BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY VOL. 15 1967-1968 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) LONDON: 1970 DATES OF PUBLICATION OF THE PARTS No. i 3 January 1967 No. 2 . . . . . -13 January 1967 No. 3 . . . . . -24 February 1967 No. 4 . . . . . -24 February 1967 No. 5 . . . . . .16 May 1967 No. 6 ..... .28 July 1967 No. 7 ..... 14 November 1967 No 8 ..... 9 January 1968 PRINTED IN GREAT BRITAIN BY ALDEN & MOWBRAY LTD AT THE ALDEN PRESS, OXFORD CONTENTS ZOOLOGY VOLUME 15 PAGE No. i. The evolution, host relationships and classification of the nematode Superfamily Heterakoidea. By WILLIAM G. INGLIS i No. 2. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae) . Part VI. By P. H. GREENWOOD 29 No. 3. The Vachell collection of Chinese fishes in Cambridge. By P. J. P. WHITEHEAD & K. A. JOYSEY 121 No. 4. Notes on asteroids in the British Museum (Natural History) V. Nardoa and some other ophidiasterids. By AILSA McGowN CLARK 167 No. 5. New species of Stellicola (Copepoda, Cyclopoida) associated with star- fishes in Madagascar, with a redescription of 5. caeruleus (Stebbing, 1900. By ARTHUR G. HUMES & JU-SHEY Ho 199 No. 6. The cirriped fauna of tropical West Africa. By H. G. STUBBINGS 227 No. 7. Polyzoa (Bryozoa) from West Africa: the Pseudostega, the Cribri- morpha and some Ascophora Imperfecta. By PATRICIA L. COOK 321 No. 8. Caligus longicaudatus Brady, 1899 (Caligidae: Copepoda) By R. R. PARKER 353 Index to Volume 15 369 THE EVOLUTION, HOST RELATIONSHIPS AND CLASSIFICATION OF THE NEMATODE SUPERFAMILY HETERAKOIDEA WILLIAM G. INGLIS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 15 No. i LONDON: 1967 THE EVOLUTION, HOST RELATIONSHIPS AND CLASSIFICATION OF THE NEMATODE SUPERFAMILY HETERAKOIDEA BY WILLIAM G. INGLIS, PLD., D.Sc. "VvMv British Museum (Natural Hi«ory) Pp. 1-28 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 15 No. i LONDON: 1967 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the 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. 15, No. I of the Zoological 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.) Trustees of the British Museum (Natural History) 1967 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 3 January 1967 Price Ten Shillings THE EVOLUTION, HOST RELATIONSHIPS AND CLASSIFICATION OF THE NEMATODE SUPERFAMILY HETERAKOIDEA By WILLIAM G. INGLIS CONTENTS Page APOLOGIA ............ 4 SYNOPSIS ............ 4 INTRODUCTION ........... 4 COMPARATIVE ANATOMY OF THE SUPERFAMILY ..... 5 The male tail ........... 6 The spicules ........... 7 SYSTEMATIC GROUPS WITHIN THE SUPERFAMILY ..... 8 Family Heterakidae . . . . . . . . . .10 Family Aspidoderidae . . . . . . . . .10 Family Ascaridiidae . . . . . . . . . .10 MORPHOLOGICAL AND EVOLUTIONARY TRENDS . . . . . .11 GEOGRAPHICAL AND HOST DISTRIBUTION . . . . . -13 SYSTEMATIC HISTORY .......... 14 ANNOTATED CLASSIFICATION OF THE SUPERFAMILY . . . . .17 Heterakoidea Railliet & Henry, 1912 . . . . . . 17 Heterakidae Railliet & Henry, 1912 . . . . . . 17 Heterakinae Railliet & Henry, 1912 . . . . . 18 Heterakis Dujardin, 1845 . . . . . 19 Odontoterakis Skrjabin & Schikhobalova, 194712 . . 20 Pseudaspidodera Baylis & Daubney, 1922 ... 20 Meteterakinae Inglis, 1958^ . . . . . .21 Meteterakis Karve, 1930 . . . . . .21 Gireterakis Lane, 1917 . . . . . .21 Spinicaudinae Travassos, 1920 . . . . . .21 Spinicauda Travassos, 1920^ ..... 22 Africana Travassos, 1920 . . . . . .22 Moaciria Freitas, 195612 . . . . . .22 Strongyluris Miiller, 1894 ...... 23 Aspidoderidae Skrjabin & Schikhobalova, 1947 . . . . 23 Aspidoderinae Skrjabin & Schikhobalova, 1947 ... 23 Aspidodera Railliet & Henry, 1912 . . . . 23 Ansiruptodera Skrjabin & Schikhobalova, 1947 . . 23 Lauroiinae Skrjabin & Schikhobalova, 1951 . -23 Lauroia Proen9a, 1938 ... .24 Pamspidodera Travassos, 1914 ..... 24 Ascaridiidae Travassos, 1919 ... • 24 KEY TO THE GENERA OF THE SUPERFAMILY ...... 24 ACKNOWLEDGEMENTS ...... 25 REFERENCES ...... -25 APPENDIX ..... • ... 27 ZOOL. 15, i. J§ 4 W. G. INGLIS APOLOGIA SEVERAL years ago I published a classification of the nematode Superfamily Heter- akoidea (Inglis, 1958) but presented no justification as I always hoped to produce a full revision. It has become increasingly clear that this is never likely to be com- pleted largely because of a shift in my interests in conjunction with a lack of material on which to base such a revision. I therefore present here reasons for the classifica- tion previously proposed and regret being unable to complete the whole work. No illustrations are given as they can be found in Skrjabin, Schikhobalova and Lagodovskaja (1961) and Chabaud (1965). SYNOPSIS The comparative anatomy of the Superfamily Heterakoidea is described and the morpho- logical sequences recognized are used as the basis for the classification proposed. The evolution of the Superfamily is interpreted in terms of (i) intra-host localisation ; (2) major host groups (i.e. amphibia, reptiles, birds and mammals) ; (3) the feeding habits and ecology of the hosts ; and (4) geographical restriction. The Superfamily is classified into three families, five subfamilies and fourteen genera, thus : Heterakidae : Heterakinae (Heterakis, Odontoterakis, Pseud- aspidodera) ; Meteterakinae (Meteterakis, Gireterakis) ; and Spinicaudinae (Spinicauda, Africana, Moaciria, Strongyluris) . Aspidoderidae : Aspidoderinae (Aspidodera, Ansirupto- dera) ; and Lauroiinae (Lauroia, Paraspidodera) . Ascaridiidae (Ascaridia). A key is given to the genera of the Superfamily. The following genera are not accepted largely because the morphology of the specimens does not correspond to the published descriptions, thus : Preterakis Freitas, 1956 ; Heterakoides Freitas, 1956 ; Pareterakis Freitas, 1956 ; Pseudaspidoderoides Freitas, 1956 ; Pseudaspidoderina Freitas, 1956 ; Raillietakis Freitas, 1956 ; Sexansodera Skrjabin & Schikhobalova, 1947 ; Bellaplectana Skrjabin, Schikhobalova & Lagodovskaja, 1961 ; Spinaspidodera Skrjabin & Schikhobalova, 1947 ; Ganguleterakis Lane, 1914 ; and Cheloniheterakis Yamaguti, 1961. INTRODUCTION No full revision of the Heterakoidea or Heterakidae has ever been carried out although several groupings have been proposed largely based on other published work. However, Travassos (1913) did produce a partial revision based largely on the species occurring in Brazil. Other minor changes in the classification were proposed later but no major alterations were suggested for almost thirty-five years when Skrjabin & Schikhobalova (1947, 19470), Freitas (1956), Inglis (1957, 1958) and Chabaud (1957) all advanced new groupings. Most recently the superfamily has been re-grouped once more by Skrjabin, Schikhobalova & Lagodovskaja (1961) and by Yamaguti (1961). All the classifications proposed, except my own, have been based on biblio-taxo- nomic studies while the results presented here are based on a study of virtually all the type material available in the world's collections together with a study of many other specimens. In anticipation of the discussion that follows it may be stressed that the following genera proposed by Freitas (1956) and Skrjabin et al. (1961) cannot stand because the descriptions they classify are contradicted by the morphology of the specimens, thus: Bellaplectana Skrjabin et al., 1961 is a synonym of Moaciria since Aplectana pharyngeodentata Belle, 1957 (of which I have studied the type series) does not THE SUPERFAMILY HETERAKOIDEA 5 possess the so-called teeth in the buccal cavity on which the genus is based. These structures are simply the retracted lips, a common artefact in poorly preserved Spinicauda and Moaciria species. Preterakis Freitas, 1956 falls as a synonym of Africana because the fusion of the caudal alae on the male tail on which it is based does not occur. Such an apparent fusion occurs in many specimens of the super- family but is an artefact due to contraction during fixation and even the artefact does not occur in all the specimens of the type series of Africana astylosterni Sand- ground, 1933. As a consequence the subfamily Preterakinae Freitas, 1956 must also disappear. Finally the following genera proposed by Freitas (1956) are not accepted by Skrjabin et al. (1961) nor by myself, thus: Heterakoides Freitas, 1956 since, although Spinicauda triaculeatus Kreis, 1933 is certainly slightly different from the other species referred to Meteterakis (Inglis, 19580) I still do not feel it warrants generic separation. Similarly the genus Pareterakis Freitas, 1956 is a synonym of Meteterakis since there are no characters which allow it to be distinguished. Pseuda- spidoderoides falls because the anastomozing cordons reported by Chakravarty (1938) appear to be due to contraction of the type specimens, which I have seen. COMPARATIVE ANATOMY OF THE SUPERFAMILY All the species referable to the Superfamily, with the exception of the genera Ascaridia and Lauroia which are discussed later, have the following features in common. The head bears three large, distinct lips, the detailed structure of which varies throughout the family and has been considered elsewhere (Inglis, 1957). In summary, however, the dorsal lip carries two doubled papillae and each ventro- lateral lip carries a single ventro-lateral doubled papilla and a single lateral papilla associated with the amphid. The inner surface of each lip carries an anterior flange which varies in relative size throughout the superfamily. The head in Lauroia, which I did not deal with before, is typical of this pattern with lobes con- necting the lips. It is, however, characterized by the development of three plate- like regions posterior to each lip. This is clearly a development of cordons from one source (see particularly Proenga's (1938) figures of L. travassosi) comparable to the condition in Gireterakis. In Ascaridia the lips are large and stout but otherwise typically Heterakoid. The anterior region of the oesophagus is divided into three separate anterior lobes, previously called " pharyngeal portions ", at the tip of each of which is a cuticular onchium, except in one case. From studies completed since 1957 it now appears that the lips are operated by a system of fibres which are not arranged in a septum as in the Ascaridoidea (Inglis, 1965) but this has not been fully analysed. In addition in the Heterakinae there is generally a pair of lateral papillae lying on the body just posterior to the lip region. The oesophagus, except in Ascaridia, is modified anteriorly into a short region generally called a pharynx and swells fairly evenly posteriorly to form a bulb-like region, containing three valves, which is not clearly offset from the remainder of the oesophagus by a constriction. The distinction between the pharyngeal region, the bulb and the remainder of the oesophagus is the presence of marginal tubes 6 W. G. TNGLIS in the radii of the lumen of the oesophagus in the intervening length between the bulb and pharynx. Into these marginal tubes project a series of paired leaf -like cuticular structures and this arrangement gives the oesophagus its typical longitudinal doubled appearance. The oesophagus is relatively long and narrow with a markedly small posterior bulb in Aspidodera, Ansiruptodera, Lauroia and Paraspidodera; the oesophagus is less markedly narrow in Africana, Gireterakis, Meteterakis, Moaciria, Spinicauda and Strongyluris, and is relatively short and stout in Heterakis, Odontoterakis, and Pseudaspidodera. The oesophagus is club-shaped without a posterior bulb in Ascaridia. The nuclei of the ventral oesophageal glands are doubled. Lateral alae, which may be prominent in the cervical region, are typically present. They are relatively broad in Heterakis, Odontoterakis, Pseudaspidodera and Giretera- kis; are present but relatively narrow in Africana, Spinicauda, Moaciria and Strongyluris — in which genera the lateral fields are very prominent — and also in Meteterakis and Ascaridia but they are lacking in Aspidodera, Ansiruptodera, Lauroia and Paraspidodera. The nerve ring encircles the oesophagus about one third of its length posterior to the pharynx and the excretory pore opens just posterior to the nerve ring. Small sessile papillae are borne on the general body surface in Africana (few), Meteterakis (few), Spinicauda (very few), Moaciria (many) and Strongyluris (many). Such papillae are, however, frequently difficult to see and they are probably more easily seen in Strongyluris species because such species are relatively larger than the others. The Male Tail A circular pre-cloacal sucker with a cuticular rim is always present (except in Lauroia). Caudal alae are fairly common in the superfamily and, when present, never meet on the ventral surface of the body anterior to the pre-cloacal sucker. In those specimens in which the tail has contracted during fixation such a junction may appear to exist but it is clearly an artefact and is, therefore, of no use as a systematic character. Freitas (1956) lays considerable stress on this " junction " of the caudal alae, going so far as to use it to differentiate genera. I would stress that not only is this an artefact but it is not present in all the specimens of most of the species of which it is considered a diagnostic character. In the genera Heterakis, Odontoterakis and Pseudaspidodera there are, typically, eleven pairs of caudal papillae, of which seven pairs are long and support broad caudal alae. Two pairs of these papillae, which are always slim, flank the pre- cloacal sucker, four pairs lie lateral to the cloacal opening (later referred to as the para-cloacal papillae), and one pair lies posterior to the cloacal opening, roughly midway between the para-cloacals and the terminal spike of the tail. The remaining four pairs of papillae are typically sessile, two pairs flank the cloacal opening, the peri-cloacal papillae, of which one pair is pre-cloacal and one pair post-cloacal. The remaining two pairs of papillae lie at the base of the terminal spike of the tail with one pair wholly ventral and one pair lateral in position. The phasmids open just anterior to the terminal group of ventral papillae and have generally been THE SUPERFAMILY HETERAKOIDEA 7 described as papillae. In some cases additional papillae occur, in particular a pair of sessile papillae frequently occur anterior to the suctorial pairs and an additional long pair sometimes occurs immediately posterior to the para-cloacals. The tail ends posteriorly in a long, narrow, evenly pointed, non-alate terminal spike. In all four genera the pre-cloacal sucker is relatively large and there is no gubernaculum. Both Spinicauda and Africana have long narrow tails bearing many small sessile papillae and a relatively small pre-cloacal sucker. There are no caudal alae in Spinicauda while those in Africana stop posteriorly at the level of the cloacal opening and are unsupported by papillae. A gubernaculum is present in Spinicauda and a gubernacular mass in Africana (gubernacular mass is a term proposed earlier for the strongly cuticularized lining of the cloaca in the genus Meteterakis (Inglis, 19580)). The tail in Strongyluris is very short with a large pre-cloacal sucker which is directed posteriorly. The caudal alae are very broad and are typically supported by seven pairs of long stout papillae of which the largest are the three most anterior pairs, with the others becoming increasingly smaller posteriorly. There are, in addition, two pairs of ventral sessile papillae peri-cloacal in position and one similar pair just anterior to the beginning of the terminal spike. There is no gubernaculum. The tail in Moaciria is similar but with a long terminal spike and a gubernaculum. The relatively long, narrow tail in Meteterakis is characterized by a small pre- cloacal sucker and narrow caudal alae supported by, typically, three, exceptionally four, pairs of short, stout papillae of which, typically two pairs, exceptionally three pairs, lie about the level of the pre-cloacal sucker and the remaining pair is roughly at the level of the cloacal opening. There are in addition many pairs of small sessile papillae (up to about twenty) on the tail which may represent the caudal complement of the similar papillae which are found scattered generally over the surface of the body. A gubernacular mass is present in all except one (possibly two) species. The tail in Gireterakis is very similar to that of Meteterakis with the same three pairs of swollen papillae supporting the caudal alae but there are much fewer small sessile papillae and there is no gubernacular mass. The tail in the genera Aspidodera, Ansiruptodera, and Paraspidodera is so uniform that one description will cover all three. It is long and thin, tapering evenly to a point. It has no caudal alae, the pre-cloacal sucker is markedly small and there is a large number of small sessile papillae arranged in roughly parallel rows. A guber- naculum is present in all three genera. The tail in Lauroia is highly modified but may be interpreted as being derived from a tail similar to that characteristic of the previous three genera by a great reduction of all its structures. Thus the pre-cloacal sucker is represented by a slight swelling and the number of sessile caudal papillae is very small. The tail in Ascaridia is relatively stout with narrow alae supported by short stout papillae. The pre-cloacal papillae are sessile and there is no gubernaculum. The Spicules The spicules are always equal in length, non-alate, simple and identical in structure in the genera Africana, Ascaridia, Aspidodera, Ansiruptodera, Moaciria, Paraspido- ZOOL. 15, i. J§§ 8 W. G. INGLIS dera, Spinicauda and Strongyluns. They are equal in length and identical in structure, with alae in some cases, in Meteterakis and Africana. They are equal in length and slightly elaborate although identical in structure in Gireterakis while in Heterakis, Odontoterakis and Pseudaspidodera they are frequently unequal in length and may be very dissimilar in Heterakis and Pseudaspidodera but are always simple, needle-like in Odontoterakis. It may be noted here, and will be referred to later, that the general facies of the male tail in Aspidodera, Ansiruptodera, and, to a lesser extent, Paraspidodera is remarkably uniform even to the same general shape of the spicules which are, almost invariably, rather stout with squarish ends. SYSTEMATIC GROUPS WITHIN THE SUPERFAMILY This section should be read in conjunction with the discussion of morphological trends given in the next section, since the classification presented reflects the sequences discussed there. In other words this classification is largely akoluthic (sensu Inglis, 1966(2) in reflecting trends in morphological modification rather than in attempting to assess over-all resemblances or simply utilizing key characters. It does appear, however that an assessment of over-all resemblances would produce a classification the same as the present since virtually the only characters available for analysis are those of the head and the male tail and a classification based on either alone is congruent with one based on the other. That is, if only tails are classified they fall into roughly the same major groupings as do the heads if classified alone. This explanatory discussion of procedure intrudes here because in classifica- tions such as this I find it impossible to separate the discussion of the process of classification from that of the establishment and recognition of morphological trends. In fact the trends were probably recognized before the classification was developed. It is, however, easier to follow the discussion of trends and host re- lationships if a foundation is given upon which they can be discussed. As stressed above most classifications of this superfamily (family) have been mainly based on published descriptions which are in many cases incomplete or inaccurate. A reliable assessment of such descriptions, and the characters upon which to form a classification can only be made after a wide and detailed knowledge of the group under consideration has been obtained. The dangers involved in bibliotaxonomy are exemplified by the gross oversplitting proposed by Freitas (1956) with a multiplicity of new names at all levels of taxa. The classification proposed by Freitas may be considered analytical in that he has given systematic recognition to any differences he found. The classification proposed here is basically synthetic with a stressing of similarities rather than differences although, obviously analysis must precede such a process. The superfamily contains three morphologically distinct groups which differ in the forms of the lips, of the oesophagus and of the male tail. One is characterized (i) by square lips which are connected by lateral lobes, (2) by a cephalic cap (i.e. a thickening of the cuticle at the anterior end of the body) , (3) by a markedly long and narrow oesophagus which expands relatively suddenly into a small posterior tri- valvulate oesophageal bulb and (4) by males with relatively long, narrow, evenly pointed tails without alae, typically bearing many small sessile papillae, (5) a THE SUPERFAMILY HETERAKOIDEA 9 relatively small pre-cloacal sucker, (6) having relatively massive, square ended spicules without alae and (7) possessing a gubernaculum. This group I recognize as the Family Aspidoderidae. The second group is characterized by (i) rounded lips without lateral lobes, (2) no cephalic cap, (3) a relatively short stout oesophagus which merges rather slowly into a relatively large posterior tri-valvulate bulb and (4) by males in which the tail (with the exception of one genus) always carries caudal alae, (5) the spicules are frequently dissimilar and alate, (6) a gubernaculum is generally lacking, and (7) there is a tendency for the pre-cloacal sucker to be prominent and for the caudal papillae to be large. This group I treat as the Family Heterakidae. The third group is characterized by (i) massive lips without anterior cuticular flanges or (2) lateral lobes or (3) onchia at the anterior ends of the pharyngeal portions which are themselves very small, (4) the oseophagus is grossly club-shaped without a posterior oesophageal bulb or valvular apparatus, (5) the male tail carries the usual circular pre-cloacal sucker with a definite cuticular rim but, although the papillae are grouped round the region posterior to this sucker, they are generally stout and rounded without marked lateral caudal alae, (6) there are frequently files of sessile papillae running down the lateral sides of the body. This group I treat as the Family Ascaridiidae. I did not previously accept the Family Ascaridiidae as referable to the Heterakoidea (Inglis, 1958; 19586) and argued that it was Ascaridoid. In this I was clearly wrong. I have now studied the Ascaridoidea (Inglis, 1965, 1965(2) and am convinced by the overwhelming weight of evidence that the Ascaridiidae are heterakoids and that their resemblances to the Ascaridoidea are simply due to the convergent development of a massive body size. The form of the male tail and the presence of paired nuclei in the ventral oesophageal glands are clear features of resemblance with the Heterakoidea and rule against ascaridoid affinities, as does the life history. Equally significant is the structure of the head which in the Ascaridiidae is heter- akoid in plan and shows no point of resemblance with that in the Ascaridoidea, other than gross size. The lips in the Ascaridoidea are operated by a septum of three systems of non-contractile fibres (Inglis, 1965) and the cheilorhabdion (i.e. the cuticle lining the inner surface of the cheilostome: Inglis, 1966, 1967) is never markedly sclerotized. In Ascaridiidae the conditions are quite different. The cheilorhabdion is modified into a definite heavily sclerotized region in each lip while the fibre system, which is present, is not organized as three subsiduary systems but is sparse except in the inter-labial regions from which it fans out into each contiguous lip. Exactly the same occurs in the heterakids and aspidoderids. Further the arcade system and associated cells bears no resemblance to that of the Ascaridoidea (Hartwich, 1957)- I continue to treat the Aspidoderidae as a distinct family in spite of Chabaud (1957; 1965) who considers it a subfamily of the Heterakidae. I did at one time agree with Chabaud over this point but further consideration leads me to conclude that to do this masks the many and manifest differences between the aspidoderids and the heterakids. The forms included within the Aspidoderidae differ in so many features from those referred to the Heterakidae, they form such a specialized, io W. G. INGLIS geographically restricted group and reach such a degree of independent specializa- tion in a form such as Lauroia, that I look on them as being as distinct from the Heterakidae as are the Ascaridiidae. Lauroia is a particularly aberrant genus at first sight and this has led to its being referred to a unique family within a different Superfamily by Skrjabin et al. (1961) but I have studied some of the syntype series of L. travassosi Proenga, 1938 (type species of the genus) and L. intermedia Caballero, 1955. The lips have the inter- connecting lobes typical of the aspidoderids and this may be considered a specialized form of the head found in Paraspidodera. Further the reduction in the structures of the male tail is so obviously secondary, and has involved all structures, that little stress can be laid on this alone. But in L, trinidadensis Cameron, 1939 there is a definite small bump anterior to the pre-cloacal opening which can be considered to be the reduced remnants of a pre-cloacal sucker while the pre-cloacal modification in L. intermedia is even more obviously sucker-like. Family HETERAKIDAE The family Heterakidae, as defined here, contains three distinct morphological groups. The first is characterized by (i) a short, rather stout oesophagus, (2) rather large lips, (3) interlabia (or at least structures which may be interpreted as homo- logous with inter-labia), (4) a rather large pre-cloacal sucker, and (5) very broad caudal alae supported by long, narrow papillae. The second is characterized by (i) rather small lips which are not set-off from the body, (2) by a relatively small pre- cloacal sucker, (3) by a flap over the vulvar opening, (4) an excretory pore which leads into a lobulate excretory vesicle and (5) by rather narrow caudal alae on the male tail which are supported by three rather small fleshy papillae. The third group is much less homogenous than the others but is characterized by (i) relatively small lips (although the anterior cuticular flange may be large) which are clearly off-set from the remainder of the body by distinct shoulders, (2) by prominent lateral lines, (3) no trace of inter-labia, (4) a relatively long, and narrow oesophagus, and (5) spicules which are always identical and relatively simple in structure. The structure of the male tail is more variable and will be discussed more fully below. These three groups are treated as subfamilies, thus: (i) Heterakinae, (2) Meteter- akinae and (3) Spinicaudinae. Family ASPIDODERIDAE The family Aspidoderidae consists of two distinct groups in one of which there are inter-labia modified as cordons while in the other there is no trace of inter-labia or cordons. These two groups are treated as subfamilies: Aspidoderinae and Lauroiinae respectively. Family ASCARIDIIDAE The family Ascaridiidae contains only one genus, Ascaridia. These taxa are diagnosed formally later. THE SUPERFAMILY HETERAKOIDEA n MORPHOLOGICAL AND EVOLUTIONARY TRENDS Species of the superfamily occur in all vertebrate groups from amphibia to birds with a few representatives in mammals, except in South American where the Family Aspidoderidae is restricted to Mammals (marsupials, edentates and histricomorph rodents). In view of the great overall similarities between the component taxa it appears reasonable to consider that the superfamily represents a mono-phyletic group. Within the family Heterakidae there is a sequence along which is a tendency towards a reduction in the number of caudal papillae, with an increase in their size and the associated appearance and increase in the size of the caudal alae. Concomitantly there is a reduction and finally a loss of the gubernaculum, an increase in the relative complexity of the spicules, an increase in the relative size of the pre- cloacal sucker and a loss of papillae on the general body surface with the development of wide lateral alae. There is also a tendency towards the development of cephalic cordons, by the modification of inter-labia associated with the lips, or by the modification of the lips alone. That these trends have developed in the order described is supported by the host distribution of the parasites. The genera Spinicauda, Africana and Meteterakis, all of which are characterized by papillae on the general surface, relatively large numbers of caudal papillae, a gubernaculum or gubernacular mass, poorly developed caudal alae (when such alae are present), a relatively small pre-cloacal sucker, equal and identical spicules and no interlabia, are restricted to reptiles and amphibia. The genera Heterakis, Pseudaspidodera and Odonototerakis, on the other hand, are characterized by no papillae on the body surface, relatively small numbers of elongate caudal papillae, no gubernaculum or gubernacular mass, well developed caudal alae, a relatively large pre-cloacal sucker, spicules which are frequently unequal and dissimilar and almost invariably complex, with inter-labia modified as cordons in Pseudaspidodera and Odontoterakis and are restricted, typically, to birds. In- dependently the genus Gireterakis has developed straight cordons, has no papillae on the general body surface, has very elaborate spicules and is known only from the mammal genus Hystrix. It is now possible to suggest that the form ancestral to the superfamily possessed most of the following characters, although not necessarily all, (i) an anterior cuticular flange which did not project anterior to the mass of the lips; (2) no interlabia; (3) papillae on the general body surface; (4) a relatively long oesophagus with a small posterior oesophageal bulb; (5) a relatively small pre-cloacal sucker on the male tail; (6) no caudal alae on the male tail; (7) equal and identical spicules; (8) many small, sessile caudal papillae on a long, narrow male tail. The genus Spinicauda possesses a combination of characters almost identical with those postulated above for an ancestral form, diverging only in that the anterior cuticular flange of the lips projects slightly anterior to the main mass of the lips, and may be considered as containing the most primitive species of the superfamily (i.e. the most generalized). Non morphological supporting evidence is given by its world-wide occurrance within tropical and subtropical regions and its restriction to reptiles. The characters of the male tail are almost identical with those 12 W. G. INGLIS postulated for an ancestral form and as it is in just those characters that Spinicauda resembles the Aspidoderidae I consider the form of the male tail in that family to be relatively unspecialized. The structure of the head in Spinicauda fits into an almost perfect series of in- creasing elaboration with Africana, in which the anterior cuticular flange is wholly within the limits of the lips, Spinicauda in which the flanges projects slightly anterior to the main mass of the lips and Strongyluris where the flange forms the major part of the lips and dominates the lip-mass completely (Inglis, 1957). It may be pointed out that Africana is a close rival to Spinicauda for the position of the most primitive genus and may be so considered without affecting the argument presented here. The connection between the genera Spinicauda and Strongyluris is accentuated by the forms referred to the genus Moaciria (formerly treated as a subgenus of Spinicauda}. In this latter genus the male tail is intermediate in form between the long narrow tail of Spinicauda and the truncate tail, with broad alae, of Strongyluris. This transitional form of the male tail is associated with a head which is identical in structure with that of Spinicauda. The genus Africana appears to be atypical in being the only genus of the super- family in which the caudal alae are not supported by caudal papillae, although this could be interpreted as an intermediate stage between the forms without alae and those with alae supported by papillae. The similarities between Spinicauda, Africana, Moaciria and Strongyluris are so great and, in addition, can be arranged in such a distinct morphological sequence with fairly clear intermediates culminating in the specialized form of Strongyluris that they are treated as a second evolutionary sequence recognized as the subfamily Spinicaudinae. The remainder of the family Heterakidae falls into two distinct groups, recognized as subfamilies, Heterakinae and Meteterakinae, of which the latter appears to be the result of radiation within a restricted geographical area. The species referred to Meteterakis may be considered more primitive than that in Gireterakis in possessing a gubernacular mass and in having papillae in the general body surface. M. tri- aculeatus without a gubernacular mass being to that extent intermediate between the two genera. The subfamily Heterakinae contains the species which are parasitic in birds and are the most highly evolved parasites, morphologically, within the family. Re- presentatives of the subfamily occur in ground feeding, grain-eating birds throughout the world but have become sufficiently distinct in two geographical regions to warrant the recognition of two genera, Odontoterakis in South American tinamous and Pseudaspidodera in Indian peafowls. The inclusion of Pseudaspidodera within the Family Aspidoderidae by Skrjabin et al. 1961 is completely unacceptable since not only does it differ from the typical aspidoderids in the structure of the head (Inglis, 1957) but also differs in the form of the male tail with its broad, typically heterakid caudal alae, and the form of the dissimilar spicules. The resemblance must in fact be dismissed as the crudest and most superficial convergence of one morphological feature and if Pseudaspidodera is grouped with the aspidoderids so must the genus Odontoterakis, which Skrjabin et al. do not do. THE SUPERFAMILY HETERAKOIDEA 13 It is also worth noting that the species referred to the genus Heterakis fall into two groups characterized by the structure of the spicules. The groups are not those in which the spicules are equal or unequal (as recognized by many authors, Lopez- Neyra (1947), Skrjabin & Schikhobalova (1947), Freitas (1956) and most recently Skrjabin et al. (1961)), but are based on whether the spicules are identical in gross structure although they can be of different lengths (e.i. non-alate without elaborate tips) or are dissimilar (i.e. one, the left, usually bears broad alae and has an elaborate tip while the right is simple and needle-like). This difference is also geographic since the first group is most frequent in the Americas while the other is most characteristic of the European and Asiatic parasites. This difference is also re- flected by the structure of the spicules in the neotropical genus Odontoterakis, in which the spicules are never alate and never have elaborate tips. The oriental genus Pseudaspidodera in contrast is characterized by dissimilar spicules in which the left spicule bears broad alae while the right is needle-like. The family Aspidoderidae represents the radiation of a stock isolated in South American mammals. In this group most of the diversity is confined to the anterior end of the body, and superficially parallels the conditions in Pseudaspidodera and Odontoterakis but there are marked differences in the details. Thus, in the heterakids the cordons, when present, never remain the same width throughout their length even when they anastomose (accepting that they do anastomose in Pseudaspidodera spinosa, see below) . In the family Aspidoderidae the cordons always remain the same width along their full length. The Aspidoderidae appear to have divided into two lines: one in which inter-labia modified as cordons are present and one in which inter-labia have not appeared. Whether the species referred to Lauroia have been derived from the forms grouped in Paraspidodera is an open question but in view of the similarities of the head structures the two genera are classified in the same subfamily, Lauroiinae. The genus Ascaridia is clearly heterakoid in affinity but appears to represent a group which diverged fairly early and probably shows its greatest affinities to the Spinicaudinae. GEOGRAPHICAL AND HOST DISTRIBUTION The superfamily as a whole is cosmopolitan in distribution and occurs in all vertebrate hosts groups from amphibia to mammals and birds, but it is in the latter group that it is most frequent and wide-spread. The distribution of the major subdivisions of the superfamily is interesting since it reflects a combination of host and geographical restrictions. Thus, the subfamily Spinicaudinae is cosmopolitan with forms occurring in all major geographical regions although restricted to reptiles and amphibia. Even the genera within the subfamily tend to be wide-spread. Spinicauda occurs in South America, Africa, Madagascar and Australia, with one species in each area, while Strongyluris is extremely widely spread in tropical and subtropical regions and is represented by a much larger number of species. In contrast Africana is restricted to Africa, and Moaciria to South America, Africa and Madagascar. Strongyluris appears to represent the culmination of the Spinicaudinae i4 W. G. INGLIS and is as successful as a group of parasites in reptiles as the genus Heterakis is in birds. The subfamily Heterakinae is equally wide-spread but is largely restricted to ground-feeding, grain-eating birds i.e. Galliformes (one species in mammals; H. spalacis and H. macrospiculum are not typical of the genus Heterakis and are best considered incertae sedis although the latter shows similarities to Africana, particularly in the structure of the male tail and spicules.) The genus Heterakis is cosmopolitan and ubiquitous but the other genera of the subfamily are geographically restricted with Odontoterakis in South America and Pseudaspidodera in the Indian region. The family Aspidoderidae is the only group of the superfamily which occurs widely in mammals and it also is geographically restricted to South America while Ascaridia, although cosmopolitan, is most commonly reported from members of the Columbi- formes and Psittaciformes although it is not uncommon in the Galliformes. The patterns suggest that the evolution of the superfamily has been due to a combination of (i) intra-host restriction to the caecum, (2) host restriction to ground-feeding grain-eating birds and (3) geographical restriction. There is no evidence to suggest any close host : parasite parallelism. The impression is rather of a group which has evolved and expanded to occupy all the space available to it within the constrains of the ecology and feeding habits of the hosts (see Inglis, 1965^. Thus the members of the superfamily exist in a specialized, selected, locality within the host, in all birds in which they occur. But they only occur in a restricted range of hosts with similar feeding habits. Then the various smaller taxa are largely determined geographically. Nevertheless the Aspidoderidae represent a special case in that they are not only geographically isolated but also occur in a long isolated group of mammalian hosts, the South American marsupials and the ground-feeding edentates. It should be noted that they do not occur in the ant-eaters or the tree- dwelling sloths. It is interesting to wonder, although impossible to know, if they continue to exist in South America because there is no competition with other nematode parasites in the large intestine where they occur or whether they represent a new group of parasites which has replaced some other group. In other words are they a remnant group or are they the result of a specialized radiation within an isolated geographical and host locality? Be that as it may the Heterakoidea is clearly an old group which is supremely successful within the hosts groups in which it occurs. SYSTEMATIC HISTORY Railliet and Henry (1914) considered that the species of the family Ascaridae for which they had created a new subfamily Heterakinae in 1912 were sufficiently distinct to warrant their further separation into a distinct family, Heterakidae. This family was diagnosed largely on the presence of a pre-cloacal sucker on the male tail and included the genera Heterakis, Ascaridia, Aspidodera, Cissophylus and Subulura, with Strongyluris as a subgenus of Heterakis and Oxynema as a subgenus of Subulura. No groupings higher than genera were proposed. THE SUPERFAMILY HETERAKOIDEA 15 Travassos (1920) introduced two new genera, Spinicauda and Africana, and later in the same year (Travassos, 19200) he divided the family Heterakidae into three subfamilies : Heterakinae, with Heterakis, Ganguleterakis , Aspidodera, Para- spidodera and Gireterakis ; Spinicaudinae nov. for Spinicauda, Africana and Strongyluris ; Subulurinae for Subulura, Heteroxynema, Oxynema and Numidica. The last subfamily differed from both the others in having in the males an elongate pre-cloacal sucker without a definite rim while the other two had a pre-cloacal sucker with a definite cuticular rim. The diagnostic differences between the other two subfamilies were vague and appear to have been mainly the restriction of the species of the first to warm-blooded hosts and those of the second to cold-blooded hosts. Yorke & Maplestone (1926) considered the subfamily Subulurinae a family and did not recognize the subfamily Spinicaudinae, thus leaving the family Heterakidae with no groups higher than genera. Baylis & Daubney, slightly later in the same year (1926), still treated the subulurids as a subfamily but this was not generally accepted. No further radical changes were proposed in the classification of the Heterakidae until Skrjabin and Schikhobalova (1947) recognized two subfamilies: Heterakinae and Aspidoderinae nov. This grouping was largely based on published descriptions and the subfamilies were considered distinct almost wholly on the presence or absence of cephalic cordons. In 1957 I described the comparative anatomy of the head in the family and argued that the classification of Skrjabin and Schikhobalova was unnatural because cephalic cordons appeared to have been developed in- dependently three times. I then proposed that the genera Aspidodera, Ansirupto- dera, Sexansodera and Paraspidodera be placed in a separate family, Aspidoderidae with two subfamilies : Aspidoderinae for the first three of the above genera and Paraspidoderinae nov. for the fourth. Independently Freitas (1956) reviewed the family and proposed four new families, four new subfamilies and five new genera (four with one species each and one with two species). This classification was, and still is, unacceptable. There are five families of which two have only one species, there are eight subfamilies of which three have only one genus each, there are twenty- two genera of which eleven have only one species each and four have only two species each. This classification was based largely on published descriptions and every error of observation and every faulty description appears to have been recognized by a distinct genus, at least, and in some cases a subfamily or even family. Then Chabaud (1957), in considering the classification of the suborder Ascaridina introduced a new superfamily Heterakoidea which he considered to contain two families and four subfamilies thus: Heterakidae, with four subfamilies Heterakinae (with sixteen genera or subgenera), Aspidoderinae (with four genera or subgenera), Lauroiinae (with one genus) and Schneidernematinae (with one genus); and the family Ascaridiidae with one genus, Ascaridia. This classification is largely that of Inglis (1957) and Freitas (1956) re-arranged. In 1958 I proposed the classification which I am justifying here and will not discuss it now accept to draw attention to the fact that Schneidernema and Morgascaridia (a genus I proposed for Paraspidodera sellsi, Morgan, 1928 : Inglis, 19586) were 16 W. G. INGLIS removed from the Heterakidae first to the Ascaridiidae (Inglis, 19586) and later to the Superfamily Seuratoidea (Inglis & Chabaud, 1958) as members of a distinct sub- family Schneidernematinae. Most recently Skrjabin, Schikhobalova & Lagodovskaja (1961) have presented a re-arrangement of the Heterakoidea, with four families, thus : Heterakidae : Heter- akinae (Heterakis, Ganguletemkis and Odontoterakis); Meteterakinae (Meteterakis; and Gireterakis}. Aspidoderidae : Aspidoderinae (Aspidodera, Ansiruptodera and Sexansodera) Spinaspidoderinae (Spinaspidodera and Pseudaspidodera}. Spinicaudidae : Spinicaudinae (Spinicauda, Africana and Paraspidodera} ; Preter- akinae (Preterakis). Strongyluridae : Strongylurinae (Strongyluris and Moaciria}. In addition they refer Lauroia to a distinct family within the Cosmocercoidea and introduce a new genus, Bellaplectana for Aplectana pharyngeodentata Belle, 1957. It should, perhaps, be pointed out that Yamaguti (1961) has grouped the Heter- akoidea in yet another way but this does not warrant serious consideration except to dismiss the new genus Cheloniheterakis. This genus was introduced for two old and inadequately described species from Testudo. No other heterakids are known from this host genus in which members of the family Kathlaniidae are fairly common parasites. As the two species referred to Cheloniheterakis possess circular precloacal suckers, which are common in Kathlaniids it is more probable they are referable to that family. Be that as it may the two species are species dubia and the genus must be ignored as unrecognizable. The Skrjabin et al. classification does warrant consideration and I will give my arguments against accepting it. Part of it can be dismissed quickly and easily as due to classifying errors of description. The arguments against the genus Bella- plectana have been adumbrated at the beginning of this paper but to recapitulate briefly the so-called teeth in the head, upon which the genus is based, do not exist they are simply the retracted lips in poorly fixed specimens. The genus Preterakis and the associated subfamily Preterakinae must disappear since the fusion of the caudal alae on the ventral surface of the male body is a fixation artefact and does not occur in all the specimens of the type series. The remaining arguments are more expressions of opinion and of assessing the weight of the morphological and other evidence, thus : The reference of the aspidoderids and Pseudaspidodera to the same family is a continuation of the position taken by Skrjabin & Schikhobalova (1947) but is still un-acceptable and morphologically indefensable. If this grouping is advocated there is no real reason for not referring Odontoterakis to the same family, but this genus is referred by Skrjabin et al. (1961) to the Heterakidae. I stress, and repeat what I wrote before, the only resemblance between the Aspidoderidae and Pseud- aspidodera is in the common possession of structures called by the same name, i.e. cordons. The family Aspidoderidae (sensu Skrjabin et al.} is otherwise characterized (in addition to cordons) by the presence of a cephalic cap, inter-connecting lobes to the lips, a long slim oesophagus and, in the male, is further characterized by equal and identical square-ended spicules, no caudal alae and the presence of many sessile THE SUPERFAMILY HETERAKOIDEA 17 papillae. In all these respects Pseudaspidodera differs from the Aspidoderids and in all the corresponding features is identical with the genera referred by Skrjabin et at. to the subfamily Heterakinae. Equally the reference of Paraspidodera to the Spinicaudinae is unacceptable since its only resemblance to the other genera of that subfamily is in lacking caudal alae and cephalic cordons. In all other respects, the structure of the head, the oesopha- gus, and the male's tail and spicules it is indistinguishable from the forms referred to the Aspidoderidae. The reference of Lauroia to the superfamily Cosmocercoidea is equally contradicated by the morphology. The structure of the anterior end of the body is very similar to that of the Aspidoderidae and the remnants of a pre- cloacal sucker are clearly present in at least two species. The separation of Strongyluris and Moaciria (which I now accept as a distinct genus) into a separate family is equally unacceptable since Strongyluris so clearly represents the end of a morphological sequence running from Spinicauda to Moaciria to Strongyluris. The structure of the head in Moaciria is identical with that in Spinicauda while the male tail forms an almost perfect intermediate between that genus and Strongyluris. I therefore propose the following classification for the Superfamily Heterakoidea. ANNOTATED CLASSIFICATION OF THE SUPERFAMILY On the basis of the arguments presented above the following classification is proposed. Notes are appendaged to each group to draw attention to points of interest and to explain some of the names used. In these notes some of the argument given above is briefly repeated so that this section is more or less complete in itself. HETERAKOIDEA (Ralliet & Henry, 1912) Chabaud, 1957 Ascaridida : three large distinct lips, one dorsal with two double papillae, two ventral with a double ventral papilla and a single lateral papilla each : lining of mouth cavity (i.e. cheilorhabdion) sclerotized and forming a flange along the anterior edge of each lip in all forms except Ascaridia ; anterior end of oesophagus divided into three projecting lobes, one to each lip, except in Ascaridia ; nuclei of ventral oesophageal glands double ; no intestinal or oesophageal appendices. MALE : circular pre-cloacal sucker with a definite cuticular rim ; two spicules. FEMALE : eggs not embryonate in utero, thick-shelled ; vulva opening about middle of body. LIFE HISTORY : Direct, with or without a migration within the final host. HOSTS : Restricted to the intestine (more specifically to the caecum in birds and the large intestine in mammals) of ground feeding birds and mammals. In amphibia (anurans) and reptiles (saurians) restricted to terrestrial feeders. GEOGRAPHICAL DISTRIBUTION : Cosmopolitan as a superfamily. i8 W. G. INGLIS HETERAKIDAE Railliet & Henry, 1912 Heterakoidea : lips rounded, not connected by lateral lobes ; cordons when present poorly developed and not remaining the same width throughout their length ; oesophagus relatively short and stout. MALES : caudal alae typically present ; number of caudal papillae relatively low ; spicules may be unequal and dissimilar ; gubernaculum generally reduced or absent ; tail generally relatively short. HOSTS : amphibia, reptiles and (mainly) birds. One species in mammals. DISTRIBUTION : Cosmopolitan as a family. TYPE GENUS : Heterakis Dujardin, 1845. HETERAKINAE Railliet & Henry, 1912 Heterakidae : lips not off-set from body ; anterior cuticular flange of lips generally not projecting beyond main lip mass ; inter-labia, or their homologues, present. MALE : precloacal sucker relatively large ; caudal alae broad, supported by long, narrow papillae ; gubernaculum lacking ; spicules frequently dissimilar. HOSTS : mainly birds, one exceptional species in mammals (rodents). DISTRIBUTION : Cosmopolitan. The subfamily is a homogeneous group in which three subgroups can be recog- nized. Odontoterakis is South America, Pseudaspidodera in India and Heterakis which is cosmopolitan. The latter genus is characterized by lacking definite cephalic cordons such as characterize the other two genera. But there is a tendency for there to be marked " bumps " in the inter-labial spaces which may be considered as precursors of the inter-labia which are modified as cordons in the other two genera. It is worth pointing out that what are here called cordons are identical in form and origin with the so-called labial grooves of some members of the Ascari- doidea, e.g. Porrocaecum and Multicaecum. These labial grooves are also formed from two sources, one the inter-labia and the other the lips, and also consist of open grooves. The morphological differences between Pseudaspidodera and Odontoterakis tend to be slight but in the former genus, in addition to the cordons being more strongly developed, the left spicule is always shorter than the right, is always broadly alate while the right is slim and needle-like, and generally has an elaborate posterior tip. In contrast the spicules in Odontoterakis are always identical in structure, never bear alae, and always end posteriorly in simple points. I therefore interpret the two groups as having developed in specialized host groups within isolated geographical areas since Pseudaspidodera appears to have arisen in peafowls in India and Odontoterakis to have arisen in tinamous in South America. Both host groups being typically ground dwelling forms with poor powers of locomotion. THE SUPERFAMILY HETERAKOIDEA 19 Heterakis Dujardin, 1845 Heterakinae : lips without definite inter-labia or cordons. MALES : spicules frequently dissimilar but may be equal and identical. TYPE SPECIES : Ascaris gallinarum Schrank, 1788. HOST AND GEOGRAPHICAL DISTRIBUTION : ground feeding birds, mainly Galli- formes, throughout the world. Several attempts have been made to divide this genus into two on the basis of the relative lengths of the spicules, species with equal spicules to one genus and with unequal to another. Both Lopez-Neyra (1947) and Skrjabin & Schikhobalova (1947^) independently attempted to do this, using the name Ganguleterakis for species with equal spicules. Madsen (1950), in a generally excellent and outstanding publication, commented upon this unfavourably pointing out the great difficulty in many cases in deciding whether the spicules should be called equal or unequal. Freitas (1956) attempted to over-come this difficulty by defining his two subgenera, Heterakis and Raillietakis, so that unequal spicules were considered to be those in which the difference in length was at least one third the length of the shorter spicules. Even with this qualification it is difficult to imagine that equality or its lack is a sufficiently fundamental character upon which to recognize genera or subgenera. Skrjabin et al. (1961) continue to recognize two groups on just this difference in the lengths of the spicules but it is still unacceptable. It is so very clearly a biblio- graphical key character resorted to in an attempt to reduce the number of species within each genus and can only have been used because it is one of the few characters which can be determined from literature rather than specimens. The structure and relative lengths of the spicules, nevertheless afford good characters in the delimitation of species within the genus used in conjunction with the number of caudal papillae, the relative size and position of the pre-cloacal sucker and the relative length of the male tail. Many authors have laid stress on the position of various caudal papillae, e.g. the number of pre-cloacal papillae, but as Maplestone (1932) and Madsen (1950), among others, have pointed out, the value of these characters is very limited. The typical arrangement of the caudal papillae is described above but it should be noted that it is relatively common for an additional pair to occur between the para-cloaca and suctorial groups or for one of the pairs of para-cloacals to be missing. There is also a tendency, in the species occurring in South America, for a pair of long papillae to be present just anterior to the peri-cloacal group, a pair which I consider homologous with the anterior pair of peri-cloacals since in specimens on which the former pair of papillae are present the latter pair is consistently absent. Both Maplestone (1932) and Madsen (1950) have pointed out that it is, in most cases, impossible to find characters by which the females of Heterakis may be distinguished. This applies throughout the entire superfamily. I have been unable to find any character, or combination of characters by which females alone can be identified. In spite of my criticism of the attempts to divide the genus Heterakis on the basis 20 W. G. INGLIS of the relative lengths of the spicules two subgroups can be recognized : (i) one in which the spicules are identical in structure and never bear alae and (2) one in which the spicules are dissimilar with the right slim and needle-like and the left broadly alate. In this latter group the posterior end of the left spicule is frequently modified into a hook or barb. In both groups the spicules may be unequal in length but in the first group when there is any modification on the posterior end it is always identical in both spicules. Thus on this basis H. gallinarum and H. isolonche, which only differ in the relative lengths of the spicules, fall together while on the basis of un- equality they are completely separated. Lane (1914) introduced the generic name Ganguleterakis for the species generally known as H. spumosa Schneider, 1866. There is a case for recognizing this genus as it contains the only species of Heterakis from mammals (H. spalacis and H. longi- spiculum incertae sedis] but, except for a reduction in the number of caudal papillae, in which it is paralleled by H. psophiae Travassos, 1913, it is a typical Heterakis. I do not, therefore, recognize this genus. Odontoterakis Skrjabin & Schikhobalova, 19470: Heterakinae : interlabia modified as non-recurrent cordons. MALES : spicules simple non-alate and identical in structure. TYPE SPECIES : Heterakis crypturi Baylis, 1944. HOST AND GEOGRAPHICAL DISTRIBUTION : Birds, mainly tinamous, in South America. Note : A restudy of the types of Heterakis interlabiata Ortlepp, 1923 has convinced me that Mendonga (1953) is correct in treating it as a synonym of H. isolonche. Pseudaspidodera Baylis & Daubney, 1922 Heterakinae : inter-labia modified as recurrent cordons. MALES : spicules dissimilar and unequal, right slim and needle-like, left with broad alae. TYPE SPECIES : Pseudaspidodera pavonis Baylis & Daubney, 1922. HOST and GEOGRAPHICAL DISTRIBUTION : Phasianid birds, India. P. jnanendrae Chakravarty, 1938 was described as having anastomosing cordons but in the type, a female, the head is very contracted and it is probable that the apparent fusion is an artefact. P. spinosa Maplestone, 1932 was described as having spines in the cordons which anastomose. As a result a genus Spinaspidodera was proposed by Skrjabin & Schikhobalova (1947) for it. There is some justification for this but in view of the small number of species involved and as Maplestone's species has never been rediscovered and the types are lost I prefer not to recognize this genus. It is worth noting that the cordons in this case, if they do definitely anastomose, vary in width along their length and do not remain the same width as in the members of the Aspidoderidae. THE SUPERFAMILY HETERAKOIDEA 21 METETERAKINAE Inglis, 19580 Heterakidae : head with three rounded lips, without interlabia ; oesophagus relatively long with a small postoesophageal bulb ; excretory pore opening into a large lobulate vesicle ; lateral alae running full length of body. MALE : spicules equal in length ; caudal alae present, supported by three pairs, exceptionally four, of large fleshy papillae — one pair lateral to the cloacal opening and typically two pairs, exceptionally three, lateral to the small pre-cloacal sucker ; a large number generally about twenty pairs, of small sessile papillae also present. FEMALE : vulvar opening covered by a flap developed from the anterior lip ; tail relatively very long. TYPE GENUS : Meteterakis Karve, 1930. HOST AND GEOGRAPHICAL DISTRIBUTION : Amphibia, reptiles and mammals (Hystrix} in the oriental region (s.l. : India, Ceylon, China, Japan, East Indies, Burma) . Meteterakis Karve, 1930 Meteterakinae : head without cordons ; body generally with many small sessile papillae scattered over surface. MALES : spicules identical ; indefinite gubernacular mass developed from the walls of the cloaca — except in M. triaculeata ; posterior lip of cloacal opening covered by small granulations. TYPE SPECIES : Meteterakis govindi Karve, 1930. HOST AND GEOGRAPHICAL DISTRIBUTION : amphibia and reptiles, as for sub- family. Gireterakis Lane, 1917 Meteterakinae : anterior cuticular flange of lips not projecting anterior to main lip mass ; three straight, simple cordons, one arising from each inter-labial space. MALE : number of sessile caudal papillae relatively small ; spicules elaborate ; no gubernacular mass present. TYPE SPECIES : Gireterakis girardi Lane, 1917. HOST AND GEOGRAPHICAL DISTRIBUTION i Hystrix species in India. SPINICAUDINAE Travassos, 1920 Heterakidae : lips off-set from body ; no inter-labia or cordons ; lateral fields prominent ; papillae on body. MALE : spicules equal, identical and simple ; papillae do not support caudal alae when alae present except when the tail is short and the pre-cloacal sucker is directed posteriorly. TYPE GENUS : Spinicauda Travassos, 1920. HOST AND GEOGRAPHICAL DISTRIBUTION : Amphibia and reptiles ; cosmopolitan in the tropics and sub-tropics. 22 W. G. INGLIS This subfamily is morphologically the least uniform, and therefore the most difficult to diagnoze, in the superfamily Heterakoidea. This is mainly because Spinicauda is morphologically generalized while Strongyluris is highly modified. A trend in the modification of the head is apparent from Africana, with a simple anterior cuticular flange, to Spinicauda and Moaciria in which the flange projects slightly beyond the lip mass to Strongyluris in which the flange forms the major part of the lip. Moaciria forms an almost perfect intermediate stage between Spinicauda and Strongyluris since it resembles Spinicauda in the presence of a gubernaculum and in having the same head form and resembles Strongyluris in the posteriorly directed pre-cloacal sucker and the presence of caudal alae (narrow) supported by longish papillae. I treated Moaciria as a subgenus of Spinicauda in 1958 but now prefer to treat it as a distinct genus. Spinicauda Travassos, 19200 Spinicaudinae : anterior cuticular flange projects slightly beyond the main mass of the lips. MALE : tail relatively long, tapering evenly to a point ; pre-cloacal sucker relatively small ; gubernaculum present ; caudal papillae small and sessile. TYPE SPECIES : Ascaris Spinicauda Rudolphi, 1819. HOST AND GEOGRAPHICAL DISTRIBUTION : Reptiles in South America, Africa, Madagascar and Australia (Queensland), i.e. tropical and subtropical. Africana Travassos, 1920 Spinicaudinae : anterior cuticular flange not projecting anterior to lips ; lateral alae present. MALES : pre-cloacal sucker small ; spicules equal and identical, alate ; guberna- cular mass present ; small sessile papillae on tail ; caudal alae not supported by papillae ; caudal alae stop about level of cloacal opening. TYPE SPECIES : Heterakis africana Gendre, 1909. HOST AND GEOGRAPHICAL DISTRIBUTION : Amphibia and reptiles in tropical and subtropical Africa. Moaciria Freitas, 19560 Spinicaudinae : anterior cuticular flange projecting slightly anterior to mass of lips. MALE : tail relatively long with relatively few stoutish papillae supporting narrow caudal alae ; posteriorly directed pre-cloacal sucker ; gubernaculum present. TYPE SPECIES : Moaciria alvarengai Freitas, 19560. HOST AND GEOGRAPHICAL DISTRIBUTION : Reptiles in South America, Africa and Madagascar. THE SUPERFAMILY HETERAKOIDEA 23 Strongyluris Mueller, 1894 Spinicaudinae : anterior cuticular flange projecting beyond main lip mass so that it forms major portion of lip ; sessile papillae scattered over body surface. MALE : posterior end obliquely truncate ; pre-cloacal sucker relatively large and directed posteriorly ; caudal alae well developed, supported by rather long, stout caudal papillae ; spicules equal, without alae ; no gubernaculum or gubernacular mass present. TYPE SPECIES : Strongyluris brevicaudata Mueller, 1894. HOST AND GEOGRAPHICAL DISTRIBUTION : Mainly reptiles, rarely amphibians, cosmopolitan within the tropics and subtropics. ASPIDODERIDAE Skrjabin & Schikhobalova, 1947 Heterakoidea : lips square connected by lateral lobes ; cuticle at anterior end of body thickened to form a cephalic cap ; oesophagus long and narrow with a small posterior oesophageal bulb ; cordons, when present, the same width along their lengths. MALES : no caudal alae ; gubernaculum generally present ; spicules equal in length and identical in structure, generally massive ; generally many small sessile papillae on long, narrow tail ; caudal papillae all about the same size. TYPE GENUS : Aspidodera Railliet & Henry, 1912. HOSTS AND GEOGRAPHICAL DISTRIBUTION : Marsupials, edentates and occasionally hystricomorphs in South America and related areas. ASPIDODERINAE Skrjabin & Schikhobalova, 1947 Aspidoderidae: cephalic cordons present, lateral lobes of lips complex. TYPE GENUS : Aspidodera Railliet & Henry, 1912. DISTRIBUTION : As family. Aspidodera Railliet & Henry, 1912 Aspidoderinae : cephalic cordons recurrent and anastomosing. TYPE SPECIES: Aspidodera scoleciformes Diesing, 1851. Ansiruptodera Skrjabin & Schikhobalova, 1947 Aspidoderinae : cephalic cordons not anastomosing. TYPE SPECIES : Aspidodera ansiruptodera Proenga, 1937. LAUROIINAE Skrjabin & Schikhobalova, 1951 Aspidoderidae : cephalic cordons lacking ; lateral lobes of lips simple. TYPE GENUS : Lauroia Proensa, 1938. 24 W. G. INGLIS HOST AND GEOGRAPHICAL DISTRIBUTION : Edentates and hystricomorphs, as the family. Lauroia Proenca, 1938 Lauroiinae : cephalic cap modified into three " plates " which are slightly undercut posteriorly. This modification can be interpreted as due to the develop- ment of cordons which do not involve inter-labia as in Girder akis. MALE : posterior end simple without a gubernaculum, relatively few papillae, and a reduced pre-cloacal sucker. TYPE SPECIES : Lauroia travassosi Proenga, 1938. Paraspidodera Travassos, 1914 Lauroiinae : cephalic cap not modified as " plates ". MALE: posterior end relatively complex, with a gubernaculum; relatively large number of caudal papillae and a well developed pre-cloacal sucker. TYPE SPECIES : Paraspidodera uncinata Rudolphi, 1819. ASCARIDIIDAE Travassos, 1919 Heterakoidea : lips large and stout ; no distinct anterior cuticular flange ; oesophagus club-shaped without a posterior bulb. MALE : caudal alae narrow supported by short stout papillae. TYPE GENUS : Ascaridia Dujardin, 1845. HOST AND GEOGRAPHICAL DISTRIBUTION : Birds, cosmopolitan with rare species in mammals. KEY TO THE GENERA OF THE SUPERFAMILY HETERAKOIDEA (1) (2) Without a tri-valvulate posterior oesophageal bulb . . . ASCARIDIA (p. 24) (2) (i) With a tri-valvulate posterior oesophageal bulb . (3) (3) (18) Head without a cephalic cap, without interconnecting lobes to lips . . (4) (4) (9) Anterior cuticular flange of lip not projecting anterior to main lip mass, no papillae on body, lateral alae present, broad caudal alae supported by long, thin papillae ........... (5) (5) (6) Head without cordons or labial grooves .... HETERAKIS (p. 19) (6) (5) Head with cordons or labial grooves . . . . . . • (?) (7) (8) Spicules dissimilar, left with broad alae, right needle-like, restricted to India and associated regions ...... PSEUDASPIDODERA (p. 20) (8) (7) Spicules similar, never alate, restricted to South America ODONTOTERAKIS (p. 20) (9) (15) Narrow lateral alae, papillae on body surface, caudal alae when present either not supported by papillae or supported by many short, stoutish papillae . (10) (10) (n) No caudal alae on male tail . . . . . SPINICAUDA (p. 22) (n) (12) Caudal alae not supported by papillae .... AFRICAN A (p. 22) (12) (15) Caudal alae supported by stout papillae, male tail obliquely truncate, and sucker directed posteriorly ......... (13) THE SUPERFAMILY HETERAKOIDEA 25 (13) (14) Male tail without long terminal spike, anterior cuticular flange major part of lip ... STRONGYLURIS (p. 23) (14) (13) Male tail with long terminal spike, anterior flange not major part of lip MOACIRIA (p. 22) (15) (9) Caudal alae supported by 3-4 stout papillae ...... (16) (16) (17) Straight cordons running posteriorly from interlabial spaces, spicules very Complex ........ . GlRETERAKIS (p. 2l) (17) (16) No cordons, spicules not very complex .... METETERAKIS (p. 21) (18) (3) Head with a cephalic cap, interconnecting lobes to lips .... (19) (19) (22) Head without cordons .......... (20) (20) (21) Head with three cuticular " plates ", pre-cloacal sucker reduced, or lacking LAUROIA (p. 24) (21) (20) Head without cuticular " plates ", sucker normal . . PARASPIDODERA (p. 24) (22) (19) Head with cordons .......... (23) (23) (24) Cordons not anastomosing ..... ANSIRUPTODERA (p. 23) (24) (23) Cordons anastomosing ....... ASPIDODERA (p. 23) ACKNOWLEDGEMENTS I wish to express my thanks to the following who kindly allowed me to study specimens under their care or helped in locating specimens : Dr. E. Belle, Macdonald College, Quebec ; Dr. P. L. G. Benoit, Musee Royal du Congo Beige, Tervuren ; Professor J. J. C. Buckley, London School of Hygiene and Tropical Medicine ; Professor E. E. Bird, University of Georgia, U.S.A. ; M. le Professeur A. G. Chabaud, Museum National d'Histoire Naturelle, Paris ; Dr. R. N. Chaudhuri, Director of the Calcutta School of Tropical Medicine ; Dr. B. S. Chauhan, Zoological Survey of India, Calcutta ; Dr. E. Diechmann, Museum of Comparative Zoology, Harvard ; Dr. L. Forcart, Naturhistorisches Museum, Basel ; Dr. E. Ford, Director of the School of Public Health and Tropical Medicine, Sydney, Australia ; Dr. H. M. Hale, Late Director of the South Australian Museum, Adelaide ; Dr. G. Hartwich, Zoological Museum, Berlin ; The late Dr. S. L. Hora, Zoological Survey of India, Calcutta ; Dr. R. Kellogg, Director of the United States National Museum, Washing- ton ; Professor W. Kershaw, Liverpool School of Tropical Medicine ; Dr. E. Kritscher, Naturhistorisches Museum, Vienna ; Dr. J. T. Lucker, U.S. Department of Agriculture, Beltsville, Maryland ; Dr. J. F. Teixeira de Freitas, Institute Oswaldo Cruz, Brazil ; Mrs. P. M. Thomas, University of Adelaide, Australia ; Professore L. Travassos, Institute Oswaldo Cruz, Brazil ; Professore Umberto d'Ancona, Institute di Zoologia, Padua ; M. le Professeur M. Vachon, Museum National d'Histoire Naturelle, Paris ; Professor H. Vogel, Tropeninstitut, Hamburg. In particular my thanks are due to the following for their kindness and assistance : M. le Professeur A. G. Chabaud and his wife during repeated visits to Paris ; Herr Dr. G. Hartwich and his wife while working in Berlin ; Mrs. P. M. Thomas while working in Adelaide, South Australia ; and Dr. B. S. Chauhan during a short stay in Calcutta. REFERENCES BAYLIS, H. A. & DAUBNEY, R. 1922. Report on the parasitic nematodes in the collections of the Zoological Survey of India. Mem. Indian Mus. 7 : 263-347. - 1926. A synopsis of the families and genera of nematoda. London : British Museum (Natural History), i-xxxvi, 1-277. 26 W. G. INGLIS BELLE, EDWARD A. 1957. Helminth parasites of reptiles, birds and mammals in Egypt. IV. Four new species of oxyurid parasites from reptiles. Canad. J. zool. 35 : 163-169. CABALLERO Y C., E. 1955. Estudios helmintologicos de la region oncocercosa de Mexico y de la Repiiblica de Guatemala. Nematoda. 9* parte. Ada zool. Mex. 1 : 1-5. CAMERON, T. W. M. 1939. Studies on the endoparasitic fauna of Trinidad mammals. VI. Parasites of edentates. Canad. J. Res. 17 D : 249-264. CHABAUD, A. G. 1957. Sur la systematique des nematodes du sous ordre des Ascaridina parasites des vertebres. Bull. Soc. zool. Fr. 82 : 243-253. - 1965. Superfamily des Ascaridoidea, in Traite de zoologie. Anatomic, systematique, biologic. Embranchement des Nemathelminthes (Nematelmia Carl Vogt 1851 — Nemathel- minthia Gegenbaur, 1859) ou Aschelminthes (Aschelmintha Grobben, 1910). 4(3), 988-1025. CHAKRAVARTY, G. K. 1938. On a nematode, Pseudaspidodera jnanendrae n.sp. from the peafowl (Pavo cristatus). Parasitology 30 : 167-170. DIESING, C. M. 1851. Systema helminthum. II. Vildebonae. i-vi, 1-588. DUJARDIN, F. 1845. Histoire naturelle des helminthes ou vers intestineaux . Paris. 1-654. FREITAS, J. F. TEIXEIRA DE. 1956. Notas sobre " Heterakidae "; Railliet & Henry. 1914. (Nematoda, Subuluroidea). Rev. bras. Biol. 16 : 461-482. - 1956(1. Novo parasitos de reptil de Ihla Fernando de Noronha : Moaciria alvarengai g.n., sp.n. (Nematoda, Subuluroidea). Rev. bras. Biol. 16 : 335-339. GENDRE, E. 1909. Notes d'helminthologie africaine (Deuxieme note). P.V. Soc. linn. Bordeaux 63 : 33-41. HARTWICH, G. 1954. Die Vorderdarmstrukturen, des Excretions-system sowie der Kopfbau der Ascariden und ihre taxonomische Bedeutung. Wiss. Ztsch. Martin-Luther Univ. Halle-Wittenb. 3 : 1171-1211. INGLIS, WILLIAM G. 1957. The comparative anatomy and systematic significance of the head in the nematode family Heterakidae. Proc. zool. Soc. Land. 128 : 133-143. - 1958. A revision of the nematode superfamily Heterakoidea. Ann. Mag. nat. Hist. (io)12 : 905-912. - 19580. A revision of the nematode genus Meteterakis Karve, 1930. Parasitology 48 : 9-31- 19586. A redescription of the nematode Paraspidodera sellsi Morgan, 1927, and its re- moval to a new genus Morgascaridia. J. Helminth. 32 : 65-72. - 1965. The function and developmental significance of the cephalic septum in the Ascaridoidea (Nematoda). Proc. linn. Soc. Lond. 176 : 23-26. - 19650. The comparative anatomy of the ascaridoid cuticle. Bull. Soc. zool. Fr. 89 : 317- 338. - 19656. Patterns of evolution in parasitic nematodes. in Evolution of Parasites, $rd Symposium, Brit. Soc. Parasit. pp. 79-124. - 1966. The origin and function of the cheilostomal complex in the nematode Falcaustra stewarti. Proc. linn. Soc. Lond. 177 : 57-64. - 19660. The observational basis of homology. Syst. Zool. 15 : 218-228. - 1967. The relationships of the Seuratoidea (Nematoda). /. Helminth, in press. - & CHABAUD, ALAIN G. 1960. Sur la position systematique des Schneidernematinae (Nematoda). Ann. Parasit. hum. comp. 35 : 428-429. KARVE, J. N. 1930. Some parasitic nematodes of frogs and toads. Ann. trop. Med. Parasit. 24 : 481-491. KREIS, H. A. 1933. Em neuer parasitischer Nematode aus Corucia zebrata (Scincidae; Reptilia). Ganguleterakis triaculeatus n.sp. Z. Parasitenk. 6 : 332-338. LANE, CLAYTON. 1914. Suckered round-worms from India and Ceylon. Indian J. med. Res. 2 : 655-669. - 1917. Gireterakis girardi (n.g., n.sp.) and other suckered nematodes. Indian J. med. Res. 4 : 754-765. Lopez-Neyra, C. R. 1947. Helmintos de los Vertebrados Ibericos. II. Granada, pp. 413- 802. THE SUPERFAMILY HETERAKOIDEA 27 MADSEN, HOLGAR. 