 mm. Type locality : None. Brady (1880, p. 170) mentioned that there is no record of the locality in which the only specimen of this species was found. REMARKS. A single complete, eroded carapace, which may represent Polycope cingulata, was found in sediment sample M-igS (which represents Challenger Stat. Port Jackson, Australia, 2-10 fathoms, April 20, 1874.) The only articulated specimen which was figured by Brady as 7a-d on pi. 35 was never deposited either in the British Museum (Natural History) or in the Hancock Museum (fide letter by Mrs O. Marshall, secretary to Mr A. M. Tynan, Curator, Hancock Museum, dated 24 July 1967 to H. S. Puri). Polycope (?)favus Brady (PI. 24, fig. 9) Polycope (?) favus Brady, 1880 : 170, pi. 36, figs 4a-b. LECTOTYPE. Left valve, BM 81.5.64. Length 0-44 mm ; height 0-40 mm. Type locality: Stat. 185, Torres' Straits, 155 fathoms. (io°35'o"S, i44°03'o"E, dredged, sand and shells, surface temp. 77-o°F, August 31, 1874.) DESCRIPTION. Shape and ornamentation as described by Brady (1880). 312 H. S. PURI AND N. C. HULINGS REMARKS. Brady (1880, p. 170) found 'one or two' valves from Stat. 185, and he figured a left valve. Topotypic material: two valves, one broken, were found from sediment sample M-237. A left valve is registered BM 1974.383. Genus CYTHERELLA Cytherella cribrosa Brady (PI. 17, fig. 13) Cytherella cribrosa Brady, 1880 : 176, pi. 26, figs 5a-c. LECTOTYPE. Left valve, BM 81.5.66. Length 0-62 mm ; height 0-34 mm. Type locality: Stat. 172, off Nuknalofa, Tongatabu, 18 fathoms. (2O°58'o"S, i75°09'o"W, dredged, coral, mud, surface temp. 75-o°F, July 22, 1874.) REMARKS. The form described and illustrated by Brady does not resemble the lectotype. Brady (1880, p. 176) observed that the 'surface of the shell is destitute of ridges or undulations, but marked with numerous rather large oblong excavations' . The left valve figured by him as figs 5a-c is not as long (0-49 mm) as the lectotype (0-62 mm) which is obviously a Cytherelloidea. We have designated this specimen the lectotype as it was the only specimen found in the collection labelled as 'Cytherella cribrosa, No. 172, D.i8, off Tongatabu'. This is the only species of Cytherella in the dredging at Stat. 172, and Brady found it only at its type locality. Cytherella dromedaria Brady (PI. 24, fig. 14) Cytherella dromedaria Brady, 1880 : 173, pi. 43, figs 6a-b. LECTOTYPE. Left valve, BM 81.5.67. Length 0-77 mm ; height 0-47 mm. Type locality: Stat. 140, Simon's Bay, South Africa, 15-20 fathoms, October 1873. REMARKS. Brady (1880, p. 173) described this species from Stat. 140 (Simon's Bay) and he figured a left valve (pi. 43, figs 6a-b). The lectotype is also a left valve from Simon's Bay. Topotypic material: sediment sample M-I&4 (Simon's Bay, Stat. 140) yielded a right valve and a left valve, BM 1974.357-8. Cytherella irregularis Brady (PI. 24, fig. 10) Cytherella irregularis Brady, 1880 : 178, pi. 43, figs 3a-c. LECTOTYPE. Left valve (damaged), Hancock Museum. (Same specimen as illustrated by Brady, pi. 43, figs 3a-c, photographed on original museum slide as the specimen is too fragile to transfer.) Length 0-72 mm ; height 0-33 mm. Type locality: Stat. 33, off Bermuda, 435 fathoms. (32°2i'3o"N, 64°35'55"W, dredged, coral mud, surface temp. 68-o°F, April 4, 1873.) DESCRIPTION. Shape and ornamentation as described by Brady (1880), see PL 24, fig. 10. OSTRACODS FROM THE CHALLENGER EXPEDITION 313 Cytherella lata Brady (PI. 24, figs 17, 18) Cytherella lata Brady, 1880 : 173, pi. 44, figs 5a-e. LECTOTYPE. Right valve, BM 80.38.172. Length 0-81 mm ; height 0-56 mm. Type locality: Stat. 75, off Azores, 450 fathoms. (38°38'o"N, 28°28'3o"W, dredged, sand, surface temp. 70-o°F, July 2, 1873.) DESCRIPTION. Shape and ornamentation as described by Brady (1880), see PI. 24, fig. 17. Central muscle scars: two curved transverse rows of six and nine elongate scars respectively. REMARKS. Brady (1880, p. 173) reported this species from the following stations : off Culebra Island, West Indies, 390 fathoms, mud (Stat. 24) ; off Azores, lat. 38°37'o"N, long. 28°3o'o"W, 450 fathoms, sand (Stat. 75) ; off Pernambuco, lat. 8°37'o"S, long. 34°28'o"W, 675 fathoms, mud (Stat. 120) ; Torres' Straits, lat. n°35'o"S, long. i44°03'o"E, 155 fathoms, sand (Stat. 185) ; off Ki Islands, 580 fathoms, lat. 5°26'o"S, long. i33°i9'o"S, mud (Stat. igia). The lectotype is from Stat. 75. Topotypic material: a left valve recovered from sediment sample M-237 (Stat. 185), BM 1974.389. Cytherella latimarginata Brady (PI. 17, figs 14, 15) Cytherella latimarginata Brady, 1880 : 178, 179, pi. 36, figs 7a-d. NEOTYPE. Disarticulated left and right valves, BM 81.5.69. Right valve : length 0-41 mm ; height 0-28 mm ; left valve : length 0-41 mm ; height 0-25 mm. Type locality: Stat. 185, Torres' Straits, 155 fathoms. (n°35'o"S, i44°03'o"E, dredged, sand and shells, surface temp. 77-o°F, August 31, 1874.) DESCRIPTION. Shape and ornamentation as described by Brady (1880), see PI. 17, figs 14 and 15. REMARKS. Specimens of this species were never deposited either in the British Museum (Natural History) or the Hancock Museum (letter by Mrs O. Marshall, secretary to Mr A. M. Tynan, Curator, Hancock Museum, dated 24 July 1967, to H. S. Puri) and have been lost. Sediment sample M-237 (Stat. 185) yielded a single articulated valve, which is designated as neotype. Cytherella venusta Brady (PI. 24, figs 11-13) Cytherella venusta Brady, 1880 : 176, pi. 43, figs 4a-d. LECTOTYPE. Right valve, BM 80.38.180. Length 0-71 mm ; height 0-34 mm. Type locality: off reefs at Honolulu, 40 fathoms, July 1875. DESCRIPTION. Shape and ornamentation as described by Brady (1880), see PI. 24, figs ii and 13. Central muscle scars: see PI. 24, fig. 12. 314 H. S. PURI AND N. C. HULINGS REMARKS. Off reefs at Honolulu is the only locality where Brady (1880, p. 176) found this species. Topotypic material: sediment sample M-32/J., which represents the type locality, yielded a right and a left valve, which are designated topotypes BM 1974.390-1. REFERENCES BATE, R. H. 1963. The Ostracoda collected during the voyage of H.M.S. Challenger. Micro- paleontology 9 : 79-84. BENSON, R. H. 1971. A new Cenozoic deep-sea genus, Abyssocythere (Crustacea : Ostracoda . Trachyleberididae), with description of five new species. Smithson, Contr. Paleobiol: 7 : 1-25, 3 pis. 1972. The Bradleya problem. With descriptions of two new psychrospheric ostracode genera, Argenocythere and Poseidonamicus (Ostracoda : Crustacea). Smithson. Contr. Paleobiol. 12 : 1-138, 14 pis, 67 figs. BRADY, G. S. 1866. On new or imperfectly known species of marine Ostracoda. Trans, zool. Soc. Lond. 5 : 359-393. Pls 57-62. 1867-1871. In : Les Fonds de la Mev, vol. i. 1878. A monograph of the Ostracoda of the Antwerp Crag. Trans, zool. Soc. Lond. 10 (8) : 379-409, pis 62-69. 1880. Report on the Ostracoda dredged by 'H.M.S. Challenger' during the years 1873- 1876. Rep. scient. Results Voy. Challenger 1 (3) : 1-184, 44 P^s- 1898. On new or imperfectly known species of Ostracoda, chiefly from New Zealand. Trans, zool. Soc. Lond. 14 : 429-452, pis 43-47. CORNUEL, J. 1846. Description des entomostraces fossils du terrain Cretace inferieur du Departement de la Haute-Marne suivie d'indications sur les profondeurs de la mer qui a depose ce terrain. Mem. Soc. gdol. Fr. ser. i, 1 (2) : 193-205, 7 pis. HARDING, J. P. & SYLVESTER-BRADLEY, P. C. 1953. The ostracode genus Trachyleberis . Bull. Br. Mus. nat. Hist. (Zool.) 2 : 1-15, 2 pis. HORNIBROOK, N. DE B. 1952. Tertiary and Recent marine Ostracoda of New Zealand. Palaeont. Bull. Wellington, 18 : 1082, pis 1-18. NEVIANI, A. 1928. Ostracodi fossil d'ltalia. i. Vallebiaja (Calabriano) . Memorie Accad. pont. Nuovi Lincei ser. 2, 11 : 1 — 120. SEGUENZA, G. 1880. Le formazioni terziarie nella provincia di Reggio (Calabria). Memorie Accad. pont. Nuovi Lincei 6 : 3-443, pis 4-17, 2 maps. STODDART, W. W. 1861. On a Microzoa bed in the Carboniferous limestone of Clifton, near Bristol. Ann. Mag. nat. Hist. ser. 3, 8 : 486-490. SWAIN, F. M. 1963. Pleistocene Ostracoda from the Gubik Formation, Arctic coastal plain, Alaska. /. Paleont. 37 (4) : 798-834. TIZARD, T. H., MOSELEY, H. N., BUCHANAN, J. Y. & MURRAY, J. 1885. Narrative of the cruise of 'H.M.S. Challenger' with a general account of the scientific results of the expedition. Rep. scient. Results Voy. Challenger. Narrative 1 : i-mo. OSTRACODS FROM THE CHALLENGER EXPEDITION 315 DR HARBANS S. PURI BUREAU OF GEOLOGY 903 WEST TENNESSEE Sx TALLAHASSEE FLORIDA 32304 U.S.A. DR NEIL C. HULINGS Faculty of Science UNIVERSITY OF JORDAN AMMAN JORDAN PLATE 2 FIGS 1-4. Bairdia villosa Brady, lectotype, BM 80.38.44. i. Exterior view of left valve ( x 25), incident light. 2. Muscle scar of right valve (x 145). 3. Right valve, incident light. 4. Right valve, transmitted light. FIGS 5, 6. Argilloecia eburnea Brady, lectotype, BM 80.38.9. 5. Exterior view of left valve ( x 60). 6. Enlargement of muscle scar pattern ( x 260). FIGS 7-10. Bythocypris reniformis Brady, lectotype, BM 80. 38. 19. 7-8. Exterior views of right valve, incident and transmitted light ( x 35). 9. Interior view of anteroventral portion of marginal area. 10. Muscle scars (both x 105), transmitted light. FIGS u, 12. Bythocypris elongata Brady, lectotype, BM 81.5.8. u. External view of right valve ( x 25). 12. Enlargement of muscle scar pattern of right valve ( x 85), transmitted light. FIGS 13—15. Macrocypris tumida Brady, lectotype, BM 80,38.17. 13. Exterior view of right valve (x55). 14. Muscle scar pattern of left valve (x 195). 15. Exterior view of left valve ( x 55). transmitted light. FIGS 16, 17. Aglaia clavata Brady, lectotype, BM 81.5.1, before separation of valves ( x 45), incident light. 16. Exterior view of left side. 17. Dorsal view of whole specimen. FIG. 18. Cytherideis laevata Brady, lectotype, BM 80.38.164 ( x 45), incident light. Exterior view of right side of whole specimen before separation of valves. Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 2 PLATE 3 FIGS 1-5. Cytherideis laevata Brady, lectotype, BM 80.38.164, transmitted light, i. External view of right valve ( x 40). 2. Internal view of muscle scars of right valve ( x 205). 3. External view of left valve ( x 40). 4, 5. Internal views of marginal areas of right valve ( x 70). FIGS 6-10. Bythocythere (?) exigua Brady, lectotype, BM 81.5. 51. 6-7. External view of posteroventral margin (x2io). 8, 9. The marginal areas and hinge of whole valve (x75). 10. The anterior hinge element ( x 210). FIGS 11 — 14. Bairdia simplex Brady, lectotype, BM 81.5.13 ( x 25), incident light. 11 — 13. External and internal views of left valve. 14 — 15. External and internal views of right valve. FIG. 15. Bairdia abyssicola Brady, lectotype, Hancock Museum ( x 25), incident light. External view of right valve. FIG. 16. Pontocypris (?) subreniformis Brady, lectotype, BM 81.5.5, Port Jackson, Australia, 2 — 10 fathoms ( x 60), incident light, unstained. Exterior view of right valve of whole specimen. FIGS 17, 18. Bairdia minima Brady, lectotype, BM 80.38.37 ( x 50), transmitted light. Exterior views of left and right valves. Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 3 PLATE 4 FIGS 1-3. Bairdia minima Brady, lectotype, BM 80.38.37. i, 3. External views of the left and right valves of what is believed to have been a whole specimen (xso), unstained. 2. Parts of the muscle scar pattern of right valve ( x 200), transmitted light, viewed from the interior. FIGS 4, 5. Bairdia hirsuta Brady, lectotype, BM 80.38.35. 4. External view of right valve with setae still attached, transmitted light ( x 25). 5. Internal view of the same valve, incident light (X25). FIGS 6- 1 1. Bairdia globulus Brady, lectotype, BM 80.38.34. 6-7. External views of the left valves ( x 35), transmitted and incident light. 8-9. External views of right valves ( x 35), transmitted and incident light. 10. Internal view of muscle scar of right valve (xgo). u. Whole specimen shown in dorsal view. FIGS 12-15. Bairdia exaltata Brady, lectotype, BM 81.5.10, transmitted light. 12-13. External and internal views of right valve ( x 25). 14. Muscle scars of left valve ( x 75). 15. External view of left valve ( x 25). FIGS 16-18. Bairdia woodwardiana Brady, lectotype, BM 80.38.46, transmitted light. 16. Exterior view of the right valve ( x 45). 17. Muscle scars of right valve ( x 95). 18. Left valve ( x 45) . Bull. BY. Mus. nat. Hist. (Zool.) 29, 5 PLATE 4 /% H^ 18 PLATE 5 FIGS 1-3. Bairdia expansa Brady, lectotype, BM 81.5.11, transmitted light, i. External view of left valve ( x 50). 2. Interior view of muscle scar of left valve ( x 200). 3. External view of right valve ( x 50) . FIGS 4-6. Bairdia attenuata Brady, lectotype, BM 80.38.27. 4-5. Exterior and interior views of right valve of a complete specimen ( x 30), incident light. 6. Muscle scar of interior of same valve ( x 85). FIGS 7-9. Bairdia for tificata Brady, lectotype, BM 81.5.12. 7. External view of left valve ( x 35)- 8. Internal view of left valve ( x 35). 9. External view of muscle scar region showing some of the scars and the finely punctate surface ( x 100). FIGS 10-12. Cythere obtusalata Brady, lectotype, BM 80.38.96. 10, n. Exterior and interior view of right valve ( x 60), incident light, stained. 12. Interior view of valve showing muscle scar and surface reticulation ( x 185), transmitted light. FIGS 13-15. Cythere vellicata Brady, lectotype, BM 81.5. 32, incident light, stained. 13. Exterior view of the right valve. 14, 15. Dorsal aspects of whole specimen ( x 170). FIGS 16-19. Cythere curvicostata Brady, lectotype, BM 80.38.64 (x5o), incident light, stained. 16. Exterior dorsal view. 17. Exterior of left valve. 18. Ventral aspect. 19. Right valve of a complete specimen. Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 5 16 17 18 PLATE 6 FIGS 1-3. Cythere moseleyi Brady, lectotype, BM 80.38.93, male (X5o), incident light, stained, i. External view of left side. 2. Dorsal view. 3. External right side. FIG. 4. Cythere falklandi Brady, BM 80.38. y8A (x5o), incident light, unstained. External view of left valve. FIGS 5-9. Cythere falklandi Brady, BM 80.38.786. 5-6. Transmitted light views of right valve ( x 50). 7. Internal view of the muscle scars ( x 155). 8. Anterior marginal area ( x 170) showing radial pore canals and vestibule. 9. External view of same specimen ( x 50), transmitted light, unstained. FIGS 10-12. Cythere inconspicua Brady, lectotype, BM 8 1.5. 22, whole specimen, inverted to illuminate ventrolateral ridges within the subtle reticulate pattern ( x 80), stained. 10. External view of left side. n. Dorsal view. 12. Ventrolateral quarter. FIGS 13-18. Cythere cumulus Brady, lectotype, BM 81.5.17. 13, 14. External views of right and left valves. 15. Dorsum of whole specimen ( x 50), incident light, stained. 16, 18. Internal views of posterior and anterior elements of hinge and radial pore canals. 17. Muscle scars and reticulate surface pattern ( x 165), transmitted light. Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 6 PLATE 7 FIGS 1-4, 6, 7. Cythere floscardui Brady, lectotype, BM 80.38.80. i, 3. External views of left and right valves (x8o), transmitted light. 2. Muscle scars with V-shaped antennal scar, and reticulate surface pattern (x65). 4, 6, 7. Views of left, dorsal, and right sides of whole specimen before separation ( x 45), incident light, stained. FIG. 5. Cythere cumulus Brady, lectotype, BM 81.5.17. Dorsal view ( x 40). FIGS 8, ii. Cythere securifer Brady, lectotype, BM 80. 38. 112 (x6o). 8, n. External and internal views of left valve of a female. FIGS 9, 10, 12-15. BM 8o.38.ii2A. 9, 10. External views of left and right sides of whole specimen (x55), showing apparently dimorphic posteroventral velate structure ('triangular or hatchet-shaped protuberance'), transmitted light views of selected portions of the valves in- cluding, 12. Muscle scars and surface reticulation (xi35). 13. Anterior marginal area (note antennule and part of antenna). 14, 15. Anterior and posterior hinge elements ( x no). FIGS 16-19. Cythere exfoveolata Neviani, lectotype, BM 80.38.81 (X55), whole carapace of male. 16. External view of lateroventral quarter. 17. Right side. 18. Dorsum. 19. Left side. Bull. Br. Mus. nal. Hist. (Zool.) 29, 5 PLATE 7 PLATE 8 FIGS 1-4. Cythere flabellicostata Brady, lectotype, BM 80.38.79 (x5o). i. External views of whole specimen from right, incident light. 2. Right side, black light. 3. Left side, incident light. 4. Left side, black light. FIG. 5. Cythere acupunctata Brady, lectotype, BM 80.38.50. External view of left valve of a complete specimen ( x 35), incident light, stained. FIGS 6, 7. Cythere scintillulata Brady, lectotype, BM 80. 38. no. 6. External view of left valve of an articulated specimen (x55), incident light, unstained. 7. External view of left valve ( x 65), incident light, unstained. FIGS 8, 9. Cythere hardingi Puri & Hulings, lectotype, BM 80.38.97. 8. External view of whole specimen as seen from left side ( x 45). 9. The top ( x 40). FIGS 10-12. Cythere fulvotincta Brady, lectotype, BM 81.5.21 ( x 55), incident light, stained. 10. External view of whole specimen as seen from left side. n. The top. 12. Right side. FIGS 13, 14. Cythere lubbockiana Brady, lectotype, BM 81.5.25. 13. External view of whole specimen as seen from left high oblique side ( x 40). 14. Right high oblique side ( x 45), incident light, stained. FIGS 15-18. Cythere parallelo gramma Brady, lectotype, BM 80.38.99. 15. External view of right valve. 16. Internal view of right valve (both x 40, incident light, stained). 17. Muscle scars as seen in transmitted light in interior of right valve (x 130). 18. Top view showing stepped anterior tooth and prominent posterior tooth of holamphidont hinge ( x 40) . Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 8 PLATE 9 FIGS 1-4. Cythere mackenziei Puri & Hulings, lectotype, BM 80.38.103. i. Interior view of left valve (X35), transmitted light, unstained. 2. Muscle scar (xiyo). 3. Hinge (xyo). 4. Radial pore canals of anterior margin ( x 155). FIGS 5, 6. Cythere cytheropteroides Brady, lectotype, BM 80.38.67 (x4o), incident light, stained. 5. External view of right valve. 6. Internal view of right valve. FIGS 7, 8. Lectotype, BM 80.38.67. 7. External view of right valve ( x 35), incident light, stained. 8. Internal view of anterior hinge element ( x 105). FIG. 9. Cythere rastromarginata Brady, paralectotype, BM 80. 38. 104 ( x 60), incident light, unstained. External lateral right valve view of probable female. FIGS 10-14. Lectotype, BM 80.38.105. 10-12. External lateral and internal views of left valve of male ( x 60), incident and transmitted light, stained. 13. Posteroventral marginal area. 14. Right side of the subcentral tubercle ( x 185). FIGS 15, 16. Cythere impluta Brady, lectotype, BM 1961.12.4.30 (X45), incident light, stained. 15. External left lateral view of whole specimen. 16. Dorsal view of whole specimen. FIGS 17-19. Cythere murrayana Brady, lectotype, BM 80.38.94 (X75), incident light, unstained. 17. External view of left valve. 18. Dorsum of whole specimen. 19. Venter of whole specimen. Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 9 18 PLATE 10 FIGS i-u. Cythere exilis Brady, lectotype, BM 81.5.20, figs 5, 6 ( x 145), transmitted light. 1-2. External lateral views of left valves. 3-4. External lateral views of right valves (all x 45, incident and transmitted light, stained). 5. Area postjacent to subcentral tubercle of left valve showing configuration of fossae. 6. Interior view of anterior margin of right valve. 7. Posterior margin of right valve. 8. Anterior hinge element of left valve. 9. Anterior hinge element of right valve. 10. Posterior hinge element of right valve. 1 1. Posterior hinge element of left valve. FIGS 12, 13. Cythere bicarinata Brady, lectotype, BM 80.38.50 ( x 50), incident light, stained. 12. External view of right valve. 13. Dorsum of whole carapace. FIGS 14-18. Cythere ericea Brady, lectotype, BM 80.38.76, figs 16-18 ( xgo), incident and transmitted light, stained. 14. External view of right valve (X3o), incident light, stained. 15. Internal view of right valve. 16. Internal anterior hinge element. 17. Muscle scars. 1 8. Posterior hinge element of right valve. Bull. BY. Mus. nat. Hist. (Zool.) 29, 5 PLATE 10 PLATE ii FIGS 1-9. Cythere irpex Brady, lectotype, BM 80.38.86. 1-3. External lateral views of left and right valves. 4. Internal view of right valve (all xyo), black light and transmitted light, unstained. 5. Enlarged internal view of muscle scars. 6. Anterior hinge element of right valve. 7-8. Posterior hinge element of left and right valves. 9. Anterior hinge element of left valve (all x 200), transmitted light. FIGS 10, ii. Cythere dasyderma Brady, lectotype, BM 1961.12.4.39, showing subreticulate, spinose surface, imperfectly developed holamphidont (penultimate stage ?) hinge and large V-shaped frontal muscle scar ( x 25), incident light, stained. 10. External lateral view of right valve, ii. Internal lateral view of left valve. FIGS 12-14. Cythere irrorata Brady, lectotype, BM 80.38. 86A (x45), incident light, un- stained. 12. External view of left valve. 13. Venter of whole specimen. 14. Dorsum of whole specimen. FIG. 15. Cythere viminea Brady, lectotype, BM 81.5.33 (x5o), incident light, unstained, photograph taken of specimen on original museum slide. External view of broken right valve. FIGS 16-18. Cythere acanthoderma Brady, lectotype, BM 8o.38.48A. 16. External view of late instar left valve. 17. Internal view of late instar left valve (both x 35, incident and transmitted light, unstained). 18. Enlargement of anterior hinge element and marginal area Bull. Br Mus. nat. Hist. (Zool.) 29, 5 PLATE ii PLATE 12 FIGS 1-3. Cythere sabulosa Brady, lectotype, BM 80.38.107. i, 2. External views of right and left valves. 3. Dorsum of whole specimen ( x 55 and x 40), incident light, stained. FIGS 4-6. Cythere packardi Brady, lectotype, BM 81.5. 26 (x6s), incident light, stained. 4. External view of left valve. 5, 6. Dorsum and right valve of whole specimen. FIGS 7-9. Cythere radula Brady, lectotype, BM 81.5.28 (x35), incident light, stained. 7. External lateral view of left valve, late instar. 8. Internal view of left valve of late instar. 9. Dorsal view. FIGS 10,11. Cythere lepralioides Brady, lectotype, BM 80.38.91 ( x 45), incident light, stained. External and internal views of right valve of probable late instar. FIGS 12, 13. Cythere torresi Brady, lectotype, BM 81.5.31 ( x 90), transmitted light, unstained. 12. Internal lateral view of right valve. 13. External lateral view of right valve. FIGS 14-16. Cythere kerguelenensis Brady, lectotype, BM 80.38.88 ( x 50), incident light, stained. 14. External view left of whole specimen. 15. Right side. 16. Dorsum. FIGS 17, 18. Lectotype, BM 80.38.88 (x5o), incident light, stained. 17. External view of separated left valve. 18. External view of separated right valve. Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 12 PLATE 13 FIGS 1-9. Cythere subrufa Brady, lectotype, BM 80.38.117. 1-5. External lateral views of left and right valves ( x 80), incident and transmitted light, unstained and stained. 6. Internal view of right valve ( x 80). 7-8. Internal views of anterior and posterior hinge elements of left valve. 9. Muscle scar pattern of right valve ( x 305), incident and transmitted light. FIGS 10-18. Cythere wyvillethomsoni (Brady), lectotype, BM 80.38.123. 10,11. External views of left and right valves ( x 40), incident light, stained. 12-17. ( x 140.) 12. Interior view of enlarged anteroventral portion of left valve showing radial pore canals. 13. Anterior hinge element of left valve. 14. Posterior hinge element of right valve. 15. Posteroventral marginal area of left valve. 16. Anterior hinge element of right valve. 17. Posterior hinge element of left valve. 18. Second antenna and penis dissected from original complete specimen (x86). Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 13 18 PLATE 14 FIGS 1-3. Cy there wyvillethomsoni Brady, lectotype, BM 80.38.123. i. External lateral view of right valve (X45), transmitted light. 2. Region postjacent to subcentral tubercle showing reticulate surface pattern in relation to normal pore canals (x 155). 