1950. Studies on species of Heterakis (Nematodes) in birds. Dan. Rev. Game Biol. 1 : 1-42. MAPLESTONE, P. A. 1932. The genera Heterakis and Pseudaspidodera in Indian Hosts. Indian J. med. Res. 20 : 403-420. MENDONCA, J. MACHADO DE. 1953. Heterakis isolonche Linstow, 1906 e Heterakis gallinae (Gmelin, 1790), agentes causais de Tiflite verrucosa em faisoes no Jardin Zoologico de Distrito Federal. Mem. Inst. Osw. Cruz 51 : 675-704. MUELLER, A. 1894. Helminthologische Beobachtungen an bekannten und unbekannten Entozoen. Arch. Naturgesch. 60, 5, 1 : 113-128. ORTLEPP, R. J. 1923. Two new nematodes collected in the Zoological Gardens of London. /. Helminth. 1 : 61-64. PROEN9A, M. C. 1937. Revisao do genero Aspidodera Railliet et Henry, 1912 (Nematoda : Subuluroidea) . Mem. Inst. Osw. Cruz. 32 : 427-438. 1938. S6bre um novo typo de Heterakinae Railliet et Henry, 1912 (Nematoda : Subulu- roidea). Livr. Jubil. Prof. L. Travassos, Rio de Janeiro, pp. 419-420. RAILLIET, A. & HENRY, A. 1912. Quelques nematodes parasites des reptiles. Bull. Soc. Path. exot. 5 : 251-259. - & HENRY, A. 1914. Essai de classification des Heterakidae. Extr. IXe Congr. Int. Zool., Monaco, pp. 674-682. RUDOLPHI, C. A. 1819. Entozoorum synopsis cui Accedunt Mantissa Duplex et Indices Locupletissimi. Berlin, i-iv, i-8n. SKRJABIN, K. I. & SCHIKHOBALOVA, N. P. 1947. (A reconstruction of the systematics of the nematode family Heterakidae). Dokl. Akad. Nauk. SSSR 58 : 718-721. (in Russian). - i947a. (A division of the nematodes belonging to Heterakis into generic components). Dokl. Akad. Nauk SSSR. 58 : 1865-1867. (In Russian). - 1951. (A reconstruction of the classification of the suborder Oxyurata Skrjabin, 1923). Trud. gel'mint. Lab. 20 : 5-8. (In Russian). - & LAGODOVSKAJA, E. A. 1961. [Principles of Nematodology. X. Oxyurata of Animals and Man.] Moscow Acad. Sci. U.S.S.R. (in Russian). TRAVASSOS, LAURO. 1913. Sobre as especies brasileiras da subfamilia Heterakinae Railliet et Henry. Mem. Inst. Osw. Cruz. 5 : 271—318. - 1914. Contribui9ao para o conhecimento da fauna helmintologica brasileira. III. Novo genero da familia Heterakidae Railliet et Henry. Mem. Inst. Osw. Cruz 6 : 137-142. 1920. Contribui9ao para o conhecimento da fauna helmintologica brasileira. IX. Sobre as especies do genero Spinicauda n.g. Mem. Inst. Osw. Cruz. 12 : 44-50. 1920^. Esb69o de uma chave geral dos nematddeos parasites. Rev. Vet., Rio de J. 10 : 59-70- YAMAGUTI, S. 1961. Systema helminthum. III. The nematodes of vertebrates. New York, London. I. pp. 1-680, II. 681-1261. YORKE, W. A. & Maplestone, P. A. 1926. The nematode parasites of vertebrates. London, i-x, 1-53°- APPENDIX While this paper was in press Chabaud (Alain G.) and Dollfus (Robert Ph.) have published the description of a new genus and species of heterakid (1966, Hatterianema hollandei N.G., N.SP., nematode heterakide parasite de rhynocephale. Bull. Mus. nat. hist. Nat. Ser. 2, 37 : 1041-1045) which they refer to the subfamily Meteterakinae. However, from the description the species is referable to the Spinicaudinae rather than Meteterakinae since i) the lips appear to be offset from the body, 2) there are no caudal alae on the male tail, 3) a gubernaculam is present, 4) there is no 28 W. G. INGLIS flap over the vulva in the female. In the key given above H. hollandei comes out at Spinicauda from which Hatterianema appears to differ largely in not possessing a cuticular flange projecting beyond the main mass of the lips and the large number of caudal papillae. In fact Hatterianema corresponds almost perfectly to the hypothetical ancestral form deduced above. PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING JAN 1967 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES (PISCES, CICHLIDAE) PART VI P. H. GREENWOOD BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 15 No. 2 LONDON: 1967 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES (PISCES, CICHLIDAE)N<^LH^ BY P. H. GREENWOOD British Museum (Natural History) ' k Pp. 29-119; 24 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 15 No. 2 LONDON: 1967 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the 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. 15, No. 2 of the Zoological 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.) Trustees of the British Museum (Natural History) 1967 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 13 January, 1967 Price £i 15$. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES (PISCES, CICHLIDAE) PART VI By P. H. GREENWOOD CONTENTS Page INTRODUCTION .......... 31 Haplochromis spekii (Blgr) ........ 32 Haplochromis pachycephalus sp. nov. ...... 39 Haplochromis maculipinna (Pellegrin) ...... 43 Haplochromis boops sp. nov. . . . . . . . 47 Haplochromis thuragnathus sp. nov. ...... 49 Haplochromis xenostoma Regan ....... 51 Haplochromis pseudopellegrini sp. nov. ...... 56 Haplochromis altigenis Regan ....... 60 Haplochromis dichrourus Regan ....... 65 Haplochromis paraguiarti sp. nov. . . . . . . 69 Haplochromis acidens sp. nov. . . . . . . . 73 Haplochromis prognathus (Pellegrin) ...... 78 Haplochromis argenteus Regan ....... 84 Haplochromis squamulatus Regan. . . . . . . 87 Haplochromis barbarae sp. nov. ....... 93 Haplochromis tridens Regan and Trcwavas ..... 97 Haplochromis orthostoma Regan . . . . . . . 100 Haplochromis parorthostoma sp. nov. . . . . . . 103 Haplochromis apogonoides sp. nov. ...... 105 DISCUSSION ........... 108 SUMMARY . . . . . . . . . . . 117 ACKNOWLEDGEMENTS . . . . . . . . . 117 APPENDIX: A statotilapia nigrescens Pellegrin, 1909 . . . . 118 REFERENCES ........... 119 INTRODUCTION THIS is the second of two papers dealing with the piscivorous species of Haplochromis in Lake Victoria. In the first part (Greenwood, 1962) representatives of the principal groups of piscivores were considered, and the main morphological trends within the trophic grade were discussed. The present paper covers the remaining species which have been studied to date ; undoubtedly more piscivorous Haplochromis species will be discovered, particularly amongst the as yet poorly sampled species of the deeper waters. Not every species considered here is a piscivore ; those of other trophic groups are included simply because the species have the morphology of a piscivore, and pre- sumably evolved from the same stem as their fish-eating relatives. ZOOL. 15, 2. 2 32 P. H. GREENWOOD Also included in this paper is a species apparently endemic to the Lake Kyoga system. This step was necessary because of its close relationship with a previously undescribed species from Lake Victoria. Some of the individual species described below, and in the previous paper, would seem to be so far removed from the generality of Victoria Haplochromis species as to justify their elevation to generic rank. Indeed, it could be argued that even some of the species-complexes have attained this level of differentiation. However, I do not think that the question can be dealt with until the whole Lake Victoria Haplo- chromis species-flock has been described. Even then, I doubt whether it will be possible to make any such divisions, at least generically. Perhaps a number of subgeneric groups could be justified on phyletic grounds, but these will be difficult to define. The situation closely resembles that encountered by Trewavas (1964) in the genus Serranochromis. However, I do not believe that her solution to the Serranochromis problem, the recognition of a gradal genus, is applicable to the situation amongst the piscivorous Haplochromis of Lake Victoria, particularly because the boundary between these species and any ancestral grade (or grades) would be even more obscure and indefinite than that separating Serranochromis from the Haplochromis of central Africa. Further complications are introduced when one considers the generic status of " Haplochromis " species outside Lake Victoria (and this includes the Lake Nyasa species in all their complexity) relative to the possibly polygeneric Haplochromis species of Lake Victoria. Haplochromis spekii (Boulenger), 1906 (Text-fig, i) Pelmatochromis spekii (part) Boulenger, 1906, Ann. Mag. nat. Hist., (7), 17, 440 ; Idem, 1915, Cat. Afr. Fish, 3, 416, fig. 285. (Lectotype.B.M. (N.H.), reg. no. 1906.^5.30.296, and pro- bably one paralectotype, 1906.5.30.307). Haplochromis spekii (part) : Regan, 1922, Proc. zool. Soc. Londn., 179 (same specimens as above) . Haplochromis serranoides Regan, 1922, op. cit. (Lectotype, B.M. (N.H.) 1911.3.27.17, and probably the two paralectotypes 1904.5.19.52-3). ? Paratilapia serranus (part) : Boulenger, 1915, Cat. Afr. Fish, 3, 334 (two specimens, 1904.5.19.52-3, see above). ? Haplochromis serranus (part): Regan, 1922, op. cit. 174 (paralectotype of P. spekii 1904.5.30.307, see above). LECTOTYPE : a male, 191-0 mm. S.L., from Bunjako, collected by Degen ; B.M. (N.H.) reg. no. 1906.5.30.296. NOTE ON THE SYNONYMY \ Certain small specimens (those indicated above with an interrogation mark) are included in the synonymy with some uncertainty. Using the diagnostic characters currently available, small preserved specimens of H. spekii cannot readily be separated from similar sized specimens of H. serranus. Regan (1922) distinguished H. spekii from H. serranoides on two characters : the maxillary extending to below the anterior quarter of the eye (barely reaches anterior orbital margin in H. serranoides}, and, the caudal peduncle longer than deep (as long as deep in H. serranoides}. Additional material shows that the A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 33 difference in caudal peduncle proportions is easily masked by intraspecific variability ; furthermore, I am unable to confirm the marked differences in caudal peduncle proportions which Regan found in the lectotypes of the two species. The difference in the posterior extension of the maxilla is valid for the lectotypes, but it must be noted that the jaws in H. spekii type are somewhat distorted because of a deformed right preorbital bone. Again, more material has shown that the maxilla has a variable posterior extension which links the extremes shown by the lectotypes of the two species. In all other characters, including the dentition and the preserved colour patterns, the two type specimens show no trenchant differences, and I consider them to be conspecific. Because of their small size (74 and 114 mm. S.L.) the paralectotypes of H. serran- oides have not been included in the redescription. I think it probable that these specimens are referable to the species. A similar problem is posed by three para- lectotypes of H. spekii (B.M. [N.H.] reg. nos. 1906.5.30.301, and 1906.5.30.297-8, of standard lengths 101-0, 91-0 and 79-0 mms. respectively). The two latter may perhaps be specimens of H. serranus, and the former is probably referable to H. spekii. However, until more is known about the characteristics of smaller specimens of H. spekii, I consider it inadvisable to give a definite identity to these three fishes. DESCRIPTION : based on 44 specimens (including the lectotype of the species, and the lectotype of H. serranoides) , 128-220 mm. standard length. Depth of body 32-8-39-8 (mean, M = 35-6) per cent of standard length, length of head 36-1-39-3 (M = 37'4) per cent. Dorsal head profile straight, sloping at an angle of 30°-35°, the premaxillary pedicels from barely to moderately prominent and interrupting the profile. Preorbital depth 18-0-24-2 (M = 20-7) per cent of head, least interorbital width 22-0-26-0 ( M = 23-3) per cent. Snout 1-2-1-3 times as long as broad, its length in fishes < 190 mm. S.L., (N = 25), 34-0-40-6 (M = 36-8) per cent of head, and in larger fishes (N = 19) 36-0-42-5 (M = 39-1) per cent. Eye diameter in fishes < 200 mm. S.L. (N = 34) 17-3-22-6 (M = 20-0), and in larger individuals (N = 10) 15-7-19-4 (M = 18-0) per cent of head ; ratio of eye/preorbital 0-8-1-3 (M == i-o). Depth of cheek 25-7-32-9 (M = 29-5) per cent of head. Caudal peduncle 16-7-19-8 (M = 17-9) per cent of standard length, 1-1-1-5 (modal range 1-2-1-3) times as long as deep. Mouth horizontal or slightly oblique, jaws equal anteriorly or the lower projecting slightly, its length 49-2-61-3 (M = 53-8) per cent of head, 1-7-2-6 (modal range 1-9-2-1) times as long as broad. Mental symphysis smooth or with a slight pro- tuberance. Premaxilla sometimes a little expanded medially but never beaked. Posterior tip of the maxilla reaching a point near the vertical through the anterior orbital margin or occasionally reaching this level (see also note on synonymy, p. 32). Gill rakers: stout or moderately stout, the lower I to 3 reduced ; 8 or 9 (rarely) on the lower part of the first gill arch. Scales ; ctenoid ; lateral line with 30 (f.i), 31 (f.8), 32 (£23), 33 (f.n) or 34 (f.i) ; 34 P. H. GREENWOOD r 1 3 -t-> rt 3 35 cheek with 3 (rare)-5 (mode 4) rows. Six to 8 (mode 7) between the upper lateral line and the dorsal fin origin, 5-9 (mode 8) between the pectoral and pelvic fin bases. Fins : Dorsal with 24 (f.i2), 25 (f-3o) or 26 (f.2) rays, comprising 14 (f.i), 15 (f.2Q) or 16 (f.i4) spinous and 8 (f.i), 9 (f.2i) or 10 (f.22) branched rays. Anal with ii (f.2), 12 (f-3o) or 13 (f.g) rays, comprising 3 spinous and 8 (f.2), 9 (f.3o) or 10 (f.g) branched elements. Pectoral 27-0-33-3 (M = 29-3) per cent of standard length. Pel vies with the first and second branched rays produced, slightly so in females but the first ray protracted and thread-like in males. Caudal subtruncate, scaled on its basal half. Teeth : In all specimens examined, both the inner and outer teeth are unicuspid, those of the outer row stout and strongly curved. The smallest fish (128 mm. S.L.) shows faint indications of lateral cusps on some teeth in the inner rows. There are 44-70 (M = 55) teeth in the outer row of the upper jaw ; inner teeth in this jaw are arranged in 3-5 (usually 3 or 4) rows, and in the lower jaw in 2 or 3 (rarely i or 5) rows. Osteology. The neurocranium of H. spekii is identical with that of H. serranus, that is, of the generalized predator type showing affinity with the skull of H. guiarti (see Greenwood, 1962). The lower pharyngeal bone is triangular, its dentigerous surface broader than long. The lower pharyngeal teeth are relatively fine, cylindrical in cross-section and weakly bicuspid ; some teeth are almost uniscuspid, with the larger cusp elongate and conical. The teeth are arranged in 22-24 rows. Vertebral counts (precaudal and caudal) for six specimens are 13 + 16 (f-3), 13 + 17 (f.2) and 12 + 17 (f.i). Coloration. Live coloration is unknown. Preserved specimens : Males (adult and sexually active) : Ground colour overall dusky, including the entire head, both jaws and the branchiostegal membrane ; very faint indications of a broad midlateral stripe visible behind the operculum to the beginning of the caudal peduncle where it merges with the dark general body colour. Dorsal fin dark except for the distal third to half of the soft part which is hyaline with dark spots and dashes. Caudal dark on its basal two-thirds, yellowish distally. Anal light dusky except for the distal quarter to third of the soft part which is hyaline ; 4 or 5 moderately large ocelli (dead white), usually arranged in two rows or one irregular row. Pelvic fins dusky. Adult (but sexually quiescent males) have a variable ground coloration which, how- ever, is always lighter than that of sexually active fishes. The snout and jaws are darker than the flanks which vary from dusky to light golden-brown ; branchiostegal membrane dark, but sometimes only in the region below the operculum. Dorsal fin dark, the lappets black, and the soft part often with close-set dark spots or dashes. Anal variable, from dusky to yellowish ; ocelli whiteish-grey, 2-5 in number and arranged as in active fishes. Pelvics usually dusky but of a variable intensity ; when light, the pigment concentrated over the spine and the first two branched rays. Females (adult and juvenile) : brownish above (and on the head and snout), shading to silvery-brown or greyish-silver on the lower flanks, belly, chest and operculum ; branchiostegal membrane greyish. A faint midlateral band (of 36 P. H. GREENWOOD variable depth and of irregular outline) runs from behind the operculum to the caudal fin origin ; there is also a very faint upper longitudinal band running slightly above the upper lateral line visible in some specimens. All fins are brownish-yellow the soft dorsal darkly maculate. Caudal dark brown on its proximal two-thirds (because of the dense maculation in that region). Immature males are coloured like females except that the longitudinal stripes are more distinct, and some specimens have very faint traces of 4 or 5 vertical bars crossing the longitudinal stripes on the flanks ; these bars extend from the back to a level about half way towards the ventral outline. The pelvic fins are faintly sooty. Ecology : Habitat. Haplochromis spekii occurs over both hard and soft substrates, but seems to show a slight preference for the former. Few specimens were collected from nets operated over exposed beaches, most coming from gill-nets set in sheltered areas where the water was 10-30 ft. deep. Some specimens were taken from more exposed areas, but not from deeper water. Food. Of the 42 fishes examined (from 24 localities), 22 contained food. Twenty- one of these had fed exclusively on small fishes (identified in 8 guts as Haplochromis species, in a further 8 as Cichlidae, and in one as a cyprinid). The exceptional fish contained unidentifiable fish remains and fragments of an ephemeropteran larva (probably Povilla adusta) . Breeding. All specimens < 150 mm. S.L. are immature, as is one specimen of 182 mm., but others > 150 mm. are mature. Both sexes reach the same maximum adult size. Affinities. The close relationship between H. spekii and H. serranus has been noted already (see above p. 33). There is complete overlap in most characters but the differential growth trends shown by two characters are such that this overlap is considerably reduced in fishes more than 120 mm. S.L. The two characters are depth of preorbital, and eye diameter as proportions of head length. In H. spekii both are, generally, larger than in H. serranus when specimens of the same size are compared. However, even in these characters there is still some overlap, and, from the sample studied, it seems likely that neither is a reliable diagnostic character when fishes < 120 mm. S.L. are compared. The difference between H. spekii and H. serranus (in the size range 120-205 mm.) is perhaps best shown by the ratio of eye diameter to preorbital depth, viz., 0-8-1-3 (mean i-o) for H. spekii, and 1-1-1-5 (mean 1-3) for H. serranus. Two other characters seem to show interspecific differences in their modal values, (i) In H. serranus the posterior tip of the maxilla usually lies below the eye or reaches to the vertical through the anterior orbital margin ; in H. spekii it rarely reaches as far posteriorly as the orbital margin (ii) Haplochromis serranus has a very prominent mental protuberance, but this bump is much weaker, if it is developed at all, in H. spekii. In many specimens of H. serranus the mental bump is so prominent that, in lateral view, the anterior margin of the dentary has a marked backward slope thus emphasizing the acuteness of the head profile ; in H. spekii the anterior margin of the dentary is, generally, almost perpendicular and so the tip of the head seems blunter than in H. serranus. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 37 Unfortunately it is impossible to compare the live colours of adult males from the two species ; preserved coloration is similar. This information, together with more field data on niche preferences, and small specimens of H. spekii, will be necessary before the precise relationships (or perhaps conspecifity) of the two species can be determined. If H. spekii and H. serranus were allopatric it would be tempting, on the information available, to consider them conspecific. However, experience with other sympatric species in Lake Victoria suggests that such slight morphological differences as are known between H. spekii and H. serranus can be the only ones manifest by biologically distinct species. Haplochromis spekii is more easily distinguished from other members of the H. serranus species complex. From H. victorianus it is recognizable by its larger adult size (some H . spekii are juvenile at a size near the upper adult limits for H. victorianus}, larger head (36-i-39'3> M =37'4% S.L.,c/. 33'5-36-o, M =34-8%), deeper cheek (257-32-9, M = 29-5% head, cf. 22-5-26-2, M — 24-6%), longer lower jaw, (49-2-61-3, M = 53-8% head, cf. 44-0-51-8, M = 47-1%), smaller eye in fishes < 200 mm. S.L. (17-3-22-6, M = 20-0% head, cf. 21-7-26-2, M— 24-6%), shorter pectoral fin (21-4-28-9, M = 25-1% S.L., cf. 26-2-32-7, M — 30-4%), and by having fewer and more curved outer teeth in the upper jaw (44-70, M = 55 teeth, cf. 64-86, M == 74). From H. maculipinna, H. spekii differs in its larger adult size, longer head (36-1-39-3, M =37-4% S.L., cf. 32-6-37-0, M =35'5%)> longer snout (34-0-40-6, M = 36-8% head, cf. 30-3-37-°. M = 337%). deeper cheek (25-7-32-9, M = 29-5% head, cf. 23-2-29-8, M —25-3%), longer lower jaw (49-2-61-3, M =53-8% head, cf. 43-3-52-8, M— 48-3%), and lower eye/preorbital ratio (0-8-1-3, M == I'O, cf. 1-3-1-6, M == 1-5). Although H. spekii resembles H. bartoni a little more closely in morphometric characters than it does H. victorianus, the species show a greater difference in neuro- cranial form. The neurocrania of H. victorianus and H. spekii are virtually identical, but that of H. bartoni is nearest the typical " prognathus "-group type (see p. 109). Morphometrically, H. spekii differs from H. bartoni in having a broader interorbital region (22-0-26-0, M = 23-3% of head, cf. 17-0-21-0, M == 18-6%), and a somewhat smaller eye (17-3-22-6, M = 20-0% head, cf. 20-3-24-1, M = 22-5%). Also H. spekii has a lower modal number of spinous dorsal fin rays (15 cf. 16). From the third member of the H. serranus species group, H. nyanzae, H. spekii differs in its larger adult size, larger head (36-1-39-3, M = 37-4% S.L., cf. 33'6-36'7> M = 35-4%), deeper cheek (25-7-32-9, M = 29-5% head, cf. 24-4-27-6, M = 25-9%), longer lower jaw (49-2-61-3, M =53-8% head, cf. 45-0-51-6, M =48-0%), and a lower modal number of spinous dorsal fin rays (15 cf. 16). Although typical specimens of H. spekii and H. gowersi are not readily confused (compare text-fig. I with text fig. 13 in Greenwood, 1962) there is one specimen whose appearance is such that I am unable to place it in one species or the other ; it is even intermediate in the two quantifiable morphological characters (body depth and inter- orbital width) showing the greatest interspecific differences. Haplochromis gowersi and H. spekii differ markedly in neurocranial form, but without dissection this character cannot be checked with sufficient precision in the unique intermediate P. H. GREENWOOD specimen. For the present, the possibility cannot be overruled that this fish is an interspecific hybrid. Phyletically, Haplochromis spekii appears to be a derivative from an H. serranus- like ancestor, the principal difference between the species being the larger adult size attained by H. spekii. Note : Gilchrist and Thompson (1917) record six specimens of Pelmatochromis spekii Blgr. from the Magalies river, Transvaal. I have not examined these speci- mens, but clearly they cannot be referred to Haplochromis spekii (Blgr.). Judging from their locality, it seems probable that they are specimens of Chetia flaviventris Trewavas. Dr. Trewavas is of a like opinion (personal communication). STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. B.M. (N.H.) 1906.5.30.296 (Lectotype) B.M. (N.H.) 1911.3.27.17 (Lectotype H. serranoides) B.M. (N.H.) 1966.3.9.1-4, 20-21, 30-35, 39-49 B.M. (N.H.) 1966.3.9.5 B.M. (N.H.) 1966.3.9.11-14 B.M. (N.H.) 1966.3.9.28-29 B.M. (N.H.) 1966.3.9.8-10 B.M. (N.H.) 1966.3.9.22 B.M. (N.H.) 1966.3.9.27 Locality UGANDA Bunjako .... Victoria Nile Napoleon Gulf, near Jinja . Beach near Nasu Point (Buvuma Channel) . Off S. tip of Ramafuta Island (Buvuma Channel) . Karenia, near Jinja (Napoleon Gulf) Pilkington Bay . Thruston Bay Buka Bay (Buvuma Channel) Collector Degen Mel land E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. B.M. (N.H.) 1966.3.9.17-19 B.M. (N.H.) 1966.3.9.7 B.M. (N.H.) 1966.3.9.15-16 B.M. (N.H.) 1928.5.24.413-5 KENYA Kisumu (Kavirondo Gulf) . Naia Bay (Kavirondo Gulf) Sagorony (Kavirondo Gulf) . Ulambwi Bay (Kavirondo Gulf) E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. Graham B.M. (N.H.) 1966.3.9.345 B.M. (N.H.) 1966.3.9.36-38 B.M. (N.H.) 1966.3.9.6 TANZANIA Beach near Majita . . . E.A.F.R.O. Between Ghogororo and Isanga River E.A.F.R.O. Mwanza (Capri Bay) . . . E.A.F.R.O. B.M. (N.H.) 1966.3.9.23-26 LAKE VICTORIA Locality unknown E.A.F.R.O. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 39 Haplochromis pachycephalus sp. nov. (Text-fig. 2) HOLOTYPE : an adult male, 199 mm. standard length, from 40 ft. of water off Kazima island, Uganda. B.M. (N.H.) reg. no. 1966.2.21.9. DESCRIPTION : based on the holotype and fourteen other specimens 150-232 mm. standard length. Depth of body 36-5-42-5 (M = 39-1) per cent of standard length, length of head 35-6-39-5 (M = 36-3) per cent. Dorsal head profile straight to moderately concave, the concavity exaggerated by the prominent premaxillary pedicels ; nuchal region prominent and gently convex, prenuchal region sloping at 30°-35°. Cephalic lateral line pores large, especially on the preorbital and preopercular bones, less so on the dentary. Preorbital depth 18-9-22-5 (M = 20-8) per cent of head, least interorbital width 24-6-31-3 (M = 27-8) per cent. Snout 1-1-1-4 (mode 1-2) times broader than long, its length 32-4-38-2 (M = 35*9) per cent of head ; eye diameter 18-8-22-2 (M = 20-6) per cent, depth of cheek 26-4-36-1 (M == 30-8) per cent. Caudal peduncle 13-2-16-0 (M == 14-8) per cent of standard length, 1-0-1-3 (mode 1-2) times as long as deep. Mouth oblique, sloping at an angle of 35°-45° (mode 40°). Jaws equal anteriorly or lower projecting slightly, its length 51-5-58-4 (M = 55-0) per cent of head, 1-5-1-9 (one specimen 2-2) times as long as broad. Posterior tip of maxilla reaching the vertical through the anterior orbital margin or nearly so. Gill rakers : stout, the lower I or 2 sometimes reduced, the upper 3 or 4 sometimes expanded ; 8 or 9 (7 in one specimen) on the lower part of the first gill arch. Scales ctenoid ; lateral line with 32 (f.3), 33 (f.4), 34 (f.4) or 35 (f.3), cheek with 5 or 6 (rarely 4) rows. Nine or 10 (less frequently 7, 8 or io|) between the upper lateral line and the dorsal fin origin, 7 or 8 (less frequently 6 or 9) between the pectoral and pelvic fin bases. Fins. Dorsal with 23 (f.i), 24 (f.6), 25 (f.6) or 26 (f.i) rays comprising 15 (i.g) or 16 (f.5) spinous and 8 (f.i), 9 (f.io) or 10 (f.3) branched rays. Anal with n (f.2) or 12 (f.i2) rays, comprising 3 spines and 8 (f.2) or 9 (f.12) branched elements. Pectoral 21-6-30-9 (M = 24-6) per cent of standard length. Pelvics with the first branched ray produced in sexually active males, slightly so in females and quiescent males. Teeth. The outer row in both jaws is composed of unicuspid, slender and slightly curved teeth. There are 60-80 (M = 70) teeth in the outer row of the upper jaw. Teeth in the inner rows are small, unicuspid, curved (strongly so in the upper jaw) and implanted obliquely. There are 4 or 5 (less frequently 2 or 3) rows in the upper jaw and 2 or 3 (rarely 4) in the lower. Osteology. No complete skeleton is available. The lower pharyngeal bone is triangular, with its dentigerous surface slightly broader than long (most markedly so in the smallest fish), or rarely, as long as broad. Lower pharyngeal teeth fairly coarse, their crowns weakly cuspidate and barely compressed ; some teeth in the two median rows are almost conical. The teeth are arranged in 18-22 rows. 4o P. H. GREENWOOD CS 1 1 N i A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 41 Vertebral counts in 5 specimens are : 13+16 (£.3), 14 + 16 (f.i) and 14 + 17 (f.i). Coloration : Live colours are unknown. Preserved material : Adult males. Ground colour variable and probably dependent on sexual state (also affected by preservation). Body greyish with black belly, chest and branchiostegal membrane, or the black replaced by a sooty-grey (in such specimens the branchiostegal membrane may be dark grey and flecked with sooty blotches) ; very faint traces of a broad midlateral stripe on the flank, originating behind the operculum and extending to the caudal fin origin. Dorsal fin greyish, dark lappets and maculae on the soft part. Caudal greyish, darkly maculate between the rays. Anal yellowish with a faintly sooty base, especially on the anterior part and around the ocelli ; the latter are dead-white, 5-9 (usually 5 or 6) in number and arranged in from i to 3 irregular rows. Pelvics blotched sooty to entirely black (the latter condition associated with the darkest body coloration). Two fishes (both from the same net haul) are more sexually active than the others. Both are a very dark brown, almost uniformly so except for a black belly, chest and branchiostegal membrane. The spinous dorsal is a very dark brown (nearly black), the soft part is lighter and has a yellowish margin. Caudal light brown with lighter maculae on the basal three-quarters. Anal very faint pink, with a narrow black basal streak which expands in the region of the proximal row of ocelli and extends in amongst them ; the ocelli are whiteish. Females (adult but quiescent ; based on two specimens only). Ground colour silver-grey becoming creamy on the chest and belly. Entire head (including the lower jaw) brownish with darker and irregular mottling. Body also mottled with sooty blotches, the effect being generally irregular except that on the flanks the blotches have some faint organisation into near vertical, broad bands extending from the dorsal outline to almost the ventral outline. There is some resemblance between this coloration and that of H. cavifrons (see Greenwood, 1962), although in H . pachycephalus the effect is less definitely that of freckling. Dorsal fin yellowish- grey with sooty freckling and blotching. Caudal densely and darkly blotched on its proximal third to half, greyish and darkly maculate distally. Anal greyish-yellow, with a narrow, sooty band along its base, and a duskiness over the spinous part ; both fishes have two large and distinct, dead white ocelli (an unusual feature in females). Pelvics hyaline with irregular sooty blotches. Ecology. Habitat. The species is known from four localities ; all are some distance off-shore but close to islands. The specimens all came from nets set on a soft bottom at depths of 100-120 ft., except in one locality where the collection was made after the use of explosives. In this instance the charge was set off in about 40 ft. of water over a rock shelf with deeper water on its off-shore side. Food. Of the 13 specimens examined (from 5 localities) six contained food in the stomach or intestines. All yielded macerated fish remains. Fragments of Haplo- chromis species were identified from three guts, a cyprinid fish in a fourth, and cichlid remains in two others. Breeding. Little information is available about the breeding habits of H. pachy- cephalus. All specimens except the smallest (a male, 150 mm. S.L.) are mature. The two largest fishes (232 mm. and 228 mm. S.L.) are males. 42 P. H. GREENWOOD Affinities. Haplochromis pacliycephalus is, at least on superficial characters and those detectable on a radiograph, related to the H. serranus species group (see p. 109). From H. serranus, H. pachycephalus is distinguished by its broader snout, broader interorbital (24-6-31-3, M = 27-8% head, cf. 20-4-26-8, M = 23-3%) and lower jaw (length/breadth ratio 1-5-1-9 cf. 1-8-2-5), and its smaller nuchal scales. From H. victorianus it differs in its broader interorbital (24-6-31-3, M = 27-8% head, cf. 21-5-24-5, M=22-6%), deeper cheek (26-4-36-1, M — 30-8% head cf. 22-5-26-2, M = 24-6%) and longer lower jaw (51-5-58-4, M = 55-0% head, cf. 44-0-51-8, M =3 47-1%) ; the lower jaw is also broader in H. pachycephalus. From H. spekii and H. maculipinna, the oblique mouth and broad snout of H. pachycephalus serve as immediately diagnostic characters, although the snout in H. maculipinna is broader than in other members of the " serranus "-group (being as much as i -i times broader than long, but generally as long as broad). As with other members of the group, H. maculipinna and H. spekii have a narrower interorbital region than H. pachycephalus ; H. maculipinna also has a larger eye (24-0-31-7, M = 26-3% head cf. 18-8-22-2, M s= 20-6% in H. pachycephalus) but the larger adult size reached by H. pachycephalus may influence this character. The same superficial characters (including the oblique mouth) serve to distinguish H. pachycephalus from H. bartoni and H. nyanzae, the former a member of the " prognathus " group, the latter a " serranus " group member. Haplochromis boops and H. thuragnathus (both " serranus "-group species) closely resemble one another (see pp. 50) and H. pachycephalus. Both differ from H. pachy- cephalus in the following characters : a narrower interorbital, shorter snout, larger eye, and larger nuchal scales. All three species have the snout broader than it is long. From the evidence available, H. pachycephalus would seem to be derived from an H. serranus-like ancestor, the principal morphological changes being an increase in mouth size coupled with greater obliquity of the mouth angle. The larger cephalic lateral line pores of H. pachycephalus are probably correlated with the deep water habitat of the species (as compared with H. serranus and its immediate allies). STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. Locality Collector UGANDA B.M. (N.H.) 1966.3.9.175-177 Off S. tip of Ramafuta Island (Buvuma Channel) . . . E.A.F.R.O. B.M. (N.H.) 1966.3.9. 174 Deep water off Dagusi Island . E.A.F.R.O. B.M. (N.H.) 1966.2.21 .9 Off Kazima Island . . . Uganda (Holotype) Fisheries Dept. B.M. (N.H.) 1966.3.9.166-169, Off Kazima Island . . . Uganda 171-173 Fisheries Dept. TANZANIA B.M. (N.H.) 1966.3.9.170 Off Godziba Island . . ' , E.A.F.R.O. LAKE VICTORIA B.M. (N.H.) 1966.3.9.178 Locality unknown ,. . . . E.A.F.R.O. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 43 Haplochromis maculipinna (Pellegrin), 1913 (Text-fig. 3) Paratilapia maculipinna Pellegrin, 1913 Bull. Soc. Zool. France, 37, 311 ; Idem, 1914, in Voyage de Ch. Alluaud et R. Jeannel en Afrique Occidental, 16, PI. i, fig. i, Paris. Paratilapia prognatha (part) : Boulenger, 1915, Cat. Afr. Fish., 3, 333. Haplochromis maculipinna : Regan, 1922, Proc. zool. Soc. Londn., 177, fig. 5. HOLOTYPE : a fish 122-0 mm. S.L. (Paris Museum No. 12-258) from Port Florence (Kavirondo Gulf) collected by Alluaud and Jeannel. This specimen differs from all others now included in the species by its much larger eye. In other characters, however, it agrees with these specimens and differs from the few other Victoria Haplochromis species characterized by large eyes. DESCRIPTION : based on 33 specimens (including the holotype), 91-5-166 mm. S.L. Depth of body 33-3-37-0 (M = 35-9) per cent of standard length, length of head 32-6-37-0 (M = 35-5) per cent. Dorsal head profile straight or slightly concave in those fishes with prominent premaxillary pedicels, sloping at 3O°-35°. Preorbital depth 16-4-20-4 (M = 18-2) per cent of head, least interorbital width 20-7-25-5 (M = 22-8) per cent. Snout as long as broad to i-i times broader than long, its length 30-3-37-0 (M = 33-7) per cent of head, eye diameter 24-0-29-2 (31-7 in the type), mean 26-3 per cent, ratio of eye/preorbital 1-3-1-6 (M = 1-5) but 1-9 in the type ; depth of cheek 23-2-29-8 (M = 25-3) per cent. Caudal peduncle 14-5-18-8 (M = 16-3) per cent of standard length, 1-2-1-8 (modal range 1-2-1-5) times as long as deep. Mouth moderately oblique, sloping upwards at 35°-40°, lower jaw projecting slightly to strongly, its length 43-3-52-8 (M = 48-0) per cent of head, 1-6-2-3 (modal 1 C M FIG. 3. Haplochromis maculipinna. Drawn by Barbara Williams. 44 P. H. GREENWOOD range 1-7-2-1) times as long as broad. Lips not noticeably thickened, premaxilla not expanded medially. Posteriorly tip of the maxilla not quite reaching the vertical through the anterior orbital margin. Gill rakers : moderately stout, the lower 1-3 reduced, the upper 2-5 flat and some- times divided ; 8-n (mode 10) on the lower part of the first arch. Scales : ctenoid ; lateral line with 31 (f.i), 32 (f.i5), 33 (f.i5) or 34 (f.2), cheek with 3-5 (mode 3) rows. Five and a half (rare) to 8 (modal range 6-7) scales between the upper lateral line and the dorsal fin origin, 5^ (rare) -9 (modal range 7-8) between the pectoral and pelvic fin bases. Fins : Dorsal with 24 (f.5), 25 (f.26) or 26 (f.2) rays, comprising 14 (f.i), 15 (f.24) or 16 (f.8) spinous and 9 (f.n) or 10 (f.22) branched rays. Anal with n (f.3), 12 (f.27) or 13 (f-3) rays, comprising 3 spines and n (f.3), 12 (^27) or 13 (f.3) branched elements. Pectoral 23-0-31-0 (M = 26-4) per cent of standard length. Pelvics with the first two branched rays produced in both sexes, but markedly elongate in adult males. Caudal truncate to subtruncate. Teeth. Except in the smallest specimen (91-5 mm. S.L.), the outer teeth are uni- cuspid ; in the exceptional fish most teeth are weakly bicuspid but a few are unicuspids. All outer teeth are slightly curved and slender, the curvature being most marked in teeth situated laterally and posterolaterally ; teeth situated posterolaterally in the upper jaw are the smallest and finest. There are 50-80 (M = 62) teeth in the outer row of the upper jaw. Teeth in the inner rows are more variable in form, and there is no clear-cut correla- tion between tooth form and the fish's size. The smallest specimen has only tricuspid inner teeth ; other and larger fishes may have only weakly tricuspids in both jaws, or tricuspid teeth predominating in both jaws but some unicuspids occurring in the lower jaw, or an admixture of tri- and unicuspids (the latter predominating) in the upper jaw and only unicuspids in the lower, or a mixture of tri- and unicuspids in both jaws, or unicuspids in the upper jaw and an admixture in the lower, or only unicuspids in both jaws. Some of the largest fishes fall in either the mixed uni- and tricuspid category or in the purely unicuspid one. The inner teeth are arranged in I (rare) — 4 (modes 2 and 3) rows in the upper jaw, and 2 (less commonly I or 3) rows in the lower. Osteology. The neurocranium of H. maculipinna is virtually identical with that of H. serranus, differing only in having a relatively lower supraoccipital crest. The premaxilla, dentary and suspensorium are also like those of H. serranus, except that the dentary is somewhat deeper and shorter in H. maculipinna. The lower pharyngeal bone is triangular and rather fine ; the dentigerous surface is slightly broader than long. Lower pharyngeal teeth are slender and distinctly cuspidate, those of the two median rows are the coarsest ; the teeth are arranged in 20-24 rows. Vertebral counts for seven specimens are : 13+16 (f.i) ; 13+17 (f.3) ; 14 +16 (f.2) and 14 + 17 (f.i), giving totals of 29 to 31. Coloration : Live colours are unknown. Preserved specimens : Males (adult and sexually active) : ground colour very dark brown, almost black, with a golden under- lay on the flanks and operculum, and a sooty overlay on the chest. Head, including A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 45 the lower jaw, dark but lips light brown. A very faint, but broad, lachrymal stripe runs from the antro ventral margin of the orbit to the angle of the lower jaw. The branchiostegal membrane is black. Dorsal fin almost uniformly dark sooty, the lappets black. Caudal dark on its proximal three-quarters, dusky distally. Anal dusky, but with a black band along its base, the band expanding anteriorly to cover most of the spinous part of the fin, which is black ; two or three large greyish ocelli are present. Pelvics black to dusky. Males (adult but sexually quiescent] : ground colour dark golden brown, lightest on the anterior flanks. Head dark brown, with a distinct, narrow lachrymal stripe from orbit to angle of lower jaw, and a narrow black vertical bar on the ascending preopercular limb ; branchiostegal membrane greyish-brown. Dorsal fin yellowish- grey, with black lappets, and the membrane between the branched rays dark grey- brown ; the pigment often broken into discrete maculae between the last three or four branched rays. Caudal dark yellowish-grey. Anal uniformly yellowish, with two or three faint, whiteish ocelli. Pelvics dark on the anterior third, otherwise yellowish to hyaline. Males (immature] : ground colour light brown on the flanks and belly, darker above the upper lateral line and on the dorsum of the head ; the branchiostegal membrane is light brown-grey, and a faint lachrymal stripe is visible. On the flanks there is a trace of an interrupted, dark midlateral band on the anterior half of the body, and a continuous band on the posterior half, extending to the caudal origin. Dorsal fin yellowish-brown, the lappets dark, as are the maculae between the branched rays. Caudal yellowish-brown, with dark elongate blotches between the rays. Anal uniformly yellowish-brown, with two or three, distinct and dusky-grey ocelli. Pelvics yellowish with a faint dusky overlay, especially over the anterior part of the fin. Females (adult and juvenile] : ground colour golden brown, darker on the upper half of the body, and the dorsal surface of the head ; faint traces of a rather broad lachrymal stripe are often visible, the stripe generally not extending to below the level of the maxilla, but reaching the angle of the lower jaw in some specimens. Faint traces of an interrupted midlateral band on the anterior half of the body, and a continuous band on the posterior half are often visible ; in some specimens no lateral band is visible, and in others the band is continuous except for a short break at about its midpoint. A few specimens show indications of a much interrupted band (really a series of 6 or 7 broad blotches) running slightly above the upper lateral line on the anterior half of the flanks, and on the lateral line posteriorly. Dorsal fin yellowish, usually darker between the posterior spines, and darkly maculate on the soft part, but uniformly yellowish with very faint maculations posteriorly in others. Anal, caudal and pelvic fins uniformly yellowish. Ecology. Although some individuals occur over sandy, exposed and wave- washed beaches, members of this species are commoner in sheltered gulfs and bays where the water is from 10-30 ft. deep and the substrate is either soft mud or sand and shingle ; a few specimens are from deeper water (35-40 ft.) near off-shore islands. Food. Eleven of the 30 specimens examined (from 16 localities) contained food ZOOL. 15, 2. 3 46 P. H. GREENWOOD in the stomach and intestines. Seven fishes yielded fragmentary fish remains (identified as a cyprinid in one, and as Haplochromis species in two others), three contained fragmentary insect remains (probably larval Ephemeroptera) , and one bottom debris. Breeding. Little information is available ; most specimens less than 140 mm. S.L. are immature, as is one slightly larger individual (145 mm.). Both sexes attain the same maximum adult size. Affinities. In both its gross and detailed morphology H. maculipinna shows affinity with the " serranus " species group, i.e. H. serranus, H. victorianus, H. spekii, and their deep water relatives H. pachycephalus, H. boops and H. thuragnathus. Criteria for distinguishing H. maculipinna from all but the first two species are considered under the descriptions of those species (see pp. 37, 42, 49 and 51 for the species respectively). From H. serranus, H. maculipinna is distinguished by its larger eye (24-0-3I-7, M = 26-3% head, cf. 20-4-26-0, M = 23-3%), shorter and more oblique lower jaw (43'3-52-8, M =48-3% head, cf. 47-7-60-0, M =54-3%), and higher eye/preorbital ratio (1-3-1-6, M = 1-5 cf. 1/1-1-5, M = 1-3). From H. victorianus, it differs in its larger eye (24-0-31-7, M = 26- 3% head cf. 21-7-25-5 M = 23-6%), higher eye/preorbital ratio (1-3-1-6, M = 1-5, cf. 1-1-1-3, M = 1-2), more oblique lower jaw (sloping at 30°-35° cf. horizontal or very slightly oblique) and its fewer and finer outer teeth (50-80, M = 62, cf. 64-86, M = 74 teeth in the upper jaw). The close resemblance between H. maculipinna and these two species is obvious, and is greater than the resemblance between H. maculipinna and other members of the " serranus " group. There are two other species, H. nyanzae and H. bartoni which, at least superficially, resemble members of the " serranus " group although H. bartoni seems to belong to a different phyletic line (see p. 109) . Haplochromis maculipinna differs from H. nyanzae in its larger eye (24-0-31-7, M = 26-3% head, cf. 19-1-24-0, M = 22-1%) and higher eye/preorbital ratio (1-3-1-6, M — 1-5, cf. 1-1-1-3, M = x*2)> an(i by its finer and less curved outer teeth. From H. bartoni, it differs in having a shorter head (32-6-37-0, M = 35-5% standard length, cf. 36-2-39-7, M=37»5%), broader interorbital (20-7-25-5, M = 22-8% head, cf. 17-0-21-0, M = 18-6%) and shorter, more oblique lower jaw (43-3-52-8, M = 48-3% head, cf. 50-8-57-0, M = 52-5%). Neurocranial form differs in the two species, that of H. maculipinna being of the " serranus " type, and that of H. bartoni being of the " prognathus " type (see p. no). Resemblances between H. maculipinna and H. acidens are discussed on p. 76. It seems unlikely that the species are closely related. Phyletically, H. maculipinna was probably derived from a H. serranus-like ancestor. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 47 STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No Locality B.M. (N.H.) 1906.5.30.263 B.M. (N.H.) 1966.3.9.145-151 B.M. (N.H.) 1966.3.9.132-134 B.M. (N.H.) 1966.3.9.135-144 B.M. (N.H.) 1966.3.9.129-131 B.M. (N.H.) 1966.3.9. 124-6 B.M. (N.H.) 1966.3.9.123 B.M. (N.H.) 1966.3.9.128 B.M. (N.H.) 1966.3.9.127 Paris Museum 12-258 (Holotype) Collector Degen E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. UGANDA Bunjako ..... Napoleon Gulf, near Jinja . Beach near Nasu Point (Buvunia channel) Near Ramafuta Island (Buvuma Channel) Between Yempita and Busiri Island (Buvuma Channel) Buka Bay Fielding Bay Kazima Island (near Entebbe) Pilkington Bay . KENYA Port Florence (Kavirondo Gulf) . Alluaud & Jeannel E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. Uganda Fisheries Dept. E.A.F.R.O. B.M. (N.H.) 1966.3.9.122 LAKE VICTORIA Locality unknown . . . E.A.F.R.O. Haplochromis boops sp. nov. (Text-fig. 4) HOLOTYPE : an adult male, 190 mm. standard length, from 120 ft. of water, off the southern tip of Buvuma island (Uganda). B.M. (N.H.) reg. no. 1966.2.21.7. DESCRIPTION : based on three specimens, 179-194 mm. standard length ; all are males. Depth of body 40-5-42-3 per cent of standard length, length of head 35-3-36-1 per cent. Dorsal head profile straight, sloping steeply at 4o°-5o° ; premaxillary pedicels not prominent. Cephalic lateral line pores enlarged, especially those on the preoperculum, preorbital and dentary. Preorbital depth 17-8-18-6 per cent of head, least interorbital width 21-7-25-7 per cent. Snout 1-2-1-3 times as broad as long, its length 32-6-32-8 per cent of head ; diameter of eye 23-9-25-7, depth of cheek 28-0-30-0 per cent. Caudal peduncle 14-8-15-6 per cent of standard length, 1-2 times as long as deep. Mouth somewhat oblique, sloping at 30°-35° (a horizontal line drawn through the tip of the lower jaw passes below the orbit). Jaws equal anteriorly or the lower projecting slightly, its length 50-0-52-5 per cent of head, 1-5-1-8 times as long as broad. Posterior tip of the maxilla reaching to a point below the anterior part of the eye. Gill rakers : variable in form, from slender to stout, even in one individual ; the upper 3 rakers branched in one fish. Eight or 9 on the lower part of the first gill arch. 48 P. H. GREENWOOD FIG. 4. Haplochromis boops. Drawn by Barbara Williams Scales : ctenoid ; lateral line with 33 (f .2) or 34 scales, cheek with 4 (f .2) or 5 rows. Seven or 8 scales between the dorsal fin origin and the upper lateral line, 7 or 8 between the pectoral and pelvic fin bases. Fins : Dorsal with 15 spines and 9 (f.i) or 10 (f.2) branched rays, anal with 3 spines and 8 (f.i) or 9 (f.2) branched rays. Pectoral 25-2-32-6 per cent of standard length. Pelvics with the first branched ray produced. Caudal subtruncate. Teeth : In the outer row of both jaws, the teeth are small, curved and slender, with about 70 in the upper jaw. Inner teeth, arranged in three rows in both jaws, are unicuspid, small and slightly curved. Osteology. No complete skeleton is available, but radiographs have been studied. The lower pharyngeal bone is triangular, with its dentigerous surface broader than long. Lower pharyngeal teeth are relatively stout with cylindrical necks, and compressed, weakly cuspidate crowns. The teeth are arranged in 20-22 irregular rows. The vertebral counts in three specimens are : 13+16 (f.i) and 13 -f- 17 (f.2). Coloration : Live colours are unknown. Preserved material : Males (adult and sexually active) : ground colour almost uniformly black (including the head, snout, branchiostegal membrane and belly) but with a brownish tinge. Dorsal with the spinous part blotched black on dark grey (black predominating), soft part black but with a hyaline band originating at the level of the tip of the last spine thence passing slightly downwards to end at a point about one third of the distance from the tip of the last branched ray ; the dark band distal to the hyaline strip is less intense than that along the basal part of the fin. Anal black over the spines and along the basal third of the whole fin, remainder sooty ; the five grey-white ocelli are arranged in two rows. Pelvic fins black. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 49 No female specimens are available. Ecology. Habitat. The two localities from which H. boops is known are in deep water (about 120 ft.) near islands and over mud substrates. Food. Two specimens provided data on feeding habits. In both, the stomach had been everted, but fragments of macerated fish were collected from the pharynx and amongst the folds of the stomach wall. Judging from the scales and fin spines collected in this way, the fishes had fed on Haplochromis. Breeding. All three specimens are adult, sexually active males. Affinities. Haplochromis boops most closely resembles H. thuragnathus ; at present the species can only be distinguished by the more oblique jaw of the latter (see p. 51). Like H. thuragnathus, H. boops appears to be a derivative of the H. serranus species group, probably from an ancestor resembling H. maculipinna. From that species H. boops is immediately distinguished by its broader snout (broader than long, cf. as long as broad), enlarged cephalic lateral line pores, and deeper body. STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. Locality Collector UGANDA B.M. (N.H.) 1966.3.9.182 Near Dagusi Island . . . E.A.F.R.O. B.M. (N.H.) 1966.2.21.7 Off southern tip of Buvuma (Holotype) Island E.A.F.R.O. B.M. (N.H.) 1966.3.9.181 Off southern tip of Buvuma Island E.A.F.R.O. Haplochromis thuragnathus sp. nov. HOLOTYPE : an adult male, 191 mm. standard length, from 120 ft. of water off the southern end of Buvuma island (Uganda) ; B.M. (N.H.) reg. no. 1966.2.21.8. DESCRIPTION : based on three specimens, 191 and 200 mm. standard length. Since so few specimens are available, only ranges for morphometric characters are given. Depth of body 39- 8-41*5 per cent of standard length, length of head 34-5-35-1 per cent. Dorsal head profile slightly concave, sloping at about 30°. Cephalic lateral line pores are enlarged, especially those on the preorbital and preopercular bones. Preorbital depth 16-4-18-8 per cent of head, least interorbital width 23-2-24-5 per cent. Snout 1-2-1-3 times as broad as long, its length 30-9-31-8 per cent of head, eye diameter 24-6-26-8, depth of cheek 27-4-29-8 per cent. Caudal peduncle 16-0-17.7 Per cent of standard length, 1-3-1-4 times as long as deep. Mouth oblique, sloping at 40°-45°, the jaws equal anteriorly or the lower projecting slightly, length of lower jaw 53-6-56-5 per cent of head, 1-7-2-2 times as long as broad. Posterior tip of the maxilla extending to the vertical through the anterior orbital margin or to below the anterior part of the eye. A horizontal drawn antero- 5o P. H. GREENWOOD posteriorly through the tip of the lower jaw passes through the lower part of the eye (cf. H. boops where the line passes below the orbit.) Gill rakers : stout, 9 on the lower part of the first gill arch. Scales : ctenoid ; lateral line with 32, 33 or 34 scales, cheek with 2-4 rows. Seven or 7^ scales between the upper lateral line and the dorsal fin origin, 7 between the pectoral and pelvic fin bases. Fins. Dorsal with 16 spines and 9 branched rays. Anal with 3 spines and 8 branched rays. Pectoral 26-o-30'0 per cent of standard length. Pelvics with the first branched ray produced, proportionately more so in males. Caudal subtruncate. Teeth. In the outer row of both jaws, the teeth are unicuspid, small and curved ; there are 70 teeth in the upper jaw. Inner teeth are unicuspid, small and slightly curved, and are arranged in 3 series in the upper jaw, and 2 or 3 series in the lower. Osteology. No complete skeleton is available, but radiographs of the three specimens were examined. The lower pharyngeal bone is triangular, its dentigerous surface broader than long. The pharyngeal teeth are relatively coarse, with cylindrical necks and compressed, weakly biscuspid crowns, and are arranged in 20-22 irregular rows. Vertebral counts for all three specimens are : 13 -f- 17. Coloration : Live colours are unknown. Preserved material : Males (adult and sexually active) : ground colour sooty over dark brown dorsally (including the head), silvery on the belly and midflank ; chest and belly darker (i.e. sootier), almost black There is a faint golden flush on the operculum, but it is confined to the centre of this bone and is outlined with a broad dark margin. On the flank of the lighter coloured fish are traces of a broad, dark midlateral stripe. The branchiostegal membrane is black except for its posterior and ventral margins which are greyish. Dorsal fin black except for the distal half of the soft part which is greyish. Caudal dark, but lighter towards the distal margin. Anal black on its basal half and over the spinous portion ; one large white ocellus is present in the fish with the smaller testes, but the other has 8 ocelli arranged in two irregular rows. Pelvic fins black. Female (quiescent) : ground colour brownish, darker on back, head and snout, lighter (with silvery background) on flanks and belly ; very faint traces of a broad (three scale rows deep) interrupted midlateral band on the flanks. Operculum silvery ; a faint, dark lachrymal stripe from the orbit to behind the posterior tip of the maxilla. All fins grey-brown, the anal with 3 small, whiteish spots in the position occupied by the ocelli in males ; pel vies more grey than brown. Ecology. Habitat. All three specimens came from nets set on the mud-bottom in water about 120 ft. deep off the southern tip of Buvuma island. Food. Two of the three specimens examined had fragments of small Haplochromis species in the stomach and intestines ; the guts of the third fish were empty. Breeding. The three specimens are adults, the two males sexually active, the female quiescent. Affinities. Haplochromis thuragnathus is most closely related to H. boops. Indeed, when more material is available it may be shown that the species are not distinct. Information on the live coloration of adult males of the two species would be extremely useful in establishing their status. From H. boops, H. thuragnathus A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 51 is distinguished by its more oblique lower jaw. If a horizontal line is drawn poster- iorly from the tip of the lower jaw (when closed) it passes through the lower part of the eye in H. thuragnathus , but below the eye in H . boops. A third member of this group, H. pachycephalus, is compared with#. thuragnathus. on p. 42. It seems probable that H. thuragnathus was derived from an H. maculipinna-like ancestor, and more particularly from one like H. boops (assuming that the more oblique lower jaw is a derived condition). Like H. pachycephalus and H. boops, Haplochromis thuragnathus differs from other members of the " serranus " phyletic assemblage by its broad snout, and from individual members of the group by various combinations of morphometric characters (see descriptions of H. serranus and H. victorianus in Greenwood, 1962, and of H. maculipinna and H. spekii on pp. 46 and 37 above). STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. Locality UGANDA B.M. (N.H.) 1966.2.21.8 B.M. (N.H.) 1966.3.9.179-180 Off S. tip of Ramafuta Island Off S. tip of Ramafuta Island Collector E.A.F.R.O. E.A.F.R.O. Haplochromis xenostoma Regan, 1922 (Text-figs. 5 and 6) Paratilapia prognatha (part) : Boulenger, 1915, Cat. Afr. Fish., 3, 333 (two specimens, one collected by Sir H. H. Johnston, the other from Entebbe and collected by Degen). Haplochromis xenostoma, Regan, 1922, Proc. zool. Soc. Londn., 185, fig. 10. LECTOTYPE : an immature fish 104-0 mm. S.L. (B.M. [N.H.]reg. no. 1901 . 6 . 24 . 90) collected by Sir H. H. Johnston from Lake Victoria (locality unspecified). FIG. 5. Haplochromis xenostoma, juvenile ; lectotype, about natural size. Regan, Proc. zool. Soc. From 52 P. H. GREENWOOD DESCRIPTION : based on 27 specimens (including the lectotype) 99-203 mm. S.L. but excluding the paralectotype. Only four specimens are less than 140 mm. S.L. (99, 104 [lectotype], 106 and 119 mm. respectively) ; on the basis of this material it appears that body depth and lower jaw length may show some slight positive allometry. Depth of body 27-0-40-0 (M = 36-5) per cent of standard length, head length 34-6-39-7 (M = 37-3) per cent. Dorsal head profile straight or slightly convex, sloping at an angle of 20°-3O°, its outline noticeably interrupted by the prominent premaxillary pedicels which give it a stepped appearance. Preorbital depth 17-5-22-7 (M = 20-3) per cent of head, least interorbital width 20-4-27-5 (M = 24-5) per cent. Snout 1-2-1-5 times as long as broad, its length 34-2-39-2 (M s= 37-4) per cent of head ; eye diameter 18-5-24-3 (M = 19*7), depth of cheek 24-3-30-8 (M = 28-1) per cent. Caudal peduncle 14-3-18-3 (M = 15-3) per cent of standard length, 1-1-1-7 (modal range 1-3-1-4) times as long as deep ; the lectotype has an unusually shallow peduncle (ratio 1-7). Mouth very oblique, sloping at an angle of 40°-45°, the lower jaw strongly pro- jecting beyond the upper, its length 50-0-62-0 (M = 57*o) per cent of head and 2-0 (rarely)-3-o times as long as broad (modal range 2-3-2-5). Posterior tip of maxilla generally not reaching the vertical through the anterior orbital margin, but reaching this point in a few specimens. Gill rakers : short and stout, or relatively slender and elongate, the lower one or two reduced ; 8-10 (mode 9) on the lower part of the first gill arch. Scales : ctenoid ; lateral line with 29 (f.i), 31 (f.io), 32 (f.i2), 33 (f.3) or 34 (f.i), cheek with 3 (mode) or 4 rows. Six or 7 (rarely 5 or 8) scales between the upper 1 CM FIG. 6. Haplochromis xenostoma adult. Drawn by Barbara Williams. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 53 lateral line and the dorsal fin origin, 5-7 (rarely 8) between the pectoral and pelvic fin bases. Fins : Dorsal with 24 (f.Q), 25 (f.17) or 26 (f.i) rays, comprising 14 (f.i), 15 (f.22) or 16 (f_4) spinous and 9 (f.n) or 10 (f.i6) branched rays. Anal with n (f.n) or 12 (f.i6) rays, comprising 3 spines and 8 or 9 branched elements. Pelvics with the first two branched rays produced in both sexes but proportionately more so in adult males. Pectoral 24-2-33-0 (M — 28-0) per cent of standard length. Caudal trun- cate, scaled on its proximal half or slightly more. Teeth. In fishes 119 mm. S.L. and above, the outer teeth in both jaws are unicuspid and moderately stout (but occasionally slender) , those in the anterior part of the jaw with a slight inward curvature, and those situated laterally and posteriorly even less curved. The lectotype (104 mm. S.L.) has an outer dentition like that of larger fishes, but in the other small fishes (88-106 mm. S.L.) the outer teeth are distinctly bicuspid anteriorly, and weakly bicuspid laterally and posterolaterally. There are 56-94 (M = 82) teeth in the outer row of the upper jaw. Fishes less than 106 mm. S.L. have either only tricuspid teeth or a mixture of uni- and tricuspids (some weakly so) in the inner series. A specimen 119 mm. S.L. has predominantly bicuspid teeth in the upper jaw, but in the lower jaw the first row of inner teeth is composed of unicuspids, and the other rows of tri- and weakly tricuspids. In all other specimens the inner rows are composed entirely of uni- cuspids. Inner teeth may be implanted somewhat obliquely so as to be medially inclined. The teeth in the outermost row of the upper inner series are often notice- ably larger than their congeners. There are 2-5 rows of inner teeth in the upper jaw, and 2 or 3 (rarely 4) in the lower. Osteology. The neurocranium of H . xenostoma is similar to that of H. victorianus (see Greenwood, 1962) but has a longer preorbital face (30-3 per cent of neurocranial length cf. 26-0 per cent ; the preorbital face being measured from the anterior tip of the vomer to the lateral ethmoid) ; the neurocrania of the two species also differ in that the supraocciptal crest of H. xenostoma is relatively higher and more pointed than in H. victorianus. It differs from the neurocranium of H. serranus (as it does from that of H. victorianus) in its less curved dorsal profile, and its longer preorbital face. The very oblique and prognathous lower jaw is reflected in certain details of the suspensorium (text-fig. 7) ; all comparisons were made with H. serranus, a species 2 MM FIG. 7. Articulatory facet for the quadrate on the articular of (A) H. xenostoma, and (B) H. serranus. 54 P. H. GREENWOOD with a moderately oblique jaw angle of 20°-30°. In H. xenostoma the articulatory surface of the articular is deeper and more nearly " U " shaped in lateral view ; it lacks the posterior prolongation of its ventral border, but has a marked, near- conical eminence developed postero-medially. All these differences seem to be associated with the oblique angle of the jaw at rest, and the wide angle through which it can be abducted when the mouth is maximally protruded. The small eminence appears to function as a control for the degree of lateral movement of the dentary, particularly when that bone is dropped almost to the horizontal. The posterior vertical limb of the articular in H. xenostoma slopes forward at a much greater angle from the perpendicular, thus providing more space between this bone and the suspensorium. On the basis of my material (two skeletons of H. xenostoma, and one each of H. serranus and H. victorianus) it appears that the horizontal length of the suspen- sorium (as measured in a horizontal plane from the mid-point of the hyomandibular to the articular surface of the quadrate) is greater in H. xenostoma than in the other two species. This could account, at least partly, for the greater prognacity of this species, whose lower jaw has the same relative length as that of H. serranus and H. victorianus. The angle of the hyomandibular relative to the perpendicular is similar in all three species. The premaxilla has a slight median expansion of its dentigerous surface, and the pedicels are relatively shorter than those of H. serranus. As compared with the dentary of H. serranus and H. victorianus, that of H. xenostoma is deeper and stouter, and the dentigerous surface has a more pronounced upward sweep towards the coronoid region. The lower pharyngeal bone is triangular, and fairly stout ; the dentigerous area is as long as broad or very slightly broader than long. Lower pharyngeal teeth are variable in form, usually with coarse, cylindrical necks and compressed, weakly cuspidate crowns ; in some fishes, however, the crowns are distinctly cuspidate. Less commonly, the teeth are slender and compressed, with very weakly cuspidate crowns. The teeth are arranged in 22-24 rows. Vertebral counts in 7 specimens are : 13+15 (f.i), 13 + 16 (f.5) and 13 + 17 (f.i), giving totals of 28-30. Coloration : Live colours are known only for a single juvenile female ; ground coloration silvery, shading to yellowish-grey dorsally, the dorsal surface of the snout dark grey. Dorsal fin dark hyaline, pelvics hyaline, anal yellow, caudal yellowish- grey. Preserved coloration : Males (adult and sexually active) have the ground coloration generally dusky over dark brown dorsally, and silver on the flanks and belly, the latter region together with the chest often with a dusky overlay ; lower jaw and entire branchiostegal membrane also sooty. Snout and preoperculum sooty, but the upper lip is dark brown ; operculum with a faint golden flush. A faint but dark and broad midlateral stripe runs along the flank from the posterior margin of the preoperculum to the caudal fin origin. Dorsal fin yellow-brown, with a sooty overlay on the spinous part, and black lappets ; soft part with dark spots and dashes. Caudal dusky on its proximal threequarters, yellowish-brown distally. Anal light brownish-yellow, with a thin dark line along the base, and black lappets ; A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 55 3 or 4 large, darker whitesh ocelli are present, each with a narrow black outline. The ocelli are arranged in one or two rows. Pel vies entirely dusky. Males (adult but quiescent] : have essentially the same coloration as active males, but some are lighter (that is, with more silvery flanks, and greyish branchiostegal membrane) . Females (adult, and at various degrees of sexual activity) . Greyish-brown above, shading through silvery-grey on the flanks to gold below ; snout and preorbital region dark. On the flanks, a faint dark midlateral band (as in males) may be visible, and in addition, a fainter upper band running just above the upper lateral line. Dorsal fin yellowish-grey with a sooty overlay on the spinous part, and with black lappets ; soft dorsal sometimes darkly spotted. Caudal dark on the proximal two-thirds, lighter distally. Anal yellowish with a faint sooty overlay, sometimes with ill- defined dark spots in the position of the ocelli in males. Pel vies hyaline, usually with a dusky overlay. Ecology. Habitat. The species is apparently confined to sheltered or relatively undisturbed water, being common in bays and gulfs where the water is less than 40 ft. deep, and the bottom is of soft mud, sand or shingle. Available records suggest that the species favours a mud substrate. Food. Of the 21 fishes examined (from 9 localities), only 6 contained food in the guts. In each case the food comprised fragmentary fish remains, unidentifiable except in one instance (a small Haplochromis species) . Breeding. All fishes less than 160 mm. S.L. are immature ; one larger individual (a female 163 mm. S.L.) is also immature. Males and females appear to reach the same maximum adult size. Affinities. The very oblique mouth, marked prognacity, and relatively deep body (at least in adults) serve to distinguish H. xenostoma from the majority of larger Haplochromis species in the lake. There is some resemblance between this species and H. macrognathus and H. plagiostoma, both species with an oblique mouth, and in the case of H . macrognathus, a prominent lower jaw. Haplochromis xenostoma is readily distinguished from H. macrognathus by its broader head (interorbital width 20-4-27-5, M = 24-5% head cf. 16-5-22-2, M = 18-6% ; snout 1-2-1-5 times as long as broad, cf. 1-5-2-2 times in H. macro- gnathus), and its more oblique mouth. The neurocranium also differs, that of H. macrognathus being of the " prognathus " type, whilst the skull of H. xenostoma is clearly of the "serranus" type (see p. in and discussion on pp. 109-113 ; also Greenwood, 1962). From H. plagiostoma, H. xenostoma differs in its larger adult size, longer and narrower snout (34-2-39-2, M = 37-4% head, cf. 28-2-35-5, M=32-5%), more prominent and longer lower jaw (50-0-62-0, M = 57-0% head, cf. 44-0-54-5, M = 49-2% ; lower jaw rarely projecting in H. plagiostoma) and the greater number of teeth in the outer row of the upper jaw (56-94, M =82, cf. 44-68, M =57). Neurocranial form in these two species is similar (see p. 113). Another species with an oblique mouth is H. cavifrons. It is distinguished from H. xenostoma by its unique mottled coloration, lack of prognacity, broader snout (as long as broad or slightly broader than long, cf. 1-2-1-5 times as long as broad), 56 P. H. GREENWOOD and fewer teeth (56-74, M = 63, cf. 56-94, M =82). The profile of the head also differs (compare fig. 6 above with fig. 20 in Greenwood, 1962). Phylogenetically, H. xenostoma could be derived from a species resembling H. plagiostoma ; its affinities seem to lie more with the " serranus " group than with the "prognathus" group to which H. macrognathus belongs. (See also discussion on PP. US)- STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. Locality B.M. (N.H.) 1906.5.30.257 (Paralectotype) B.M. (N.H.) 1966.3.9.92-93 B.M. (N.H.) 1966.3.9.88, 96-106 B.M. (N.H.) 1966.3.9.87 B.M. (N.H.) 1966.3.9.90-91 B.M. (N.H.) 1966.3.9.89 B.M. (N.H.) 1966.3.9.86 B.M. (N.H.) 1966.9.9.85 B.M. (N.H.) 1901.6.24.90 (Lectotype) B.M. (N.H.) 1966.3.9-94-95 UGANDA Entebbe .... Napoleon Gulf, near Jinja . Ekunu Bay Pilkington Bay Off Ramafuta Island (Buvuma Channel) . KENYA Naia Bay (Kavirondo Gulf) . Nanga Bay (Kavirondo Gulf) Off mouth of Nzoia River . LAKE VICTORIA Locality unknown Locality unknown Collector Degen E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. Sir H. H. Johnston E.A.F.R.O. Haplochromis pseudopellegrini sp. nov. (Text-fig. 8) HOLOTYPE : an adult male 139 mm. standard length, from Pilkington Bay (Uganda). B.M. (N.H.) no. 1966.2.21.2. Named pseudopellegrini because of its resemblance to H. pellegrini Regan. DESCRIPTION : based on 17 specimens (including the holotype) 98 to 150 mm. standard length. Depth of body 29-0-33-5 (M = 30-9) per cent of standard length, length of head 32-4-37-0 (M = 33-8) per cent. Dorsal head profile gently curved (rarely straight) but interrupted by the prominent premaxillary pedicels. Preorbital depth 19-1-22-4 (M = 20-4) per cent of head length, least interorbital width 20-4-27-1 (M = 23-9) per cent. Snout a little longer than broad (1-2-1-3 times), its length 35-1-39-0 (M =37-1) per cent of head; eye diameter 18-5-25-4 (M = 20-6), depth of cheek 24-0-29-6 (M = 26-8) per cent. Caudal peduncle 16-3-20-8 (M = 18-3) per cent of standard length, 1-3-2-0 (modal range 1-6-1-7) times as long as deep. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 57 1 CM Haplochromis pseudopellegrini ; this specimen shows extreme development of an obliquely truncate caudal fin. Drawn by Barbara Williams. Mouth slightly oblique, lower jaw projecting a little. Premaxilla slightly ex- panded medially. Lower jaw 46-8-53-4 (M = 49-1) per cent of head, 2-0-2-6 (modal range 2-1-2-4) times as long as broad. Posterior tip of the maxilla reaching, or almost reaching the vertical through the anterior orbital margin, extending a little beyond this point in a few specimens. Gill rakers : of variable form, from relatively slender to moderately stout (reduced to short knobs in one specimen), the upper four sometimes flat ; 9 (rarely 8 or 10) on the lower part of the first gill arch. Scales: ctenoid; lateral line with 32 (f.4), 33 (f.7), 34 (f.4) or 35 (f.i) scales. Cheek with 4 (less commonly 3, rarely 5) rows. Six or 7 (rarely 7^) scales between the dorsal fin origin and the upper lateral line, 7 (occasionally 6, rarely 8) between the pectoral and pelvic fin bases. Fins. Dorsal with 23 (f.i), 24 (f.4) or 25 (f.i2) rays, comprising 15 (f.g) or 16 (f.8) spinous and 8 (f.i), 9 (f.i2) or 10 (f-4) branched rays. Anal with n (f.io) or 12 (f.7) rays, comprising 3 spines and 8 (f.io) or 9 (f.7) branched elements. Pectoral 22-7-28-0 (M = 24-9) per cent of standard length. First branched ray of pelvic fin slightly produced in both sexes, proportionately more so in adult males. Caudal scaled over its proximal half ; in most specimens the fin is truncate or subtruncate but in a few the lower half, or the ventro-posterior corner, slopes obliquely forward and may also be rounded. Teeth. The outer row of teeth in both jaws is composed of slender and slightly recurved unicuspids ; there are 35-52 (M = 44) teeth in the outer row of the upper jaw. Inner teeth are also unicuspid, and are implanted at an angle varying from near vertical to almost horizontal, the latter condition being most common. There are 2 (rarely 3) rows of inner teeth in the upper jaw, and I or 2 rows in the lower. Osteology. The neurocranium of H. pseudopellegrini shows some similarity with 58 P. H. GREENWOOD that of H. mento and other species in the " prognathus " group (see Greenwood, 1962, and p. no), but at the same time it retains characteristics of the more generalized skull seen in H. serranus and its allies. In this respect it resembles the neurocranium of H. prognathus, but is slightly less " mento "-like. In general appearance and proportions it is similar to the neurocrania of H. bayoni and H. dentex but lacks the characteristically decurved ethmo-vomerine region of these species (see Green- wood, op. cit.