3. Lateral view of left valve ( x 45) . FIG. 4. Cythere stolonifera Brady, lectotype, BM 80.38.115 ( x 50), incident light, unstained. External lateral left valve view of broken complete specimen. FIGS 5-8. Cythere lauta Brady, lectotype, BM 81.5.24. 5-6. External views of right valve (x65), incident light in air and immersed in water, stained. 7. Dorsal view. 8. Left valve (both x 65), incident light, immersed in water. FIGS 9-12. Cythere craticula Brady, lectotype, BM 81.5.16. 9-10. External lateral views of left valve ( x 45), incident and transmitted light, unstained, n. Internal view showing hinge and portions of marginal area ( x 70). 12. Area postjacent and ventral to subcentral tubercule ( x 120). FIG. 13. Cythere scalaris Brady, lectotype, BM 80.38.109 ( x 35), incident light, photograph taken of specimen on original museum slide. External view of broken right valve. FIGS 14-18. Cythere quadriaculeata Brady, lectotype, BM 80.38.50. 14. External lateral view of whole specimen. 15. Dorsal view of whole specimen (both x 50), incident light, stained. 16, 17. Internal views of hinges of right and left valves. 18. Anterior marginal area showing radial pore canals (16-18, x 105), transmitted light. Bull. BY. Mus. nat. Hist. (Zool.) 29, 5 PLATE 14 18 PLATE 15 FIGS 1-4. Cythere dorsoserrata Brady, lectotype, BM 81.5.19. i. External lateral view of right valve (X45), incident light, stained. 2. Internal view of anterior hinge element. 3. Internal view of posterior hinge element. 4. Anterior portion of marginal area (2-4, x 105), incident light, stained. FIGS 5-8. Cythere patagoniensis Brady, lectotype, BM 81.5.27. 5. External view of left valve. 6. External view of right valve. 7. Dorsum view of whole specimen. 8. Venter view of whole specimen. FIGS 9-16. Cythere velivola Brady, lectotype, BM 80.38.122. 9. External lateral view of left valve ( x 45), black light, immersed in water. 10. Interior view of left valve ( x 45), trans- mitted light, ii. Interior view of muscle scar region showing normal pore canals. 12. Anterior marginal area showing radial pore canals (both x 160), transmitted light ; the sieve-like nature of the normal pore canals may be an optical artifact caused by air trapped in the canals in contact with the parallel oriented C-axes of the calcite crystals of the carapace wall. 13, 14. Dorsal views of posterior and anterior teeth of hinge of right valve. 15, 16. Plate-shaped spines of ventrolateral velate flange, and plates of dorsal crest (13-16, about X2O5), transmitted light. FIGS 17, 18. Cythere tricristata Brady, lectotype, BM 80.38.121 ( x 50), incident light, stained. 17. External view of right valve. 18. Dorsum of whole carapace. Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 15 PLATE 16 FIGS i, 2. Cythere clavigera Brady, lectotype, BM 80.38.59. i. External view of right valve (X35), incident light, unstained. 2. Internal view of same right valve in transmitted light (X45). FIGS 3-5. Cythere squalidentata Brady, lectotype, BM 81.5.29 (x65), incident light, un- stained. 3. External view of whole carapace as seen from left side. 4. External view from right side. 5. External view from top. FIGS 6-8. Cythere dictyon Brady, lectotype, BM 1961.12.4.32. 6. External view of left valve. 7. Internal view of left valve (both X35), incident light, stained. 8. Enlargement of anteromedian portion of valve in region of subcentral tubercule showing bifurcation anteriorly of posteromedian ridge ( x 105). FIGS 9, 10. Cythere arata Brady, lectotype, BM 80.38.52, penultimate instar ( x 60), incident light, stained. 9. External view of right valve. 10. Internal view of right valve. FIGS 11-18. Cythere papuensis Brady, lectotype, BM 80.38.98. n, 12. External views of left and right valves of an articulated specimen. 13, 14. Internal views of same right and left valves. 15. Top view of left valve ( x 45), incident light, stained. 16. Internal view of anterior hinge element. 17. Muscle scars. 18. Posterior hinge elements of right valve (16, 17, x 145), transmitted light. Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 16 PLATE 17 FIGS 1-2. Cythere sulcatoperforata Brady, lectotype, BM 81.5.30. i. External lateral view of left valve (X35), incident light, stained. 2. Internal view of left valve showing merodont hinge and trachyleberid muscle scar ( x 25), incident light, stained. FIGS 3-6. Cythere circumdentata Brady, lectotype, BM 80.38.58. 3-4. External lateral views of left valve (x3o), incident light, stained, immersed in water and dried. 5. Internal view of left valve showing hinge and marginal area of probable penultimate instar ( x 30) . 6. Muscle scar ( x 80) . FIGS 7-12. Cythere suhmi Brady, lectotype, BM 80. 38. 119 ( x 30), incident light, stained. 7-8. External lateral views of left and right valves. 9. Internal view of right valve. 10. External view of left valve, n. Internal view of left valve. 12. Internal view of right valve ( x 3S)> transmitted light, unstained. FIG. 13. Cytherella cribrosa Brady, lectotype, BM 81.5.66 (x55), incident light, stained. External lateral view of left valve. FIGS 14, 15. Cytherella latimarginata Brady, neotype, BM 81.5.69 (X75), transmitted light. 14. External lateral view of right valve. 15. External view of left valve. FIGS 16-18. Krithe producta Brady, BM 80.38.128. 16. External lateral view of left valve (X40), transmitted light. 17. Muscle scars (xiis). BM 80.38.127 (xso), incident light. 1 8. External view of left valve of whole specimen. Bull. BY. Mus. nat. Hist. (Zool.) 29, 5 PLATE 17 PLATE 18 FIGS i, 2. Krithe hyalina Brady, lectotype, BM 81.5.34 (x6o), incident light, unstained, i. External view of left valve. 2. Dorsum of whole carapace. FIGS 3-5. Krithe tumida Brady, lectotype, BM 81.5. 36 (X55), incident light, unstained. 3. External view of left valve. 4. Dorsum of whole carapace. 5. Venter of whole carapace. FIGS 6-9. Loxoconcha anomala Brady, BM 80.38.132. 6, 7. External lateral views of left valve ( x 55), incident and transmitted light, stained. 8. Muscle scar region showing adductor scars and aligned subreticulate structure of surface ornament. 9. Lateral view of right valve. (6, 7, 9, x 55 ; 8, x 100), transmitted light. FIGS 10-12. Loxoconcha pumicosa Brady, lectotype, BM 81.5.37. Io- External view of left valve ( x 60), incident light, unstained. 1 1. Right valve, external view ( x 70), transmitted light. 12. Dorsum of whole specimen ( x 60). FIGS 13, 14. Loxoconcha africana Brady, lectotype, BM 80.28.130 (xso), incident light, stained. 13. External lateral view of right valve. 14. External lateral view of left valve. FIGS 15, 16. Loxoconcha subrhomboidea Brady, lectotype, Hancock Museum ( x 70), incident light, stained. 15. External view of left valve showing enlarged row of pits along dorsal side of ventrolateral ridge. 16. Dorsum showing dorsal marginal ridges. FIGS 17, 18. Loxoconcha australis Brady, lectotype, BM 80.38.133. 17. External view of left valve (xgs), incident light, stained. 18. Internal view of right valve (x95), incident light, unstained. Bull. Br. Mus. nat. Uisl. (Zool.) 29, 5 PLATE 18 PLATE 19 FIGS 1—4. Loxoconcha australis Brady, lectotype, BM 80.38.133. i, 2. External lateral views of left valve ( x 50), incident and transmitted light. 3. Internal view of left valve ( x 65). 4. Muscle scar region showing reticulate surface pattern ( x 75) from exterior. FIGS 5, 6. Loxoconcha honoluliensis Brady, lectotype, BM 80.38.136 (x55), incident light, stained. 5. External view of right side of a whole carapace. 6. View from above. FIGS 7-10. Cytherura clavata Brady, lectotype, BM8o.38.i48 ( x 60), incident and transmitted light, stained. 7. External view of left valve. 8. External view of right valve. 9. Internal view of right valve. 10. Internal view of left valve. FIGS ii, 12. Xestoleberis foveolata Brady, lectotype, BM 80.38.141 ( x 65), incident light, stained, n. External view of left valve. 12. Venter of whole carapace. FIGS 13, 14. 'Xestoleberis expansa' Brady, BM 81.5.41 (x85), incident light, unstained. 13. External view of left valve. 14. Venter of whole carapace. FIGS 15, 16. Xestoleberis africana Brady, lectotype, BM 81.5.40 ( x 80), incident light, unstained. 15. External view of right valve. 16. Dorsum of whole carapace. FIGS 17, 1 8. Xestoleberis granulosa Brady, lectotype, Hancock Museum ( x 60), incident light, unstained. 17. External view of left valve. 18. Venter of whole carapace. Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 19 PLATE 20 FIGS 1-3. Aglaia (?) pusilla Brady, lectotype, BM 81.5.2, transmitted light, i. External lateral view of left valve ( x 75). 2. Muscle scars of right valve ( x 250). 3. Lateral view of right valve ( x 75). FIGS 4-6. Aglaia (?) meridionalis Brady, lectotype, BM 1961.12.4.63. 4, 5. External lateral views of left valve ( x 80), incident and transmitted light, unstained. 6. Muscle scars (x 190). FIGS 7, 8. Aglaia (?) obtusata Brady, lectotype, BM 80.38.4 ( x 90), incident light, unstained. 7. External view of left side of whole carapace. 8. Dorsum of whole carapace. FIGS 9-11. Xestoleberis setigera Brady, lectotype, BM 80.38.145. 9. External view of whole carapace as seen from left side. 10. Dorsum (x4o). n. Venter showing marginal area and radial pore canals ( x 50), incident light, unstained. FIGS 12, 13. 'Xestoleberis tumefacta' Brady, BM 81.5.44 (x<^5)< incident and transmitted light, stained. External views of broken left valve. FIGS 14, 15. Xestoleberis nana Brady, lectotype, BM 80.38.143 (x75), transmitted light, unstained. 14. External view of right valve. 15. Internal view of right valve. FIGS 16-18. Xestoleberis variegata Brady, lectotype, BM 80. 38. 146 (x75), incident and transmitted light, unstained. 16. External view of whole carapace as seen from right side. 17. Venter. 18. Dorsum. Bull. BY. Mus. nat. Hist. (Zool.) 29, 5 PLATE 20 18 PLATE 21 FIG. i. Cytherura cryptifera Brady, Hancock Museum (x8o), incident light, unstained. External lateral view of broken left valve photographed on museum slide. FIG. 2. Cytherura cribrosa Brady, lectotype, BM 80.38.150 ( x 70), incident light, stained. External lateral left valve view of complete carapace. FIGS 3-6. Cytherura lilljeborgii Brady, lectotype, BM 80.38.151. 3. External view of whole carapace as seen from left side. 4. Right side (both x 80). 5. Venter. 6. Dorsum (5, 6, x 60, incident light, stained). FIGS 7, 8. Cytherura costellata Brady, lectotype, BM 80.38.149 ( x 60), incident light, stained. 7. External lateral view of right valve. 8. Internal lateral view of right valve. FIGS 9, 10. Cytherura clausi Brady, lectotype, BM 81.5. 17, incident light, stained. 9. External lateral view of left valve ( x 80). 10. Dorsum view of left valve ( x 55). FIGS n, 12. Cytherura mucronata Brady, lectotype, BM 81.5.45 (X7o), incident light, unstained, n. External lateral view of whole carapace. 12. Dorsum of whole carapace. FIG. 13. Cytherura curvistriata Brady, lectotype, BM 81.5.46 ( x 60), incident light, unstained. External lateral view of complete specimen right valve. FIGS 14—18. Cytheropteron scaphoides Brady, lectotype, BM 80.38.159. 14. External lateral view of right valve. 15. External lateral view of left valve (both of a former single specimen, x 100), transmitted light. 16. Internal view of muscle scar of left valve. 17. Hinge of left valve. 18. Hinge of right valve (both x 270), transmitted light. Bull. Bv. Mus. nat. Hist. (Zool.) 29, 5 PLATE 21 17'. 18 PLATE 22 FIGS 1-5. Bythocythere arenacea Brady, lectotype, BM 81.5.50, transmitted light, i. Exterior of left valve ( x 45). 2. Interior of left valve. 3. Anterior marginal area. 4. Posterior hinge element. 5. Anterior hinge element (3-5, x 150). FIGS 6-8. Bythocythere pumilio Brady, lectotype, BM 81.5.52 ( x 40), incident light, stained. 6. Exterior view of whole specimen, left side. 7. View of venter. 8. View of dorsum. FIGS 9-11. 'Bythocythere velifera' Brady, BM 81.5.53. 9-10. Exterior and interior views of left valve ( x 60), transmitted light, n. View of venter ( x 65), incident light, stained. FIGS, 12, 13. Cytheropteron patagoniense Brady, lectotype, BM 80.38.158 (x65), incident light, stained. 12. Exterior view of left valve. 13. Interior view of left valve. FIGS 14-18. Cytheropteron mucronalatum Brady, lectotype, BM 80.38,157, all (x3o), incident light, stained. 14, 15. Exterior views of right and left valves. 16, 17. Interior views of right and left valves of a former complete specimen, showing hinge with 'accommodation groove' in left valve and V-shaped frontal scar in right valve. 18. Dorsal view of right valve showing stepped anterior tooth and crenulate posterior tooth. Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 22 PLATE 23 FIGS 1-7. Cytheropteron assimile Brady, lectotype, BM 80.38.153, all with transmitted light. 1,2. External views of left valves ( x 55). 3. Dorsal view of posterior hinge element of left valve. 4, 5. Internal views of anterior and posterior hinge elements of right valve. 6, 7. Internal view of anterior and posterior hinge elements of valve (3-7, x 155). FIG. 8. Cytheropteron abyssorum Brady, lectotype, BM 81.5.49 (x55). Dorsal view of crushed specimen with valves joined. FIGS 9-14. Cytheropteron wellingtoniense Brady, lectotype, BM 80.38.160. 9, 10. External views of left and right valves of whole specimen ( x 70). n. Internal view of left valve. 12. Dorsal view of hinge of left valve ( x 75). 13. Interior view of anterior hinge element of right valve ( x 225). 14. Interior view of muscle scars of right valve ( x 250). FIGS 15-17. Cytheropteron (?) angustatum Brady, lectotype, BM 80.38.152, transmitted light. 15, 16. Interior and exterior view of left valve ( x 75). 17. Muscle scars ( x 130). FIG. 18. Cytheropteron fenestratum Brady, lectotype, BM 80.38.154-155 (x4o). External view of right valve as seen in black-light illumination. Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 23 PLATE 24 FIGS 1-6. Cytheropteron fenestratum Brady, lectotype, BM 80.38.154-155. i, 2. External lateral views of right valve. 3. External lateral view as seen from above. 4. External lateral view of left valve (all X4o), incident light, unstained. 5. Hinge as seen from above (x8o). 6. Posterior element as seen internally in transmitted light showing the tooth and its junctive with confining bars of median groove ( x 250) . FIGS 7, 8. Xiphichilus (?) arcuatus Brady, lectotype, BM 81.5.55 ( x 65), incident light, unstained. 7. External view of left valve. 8. Dorsum of whole carapace. FIG. 9. Polycope (?) favus Brady, lectotype, BM 81.5. 64 (X75), transmitted light, stained. External view of left valve. FIG. 10. Cytherella irregularis Brady, lectotype, Hancock Museum ( x 45), photograph taken of specimen on museum slide. External lateral view of damaged left valve. FIGS 11-13. Cytherella venusta Brady, lectotype, BM 80. 38.180 ( x 55), incident and trans- mitted light, ii, 12. External lateral views of left valve. 13. External lateral view of right valve. FIG. 14. Cytherella dromedaria Brady, BM 81.5.67. External lateral view of left valve. FIGS 15, 16. Cythere (?) serratula Brady, lectotype, BM 80. 38. 113 ( x 35), incident light, stained. External and internal views of right valve. FIGS 17, 18. Cytherella lata Brady, lectotype, BM 80.38.172 (x25), incident light, stained. External and internal views of right valve. Bull. BY. Mus. nat. Hist. (Zool.) 29, 5 PLATE 24 . 25 ^^^BBpB^^^^^^^^^^^imP 10 PLATE 25 FIGS i, 2. Cythere melobesioides Brady, 1869, BM 80.38.92 ( x 55), incident light, stained. Station 142, off Cape of Good Hope, 150 fathoms. Originally described by Brady, 1869, in Les Fonds de la Mer. The specimen photographed is very similar to that illustrated in the Challenger Report (pi. XVIII, figs le-g), which he states (p. 162) most closely agrees with the original specimens described from Mauritius in 1869. i. External lateral view of left valve. 2. Internal lateral view of left valve. FIGS 3-6. Cythere cymba Brady, 1869, BM 1961.12.4.44 ( x 35), incident light, stained. Station 233b, Inland Sea of Japan, 14 fathoms. Described by Brady (1869 : 157) in Les Fonds de la Mer, from Hong Kong. 3. Exterior view of a whole carapace as seen from left side. 4. Exterior view of whole carapace as seen from right side. 5. Left lower oblique view of whole carapace. 6. Dorsum. FIGS 7-13. Cythere polytrema Brady, 1878, BM 80.38.100, off Prince Edward's Island, 50-150 fathoms. Originally described by Brady from fossil specimens from the lower sands and the Isocardium bed of the middle sands of Antwerp Crag. This is undoubtedly another species. The present form may be most closely identified to Cativella bensoni Neale, 1967. 7. External lateral view of left valve. 8. Internal lateral view of left valve. 9. External lateral view of right valve. 10. Internal lateral view of right valve (all x 45), incident light, n. Left valve hinge. 12. Right valve hinge. 13. Muscle scars (11-13, x 55)- FIGS 14-18. Cythere euplectella Brady, 1869, BM 80.38.77, station 189, Arafura Sea, 28 fathoms. Originally described by Brady (1869 : 157) in Les Fonds de la Mer from Hong Kong. 14, 15. Exterior views of right valve. 16. Interior view of right valve ( x 65), incident and trans- mitted light. 17. Muscle scars. 18. Anterior hinge element (both x 120). Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 25 PLATE 26 FIGS 1-3. Xestoleberis tumefacta Brady, neotype, BM 1974.370 (xQ4). i. External right lateral view. 2. External left lateral view. 3. Dorsum. FIG. 4. (?) Polycope cingulata Brady, BM 1974.384 ( x 114). External left lateral view. FIGS 5, 7. Cythere cristatella Brady, BM 80.38.63 (X77). 5. External right lateral view. 7. External left lateral view. FIGS 6, 8. Cythere scabrocuneata, Brady, topotype, BM 1974.342 (x64). 6. Internal view left valve. 8. External view left valve. Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 26 PLATE 27 FIGS i, 2. Argilloecia badia Brady, topotype, BM 1974.252 A (x 139). i. Internal view of right valve. 2. External view of right valve. FIG. 3. Bythocythere velifera Brady, neotype, BM 1974.381 (x85). External view of left valve. FIGS 4-6. Cythere tetrica Brady, topotype, BM 1974.338 (x78). 4. External view of left side of complete carapace. 5. External view of right side of complete carapace. 6. Dorsum of whole carapace. FIGS 7-9. Cythere laganella Brady, BM 81.5. 23 (xioy). 7. Dorsum of whole carapace. 8. External view of right side of complete carapace. 9. External view of left side of carapace. Bull. Br. Mus. nat. Hist. (Zool.) 29, 5 PLATE 27 A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) r. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96 ; 14 Plates. 1965. (Out of Print.) 2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125 ; 29 Plates 77 Text-figures. 1969. £4.50. 4. HAYNES, J. R. Cardigan Bay Recent Foraminifera (Cruises of the R.V. Antur) 1962-1964. Pp. 245 ; 33 Plates, 47 Text-figures. 1973. £10.80. 5. WHITEHEAD, P. J. P. The Clupeoid Fishes of the Guianas. Pp. 227 ; 72 Text-figures. 1973. £9.70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : the Biology and Evolution of a Species Flock. Pp. 134 ; i Plate, 77 Text-figures. 1974. £3.75. Hardback edition £6. Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 ;NU SOME TERTIARY AND RECENT CONESCHARELLINIFORM BRY6"^ f °> P. L. COOK AND R. LAGAAIJ BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 29 No. 6 LONDON: 1976 SOME TERTIARY AND RECENT CONESCHARELLINIFORM BRYOZOA BY PATRICIA L. COOK British Museum (Natural History) AND ROBERT LAGAAIJ Shell International Petroleum Pp. 317-376 ; 8 Plates ; 7 Text-figures ; 3 Maps BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 29 No. 6 LONDON: 1976 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Scientific Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 29, No. 6, of the Zoology series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation : Bull. Br. Mus. nat. Hist. (Zool.) ISSN 0007-1498 Trustees of the British Museum (Natural History), 1976 BRITISH MUSEUM (NATURAL HISTORY) Issued 27 May 1976 Price £5-40 SOME TERTIARY AND RECENT CONESCHARELLINIFORM BRYOZOA By PATRICIA L. COOK & ROBERT LAGAAIJ1 CONTENTS Page SYNOPSIS ........... 319 INTRODUCTION ........... 320 TERMINOLOGY ........... 321 STRUCTURE AND BUDDING ......... 323 ORIENTATION OF THE COLONY, ROOTLETS AND EARLY ASTOGENY . . 327 COMPARISON OF ASTOGENETIC SERIES . . . . . . 331 CODING OF CHARACTERS AND RESULTS OF POLYTHETIC CLUSTERING . . 333 EVOLUTIONARY TRENDS IN MORPHOLOGY ...... 342 ECOLOGY AND PALAEOECOLOGY ........ 343 DISTRIBUTION IN TIME AND SPACE ....... 346 CONCLUSIONS ........... 348 DESCRIPTIONS OF SPECIES ......... 349 Family ORBITULIPORIDAE ........ 349 Genus Batopora ......... 349 Batopora stoliczkai Reuss ....... 352 Batopora grandis sp. nov. . . . . . . . 353 Batopora aster izans sp. nov. . . . . . . . 354 Genus Lacrimula ......... 355 Lacrimula burrowsi Cook ....... 356 Lacrimula visakhensis Rao & Rao . . . . . . 357 Lacrimula perfecta (Accordi) ....... 358 Lacrimula borealis sp. nov. ....... 360 Lacrimula asymmetrica sp. nov. ...... 361 Lacrimula grunaui sp. nov. ....... 363 Lacrimula similis sp. nov. . . . . . . -363 Genus Atactoporidra ........ 365 Atactoporidra bredaniana (Morren) ...... 365 ACKNOWLEDGEMENTS ......... 367 SUMMARIES IN FRENCH AND GERMAN ....... 367 APPENDICES ........... 368 REFERENCES ........... 