}. The premaxilla is moderately beaked, that is, the dentigerous part of the bone is somewhat expanded medially ; the pedicels are short, being about two-thirds the length of the dentigerous arm. The lower pharyngeal bone is triangular, its dental surface is as long as broad or slightly longer than broad. Lower pharyngeal teeth are slender and compressed, those in the two median rows are slightly coarser than their lateral congeners ; the teeth are arranged in 22-24 rows. Vertebral counts in 14 specimens are : 13+17 (f.4), 13 + 18 (f.i), 14 + 17 (f.g), giving totals of 30 and 31. Coloration : Live colours are known only for adult and sexually active males, which have the dorsal surface of the head and body dark brown, the flanks and belly golden-yellow overlain by an orange-red flush on the chest and anterior flanks, and also on the operculum. Dorsal fin dark neutral with a slight orange flush. Caudal dark neutral, with a reddish flush at the base and over the ventral quarter of the fin. Anal neutral, with yellowish-red ocelli. Pelvics are sooty. Preserved material : Males (adult and active] : ground colour brownish overlying silver, silvery-yellow on the chest and belly, and on the operculum ; rest of head (including the branchiostegal membrane) brownish with very faint traces of a lachry- mal blotch. Dorsal fin hyaline, with a faint, narrow black band running from the tip of the eleventh spine to about the middle of the last branched ray (i.e. curving gently downwards). Caudal brownish to hyaline, dark on the proximal half. Anal hyaline, with faint traces of one or two whiteish-grey ocelli. Pelvics sooty, darkest on the anterior half. Adult but quiescent males are light brown dorsally, shading to silvery below, some showing a faint but broad and dark midlateral stripe, and a fainter upper lateral band above the upper lateral line. The two lateral bands are connected by 4-6 vertical bars, which extend ventrally a little below the midlateral band ; where the lines intersect, the lateral one is diffusely expanded. A faint lachrymal blotch is present below the anterior part of the orbit. All fins are yellowish-brown, the soft dorsal and the proximal part of the caudal are often darkly maculate. Anal with 2 or 3 faint, whiteish ocelli. Pelvics variable, from yellowish-brown to sooty. Ecology. Habitat. The species is recorded from four localities only. Two of these are shallow, sheltered bays, one is a fairly exposed, offshore and deep (90 ft.) channel, and the fourth is not fully documented except for a note that the nets were set in water about 20 ft. deep. In all, the substrate is of organic mud. Food. Fourteen specimens were examined, and of these only two contained food, very fragmentary and generically unidentifiable fish remains. Breeding. Little information is available on the reproductive biology of this species. The sex of the smallest specimen (98 mm. S.L.) is indeterminable ; the A REVISION OF THE LAKE VICTORIA H A PLOCHRO M I S SPECIES 59 others (132-153 mm.) are all adults, and only two are females. The two largest fishes are males. Affinities. In general appearance H. pseudopellegrini closely resembles H. pellegrini ; however, it reaches a much larger adult size, and the preserved coloration of adult males is much lighter (uniformly dark brown, nearly black in H. Pellegrini, light brown over silver in H. pseudopellegrini}. The species also differ in certain morphometric characters. Haplochromis pseudopellegrini has a longer snout (35-1-39-0, M = 37-1% head, cf. 30-8-36-0, M = 34-0%), a longer and more slender caudal peduncle (16-3-20-8, M = 18-3% standard length, cf. 13-2-17-8, M = 15-4% ; length/depth ratio 1-3-2-0 [modal range 1-6-1-7], cf- I M=34'4%), slightly narrower interorbital region (20-4-27-1, M = 23-9% head, cf. 23-4-30-2, M = 27-4%), longer and narrower lower jaw (48-6-53-4, M =49-1% head, cf. 39-2-48-2, M = 44-4% ; length/breadth ratios 2-0-2-6 [modal range 2-1-2-4] cf. 1-5-2-3 [mode 2-0]), fewer teeth in the outer row of the upper jaw (35-52, M = 44, cf. 48-74, M = 62). From H. gowersi it differs in head shape (and neurocranial form), and in having a much shorter head (34-2-37-0, M = 33-8% standard length, cf. 35-8-38-4, M = 37-0%), a broader snout, a larger eye (18-5-23-4, M = 20-6% head, cf. I5'5-I9'3, M = 17-5%), a slightly shallower cheek (24-0-29-6, M = 26-8% head, cf. 27-8-33-3, M = 29-5%), and a longer caudal peduncle (16-3-20-8, M = 18-3% standard length, cf. I3-3-I7-6, M = i4-8%). From H. dentex, H. pseudopellegrini differs, superficially, by its less strongly de- curved dorsal head profile. At a deeper level, there are differences in the shape of the neurocranium, that of H. dentex having a sharply decurved ethmoid-vomer region (see Greenwood [1962], p. 168 and fig. 25) ; but in other respects, the neurocrania of the two species are similar. Other interspecific differences lie in the more numerous and closely set teeth of H. pseudopellegrini (35-52, M =44, cf. 32-48, M =36), its deeper body (29-0-33-5, M = 30-9% of standard length, cf. 24-6-29-5, M = 26-7), and the greater posterior extension of the maxilla in this species (posterior maxillary tip reaching anterior orbital margin or to below the eye, cf. not reaching the orbital margin) . From H. estor, H. pseudopellegrini differs, principally, in having a shorter head 60 P. H. GREENWOOD (32-4-37-0, M = 33'8% standard length, cf. 37-2-38-5, M = 37'8%), a shorter lower jaw (46-8-53-4, M =49-1% head, cf. 54-2-57-5, M =55-5%), and a longer caudal peduncle (16-3-20-8, M = 18-3% standard length, cf. 14-3-16-1, M = 15-8%). Neurocranial form in the two species differs, that of H. estor belonging to the " prognathus "group, whilst that of H. pseudopellegrini has stronger affinities with the " altigenis "-type (see p. no). The resemblance between H. pseudopellegrini and H . mento is probably the most distant of all. Osteologically, there is a clear-cut difference in neurocranial form (like that distinguishing H. pseudopellegrini and H. estor}, and in most specimens the external head shape is distinctive (cf. fig. 12 in Greenwood [1962] with Text-fig. 8 above) . Nevertheless, most cephalic morphometric characters are similar in the two species, although the snout of H . pseudopellegrini is broader (length/breadth ratio I-I-I-3 cf. 1-5-1-8 in H. mento). The outer teeth in H. mento are stouter and more strongly curved than those of H. pseudopellegrini ; the range for the number of outer upper jaw teeth overlaps in the two species, but the mean for H. pseudopellegrini is lower (44 cf. 52). Phyletically, H. pseudopellegrini appears to be a derivative from an H, guiarti- like stem, and thus shows relationship with H. bayoni and H. dentex. However, unlike these species it also shows relationship with both H . altigenis and H. pellegrini. Structurally, H. pseudopellegrini could represent an ancestral level in the evolution of H. altigenis. Museum and Reg. No. Locality Collector UGANDA B.M. (N.H.) 1966.3.9.299-310 Ekunu Bay .... E.A.F.R.O. B.M. (N.H.) 1966.3.9.296 Trawl in Buvuma Channel off Nasu Point . . . .- E.A.F.R.O. B.M. (N.H.) 1966.3.9.297 Pilkington Bay . . . . ~ E.A.F.R.O. B.M. (N.H.) 1966.2.21.2 Pilkington Bay . . . E.A.F.R.O. (Holotype) B.M. (N.H.) 1966.3.9.298 Sesse Islands . . . .-..-:; E.A.F.R.O. Haplochromis altigenis Regan, 1922 (Text-figs. 9 and 10) Paratilapia longirostris (part) : Boulenger, Cat. Afr. Fish., 3, 332. Pelmatochromis spekii (part) : Boulenger, 1915, op. cit., torn, cit., 417. Haplochromis altigenis Regan, 1922, Proc. zool. Soc. Londn., 175, PI. i. LECTOTYPE : a male, 186 mm. standard length from Bunjako (Uganda), collected by Degen. B.M. (N.H.) reg. no. 1906.5.30.294. DESCRIPTION : based on 25 specimens (including the lectotype and paralectotype), 100-202 mm. standard length. Depth of body 28-4-34-7 (M = 31-5) per cent of standard length, length of head 36-7-39-5 (M = 38-2) per cent. Dorsal head profile gently curved, rather variable in its shape, tending to slope more steeply in large fishes which therefore have more rounded profiles ; the two type specimens have the most strongly sloping head A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 61 FIG. 9. Haplochromis altigenis ; lectotype, about -57 times natural size. From Regan, Proc. zool. Soc. profiles (ca 40°) ; most other specimens lie in the range 30°-35°. The premaxillary pedicels are prominent, and clearly break the outline of the profile. Altogether, one is left with the impression of a heavy-headed fish. Preorbital depth 17-6-23-7 (M = 19-9) per cent of head length, least interorbital width 17-4-25-0 (M — 20-2) per cent. Snout 1-1-1-3 times as long as broad, its length 36-6-42-2 (M =39-3) per cent of head, eye diameter 16-9-21-7 (M = 19-2), depth of cheek 25-6-34-5 (M = 30-5) per cent. Cheek depth may show positive allometry with standard length in fishes > 180 mm. ; the three largest fishes have the deepest cheeks. Caudal peduncle 12-4-16-0 (M = 14-7) per cent of standard length, 1-2-1-4 (mode 1-3) times as long as deep. FIG. 10. Haplochromis altigenio, to show the usual head profile in fishes less than ca. 1 80 mm. S.L. Drawn by Barbara Williams. ZOOL. 15, 2. 4 62 P. H. GREENWOOD Mouth slope variable, from near horizontal to moderately oblique (20°-30°). Lips slightly thickened, the premaxilla expanded medially. Lower jaw generally projecting (but jaws equal anteriorly in some specimens), its length 51-0-55-8 (M = 52-7) per cent of head, 1-8-2-4 (modal range 1-9-2-0) times as long as broad. Posterior tip of the maxilla almost reaching the vertical through the anterior orbital margin (reaching this point in a few specimens) . Gill rakers : stout (finer in the two smallest fishes), the upper 1-4 sometimes flattened and anvil-shaped ; 8 or 9 (rarely 7 or 10) on the lower part of the first arch. Scales : ctenoid ; lateral line with 31 (f.7), 32 (£13) or 33 (£.4), cheek with 4-6 (mode 5) rows. Six to 8 scales between the upper lateral line and the dorsal fin origin ; chest and anterior belly scales small, 7 or 8 (rarely 9) between the pectoral and pelvic fin bases. Fins : Dorsal with 24 (f.8), 25 (f.i6) or 26 (f.i) rays, comprising 15 (f.22) or 16 (f.3) spinous and 9 (f.io) or 10 (f.15) branched rays. Anal with n (f.2), 12 (f.22) or 13 (f.i) rays, comprising 3 spines and 8 (f.2), 9 (f.22) or 10 (f.i) branched elements. Pectoral 20-1-26-2 (M = 22-6) per cent of standard length. Pelvics with the first branched ray produced, usually more so in males but almost as elongate in some females. Caudal subtruncate, scaled on its proximal half. Teeth. In the size range of fishes studied, all teeth in the outer row of both jaws are stout, unicuspid and curved. There are 40-60 (M = 50) teeth in the outer row of the upper jaw. The inner teeth of the two smallest fishes (90 and 119 mm. S.L.) are predominantly unicuspids, but some weakly tricuspid teeth occur in both jaws. In all other speci- mens, only unicuspids are found ; these are moderately large, curved and implanted obliquely. The inner tooth rows are often irregularly arranged, with 3 (rarely 4) rows in the upper jaw, and 2 or 3 in the lower. Osteology. The neurocranium of H. altigenis resembles that of H. bayoni, but is relatively broader in the otic region. Thus, although it shows some of the characters associated with the H. mento-type skull (see Greenwood, [1962], fig. 25) it still retains the curved preorbital profile, greater preorbital skull depth, and broad otic region of the more generalized neurocranium. In these characters it also resembles the neurocranium of H. pseudopellegrini. The dentary and premaxillary show no outstanding characters. The former is a stout bone, and the dentigerous surface of the latter is but moderately expanded medially (less so, for example, than in H. bayoni). The dentigerous surface of the lower pharyngeal bone is as broad as long but is broader than long in large fishes. Anteriorly this surface narrows rather abruptly so that the apex of the dentigerous triangle is produced into a narrow " stem ". The lower pharyngeal teeth are rather coarse, somewhat compressed and clearly but weakly cuspidate. Vertebral counts in 7 specimens are : 13+16 (£.5) and 13 + 17 (f.2), giving totals of 29 and 30. Coloration. Live colours are unknown. Preserved material : Males (adult and sexually active): ground coloration a very dark brown, almost black on the dorsal A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 63 surface and snout, and with a sooty overlay on the chest. Lower jaw and branchio- stegal membrane pinkish-brown ; margin of the preoperculum outlined in dark brown. Dorsal fin yellow-brown, lappets black, and with a faint black outline to the margin of the soft part. Caudal darker yellow-brown (darkest proximally). Anal yellow-brown, with 2 large, dead-white ocelli. Pelvics very dark brown, appearing black when folded. Pectorals dark brown (dark pectorals are unusual). Males (adult but sexually quiescent) have a light brown ground coloration, darker on the dorsal surface of the snout, head and body. A prominent, but narrow, dark lachrymal stripe runs from the lower anterior border of the eye, passes almost vertically downwards behind the posterior tip of the maxilla to end on the dentary ; a slightly broader dark bar lies immediately anterior to the vertical limb of the preoperculum. The lower jaw and branchiostegal membrane are very light brown ; the chest is somewhat dusky. The flanks and caudal peduncle are crossed by seven very faint, moderately broad dark bars ; these extend (on the flanks) from the dorsal fin base almost to the ventral body outline. Dorsal fin yellow-brown, the lappets black and the soft part darkly maculate. Caudal yellowish-brown, darkest on the proximal half. Anal dark yellow-brown. Pelvics dark brown, nearly black, on the anterior half, otherwise light yellow-brown. Male (immature) : essentially as for females (see below), but with black lappets on the dorsal fin, the caudal uniformly light but with some dark spots on the upper fifth, 3 faint whiteish ocelli on the anal, and a dusky overlay on the pelvics. Note on the coloration of the type specimens. Both these specimens are males, but both are now a pale silver, shading to white. One specimen (that illustrated by Regan, see fig. 9 above) has a very faint midlateral stripe, which is now much fainter than is shown in the figure. Both fishes have dusky pelvics. In one, there are five, large, dead- white ocelli (arranged in three rows) on the anal fin, and in the other there are four (in two rows). The difference in coloration between these specimens and those described above is, presumably, due both to time and to the fact that the types were not fixed in formol but in alcohol. Females (immature and adult) : ground coloration light brown, darkest dorsally, and shading to yellowish-silver on the chest and belly. Lachrymal and preopercular stripes are as described for males, but in addition there are two, faint, parallel dark bands across the snout. The lower jaw and branchiostegal membrane are light yellow-brown. A faint dark midlateral band runs along the flank from slightly behind the opercular margin to the base of the caudal fin (sometimes extending onto the fin itself), and is crossed by 8 to 10, moderately broad vertical bars which extend from the dorsal fin base to about the level of the pectoral fin. Dorsal fin light greenish-brown, the soft part darkly maculate. Caudal yellow-brown on its distal quarter to third, dark brown basally. Anal greenish-brown. Pelvics light yellow-brown, somewhat darker along the anterior margin. Three fishes (140, 148 and 149 mm. S.L.) caught on two occasions off Kisigala; Point, North Rome Island, show typical piebald coloration of black on silver; all three are immature females. These are the first known examples of piebald polychromatism amongst the piscivorous species-groups of Lake Victoria Haplo- chromis, although the phenomenon is recorded from other trophic and phyletic lines 64 P. H. GREENWOOD (see Greenwood, 1956, 1957, 1959, and p. 95 below). Ecology. Habitat. The species occurs over both hard and soft substrates in sheltered bays and gulfs, as well as off-shore in places where the water is relatively undisturbed ; apparently it does not occur at depths of over 50 ft. Food. Eleven of the 26 specimens examined (from 15 localities) contained food in the gut. In all cases this consisted of finely macerated fish remains, identifiable as cichlids in three specimens, and as Haplochromis species in six others. Breeding. Haplochromis altigenis is a female mouth breeder. Fishes less than 145 mm. S.L. are immature, or, in the upper levels of the range, show early signs of maturation. The largest specimen (202 mm. S.L.) is a female, but both sexes occur in the size range 170-194 mm. Affinities. At least superficially, four species, H. pseudopellegrini, H. estor, H. dichrourus and H. gowersi, closely resemble H. altigenis. A fifth species, H. squamulatus shows a more distant resemblance. Similarities between H. altigenis and H. dichrourus are considered elsewhere (see p. 68) ; the resemblances between these species may indicate a fairly close phyletic relationship. From H. pseudopellegrini, H. altigenis is distinguished by its longer head (36-7- 39-5, M = 37-2% of standard length, cf. 32-4-37-0, M=33-8%), deeper cheek (25-6-34-5, M =30-5% head, cf. 24-0-29-0, M = 26-8%), longer lower jaw (51-0- 55-8, M = 52-7% head, cf. 46-8-53-4, M = 49-1%) and its shorter and deeper caudal peduncle (12-4-16-0, M = 14-7% standard length, cf. 16-3-20-8, M = 18-3%, length/depth ratio 1-2-1-4 [mode 1-3] cf. 1-3-2-0 [modal range 1-6-1-7]). The neurocranium in the two species is generally similar. It seems that H. altigenis could have evolved from an H. pseudopellegrini-\ike ancestor. From H. estor, H. altigenis differs in its longer snout (36-6-42-2, M = 39'3% head, cf. 34-5-37-0, M =36-2%), smaller eye (16-9-21-7, M = 19-2% head, cf. 22-8-25-4, M = 24'5%), much deeper cheek (25-6-34-5, M = 30-5% head, cf. 22-8-25-4, M = 24-5%) and slightly shorter lower jaw (51-0-55-8, M = 52-7% head, cf. 54-2-57-5, M =55'5%). There are fairly marked differences between the neuro- crania of the two species, that of H. estor being of the " prognathus " type. Superficially, the resemblances between H. gowersi and H . altigenis are great, and some difficulty may be experienced in separating certain specimens of the two species. However, there are distinct differences between modal specimens of the two species, and there is a clear difference in neurocranial form ; the skull of H. gowersi is of the " prognathus " type. This difference in neurocranial shape probably accounts for the more declivous snout and dorsal head profile of H. altigenis, and the greater prominence of the premaxillary pedicels in this species. In addition to these qualitative differences, H. altigenis differs from H. gowersi in having the interorbital width equal to or slightly greater than the preorbital depth (interorbital less than preorbital in H. gowersi), a broader head (greatest width, measured at about the middle of the operculum, 41-5-45-5, M = 42-7% head, cf. 35-6-39-0, M = 36-8) ; the greater head breadth is also reflected in the broader lower jaw of H. altigenis. Haplochromis squamulatus and H. altigenis both have noticeably rounded head profiles, and very small chest and anterior belly scales. However, they differ in A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 65 several morphometric characters. Haplochromis altigenis has a longer head, narrower interorbital, longer snout, deeper cheek and a longer lower jaw (see p. 87). The neurocrania show several interspecific differences (that of H. squamulatus showing affinities with the neurocrania of H. martini and H. michaeli) and there are marked differences in preserved coloration. Considering the evidence available, both morphometric and osteological, it seems H. altigenis could have evolved from an H. pseudopellegrini-like. stem ; certainly its neurocranial form does not favour a closer association with H. estor and H. gowersi than with H. pseudopellegrini. STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. Locality Collector UGANDA B.M. (N.H.) 1906.5.30.294 Bunjako Degen (Lectotype) B.M. (N.H.) 1906.5.30.295 Bunjako ..... Degen (Paralectotype) B.M. (N.H.) 1966.3.9.220-227 Napoleon Gulf, near Jinja . . E.A.F.R.O. B.M. (N.H.) 1966.3.9.228-235 Karenia Beach, near Jinja . . E.A.F.R.O. B.M. (N.H.) 1966.3.9.215-216 Near Ramafuta Island (Buvuma Channel) . . E.A.F.R.O. B.M. (N.H.) 1966.3.9.219 Pilkington Bay . . . . E.A.F.R.O. B.M. (N.H.) 1966.3.9.218 Manadu Island . . . . E.A.F.R.O. B.M. (N.H) 1966.3.9.213-214 Off Entebbe harbour . . . E.A.F.R.O. TANZANIA B.M. (N.H.) 1966.3.9.217 Mwanza Harbour . . . E.A.F.R.O. LAKE VICTORIA B.M. (N.H.) 1966.3.9.211-212 Locality unknown . . . E.A.F.R.O. Haplochromis dichrourus Regan, 1922 (Text-fig, n) Paratilapia serranus (part) : Boulenger, 1915, Cat. Afr. Fish., 3, 334. Haplochromis dichrourus Regan, 1922, Proc zool. Soc. Londn., 178, fig. 6. HOLOTYPE : a juvenile female, 113-0 mm. S.L., from Buganga ; B.M. (N.H.) reg. no. 1906.5.30.265. DESCRIPTION : based on eight specimens (including the holotype), 84-186 mm. S.L. Depth of body 28-6-35-5 (M = 32-1) per cent of standard length, length of head 35.3-37-7 (M = 36-6) per cent. Dorsal head profile convex, sloping steeply (ca 40°) especially in the snout region ; premaxillary pedicels prominent and breaking the outline of the profile. Preorbital depth 16-4-21-0 (M = 18-7) per cent of head, least interorbital width 18-0-24-6 (M = 21-3) per cent. Snout 1-10-1-25 times as long as broad, its length 66 P. H. GREENWOOD 34-1-41-0 (M — 37-4) per cent of head, eye diameter 19-4-24-6 (M = 21-1), depth of cheek 23-0-30-6 (M = 27-6) per cent. Caudal peduncle 12-4-17-2 (M = 15-2) per cent of standard length, 1-1-1-5 times as long as deep. Mouth slightly oblique (20°-30°), lips thickened, premaxilla slightly expanded medially. Lower jaw projecting, its length 49-0-54-0 (M = 52-7) per cent of head, 1-8-2-6 (no distinct mode) times as long as broad. Posterior tip of maxilla reaching the vertical through the anterior margin of the orbit, or to below the anterior part of the eye. FIG. ii. Haplochromis dichrourus ; holotype, about -93 times natural size. From Regan, Proc. zool. Soc. Gill rakers : stout, the lower 2 or 3 reduced, the upper 3 or 4 flat and expanded in some specimens ; 8 or 9 (mode) on the lower part of the first gill arch. Scales : ctenoid ; lateral line with 32 (f.2), 33 (f.4) or 34 (f.2), cheek with 3 (f.i), 4 (f.i), 5 (f.5), or 6 (f.i) rows. Seven to 9 scales between the upper lateral line and the dorsal fin origin, 6-9 (mode 8) between the pectoral and pelvic fin bases ; scales on the chest and belly small. Fins : Dorsal with 25 rays, comprising 15 (f.i) or 16 (f-7) spinous and 9 (f-7) or 10 (f.i) branched rays. Anal with 11 (f.2), 12 (f.5) or 13 (f.i) rays, comprising 3 spines and 8 (f.2), 9 (f.5) or 10 (f.i) branched elements. Pectoral fin 21-2-27-3 (M = 23-7) per cent of standard length. Pelvics with the first, and to a lesser degree, the second branched rays produced, slightly so in females and markedly elongate in males. Caudal truncate or subtruncate. Teeth. In all specimens the outer teeth (numbering 48-70, M = 58 in the upper jaw) are unicuspid, slender and very strongly curved (the tips of the anterolateral premaxillary teeth are not visible when the specimen is viewed laterally) . The inner teeth in fishes > 149 mm. S.L. are all unicuspid, curved and obliquely implanted. In the four smaller specimens, the 113 mm. fish (holotype) has mostly tricuspids in the upper jaw and an admixture of tri- and unicuspids (the latter A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 67 predominating) in the lower jaw ; the 84, 101 and 121 mm. specimens have a mixture of unicuspids and weakly bicuspids in both jaws ; as in the larger fishes, these teeth are curved and obliquely implanted. The inner teeth are arranged in 1-3 series in the upper jaw, and in 2 or 3 in the lower (in a single row in one fish). Osteology. With so few specimens available, it has not been possible to prepare a complete skeleton, but radiographs have been studied. The lower pharyngeal bone is fine, with a triangular dentigerous surface which is equilateral or slightly broader than long. The lower pharyngeal teeth are slender, with cylindrical necks and slightly compressed weakly cuspidate crowns, and are arranged in 20-22 rows. Vertebral counts in 5 specimens are : 13+16 (f.i) and 13 + 17 (f. 4), giving totals of 29 and 30. Ecology. Habitat. The species has a wide depth range, from 10-90 feet. It is found in both sheltered and exposed places (including beaches) over sand and shingle substrata. Food. Of the five specimens examined (from four localities) four contained food in the gut. In three fishes this consisted solely of fish remains (Haplochromis sp.), and in the fourth fish-remains (a cichlid) and fragments of larval insects (one probably a dragon-fly, the other Ephemeroptera) . Breeding. Little information is available ; three of the smallest fishes (101, 113 and 121 mm. S.L.) are immature females. An unusual feature of this species is the coloration of females, which, at least in preserved specimens, seems to be as polychromatic as that of males, and certainly more complex than the female coloration of other species (except, perhaps, H. chromogynos) . Coloration. The only information on live colours is provided by brief field notes made on a sexually active male caught in deep water near Soswa island. In this fish the median fins and back were described (by Mr. J. D. Kelsall) as vivid flame red, the belly as jet black. Preserved Colours : Males. The most extreme pattern is shown by a fish which, although adult, is in an early stage of sexual activity. The upper part of the head (above the level of the lower orbital rim) brownish, snout yellowish and crossed by a thin, well-defined and dusky bar ; running parallel with the premaxillary pedicels are a pair of short, dark bars, each of which (at about the level of the pedicel tips) broadens somewhat and curves sharply at right angles to meet the orbit. The lower part of the head (cheeks, preorbital and the entire operculum) jet black. This colour extends onto the chest but does not reach above the level of the pectoral fin base. Immediately behind the pectoral fin, the margin of this black area curves ventrally but rises again above the vent so that the posterior half of the body is black on its lower half. On the caudal peduncle the dark area rises again to cover about the entire lateral aspect. Above the black areas the body is brownish. Dorsal and caudal fins are dark yellow-orange, with a narrow black crescent at the caudal base. The anal fin is a similar colour, with two ocelli, each outlined by a narrow black margin. Pelvics jet black, except for the yellowish innermost ray of each side. In two other males (both sexually active, with convoluted testes and therefore thought to be more mature than the specimen above) the head and anterior half of 68 P. H. GREENWOOD the body are light orange-brown the posterior half and the caudal peduncle dusky brown. A well-defined black band crosses the snout (just above the upper lip) to the anterior margin of the preorbital bone. Another transverse black band at the level of the pedicel tips, runs from orbit to orbit. An interrupted band extends from the upper posterior margin of the orbit across the nape. A dusky, nearly vertical lachrymal stripe is present. The branchiostegal membrane and chest are black, but the belly is brown. Dorsal fin yellowish, the soft part is maculate, the spots being clear. Anal yellowish but dusky along its base, and with two, hyaline ocelli (set, in one specimen, in the dark basal zone). Caudal yellowish-grey, but with a broad, dusky band at its base. The pelvics are black except for the distal half of the membrane between the last two rays. Females (adult and immature] : body and head dark grey-brown with a blueish hue. Branchiostegal membrane and chest dusky as are the lower parts of the operculum, preoperculum and interoperculum. Faint traces of snout and lachrymal bands (like those of males) are visible, as is a nuchal bar. Dorsal fin greyish to light sooty, the lappets dark. Anal pale orange with a single, well-defined, dead- white ocellus. Caudal with a dark base, the upper half grey and the lower orange. Pelvics light sooty, the pigment most concentrated on the anterior half of the fin. Affinities. The anomalous coloration of female H. dichrourus was commented upon above. Apart from the relatively infrequent piebald females in certain species, and the occurrence of a piebald coloration as the usual one in females of H. chromo- gynos (see Greenwood, 1959), I know of no other species in Lake Victoria with colourful females. In its gross morphology, H. dichrourus closely resembles H. altigenis although there are marked differences in the preserved coloration of the females. Both species have very small chest scales. They differ in that the maxilla of H. dichrourus reaches further posteriorly (rarely reaching the orbit in H. altigenis) and the pre- maxilla is not markedly expanded medially. Also, the teeth in H. dichrourus are more strongly curved, so that when viewed laterally, their tips are hidden ; in H. altigenis, the tips can always be seen. The two smallest fishes resemble specimens of H. Pellegrini of a similar size. They are, however, distinguished by their more slender and much more strongly recurved outer jaw teeth (in H. pellegrini these teeth have a gentle curvature confined to the distal part), and by having the maxilla extending further posteriorly (to below the anterior orbital margin). Also, in H. dichrourus the premaxillary pedicels are more prominent, and there are differences in preserved coloration. Phyletically, H. dichrourus could be related to H. altigenis and H. pellegrini. STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. Locality Collector UGANDA B.M. (N.H.) 1906.5.30.265 Buganga ..... Degen (type) B.M. (N.H.) 1966.3.9.186 J in j a, off golf course . . . E.A.F.R.O. B.M. (N.H.) 1966.3.9.187 Katebo . ;.:* - . gj$* . E.A.F.R.O. B.M. (N.H.) 1966.3.9.188-9 Karenia, Napoleon Gulf . . E.A.F.R.O. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 69 TANZANIA B.M. (N.H.) 1966.3.9.185 B.M. (N.H.) 1966.3.9.183-4 Off Soswa Island Beach N. of Majita J. D. Kelsall E.A.F.R.O. Haplochromis paraguiarti sp. nov. (Text-fig. 12) HOLOTYPE : an adult male 130 mm. standard length, from a beach near Nasu Point, Buvuma Channel (Uganda); B.M. (N.H.) reg. no. 1966.2.21.6. Named paraguiarti because of its close superficial resemblance to H. guiarti. DESCRIPTION: Based on 31 specimens (including the holotype), 70-156 mm. standard length. Depth of body 31-7-36-6 (M = 34-1) per cent of standard length, length of head 33-0-36-8 (M = 34*8) per cent. Dorsal head profile straight or very slightly curved, the premaxillary pedicels forming a slight prominence. Preorbital depth 14-5-19-5 (M = 16-9) per cent of head length, least interorbital width 22-9-27-7 (M = 25-3) per cent. Snout as long as broad or slightly longer (1-2 times), its length 31-0-36-2 (M = 33-5) per cent of head, eye diameter 23-8-28-6 (M = 26-2), depth of cheek 20-0-26-1 (M = 23-8) per cent. Caudal peduncle 14-7-18-6 (M = 16-6) per cent of standard length, 1-3-1-9 (modal range 1-3-1-5) times as long as deep. Mouth horizontal or slightly oblique, the jaws equal anteriorly. Lower jaw 41-0-48-6 (M = 45-0) per cent of head, 1-7-2-4 (modal range 1-9-2-1) times as long as broad. Posterior tip of the maxilla reaching the vertical through the anterior orbital margin, occasionally extending to below the eye. Premaxilla slightly expanded medially. 1 CM FIG. 12. Haplochromis paraguiarti. Drawn by Barbara Williams. 70 P. H. GREENWOOD Gill rakers : moderately stout (relatively slender in fishes less than 90 mm. S.L.), the upper 4-6 flat and sometimes branched in fishes over 115 mm. S.L., the lower i or 2 reduced ; 8-10 (mode 9), rarely ir, on the lower part of the first gill arch. Scales : ctenoid ; lateral line with 31 (f.4), 32 (f.i2), 33 (f.i2) or 34 (£3) scales, cheek with 3 or 4 (rarely 2 or 5) rows. Five or 6 (rarely 6|) scales between the upper lateral line and the dorsal fin origin, 6 or 7 between the pectoral and pelvic fin bases. (Because many specimens were damaged during capture by gill-nets, the last two counts are based on only 17 specimens). Fins : Dorsal with 23 (f.i), 24 (f.8), 25 (f.ig) or 26 (£.3) rays, comprising 15 (f.2i) or 16 (f.io) spinous and 8 (f.i), 9 (f.i-5) or 10 (f.i5) branched rays. Anal with n (f.6), 12 (f.23) or 13 (f.2) rays, comprising 3 spines and 8 (f.6), 9 (f.23) or 10 (f.2) branched elements. Pelvic fins with the first unbranched ray slightly produced in the adults of both sexes, but proportionately more so in males. Pectoral 24'3~30'0 (M=26-3) per cent of standard length. Caudal truncate or subtruncate, scaled on its basal half to two-thirds. Teeth : The form of the teeth in the outer row of both jaws shows some correlation with standard length. Only bicuspids (with a few tricuspids posterolaterally in the upper jaw) are found in fishes 70-75 mm. S.L. Unicuspids anteriorly, with bicuspids laterally are characteristic of most fishes 84-113 mm. S.L. (and in one exceptional individual 144 mm. S.L.). In most specimens over 112 mm. S.L., only unicuspids occur, but this is also the condition in a few fishes between 99 and 112 mm. S.L. The unicuspids are moderately stout teeth, and are only slightly curved. There are 42-62 (M = 54) outer teeth in the upper jaw. Teeth in the inner rows are relatively large ; tricuspids predominate in fishes of all sizes, but in specimens over 120 mm. S.L. weakly tricuspids are commoner than the distinctly tricuspid ones, and some unicuspid teeth also occur (especially in the outer- most row of the inner series) . There are 3 or 4 inner rows in the upper jaw, and 2 or 3 (less frequently I row) in the lower. Osteology : The neurocranium of H . paraguiarti is identical with that of H. acidens (see p. 75). It differs somewhat from the presumed generalized piscivore skull of H. guiarti, and shows some of the characters found in the more specialized type of H. prognathus. It is, in fact, almost intermediate between the two types. The premaxilla has a less pronounced medial expansion than in H. acidens, but as in that species it has long pedicels (as long as the horizontal dentigerous arms) which meet the horizontal arms at an appreciable angle. In these characters the premaxilla of H, acidens differs from that bone in the generalized species like H. guiarti and H. serranus. The lower pharyngeal bone is slender, its dentigerous surface slightly broader than long. The teeth are fine, slender, compressed and clearly bicuspid, only those in the posterior one or two rows of the median series are enlarged. There are 26-30 (mode 28) rows of lower pharyneal teeth. Vertebral counts for 6 specimens are : 13+17 (f-4) and 14 + 16 (f.2), giving a total of 30. Coloration : live colours unknown. Preserved material : Males (adult and sexually active) : ground colour dark sooty, almost uniformly so except for a lighter A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 71 (dusky gold) patch on the midflank, crossed by four, faint and narrow vertical stripes. Branchiostegal membrane dark (darker below the opercular series than between the jaws). A very faint, near vertical lachrymal stripe runs from the anterior orbital margin to behind the posterior tip of the maxilla. Dorsal fin dusky, lappets black, the soft dorsal maculate. Caudal dark. Anal dusky, but lighter than the dorsal except for a narrow band along the base, and over the spinous part ; two or three greyish ocelli present. Pel vies dusky, the proximal threequarters almost black, the distal quarter lighter. Males (adult but quiescent] : ground colour dusky silver-grey, darkest dorsally ; cheek bright silver. A faint lachrymal stripe present. Dorsal fin greyish-dusky, darkly maculate between the last four spines and all the branched rays. Caudal dark. Anal dark hyaline, with two or three greyish ocelli. Pel vies sooty, darkest on the leading edge. Females (adult) : ground colour silver grey, darker dorsally : cheek bright silver (i.e. like quiet males but lighter and brighter). Dorsal fin dark hyaline, with dark spots on the soft part. Caudal dark hyaline, weakly to distinctly maculate. Anal and pelvic fins hyaline. Females (juvenile] : Two small (75 mm. S.L.) specimens have colours like those of adult females, but with traces of 7 or 8 vertical bars on the flanks and caudal ped- uncle ; these bars reach the dorsal outline but do not extend ventrally below the level of the pectoral fin base. Ecology. Habitat. Most records of H. paraguiarti are from exposed, wave- washed beaches where the substrate is of sand, rock or shingle. There are, however, a few records from more sheltered beaches and areas, but with one exception (a mud bottom) the substrate was hard, and the depth invariably never more than 30 ft. Food. Twenty specimens (from 12 localities) were examined. Of these, 17 con- tained ingested material. One individual contained fish remains (very fragmentary and unidentifiable, even to family) together with a little macerated phanerogam, tissue. Eleven others yielded either macerated plant tissue (f-4), or a mixture of plant tissue and insect fragments (larval Ephemeroptera) ; five specimens contained only fragments of larval Ephemeroptera. Breeding. Little information is available on reproduction in this species. Fishes less than 95 mm. S.L. are immature, as is one larger specimen (a male, 100 mm. S.L.). The largest fish (156 mm. S.L.) is a male. Affinities. Superficially, H. paraguiarti resembles H. guiarti both morphologically and trophically, although H. guiarti apparently includes a greater proportion of fish in its diet. Morphologically the species may be distinguished by the straight dorsal head profile of H. paraguiarti (gently curved in H. guiarti], its longer head (33-0-36-8, M =34-8 per cent standard length cf. 29-5-33-8, M =31-4 per cent), somewhat broader snout, and fewer, coarser outer teeth (42-60, M = 53, cf. 48-74, M = 62 is the upper jaw). Live colours are unknown for H. paraguiarti, but the coloration of preserved, sexually active males is noticeably darker than that of H. guiarti males. Osteological differences, especially in neurocranial shape, are discussed above (p. 70). 72 P. H. GREENWOOD Haplochromis paraguiarti also resembles, rather closely, H. acidens ; the species are compared on p. 76. Some specimens of H. prognathus, a rather variable species, resemble H. para- guiarti but there are several differences which serve to distinguish even these superficially similar individuals ; H. paraguiarti has a shorter head (33-0-36-8, M = 34-8 per cent of standard length, cf. 35-5-38-4, M — 36-9 per cent), a shallower preorbital (14-5-19-5, M = 16-9 per cent head, cf. 18-8-23-1, M =20-8 per cent), shorter snout (31-0-36-2, M = 33-5 per cent head, cf. 33-4-39-0, M = 37-0 per cent), and a larger eye (23-8-28-6, M = 26-2 per cent head, cf. 20-0-25-0, M = 22-6 per cent). Differences in neurocranial form between the species are also distinctive, although the neurocranium of H. paraguiarti is intermediate between the more generalized H. guiarti type and the relatively elongate skull of H. prognathus (see p. in). When all characters are considered, H. paraguiarti, despite its superficial resem- blance to H . guiarti, is probably more closely related to H. prognathus. It appears to possess the structural characters of the ancestral species or species group from which piscivorous predators like H. prognathus, H. bartoni and H. mandibularis were derived. Outside the piscivorous predator group, it shows very close relation- ship with H. acidens (see p. 76). STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. B.M. (N.H.) 1966.3 B.M. (N.H.) 1966.2 (Holotype) B.M. (N.H.) 1966.3 B.M. (N.H.) 1966.3 B.M. (N.H.) 1966.3 B.M. (B.H.) 1966.3. B.M. (N.H.) 1966.3 B.M. (N.H.) 1966.3 B.M. (N.H.) 1966.3 9-324 21.6 9-338-344 9.313-314 9.320-323 9-333-337 9.317-319 9.312 9-325-330 Locality UGANDA Beach near Jinja Beach near Nasu Point (Buvuma Channel) . Beach near Nasu Point Fisherman's point near Jinja Near Grant Bay Entebbe Harbour Buka Bay Katebo .... Beach near Hannington Bay Collector E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. B.M. (N.H.) 1966.3.9.316 KENYA Kamaringa (Kavirondo Gulf) E.A.F.R.O. B.M. (N.H.) 1966.3.9.311 B.M. (N.H.) 1966.3.9.315 TANZANIA Beach N. of Majita Majita Beach E.A.F.R.O. E.A.F.R.O. B.M. (N.H.) 1966.3.9.331-332 LAKE VICTORIA Locality unknown E.A.F.R.O. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 73 Haplochromis acidens sp. nov. (Text-fig. 13) Haplochromis percoides (part) : Boulenger, 1915, Cat. Afr. Fish., 3, 296 (specimens from Kakindu, Victoria Nile, B.M. [N.H.] reg. no. 1911.3.3.82-3). Paratilapia serranus (part) : Boulenger, 1915, op. cit., torn, cit., 334 (specimens from Ripon Falls, nr. Jinja, B.M. [N.H.] reg. no. 1911.3.3.20-1). Haplochromis nigrescens (part) : Regan, 1922, Proc. zool. Soc. Londn., 172 (excluding his tentative synonymy of Astatotilapia roberti Pellegrin, and the synonymy of Paratilapia parvidens Blgr., for which see Greenwood, 1959, p. 194). NOTES ON SYNONYMY. It is obvious that Regan's (1922) redescription of H. nigrescens (Pellegrin), 1909 was influenced by Pellegrin's figure of that species (Pellegrin, 1910, Mem. soc. Zool. France, 32, pi. 14, fig. 3). The fish depicted certainly does resemble a specimen of the species here described as H. acidens, but the type specimen of Pellegrin's Astatotilapia nigrescens more closely resembles Haplochromis flavipinnis (personal observations). Boulenger (1915) actually synonymized A . nigrescens Pellegrin with H. percoides Blgr., 1906 but this action was unacceptable to Regan, who resurrected the species as Haplochromis nigrescens. Certainly there are similarities between A . nigrescens type specimen and H. percoides, but my study of the type was revealed several differences (see appendix for a re- description of this specimen and further comments on its affinities). In earlier papers I had accepted Regan's resurrection of H. nigrescens, and used specimens identified by him as a basis for comparing other Lake Victoria species with H. nigrescens ; thus, where H . nigrescens is mentioned in those papers, the name should now be corrected to read H. acidens. Two of the three Lake Victoria specimens examined by Regan undoubtedly can be referred to H. acidens, but I am uncertain about the identity of the third fish. It is the smallest specimen, and because so little is known about small fishes of this and related groups, diagnostic characters applicable to larger individuals cannot be used with confidence. In many respects, this specimen resembles larger individuals of H. prognathus. HOLOTYPE : an adult male, 108 mm. standard length, from Kisumu Kavirondo Gulf (Kenya), B.M. (N.H.) reg. no. 1966.2.21.1. Named acidens from its sharp, needle-like teeth. DESCRIPTION : based on 17 specimens 67-128 mm. S.L., including the holotype and two of the specimens examined by Regan (B.M. [N.H.] reg. no. 1911.3.3.20-1). Depth of body 34-6-40-3 (M — 38-1) per cent of standard length, length of head 34-0-36-7 (M = 35-4) per cent. Dorsal head profile gently concave (clearly so in most specimens but weakly concave in a few), sloping fairly steeply ; premaxillary pedicels not prominent. Preorbital depth 15-2-19-2 (M = 17-1) per cent of head, least interorbital width 21-4-28-2 (M = 24-5) per cent. Snout as long as broad or 1-1-1-2 times longer, its length 31-8-37-2 (M =34-2) per cent of head, eye diameter 23-0-29-0 (M =26-4), depth of cheek 23-0-28-2 (M = 25-7) per cent. 74 P. H. GREENWOOD Caudal peduncle 14-8-17-9 (M — 16-6) per cent of standard length, 1-1-1-5 (mode 1-3) times as long as deep. Mouth slightly oblique, lips somewhat thickened, premaxilla with a slight median expansion. Jaws equal anteriorly, the lower 44-0-50-2 (M = 46-9) per cent of head, 1-9-2-4 (rare), mode 2-1, times as long as broad. Posterior tip of the maxilla reaching the vertical through the anterior orbital margin, occasionally not quite reaching this point. FIG. 13. Haplochromis acidens. Drawn by Barbara Williams. Gill rakers : moderately slender to slender, the lower 1-4 reduced, the upper 3 or 4 flattened and lobed in some fishes ; 10 (rarely 9 or n) on the lower part of the first gill arch. Scales : ctenoid ; lateral line with 31 (f.i), 32 (f.8), 33 (f-7) or 34 (f.i), cheek with 3 (rarely 4) rows. Five and a half to 7 (mode 6) scales between the dorsal fin origin and the upper lateral line, 5 or 6 (mode), rarely 6| or 7 between the pectoral and pelvic fin bases. Fins : Dorsal with 23 (f.i), 24 (f.7) or 25 (f.g) rays, comprising 15 (f.12) or 16 (f.5) spinous and 7 (f.i), 9 (f.n) or 10 (f.5) branched rays. Anal with n (f.7) or 12 (f.io) rays comprising 3 spines and 8 or 9 branched elements. Pectoral 22-7-28-0 (M = 24-9) per cent of standard length. Pelvics with the first two branched rays slightly produced, relatively more so in adult males. Caudal truncate to subtruncate, scaled on its proximal half to two-thirds. Teeth. In the outer row of both jaws the teeth are tall, slender and slightly curved. Fishes between 67 and 90 mm. S.L. have unicuspid teeth anteriorly, and weakly bicuspid teeth laterally and posterolaterally. Larger fishes have only unicuspids in the outer row. The anterior teeth in the upper jaw of the largest specimens may be relatively enlarged (i.e. stouter and longer than the lateral teeth). There are 40-64 (M = 48) teeth in the outer, upper row. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 75 The inner rows of the upper jaw are composed of tricuspid teeth in fishes < 90 mm. S.L. and a mixture of uni- and tricuspids in larger fishes ; there is an increased proportion of unicuspids in specimens > 115 mm. S.L. In fishes over TOO mm. S.L. the outermost row of the inner series is often composed of unicuspids and the re- maining rows of tricuspids. The inner rows are widely spaced in most fishes so that the teeth form a broad band anteriorly and anterolaterally. There are 3 or 4 rows of teeth in the upper jaw. In the lower jaw of most specimens less than 120 mm. S.L., only tricuspid teeth are found, but some unicuspids also occur in fishes of this size group. Larger in- dividuals have a mixture of tri- and unicuspid or tri- and weakly tricuspid teeth ; only rarely are all the lower, inner teeth unicuspid. There are 2 (mode) or 3 rows of inner teeth in this jaw. Osteology. The neurocranium of H. acidens departs from that of H, guiarti towards the H. mento type (see Greenwood, 1962) . The slope of its anterior dorsal profile (from vomer tip to the origin of the supraoccipital crest) is less steep than in H. guiarti, and the maximum width (across the otic region) is relatively less, as is its greatest depth (exclusive of the supraoccipital crest). In all these characters, H. acidens is intermediate between H. guiarti and H. prognathus, a species whose neurocranium could provide a basic " bauplan " from which the more extreme " mento "-type evolved. The premaxilla is somewhat beaked, with the dentigerous surface expanded and protracted medially ; the pedicels are elongate, being almost as long as the horizontal arms of the bone. The lower pharyngeal bone is fine, and has the dentigerous surface broader than long. Lower pharyngeal teeth are arranged in 24-28 rows, and are slender, com- pressed and distinctly cuspidate, with those in the two median rows somewhat coarser than the others. Vertebral counts from 12 specimens are : 13+16 (f-4), 13 + 17 (f-7) and 14 + 16 (f.i), giving totals of 29 and 30. Coloration. Live colours : Males (sexually active) have a slatey-blue ground colour, darkest dorsally. Dorsal fin smokey-grey with a pale red margin and a deep red flush on the soft part. Caudal dark grey with deep red streaks between the rays. Anal dark hyaline with a dull maroon flush, and orange-red ocelli. Pel vies black. Females (adult and quiescent) : dorsal surface yellowish-grey shading to silver on the flanks and belly. Dorsal fin hyaline but with a yellowish base. Anal and caudal dark hyaline. Pelvics hyaline. Preserved material : Males (adult and sexually active) : ground colour brown (darker than in females and juveniles), darkest dorsally and on chest, belly and ventral half of the caudal peduncle ; six to eight vertical bands cross the flanks and caudal peduncle, each band broadening slightly below the level of the upper lateral line, and merging ventrally with the dark body coloration, but remaining discrete dorsally. A narrow, dark lachrymal stripe runs from the anterior border of the eye, sloping backwards to pass behind the posterior tip of the maxilla ; a dark narrow stripe outlines the vertical limb of the preoperculum. The spinous dorsal is greyish, the soft part darker on its basal two-thirds, maculate over the posterior half. Caudal 76 P. H. GREENWOOD maculate on its proximal half to two-thirds. Pelvics dusky, particularly on the basal half ; membrane covering the spine colourless. Anal greyish to hyaline, with two, ill-defined and dark grey to dusky ocelli. Males (sexually quiescent) : as above but all stripes and bars are much fainter and less well-defined, and the ventral body surface is not noticeably darker than the dorsum, although still darker than the mid-flank region. Only the basal half of each pelvic fin is dusky, and the anal ocelli are almost invisible. Immature fishes have the same coloration as females. Ecology. Habitat. The majority of specimens is from a sheltered habitat in the Kavirondo Gulf near Kisumu. At this place the water was about 10 ft. deep, and the substrate of mud ; fairly dense stands of submerged plants were common in the area. The other specimens are recorded as being from " Jinja, Ripon Falls ". I am presuming that this locality is in the Napoleon Gulf, that is, above the falls. The area, before the submergence of the Falls, was sheltered, with a hard substrate and, at least close inshore, fairly dense plant stands. Food. All sixteen specimens examined came from the Kisumu locality, but were caught on different occasions ; fifteen specimens contained ingested material in the stomach and intestines. Every gut had, as its major content, finely macerated phanerogam tissue and varying amounts of epiphytic diatoms. Some specimens also contained a flocculent, grey-green mass (as seen in preserved material) thought to be bottom debris, principally blue-green algae. In addition, one gut yielded a few fish bones, one a number of insect eggs (apparently undigested), one a mass of tissue (thought to be the foot of a gastropod) and some fragments of larval Ephemeroptera, and another, fragments of unidentifiable insects. The phanerogam tissue is very finely divided, and a larger proportion seems to be digested than is the case in other phytophagous species examined. The apparently vegetarian diet of H. acidens is at variance with the dentition and general facies which are essentially those of a piscivorous predator. The length of the gut (i \ to 2 times standard length) is greater than in the piscivores, but is not as long as the gut in purely phytophagous species. Breeding. Specimens < 90 mm. S.L. are immature ; both sexes reach the same maximum adult size. Affinities. In its general appearance, H. acidens resembles a number of the deeper- bodied piscivorous species, and one of the specialized predators on larval and embryo fishes, H. parvidens (Greenwood, 1959). Perhaps the greatest resemblance is between H. acidens and H. paraguiarti. Both species have similar neurocrania and jaw structure, but H. acidens has finer teeth, a deeper body (34-6-40-3, M = 38-1 per cent of standard length, cf. 31-7-36-6, M = 34-8%) and a concave as opposed to a straight dorsal head profile. Also showing an overall similarity with H. acidens is H. maculipinna. The con- cavity of the dorsal head profile in H. acidens again serves as one differentiating character. Others are its less oblique jaws, fewer (40-64, M = 48 cf. 50-80, M =62), more slender and longer teeth, and its larger chest scales (5-6 between pectoral and pelvic fin bases, cf. 6-9, mode 7 or 8, in H . maculipinna) . The neuro- crania of the two species also differ, that of H . maculipinna being deeper and having A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 77 a steeper dorsal profile ; the premaxilla in this species lacks a pronounced median expansion, and the pedicels meet the horizontal arms almost at right angles (and not at the more acute angle found in H. acidens). From other species of the H . serranus group (H. victorianus, H. serranus, H. spekii and H. nyanzae], H. acidens is distinguished by its concave dorsal head profile, smaller eye, finer, longer and fewer teeth (at least as shown by the mean number, the ranges overlap), less oblique and non-prognathous lower jaw, and by the absence of a prominent mental bump at the symphysis of the dentaries. Other differences also serve to distinguish H. acidens from individual species of the H. serranus complex ; for these see the species descriptions in Greenwood (1962) and p. 32 above for H. spekii. At about the same level of similarity with H. acidens is H. prognathus. This species differs from H. acidens chiefly in its shallower body (30-1-37-1, M = 33-3% of standard length, cf. 34-6-40-3, M=38-i%), deeper preorbital (18-8-23-1, M = 20-8% of head, cf. 15-2-19-2, M = 17-1%) and smaller eye (20-0-25-0, M = 22-6% of head, cf. 23-0-29-0, M = 26-4%). The dentition and neurocrania of the species differ quite markedly, with H. prognathus having a skull nearer the H. mento type (see Greenwood, 1962, and p. 81) and stouter teeth. It is difficult to assess the phyletic position of H. acidens. The level of anatomical specialization, especially of the syncranium and dentition, suggests affinity both with the H. serranus species group and with H. prognathus, a species not far removed from that group but probably representative of the ancestral type from which such specialized forms as H. mento and H. macrognathus evolved. Its plant diet suggests a possible trophic specialization paralleling that achieved by a species (H. phyto- phagus) related to the small, anatomically generalized, and usually insectivorous species (Greenwood, 1966). In an earlier paper (Greenwood, 1959), I suggested that H. parvidens might have evolved from a species anatomically like H. acidens (called H. nigrescens in that paper) ; this relationship is discussed further on page 114. Phyletically, H. acidens was probably derived from the same stem as H. para- guiarti ; this stem could have been related to either an H. guiarti-\ike lineage, or to an H. serranus-like one. STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. Locality Collector UGANDA B.M. (N.H.) 1911.3.3.20-21 Jinja, Ripon Falls . . . Bayon KENYA B.M. (N.H.) 1966.3.9.107-121 Kavirondo Gulf, near Kisumu . E.A.F.R.O. B.M. (N.H.) 1966.2.21.1 Kavirondo Gulf, near Kisumu . E.A.F.R.O. (Holotype) ZOOL. 15, 2. 78 P. H. GREENWOOD Haplochrotnis prognathus (Pellegrin), 1904 (Text-figs. 14 and 15) Paratilapia prognatha Pellegrin, 1904 (probably in part ; the identity of the three smallest syntypes is still in doubt), Bull. Soc. Zool. France, 29, 185 ; Idem, 1905, Mem. Soc. Zool. France, 17, 181, pi. 16, fig. 4. Boulenger, 1915, Cat. Afr. Fish., 3, 333, fig. 224 (in part, the syntype B.M. [N.H.] reg. no. 1905.2.28.1, ex Kavirondo Gulf). Pelmatochromis spekii (part) : Boulenger, 1915, Cat. Afr. Fish., 3, 417. Haplochromis prognathus: Regan, 1922, Proc. zool. Soc. London, 177. Haplochromis taeniatus Regan, 1922, op. cit., 170, text-fig. 3. Haplochromis macrodon Regan, 1922, op. cit., 176, text-fig. 4. Haplochromis lamprogenys Fowler, 1936 Proc. Acad. Nat. Sci. Philad, 88, 330, fig 137. Haplochromis steindachneri Lohberger, 1929. Anz. Akad. Wiss. Wein, no. 17, 207. Haplochromis versluysi Lohberger, 1929, Anz. Akad. Wiss. Wein no. 17, 206. Haplochromis rebeli Lohberger, 1929, Anz. Akad. Wiss. Wein., no. n, 94 (probably in part; the identity of one paratype, a female 116 mm. S.L., reg. no. 18768 of the Vienna Museum, is still doubtful). LECTOTYPE : a fish 136-0 mm. standard length, B.M. (N.H.) reg. no. 1905.2.28.1 (presented by the Paris Museum), collected by C. Alluaud from Kavirondo Bay, Kenya. This specimen appears to be the fish figured by Pellegrin (1905), and is chosen as lectotype principally for that reason. Note : Defining this species has proved particularly difficult, mainly because of the condsiderable intraspecific variability in head shape (see text-fig. 15). This variability, although apparent to the eye, is not readily quantifiable. Intergrades exist between the most outstanding variants, and now that a large series of specimens is available it appears that the variation is, to a large extent, size correlated. In fishes less than no mm. S.L., the snout seems protracted and the dorsal head profile has a marked and extended concavity above the eye. In larger fishes the elongate " face " is less noticeable, and the dorsal profile is straighter and slopes steeply (but still with a slight supraorbital concavity). DESCRIPTION : based on 43 specimens, 70-141 mm. S.L., and including the lectotype, the syntypes of H. taeniatus and H. macrodon and the holotypes of H . steindachneri and H. lamprogenys. The types of H. rebeli and H. versluysi were examined but are not included in this redescription. Depth of body 30-1-37-1 (M = 33*3) per cent of standard length, length of head 35-3-38-4 (M = 36-9) per cent ; head profile variable (see note above), usually with a supraorbital concavity, the dorsal outline sloping at an angle of 30°-40° with the horizontal. Preorbital depth 18-8-23-1 (M = 20-8) per cent of head, least interorbital width 18-1-23-7 (M = 21-0) per cent. Snout length 33-4-39-6 (M = 37'0) per cent of head, i £ to i^ times as long as broad (rarely if times). Eye diameter with slight negative allometry, 20-0-25-0 (M = 22-6) per cent of head ; in fishes 70-90 mm. S.L., the eye is clearly larger than the cheek is deep but in specimens 90-110 mm. it is equal to or slightly smaller than the cheek, a relationship that holds for some larger in- dividuals (110-125 mm. S.L.) although in most individuals in this size range the eye diameter is manifestly less than the cheek depth. Depth of cheek 20-6-28-6 (M = 24-5) per cent of head (18-9% in the smallest specimen). A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 79 1-1 . •« bo II >§ -1- M ei 8o P. H. GREENWOOD 1 CM FIG. 15. Haplochromis prognathus, showing variability of head profile. Caudal peduncle 14-6-19-0 (M = 16-0) per cent of standard length, 1-2 (rarely)-i-7 times is long as deep (modal range 1-3-1-5). Mouth moderately oblique, the jaws equal anteriorly or the lower projecting slightly ; lower jaw with a slight mental protuberance, its length 42-5-51-1 (M = 46-2) per cent of head, 2-0-2-8 (mode 2-4) times as long as broad. Posterior tip of the maxilla not reaching the vertical through the anterior orbital margin (except in one specimen), but usually reaching a point nearer this line than one through the nostril. Gill rakers : 8-10 (mode 9) on the lower part of the first gill arch, the lower 1-3 rakers reduced, the upper 3 or 4 often flat or flat and lobed, the remainder slender. Scales : ctenoid. Lateral line with 30 (f.2), 31 (f.12), 32 (f.2i) or 33 (f.8) scales ; some individuals lack pores in the scales of this series. Cheek with 3 (occasionally 4) rows. Five to 7 (mode 6) scales between the dorsal fin origin and the upper lateral line, 5-7 (mode 6) between the pectoral and pelvic fin bases. Fins. Dorsal with 23 (f.6), 24 (f.26) or 25 (f.io) rays, comprising 14 (f.n), 15 (f.28) or 16 (f.3) spines and 8 (f.i), 9 (f.28) or 10 (f.i3) branched rays. Anal with n (f.2), 12 (f.2i) or 13 (f.n) rays, comprising 3 spines and 8 (f.2), 9 (f.2i) or 10 (f.n) branched elements. Pectoral fin shorter than head, 22-0-29-3 (M = 25-7) per cent of standard length . First pelvic branched ray slightly produced in both sexes but proportionately more so in males. Caudal truncate or subtruncate, scaled on its basal half to two- thirds. Teeth. In the smallest fish examined (70 mm. S.L.) the outer teeth in both jaws are mostly unequally bicuspid, but a few unicuspids are present. Fishes between this size and 105 mm. S.L. have mostly unicuspids in the outer row, but some bi- cuspids occur posterolaterally in the upper jaw. Fishes > 105 mm. have only uni- cuspid outer teeth, a condition found occasionally in individuals as small as 90 mm. S.L. The unicuspids are slightly curved inwards, and vary in form from relatively slender to moderately stout. There are 30-60 (M = 45) teeth in the outer row of the upper jaw, the number not showing any correlation with the fish's size. Regan's (1922) " key " character separating H. taeniatus from H. macrodon (outer teeth numerous, close together cf. outer teeth rather strong, set well apart) is hardly A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 81 trenchant even when only the type specimens are compared, and is inapplicable when a large series is examined. The inner teeth are mostly tricuspids in fishes < no mm. S.L., but an admixture of tri- and unicuspids (or weakly tricuspids) is found in fishes at the upper end of this size range. Some tricuspids also occur in large fishes, but in the majority of these all inner rows are composed of unicuspids. When tri- and unicuspids occur together in fishes > 120 mm. S.L. the tricuspids are confined to the innermost rows. Inner teeth are arranged in 2 (mode) or 3, rarely I, rows in the upper jaw, and in I (mode) or 2 (rarely 3) rows in the lower. Osteology. The neurocranium of H. prognathus is approximately intermediate between that of H. paraguiarti and H. mento (see p. 70, and Greenwood, 1962). It thus closely resembles the neurocrania of H. bartoni, H. mandibularis , H. argenteus and H. longirostris (see p. 112 and Greenwood, op. cit), in Lake Victoria, and H. venator in Lake Nabugabo (Greenwood, 1965). From the four former species it differs most noticeably in its deeper supraoccipital crest, and from H. argenteus also by its being somewhat broader anterior to the orbits. There are also resemblances in neuro- cranial form between H. prognathus and H. xenostoma (see p. 53) but the differences here are somewhat more pronounced than in the other species mentioned. The premaxilla has an expanded median dentigerous surface which gives the bone a beaked appearance ; this is a feature of those piscivorous species which I consider to be more specialized in body-form than the members of the H. guiarti and H. serranus species complexes (see Greenwood, 1962). In contrast, the dentary of H. prognathus is of the stout, deep-bodied type found in the latter groups. The lower pharyngeal bone is triangular, with the dentigerous surface as long as broad or slightly broader (both conditions are of equal frequency). The lower pharyngeal teeth are fairly coarse, with cylindrical necks and compressed, weakly bicuspid crowns ; these teeth are well-spaced on the bone in 18-24 rows. Vertebral counts in 8 specimens are : 12+17 (f.2), 13 + 15 (f.i), 13 -f 16 (£.4) and 13 + 17 (f.i) giving totals of 28-30. Coloration. Live colours are unknown. Preserved material : Males (adult and sexually active). Body and head brownish above the midline, silvery-yellow below ; two faint lateral bands are visible, one running midlaterally from behind the eye to the origin of the caudal fin, the other along the upper lateral line. There is a faint, ill-defined but broad lachrymal band, but it does not extend ventrally to the margin of the preorbital. Dorsal fin greyish, the dark pigment most intense basally on the spinous region, lappets dusky ; soft dorsal maculate. Caudal dark grey, darkest between the rays on the proximal two-thirds of the fin. Anal greyish, lappets black, as may be the basal part of the membrane between the spines ; 2 or 3 large but faint, greyish ocelli. Pel vies black. Females (adult and juvenile) . Light grey-brown above, silvery below ; very faint traces of longitudinal bands as in males, the lower often visible on the posterior third of the body. All fins hyaline, but the caudal is dark on its proximal two- thirds. Ecology. Habitat. Haplochromis prognathus is apparently confined to water less than 20 ft. deep, and to hard substrates ; it occurs in both sheltered and exposed 82 P. H. GREENWOOD localities, including wave-washed beaches. Food. Of the 33 specimens examined (from n localities), 22 contained food. Ten specimens contained only remains of fishes (predominantly small Cyprinidae, but Haplochromis were also identified), 3 yielded fish and insect remains (the fish unidentifiable, the insects adult Isoptera and larval Ephemeroptera) , 7 contained only insects (adult Isoptera and larval Ephemeroptera), i contained unidentifiable fish remains and some plant fragments (undigested), and 2 specimens contained unidentifiable material (? bottom debris) and some plant remains. Breeding. Little is known about the breeding habits of this species ; no brooding individuals have been recorded. Sexual maturity is attained at a length of about 100 mm., but larger juvenile individuals (up to 120 mm.) are known. Both sexes reach the same maximum adult size. Affinities. There is nothing particularly outstanding about the general appearance of H. prognathus when it is compared with the other relatively deep-bodied predatory species of Lake Victoria. Thus, it bears a superficial resemblance to several pisci- vorous species, especially H. serranus, H. bartoni, H. nyanzae and H. mandibularis . From H. serranus, H. prognathus differs in its slightly shallower body (30-1-37-1, M = 33'3% standard length cf. 32-7-39-2, M = 36-0%), deeper preorbital (18-8-23-1, M = 20-8% head, cf. 14-6-20-0, M = 17-7%), slightly longer snout (33-4-39-0, M=37-o% head, cf. 30-8-37-0, M=34'O%), shorter lower jaw (42-5-51-1, M = 46-2% head, cf. 47-0-60-0, M = 54-3%) and slightly larger chest scales (5-7, mode 6, between the pelvic and pectoral fin bases, cf. 7 or 8 [rarely 6]). The pre- served coloration of the two species suggests a fairly marked difference in live colours, and there are clear-cut differences in neurocranial form (see p. no). From H. bartoni (probably its nearest extant relative), H. prognathus differs in having a broader and shorter lower jaw (42-5-51-1, M = 46-2% head, cf. 50-8-57-0, M = 52-5%), fewer outer teeth in the upper jaw (30-60, M = 45, cf. 50-80, M = 62), the maxilla not extending so far posteriorly (reaching or almost reaching the orbit in H. bartoni), and its smaller chest scales. Neurocranial form in the two species is similar. The characters separating H. prognathus from H. nyanzae are its longer snout (33'4-39'6» M =37'0% head, cf. 33'4-35'8, M =34-5%). more oblique lower jaw, its maxilla not reaching the orbit, and its larger chest scales. Unfortunately it has not been possible to compare directly the neurocrania of the two species. But, from radiographs it seems probable that the skull of H. nyanzae is like that of H. serranus (see Greenwood, 1962). Haplochromis prognathus differs from H. mandibularis in having a broader snout, shorter lower jaw (42-5-51-1, M=46-2% head, cf. 47-3-56-8, M=5i-5%), and longer caudal peduncle (14-6-19-0, M = 16-6% standard length, cf. 12-2-15-2, M = 14-2%). Neurocranial form in the two species is similar. Haplochromis prognathus is, apparently, closely related to H. argenteus. At first sight, the great variability in head shape of H. prognathus obscures the relationship. But, if smaller specimens of H. prognathus are compared with larger individuals of H. argenteus the likeness is striking. Morphometrically, even superficially dissimilar specimens of the two species are not readily separable. Characteristics of the lower A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 83 jaw serve to distinguish the species. The lower jaw of H. prognathus is broader (length/breadth ratio 2-0-2-8 [modal range 2-0-2-4], cf- 2'3~3'1 [modal range 2-8- 3-0]), shorter (42-5-51-1, M = 46-2% of head, cf. 45-0-59-0, M— 50-2%), less oblique and less prognathous. The possible phyletic relationship of H. prognathus within the Lake Victoria species-flock will be discussed later (p. no) ; outside Lake Victoria, H. prognathus is probably related to H. venator of Lake Nabugabo (Greenwood, 1965). Indeed H. venator could well have been derived from populations of H. prognathus cut off when the sand bar which isolated Lake Nabugabo was formed about 4,000 years ago. STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. B.M. (N.H.) 1906.5.30.304 (Syntype H. macrodon) B.M. (N.H.) 1906.5.30.305-6 (Syntypes H . macrodon) B.M. (N.H.) 1906.5.30.250 (Syntype H. taeniatus) B.M. (N.H.) 1966.3.9.50 B.M. (N.H.) 1966.3.9.64-65 B.M. (N.H.) I966-3-9.53-59, 66-67, 72~75 B.M. (N.H.) 1966.3.9.68-71 B.M. (N.H.) 1966.3.9.79-83 B.M. (N.H.) 1966.3.9.51-52 B.M. (N.H.) 1966.3.9.60-63 Vienna Museum 17872 (Type of H . steindachneri) Academy of Nat. Sciences Philadelphia ANSP 66131 (Holotype of H . lamprogenys) B.M. (N.H.) 1905.2.28.1 (Lectotype) B.M. (N.H.) 1905.12.11.4 (Syntype H. taeniatus) B.M. (N.H.) 1966.3.9.76-78 B.M. (N.H.) 1966.3.9.84 B.M. (N.H.) 1906.5.30.251-2, 256 Vienna Museum 17876-7 (Types of H. versluysi) Vienna Museum 18830 (Holotype of H. rebeli) Vienna Museum 18769-71 (Paratypes of H. rebeli) Locality UGANDA Entebbe . Bugonga Entebbe . Entebbe, Airport Beach Napoleon Gulf, near Jinja . Beach near Nasu Point (Buvuma Channel) . Katebo . Bugonga, near Entebbe Kagera Bay, near mouth of Kagera River Bukafu Bay Sesse Islands Kitala KEYNA Kavirondo Gulf Kavirondo Gulf TANZANIA Mwanza Harbour Capri Bay, Mwanza . LAKE VICTORIA Locality unknown Locality unknown Locality unknown Locality unknown Collector Degen Degen Degen E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. Rolle Alluaud Alluaud E.A.F.R.O. E.A.F.R.O. Degen Rolle Rolle Rolle 84 P. H. GREENWOOD Haplochromis argenteus Regan, 1922 (Text-fig. 16) Paratilapia longirostris (part): Boulenger, 1915, Cat. Afr. Fish., 3, 332. Haplochromis argenteus Regan, 1922, Proc. zool. Soc. Londn., 186, fig. 12. LECTOTYPE : a juvenile female 114-0 mm. standard length, from Bunjako, Uganda (B.M. [N.H.] reg. no. 1906.5.30.266 ; the specimen figured by Regan). FIG. 1 6. Haplochromis argenteus ; the lectotype, a juvenile, about -75 times natural size. From Regan, Proc. zool. Soc. DESCRIPTION : based on 23 specimens (including the lecto- and paralectotype) 93-0-202-0 mm. standard length. Depth of body 26-8-34-8 (M = 31-2) per cent of standard length, length of head 35-5-39-6 (M = 36-8) per cent. Dorsal head profile slightly concave, with prominent premaxillary pedicels, and sloping at an angle of 20°-25°. Preorbital depth 19-6-24-0 (M = 21-7) per cent of head, least interorbital width 17-7-22-3 (M = 20-0) per cent. Snout 1-25-1-40 times as long as broad, its length 34-8-41 '6 (M = 37*8) per cent of head, diameter of eye 19-4-23-5 (M = 21-5), depth of cheek 22-3-28-7 (M = 24-7) per cent. Caudal peduncle I3*5-I7'7 (M = 16-1) per cent of standard length, 1-3-1-7 (modal range 1-3-1-5) times as long as deep. Mouth oblique (35°-40°), the lower jaw projecting moderately in some specimens, its length 45-0-59-0 (M = 50-2) per cent of head, 2-3-3-1 (modal range 2-8-3-0) times as long as broad. Lips slightly thickened. Premaxilla expanded slightly in the midline. Posterior tip of the maxilla reaching a point about midway between the orbit and the nostril, or almost reaching the orbit. Gill rakers : generally slender but moderately stout in a few fishes ; lower i or 2 reduced, the upper 3 or 4 (in one fish, the upper 6) flat and lobed. Eight to 10 (mode 9) on the lower part of the first arch. Scales : ctenoid ; lateral line with 29 (f.2), 30 (1.2), 31 (f-5), 32 (f.n), 33 (f.i), 34 (f.i) or 35 (f.i) scales. Cheek with 3 or 4 (rarely 5) rows. Five or 6 (rarely 7) scales between the dorsal fin origin and the upper lateral line, 6 or 7 (less frequently 5) between the pectoral and pelvic fin bases. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 85 FIG. i6A. Haplochromis argenteus, adult. Drawn by Lavinia Beard. Fins. Dorsal with 23 (f.2), 24 (f.i2) or 25 (f.g) rays, comprising 13 (f.i), 14 (f.i), 15 (f.ig) or 16 (f.2) spines and 8 (f.2), 9 (f.i2) or 10 (f.g) branched rays. Anal with II (f.3), 12 (f.ig) or 13 (f.i) rays, comprising 3 spines and 8, 9 or 10 rays. Pectoral 24-1-29-7 (M = 25-0) per cent of standard length. Pelvics with the first branched ray somewhat elongate, proportionately more so in adult males. Caudal truncate. Teeth. Except for the smallest fish examined (the paralectotype 93 mm. S.L.) the outer teeth in both jaws are unicuspid, slender and slightly curved (those situated posterolaterally in the upper j aw more so than the others) . In the smallest individual most outer teeth are like those described above but there are also some weakly bicuspid and slender teeth. There are 44-80 (M = 58) teeth in the outer row of the upper jaw, the number showing slight positive correlation with standard length. In all except four specimens (the two types, 93 and 114 mm. S.L., and two others 88 and 112 mm. S.L.) the inner teeth are unicuspid. Of the exceptional specimens, the largest (lectotype) has an admixture of tri- and unicuspids, the 93 mm. fish (paralectotype) only tricuspids, while the 88 and 112 mm. individuals have tricuspids in the upper jaw and a mixture of tri- and some unicuspids in the lower. Inner teeth are arranged in 2-4 (rarely 5) rows in the upper jaw, and 1-3 in the lower. One exceptional fish (182 mm. S.L.) has a single, irregular row of widely separated teeth in both jaws. Osteology. The neurocranium of H. argenteus is virtually identical with that of H. longirostris (see Greenwood, 1962), and also resembles the skull of H. prognathus. The premaxilla is more beaked than that of H. longirostris. The lower pharyngeal bone is narrow, but its dentigerous surface is generally broader than long ; less frequently it is equilateral. The teeth are rather coarse, although they are finer and more compressed in a few fishes. These teeth are arranged in 20-24 (usually 22) rows. 86 P. H. GREENWOOD Vertebral counts in 8 specimens are : 12+17 (f.3) and 13 -f- 16 (f.5) giving a total of 29. Coloration. Live colours are unknown. Preserved material : Males (adult and sexually active) : ground colour grey-silver, darker (almost brown) on back and dorsal head surfaces ; tip of lower jaw and anterior part of each ramus dusky. Branchiostegal membrane sooty in the opercular region, dark (but not black) anteriorly. Sub- and interopercula with a golden-yellow flush. An ill-defined to distinct lachrymal stripe runs from the anterior orbital margin to behind the posterior tip of the maxilla. Dorsal fin greyish-yellow, with black lappets, and in some specimens a dusky, irregular banding or marbling on the entire spinous and anterior soft parts. Caudal greyish-yellow, dark grey-brown on the proximal half. Anal greyish with black lappets, and 2 or 3 dead- white ocelli. Pelvics dusky. Males (sexually quiescent, and immature) : Ground colours like those of active individuals but somewhat lighter, often with very faint traces of 4 or 5 bars mid- laterally on the flanks ; the lachrymal stripe is of variable intensity. Dorsal fin as above or without the dark marbling. Other fins also as above, but the anal ocelli may be weakly defined and small, and the pelvics lighter. Females (adult and immature). Ground coloration as in males, but with a faint, interrupted and narrow midlateral band running from the posterior opercular margin to the basal part of the caudal fin. Even fainter traces of a more dorsal longitudinal band (following the course of the upper lateral line) are seen in some specimens ; the two bands may be linked by 3 or 4 extremely faint, short and narrow vertical bars. All fins are yellowish, the caudal dark basally. Ecology. Habitat. Most specimens are from sheltered localities where the bottom is composed of organic mud, and at depths of less than 40 ft. Other localities are, however, more exposed and the substrate is of rock, sand or shingle, but only a few (and juvenile) fishes were obtained from nets operated over exposed beaches. Food. Of the 20 individuals examined (from n localities) 8 had food in the gut. Four fishes contained only insect remains (in 2 thought to be terrestrial species, and in the others identified as larval Ephemeroptera) , and 4 had fragmentary fish remains (identified as the cyprinid Engraulicypris argenteus) . Breeding. Fishes < 114 mm. S.L. are immature, as is one exceptional specimen of 146 mm. Both sexes reach the same maximum adult size. Affinities. The species most like H. argenteus is H. longirostris. There are, however, a number of morphological differences which serve to separate them, in- cluding the larger adult size reached by H. argenteus. From H. longirostris, H. argenteus also differs in its less oblique and longer lower jaw (45*0-59-0, M = 50-2% of head, cf. 42-2-51-4, M =46-0%), shorter and deeper caudal peduncle (13-5-17-7, M = 16-1% of standard length, cf. 17-2-22-2, M = 19-2% ; length/depth ratio I'3~1'7 [modal range 1-3-1-5], cf. 1-7-2-3 [modal range 1-9-2-0]), somewhat deeper body (26-8-34-8, M = 31-2% of standard length, cf. 24-6-30-4, M = 27-2%) and slightly longer head (35-5-39-6, M = 36-8% standard length, cf. 29-2-36-2, M = 33'0%) ; also, the premaxilla of H. argenteus is somewhat more beaked. In an earlier paper (Greenwood, 1962) I used the ratio of eye diameter to interorbital A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 87 width as a diagnostic character ; more material of H. argenteus has shown, however, that this difference is too slight to be of value. The similarities between H. argenteus and H. prognathus are discussed elsewhere (p. 82). When these two species and H. longirostris are considered together, it seems probable that H. argenteus and H. longirostris were derived from an H. prognathus-like ancestor, probably as distinct lines developing almost in parallel. STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. B.M. (N.H.) 1906.5.30.266 (Lectotype) B.M. (N.H.) 1906.5.30.267 (Paralectotype) B.M. (N.H.) 1966.3.9.194-195 B.M. (N.H.) 1966.3.9.201-202 B.M. (N.H.) 1966.3.9.193 B.M. (N.H.) 1966.3.9.196-198 B.M. (N.H.) 1966.3.9.192 B.M. (N.H.) 1966.3.9.208-210 B.M. (N.H.) 1966.3.9.191 B.M. (N.H.) 1966.3.9.199-200 B.M. (N.H.) 1966.3.9.203-204 B.M. (N.H.) 1966.3.9.207 B.M. (N.H.) 1966.3.9.205-206 B.M. (N.H.) 1901.6.24.90 B.M. (N.H.) 1966.3.9.190 Locality UGANDA Bunjako .... Bunjako .... Ekunu Bay Pilkington Bay Beach near Nasu Point Ramafuta Island (Buvuma Channel) Off S. end of Buvuma Island Beach near Grant Bay (Buvuma Channel) . Channel between Yempita and Busiri Islands Kazima Island Off Entebbe Harbour Entebbe Harbour Bukakata Bay . LAKE VICTORIA Locality unknown Locality unknown Collector Degen Degen E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. Uganda Fisheries Dept. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. Sir. H. Johnston E.A.F.R.O. Haplochromis squamulatus Regan, 1922 (Text-fig. 17) Paratilapia pectoralis (non Ctenochromis pectoralis Pfeffer) Boulenger, 1911, Ann. Mus. Genova (3), 5, 66, pi. i, fig. 2. Idem, 1915, Cat. Afr. Fish, 3, 339, fig. 229. Paratilapia serranus (part) : Boulenger, 1915, op.cit., torn, cit., 334. Haplochromis guiarti (part): Regan, 1922, Proc. zool. Soc. Londn., 174. Haplochromis squamulatus Regan, 1922, op. cit., 175 (nom. nov. for Paratilapia pectoralis Blgr., 1911). HOLOTYPE : a male (probably adult), 149 mm. standard length, from Jinja (Uganda). Genoa Museum reg. no. C.E. 12977. DESCRIPTION : based on 54 specimens (including the holotype), 66-198 mm. S.L. Depth of body 23-5-35-1 (M = 30-3) per cent of standard length, length of head 29.4-39.5 (M == 34-3) per cent. Dorsal head profile moderately to strongly sloping, 88 P. H. GREENWOOD the premaxillary pedicels prominent and breaking the outline of the profile, snout region sloping at an angle of 40°-5o° with the horizontal. Preorbital depth in fishes < 100 mm. S.L. (N = n), 12-5-16-5 (M = 15-5) per cent of head, in larger fishes (N = 43) 15-6-21-6 (M = 18-9) per cent, least inter- orbital width 21-2-28-3 (M = 25-0) per cent. Snout as long as broad to 1-3 times longer, its length in fishes < 100 mm. S.L., 26-0-32-1 (M = 29-1) per cent of head, in larger individuals 20-6-25-9, M = 23-3 per cent. Cheek depth 19-5-29-3 (M = 25-7) per cent of head. Caudal peduncle 14-3-22-3 (M = 17-9) per cent of standard length, 1-2 (rare)-2-o (mode 1-7) times as long as deep. Mouth horizontal or very slightly oblique, jaws equal anteriorly, or infrequently, lower projecting slightly. Length of lower jaw in fishes < 170 mm. S.L. (N = 46), 39-0-48-0 (M =44-1) percent of head, 44-2-51-0 (M = 47-5) per cent in larger fishes (N = 8) ; 1-5-2-5 (mode 1-8, modal range 1-5-2-0) times as long as broad in fishes of all sizes. Posterior tip of the maxilla generally reaching the vertical through the anterior orbital margin or to below the eye, but not reaching either point in a few individuals. Lips slightly thickened ; premaxilla not expanded medially. Gill rakers : short and stout, often flat, in most specimens ; in fishes < 120 mm. S.L., but also in a few larger individuals, most of the rakers are relatively slender. Eight-io (mode 9), rarely n, on the lower part of the first arch, the lower 1-3 rakers reduced. Scales : ctenoid. Lateral line with 32 (f.2), 33 (f.12), 34 (f.8), 35 (f.15), 36 (f.io), 37 (f.6) or 38 (f.i). Pore distribution in this series is irregular, with some pore scales occurring in the horizontal row above that in which the majority lies ; also, some scales are without pores and are often smaller than those before and behind them. Cheek with 3 (f.8), 4 (£32), 5 (f.12) or 6 (f.i) rows of scales. Scales on the nape, chest and belly are small ; 6 (£.3) 6| (£3), 7 (f.n), 7i (1.5), 8 (£14) or 9 (f.i6) between the FIG. 17. Haplochromis squamulatus, holotype, about -72 times natural size. Cat. Afr. Fish. From Boulenger, A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 89 dorsal fin origin and the upper lateral line, 6 (f.i), 7 (f.8), 8 (f.i6), 9 (£.25) or 10 (£.3) between the pectoral and pelvic fin bases. Fins. Dorsal with 24 (f.g), 25 (f.37) or 26 (f.6) rays, comprising 14 (f.2), 15 (f.26) or 16 (f.24) spines, and 8 (f.2), 9 (£23), 10 (£.25) or n (f.2) branched rays. Anal with II (f.3), 12 (f.4o) or 13 (f.5) rays, comprising 3 spines (except for one specimen with 2), and 8 (f.3), 9 (f-4o) or 10 (f.5) branched elements. Pectoral 21-8-27-0 (M =24-7) per cent of standard length. Pelvics with the first branched ray produced in both sexes but proportionately more so in adult males. Caudal subtruncate to truncate. Teeth. Most fishes < 105 mm. S.L. have either an admixture of uni- and bicuspid teeth in the outer row of both jaws, or only bicuspids. One specimen (89 mm. S.L.) has predominantly bicuspids and a few tricuspids in the upper jaw, but mostly uni- cuspids in the lower. In larger fishes of the < 105 mm. group, unicuspid teeth predominate in both jaws, and in fishes > 105 mm. S.L. only unicuspids are found. All outer teeth are relatively slender and gently curved, the unicuspids more so than the bicuspids. The mean number of teeth in the outer row of the upper jaw shows some positive correlation with standard length although there is little difference in the ranges for the various groups ; fishes < 120 mm. S.L. have 50-72, M = 57 teeth in this row, while larger specimens have 52-80, M = 68. There is less obvious correlation between tooth form and the fish's size when the inner rows of teeth are considered. All fishes < 105 mm. S.L. have only tricuspids in the inner rows. Above this length many fishes have only unicuspids, but a mixture of uni- and tricuspids (or weakly tricuspids) is common even in the largest individuals. Unicuspids occur most frequently in the outer rows of the inner series. Inner teeth are arranged in 2-6 rows (usually 3 or 4) in the upper jaw, and in 1-6 (usually 2 or 3) in the lower jaw. There is perhaps some correlation between the number of rows and the fish's length because the largest specimens (179-198 mm. S.L.) have the greatest number of inner rows. Also, in larger fishes the inner teeth are implanted very obliquely but are almost vertical in fishes < 105 mm. S.L. Osteology. The neurocranium of H. squamulatus combines characteristics of both the H. guiarti and the H. serranus types (see Greenwood, 1962). The dorsal pre- orbital profile rises steeply and is gently curved ; its line is continuous with that of the supraoccipital crest whose anterior point lies further forward than in the skulls of H. guiarti and H. serranus. In general there is a great similarity between the neurocrania of H. squamulatus, H. michaeli and H. martini (see Greenwood, op. cit). The premaxilla is of the generalized type, and thus lacks a pronounced anterior extension of its medial dentigerous surface. The lower pharyngeal bone has the dentigerous surface broader than long, from slightly to markedly so. The lower pharyngeal teeth are relatively slender, com- pressed and distinctly cuspidate ; the teeth are arranged in 22-26 (mode 24) rows. Vertebral counts in 10 specimens are : 13+16 (f.2), 13 + 17 (f.2), 13 + 18 (f.2), 14 + 16 (f.i), 14 + 17 (f.3) giving totals of 29-31. Coloration. Live colours are known for females (adult) and quiescent males. Females : ground colour yellow-silver above shading to yellowish-white ventrally, the dorsal body and head surfaces are darkest. Two distinct longitudinal bands are 90 P. H. GREENWOOD invariably present, the lower running from the operculum to the caudal fin origin along the level of the lower lateral line, the upper extending from the nape to the end of the dorsal fin at a level about two scale rows below the dorsal fin insertion. Both bands have a finely zig-zagged outline ; the upper band may be faint. Dorsal fin dark neutral. Caudal dark neutral with a yellow flush. Pelvic and anal fins yellow. Quiescent males have a coloration like that of the females described above. Preserved material : Males (adult, sexually active but not ripe). The general im- pression is one of dusky greyness, with small areas of light brown on the flanks. The entire head is dark but is lighter on the operculum and lower lip (the lower jaw is dark). A broad black band (faint in some specimens) runs from the posterior opercular margin to the caudal fin origin ; it is crossed by 3 or 4 broad but faint vertical bars in the zone of lighter flank coloration. These bars merge with the dark dorsum and sooty-grey chest and belly. The ventral aspects of the caudal peduncle are also sooty-grey. Dorsal fin yellow-brown with a sooty overlay, the lappets are black, and a few ill-defined dark maculae may be visible on the posterior part of the soft dorsal fin. Caudal dark grey-brown on its proximal three-quarters, yellowish-brown distally. Anal coloured like the dorsal but greyer over the spinous part ; ocelli either not clearly defined, or white with a sooty surround. Pelvic fins dusky over a yellowish ground. Males (adult but quiescent}. Ground colour light brown, shading to golden on the ventral surfaces which are, however, overlaid with greyish-black. Dorsal surface of the head and body dark brown. A fairly broad, distinct and almost vertical lachrymal stripe runs from the anteroventral orbital margin to the dentary. A very distinct, broad lateral stripe runs from the opercular margin to the caudal fin base ; the upper margin of this band is irregularly serrate. There are 3 or 4 in- complete but broad and distinct vertical blotches on the ventral half of the flanks, each blotch originating from the midlateral band but not extending to a point more than half way between the lateral band and the ventral body outline. A very faint dark upper lateral band may be distinguished running at a level about midway between the dorsal fin base and the upper lateral line ; this band is often in- distinguishable from the generally dark coloration of the dorsum. A third band at the base of the dorsal fin may be visible. Dorsal fin pale brownish-yellow, with a faint sooty overlay ; lappets black, dark blotches present on the soft fin. Anal pale yellowish, with a very faint sooty overlay. Pelvics yellowish with an intense sooty overlay on the anterior half. Caudal dark greyish-yellow. Immature males are coloured like females (see below), but have a darker ground coloration (almost grey on the chest and belly) and faintly sooty pel vies. Females (adult and immature}. Female coloration is very distinctive because of the well-marked lateral bands. The ground coloration is a yellowish-silver, some- what greyish above the upper lateral line and on the dorsal head surface ; the cheek and operculum are yellow-silver. The midlateral band is black and has character- istically serrate upper and lower margins ; the depth of this band is rather variable in any one fish, and is generally deepest above the anal fin. The upper lateral band also has irregularly serrate margins but it is generally less obvious because of its A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 91 position on the dark colour of the dorsum. A third longitudinal band, narrower and interrupted, lies along the base of the dorsal fin as a series of narrow, elongate blotches. Dorsal fin yellowish with a faint sooty overlay, and sooty lappets ; the soft part often has dark spots and blotches between the rays. Caudal yellowish with dark maculae on the proximal quarter to third. Anal and pelvic fins yellow. Ecology. Habitat. The species is known from a wide variety of habitats, in- cluding sheltered bays and gulfs, exposed beaches, and from certain off-shore localities near small islands. In most places the substrate is hard (rock, sand or shingle) but H . squamulatus is known to occur over mud bottoms. Food. Thirty-six specimens were examined (covering the entire size-range, and from 18 localities) ; of these, 19 contained food in the gut. In 16 fishes the food consisted entirely of fishes, the remains so macerated that certain identification was difficult ; in two cases the remains were identified as Haplochromis species, in one as Engraulicypris , and in another as a cichlid. In two female fishes the stomach con- tained many small embryos of cichlid fishes ; these could well be the fishes' own broods swallowed at the time of capture, although it is not known whether H. squamulatus is a mouth-brooder. A third female fish had several small post-larval cichlids in the stomach ; judging from the gonad state of the predator it seems un- likely that the larvae were its own brood. In addition to the sixteen piscivorous fishes, one other yielded only crustacean remains, one several insect egg-masses, and a third the remains of boring mayfly larvae (Povilla adusta). Breeding. All specimens, except one, below 135 mm. S.L. are immature ; the exceptional fish (93 mm.) is a ripening female. The largest fishes (180-198 mm. S.L.) are females but there is one male of 179 mm. S.L. Affinities. The colour pattern, small chest and nuchal scales and the sharply decurved head profile of H. squamulatus make at least large specimens readily identifiable. But, smaller individuals are less easily distinguished from specimens of H . martini (although the latter are adult at a size when most H. squamulatus are still immature). Unfortunately, little is known about the live coloration of sexually active male H. squamulatus but there are several similarities in the coloration of female H. martini and H. squamulatus, particularly in the striping and the yellowish coloration. In fishes of all sizes, H. squamulatus differ from H. martini in their less strongly decurved head profile, maxilla not extending so far posteriorly (never reaching a point below the pupil as is general in H. martini], in having a higher modal number of lateral line scales (35 cf. 33) and of scales between the pectoral and pelvic fin bases (9 cf. 7 or 8), and in the somewhat shallower body (23-5-35-1, M = 30-3% standard length, cf. 30-8-38-0, M =34-4%). When specimens over 100 mm. S.L. are compared, a number of additional mor- phometric differences are apparent : H. squamulatus has a longer snout, smaller eye and a longer lower jaw. Specimens less than 100 mm. S.L. have only one trenchant morphometric difference, the size of the eye (eye diameter in H. squamulatus is 25-9-30-4 [M = 28-3]% of head, cf. 29-4-37-5 [M = 317]% in H. martini}. Also resembling H. squamulatus (and H. martini, see Greenwood, 1962) is Haplo- chromis michaeli. The species differ in their preserved coloration (see above, and 92 P. H. GREENWOOD Greenwood, op. cit., p. 205), in the straighter, slightly concave, dorsal head profile of H . michaeli, and the greater posterior extension of the maxilla in that species (gener- ally to below the pupil). Since specimens of H. michaeli over 100 mm. S.L. only are known, interspecific morphometric comparisons are restricted. These, however, show that H. squamulatus has a slightly shallower body (23-5-35-1, M=30-3% of standard length, cf. 30-8-37-6, M=34«3%), and a smaller eye (20-6-25-9, M = 23-3% head, cf. 24-0-29-1, M =27-0% in H. michaeli). The nuchal scales of H. squamulatus are somewhat smaller (modal number of scales between the dorsal fin origin and the upper lateral line 8 or 9, cf. 6 or 7 in H. michaeli) as are those of the lateral line series (mode 35 cf. 33) ; the upper limit of the range for the lateral line scale count is also higher in H. squamulatus (38 cf. 35) . All in all, the resemblances between H. squamulatus and H. martini are greater than than those between H. squamulatus and H. michaeli, but the three species seem to form a closely related group within the more generalized piscivorous predators of Lake Victoria. There is a certain similarity, albeit superficial, between H. squamulatus and H. altigenis. On closer examination, however, it does not seem likely that the re- semblance can be construed as implying a close phyletic connection between the species. Haplochromis altigenis is probably a derivative of the H . guiarti stem. In its general facies, H. squamulatus shows some Similarity with H. dichrourus, but there is a very pronounced interspecific difference in coloration (cf. p. 90 with p. 67), as well as differences in some morphometric characters. Haplochromis squamulatus has a larger eye (20-6-25-9, M = 23-3% of head, cf. 19-4-22-6, M = 21-1) a shorter lower jaw (44-2-51-0, M = 47-5% of head, cf. 51-3-54-0, M =52-7%) and a higher modal number of lateral line scales (35 cf. 33). Phyletically, H. squamulatus and H. michaeli may represent slightly divergent developments (less so morphologically and ecologically in the former species) from an H. martini-like stem, an increase in adult size being a common factor in the two lines. Museum and Reg. No. B.M. (N.H.) 1966.3.9.262-272 B.M. (N.H.) 1966.3.9.282-284 B.M. (N.H.) 1966.3.9.285-288 B.M. (N.H.) 1966.3.9.289-295 B.M. (N.H.) 1966.3.9.259-260 B.M. (N.H.) 1966.3.9.261 Genoa Museum C.E. 12977 (Holotype) B.M. (N.H.) 1906.5.30.233-9 B.M. (N.H.) 1909.5.4.4-5 B.M. (N.H.) 1966.3.9.278-281 Locality Collector UGANDA Napoleon Gulf near Jinja . . E.A.F.R.O. Fisherman's Point near Jinja . E.A.F.R.O. Beach near Nasu Point (Buvuma Channel) . . . E.A.F.R.O. Ramafuta Island (Buvuma Channel) . . " . . E.A.F.R.O. Off Buvuma Island (Buvuma Channel) . . : 3,,=, .... E.A.F.R.O. Grant Bay . . . . E.A.F.R.O. Jinja (Napoleon Gulf) . . . Bayon Nsonga . . . . Degen Sesse Islands .... Bayon Beach in Entebbe Harbour E.A.F.R.O. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 93 B.M. (N.H.) 1966.3.9.258 B.M. (N.H.) 1966.3.9.273-277 B.M. (N.H.) 1966.3.9.255 B.M. (N.H.) 1966.3.9.257 B.M. (N.H.) 1966.3.9.256 B.M. (N.H.) 1901.6.24.86 Entebbe, Airport Beach Kazima Island near Entebbe Old Bukakata Bay . TANZANIA Capri Bay, Mwanza . Mwanza Harbour LAKE VICTORIA Locality unknown E.A.F.R.O. Uganda Fisheries Dept. E.A.F.R.O. E.A.F.R.O. E.A.F.R.O. Sir H. Johnston Haplochromis barbarae sp. nov. (Text-figs. 18 and 19) HOLOTYPE : an adult female, 97 mm. standard length, from a beach near Nasu Point (Buvuma Channel) ; B.M. (N.H.) reg. no. 1966.2.21.5. Named in honour of Mrs. Barbara Williams, whose drawings illustrate this and others of my papers. DESCRIPTION : based on 9 specimens (including the holotype) 89-0-106-0 mm. standard length. Depth of body 32-1-35-5 (M = 34-0) per cent of standard length, length of head 31-4-35-0 (M = 33-5) per cent. Dorsal head profile very slightly decurved or straight. Preorbital depth 15-1-18-0 (M = 16-9) per cent of head length, least interorbital width 24-2-26-7 (M = 25-5) per cent. Snout as long as broad, its length 30-0-32-4 (M=3i-5) per cent of head, eye diameter 22-9-27-4 (M=25«8), cheek depth 20-0-25-0 (M = 22-2) per cent. Caudal peduncle 15-7-18-0 (M = 16-8) per cent of standard length, 1-4-1-7 times as long as deep (no well-defined mode). Mouth very slightly oblique, the jaws equal anteriorly ; lower jaw 36-6-41-2 (M = 38-0) per cent of head, 1-5-1-7 (in one fish 2-0) times as long as broad. Posterior tip of the maxilla reaching a point near the vertical through the anterior orbital margin. FIG. 18. Haplochromis barbarae. Drawn by Barbara Williams. ZOOL. 16, 2. 94 P. H. GREENWOOD Gill rakers : of variable form, from moderately short and stout to slender and relatively elongate, but of constant form (except for the reduced lower rakers) in any one individual ; 8 or 9 rakers on the lower part of the first gill arch. Scales : ctenoid ; lateral line with 33 (f.3), 34 (f.2) or 35 (f.4), cheek with 3 or 4 (rarely 2) rows. Seven (rarely 6^ or 8) scales between the upper lateral line and the dorsal fin origin, 7 or 8 (rarely 6 or 9) between the pectoral and pelvic fin bases. Scales on the ventral aspects of the chest are noticeably smaller than those situated laterally. Fins. Dorsal with 25 (f.7) or 26 (f.2) rays, comprising 16 spinous and 9 (f.7) or 10 (f.2) branched rays. Anal with n (f.2), 12 (f.6) or 13 (f.i) rays, comprising 3 spines and 8 (f.2), 9 (f.6) or 10 (f.i) branched elements. Pectoral 25-0-29-2 (M = 26-7) per cent of standard length. Caudal truncate, scaled on its proximal half. Teeth. In the outer row of both jaws, the teeth are relatively stout, bicuspid and slightly curved ; in a few specimens some posterolateral upper teeth are unicuspid and enlarged. There are 40-65 (M = 50) teeth in the outer row of the upper jaw. Teeth forming the inner rows are relatively large and tricuspid, and are implanted somewhat obliquely so that the crowns point inwards. There are 2 or 3 inner rows in the upper jaw, and 2 (3 in one specimen) in the lower. Osteology. No complete skeleton is available. The lower pharyngeal bone is, compared with that of similar species, small. Its dentigerous surface is broader than long, and narrows rapidly at about the midpoint so that not only is the whole bone relatively small, but so is the area of pharyngeal teeth. The teeth are fine, com- pressed and distinctly cuspidate ; they are rather sparsely distributed in from 22-24 rows. Vertebral counts in 9 specimens are : 13+17 (f.8) and 13 -f 18 (f.i) giving totals of 30 and 31. Coloration : Live colours are unknown. Preserved colours : Males (adult but quiescent). Ground colour dark yellowish-brown. A well-defined lachrymal stripe is present, as are faint traces of two transverse, parallel stripes across the snout. There is some darkening over the preoperculum but this is not concentrated into a FIG. 19. Lower pharyngeal bones (in occlusal view, with dentition shown on one side only) of (A) H. barbarae, and (B) H. chromogynos. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 95 bar or stripe. The branchiostegal membrane is dark, most intensely so below the opercular bones. On the flank is a faint midlateral stripe, and four even fainter and incomplete vertical bars. Dorsal and anal fins are yellowish with a sooty overlay, the anal being darkest along its proximal third ; anal ocelli very faint. Pelvics sooty over a yellowish ground. Caudal yellowish on the margins and distal third, dark brown proximally. Females (adult.} Three distinct colour patterns occur ; two of these are probably identical with the polychrome patterns described for live Hoplotilapia retrodens (see Greenwood, 1956). The colour pattern corresponding with the modal coloration in other polychromatic species has a yellow-brown ground colour with a distinct midlateral dark stripe running from the opercular margin to the caudal origin, and extending onto the caudal fin itself for about half its length ; in two specimens there are four, narrow vertical bars on the flanks, extending from the dorsal fin base almost to the ventral outline. In all specimens a very faint upper line is situated slightly above the upper lateral line. All fins are yellowish-brown. Three out of the eight female specimens have this type of coloration. The second pattern is a piebald, black on silvery-yellow (to yellow brown), the black pigment arranged in blotch-like bars of variable width and dorso-ventral extent. Some blotches extend onto the dorsal, caudal, anal and pelvic fins. The cheeks and snout may be blotched or clear. All fins are yellowish. This pattern does not differ from that shown by other species with a piebald coloration in females. Four of the eight H. barbarae females are piebald. The third pattern, found only in one specimen, has an orange-yellow background peppered with fine melanophores which are, in places, aggregated into blotches (especially on the dorsum) with an irregular distribution. Some dark blotches occur on all fins (but especially the dorsal and caudal) ; these spots are smaller than those on the body. The ground colour of all fins is light orange-yellow. Ecology. Habitat. The nine known specimens came from three different sites. Two of these are shallow, exposed and sandy beaches, the third, shallow water (ca. 10 ft. deep) over a hard substrate a short distance off-shore and near the water- lily zone fringing a papyrus swamp. Food. Seven of the 9 specimens examined contained ingested matter in the guts. In each case this consisted of from 1-8 (mode 5) recently fertilized cichlid ova. In two specimens these could have been part of the fish's own brood swallowed during capture, because the fishes have recently spent ovaries. But, judging from the advanced stages of oogenesis shown by four other specimens, and the fact that the fifth is a male, it seems reasonable to conclude that the embryos were taken as food (see Greenwood, 1959, for a discussion of other paedophagous Haplochromis species). Breeding. Nothing is known about the breeding habits of H. barbarae. All nine specimens are sexually mature ; the sole male (102 mm. S.L.) is the second largest specimen. Affinities. Anatomically, H. barbarae is very like H. brownae, a generalized species exhibiting many characters suggestive of affinity with anatomically generalized predators such as H. guiarti (Greenwood, 1962). 96 P. H. GREENWOOD Haplochromis barbarae differs from H . brownae in having a narrower interorbital (24-2-26-7, M = 25-5% of head, cf. 26-0-34-0, M = 29-8%), a smaller eye (22-9-27-4, M = 25-8% of head, cf. 26-0-31-3, M = 28-6%), fewer gill rakers (8 or 9, cf. 9-12 [modes 10 and n]), and smaller scales, especially on the ventral aspects of the chest. There is also a difference in dention in that the outer teeth of this species are stout bicuspids whereas in H. brownae of a similar size the teeth are slender unicuspids. It differs from H. guiarti in having a shorter snout (30-0-32-4, M = 31*5% of head, cf. 31-7-37-5, M = 34-4) and lower jaw (36-6-41-2, M = 38-0% of head, cf. 39-2-48-2, M = 44-4%), fewer gill rakers (8 or 9, cf. 9-11, mode 10) and, at com- parable sizes, the presence of bicuspid teeth anteriorly in both jaws (unicuspid in H. guiarti} . Because of its generalized anatomy and unspecialized dentition, H. barbarae closely resembles a number of other species in the Lake Victoria flock, viz. H. cinereus, H. macrops, H. lacrimosus and H. chromogynos (see Greenwood, 1959 for the latter species and Greenwood, 1960 for the three former). In addition to their anatomical similarities, H. barbarae and H. chromogynos also share (with several other and structurally unrelated species) the piebald coloration in females. However, H. barbarae differs from H. chromogynos in having a longer lower jaw (36-6-41-2, M = 38-0% of head, cf. 30-0-34-4, M = 32-5%), and, when specimens of equal size are compared, bicuspid instead of slender unicuspid teeth (specimens of H . chromo- gynos < 95 mm. S.L. have a mixed bi- and unicuspid dentition but larger individuals have only unicuspid teeth). The shape of the pharyngeal bone differs in the two species, and it is this character (see p. 94 and fig. 19) which most readily distinguishes H. barbarae from the other three species mentioned above, none of which is known to have piebald females or paedophagous habits. The peculiar feeding habits of H. barbarae immediately suggest some affinity with the other paedophagous Haplochromis species, and because H. barbarae is anatomic- ally unspecialized, particularly with the more " generalized " paedophages, H . obesus and H. cronus. However, in many morphological details these two differ consider- ably from H. barbarae. For instance, the peculiar dental morphology of H. obesus (with the small teeth deeply embedded in the gums), and the stout unicuspid teeth of H. cronus, serve as immediately diagnostic characters, as do the several mor- phometric differences between the species. It is of interest, however, to recall that piebald females occur in all three species ; but, since this character appears in other and widely different species, it is not thought to have any phyletic significance. Any supposed close phyletic relationship between H. barbarae and the other members of the paedophagous species group would also be difficult to substantiate. At present all that can be suggested is that H. barbarae might represent an isolated line, derived from a generalized and probably insectivorous stem, paralleling trophically the paedophagous species group (itself probably of diphyletic origin). Alternatively, H. barbarae might represent a survivor of the stem from which such species as H. cryptodon and H. microdon arose ; more will have to be learned about the anatomy of H. barbarae before this possibility can be substantiated. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 97 STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. Locality Collector UGANDA B.M. (N.H.) 1966.3.9.243 Napoleon Gulf, off Jinj a . . E.A.F.R.O. B.M. (N.H.) 1966.2.21.5 Beach near Nasu Point (Holotype) (Buvuma Channel) . . . E.A.F.R.O. TANZANIA B.M. (N.H.) 1966.3.9.244-251 Beach near Majita . . . E.A.F.R.O. Haplochromis tridens Regan and Trewavas, 1928 (Text-fig. 20) Haplochromis tridens Regan & Trewavas, 1928, Ann. Mag. nat. Hist, (io), 2, 226. LECTOTYPE : a fish 116-0 mm. standard length (caudal fin damaged), B.M. (N.H.) reg. no. 1928.6.2.41, collected in Tanzanian waters at Michael Graham's station 234 (i° 4' S, 32° 13' E), at a depth of over 100 ft. (Graham, 1929). DESCRIPTION : based on 16 specimens (including the lectotype and the paralecto- type) 72-119 mm. standard length. Depth of body 30-1-36-2 (M = 33-5) per cent of standard length, length of head 32-0-37-2 (M = 35*4) per cent. Dorsal head profile straight, but interrupted by the prominent premaxillary pedicels, sloping steeply at 4o°-45°. The cephalic lateral line system with prominent pores, especially those of the preopercular and preorbital canals. These pores are probably larger than in any Lake Victoria Haplochromis (including H . pachycephalus and H. hoops] . Preorbital depth 16-0-20-9 (M = 17*2) per cent of head, least interorbital width 15-0-19-5 (M = 16-7) per cent. Snout 1-0-1-3 (mode i-i) times as long as broad, its length 28-0-34-8 (M = 30-4) per cent of head. Eye with an oval, horizontally aligned pupil, eye diameter 25-6-34-0 (M = 30-7) per cent of head, depth of cheek 17-5-22-8 (M = 19-2) per cent. Caudal peduncle 16-7-19-8 (M = 17-9) per cent of standard length, 1-4-1-9 (mode 1-6) times as long as deep. Mouth horizontal or slightly oblique, lower jaw projecting slightly and with a distinct mental projection ; length of lower jaw 43-3-51-8 (M = 47-5) per cent of head, 2-0-2-8 (mode) times as long as broad. Premaxilla slightly expanded in the midline. Posterior tip of the maxilla generally reaching to below the anterior part of the eye, and occasionally to below the pupil ; rarely reaching only to the vertical through the anterior margin of the orbit. Gill rakers : relatively slender, except for the lower 1-3 which are reduced, and the upper 2-4 which are usually flat ; 8-n (mode 9) on the lower part of the first gill arch. Scales : strongly ctenoid, especially those on the chest region. Lateral line with 31 (f.4), 32 (f.5), 33 (f.4) or 34 (f.2) scales. Cheek with 3 (mode) or 4 rows. Five and a half to 6^ (modes 6 and 6|) scales between the upper lateral line and the dorsal fin origin, 5-6^ (mode) between the pectoral and pelvic fin bases. 98 P. H. GREENWOOD FIG. 20. Haplochromis tridens, paralectotype, about -73 times natural size. Drawn by Miss M. Fasken. Fins. Dorsal with 23 (f.y), 24 (f.y) or 25 (f.2) rays, comprising 15 (£.4) or 16 (f.12) spinous, and 7 (f.6), 8 (f.6), 9 (f.3) or 10 (f.i) branched rays. Anal with 10 (f.io), II (f.5) or 12 (f.i) rays, comprising 3 spines and 7 (f.io), 8 (f.5) or 9 (f.i) branched elements. (N.B. More than usual difficulty was experienced in deciding if the last dorsal and anal ray was a single, deeply divided element, or two distinct rays ; this may account for the number of specimens with low (7) branched ray counts). Pectoral 27-0-33-3 (M = 29-3) per cent of standard length. Pelvics with the first branched ray produced, proportionately more so in adult males. Caudal truncate. Teeth. The trivial name " tridens " was given to the species because the larger of the two syntypes had only tricuspid teeth in the outer tooth row of both jaws, an unusual (probably unique) feature for a Haplochromis species. This condition is, however, rare in the species. Uni-, bi- and tricuspid outer teeth may all occur in a single specimen. Tricuspid teeth, at least in the upper jaw, are usually found posterolaterally, the bi- and unicuspids laterally and anteriorly. This arrangement is by no means constant ; in some specimens tricuspids occur anteriorly and antero- laterally. One fish (72 mm. S.L.) has only bicuspids in the upper jaw, and another (81 mm.) only unicuspids. There is no apparent correlation between the predomin- ance of any one tooth form and the size of the fish. In three out of the sixteen specimens examined, no tricuspid outer teeth were found in either jaw. There are 58-80 (M = 66) teeth in the outer row of the upper jaw. Variation in the type of outer teeth occurring in the lower jaw follows the same pattern as in the upper jaw. Teeth forming the inner rows in both jaws are predominantly tricuspids. These teeth are relatively large, are stout, and have the median cusp larger than the lateral ones. There are usually 2 inner rows (occasionally 3, rarely 4) in the upper jaw, and 2 (rarely i) in the lower. Osteology. No entire skeleton is available. The lower pharyngeal bone has its dentigerous surface equilateral or slightly broader than long. The lower pharyngeal teeth have cuspidate, compressed crowns, and cylindrical necks ; the teeth are arranged in 22-26 rows. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 99 Vertebral counts in 10 specimens are : 13+16 (f.y) and 13 -f 17 (£3), giving totals of 29 and 30. Coloration. Live colours are unknown. Preserved specimens : Males (adult and sexually active) : ground colour grey-blue (gun-metal) above the upper lateral line, greyish silver on the flanks, and light dusky silver on the chest and belly. Dorsal and caudal fins hyaline. Anal hyaline except for a dusky area between the spines, and black lappets ; the dusky area extends as a fairly distinct line at the base of the spinous part, and may be expanded basally onto the soft fin. There are two, large, grey- white anal ocelli. Pelvics dusky, darkest on the lateral four-fifths of the fin. Males (adult but quiescent} have a ground coloration more like that of females. All fins are hyaline except for a faint darkening between the anal spines, and on the pelvic fins. Females (juvenile and adult) : greyish silver above, shading to silver below. One specimen (ovaries ripening) is darker dorsally (almost brown), and has a broad, interrupted midlateral stripe running from behind the operculum to about half the length of the caudal fin (on which the stripe narrows) ; the band is interrupted at about the middle of its length. All fins yellowish to hyaline. Ecology. Habitat. Excepting the two types, all the material came from one trawl haul at an unknown locality (thought to be off the Kenya coast). The types came from deep water (more than 100 ft.) some distance off-shore, and from over a soft bottom. The large eyes and hypertrophied pores of the cephalic lateral line system certainly suggest adaptations to a deep water habitat. Food. Only 6 of the 10 specimens examined (all from the same, and unknown, locality) contained ingested material in the gut. In each, the predominant contents are undigested blue-green algae, and empty diatom frustules. Two specimens have, in addition, a few fragments of Crustacea, and some unidentifiable insect remains. Breeding. All fourteen of the non-typical specimens are adults. The types are not well-preserved internally, but both appear to be females. Affinities. The admixture of tri-, uni- and bicuspid outer teeth together with the hypertrophy of the cephalic lateral line pores, provide a trenchant means of distin- guishing H. tridens from other species in the lake. Indeed, it is difficult to suggest any close relationship between this species and any other so far considered. Perhaps some relationship will become apparent when more is known about the numerous and small species of Haplochromis which make up the bulk of fishes caught by trawling in the deeper waters of Lake Victoria (Greenwood, unpublished). Anticipating these results, it is possible to say that the elongate body and general " predatory " facies (especially the large mouth) of H . tridens are not common amongst these species, and nor is the H. tridens dental type. Amongst the larger species with a predatory facies, H. tridens most closely resembles H . victorianus. There are, however, many differences between the species, not least of which are their differences in ecology and the much smaller adult size of H. tridens. Haplochromis victorianus differs also in its dentition, broader interorbital region (21-5-24-5, M = 22-6% of head, cf. 15-0-19-5, M = 16-7%) smaller and rounder eye (21-7-25-5, M = 23-6% head, cf. 25-6-34-0, M =30-7%), and deeper P. H. GREENWOOD cheek (22-5-26-2, M = 24-6% head, cf. 17-5-22-8, M = i9'9%). It should be remembered, however, that the data for H. victorianus are derived from larger speci- mens than are available for H. tridens, and that the most trenchant morphometric differences are in characters most affected by allometric growth. Nevertheless, it seems very unlikely that H. victorianus and H . tridens are close relatives. STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. B.M. (N.H.) 1928.6.2.41 (Lectotype) B.M. (N.H.) 1928.6.2.42 (Paralectotype) B.M. (N.H.) 1966.3.9.152-165 Locality TANZANIA 104' 8,32° 13' E 104' S, 32° 13' E LAKE VICTORIA Locality unknown (? Kenya coast) Collector M. Graham M. Graham E.A.F.R.O. Haplochromis orthostoma Regan, 1922 (Text-fig. 21) Pelmatochromis spekii (part): Boulenger, 1915, Cat. Afr. Fish., 3, 417. Haplochromis orthostoma Regan, 1922, Proc. zool. Soc. Londn., 184, fig. 9. HOLOTYPE : a male (probably adult), 91 mm. standard length (B.M. [N.H.] reg. no. 1912.10.15.67) from Lake Salisbury (Kyoga system). The specimen is now in very poor condition, and has lost most of its scales. This species, possibly one of the two most distinctive looking members of the Victoria-Kyoga flock, was described from a single specimen. Since then, two further specimens have been collected from Lake Salisbury, and three superficially similar specimens have been caught in Lake Victoria. The latter specimens differ from the Salisbury fishes in their dentition (and some other characters) and are thought to represent a distinct species. It is for this reason that I am redescribing the Lake Salisbury species in this paper. The unusual physiognomy of H. orthostoma is well shown in text-figure 21 ; the snout and nuchal region meet at a distinct angle, the nuchal musculature not bulging anteriorly and laterally. Since there are only three specimens available, morphometric characters are tabulated below. The holotype is indicated with an asterisk. S.L. (mm.) 83-0 *9i-o Depth t 36-3 Head Preorb. Interorb. Snout Eye Cheek Caudal t /o /o /o /o /o Peduncle f 36-0 19-6 20-4 32-7 24-5 24-5 19-3 36-1 2O-O 23-3 30-0 21-7 30-0 J7'5 35'5 18-5 21-5 30-8 22-8 30-8 16-5 t Per cent of standard length. % Per cent of head length. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 101 Mouth very oblique, sloping at ca 5o°-7o°, the lower jaw projecting, its length 51-0-56-6 per cent of head length, and 2-5-3-4 times as long as broad. Lips not thickened, the lower jaw with a distinct mental protuberance. Posterior tip of the maxilla reaching a point about midway between the vertical through the anterior orbital margin and that through the nostril. Snout 1-1-1-3 times as long as broad, its dorsal surface slightly rounded, the premaxillary pedicels not prominent. Caudal peduncle 1-5-1-6 times as long as deep. Gill rakers : moderately stout, the lower 1-3 reduced, the upper 2 or 3 flattened (anvil-shaped in one fish) ; 9 or 10 on the lower part of the first gill arch (n on one arch of a fish with 9 rakers on the other arch) . FIG. 21. Haplochromis orthostoma, holotype. From Regan, Proc. zool. Soc., about •86 times natural size. Scales : ctenoid ; lateral line with 30 or 31 scales (Regan gives 33 for the now scaleless type), cheek with 3 or 4 rows (4 or 5 in the type according to Regan). Six or 6| between the upper lateral line and the dorsal fin origin (6 or 7 in the type ace. Regan), 6 or 7 between the pectoral and pelvic fin bases, the chest scales small. Fins. Dorsal with 14 (f.i) or 15 (f.2) spines and 9 branched rays, anal with 3 spines and 8 (f.2) or 9 branched rays. First branched pelvic ray not produced in the type, but elongate in the two other specimens (male and female) . Caudal subtrucate. Teeth. In the two larger specimens (83 and 91 mm. S.L.), the outer teeth in both jaws are a most distinctive feature. These teeth are slender and unicuspid with sharply curved tips directed medially ; those situated anteriorly are somewhat larger than the others. There are 66 and 61 teeth in this row for the two specimens respectively. In the lower jaw, the outer teeth, at least anteriorly, are somewhat stouter than their opposites in the upper jaw. The smallest specimen (67-5 mm. S.L.) has outer teeth quite unlike those of the larger individuals. In the upper jaw, the anterior and lateral teeth are stout and bicuspid, those situated laterally and posteriorly are stout and tricuspid. All outer 102 P. H. GREENWOOD teeth in the lower jaw are bicuspid and stout, and are a little stouter than the upper jaw teeth. There are 68 teeth in the outer row of the upper jaw. The inner teeth in the two larger fishes are unicuspid, small, slender and implanted obliquely ; in the smallest fish they are small and tricuspid. The dental arcade in all specimens is V shaped with a rounded apex ; there are 2 inner tooth rows in the upper jaw, and I or 2 rows in the lower jaw. Osteology. No complete skeleton is available. The lower pharyngeal bone has a triangular and equilateral dentigerous area. The teeth are relatively slender, with bicuspid and weakly compressed crowns, and are arranged in 20-22 rows ; except in the smallest fish the teeth of the two median rows are coarser than the lateral rows. Vertebral counts for 2 specimens are : 13+16 (type) and 12 + 16. Coloration : Live colours are unknown. The type is now completely colourless ; originally it was described as greyish, with a dark lachrymal stripe, and blackish pelvic fins. A juvenile female has a brown ground coloration, with very faint traces of four broad vertical bars on the flanks running from the dorsal fin origin to about the level of the ventral margin of the pectoral fin. No lachrymal stripe is visible. The lower jaw (especially over its anterior half) is rather dusky. The dorsal and caudal fins are yellowish, the former with black lappets. Anal dusky yellow. Pel vies faintly dusky, especially at their tips. Pectorals yellowish-grey. Adult Male (probably fixed in alcohol) : light brown above, shading to silver on the mid-flanks, and silvery white on the belly. Snout dark, as are the lips ; branchio- stegal membrane pale. A faint lachrymal stripe is visible. Dorsal fin greyish, with black lappets. Anal greyish, with two white ocelli. Caudal grey, the melanophores most concentrated along its midline. Pelvics dark along the anterior quarter, hyaline elsewhere. Ecology. No information is available on the habitat or food of H. orthostoma, nor is there any information on the breeding habits of the species. The two males (91 and 83 mm. S.L.) are adult, and the female (67-5 mm.) is apparently immature. Affinities. The peculiar head shape, large and very oblique mouth, coupled with the peculiar tooth form (at least in the larger fishes), serve to distinguish the species from all others in the Lake Victoria-Kyoga flock. The nearest relative is H. par- orthostoma from Lake Victoria (see below) . Further material, and field observations, may yet show that the two species are not distinct at that level. STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. Locality Collector B.M. (N.H.) 1912.10.15.67 Lake Salisbury . . . . Presented by (Holotype) F. J. Jackson B.M. (N.H.) 1958.12.5. 173 Ongino, Lake Salisbury . . Pitman B.M. (N.H.) 1966.3.9.252 Lake Salisbury . . . . E.A.F.R.O. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 103 Haplochromis parorthostoma sp. nov. (Text-fig. 22) HOLOTYPE : an adult male, 117 mm. standard length, from near Zero Island (Buvuma Channel), Uganda. B.M. (N.H.) reg. no. 1966.2.21.4. DESCRIPTION. The overall similarity between this species and H. orthostoma is great, particularly since both share a peculiar head profile not seen in any other Haplochromis species from Lakes Victoria or Kyoga. The dorsal head profile is strongly concave, with the nuchal region meeting the snout at a noticeable but rounded angle, the junction emphasized by an anterior bulge of the cephalic epaxial body musculature. This muscular protuberance gives the fish a pronounced " forehead ", especially in the frontal plane. Since only three specimens are available, morphometric data are tabulated below ; the holotype is marked with an asterisk. Preorb. 16-7 19-5 t Per cent of standard length. % Per cent of head length. Mouth oblique, sloping upwards at ca. 40°-5o°, jaws equal anteriorly, or the lower projecting slightly. Lower jaw length 48-3-53-5 per cent of head, 2-3-2-4 times as long as broad ; chin with a distinct protuberance. Posterior tip of the maxilla reaching a point nearer the vertical through the anterior orbital margin than one through the nostril. Snout 1-2 times as long as broad, with a convex dorsal surface ; premaxillary pedicels not prominent. Lips moderately thickened. Caudal peduncle 1-1-1-3 times as long as deep. S.L. Depth Head (mm.) t t 86-0 38-4 35-o IIO-O 42-3 36-4 117-0 41-0 35'° Interorb. Snout Eye Cheek Caudal o/ /o % o/ /o /o Peduncle f 2O-O 31-6 27-7 25-0 17-5 17-5 33-8 25-0 27-5 12-7 19-5 34-9 24-4 26-8 14-5 FIG. 22. Haplochromis parorthostoma. Drawn by Lavinia Beard. 104 p- H- GREENWOOD Gill rakers : moderately stout (relatively stouter in one specimen) , the lower 1-3 reduced, the upper 2 or 3 expanded and anvil-shaped ; 9 (f.2) or 10 rakers on the lower part of the first gill arch. Scales : ctenoid, lateral line with 30 or 32 (f.2) scales, cheek with 2 or 3 rows (in the former specimen, the scales not covering the ventral aspects of the cheek). Six or 7 scales between the upper lateral line and the dorsal fin origin, 6 or 7 between the pectoral and pelvic fin bases. Fins. Dorsal with 15 (f.2) or 16 spines and 8, 9 or 10 branched rays. Anal with 3 spines and 8 or 9 (f.2) branched rays, the spines short and stout. Pectoral 24-5-26-2 per cent of standard length. First branched pelvic ray not or slightly produced (all specimens are males) . Caudal almost rounded, scaled on its basal two-thirds. Teeth. The outer row in both jaws is composed of unicuspid, slender and slightly curved teeth, the curvature being gentle and not confined to the distal part of the tooth (cf. H. orthostoma, p. 101) ; in one fish, the last three teeth in the upper jaw are larger, stouter and straighter than the anterior ones. A few bicuspid, moderately stout teeth occur posteriorly in the lower jaw of the smallest (86 mm.) fish. In no specimen are the lower jaw teeth stouter than those in the upper jaw. There are 38, 40 and 52 teeth in the outer row of the upper jaw. Teeth forming the inner series are small, tricuspid and weakly tricuspid, and are arranged in 3 or 4 rows in the upper jaws and in 2 rows in the lower. The dental arcade is V shaped, with the apex broadly rounded. Osteology. No complete skeleton is available. The lower pharyngeal bone has its triangular dentigerous surface as long as broad, or slightly broader than long. The teeth, arranged in 24-30 rows, are fine, with weakly compressed bicuspid crowns ; those in the two median rows are but slightly larger than the lateral teeth. Vertebral counts in 3 specimens are : 13+15 and 13 -j- 16 (f.2). Coloration : Live colours are known for a sexually active (but not ripe) male. Ground colour dark slate-grey, with faint vertical bars of a darker shade. Dorsal fin dark grey with crimson lappets on the posterior two-thirds of the spinous part, and a dark crimson margin to the entire soft part. Caudal dark grey with a crimson flush, particularly intense on its lower half. Entire anal fin, except for a dark base, crimson. Pel vies black on the anterior third, remainder dull crimson. Preserved material : Males (adult) . Ground colour light brown (including the branchiostegal membrane in two fishes ; this membrane blackish in the third specimen) ; flank crossed by 5 or 6 dark but incomplete bars, each bar originating just above the upper lateral line and extending to about the level of the ventral margin of the pectoral fin. A fairly distinct vertical lachrymal stripe runs from the antero ventral margin of the orbit to the angle of the lower jaw. Dorsal fin yellowish- brown along its margin, but dark brown between the rays. Anal yellowish, sooty or dark brown between the branched rays ; very faint indications of 2 or 3 whiteish ocelli. Caudal light to dark brown (almost black). Pelvics dusky on the anterior third to half, otherwise hyaline. Ecology. One specimen came from an exposed, sandy beach, another from a rocky outcrop in about 20 ft. of water near an off-shore island, and the third from over a rocky shelf in about 40 ft. of water, also near an island. A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 105 No information is available on the food of H. parorthostoma. The three specimens are adult males. Affinities. Haplochromis parorthostoma seems to be very closely related to H. orthostoma of the Kyoga system, at least in its peculiar head-shape. There is, however, a marked interspecific difference in the shape of the outer teeth. In H. parorthostoma these teeth are gently curved, whereas in H. orthostoma they have sharply recurved crowns but relatively straight necks. Furthermore, the inner teeth of H. orthostoma are unicuspid and arranged in one or two rows, but in H. parorthostoma are tricuspid and arranged in three or four rows in the upper jaw (two rows in the lower). An exception (at least with regard to outer tooth shape) is provided by the smallest specimen of H. orthostoma whose stout, erect and bicuspid teeth do not resemble those of its larger congeners (or, for that matter, H. parortho- stoma}. But, since the cranial morphology of this small fish is so like that of larger H. orthostoma individuals it is included in that species. Haplochromis parorthostoma and H . orthostoma differ in characters other than dental ones. The bulging cranial epaxial muscles of H. parorthostoma impart a different shape to the dorsal head profile, which is further modified by the less oblique mouth. The available samples also suggest that H. parorthostoma has a bigger eye than H. orthostoma (and this despite the fact that the specimens of H. parorthostoma are larger) and probably a shorter and broader lower jaw. Clearly, much more material is required before it will be possible to reach more definite conclusion about the status of the two species. Even then the decision will be complicated by the fact that they are allopatric, and their areas of distribution are physically isolated (now by the virtually impenetrable Owen Falls dam, and previously by the Ripon Falls, perhaps not a complete barrier to migration from Lake Victoria to the Kyoga system). For the moment it is not possible to speculate on the wider relationship of these two species. STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. Locality Collector Uganda B.M. (N.H.) 1966 .2.21 Near Zero Island (Holotype) (Buvuma Channel) . . . E.A.F.R.O. B.M. (N.H.) 1966.3.9.253 Near Zero Island (Buvuma Channel) . . . E.A.F.R.O. B.M. (N.H.) 1966.3.9.254 Near Kazima Island . . . Uganda Fish. Dept. Haplochromis apogonoides sp. nov. (Text-fig. 23) HOLOTYPE an adult male, 118 mm. standard length, from Ekunu Bay, Uganda. B.M. (N.H.) reg. no. 1966.2.21.3. Named " apogonoides " because of its fancied resemblance to the genus Apogon. io6 P. H. GREENWOOD DESCRIPTION : based on eight specimens (including the hole-type) 112-132 mm. standard length ; with one exception (a quiescent female) all are males. Depth of body 36-0-39-6 (M = 38-1) per cent of standard length, length of head 35-0-37-2 (M = 36-1) per cent. Dorsal head profile curved (strongly so in some specimens), sloping fairly steeply (ca. 40°), the premaxillary pedicels not prominent. Preorbital depth 13-6-16-7 (M = 15*2) per cent of head length, least interorbital width 27-5-31-0 (M = 29-5) per cent. Snout 1-3-1-5 (mode 1-4) times as broad as long, its length 29-3-33-4 (M =31-2) per cent of head, eye 25-3-27-5 (M = 26-6), depth of cheek 27-8-31-0 (M — 29-3) per cent. Caudal peduncle 15-3-19-8 (M = 18-0) per cent of standard length, 1-5-1-7 times as long as deep. Mouth slightly to moderately oblique, lips somewhat thickened, the jaws equal anteriorly. Lower jaw 45-0-51-0 (M = 47-9) per cent of head, 1-2-1-5 (modal range 1 CM. FIG. 23. Haplochromis apogonoides. Drawn by Barbara Williams. 1-4-1-5) times as long as broad. Posterior tip of the maxilla reaching to below the pupil. Premaxilla not expanded medially. Gill rakers : stout, the lower 1-3 reduced, the upper 3 or 4 flat ; 8-10 on the lower part of the first gill arch. Scales : ctenoid ; lateral line with 32 (f.4), 33 (f.3) or 34 (f.i) scales. Cheek with 3 or 4 (mode) rows. Six to 8 (mode 6) scales between the upper lateral line and the dorsal fin origin, 6-8 (modes 6 and 7) between the pectoral and pelvic fin bases. Fins. Dorsal with 22 (f.i), 24 (f.6), or 25 (f.i) rays, comprising 14 (f.2) or 15 (f.6) spinous and 8 (f.i), 9 (f.5J or 10 (f.2) branched rays. Anal with 10 (f.i), n (f.5) or 12 (f.2) rays, comprising 3 spines and 7 (f.i), 8 (f.5) or 9 (f.2) branched elements. Pectoral 29-0-34-1 (M = 30-8) per cent of standard length. Pelvics with the first ray slightly produced. Caudal truncate to subtruncate, scaled on its basal half to two-thirds. Teeth. The outer teeth in both jaws are a characteristic feature of the species, being unicuspid, moderately stout and with very strongly recurved tips. Such teeth A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 107 are otherwise found only in the H. sauvagei species group (see Greenwood, 1957). There are 50-60 (M = 58) teeth in the outer row of the upper jaw. The inner teeth are also unicuspid, large and recurved, and are implanted obliquely. There are 2 rows in the upper jaw, and I or 2 rows in the lower jaw. A distinct space separates the inner series from the outer row. Osteology. No complete skeleton is available. The lower pharyngeal bone is relatively stout, the dentigerous surface noticeably broader than long (i^ to ij times) . The teeth are stout and cuspidate, and are arranged in from 16-20 rows, those of the two median rows being slightly coarser than the others. Vertebral counts in 7 specimens are : 13+15 (f.i), 13 + 16 (f.5) and 14 + 16 (f.i), giving totals of 28-30. Coloration. Live colours are unknown. Preserved coloration : Males (adult and sexually active, but probably not ripe). Ground colour light yellow-brown, with a silvery underlay on the flanks. Belly, chest and branchiostegal membrane dusky, the branchiostegal membrane darkest below the opercular region. On the flanks there are faint traces of 4 or 5 fairly broad vertical bars which become very faint dorsally and ventrally ; the anterior 3 or 4 bars merge ventrally with the dark belly coloration. A dark lachrymal stripe is present ; it does not reach the ventral margin of the preorbital, but appears to pass upwards across the eye. Dorsal fin yellowish, the soft part maculate in some specimens ; also in some fishes the lappets are black. Caudal yellowish marginally and on the distal third to quarter, dark yellow-brown proximally. Anal yellowish, with one large greyish to whiteish ocellus faintly outlined in black. Pel vies sooty, the colour less intense between the last two or three rays. Female (adult and quiescent) . Ground colour greyish-silver. A very faint lachry- mal bar extends from below the orbit to below the posterior tip of the maxilla ; a dark spot on the upper part of the eye suggests that this bar may pass across the eye. All fins are yellowish, the dorsal with dusky lappets, and the pelvics with a faint duskiness over the anterior rays. Ecology. Habitat. The species has been caught in two localities only. In one, a sheltered bay, the water was between 20 and 30 feet deep, and the substrate of organic mud. The second locality was at a depth of about 80 ft in the Buvuma Channel, near Buvuma Island ; again the substrate (on which the nets were set) was organic mud. Food. Regrettably, the gut was empty in all except one specimen ; this fish contained only a little, unidentifiable sludge. Breeding. Apart from the sexually inactive female (132 mm. S.L.), all the speci- mens are adult and active males. Affinities. The stout, unicuspid outer teeth with sharply recurved crowns immediately suggest affinity with H. sauvagei, H.prodromusa.ndH.granti (Greenwood, 1957). Like these species, the dorsal head profile of H. apogonoides is strongly rounded. However, unlike these species, there are fewer rows of inner teeth in H. apogonoides, there is a distinct interspace between the inner and outer tooth rows, and the lower jaw is much longer (45-0-51-0, M =47-9% head cf. 30-6-37-7, M = 34-5% for H. sauvagei, 30-5-37-8, M = 34-3% for H. prodromus, and 22-2-30-6, io8 P. H. GREENWOOD M = 26-8% for H. granti). Haplochromis apogonoides also differs from these species in its broader snout, and from H. granti in the anatomy of the jaws (see Greenwood, 1957). Superficially, H. apogonoides resembles H. cronus (Greenwood, 1959) and, to a lesser degree, H. empodisma and H. michaeli (see Greenwood, 1960 and 1962 for the species respectively). From H . cronus, it is differentiated by its dental morphology and its longer lower jaw. From H. empodisma, H. apogonoides differs in its longer lower jaw (45-0-51-0, M = 47-9 per cent of head, cf. 39-1-48-7, M = 43-9%), dental morphology (strongly cf. gently curved teeth), broader interorbital (27-5-31-0, M = 29-5% head, cf. 20-6-28-6, M =24-3), broader snout, and noticeably wider dentigerous surface on the lower pharyngeal bone( if-ij times as broad as long, cf. longer than broad). Haplochromis apogonoides differs from H. michaeli in dental morphology (see above), in its slightly deeper body, shallower preorbital, broader interorbita , markedly broader snout, and deeper cheek (27-8-31-0, M = 29-2% of head, cf. 22-9-27-7, M = 25-8%). The lower pharyngeal bone is similar in both species, but the teeth in H. apogonoides are stouter. The marked similarity between the outer jaw teeth of H. apognoides and those in species of the H. sauvagei complex (which includes H. xenognathus, a species not mentioned above because of its distinctive jaw morphology, tooth pattern, and head shape) suggests that H. apogonoides might be an off-shoot from this species group. Unfortunately, nothing is known about the feeding habits of this species ; all members of the H. sauvagei group are snail-eaters with the unusual habit of removing the snail from its shell before ingesting it. The large mouth and not especially strong jaws of H. apogonoides, do not, however, suggest similar feeding methods ; rather, these characters indicate piscivorous habits. STUDY MATERIAL AND DISTRIBUTION RECORDS Museum and Reg. No. Locality Collector UGANDA B.M. (N.H.) 1966.2.21.3 Ekunu Bay .... E.A.F.R.O. (Holotype) B.M. (N.H.) 1966.3.9.238-242 Ekunu Bay .... E.A.F.R.O. LAKE VICTORIA B.M. (N.H.) 1966.3.9.236-237 Locality unknown . . . E.A.F.R.O. DISCUSSION Phytogeny In an earlier paper (Greenwood, 1962) I outlined the various morphological trends shown by the piscivorous species, and suggested a possible ancestral morphotype (represented today by H. brownae) from which the different lines could have evolved. Also in that paper I indicated two major, and two minor, possibly phyletic groups of fish-eating predators. Additional information provided by the species described A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 109 above does not affect the suggested trends, but does alter the phyletic picture. It now seems likely that there are three major phyletic lines, and probably three minor ones as well. As mentioned before (Greenwood, op. cit.}, possible phyletic lines amongst pisci- vorous species are less readily detected and defined than those of other trophic groups. In these latter there are dental as well as somatic characters which may be used for this purpose, but amongst the piscivores (at least when adult) the teeth are invariably unicuspid and of a very similar form. The principal " group " characters in these fishes are neurocranial shape, and body form ; the latter character often shows a greater or lesser degree of intergroup convergence. Using these two character complexes, I at first recognized two species aggregates, the " sen anus " group, and the " mento-macrognathus " group. The former comprises the supposedly more generalized, broad-headed and deep-bodied species, and the latter group the more specialized, slender-bodied forms. Information obtained from the species described in this paper suggests that my " mento-macrognathus " group consists of two groups, neither of which is as readily defined as the " serranus " group, but both being easily distinguished from that group. One of the minor groups (that of H. percoides, H. flavipinnis and H. cavifrons) is now thought to be polyphyletic (see p. 113). For convenience the groups will be referred to by the trivial epithet of a con- stituent species, which species, however, is not necessarily to be considered a " typical " member of the group. Indeed it is often difficult to determine just what a " typical " species would be ; at the most, the nominate species of a group is representative of a structural type found in three or four species of the group. The three major groups are : (i) The " serranus " group, consisting of H. serranus, H. victorianus, H. nyanzae, H. spekii, H. maculipinna, H. boops, H. thuragnathus and H. pachycephalus : morphologically, this is a relatively homogeneous group (but, see also p. no). (ii) The " altigenis " group, comprising H. guiarti, H. bayoni, H. dentex, H. pseudo- pellegrini, H. altigenis, H. pellegrini and H. dichrourus ; a number of subgroups (some monotypic) can be recognized, and are discussed later. (iii) The " prognathus " group, comprising H. paraguiarti, H. acidens, H. prog- nathus, H. bartoni, H. estor, H. gowersi, H. mento, H. mandibularis , H. macrognathus , H. longirostris and H. argenteus. This, the largest species aggregate shows several intragroup trends of which the most distinctive are the H. longirostris— H . argenteus, and the H. mandibularis — H. macrognathus subgroups. Members of the "serranus" group differ from those of the other two groups in having shorter snouts* (one species out of seven with the snout > 36 per cent of head length, cf. fourteen species out of eighteen), deeper bodies (one species out of seven with the body depth < 36 per cent of standard length, cf. seventeen out of eighteen), and broader heads (no species with the interorbital width < 22-6 per cent of head, cf. thirteen out of eighteen species.) Osteologically, the group is character- * The figures given are derived from those for the mean value of a particular character in species of the groups under consideration. ZOOL. 16, 2. no P. H. GREENWOOD ized by having a neurocranial shape nearest that of the generalized skull (see Greenwood, 1962), but with the preotic part elongate. The dorsal skull roof is straight and slopes fairly steeply, is broad both interorbitally and across the otic region, and the supraoccipital crest is high and presents a substantial area for muscle insertion (see fig. 25 in Greenwood, op. cit.}. The " serranus " group could have evolved directly from an H. brownae-like ancestor, the principal morphological changes being an increase in adult size, and those alterations in neurocranial proportions already mentioned. Within the group, the most differentiated species are H. hoops, H. thuragnathus and H. pachycephalus, all three being confined to deep water (see pp. 49, 50 and 41) . Haplochromis boops and H . thuragnathus were apparently derived from an H. maculipinna-like. ancestor, whilst H. pachycephalus seems to show greater affinity with the H. serranus — H. spekii level of organization. Haplochromis maculipinna is also essentially of this affinity, but has markedly larger eyes. In turn, H. serranus is clearly derived from an H. brownae-like stem. The " altigenis " and " prognathus " groups probably evolved from an H. guiarti- like ancestor or ancestors, the latter species also showing affinities with H. brownae. Although both the "altigenis" and "prognathus" groups have included species with a relatively deep body, the main trend shown by both groups is towards a slender, somewhat compressed body-form and a correlated head shape. It is difficult to characterize these two groups, particularly their more basic members. The most trenchant group characters are probably in neurocranial form. In lateral view there is little to differentiate the skull form in the two groups ; both have the preorbital part relatively more elongate than in the " serranus "-type skull, the dorsal profile slopes upward at a rather slight angle (as compared with the angle in skulls of the " serranus " group), and the supraoccipital crest is relatively low. (These generalizations must, however, be modified somewhat for those species which seem to be structurally basal for the groups [H. guiarti and H. pseudopellegrini for the " altigenis " group, and H. paraguiarti and H. acidens for the "prognathus " group]. In these species the preorbital face is less protracted and consequently [since relative neurocranial depth varies little amongst all members of both groups] the dorsal skull roof slopes more steeply and the supraoccipital crest is higher and has a fairly extensive area). However, when the neurocranium is viewed dorsally, a difference between the groups (including their basal species) is apparent (see text-fig. 24). In members of the " altigenis " group, the otic region is relatively broader than in those of the " prognathus " group so that the outline narrows more rapidly (from a point immediately behind the orbit) than in " prognathus " skulls. In these the outline is that of a narrow wedge with the margins closing gradually from a point further behind the orbits. In supposedly basal members of both groups, the otic region is of about equal relative breadth but basal " prognathus " members nevertheless have a more gradual medial inclination of the lateral margins. Further- more, in these species the dorsal skull profile is straighter than in the most basic " altigenis " group member, H. guiarti. Indeed, in most members of the " altigenis " group the profile is more curved than in species of the " prognathus " group. Reasons for considering H. guiarti as a basic morphotype in the radiations of A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES in U FIG. 24. Outlines of the dorsicranium in species of : line A, the " serranus-group ", line B, 4-1, ,» " ^//^^Vc-o-rrmr. " anrl line C the " •broE'nathus-erou'p" . the " altigenis-group ", and line C, the " prognathus-group ZOOL. 15, 2. 