374 SYNOPSIS The characters of conescharelliniform and orbituliporiform colonies of Bryozoa, and the occurrence in both groups of two types of astogeny, 'normal' and 'frontal', are described. The genera Conescharellina and Trochosodon are known to be anchored to their substrata by rootlets. They are compared with the genera Batopora, Lacrimula and Atactoporidra. A hypothetical model for the primary colony development of these genera is suggested. The characters and distribution in time and space of the genera Batopora, Lacrimula, Atactoporidra and Cones- charellina are discussed, and the available information on their ecology and palaeoecology is 1 This paper was in the final stages of preparation at Dr Lagaaij's death in January, 1975. The sec- tions on the results of polythetic clustering and the conclusions had been discussed but were not then completed; they therefore represent principally my own interpretations. P. L. COOK 320 P. L. COOK AND R. LAGAAIJ recorded. Full descriptions are given of three species of Batopora, two of which are considered to be new, and of seven species of Lacrimula, four of which are considered to be new. The combined effects of genetics and environment outweigh microenvironmental influences within the colony. Integration within colonies is considerable, and is demonstrated by the interzooidal communications, astogenetic zonation and polymorphism. The specialized mode of life allows palaeoecological inferences to be made as to depth and type of sea-bottom from the Eocene to the Recent. INTRODUCTION MARINE cheilostomatous Bryozoa have evolved colony forms capable of inhabiting many environments. Members of the ascophoran families Orbituliporidae and Conescharellinidae particularly appear to be adapted to conditions unsuitable for many other forms. The members of the two families are closely similar in many characters, not least in the possession of a form of astogeny which may be unique among Bryozoa (see below). Both families exhibit parallel groups of genera with distinct colony forms, and it must be stressed that those which are here considered to be 'orbituliporiform' colonies are found both in the Orbituliporidae (e.g. some species of Orbitulipora) and in the Conescharellinidae (e.g. Flabellopora). Con- versely, 'conescharelliniform' genera are found in the Conescharellinidae (e.g. Conescharellina) and in the Orbituliporidae (e.g. Batopora). Other genera exhibiting similar colony forms have no close systematic relationship with these families and have an entirely different astogeny (see below). For example, 'orbituliporiform' colonies occur in Lanceopora and 'conescharelliniform' colonies in Fedora. This paper is principally concerned with the colony structure, diversity, distribu- tion and relationships of three conescharelliniform genera, Batopora (Eocene to Recent), Lacrimula (Eocene to Recent) and Atactoporidra (Eocene to Oligocene). The relationships and distribution of a fourth genus-group, comprising some representatives of the genera Trochosodon and Conescharellina, are briefly compared. Conescharelliniform colonies are small, rarely reaching 8 mm in height or diameter. They are conical, usually with no large basal concavity, and apparently without substratum. The ancestrula or ancestrular complex is concealed by secondary kenozooidal or extrazooidal tissue in later astogenetic and ontogenetic stages. Orbituliporiform colonies are frequently larger, often reaching 20-30 mm in height or diameter. They are actually or apparently bilaminar, and may be disc-shaped, sagittate or trilobate. One of the disadvantages in defining colony form in terms derived from names of genera which illustrate a distinct type of structure is that the terms themselves may begin to carry with them a systematic connotation. The term 'lunulitiform' (from Lunulites) is, however, now generally used for cup-shaped or conical colonies which are free-living, and which at some stage in their astogeny have a basal cavity. Although the overall form of these colonies is similar, the budding pattern, microstructure and interrelationships of polymorphs are totally dissimilar. Unrelated genera included in this grouping are, for example, Lunulites, Cupuladria, Selenaria and Cyttaridium. Conescharelliniform and orbituliporiform colonies have, in common with lunuliti- form colonies, an association with fairly calm to calm, often deeper shelf waters, and a soft unstable sea-bottom, with the concomitant problems to the bryozoan of CONESCHARELLINIFORM BRYOZOA 321 deposition and restricted availability of substratum for settlement of larvae. Unlike lunulitiform colonies, orbituliporiform and conescharelliniform colonies may inhabit abyssal depths and have been observed, or may be inferred, to possess cuticular rootlets as an essential part of the colony structure. The astogeny of both types of colony may be similar to that found in encrusting cheilostome species ; i.e. new zooid buds arise from the distal or distal-lateral walls of existing zooids, forming linear series of increasing astogenetic age in the direction leading away from the ancestrular area ('normal astogeny'). In other colonies of both groups new zooids are budded entirely from the frontal walls of existing zooids in a succession which is described in detail below (p. 324) as 'frontal astogeny'. Conescharelliniform and orbituliporiform genera exhibiting these forms of growth include : Conescharelliniform 'frontal astogeny' Conescharellina, Trochosodon, Batopora, Lacri- colonies mula, Atactoporidra, Fedorella 'normal astogeny' Fedora, Kionidella, Mamillopora Orbituliporiform 'frontal astogeny' Orbitulipora, Flabellopora, Zeuglopora colonies 'normal astogeny' Lanceopora TERMINOLOGY Classical terms in bryozoan morphology have rather wide definitions, but in practice some have been found to be satisfactory. There are considerable difficulties, however, in applying these terms to the genera discussed here. These stem in part from the peculiar structure of the colonies themselves, and in part from the fact that their orientation in life is unknown. Distal and basal walls as such do not exist in these colonies, and the classical 'distal' direction of the astogenetic process is apparently 'proximal'. The orientation of the colony in relation to the substratum is not known from direct observation. The term 'distal' usually refers to that direction, and by morphological analogy, to those zooidal walls, and parts of the whole colony, which are astogenetically 'away from' the ancestrular area. Similarly the conventional representation of conical, lunulitiform colonies assumes that the geometric apex of the cone is uppermost, as in these colonies the basal walls are, by direct observation of living forms, directed downwards (see Cook, 1963 ; Greeley, 1967). The terms used here are denned below (see Figs i and 2). Adapical - directed toward the apical region of the colony (the classically 'distal' part of the orifice is adapical). Antapical - directed away from the apical region of the colony. Apical region - the region of the colony in which the ancestrula and primary zooids may be observed or inferred to occur. Concealed frontal wall - that part of the frontal wall of each zooid which, except in the proliferal region, is hidden from view. Primary series of frontal buds originate from this part of the wall. 322 P. L. COOK AND R. LAGAAIJ ant FIG. i. Terms used in describing conescharelliniform colonies with 'frontal' budding. Note that the colony orientation in life is not known, ad, adapical direction ; ant, antapi- cal direction ; ap, apical region ; ax, colony axis ; pr, proliferal region ; s, colony surface ; zs, zooid series ; zw, zooid whorl. Exposed frontal wall - that part of the frontal wall of each zooid which surrounds the orifice and which contributes to the exposed surface of the colony between its geometrical apex and its geometrical base. Secondary series of frontal buds may originate from this part of the wall. Proliferal region - the region of the colony in which the most recently formed primary zooid buds (i.e. those of the primary zone of astogenetic change) may be observed or inferred to occur. Rootlets - long cuticular kenozooids or zooidal extensions arising from specific positions in a colony, actually or hypothetically functioning as anchoring structures. Rootlet pores - small areas, which may be kenozooids derived by frontal budding from frontal septulae, from which rootlets may be observed or be inferred to originate. Abbreviations used: AxL axial length of colony Prl proliferal region width of colony Lfw length of exposed frontal wall Ifw width of exposed frontal wall Lo orifice length lo orifice width Lov ovicell length lov ovicell width Lt length of apical tube BM British Museum (Natural History) NMV Naturhistorisches Museum, Vienna USNM United States National Museum CONESCHARELLINIFORM BRYOZOA 323 STRUCTURE AND BUDDING Apparent reversal of the orientation of the zooidal orifice occurs in two distinct forms in Bryozoa. In the Inversiulidae the operculum opens in a distal direction, but all other zooidal relationships appear to be normal (see Harmer, 1957 : 956). In the Conescharellinidae and Orbituliporidae the whole zooidal orientation is apparently reversed in relationship to the direction of budding of the colony. The classically 'distal' part of the orifice is thus directed towards and not away from the ancestrular region. Hypothetical models illustrating the methods involved in this reversal have frequently been made in the past, two of the most recent being those of Silen (1947) and Harmer (1957). They have been based on the assumption that budding in Bryozoa is primarily a function of the distal and/or lateral zooidal body walls. In the Cheilostomata, the astogeny of the great majority of species consists of the production of a primary uncalcified bud by the expansion of cuticle and underlying epidermis distally from an existing zooid. The bud usually proceeds to secrete calcified basal and lateral walls, and becomes limited distally by the growth of a transverse distal wall. Coelomic connection between and among zooids is through septulae in the lateral and distal walls. Recently, studies have been made on another type of astogenetic series in the Ascophora, the formation of 'frontal' buds (see Banta, 1972). Hastings (1964), in AD -K ADV AV. EXP ANT FIG. 2. Generalized morphology of a conescharelliniform colony with one zone of asto- genetic change. Colony has four zooid whorls and six zooid series budded alternately. Note condyles of zooid orifices, marginal pores, avicularia and ovicells. AD, adapical direction ; ANT, antapical direction ; CON, concealed frontal wall ; EXP, exposed frontal wall ; T, position of internal or external apical tube ; K, apical area of kenozooids, and/or closed zooids or extrazooidal tissue ; AV, interzooidal avicularium ; ADV, adventitious avicularium ; ov, ovicell. 324 P. L- COOK AND R. LAGAAIJ describing the colony structure of Reginella doliaris (Maplestone), a cribrimorph species with a free-living conical colony form, postulated a kind of frontal budding, and compared the astogeny with that of Conescharellina. Although there may be several methods by which these frontal buds are formed, the sequence of development is basically as follows. There is an expansion of cuticle and underlying epidermis above the calcified frontal wall of an existing zooid. Coelomic connection is provided by frontal septulae, which seem to be represented by some or all of the marginal pores. The buds so produced have no cuticle basally, and some do not appear to have lateral walls. Marginal septulae are present. Some colonies may show long series of frontal buds which develop in a plane at right angles to that of the primary growth (see also Boardman, Cheetham & Cook, 1969, fig. 5). Other colonies may produce small groups of frontal buds, which then proceed to bud distally and laterally, forming a secondary layer of zooids above and parallel to that of the primary growth (similar to an 'overgrowth' of Boardman et al., 1969, fig. 6). Zooids of this kind of secondary layer may therefore be distinguished from frontally budded layers by the possession of a basal wall and basal cuticle. In the Conescharellinidae and Orbituliporidae normal distal or distal-lateral budding appears to have been completely abandoned in favour of frontal budding. Each zooid may be considered to have a calcified frontal wall which has extended at the expense of the distal, lateral and basal walls. At the same time, the frontal wall has become divided into two elements. The first element is a flat or inflated, often hexagonal portion which represents the projection of the geometrical base of a cone at the surface of the colony. The second portion consists of part of the remainder of the cone. The 'basal' wall of each zooid completes the cone, but this is composed of the frontal wall, or walls, of the parent zooid or zooids (see Figs 2 and 3). Marginal pores, which are inferred to be septulae, are placed round the periphery of the hexagonal portion of the frontal wall and in two converging rows on its remaining surface ('exposed' and 'concealed' frontal wall elements, see p. 321). The next generation of frontal buds is produced either directly from one or from between two existing zooids, arising in a line extending along the 'concealed' part of the frontal wall toward the centre of the colony, and thus including two series of marginal pores. As the bud enlarges it completes the conical shape, and the en- larged frontal wall calcifies, the geometrical base of the zooid cone again forming the hexagonal exposed frontal wall at the surface of the colony, and surrounding a centrally placed orifice. In both families, buds are produced in direct or in alternating series, i.e. only one, or two to three zooids of the previous generation contribute to the next generation. In the Conescharellinidae the type of budding is usually direct and specific. It was illustrated by Harmer (1957) for most species of Conescharellina and Trochosodon (but C. ovalis, p. 743, buds in alternating series). In the Orbituliporidae, most species bud in alternating series, and there is a good deal of intracolony variation. Even in zooids budded directly, there may be a variable contribution from other zooids, potentially at least, as their marginal pores are incorporated beneath the frontal wall of the new bud. Some colonies, for example the largest of Lacrimula burrowsi, show an initial secondary zone of alternating budding, which, as it gives CONESCHARELLINIFORM BRYOZOA 325 rise to a few intercalary series of zooids, becomes somewhat irregularly direct at the later astogenetic stages. A few species do appear to be regularly directly budded ; these are Batopom ernii as figured by Dartevelle (1948) and Lacrimula visakhensis. Batopora murrayi also has strong tendencies to directly budded series of zooids. The very regularly directed budded zooids figured by Reuss (1867), in Batopora rosula are not in fact of this type (see p. 351). If the first circle of buds is regarded as consisting of frontally budded zooids from an ancestrular complex (see below), the apparent reversal of normal zooid orientation is explicable. Theoretical explanations of reversal of the operculum and viscera, or of the proportions and roles of the basal, lateral and distal walls (see Harmer, 1957), are not necessary. In other, unrelated genera colonies do exist in which instead of a frontal wall development at the expense of other walls, the frontal wall is very restricted, and the lateral and distal walls greatly increased in extent, as postulated by Harmer (1957) for Conescharellina. It is interesting that these colonies have a conical shape, and may have a mode of life similar to that of the frontally budded colonies described here. For example, the lunulitiform genera Anoteropora and Actisecos are associated with ecological conditions similar to those in which the orbituliporiform and conescharelliniform genera are found. Conescharelliniform colonies belonging to the Conescharellinidae and Orbituli- poridae may be considered to have a primary zone of astogenetic change which never develops into one of astogenetic repetition (see Boardman et al., 1969), because the zooids become progressively larger throughout the budding series. In some forms, such as Atactoporidra and some species of Lacrimula and Batopora, the primary zone may develop almost concurrently with, or be replaced by, a secondary zone of change. This secondary zone comprises a secondary series of buds which are produced from the exposed frontal walls of zooids of the first zone of change (see Fig. 3). In Atactoporidra, further series of such frontal buds may eventually form a primary zone of astogenetic repetition. In the genera studied here, interpretation of colony structure, and even recognition of species, is complicated by the development of secondary and occasionally tertiary zones of change. These may involve part of the colony or the whole colony, and are accompanied by astogenetic and ontogenetic changes in the apical region. During the course of this work we have been fortunate in having been able to examine several populations which show almost complete astogenetic series. This has not only enabled us to postulate a model for the earliest astogenetic stages, but has made it possible to infer parallel series of changes in less representative or less well-preserved populations. Secondary, frontally budded zones of change are present in some species of all three genera principally studied. The secondary zone originates adapically in Lacrimula and Batopora multiradiata, and follows a regular, but entirely different pattern in the two genera. In Lacrimula (see Fig. 36) each apical zooid produces one, or occasionally two, frontal buds directly from its exposed frontal wall. Further secondary zone buds arise in a similar fashion and in a regular sequence in an antapical direction. The adapical zooids frequently have closed orifices at the stage when the secondary buds are produced, but in more antapically placed zooids of the 326 P. L. COOK AND R. LAGAAIJ ad ant FIG. 3. Patterns of secondary budding in conescharelliniform colonies with 'frontal' astogeny. pz, primary zone of astogenetic change ; sz, secondary zone of astogenetic change ; sr, secondary zone of astogenetic repetition, ad, adapical direction ; ant, antapi- cal direction. A. Primary zone followed by antapically directed overgrowth of frontally budded secondary zone of change (e.g. Batopora multiradiata). B. Primary zone followed by a secondary zone of directly frontally budded zooids. Sequence of budding antapically directed (e.g. Lacrimula). C. Primary zone followed by isolated interzooidally budded secondary zooids (e.g. B. clithridiatd) . D. Primary zone followed by directly or inter- zooidally budded secondary zooids eventually forming a zone of repetition. Secondary buds often arising antapically, and sequence of budding adapically directed (e.g. Atacto- poridrd) . primary zone, the orifices are not closed by calcification, and may be seen through the orifice of the secondary frontal bud. In B. multiradiata (see Fig. 3A) the second- ary zone arises in the same manner as in Lacrimula, as a circle of apical frontal buds. These zooids then proceed to bud the next generation antapically from their con- cealed frontal walls. This secondary zone advances, often very regularly, over the primary zone zooids. It is thus an overgrowth, but one which consists of frontally budded zooids, not those of 'normal' astogeny. In all other species of Batopora secondary zooids may be produced randomly and irregularly between primary zooids as interzooidal frontal buds (see Fig. 36). In Atactoporidra, the secondary zone arises antapically, in contrast to Lacrimula, and may extend adapically. Zooids arise as interzooidal frontal buds and in some colonies this process is continued through three generations of zooids and results in a zone of repetition (see Fig. 3D). The astogeny of Conescharellina is basically of a similar pattern but differs in that the axial part of the proliferal region is filled with kenozooids, and sometimes avicularia, budded between zooids from the axial part of the concealed frontal walls. As the colony increases in size, avicularia and kenozooids arise between zooids along CONESCHARELLINIFORM BRYOZOA 327 the whole length of the concealed frontal wall. When colony growth ceases (at various astogenetic stages which may be correlated both specifically and with unknown environmental conditions) the entire antapical area becomes covered by kenozooids and avicularia. These form a secondary zone of astogenetic change and eventually of repetition, but secondary zones of autozooids do not seem to occur in Conescharellina. A form of growth somewhat analogous to that of Atactoporidra is found in some 'celleporiform' colonies which grow on hydroids. Here the primary zone of change is formed of a small number of zooids encrusting a very restricted substratum. The major part of the colony consists of zooids of a secondary zone of change and repetition formed by frontally budded erect interzooidal zooids. In orbituliporiform colonies a primary zone of astogenetic repetition is usually established after a variable number of generations. The zooids do not continue to increase in size, but intercalary frontally budded series are produced in both laminae. Conescharelliform and lunulitiform genera have somewhat similar shapes and it is interesting to compare the role played by zooidal size in colony structure. In lunulitiform colonies, especially those belonging to the Cupuladriidae, which have a triadic ancestrular complex, the conical shape is achieved by the budding of inter- calary rows. After the zone of repetition is established, there is little or no increase in zooid size. Even in Cupuladria doma, where the highly conical colony resembles that of Conescharellina, and where there is a small but continuous increase in size, a colony which begins with a zooidal triplet will have 30 closed peripheral zooids when it reaches full size (see Cook, 1965, fig. 3). In conescharelliniform colonies, relatively few intercalary series of zooids are produced, the conical shape resulting from a steady increase in size of the zooids of successive whorls. There are, of course, exceptions to this rule (see p. 366). ORIENTATION OF THE COLONY, ROOTLETS AND EARLY ASTOGENY The colonies considered here all have structures which have been observed, or may be inferred to have been associated with cuticular rootlets, which served as organs of attachment. Orientation in life of conical and bilaminar colonies with rootlets has been the subject of a great deal of speculation in the past (see Silen, 1947 ; Harmer, I957 : 724-725) • It is possible that some colonies may live suspended by rootlets from algal fronds, or from hydroids, worm tubes or other Bryozoa. It is equally possible that they may be supported at or above the surface of a soft and unstable sea-bottom by rootlets which function as an anchor. These alternative theories of mode of life are not mutually exclusive, and the rootlets could perform either function. Until observations are made upon living colonies, and their larval life and settle- ment preferences are known, discussion of orientation and mode of life must remain purely speculative. Generally, those rootlets which have been observed have very thin cuticular walls, and may therefore not be comparable in function with those found in the Scrupocellariidae and Petraliellidae, which are strong enough to support the colonies above a substratum. Numbers of thin-walled rootlets would, however, have 328 P. L. COOK AND R. LAGAAIJ sufficient strength to support the smaller orbituliporiform colonies. Colonies of Flabellopora and Lanceopora, which are flattened and sagittate, and may be more than 30 mm in height or diameter, are known to have numerous rootlets which originate in the ancestrular region, and which are associated with kenozooids and extra- zooidal tissue. From evidence of adherent Foraminifera and sand grains, these colonies are inferred to be anchored by their rootlets at the surface of the soft and unstable sea-bottoms with which they are associated. Many of the stations which have yielded specimens of these genera have also provided numerous specimens of Conescharellina (see Canu & Bassler, i92Qa ; Silen, 1947 ; Harmer, 1957). Rootlets have been seen in Recent specimens of Conescharellina and the closely related genus Trochosodon. These arise, principally, in the apical region, from specialized rootlet pores ('lunoecia'), which are apparently modified kenozooids budded frontally among the autozooids (see PL i, figs 4 and 5), and which them- selves may bud frontal avicularia (see Harmer, 1957 : 742). Not all forms of Conescharellina have distinct rootlet pores, however, but have complexes of keno- zooids in the apical region which may form a solid structure, surrounded by avicularia, as in C. africana (see Cook, 1966), or a distinct tube, as in colonies of an unnamed species from Zanzibar (see Appendix 3, p. 372). These apical structures are very similar to those found in Lacrimula and some colonies of Batopora. The apical region in Batopora has usually been described as a 'pit'. In many colonies, a round cavity with thick calcified walls is present, surrounded by a circlet of zooids or keno- zooids. In other colonies the cavity is surrounded by a raised series of kenozooids forming an external, often elongated tube. The sequence of astogenetic and onto- genetic changes is not fully known, and may be environmentally influenced, as different forms of apical structure occur among colonies of similar astogenetic age in a single population. In Batopora and Lacrimula, subsequent ontogenetic changes include a thickening of the calcified walls, and astogenetic changes consist of a proliferation of frontally budded kenozooids, presumably by adapically directed growth of units from the exposed frontal walls of the primary kenozooid and zooids. The kenozooids each have a small uncalcified area, from which it is inferred that rootlets could originate. In L. asymmetrica, the primary kenozooid is itself very elongated and tubular (see Fig. 6, p. 362). It appears to enlarge and become thicker walled during colony development, but whether this is an ontogenetic change, an astogenetic change or a change involving extrazooidal colony-wide calcification is not known (see PI. 5, fig. 4). Small uncalcified areas are present upon both the external and internal surfaces of the tube. In some colonies of B. rosula, and in B. scrobiculata, the tube is apparently formed by a secondarily-budded kenozooid, which has similar uncalcified areas on its outer walls. In other species of Lacrimula, and in Atactoporidra bredaniana, the tube is composed of small kenozooids, which arise adapically as a regular series from the primary zooids (see PI. 5). This also occurs in some colonies of B. multiradiata. Other changes in the apical region, found particularly in Lacrimula, include progressive, antapically directed closure of the orifices of the primary and later-budded zooids. At the same time, the ex- posed