7§ H2 P. H. GREENWOOD predatory piscivores have been discussed already (see Greenwood, 1962). Starting from a species similar to H. guiarti, the principal morphological changes seen amongst species of the " altigenis " group involve relative elongation of the pre- orbital face, a slight decrease in skull height (especially in the otic region) and a consequent flattening in the slope of the dorsal skull profile. In these respects H. bayoni represents a fairly marked departure from the basic " guiarti " skull form, but one less marked than that shown by H. dentex, H. altigenis or H. pellegrini, particularly the latter (see fig. 25 in Greenwood, 1962). A link between these forms is provided by the skull of H. pseudopellegrini which is intermediate between the " bayoni " and " altigenis " types. There is not a great deal of difference in body form or jaw morphology among members of the " altigenis " group. Perhaps the most extreme member is H. altigenis itself, a species with the deepest head and con- sequently the largest mouth. Haplochromis pellegrini is, because of its small adult size and relatively small mouth, atypical for the group. There is far greater variation in body form and cranial morphology within the " prognathus " group. Here the basic species, H. paraguiarti and H. acidens, are morphologically similar to H. bayoni of the " altigenis " group and could be derived either from a " bayoni " or a " guiarti "-like ancestor. Haplochromis acidens is, of course, peculiar in that despite its predatory facies, it is apparently a phytophage (see p. 76). Haplochromis prognathus and H. bartoni are, in most respects, very similar to one another and represent the next morphological step in the evolution of such species as H. longirostris and H. mandibularis from a " paraguiarti "-like stem. That is to say, the neurocranium clearly shows narrowing and elongation, and there is a related refinement of body proportions. Haplochromis estor and H. gowersi continue this trend with, in addition, the development of a more oblique angle to the jaws, and in H. gowersi a deepening of the cheek which contributes to a larger buccal cavity. Haplochromis mento probably evolved from a "prognathus "-like ancestor, and shows many of the trends exhibited by H. gowersi and H. estor. However, in this species the mouth is almost horizontal, and the neurocranium is somewhat nearer that of H . prognathus. Also apparently stemming from a " prognathus "-like ancestor is H. mandibularis. Here the trend is towards further narrowing of the skull, deepening of the cheek, increased obliquity of the jaws and lengthening of the lower jaw. This trend culminates in H. macrognathus (see Greenwood, 1962, pp. 180 and 186), a species which could well be a direct descendant of H. mandibularis. A third derivative from a "prognathus " or " bartoni "-like ancestor is H. longi- rostris. In this line there has been little change in neurocranial shape but there is a marked increase in jaw obliquity (much greater, too, than in the " mandibularis "- " macrognathus " line), and a marked refinement in body proportions which results in one of the most slender bodies found amongst Lake Victoria Haplochromis species. These characters are shared by H. argenteus, although the elongate body-form seems less obvious in that species. Haplochromis argenteus could be derived either from a " longirostris "-like stem, or perhaps directly from a " prognathus "-like ancestor. Thus, the " prognathus " group seems to show at least three radiations from a basal " prognathus "-" bartoni " stem, viz. the H. estor, H. gowersi, H. mento sub-group, A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 113 and the H. mandibularis-H . macrognathus , and H. longirostris-H . argenteus species pairs (but see above for possible reservations about the last named pair) . It seems reasonable to assume that the H. prognathus level of organization was derived from a level similar to that shown by the extant species H. paraguiarti. The relationships of that species are, however, less clear-cut. As was mentioned above, H. para- guiarti shows several structural affinities with H. bayoni, a species probably derived from an H. guiarti-like ancestor. But, it is difficult to overrule the possibility that H. paraguiarti evolved independently from an H. brownae-\ike stem. (The status of H. brownae in relation to the piscivorous predators is discussed in Greenwood, 1962). Two species, H . plagiostoma and H. xenostoma, have not been included in the dis- cussion so far. Both are characterized by having the mouth set at a very steep angle to the horizontal (ca. 40°-5o°) ; H. plagiostoma is further characterized by its obliquely truncate caudal fin, and H. xenostoma by its pronounced prognathism. Because of these characters, especially the oblique jaws, neither species shows any obvious superficial similarity with members of the groups discussed above. The neurocranium of H. plagiostoma is of the " serranus " type (see also Greenwood, 1962) but also shows certain " guiarti "-like features. Thus, on this character complex H. plagiostoma could either be associated with the " serranus " group or be looked upon as an isolated offshoot of the " altigenis " group arising from an ancestor near the stem of that complex. In either eventuality, H. plagiostoma is not linked with the basal group by any extant, structurally intermediate species. Haplochromis xenostoma, both in its overall organization and in its neurocranial form, seems to represent a further development from a " plagiostoma " stem. The chief trend involves an increase in adult size, and a relative enlargement of the mouth and jaws. The neurocranium of H. xenostoma reflects these changes, especially in the longer preorbital region ; it is thus essentially a " serranus " group neurocranium. From the available evidence it is impossible to determine whether H. plagiostoma and H. xenostoma are part of the same phyletic lineage or whether the two species are end-points of parallel evolution from " guiarti " and " serranus "-like stems respectively. The two smaller species complexes, the H. michaeli, H. martini, H. squamulatus group and the H.percoides, H . flavipinnis , H. cavifrons group, will now be considered. The relationships of H. michaeli and H. martini are discussed in my paper of 1962, and there is nothing further to add. Haplochromis squamulatus is included with these species because of its several similarities with H. martini, similarities which include a basically similar coloration of a type not otherwise found in species of the Lake Victoria Haplochromis flock. In the "michaeli" group, the relationship between H. martini and H. squamulatus seems to be closer and more direct than that between either species and H. michaeli (see also Greenwood, 1960, pp. 245-8 ; idem, 1962, p. 206, and p. 91 above). When discussing the relationships of the H. percoides-H. cavifrons group (Green- wood, 1962), I suggested that H. percoides could have evolved from an H. pellegrini- like stem, and that H. flavipinnis was a derivative of an H. percoides-like ancestor. Also in that paper I noted the less certain relationships of H. cavifrons, but considered it to be part of the H. percoides phyletic line. On revising this complex, I began to H4 P. H. GREENWOOD doubt my earlier conclusions about the affinities of H. cavifrons, which now seems to have greater relationship with the " serranus " group. This review provided no evidence to negative my conclusions about the interrelationships of H. percoides and H. flavipinnis, or the association of H. percoides with an H. pellegrini-\ike ancestor. However, I am not at all certain about the validity of my previous suggestion that the neurocrania of H. percoides and H. flavipinnis show affinity with those of H. bartoni and H. longirostris (i.e. with the " prognathus " group as it is now denned). With a better appreciation of neurocranial form in the piscivores as a whole, I now think that the skull of H. percoides is basically an " altigenis " group one, as is the skull of H. pellegrini. Similar re-evaluation leads me to think that the syncranial organization and form shown by H. cavifrons links that species more closely with the " serranus " group than with the " altigenis " group and its H. percoides-like off-shoots. The freckled coloration of H. cavifrons remains unique (Greenwood, 1962), and nothing ap- proaching it is seen in the " serranus " group. If H. cavifrons is a " serranus "-group derivative, then it stands in the same relationship to that group as does H. plagio- stoma, namely as an isolated off-shoot without any extant intermediates bridging the gap. Thus, the H. percoides-H. cavifrons " group ", although a valid one on grounds of general similarity between the constituent species, is probably polyphyletic in origin. Because so little material or information is available on three species described above (H. tridens, H. orthostoma, and H. parorthostoma) , they cannot as yet be in- cluded in a discussion on phylogeny. Haplochromis tridens does not seem to be re- lated to any of the piscivorous species groups ; its affinities probably lie with the complex of small, bottom-living species which form the bulk of trawl catches in sheltered, mud-bottomed areas of the lake. Haplochromis orthostoma and H. paro- thostoma are closely related to one another but cannot readily be associated with any other species. Leaving for the moment those species which as adults feed on post-larval fishes, consideration will be given to species which prey on larval and embryo fishes, the paedophagous species. Only one paedophage, H. barbarae, is described in this paper ; the others are dealt with in an earlier publication (Greenwood, 1959) . Haplochromis barbarae resembles small specimens of H. guiarti and adults of H. brownae, and does not show any close similarity in body form, jaw morphology or dentition with other paedophagous species ; unfortunately it has not been possible to study its osteology in detail. The probable phylogeny of the larval and embryo fish eating species was discussed in the paper cited above ; the conclusion reached was that the group had a polyphyletic origin. Little more can be added to these thoughts, except to reconsider the possible relationships existing between H. par- videns and H. acidens (previously misidentified as H. nigrescens; see above, p. 73). The morphology of the lower jaw in H. parvidens differs considerably from that of H. acidens (as does the diet, paedophage cf. herbivore) ; but, in other syncranial characters, and in body-form, the species are very similar. In these latter characters the two species are more similar than are H. microdon and H . cryptodon, the only A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 115 known paedophages showing a morphotype which could be ancestral to that of the " parvidens " level. Haplochromis cryptodon could have evolved from an H . brownae- like stem (possibly one like H. barbarae which had already adopted paedophagous habits?), the chief morphological changes involving the dentition (reduction), jaw form (to give greater distensibility) and a differential growth of the preorbital neuro- cranium (also leading to greater jaw motility). The changes in neurocranial form would lead to a skull essentially like that found in basal species of the " altigenis " and " prognathus " groups, in other words one like that in H. acidens. Thus, the origin of the " parvidens " structural grade is equivocal. Reconsidering the relationship of this grade (as represented by H. cryptodon, H. parvidens and H. microdon) with the other distensibly-mouthed grade of paedo- phages (represented by H. obesus and H. maxillaris), I can find, as before, few reasons to support a close phyletic linkage between them. Osteologically neither H. obesus nor H. maxillaris has a dentary like that occurring in the H. parvidens group, nor in the case of H. obesus is its form one from which a " parvidens " type might evolve. The neurocranium of H. obesus is most unlike that of the "parvidens " group, but that of H. maxillaris is virtually identical with the skull of H. cryptodon. The character which most clearly distinguishes these two species from any member of the " parvidens " group is the occurrence of teeth in which the crowns are curved labially (and not buccally as is usual in unicuspid teeth). This dental character is so marked, and restricted to these two species, that I am inclined to give it con- siderable weight when speculating on phylogenies, particularly since this tooth-form seems to have no adaptive significance. If the peculiar teeth in H. maxillaris and H . obesus do indicate a fundamental relationship between the species, then their syncranial differences would suggest an independent origin from a common stem, possibly a form like H. cronus (see Greenwood, 1959). Dentally, and also in its general level of organization, H. cronus is like those generalized Haplochromis species that attain a larger adult size than most members of that group ; it could be derived from an H. empodisma-like stem (see Greenwood, 1960). Natural History Only broad generalizations can be made about the natural history of the pisci- vorous predators. To date, seventy-nine species of Lake Victoria Haplochromis (and related monotypic genera) have been revised. Of these, forty-two species can be classed as piscivores, thirty-four species preying on free-swimming fishes, and eight species on cichlid embryos and larvae presumably taken from the mouths of parent fishes. Anticipating results still to be obtained from those species as yet unanalyzed, it seems probable that the number of non-piscivorous species will be increased sub- stantially. I suspect that ultimately the number of piscivorous species will be about forty per cent of the total. These figures apply to sub-adult and adult members of the species only since no data are available on the feeding habits of younger stages. The same restrictions apply to considerations on habitat preferences among the piscivores. Piscivorous species have been found in all the localities so far sampled, but much of the deeper water in Lake Victoria remains unexplored, especially at the levels n6 P. H. GREENWOOD occupied by bathypelagic species. In order to give a general picture of habitats in the lake, these can be divided, rather crudely, into three types : (i) sheltered areas such as bays and the smaller gulfs, (ii) Exposed areas, especially wave-washed beaches, (iii) Relatively undisturbed, open, off-shore waters (undisturbed that is, relative to the rather turbulent conditions prevailing over exposed beaches), not enclosed in gulfs or bays. Such a subdivision is, in many respects, unsatisfactory because conditions like those in habitat (iii) do occur in the larger gulfs and bays. More species are found in sheltered areas (habitat [i]) than in the other two habitats ; 23 species have been recorded frequently in habitat (i), 14 from habitat (ii), and 15 from habitat (iii), with, in the habitats respectively, 2, 3, and i species occurring infrequently. However, these various species are by no means confined to a particular habitat. For example, ii species were found in both sheltered and exposed areas, 9 in sheltered and off-shore habitats (3 of these species also occurring over beaches) ; only 6 species are apparently confined to sheltered areas. Two species occur both over beaches and in the quieter off-shore areas, and only 4 are apparently confined to the latter habitat (but 3 of these species are known only from deep water) . The nature of the substrate seems to exert a less restricting influence on the piscivores than on members of other trophic groups. Twenty-four species are recorded as occurring over hard substrata (sand, shingle or rock), and 20 over a soft substrate (organic mud) ; 10 of these species are found over both kinds of substrate, 6 are thought to be confined to a soft bottom, and 13 to a hard substrate. The eight strictly paedophagous species (see Greenwood, 1959, and p. 114 above) are not included in the analysis above. These species appear to be rather more re- stricted in their distribution, particularly from the viewpoint of depth range. It seems that the paedophages are confined to the littoral and immediately sublittoral zone, and are probably restricted to sheltered bays and exposed beaches. The depth range of the other 34 piscivores is, in general, confined to water less than 60 ft. deep (with of course, the exception of H. boops, H. pachycephalus , H. thuragnathus and H. dichrourus which have been caught in water about 120 ft. deep). Most species have a wide range within these depth limits, and few if any are restricted to purely littoral areas. However, it must be emphasized that this picture may be unduly biased by sampling limitations. The horizontal distribution of the species has not been studied critically ; in water less than 20 feet deep, the spatial distribu- tion of piscivores caught in nets set to cover about the first five feet of water below the surface, and the five feet above the bottom, does not show any obvious horizontal stratification of the particular species. But, these observations were not tested statistically. From the relatively few gut analyses available it would seem that the majority of piscivores prey on other Haplochromis species, and to a much lesser extent on small cyprinid fishes (especially Engraulicypris argenteus). Some species have a mixed insect-fish diet, and there are several records of otherwise exclusively piscivorous species eating insects when these are periodically and suddenly super-abundant, as for instance after a heavy termite hatch. The paedophagous species appear to feed mostly on cichlid embryos and larvae (it is presumed those of Haplochromis species) , but insects are also recorded from the gut contents of these species. A REVISION OF THE LAKE VICTORIA HA PLOCHROMIS SPECIES 117 Information on the breeding habits of piscivores is almost non-existent ; in those cases where some data are available, the species are known to be female mouth brooders. The turbidity of the water in most parts of Lake Victoria has precluded field observations on the actual spawning sites of both predators and prey alike. There has not yet been enough collecting on a lake-wide basis to establish whether or not any of the fish-eating species has a geographically restricted distribution. For eight species this possibility can definitely be overruled, (and in ten others it seems very unlikely) but for sixteen species there are suggestions of geographical restriction because they have not been caught in some regions where, on the basis of habitat and niche suitability, they should be present. SUMMARY (1) Ten species are redescribed on the basis of new material. (2) Nine new species (H. pachycephalus, H. boops, H. thuragnathus, H. pseudopelle- grini, H. paraguiarti, H. acidens, H. barbarae, H. parorthostoma and H. apogo- noides) are described. (3) Although all these species have a general facies and dentition usually associated with piscivorous habits, some do not belong to this trophic group ; one species (H, acidens] is apparently herbivorous. (4) The possible phyletic interrelationships of the piscivorous species are discussed. Three major morphological groups can be detected, at least on the basis of their more extreme members, but the boundaries are ill-defined. Some minor groups are also considered, as are the larval and embryo fish-eating species groups. (5) Broad summaries of the natural history of piscivorous species are given. (6) The holotype of Astatotilapia nigrescens Pellegrin is redescribed and its possible synonymy discussed. ACKNOWLEDGEMENTS I am deeply indebted to many people for their assistance in preparing this paper. To my colleagues Mr. A. C. Wheeler and Dr. E. Trewavas are due my thanks for, respectively, providing numerous and excellent radiographs, and for many profitable discussions. Through the cooperation of Dr. M. Blanc of the Paris Museum, I have been able to examine Pellegrin's type specimens and thus to settle several problems. Dr. Paul Kahsbauer of the Vienna Museum has graciously lent me Lohberger's type material ; I am deeply indebted to him for this privilege. In east Africa, Dr. M. Gee and Mr. R. Welcomme of E.A.F.F.R.O. aided my studies by proving additional material and field observations ; material collected by the Uganda Fisheries Department has provided many specimens from localities which I was unable to sample personally. The cooperation of E.A.F.F.R.O. and the Uganda Fisheries Department is warmly appreciated. n8 P. H. GREENWOOD APPENDIX The disputed identity of Astatotilapia nigrescens Pellegrin, 1909, (Bull. Soc. Zool. France, 34, 157) was mentioned on page 73. Boulenger (1915) synonymized this species with Haplochromis percoides Blgr. 1906. Regan (1922), however, resurrected the species as Haplochromis nigrescens, and included in his redescription a number of specimens which I have placed in a new species, H. acidens. Pellegrin's figure of A. nigrescens is misleading and does not convey an accurate impression of the holotype and unique specimen (Paris Museum, number 09-508). Recently, I examined this specimen, an immature male 71 mm. standard length, collected by Alluaud from the Kavirondo Gulf, Kenya. It does not agree closely with any other specimens I have handled, but is does show affinity with both H. percoides Blgr., 1906 and H . flavipinnis (Big.), 1906, especially the latter. Before considering its identity further, a redescription of the holotype will be given. mm. Proportional percentage Standard length . . . . .71-0 Depth of body ..... 24-0 33-8 standard length Length of head ..... 26-0 36-6 standard length Depth of preorbital . . . . 4-3 16-5 head length Width of interorbital . . . . 6-0 23-1 head length Length of snout ..... 8-0 30-8 head length Diameter of eye . . . . . 7-0 27-0 head length Depth of cheek ..... 6-5 25-0 head length Length of lower jaw . . . .12-0 46-2 head length Length of caudal peduncle . . . 13-0 18-3 standard length Length of pectoral fin . . . . 18-0 25-3 standard length Caudal peduncle 1-4 times as long as deep. Lower jaw slightly oblique and very slightly projecting ; twice as long as broad. Posterior tip of the maxilla almost reaching the vertical through the anterior orbital margin. Gill rakers : moderately stout, 9 on the lower part of the first gill arch. Scales : ctenoid ; lateral line with 31 scales, cheek with 4 rows (the rows short, so that the anterior part of the cheek is naked). Seven scales between the upper lateral line and the dorsal fin origin ; 7 between the pectoral and pelvic fin bases. Chest, belly and nuchal scales small. Teeth : in the outer row of both jaws relatively slender, slightly curved ; about 50 in the upper jaw. Inner teeth tricuspid or weakly tricuspid, arranged in 2 and I rows in the upper and lower jaw respectively. Coloration. The preserved colour pattern (in shades of brown) closely resembles that of H. percoides and H. flavipinnis (see Greenwood, 1962). The dorsal fin is marbled, and the caudal both marbled and maculate ; the pelvics are dark. DISCUSSION : In its general facies, and particularly its head shape, the holotype of A. nigrescens resembles both H. percoides and H '. flavipinnis , especially the former because the angle of the mouth is less oblique than in most specimens of H. flavi- pinnis. However, when morphometric characters are considered a number of A REVISION OF THE LAKE VICTORIA H A.PLOCHRO MI S SPECIES 119 differences between H. percoides and A.nigrescens holotype are apparent. In six characters (body depth, preorbital depth, interorbital width, snout length, eye dia- meter, and caudal peduncle length) the values fall outside the known range for H. percoides, and the lower jaw is relatively longer than in specimens of H. percoides of a comparable size. There is greater correspondence between these characters in A. nigrescens and H . flavipinnis, since only two (the shallower preorbital and larger eye) fall outside the range for H . flavipinnis, but there is a greater difference in head shape. Thus, A . nigrescens could be an aberrant specimen of either H. flavipinnis (differing especially in head shape) or H. percoides (numerous morphometric differences) . For the time being, however, I do not think that A. nigrescens can be formally synonymized with either species. I would prefer to recognize it as the purely nominal species Haplochromis nigrescens (Pellegrin) until more is known about the range of variation in small specimens of H. flavipinnis, the species I think it most closely resembles. REFERENCES GILCHRIST, J. D. F., & THOMPSON, W. W. 1917. The freshwater fishes of South Africa. Ann. S. Afr. Mus., 11 : pt. 6, 465-575. GRAHAM, M. 1929. A Report on the Fishing Survey of Lake Victoria, 1927-1928, and Appendices. Crown Agents, London. GREENWOOD, P.H. 1956. The monotypic genera of cichlid fishes in Lake Victoria. Bull. BY. Mus. nat. Hist., Zool. 3 : No. 7, 295-333. - 1957. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae) Part II. Bull. Br. Mus. nat. Hist., Zool. 5 : No 4, 76-97. - 1959. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae) Part III. Bull. Br. Mus. nat. Hist., Zool. 5 : No. 7, 179-218. 1960. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae) Part IV. Bull. Br. Mus. nat. Hist., Zool. 6 : No. 4, 227-81. - 1962. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae) Part V. Bull. Br. Mus. nat. Hist., Zool. 9 : No. 4, 139-214. 1965. The cichlid fishes of Lake Nabugabo, Uganda. Bull. Br. Mus. nat. Hist., Zool. 12 : No. 9, 313-57. - 1966. Two new species of Haplochromis (Pisces, Cichlidae) from Lake Victoria. Ann. Mag. nat. Hist., (13), 8 : 303-318. TREWAVAS, E. 1964. A revision of the genus Serranochromis Regan (Pisces, Cichlidae). Ann. Mus. Roy. Afr. Cent. Tervuren, ser in 8°, Zool. No. 125, 1-58. PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING / 27 FEB1967 THE VACHELL COLLECTION OF CHINESE FISHES IN CAMBRIDGE P. J. P. WHITEHEAD & K. A. JOYSEY BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 15 No. 3 LONDON: 1967 27 FEBH THE VACHELI^ COLLECTION OF CHINESE \^ FISHES IN CAMBRIDGE BY P. J. P. WHITEHEAD (British Museum, Natural History) "\ and K. A. JOYSEY ( University Museum of Zoology, Cambridge) Pp. 121-165 ; 3 Plates BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 1 5 No. 3 LONDON: 1967 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the 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. 15, No. 3 of the Zoological series. The abbreviated titles of the periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation : Bull. Br. Mus. nat. Hist. (Zool.) Trustees of the British Museum (Natural History) 1967 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 24 February, 1967 Price £i is. THE VACHELL COLLECTION OF CHINESE FISHES IN CAMBRIDGE By P. J. P. WHITEHEAD & K. A. JOYSEY CONTENTS Page 1. INTRODUCTION .......... 123 2. RICHARDSON'S " REPORT "........ 123 3. THE VACHELL COLLECTION ........ 124 4. SYSTEMATIC LIST ......... 127 5. EXTANT VACHELL SPECIMENS ....... 147 6. REFERENCES .......... 158 7. INDEX ........... 160 ABSTRACT The Rev. G. Vachell of Macao presented about a hundred Chinese fishes to the Cambridge Philosophical Society in the early part of the last century. In his Report on the Ichthyology of the Seas of China and Japan, Richardson based 22 new species on Vachell material. Although 80 species are listed here, the greater part of the Vachell collection is no longer extant and only 15 specimens survive ; these include the types of Anguilla clathrata Rich., Pelor tigrinum Rich., and Seserinus vachellii Rich. i. INTRODUCTION SOME i8th and igth century fish collections are well-known and their contents well documented. Others have lapsed into obscurity and provide considerable difficulties when the need arises to establish types or validate old names. One of these is the Vachell collection of Chinese fishes, the remains of which are housed in the University Museum of Zoology in Cambridge. The Vachell collection, which appears to have contained about 80 species of fish from Macao, formed a significant part of the material on which Sir John Richardson based his " Report on the Ichthyology of the Seas of China and Japan " published in 1846. As often happens with old collections, the Vachell material included a large proportion of types. Thus 22 of the new species described by Richardson in the " Report " were based on Vachell fishes. Unfortunately, only a fraction of the Vachell collection is now extant. But this collection is of sufficient importance for a complete list to be given, particularly in order to record those specimens which survive (including the types of 3 Richardson species) those which are missing (including 7 types), and those specimens which are now known with certainty to have been destroyed (including 12 types). The principal author, P.J.P.W., is responsible for all the systematic zoology, and K.A.J. undertook the investigation of the records in Cambridge. 2. RICHARDSON'S "REPORT" Richardson (1846) listed 665 species of fishes from the seas of China and Japan, an enormous increase on any previous list ; 142 of these were described as new species or varieties. Over three hundred of the species listed were represented by an original ZOOL. 15, 3 8 124 P. J- P- WHITEHEAD & K. A. JOYSEY coloured drawing from a collection of illustrations of fishes from Macao and Canton compiled by John Reeves (1774-1856), an Inspector of Tea for the East India Company at Canton ; 83 new species introduced by Richardson were based solely on a Reeves illustration. Three sets of these illustrations are now in the Zoology Library of the British Museum (Natural History). Since half (40 out of 80) of the species listed here (including n of the types) are illustrated by a Reeves drawing, the Reeves illustration number is cited in the synony- mies. This may assist in identifying some of the missing Vachell specimens. The illustrations of the types are reproduced here, apparently for the first time (except Chatoessus maculatus — see Whitehead, 1966). John Reeves, and his son J. R. Reeves, also sent a collection of fishes from Macao and Canton to the British Museum. These were examined by Richardson and Giinther, and since they came from the same area as the Vachell fishes, reference is made to them in the text. Richardson evidently examined the Vachell collection in the years up to 1845. Four of the species listed here (Batistes vachellii, Anguilla ctathrata, Aploactis breviceps, and Congrus lepturus) had already been described as new by Richardson in the " Zoology of the Voyage of H.M.S. Sulphur " published in three parts between 1844 and 1845 (see " Report " p. 316 for dating). In general, however, Richardson left description of the Vachell material until the " Report ". 3. THE VACHELL COLLECTION According to Richardson's " Report " (1846, p. 189), the Rev. George Vachell was Chaplain to the India Company at Macao in about 1830. He made a collection of about a hundred fishes from that region, and these were presented to the Philosophical Institution in Cambridge, preserved in spirits, and mostly in good condition. None- theless, Richardson himself described at least one fish as " not in very good condition " (p. 204), another that " the colours have suffered from long maceration in spirits " (p. 208), and another that " it is flaccid and may have lost its exact shape " (p. 267). In 1865 the collections of the Cambridge Philosophical Society were transferred to the University of Cambridge and housed in the newly built Museum of Comparative Anatomy and Zoology. Indeed, the Philosophical Society Collection formed the nucleus of the Museum of Zoology, and although under the same roof as the Museum of Comparative Anatomy, the two collections were kept in separate rooms and remained distinct for several years. In 1867 J. W. Clark, who was Superintendant of the Museums, reported that, " Two presses have been provided to contain the collections of Fishes presented by the Philosophical Society. These consist of the following : the series of Fishes of Madeira, collected by Mr. Lowe, many of which are unique, and nearly all the types of his descriptions in the transactions of the Cam- bridge Philosophical Society : a considerable number of specimens procured by Mr. Darwin during the voyage of the ' Beagle ', also the types of the descriptions published by the late Sir John Richardson : a collection sent from China by Mr. Vachell ; and a collection of Fishes of Great Britain formed by the late Professor Henslow and Mr. L. Jenyns. These will at no distant period be named and catalogued by Dr. Giinther, of the British Museum, who has most generously offered to undertake VACHELL COLLECTION OF CHINESE FISHES 125 this work ". (It should be noted that the punctuation of this passage suggests that Clark did not realize that the Richardson types were included within the Vachell collection.) The following year Clark (1868) reported that during 1866-67 " Dr. Gunther, of the British Museum, one of the first ichthyologists in Europe, has been so kind as to examine the collections of Fish in spirits, and to determine those that were unnamed." The major part of Gunther 's " Catalogue " had by then been published, but Gunther makes only rare reference to specimens in Cambridge in the subsequent volumes (7 and 8), and no mention at all of having seen Vachell fishes. It is probably for this reason that the Vachell collection has been generally ignored. A few years later, Clark (1871) again referred to the identification of the spirit collection of Fish, Amphibia and Reptiles and reported that, " I had hoped to have announced the completion of an arrangement by which the services of Dr. A. Gunther, of the British Museum, the best living authority on the subject, might have been secured for this work ; but though the proposal made by the Museums' Syndicate to the Trustees of the British Museum was a most liberal one, that body did not think proper to accede to it. It will therefore be necessary to select some other competent person ". Apparently this statement had the desired effect and only a year later Clark (1872) reported that these collections had been examined and determined by Dr. Gunther! It is clear that in both 1866-67 and 1870-71, Gunther worked on the fish collections in Cambridge. Shipley (1913) even claims (p. 265) that the fishes were catalogued by Dr. Gunther, but we have been unable to trace any other record of such a catalogue, either in London or in Cambridge. In 1893, S. F. Manner catalogued the entire fish collection at Cambridge, indexing the species according to the volume and page numbers of Giinther's " Catalogue". The names used by Harmer are generally identical to those of Gunther, but in the case of the Vachell specimens the names often differ from those given in Richardson's " Report ". This suggests that Gunther re-identified and relabelled many of the Vachell specimens, despite the fact that he did not refer to them in the " Catalogue ". Altogether, 80 species (plus 3 names here considered synonyms) are listed as appearing either in Richardson's " Report " or in Harmer's catalogue, and often in both. The vast majority are now either missing or known to have been destroyed. A total of 32 species which were listed by Richardson as including Vachell speci- mens are not so listed by Harmer (including the types of 10 Richardson species). These specimens may have been lost, exchanged or destroyed without record either in the Cambridge Philosophical Institute between 1845 and 1865, or in the Museum of Comparative Anatomy and Zoology between 1865 and 1893. It is possible that Richardson worked on some of the Vachell material at the British Museum, that it was never returned to the Philosophical Institution, and that it became incorporated into the British Museum collections without ever being registered. There are in fact a number of unregistered bottles, including some labelled as presented (or collected) by Richardson. In some cases these involve Vachell species which were never included in Harmer's Catalogue of Cambridge material, and each of these have been carefully checked. Unfortunately in no case is there supplementary evidence which would prove the case either way. 126 P. J. P. WHITEHEAD & K. A. JOYSEY It is also known that J. W. Clark spent a good deal of the summer of 1866 over- hauling the Physiological Series, which had just been moved into the newly built Museum in Cambridge, and it is recorded that useless and decayed specimens were thrown away (Clark, 1867 ; Shipley, 1913). Although this activity does not refer explicitly to the fish collection it does give a clear indication of the policy that was being implemented in the Museum during the same period that Gunther was working on the fish, and this might account for some of the 32 species which were missing before Harmer's 1893 catalogue. The possibility that some of the Vachell material was acquired by the British Museum between 1865 and 1893 has also been checked. Such an acquisition seems most likely to have occurred during or after Giinther's visit in 1866-67, or as a Part of the arrangement made in 1870-71, the terms of which are not known. But the British Museum registers from 1866-93 show no gifts, purchases or exchanges in- volving the Cambridge Museum. It is certain that no Vachell material went to London officially, and in some doubtful cases the British Museum material has been checked and no evidence found that any arrived unofficially. Conversely, there are 16 species listed by Harmer as being Vachell material which are not so mentioned by Richardson. Either Richardson missed these, or perhaps they were not true Vachell specimens but were erroneously included by Harmer. Among these, one Cambridge specimen of Boleophthalmus campylostomus (see p. 147) was transferred to the British Museum in 1917, but although Harmer lists it as a Vachell fish, Richardson (Report, p. 209) states that he had seen no specimens. Harmer's 1893 catalogue is in two parts, one listing the stored material and the other listing those specimens which were on exhibition at that time, the latter being given separate registration numbers. A large number of Harmer's entries in the catalogue of stored material have since been crossed out, and these specimens are no longer extant. A note in the front of the catalogue states : " The collection of fishes in store was overhauled during May, 1939, when specimens thought to be of little or no value were discarded." We understand from the present Director, Dr. F. R. Parrington, that this clearance included many dried-up, rotten and dis- integrated specimens, and he recalls that his predecessor, Sir Clive Forster-Cooper, once recounted that some of the fishes which had matured beyond repair were found to have been pickled in rum! It seems certain that those specimens which are crossed off Harmer's catalogue were destroyed, in contrast to those which are just missing insofar as they do not appear in Harmer's list. It is right and proper to ask why so little of the Vachell collection survives today, despite the fact that it was presented to an Institution which could well have been expected to provide security. Only by such enquiry can it be hoped to avoid similar disappointments in the future. In this case, there seems to have been a combination of circumstances. Richardson provides some evidence of poor fixation and preserva- tion, which continued to take its toll more than a century later. Clark apparently did not appreciate that the Vachell collection included the Richardson types, and for some inexplicable reason Giinther drew no attention to them. Hence, in the majority of cases, Harmer was unaware of the type specimens, and did not indicate them as such when he prepared his catalogue. In consequence, when the collection VACHELL COLLECTION OF CHINESE FISHES 127 was overhauled nearly 50 years later, many types were discarded. Until relatively recently it has been accepted practice in most museums that material which had suffered beyond repair was destroyed and crossed off the cata- logue, unless, of course, it was recorded as type material. In retrospect this policy is to be regretted in the present case, and this has caused us to give some thought to the problem. For one reason or another, whether it be faulty fixation, poor storage conditions, sheer neglect or genuine accident, nearly all Museums possess some important material that has suffered, apparently beyond repair. We now hold the view that those responsible for such collections must accept this as almost inevitable, and without shame should retain such material, rather than totally destroying it. Advances in technology are now providing new methods of obtaining information from such material, and although apparently useless to one generation, it may be capable of yielding information to the next. Methods now exist for reconstituting dried up specimens, and for " developing " labels which have become illegible in ordinary light. In the present instance, X-ray photography of even the rotten material might have yielded sufficient information about the skeleton to establish the identity of the species, but unfortunately none of these techniques can now be applied, because most of the material has been destroyed. 4. SYSTEMATIC LIST In the following list of Vachell species, the synonyms of Richardson and Giinthei have been given, headed by the modern name for the species. The latter have posed considerable difficulties, particularly when no specimens survive and where the identification must be made on a brief description by Richardson, sometimes supplemented by a Reeves illustration. Richardson's specimens were evidently not always consistent with the Reeves drawing (e.g. in the case of Sebastodes vachellii — p. 140), a fact not always appreciated by later authors. In many cases, therefore, it has been possible to do no more than to follow the comprehensive synonymies of Chu (1931), Herre (1953), Chu, Tchang & Chen (1964) and Fowler (Synopsis of fishes of China, from 1930 onwards) ; only Chu and Fowler have listed all or almost all Richardson names. Sometimes, a recent study of a particular group has helped to identify the Vachell material, but even then the Vachell specimens themselves do not appear to have been examined, perhaps since the time of Giinther's visits to Cambridge nearly a century ago. In addition, very few authors seem to have examined the Reeves illustrations. We have listed here fifteen specimens representing eleven species which appear to be part of the original Vachell collection. Even these numbers are by no means certain since several are listed only by Harmer and not by Richardson as being Vachell material. Notes on these extant specimens and their identifications are given separately in Section 5 (p. 147). Those of Richardson's species which were based wholly or partly on Vachell material are marked with an asterisk and are listed separately in Table I. We are particularly grateful to Mr. W. L. Chan, of the Fisheries Research Station in Hong Kong, for his help in identifying the fishes shown in the Reeves illustrations reproduced here and his comments on the Chinese names applied to these fishes. 128 P. J. P. WHITEHEAD & K. A. JOYSEY Class CHONDRICHTHYES Family ORECTOLOBIDAE i. Chiloscyllium plagiosum (Bennett) Chiloscyllium plagiosum: Richardson, 1846: 194 (Vachell material) (Reeves illustr. No. 252). Chiloscyllium indicum: Giinther, 1870, 8 : 413 (material listed under six varieties). As Chiloscyllium indicum " (several) China Rev. G. Vachell " in Harmer's list — destroyed, 1912 according to catalogue. A half-grown Richardson specimen and a Reeves juvenile in British Museum. Giinther does not mention any specimens in Cambridge, although he had inspected the Vachell material by this time. The identification of the three sharks listed here is based on the review of Taiwan species by Chen (1963). Family CARCHARHINIDAE 2. Carcharhinus melanopterus Quoy & Gaim. Carcharias (Prionodon) melanopterus: Richardson 1846: 194 (not stated as Vachell specimen) (Reeves illustr. No. 23). Carcharias melanopterus: Giinther, 1870, 8 : 369 (South Africa and Amboyna material only). As Carcharias (Prionodon) melanopterus, juvenile " China, Rev. G. Vachell " in Harmer's list — destroyed. No Cambridge material mentioned by Giinther. Family TRIAKIDAE 3. Triakis scyllia Miiller & Henle Triakis scyllium: Richardson, 1846: 195 (no specimens). Triads scyllium: Giinther, 1870, 8 : 384 (no specimens). As Triads ? scyllium, juvenile, " China Rev. G. Vachell " in Harmer's list — destroyed. Vachell specimen presumably overlooked by Richardson, if indeed it did belong to the Vachell collection. Again, Giinther makes no mention of Cambridge material. Class OSTEICHTHYES Family CLUPEIDAE 4. Clupanodon thrissa (Linnaeus) *Chatoessus maculatus Richardson, 1846 : 308 (a single Vachell specimen, TYPE) (Reeves illustr. No. 109). Chatoessus maculatus: Giinther, 1868, 7 : 409 (Formosa specimens only). Harmer lists " Clupea thrissa. (several). No histy.", but these were subsequently destroyed. Since Chatoessus maculatus was the only clupeid represented in the Vachell collection, it is possible that one of the Harmer specimens was the type. Giinther mentions only three Formosan specimens but not the Vachell material in Cambridge. For identification of C. maculatus, see Whitehead (1966). Reeves illustration reproduced here (Plate I, fig. i). VACHELL COLLECTION OF CHINESE FISHES 129 Family ENGRAULIDAE 5. Coilia playfairii (McClelland) Coilia playfairii: Richardson, 1846 : 309 (see note below). Coilia clupeoides: Giinther, 1868, 7 : 404 (" Chinese Seas ", no mention of Cambridge material). Listed as Coilia (mystus), with the name clupeoides written above, " ' Fishes from China ' Canton ", not in Harmer's hand — EXTANT. A single specimen now exists, but with only a modern label indicating that it is a Canton fish. However, Richard- son states " specimens exist in all the collections of Chinese fishes that we have seen ". Giinther does not mention having seen any Cambridge material. Measurements and notes on the extant specimen are given in Section 5 (p. 149), and further notes on Richardson material are given by Whitehead (1966) . Family SALANGIDAE 6. Salanx chinensis (Osbeck) Leucosoma chinensis: Richardson, 1846 : 303 (Reeves and Vachell specimens). Salanx chinensis: Giinther, 1866, 6 : 205 (China, including type of Leucosoma reevesii Gray). Listed by Harmer as " Salanx chinensis (2) ? History "—destroyed. The British Museum has a specimen from the Haslar collection, as well as the type of Leucosoma reevesii. Identification based on Chu (1931). Family ANGUILLIDAE 7. Anguilla japonica (Temminck & Schlegel) *Anguilla clathrata Richardson, 1844 : 104 (one Vachell specimen, TYPE) ; Idem, 1846 : 312. Anguilla clathrata: Giinther, 1870, 8 : 23 (doubtful species No. 4, name only). A single specimen listed by Harmer as A. vulgaris " China. Rev. G. Vachell. "- EXTANT. This specimen was subsequently transferred as F.2002 to the Exhibited series. Giinther had already examined the Vachell collection but did not acknow- ledge having seen this fish. Richardson's description (Zoology of the Sulphur, p. 104) was based on a Vachell fish of 8-8 inches from Canton. See Section 5 for description and notes on this HOLOTYPE. Family MURAENESOCIDAE 8. Muraenesox cinereus (Forsskal) Congrus tricuspidatus : Richardson, 1846 : 312 (Vachell specimens). Muraenesox cinereus: Giinther, 1870, 8 : 46 (Reeves specimens and type of C. tricuspidatus}. Listed as M, cinereus " China Rev. G. Vachell " by Harmer — destroyed. Cam- bridge material not mentioned by Giinther. Identification based on Chu, Tchang & Chen (1963). 13° P. J. P. WHITEHEAD & K. A. JOYSEY Family CONGRIDAE 9. Uroconger lepturus (Richardson) Congrus lepturus Richardson, 1844 : 106, and 1846 : 132 (Canton, but no reference to Vachell specimens) . Uroconger lepturus: Giinther, 1870, 8 : 44 (Reeves specimens, the TYPE, and another Chinese specimen) . Listed as Uroconger lepturus, 2 specimens " China Rev. G. Vachell " by Harmer — destroyed. Giinther does not refer to Cambridge material. The species was des- cribed by Richardson (Zoology of the Sulphur, p. 106, PL 56, figs. 1-6) with a single set of measurements relating to a specimen of 9 inches from Canton, but no indication of the collector (presumably Reeves). There is a specimen of 160 mm. S.L. (io£ inches) labelled as type in the British Museum (unregistered but with a metal tag " 821 ") ; the second Chinese specimen is even larger (322 mm.). The first is labelled "Reeves " and the second was purchased from a Mr. Warwick. Present identifica- tion based on Chu, Tchang & Chen (1963). Family OPHICHTHIDAE 10. Pisodonophis boro (Ham. Buch.) Ophisurus harancha: Richardson, 1846 : 313 (one fish of 14^ inches in Camb. Phil. Inst., a Reeves specimen and an Indian specimen). Ophichthys pattens: Giinther, 1870, 8 : 61 (the Reeves type only). Harmer listed 2 specimens of Ophichthys cancrivorus " China Rev. G. Vachell " — destroyed. Although Richardson (1846) believed the Vachell, Reeves, and Indian specimens to be conspecific, he elsewhere (Erebus & Terror, p. 10) proposed the name pattens for the Reeves fish, while still claiming the Vachell specimen to be true harancha. Giinther (1868, p. 61) considered Richardson's harancha to be pattens, as also did Chu (1931). Fowler (19320, p. 126) agreed, but placed both under an earlier Hamilton-Buchanan name, Pisodonophis boro, and this course has been adopted here. Family BAGRIDAE IT. Pseudobagrus vachellii (Richardson) *Bagrus vachellii Richardson, 1846 : 284 (one Vachell specimen of 5 inches, the TYPE). Pseudobagrus vachellii: Giinther, 1864, 5 : 85 (Chinese specimens). Listed as P. aurantiacus " China. Rev. G. Vachell " in Harmer's catalogue — destroyed. Giinther had kept the closely related P. vachellii and P. aurantiacus separate, but he may have reconsidered this when he came to identify the Cambridge material. Since Harmer listed two Vachell Pseudobagrus species, and Richardson two Vachell Bagrus species, it must be assumed that the two species correspond. Present identifications follow Chu (1931). VACHELL COLLECTION OF CHINESE FISHES 131 12. Pelteobagrus fulvidraco (Richardson) *Bagrus limbatus Richardson, 1846 : 283 (one Vachell specimen, the TYPE). Pseudobagrus fulvi-draco: Giinther, 1864, 5 : 85 (one Chinese fish presented by Giinther himself). Listed as Pseudobagrus fulvi-draco " China. Rev. G. Vachell " by Harmer — destroyed. Fowler (19326) identified Richardson's species as Plotosus anguillaris (Bloch), but curiously did not list Richardson's B. vachellii. Family PLOTOSIDAE 13. Plotosus anguillaris (Bloch) Plotosus lineatus: Richardson, 1846 : 286 (specimens in the Camb. Phil. Inst.) (Reeves illustr. No. /Sn). Plotosus anguillaris: Giinther, 1864, 5 : 24 (Chinese specimens). Listed as Plotosus anguillaris " (several) Phistory " in Harmer's Catalogue — des- troyed. Possibly the Cambridge Philosophical Institution had other Chinese material in addition to the Vachell specimens. Present identification based on Chu (1931) and Fowler (19326). Family BELONIDAE 14. Strongylura strongylura (van Hasselt) Belone caudimaculata: Richardson, 1846 : 264 (no Vachell material) (Reeves illustr. No. ^33). Belone caudimaculata: Giinther, 1866, 6 : 245 (no China specimens). Listed as B. strongylurus, " ? China " by Harmer — destroyed. Richardson gave no description but mentioned specimens from Canton (Reeves) and from Port Essington. Giinther (1866) placed the former in B. strongylurus and the latter in B. caudimaculata. Present identification based on Fowler (19326). Family SOLEIDAE 15. Microbuglossus ovatus (Richardson) *Solea ovata Richardson, 1846 : 279 (a single Vachell specimen, 3^ inches, the TYPE) (no Reeves illustr.). Solea ovata: Giinther, 1862, 4 : 472 (Chinese specimens). Listed by Harmer under Solea ovata, 4 specimens " China Rev. G. Vachell " destroyed. Giinther (1862) lists an Amoy and a Haslar Collection specimen, and 4 fishes presented by Belcher. The latter are labelled as types, but they are too small and Richardson mentions only Vachell material. The single Haslar fish, however, may well have been 3^ inches in length (caudal now damaged) but there are 59 dorsal rays (65 described). Identification based on Chu, Tchang & Chen (1963). 132 P. J. P. WHITEHEAD & K. A. JOYSEY 16. Zebrias zebra (Bloch) *Solea ommatura Richardson, 1846 : 279 (two Vachell specimens, the TYPES) (Reeves illustr. No. £13). Synaptura zebra: Giinther, 1862, 4 : 484 (Chinese specimens). Listed by Harmer, as Synaptura zebra, four fishes, " No histy." — destroyed. No British Museum material which could be the lost Vachell types. Reeves illustration reproduced here (Plate I, fig. 4). Identification follows Chu, Tchang & Chen (1963). Family CYNOGLOSSIDAE 17. Cynoglossus grammicus (Richardson) *Plagiusa grammica Richardson, 1846 : 280 (two Vachell specimens 3! inches, the TYPES) (no Reeves illustr.). Plagiusa grammica: Giinther, 1862, 4 : 492 (doubtful species No. 5, name and reference to the " typical specimen " in Camb. Phil. Soc. collection). Listed as Cynoglossus trigrammus " (several). China. Rev. G. Vachell " by Harmer (presumably having been re-identified by Giinther) — destroyed. No evidence of Vachell material in British Museum. Richardson's species is ignored by Chu, Tchang & Chen (1963), and the present identification follows Fowler (1934). Family HOLOCENTRIDAE 18. Holocentrus ruber (Forsskal) Holocentrum albo-rubrum: Richardson, 1846 : 223 (Vachell specimens) (Reeves illustr. 019.) Holocentrum rubrum: Giinther, 1859, 1 : 35 (Reeves and other Chinese material). Two specimens are given in Harmer's list as Holocentrum rubrum, " China. Rev. G. Vachell." — EXTANT. There are also two dry specimens listed by Harmer but stated to have " no history ". These are also extant and were presumably identified by Giinther. See Section 5 (p. 151) for notes on the two extant spirit specimens. Family CHANNIDAE 19. Channa maculata (Lacepede) Ophicephalus maculatus: Richardson, 1846 : 251 (two Vachell specimens) (Reeves illustr. Nos. 148 and /3ig). Ophiocephalus maculatus: Giinther, 1861, 3 : 480 (Reeves and China specimens). Listed as 0. argus in Harmer's list, 2 + I specimens, " China. Rev. G. Vachell " destroyed. Giinther considered 0. maculatus to be close to 0. argus, but seems to have re-identified the Cambridge material as the latter. Present identification based on Chu (1931). VACHELL COLLECTION OF CHINESE FISHES 133 Family ATHERINIDAE 20. Atherina bleekeri (Gunther) Atherina bleekeri Giinther, 1861, 3 : 398 (Reeves specimens). Listed as A. bleekeri "China. Rev. G. Vachell " by Harmer — destroyed. No species of Atherina are listed by Richardson, so Manner's specimens were either overlooked or were not in fact Vachell fishes. Family MUGILIDAE 21. Mugil cephalus (Linnaeus) Mugil japonicus: Richardson, 1846 : 247 (no Vachell material). Mugil cephalus: Giinther, 1861, 3 : 419 (Reeves and Chinese specimens). Listed as M. cephalotus by Harmer, 2 specimens " China. Rev. G. Vachell "• destroyed. Richardson included seven species of Mugil in the " Report ", none based on Vachell specimens. Richardson's M. macrolepidotus is another synonym of M. cephalus, according to Fowler (1935), who has been followed here. Family SPHYRAENIDAE 22. Sphyraena obtusata (Cuvier) Sphyraena chinensis: Richardson, 1846 : 266 (one Vachell specimen) (Reeves illustr. No. 62). Sphyraena chinensis: Giinther, 1860, 2 : 334 (doubtful species No. i). Not listed by Harmer. No British Museum specimens labelled S. chinensis. Present identification follows Chu (1931). Family TRICHIURIDAE 23. Leptur -acanthus savala (Cuvier) Trichiurus intermedius: Richardson, 1846 : 268 (one British Museum specimen, but no Vachell material) (Reeves illustr. No. ^56) . Trichiurus savala: Giinther, 1860, 2 : 347 (East Indian and Chinese specimens). One specimen listed by Harmer, as T. muticus " China. Rev. G. Vachell ", now transferred to Exhibited Series No. F.2685 — EXTANT. Richardson placed T. muticus Gray in his synonymy of " T. lepturus, japonicus " (" Report ", p. 268), a record which he based solely on T. lepturus Temm. & Schl. from Japan. He included Trichiurus savala Cuvier in his synonymy of T. armatus Gray, but did not list any material at Cambridge. The present specimen may have been one that he overlooked. See Section 5 (p. 152) for discussion of this specimen. 134 P- J- P. WHITEHEAD & K. A. JOYSEY Family NOMEIDAE 24. Psenopsis anomala (Temm. & Schl.) *Trachinotus melo Richardson, 1846 : 270 (one Vachell specimen, the TYPE) (Reeves illustr. No. 97). Trachinotus melo: Giinther, 1860, 2 : 485 (on Richardson's description, no specimens). Not included in Harmer's list, presumed lost before 1893. The specimen is not in the British Museum collections. Reeves illustration reproduced here (Plate i, fig. 3). Identification follows Chu (1931) and Fowler (1936). Family FORMIONIDAE 25. Parastromateus niger (Bloch) *Seserinus vachellii Richardson, 1846 : 273 (two Vachell specimens, the larger 3-75 inches, the TYPES). Stromateus niger: Giinther, 1860, 2 : 401 (one Reeves specimen). Harmer lists two specimens under the name Platax teira " China. Rev. G. Vachell." — EXTANT. These specimens have been identified as the types of Seserinus vachellii. See Section 5 (p. 153) for description and notes. Family CARANGIDAE 26. Alectis indica (Riippell) Gallichthys major: Richardson, 1846 : 271 (one Vachell specimen) (Reeves illustr. No. 189). Caranx gallus: Giinther, 1860, 2 : 455 (one Reeves specimen and one other Chinese specimen). As Caranx gallus in Harmer's catalogue, two fishes " China. Rev. G. Vachell " — destroyed. Identification follows Fowler (1936, p. 297), who used Cuvier's generic name Scyris. 27. ? Alectis ciliaris (Bloch) Blepharis fasciatus: Richardson, 1846:271 (one Vachell specimen) (Reeves illustr. No. 269). Blepharis fasciatus (non Riipp.): Giinther, 1860, 2 : 422 (doubtful species No. 19, typical specimen in Camb. Phil. Inst.). Not listed by Harmer, presumed lost before 1893. Richardson identified his specimen with B. fasciatus Riippell, a synonym of A. ciliaris (Bloch), but Giinther disagreed (1860, p. 454), presumably on Richardson's description. Not in British Museum collections. Tentative identification of Fowler (1936, p. 295) followed here. 28. Caranx (Atule) kalla (Cuvier) * Caranx cancroides Richardson, 1846 : 274 (one Vachell specimen, the TYPE) (Reeves illustr. No. £30). Caranx cancroides: Giinther, 1860, 2 : 422 (doubtful species No. 12). Not mentioned in Harmer's list, presumed lost or destroyed before 1893. Reeves VACHELL COLLECTION OF CHINESE FISHES 135 illustration reproduced here (Plate I, fig. 2). W. L. Chan (in litt.} states that in Hong Kong the vernacular name Ha-tsee on the Reeves illustration (Hwa tsze in the " Report ") refers (prefix) to shrimp, and (suffix) to " a carangid fish either typically of the genus Decapterus, or of the subgenus Atule Jordan." Decapterus can be ruled out as being too slender (depth about 5 or more times in total length ; about 3 in Richardson's description and in the drawing). Caranx (Atule) kalla Cuv. is the most likely species and one that is caught in fair numbers by the Hong Kong shrimp trawlers. Richardson states " No spots are shown on the operculum " (present in C. kalla), but the figure shows a fish of only 4^ inches. 29. Caranx malabaricus (Bloch & Schneider) Caranx malabaricus: Richardson, 1846 : 275 (two Vachell specimens) (Reeves illustr. No. /?2i). Caranx malabaricus: Gunther, 1860, 2 : 436 (one Reeves and one other Chinese specimen ; BMNH. 1851.12.27.118). Not listed by Harmer, presumed destroyed or lost before 1893. An unregistered specimen merely labelled " Caranx malabaricus " in British Museum collection, but no indication of donor. Identification based on Fowler (1936, p. 293), who placed the species in Carangoides. 30. Citula armata (Forsskal) Caranx ciliaris: Richardson, 1846 : 276 (" spec. C. Ph. Inst."). Caranx armatus: Gunther, 1860, 2:453 (one Chinese specimen, BMNH. 1851.12.27.129, no donor given). Listed by Harmer as Caranx armatus " China. Cambridge Philosoph. Society's Collection " — EXTANT, Exhibited Series No. F.2755. There is also an unregistered British Museum specimen with an old label " Caranx ciliaris " amended to " arma- tus ". The jar is unusual, being oval in cross-section and sealed with parchment ; such jars were initially suspected of being part of the Vachell collection, but the present case shows this to be incorrect. The Cambridge specimen is discussed further in Section 5 (p. 154). Family LEIOGNATHIDAE 31. Leiognathus brevirostris (Valenciennes) Equula nuchalis: Richardson, 1846 : 276 (two Vachell specimens) (Reeves illustr. Nos. Ggo and £85). Equula nuchalis: Gunther, 1860, 2 : 500 (Reeves and other Chinese specimens). Not included in Harmer's list. Reeves illustration No. /? 85 is not a leiognathid, but No. G 90 is almost certainly L. brevirostris, having a distinct black pre-dorsal blotch, a body depth 21/5 times in S.L., and the lateral line not reaching the base of the caudal. Richardson gives no description. 136 P. J. P. WHITEHEAD & K. A. JOYSEY Family GERRIDAE 32. Gerres erythrourus (Bloch) Gerres equula: Richardson, 1846 : 239 (one Vachell fish) (Reeves illustr. No. 215). Gerres oyena: Giinther, 1859, 1 : 352 and 1862, 4 : 261 (no Chinese specimens). Not included in Harmer's list. Identification follows Chu (1931). Family AMBASSIDAE 33. Ambassis commersonii (Cuvier) *Ambassis vachellii Richardson, 1846 : 221 (one Vachell specimen, the TYPE) (no Reeves illustr.). Ambassis vachellii: Giinther, 1859, 1 : 227 (no British Museum specimens). Listed as Ambassis commersonii, " ? China Rev. G. Vachell " in Harmer's catalogue, two specimens — destroyed. Fowler (1937) tentatively recognized Richardson's species because of its deep body, but without specimens or illustration the species must remain doubtful. Family SERRANIDAE 34. Epinephelus fario (Thunberg) Serranus trimaculatus : Richardson, 1846 : 232 (Vachell specimens). Serranus trimaculatus: Giinther, 1859, 1 : 109 (Reeves fish and another Chinese specimen). As S. trimaculatus " China. Rev. G. Vachell " in Harmer's catalogue — destroyed, with date 21.4.1939. Identification based on Chu (1931) and Fowler (19380, P- 273)- 35. Epinephelus akaara (Temm. & Schl.) * Serranus shihpan Richardson, 1846 : 231 (Vachell specimens, also Reeves and Hyde Park collections all part of TYPE SERIES) (Reeves illustr. No. 71). Serranus diacanthus: Giinther, 1859, 1 : no (Chinese specimens, presented by Reeves, Richard- son and the East India Co.). Not included in Harmer's list. Giinther placed Richardson's species in the synonymy of 5. diacanthus. There are several unregistered Chinese specimens in the British Museum (including a Reeves fish) labelled 5. diacanthus, but none labelled 5. shihpan. Reeves illustration reproduced here (Plate 2, fig. i). Richardson was strongly inclined to refer this species to E. akaara, but decided to keep it separate because of the " dark bars which cross the body ". Matayama (1960), Chu, Tchang & Chen (1963), as well as earlier authors, have placed Richard- son's species in the synonymy of E. akaara, but Fowler & Bean (1930) placed it in the synonymy of E. malabaricus (Bloch & Schn.). W. L. Chan (in litt.) has pointed out that the vernacular name Sek-dang-paan is used in Hong Kong for one of the commonest species, E. fasciatomaculatus (Peters), whereas E. akaara is known as Hung-pan. Also, the absence of definite spots on the dorsal fin in the Reeves figure, and the inclination of the vertical bars on che flanks and their tendency to VACHELL COLLECTION OF CHINESE FISHES 137 fork ventrally, all point to E. fasciatomaculatus. However, Matayama (loc. cit.} placed E. fasciatomaculatus (Peters) as a synonym of E. fario (Thunberg), but con- sidered E. fasciatomaculatus of Fowler & Bean to have been E. diacanthus (Val.). Because of such uncertainties in the nomenclature we have preferred to let Richard- son's species remain in the synonymy of E. akaara for the time being. Family PRIACANTHIDAE 36. Priacanthus tayenus (Richardson) *Priacanthus tayenus Richardson, 1846 : 237 (one Vachell specimen and one Reeves specimen, the TYPES) (Reeves illustr. No. /3i4). Priacanthus tayenus: Giinther, 1859, 1 : 221 (a single specimen, collected by Reeves). Listed as P. japonicus " China. Rev. G. Vachell " by Harmer — destroyed. A single Reeves specimen in British Museum (BMNH. 1965.8.12.50) labelled as type. No evidence that the Vachell specimen is present. Reeves illustration reproduced here (Plate 2, fig. 2). Identification follows Chu (1931) and Fowler (19386, p. 67). Family PEMPHERIDAE 37. Pempheris otaitensis (Cuvier) Pempheris otaitensis: Giinther, 1860, 2 : 508 (one Chinese specimen). Listed by Harmer as P. otaitensis " China. Rev. G. Vachell " — destroyed. Richardson (p. 244) only mentions the related P. moluca Cuvier without reference to any specimens at all. Family LUTJANIDAE 38. Lut -janus erythropterus (Bloch) Mesoprion annularis: Richardson, 1846 : 229 (one Vachell specimen). Mesoprion annularis: Giinther, 1859, 1 : 204 (Reeves specimens). Two specimens included in Harmer's list as M. annularis " ? History "—destroyed. Identification follows Chu (1931) and Fowler (19386, p. 83). Family POMADASYIDAE 39. Pomadasys argenteus (Forsskal) Pristipoma nageb: Richardson, 1846 : 227 (one Vachell specimen) (Reeves illustr. No. 244). Pristipoma hasta: Giinther, 1859, 1 : 289 (Reeves and other Chinese material). Listed as P. hasta " China. Rev. G. Vachell " in Harmer's catalogue — destroyed. Giinther retained P. nageb as a separate species (1859, p. 290), but mentioned no Cambridge material and only tentatively placed Richardson's record in the synonymy. Fowler (19390) considered Richardson's record to refer to P. argenteus (Forssk.) and the Reeves illustration is consistent with this. ZOOL. 15, 3. 9 138 P. J. P. WHITEHEAD & K. A. JOYSEY Family THERAPONIDAE 40. Therapon jarbua (Forsskal) Therapon servus: Richardson, 1846 : 238 (no Vachell specimens mentioned) (Reeves illustr. No. 044). Therapon servus: Gunther, 1859, 1 : 278 (Reeves and other Chinese specimens). Listed as T. servus " China. Rev. G. Vachell " by Harmer, several specimens — destroyed. Richardson included three other species of Therapon (T. theraps, T. oxyrhynchus and T. quadrilineatus] , none of which were represented by Vachell material. Identification based on Fowler (19396, p. 204). Family SPARIDAE 41. Chrysophrys major (Temminck & Schlegel) Pagrus unicolor: Richardson, 1846 : 242 (Vachell specimen) (Reeves illustr. No. 160). Pagrus unicolor: Gunther, 1859, 1 : 468 (One Hong Kong fish presented by Richardson). Listed as P. major by Harmer " [A specimen from China, Rev. G. Vachell, was lost by evaporation of spirit] " Identification based on Fowler (1940, p. 53). 42. Spams latus (Houttuyn) ^Chrysophrys berda: Richardson, 1846 : 240 (no Vachell material) (Reeves illustr. No. 223). Chrysophrys hasta: Gunther, 1859, 1 : 491 (the Reeves type and other Chinese specimens). Listed as C. hasta, " China (? Rev. G. Vachell) " by Harmer — destroyed. Richard- son gives nine species of Chrysophrys, none of which included Vachell material. He placed Sparus hasta Bloch & Schn. in his synonymy of C. berda. Identification follows Chu (1931). Family MULLIDAE 43a. Upenoides subvittatus (Temm. & Schl.) Upeneus subvittatus: Richardson, 1846 : 219 (one Vachell fish, 4 inches). Mullus subvittatus: Gunther, 1859, 1 : 397 (doubtful species No. 2). Not listed in Harmer's catalogue. Fowler (1941) listed this Richardson record under Upeneus subvittatus. 43b. Upenoides subvittatus(Temm. & Schl.) Upeneus russelii: Richardson, 1846 : 220 (one injured Vachell specimen) (Reeves illustr. 036). Upeneus indicus: Gunther, 1859, 1 : 406 (two Reeves stuffed specimens, no others). Not listed by Harmer. Fowler (1941) identified Richardson's record as Pseudu- peneus indicus (Shaw). VACHELL COLLECTION OF CHINESE FISHES 139 Family SCIAENIDAE 44. Otolithes argenteus (Cuvier) Otolithus argenteus: Richardson, 1846 : 225 (one Vachell specimen, 6-55 inches). Otolithus argenteus: Giinther, 1860, 2 : 310 (Reeves and East India Co. specimens). Not included in Manner's list. The identifications of the three sciaenids listed here are based on the recent work by Chu, Lo & Wu (1963). 45. Collichthys lucidus (Richardson) Sciaena lucida Richardson, 1844 : 87, and 1846 : 224 (see below). Collichthys lucida: Giinther, 1860, 2 : 312 (Reeves and other Chinese specimens). Listed as Collichthys lucida " China. Rev. G. Vachell " in Harmer's catalogue — destroyed. Richardson made no direct reference to Vachell material but states " forms part of all the collections of Chinese fish that we have examined ". Richard- son described the species (Zoology of Sulphur, p. 87) on material in the British Museum, Hasler Museum and Camb. Phil. Soc. He measured a single specimen, 6-45 inches in length. There are two British Museum fishes labelled as types (BMNH. 1848.3.18.107-8) ; the larger of the two (163 mm. tot. 1., 6-4 inches) is most likely the specimen measured by Richardson. There are also two Haslar specimens (BMNH. 1855.9.19.195-7). 46. Dendrophysa russelii (Cuvier) Umbrina russelii: Richardson, 1846 : 226 (one Vachell fish) (Reeves illustr. No. /?37). Umbrina russellii: Giinther, 1860, 2 : 278 (Malaya and " China Seas? "). Listed by Harmer as Umbrina russellii " ? E. Indies " — destroyed. Family SILLAGINIDAE 47. Sillago japonica (Temm. & Schl.) Sillago japonica: Richardson, 1846 : 223 (Vachell and Reeves specimens) (Reeves illustr. No. £40). Sillago sihama: Giinther, 1860, 2 : 243 (Reeves and China specimens). Listed as 5. japonica " China. Rev. G. Vachell " by Harmer, three specimens — destroyed. Giinther (1860) referred all his Chinese material to S. sihama and not S. japonica ; presumably he changed his mind by the time he came to re-identify the Cambridge specimens. Identification based on Chu (1931). Family EPHIPPIDAE 48. Platax orbicularis (Forsskal) Platax teira: Giinther, 1860, 2 : 492 (China specimens). Harmer lists Platax teira, 2 specimens " China. Rev. G. Vachell." but, as shown earlier, the actual specimens are the types of Seserinus vachellii. Richardson (1846, I4o P. J. P. WHITEHEAD & K. A. JOYSEY p. 245) lists P. ehrenbergii Cuvier and P. vespertilis Bloch, both of which he identified from Reeves drawings ; in neither case does he mention specimens. Although the present specimens were mislabelled, the name Platax teira is included in this list of possible Vachell material. 49. Drepane punctata (Linnaeus) Drepane longimana: Richardson, 1846 : 245 (one Vachell fish) (Reeves illustr. No. 241). Drepane punctata: Gunther, 1860, 2 : 62 (two China specimens). Not listed by Harmer. Identification follows Fowler (1953, p. 12). Family SGATOPHAGIDAE 50. Scatophagus argus (Linnaeus) Scatophagus argus: Richardson, 1846 : 245 (two Vachell fishes) (Reeves illustr. No. " 272? "). Scatophagus argus: Gunther, 1860, 2 : 58 (one Reeves and other Chinese specimens). Listed by Harmer as " ? Fam. ? Scatophagus argus (2) PHistory " — destroyed. Identification follows Fowler (1953, p. 14). Family AGANTHURIDAE 51. Teuthis fuscescens (Houttuyn) Amphacanthus margaritiferus : Richardson, 1846 : 243 (one Vachell specimen, 7 inches) (Reeves illustr. No. 259). Teuthis albopunctata: Gunther, 1861, 3 : 318 (Reeves and other Chinese specimens). Not listed by Harmer. Identification based on Chu (1931). Family SCORPAENIDAE 52. Vespicula trachinoides (Cuvier) Apistes trachinoides: Richardson, 1846 : 213 (one Vachell fish). Prosopodasys trachinoides: Gunther, 1860, 2 : 139 (one Richardson specimen). Listed by Harmer as Prosopodasys trachinoides " ? East Indies " — destroyed. Richardson (p. 213) also lists the British Museum specimen, collected by Sir Edward Belcher. Identification follows Chu (1931). 53. Sebastodes vachellii (Richardson) *Sebastes vachellii Richardson, 1846 : 214 (one Vachell specimen, the TYPE) (Reeves illustr. No. 69?). Sebastes vachellii: Gunther, 1860, 2 : 95 (doubtful species No. i). Not listed by Harmer ; not amongst type material in British Museum. Reeves illustration reproduced here (Plate 2, fig. 3). W. L. Chan (in lift.) states that the caption to the Reeves illustration, pronounced in Hong Kong as Sek-gwau- VACHELL COLLECTION OF CHINESE FISHES 141 kwung (Shih kow kung in the " Report "), refers to one of the commonest of Hong Kong inshore fishes, Sebastiscus marmoratus (Cuv.), and that the drawing agrees with this. However, Richardson listed Sebastes marmoratus (" Report ", p. 215), based on a Biirger specimen in the British Museum, but was not able to identify a Reeves drawing with it. Further, he speaks of the " uncertainty of the drawing " of S. vachellii, citing it as " Icon. Reeves, 69? ". Since there are two intergrading colour forms of 5. marmoratus (Matsubara, 1943, p. 256), it is possible that Richardson did not recognize the Reeves drawing as that species. Neither Matsubara (loc. cit.) nor Chu, Tchang & Chen (1963) mention Richardson's 5. vachellii, and in the absence of the single Vachell specimen we have merely followed the synonymy of Chu (1931), one of the few authors to list this name. Family SYNANCEJIDAE 54 . Inimicus japonicus (Cuvier) *Pelor tigrinum Richardson, 1846 : 212 (one Vachell specimen, the TYPE) (Reeves illustr. No. M- Pelor japonicum: Giinther, 1860, 2 : 151 (two Reeves specimens). Listed as P. japonicum " China. Rev. G. Vachell " by Harmer — EXTANT. Reeves illustration reproduced here (Plate 3, fig. i). See Section 5 (p. 154) for notes on extant specimen. 55. Minous monodactylus (Bloch) Minous woora: Richardson, 1846 : 213 (one Vachell fish). Minous monodactylus: Giinther, 1860, 2 : 148 (Chinese specimens). Listed as M. monodactylus, " = Apistus minous, Cuv. (2) China. Rev. G. Vachell." by Harmer — destroyed. Identification follows Chu, Tchang & Chen (1963). 56. Polycaulus uranoscopus (Bloch & Schneider) Synanceia breviceps Richardson, 1844 : 71 (one Reeves and three Vachell fishes). Aploactis breviceps: Richardson, 1846 : 212 (three Vachell fishes). Polycaulus elongatus: Giinther, 1860, 2 : 175 (one Reeves fish, later cancelled, presumed des- troyed) . Listed as " Polycaulus elongatus (3) = Synanceia breviceps, Richards. China Rev. G. Vachell." by Harmer — destroyed. Identification follows Chu (1931). Family PLATYCEPHALIDAE 57. Platycephalus indicus (Linnaeus) Platycephalus insidiator: Richardson, 1846 : 216 (one Vachell specimen). Platycephalus insidiator: Giinther, 1860, 2 : 177 (Reeves and other China specimens). Listed as P. insidiator " China. Rev. G. Vachell " by Harmer, three specimens- destroyed. Identification based on Chu (1931). 142 P. J. P. WHITEHEAD & K. A. JOYSEY 58. Thysanophrys crocodilus (Tilesius) Platycephalus guttatus: Richardson, 1846 : 217 (one Vachell fish). Platycephalus guttatus: Gunther, 1860, 2 : 183 (no specimens — doubtful species). Not listed by Harmer. Chu, Tchang & Chen (1963) place Richardson's record in Inegocia guttata (Cuvier). The synonymy of Chu (1931) is followed here. Family POMACENTRIDAE 59. Amphiprion bicinctus (Riippell) Amphiprion chrysargyrus Richardson, 1846 : 254 (on Reeves drawing only) (Reeves illustr. No. L26). Amphiprion clarkii: Gunther, 1862, 4 : 5 (Reeves and other China material, including one stuffed fish collected by Reeves, the TYPE of A. chrysargyrus). A. chrysogaster " China. Rev. G. Vachell " is listed by Harmer (possibly an error since that is an Indian Ocean species) 2 fishes — destroyed. Richardson mentions no specimens but based the species on a Reeves drawing ; in fact a Reeves fish (stuffed) is present in the British Museum collections. Fowler (1954) placed Richardson's A. chrysargyrus in the synonymy of A. xanthurus Cuvier, but the pattern of white bands in the Reeves drawing corresponds much more nearly with Fowler's figure I for A . bicinctus. Family LABRIDAE 60. Halichoeres nigrescens (Bloch & Schneider) *Julis exornatus Richardson, 1846 : 258 (Bankier specimens from Hong Kong, and Vachell and Reeves material — TYPES) . Platyglossus dussumieri: Gunther, 1862, 4 : 143 (Reeves and other China specimens, including TYPES of Julis exornatus). Not listed by Harmer. Gunther listed China specimens as types of /. exornatus (BMNH. 1851.12.27.1634). The Vachell fishes can be considered to have been syntypical material. Two Richardson specimens in British Museum from Hong Kong (BMNH. 1848.3.8.6 and 1868.3.19.833). It is possible that these are Vachell specimens, but there is no evidence to show it. Reeves illustration re- produced here (Plate 3, fig. 4). Identification based on Fowler (1956, p. 230). Family GOBIIDAE 61. Glossogobius giuris (Ham. Buch.) Gobiusfasciato-punctatus: Richardson, 1846 : 204 (Vachell fish or fishes) (Reeves illustr. No. 146). Gobius giuris: Gunther, 1861, 8:21 (Reeves type of G. fasciato-punctatus in British Museum and other China specimens). Not listed by Harmer. Identifications and synonymies for the eleven gobioid species listed here have been based on Koumans (1931, 1953). VACHELL COLLECTION OF CHINESE FISHES 143 62. Glossogobius brunneus (Tern. & Schl.) *Gobius platycephalus Richardson, 1846 : 204 (one Vachell fish, the TYPE, " not in very good condition ") (Reeves illustr. No. LQ4). Gobius platycephalus: Giinther, 1861, 3 : 5 (doubtful species No. 39, related to G. brunneus). Not listed by Harmer. In the addenda to the " Report " Richardson states (p. 318) that he had just seen the description of G. brunneus by Temminck and Schlegel and he considered his G. platycephalus probably identical to that species. Vachell fish not amongst types in British Museum. Reeves illustration reproduced here (Plate 3, fig. 3). Fowler (1960) considered G. brunneus a synonym of G. giuris. 63. Cryptocentrus filifer (Valenciennes) Gobius filifer: Richardson, 1846:205 (Vachell material). Gobius knutteli: Giinther, 1861, 3 : 73 (Reeves and other China specimens). Listed as G. knutteli " China. Rev. G. Vachell." by Harmer — destroyed. A second Vachell specimen listed by Harmer as " [Origl. label = G. filifer] (probably the speci- men alluded to by Richardson, in Brit. Assn. Report, 1845, p. 205) " — destroyed also. 64. Rhinogobius margariturus (Richardson) *Gobius margariturus Richardson, 1846 : 205 (one, perhaps more, Vachell specimens — TYPE) (no Reeves illustr.). Gobius margariturus: Giinther, 1861, 3 : 47 (on Richardson, no specimens listed). As G. margariturus " China. Rev. G. Vachell " in Harmer's list — destroyed. Vachell specimen not amongst types in British Museum. Fowler (1960) placed this species in Ctenogobius and commented " A rare species, apparently not seen since described in 1846." 65. Drombus ripilepis (Richardson) *Gobius ripilepis Richardson, 1846 : 205 (one Vachell fish, the TYPE, 3^ inches) (no Reeves illustr.). Gobius ripilepis: Giinther, 1861, 3 : 5 (doubtful species No. 28, apparently allied to G. kokius "). Not included in Harmer's list. Placed in Ctenogobius by Fowler (1960). 66. Acanthogobius stigmothonus (Richardson) *Gobius stigmothonus Richardson, 1844 : 147, and 1846 : 205 (two Vachell fishes of 5 or 6 inches — the TYPES) (no Reeves illustr.). Gobius stigmothonus: Giinther, 1861, 3 : 76 (no specimens). Listed as G. stigmothonus (3 fishes) " China. Rev. G. Vachell " by Harmer— destroyed. The species was described by Richardson in the Zoology of the Sulphur (1844, p. 147) and was based on "... several specimens . . . brought from Canton by the Rev. George Vachell ". The Vachell specimens are not amongst the types in the British Museum. M4 p- J- P- WHITEHEAD & K. A. JOYSEY 67. Apocryptes serperaster (Richardson) *Apocryptes serperaster Richardson, 1846 : 206 (two Vachell specimens, 6 inches in length — the TYPES) (Reeves illustr. No. £55) . Apocryptes serperaster: Giinther, 1861, 3 : 82 (Reeves and other China specimens). Listed by Harmer as A. serperaster " ? China Rev. G. Vachell [labelled " Reeves "]" — destroyed. The words " Rev. G. Vachell " are crossed out, perhaps when the reference to Reeves was added. A Reeves specimen of 6 inches is in the British Museum, now labelled as a type, BMNH. 1965.8.12.51. It has a metal tag tied to the jaw "19.2.3.". Reeves illustration reproduced here (Plate 3, fig. 2). Family PERIOPHTHALMIDAE 68a. Boleophthalmus pectinirostris (Linnaeus) Boleophthalmus boddaerti: Richardson, 1846 : 208 (Vachell material) (Reeves illustr. No. ^38). Boleophthalmus boddaerti: Giinther, 1861, 3 : 102 (Indian Ocean specimens only). Not included in Banner's list. Fowler (1962) distinguished B. boddaerti (Pallas) from B. pectinirostris mainly on the presence in the former of dark transverse badns on the body. The Reeves drawing does not show these bands. 68b. Boleophthalmus pectinirostris (Linnaeus) Boleophthalmus pectinirostris: Richardson, 1846 : 208 (one Vachell fish, 2-80 inches). Boleophthalmus pectinirostris: Giinther, 1861, 3 : 102 (Reeves specimen). Not included in Harmer's list. Unregistered Reeves specimen in British Museum of 132 mm. tot. 1., and an unregistered Haslar specimen of 106-5 min- "tot. 1., labelled merely " 158 ". Both are too large to have been the Vachell fish. 6ga. Scartelaos histophorus (Valenciennes) Boleophthalmus aucupatorius Richardson, 1844 : 148, and 1846 : 208 (Vachell material). Boleophthalmus viridis: Giinther, 1861, 3 : 104 (" b,c. Half-grown. China. Presented by Sir J. Richardson — Types of B. aucupatorius."). Not included in Harmer's list. Described by Richardson in the Zoology of the Sulphur (1844, P- T4&}> based on fishes collected by Sir Everard Home, John Reeves and the Rev. George Vachell. Two Chinese specimens in the British Museum (BMNH. 1965.8.12.52-3) presented by Richardson; the larger of the two (2-65 inches total length) is undoubtedly Richardson's measured and figured specimen. There is no indication, however, that these are Vachell specimens. 