frontal walls of these apical zooids become thickened, perhaps extrazooidally. Evidence of some developmental sequences has been seen in very young colonies. CONESCHARELLINIFORM BRYOZOA 329 In B. stoliczkai, B. murrayi, L. asymmetrica and Trochosodon sp. a kenozooid appears to be one of a primary group of zooids. The walls of the kenozooid are calcined, but there is always a large, round, uncalcified area adapically. It is inferred that this marks the position of origin of an uncalcified rooting or anchoring element. In L. asymmetrica the primary complex consists of five zooids and a long, tubular kenozooid (see Fig. 6) . One of the five zooids is asymmetrical and may have been developed before the remaining four. If the kenozooid was involved in the growth of a rootlet, it was present from the earliest stage, and remained prominent, becoming larger in subsequent growth stages of the colony (see p. 362). In B. murrayi there is a primary pair of zooids, followed by a triad. The exposed frontal walls of the primary pair and two of the subsequent triad surround a large rounded apical cavity which itself has calcified walls. This is inferred to mark the site of an im- mersed primary kenozooid. In B. stoliczkai (see PI. I, fig. i) there is a primary zooid pair and a large kenozooid with a prominent round aperture. Later stages show a 'pit' surrounded by calcification or kenozooids at this point, and it is inferred that the kenozooid is part of the primary group. A young colony of Trochosodon sp., a genus known to have rootlets, shows a zooidal triad and a large rounded cavity which is inferred to mark the position of a primary kenozooid. Although this colony comprises only ten zooids, it has secondarily budded apical avicularia and three semilunar rootlet pores. The general similarity of the modes of growth of these colonies is shown in Fig. 6 and on PI. i, figs 1-6. Harmer (1957 : 748, fig. 78) made some interesting observations on the early development of Trochosodon. He remarked : There is doubtless some variation in the details of the early development.' He concluded that in some specimens of T. optatus there was an ancestrula and paired primary zooids. In view of its prominence, it is possible that the 'ancestrula' was an apical kenozooid. The specimen of T. linearis figured by Harmer (1957, fig. 75) was drawn from the 'basal' side by transmitted light. In reflected light from the adapical side it shows a large central cavity, with calcified walls, filled with darkly stained tissue, unlike that of the surrounding zooids. It is surrounded by one pair of very small zooids on one side, and a pair of slightly larger zooids on the other side. It is possible that the central cavity represents a primary rootlet element arising from a kenozooid. The presence of rootlet pores in the early astogenetic stages of Trochosodon suggests that the analogous series of apical structures and the primary kenozooid in species without rootlet pores fulfil a similar function, especially as the colonies inhabit similar environments (see also p. 346). The ancestrular area of conescharelliniform colonies is considered apical in the geometric sense. In the bilaminar orbituliporiform colonies the theoretical colony cone may be regarded as bilaterally compressed, with a consequent distortion of the apical region to one side of the colony. Development of kenozooidal and/or extra- zooidal tissue complicates the structure of the apical region in both groups. The exact nature of the ancestrula is not known, and as the calcified parts of the colony are not attached to, nor incorporate a recognizable substratum (as, for ex- ample, in the Cupuladriidae, which incorporate a small sand grain or foraminiferan), the earliest astogenetic changes may only be inferred. It is probable that a single P. L. COOK AND R. LAGAAIJ ANT AD FIG. 4. Hypothetical early astogeny of a conescharelliniform colony, assuming that the apical tube produces rootlets anchored in a soft substratum. The orientation in life is unknown, but this form of astogeny would be applicable if the colonies were suspended by rootlets. AD, adapical direction ; ANT, antapical direction ; PK, primary kenozooidal rootlet ; SK, secondary kenozooidal rootlet ; i, first zooid whorl ; 2, second zooid whorl. A. Metamorphosed larva producing an ancestrular complex comprising one kenozooid and three zooid elements. B. First whorl of zooids complete, kenozooidal tube developing and forming primary rootlet. C. Second whorl of zooids complete, apical kenozooids forming round apical kenozooidal tube. D. Third whorl of zooids developing, apical tube now comprising several secondary kenozooids with secondary rootlets. ancestrula with calcified walls is not developed at metamorphosis. Instead the primary whorl of zooids or kenozooids may be simultaneously differentiated within an uncalcified ancestrular complex which also includes a rootlet element. Develop- ment of such a kenozooidal rootlet or rootlets would rapidly separate the budding locus from the substratum, into which the rootlet system would become attached or anchored. Later development of apical kenozooids could also provide further rootlets, while the primary zooid whorl budded successive generations of zooids frontally in the opposite direction (see Fig. 4). This would account theoretically both for the inferred method of life, and for the absence of a substratum attached to a single, calcified ancestrula. Ancestrular complexes of 2-4 zooids are known (see Eitan, 1972, and Cook, 1973). Rootlet development may be very rapid. For CONESCHARELLINIFORM BRYOZOA 331 example, living colonies of Hippopetraliella africana from Ghana have been seen to develop basal rootlets 5 mm long in 48 hours. Opposing directions of growth from an ancestrular focus were described by Harmer (1957 : 794-795) for colonies of Adeona with complex systems of kenozooidal rootlets. Bryozoan colonies may also be established by kenozooids, feeding zooids not appearing until later astogenetic stages, as in Scruparia. Thus the necessary elements for the hypothetical astogeny given in Fig. 4 do occur in Bryozoa, although they have not all been directly observed in such a combination. A demonstration by Dr G. Eitan, exhibited in September 1974 at the 3rd Inter- national Conference of the International Bryozoology Association (Department of Geology, Universite Claude-Bernard, Lyon), showed that the early astogeny of Margaretta, an erect jointed form, was very similar to that postulated for cone- scharelliniform colonies. The ancestrular complex included kenozooidal rootlets, and was adhesive, but not adherent to the substratum. Lagaaij (ig63b : 203-207), discussing the possible mode of life of Fedora, postulated anchoring rootlets arising from 'special chambers' associated with the zooids. One such chamber was observed with a fine, cuticular rootlet intact. He also described colonies which had incorporated fine grains of substratum material. One colony of Batopora clithridiata (see p. 351) from the Eocene (Hampstead, London Clay, BM 69554 pt.) shows a similar growth form. At some point in its astogeny the colony has incorporated a small sand grain, 0-60 mm in diameter. The apical zooids surround part of the sand grain, but are very irregular in shape and may belong to a secondary zone of change. The remaining zooids are budded frontally in regular alternating series. COMPARISON OF ASTOGENETIC SERIES Astogenetic series within a single population are rare, but it has been possible to trace the astogeny almost completely in Lacrimula asymmetrica (see p. 361) and partially in L. grunaui. Comparison of the number of zooids per whorl with the number of whorls and the total number of zooids in a colony has shown that there are some fundamental differences among species which are related to their astogeny. For example, if there is little increase in the number of zooids per whorl throughout astogeny in the primary zones of change, the number of zooids in a colony should increase arithmetically. Differences between estimated and actual totals should be attributable to other observable astogenetic changes. This hypothesis was tested in samples which contained several colonies at different astogenetic stages (see Table i). In Batopora murrayi the number of zooids falls below that estimated because avicularia and kenozooids actually take the place of entire zooids in series. In B. stoliczkai the primary triad comprises one kenozooid and two zooids, and in both B. stoliczkai and B. clithridiata the actual number increases rapidly above the esti- mated figure as scattered, secondary frontally budded zooids occur with greater frequency. The same reason for increase occurs in one B. rosula colony from Crete. The larger colonies from Baden appear to have only four zooids per whorl in the 332 P. L. COOK AND R. LAGAAIJ TABLE i Comparison of astogenetic series Batopora murrayi B. stoliczkai (first whorl has two zooids only) B. dithridiata B. rosula (Crete) B. rosula (Baden) B. multiradiata Lacrimula asymmetrica L. grunaui L. perfecta No. of zooids No. of per whorl whorls 5 3 5 4 5 5 3 3 3 4 3 5 3 6 3 7 4 3 4 4 4 5 4 6 5 4 5 8 4 4 5 4 7 4 7 7 7 6 7 7 7 9 7 12 4 6 4 8 4 ii 5 i 4 2 5 3 4 4 5 5 4 6 4 5 4 6 4 9 5 6 6 6 6 8 (estimated) 9 ii 9 20 „ Estimated no. Total observed of zooids no. of zooids 15 15 20 18 25 20 8 8 II 15 14 18 17 22 20 30 12 13 16 18 20 28 24 33 20 21 40 38 16 17 20 21 28 30 49 48 42 43 49 49 63 66 84 80 24 56 32 60 44 66 5 5 9 9 14 H 18 18 23 23 27 27 20 20 24 24 36 36 30 30 36 36 48 46 99 86 1 80 160 CONESCHARELLINIFORM BRYOZOA 333 first whorl, but there does seem to be a production of intercalary series of zooids in the primary zone of change, which accounts for the larger totals at the same asto- genetic stage as the remaining colonies from Crete. The actual numbers may fall below the estimated numbers because the production of intercalary series is not absolutely regular. Young stages of B. multiradiata resemble some of the larger colonies of B. rosula from Baden in number of zooids, which is very close to the estimated figure. They may be distinguished from nearly all other specimens of Batopora by the number of zooids per whorl. The colony and zooid size is, however, small. Young colonies with six to seven zooid whorls comprise 43-49 zooids ; a colony of B. scrobiculata of comparable size (about i mm high and 2 mm wide) has only 24 zooids, and one of B. murrayi only 15. The colonies of B. grandis differ completely in their very large number of zooids yet apparently low number of zooids per whorl. In L. burrowsi and L. perfecta, the proliferal region, comprising relatively few zooids of the primary zone of change, merges with a secondary zone of change, including intercalary rows, and contains a larger number of zooids. In L. perfecta the apical region is rapidly covered by a proliferation of kenozooids, and the number of whorls in this region has been estimated by comparison with younger colonies. The actual number of zooids is less than that estimated, as the number of zooids in the primary whorl is unknown in large colonies, but due to the introduction of intercalary rows, is less than that in the proliferal region, on which the estimate for the whole colony is based. In L. asymmetrica the number of zooids per whorl regularly alternates between four and five, because one zooid is always developed slightly earlier than the others in alternate whorls. The number does not increase arithmetically, but the agreement between the estimated and actual numbers in these colonies and those of L. grunaui is a measure of the regularity of growth and the absence of any secondary zooids at these astogenetic stages. CODING OF CHARACTERS AND RESULTS OF POLYTHETIC CLUSTERING The advantages of studying morphological relationships of bryozoan populations by polythetic clustering have recently been discussed by Cheetham (1968) and by Boardman et al. (1969). Apart from supplying a visual display of degrees of morphological similarity it may reveal new aspects of problems which have been hidden by previously accepted taxonomic concepts. In addition, the discipline of producing a coding for comparison of characters means that the basic nature of the 'characters' themselves is re-examined and that the specimens are subjected to a consistent series of observations and analysis (see Boardman et al., 1959, fig. i). After analysis and clustering, under ideal conditions, the morphological relation- ships displayed can be tested for systematic relevance by plotting the clusters in a time-space context (see Boardman et al., 1969, figs n and 12). During work on the conescharelliniform species described here, we became aware of a number of exceptions to the generally accepted character-correlations within the populations studied. This was particularly noticeable when specimens were being 22 334 P. L. COOK AND R. LAGAAIJ assigned to nominal genera using the classical concepts of such groups. Great variation within samples of what appeared to be 'species' was also found, although to some extent this could be attributed to astogenetic and ontogenetic differences. Some characters are listed in Table 2, and their occurrence compared among species assigned to the genera illustrates how arbitrary some of our decisions have been. TABLE 2 Occurrence of characters among species. i. External apical tube 2. Colony- wide secondary zones of zooids 3. Rootlet pores (lunoecia) 4. Axial kenozooids 5. Avicularia 6. Condyles 7. Sinus 8. Ovicells laterally displaced 9. Ovicells apical Conescharellina i species absent nearly all species nearly all species all species all species all species C. catella C. africana Batopora B. rosula B. multiradiata B. grandis B. multiradiata absent B. rosula B. scrobiculata B. asterizans B. murrayi B. clithridiata absent B. grandis B. grandis not apical Lacrimula all species L. asymmetrica L. similis absent L. perfecta L. visakhensis L. similis nearly all species nearly all species L. asymmetrica L. grunaui L. similis not displaced L. visakhensis Atactoporidra A, bredaniana all species absent absent A. bredaniana absent not displaced not apical In order to test concepts of both 'species' and 'genera', characters were chosen, analysed and used to assess similarity among the specimens examined. One of the most useful results of this analysis has been the demonstration of factors which must be considered both when deciding character states and when interpreting the clusters. The biology of living specimens of conescharelliniform species is virtually unknown, and the characters used here, both quantitative and qualitative, represent only a small part of those potentially available in Bryozoa. Much finer examination of plentiful colonies may reveal, for example, characters associated with calcification of walls, intercommunication of zooids, and detailed structure of orifices, avicularia and ovicells. Further information may come from investigation of the astogenetic and ontogenetic changes occurring in the apical region. CONESCHARELLINIFORM BRYOZOA 335 We are here dealing with Bryozoa in which both the sample and colony size are often very small. The interaction of genetic relationships and environmental influences has apparently evolved great similarity in colony structure and zooid form among samples. Conversely, astogenetic and ontogenetic changes produce large morphological differences within samples. For example, one possibility arising from comparison of colonies at different astogenetic stages, or where relative asto- genetic ages cannot be inferred, is that clusters may reflect age similarities more than 'taxonomic' similarities. Comparison of characters of similar astogenetic age is difficult, as a zone of repetition is not usually present. Although both absolute and relative rates of astogenetic and ontogenetic change are unknown, one approximate guideline available is the comparison of colonies with the same, or nearly the same, number of zooid whorls. Because colonies rarely bud in regularly alternating or direct patterns (see p. 324), it is not always easy to decide how many zooid generations are present, and because the numbers of whorls is often low, errors are correspondingly significant. Much of the material examined is fossil, and differences among and within samples may therefore be partly the result of differential conservation. Characters and character states used are given in Table 3, and a list of the specimens in Table 4 (see also Appendices 1-3). The following notes explain some of the concepts used in defining the characters, and why difficulties were sometimes encountered in deciding which of the character states was present. Characters i and 2. The axial length and proliferal region width of a colony not only give a measure of absolute size, but describe its shape. In conjunction with the number of zooid whorls it also gives a secondary measure of zooid size range and arrangement. Characters 3, 4, 14, 27 and 28. The type, number, position and nature of astogenetic zones. All colonies comprise at least one zone of change. Coding the presence of secondary zones was confined for these characters to those consisting of zooids only. The kenozooids, interzooidal avicularia and apical structure are considered separ- ately, as it is not known exactly when and in what sequence these may be secondary in occurrence. The numbers of whorls of zooids is counted as the number of series which it is inferred were budded simultaneously, whether alternately or directly. The number of zooids in each whorl can be inferred from the proliferal region, and in many colonies is the same or very few more than the number in the primary whorl. Character 5. Number of zooids at the surface. Here only forms with one zone of change are strictly comparable, but the presence of very large numbers of zooids is often directly correlated with the presence of a secondary zone of change or zone of repetition. Characters 6, 7, 10 and u. The maximum length and width of exposed frontal wall. This is the equivalent of the 'classical' zooid length and width usually measured for cheilostomes. Where possible, the measurement of the third whorl zooids was taken as this gives one of the few estimates of size among colonies at a comparable asto- genetic stage. The size of the subproliferal zooids is correlated with the number of whorls and the size and astogenetic age of the colony, and gives an estimate of the 336 P. L. COOK AND R. LAGAAIJ <*-i r1 ° o *_1 S S 'f 3s.iili 'O 5 3 § ex 3 CD I.FB.BJ 2 _rt 35 CD (3 C eg CD S*f rt Q ^^ £ o 2 I-! H 0 N 0 C/) O CD « r5 CUD § 5 M CD O -(-> c "S, CO d IH CD 4-> 13 rt 0 w ^j -(-< - o \ji PQ T) ^5 a H rt CO C CD O d 13 T3 B d i-i 3 en CU -t-> s O 3 s O en < u O 12 "O O o o ^ N 3 O % G W A) O (yl O ;£ Xi :r£ rt-'S o g < ^ ?f R §- .3 o h - \ CJ I! o o o o § § o ^ "S 'o C o CD N | O ^§ 1 1 i •* I •S - o .« g rtl C/3 ^ o « 2 3 13 £ ^ § y -8 o o ^ ^ Xi '* ic 3 «„ SH W O IH bo 10 O CONESCHARELLINIFORM BRYOZOA 337 Ill ill •s a g is >> 2 ^ rj. 'R +5 ft >> f- re •s •§ 1 ? + £ re ro A £8 a lu IH ni G Jj • I *w ;les finely granul; 5-20 (i I narrower tha half width 1 o •8 T3 c re 13 0 'ft re scattered adventitious 1 13 G HH "re 42 re o ' ' -o 3 (-{ 3- -4-1 0 W§ ° v $ S IH (0 ft -M U 0) 12 1 0 0 d O 13 ^0 re interzooi •»* G 0 ft 1 ed ft ri 1 I o ~ O a IH O a IH 0 O § 1 J fl ^ §^ o ^ .2 S sl ft b If ?— i \i/ re w ir! **H «£ o d 'So PH w O G N o o <•£>£•"£ o •r1 -r" G ^ N ^ > ° re T3 338 P. L. COOK AND R. LAGAAIJ TABLE 4 List of specimens analysed (see also Fig. 5) Asto- genetic Number group i A 2 B 3 B 4 A 5-6 B 7 D 8 B 9-10 C ii B 12 B 13 A 14 & 16 A 15 B 17-18 A 19 B 2O-2I A 22 C 23 B 24 D 25-26 C 27 B 28-30 C 3i &33 C 32 B 34-35 D 36-37 C 38 D 39 C 40 B 41 D 42 D 43 C 44 D 45 D 46 D 47 C 48-49 D 50-52 D 53 D 54 C 55-56 D 57 A 58 C 59 D 60 D 61 C Name Batopora multiradiata Batopora scrobiculata Batopora grandis Batopora multiradiata A tactoporidra bredaniana Lacrimula burrowsi Lacrimula pyriformis Lacrimula sp. Lacrimula visakhensis Lacrimula similis Lacrimula asymmetrica Lacrimula grunaui Lacrimula borealis Batopora sp. Batopora rosula Batopora murrayi Batopora rosula Batopora multiradiata Trochosodon sp. Batopora clithridiata Reference Locality Age K 48-50 Spain Oligocene USNM 71205 Italy Eocene USNM 88881 M M USNM 71205 ,, ,, USNM 88881 tt . USNM 88882 Bavaria BM 031117 USNM 71205 BM 07864 BM 1965.8.24.11 BM Stn 103 BM 1965.8.24.13 BM Stn 98 BM Stn 126 BM 1970.8.8.1 BG3I2 G567I Italy Belgium Zanzibar Recent E. India Indonesia Holocene Miocene 43/3-1 North Sea Oligocene SM55 Italy Miocene 6-27 Crete " Huelva Spain Pliocene PLR 4607 Malta Miocene Rii03 Spain ,, BMI965.8.24.6 Zanzibar Recent BM Chall. 1740 Fiji ,, NMV fig. 7 Austria Miocene NMV fig. i ,, „ NMV M " BM Italy Eocene BM Stn 126 Zanzibar Recent BM BI357C England Eocene BM 040339 ,, ,, BM 61357 ,, ,, CONESCHARELLINIFORM BRYOZOA 339 TABLE 4 (cont.) Asto- genetic Number group Name Reference Locality Age 62 C Batopora asterizans K52 Netherlands Oligocene 63 D » » t f )( M 64 D ,, ,, 43/3-1 North Sea M 65 D Lacrimula perfecta BM Italy Eocene 66-67 C ,, lt it 68 C Conescharellina africana BM 1949.11.10.639 Durban Recent 69 B » » M 70-72 C Batopora stoliczkai NMV Biinde, Oligocene Germany 73-74 C ,, ,, USNM Calbe, ,, Germany 75&77 C ,, ,, NMV Biinde, ,, Germany 76 78 C Batopora sp. „ M M 79 D 80 A Atactoporidra glandiformis BM 030077 England Eocene rate of astogenetic change when compared with the third whorl measurements. In a few cases, both sets of measurements are identical, as only four whorls were present. Characters 8, 9, 12 and 13. The size of the orifice generally increases with each whorl ; the characteristics of these measurements are similar to those of the exposed frontal wall. Characters 15, 16, 17 and 22. The astogeny of the apical region. This is not easily inferred and the choice of states is somewhat arbitrary. In some forms of Batopora, Lacrimula and Atactoporidra, distinct units of calcification, presumably kenozooids, each with a small central pore, form the apical region as an external tube. In other forms such kenozooids are absent and the tube appears to be extrazooidal in structure, or an internal tube is present, which may also be composed of either kenozooidal or extrazooidal tissue. The correlation of types of structure with asto- genetic age or sample is not clear. In Batopora, the internal tube may be obscured by kenozooids, or replaced by a secondarily budded kenozooid which may or may not be surrounded by other small kenozooids. In some samples an astogenetic series is present which enables the sequence of appearance of the structures to be inferred, but generally there is a great deal of variation. Kenozooids and extra- zooidal tissue tend to merge with closed zooids in the apical region. In Lacrimula, closure consists of a calcined lamina filling the zooidal orifice, often leaving a small pore or slit centrally. Small kenozooids are often found among the zooids in Batopora, presumably budded frontally, but whether secondary to the primary zone of zooids is not known. They may occur adapically to antapically, and in some forms appear antapically in the axial region of the proliferal zooids. 34o P. L. COOK AND R. LAGAAIJ Character 18. Inflation of the frontal walls. Inflated walls are positively correlated with long, tubular peristomes in Recent forms. Fossil specimens are usually worn but the subproliferal zooids may show distinct states of inflation and sometimes elongated peristomes. Character 19. The degree of tuberculation of the exposed frontal wall may be affected by preservation. The majority of colonies thus appear to be 'finely granular,' but some are consistently smooth and others coarsely granular. Character 23. Avicularia are not generally common and are usually interzooidal, budded frontally among zooids in a similar manner to the kenozooids. Structures inferred to be adventitious avicularia are very rare and occur in one species only ; they are associated with the edge of the peristome. Characters 24, 25 and 26. All ovicells seen are hyperstomial and usually appear to have been closed by the operculum. Some are large and prominent, others small and immersed, a few are laterally displaced. The presence of ovicells may indicate astogenetic maturity, and they are often present only in the proliferal region. Ideally, comparison should be made only among colonies from each sample which are at exactly the same astogenetic stage, but this was not possible with the material available. Eighty colonies were therefore divided into four groups of roughly com- parable astogenetic age. The division was not completely arbitrary, but endeavoured to include colonies from as many samples as possible in each group (see also Table 4) . Group A comprised n colonies with 15-22 astogenetic generations from 7 samples. 3 of which were also represented in Group B. (Nos i, 4, 13, 14, 16, 17, 18, 20, 21, 57, 80.) Group B comprised 14 colonies with 9-13 astogenetic generations from n samples, 3 of which were also represented in Group A, 6 in Group C and i in Group D. (Nos 2, 3, 5, 6, 8, n, 12, 15, 19, 23, 27, 32, 40, 69.) Group C comprised 31 colonies with 6-8 astogenetic generations from 18 samples, 5 of which were also represented in Group B, and 10 in Group D. (Nos 9, 10, 22, 25, 26, 28-31, 33, 36, 37, 39, 43, 47, 54, 58, 60-62, 66, 67, 68, 70-75, 77. 78.) Group D comprised 24 colonies with 3-5 astogenetic generations from 18 samples, 10 of which were also represented in Group C, and i in Group B. (Nos 7, 24, 34, 35, 38, 41, 42, 44~46, 48-53, 55, 56, 59, 63~65, 67, 79.) FIG. 5. Ordination diagrams, prepared by the principal co-ordinates algorithm (Gower, 1966). Squared distance in proportion to (loo-similarity). See Table 4 for key to numbering ; data and co-ordinates stored at BMNH. A. Group A colonies with 15-22 zooid whorls. Co-ordinates i & 2, i & 3 and 3 & 2. Note complete separation of clusters of B. multiradiata (1,4,13,57), Atactoporidra (14, 16, 80) and Lacrimula (17, 18, 20, 21) in three dimensions. B. Group B colonies with 9-13 zooid whorls. Co-ordinates as above. Note degree of separation similar to that in Group A of clusters of B. multiradiata (2,3,5,6) Lacrimula (19,23,27) and Atactoporidra (15). B. grandis (8,11,12), L. grunaui (32,40) and Cone- scharellina africana (69) are also completely separated in three dimensions. CONESCHARELLINIFORM BRYOZOA 1 I i 4 5 B *• o O o O H o o O 342 P. L. COOK AND R. LAGAAIJ Results In Group A, separation of three-dimensional clusters of the large colonies in samples of B. multiradiata, L. burrowsi and Atactoporidra was distinct. Within Atactoporidra, both species clustered together (see Fig. 5A). In Group B, smaller colonies of B. multiradiata, A. bredaniana and Lacrimula maintained a similar degree of separation to that of Group A, except that within Lacrimula, two clusters, one of L. burrowsi and L. pyriformis, the other of L. similis were formed. Colonies of B. grandis also formed a distinct cluster (see Fig. 56). In Group C, a much larger number of still smaller colonies were compared, and the resulting clusters were far less distinct. B. grandis, B. clithridiata and B. rosula formed consistently separated clusters, but the variation within several of the other samples was apparently greater than the difference among samples of different nominal species. For example, three widely separated and variably constituted clusters of several species of Lacrimula were formed, and two distinct clusters of B. stoliczkai, one of which included B. asterizans. In Group D, this within-sample diversity was maintained in Lacrimula. B. rosula, however, formed a distinct cluster, but included L. borealis, just as the very loose cluster of B. murrayi included B. clithridiata. B. asterizans again clustered with B. stoliczkai. Some of this apparent confusion can be attributed to the general similarity of these very small colonies, the differences in total numbers and overall dimensions of zooids etc. being much less in groups C and D than in A and B. Ontogenetic differences such as presence or absence of ovicells also produced relatively large differences within samples which outweighed similarities in other characters. It is probable that if additional characters, and a larger number of colonies from each sample were available, the resulting clusters might be less diffuse. Generally, the interpretation of these three-dimensional clusters has confirmed the degrees of difference among the samples reached by inspection. Until further work can be done, using a more thorough analysis of larger samples, few systematic relationships in time can be demonstrated here. The Eocene to Oligocene samples of B. multiradiata may be confidently inferred to be genetically continuous, as may the Miocene to Pliocene samples of B. rosula. There is a very close relationship between the Oligocene B. stoliczkai and B. asterizans, and a more tenuous link may perhaps be indicated between these populations and the Eocene B. clithridiata. Relationships within Lacrimula are obviously very complex, and intermediate forms in time and space are required to establish any pattern of relationship both among the Eocene to Recent forms, and between Lacrimula and Conescharellina. EVOLUTIONARY TRENDS IN MORPHOLOGY Batopora, appropriately named after the blackberry, shows little evolutionary change with time. At most there is a tendency towards increasing regularity in the arrangement of the radial rows of zooids around the colony axis, but on the other hand good regularity is already present in an early, Middle Eocene (Lutetian), form CONESCHARELLINIFORM BRYOZOA 343 like B. scrobiculata Koschinsky. B. multiradiata would seem to be an exception, with its very irregular, double-layered arrangement, and large number (120 or more) of zooids per adult colony. This irregularity is however more apparent than real (see p. 350). In most other species of Batopora the number of zooids fluctuates from 13 to 48 (see Table i). There is not much of an evolutionary trend to be found in the development of either kenozooids or heterozooids. Kenozooids on the antapical surface are already mentioned in the original descriptions of the Late Eocene B. multiradiata and of the Oligocene B. stoliczkai. In the latter species other kenozooids occur among the zooids on the opposite, conical surface, just as they do in the Recent B. murrayi. Avicularia with a cross-bar, which occasionally occur among the zooids in the last species, are rare in the genus but also occur in one of the earliest forms, the Eocene B. clithridiata. In Lacrimula, the principal change with time appears to be the loss of axial kenozooids, which are present in the Upper Eocene L. perfecta but absent in some Miocene and all Recent forms. This trend is not a simple one, however, as axial kenozooids are present in the Holocene L. visakhensis and the Miocene L. similis, but absent in the Oligocene L. borealis. Although one sample of L. perfecta contains fairly large colonies, there does also seem to have been a general increase in maximum colony size with time. ECOLOGY AND PALAEOECOLOGY Conescharelliniform, orbituliporiform and generally lunulitiform colonies are classically associated with calm, often deeper water, and sands, i.e. coarser-grained sediments. The evidence given below includes lithology, known and inferred depth and assemblages of Recent and fossil forms. Nearly all Recent records of species with these three colony forms are associated with sea-bottoms of sand or mud. Lagaaij (ig63a) gave details of the limits of tolerance in Cupuladria, and Cook (1963, 1966) has noted similar associations. The distribution of Recent Conescharellina is almost confined to sandy and muddy sea- bottoms (see Station data given by Canu & Bassler, 1929 ; Silen, 1947 ; Harmer, 1957). The lithology of fossil colonies shows a similar range of grain size, from calcarenite to clay (see Table 5). Recent Conescharellina occurs in depths which vary from 12 m to more than 2000 m. Most records are from the range 140-450 m. Recent Lacrimula occurs from approximately 100 to 200 m, and Batopora from 285 to 805 m. Other cone- scharelliniform genera such as Fedora have been found at 2018 m (see Jullien, 1883), and Trochosodon occurs from 88 to 2081 m, the majority of records being from the deeper end of this range (see Harmer, 1957 : 744 for details). Fossil records of Batopora carry with them the connotation of occurrence at rela- tively great depths. Namias (1891 : 506), for example, reported B. rosula from 'stradi di mare profondo' and Seguenza (1880 : 130) reported B. rosula as one of the commonest, and always perfectly preserved species of Bryozoa in the Tortonian clays at Benestare, Calabria, which he considered (p. 90) 'un deposito di mare considerevolmente profondo'. Le Saint (1961 : 96) referred to 'la preference de ses 344 P. L. COOK AND R. LAGAAIJ formes [Batopora] pour les eaux relativement profondes'. Some occurrences of Batopora, however, may be inferred to be relatively shallow and neritic, two of the shallowest being (i) that of B. rosula with numerous Cupuladria haidingeri, some C. canariensis and some Biflustra texturata in the upper part of the Middle Tortonian Tefeli Formation, Almiri Section, Iraklion Province, Crete ; Sissingh (1972 : 33) referred to these beds as 'deposited in a marine environment of shallow to moderately deep water' ; (2) that of B. multiradiata in the Lower Oligocene calcarenites of Moli de Llinares, north of Villajoyosa, Alicante Province, Spain. The associated rich and diversified larger Foraminifera assemblage was listed by Cosijn (1938 : 13) and guarantees non-bathyal depth, although MacGillivray (1971 : 236) remarked, 'the formations have characteristics of gravitational deposits', i.e. may have been secondarily displaced into deeper waters. The deepest bathyal occurrences which may be inferred, on the other hand, are found in (i) the Torre Veglio Section, nor- thern Italy, in a wash residue consisting almost entirely of planktonic Foraminifera (Schuttenhelm, pers. comm.) ; (2) the INI-Coparex Huelva-i well, at 648 m below surface, in the Cadiz Basin, Spain, associated with a flood of planktonic Foraminifera and with common Liebusella soldanii. TABLE 5 Correlation of sediment type, depth and assemblage of conescharelliniform colonies Name Reference Type of Type of Depth (known Age sediment assemblage or inferred) Batopora clithridiata Gregory, 1893 B. stoliczkai Labracherie, 1961 Atactoporidra globosa B. scrobiculata A. glandiformis B. multiradiata B. rosula B. stoliczkai Lacrimula perfecta A. bredaniana B. multiradiata L. borealis B. aster izans B. stoliczkai Batopora sp. Labracherie, 1961 Koschinsky, 1885 Cheetham, 1966 Reuss, 1867 Malecki, 1963 Malecki, 1963 Accordi, 1947 Dartevelle, 1933 K 48-51, Spain 43/3 -1. North Sea Middle Oligocene Boom Clay Middle Oligocene Early Eocene C 0 s Early to Middle Eocene PAS (M) s Early to Middle Eocene PAS O s Middle Eocene O s Middle Eocene C O s Late Eocene M 0 s Late Eocene CS O s Late Eocene CS 0 s Late Eocene 0 s Late Eocene to Early Oligocene PAS 0 s Early Oligocene CR 0 Reuss, 1867 Cheetham & Hakansson, 1972 Late Oligocene Late Oligocene 0 s (M) s O s O s O s 55-90 m 0 s O s 0 s O s 0 s 0 Shallow to moderately deep (M) s M 150-250 m (Boekscho ten, 1967) S O 150-200 m CONESCHARELLINIFORM BRYOZOA 345 Name B. rosula B. rosula B. multiradiata B. rosula L. asymmetrica L. similis B. rosula B. rosula B. rosula B. rosula B. rosula L. visakhensis B. murrayi L. burrowsi L. pyriformis Conescharellina africana C. angustata Trochosodon radiatus Reference R 1103, Spain SM 55, Italy Ceretti & Poluzzi, 1970 MU 214, Sardinia BG3i2 BG3i2 PLR 4067, Malta Reuss, 1848 TABLE 5 (cont.) Age Early Miocene Early Miocene Early Miocene Early Miocene Early Miocene Early Miocene Middle Miocene Middle to Late Miocene Seguenza, 1880 Middle to Late Miocene 6-27, Crete Late Miocene Huelva, Spain Pliocene Rao & Rao, 1973 Holocene Cook, 1966 Recent Cook, 1966 Recent Cook, 1966 Recent Cook, 1966 Harmer, 1957 Harmer, 1957 Recent Recent Recent Type of Type of Depth (known sediment assemblage or inferred) C (M) M M Bathyal CR O M M S s C M Bathyal C (M) : S (Haug, 1920 : 1637) M M shallow to moderately deep FS M Bathyal CS O:S 89 m C M 805 m CS O:S 101—207 m CS 0:S 310 m Md Md Md O : S 102 m O : S 88 m M 1944 n Sediment : CR = Calcarenite ; C = Clay ; Md = Mud ; M = Marl ; FS = Fine-grained sand ; AS = Argillaceous sand ; CS = Coarse-grained sand. A ssemblage : M = Monospecific ; (M) = Almost monospecific ; O = Multispecific assemblage ; S = Other specialized colony forms present in assemblage, i.e. orbituliporiform or lunulitiform colonies. Generally, both fossil and Recent bryozoan assemblages associated with soft and unstable sea-bottoms show first an increasing number of specially adapted forms with increase in depth. This is followed by a decrease in all forms ending in mono- specific occurrences at very great depths. Harmer (1957 : 649-650) analysed the species obtained by the 'Siboga' from a few stations in the Java Sea where the sea-bottom was mud at 82-88 m ; he remarked : There is no group of Stations ... in which the correlation between mode of growth and nature of the bottom is more clearly established.' Of the 39 species found, many were new, and at least 15 showed some recognizable adaptation to the specialized conditions. Eight had some form of rootlet system and six were 346 P. L. COOK AND R. LAGAAIJ lunulitiform species. As Conescharellina and Flabellopora are known to have rooting systems which can be associated with foraminiferal ooze, it is interesting to note the close correlation of their distributions. These two genera, often represented by several species, occur together at 20 of 36 stations listed by Canu & Bassler (1929), Silen (1947) and Harmer (1957). In addition, Lanceopora occurred with the above genera at eight of these stations. Recent and Holocene records of Lacrimula are correlated with Cupuladria and Conescharellina. Conescharellina and Trochosodon have been found in association with as many as seven other similarly adapted species at the shallower end of their range. At great depths, both these genera tend to have monospecific occurrences, and, as noted above, one record of Recent Batopora is from very deep water. Evidence from fossil assemblages gives a similar picture. Maplestone (1904 : 207, 209) gave tabulations which noted the correlation of fossil Conescharellina and Lanceopora (as Schizoporetta flabellata, see also Maplestone, 1910). Among the specially adapted forms such as Kionidella, Stichoporina and Lunulites, Orbitulipora is one of the commonest genera found in association with specimens of Atactoporidra and Batopora (see, for example, Ossat & Neviani, 1897 ; Gregory, 1893 ; Malecki, 1963 ; Labracherie, 1961). Batopora multiradiata is generally associated with diversified bryozoan faunas, but the three very deep records of Batopora mentioned above, two fossil and one Recent, are conspicuously monospecific, and it is only one step further to suggest that all monospecific occurrences come from the deep end of its bathymetric range. There is nothing in the data at hand (Table 5) to contradict this suggestion. The Lower and lower Middle Miocene of the Mediterranean area seem to be particularly marked by such monospecific deep-water occurrences, and in keeping with the inferred depth, the associated lithologies tend towards the finer grain-sizes ; either marl or calcareous clay (Table 5). DISTRIBUTION IN TIME AND SPACE Historically, the oldest published record of a conescharelliniform colony is that of Batopora, in Soldani's figure of 'Historices marinas minimas' (Soldani, 1780, pi. 16, figs 83(3, R ; figures copied by Annoscia, 1968, pi. i, figs 8a, b). According to the original plate explanation (p. 130) these fossils originate from 'the valley East of San Quirici'. Dr Annoscia kindly informs us that 'The location of the samples containing Batopora, according to Soldani's words, is "not far from Caitro [nowa- days unknown] S. Quirico, in a place above the Fosso di Rifigliuoli [now 'Fosso Refiglioli' see map] named Poggi di Rifigliuoli (Soldani, 1780, pp. 130 e 35, art. Ill), i mile from Castello [now unknown or destroyed] beyond the Fosso di Rifigliuoli toward S. Quirico" [some 35 km S.E. of Siena]. I think it might be the circled place on the enclosed map, named today "Podere Favorite", or not far from it.' 'The formation "Upper marine clay and lower ocreaceous shale" by Soldani might belong to the formation "P^"1" in the new edition (1968) (sheet 121) of the official geological map printed by the Italian Geological Survey. This formation is made by clay, sandy clay, also in continental facies ("Piano del Sentino" Fm.) associated CONESCHARELLINIFORM BRYOZOA 347 with small lenses or layers of puddingstone, locally with Lower -Middle Pliocene microfauna.' Time-stratigraphically, Batopora clithridiata (Gregory) from the London Clay (Ypresian) of the London Basin and the forms described by Labracherie (1961, pi. 16, figs 2 and 4) as Batopora stoliczkai from the subsurface Lower Eocene of the Bordeaux area, France, vie for the distinction of being the oldest Batopora on record. From then onwards the genus occurs in all younger Tertiary stages in Europe (Table 5) with the exception of the Uppermost Miocene. Data are too few, however, to assess whether this absence is fortuitous or indeed reflects the Late Miocene (Messinian) 'Crise de salinite' in the Mediterranean (Hsu, Ryan & Cita, 1972). For more than a century Batopora has been considered an extinct genus, until recently one of us found a living representative in the western Indian Ocean, near Zanzibar (Cook, 1966). It appears that Batopora had disappeared from western Europe after the Oligocene, and that most Miocene and Pliocene records are from the Mediter- ranean and Paratethys (Map i, p. 369). In the Pliocene the genus had even ventured out into the western Atlantic. Its present occurrences near Zanzibar and Fiji are curiously remote from the palaeodistribution even in the not-so-distant past, although naturally this may only reflect our ignorance of deeper-water Tertiary faunas from the Indian and the Pacific Oceans. In contrast to Batopora, the genus Lacrimula was first described by one of us (P. L. C.) from the Recent, east African coast, and was then found by the other (R. L.) in fossil assemblages from western Europe and the East Indies. Specimens from the Holocene of the northern Indian Ocean have also since been described by Rao & Rao (1973) (see Map 2, p. 370). Atactoporidra apparently never had a wide distribution, and there are no records after the Oligocene. The three species recorded were from the English and French Eocene, and from the Belgian Eocene and Oligocene. Conescharellina may have been present in the Eocene of western Europe, although records of species may prove to be referable to Lacrimula (see p. 359). The genus seems to have had an almost constant presence in the Australasian region from the late Oligocene to the Recent. (Dr R. Wass, in an unpublished report on stratigraphic ranges of some Bryozoan species from the Tertiary of south-eastern Australia (unpublished report in the Geological Sciences, University of Sydney, 1973 - derived in part from Cockbain, 1971) gives the age of the Mount Gambier fauna as late Oligocene -early Miocene.) Conescharellina was common during the Australian Miocene, and there are records from the Pleistocene of the East Indies and Japan. Kataoka (1957) found no Conescharellina in a cold-water, Pliocene fauna from Northern Honshu, Japan (approximately 40°N, I40°E). However, by Pleistocene times, probably during an interglacial period, the genus was present further south, at Kikai Jima (approximately 28°N, i3O°E, see Kataoka, 1961 : 259). The accompanying large bryozoan fauna included many tropical and subtropical species, including the orbituliporiform Flabellopora, and the lunulitiform Cupuladria and Actisecos. Recent records extend even further north, up to 41 °N, I40°E (see Canu & Bassler, 1929^ and west to East Africa (see Map 3 and Appendix 3, pp. 371-373)- Generally, the distributions given here may be interpreted as wide, warm water 348 P. L. COOK AND R. LAGAAIJ occurrences during the Eocene and Oligocene, which have either shifted or become progressively restricted in a southerly and easterly direction up to the Recent. CONCLUSIONS It has become apparent during this study that the classical concepts both of structure and taxonomy of conescharelliniform colonies require much further work. Analysis of many more samples from additional areas and horizons may begin to throw light upon their evolutionary systematics. At present, we may reach the following conclusions and indicate some of the directions of future research which may prove fruitful. The overall effects of genetic constitution and environmental selection outweigh microenvironmental effects in colony structure. The conescharelliniform colony has a rigid astogenetic pattern allowing for little variation in the primary zone of change, but allows more in secondary zones, especially in the apical region. Here environmental influences may be the reason for the latitude in astogenetic and ontogenetic expression both within and among populations. It is possible that convergent lines of evolution with parallel development have occurred, and this possibility requires further investigation. The obvious physical separation and finiteness of colonies make them very susceptible to statistical analysis of samples. Comparative astogenetic age of colonies within samples is essential. The within-population variation of species requires analysis in the hope of following environmental changes in population characteristics. Integration within colonies is considerable. A low degree of integration is expressed in colonies where zooids have little interdependence and function in a manner similar to that of solitary animals. Boardman & Cheetham (1973 : 132-134) have suggested several characters with sequences of states showing increasing integration among bryozoan colonies. Using these sequences the wall structure, interzooidal communication, astogenetic zonation and polymorphism of conescharel- liniform colonies illustrate a high degree of integration. The devolution of roles among zooids and kenozooids at the earliest astogenetic stages, and the continued development of patterned polymorphs and extrazooidal tissues, are particular examples of colony-wide control. The specialized mode of life allows palaeoecological inferences to be made as to the depth and nature of the sea-bottom through a considerable range of time. Future studies on larger samples of more populations should include further examination of type or topotype material of described species. A systematic search of fine-grained sediments, both fossil and Recent, should be made, to enlarge knowledge of distribution and abundance in time and space. Eventually, this should include parallel studies on other orbituliporiform and conescharelliniform species both with 'normal' and 'frontal' astogeny. A study of similar colony forms such as that exhibited by Sphaeropora, which has a Tertiary to Recent range, may throw some light on the evolution of the colony form (see also Waters, 1919 : 80). CONESCHARELLINIFORM BRYOZOA 349 The possibility of a study of living colonies may be remote but should not be neglected. The breeding and above all the settlement and early astogeny of a single species would provide guidelines applicable with confidence to other forms. DESCRIPTIONS OF SPECIES Family ORBITULIPORIDAE Orbituliporidae Canu & Bassler, 1923 : 186. Ascophora with discoidal and bilaminar, or conical colonies, which may be multi- laminar. Zooidal calcified frontal wall extensive, cryptocystidean, consisting of two elements, one of which contributes to the exposed surface of the colony and which surrounds a centrally placed orifice. Orifice oval, frequently with paired condyles, occasionally with a distinct antapical sinus. Ovicells usually large, hyperstomial, directed adapically, occasionally immersed. There is constantly a tube, comprising kenozooids and/or extrazooidal tissue, which in some cases extends beyond the colony surface, and which is in contact with the ancestrula or ancestrular complex internally. It is inferred that the tube marks the origin of rootlets which anchored the colony to its substratum. The tube is radially placed in bilaminar (orbituli- poriform) colonies, and