6«, 134, 139, 153 vespertilis, 140 PLATYCEPHALIDAE, 141 Platycephalus guttatus, 142 indicus, 141 insidiator, 141 platy cephalus, Gobius, 143 Platyglossus dussumieri, 142 playjairii, Coilia, 129, 149 PLOTOSIDAE, 131 Plotosus anguillaris, 131 lineatus, 131 Polycaulus elongatus, 141 uranoscopus, 141 POMACENTRIDAE, 142 POMADASYIDAE, 137 Pomadasys argenteus, 137 PRIACANTHIDAE, 137 Priacanthus japonicus, 137 tayenus, 137 Pristipoma hasta, 137 nageb, 137 Prosopodasys trachinoides , 140 Psenopsis anomala, 134 Pseudobagrus aurantiacus, 130 fulvi-draco, 131 vachellii, 130 Pseudupeneus indicus, 138 pulchella, Parapercis, 146 Percis, 146 punctata, Drepane, 140 quadrilineatus , Therapon, 138 reevesii, Callionymus, 145 Leucosoma, 129 Rhinogobius margariturus, 143 164 P. J. P. WHITEHEAD & K. A. JOYSEY ripilepis, Drombus, 143 Gobius, 143 ruber, Holocentrus, 132, 151 rubrum, Holocentrum, 132 rugosus, Amblyopus, 145 russelii, Dendrophysa, 139 Umbrina, 139 Upeneus, 138 russellii, Umbrina, 139 SALANGIDAE, 129 Salanx chinensis, 129 savala, Leptur acanthus, 133, 152 Trichiurus, 133 Scartelaos histophorus, 144, 155 viridis, 156 SCATOPHAGIDAE, 140 Scatophagus argus, 140 Sciaena lucida, 139 SCIAENIDAE, 139 scripta, Alutera, 147 scriptus, Monacanthus, 147 SCORPAENIDAE, 140 scyllia, Triakis, 128 scyllium, Triakis, 128 Seym, 134 Sebastes marmoratus, 141 vachellii, 140 Sebastiscus marmoratus, 141 Sebastodes vachellii, 140 serper aster, Apocryptes, 144 SERRANIDAE, 136 S err anus diacanthus, 136 shihpan, 136 trimaculatus, 136 servus, Therapon, 138 Seserinus vachellii, 134, 139, 153 shihpan, Serranus, 136 sihama, Sillago, 139 SILLAGINIDAE, 139 Sillago japonica, 139 sihama, 139 sinense, Pelor, 155 Solea, ommatura, 132 ovata, 131 SOLEIDAE, 131 SPARIDAE, 138 Spams hasta, 138 latus, 138 Sphaeroides, 157 Sphyraena chinensis, 133 obtusata, 133 SPHYRAENIDAE, 133 spinosissimus, Holocentrus, 151 stellatus, Abalistes, 146 Balistes, 146 stigmothonus, Acanthogobius, 143 Gobius, 143 Stromateus niger, 134 Strongylura strongylura, 131 strongylura, Strongylura, 131 Strongylurus , B clone, 131 subvittatus, Mullus, 138 Upeneus, 138 Upenoides, 138 Synanceia breviceps, 141 SYNANCEJIDAE, 141 Synaptura zebra, 132 Taenioides anguillaris, 145 tayenus, Priacanthus, 137 fern*, Platax, 134, 139, 153 TETRAODONTIDAE, 147 Tetrodon ocellatus, 147, 157 Teuthis albopunctata, 140 fuscescens, 140 Therapon jarbua, 138 oxyrhynchus, 138 quadrilineatus , 138 servus, 138 theraps, 138 THERAPONIDAE, 138 theraps, Therapon, 138 thrissa, Clupanodon, 128 Clupea, 128 Thysanophrys crocodilus, 142 tigrinum, Pelor, 141, 154 Torquigener, 157 trachinoides, Apistes, 140 Prosopodasys, 140 Vespicula, 140 Trachinotus melo, 134 TRIACANTHIDAE, 146 Triacanthus biaculeatus, 146, 156 brevirostris, 146, 156 weberi, 157 Triacanthus (Trixiphichthys), 157 TRIAKIDAE, 128 Triakis scyllia, 128 scyllium, 128 TRICHIURIDAE, 133 Trichiurus armatus, 133, 152 intermedius, 133, 152 lepturus, 133, 152 muticus, 133, 152 savala, 133, 152 INDEX 165 tricuspidatus , Congrus, 129 trigrammus, Cynoglossus, 132 trimaculatus , Serranus, 136 Trypauchen vagina, 145 TRYPAUCHENIDAE, 145 unicolor, Pagrus, 138 Umbrina russelii, 139 russellii, 139 Upenoides subvittatus, 138 Upeneus indicus, 138 russelii, 138 subvittatus, 138 URANOSCOPIDAE, 146 Uranoscopus asper, 146 japonicus, 146 uranoscopus, Polycaulus, 141 Uroconger lepturus, 130 vachellii, Ambassis, 136 Bagrus, 130 Batistes, 124, 146 Pseudobagrus, 130 Sebastes, 140 Sebastodes, 140 Seserinus, 134, 139, 153 vagina, Trypauchen, 145 vespertilis, Platax, 140 Vespicula trachinoides, 140 viridis, Boleophthalmus, 144 Gobius, 156 Scartelaos, 156 vulgaris, Anguilla, 129, 150 weberi, Triacanthus , 157 woora, Minous, 141 xanthurus, Amphiprion, 142 zebra, Synaptura, 132 Zebrias, 132 Zebrias zebra, 132 PLATE i FIG. i. Chatoessus maculatus ( — Clupanodon thrissa). Reeves p. 91, No. 109. FIG. 2. Caranx cancroides ( = Caranx (Atule) hallo). Reeves p. 35, No. ^30. FIG. 3. Tmchinotus melo (=Psenopsis anomala). Reeves p. 68, No. 97. FIG. 4. Solea ommatura (=Zebrias zebra). Reeves p. 71, No. ^13. Bull. Br. Mus. nat. Hist. (Zool.) 15, 3 PLATE i ZOOL. 15, 3 PLATE FIG. i. Serranus shihpan (=Epinephelus akaara). Reeves p. 14, No. 71. FIG. 2. Priacanthus tayenus (=Priacanthus tayenus). Reeves p. 71, No. ^1 FIG. 3. Sebastes vachellii ( = Sebastodes vachellii). Reeves p. 14, No. 69. Bull. Br. Mus. nut. Hist. (Zool.) 15, 3 PLATE 2 PLATE 3 FIG. i. Pelor tigrinum ( = Inimicus japonicus). Reeves p. 28, No. ^42. FIG. 2. Apocryptes serperaster (=Apocryptes serperaster). Reeves p. 96, No. /?55. FIG. 3. Gobius platycephalus (=Glossogobius brunneus). Reeves p. 124, No. 194. FIG. 4. Julis exornatus (=Halichoeres nigrescens). Reeves p. 56, No. /Sio. Bull. By. Mus. nat. Hist. (Zool.) 15, 3 PLATE 3 PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING 27 FEB1967 k NOTES ON ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) V. NARDOA AND SOME OTHER OPHIDIASTERIDS AILSA MCGOWN CLARK BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 15 No. 4 LONDON: 1967 NOTES ON ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) V. NARDOA AND SOME OTHER OPHIDIASTERIDS BY AILSA MCGOWN CLARK British Museum (Natural History) >c Pp. 167-198 ; 6 Plates BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 15 No. 4 LONDON: 1967 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted, in 1949, is issued in five series corresponding to the 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. 15, No. 4 of the Zoological 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.) Trustees of the British Museum (Natural History) 1967 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 24 February, 1967 Price NOTES ON ASTEROIDS IN THE BRITISH MUSEUM (NATURAL HISTORY) V. NARDOA AND SOME OTHER OPHIDIASTERIDS By AILSA McGOWN CLARK THE present study includes notes on the provenance of Nardoa variolata (Lamarck) the type-species of Nardoa, details of the true holotype of N. gomophia (Perrier), description of a new species from northern Australia and a discussion of the remain- ing species of the genus coupled with notes on the validity of the monotypic Gomo- phia, a new subspecies of which is described. In the course of this, the following conclusions are reached : — Ophidiaster watsoni Livingstone (1936) is referred to Gomophia and is probably conspecific with G. egyptiaca Nardoa mamillifera Livingstone (1930) and N. tumulosa Fisher (1917) are prob- ably synonyms of N, frianti Koehler Nardoa obtusa (Perrier) is conspecific with N. tuberculata Gray Nardoa lemonnieriKoehler is revived from the synonymy of N. mollis de Loriol and Nardoa faouzii Macan is designated as type-species of a new subgenus. A key to the species of Nardoa is included. Other Ophidiasterids dealt with include :— Fromia indica (Perrier), with F. tumida Bell newly referred to synonymy but F. elegans H. L. Clark revived as a valid species A new species of Fromia from the Indian Ocean . Ferdina Gray, re-diagnosed and restricted to a monotypic genus by removal of F. heffernani Livingstone to a new genus ; Ferdina flavescens Gray partially redescribed Tamaria tumescens (Koehler), with T. ajax Livingstone, 19320, newly referred to synonymy and Ophidiaster helicostichus Sladen now referred back to Ophidiaster from Hacelia with possible synonym 0. astridae Engel, 1938. Gomophia egyptiaca egeriae1 subsp. nov. (PI. i, figs. 1-5) Nardoa tuberculata (pt.) Bell, 1894 : 396. ? Nardoa aegyptiaca : Koehler, 1910 : 157 [Non Gomophia egyptiaca Gray]. MATERIAL. Macclesfield Bank, South China Sea ; Admiralty : — 23 metres, 2 specimens — the holotype and paratype, B.M. reg. no. 92.8.22.52 and 53 ; 66-76 metres, 3 specimens, nos. 92.8.22.74, 259 and 260 ; 58 metres, I specimen, no. 92.8.22.29 ; 76-84 metres, I specimen, no. 93.8.25.203. 1 Named after H.M.S. " Egeria " which surveyed Macclesfield Bank. ZOOL. 15, 4 12 iyo A. M. CLARK DESCRIPTION OF HOLOTYPE. R/r = 70 mm./n-5 mm. = 6-1/1. The arms taper fairly evenly to attenuated tips ; br at the base = 10-5 mm., at half R = 9-0 mm. and at 10 mm. from the tip = 4-0 mm. The aboral skeleton is reticulate but the oblong secondary plates linking the five- or six-lobed primary abactinal ones are relatively inconspicuous and the reticulum is more compact than in the holotype of G. egyptiaca egyptiaca, the pore-areas being relatively small, mostly with only 3-6 pores in each. Three plates around the centre of the disc and fifteen to twenty other primary ones scattered along the upper side of each arm are abruptly elevated, the larger ones 2-4-2-6 mm. in basal diameter and 2-0-2-4 mm. in height. Their shape is conical but some have the apex blunted. The supero-marginal series of plates similarly includes both flat and markedly convex plates, the former being mostly equal in size to the largest of the flat abactinal plates. The convex plates arise at intervals of one to four flat ones, and are more widely-spaced on the distal half of the arm. The entire surface is coated with fine granules concealing the underlying plates. On the flatter parts of the aboral skeleton these number c. 6o/sq. mm. but on the convexities they become distinctly coarser, especially towards the tips of the plates. Most of these tips have been rubbed but on those that remain intact the apex is crowned by several enlarged granules among which a single central one may reach twice the diameter of the granules encircling it. Though slightly higher than the other granules, this single one is throughout too small to evoke comparison with a nipple. It is more often developed on the supero-marginal plates than the abactinal ones. The infero-marginal plates form a much more regular series than the supero- marginals, all being similar and gradually reducing in size distally ; the proximal ones are conspicuous by their size and measure almost 3 mm. in diameter. The number in one series partially denuded is c. 30. Between the two main marginal series proximally is a row of intermarginal plates. Beyond the basal third of the arm length these are concealed by the granulation but spaced ones are revealed on the denuded arm to c. half R. Small pore-areas are present both above and below the intermarginal plates continuing distally beyond them but there are none on the lower surface of the body below the infero- marginals. The actinal series of plates is poorly developed, the one cleared consisting of 18 plates, about three corresponding to each infero-marginal where they lie adjacent ; the distalmost actinal plate lies interstitially between the seventh and eighth infero- marginals and the adjoining adambulacrals, i.e. at just over one-third R from the mouth. The adambulacral plates for much of the arm each bear four, rarely five, prismatic furrow spines c. I mm. long, the fans of which are slightly oblique so that consecutive ones tend to overlap. Behind these are series of usually three shorter subambulacral spines, the middle one becoming progressively stouter and relatively longer on the distal part of the arm, while the proximal one disappears or becomes granuliform. The outer part of the plate bears polygonal granules indistinguishable from those of the adjacent actinal and infero-marginal plates so that the two rows of spines project NOTES ON ASTEROIDS: OPHIDI ASTERIDS 171 abruptly. In comparison with the species of Nardoa, the adambulacral spines are more slender and spaced farther apart. PARATYPES. [Treated in the same order as in table i on p. 177]. As indicated in the table, the raised abactinal and supero-marginal plates are more or less rounded at the tips and may lack the single enlarged granule altogether. In the specimen from the same depth as the holotype (R 65 mm., PL i, fig. 4) the enlargement of the supero-marginal plates is more frequent, there being a marked tendency for every second one to be convex. In this specimen the furrow spines number five on most of the plates of the proximal half of the arm ; also the granules adjacent to the subambulacral spines are slightly elongated. No. 93.8.25.203 (R7omm.) has alternate proximal supero-marginal plates tubercular but distally the frequency of these is less. Again there are five furrow spines on most of the proximal plates. No. 92.8.22.29 (R6omm.) has very regular alternation of convex and flat supero-marginals for almost the entire length of all the arms. There are four, occasionally five, furrow spines. In no. 92.8.22.74 (R 52 mm.) the single apical granule of some of the supero- marginal plates at least (most of the aboral ones being rubbed) approaches nipple- like proportions. The furrow spines usually number four. Finally, nos. 92 . 8 . 22 . 259 and 260 are smaller, R 34 and 32 mm. (The latter has only two arms intact and is not included in the Table). The former (PI. i, fig. 5) has markedly conical tubercular plates, of which the two midradial ones near the base of each arm are so large as almost to meet the supero-marginal plates each side, only one small flat plate intervening. This is in marked contrast to the specimens of G. egyptiaca egyptiaca of similar size from Christmas Island, where the abactinal plates are relatively smaller and more numerous (PL i, fig. 6). Another difference is that the intermarginal plates are very narrow in the Macclesfield Bank specimen and indeed are only visible when the granulation is removed, being sandwiched closely between the two main marginal series. The Christmas Island specimens have conspicuous broad series of intermarginal plates extending for about half R. However, the two from Macclesfield Bank cannot be referred to Nardoa frianti since they both have unusually well-developed, almost nipple-sized and sharp apical tubercles on the conical abactinal and supero-marginal plates. It may be noted that a few of the supero-marginals are crowned by two or even three tubercles, which may also occur in G. egyptiaca egyptiaca, as shown by a specimen from Samoa (poorly preserved and not included in Table i) though here the tubercular plates tend to have a double apex like a cow-bell and the tubercles are separated at the ends of the ridge. Comparative remarks between egeriae and egyptiaca as well as with Nardoa are included in the general discussion of Nardoa and Gomophia. Nardoa variolata (Retzius) Tetractis cruciata variolata Linck, 1733 : 19, pi. i, no. i. Pentadactylosaster variolatus Linck, 1733 : 34, pi. viii, no. 10. Hexactis variolata Linck, 1733 : 37, pi. xiv, no. 24. 172 A. M. CLARK [Un-named] Encyclopedie Methodique, 1792 : pi. cxix, figs. 4, 5 [" Asterias variolata : Lamarck " in captions, 1827]. Asterias variolata Retzius, 1805 : 19 ; Lamarck, 1816 : 565. Linkia variolosa (sic) : Nardo, 1834 : 717. Nardoa variolata : Gray, 1840 : 286 ; 1866 : 15 ; Fisher, 1919 : 379, pi. cviii, fig. 4 ; H. L. Clark, 1921 : 51, pi. xxviii, figs. 3, 4. [Non Domantay & Roxas, 1938 : 224-225, probably = N. novaecaledoniae]. Nardoa Agassizii Gray, 1840 : 287 ; 1866 : 15. Scy 'taster variolatus : Miiller & Troschel, 1842 : 34-35 ; Michelin, 1845 : 21 ; Peters, 1852 : 178 ; Liitken, 1865 : 41 ; Perrier, 1875 : 159-162 (423-426) ; Smith, 1879 : 566 ; Mobius, 1880 : 50 ; de Loriol, 1885 : 43-44. [Non Bell, 1882 : 220 ; 1887 : 647 ; ?i9O2 : 226 ; Pigog : 19]. This species was formally designated as type-species of Nardoa by H. L. Clark (1921). Unfortunately, although there has long been agreement that the common Nardoa of Mauritius should be called variolata, the basis for this is not perfectly secure since no holotype is extant. One solution would be for a neotype to be chosen. Alternatively a lectotype could be selected from among Linck's figures (1733) which were cited by Retzius, the first post-Linnean author to use and thereby validate the name. As Linck's specimens are not preserved at the present museum in Trier the figures alone will have to suffice. Those of the normal five-armed specimen (Linck, pi. viii, no. io)reproducedin the" Encyclopedie Methodique", are to be preferred; they certainly have a strong resemblance to the common Nardoa of Mauritius. Linck, Retzius and Nardo all failed to give a locality for the species but Lamarck put " mers d'Europe " from which Gray appears to have derived " Mediterranean Sea " at the same time as he gave " Isle of France " (Mauritius) for Nardoa Agassizii, subsequently referred to the synonymy of variolata by Miiller and Troschel, who also record specimens from Mauritius. Some remarks about the geographical and morphological limits of the species are included in the general discussion of Nardoa. Nardoa gomophia (Perrier) (PI. 2, figs. 1-4) Scy taster gomophia Perrier, 1875:167-169 (431-433). [Non Nardoa " gamophia " : Fisher, 1919 : 380, which is based on a specimen of N. novaecaledoniae]. MATERIAL. New Caledonia ; Cuming ; the holotype, B.M. reg. no. 59.4.25.3. As Fisher noted in 1919, the specimen (PL 5, fig. 2) in the British Museum col- lections hitherto labelled as the holotype of Perrier's Scytaster gomophia does not particularly resemble Gomophia egyptiaca with which Perrier allied and compared his new species but is close to Nardoa novaecaledoniae. Its number is 59.4.25.4 and a search through the dry collections yielded a second specimen not conspecific with the first but also labelled as from New Caledonia and having the consecutive number 59.4.25.3 (PI. 2, figs. 1-4). This specimen bears a label" Scytaster n. sp. ", though not in Perrier's handwriting. A later printed label names it Nardoa perrieri, which I think must be an MS name given it by Bell prior to placing it on exhibition. This second specimen does have a considerable resemblance to Gomophia egyptiaca NOTES ON ASTEROIDS: OPHIDI ASTERIDS 173 owing to the presence of some spaced, abruptly projecting, tubercular plates on the upper side. As in Perrier's description, these are hemispherical rather than conical, unlike G. egyptiaca, and tend to form three alternating series along each arm. No such projections are present in the first specimen, 59.4.25.4, but this does agree in diameter with Perrier's measurement of 135 mm., the mean of its five possible diameters being 131 mm., whereas the same mean in no. 59.4.25.3 is 167 mm. However, the R/r ratio of 7-7/1 in the latter compared with 5-5/1 in no. 4 agrees so closely with Perrier's estimate of " nearly 8/1 " that I am convinced the wrong specimen has been labelled as the holotype of N. gomophia. Accordingly the assessment of the species must be reviewed since H. L. Clark referred gomophia to the synonymy of N. novaecaledoniae following Fisher's comments. On the one hand it is clearly related to those species of Nardoa which have some of the abactinal plates distinctly tubercular and on the other to Gomophia egyptiaca, as Perrier stated. To avoid repetition, the comparison is included in the general discussion of Nardoa and Gomophia. Nardoa sphenisci1 sp. nov. (PI. 3, figs. 1-3) MATERIAL. Holothuria Bank, N.W. Australia, 27 metres ; H.M.S. " Penguin ". i specimen, the holotype, B.M. reg. no. 92.1.14.27. Parry Shoal (N.W. of Darwin), 22 metres ; H.M.S. " Penguin ". 2 specimens, no. 92.4.4.6 and 7. DESCRIPTION OF THE HOLOTYPE. R 115-120 mm. ; r c. 13 mm. ; R/r = 9-0/1. The arms are slender and attenuated, br at the base = 13-14 mm., at half R = c. 10 mm., at 10 mm. from the tip = 4-5-5-0 mm. and at the very tip = c. 2 mm. One arm has been broken near the base and is regenerating. Most of the abactinal plates are more or less convex, especially the midradial ones near the bases of the arms, which are 2-5-3-5 mm. in transverse diameter (slightly less longitudinally) and up to 1-5 mm. in height ; their tops are rounded. Beyond the bases of the arms the more convex plates are scattered over the whole upper side. Across the base of each arm between the two supero-marginal series the abactinal plates number five to seven ; when there are seven the two outermost plates are small, for most of the proximal part of the arm there are about five plates across the width though the arrangement is irregular. Distally the plates gradually decrease in size and become elongated, the more convex ones measuring 2-0-2-5 mm. in length and 1-0-1-5 mm- m breadth. Only on the regenerating arm is there an abrupt reduction in the size of the plates at the point of cleavage. The entire aboral surface is coated with polygonal granules which gradually increase in size on the convexities of the skeleton. The smaller granules between and towards the edges of the plates number 25-30 per square mm. ; they are flat-topped and each granule bears a small boss in the centre except for the coarser ones near the apices of the more conical plates, which tend to have their whole free surface 1 Of the " Penguin ". 174 A. M. CLARK convex and smooth. The smaller distal convex plates particularly have the few enlarged central granules distinctly projecting and on some of these plates there is one central granule which is markedly larger than the seven or eight immediately encircling it. This gives a rough texture to the distal parts of the arms although the central granules are barely visible with the naked eye and never so abruptly different from the other granules as to recall the nipple-like tubercles found on the swollen plates of most specimens of Gomophia egyptiaca. The supero-marginal plates also are more or less convex in form though never to the same extent as the proximal mid-radial plates. There is a strong tendency for alternate supero-marginals beyond the basal five to ten to be larger and more convex than the rest so that the outline of the arms in perpendicular view is distinctly and fairly regularly scalloped. The larger plates of the basal half of the arm are 2-0-2-5 mm. in diameter, the alternate plates being only c. i-o mm. long though still c. 2 mm. wide. In two supero-marginal series counted the total number of plates is 58. In contrast the infero-marginal plates are fairly equal in size, gradually diminishing distally. They number c. 54. Both series of marginal plates are covered with granulation similar to that on the abactinal plates ; this again becomes increasingly coarse on the more convex surfaces. On some of the distalmost plates of both series when only a single granule is enlarged the height of this may exceed its basal breadth though the tip is rounded. In each interradius the two marginal series are separated by a progressively narrowing series of irregularly-placed and irregularly-shaped intermarginal plates. On one area cleared these number about 40 and extend for c. 40 mm. along the arm, although when the granule-covering was intact the plates of the distal half of the series were concealed. On the lower side, the main series of actinal plates is visible through the covering granulation for about half the arm length, (i.e. to about the twenty-fifth infero- marginal) ; further plates extend interstitially concealed by the granulation to about the thirty-fifth infero-marginal or two-thirds of the arm length. The actinal plates are twice as numerous as the adjoining infero-marginals. On the disc there is a second series consisting of only two or three plates. The granulation is again coarser on the centres of the plates than around their edges. The adambulacral plates correspond in number to the actinal plates. Each bears four or five blunt angular furrow spines, the distal plates more often four, though the number may be only three on the terminal fifth of the arm. When there are five furrow spines, usually the proximalmost one is slightly smaller than the rest and inset from the furrow so that the series is curved and adjacent ones tend to overlap. The subambulacral row consists usually of three spines, shorter but somewhat stouter than the furrow spines and often with a smaller fourth proxi- mal spine almost in series with them. The outer part of the plate bears polygonal granules some of which, close behind the subambulacral spines are slightly elongated so that there is no abrupt transition from spines to granules. The pore-areas are small in comparison to the size of the plates. The number of pores in each area is difficult to estimate, perhaps owing to the dried condition ; NOTES ON ASTEROIDS: OPHIDI ASTERIDS 175 on the upper side it is probably 4-8 in most areas. Smaller pore-areas are also present intermarginally and below the infero-marginals on the proximal half of the arm. PARATYPES. Both have R/r 75-85 mm./io mm., approximately 8/1. The arms are slender and attenuated as in the holotype, which they closely resemble. Both have the intermarginal series of plates well-developed on most arms and only lacking on those arms which appear to have regenerated from the base. This can be assumed from the abrupt break in all three marginal series close to the interradius and to coincidence in the occurrence of intermarginal series on both sides of a single arm rather than on the two continuous series of adjacent arms across an interradius. The actinal plates are more extensive than in the holotype, being visible for about five- eighths R with further plates extending to three-quarters R concealed to the naked eye beneath the granulation. The furrow spines number four on most plates of the proximal half of the arm, then three. REMARKS. All three specimens were identified as Nardoa tuber culata by Bell. In 1953 I re-named them Gomophia egyptiaca on account of the intermarginal series of plates and the attenuated arms, disregarding the rounded rather than conical tubercular abactinal plates lacking in central conical nipples. However, a recent study of material of these two genera convinces me that the three specimens are referable to neither species. Their relationships are dealt with below in the general discussion of Nardoa and Gomophia. NARDOA Gray Gray, 1840 : 268 ; 1866 : 15 ; Fisher, 1919 : 378-382 ; H. L. Clark, 1921 : 49-50. The species of Nardoa are usually divided into two groups, those in which some of the abactinal plates are distinctly more convex than the rest and those with similar low plates. The relationship between the first group and the monotypic genus Gomophia should be reviewed, the latter having been referred to the synonymy of Nardoa by several specialists including Sladen (1889), Koehler (1910) and Fisher (1919) but restored to generic rank by H. L. Clark (1921) and following workers. Unfortunately, most of the species involved seem to be solitary and the available samples are very inadequate, rarely more than one or two specimens being taken at a time so that the range of variation is poorly known. Following the restoration here of N. gomophia as a valid species and the addition of N. sphenisci as a new one, the nominal species to be included in such a survey are :— Gomophia egyptiaca Gray, 1840 Ophidiaster " watsoni Livingstone, 1936 Nardoa frianti Koehler, 1910 Nardoa tumulosa Fisher, 1917 Nardoa mamillifera Livingstone, 1930 Nardoa gomophia (Perrier), 1875 Nardoa sphenisci sp. nov. 176 A. M. CLARK Nardoa tuberculata Gray, 1840 Nardoa oUusa (Perrier), 1875 and possibly also Nardoa rosea H. L. Clark, 1921. Some of the distinguishing characters between these are brought out by the following table. In this the skeletal structure is indicated as either open, when the papular areas are sufficiently large to give a reticular appearance, as opposed to compact, with the spaces small and irregular in arrangement. A comparison of PL i figs. I and 3 with 6 and 8 may serve to illustrate this point). Negative entries for the abactinal tubercles signify that they are conical. The degree of fineness of the covering granulation is shown by an assessment of the number per square mm. taken on the flatter parts of the aboral skeleton where the granulation is at its finest. The other columns should be self evident. Intermediate conditions are indicated by " +/— " and a tendency one way or infrequent occurrence by brackets. H. L. Clark distinguished Gomophia from Nardoa by the unspecialized supero- marginal plates resembling the abactinal ones (this being enhanced by the series of intermarginal plates separating the supero-marginals from the markedly enlarged infero-marginals proximally) and by the absence of actinal papulae below the infero- marginals (this character implicit in his key) coupled with the conical form and apical " nipples " of the tubercular abactinal plates ; an additional distinction is provided by the rather open, apparently reticulate, aboral skeleton. However, although in the holotype of Gomophia egyptiaca and in a few other specimens closely resembling it, notably the one from Samoa shown in PL I, figs. 7 and 8, none of the supero-marginal plates are enlarged or projecting, this is not always the case ; for instance the fine large specimen from the Gulf of Aqaba (see A. M. Clark, 1952), the second one in the table, has a conspicuous conical tubercular plate in the position of every third or fourth supero-marginal. Similarly some much smaller specimens from Christmas Island in the Indian Ocean have some elevated supero-marginals; in this case alternate plates are so modified, as in the holotype of Livingstone's Ophidiaster watsoni to which the Christmas Island specimens bear great resemblance. In fact, despite the extent of some of the actinal series of plates beyond half the arm length and the slight enlargement of the granules adjoining the subambulacral spines, I have no hesitation in declaring 0. watsoni to be congeneric, possibly even conspecific, with Gomophia egyptiaca. It is significant that Endean (1965) has recorded G. egyptiaca from Queensland. Seven specimens from Macclesfield Bank, South China Sea, which I have referred to Gomophia, also have some of the supero-marginals, often alternate ones, tuber- cular ; such a frequency of the tubercular plates coupled with their linear arrange- ment gives a greater appearance of specialization to the supero-marginals than is evident in the holotype of G. egyptiaca. Bell included these China Sea specimens, together with three others which I believe are Nardoa frianti, in N. tuberculata. However, the presence of intermarginal series of plates, the much more prominent tubercular plates of the aboral side often crowned with enlarged granules or even a single small tubercle, the absence of actinal papulae, the reduction of the actinal series of plates and the abrupt transition from the granulation of the lower side to NOTES ON ASTEROIDS: OPHIDI ASTERIDS 177 ' t; ^3 P G 4) ll its 1111 s~§a i . i i i i+ i i . £ i £ Interstitial granulation no./sq. mm. 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CLARK the two rows of adambulacral spines all agree with G. egyptiaca. Nevertheless, when the fine coat of granules is removed from the upper side, the underlying skeleton appears to be much more compact in these specimens from Macclesfield Bank than in the others included in the table from the Red Sea, Mauritius, Samoa and Christmas Island. Possibly this can be attributed in some degree to contraction in preservation since an unnaturally flattened specimen from Eilat in the Red Sea similarly shows compaction of the abactinal skeleton. The more or less rounded form of the swollen abactinal and supero-marginal plates in most of the specimens from Macclesfield Bank, coupled with the poor development of the apical nipples (these often being confined to the supero-marginals) provides another distinction and three of the specimens have a slight transition from the granulation of the lower side to the subambulacral spines. Also the granulation of the abactinal plates seems to be slightly coarser than in those specimens of G. egyptiaca which most resemble the holotype. I do not consider that these differences carry sufficient weight to merit more than a subspecific distinction for the Macclesfield Bank specimens which are accordingly described here as Gomophia egyptiaca egeriae. The relative specialization of their supero-marginal plates weakens the generic distinction of Gomophia on the basis of this character. Of the other supposedly diagnostic characters, the absence of actinal papulae is certainly constant in G. egyptiaca but unfortunately is matched by individuals of several species of Nardoa, for instance the types of both Nardoa frianti and Nardoa tumulosa, also the specimen from the Ogasawara (Benin) Islands referred to frianti by Hayashi (19386) ; certainly the actinal papulae are few and inconspicuous in the specimens from Macclesfield Bank which I have attributed to frianti. As for the occurrence of intermarginal plates, these are not restricted to G. egypti- aca. Fisher notes that there are a few intermarginal plates at the base of each arm of the holotype of N. tumulosa, while Koehler writes to the effect that in the two syntypes of N. frianti " sometimes the two marginal series are fully contiguous but sometimes there are small plates intercalated between them, though these rarely form a regular and distinct series but rather are inserted in the intervals between the larger plates ". No such intercalary intermarginals are developed in the Macclesfield Bank specimens of Nardoa, while Hayashi (1938 and 19380) mentions only intermarginal papulae, not plates, in the specimens from the Caroline and Ryu Kyu Islands which he referred to N. tumulosa. However, the specimen from the Ogasawara Islands (Hayashi, 19386), figured under the name of N. frianti, not only has well-developed series of intermarginal plates but also has the tubercular abactinal plates restricted to the proximal parts of the arms (like Hayasaka's speci- men from Taiwan [Formosa], 1949, PI. ii, fig. 4, named tumulosa), combining to give it a strong resemblance to the holotype of N. tumulosa. Hayashi gave no reasons for naming his specimen frianti rather than tumulosa. Intermarginal series of plates are also developed in the type material of N. sphenisci. It should be noted that Nardoa frianti and Gomophia egyptiaca are sympatric for at least part of their ranges ; both have been taken in the Andaman Islands (Koehler), on Macclesfield Bank and in the Loyalty Islands. Koehler gave insuffi- cient information about his material of Gomophia from the Andamans for me to be NOTES ON ASTEROIDS: OPHIDI ASTERIDS 179 certain whether it should be left in egyptiaca or referred to subspecies egeriae, though his photograph suggests that the aboral skeleton is fairly open giving a reticulate appearance and thus linking it with egyptiaca sensu stricto. Following from the variation in shape of the abactinal tubercular plates in G. egyptiaca egeriae, I cannot help suspecting that Koehler's unfigured syntype (or paratype) of N. frianti should have been referred to Gomophia. This would account for the discrepancies between his description and the Macclesfield Bank specimens which I have referred to frianti with regard to the occasional presence of intermarginal plates and the absence of actinal papulae (though the latter could well evade detection if they are few in number). The generic status of Gomophia is therefore in some doubt but, since the range of variation and specific limits of most of the species involved in the comparison have yet to be properly determined, I hesitate at present to refer it yet again to the synonymy of Nardoa. [In this it must be confessed that I am influenced to some extent by respect for Gray's eye for a genus, so often vindicated]. Included in the table is a column dealing with the shape of the arm tips, whether blunt or attenuated. However, since several of the species of Nardoa appear to be particularly susceptible to loss of parts of the arms, the subsequent regeneration, when incomplete, may give a false impression of the true shape. However, the mutilated holotype of Nardoa mamillifera Livingstone has all three of there main- ing arms ending abruptly, R/r being 5-7/1. Similarly abbreviated arms are found throughout in the two specimens I have seen from the Loyalty Islands (R/r 5-7/1 and 5-5/1), which otherwise agree closely with the three of N. frianti from Macclesfield Bank. The latter all have markedly attenuated arms with R/r 7-75-8-2/1. In the syntypes of N. frianti the ratio is 7-3 and 7-1/1, while Domantay and Roxas (1938) recorded a specimen from the Philippines with a value as high as 8/1. Doder- lein (1926) recorded as N. frianti one from the Caroline Islands with R/r 6-4/1 but Hayashi (1938) identified as N. tumulosa his tuberculated Carolinian specimen ; this has relatively short though tapering arms, R/r being only 5/1. A specimen from the Ryu Kyu Islands which Hayashi (19380;) similarly named tumulosa has R/r 5-3/1. In Fisher's holotype of N. tumulosa the ratio is 6-4/1. There is con- siderable resemblance between the Ryu Kyu specimen and the individual taken by the Barrier Reef Expedition which Livingstone (1932) referred to Nardoa rosea. Despite Livingstone's assurance that the latter agrees perfectly with the original description of rosea and shows no variation, a direct comparison with a paratype of N. rosea (PL 5, figs. 3 and 4) reveals a marked disparity in the size and convexity of the larger abactinal plates, many of which in the Barrier Reef specimen are c. 3 mm. in transverse diameter and c. 1-5 mm. in height (compared with c. 2 mm. and 50 mm. ........ tumulosa Fisher, 1917 3' Tubercular plates relatively small, rarely much exceeding 2 mm. in diameter, even at R > 70 mm., often present distally as well as (or rather than) proximally . 4 4 Arms not attenuated but more or less blunt at the tip, R/r 5-0-7-0/1 ; the main actinal row of plates extending the whole length of the arm ; tubercular abactinal plates inconspicuous, tending to be fewer proximally and restricted to the lateral areas of the distal parts of the arms ; supero-marginal plates all similar in size, not alternating . tuberculata Gray, 1840 4' Arms very attenuated distally, R/r 7-5-9-2/1 (in the only four specimens known) ; the main actinal row of plates terminating at half to two-thirds R from the base of the arm ; tubercular abactinal plates well-developed, proximally as well as distally ; some of the supero-marginals, often alternate ones, reduced in size . 5 5 Intermarginal plates present in the arm angles ; the majority of abactinal plates markedly convex, particularly the midradial ones . . . sphenisci sp. nov. 5' No intermarginal plates ; tubercular abactinal plates widely separated, as well- developed laterally as mid-radially . . . gotnophia (Perrier), 1875 6 Supero-marginal plates tending to alternate large and small, especially distally rosea H. L. Clark, 1921 * An anagram of Nardoa. 1 If the abruptly terminating arms of the mutilated holotype of N. mamillifera are shared by other specimens from the type-locality this might provide a valid distinction from N. frianti, in which case specimens from the Loyalty Islands could also be referred to mamillifera. :88 A. M. CLARK 6' No regular alternation in size of the supero-marginals ..... 7 7 Abactinal plates of the proximal third to half of the arm forming distinct regular longitudinal and transverse series, though this breaks down distally Nardoa (Andora) fouazii Macan, 1938 7' Abactinal plates irregular in arrangement throughout ..... 8 8 Adambulacral plates each bearing only four furrow spines and four subambulacral ones (though in my interpretation of Liitken's danish the granulation backing on to these spines is transitional) ..... galatheae (Lutken), 1865 8' Granulation of the outer part of each adambulacral plate usually modified to form a third row of spines, numbering three like the two inner rows, though the furrow series may consist of four spines ......... 9 9 Many of the abactinal plates more or less conspicuously broadened and alternating to form an almost continuous pavement, the interstitial pore-areas being small ; the larger plates markedly bigger than the supero-marginals, exceeding 3 mm. in diameter, even at R only c. 40 mm., numbering only about five across the width of the arm basally at R < 50 mm., seven at R 50-70 mm. or up to nine in the rare specimens that exceed 70 mm. ; the distal abactinal plates similar to the proxi- mal ones though gradually becoming smaller . . variolata (Retzius), 1805 9' Few if any of the abactinal plates conspicuously larger than the supero-marginals and rarely broadened transversely to exceed 2-5 mm. in diameter ; if some large plates are present proximally then the distal plates are relatively much smaller, often with an abrupt change in magnitude at about two-thirds R . . . 10 10 All the abactinal plates, both proximally and distally, small in comparison with the marginal plates, numbering about 13 across the arm width basally, at least at R >go mm. ; granuliform pedicellariae often present between the aboral plates and distinctly coarser than the granules ; ventral papulae few and inconspicuous pauciforis (von Martens), 1866 10' The proximal abactinal plates, especially midradially, distinctly larger than the distal plates and sometimes equalling or even slightly exceeding the supero- marginals in diameter, rarely more than eleven plates across the width, more often seven or nine ; pedicellariae not recorded ; usually a number of pore-areas developed between infero-marginal and actinal plates . . . . . n 11 R/r usually 5-0-6-0/1 (see Table 2) . . novaecaledoniae (Perrier), 1875 n' Arms more attenuated, R/r usually 5*5-8-5/1 . . . . . . . 12 1 2 Distal abactinal plates conspicuously smaller than the proximal ones, not exceeding i mm. in transverse diameter, their positions shown by rounded groups of coarse granules among the fine interstitial granulation . . mollis de Loriol, 1891 12' Distal abactinal plates becoming gradually somewhat smaller than the proximal ones and usually 1-2-1-5 rnm. in diameter but markedly elongated in form with ovate groups of coarse granules ..... lernonnieri Koehler, 1910 Fromia indica (Perrier) Scytaster indicus Perrier, 1869 : 255 (?63). Fromia indica : Perrier, 1875 : 177-178 (441-442) ; Koehler, 1910 : 140, pi. xvii, figs. 7, 8 ; H. L. Clark, 1921 : 42 ; Hayashi, 19380 : 207 ; 19386 : 59-62, figs, i, 2, pi. iv, figs. 1-4 ; 1938^ : 279-280. Fromia tumida Bell, 18820 : 124, pi. vi, fig. 4 ; H. L. Clark, 1921 : 41. Fromia andamanensis Koehler, 1909 : 105-108, pi. vii, figs. 5, 6 ; H. L. Clark, 1921 : 41. Fromia elegans : Engel, 1938 : n, fig. i, pi. iii, fig. i [Non F. elegans H. L. Clark, 1921]. Fromia indica forma andamanensis ; Hayashi, 1938 : 428-430, pi. ii, figs. 7, 8, NOTES ON ASTEROIDS: OPHIDI ASTERI DS 189 MATERIAL. Ceylon ; Mr. Kelaart ; two syntypes of Fromia tumida Bell, B.M. reg. no. 52.5.15.62 and 63. Ceylon ; Dr. Ondaatje ; I specimen, no. 88.4.13.3. Andaman Islands ; Dr. Anderson ; 5 specimens, no. 86.6.26.20-22 and 79. Fiji Islands ; Admiralty ; I specimen, no. 56.9.24.26. SYNONYMY. I agree with Hayashi (1938 and 19386) that Fromia andamanensis Koehler is better ranked as no more than a forma of F. indica distinguished by the similar dimensions of the abactinal plates as opposed to forma indica with two series of enlarged plates along each arm. The material in the British Museum collections from the type-locality of andamanensis supports this synonymy. However, I do not agree that F. elegans is also a synonym, since Mr. James F. Clark has re- examined the holotype in the Museum of Comparative Zoology at my request and found that the granules of the actinal plates number seven to nine on a plate and are no higher than wide, differing from the very characteristic elongated granules or short blunt spines, usually in groups of about five, found in F. indica. My thanks are due to Mr. Clark for his observations. Both Domantay and Roxas (1938) and A. H. Clark (1949) have extended the geographical range of F. elegans from Torres Strait to the Philippines. Judging from the photographs given by the former, their specimens could well be elegans but the specimen from Banda Neira referred to elegans by Engel (1938) has the distinctive elongated armament of the actinal plates characteristic of F. indica. To the synonymy of F. indica must also be added Fromia tumida Bell. The syntypes are dried and have suffered some distortion so that the lower surface has become concave and the arms are probably unnaturally wide. The hollowed under side probably evoked Bell's erroneous statement that the adambulacral spines are smaller than the spines of the actinal plates. In fact the armament is just like that of F. indica with short spines on the actinal plates and slightly longer ones on the adambulacrals. Fromia nodosa sp. nov. (PI. 6, figs. 1-3) Ferdina Offreti (pt.) Koehler, 1910 : 143, 147, pi. xvi, figs. 4, 5 only. MATERIAL. Amirante Islands, W. Indian Ocean ; Gardiner collection ; the holo- type and one paratype. B.M. reg. no. 1907.7.1.95 and 96. South Nilandu, Maldive Islands ; Gardiner collection ; I specimen. Cambridge Museum. DIAGNOSIS. A species of Fromia with very conspicuous alternation in size and shape of the supero-marginal plates, the larger ones being abruptly convex and separated by less than their own width from the series of similarly enlarged and convex spaced carinal plates ; no crystal-bodies embedded in the surface of the plates. DESCRIPTION OF THE HOLOTYPE. R/r = 32 mm./7'5 mm- = 4'3/i ', br at base 7-5 mm. and at arm tip 3-5 mm. The body is flattened above and the arms taper evenly to blunt tips. i go A.M. CLARK The abactinal plates are of two distinct kinds, the majority being flat and poly- gonal while nine to eleven spaced carinal plates on each arm, starting with the primary radial plates, are enlarged, up to 2*75 mm. in transverse diameter (slightly less longitudinally), rounded in outline and markedly convex, standing out sharply from the surface though not exceeding I mm. in height. When denuded, the plates between the convex carinal ones are revealed as not part of the series, some being adradial in position though others may be midradial. All the plates are slightly notched at intervals peripherally to accommodate the single interstitial papular pores. There are four to six pores around each of the flatter plates but seven to nine round most of the enlarged ones. The surface of the denuded plates is almost smooth, there being no embedded crystal-bodies. The supero-marginals are similarly of two forms with almost perfect alternation, the larger ones equal to or even slightly exceeding the diameter of the enlarged carinal plates. The swollen supero-marginals alternate in position obliquely with the swollen carinals, the three series on each arm being separated by only single small flat plates for most of the arm length while from the tenth supero-marginal all three rows of swollen plates are partially in contact laterally or obliquely. There are fifteen supero-marginal plates in one series denuded, the first, second, fourth, sixth, eighth, tenth, twelfth, fourteenth and fifteenth being convex and dwarfing the intervening plates. The infero-marginals number sixteen and are all similar in form though gradually decreasing in size from a proximal breadth of c. 2 mm. There are two series of actinal plates, the outer consisting of ten plates and extend- ing to the fifth infero-marginal, two actinal plates corresponding almost exactly to one marginal. The inner row consists of 24 plates (in the series denuded) extending to the middle of the eleventh infero-marginal or 25 mm. distance from the mouth. All these plates are covered with a continuous coat of polygonal granules which are slightly larger on the swollen abactinal and supero-marginal plates, numbering c. 35/sq. mm. as opposed to c. 55/sq. mm. on the flatter plates. Adjoining many of the papular pores one or sometimes several of the granules are distinctly enlarged. There are about 46 adambulacral plates in each series. Four or five of the basal ones bear four furrow spines, the adoral one of these usually smaller and inset on the plate, like a thumb. The remaining plates, except at the very tip of the arm, each have three furrow spines. These are flattened and spatulate, measuring up to 0-75 mm. in length. Separated from the furrow spines by a distinct gap come two subambulacral spines, which are about two-thirds as long as the furrow spines but slightly stouter. Between the twenty-fifth and thirtieth adambulacral plates the number of subambulacral spines falls to one. The outer part of each plate is covered with small low polygonal granules similar to those on the actinal plates and abruptly contrasting with the projecting subambulacral spines. There are two series of single actinal papulae corresponding in extent to the actinal plates. PARATYPES. The second specimen from the Amirantes has R/r = 19 mm./5 mm, = 3-8/1 and closely resembles the holotype except that the arms maintain the NOTES ON ASTEROIDS: OPHIDI ASTERIDS 191 same width for about two-thirds of their length. Also the enlarged supero-marginal plates are distinctly bigger than the enlarged carinals. The second series of actinal plates consists of only one or two plates. There are only about five actinal papulae between the proximal infero-marginals and the main series of actinal plates. The Maldive specimen (PI. 6, fig. 3) has R/r 34 mm./8-5 mm. = 4-0/1 ; br is 9-5 mm. basally but only 2-5 mm. at the tip, the arms tapering markedly. The swollen carinal and supero-marginal plates, particularly the latter, are even larger than in the holotype, producing a very mosaic-like effect, the low interstitial plates occupying little area. The largest supero-marginals are as much as 4-0 mm. in breadth and 4'5 mm. in length. Again there is a distinct enlargement of one or more granules near many of the papular pores and there are two rows of actinal papulae ; in one infero-marginal series there are 17 plates. REMARKS. The specimen with R 13-5 mm. from Ceylon thought by Koehler (1910) to be the immature fully granule-covered form of his Ferdina offreti is probably referable to Fromia nodosa. The holotype of Ferdina offreti (R 37-38 mm.) from the Andaman Islands likewise has enlarged swollen supero-marginal and carinal plates but these are completely bare and contrast conspicuously in colour with the granular areas. Livingstone (1931) has since referred offreti to Neoferdina, which he characterized by the presence of some bare enlarged aboral and marginal plates, as opposed to Ferdina sensu stricto with complete granulation even though some of the plates may be convex. Ferdina heffernani Livingstone (referred below to a new genus) has a superficial resemblance to Fromia nodosa but differs in the single row of adambulacral spines and the absence of papulae from the under side. Since these two features also apply to Ferdina I have no doubt that nodosa is most appropri- ately placed in Fromia. Nevertheless it does provide an interesting link between these genera. Only two other species of Fromia agree with F. nodosa in having a tendency for alternation in size and more or less marked convexity of the supero-marginal plates, namely Fromia monilis Perrier and F. ghardaqana Mortensen. The latter particu- larly may also have some slightly convex carinal plates but nowhere does the size and degree of projection of these enlarged plates approach the condition found in F. nodosa where these plates are separated laterally from each other by less than their own width, while in the distal third of the arm they are partially in contact with each other. In Fromia nodosa too the surface of the denuded plates is almost smooth, whereas in the other two species there are numerous glassy knobs or crystal-bodies embedded in them. Among the other species of Fromia, crystal-bodies are present in F. milleporella, armata and balansae but absent in indica and elegans ; their occurrence in F. hemiopla, hadracantha and pacifica is unknown. FERDINA Gray Ferdina Gray, 1840 : 282 ; 1866 : 12-13 ; Fisher, 1919 : 370 ; H. L. Clark, 1921 : 58 ; Living stone, 1931 : 305. TYPE SPECIES, Ferdina flavescens Gray, 1840 ; designated by Fisher, 1919. 192 A. M. CLARK REMARKS. In 1931 Livingstone restricted Ferdina by splitting off F. cumingi Gray and several other species with some conspicuously naked plates as a new genus Neoferdina. At the same time he re-diagnosed Ferdina itself and added a new species, Ferdina heffernani. Unfortunately his generic diagnosis appears to have been based more on the latter than on the type-species, which he knew only from photo- graphs of the holotype and the very inadequate published descriptions. Re- examination of the holotype of F. flavescens (PI. 6, fig. 6) and the removal of the aboral granulation of one arm reveals that the skeleton is of similar form to that of Gomophia and some species of Nardoa, with larger abactinal plates placed at irregular intervals and superimposed on a reticulum of smaller plates leaving meshes of sufficient size to accommodate small groups of slightly spaced papulae. Probably because the specimen has R only 26-28 mm. the number of pores in each area is small, mostly two to four, which may form linear series around the larger plates so that the grouping is not obvious. Nevertheless, some grouping can be seen even with the granulation intact, this showing up both in the photographs of the holotype published by Livingstone and in de Loriol's drawings of a specimen from Mauritius, the type-locality. This is nothing like the condition in F. heffernani, which resembles Fromia in its polygonal close-fitting pavement of abactinal plates with isolated papulae at the angles. In addition, in flavescens the supero-marginals are indistinguishable from the abactinal plates on the one arm cleared, only a few large ones near the arm tip being arranged in a linear series. This may also be the case in some larger specimens of Gomophia egyptiaca where the degree of specialization of the supero-marginals is variable. Another unusual feature of Ferdina flavescens is that the actinal and adambulacral plates are embedded in tissue so that the uniform granulation covering them is almost completely smooth and conceals the limits of the underlying plates. In F. heffernani these plates are much more easily dislodged when the skin is dissolved by bleach and their presence is evident without denuding due to their contours and to the coarsening of the granulation in the middle of each one. Finally, the armament of the adambulacral plates differs from that of F. heffernani in that the even granulation of the lower surface runs up the outer side of the single row of furrow spines, having shrunk in drying, leaving only the tips of the spines projecting. This produces somewhat the same effect as found in Linckia where the furrow spines appear to be sunk into the furrow. However, in Linckia the tips of the furrow spines hardly project at all from the surface. Possibly the same will prove to be true of live and spirit specimens of Ferdina flavescens, as opposed to dried ones. Thus, although F. flavescens appears to agree with heffernani in having complete granulation, no actinal papulae and only a single series of adambulacral spines, in fact the two species are not at all closely related and are certainly not congeneric. It remains then to re-diagnose Ferdina and to establish a new genus to accommodate F. heffernani. DIAGNOSIS. A genus of Ophidiasteridae with the aboral skeleton consisting of NOTES ON A STEROID S:OPHIDIASTERIDS 193 irregularly-placed larger plates superimposed on a reticulum of smaller ones leaving space between for small papular areas with poorly-integrated groups of usually two to four pores in each ; the limits of the underlying plates concealed by continuous even granulation ; supero-marginal plates irregular and only distinguishable from the abactinal plates near the tips of the arms ; actinal and adambulacral plates embedded in soft tissue so that their limits are concealed by the uniform granulation covering them ; no actinal papulae ; adambulacral plates armed with only a single row of short blunt furrow spines, only the outer halves of which project from the continuous granulation of the under side. AFFINITIES. Ferdina thus becomes a monotypic genus. It seems to occupy an isolated position within the family but may have some affinity with Gomophia. CELERINA gen. nov. TYPE-SPECIES. Ferdina heffernanii Livingstone, 1931. DIAGNOSIS. A genus of Ophidiasteridae with polygonal aboral plates, irregular in size, convexity and arrangement, having isolated papulae at the angles between ; supero-marginal plates tending to alternate large and convex with small and flat ; actinal plates not embedded in tissue ; granulation continuous all over, coarser on the centres of the actinal plates ; no actinal papulae ; a single row of short but projecting furrow spines. Celerina heffernani (Livingstone) (PI. 6, figs. 4 and 5) Ferdina heffernanii Livingstone, 1931 : 306-307, pi. xxiv, figs. 1-5. MATERIAL. Macclesfield Bank, South China Sea ; 40-55 metres ; Admiralty ; i specimen, B.M. reg. no. 93.8.25.90. This specimen was named Fromia milleporella by Bell. DESCRIPTION. R/r — 29 mm./6 mm. = 4-8/1 ; br basally = 6-5 mm. The abac- tinal plates are mostly about i-o mm. in diameter and are irregular in arrangement, even the carinal series being indistinct proximally, though four to seven spaced carinal plates in the distal half of each arm are enlarged up to c. 1-5 mm. diameter and are slightly convex. At the base of the arm there are three or four plates across the width. The supero-marginal plates number about 17 in each series and exhibit a fairly regular alternation of large convex plates, up to 2-0 mm. in breadth and 2-4 mm. in length, with small flat plates in between. Consequently, the upper side superficially has considerable resemblance to that of Fromia monilis. The whole aboral surface is covered with a close coat of very fine granules, c. I5o/sq. mm. These are very slightly coarser on the convexities of the skeleton but much less so than in the holotype of F. heffernani. However, on the distal marginal plates, especially the infero-marginal ones, the granulation is distinctly coarser centrally. There are about 19 similar infero-marginals in each series, decreasing gradually 194 A. M. CLARK in size distally. In each arm angle there are about six narrow intermarginal plates, three each side of the inter-radius. The main series of actinal plates numbers about 20 with an odd inter-radial plate adjoining each pair of oral plates. The granulation of these is distinctly coarser than the aboral granulation and in the centre of each actinal plate one to four granules are both broader and higher than the rest. Similarly the adambulacral plates, after the basal three or four, have one or more enlarged granules in the centre, a single one on the distal plates becoming increasingly prominent though without reaching the eminence of a spine. Along the furrow margin runs a single series of small sharply-pointed spines up to 0-6 mm. long. On the first 15-18 plates these usually number three ; the following plates normally bear two furrow spines. There are single papular pores between the angles of the abactinal plates and intermarginally but none on the lower surface. REMARKS. The holotype of Ferdina heffernani from the Santa Cruz Islands (east of the Solomons) has R4omm., which may account for its more numerous enlarged abactinal plates and the coarser granules on the centres of these ; however, Livingstone counted only two furrow spines, despite the larger size. He made no mention of intermarginal plates but these could easily be overlooked if they are as inconspicuous as in the British Museum specimen. Tamaria tumescens (Koehler) Linckia megaloplax (pt.) Bell, 1894 : 395. Ophidiaster tumescens Koehler, igioa : 281-283, P^ xyi. fig5- 3» 4- Tamaria tumescens : H. L. Clark, 1921 : 94 ; 1938 : 141 ; 1946 : 123-124. Tamaria propetumescens Livingstone, 19320; : 369-371, fig. i, pi. xlii, figs. 1-5. Tamaria ajax Livingstone, i932« : 371-372, fig. 2, pi. xli, figs. 1-5. MATERIAL. Parry Shoal, Arafura Sea ; 22 metres ; Admiralty ; 4 specimens including the holotype of T. propetumescens Livingstone, B.M. reg. no. 92.4.4.2-5. i specimen, no. 92.4.4.27. Holothuria Bank, northern Australia ; 27 metres ; Admiralty ; i specimen, no. 91.8.26.83. 2 specimens, no. 92.1.14.326-327. Broome, N.W. Australia ; presented by Museum of Comparative Zoology, Har- vard ; i specimen, no. 1939.6.15.65. Dammer I., Banda Sea ; 16-27 metres ; Admiralty ; the holotype of T. ajax Livingstone, no. 92.4.4.24. SYNONYMY. Unfortunately Livingstone had only photographs upon which to base his two nominal species, Tamaria propetumescens and ajax, and was unable to handle personally these and the other relevant specimens in the British Museum collections. The five individuals from Parry Shoal, including the holotype of T. propetumescens, show a range of variation in the armament of the central parts of the convex abactinal plates from almost as fine as the peripheral granules (as in the type of T. propetumescens} through somewhat coarser granulation (as in the type of tumescens) to markedly coarser with often a single enlarged flattened or conical tubercle (as in the type of ajax from the Banda Sea). H. L. Clark (1938) has already referred T. propetumescens to the synonymy of T, tumescens and I NOTES ON ASTEROIDS: OPHIDI ASTERIDS 195 consider that the holotype of T. ajax also comes within the range of variation of the species. Livingstone distinguished T. propetumescens from the holotype of tumescens by the more obvious demarcation of the marginal plates, the greater extent proximally of the subambulacral spines, their separation from the furrow spines by granulation and by the development of stout infero-marginal tubercles in the type of propetu- mescens. H. L. Clark found the infero-marginal armament to be very variable, while the marked separation of the marginal plates is due to shrinkage in the drying of Livingstone's type. Also I suspect that Koehler's artist omitted to draw the granules between the furrow and subambulacral spines since these are present in the other specimens in the British Museum collections but may be inconspicuous owing to contraction of the adambulacral plates. The extent and regularity of the sub- ambulacral spines proximally are variable in other species of Tamaria. As for Tamaria ajax, Livingstone believed this to be related to T. triseriata (Fisher) from the Hawaiian Islands and did not compare it with T. tumescens. However, it shares the distinctive broadened carinal row of plates so characteristic of tumescens with which the armament is much more closely comparable than is that of triseriata. The holotype of ajax is in spirit, which accounts for its smoother appearance in photographs than the type of propetumescens. The apparent difference in the pedicellariae of the two shown by C. C. A. Monro's drawings in Livingstone's paper is not constant, the valves being equally short and with inconspicuous teeth all round the expanded end in some pedicellariae of the type of propetumescens as in the one shown for ajax. Ophidiaster helicostichus Sladen Linckia nodosa : Bell, 1884 : 124 [Non L. nodosa Perrier, 1875, which is a synonym of L. bouvieri.] Ophidiaster helicostichus Sladen, 1889 : 405-407, pi. Ixix, figs. 5-7. Hacelia helicostichus : H. L. Clark, 1909 : in ; 1921 : 86 ; 1938 : 139 ; 1946 ; 122-123. Tamaria sp. Livingstone, 1932 : 261-262 (at least the specimen in pi. xi, figs. 5, 6). ^Ophidiaster astridae Engel, 1938 : 12-14, figs. 2-4, pi. iii, fig. 3. MATERIAL. " Challenger " station 187, Booby L, Torres Strait ; n metres ; the holotype, B.M. reg. no. 90.5.7.618. Cape York, N. Queensland ; Mr. Darnel ; i specimen, no. 67.5.27.1. Prince of Wales Channel, Torres Strait ; 13 metres ; Admiralty (H. M.S." Alert"); 2 specimens, no. 82.2.22.22 and 23. N.W. Australia; Admiralty ; I specimen, no. 92.1.14.76. Holothuria Bank, northern Australia ; Admiralty (H.M.S. " Penguin ") ; I specimen, no. 92.1.14.69. Holothuria Bank ; 40 metres ; i specimen, no. 92.1.14.238. Port Molle, Queensland ; 26 metres ; Admiralty (H.M.S. " Alert ") ; i specimen, no. 81.10.26.125-127. Port Molle ; presented by Sydney Museum ; i specimen, no. 83.12.9.88. AFFINITIES. In 1909, following receipt of a British Museum specimen from Prince of Wales Channel (" Alert "), H. L. Clark commented " There can be no ig6 A. M. CLARK question, I think, that this species is congeneric with Hacelia attenuata Gray from the Mediterranean ". In his key to the genera of Ophidiasteridae (1921) he dis- tinguished Hacelia as having ten series of pore-areas on each arm, the lowest one each side situated between the adjacent rows of actinal plates and so more numerous than the others. This is true enough of H. attenuata and evidently also of Koehler's variety inarmatus of helicostichus (1885) from the Sunda Islands, which was raised to specific rank by H. L. Clark (1921). However, in 1938 Dr. Clark referred another Australian specimen to helicostichus and noted that " the actinal papulae are poorly developed and only here and there at isolated spots can one count ten (or even nine!) longitudinal series of papulae". There are nine specimens of helicostichus in the British Museum collections, six of them with R > 100 mm., including two from the same station as the one sent to H. L. Clark in 1909. In none of these can I find more than eight series of papular areas, the lowest of which are between the infero-marginal plates and the outermost series of actinals. Similarly in 1946, D. D. John of this museum could find no interactinal papulae in the six larger specimens when he examined them at the request of Dr. Engel. Nothing appears to have been published by Engel concerning this but in 1938 he had described a large sea-star (R up to 220 mm.) from the East Indies as Ophidiaster astridae. This is clearly closely related to, if not conspecific with, helicostichus ; his description and figures bring out no way in which the two can be distinguished. In view of this, I consider that there are insufficient grounds for treating heli- costichus as congeneric with Hacelia attenuata and accordingly refer it back to Ophidiaster. It differs from the Mediterranean and Atlantic type-species, 0. ophi- dianus, in having more numerous series of actinal plates, about four proximally when R = 100-150 mm., rather than two, and the area covered by these plates is correspondingly more extensive. However, several of the Indo-Pacific species of Ophidiaster, such as 0. hemprichi, approximate to helicostichus in the number and extent of the actinal plates, though they do not reach nearly such a large size. Immature specimens of 0. helicostichus (R < c. 70 mm.) with the lowest series of pore-areas below the infero-marginals barely developed or even lacking, may be confused with Tamaria but can be distinguished from the Australian species of that genus by the more attenuated arms. The specimen with R 103-5 mm- from Port Curtis, Queensland, shown in Livingstone's pi. xi, figs. 5 and 6 (1932) under the caption of " Tamaria sp. " is almost certainly conspecific with helicostichus, resemb- ling closely the two specimens from Port Molle, Queensland (R 50-57 mm.) and a smaller one (R36mm.) from Holothuria Bank, which I have referred to this species. The incipient pore-areas below the infero-marginals (which prompted Livingstone's doubts about the validity of Tamaria as a genus distinct from Ophi- diaster}, the faint grooving of the furrow spines and their alternation in size (though this is admittedly a general tendency in Tamaria) as well as the form of the skeleton with its even armament of granules, are all as would be expected in helicostichus at this size. NOTES ON ASTEROIDS: OPH IDI ASTERI DS 197 REFERENCES BELL, F. J. 1882. Note on the Echinoderm-Fauna of the Island of Ceylon, together with some observations on Heteractinism. Ann. Mag. nat. Hist. (5) 10 : 218-225. - 18820. Description of new or rare species of Asteroidea in the collection of the British Museum. Proc. zool. Soc. Lond. 1882 : 121-124, pi- yi- - 1884. [In] Report on the zoological collections made in the Indo-Pacific Ocean during the voyage of H.M.S. " Alert ", 1881-2. pp. 117-177 & 509-512, pis. viii-xvii & xlv. London. - 1887. The Echinoderm fauna of the island of Ceylon. Scient. Trans. R. Dubl. Soc. (2) 3 (14) : 643-658, 2 pis. — 1894. On the Echinoderms collected during the Voyage of H.M.S. " Penguin " and H.M.S. " Egeria ", when surveying Macclesfield Bank. Proc. zool. Soc. Lond. 1894 : 392- 413, pis. 23-27. - 1902. The Actinogonidiate Echinoderms of the Maldive and Laccadive Islands. [In] Gardiner, J. S. The Fauna and Geography of the Maldive and Laccadive Archipelagoes. vol. I: 223-233. Cambridge (University Press). - 1909. Report on the Echinoderma (other than Holothurians) collected by Mr. J. Stanley Gardiner in the western parts of the Indian Ocean. Trans. Linn. Soc. Lond. Zool. (2) 13 (i) : 17-22, pi. 3. BRUGIERE, M. 1791-1827. Encyclopedic Methodique. Zoologie 7. Vers, Coquilles, Mol- lusques, Polypiers. Tableaux, vii -f 180 pp., 488 pis., 3 vols. Paris. CLARK, A. H. 1949. On a collection of sea-stars from the Philippine Islands. Proc. biol. Soc. Wash. 62 : 73-78. CLARK, A. M. 1952. The " Manihine " Expedition to the Gulf of Aqaba, 1948-1949. VII. Echinodermata. Bull. Br. Mus. nat. Hist., (Zool). 1 : 203-214, pis. xxxi-xxxii. CLARK, A. M. & DAVIES, P. SPENCER. 1966. Echinoderms of the Maldive Islands. Ann. Mag. nat. Hist. (13) 8: 597-612, pi. xviii. CLARK, H. L. 1909. Notes on some Australian and Indo-Pacific Echinoderms. Bull. Mus. comp. Zool. Harv. 52 : 107-135, i pi. — 1921. The Echinoderm fauna of Torres Strait : its composition and its origin. Pap. Dept mar. Biol. Carnegie Instn. Wash. 10 : viii + 223, 38 pis. - 1938. Echinoderms from Australia. Mem. Mus. comp. Zool. Harv. 55 : viii + 596, 63 figs., 28 pis. - 1946. The Echinoderm Fauna of Australia. Publ. Carnegie Instn 566 : 1-567. DODERLEIN, L. 1926. Uber Asteriden aus dem Museum von Stockholm. K. svensk. Vetensk- Akad. Handl. (3) 2 (6) : 1-22, 4 pis. DOMANTAY, J. S. & ROXAS, H. A. 1938. The littoral Asteroidea of Port Galera Bay and adjacent waters. Philipp. J. Sci. 65 (3) : 203-237, 17 pis. ENDEAN, R. 1965. Queensland Faunistic Records. Part VIII. Further records of Echino- dermata (excluding Crinoidea) from southern Queensland. Pap. Dep. Zool. Univ. Qd. 2 : 227-235. ENGEL, H. 1938. Asteries et Ophiures. [In] Resultats scientifiques du voyage aux Orient- ale Neerlandaises de le Prince et la Princesse Leopold de Belgique. Mem. Mus. r. Hist, nat. Belg. 3 (18) : 1-31, 5 figs., 4 pis. FISHER, W. K. 1917. New starfishes from the Philippines and Celebes. Proc. biol. Soc. Wash. 30 : 89-93. - 1919. Starfishes of the Philippine Seas and adjacent waters. Bull. U.S. natn. Mus. 100 (3) : 1-546, 156 pis. GRAY, J. E. 1840. A synopsis of the genera and species of the class Hypostoma (Asterias, Linn.) Ann. Mag. nat. Hist. 6 : 175-184 ; 275-290. — 1866. Synopsis of the species of starfish in the British Museum (with figures of some of the new species), iv + 17 pp., pis. 1-16. London. HAYASAKA, I. 1949. On some starfishes from Taiwan. Bull, oceanogr. Inst. Taiwan. No. 5: 11-19, pis. i-iii. ig8 A. M. CLARK HAYASHI, R. 1938. Sea-stars of the Caroline Islands. Palao trop. biol. Stn Stud. 3 : 417-446, 5 figs., pis. 2-4. — iQ38a. Sea-stars of the Ryukyu Islands. Bull, biogeogr. Soc. Japan 8 (14) : 197-222, i fig., 2 pis. — 19386. Sea-stars of the Ogasawara Islands. Annotnes zool. jap. 17 (i) : 59-68, 5 figs., i pi. — i938c. Sea-stars in the vicinity of the Seto Marine Biological Laboratory. Bull, bio- geogr. Soc. Japan 8 (19) : 271-292, 4 figs., 3 pis. KOEHLER, R. 1895. Catalogue raisonne des Echinodermes recueillis par M. Korotnev aux iles de la Sonde. Mem. Soc. zool. Fr. 8 : 374-423, pi. ix. — 1909. Echinoderma of the Indian Museum. Part V. An account of the deep-sea Aster- oidea collected by the R.I. M.S. Investigator. 143 pp., 13 pis., Calcutta. — 1910. Echinoderma of the Indian Museum. Part VI. An account of the shallow-water Asteroidea. 192 pp., 20 pis. Calcutta. — igioa. Asteries et Ophiures des iles Aru et Kei. Abh. senckenb. naturforsch. Ges. 33 : 265-295, pis. xv-xvii. LAMARCK, J. B. P. A. DE. 1816. Les Stellerides. Histoire naturelle des animaux sans vertebres. Ed. i. vol. II, pp. 528-568. Paris. LINCK, J. H. 1733. De Stellis Marinis. xxiv + 107 pp., 42 pis. Lipsiae. LIVINGSTONE, A. A. 1930. On some new and little-known Australian Asteroids. Rec. Aust. Mus. 18 (i) : 15-24, 5 pis. — 1931. On the restriction of the genus Ferdina Gray. Aust. Zool. 6 : 305-309, pis. xxi- xxiv. — 1932. Asteroidea. Sci. Rep. Gr. Barrier Reef Exped. 4 : 241-265, 2 figs., 12 pis. — 1932^. Some further notes on species of Tamaria. Rec. Aust. Mus. 18 : 368-372, 2 figs., pis. xl-xlii. 1936. Descriptions of new Asteroidea from the Pacific. Rec. Aust. Mus. 19 : 383-397, 2 pis. LORIOL, P. DE. 1885. Catalogue raisonn6 des Echinodermes recueillis par M. V. de Robillard a File Maurice. II. Stellerides. Mem. Soc. Phys. Hist. nat. Geneve 29 (4) : 1-84, pis. vii-xxii. — 1891. Notes pour servir a 1'etude des Echinodermes. Mem. Soc. Phys. Hist. nat. Geneve vol. suppl. 1891 (8) : 1-31, 3 pis. LUTKEN, C. F. 1865. Kritiske Bemaerkninger om forskellige Sostjerner (Asterider), med Beskrivelse af nogle nye Arter. Vidensk. Meddr dansk. naturh. Foren. 1864 : 123-169. MARTENS, E. VON. 1866. Ueber ostasiatische Echinodermen. Arch. Naturgesch. 32 : 57-88, 133-189. MICHELIN, H. 1845. Zoophytes, Echinodermes et Stellerides de 1'ile Maurice. Magasin Zool., Paris (2) 1845 Zoophytes: 1-27, pis. 7-12. MOBIUS, K. A. 1880. Beitrdge zur Meeresfauna der Inseln Mauritius und der Seychellen. 352 pp., pis. i-xxii. Berlin. MULLER, J. & TROSCHEL, F. H. 1842. System der Asteriden. i Asteriae. 2 Ophiuridae. xx -f 134 PP-, 12 pis. Braunschweig. NARDO, J. D. 1934. De Asteriis. Isis, Jena. Encyclopaedische Zeitung: 716-717. PERRIER, E. 1869. Recherches sur les Pedicellaires et les Ambulacres des Asteries et des Our sins. 188 pp., 2 pis. Paris [Also published in : Annls Sci. nat. 12 (1869) : 197-304 ; 13 (1870) : 1-81]. — 1875. Revision de la collection de Stellerides du Museum d'Histoire Naturelle de Paris. 384 pp. Paris. [Also published in : Archs. Zool. exp. gen. 4 (1875) : 265-450, 5 (1876) : 1-104, 209-309]. PETERS, W. 1852. Ubersicht der Seesterne (Asteridae) von Mossambique. Mber. K. preuss. Akad. Wiss. 1852 : 177-178. RETZIUS, A. J. 1805. Dissertatio sistens species cognitas Asteriarum. 37 pp. Lundae. SLADEN, W. P. 1889. Asteroidea. Rep. scient. Res.Voy. Challenger, Zool. 30 : 1-935, JI8 pis. TORTONESE, E. 1955. Notes on Asteroidea. Ann. Mag. nat. Hist. (12) 8 : 675-684. ZOOL. 15, 4 PLATE i FIGS. 1-3. Gomophia egyptiaca egeriae subsp. nov., holotype, B.M. reg. no. 92.8.22.52, aboral (i) and oral (2) views, with detail of denuded arm (3). FIGS. 4, 5. The same, paratypes, no. 92.8.22.53 and 259, aboral views ; all from Maccles- field Bank. FIG. 6. Gomophia egyptiaca egyptiaca Gray, no. 98.10.17.12, Christmas I., Indian Ocean, aboral view. FIGS. 7, 8. The same, no. 67 . i . 7 . 42, Samoa, side view of denuded arm (7) and aboral view (8), in (7) the lowest series of plates is the infero-marginal one. FIG. 9. Nardoa frianti Koehler, no. 92.8.22.264, Macclesfield Bank, aboral view. FIG. 10. The same, no. 98.8.8.53, Loyalty Is., aboral view. All approximately x f except (2) x 2^ and (7) x i£. Bull. Br. Mus. nat. Hist. (Zool.) 15, 4 PLATE i ZOOL. 15, 4 PLATE 2 FIGS. 1-4. Nardoa gomophia (Perrier), holotype, B.M. reg. no. 59.4.25.3, New Caledonia, aboral (i) and oral (2) views, both x f, detail of denuded arm (3), x 2\ and side view of arm (4), X li. Bull. Br. Mus. nat. Hist. (Zool.) 15, 4 PLATE 2 ZOOL. 15, 4 PLATE 3 FIGS. 1-3. Nardoa sphenisci sp. nov., holotype, B.M. reg. no. 92.1.14.27, Holothuria Bank, N.W. Australia, aboral (i) and oral (2) views, both x f and side view of partly denuded arm (3), x 2. Rull. Br. Mus. nat. Hist. (Zool.) 15, 4 PLATE 3 PLATE 4 FIG. i. Nardoa tubevculata Gray, holotype, B.M. reg. no. 1938.6.23.4, Sual, Philippine Is., aboral view, x f . FIGS. 2, 3. The same, holotype of Scytaster obtusus Perrier, no. 43.3.29.6, Siguijor, Philip- pine Is., aboral view (2), x f and side view of arm (3), x i|. FIG. 4. Nardoa sp. aff. tuber culata, no. 1949.2.14.16, Batjan, Molucca Is., aboral view, x f. Bull. Br. Mus. nat. Hist. (Zool.) 15, 4 PLATE 4 PLATE 5 FIG. i. Nardoa novaecaledoniae (Perrier), Oslo Museum collection, New Caledonia, aboral view, x f . FIG. 2. The same, B.M. reg. no. 59.4.25.4, New Caledonia, previously labelled as holotype of N. gomophia, aboral view, x f . FIG. 3. Nardoa rosea H. L. Clark, paratype, no. 1966.7.14.1, Mer, Torres Strait, side view of the single arm, x ij. FIG. 4. The same, paratype, Museum of Comparative Zoology, Harvard collection, no. 2326, Mer, detail of partly denuded arm, X ij. Bull. Br. Mus. nat. Hist. (Zool.) 15,4 PLATE 5 PLATE 6 FIGS, i, 2. Fromia nodosa sp. nov., holotype, B.M. reg. no. 1907.7.1.95, Amirante Is., aboral (i) and oral (2) views, both x i£. FIG. 3. The same, Cambridge Museum collection, S. Nilandu, Maldive Is., aboral view, x f . FIGS. 4, 5. Celerina heffernani (Livingstone), no. 93.8.25.90, Macclesfield Bank, S. China Sea, aboral view (4), x i£ and detail of denuded arm (5), x 2§. FIG. 6. Ferdina flavescens Gray, holotype, no. 1938.6.23.7, Mauritius, detail of denuded arm and part of disc, x ^\. Bull. fir. Mus. nat. Hist. (Zool.) 15, 4 PLATE 6 PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING NEW SPECIES OF STELLICOLA (COPEPODA, CYCLOPOIDA) ASSOCIATED WITH STARFISHES IN MADAGASCAR, WITH A REDESCRIPTION OF S. CAERULEUS (STEBBING, 1900) ARTHUR G. HUMES and JU-SHEY HO BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 15 No. 5 LONDON: 1967 oSV NEW SPECIES OF STELLICOLA (COPEPODA, % CYCLOPOIDA) ASSOCIATED WITH STARFISHES IN MADAGASCAR, WITH A REDESCRIPTION OF S. CAERULEUS (STEBBING, 1900) BY ARTHUR G. HUMES and TU-SHEY HO v J ' VAj Boston University, Boston, Massachusetts, _U.S. A. Pp. 199-225 ; 17 Plates BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 15 No. 5 LONDON: 1967 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the 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. 15, No. 5 of the Zoological 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.). Trustees of the British Museum (Natural History) 1967 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 16 May, 1967 Price £i 4$. NEW SPECIES OF STELLICOLA (COPEPODA, CYCLOPOIDA) ASSOCIATED WITH STARFISHES IN MADAGASCAR, WITH A REDESCRIPTION OF S. CAERULEUS (STEBBING, 1900) By ARTHUR G. HUMES & JU-SHEY HO INTRODUCTION Two species of the lichomolgid genus Stellicola (a genus usually living in association with asteroid echinoderms) are known from starfishes in the region of Nosy Be, in northwestern Madagascar. 5. oreastriphilus Kossmann, 1877, occurs commonly on Protoreaster lincki (Blainville) , Culcita schmideliana (Retzius), Pentacer aster mammil- latus (Audouin), and Pot 'aster superbus (Mobius) (Humes & Cressey, 1961), and more rarely on Choriaster granulatus (Liitken) (Humes & Ho, in press). A second species of Stellicola, whose description by Humes and Ho is in press, also occurs on Choriaster granulatus. Five new species from the region of Nosy Be are now to be added, bringing the total number of species in that area to seven. The field work was done by the first author, in 1960 while a member of an expedi- tion sent to Madagascar by the Academy of Natural Sciences of Philadelphia, and in 1963-64 while participating in the U.S. Program in Biology of the International Indian Ocean Expedition. The study of the material has been aided by a grant (GB-iSog) from the National Science Foundation of the U.S. We wish to thank Dr. H. Barraclough Fell and Mr. James F. Clark of the Museum of Comparative Zoology, Harvard University, for the identifications of the star- fishes collected in 1963-64 (with the exception of the Retaster cribrosus from Ankify which was determined by Dr. G. Cherbonnier of the Museum National d'Histoire Naturelle in Paris) and Dr. Elisabeth Deichmann, also of the Museum of Comparative Zoology, for determining the starfishes collected in 1960. It is a pleasure also to acknowledge the assistance given to the field work by the staff at the Centre d'Oceano- graphie et des Peches et Nosy Be during the eighteen months while the first author was resident there. We are also indebted to Dr. J. P. Harding of the British Museum (Natural History) for information on the whereabouts of Thompson and A. Scott's Ceylonese copepods and to Miss Patricia D. Lofthouse of the same museum for arranging the loan of syntypes of 5. caeruleus with permission to dissect a male and a female. The material covered in this paper comprises : Stellicola kossmanni, new species Stellicola affinis, new species Stellicola longiseta, new species Stellicola femineus , new species Stellicola pollex, new species Stellicola oreastriphilus Kossmann, redescription in part Stellicola caeruleus (Stebbing), redescription of syntypes ZOOL. 15, 5. 16 202 ARTHUR G. HUMES & JU-SHEY HO STELLICOLA Kossmann, 1877 Stellicola kossmanni n. sp. PL i, figs. 1-6; PL 2, figs. 7-15; PL 3, figs. 16-21; PL 4, figs. 22-24. TYPE MATERIAL. 4 $ and 14 <$ from washings of 83 starfishes, Protoreaster lincki (Blainville) , in a depth of 0-5 m. at Pte. Antsamantsara, west of Madirokely, Nosy Be, Madagascar. Collected 7th July, 1963. Holotype female, allotype, and 13 paratypes (2 $, n <$) deposited in the British Mus. (Nat. Hist.) Reg. No. 1966.12.7.1-3; the remaining 3 paratypes (dissected) in the collection of A. G. Humes. Other specimens (all from Protoreaster lincki in 0-5 m.). 6 $, 6 $ from 7 hosts, Pte. Mahatsinjo, Nosy Be, 4th July, 1963 ; i $, 6 $ from 60 hosts at Pte. Lokobe, Nosy Be, nth July, 1963. FEMALE. Body (PL i, fig. i) with moderately broadened prosome. Length (not including setae on caudal rami) i-o mm. (0-95-1-03 mm.) and greatest width 0-46 mm. (0-44-0-47 mm.), based on 10 specimens (holotype, 3 paratypes, and 6 specimens from Pte. Mahatsinjo, 4th July, 1963). Ratio of length to width of prosome 1-45 : i. Segment of leg I separated dorsally and laterally from head by a furrow. Epimeral areas of segments of legs 1-4 rounded ; lateral margins of segments of legs 1-3 with membrane (PL i, fig. 2). Segment of leg 5 (PL i, fig. 3) 83 ju, long and 105 fi in greatest width ; fifth legs borne slightly dorsally. Between this segment and genital segment a short inter- segmental sclerite ventrally. Genital segment longer than wide, 146 fi in length, 127 fji in width in its anterior three-fourths where its lateral margins are somewhat irregular, and 85 /* in width in its constricted posterior fourth. Areas of attachment of egg sacs situated dorsolaterally just in front of posteriormost fourth ; each area (PL I, fig. 4) with 2 long setae (outer 82 /* long and bearing minute unilateral spinules, inner 36 /£ and feathered) and between them a small spinous process 6 fi long. Posterior margin of segment with a membrane having a ragged edge simulating small spinules especially laterally. Three postgenital segments, first 44 X 75 fi with a similar posterior membrane, second 18 X 68 fi, and third 28 X 65 /*. Caudal ramus (PL I, fig. 5), inserted ventrally on anal segment, quadrate, 31 x 29 /£. Long naked setule (34 /^) midway on outer margin. Pedicellate dorsal seta 65 fji and haired. Outer lateral seta 109 /£ and naked, inserted close to outer- most terminal seta (156 /£, with a few spinules on each side in its proximal third) . Innermost terminal seta 216 /£ and naked. Two long terminal setae, inserted between dorsal (smooth) and ventral (with a row of prominent setules) flaps, 429 /* (outer) and 600 /* (inner) and both with lateral spinules ; " pegged " region of these two setae finely punctate. A small hair on ventral surface of ramus. Dorsal surface of prosome with scattered hairs (sensilla). Dorsal surface of urosome with hairs as shown in PL I, fig. 3 ; ventral surface less ornamented with hairs. Ratio of length of prosome to that of urosome 1-87 : i. Egg sac (PL i, fig. i) elongated, 520 x 275 fi, extending far beyond caudal rami, and containing many eggs about 50 ju, in diameter. NEW SPECIES OF STELLICOLA IN MADAGASCAR 203 Rostral area (PI. I, fig. 6) rounded, not strongly sclerotized. First antenna (PL 2, fig. 7) 7-segmented, lengths of segments (measured along their posterior non-setiferous margins) being 39 (50 fi along anterior margin), 74, 31, 41, 34, 18, and 16 /* respectively. Formula for armature : 4, 13 (5 + 2 +6), 6, 3, 4 + i aesthete, 2 + 1 aesthete, and 7+1 aesthete. All setae naked except i on second segment and 4 on last 3 segments which are delicately haired. Terminal seta on segment 7 very long (260 /t), equal to length of entire first antenna. Aesthete on segment 7 arising from common base with adjacent long naked seta. Ventral surface of segment 3 with sclerotization suggesting an intercalary segment. Second antenna (PI. 2, fig. 8) relatively slender and 3-segmented, third segment showing a partial division on anterior surface but entire posteriorly. First segment with a smooth seta, second with a haired seta, and third with 3 haired setae near the incomplete division and terminally with 6 elements : a stout claw 42 (i along its axis, 4 long naked setae, and a short blunt naked seta. Anterior surface of second antenna pilose, with 4 slender setules subterminally on last segment. Long setae annulated. Labrum (PI. 2, fig. 9) with its two posteroventral lobes somewhat angular and unornamented. Mandible (PI. 2, fig. 10) with an outer row of slender spinules and an inner row of rather stout blunt teeth preceded proximally by a prominent dentiform process ; both rows diminishing to minute spinules along the slender terminal lash. Paragnath a small lobe hidden under lobe of labrum in ventral view. First maxilla (PI. 2, fig. n) an elongated lobe with 3 naked setae and i stout spiniform element strongly spinose on its posterolateral surface. Second maxilla (PI. 2, fig. 12) 2-segmented ; first segment unarmed, its proximal ventral margin projecting strongly ; second segment with 2 postero-inner elements (a slender seta with short lateral spinules and a spine with rows of spinules) and terminating in a spiniform process bearing rows of teeth as in the figure (proximal tooth larger than others, producing a bifur- cated appearance). Maxilliped (PI. 2, fig. 13) 3-segmented ; first segment unarmed, second with 2 setae and anterior surficial spinules, and third with 2 setae (i with lateral spinules, other naked) and terminating in a spiniform process bearing lateral spinules. Area between maxillipeds and first pair of legs (PI. 2, fig. 14) not protuberant, with a distinct sclerotized line between bases of maxillipeds. Kami of legs 1-4 (PI. 2, fig. 15, PI. 3, figs. 16, 17, and 18) 3-segmented, except endopod of leg 4 which is weakly 2-segmented. Spine and setal formula as follows (Roman numerals indicating spines, Arabic numerals setae) : PI protopod o-i ; i-o exp I-o ; I-i end o-i ; o-i ?2 protopod o-i ; i-o exp I-o ; I-i end o-i ; 0-2 PS protopod o-i ; i-o exp I-o ; I-i end o-i ; 0-2 ?4 protopod o-i ; i-o exp I-o ; I-i end o-i ; II, i III, 1,4 1,5 III, I, 5 I, II, 3 HI, I, 5 I, II, 2 II, I, 5 204 ARTHUR G. HUMES & JU-SHEY HO Inner seta on coxa feathered in legs 1-3 but naked (42 [i long) in leg 4. Hairs on inner margin of basis in legs 1-3 absent in leg 4. Expansion of basis over anterior surface of first segment of exopod rather acute. Between 2 terminal spines on last segment of endopod of leg 2 a bifurcated spinous process. On second segment of exopod of leg 4 relatively short inner seta lying (in alcoholized specimens) nearly parallel to ramus and having short lateral spinules instead of the usual long hairs. Endopod of leg 4 two-segmented, with a line of separation visible on both anterior and posterior surfaces, but with the lateral sclerotization continuous between the two segments. First segment 26 X 24 /i (greatest dimensions) with inner distal seta 84 fi long, spiniform, with short lateral spinules. Second segment 46 X 21 fi (greatest dimensions) , the feathered inner seta 67 fi, the 2 terminal spines 105 fi (inner) and 58 /t (outer) both with delicately serrate lamellae ; a row of minute spinules anteriorly across end of segment near insertions of 2 spines, and a row of hairs on proximal half of outer margin of segment. Leg 5 (PL 3, fig. 19) with free segment elongated, 32 x 14 /A. Two very unequal terminal setae, outer 102 ju long, with minute lateral spinules, inner 55 f.t, with delicately serrate lamellae. Seta on body near insertion of free segment 40 fi and feathered. Leg 6 probably represented by the 2 setae on area of attachment of egg sac (see PL i, figs. 3 and 4). Colour in life in transmitted light faintly reddish, eye red, intestine dark brown, ovary reddish brown. MALE. Body form (PL 3, fig. 20) resembling that of female, though prosome somewhat narrower. Length (excluding setae on caudal rami) 0-71 mm. (o-69-0'74 mm.) and greatest width 0-27 mm. (0-26-0-28 mm.), based on 10 specimens. Ratio of length to width of prosome 1-72 : i. Segment of leg 5 (PL 3, fig. 21) 33 X 70 jn. Ventral intersegmental sclerite absent. Genital segment about as long as wide, 99 X 94 /*, with slightly rounded lateral margins in dorsal view. Four postgenital segments 42 x 66, 34 X 58, n x 50, and 21 X 52 ju, from anterior to posterior. Caudal ramus like that of female but smaller, 23 x 23 jn. Surfaces of prosome and urosome, as in female, ornamented with hairs. Ratio of length of prosome to that of urosome 1-85 : i. Rostral area, first antenna, second antenna, labrum, mandible, paragnath, first maxilla, and second maxilla like those of female. Maxilliped (PL 4, fig. 22) slender and 4-segmented. First segment unarmed, second with 2 naked setae and 2 rows of spinules, and third small and unarmed. Proximal part of terminal claw probably representing fourth segment. Claw 101 ju, in length (measured along its axis), recurved, showing a slight evidence of division about midway and having a fringe of delicate hyaline spinules along its concave edge ; bearing 2 setae proximally, one small naked and hyaline 12 /i long, other long (50 ju,} with a few distal spinules. Area between maxillipeds and first pair of legs like that in female. Legs 1-4 resembling those of female, with same spine and setal formula. Endopod of leg 4 with 2 terminal spines differing in their proportional lengths. Measurements NEW SPECIES OF STELLICOLA IN MADAGASCAR 205 for this ramus : first segment 19 X 18 //• with seta 66 /*, second segment 33 X 17 /i with seta 45 fi and 2 terminal spines 80 fi and 47 /* (ratio of these 2 spines 1-7 : i, instead of 1-8 : i in female). Leg 5 resembling that of female, with free segment 21 X 8 [i and 2 terminal elements 70 /* and 34 ju, respectively. Leg 6 (PI. 4, fig. 23) consisting of posterolateral flap on ventral surface of genital segment and bearing a minute spiniform process and 2 unequal setae, one naked (69 fi), other feathered (33 /*). In view of entire urosome these setae conspicuous (see PL 3, fig. 21). Spermatophore (PL 4, fig. 24), inside body of male, elongated, 86 X 31 [i without neck. Colour in life in transmitted light resembling that of female. (This species is named for Dr. Robby Kossmann, who originally described the genus Stellicola}. COMPARISON WITH RELATED SPECIES. There are at present fifteen species known with some certainty to belong to the genus Stellicola (14 with published descriptions and I whose description by Humes and Ho is in press). Stock (1957) included S. thorelli Kossmann, 1877, 5. pleurobranchi Kossmann, 1877, 5. oreastriphilus Kossmann, 1877, 5. caeruleus (Stebbing, 1900), 5. curticaudatus (Thompson & A. Scott, 1903), 5. gracilis (Thompson & A. Scott, 1903), 5. lankensis (Thompson & A. Scott, 1903), and 5. asterinae (Bocquet, 1952), a species later synonymized with 5. dausi (Rosoll, 1889) by Bocquet and Stock (1962). The four West African species (S. frequens, S. astropectinis, S. luidiae, and 5. lautus} originally described by Humes & Cressey (1958) in the genus Lichomolgus also belong to Stellicola, as pointed out by Bocquet and Stock (1962). In Stellicola there should also be included S. holothuriae (Ummerkutty, 1961) and probably 5. longicaudatus (Thompson & A. Scott, 1903). 5. kossmanni may be separated from these fifteen species as follows : in gracilis, lankensis, and the new species of Humes & Ho (in press) there are two postgenital segments in the female and three in the male ; in frequens, astropectinis, luidiae, and lautus there are three claws on the second antenna ; in clausi there are two claws on the second antenna ; and in oreastriphilus, holothuriae, thorelli, pleuro- branchi, curticaudatus, longicaudatus, and caeruleus, although the second antenna has only a single claw, the length ratio of prosome to urosome in the female (ranging from 2-3-3-3 : I) is distinctly more than 2:1. S. kossmanni shows several features which may be used as identifying characters : the very long terminal seta on the first antenna (as long as the antenna itself), the four slender subterminal setules on the second antenna (these in addition to the six terminal elements), the strongly spinose element on the first maxilla, and in leg 4 the detailed ornamentation of the four elements on the endopod and the nature of the inner seta on the second segment of the exopod. 206 ARTHUR G. HUMES & JU-SHEY HO Stellicola affinis n. sp. PI. 4, figs. 25-31; PL 5, figs. 32-41; PL 6, figs, 42-48. TYPE MATERIAL. 84 $ and 26 £ washed from 2 starfishes, Maculaster maculata maculata (Mtiller & Troschel), exposed on sand bar at low tide on the eastern side of Ankify, on the mainland of Madagascar, opposite Nosy Komba, and about 15 kilometers south of Nosy Be. Collected 8th August, 1964. Holotype female, allotype, and 51 paratypes (40 $, n g) deposited in the British Mus. (Nat. Hist.) Reg. No. 1966.12.7.4-6 ; 27 paratypes (20 $, 7 <$} in the United States National Museum, Washington ; and the remaining paratypes in the collection of A. G. Humes. Other specimens (all in the vicinity of Nosy Be). From Maculaster maculata maculata : 16 $, 8 <$ from I host, exposed on sand bar at Ankify (the type locality), 9th September, 1964. From Maculaster savignyi (Audouin) : 38 $, 6 <$ from 9 hosts, exposed on sand bar at Ankify, 8th August, 1964 ; 60 $, 27 $ from i host, in 2 m., eastern shore of Ambariobe, a small island between Nosy Komba and Nosy Be, 4th October, 1964 ; 12 $, 3 ^ from i host, on sand exposed at low tide, Ambari- obe, 2ist July, 1963. From Retaster cribrosus von Martens : II $, 5 <$ from i host, exposed on sand bar, Ankify, 2nd November, 1963. From Luidia maculata Miiller & Troschel : 35 $, 16 $ from 2 hosts, dredged in 17 m., 13° 16' S, 48° 31' 30" E, northeast of Nosy Faly, I4th September, 1960. FEMALE. Body (PI. 4, figs. 25 and 26) relatively slender. Length (not including setae on caudal rami) i-oi mm. (0-98-1-04 mm.) and greatest width 0-35 mm. (0-32-0-37 mm.), based on 10 specimens. Ratio of length to width of prosome 1-56 : i. Epimeral areas of segment of leg 4 pointed. Segment of leg 5 (PL 4, fig. 27) 61 x 97 /£. Between this segment and genital segment a short intersegmental sclerite ventrally. Genital segment elongated, 156 /i in length, being somewhat wider in its anterior two-thirds (109 /*) than in its posterior third (80 /*) . Areas of attachment of egg sacs located dorsolaterally on anterior part of middle third ; each area (PL 4, fig. 28) bearing 2 short naked setae (outer 13 {i long, inner 17 fi), with a small spinous process about 5 fi long between them. Two postgenital segments, first 56 X 63 fi, second (anal) 43 /* long and 72 ii in greatest width (this segment deeply incised). Caudal ramus (PL 4, fig. 29) divergent, inserted a little ventrally on anal segment, elongated and somewhat narrowed distally, in one female 106 fi (greatest length) X 15 /* (width taken at level of outer seta) making a ratio of 7 : I, in another female 108 X 17 /*, ratio of 6-3:1. Minute setule (4/4 long) on outer proximal margin. Another small setule (8 /*) on outer margin just beyond midpoint of ramus (perhaps corresponding to long marginal setule in S. kossmanni}. Pedicellate dorsal seta 25 p. Outer lateral seta 74 /£, inserted near end of ramus. Innermost terminal seta 130 fi, outermost terminal seta 88 [i, and 2 long median terminal setae, inserted between dorsal (smooth) and ventral (with a row of minute spinules) flaps, 242 /* (outer) and 363 JLL (inner). All setae naked. Both outer and inner walls of ramus with an interruption in sclerotization as shown in figure. Two minute hairs on ventral surface of ramus. NEW SPECIES OF STELLICOLA IN MADAGASCAR 207 Dorsal surface of prosome with scattered hairs. Dorsal surface of urosome with hairs as shown in PL 4, fig. 27 ; ventral surface much less ornamented with hairs. Ratio of length of prosome to that of urosome 1-30 : i. Egg sac (PL 4, fig. 25) fairly elongated, 374 x 143 /*, extending not quite to the ends of caudal rami, and containing a moderate number of eggs about 50 /* in diameter. Rostral area (PL 4, fig. 30) rounded. First antenna (PL 4, fig. 31) y-segmented, lengths of segments (measured along their posterior non-setiferous margins) being 30 (47 /* along anterior margin), 67, 23, 25, 31, 19, and 14/4 respectively. Formula for armature as in 5. kossmanni. Setae naked except for I on antepenultimate segment, I on penultimate segment, and 2 on last segment which are delicately haired. Long terminal seta 213 fi in length, nearly as long as entire first antenna. Aesthete on last segment having a common base with adjacent seta. Sclerotization on ventral surface of segment 3 suggesting an intercalary segment. Second antenna (PL 5, fig. 32) 3-segmented, with no evidence of division of third segment. Armature i ; i ; 3+5+1 claw, as in 5. kossmanni, though relative lengths and fine ornamentation of setae different (compare PL 2, fig. 8 and PL 5, fig. 32). Claw 41 fi along its axis. Anterior surface of second antenna not pilose as in 5. kossmanni ; posterior surface with very fine punctations. Labrum (PL 5, fig. 33) with subtriangular unornamented lobes. Mandible (PL 5, fig. 34) in general resembling that of S. kossmanni, but first element in inner row of teeth less prominent and terminal lash relatively shorter. Paragnath a small indistinct lobe medial to insertions of mandible and first maxilla (PL 5, fig. 38). Four elements on first maxilla (PL 5, fig. 35), including spine with lateral rows of spinules. Second maxilla (PL 5, fig. 36) 2-segmented, with proximal ventral margin of unarmed first segment rather angularly produced ; second segment with long inner spine having serrated flanges (with serrations especially prominent along distal flange) and with terminal process bifurcated as shown in figure. Maxil- liped (PL 5, fig. 37) 3-segmented, with armature resembling that of 5. kossmanni ; second segment lacking rows of small spinules seen in that species. Area between maxillipeds and first pair of legs only slightly protuberant (PL 4, fig. 26) ; a sclerotized line connecting bases of maxillipeds (PL 5, fig. 38). Rami of legs 1-4 (PL 5, figs. 39, 40,41, and PL 6, fig. 42) with same segmentation and spine and setal formula as in 5. kossmanni. Inner seta on coxa relatively long and feathered in legs 1-3 but short (19 /*) and naked in leg 4. Hairs on inner margin of basis absent in leg 4 and apparently also in leg i. In legs 1-3 expansion of basis over anterior surface of first segment of exopod with membranous tip ; this expansion absent in leg 4. Second segment of exopod of leg 4 with inner seta like that in 5. kossmanni. Endopod of leg 4 two-segmented, with distinct articulation between these segments. First segment 31 X 25 ju (greatest dimensions) with finely barbed inner seta 91 ja long. Second segment 50 x 24 /i (greatest dimensions), the inner seta finely barbed and 48 /*, the 2 terminal spines 62 jn (inner) and 51 /* (outer), both with delicately serrate lamellae. Rows of long hairs along outer margins of both segments, with row on second segment interrupted. Row of minute spinules 208 ARTHUR G. HUMES & JU-SHEY HO on anterior surface across end of segment near insertions of 2 spines. Leg 5 (PI. 6, fig. 43) with elongated free segment, 28 X 13 /*. Two unequal terminal setae, outer 61 /* long with a few minute barbs distally, inner 39 [JL with outer serrated fringe. Seta on body near insertion of free segment 25 fi long and delicately feathered. Leg 6 probably represented by the 2 setae on area of attachment of egg sac (see PI. 4, fig- 28). Colour in life in transmitted light slightly opaque, ovary dark gray, eye dark blackish red, egg sacs dark gray. MALE. Body form (PI. 6, fig. 44) resembling that of female but more slender. Length (without ramal setae) 0-78 mm. (0-75-0-82 mm.) and greatest width 0-22 mm. (0-21-0-23 mm.), based on 10 specimens. Ratio of length to width of prosome 1-84 : i. Segment of leg 5 (PI. 6, fig. 45) 31 X 65 ju,. Ventral intersegmental sclerite absent. Genital segment subquadrate, 133 x 117 fi, a little longer than wide, with lateral borders nearly parallel in dorsal view. Three postgenital segments 47 X 59, 43 X 46, and 26 x 46 /* from anterior to posterior. Caudal ramus (see PL 6, fig. 45) resembling that of female, but proportionately more slender, 90 X 12 fi, with ratio of 7-5 : I, and a little more narrowed in its distal half. Surfaces of prosome and urosome, as in female, ornamented with hairs. Ratio of length of prosome to that of urosome 1-16 : i. Rostral area, first antenna, second antenna, labrum, mandible, paragnath, and first maxilla like those of female. Second maxilla also resembling that of female but proximal ventral margin of first segment less produced and more rounded. Maxilliped (PL 6, fig. 46) slender and 4-segmented. First segment unarmed. Second segment with 2 slender naked setae and 2 rows of spinules (i row short, consisting of only about 5 spinules.) Third segment short and unarmed. Proximal part of claw probably representing fourth segment. Slender gently recurved claw in [i along its axis, with evidence of division about midway and having a fringe of delicate hyaline spinules along its concave margin ; bearing proximally 2 setae, one n /* and naked, other 29 fi long with a few minute barbs near its tip. Area between maxillipeds and first pair of legs like that in female. Legs 1-4 resembling those of female, with same spine and setal formula. Endopod of leg 4 much like that in female, with 4 elements from inner to outer 72, 35, 43, and 34 fi long. Leg 5 (PL 6, fig. 47) with free segment 21 x 8 /i, more slender than in female, and 2 terminal setae more unequal, 53 /* and 22 ju, (ratio of 2-4 : i, instead of 1-56 : i in female). Leg 6 (PL 6, fig. 48) consisting of posterolateral flap on ventral surface of genital segment and bearing 2 slender naked setae 25 and 38 /£ long. Spermatophores not observed. Colour in life in transmitted light resembling that of female. [The specific name affinis, Latin = adjoining or related, refers to the close similarity of this species with Stellicola gracilis (Thompson & A. Scott) as discussed below]. NEW SPECIES OF STELLICOLA IN MADAGASCAR 209 COMPARISON WITH OTHER SPECIES. At present there are three species known in the genus Stellicola which, like 5. affinis, have two postgenital segments in the female and three in the male. These are 5. gmcilis (Thompson & A. Scott, 1903), 5. lanken- sis (Thompson & A. Scott, 1903), and the species whose description by Humes and Ho is in press. 5. lankensis (of which only the female is known), from washings of dredged invertebrates in Ceylon, has a much shorter caudal ramus (proportions of about 2-37 : i in T. & S.'s pi. XV, fig. 25). The new species of Humes and Ho differs in having a quadrate caudal ramus, in its smaller size (female 0-79 mm., male 0-64 mm.), and in many other features. 5. gracilis, from washings of dredged invertebrates in Ceylon, is known only from the original description and figures by Thompson & A. Scott. The existence of type material (the " several males and females " of Thompson & A. Scott) is ex- tremely doubtful. The type specimens of S. gracilis were apparently deposited in the University of Liverpool, where Dr. W. A. Herdman, the author of the Report on Pearl Oyster Fisheries, was professor of natural history. An inquiry recently made by Dr. J. P. Harding of the British Museum to the Department of Oceano- graphy of Liverpool University drew the reply that the Andrew Scott collection had been discarded. Our only available information on S. gracilis is, then, to be found in the brief original description. When S. affinis is compared with 5. gracilis certain similarities are obvious. The two species are of about the same length and have the same number of post- genital segments. In the female of both the anal segment is cleft deeply, the caudal rami are divergent, the genital segment has a similar shape, the second antenna appears to have the same general form, the base of the mandible is somewhat angular, and the first segment of the second maxilla seems to project. In the male of both the maxilliped is rather similar. There exist, however, differences between the two species which in our opinion make it impossible to regard them as one and the same species. In the female of S. gracilis the proportions of the caudal ramus are about 4 : I (instead of 6-3 : I or 7 : i as in S. affinis), the two inner setae on the endopod of leg 4 are distinctly feathered (while in S. affinis these setae have very minute barbs), and (based on measurements of T. & S.'s pi. XV, fig. i) the ratio of the length of prosome to urosome is i : I (1-30 : i in S. affinis} and the ratio of length to width of the prosome is 1-75 : i (1-56 : i in 5. affinis}. T. & S.'s pi. XV, fig. 4, shows only three elements on the first maxilla, but a fourth minute seta may have escaped their notice. They describe the tip of the second maxilla as " triangular, sharp and wedge-shaped ", illustrated on pi. XV, fig. 5, but their description is wanting in detail. We conclude that 5. affinis, while related to 5. gracilis, is a separate species. Stellicola longiseta n. sp. PL 6, figs. 49-50; PI. 7, figs. 51-60; PI. 8, figs. 61-66. TYPE MATERIAL. 2 $ and 3 <§ from washings of i starfish, Mithrodia clavigera (Lamarck), under dead coral (Acropora) in a depth of i m., Pte. Ambarionaomby, Nosy Komba, near Nosy Be, Madagascar. Collected 3rd September, 1963. Holo- 210 ARTHUR G. HUMES & JU-SHEY HO type female, allotype, and 2 paratypes (i ?, I $) deposited in the British Mus. (Nat. Hist.) Reg. No. 1966.12.7.7-9 ; the remaining paratypic male (dissected) in the collection of A. G. Humes. Other specimens (from Mithrodia davigem in the same locality). 2 $ from I host in i m., I4th December, 1963. FEMALE. Body (PI. 6, fig. 49) with prosome relatively thickened as in S. affinis, but broader than in that species. Length (excluding setae on caudal rami) 1-18 mm. (1-14-1-22 mm.) and greatest width 0-60 mm. (0-58-0-62 mm.), based on 4 specimens (holotype, I paratype, and 2 specimens collected on I4th December, 1963). Ratio of length to width of prosome 1-36 : i. Segment of leg i separated from head by a dorsal and lateral furrow. Epimeral areas of segment of leg 2 rather pointed posteriorly ; those of segments of legs i, 3, and 4 rounded. Segment of leg 5 (PI. 6, fig. 50) 88 X 146 ju. Between this segment and genital segment a short intersegmental sclerite ventrally. Genital segment a little longer than wide, 177 /* in length, 159 fi in width in its anterior two-thirds, and 114 ju, in width in its constricted posterior third, and showing a slight middorsal hump anteriorly. Areas of attachment of egg sacs situated dorsolaterally on anterior part of posterior third ; each area (PI. 6, fig. 50) with 2 setae (outer 77 fi long and naked, inner 23 ju, and feathered) and between them a small spinous process 6 ju, long. Posterior margin of genital segment with a membrane which simulates spinules laterally. Two postgenital segments 52 X 95 and 34 X 88 /i respectively. Caudal ramus (PL 7, fig. 51), inserted ventrally on anal segment, a little wider than long, 28 X 35 /*, taking average length as 28 /*, the ramus being longer ventrally (31 ju,} than dorsally (25 /*). Long naked setule on outer margin 31/4 long. Outer lateral seta 120 /* and naked. Outermost terminal seta 180 /* with lateral spinules. Innermost terminal seta long, 451 {i, and naked. Two long median terminal setae 460 (i (outer) and 660 fi (inner), both with lateral spinules and their " pegged " regions punctate. These 2 setae inserted between ventral flap (with rows of spinules) and dorsal flap of form shown in figure and bearing very small spinules near pedi- cellate dorsal seta (33 /* long and naked). Group of 2-3 minute hairs on dorsal surface of ramus. Dorsal surface of prosome with only a few hairs. Dorsal surface of urosome with more abundant hairs as in PI. 6, fig. 50 ; ventral surface with very few hairs. Ratio of length of prosome to that of urosome 2-45 : i. Egg sac (PI. 6, fig. 49) elongated and rather stout, about 810 X 300 /*, more than twice as long as the urosome, and containing many eggs about 52 fi in diameter. Rostral area (PI. 7, fig. 52) rather pointed. First antenna with armature as in 5. kossmanni and S. affinis. All setae apparently naked. Lengths of segments (measured as before) 30 (67 ju, along anterior edge) , 107, 29, 53, 46, 23, and 15 fi respectively. Terminal aesthete and adjacent seta with common base. Sclerite on ventral surface of third segment. Long terminal seta 234 fi in length, distinctly less than length of antenna (340 fi). Second antenna (PI. 7, fig. 53) 3-segmented, third segment showing no evidence of division. Armature i ; i ; 3+5+1 claw, as in 2 previous species. Seta on NEW SPECIES OF STELLICOLA IN MADAGASCAR 211 segment i unusually long (165 /i) and setose. Seta on segment 2 relatively short (44 ft] and naked. Claw 68 ju, along its axis. On anterior surface near base of claw a short vermiform protrusion (aesthete ?). Anterior surface of second antenna pilose as in figure. Labrum (PI. 7, fig. 54) with attenuated unornamented lobes. Mandible (PI. 7, fig. 55) with elements of outer row beginning as stout spines and diminishing to slender spinules ; first element in inner row not much larger than rest ; a patch of spinules on posterior proximal surface of mandible. Paragnath a small lobe as in 5. affinis. First maxilla with 4 elements as in 2 previous species, but ornamented spiniform element only very finely spinulose. Second maxilla (PI. 7, fig. 56) with first segment ornamented with a few spinules on both anterior and posterior surfaces and with its proximal ventral margin angularly produced. Second segment with postero-inner finely barbed seta, long inner spiniform seta with its posterior surface spinulose but anterior surface smooth, and with terminal spini- form process spinulose and showing 2 prominent spines near base. Maxilliped (PI. 7, fig. 57) with first segment unarmed, second with usual 2 setae and with spinules as in figure, and third with 2 setae (one spinulose, other naked) and terminal spiniform process. Area between maxillipeds and first pair of legs not protuberant and resembling that in 5. kossmanni. Kami of legs 1-4 like those in 2 previous species, with same spine and setal formula. Fine ornamentation much like that in 5. kossmanni. Last segment of endopod of leg 2 without a bifurcated spinous process (PI. 7, fig. 58), and having a form somewhat different from that in 5. kossmanni. Last segment of endopod of leg 3 (PL 7, fig. 59) of a form slightly different from 5. kossmanni. Leg 4 (PI. 7, fig. 60) with inner seta on coxa relatively short (26 fi) and naked. Slightly barbed inner seta on second segment of exopod shorter than setae on third segment, recurved, and lying (in alcoholized specimens) over the posterior surface of ramus. Endopod (PI. 8, fig. 61) with 2 segments clearly separated. First segment 31 x 28 /*, with inner seta 58 fi long and feathered. Second segment 69 /£ long, 28 ju, in greatest width, and 18 fi wide at narrowest point, its feathered inner seta 74 fi, inner terminal spine 76 fi with a very narrow lamella along one edge, and outer terminal spine 55 ju, with finely serrated lamellae and with a delicately trifurcate tip. Row of hairs on outer margin of this segment interrupted at slight indentation. Leg 5 (PI. 8, fig. 62) with free segment swollen on inner margin, its greatest dimen- sions 39 x 19 ju. Two terminal naked setae extremely unequal, inner 34 fi, outer 170 fi. Seta on body near insertion of free segment about 35 fi and feathered. Leg 6 probably represented by the 2 setae on area of attachment of egg sac (see PI. 6, fig. 50). Colour in life in transmitted light translucent to grayish, eye red, egg sacs gray. MALE. Body (PI. 8, fig. 63) with prosome more slender than in female. Length (without ramal setae) 0-73 mm. (0-72-0-73 mm.) and greatest width 0-29 mm. (0-28-0-29 mm.), based on 3 specimens (allotype, I paratype, and I specimen col- lected I4th December, 1963). Ratio of length to width of prosome 1-62 : I. 212 ARTHUR G. HUMES & JU-SHEY HO Segment of leg 5 (PI. 8, fig. 64) 46 x 79 /i. Ventral intersegmental sclerite absent. Genital segment a little longer than wide, 125 X 104 /*, with sides in dorsal view nearly parallel. Three postgenital segments 42 X 69, 33 X 55, and 20 x 53 fi from anterior to posterior. Caudal ramus like that of female. Surfaces of prosome and urosome, as in female, sparsely ornamented with hairs. Ratio of length of prosome to that of urosome 1-70 : i. Rostral area, first antenna, second antenna, labrum, mandible, paragnath, and first maxilla resembling those in female. Second maxilla much like that of female, with same process on first segment, but with long inner seta less strongly developed and with terminal spiniform process weaker and showing near base only I prominent spine, the second being greatly reduced in size. Maxilliped (PL 8, fig. 65) slender. Spinules on second segment arranged differently than in 2 previous species. Claw 117 fji long (measured along its axis) and showing 2 slight flexures, with only indica- tion of division being an interruption in fringe of delicate hyaline spinules along concave margin. Two setae near base of claw 67 ju, (with a few barbs near tip) and 8 fi (naked) . Area between maxillipeds and first pair of legs resembling that of female. Legs 1-4 resembling those of female, with same spine and setal formula. Leg 5 (PI. 8, fig. 66) with free segment lacking inner swelling, 18 x 9 /*. Two terminal setae 20 ju, (inner) and 58/4 (outer), ratio between them being 2-9:1, instead of 7 : I as in female. Seta on body near insertion of free segment about 20 fi long and slightly feathered. Leg 6 (see PI. 8, fig. 64) resembling that of S. kossmanni, with a minute spinous process and 2 unequal setae, one naked (64^), other feathered (20;*). Spermatophores not observed. Colour in life in transmitted light as in female. (The specific name longiseta, Latin = long seta, alludes to the unusually long terminal seta on the fifth leg in the female). COMPARISON WITH RELATED SPECIES. From the four species of Stellicola known to have two postgenital segments in the female, S. longiseta may readily be separated by its very long seta on leg 5. In addition, this species from Mithrodia has a caudal ramus which is a little wider than long, instead of 4 : I as in 5. gracilis (Thompson & A. Scott, 1903), about 2 : i as in 5. lankensis (Thompson & A. Scott, 1903), and 6-3 : i or 7 : I as in S. affinis. The female of the species from Madagascar whose description by Humes and Ho is in press, although having a nearly quadrate caudal ramus, differs from 5. longiseta in having a rounded rostrum, in the seta on the first segment of the second antenna being not unusually long, in the two setae on the endopod of leg 4 being finely barbed instead of feathered, in the two terminal setae on leg 5 having a ratio of 3 : i instead of 7 : i, and in the ratio of the length of the prosome to that of the urosome being 2-0:1 instead of 2-45 : i. In the male, the species of Humes and Ho differs from S. longiseta in having the genital segment somewhat tapered anteriorly instead of having nearly parallel sides, and in the claw of the maxilliped being evenly recurved instead of showing two flexures. NEW SPECIES OF STELLICOLA IN MADAGASCAR 213 From all four of these species 5. longiseta may be further distinguished by details in ornamentation of the appendages. S. longiseta appears to be closest to the species to be described by Humes and Ho, both having a very short caudal ramus and a somewhat similar genital segment in the female. Stellicola femineus n. sp. PI. 8, fig. 67 ; PL 9, figs. 68-75 ; PL 10, figs. 76-84 ; PL ir, figs. 85-92 TYPE MATERIAL. 8 $, 9 <$, and I copepodid from i starfish, Leiaster leachi Gray, in a depth of 0-5 m., Pte. Mahatsinjo, Nosy Be, Madagascar. Collected 8th August, 1960. Holotype female, allotype, and u paratypes (5 $, 6 $) deposited in the British Mus. (Nat. Hist.) Reg. No. 1966.12.7.10-12, and the remaining paratypes in the collection of A. G. Humes. Other specimens. 4 $, 9 <$ from i starfish, Leiaster speciosus von Martens, in 0-5 m., Pte. Ambarionaomby, Nosy Komba, near Nosy Be, Madagascar, 2nd October, 1960. FEMALE. Body (PL 8, fig. 67) with broad flattened prosome. Length (not including setae on caudal rami) 1-41 mm. (1-32-1-49 mm.) and greatest width 0-94 mm. (0-88-0-99 mm.), based on 10 specimens (holotype, 7 paratypes, and 2 specimens from Leiaster speciosus). Ratio of length to width of prosome 1-06 : i. Segment of leg i completely fused with head. Epimeral areas of segments of legs I and 2 pointed posteriorly ; those of segment of leg 3 also pointed but indented as in figure. Segment of leg 4 small, with its tergum and rounded epimeral areas partly overlapped by preceding segment. Segment of leg 5 (PL 9, fig. 68) 164 X 257 fi. Intersegmental sclerite not evident. Genital segment broad and flattened, 135 X 239 /i, wider than long, with expanded rounded lateral margins. Areas of attachment of egg sacs located posteriorly and slightly dorsally ; each area (PL 9, fig. 69) with 2 unequal feathered setae 34 and 17 fi in length, each arising from a pedicel. Three postgenital segments 44 X 99, 39 X 89, and 59 X 77 /£ from anterior to posterior. Caudal ramus (PL 9, fig. 70), inserted slightly ventrally, subquadrate, 30 X 33 ft, a little wider than long. Long naked setule (55 ju) midway on outer margin. Pedi- cellate dorsal seta 60 fi and feathered. Outer lateral seta 101 /£ and naked, inserted close to outermost terminal seta (208 fi with straight lateral spinules) . Innermost terminal seta 234 ju, with straight lateral spinules. Two long terminal setae, inserted between dorsal (smooth) and ventral (with row of minute spinules) flaps, 396 fi (outer) and 540 /* (inner) and both with unusually long hairlike recurved lateral spinules. Dorsal surface of prosome with short hairs. Dorsal and ventral surfaces of urosome with many hairs as shown in PL 9, fig. 68. Ratio of length of prosome to that of urosome 2-32 : i. Egg sac (see PL 8, fig. 67) moderately elongated and stout, 682 X 341 ju, extending far beyond ends of caudal rami, and containing many eggs about 70 p in diameter. 214 ARTHUR G. HUMES & JU-SHEY HO Rostral area (PL 9, fig. 71) prominent, its posterior margin a little truncated. First antenna (PI. 9, fig. 72) 7-segmented, lengths of segments (measured along their posterior non-setif erous margins) being 46 (35 ju, along anterior margin) , 107, 34, 36, 25, 13, and 12 fi respectively. Formula for armature as in 3 previous species. Certain setae with short hairs as in figure. Ventral sclerite on third segment. Long terminal seta 203 ju in length, distinctly less than length of antenna (270 ju,). Second antenna (PL 9, fig. 73) 3-segmented, 495 /i in length, with no evidence of division in segment 3. Armature I ; I ; 3+5+1 claw as in 3 previous species. Third segment elongated, 300 /* along outer edge, 187 /i along inner edge, with a sclerotized strongly bent pointed process near base of claw (PL 9, fig. 74). Claw 96 fi along its axis. All setae naked. Labrum (PL 9, fig. 75) with moderately rounded unornamented lobes. Mandible (PL 10, fig. 76) in general resembling that of S. longiseta but without proximal patch of spinules. Paragnath (see PL 9, fig. 71) a small lobe. First maxilla (PL 10, fig. 77) with 4 elements. Second maxilla (PL 10, fig. 78) showing second segment with long inner spiniform seta bearing a row of spinules, those in middle of row much longer and stronger ; terminal spiniform process with a single large tooth-like proximal spine (2 such spines seen on left second maxilla in I female) . Maxilliped (PL 10, fig. 79) with first segment having a distal patch of spinules, second with i naked and i feathered seta and rows of spinules as in figure, and third with 2 small unequal setae and terminal spiniform process. Area between maxillipeds and first pair of legs not protuberant and formed as in PL 9, fig. 71. Rami of legs 1-4 (PL 10, figs. 80, 81, 82, and 83) like those in 3 previous species with same spine and setal formula. Endopods of legs 1-3 rather elongated ; last segment of endopod of leg 2 without a bifurcated spinous process. In leg 4 coxa with inner seta apparently reduced to a mere vestige. Feathered inner seta on second segment of exopod short and lying free of ramus. Endopod (PL 10, fig. 84) slender, 2 segments clearly separated. First segment 38 X 15-5 /* with its inner seta short, 30 fi, and finely barbed distally. Second segment 50 X 12 /* (greatest dimensions), its feathered inner seta 52 ju,, inner terminal spine 76 fi, and outer terminal spine 31 /*, both spines finely barbed. (In another female first segment 33 x 20/1, and second segment 58 X 14-5 /*). Marginal hairs on segments as in figure. Leg 5 (PL n, fig. 85) with elongated slightly recurved slender free segment, 71 x 22 /£ (greatest dimensions), tapering slightly distally. Two terminal setae naked and very unequal, inner 22 /*, outer 113 ju,. Seta on body near insertion of free segment about 48 /* and feathered. Leg 6 probably represented by the 2 setae on area of attachment of egg sac (see PL 9, fig. 69). Colour in life in transmitted light orange-red to deep red, eye red, egg sacs red. MALE. Body (PL n, fig. 86) with broad flattened prosome as in female. Length (excluding setae on caudal rami) 0-85 mm. (0-82-0-88 mm.) and greatest width 0-48 mm. (0-46-0-49 mm.), based on 10 specimens (allotype, 8 paratypes, and I NEW SPECIES OF STELLICOLA IN MADAGASCAR 215 specimen from Leiaster speciosus). Ratio of length to width of prosome i-n : i. Segment of leg 5 completely fused dorsally with genital segment (PL n, fig. 87), and only a faint indication of separation between these 2 segments ventrally ; segment of leg 5 and genital segment combined a little longer than wide, 221 X 195 /«. No intersegmental sclerite. Four postgenital segments 28 x 68, 27 x 62, 24 x 57, and 29 X 52 fi, from anterior to posterior. Caudal ramus similar to that of female but smaller, 18 x 21 ju,. Surfaces of prosome and urosome, as in female, ornamented with hairs. Ratio of length of prosome to that of urosome 1-73 : i. Rostral area, first antenna, and second antenna as in female. Labrum (PI. n, fig. 88) with 2 lobes apparently somewhat more pointed than in female. Mandible, paragnath, first maxilla, and second maxilla like those of female. Maxilliped (PI. n, fig. 89) without long terminal claw, but instead having a form rather like that of female. First segment without ornamentation, second with 2 small naked setae (one hyaline and with a blunt tip) and bearing a patch of spinules, and third with one of 2 setae at base of naked terminal spiniform process greatly elongated. Area between maxillipeds and first pair of legs as in female. Legs 1-4 resembling in major features those of female, having same spine and setal formula. In legs 2 and 3 apparently a slight sexual dimorphism in lengths of 3 spines on last segment of endopod (from proximal to distal in leg 2 of male 12, 13, and 24 [i, but in female 22, 21, and 34 /£ ; in leg 3 of male 12, 13, and 25 ju, but in female 23, 24, and 40 ft). Dimensions of endopod of leg 4 : first segment 22 x 13 fi, with its seta 21 p ; second segment 31 x n /i, with its seta 34 fi and spines 50 fi (inner) and 24 fi (outer) . Leg 5 (PI. n, fig. 90) with rectangular free segment relatively shorter than in female, 19 X 9 /*, with 2 terminal setae 16 /* (inner) and 56 fi (outer), with ratio of 3-5 : i, instead of 5 : i as in female. Seta on body near insertion of free segment about 22 11 long and feathered. Leg 6 (PI. 11, fig. 91) consisting of a posterolateral flap on ventral surface of genital segment and bearing at either side of a small spinous process 2 slender naked setae 46 and 29 /* long. Spermatophore, observed only inside body of male, of elongated form (PI. n, fig. 92). Colour in life in transmitted light paler orange-red than in female, eye red. (The specific name femineus , Latin = womanly or feminine, alludes to the weakly prehensile, female-like maxilliped in the male of this species). COMPARISON WITH RELATED SPECIES. The female of S. femineus differs from almost all other known species in the genus in having the free segment of leg 5 at least three times longer than wide. The only species which has similar proportions in leg 5 is the species of Humes and Ho (in press) where the free segment is 65 x 21 fi. In this species, however, there are only two postgenital segments in the female. The male of S. femineus is unique in the genus in having a female-like maxilliped, without a well-formed prehensile claw. ZOOL. 15, 5. 17 216 ARTHUR G. HUMES & JU-SHEY HO Stellicola pollex n. sp. PI. n, fig. 93 ; PL 12, figs. 94-101 ; PI. 13, figs. 102-109 ; PI. 14, figs. 110-114. TYPE MATERIAL. 5 $ and 4 <$ from 2 starfishes, Linckia laevigata (L.), in a depth of 10 m., at Nosy Ovy (=Berafia), 13° 59' S, 47° 46' 30" E, to the southwest of Nosy Be, Madagascar. Collected ist October, 1964. Holotype, allotype, and 5 paratypes (3 $, 2 <£) deposited in the British Mus. (Nat. Hist.) Reg. No. 1966.12.7.13-15, and the remaining 2 paratypes (dissected) in the collection of A. G. Humes. FEMALE. Body (PI. n, fig. 93) with broad flat prosome. Length (without setae on caudal rami) 0-99 mm. (0-94-1-03 mm.) and greatest width 0-78 mm. (0-68-0-88 mm.), based on 5 specimens. Ratio of length to width of prosome 0-95 : i, prosome being slightly wider than long. Segment of leg I separated dorsally and laterally from head by a distinct furrow. Epimera of metasomal segments as in 5. femineus. Segment of leg 5 (PI. 12, fig. 94) 91 X 180 ju. Ventrally between this segment and genital segment a very short intersegmental sclerite. Genital segment broad and flattened, 161 X 198 /*, wider than long, with expanded rounded lateral margins lobed posteriorly. Areas of attachment of egg sacs located ventrally and hidden in dorsal view by the lobes ; each area (PI. 12, fig. 95) with 2 naked setae, 39 and 35 IL in length, borne on pedicels. Three postgenital segments 35 X 64, 36 X 62, and 34 X 57 /t from anterior to posterior. Caudal ramus (PI. 12, fig. 96), inserted ventrally, quadrate, 20 X 20 /*, with naked setule on outer margin 19 ju, long. Pedicellate dorsal seta 36 /* and feathered. Outer lateral seta 60 /*, and naked. Outermost terminal seta 127 /* and innermost 96 [i, both with lateral spinules. Two long terminal setae, inserted between flaps as in S. femineus, 320 fi (outer) and 495 /i (inner), both with short spinules. Dorsal surface of prosome with scattered hairs (not shown in PI. 11, fig. 93, because of the reduced scale). Dorsal surface of urosome with hairs and refractile points as in PI. 12, fig. 94 ; ventral surface with very little ornamentation. Ratio of length of prosome to that of urosome 2-5 : I. Egg sacs not present on any of females collected. Rostral area as in S. femineus, though posterior margin less truncated. First antenna (PI. 12, fig. 97) more slender than in 5. femineus, but with same segmentation and armature. Lengths of segments (measured as before) 44 (35 fi along anterior margin), 116, 45, 50, 33, 20, and 16 ju, respectively. One seta on segment 6 and 4 setae on segment 7 with lateral hairs ; rest of setae naked. Terminal seta 117 /*, about one-third length of antenna (330 ju). Second antenna (PI. 12, fig. 98) slender as in 5. femineus, with third segment showing no indication of division. Armature I ; I ; 3+5+1 claw as in 4 previous species. Near base of claw a small spinous process. Claw 61 /* along its axis. All setae naked. Labrum as in S. femineus, though lobes a little more rounded. Mandible (PI. 12, fig. 99) with slender spinules along outer margin and row of dentiform spinules along inner margin (proximal spinule enlarged and preceded by NEW SPECIES OF STELLICOLA IN MADAGASCAR 217 a slight flange). Paragnath (PL 12, fig. 100) a small lobe with very slender spinules. First maxilla in general like that of S. femineus. Second maxilla (PI. 12, fig. 101) with first segment unarmed, second with 2 postero-inner elements (a slender barbed seta and a long spiniform seta ornamented with a row of spinules along its distal edge and showing a very weak line of division proximally) and terminating in a spiniform process bearing rows of teeth as in figure. Maxilliped (PL 13, fig. 102) resembling in major features that of S. femineus, though details of ornamentation somewhat different. Area between maxillipeds and first pair of legs as in 5. femineus. Kami of legs 1-4 (PL 13, figs. 103, 104, 105, and 106) segmented as in previous 4 species, with similar spine and setal formula, except for endopod of leg 4 which is o-o ; II. Endopods of legs 1-3 rather elongated as in 5. femineus ; last segment of endopod of leg 2 without a bifurcated spinous process. Leg 4 with only a vestige of inner coxal seta. Inner seta on second segment of exopod short (22 /*) and apparently naked. Endopod (PL 13, fig. 107) 41 x 10 fi, only incompletely divided into 2 segments, its inner margin entire. Two inner setae usually seen in Stellicola here absent, and only armature of endopod consisting of 2 terminal smooth spines 60 IJL (inner) and 31 ju, (outer) in length. Rows of hairs along outer and inner margins of endopod, outer row interrupted at incomplete division of ramus. Leg 5 (PL 13, fig. 108) with free segment 55 x 27 ju, (greatest dimensions, its inner proximal margin somewhat irregular and produced) . Two terminal setae naked and unequal, inner 43 fi, outer 56 /i. Seta on body near insertion of free segment about 37 fi and slightly feathered. Leg 6 probably represented by the 2 setae on area of attachment of egg sac (see PL 12, fig. 95). Colour in life in transmitted light slightly opaque, eye red, egg sacs gray. MALE. Body (PL 13, fig. 109) resembling that of female. Length (excluding setae on caudal ramus) 0-55 mm. (0-53-0-56 mm.) and greatest width 0-36 mm. (0-35-0-37 mm.), based on 4 specimens. Ratio of length to width of prosome 1-07 : i. Segment of leg 5 and genital segment fused (PL 14, fig. no) as in 5. femineus ; these fused segments 127 fi long, 115 p wide just in front of leg 5, and 105 fi wide behind leg 5. No intersegmental sclerite. Four postgenital segments 28 x 46, 23 X 43> 22 X 40, and 18 X 36 IJL from anterior to posterior. Caudal ramus similar to that of female but smaller, 12-5 X 13 (i. Surfaces of prosome and urosome, as in female, ornamented with hairs. Ratio of length of prosome to that of urosome 2-07 : I. Rostral area and first antenna as in female. Second antenna also resembling that of female, but having a conspicuous thumb-like process on inner margin of third segment (PL 14, fig. in). Labrum, mandible, paragnath, first maxilla, and second maxilla like those in female. Maxilliped (PL 14, fig. 112) slender, with long terminal claw. First segment unarmed, second with 2 naked setae and 2 patches of spinules, third short and unarmed. Proximal half of claw probably representing fourth segment. Claw only slightly recurved, 130 /* in length along its axis, showing ZOOL. 15, 5. I?§ 2i8 ARTHUR G. HUMES & JU-SHEY HO faint indication of division about midway, with a fringe of small spinules along its concave margin and 2 unequal setae 8 /* (naked) and 26 fi (slightly barbed) near its base. Area between maxillipeds and first pair of legs as in female. Legs 1-4 resembling those of female, with same spine and setal formula. As in 5. femineus a slight sexual dimorphism in lengths of 3 spines on last segment of endopod of legs 2 and 3 (from proximal to distal in leg 2 of male 9, 10, and 22 /£, but in female 16-5, 17-5, and 35 JLI ; in leg 3 of male 8, 10, and 20 fi, but in female 16-5, 18-5, and 35 /*). Dimensions of endopod of fourth leg 22 X 8 /*, with inner spine 31 ju, and outer spine 17 /^. Leg 5 (PI. 14, fig. 113) with free segment smaller and having nearly parallel sides, 20 X 8 [i, the terminal setae 18 ju, (inner) and 52 ju (outer), with ratio of 2-9 : I, instead of 1-3:1 as in female. Leg 6 (PI. 14, fig. 114) consisting of a posterolateral flap on ventral surface of genital segment and bearing 2 slender naked setae 21 and 36 jn. Spermatophores not observed. Colour in life in transmitted light as in female. (The specific name pollex, Latin — a thumb, refers to the thumb-like process on the third segment of the second antenna of the male). COMPARISON WITH OTHER SPECIES. 5. pollex has two readily observable characters by which it may be distinguished from all other species in the genus, namely, the thumb-like process on the third segment of the second antenna in the male and the armature of the endopod of leg 4 (o-o ; II) in both sexes. In the fifteen previously known species and in the four species described above the formula for the endopod of leg 4 is o-i ; II, i. Such an armature seems to be a stable and constant feature in the genus. The unusual armature in 5. pollex might suggest at first glance that this species should be removed to a separate genus. However, other features of 5. pollex (first antenna, second antenna, mouthparts, legs 1-5, and body form) conform very closely to the pattern of Stellicola. The structure of the mandible (which is an important generic character in the Licho- molgidae) is very much like that in other species of the genus. Therefore, in spite of the rather remarkable difference in the armature of this endopod, we are consider- ing the species to belong to Stellicola. Stellicola oreastriphilus Kossmann, 1877 PI. 14, figs. 115-120 Two females and one male were recovered from washings of a single starfish, Protoreaster nodosus (L.), in a depth of 2m., on Cymodocea, at Ambatoloaka, Nosy Be, I5th October, 1960. This represents a new host record, the species having been previously found on six other starfishes — Asterope carinifera (Lamarck) in the Red Sea by Kossmann (1877), on Protoreaster lincki (Blainville) , Culcita schmideli- ana (Retzius) , Pentaceraster mammillatus (Audouin) , and Poraster superbus (Mobius) in Madagascar by Humes & Cressey (1961), and on Choriaster granulatus (Liitken) in Madagascar by Humes & Ho (in press) . NEW SPECIES OF STELLICOLA IN MADAGASCAR 219 In connection with the description of the five new species above, specimens of 5. oreastriphilus (from a lot of approximately 1,000 individuals washed from 200 Protoreaster lincki, in a depth of 10 cm., at Pte. Mahatsinjo, Nosy Be, 6th September, 1964) were restudied. The dissections were done in lactic acid (see Humes & Good- ing, 1964), enabling us to see certain details not easily observed in the original dis- sections in glycerine. Notes on these features are offered here as a supplement to the work of Humes & Cressey (1961). FEMALE. The flattened genital segment (PI. 14, fig. 115) expanded laterally, with subparallel lateral margins. A pair of ventral refractile sclerotizations present on sides of segment. Tip of second antenna (PI. 14, fig. 116) bearing 5 setae and I claw. Paragnath (PI. 14, fig. 117) a hairy lobe bearing a sclerotized process. Maxilliped (PL 14, fig. 118) with second segment having 2 setae (one surficial, other on a hyaline base at inner marginal angle) and a crescentic row of spinules ; third segment with 2 very unequal small setae at base of terminal spiniform process. Endopod of leg 4 (PI. 14, fig. 119) with marginal hairs as in figure. MALE. Segment of leg 5 and genital segment (PI. 14, fig. 120) fused completely dorsally and their ventral separation suggested by a very indistinct line. Fused segments 120 JJL long, 109 fi wide just in front of leg 5, and 101 /* wide posterior to leg 5. (The rather distinct separation of the two segments shown by Humes & Cressey, 1961, fig. 25, is not present in the material which we have re-examined). Stellicola caeruleus (Stebbing, 1900) PI. 15, figs. 121-128 ; PL 16, figs. 129-134 ; PL 17, figs. 135-141 This species was placed by Stebbing (1900, pp. 664-666, plate LXXIVB) in a new genus Linckiomolgus , but, as Stock (1957) has pointed out, and as will be seen from the redescription below, it belongs more properly to Stellicola. Through the kindness of Miss Patricia D. Lofthouse we have been able to study syntypes (i female and I male) of 5. caeruleus [Brit. Mus. (Nat. Hist.) reg. no. 1928.12.1.3020]. These are labelled "China Sea on blue Linckia", though the type locality given by Stebbing was Feather Island, China Straits, New Guinea. The dissections of these specimens, mounted on two slides, have been returned to the British Museum (Nat. Hist.). Although the condition of the specimens (which were in amplexus) was less than perfect, with certain setae broken or missing, the major features could be observed in the dissections. Stebbing's original description being inadequate in many respects, especially regarding the mouthparts, a redescrip- tion of the species based on syntypic material is desirable. FEMALE. Body (PL 15, fig. 121) with greatly expanded prosome. Length (without setae on caudal rami) 1-23 mm. and greatest width 0-88 mm. Ratio of length to width of prosome 0-97 : I, being a little wider than long. Segment of leg I completely fused with head. Epimeral areas of metasomal segments resembling those of 5. femineus. Genital segment (PL 15, fig. 122) wider than long, 156 X 208 /*, broadly rounded laterally in its anterior half and tapering in its posterior half where the areas of 220 ARTHUR G. HUMES & JU-SHEY HO attachment of the egg sacs are located laterally. Dorsally with two longitudinal posterior elevations with sclerotization as in figure. Each egg sac attachment area with 2 unequal setae, one arising from a prominent slender pedicel 31 x 8 /t. Three postgenital segments 60 X 86, 42 X 78, and 52 X 72 /* from anterior to posterior. Caudal ramus (PI. 15, fig. 123) only a little longer than wide, 32 X 28 /t. Ratio of length of prosome to that of urosome 2-32 : I. Egg sac broken on female examined ; an isolated egg about 74 ju, in diameter. Rostral area (PI. 15, fig. 124) incomplete posteriorly. First antenna (PI. 15, fig. 125) 7-segmented, lengths of segments (measured along their posterior non-setiferous margins) being 55 (44 /i along anterior margin), 113, 39> 43> 33> r5> and *4 /* respectively. Formula for armature 4, 13 (5 -f- 2 +6), 6, 3, 4 + i aesthete, 2+1 aesthete, and 7+1 aesthete, as in all 5 new species de- scribed above and as in S. oreastriphilus. Ventral sclerite present on third segment. Second antenna (PI. 15, fig. 126) with a single claw 79 /* along its axis and with armature as in 5. femineus. Third segment 166 /i along outer side, 91 fi along inner side, with no evidence of division. Labrum as in PL 15, fig. 124. Mandible (PL 15, fig. 127) with first 2 elements of inner row only slightly larger than succeeding ones. Paragnath a small lobe with coarse hairs. First maxilla with 4 elements. Second maxilla (PL 15, fig. 128) with second segment with long inner spiniform seta bearing a row of spinules, those in middle of row somewhat longer ; terminal spiniform process with strong graduated teeth. Maxilliped (PL 16, fig. 129) 3-segmented, with second segment apparently having only i seta, and with 2 very unequal spinules near base of short terminal claw-like process. Area between maxillipeds and first pair of legs resembling that of 5. femineus. Rami of legs 1-4 (PL 16, figs. 130, 131, 132, and 133) segmented and armed as in S. kossmanni, S. affinis, S. longiseta, and 5. femineus, except for last segment of exopod of leg 3 which has formula of II, I, 5. In leg 4 coxa with only a minute knob probably representing a vestige of inner coxal seta seen in other species. Inner seta on second segment of exopod 25 ft long and naked. Endopod 2-segmented, 2 segments distinctly separated. First segment 27 X 15 /£, with its inner seta 36 ju, and finely barbed. Second segment 31 x 12 ju,, its inner seta 42 /£ and feathered, inner terminal spine 66 fi and barbed, outer spine 29 fi with only a few minute barbs. (Marginal hairs on endopod segments not discernible). Leg 5 (PL 16, fig. 134) with free segment about 44 x 17 /*, bearing 2 very unequal terminal setae. Leg 6 probably represented by the 2 setae on area of attachment of egg sac (see PL 15, fig. 122). MALE. Body (PL 17, fig. 135) with prosome somewhat less expanded than in female. Length (excluding ramal setae) 0-52 mm. and greatest width 0-32 mm. Ratio of length to width of prosome 1-09 : I, being a little longer than wide. Segment of leg 5 fused with genital segment (PL 17, fig. 136), as in 5. femineus, S. pollex, and 5. oreastriphilus. These 2 segments combined longer than wide : no 11 in length, 83 fi wide at level of fifth legs, and 75 /* wide posterior to these NEW SPECIES OF STELLICOLA IN MADAGASCAR 221 legs, where lateral margins are subparallel. Four postgenital segments 30 X 47, 30 x 44, 20 x 40, and 22 X 37 fi from anterior to posterior. Caudal ramus similar to that of female but smaller, 19 x 15 fi. Ratio of length of prosome to that of urosome 2-05 : i. Rostral area (PL 17, fig. 137) well-defined posteriorly, with its border there slightly indented. First antenna, second antenna, labrum, mandible, paragnath, and first maxilla like those of female. Second maxilla (PI. 17, fig. 138) with second segment much like that of female, but with first segment showing 3 peculiar sclerotized processes arising ventrally. (Possibly these are adhesive pads). Maxilliped (PI. 17, fig. 139) small, 2-segmented, with terminal claw 22 fi along its axis. Both segments without ornamentation or armature ; a small knob on basal area of claw. Legs 1-4 like those in female, with same spine and seta formula. Leg 5 (PI. 17, fig. 140) with a small free segment, 12 x 7 /a, bearing 2 terminal setae 18 /* (inner) and 55 /* (outer). Leg 6 (PI. 17, fig. 141) consisting of the usual posterolateral flap on ventral surface of genital segment and bearing 2 unequal setae. Spermatophores not observed. COMPARISON WITH OTHER SPECIES. Based on our study of syntypes, 5. caeruleus shows three features not found in the other nineteen species of the genus, as far as known : the two posterior sclerotized elevations on the dorsal surface of the genital segment of the female, the three peculiar sclerotized processes on the first segment of the second maxilla in the male, and the formula of II, I, 5 on the last segment of the exopod of leg 3 in both sexes. A small 2-segmented maxilliped with a short claw is also found in the male of 5. holothuriae (Ummerkutty) , but this Indian species differs from 5. caeruleus in having the formula III, I, 5 on the last segment of the exopod in leg 3 and in the shape of the caudal ramus (much wider than long). 5. caeruleus may thus readily be distinguished from other members of the genus. NOTES ON THE VARIABILITY OF CERTAIN FEATURES IN THE GENUS STELLICOLA The twenty species here recognized in the genus Stellicola exhibit certain variations in external structure worthy of note. The second antenna may bear terminally a single claw, two claws, or three claws. Those with three claws are the four West African species : 5. frequens, S. astropectinis, S. luidiae, and 5. lautus. (In all of these the middle spine on the last segment of the endopod of leg 2 is unusually short, whereas in other species, as far as known, this spine is not thus shortened ; the second segment of the endopod of leg 4 has a sharp spinous process midway on its outer margin opposite the inner seta). The only species with two claws is 5. clausi. The remaining fifteen species have a single claw. The third segment of the second antenna in some species may show an incomplete division (as in the species to be described by Humes and Ho, in press, and in S. kossmanni). The armature of legs 1-4 is similar in all species (where it has been described) except for 5. caeruleus, where the formula for the last segment of the exopod of leg 3 is II, I, 5, and S. pollex, where the endopod of leg 4 has a formula of o-o ; II. 222 ARTHUR G. HUMES & JU-SHEY HO In a few species the basis of leg 4 is greatly elongated and the inner coxal seta is absent, this seta being perhaps represented by a small knob, as in 5. femineus, S. pollex, and S. caeruleus. S. thorelli, S. pleurobranchi, S. curticaudatus , S. longi- caudatus, and 5. holothuriae appear to be similarly constructed, but the exact nature of the protopodal segments of leg 4 is impossible to determine from the descriptions and figures of these species. S. oreastriphilus represents an intermediate form between those species with an elongated basis and those with a short basis (as in 5. kossmanni, for example). The maxilliped of the male in most species is elongated and bears a long prehensile claw, but in S. femineus it is female-like and only weakly prehensile, and in 5. caeruleus it is small, 2-segmented, with a short claw. The second maxilla of the male of S. caeruleus is remarkable in the development of three peculiar sclerotized processes on the first segment. In most other species, as far as known, there is no sexual dimorphism in this appendage, but in S. longiseta and 5. affinis the second maxilla of the male shows slight differences from that of the female. In the males of certain species the segment of leg 5 and the genital segment are almost completely fused, as in 5. oreastriphilus, S. femineus, S. pollex, and S. caeruleus. It may be noted that the tendency in leg 4 toward elongation of the basis and loss of the inner coxal seta occurs in the same four species. The males of five species (S. thorelli, S. pleurobranchi, S. lankensis, S. curticaudatus, and 5. longicaudatus} are unknown, and the available descriptions of the females are lacking in detail. Hence, it is impossible at present to determine whether or not these variations exist in them. The significance of such variations as mentioned above is not clear. The four West African species seem to be closely related and set apart from the rest, perhaps constituting an evolutionary line within the genus. It will be necessary, however, to have much more information on the species within the genus before well-founded conclusions regarding evolutionary relationships can be made. Undoubtedly many more species of Stellicola remain to be discovered, since relatively few species of potential asteroid hosts have been examined. KEYS TO THE SPECIES OF STELLICOLA The preparation of keys for the identification of the species of Stellicola presents certain difficulties, since detailed information regarding various structures in several species is not available in the existing descriptions or figures. Furthermore, the males of five species (thorelli, pleurobranchi, lankensis, curticaudatus, and longicaudatus} are unknown and consequently cannot be included in the key. The keys given here are entirely artificial and are presented only as aids in identification. Final determination of a species should be made by reference to the original descriptions and figures, or to actual specimens, if available. FEMALES 1 Two postgenital segments . . ........ 2 Three postgenital segments .......... 6 2 Caudal ramus 2 x or more than 2 x longer than wide . . . . • . . 3 Caudal ramus short, nearly quadrate . . . . .* • . 1 1; 5 NEW SPECIES OF STELLICOLA IN MADAGASCAR 223 3 Caudal ramus at least 4 x longer than wide ....... 4 Caudal ramus about 2 x longer than wide ...... lankensis 4 Caudal ramus 6-3-7 x longer than wide ....... affinis Caudal ramus 4 x longer than wide ........ gracilis 5 Seta on first segment of second antenna unusually long, reaching nearly to end of appendage ; free segment of leg 5 swollen on inner margin ; caudal ramus slightly wider than long, 28x35^ . . . . . . . . longiseta Seta on first segment of second antenna not unusually long ; free segment of leg 5 with sides nearly parallel ; caudal ramus slightly longer than wide, 28 x 23 /* species of Humes and Ho, in press 6 With more than i claw on last segment of second antenna ..... 7 With only i claw on last segment of second antenna . . . . . 1 1 7 With 3 claws on last segment of second antenna ...... 8 With 2 claws on last segment of second antenna ...... clausi 8 Length about i-o mm. ; genital segment a little longer than wide ... 9 Length 1-3-1-6 mm. ; genital segment about as long as wide . . . . 10 9 Caudal ramus 48 x 36 /LI, ratio of 1-33 : i . . . . . . . frequens Caudal ramus 60 x 28 //, ratio of 2-14 : i . . . . . . astropectinis 10 Caudal ramus 61 x 38 /u, ratio of 1-61 : i . . . . . . . luidiae Caudal ramus 96 x 43 //, ratio of 2-23 : i . . . . . . . lautus 1 1 Formula of endopod of fourth leg o-o ; II ...... pollex Formula of endopod of fourth leg o-i ; II, i . . . . . . . 12 12 With 2 posterior dorsal sclerotized elevations on genital segment . caeruleus Without such elevations ........... 13 13 Free segment of leg 5 a little more than 3 x longer than wide and slightly recurved ; length 1-41 mm. ........... femineus Free segment of leg 5 less than 3 x longer than wide and not recurved ; length 1-2 mm. or less ............ 14 14 Ratio of length of prosome to urosome 1-87 : i, ratio of length to width of prosome 1-45 : i ........... kossmanni Ratio of length of prosome to urosome 2-4-3-3 : i ; ratio of length to width of prosome 0-94-1-22 : i .......... 15 15 Ratio of body length to width about 1-28 : i . . . . . longicaudatus Ratio of body length to width about 1-5:1 . . . . . . . 16 1 6 Two setae on free segment of leg 5 about equal in length . . . . . 17 Two setae on free segment of leg 5 distinctly unequal in length . . . . 18 17 Basis of leg 4 greatly elongated, ratio of its length to length of exopod about 13-5 .