axially placed in conical (conescharelliniform) colonies. Avicularia and kenozooids often present. Genus BATOPORA Batopora Reuss, 1867 : 233. Type species B. stoliczkai Reuss. Reuss formally introduced Batopora as a new genus, when describing B. stoliczkai. Other species were referred to the genus in the discussion, but were not formally described. They included Cellepora rosula, which Reuss had described in 1848, and Conescharellina angustata d'Orbigny. Waters (1919 : 93) formally listed B. stoliczkai as type species of Batopora. Canu & Bassler (1917 : 75), however, had already listed 'Batopora rosula Reuss 1847' as the type species of the genus, which they quoted erroneously as 'Batopora, Reuss 1847'. This was emended in later works, and Canu & Bassler (1920 : 629) and Bassler (1935 : 54 ; 1953 : 0230) gave the type species as B. stoliczkai. Some previous attributions of species to Batopora need examination of specimens before they can be assigned to the genus with any confidence. Among these B. aviculata Hejjas (1894 : 214, 251, pi. 6, fig. n) is problematical, and Batoporella eocaenica Hejjas (1894 : 215, 252, pi. 7, fig. 13) seems to be related to Orbitulipora. Both forms were reported with records of 'B. conica Hantken' (see p. 359), B. multi- radiata, B. rosula and B. scrobiculata from the Eocene of Hungary. Colony conescharelliniform, with an external or internal adapical tube. Zooids with antapical edge of orifice flattened, peristomes frequently raised, tubular and prominent, exposed frontal walls of zooids often inflated, especially in the proliferal 23 350 P. L. COOK AND R. LAGAAIJ region. Avicularia sometimes present, mandibles hinged on a bar. Ovicells large, hyperstomial but often immersed and obscured by the peristome. Secondary zones of change formed by frontal budding. Secondary series of zooids forming over- growths in some species. The species previously described in this genus fall into two groups ; those in which there are a large number of zooids (maximum seen 200) in a colony, and in which frontal budding of secondary overgrowth is found, and those with a smaller number of zooids (rarely more than 50), in which secondary zooids are irregular in occurrence and in which overgrowth does not occur. The apical region may be marked by an internal kenozooidal or extrazooidal tube (the 'pit' of authors) or by an external tube. In some colonies the tube appears to be replaced by a secondarily budded kenozooid. The first group is represented by B. multiradiata and B. grandis. In B. multi- radiata the colonies often reach a considerable size (3 mm in diameter), and are marked by the development of a distinct type of secondary zone of astogenetic change. Secondary zooids arise adapically, and apparently form an overgrowth of zooids advancing over the primary zone in a regular manner. Each zooid is budded frontally from the concealed part of the frontal wall or walls of zooids of the new prolif eral region (see p. 326) . This secondary zone is a modified form of overgrowth and thus differs radically both from 'normal astogeny' overgrowth and from frontal budding (see Banta, 1972). It also differs from the type of secondary zones of change found in Atactoporidra and Lacrimula (see p. 326). The secondary zone of change does not always develop regularly, especially during its later astogeny. Few populations of B. multiradiata are well preserved, and wear increases their irregular appearance. We have been able to examine two well- preserved populations, which show the astogeny both of the primary and secondary zones of change particularly clearly. Specimens from the Val di Lonte and Montecchio Maggiore (Eocene, Bartonian, Italy, USNM 71205, 7 colonies and 71196, 4 colonies) average 1-25 mm in axial length and 2-75 mm in prolif eral region width. Nearly all these colonies possess a distinct raised apical tube apparently composed of extrazooidal tissue, and sur- rounded by a circlet of kenozooids. The secondary astogenetic zone of zooids can be seen to arise in two colonies as a circlet of eight very small zooids just below the tube. The primary zone consisted of regularly alternating whorls of six zooids. The number in the prolif eral zone of secondary zooids varies from 30 to 40 (see PI. 3, fig. i; PI. 4, figs 5-6). Ovicells are present throughout the primary zone and in the last 1-4 whorls of the secondary zone. They are large, very prominent, and have a regularly pitted surface, which may indicate the presence of pseudopores in life. Orifices of brooding zooids are wide, with a distinct antapical peristome. Other colonies from Gotzreuth (USNM 88881, Eocene, 65 colonies) may not all belong to B. multiradiata but several show clearly that the secondary zone consists of zooids budded frontally in alternat- ing series as an overgrowth of the primary zone (see PI. 4, fig. 4). Here, too, ovicells are present in both primary and secondary zones. These specimens include some very young colonies (see Table i). CONESCHARELLINIFORM BRYOZOA 351 B. grandis also possesses large colonies with numerous zooids. It may produce intercalary rows of zooids, but does not develop a distinct secondary zone of over- growth. The second group includes all other known species of Batopora. The colonies are never very large, and secondary zones of change are limited to isolated zooids budded frontally from exposed frontal walls of primary zone zooids. This group may be subdivided into species with globular colonies with small zooids (B. stoliczkai, B. clithridiata and B. asterizans) and those with a more conical shape and larger zooids (B. scrobiculata, B. rosula and B. murrayi). Specimens of B. stoliczkai from the Lower Oligocene of Saale (see p. 352), have small colonies with very small zooids and frequently occurring interzooidal kenozooids. Avicularia are absent, but the ovicells are very well preserved and quite prominent. Isolated secondary zooids are fairly frequent in the larger, astogenetically older colonies (see PL i, fig. i ; PL 2, fig. 2 ; PL 3, fig. 4). B. clithridiata was described as Conescharellina clithridiata by Gregory (1893 : 252, pi. 31, figs 10 and n), from specimens from the London Clay at Sydenham (South London) and Hampstead (North London). The figured colony was separated from a large number from Sydenham (BM 1357) which remained in the Collection under a manuscript name. Nearly all the specimens possess some interzooidal avicularia with a complete bar, many also have small interzooidal kenozooids and isolated secondary zooids (see PL 2, fig. i ; PL 5, fig. 5). B. clithridiata has slightly larger zooids than B. stoliczkai, in other characters they are very similar. B. rosula was first described by Reuss (1848 : 78, pi. 9, fig. 17) as Cellepora rosula, from the Miocene of Baden, Austria. These specimens have not been examined, but those he described later (1867 : 225, pi. i, figs 7a-c ; pi. 2, figs la-c) have been seen. Reuss apparently illustrated two distinct astogenetic stages of B. rosula. Both possess a small, external apical tube, apparently composed of kenozooids, and in both the zooid series were figured as budding directly. The two figured specimens differ in their alternate budding pattern from Reuss's drawings. The remaining three specimens in Reuss's material show a fairly high range of variation. Two of them are irregular in development, and one has a secondarily budded conical keno- zooid apically instead of the complex kenozooidal tube of the other colonies. The calcification of the zooids varies from medium to coarsely granular. Generally all five colonies are well preserved and most have ovicells. These are not prominent, and the frontal surface has a distinctly punctate appearance. Axial kenozooids are present in some colonies, and the apical region shows either a small aperture (presumably a pit) surrounded by slightly raised kenozooids or a narrow, raised tube with rows of radially arranged pores on its surface. This tube greatly resembles a zooid in size and shape and seems to be a product of later astogenetic changes in the colony. One colony is worn adapically, and shows traces of the walls of the ances- trular region in section. The primary zooid tetrad surrounds a circular area which presumably marks the position of the apical kenozooid or kenozooidal tube (cf. B. stoliczkai, p. 352, and see PL 3, figs 2 and 3). The other Miocene specimens from the Mediterranean area assigned here to B. rosula are well within the range of variation shown by Reuss's specimens. The 23* 352 P. L. COOK AND R. LAGAAIJ colonies from Malta and Crete (see PI. 4, figs i and 2) are slightly smaller, and have a secondary kenozooid apically. One specimen from Spain (Roep 1103), which has ovicells, is almost exactly like the colony figured by Reuss (1867, pi. I, fig. 7). B. scrobiculata was described by Koschinsky (1885 : 63, pi. 6, figs 2a-c, 3a-c) from the Eocene of Gotzreuth. A specimen from this locality (USNM 88882 pt), labelled B. scrobiculata, has a flatter colony shape than B. rosula, the zooids are very large, comparable with those of B. grandis and B. murrayi, and easily distinguishable from those of B. rosula at a similar astogenetic age. The calcification of the zooids is coarsely granular and axial kenozooids are present (see PL 3, figs 5, 6 and 7). The Pliocene Batopora from Huelva, Spain, has somewhat similar characters. It has a flattened colony, and the calcification of the large zooids is coarsely granular. In view of its age and locality, it should, however, perhaps be assigned to B. rosula. Batopora stoliczkai Reuss (PI. i, fig. i ; PI. 2, fig. 2 ; PI. 3, fig. 4) Batopora stoliczkai Reuss, 1867 : 223, pi. 2, figs 2-4. MATERIAL EXAMINED. Lower Oligocene, Bunde, Germany, 27 colonies, NMV, J.867.XII, I3a-d. Lower Oligocene, Calbe, Saale, Germany, 5 colonies, USNM. DESCRIPTION. Colony small, globular. Maximum number of whorls 7. Maxi- mum number of primary zooids in proliferal whorl 3. Maximum number of primary buds 2. AxL 0-63-0-90 mm Prl 0-87-1-06 mm Lfw 0-21-0-23 mm Ifw 0-23-0-25 mm Lo 0-09-0-10 mm lo 0-08-0-09 mm Lov 0-16-0-17 mm lov 0-18-0-23 mm REMARKS. The five colonies from Calbe have almost the same characteristics as those from Bunde, except that interzooidal kenozooids are more frequently and regularly developed. The kenozooids do not appear to have any uncalcified central portion, and the small zooid-like structures figured by Reuss (1867, pi. 2, figs 2a, b) as occurring between zooids are thus not exactly as depicted. Among the small kenozooids, the partially closed, half-submerged orifices of primary zone zooids are often visible, below the surface of the secondary zone interzooidal zooids. The antapical axial kenozooids are exactly as figured by Reuss (pi. 2, fig. 3). Of the 32 colonies examined, seven of the specimens from Bunde differ in several ways from the majority. Three colonies are worn, regularly oval in shape and larger than all the others. They appear to have been composed of alternating series, and more than one zone of zooids. The largest colony measures 1-40 mm in axial length and 0-90 mm in proliferal width and comprises approximately 49 zooids. It is very probable that these three colonies are late astogenetic stages of B. stoliczkai. Two of the remaining colonies are probably attributable to Batopora, but differ from B. stoliczkai. They consist of 22 and 30 zooids respectively, arranged in rather irregular whorls of 5-6 zooids each. The colonies are flatter than those of B. CONESCHARELLINIFORM BRYOZOA 353 stoliczkai (AxL, 0-52-0-64 ; Prl, 1-12-1-30) and have a wide, internal apical tube, surrounded by kenozooids. There are no antapical kenozooids. It is possible that they are representatives of yet another form of Batopora, but further specimens would be required to ascertain their specific position. The two remaining colonies resemble B. stoliczkai in zooidal characters, but appear to be at least partially bilaminar. The focus of budding is at the periphery of each colony, from which point alternating zooids are produced facing in opposite direc- tions. These form an astogenetic gradient, becoming larger and more regularly bilaminar so that the colonies form rounded wedges. It is possible that these colonies are very young astogenetic stages of Orbitulipora. Although these last four colonies differ widely from the others, it is just possible that they are variants of B. stoliczkai induced by unknown environmental factors, and much more analysis of larger populations would be needed before any further conclusions as to their relationships could be made. One colony from Bunde shows a very early stage in the astogeny, which may be compared with those described for Lacrimula asymmetrica (p. 361). The primary, almost certainly ancestrular, complex consists of a triad of two zooids and a keno- zooid. The zooids have elongated peristomes, the kenozooid a wide rounded orifice hardly raised above the remainder of its exposed frontal wall. Two further whorls are present, each consisting of alternating triads of zooids (see PI. i, fig. i). The antapical surface at this stage is flat, and consists of the concealed frontal walls of the last triad only, no kenozooids are present. The next stage present (from Calbe) has 15 zooids and only four whorls of zooids. The four zooids in excess of the estimated number (see p. 333) are secondarily budded zooids arising between those of the first and second whorls. In all larger colonies, secondary buds are more regu- larly triadic than those in B. clithridiata, and tend to be produced in sequence, about one astogenetic generation behind the zooids of the primary zone. The presence of a narrower proliferal region produces the globular appearance of the colonies, which also have a more regular aspect than those of B. clithridiata after the 12-14 zooid colony stage, as the small kenozooids are budded between the zooids. The similarity among young stages of B. stoliczkai, B. murrayi, and Trochosodon sp. is illustrated on Plate i, figs 1-6. Batopora grandis1 sp. nov. (PI. 2, figs 5, 6 ; PI. 3, fig. 8) HOLOTYPE. Eocene, Gotzreuth, Bavaria, BM 031117. OTHER MATERIAL EXAMINED. Lutecian, Eocene, Gotzreuth, Bavaria, with B. scrobiculata, B. multiradiata and K, excelsa, 4 colonies, USNM 88882 ; paratypes. DESCRIPTION. Colony large, elongated, conical. Maximum number of whorls 12. Maximum number of zooids in proliferal whorl 4. Maximum number of primary buds unknown, probably 4. 1 grandis — (L) - large - referring to the large size of the zooids. 354 P- L- COOK AND R. LAGAAIJ AxL 1-88-2-72 mm Prl 1-70-2-00 mm Lfw 0-40— 0-45 mm Ifw 0-35-0-60 mm Lo 0-14-0-20 mm lo 0-10-0-12 mm Lov 0-32-0-50 mm lov 0-35-0-55 mm Apical region an internal tube surrounded by small kenozooids. Zooids very large, arranged in directly budded series with secondary zooid series apparently budded regularly between them, forming a spiral pattern. Exposed frontal walls of zooids not inflated, finely granular. Approximately 12 marginal pores in each exposed frontal wall and 8 in each concealed frontal wall. Orifices not raised, apparently with an antapical sinus. Avicularia absent. Ovicells very large, asymmetrically displaced laterally. REMARKS. The colonies are elongated and very large, and form an astogenetic series. The zooids are very wide and have a spiral arrangement. It is not known whether the orifices are primary or secondary, and most are worn, but some show a distinct and elongated antapical sinus. The apical region consists of a very narrow tube surrounded by 2-3 series of small kenozooids or secondary closed zooids. The number of zooids is very high, especially as there are only four (occasionally an asymmetrically-arranged fifth zooid is present) in the proliferal region. This strongly suggests that the intercalary series of zooids found on the colony surface are budded as in Lacrimula burrow si, which also has a very regular appearance. There is no observable second layer of zooids in the subproliferal region of either of these species, in contrast to L. similis and B. multiradiata, where the limits of the secondary zone are strikingly obvious. The ovicells are broken in all specimens and occur on proliferal region zooids only. They are very large, with thick, apparently two-layered walls. Each ovicell is asymmetrical and they resemble those described by Harmer (1957 : 733) in Cone- scharellina catella. The characters of B. grandis are distinct from all other species, but it has features in common with Lacrimula, Atactoporidra and Conescharellina. In size and shape and the possession of an apical tube it resembles L. burrowsi and Atactoporidra bredaniana. The apparently sinuate orifice and asymmetrical ovicells are super- ficially, at least, similar to those found in some Recent forms of Conescharellina. B. grandis does not appear to have been described before. It is part of a very interesting fauna of conescharelliniform and similar colonies, all from the Gotzreuth locality, which includes Kionidella, B. multiradiata and B. scrobiculata. Batopora asterizans1 sp. nov. (PL 2, figs 3 and 4) HOLOTYPE. Middle Oligocene, 45°95'3'N, i°3i-6'E, North Sea 43/3-1 well, 960-990 ft, BM 052567. OTHER MATERIAL EXAMINED. As above, 840-870 ft, 3 colonies ; 870-900 ft, 5 colonies ; 1110-1140 ft, i colony ; 1530-1560 ft, I colony ; paratypes. Rupelian, 1 aster - (L) - a star - referring to the stellate appearance of the apical and axial kenozooids. CONESCHARELLINIFORM BRYOZOA 355 probably Middle Oligocene, Ijzendijke, Zeeland, boring K62, Boom Clay, 5 colonies ; paratypes. DESCRIPTION. Colony small, conical to globular. Maximum number of whorls 6. Maximum number of zooids in proliferal whorl 3. Maximum number of primary buds 3. AxL 0-70-0-75 mm Prl 0-90-1-00 mm Lfw 0-20-0-25 mm lfw 0-25 mm Lo 0-06-0-09 mm lo 0-07-0-09 mm Apical region apparently consisting of a secondarily budded kenozooid. Zooids arranged in alternating series, with small globular kenozooids arising interzooidally and usually alternating with the zooids. Circles of apical kenozooids and a group of axial, antapical kenozooids present. Frontal walls of zooids distinctly, almost coarsely granular. Exposed frontal walls inflated, with six marginal pores. Primary orifices not seen, secondary orifices oval. Ovicells and avicularia not seen. The apical region is marked by an inner circle of five, and an outer circle of eight, alternating, globular kenozooids (diameter 0-14 mm). These presumably have over- grown the primary zooids, and also occur, budded interzooidally, fairly regularly in alternating series among the zooids. A further, axial group of five kenozooids is also present. REMARKS. B. asterizans has a similar growth form to that of B. stoliczkai and B. clithridiata. The arrangement of the zooids and alternating kenozooids is, however, much more regular than in either of these species. It further differs from B. clithridiata in the absence of avicularia. The colonies are all very small and none have ovicells. It is therefore possible that larger colonies may eventually be found, although ovicells occur in both B. stoliczkai and Trochosodon (see PI. i, figs 5 and 6 ; PI. 2, fig. 2) in colonies of comparable size to those of B. asterizans. Generally the colonies are regularly constructed, although one from the North Sea shows several frontally budded secondary interzooidal zooids which results in an appearance very similar to that of B. clithridiata. The apical tube is raised and appears to consist of a secondarily budded kenozooid. It is regularly surrounded by small, rounded kenozooids, which are inferred to be secondarily budded, as one very small colony from the North Sea shows only three kenozooids, whereas larger examples show two series of 5-8 kenozooids. Genus LACRIMULA Lacrimula Cook, 1966 : 217. Type species L, burrowsi Cook. Colony conescharelliniform, with an external tube. Zooids with rounded orifices, and often well-developed paired condyles, sinus occasionally present. Interzooidal avicularia sometimes present. Ovicells large, hyperstomial, prominent. Secondary zone of frontal buds arising directly from exposed frontal walls of primary zone zooids, beginning adapically. 356 P. L. COOK AND R. LAGAAIJ Specimens ascribed here to Lacrimula vary from small elongated colonies which have some morphological affinity to the Batopora rosula group (L. borealis) to complex colonies with well-developed zones of secondary zooids which approach, superficially at least, those of Atactoporidra bredaniana and Batopora multiradiata. Some species also show an interesting similarity in character with Conescharellina. Both L. visakhensis and L. perfecta possess regular axial series of kenozooids, and L. visakhensis resembles C. africana in producing adapical ovicells. Generally Lacrimula includes species in which the colony is elongated and the apical region consists of a prominent kenozooidal tube. Lacrimula burrowsi Cook (PI. 5, figs i, 6 ; PI. 6, fig. 3 ; PI. 8, figs 1-6) Lacrimula burrowsi Cook, 1966 : 218, pi. 2, figs 2-4 ; fig. 4A. MATERIAL EXAMINED. Recent, Zanzibar, 101 m, 207 m, 37 colonies, BM John Murray Coll. 1965.8.24.7-10, 1965.8.24.11 ; paratypes. Off Umvoti River, S. Africa, 102 m, 34 colonies, BM Burrows Coll., 1949.11.10.642. DESCRIPTION. Colony elongated, often slightly flattened in one direction. Maximum number of whorls 19. Maximum number of zooids in proliferal whorl 6. Maximum number of primary buds 6. AxL 2-00-3-20 mm Prl 1-24-1-88 mm. Lfw 0-30-0-37 mm Ifw 0-30-0-45 mm Lo 0-10-0-15 mm 1° o-io-o-n mm Lov 0-14-0-30 mm lov 0-15-0-17 mm Apical region a tube with closely spaced external pores. Zooids primarily budded in alternating series, with avicularia frequently, but often irregularly interspersed. Frontal wall of zooids finely granular. Exposed part of frontal wall with 4-6 marginal pores, 4-6 pores in concealed part. Primary orifice divided at the mid-line by large condyles. Ovicells large, occurring in the proliferal and subproliferal zone zooids. Fertile orifices not dimorphic, closed by the operculum. Avicularia arising as frontal buds between adjacent primary zooids. Chamber large, but not reaching the axial region of the colony. Exposed part of frontal wall of chamber with marginal pores. Semicircular mandible hinged to large, paired condyles. Rostra directed adapically and slightly laterally, sometimes alternating in radial series. The first whorl of individuals below the apical region consists of avicularia ; these are secondary in origin. Secondary calcification affects zooids progressively from the apical region in an antapical direction. In some colonies the first seven whorls are comprised of zooids with closed orifices, which may have a small central rounded opening. REMARKS. A large number of colonies of Lacrimula burrowsi have recently been found in bottom sediment samples kept by the Mineralogy Department of the British Museum (Natural History). The samples were all from the Zanzibar area and were collected by the John Murray Expedition to the Indian Ocean. The colonies were CONESCHARELLINIFORM BRYOZOA 357 dried and were therefore restored in trisodium phosphate solution and stained before being mounted in epoxy-resin and sectioned. These sections have shown details which generally support some of the inferences about colony structure suggested here. The marginal pores appear in longitudinal section as small tubules, often filled with stained tissue, passing through the thick calcification of the concealed frontal wall of one zooid into the living chamber of the next successive zooid. Tubules tend to be concentrated towards the axial end of zooids (see PI. 8, figs i and 2). They are very similar in appearance to the extended 'areolar' tubules, which are derived from frontal septulae, in other Bryozoa. The apical region shows the ancestrular whorl of zooids to be surrounded by very thick calcification. A series of large tubular pores, each surrounded by calcification, forms the apical tube (see PI. 8, fig. 5). The pores are inferred to be the coelomic cavities of kenozooids ; they are often filled with stained tissue and pass from the exterior of the apical tube into a large interior cavity. This cavity is lined with stained tissue and it is inferred from analogy with the marginal pore tubules that neither the axial kenozooid nor the smaller kenozooids forming the tube had any connection directly with the exterior environment in life, but may have been the site of rootlets. Although elongated tubules extend from the primary zooids to the secondarily budded avicularian chambers at the base of the kenozooidal tube, there does not appear to be any communication among the small kenozooids. If the axial cavity marks the position of the coelomic cavity of a kenozooid, the smaller cavities would have been able to intercommunicate. Generally, all these structures need much more work, in this and other species, particularly those which can be demon- strated to have rootlets. The apical structures in L. burrow si, however, appear to be consistent with the postulate that they were the site of rootlets arising from highly modified kenozooids which were budded successively in an adapical direction from a large axial kenozooid. Lacrimula visakhensis Rao & Rao (PL 6, fig. 4) Lacrimula visakhensis Rao & Rao, 1973 : 506, fig. i. MATERIAL EXAMINED. Holocene, Bay of Bengal, E. India, 89 m, 4 colonies, BM Subba Rao Coll. 1970.8.8.1 A-D. DESCRIPTION. Colony small, conical. Maximum number of whorls 7. Maxi- mum number of zooids in proliferal whorl 8. Maximum number of primary buds 8. AxL 1-12-2-20 mm Prl 1-60-2-00 mm Lfw 0-23-0-25 mm Ifw 0-25-0-40 mm Lo 0-12-0-18 mm lo 0-10-0-16 mm Lov 0-25-0-32 mm lov 0-30-0-40 mm Apical region a tube with large external pores. Primary zooids budded in direct series. Secondarily budded series of zooids confined to the adapical end of the zoarium. Axial region composed of small kenozooids. Frontal wall of zooids very 358 P. L. COOK AND R. LAGAAIJ finely granular. Marginal pores 2-6 in exposed part of frontal wall, 8-n in concealed part. Primary orifice oval, divided adapically to the mid-line by small, paired condyles. Both adapical and antapical parts curved ; peristome absent. Ovicells very large and wide, orifice wider than long. Brooding zooids budded secondarily from zooids at the adapical end of the colony. Frontal surface of ovicell granular, marginal pores present. Avicularia arising as frontal buds between adjacent zooids in the adapical region ; mandibles inferred to have been rounded. Avicularia are not common, occurring laterally beside a zooidal orifice in a few zooids only ; they have no bar or condyles and are inferred to be avicularia only by their size and position. The first whorl of zooids in the young colony is hidden by a whorl of avicularia which are budded between the primary zooids. REMARKS. The condyles of the autozooidal orifices are placed distinctly adapic- ally, those of brooding zooids are not visible. The ovicells occur in a single whorl at the adapical end of the colony, and all arise as secondary zooid buds between primary zooids. Antapically the concealed frontal walls of the youngest proliferal region zooids show that the marginal pores do not extend axially to the limit of each zooid. They form a row half-way across the wall. The axial region is filled with 6-8 very small kenozooids. L. visakhensis differs from other species of Lacrimula in the position of the con- dyles, and in the astogeny, position and frontal wall characteristics of the ovicells. It resembles L. perfecta in possessing axial kenozooids, and Coneschardlina africana in the apical position of the ovicells. Lacrimula perfecta (Accordi) (Fie. 7A ; PL 4, fig. 3 ; PI. 7, fig. i) Conescharellina perfecta Accordi, 1947 : 105, figs 1-7 ; Braga & Munari, 1972. MATERIAL EXAMINED. Priabonian, Upper Eocene, Cunial Quarry, Possagno, N. Italy, 16 colonies and several fragments, collected by Drs E. Annoscia and P. Ascoli, 1968. Priabonian, Upper Eocene, Forte di San Leonardo, Verona, N. Italy, 31 colonies, collected by Dr G. Braga. DESCRIPTION. Colony conical to pyriform. Maximum number of whorls 20. Maximum number of zooids in proliferal whorl 9. Maximum number of primary buds 6. AxL 0-75-3-40 mm Prl 0-86-2-80 mm Lfw 0-19-0-40 mm Ifw 0-20-0-45 mm Lo 0-09-0-12 mm lo 0-09-0-14 mm Lov 0-17-0-20 mm lov 0-25-0-30 mm Colony with a distinct kenozooidal apical tube with external and internal un- calcified pores. Tube becoming very large, bulbous and thick by accretion of kenozooids during astogeny (i-oo mm in diameter), with a very small central CONESCHARELLINIFORM BRYOZOA 359 aperture at the apex (o-io mm in diameter). Axial kenozooids also budded regularly in series forming a central core (1-60 mm in diameter). Zooids with a non-sinuate orifice, wider antapically, with paired lateral condyles. Zooids budded in alternating series ; later, intercalary series also budded. Avicularia small, regularly budded between two orifices, oval with a complete bar, mandible directed laterally. Ovicells present in larger colonies, fairly prominent and globular. REMARKS. The attribution of this form to Conescharellina in the past was depen- dent on the possession of regularly patterned avicularia and axial kenozooids. Both these characteristics now appear not to be exclusive to Conescharellina. Regularly patterned avicularia also occur in L. grunaui and in many colonies of L. burrowsi, although in both this last species and L. perfecta there are colonies in which the occurrence and distribution of avicularia are much less regular. Axial kenozooids are present in L. visakhensis, L. similis and L. perfecta. The general distinction between the two genera has thus become progressively restricted to the nature of the primary orifice and, to a lesser extent, to the nature of the apical region of the colony. Sinuate orifices occur in L. asymmetrica and L. grunaui, but do not have the very distinct, narrow sinus usually associated with species of Conescharellina. All other species assigned to Lacrimula have rather large, non-sinuate orifices, which tend to be wider rather than narrower antapically. The apical region in Lacrimula is typically formed by a kenozooidal tube ; that in Conescharellina by kenozooids, avicularia and lunoecia' (rootlet pores). Forms of Conescharellina from E. Africa (C. africana and an unnamed species of Conescharellina, see Appendix 3, p. 372), apparently have no rootlet pores. C. africana has an accumulation of kenozooids, ringed by avicularia, but no axial aperture (see Cook, 1966 : 215). The other species of Conescharellina has a small but distinct kenozooidal tube apically. The correlation of apical region with orifice shape, although not exclusive, has decided the attribution of C. perfecta to Lacrimula. Accordi (1947 : 108, figs 8-10) also described another form, C. veronensis, from the same Italian deposits. An analysis of further samples from the area was made recently by Braga & Munari (1972), who concluded that C. veronensis was a synonym of C. perfecta. The nature of the larger Upper Eocene specimens does, however, raise another problem. A species with a conescharelliniform colony was described as C. eocoena by Neviani (1895 : 122, fig.) from the Eocene of Mosciano, near Firenze. Only one specimen was found and that was not well preserved. Its principal characters were rounded orifices and, possibly, small avicularia between zooids ('piccolo aperture vibracolifere (?) sul solco superficial che divide i vari zoeci'). The colony had an axial length of 2-33 mm, and a proliferal width of 1-47 mm. Neviani con- sidered that C. eocoena was close to 'Batopora conica Seguenza (non Hantken)' which had been described and figured earlier in very similar terms (see Seguenza, 1880 : 42, pi. 4, fig. 10). Seguenza's specimens were from the Tongrian (Oligocene) of Antonimina, Reggio Calabria. Waters (1921 : 424) regarded Neviani's, Seguenza's and Hantken's species as synonymous, but pointed out that 'B. conica Hantken' was almost certainly a manuscript name. 360 P. L. COOK AND R. LAGAAIJ The lack of well-preserved type specimens for examination, together with the nomenclatural confusion inherent in using Seguenza's name, which antedates that of the Recent form, C. conica Haswell (1880 : 42, pi. 3, figs 7 and 8), suggests that, for the present at least, the name used for his Eocene record should be C. eocoena Neviani. Specimens in the British Museum, Palaeontology Department, labelled 'Batopora conica Hantken' from the Hantken Collection (63724 Buda, Szaboi beds, Lower Clay, Eocene) were mentioned by Waters (1921 : 424). These colonies are very large (AxL 4-00-6-50 mm, Prl 3-50-4-00 mm) and very worn. Their appear- ance is, however, similar to that of the far better preserved large specimens of L. perfecta from San Leonardo. The orifices are rounded, and there are small pores between them placed regularly as are the avicularia in L. perfecta. Series of axial kenozooids are present, and the bulbous apical region (diameter 2-00 mm) has a very small central aperture. These colonies are associated with many other, mainly erect species of Bryozoa, and with Lunulites and B. multiradiata. It thus appears possible that some, perhaps all, fossil records of Conescharellina from western Europe may prove to belong to one species-complex, attributable to Lacrimula. Lacrimula borealis1 sp. nov. (PL 7, figs 4, 5) HOLOTYPE. Middle Oligocene, 45°95'3'N, i°3i-6'E, North Sea 43/3-1 well, 1430-1470 ft, BM 052568. OTHER MATERIAL EXAMINED. As above, 840-870 ft, i colony; 1860-1890 ft, i colony ; paratypes. DESCRIPTION. Colony small, conical. Maximum number of whorls 6. Maxi- mum number of zooids in proliferal whorl 3. Maximum number of primary buds 3. AxL 0-50-1-20 mm Prl o-go-i-15 mm Lfw 0-38-0-50 mm Ifw 0-40-0-55 mm Lo 0-16-0-22 mm lo 0-15-0-17 mm Lt 0-28—0-50 mm Apical region a long narrow, prominent tube with granular calcification and occasional pores. Zooids large, with frontal wall very slightly inflated, with 6-8 marginal pores. Peristome absent, primary orifice large, rounded adapically, straight antapically, apparently without condyles. Avicularia and ovicells not seen (see below). REMARKS. The colonies are obviously at a very early astogenetic stage, but the zooids are very large in comparison with other species of comparable age. The kenozooidal tube is long, with 4-8 regularly spaced pores on its outer surface, and a very small apical aperture. Each of the three colonies includes one peripheral zooid which has a small, rounded foramen (approximately 0-07 mm in diameter) in the wall adapical to the orifice 1 borealis - (L) - northern - referring to the distribution of the species. CONESCHARELLINIFORM BRYOZOA 361 (see PI. 7, fig. 5). The cavity behind the foramen does not appear to be confluent with the zooid living chamber. By analogy with other species, this would therefore appear not to be a brooding zooid with adapical ovicell. The lack of other structures such as condyles, etc., also make it unlikely to be an avicularium, and as the chamber is apparently part of a zooid, it cannot be interpreted as a kenozooid. Until more specimens can be found, the nature and possible function of these distinctive structures is unknown. Lacrimula asymmetrica1 sp. nov. (Fie. 6 ; PL 5, ng. 4 ; PI. 7, fig. 3) HOLOTYPE. Miocene, O7°oo'S, H3°oo'E, Kombangan, W. Madura, Indonesia, Tertiary f1, Globigerinatella insueta zone (see van der Vlerk & Postuma, 1967, fig. i), BG 312, BM 052569. OTHER MATERIAL EXAMINED. As above, 39 colonies ; paratypes. DESCRIPTION. Colony pyriform, asymmetrical. Maximum number of whorls 7. Maximum number of zooids in proliferal whorl 4-5 alternating. Maximum number of primary buds 1 + 4. AxL 0-55-1-50 mm . Prl 0-42-0-88 mm Lfw 0-17-0-25 mm Ifw 0-15-0-25 mm Lo 0-08-0-13 mm 1° 0-05-0-10 mm Lt 0-15-0-30 mm Apical region a tube, with closely spaced, small external pores. Whorls of zooids alternate in number from 4 to 5 so that the whorls are only approximately arranged in a plane perpendicular to the colony axis, and the outline is asymmetrical. Secondary frontal buds forming a secondary zone of change are first produced at the adapical end of the colony. Frontal walls of zooids very finely granular and apparently non-porous, except for occasional marginal pores. Primary orifices of primary zooids with a very narrow adapical shelf which terminates abruptly to form paired condyles. Adapical part of the orifice sub-circular, the antapical part rounded-triangular, slightly narrower. No peristome present. Primary orifice of secondarily budded zooids ovate, apparently without condyles. Ovicells and avicularia not observed. REMARKS. Secondary calcification has affected at least two whorls below the apical region, completely closing the orifices, or leaving a semi-lunar slit. The specimens of L. asymmetrica comprise an almost complete astogenetic series, from the early stage of one whorl of zooids to the development of the secondary zone of astogenetic change. The earliest stage present has only five zooids. The colony measures 0-45 mm in axial length and 0-40 mm in proliferal region width. Among these zooids there is one, which may have been slightly earlier in development than the others, slightly asymmetrical in position. The ancestrular region is completed by an apical, axial 1 asymmetros - (G) - asymmetrical - referring to the shape of the colony. 24 362 P. L. COOK AND R. LAGAAIJ B 0-50 mm FIG. 6. Early astogenetic stages in Lacrimula asymmetrica sp. nov. A. Colony comprising apical kenozooid and first whorl of 5 zooids (i is asymmetrically developed). This colony has slightly smaller zooids than those of others at the same stage. K, kenozooid. B. Colony comprising apical kenozooid, first whorl of 5 and second whorl of 4 zooids. K, kenozooid. C. Colony comprising apical kenozooidal tube, enlarged by astogenetic development of kenozooids, or by ontogenetic thickening of primary tube, or by extrazooidal tissue, or a combination of any or all methods. Note three whorls of zooids present (5, 4, 5), total 14 ; the first whorl zooid orifices are occluded. T, apical kenozooidal tube. elongated kenozooid, its tube extending 0-15 mm above the colony surface. Other young colonies show slight variations on this theme, but it is interesting that the alternating whorls of 4-5 zooids are present in the earliest stages, as is the keno- zooidal tube, which in some cases reaches 0-20 mm in length at this stage. Later development consists of the budding of further alternating whorls and of the appearance of rugosities and pores on the surface of the kenozooidal tube, which becomes both thicker and longer. It is not known whether this is the result of kenozooidal budding, growth of extrazooidal tissues or both. By the time the colony has reached a size of 1-50 x 0-50 mm, and comprises 18 zooids, the tube may be 0-30 mm in length and may have expanded to a width of nearly 0-20 mm and be covered by 15-20 pores or depressions. The first whorl of orifices is often closed by calcification at this stage, and appears to have become incorporated into the calcifi- cation of the kenozooidal tube. At this stage, or even earlier, growth of the second- ary zone begins. It starts with the production of frontal buds almost simultaneously CONESCHARELLINIFORM BRYOZOA 363 from the second and third zooid whorls and extends, apparently fairly rapidly, to the remaining whorls. The primary zooids may also produce frontal buds, but at a slightly later stage. Some, but not all, of the primary zooids may have orifices closed by calcification when the buds are produced ; in others the orifices of the primary zone zooids may be seen through the orifice of the secondary zooid. Lacrimula grunaui1 sp. nov. (FiG. 7B ; PI. 7, fig. 2) HOLOTYPE. Miocene, 07°oo'S, U4°oo'E, Batuputih, E. Madura, Indonesia, Tertiary f1, Globigerinatella insueta zone (see van der Vlerk & Postuma, 1967, fig. i), G 5671, BM D52570. OTHER MATERIAL EXAMINED. As above, 6 colonies ; paratypes. DESCRIPTION. Colony elongated, pyriform, slender. Maximum number of whorls 10. Maximum number of zooids in proliferal whorl 4. Maximum number of primary buds 4. AxL 0-81-1-60 mm Prl 0-70-0-88 mm Lfw 0-20-0-25 mm Ww 0-20-0-25 mm Lo 0-11-0-13 mm lo 0-07-0-10 mm Lt 0-10-0-20 mm Apical region a tube, with closely spaced, small external pores. Each zooid with a consistently placed antapical avicularium. Frontal walls of zooids very finely granular, apparently non-porous, except for occasional marginal pores. Primary orifice with distinct, paired condyles, which separate a sub-circular adapical portion from a large, semicircular but narrower antapical part. No protruding peristome present, but the adapical side of the orifice is slightly raised. Ovicells not observed. Avicularian chambers situated adjacent to the central part of the concealed frontal wall, not reaching the colony axis. Exposed part of avicularian chamber very small, triangular, with a delicate cross-bar, the rostrum rounded-triangular and antapically directed. The orifices increase rapidly in size (1-5 times in 5 whorls). Secondary calcifi- cation affects the first two whorls below the apical region. It has almost closed the orifices, leaving a vertical slit. REMARKS. L. grunaui differs from L. asymmetrica in the very symmetrical growth of the zooid whorls, and in the possession of avicularia. Lacrimula similis2 sp. nov. (FiG. 70 ; PI. 6, figs i, 2) HOLOTYPE. Miocene, o7°oo'S, ii3°oo'E, Kombangan, W. Madura, Indonesia, Tertiary f1, Globigerinatella insueta zone (see van der Vlerk & Postuma, 1967, fig. i), BG 312, BM 052571. 1 Named for Dr H. R. Grunau, who collected the specimens. 2 similis - (L) - resembling - referring to the similarities of this species with both L. grunaui and Conescharellina. P. L. COOK AND R. LAGAAIJ 0 • 20mm FIG. 7. Orifice and interzooidal avicularium in Lacrimula. A. L. perfecta. B. L. grunaui. C. L. similis. Scale = 0-20 mm. OTHER MATERIAL EXAMINED. As above, 12 colonies ; paratypes. DESCRIPTION. Colony conical, irregular. Maximum number of whorls (second- ary zone) 10. Maximum number of zooids in primary proliferal whorl 4. Maximum number of zooids in secondary proliferal whorl 10. Maximum number of primary buds 4. AxL 1-16— i-36mm Lfw 0-25 -0-26 mm Lo 0-12-0-14 mm Lt 0-08-0-15 mm Prl 1-20-1-44 mm Ifw 0-25-0-26 mm lo 0-08-0-12 mm Apical region originally a short tube, later surrounded by secondary and tertiary zone zooids and transformed into a shallow pit. Zooids budded in alternate series. Frontal wall smooth. Primary orifice with minute condyles, delimiting a distinct antapical sinus. Avicularia arising as frontal buds, placed asymmetrically on one side of the antapical part of the orifice which is raised as a peristome. Avicularian rostrum rounded, with a complete bar, directed laterally and antapically. Subrostral chamber large. Ovicells not seen. Small, rounded axial kenozooids formed late in astogeny. REMARKS. The size and position of the avicularia distinguish this species from L. grunaui, to which it appears to be closely related. It also differs in the very early development of secondary and even tertiary zones of change, neither of which has been found in colonies of L. grunaui of similar size and astogenetic age. The primary zone of change apparently consists of whorls of four zooids, but as few as three to four whorls are present when the secondary zone of change appears. It begins as a series of small buds arising apically and surrounding the small, shallow kenozooidal CONESCHARELLINIFORM BRYOZOA 365 tube. The secondary zone consists of zooids budded directly frontally from primary zooids and is not an overgrowth. Primary zone orifices may be seen through those of secondary zone zooids as in L. asymmetrica. The zone is very regular, a complete whorl being budded simultaneously and the sequence is in an antapical direction. Superficially, the colonies may resemble those of B. multiradiata as the secondary zone zooids advance on a front over those of the primary zone. In some colonies a third zone arises antapically. The avicularia are enlarged in comparison to the zooids and seem to form a special group comparable with those found in Conescharellina africana (see Cook, 1966). In fact, L. similis superficially resembles some species of Conescharellina with its adapical avicularia and sinuate orifices. It also develops axial antapical kenozooids which cover the concealed frontal walls of the proliferal region zooids. Genus ATACTOPORIDRA Atactoporidra Canu & Bassler, 1931 : 22 ; new name for Atactopora Canu & Bassler, 19290 : 50 (preoccupied). Type species Atactopora bredaniana (Morren). Canu & Bassler placed this genus in the Orbituliporidae and described the colony as 'libre'. Their generic description described the zooids as 'amoncelees en desordre les unes sur les autres'. Additional information is now available as to the character of the genus, and the description has therefore been somewhat modified. Colony conescharelliniform, with an apical kenozooidal tube. Primary zooids budded in alternating series. Secondary to quaternary series of zooids budded concurrently with the later zooids of the primary series, originating from the exposed frontal walls of the primary zone zooids, either directly or in alternating series. Three species of Atactoporidra have been described, A. bredaniana, A. glandiformis and, more recently, A. globosa (see Labracherie, 1961). A. glandiformis was dis- cussed by Cheetham (1966 : 106, fig. 81). In this species there seems to be an actual increase in the number of zooid series in the primary zone, as well as an increase in size. Similarly budded series of zooids arise antapically quite early in the astogeny, but the apical region is rarely affected. Atactoporidra bredaniana (Morren) (PI. 5, fig. 2 ; PI. 6, fig. 6) Atactopora bredaniana (Morren) Canu & Bassler, 19290 : 51, pi. 4, figs 1-6. Atactoporidra bredaniana (Morren) Canu & Bassler, 1931 : 22, pi. 4, figs 5 and 6 ; Dartevelle, 1933 : 85, 108. MATERIAL EXAMINED. Eocene, Wemmel sands, Laeken, Belgium, 4 colonies, BM Dartevelle Coll., 033249-59. ? Eocene, Belgium, 20.11.1905, 50 colonies, BM Vassall Coll., 07864. ' 366 P. L. COOK AND R. LAGAAIJ DESCRIPTION. Colony elongated, pyriform, irregular. Maximum number of whorls 22. Maximum number of zooids in proliferal whorl 6. Maximum number of primary buds 4. AxL 2-20-4-60 mm Prl 0-80-1-20 mm Lfw 0-30-0-35 mm Ifw 0-30-0-32 mm Lo 0-13-0-16 mm lo o-io-o-n mm Lov 0-14-0-16 mm lov 0-15-0-17 mm Lt 0-15-0-30 mm Apical region with a short tube with external pits and ridges of calcification. Frontal wall of zooids finely granular, marginal pores not seen. Primary and secondary zooidal orifices apparently without condyles. Ovicells hyperstomial, prominent, associated with tertiary and quaternary zooidal series. Avicularia not budded frontally. At the sides of the raised peristomes of some ovicelled zooids, there is a small pore, which is inferred to have been an adventitious avicularium. REMARKS. The four specimens previously examined by Cook (1966 : 217) are worn, and had developed several series of secondarily frontally budded zooids before preservation. The basic structure of the colony was therefore not readily apparent. Examination of the more plentiful, astogenetically younger, and better preserved material from the Vassal Coll. not only enables the astogeny of the colony to be inferred but shows that Atactoporidra and Lacrimula have many more charac- ters in common than was first realized. The specimens (07864) include colonies in which the irregular series of secondary buds are not much developed, and in which a distinct apical kenozooidal tube is present. The arrangement of the primary zooids is regular and alternating. The orifices of the zooids are small, rounded adapically and nearly straight antapically. The ovicells have been seen in a few specimens only, and all have broken frontal walls. They are present only on the tertiary to quaternary zooidal series, are randomly placed and irregularly orientated. The greatest concentration of secondary to quaternary frontal budding is in the antapical part of the colony, causing irregularity of outline and a confused appear- ance of the surface. The budding apparently occurs at the apical end of the colony at a late astogenetic stage. No progressive closure of zooidal orifices from the apical end has been seen as in Lacrimula. Secondary zooidal series appear to arise directly from the exposed frontal walls of primary zooids, but tertiary and quaternary series often arise alternately between zooids of the secondary series, and their orientation is irregular. One colony of A. bredaniana from 07864 is rod-shaped and measures 3-00 mm long by 1-00-1-20 mm wide. The zooids of the primary zone of change are visible adapically. Secondary zooids comprise the antapical part of the colony, which is hardly wider than the adapical part. Some colonies may therefore show very little increase in size of zooids in the primary zone of change. It is not possible to see whether a primary zone of repetition occurred. Dartevelle (1933) reported A. bredaniana not only from the Wemmel sands (Bartonian) but from the earlier Ledien ('gravier de base, sables a N. variolarius'} CONESCHARELLINIFORM BRYOZOA 367 which he equated with the Upper Bracklesham of England. Many other bryozoan species, including Lunulites, were present. ACKNOWLEDGEMENTS Permission of Shell Internationale Petroleum Maatschappij , B.V. to publish this paper is gratefully acknowledged. I should like to thank particularly Mr J. W. C. van der Sijp, Dr H. R. Eckert, Dr J. Keij and Dr R. A. Pohowsky of Shell for all their help during the final stages of preparation. Acknowledgements are also made to Rothamsted Experimental Station for use of CLASP program and computer time. A large number of colleagues and correspondents have contributed over a period of years to the observations made here. The names of some of those who assisted us were known only to Dr Lagaaij, and I therefore apologize for any omissions. Foremost among those who have lent or presented specimens, and given advice on stratigraphical problems, etc. are : Dr E. Annoscia (Paleontologo dell'A.G.I.P., Milan), Dr G. Braga (Universita di Padova), Dr A. H. Cheetham (U.S. National Museum), Dr J. J. Hermes (Geologische Institut de Universiteet, Amsterdam), Dr J. Keij (Koninklijke Shell Exploratie en Productie Laboritorium), Dr M. Subba Rao (Geology Department, Andhra University, Waltair), Dr O. Schultz (Natur- historisches Museum, Wien), Dr N. Vavra (Palaontologisches Institut der Universitat, Wien) and Prof. Dr E. Voigt (Geologisches und Palaontologisches Institut, Hamburg). A demonstration of some aspects of conescharelliniform morphology and distri- bution was presented in September 1974 at the 3rd conference of the International Bryozoology Association, held at the Department of Geology, Universite Claude Bernard, Villeurbanne, Lyon. Discussions with and suggestions from colleagues attending the conference are gratefully acknowledged. I should also like to thank colleagues at the British Museum (Natural History), particularly Dr M. Hills for help and advice on statistical analysis, and Dr D. R. C. Kempe and Mr H. A. Buckley (Department of Mineralogy) and Dr B. Rosen and Mr R. Wise (Department of Palaeontology) for their help, and for access to the collections in their care. Finally, I am deeply grateful to Mr P. J. Chimonides of the Bryozoa Section for all his help, particularly in preparation of specimens and thin-sections, and for scanning microscopy and photography. P.L.C. SUMMARIES IN FRENCH AND GERMAN Les colonies conescharelliniformes et orbituliporiformes de Bryozoaires, et 1'occurrence dans les deux groupes de deux formes de bourgeonnement, le 'normal' et le 'frontal', sont decrits. Les genres Conescharellina et Trochosodon sont connus d'etre fixes aux leurs substrata par les petits racines. Us sont compares avec les genres Batopora, Lacrimula et Atactoporidra. On suggere un modele hypothetique pour la croissance primaire de la colonie dans ces genres. Les caracteres et le distribution dans le temps et dans 1'espace de genres Batopora, Lacrimula, Atactoporidra et Conescharellina sont discutes, et on enregistre 1'information a 1'egard de leur 368 P. L. COOK AND R. LAGAAIJ Geologic et leur paleoecologie. Les descriptions completes sont donnes de trois especes de Batopora, dont deux especes sont considerees d'etre nouvelles, et de sept especes de Lacrimula, dont quatre especes sont considerees d'etre nouvelles. Les efforts combines de genetique et d'environment surpassent les influences micro-environmentales dans les limites de la colonie. L'integration dans les colonies meme est considerable, elle est demontr6e par les communications interzooidales, la zonation astogenetique et le polymorphisme. La maniere de vivre particuliere permets que les conclusions paleoecologiques soient faites quant a la profondeur et le type du fond de la mer, depuis le Eocene jusqu'au Recent. Conescharelliniforme und orbituliporiforme Kolonien von Bryozoen und das Auftreten von zwei Typen von Astogenie, 'normaler' und 'frontaler', in beiden Gruppen werden beschrieben. Die Gattungen Conescharellina und Trochosodon, die durch Wlirzelchen mit ihrem Substrat verankert sind, werden mit den Gattungen Batopora, Lacrimula und Atactoporidra verglichen. Ein hypothetisches Modell fur die Anfangsentwicklung der Kolonien dieser Gattungen wird vorgeschlagen. Die Merkmale und die zeitliche und raumliche Verbreitung der Gattungen Batopora, Lacrimula, Atactoporidra und Conescharellina werden diskutiert und die verfiigbaren Daten iiber ihre Okologie und Palao-Okologie werden angefiihrt. Drei Arten der Gattung Batopora, zwei davon neu, und sieben Arten der Gattung Lacrimula, vier davon neu, sind aus- fiihrlich beschrieben. Der kombinierte Einfluss von Genetik und Umwelt iiberwiegt die Mikroeinflusse innerhalb der Kolonie. Integration innerhalb der Kolonien ist betrachtlich und driickt sich aus in interzooidaler Kommunikation, astogenetischer Zonenbildung und Poly- morphismus. Die spezialisierte Lebensweise gestattet palao-okologische Schliisse iiber die Tiefe und Beschaffenheit des Meeresbodens vom Eozan bis zur Jetztzeit. APPENDICES r Appendix i The following records of Batopora have been plotted on Map i. Many records include more than one species ; previously unpublished observations are noted thus '(R. L. obs.)'. RECENT. Zanzibar, 805 m (Cook, 1966) : B. murrayi. Fiji, Challenger Stn 1740, 384 m (P. L. C. obs.) : B. murrayi. PLIOCENE. Sassuolo, near Modena, Italy (Namias, 1891) : B. rosula. Huelva-i well, near Huelva, Spain, 648 m (ditch cuttings) (R. L. obs.) : B. rosula. MIOCENE. Crete, Sample 6-27, Almiri section (R. L. obs.) : B. rosula. Malta, Sample PLR 4067, Blue Clay Formation (R. L. obs.) : Batopora sp. Arzeboun II, Prov. du Mazanderan, Iran (Dartevelle, 1948) : B. ernii. Benestare, Calabria, Italy (Seguenza, 1880) : B. rosula. N. Italy, Sample SM 55, Torre Veglio Section (R. L. obs.) : Batopora sp. Abruzzi, Italy (Ceretti & Poluzzi, 1970) : B. multiradiata. Sardinia, Sample MU 214, Gesturi-Furtei, Central N. Campidano (R. L. obs.) : Batopora sp. Baden, near Vienna (Reuss, 1848) : B. rosula. W. Aquitaine Basin (Le Saint, 1961) : B. rosula and B. multiradiata. R.H03, Cuidad Granada Formation (Globorotalia kugleri zone), Velez Rubio, T. B. Roep Coll. (R. L. obs.) : B. rosula. CONESCHARELLINIFORM BRYOZOA 369 Recent -Holocene © Plio/Pleistocene O Miocene Oligocene Eocene MAP i. Distribution of species of Batopora in time and space (see Appendix i). OLIGOCENE. Rockall Plateau, Site 117, 1038 m (Cheetham & Hakansson, 1972) : Batopora sp. Boring K 62, 23.90-24, 80 m surface, Ijzendijke, Netherlands (presumably Boom Clay, Rupelian) (R. L. obs.) : B. asterizans sp. nov. North Sea, Whitehall 43/3-1 well, 840-870 ft ditch cuttings (R. L. obs.) : B. asterizans sp. nov. Spain, Samples K 48-50, Moli de Llinares, Villajoyosa, Alicante Province (R. L. obs.) : B. multiradiata. Calbe and Saale, Germany (Reuss, 1867) : B. stoliczkai. EOCENE. Skalnik, Central Carpathians (Malecki, 1963) : B. multiradiata and B. stoliczkai. Buda, Marne de Buda (Couches a Clavulinoides szaboi] (Cook, 1966) : B. multi- radiata. Gotzreuth, Bavaria (Koschinsky, 1885) : B. scrobiculata and B, multiradiata, and (P. L. C. obs.) USNM and BM : B. grandis. 370 P. L. COOK AND R. LAGAAIJ Priabona and other northern Italian localities (Reuss, 1869 ; Waters, 1891 ; Braga, 1963) : B. multiradiata, B. stoliczkai and B. rosula. Bordeaux area (Labracherie, 1961) : B. stoliczkai. Sydenham and Hampstead, London Clay (Gregory, 1893) : B. clithridiata. Appendix 2 The following records of Lacrimula and Atactoporidra have been plotted on Map 2. RECENT. Zanzibar, 101 and 207 m (Cook, 1966) : L. burrowsi, and L. pyriformis. S.E. Africa, near Durban, 102 m (Cook, 1966) : L. burrowsi. S. China Sea, 677 m (P. L. C. obs.) : Lacrimula sp. HOLOCENE. Bay of Bengal, E. India, 89 m (Rao & Rao, 1973) : L. visakhensis. MIOCENE. Sample BG 312, Kombangan, Madura, Indonesia (N.E. Java) (R. L. obs.) : L. similis sp. nov., and L. asymmetrica sp. nov. Recent - Holocene O Miocene Oligocene Eocene MAP 2. Distribution of species of Lacrimula and Atactoporidra in time and space (see Appendix 2). CONESCHARELLINIFORM BRYOZOA 371 Sample G 5671, west of Batuputih, Madura, Indonesia (R. L. obs.) : L. grunaui sp. nov. OLIGOCENE. North Sea, Whitehall 43/3-1 well, 840-870 ft ditch cuttings (R. L. obs.) : L. borealis sp. nov. Belgium, Tongrian, Rupelian (Canu & Bassler, 1931) : Atactoporidra bredaniana. EOCENE. Near Verona, and near Possagno, N. Italy (Accordi, 1947, and P. L. C. obs.) : L. perfecta. Belgium, Wemmelian (Canu & Bassler, 1931) : A. bredaniana. Selsey, Barton Clay, England (Cheetham, 1966) : A. glandiformis. Marcheprime, Gironde (Labracherie, 1961) : A. globosa. Appendix 3 The following records of Conescharellina have been noted, asterisk (*) are plotted on Map 3. Those marked with an RECENT Japan Hong Kong Philippines Celebes Sea *4i°36'N, I40°36'E, no depth 4i°3i'N, I40°36'E, 80-5 m *35°N, I39°E, 300 m 35°N, i3i°E 33°N, I29°E, no depth *33°05'N, i30°03'E, 40 m *2i°33'N, n6°i5'E, 161 m i3°2i'N, i22°i8'E, 970 m I2°I5'N, I23°57'E, 146 m I2°O4'N, I24°O4'E, 193 m n°O9'N, i23°5o'E, 60-5 m io°oi'N, 42'E, 216 m , 46 m 6°u'N, i2i°o8'E, 295-5 6°09'N, i2o°58'E, 53 m 6°05'N, i2i°02'E, 35 m 6°04'N, I20°58'E, 37 m 5°4i'N, I2O°27'E, 44m 5°4i'N, I20°47'E, 38m 5°3o'N, I20°O7'E, 612 m 5°24'N, i20°27'E, 44 m (Canu & Bassler, 1929) C. catella (Silen, 1947) (Harmer, 1957) (Silen, 1947) (Silen, 1947) (Canu & Bassler, 1929) (Canu & Bassler, 1929) (Canu & Bassler, 1929) (Canu & Bassler, 1929) (Canu & Bassler, 1929) (Canu & Bassler, 1929) (Canu & Bassler, 1929) (Canu & Bassler, 1929) (Canu & Bassler, 1929) (Canu & Bassler, 1929) (Canu & Bassler, 1929) (Canu & Bassler, 1929) (Canu & Bassler, 1929) (Canu & Bassler, 1929) (Canu & Bassler, 1929) C. parviporosa C. striata C. catella Conescharellina spp. C. striata C. concava C. breviconica C. breviconica C. milleporacea C. catella C. breviconica C. milleporacea C. breviconica C. breviconica C. jucunda C. delicatula C. obliqua C. jucunda C. delicatula C. milleporacea C. elongata C. catella C. grandiporosa C. elongata C. milleporacea C. elongata C. milleporacea C. lunata C. elongata C. radiata C. elongata 372 P. L. COOK AND R. LAGAAIJ RECENT (cont.) Celebes Sea 5°2o'N, II9°58'E, 440 m (Canu & Bassler, 1929) C. milleporacea (cont.) 5°io'N, H9°47'E, 421 m (Canu & Bassler, 1929) C. elongata C. catella 4°54'N, H9°O9'E, 3iom (Canu & Bassler, 1929) C. radiata Malacca Str. *4°2o'N, 99°35'E, 50 m (Sil6n, 1947) C. striata N. Celebes i°N, I23°E, 72 m (Harmer, 1957) C. elongata N. New Guinea *o°, I30°E, 18-32 m (Harmer, 1957) C. jucunda Makassar Straits *I°I9'S, n8°43'E, 2161 m (Canu & Bassler, 1929) C. radiata & Java Sea C. transversa 2°S, ii5°E, 59 m (Harmer, 1957) C. catella 2°3o'S, io7°io'E, 15-27 m (Silen, 1947) C. brevirostris C. longirostris C. laevis off Java *7°S, ii5°E, 88m (Harmer, 1957) C. angustata 7°S, H5°E, 1060 m (Harmer, 1957) C. distalis Java Sea 6°05'S, II4°07'E, 82 m (Harmer, 1957) C. ovalis Arafura Sea 7°S, I32°E, 58-5 -66m (BM) C. crassa Pt Moresby, *9°S, i47°E, no depth (BM) C. crassa Papua Torres Str. 9°, I40°E, 27-5-37 m (BM) C. crassa Murray Is. *io°S, I44°E, 27-5 m (BM) C. conica Torres Str. Baudin Is. I2°S, i25'E, 27-5 m (BM) Conescharellina spp. Timor Sea Holothuria Bank *I3°09'S, I26°22'E, 66-71 m (BM) Conescharellina spp. i3°oi'S, I25°58'E, 32 m (BM) Conescharellina spp. E. Australia I4°S, I44°E, 24 m (Waters, 1921) C. philippinensis (P. Charlotte Bay) I9°42'S, i48°2i'E, 42 m (Waters, 1921) C. conica (Holborn Is.) 27°3o'S, i52°3o'E, no depth (Waters, 1921) C. philippinensis *32°3o'S, i52°3o'E, 40-46 m (Waters, 1921) C. cancellata (Pt Stephens) C. philippinensis C. angulopora 33°S, I52°E, no depth (Waters, 1921) C. philippinensis 35 km E. of Pt Jackson, (BM) C. biarmata 146 m C. angulopora C. eburnea S. E. Australia 36°S, i35°3o'E, 190 m (BM) (56 km S.W. of Neptune Is.) *40°S, I45°E, no depth (Waters, 1921) (Bass Straits) S. E. Africa *near Durban, 102 m (Cook, 1966) E. Africa *Zanzibar, 'Dalrymple' (P. L. C. obs.) Stn 98, 69-5 m Conescharellina sp. C. angulopora C. africana Conescharellina sp. CONESCHARELLINIFORM BRYOZOA 373 PLIOCENE-PLEISTOCENE S. Japan *a8°N, i3o°E (Kikai Jima) (Kataoka, 1961) Sarawak *3°35'N, ii2°55'E (R. L. obs.) (Patricia Is.) Australia *Weymouth Bore, Adelaide, (BM, P. L. C. obs.) 310-330 ft Conescharellina sp. Conescharellina sp. Conescharellina sp. 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N.S.W. 5:33-44. HEJJAS, I. 1894. Adatok erdely Tertiar Bryozoa -Faunajahoz. (Beitrage zur kenntniss der Tertiare Bryozoenfauna Siebenburgens.) Ert. erdel. Mus.-Egyes. Orvos. Term.-Tudom. Szakoszt. 19, 2, 16 : 113-152 ; 19, 2, 17 : 201-216 ; 19, 3, 16 : 217-260. Hsu, K. J., RYAN, W. B. F. & CITA, M. B. 1972. Late Miocene desiccation of the Mediter- ranean. Nature, Lond. 242 : 240-244. JULLIEN, J. 1883. Dragages du 'Travailleur'. Bryozoaires. Especes draguees dans 1'Ocean atlantique en 1881. Bull. Soc. zool. Fr., 7 : 497-529. KATAOKA, J. 1957. Bryozoa from the Daishaka Formation (Pliocene) .... Trans. Proc. paleont. Soc. Japan, n.s., 28 : 143-153. 1961 . Bryozoan fauna from the 'Ryukyu Limestone' of Kikai-jima, Kagoshima Prefecture, Japan. Sci. Rep. Tohoku Univ. Geol. 32, 2 : 213 — 272. KOSCHINSKY, C. 1885. Bryozoen Fauna der alteren Tertiarschichten des siidlichen Bayerns. I, Cheilostomata. Palaeontographica, 32 : 1-73. LABRACHERIE, M. 1961. Les Bryozoaires eocenes du sous-sol du Bordelais. Observations paleontologiques et biostratigraphiques. These, 3me Cycle d'Enseignement superieur, pp. 1-210. Laboratoire de Geologic approfondie, Universite de Bordeaux. 1970. Les Bryozoaires dans 1'Eocene Nord-Aquitain. Signification biostratigraphique et paleoecologique, 1 : 1-141, i-xxxiv ; 2: 142-314; 3, tables 1-36. These de doctorat d'etat des Sciences naturelles . . . de 1'Universite de Bordeaux No. 316. 1971. Evolution generate des assemblages de Bryozoaires dans 1'Eocene du bassin nord- aquitain. Somm. Sea. Soc. geol. 21 : 388-389. LAGAAIJ, R. i963a. Cupuladria canariensis (Busk) -portrait of a bryozoan. Palaeont. 6, i : 172-217. i963b. New Additions to the Bryozoan fauna of the Gulf of Mexico. Publ. Inst. mar. Sci. Univ. Texas, 9 : 162-236. LE SAINT, A. 1961. Etude paleontologique et biostratigraphique des Bryozoaires girondiens du Bassin ouest-Aquitain. These, 3me Cycle d'Enseignement superieur. No. 123 : 1-223. 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DE & NEVIANI, A. 1897. Corallarii e Briozoi Neogenici di Sardegna. Boll. Soc. geol. Ital. 15, 4 : 571-595. RAO, M. S. & RAO, T. K. 1973. A new bryozoan species from the shelf sediments off the East Coast of India. Curr. Sci. 42, 14 : 506-507. REUSS, A. E. 1848. Die fossilen Polyparien des Wiener Tertiarbeckens. Haidingers Natur- historisches Abhandl. II. Vienna. 1867. t)ber einige Bryozoen aus dem deutschen Unterologocan. Sber. Akad. Wiss. Wien, 55, i : 216-234. 1869. Palaontologische Studien iiber die alteren Tertiarschichten der Alpen. II. Die fossilen Anthozoen und Bryozoen der Schichtengruppe von Crosara. Sber. Akad. Wiss. Wien, 58 : 288-292. SEGUENZA, G. 1880. Le formazioni Terziare nella provincia di Reggio (Calabria). Atti A cad. Lincei, ser. 3, 6 : 1—446. SILEN, L. 1947. Conescharellinidae (Bryozoa Gymnolaemata) collected by Prof Dr Sixten Bock's Expedition to Japan and the Bonin Islands, 1914. Ark. Zool. 39A, 9 : 1-61. SISSINGH, W. 1972. Late Cenozoic Ostracoda of the South Aegean Island arc. Utrecht Micropal. Bull. 6 : 1-187. SOLD AMI, A. 1780. Saggio orittografico ovvero osservazioni sopra le terre nautiliche ed ammonitiche della Toscana. Sienna. VAN DER VLERK, I. M. & POSTUMA, J. A. 1967. Oligo-Miocene Lepidocyclinas and planktonic Foraminifera from East Java and Madura, Indonesia. Proc. K. ned. Akad. Wet. ser. B, 70, 4: 391-398. WATERS, A. W. 1891. North-Italian Bryozoa. Pt i, Cheilostomata. Q. Jl. geol. Soc. 47 : 1-34- 1919. Batopora and its allies. Ann. Mag. nat. Hist. (9), 3 : 79-94. 1921. Observations on the relationships of the Selenariidae. /. Linn. Soc. Zool. 34 : 399- 427. P. L. COOK Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD PLATE i Batopora and Trochosodon - early astogenetic stages. Photographs by scanning electron microscope. FIG. i. B. stoliczkai Reuss. NMV 1867. XII, isa-d, Biinde, Germany, Oligocene. Diameter of colony 0-72 mm. xyi. FIG. 2. B. murrayi Cook. BM 'Dalrymple' Stn 98, Zanzibar, Recent. Diameter of colony 0-63 mm. X95. FIG. 3. Trochosodon sp. i. BM 'Challenger' Stn 185, off Cape York, N. Australia, Recent. Diameter of colony 0-93 mm. xyi. FIG. 4. Trochosodon sp. i. BM as above, showing specialized rootlet pores (lunoecia, arrowed), x 178. FIG. 5. Trochosodon sp. 2. BM as above, very young colony with ovicell and lunoecia (arrowed). Diameter of colony 0-84 mm. x 85. FIG. 6. Trochosodon sp. 2. BM as above, lateral view of colony ; note extensive develop- ment of apical tissues, x 85. Bull. Br. Mus. nat. Hist. (Zool.) 29, 6 PLATE i PLATE 2 Batopora. Photographs by scanning electron microscope. FIG. i. B. clithridiata (Gregory). BM 61357, Sydenham, London Clay, Eocene. See also PI. 5, fig. 5. X45. FIG. 2. B. stoliczkai Reuss. USNM, Calbe, Germany, Oligocene. Lateral view, note ovicells. x 73. FIG. 3. B. asterizans sp. nov. North Sea, Middle Oligocene. Lateral-frontal view. x 72. FIG. 4. B. asterizans sp. nov. As above, view of apical region, x 128. FIG. 5. B. grandis sp. nov. USNM 88882 pt., Gotzreuth, Bavaria, Eocene, x 12. FIG. 6. B. grandis sp. nov. As above, zooids with sinuate orifices and a broken ovicell. x 72. Bull. Br. Mus. nat. Hist. (Zool.) 29, 6 PLATE 2 PLATE 3 Batopora. FIG. i. B. multivadiata Reuss. USNM 88881 pt., Gotzreuth, Bavaria, Eocene. Young colony with one zone of change. x 21. FIG. 2. B. rosula (Reuss). NMV Baden, Austria, Miocene, x 36. FIG. 3. B. rosula (Reuss). NMV Baden, Austria, Miocene, x 36. FIG. 4. B. stoliczkai Reuss. NMV, Biinde, Germany, Oligocene. x 36. FIG. 5. B. scrobiculata. Koschinsky. USNM 88882 pt., Gotzreuth, Bavaria, Eocene. Note ovicells. X2i. FIG. 6. B. scrobiculata. Same specimen, antapical side of colony. x 21. FIG. 7. B. scrobiculata. Same specimen, showing antapical axial kenozooids. X2i. FIG. 8. B. grandis sp. nov. USNM 88882 pt., Gotzreuth, Bavaria, Eocene, x 16. Bull. Br. Mus. mat. Hist. (Zool.) 29, 6 PLATE 3 PLATE 4 Batopora and Lacrimula. Photographs by scanning electron microscope (except Fig. 5). FIG. i. B. rosula (Reuss). Crete, Miocene. Lateral view, showing prominent secondarily budded apical kenozooid. x 44. FIG. 2. B. rosula, as above, view from apical side, x 44. FIG. 3. L. perfecta (Accordi). BM, Possagno, N. Italy, Eocene. Proliferal region zooids showing orifice with lateral condyles (arrowed) and marginal pores. See also PI. 7, fig. i. xio5. FIG. 4. B. multiradiata Reuss. USNM 88881 pt., Gotzreuth, Bavaria, Eocene. Lateral view showing overgrowth of secondary zone zooids and broken ovicells (arrowed). x 28. FIG. 5. B. multiradiata. USNM 71205, Val di Lonte, N. Italy, Bartonian, Eocene. Basal view. X2i. FIG. 6. B. multiradiata, as above. Lateral view showing ovicells and tertiary series of apical zooids and kenozooids. x 25. Bull. Br. Mus. nat. Hist. (Zool.) 29, 6 PLATE 4 PLATE 5 Lacrimula, Atactoporidra and Batopora. Photographs by scanning electron microscope. FIG. i. L. burrowsi Cook. BM 1965.8.24.9, Zanzibar, Recent. Apical region ; note kenozooidal tube and secondarily budded zooids and avicularia. See also PL 6, fig. 3. x8i. FIG. 2. A. bredaniana (Morren). BM 07864, Belgium, Eocene. Apical region; note similarity with L. burrowsi. See also PI. 6, fig. 6. x 58. FIG. 3. L. pyriformis Cook. BM 1965.8.24.12, Zanzibar, Recent. Apical region ; note the reticulation of the surface of the tube, and the regularly placed pores. See also PI. 6, fig. 5. xi56. FIG. 4. L. asymmetrica sp. nov. BG 312, W. Madura, Indonesia, Miocene. Apical region showing one of the pores. See also PI. 7, fig. 3. x 104. FIG. 5. B. clithridiata (Gregory). BM 61357, Sydenham, London Clay, Eocene. Inter- zooidal avicularium with complete bar (av). See also PL 2, fig. i. i. x 126. FIG. 6. L. burrowsi Cook. BM 1965.8.24.9, Zanzibar, Recent. Ovicells and avicularia. X74- Bull. Br. Mus. nat. Hist. (Zool.) 29, 6 PLATE 5 PLATE 6 Lacrimula and Atactoporidra. Photographs by scanning electron microscope. FIG. i. L. similis sp. nov. BG 312, Madura, Indonesia, Miocene. Colony with a secondary zone of change. x 57. FIG. 2. L. similis sp. nov. As above, showing primary and secondary zone zooids with avicularia. x 57. FIG. 3. L. burrowsi Cook. BM 1965.8.24.9, Zanzibar, Recent. View from the apical end of a colony. See also PI. 5, fig. i. x 34. FIG. 4. L. visakhensis Rao & Rao. BM 1070.8.8.1, Bay of Bengal, India, Holocene. Colony with apical ovicells. x 38. FIG. 5. L. pyriformis Cook. BM 1965.8.24.12, Zanzibar, Recent. Colony with ovicelled zooids in the proliferal whorl. See also PL 5, fig. 3. x 23. FIG. 6. A . bredaniana (Morren). BM 07864, Belgium, Eocene. See also PI. 5, fig. 2. x 30. Bull. Br. Mus. nat. Hist. (Zool.) 29, 6 PLATE 6 PLATE 7 Lacrimula. Photographs for Figs. 1-3 by scanning electron microscope. FIG. i. L. perfecta (Accordi). BM, Possagno, N. Italy, Eocene. View of proliferal region, note axial kenozooids. See also PI. 4, fig. 3. x 44. FIG. 2. L. grunaui sp. nov. G 5671, Madura, Indonesia, Miocene. x8i. FIG. 3. L. asymmetvica sp. nov. BG 312, Madura, Indonesia, Miocene. See also PI. 5, fig. 4. x 60. FIG. 4. L. borealis sp. nov. 43/3-1, North Sea, Oligocene. x 37. FIG. 5. L. borealis. Same specimen, x 37. Bull. Br. Mus. nat. Hist. (Zool.) 29, 6 PLATE 7 PLATE 8 Lacrimula burrowsi Cook - sections through restored colonies. FIG. i. Transverse section showing alternating zooid whorls. The zooids nearer the axis belong to the whorl adapical in position to the zooids at the periphery. Note the axial con- centration of communicating tubules derived from marginal pores. Opercula and avicularian chambers arrowed. x 41. FIG. 2. Transverse section near antapical end of a large colony. Ovicells (large cavities) and avicularian chambers (small cavities) arrowed. x 28. FIG. 3. Tangential longitudinal section through zooid orifices. Note the increase in size with astogenetic position. Marginal pores and condyles arrowed. x 41. FIG. 4. Thick transverse section through an apical tube. Note passage of tubular keno- zooidal cavities at an angle through the calcification from exterior to interior (arrowed), x 91. FIG. 5. Deep tangential longitudinal section of the apical region. Note tubular kenozooidal cavities passing through the calcification at various angles and tubules from pores connecting avicularian chambers with primary zooids. Avicularian chambers arrowed, a, a1 zooids of ancestrular whorl. x 86. FIG. 6. Thick section through apical region of an astogenetically older colony. Note kenozooidal cavities and axial tube cavity with stained tissue. Kenozooidal cavities arrowed, x 91. Bull. Br. Mus. nat. Hist. (Zool.) 29, 6 PLATE 8 4 A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) 1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96 ; 14 Plates. 1965. (Out of Print.) 2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125 ; 29 Plates, 77 Text-figures. 1969. £4-50. 4. HAYNES, J. R. Cardigan Bay Recent Foraminifera (Cruises of the R.V. Antur) 1962-1964. Pp. 245 ; 33 Plates, 47 Text-figures. 1973. £10-80. 5. WHITEHEAD, P. J. P. The Clupeoid Fishes of the Guianas. Pp. 227 ; 72 Text-figures. 1973. £9*70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : the Biology and Evolution of a Species Flock. Pp. 134 ; i Plate, 77 Text-figures. 1974. £3-75. Hardback edition £6. Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol B§4 jNU