-5 pleurobranchi Basis of leg 4 not as elongated, ratio of its length to length of exopod about 9 : 5 oreastriphilus 1 8 Postgenital segments and caudal rami together more than 4 x as long as genital segment ............ thorelli Postgenital segments and caudal rami together only a little longer than genital segment or shorter than that segment . . . . . . . . 19 19 Postgenital segments and caudal rami together a little longer than genital segment curticaudatus Postgenital segments and caudal rami together a little shorter than genital segment holothuriae MALES 1 Three postgenital segments .......... 2 Four postgenital segments .......... 5 2 Caudal ramus distinctly elongated ......... 3 Caudal ramus nearly quadrate ......... 4 224 ARTHUR G. HUMES & JU-SHEY HO 3 Caudal ramus 7-5 x longer than wide . . . . . . . affinis Caudal ramus about 4 x longer than wide . . . . . . gracilis 4 Caudal ramus slightly longer than wide ; claw of maxilliped evenly recurved ; free segment of leg 5 nearly 4 x longer than wide . species of Humes and Ho, in press Caudal ramus a little wider than long ; claw of maxilliped with 2 slight flexures ; free segment of leg 5 about 2 x longer than wide .... longiseta 5 With more than i claw on last segment of second antenna . . . . 6 With only i claw on last segment of second antenna . . . . . 10 6 With 3 claws on last segment of second antenna ...... 7 With 2 claws on last segment of second antenna ..... clausi 7 Length less than i mm. ; free segment of leg 5 about 24-25 x 12 fj, . . 8 Length somewhat more than i mm. ; free segment of leg 5 about 36-38 x 17 /* . 9 8 Caudal ramus a little longer than wide, ratio about 1-3:1 . . frequens Caudal ramus more elongated, ratio about 2-1:1 . . . . astropectinis 9 Caudal ramus somewhat elongated, ratio about 1-6 : i . . . . luidiae Caudal ramus more elongated, ratio about 2-2:1 . . . . . hint us 10 With thumb-like process on third segment of second antenna ; formula of endopod of fourth leg o-o ; II ......... pollex Without thumb-like process on third segment of second antenna ; formula of endo- pod of fourth leg o-i ; II, i . . . . . . . . . ii 11 Maxilliped not elongated, 2- or 3 -segmented, with relatively short claw . . 12 Maxilliped elongated, 4-segmented, with long prehensile claw . . . . 14 12 Maxilliped small, 2 -segmented, with relatively small but distinctly prehensile claw 13 Maxilliped female-like, 3-segmented, with terminal spiniform process instead of a claw femineus 13 First segment of second maxilla with 3 peculiar sclerotized processes . caeruleus First segment of second maxilla without such processes . . . holothuriae 14 Segment of leg 5 and genital segment separated ..... kossmanni Segment of leg 5 and genital segment almost completely fused . oreastriphilus REFERENCES BOCQUET, C. 1952. Copepodes semi-parasites et parasites des echinodermes de la region de Roscoff. Description de Lichomolgus asterinae n. sp. Bull. Soc. Zool. France, 77 (5-6) : 495-504- BOCQUET, C. & STOCK, J. H. 1962. Copepodes parasites d'invertebres des cotes de la Manche. IX. Cyclopoi'des associ6s a Marthasterias glacialis (L.). Arch. Zool. Exp. Gen. 101 (notes et revue) (2) : 79-91. HUMES, A. G. & CRESSEY, R. F. 1958. Four new species of lichomolgid copepods parasitic on West African starfishes. Butt. Inst. Fr. Afrique Noire, 20, ser. A, no. 2, pp. 330-341. 1961. Lichomolgus oreastriphilus (Kossmann) copepode cyclopoi'de parasite des etoiles de mer a Madagascar. Mem. Inst. Sci. Madagascar, 1959, ser. F, 3 : 83-92. HUMES, A. G. & GOODING, R. U. 1964. A method for studying the external anatomy of copepods. Crustaceana, 6 (3) : 238-240. HUMES, A. G. & Ho, J.-S. (in press). Cyclopoid copepods associated with the starfish Choriaster granulatus (Liitken) in Madagascar. Cahiers ORSTOM Oceanographie. KOSSMANN, R. 1877. Entomostraca (i. Theil : Lichomolgidae) . In : Zool. Ergeb. Reise Kustengeb. Rothen Meeres, Erste Halfte, IV, pp. 1-24. ROSOLL, A. 1889. Uber zwei neue an Echinodermen lebende parasitische Copepoden : Ascomyzon comatulae und Astericola clausii. S.-B. Math.-Naturw. Cl. Ah. Wiss. Wien, 97, Abt. I, Heft 4, pp. 188-202. STEBBING, T. R. R. 1900. On Crustacea brought by Dr. Willey from the South Seas. Zoologi- cal Results based on material from New Britain, New Guinea, Loyalty Islands and elsewhere collected during the years 1895, 1896 and 1897, part 5, pp. 605-690. NEW SPECIES OF STELLICOLA IN MADAGASCAR 225 STOCK, J. H. 1957. Some notes on the genus Macrochiron Brady, 1872 (Copepoda, Cyclo- poida). Ann. Mag. Nat. Hist., ser. 12, 10 : 378-382. THOMPSON, I. C. & SCOTT, A. 1903. Report on the Copepoda collected by Professor Herdman, at Ceylon, in 1902. Kept. Govt. Ceylon Pearl Oyster Fish. Gulf of Manaar, part I, suppl. rept. 7, pp. 227-307. UMMERKUTTY, A. N. P. 1961. Studies on Indian copepods 5. On eleven new species of marine cyclopoid copepods from the south-east coast of India. /. Mar. Biol. Ass. India, 3 (i & 2) : 19-69. EXPLANATION OF FIGURES All figures were drawn with the aid of a camera lucida. The letter after the explanation of each figure refers to the scale at which it was drawn. Abbreviations used : Ax = first antenna, A2 = second antenna, MD — mandible, P = paragnath, MXt = first maxilla, MX2 = second maxilla, MXPD = maxilliped, PI = leg i. PLATE i Stellicola kossmanni n. sp., female FIG. i. Dorsal (A). FIG. 2. Marginal membrane on prosomal segments, dorsal (B). FIG. 3. Urosome, dorsal (c). FIG. 4. Area of attachment of egg sac, dorsal (D). FIG. 5. Caudal ramus and part of anal segment, dorsal (E). FIG. 6. Rostral area, ventral (c). Bull. Br. Mus. mat. Hist. (Zool.) 15, 5 PLATE i ZOOL. 15, 5 PLATE 2 Stellicola kossmanni n. sp., female (continued) FIG. 7. First antenna, anterodorsal (c). FIG. 8. Second antenna, anterior (F). FIG. g. Labrum, ventral (D). FIG. 10. Mandible, posterior (D). FIG. ii. First maxilla, posterolateral (G). FIG. 12. Second maxilla, posterior (D). FIG. 13. Maxilliped, anterior (D). FIG. 14. Area between maxillipeds and first pair of legs, ventral (c). FIG. 15. Leg i and intercoxal plate, anterior (c). Bull. BY. Mus. nat. Hist. (Zool.) 15, 5 PLATE 2 PLATE 3 Stellicola kossmanni n. sp., female (continued) FIG. 1 6. Leg 2, anterior (c). FIG. 17. Leg 3, anterior (c). FIG. 1 8. Leg 4 and intercoxal plate, anterior (c). FIG. 19. Leg 5, dorsal (D). Stellicola kossmanni n. sp., male FIG. 20. Dorsal (A). FIG. 21. Urosome, dorsal (c). Bull. Br. Mus. nat. Hist. (Zool.) 15, 5 PLATE 3 20 V PLATE 4 Stellicola kossmanni n. sp., male (continued) FIG. 22. Maxilliped, anterior (F). FIG. 23. Leg 6, ventral (F). FIG. 24. Spermatophore, in body of male, dorsal (c). Stellicola affinis n. sp., female FIG. 25. Dorsal (A). FIG. 26. Lateral (A). FIG. 27. Urosome, dorsal (B). FIG. 28. Area of attachment of egg sac, dorsal (D). FIG. 29. Caudal ramus, dorsal (F). FIG. 30. Rostrum, ventral (c). FIG. 31. First antenna, anterodorsal (c). Bull. Br. Mus. nat. Hist. (Zool.) 15, 5 23 PLATE 4 PLATE 5 Stellicola affinis n. sp., female (continued) FIG. 32. Second antenna, posterior (F). FIG. 33. Labrum, ventral (D). FIG. 34. Mandible, posterior (D). FIG. 35. First maxilla, posterior (E). FIG. 36. Second maxilla, posterior (D). FIG. 37. Maxilliped, posterior (D). FIG. 38. Oral and postoral areas, ventral (c). FIG. 39. Leg i and intercoxal plate, anterior (c). FIG. 40. Leg 2, anterior (c). FIG. 41. Leg 3, anterior (c). Bull. Br. Mus. nat. Hist. (Zool.) 15,5 PLATE 5 PLATE 6 Stellicola affinis n. sp., female (continued) FIG. 42. Leg 4 and intercoxal plate, anterior (c). FIG. 43. Leg 5, dorsal (D). Stellicola affinis n. sp., male FIG. 44. Dorsal (A). FIG. 45. Urosome, dorsal (B). FIG. 46. Maxilliped, anterior (F). FIG. 47. Leg 5, ventral (D). FIG. 48. Leg 6, ventral (F). Stellicola longiseta n. sp., female FIG. 49. Dorsal (H). FIG. 50. Urosome, dorsal (B). Bull. Br. Mus. nat. Hist. (Zool.) 15, 5 PLATE 6 47 PLATE 7 Stellicola longiseta n. sp., female (continued) FIG. 51. Caudal ramus, dorsal (E). FIG. 52. Rostrum, ventral (c). FIG. 53. Second antenna, anterior (F). FIG. 54. Labrum, ventral (F). FIG. 55. Mandible, posterior (D). FIG. 56. Second maxilla, posterior (D). FIG. 57. Maxilliped, anterior (D). FIG. 58. Last segment of endopod of leg 2, anterior (F). FIG. 59. Last segment of endopod of leg 3, anterior (F). FIG. 60. Leg 4 and intercoxal plate, anterior (c). Bull. BY. Mus. nat. Hist. (Zool.) 15, 5 PLATE 7 55 PLATE 8 Stellicola longiseta n. sp., female (continued) FIG. 61. Endopod of leg 4, anterior (F). FIG. 62. Leg 5, dorsal (D). Stellicola longiseta n. sp., male FIG. 63. Dorsal (A). FIG. 64. Urosome, dorsal (c). FIG. 65. Maxilliped, anterior (F). FIG. 66. Leg 5, dorsal (E). Stellicola femineus n. sp., female FIG. 67. Dorsal (H). Bull. Br. Mus. nat. Hist. (Zool.) 15, 5 PLATE 8 66 PLATE 9 Stellicola femineus n. sp., female (continued) FIG. 68. Urosome, dorsal (i). FIG. 69. Area of attachment of egg sac, dorsal (F). FIG. 70. Caudal ramus, dorsal (E). FIG. 71. Rostrum, oral and postoral areas, mouthparts not labelled but in same relative positions as in Plate 5, fig. 38, ventral (A). FIG. 72. First antenna, dorsal (c). FIG. 73. Second antenna, posterior (B). FIG. 74. Tip of second antenna, anterior (F). FIG. 75. Labrum, ventral (c). Bull. Br. Mus. nat. Hist. (Zool.) 15, 5 68 PLATE 9 71 ZOOL. 15, 5. PLATE 10 Stellicola femineus n. sp., female (continued) FIG. 76. Mandible, posterior (F). FIG. 77. First maxilla, anterolateral (F). FIG. 78. Second maxilla, posterior (F). FIG. 79. Maxilliped, posterior (F). FIG. 80. Leg i and intercoxal plate, anterior (B). FIG. 81. Leg 2, anterior (B). FIG. 82. Leg 3, anterior (B). FIG. 83. Leg 4 and intercoxal plate, anterior (B). FIG. 84. Endopod of leg 4, anterior (F). Bull. Br. Mus. nat. Hist. (Zool.) 15, 5 PLATE 10 84 19§ PLATE n Stellicola femineus n. sp., female (continued) FIG. 85. Leg 5, dorsal (F). Stellicola femineus n. sp., male FIG. 86. Dorsal (A). FIG. 87. Urosome, dorsal (B). FIG. 88. Labrum, ventral (F). FIG. 89. Maxilliped, postero-inner (D). FIG. 90. Leg 5, dorsal (E). FIG. 91. Leg 6, ventral (F). FIG. 92. Spermatophore, inside male, dorsal (B). Stellicola pollex n. sp., female FIG. 93. Dorsal (H). Bull. Br. Mus. nat. Hist. (Zool.) 15, 5 PLATE ii 87 PLATE 12 Stellicola pollex n. sp., female (continued) FIG. 94. Urosome, dorsal (c). FIG. 95. Area of attachment of egg sac, ventral (D). FIG. 96. Caudal ramus, dorsal (E). FIG. 97. First antenna, ventral (c). FIG. 98. Second antenna, posterior (c). FIG. 99. Mandible, posterior (F). FIG. 100. Paragnath, ventral (G). FIG. 101. Second maxilla, posterior (D). Bull. Br. Mus. nat. Hist. (Zool.) 15, 5 PLATE 12 97 PLATE 13 Stellicola pollex n. sp., female (continued) FIG. 102. Maxilliped, posterior (D). FIG. 103. Leg i and intercoxal plate, anterior (c). FIG. 104. Leg 2, anterior (c). FIG. 105. Leg 3, anterior (c). FIG. 1 06. Leg 4 and intercoxal plate, anterior (c). FIG. 107. Endopod of leg 4, anterior (E). FIG. 108. Leg 5, dorsal (D). Stellicola pollex n. sp., male FIG. 109. Dorsal (i). Bull. Br. Mus. nat. Hist. (Zool.) 15, 5 PLATE 13 104 PLATE 14 Stellicola pollex n. sp., male (continued) FIG. no. Urosome, dorsal (F). FIG. in. Third segment of second antenna, anterior (E). FIG. 112. Maxilliped, postero-inner (D). FIG. 113. Leg 5, dorsal (G). FIG. 114. Leg 6, ventral (D). Stellicola oreastriphilus Kossmann, 1877, female FIG. 115. Genital segment, dorsal (c). FIG. 1 1 6. Tip of second antenna, posterior (F). FIG. 117. Paragnath, ventral (E). FIG. 1 1 8. Maxilliped, anterior (D). FIG. 119. Endopod of leg 4, anterior (F). Stellicola oreastriphilus Kossmann, 1877, male FIG. 1 20. Segment of leg 5 and genital segment, dorsal (c). Bull. BY. Mus. nat. Hist. (Zool.) 15, 5 PLATE 14 118 PLATE 15 Stellicola caeruleus (Stebbing, 1900), syntypic female FIG. 121. Dorsal (H). FIG. 122. Urosome, dorsal (B). FIG. 123. Caudal ramus, dorsal (E). FIG. 124. Rostrum and labrum, ventral (i). FIG. 125. First antenna, ventral (c). FIG. 126. Second antenna, anterior (B). FIG. 127. Mandible, posterior (F). FIG. 128. Second maxilla, posterior (F). Bull. Br. Mus. nat. Hist. (Zool.) 15, 5 PLATE 15 122 PLATE 16 Stellicola caeruleus (Stebbing, 1900), syntypic female (continued) FIG. 129. Maxilliped, posterior (F). FIG. 130. Leg i and intercoxal plate, anterior (c). FIG. 131. Leg 2, anterior (c). FIG. 132. Leg 3, anterior (c). FIG. 133. Leg 4, anterior (F). FIG. 134. Leg 5, dorsal (D). Bull. Br. Mus. nat. Hist. (Zool.) 15, 5 PLATE 16 134 PLATE 17 Stellicola caeruleus (Stebbing, 1900) syntypic male FIG. 135. Dorsal (i). FIG. 136. Urosome, dorsal (F). FIG. 137. Rostrum, ventral (c). FIG. 138. Second maxilla, dorsal (E). FIG. 139. Maxilliped, dorsal (G). FIG. 140. Leg 5, dorsal (G). FIG. 141. Leg 6, ventral (E). Bull. BY. Mus. nat. Hist. (Zool.) 15, 5 PLATE 17 135 140 136 137 138 139 PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING H. G. STUBBINGS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 15 No. 6 LONDON: 1967 THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA BY H. G. STUBBINGS Admiralty Materials Laboratory, Holton Heath, Poole Pp. 227-319 ; 28 Text-figures, I Plate BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 15 No. 6 LONDON: 1967 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the 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. 15, No. 6 of the Zoological 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.). Trustees of the British Museum (Natural History) 1967 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 28 July, 1967 Price £2 THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA By H. G. STUBBINGS CONTENTS Page SYNOPSIS ........... 229 1. INTRODUCTION .......... 229 2. SYSTEMATIC ACCOUNT ......... 232 3. THE WEST AFRICAN CIRRIPED FAUNA ...... 307 4. SUMMARY ........... 313 5. REFERENCES .......... 314 SYNOPSIS Sixty-nine species of Cirripedia are known from the tropical and warm-temperate waters of West Africa. This number includes littoral and shallow-water species and a very few from deep water. Further deep-water species, particularly of Scalpellum and Verruca may be expected when the deep basins off tropical West Africa are investigated more thoroughly. For this reason several species of Scalpellidae from the warm-temperate waters off north-west Africa, e.g. Smilium longirostrum (Gruvel) and Scalpellum velutinum Hoek, are included. The status of many rare or poorly recorded species requires further elucidation. The absence of the genus Acasta is remarkable. The fauna contains elements from the north, Scalpellum Scalpellum (L.) and Chthamalus stellatus (Poli) and from south-west Europe and the Mediterranean as would be expected from the published knowledge of the distribution of other groups of animals on this coast. Some of these species extend no farther south than Cape Verde, others extend to the south of the Gulf of Guinea. There is a very small South African component, of which Chthamalus dentatus Krauss is the most note-worthy. Four species, Smilium renei (Gruvel) Chthamalus aestuarii Stubbings, Balanus pallidus Darwin and Chelonibia manati Gruvel appear to be 'indigenous to tropical West Africa south of Cape Verde. The faunistic boundary in the region of Cape Verde noted by workers on other groups applies to many species of barnacle also. I. INTRODUCTION IN considering the fauna of tropical West Africa it is appropriate to set limits to the region other than those imposed by the tropics of Cancer and Capricorn, to exclude those areas of coast, north and south, under the influence of the Canary and Benguela currents. Nickles (1950) denned the " West Coast of Africa " for purposes of listing the Molluscan fauna, as the area between Cap Blanc in Mauretania and Mossamedes, Angola, that is approximately from 21° N. Lat. to 15° 30' S. Lat. Longhurst (1962), reviewing the oceanography of the Gulf of Guinea has concluded that 14° N. and 14° S. Lat. present important faunistic boundaries corresponding to oceanographic frontal zones in these latitudes. These limits are in close agreement with those proposed by Ekman (1953) namely 15° N. and 15° S. (or possibly i6°-i7°S.). This northern limit at 14-15° N. corresponds approximately to Cape Verde, Senegal, and there is evidence of a change of Cirriped fauna in this area, north temperate warm water ZOOL. 15, 6. 20 230 H. G. STUBBINGS species giving place to sub-tropical or tropical forms. In the present work Long- hurst's (1962) limits of 14° N. and 14° S. are regarded as denning the tropical coast of West Africa but species from both farther north and south, which it seems likely may occur within these latitudes, have been included. There is adequate material available from several sources linking the West African tropical fauna with that of the Atlantic north temperate zone. To the southward there is still a gap in our knowledge due to the virtual absence of information from south west Africa. In consequence the extent to which South African species pene- trate northwards towards Angola is uncertain. Broch (1924) summarized knowledge of the west African shallow-water barnacle fauna to that date. Few species had been recorded and many of the records gave vague localities, e.g. " West Africa ". Later works added only a little new informa- tin, but exploration since 1945 has increased our knowledge greatly. The " Atlan- tide " expedition, " Calypso " expeditions to the Gulf of Guinea and to the Cape Verde Archipelago and the Belgian Atlantique Sud expedition have all contributed. In addition collections made by the Museum d'Afrique Centrale (Tervuren), the Institut Fran$ais d'Afrique Noire, Dakar and the, now defunct, West African Fisheries Research Institute, Sierra Leone have all contributed specimens. The present work reports on an extensive collection made in Ghana by Mr. R. Bassindale in 1949-51 and on a smaller one made by the author in Nigeria in 1957-8. Unpublished West African material in the British Museum is included. In order to make this report fully representative of the Cirriped fauna of the region, all species recorded by previous authors are included, provided the records are considered valid. The few species not included individually appear under the synonymy of species with which they have been merged since the original publication of the record. The species and varieties omitted are as follows : Chthamalus stellatus depressus: Nilsson-Cantell (1938), Gauld (1957), C. cirratus: Kolosvary (19416, 19430), C. withersi: and C. rhizophorae: Longhurst (1958) = C. aestuarii Stubbings. Balanus amphitrite var. hawaiiensis: Stubbings (1963) and B. amphitrite var. denticulata: Stubbings (19610) = B. amphitrite amphitrite Darwin. B. dybowskii Gruvel (1903) and Broch (19240), B. amphitrite var. stutsburi Darwin (1854) ^ auct., B. pallidus stutsburi: Harding (1962) = B. pallidus Darwin. B. dollfusi Broch (1927) and Nilsson-Cantell (1938, 19396) = B. spongicola Brown. B. occidentalis Stubbings (19610, b) = B. fallax Broch. It is probable that the littoral and shallow water Cirriped fauna of tropical Africa is now fairly well known and that few species remain to be discovered at least in the Lepadomorpha and Balanomorpha. There is still need for more information on the occurrence of the species of intermediate depths, i.e., of the continental shelf. The geographical range of Scalpellum renei Gruvel is uncertain and that of Ibla atlantica sp. n. virtually unknown. It is not improbable that intensive investigation of these intermediate depths will reveal further species. THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 231 Post-war investigations into the Acrothoracica have revealed several new West African species (Stubbings, 19616, 19646; Tomlinson, 1960) and there would appear to be a favourable field here for further work. The Ascothoracica have not been investigated so far. The considerable development of Zoanthid colonies in some areas, as on parts of the Ghana coast, would suggest the presence of these parasitic Cirripeds. The deep-water fauna off the west coast of Africa has not been investigated to any extent. This is in marked contrast to the warm temperate region to the northward in which extensive work in the area of Madeira and the Azores has yielded many species notably of Scalpellum and Verruca (Aurivillius, 1898; Gruvel, 19000, b). The German Tiefsee-Expedition made a number of stations down the coast in deep water from which which the deepwater Scalpellids, Smilium longirostrum and Scalpellum trapezoideum, were obtained (Weltner, 1922). As Gruvel (1905) could list 25 species of Scalpellum and 15 of Verruca from the northern warm temperate area it is probable that more species will be found when the deep waters of the Cape Verde, Sierra Leone, Guinea and Angola basins are worked systematically. The following list of 70 species and varieties includes all those known to occur within the specified area plus a few that have been found outside it but may well be found there : Mitella potticipes (Gmelin) Smilium longirostrum (Gruvel) S. renei (Gruvel) Scalpellum scalpellum (Linn.) 5. imperfectum Pilsbry 5. trapezoideum Hoek 5. velutinum Hoek Ibla atlantica sp. n. Lepas anatifera Linn. L. anserifera Linn. L. hillii Leach L. fascicularis Ellis and Solander L. pectinata Spengler Heteralepas cornuta (Darwin) Paralepas minuta (Philippi) Conchoderma auritum (Linn.) C. virgatum (Spengler) C. virgatum var. chelonophilum Leach Trilasmis (Poecilasma) crassum (Gray) T. (P.) kaempferi (Darwin) Octolasmis tridens (Aurivillius) 0. lowei (Darwin) 0. hoeki (Stebbing) Verruca striata Gruvel Chthamalus stellatus stellatus (Poli) C. stellatus bisinuatus Pilsbry C. dentatus Krauss C. aestuarii Stubbings C. fragilis Darwin Pachylasma giganteum (Philippi) Balanus (Megabalanus) tintinnabulum tintinnabulum Linn. B. (M.) tintinnabulum maroccana Broch B. (M.) tintinnabulum zebra Darwin B. (M.} tintinnabulum concinnus Darwin B. (M.) tintinnabulum spinosus (Gmelin) B. (M.) tulipiformis Darwin B. (M.} nigrescens Lamarck B. (M.) maxillaris (Gronovius) B. (Balanus) trigonus Darwin B. (B.) spongicola Brown B. (B.) perforatus Brug. B. (B.) perforatus var. angustus Gmelin B. (B.) perforatus var. fistulosus Poli B. (B.) eburneus Gould B. (B.) improvisus Darwin B. (B.) improvisus var. assimilis Darwin B. (B.) amphitrite amphitrite Darwin B. (B.) amphitrite albicostatus Pilsbry B. (B.) pallidus Darwin 232 H. G. STUBBINGS B. (B.) venustus venustus Darwin B. (B.) venustus niveus Darwin B. (Hesperibalanus) fallax Broch B. (Conopea) calceolus Darwin Tetraclita divisa Nilsson-Cantell T. purpurascens Wood T. squamosa squamosa (Brug.) Pyrgoma anglicum Sowerby Chelonibia testudinaria Ellis C. caretta Spengler C. manati Gruvel C. patula (Ranzani) Coronula complanata (Morch) Platylepas hexastylos (O. Fabr.) Stomatolepas elegans (O. G. Costa) Xenobalanus globicipitis Steenstrup Cryptophialus coronatus Tomlinson C. variabilis Stubbings Kochlorine hamata Noll K. inermis Stubbings Kochlorinopsis discoporellae gen. sp. n. et. References under species synonymy have been reduced to as few as possible and are only given when the species is but little known, though originally described long ago, e.g. Smilium renei (Gruvel, 1902) or when the nomenclature is questioned or has been recently, e.g. Chthamalus aestuarii Stubbings, 1963. References to geo- graphical distribution are given only in the text. Locality citations for material reported here for the first time are arranged geo- graphically according to country of origin from North Africa to the Cape. Mr. Bassindale carried out a large number of trawl and dredge hauls in a circumscribed area off Accra, Ghana, in addition to collecting at many named shore localities. The Accra stations are referred to here by their numbers. For full available details of these collecting stations reference should be made to Bassindale (1961). Mr. Bassindale's material comprises the largest part of that described here. To avoid excessive repetition his name does not appear in the locality citations. Other sources of materials are distinguished by citation of the collecter's name and where appropriate the British Museum registration number. The author's own collections are followed by his initials (H.G.S.) with or without a serial collector's number as appropriate. Samples collected by the Staff of the Tropical Testing Establishment of the Ministry of Supply in 1953-4 are designated by the initials T.T.E. The " dolphin " mentioned in some references to Port Harcourt, Nigeria, is a concrete structure erected in the river to aid in warping out ships from the quay but no longer used. 2. SYSTEMATIC ACCOUNT LEPADOMORPHA MITELLA Mitella pollicipes (Gmelin) Mitella pollicipes: Stubbings, 1965, 877. LOCALITIES : Cape Verde Is.: (i) St. Vincent, coll. Mrs. M. G. Bannerman. (2 spec.) B.M. 1915.1.11.1-2. (2) coll. C. R. Stonor. B.M. 1939.7.22.9-13 (21 large and 6 small spec.). THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 233 The distribution of M. pollicipes is rather clearly defined. On the European Atlantic coast it is found from Finisterre to Spain and Portugal (Bishop et al., 1957 ; Gruvel, 1920 ; Fischer-Piette & Prenant, 1956, 1957). In the Mediterranean, Llabador (1937) has established its occurrence on the Algerian coast from Algiers westward. From the northern shore there is a doubtful reference to the species from Nice (Caziot, 1921) based on a century-old record of Verany (1862) but it is not recorded from Banyuls (Utinomi, 19590). Kolosvary (1940) records it from the Mediter- ranean without precise locality. Most recently Barnes & Barnes (1964 : 9) record it from Catalan Bay, Gibraltar. The textual name polymerus in this paper is a MS error (Auct. in litt.). From Algeria it extends westward and southward on the African coastline and is found in suitable situations in Morocco, Rio de Oro, Mauretania and Senegal (Broch, 19270 ; Gruvel, 1912 ; Nilsson-Cantell, 19396 ; Stubbings 1965). Sourie (1954) states that it is well represented at Dakar and fairly common at Cap Blanc (Maure- tania) . He did not find it at Conakry, French Guinea and there seems no doubt that Dakar defines the southern limit of the species in West Africa. From the Atlantic islands there are very few records. The above noted material in the British Museum confirms the presence of M. pollicipes in the Cape Verde archipelago. Its absence from the " Calypso " Cape Verde collections (Stubbings, 19640) suggests, however, that it is not very common there. From the Canary Is. there are no recent records, but Darwin (1851) clearly handled material from Teneriffe. Its presence, or absence, in the Azores likewise needs elucidation. Weltner (1897) records specimens from " Portugal oder Acoren " but there is no certain record from these islands. The inclusion of England, Scotland, Ireland, the North Sea and even Jan Mayen within the area of distribution of M. pollicipes (Darwin, 1851 ; Weltner, 1897 ; Gruvel, 1905) would seem highly improbable except for chance specimens on drift, an unusual habitat for the surf -loving Mitella. The Indonesian record of Tonkin (Gruvel, 1912) likewise is improbable. Smilium renei (Gruvel) Scalpellum renei Gruvel, 19026, 229-233, pi. 12, figs. 5-7. Scalpellum (Smilium) renei: Pilsbry, 1907, 13. Smilium renei: Stubbings, 1961 a, 9—11, text-fig, i. Smilium renei: Stubbings, ig6ib, 181. LOCALITY : Ghana : Off Accra Sta. 72, 38m, on fine branches of an Alcyonarian (2 spec.). One specimen measured, capitulum 3-0 mm., peduncle 1-5 mm., and the other capitulum 2-0 mm., peduncle 0-75 mm. These are small compared with Gruvel's type specimen which has capitular and peduncular lengths each 5-25 mm., and the largest Atlantide specimens which had a capitular length of about 7-0 mm. The species has now been recorded from French Guinea, Ivory Coast, Ghana, Nigeria and Angola, thus establishing its presence as a shallow-water species through- out tropical West Africa from approximately 10° N. to 10° S. Lat. 234 H. G. STUBBINGS Smilium longirostrum (Gruvel) Weltner (1922) records this species in 2480 m. south of the Canary Islands, Lat. 24° 35' N., Long 17° 4' W., i.e., somewhat to the north of the area under discussion. It is thus a species that may be expected to occur in the deep water basins off tropical West Africa. Scalpellum scalpellum (Linn.) LOCALITIES: (i) Gambia, on carapace of the crab Acanthocarpus africanus, coll. A. R. Longhurst, B.M. 1956.1. 7. i.(i spec.); (2) Sierra Leone, W.A.F.R.I. sta. MBi/A4, on continental shelf 88 m., on a spine of Cidaris cidaris meridionalis, coll. A. R. Longhurst, B.M. 1956.4.27.1. (i small spec.). The southward distribution of Sc. scalpellum in coastal waters of the sub-tropical and tropical eastern Atlantic is now fully documented. It has been reported from the western Mediterranean (Utinomi, 19590, b) and Adriatic (Broch, 19536), Morocco and Rio de Oro (Broch, 19270 ; Gruvel, 190212), Canary Is. (Gruvel, 1920), Cape Verde Is. (Stubbings, 19640), Senegal and Gambia (Stubbings, 1965), Sierra Leone (Stubbings, 19610), Nigeria (Stubbings, 19616), Gabon and the Congo estuary (Stub- bings, 1963). This species is now known to have a vast range in latitude from Iceland and Lofoten, Norway (Lat. 68° N.) (Broch, I924c) to the Congo (Lat. 6° S.). Scalpellum imperfectum Pilsbry Scalpellum imperfectum Pilsbry, 1907, 75-77, pi. 4, figs. 15-18, text-fig. 30. Scalpellum imperfectum: Stubbings, ig6ia, 11-13, text-fig. 2. Not represented in the Ghanaian and Nigerian collections. The few records of this species indicate a very wide geographical range. In the Atlantic there are three records from eastern North America (Pilsbry, 1907), one from south of Iceland (Broch, 19530), one from off Spanish Guinea (Stubbings, 19610) and one from the Cape (Barnard, 1924). There is a further record from the Galapagos Is. (Macdonald, 1929). Scalpellum trapezoideum Hoek A single specimen of this species is recorded by Weltner (1922) from the Bight of Biafra in 2492 m. Lat. 2° N. Long. 8° 4' E. It remains the single deep water species recorded from tropical West Africa as here defined. Scalpellum velutinum Hoek Scalpellum velutinum has not been found within the area but the " Talisman " took it in 882 m. off Pilones, Rio de Oro, as well as at two stations in Morocco and at Fuerteventura, Canary Islands (Gruvel, 19020). It is, therefore, to be expected that S. velutinum will be found in the deep basins south of Cape Verde. THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 235 Ibla atlantica sp. n. LOCALITY: Sierra Leone: (i) W.A.F.R.I. Sta. MBi/B4, o8°45'N., 14° 38' W., 220-440 m. (i spec.) : Sta. MBi/B5, same position, 800 m. in burrows of Pholadidea prox. loscombiana Turton (Lamell., Pholadidae) (3 spec.). DIAGNOSIS : Unisexual : female with peduncle clothed in golden spines ; opercular valves golden brown with prominent growth ridges : no male organs : caudal appendages minute, one-segmented : cirri with numerous segments, increasing in number from Ci to C6. Male differentiated into peduncle and capitulum, attached to mantle wall by the antennae. TYPE LOCALITY: as above, Guinea shelf off Sierra Leone, West Africa (see Longhurst, 1958, p. 73 and text-fig. 2). Holotype in British Museum B.M. 1956.4.27.3, a specimen with long peduncle from sta. MBi/B5: paratypes: i) 2 other specimens from MBi/B5, one dissected registered number B.M. 1956.4.27.4. and slides reg. no. B.M. 1956.4.27.6 and 2) i specimen from sta. MBi/B4, reg. no. B.M. 1956.4.27.5. FEMALE : Valves golden brown with prominent growth ridges : scutum curved slightly towards tergum, the occludent margin slightly convex. Peduncle paler in colour, densely clothed in short golden hairs curved towards the valves. Length of peduncle variable, the lower portion narrowed and embedded in the wall of the mollusc burrow. Size 9-5 (5-0) mm. (MBi/B4) ; 14-5 (4-0) (Text-fig. la), 8-5 (3-5) FIG. i. Ibla atlantica sp. n. : a, type specimen, 14-5 mm. overall, with long peduncle (x c. 5) ; b, paratype, 8-5 mm. overall from same haul (x8-5) ; c, labrum (X47) ; d, mandible; e, maxilla I (both Xi35) ; /, paired maxilla II (X47). 236 H. G. STUBBINGS (Text-fig. ib) and 6-5 (3-0) mm. (MBi/B5) overall. Figures in brackets are the respective lengths of the valves. Labrum (Text-fig, ic) smoothly concave without teeth or hairs on margin. The palps are small, oval with a few short terminal setae. The mandible (Text-fig, id), bears denticles on the upper margin of the second and third teeth. The lower angle carries a tuft of short stout setae. The first maxilla (Text-fig. le) has a convex margin with ten stout spines. Maxilla II (Text-fig, if) has rather short terminal setae and very short setae on both inner and outer margins. The lower lobe is almost as large as the main lobe. The two appendages are set wide apart and there is a raised fold posterior to them " closing the gap " as described by Darwin. The cirri have short segments rather broader than long near the base but elongating somewhat distally. There are one or two setae distally on the posterior margin. On the anterior margin each segment bears one pair of large setae distally with a single fine seta beside or between them. A smaller, often very small, pair of setae lies below the main pair. The number of segments in the cirri is high : Cirrus I II III IV V VI Segments 14-22 27-31 36-35 35-37 39'39 4°'42 The caudal appendages are minute consisting of one oval lobe less than half the length of the pedicel of cirrus VI. The mantle cavity contained nine minute individuals all apparently attached to the mantle wall, though six of them were early embryos. Two embryos had a single well-developed pigmented eye (? nauplii) and only one was recognizable as a young male probably recently metamorphosed as the single eye was still visible. The male has clearly differentiated capitulum and peduncle and is attached by the antennae embedded in a mass of cement in the mantle wall. The black nauplius eye is still visible. The lobes of the cement gland are clearly visible within the capitulum and adjacent to the peduncle. The prosoma was poorly preserved and did not stain well. Little can be made out but an elongated slightly dumb-bell shaped structure, staining deeply probably represents the vesicula seminalis and male reproductive organs. The portion towards the base of the capitulum may be double, the two lobes lying one over the other, in which case two testes are present. This struc- ture is obscured further by the presence of the cement gland. No valves are discernible. Including Ibla atlantica, five species of Ibla are now recognized namely, I. quadrivalvis Cuvier 1817 /. cumingi Darwin 1851 I. segmentata (Studer 1889) syn. I, pygmaea Broch 1922 I. idiotica Batham 1945 /. atlantica sp. n. Two of these species, I. quadrivalvis and /. segmentata are hermaphrodite. /. quadrivalvis has extremely long caudal appendages composed of 32 segments and up THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 237 to three-quarters the length of the rami of cirrus VI. /. segmentata has fairly long caudal appendages of about 10 segments, longer than the pedicel of cirrus VI. Of the other three, female, forms without penis, only /. cumingi has long caudal appen- dages. /. idiotica resembles the new species in the minute caudal appendages, but differs in having only a few segments, up to 8, in the short rami of the cirri and in its very small size. There is no doubt that /. atlantica represents a new species of Ibla. This is the first record of the genus from the Atlantic ocean. Although many records of Ibla spp. note the association with larger Cirripedia (Batham, 1945 ; New- man, 1960) this association has been shown not to be obligatory. The occurrence of Ibla in the burrows of an unrelated organism, as here, seems to be a hitherto un- recorded phenomenon. Lepas anatifera Linn. Lepas anatifera: Gauld, 1957, 10. Lepas anatifera: Stubbings, 19610, 13-14. Not represented in the Ghanaian and Nigerian collections. This species was collected from the hull of the Danish Expedition ship " Atlantide " at ports in Liberia, Ghana and Nigeria (Stubbings, 1961^). The duration of the ship's stay in West African waters, six weeks before the earliest of these collections, leaves no room for doubt that L. anatifera is present in those waters and was not brought from further north by the ship. There are few other records from the west coast. Gauld (1957) records this species from beaches in Ghana and Weltner (1897) specimens from the Cameroons. Evans (1958) has discussed the growth rate of specimens on the hull of " Petula " which must have settled in the vicinity of the Cape Verde Is. From further north there are records by Gruvel (1920) and Broch (19270) from Morocco. There are many records from the Azores (Gruvel, 1920). The British Museum has recent material from the Azores (B.M. 1955.9.2.2-3, coll. G. Chapman) and from Funchal, Madeira (B.M. 1954.9.15.4 coll. D. W. Tucker). L. anatifera is much less common on the West Coast than L. anserifera and from the greater number of records from more temperate waters it is possible that L. anatifera prefers rather lower temperatures than obtain in the tropical Eastern Atlantic. Lepas anserifera Linn. 1767 Lepas anserifera Gruvel 1912, 344-345. Lepas anserifera "" Lepas anserifera Gruvel 1912, 344-345. Broch 19240, 202. Stubbings 19610, 14 ; 1963, 4-5 ; 19640, 104 ; 19646, 330. LOCALITIES: (i) Sierra Leone, on a floating plant stalk, — .2.55, coll. A. R. Longhurst, B.M. 1956.1.7.2 (28 almost full-grown spec.); (2) " off West Africa", coll. A. R. Longhurst, B.M. 1957.6.3.2 (3 part-grown spec.). Ghana: (i) Accra, on cuttlefish "bone", 1935 coll. F. R. Irvine (5 spec.); (2) Takoradi, coll. Mrs. Carlyle Bell, B.M. 1948.1.17.1 (i spec.); (3) Takoradi, on 238 H. G. STUBBINGS floating seeds, 6.1.55, coll. D. T. Gauld (5 spec.); (4) Prampram, — .11.37, C°U- Miss V. J. Foote, B.M. 1952 .5.7.1 (many very small spec.) ; (5) Senegal, on Sargassum coll. Captain Moloney, B.M. 85-5 (9 small spec.). Nigeria, Lagos : (i) on cuttlefish " bone " coll. Miss E. Trewavas, B.M. 1948 . 3 . 20 . i (27 nearly full grown spec, and some very small ones) ; (2) coll. Captain Moloney, B.M. 91.4.1.59-63 (10 large and 12 very small spec.). Cameroons: (i) Off Fernando Po. coll. R. C. Ward, B.M. 1962.4.17.1 (34 spec.); (2) Debunscha Beach, Victoria, from a log 21.5.61, coll. J. T. Swarbrick, B.M. 1962.8.1.1 (about 90 spec, in small clusters). Specimens in each of these collections have been checked for the possession of five filamentary appendages. The Debunscha beach specimens are of interest in respect of the very white shells with no bluish shadowing from the pigmented mantle and for the deep orange colour of the edge of the mantle. In addition to the authors cited above, Weltner (1897) and Nilsson-Cantell (1921) record L. anserifera from the Cameroons. It is evidently the most widely distributed and frequently occurring species of Lepas in Tropical West Africa. Weltner (1897) records " L. pectinata Spengler var. Darwin " from " West Africa " and Brian and Dartevelle (1954) record L. pectinata from Moita Seca at the mouth of the R. Congo. As reported elsewhere (Stubbings 19646) I can find no L. pectinata in Dr. Dartevelle's material from Congo that I have examined all of which appears to be L. anserifera. Lepas hillii (Leach) 1818 LOCALITY: Ghana, Tenpobo : on cuttlefish "bone", 1.2.50, coll. R.Bassindale (many small spec.). These specimens, preserved in Bouins fluid, were completely decalcified and spoiled. From the presence of three filamentary appendages, a pair at the base of cirrus I and a single one on the prosoma, and apparently a gap between carina and scutum, they have been referred to L. hillii. The above specimens appear to be the only ones recorded from the West African coast. Evans (1958) recorded them on the hull of " Petula " and deduced that they must have settled there shortly after the vessel left Dakar. They may be presumed to have settled during the voyage between that port and the Cape Verde Is. Hoek's (1883) specimens taken from the screw of H.M.S. " Challenger " at St. Vincent, Cape Verde Is. and those removed from " Atlantide " in Le Havre (Stubbings, 1961^) have a less certain provenance. Specimens have been taken off Morocco (Aguilar-Amat, 1927) and at Tangier (Gruvel, 1912) and many off the Azores (Gruvel, 19076, 1920) and in the W. Mediterranean (Utinomi, 1959^). L. hillii is thus distributed over the eastern tropical and sub-tropical North Atlantic, but evidently rather few specimens occur in-shore. It is not represented in the collections of the Institut Fran$ais d'Afrique Noire in Dakar or in those of the Musee Royal de 1'Afrique Centrale, Tervuren. THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 239 Lepas pectinata Spengler non Lepas pectinata: Brian & Dartevelle 1954, 150. There is no reliable evidence of the occurrence of Lepas pectinata on the tropical West African coast. The only precise record in the literature is that of Brian and Dartevelle (1954) from the Congo estuary which the author has given reasons for referring to L. anserifera (S tubbings 19646). Weltner (1897) recorded the " var. Darwin " from W. Africa without precise locality. Further, Gruvel (igoja) recorded the species from " West of the coast of Guinea " and (1910) gave the same locality statement and also Porto Grande, St. Vincent, C. Verde Archipelago. The author has seen no material in the several collections handled that is referrable to L. pectinata and so considers it at best a doubtfully West African form. In 1966 R.R.S. " Discovery " took some recently metamorphosed Lepas off Fuerteventura, Canary Islands. These can be assigned with some degree of confidence to L. pectinata. Thus, after a lapse of over 50 years reliable L. pectinata material has been obtained from the eastern Atlantic. There is a reasonable possibility, therefore, that it will be found sooner or later in inshore or estuarine waters of tropical West Africa. Lepas fascicularis Ellis and Solander Not represented in the Ghanaian and Nigerian collections. The paucity of records of L. fascicularis from the west coast has been noted (Stubbings, 1965). That so few specimens are contained in the Institut Francais d'Afrique Noire, in Dakar can only indicate its infrequent occurrence in the Senegal area. Its scarcity on the Ghana coast and off Sierra Leone must be presumed for the same reasons. It has been recorded once from Morocco (Broch, 1927^) and once west of the Canary Islands (Gruvel, 1920). Available evidence thus suggests that L. fascicularis is not common on the north-west coast or west coast north of the equator. Its apparent abundance off the Congo estuary (Stubbings, 1963) is in complete contrast. Heteralepas cornuta (Darwin) Not represented in the Ghanaian and Nigerian collections. The distribution of H. cornuta is still very imperfectly known. The records given by Broch (1927*1) and Stubbings (1964(2, 1965) suggest that it is present over a wide area of the north-western part of the coast of West Africa from the mouth of the Mediterranean to Cape Verde and the Cape Verde Archipelago. As it was not represented in collections from Ghana, Nigeria, the Gulf of Guinea, the Congo estuary and Angola it may well be that it is absent south of Cape Verde. This, however, would be surprising in view of its occurrence much further afield in the West Indies (Darwin, 1851) and the Andaman Sea (Nilsson-Cantell, I938a). H. cornuta is apparently a deep-water species, all records being from below 100 m. and one at least from 750 m., so the lack of records from the tropical and southern sub- tropical coasts of West Africa may be due to the comparatively small number of deep-water collections so far made there. 240 H. G. STUBBINGS Paralepas rninuta (Philippi) LOCALITY: Sierra Leone: (i), W.A.F.R.I. Sta. MBI/A5, 132 m., 22.2.56, from near base of primary spines of Cidaris cidaris meridionalis B.M. 1956.4.27.2 : (2) the same, B.M. 1956. 4. 27. 2A. (33 juv. spec.). The recorded localities for P. minuta were listed by Stubbings (19616) . It occurs from Sicily in the central Mediterranean westwards to Gibraltar and down the African coast as far as Liberia (4° 40' N.), the most southerly record being that made by the " Gazelle " (Studer, 1882, 1889). According to Broch (19270) it occurs also in Madeira. The geographical distribution of P. minuta off north-west Africa is, therefore, somewhat greater than that of H. cornuta which is as yet unknown from the Mediterranean or from south of Cape Verde. Conchoderma auritum (Linn.) Not represented in the Ghanaian or Nigerian collections, C. auritum has been recorded from West African waters by Gruvel (1910), Weltner (1922), Kolosvary (19430) and Stubbings (19610). Three of these reports record material from a ship's hull, the fourth (Kolosvary, 19430) from a whale. Where precisely in tropical Atlantic waters settlement occurred cannot be determined. Broch (19240, 19270) summarized the then known occurrence of the species in the tropical and sub-tropical Atlantic but added no new localities. As there are records from the Mediterranean (Monod, 1938 ; Nilsson-Cantell, 19320) and South Africa (Barnard, 1924 ; Nilsson- Cantell, 19300) and from off-shore West African waters (Weltner, 1887) a general distribution of this pelagic species in warm-temperate and tropical eastern Atlantic waters may be presumed. Conchoderma virgatum (Spengler) LOCALITY : Madeira, Funchal, 12 juv., Aug. 1953, coll. D. W. Tucker, B.M. 1954.9.15.2 (12 juv. spec.). Not represented in the collections from Ghana and Nigeria. Records of this cosmopolitan species refer almost entirely to specimens recovered from the hulls of ships either in tropical or warm temperate waters or shortly after moving into cooler waters. The major exception is that series of specimens collected mainly from fishes at Goree, Senegal in the collection of I.F.A.N., Dakar (Stubbings, 1965)- Records north of the equator range from 2° N. (Nilsson-Cantell, 19300) to nearly 42° N. (Gruvel, 1920). South of the equator there are fewer records except for South Africa (Barnard, 1924, 1925). It may be expected anywhere off the West African coast where flotsam comes ashore or craft are moored or ply for long periods in warm waters. var. chelonophilum Leach Not represented in the collections from West Africa. There are a number of records from the Atlantic in Lat. 30°-40° N. (Hoek, 1883 ; Leach, 1818 ; Weltner, 1897 ; Gruvel, 1920) and Chevreux and de Guerne (1893) have recorded it from the western Mediterranean. The specimens reported by THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 241 Stubbings (19610) were very small and could also have come from these warm temperate waters and not from tropical West African waters. Accordingly, the presence of this variety in West African waters requires confirmation. Trilasmis (Poecilasma) crassum (Gray) Weltner (1922) records this species from Grosse Fischbai (= Tiger Bay), Angola. As it was originally described from Madeira (Darwin, 1851) it is conceivable that it may yet be found within the report area. Trilasmis (Poecilasma) kaempferi (Darwin) Not represented in the Ghanaian and Nigerian collections. There are numerous records of this species from the warmer North Atlantic, particularly from the region of the Azores (Gruvel, 19026, 1920). It is also known from Madeira (Darwin, 1854, Weltner, 1897). From the West Coast of Africa there are comparatively few records : from Cape Bojador, Morocco (Gruvel, 19020), Senegal (Stubbings, 1965) ; Spanish Guinea (Stubbings, 19610) and in the South Atlantic from Grosse Fischbai (Weltner, 1922). Barnard (1924) records the species from Cape Point and off the Buffalo River, South Africa. T. (P.) kaempferi is therefore widely distributed in the eastern Atlantic at depths of 100 to 1,500 m. and can be expected off the West African coast wherever local conditions are favourable. Octolasmis tridens (Aurivillius) The only record of this species from West Africa is that from the Cape Verde Islands collected by the Danish " Atlantide " Expedition (Stubbings, 19610). About 100 specimens of 0. tridens were found on that occasion on a single Palinurid crawfish, yet no other specimens seem to have been collected from either the Islands or the African mainland. Octolasmis lowei (Darwin) 1851 LOCALITIES : Mauretania : Cap Blanc, on gills of a crab, coll. M.H. Routh, B.M. 1952 .10.2.7 (many spec.) . Ghana: (i) Accra, on the Crab Callinects gladiator coll. D. J. Gauld (28 spec.); (2) off Chorkor, on gills and in branchial chamber of the crab Apiomithrax bocagei (Osorio) (numerous spec.) ; (3) off Chorkor, in branchial cavity of Calappa rubroguttata Herklots (i spec.). The largest Ghanaian specimen had a capitular length of 2-5 mm. and a peduncle of 3-0 mm. A number contained developing eggs. The species was described by Darwin from Madeira and is recorded by Broch (19246) from Mauretania and by Brian & Dartevelle (1954) from Luanda, Angola. It is evidently distributed over both the tropical and subtropical West African coast-line. Gauld (1957) records D. lowei from Callinectes gladiator and has found it also on species of Neptunus, Cronius, Micropisa, Dromia and Palinurus (Buchanan, 1958 : 23). Brian & Dartevelle (1954) cite also Echinoplax and Maia. The presence of a single specimen of 0. lowei in the branchial space of Calappa rubroguttata is note- worthy as Gauld states that this crab is the only one examined by him in large num- 242 H. G. STUBBINGS bers without finding this cirripede. A number of Callinectes collected at Port Harcourt were quite free from Octolasmis, perhaps due to the lowered salinity, which may fall to only 12 %0 in the wet season. Octolasmis hoeki (Stebbing) Dichelaspis hoeki Stebbing, 1895, 18-19, pi- H, figs. A-D. non Dichelaspis antiguae Stebbing, 1895, 19-20, pi. II, figs. E-G. Octolasmis hoeki: Nilsson-Cantell, 1927, 763-6, text-fig. 9. LOCALITY : Ghana, Tema ; on Scyllarides sp. 2.9.59, coll. D. T. Gauld (n spec.). A suggestion from Mr. W. A. Newman of the Scripps Institution to the effect that some 0. hoeki from Cape Verde Is. (Stubbings, 19640) were probably not this species on the grounds of differences in mouth appendages, prompted a re-examination of Stebbing's types of D. hoeki and D. antiguae. Stebbing (1895) described the two species from material from Antigua, W.I., and listed a number of points in which they differed, both in valve form and internal morphology. His figures are, unfor- tunately, inadequate. Annandale (1910 : 217) studying further West Indian material admitted the occurrence of these differences, but found they were not correlated one with the other and united Stebbing's two species under D. hoeki which has page priority. Nilsson-Cantell (1927) was of the opinion that too much emphasis had been laid on differences in valve form in Octolasmis and followed Annandale, at the same time including D. aurivillii (Gruvel, 19026) as a further synonym. He did not consider the differences in number of segments in the cirri of the two forms to be significant. Neither were the even smaller differences in caudal appendages con- sidered of sufficient importance to justify the maintenance of the three species. Nilsson-Cantell (1927) does not mention the difference in setation of the segments noted by Stebbing. This was unfortunate as there seems here to be a constant difference between 0. hoeki and 0. antiguae. In order to clarify the position, specimens variously named Dichelaspis, or Octolas- mis, hoeki and antiguae from the West Indies, Cape Verde Islands and West Africa have been examined. They are referable to one species or the other and will be discussed below under these two species headings. MATERIALS EXAMINED : Tema, Ghana, W. Africa, coll. D. T. Gauld ; Cape Verde Islands (" Calypso " Exped. 1959 : Stubbings 19640) ; Cape Verde Islands (B.M. 79-10 : publ. Nilsson-Cantell 1927) ; Dry Tortugas, W. Indies (B.M. 1952.6.10.2) ; Antigua, W. Indies (B.M. 1928.12.1.2988-2993, syntypes). The shape of the opercular plates has been considered in some detail by Nilsson- Cantell (1927). He concluded that the differences between scutum and tergum in 0. hoeki and in 0. antiguae, described by Stebbing, were merely variations associated with age. The shape of these valves does, undoubtedly, vary with age as does also the proportion of the capitulum covered by the valves. Text-fig. 3 shows the scutum and tergum of a series of specimens of increasing capitular length from 2-03 to 3-44 mm. from Tema, Ghana. The gap between base of tergum and scutum becomes proportionately greater as the capitulum increases in size. The Cape Verde material (B.M. 79-10) shows a similar proportional reduction in the area covered by the valves. Particular regard should be paid to the shape of the tergum (Text-figs. 20, 3). THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 243 Owing to the deep emargination of the scutal margin of the tergum to accommodate the occludent lobe of the scutum the basi-occludent angle of the tergum has a " hooked " or " beaked " appearance. Also, the portion of the scutal margin opposite the basal segment of the scutum is marked off distinctly from the remainder so that a distinct corner is produced on the valve at this point. This angle is marked equally clearly in young and old specimens. The sharpness of the angle varies considerably. The appendages have been compared in specimens from Stebbing's syntypes (Text-fig. 2b-e), from Tema (Text-fig. 40-0) from the Cape Verde Islands and from the Tortugas. These may be compared with the published illustrations of the Cape FIG. 2. Octolasmis hoeki (Stebbing) : syntype (B.M. 1928.12.1) : a, whole animal (X2o) ; b, mandible (x 183) ; c, maxilla I ( xsoo) ; d, segment 4 of posterior ramus of cirrus VI ( X2O5) ; e, base of cirrus VI and caudal appendages ( x 100). ZOOL. 15, 6. 21 244 H. G. STUBBINGS Verde material (Nilsson-Cantell, 1927) and the author (i964«) and also with Stebbing's (1895) rather inadequate figures. They are all very similar and referable to the same species, 0. hoeki (Stebbing). The following points are noteworthy : The mandible (Text-figs. 2b, 40) has fourteeth, the third and fourth, and sometimes the second, with a subsidiary cusp. The lower angle is produced into two acute teeth, the upper of which has a subsidiary cusp. All the subsidiary cusps are weakly developed. The first maxilla (Text-figs. 2c, 46) has two large spines above the notch with a third FIG. 3. Octolasmis hoeki (Stebbing) : right scutum and tergum of specimens of increasing size from Tema, Ghana to show change in shape of valves and in proportion of capitulum covered : capitular lengths a, 2-03 mm., b, 2-09 mm., c, 2-71 mm., d, 2-89 mm., e, 3-01 mm., /, 3'44 mm- THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 245 smaller spine lateral to the second spine. This third spine has not been reported hitherto. The notch is distinct and fairly deep. About three stout setae project from the notch. The lower part of the maxilla stands a little forward of the upper part. It bears a variable number of stout spines, eight or nine in those figured. Nilsson-Cantell (1927, Text-fig, gc) shows only six. The second maxilla (Text-fig. 40) is rounded with the front edge more or less flattened with spines along the whole border, as described and figured by Nilsson-Cantell (1927, Text-fig. 9^). The number of segments in cirri II-VI varies between 8 and 12, the average falling between 9 and 10 for both rami. The numbers found in specimens from several localities and reported in the literature or newly examined are presented in Table I. TABLE i Source I II III IV V VI Antigua, Stebbing 1894 6 or 7 < — 8-10 >• 8 „ syntypes (B.M. 1928.12.1 .2988- . 6-6 9-9 8-9 10-9 9-10 9-8 2993) new preparation Tortugas (B.M. 1952.6.10.2) . . . . 6-6 10-10 10-10 10-10 n-io 10-11 Cape Verde Is. (B.M. 79-10) . . . >f€>-7 9'9 9'9 9'9 9'9 9'io Nilsson-Cantell (1927) .... . \6-8 9-9 9-10 9-9 9-10 --10 Cape Verde Is. new preparation . . . 6-7 n-io 7+-io 8-10 9-10 12 -n ,, ,, Calypso Exped. 1959 . . . 6-7 11-10 ii-n 12-12 12-12 lo-n (Stubbings 19640) Tema, Ghana, D. T. Gauld 2.9.59 . . . 6-6 9-9 7+ -9 8-10 8-9 9-10 TABLE i. Number of segments in the rami of the six cirri in specimens of O. hoeki from various sources. The individual segments are less than twice as long as broad (Text-figs. 2d, 4^), sometimes longer by as little as a fifth only. The ratio of length to breadth of segments from cirri of a number of specimens of 0. hoeki are given in Table 2, together with similar figures for 0. antiguae : TABLE 2 No. of pairs of setae Species and source per segment L/Br. O. hoeki B.M. 1928.12.1.2988-2993 syntypes (Text-fig, zd) . 4 1-50 B.M. 1952.6.10.2 Tortugas. ..... 4 1-83 B.M. 79-10 Cape Verde 4 1-58 Calypso Cape Verde ....... 3 1-21 Tema, Ghana (Text-fig. 4^) 3 1-41 m = 1-51 O. antiguae D. antiguae syntype (Text-fig. 6/) .... 6 3-0 B.M. 1952.2.14.1 Georgetown (Fig. jc) ... 8 3-22 B.M. 1951.7.2.6 „ .... 6 2-15 m — 2-79 TABLE 2. Number of pairs of setae on segments of the posterior cirri and length /breadth ratio in these segments in O. hoeki and O. antiguae. 246 H. G. STUBBINGS The number of pairs of setae on each segment of the posterior cirri is three or four. The caudal appendage is variable in length and in the length of its setae. It may not reach to the distal end of segment I of the protopodite of cirrus VI, as in a Tortu- gas specimen or may extend some way along the second segment as in the syntype (Text-fig. 2e), the Tema specimen (Text-fig. 40) and a Cape Verde specimen (B.M. 79-10). The setae may reach to the distal end of segment i of the rami of cirrus VI (syntype) or scarcely to the base of that segment (Tortugas specimen). Stebbing was probably right in adducing this as a point of distinction from his D. antiguae as in the latter the setae do seem to be consistently longer (see Text-figs. 5g, jd) but the character is not constant in 0. hoeki. The penis is of moderate length and tapers to a rounded tip. It is sparsely covered with soft hairlike setae arranged in rather diffuse rows. The tip bears a FIG. 4. Octolasmis hoeki (Stebbing) : appendages of a specimen from Tema, Ghana ; a, mandible ; b, maxilla I (both x 300) ; c, maxilla II ; d, segment 6 of anterior ramus of cirrus VI ; e, base of cirrus VI and caudal appendage (all x 183). THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 247 terminal tuft of rather stouter and longer soft setae. There is no terminal languet as in, for example, 0. lowei. In several specimens examined, from different localities, the penis was of uniform width tapering toward the distal extremity. There was no widening of the middle portion as noted by Stebbing. This presumed character in two of Stebbing's specimens may be regarded as fortuitous and due to the state of contraction when preserved. Octolasmis antiguae (Stebbing) Dichelaspis antiguae Stebbing 1895, 19-20, pi. 2, figs. E-G. MATERIALS EXAMINED : Dichelaspis antiguae Stebbing syntype, from Antigua, W.I. (B.M. 1928.12.1.2985-2987); D. antiguae, from Georgetown, Br. Guiana, (B.M. 1951.7.2.6) ; D. antiguae from Georgetown, Br. Guiana, (B.M. 1952.2.14.1). Not known from the West African Cirriped fauna. Stebbing (1895) separated D. antiguae from D. hoeki on the basis of the larger proportion of the capitulum covered by the valves, the contraction instead of widening of the tergum below, the slight emargination of the base of the carina and several characters of the appendages. The cirri were said to be more elongate than in D. hoeki and to have from 12 to 14 segments in all except the first pair ; the segments being more slender and elongate and with more numerous spines. The caudal appendages also were said to be larger and more slender with longer terminal setae. Outline drawings of scutum and tergum of British Guiana specimens are shown in Text-fig. 5 (B.M. 1952.2 . 14. i) from which the shape of these valves at different ages is apparent. The downward prolongation of the tergum noted by Stebbing is well seen in these figures of specimens in which the capitulum was well covered by scutum and tergum. The scutal border of the tergum is much straighter than in 0. hoeki and in consequence the hook-like occludent lobe to the valve seen in that species is absent or at the most poorly developed. Furthermore the scutal margin becomes straighter with age. If specimens of similar size are compared, e.g., Text-figs. 30, 5c or Text-figs, y, 5/, the difference in tergal form between the two species will be apparent. 0. antiguae appears to be rather smaller than 0. hoeki. The greatest capitular length found was 3-44 mm. for 0. hoeki but only 2-79 mm. for 0. antiguae. The labrum (Text-fig. 6a, syntype) has a series of well-developed teeth. The palp is oval and tapers slightly (Text-fig. 6b syntype). The mandible (Text-figs. 6c, ja] resembles that of 0. hoeki closely but the fourth tooth and the lower angle have more strongly developed subsidiary cusps. The maxilla (Text-figs. 6d, 76) also resembles that of 0. hoeki. The lower part of the biting edge is almost in line with that above the notch (Text-figs. 6d syntype and B.M. 1951.7.2.6) whereas in 0. hoeki (Text-fig. 2c) the lower portion projects somewhat. This character again is not entirely reliable diagnostically as the lower angle may project in 0. antiguae as shown in Text-fig. 76 (B.M. 1952 . 2 . 14. i). The second maxilla (Text-fig. 6e, syntype) is square and shows no distinguishing characters. H. G. STUBBINGS FIG. 5. Octolasmis antiguae (Stebbing) from Georgetown, British Guiana (B.M. 1952. 2.14.1) : right scutum and tergum of specimens of increasing size showing greater coverage of capitulum and more elongated shape of tergum : for comparison with similar valves of O. hoeki shown in Fig. 3 : capitular lengths a, 1-24 mm., b, 1-92 mm., c, 1-98 mm., d, 2-16 mm., e, 2-54 mm., /, 2-79 mm. Re-examination of the cirri supports Stebbing's statement that they have more segments than do those of 0. hoeki. The numbers found or recorded previously are shown in Table 3. Stebbing 1895 .... D. antiguae syntype B.M. 1952.2.14.1 Georgetown B.M. 1951.7.2.6 „ TABLE 3. Number of segments in the cirri of four O. antiguae including Stebbing's original statement (1895) and those in a syntype. TABLE 3 I *7 or ft II III IV -T 1 T A- V VI • 1 \J1. CJ — I4-IO . 6-7 12-11 I2-I2 12-12 12-12 II -IO . 6-7 12-11 10-12 I2-I4 11-13 IO+-I3 THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 249 The mean number of segments in cirri II- VI is nearly 12. As Table 2 shows the length /breadth ratio of individual segments is much greater than in 0. hoeki, being more than 2 and often more than 3. The number of pairs of setae is consistently higher at 6-8 pairs per segment. The single- jointed caudal appendage may extend to just beyond the distal end of the first segment of the pedicel of cirrus VI (Text-fig. 6g, yd) or may be a little shorter. In this it does not differ clearly from the condition in 0. hoeki. The terminal setae are appreciably longer as normally they extend to the distal end of segment i of the rami and often appreciably on to the second segment. The penis is of moderate length, tapers towards the tip and has sparse hairs along its length and a terminal tuft of rather longer hair-like setae. It does not differ from that of 0. hoeki. It would appear, therefore, that the differences between D. hoeki and D. antiguae originally described by Stebbing (1895) and now re-examined are distinguishable. They are not so strongly marked that variability within the two species cannot bring FIG. 6. Octolasmis antiguae (Stebbing) : appendages of a syntype (B.M.) : a, labrum ; b, palp (both x 206) ; c, mandible ( x 294) ; d, maxilla I ( x 397) ; e, maxilla II; /, segment 7 of anterior ramus of cirrus VI (both X2o6) ; g, lower part of cirrus VI and caudal appendage (X45). 250 H. G. STUBBINGS about an overlap of individual characters. In consequence characters such as the shape of the tergum, the degree of dentation of the lower angle of the mandible, the straight or stepped edge of the maxilla and the length of the caudal appendage and its setae are individually unreliable as diagnostic characters. Taken collectively or when a series of specimens is available from the same source they afford a reasonably accurate guide to the species. The most reliable features are undoubtedly the pro- portions and setation of the cirral segments. In 0. hoeki the length of the segments is less than twice the breadth and there are three or four pairs of setae. In 0. antiguae the segments are more than twice, often over three times, as long as broad and there are from six to eight pairs of setae. These characters do not intergrade. Accordingly Dichelaspis antiguae Stebbing, 1895, is reinstated as a distinct species under the modern name Octolasmis antiguae (Stebbing, 1895). The present investigation has shown that 0. hoeki occurs on both sides of the tropi- cal Atlantic whereas 0. antiguae is known only from the much more restricted area of the Leeward Islands and British Guiana. FIG. 7. Octolasmis antiguae (Stebbing) : Appendages of a specimen from Georgetown, British Guiana (B.M. 1952 . 2 . 14 . i) : a, mandible ; b, maxilla I ; c, segment 7 of posterior ramus of cirrus VI (all x 300) ; d, lower part of cirrus VI and caudal appendages ( X 90) • THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 251 VERRUCOMORPHA VERRUCA Verruca striata Gravel Taken by the " Talisman " south of the Cape Verde Islands, Lat. 16° 51' N., Long. 27° 30' W. in 598-633 m. (Gravel, 19020) this is the only species of Verruca so far recorded from tropical West African waters, though many species are known from the warm temperate waters of Madeira and the Azores. As yet, no species has been found in West African coastal waters. BALANOMORPHA CHTHAMALUS Chthamalus stellatus (Poli) Four varietal forms of C. stellatus are recorded in the literature from West Africa, vars. stellatus, depressus, fragilis and bisinuatus, in addition to records where no varieties are specified. The records of fragilis and bisinuatus are considered below, the former as C. fragilis. Records of " depressus " refer to C. aestuarii Stubbings. Of the remaining records of " C. stellatus " and " C. stellatus var. stellatus " it is highly improbable that all refer to the var. stellatus. C. stellatus var. stellatus (Poli) LOCALITY: Ghana, Winneba : shore Section, £ m.2, no. i A, 4.3.50 (i spec.). The southward extent of var. stellatus is difficult to determine from records owing to the confusion of forms in earlier reports. Records in the literature up to 1943 were cited in Stubbings (19610). The more northerly records, namely Morocco (Broch, 1927), Rio de Oro (Nilsson-Cantell, 19396), Mauretania (Broch, 19246) and Madeira and Cape Verde Is. (Darwin, 1854) are acceptable. The Cape Verde record is confirmed by " Calypso " material (Stubbings, 19640). Sourie (1954) records only C. stellatus in his study of the ecology of rocky shores from Mauretania to Guinea, thus including Senegal. Yet there were no C. stellatus in the cirripede collections from I.F.A.N., Dakar, studied by the writer, only two samples of C. dentatus. Assuming Sourie's identifications to be correct, some at least of which were made by Professor Hj. Broch of Oslo, this implies an abrupt change of predominant species in the region of Cape Verde. The northern species C. stellatus is predominant in Mauretania and south of the Cape the tropical and southern hemisphere species C. dentatus. Records from further south are suspect because of the possibility of confusion with C. aestuarii and C. dentatus. Thus Gruvel (1912) records C. stellatus from Libreville (Gabon) and Kitombe, Banana, at the mouth of the Congo. Weltner (1897) gives Victoria, Cameroons and Kolosvary (19430) Fernando Po, as localities. There are no C. stellatus in the extensive collections from the Congo estuary, including Kitombe, in the collections of the Musee Royal de TAfrique Centrale, Tervuren 252 H. G. STUBBINGS (Stubbings, 19646). Whilst " Calypso " did not visit Fernando Po, collections were made at the other islands in the Gulf of Guinea and no C. stellatus were collected (Stubbings, 19616). In both areas C. dentatus was collected, and C. aestuarii is common in the Congo estuary. Weltner's record from Victoria is presumably correct as specimens were taken there by " Atlantide " (Stubbings, 1961 a). It would seem, therefore, that in West Africa C. stellatus stellatus is only common on the north-west coast (Morocco, Mauretania) and in Madeira and the Cape Verde Islands. It may occur sporadically in small numbers further south, certainly as far as the Cameroons, but in general it is replaced south of Cape Verde by C. dentatus in more exposed conditions and by C. aestuarii in the more sheltered conditions of estuaries. C. stellatus var. bisinuatus Pilsbry C. stellatus var. bisinuatus: Stubbings, 19610, 18-19, text-fig. 3. Specimens from Lagos, Nigeria in the University Zoological Museum, Copenhagen (Stubbings, 19610;) belong to this variety. There are no other WTest African records of this otherwise Brazilian variety. Chthamalus dentatus Krauss LOCALITIES: Gambia: No. 3 Gambia Lightship, 22.9.48, pres. M. W. H. Bishop, B.M. 1950.7.19.1 (several spec.). Sierra Leone: Kissy, Sierra Leone River, on rocks above littoral fringe, coll. A. R. Longhurst, B.M. 1956.1.7.4 (32 spec.). Ghana: (i) Ada, on launch " Akuse " (21 spec.); (2-9) Accra, (2) Christiansborg shore, 15 . 1 .49 (several young uneroded spec.) ; (3) on Thais haemastoma L., 14.2 .49 (several hundred spec.); (4) from splash zone, 14.2.49 (no +22 juv. +5 dead spec.); (5) on T. haemastoma, T. nodosa L. and Patella safiana Lam., 17.3.49 (c. 100 + 50 juv. + 29 dead spec.); (6) on a limpet, 5.11.49 (123 + 20 juv. + 45 dead spec.) ; (7) on a large B. t. tintinnabulum, 19 . n .49 (few spec.) ; (8) on a rubber tyre fender, 14 . 3 . 51 (several hundred spec.) ; (9) HWM on sandstone rock, 19.3. 58, coll. H.G.S. (91 -f 45 juv. spec.) ; (10-12) Winneba: (10) on rock and shells including T. haemastoma, P. safiana and Mytilusperna L., 15 . n .49 (453 -f 264 juv. -f 32 dead spec.); (n) on similar substrate, 22.11.49 (several hundred spec.); (12) shore section made on 3/4.3 .50 (over 1500 spec.) ; (13-16) Apam; (13) " high on shore ", 16.2.49 (many spec.); (14) on Gadinia afra Gmelin (44 + 25 juv. spec.); (15) embedded in a sponge (i spec.); (16) on Gastropods including T. haemastoma, T. nodosa and Nerita senegalensis Gmelin (several hundred spec., about half juv.); (17) Sekondi, High Land Plateau, on sandstone rock, 21.2.49 (numerous spec.); (18) Prince's Town shore, 15.4.49 (r spec.); (19) Axim, Hospital reef, H.W.M., 13.4.49 (several hundred spec.); (20) Axim, on reef, B. t. tintinnabulum and a limpet Fissurella nubecula L., 17.1.51 (184 spec.); (21) Ankobra ferry, on log, 19.2.49 (many spec.); (22) Half Assini, on wreck, — 2.49 (217 + 2 juv. + 7 dead spec.); (23) Prampram, on rock fragment, — .11.37, Pres- Miss V. J. Foote, B.M. 1952.5.7.2 (numerous spec.). THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 253 Nigeria: (i) Bonny River, fairway buoy in river mouth, in splash zone and down to i ft. below float line, with B. pallidus (45 spec.) ; (2) Bonny, on New Pier on concrete at HW and above with C. aestuarii, 28.11.57 (4 spec.) (H.G.S. 331). I have seen further material from Labadi Beach, Accra, and Tema, Ghana, the west mole, Lagos and from Victoria, Cameroons, all collected by Dr. Eyvor Sandison. This extensive material includes young and old, uneroded and eroded specimens, single individuals and densely packed colonies, the last sometimes showing some shell elongation. Where erosion is heavy shells become low and featureless and when densely packed sutures and even boundaries between specimens become indistinguish- able. Young uneroded specimens are fawn or pale-brown with a thin epidermis. The compartments are practically smooth, with only feeble longitudinal furrows on some of them. Growth lines are visible intermittently. The sutures are rather irregularly sinuous externally and not dentate. With erosion the worn areas become marbled fawn or brown and white with the brown epidermis restricted to the newer basal parts of the compartments. In more heavily eroded specimens the shells may be wholly white. The Nigerian specimens from a buoy at Bonny are large isolated broadly conical individuals, the largest 12 X 9-5 mm. diameter x 5 mm. high. In most the lower part of the paries is " narrowly and regularly folded " as described by Darwin (1854 : 463) for shipborne specimens. The development of the characteristic dentate sutures is variable. When well- FIG. 8. Chthamalus dentatus Krauss : specimen from rubber tyre fender at Accra, a, internal view of rostro-lateral compartment with well-developed dentate sutures ( X 14) ; b, c, external and internal views of the scuta and d, e, of the terga of the same indi- vidual (all X23). 254 H. G. STUBBINGS formed and uneroded the zigzag sutures are clearly visible externally. The sutural surfaces or " teeth " are short ridges with sharp, angular crests, the sutures then appearing sharply dentate, or with smoothly rounded crests in which case the sutures become sinuous. The broader and lower these ridges the more the sutural line departs from a zigzag and approaches a straight line. Both angular and rounded sutural teeth are seen in the left rostrolateral compartment in Text-fig. 8a. In eroded shells it is often impossible to discern the dentate margin to the compartments. They can then be seen only from the inside of the shell or after dissociation of the compartments. The dentations may be obliterated by distortion during the growth of close-packed low-growing specimens. In elongated specimens the teeth may be very small or obscure on the lower part of the sutures which become more or less sinuous in outline. Internally the dentations are often in a narrow groove which renders them less visible. In close-packed specimens the groove may be almost closed in the lower part and the sutures show only i or 2 dentations in the older part. These were presumably produced before the barnacles became contiguous. The clearest dentations are shown on specimens grown singly in sheltered places free from erosion, such as on piers or buoys. Two forms of the shell and an internal view of the tergum were shown by Darwin (1854, pl- I^>> %s- 3a~c)- No figure of the scutum has been published. Internal and external views of both opercular valves are given here. The scutum (Text-fig. 86, c) agrees well with Darwin's description in the very prominent articular ridge, which is most protuberant about the middle of the articular border and in the deep adductor pit and the row of small pits for the depresser muscles referred to by Darwin as " distinct, though minute, pits ..." (1854 : 464). In some young specimens the broadest part of the articular ridge may be nearer the apex when the valve comes to resemble that figured by Pilsbry (1916, fig. 83) as C. fragilis Darwin. Specimens with this type of scutum but with well developed dentate sutures are in the collection so these are presumably C. dentatus though somewhat divergent from the norm. Externally uneroded scuta bear fairly prominent regular growth lines. The tergum (Text-fig. 8d, e) agrees with Darwin's figure in the strong articular ridge and strongly curved carinal margin. The depressor muscle crests are on a projecting portion — " Auf einer vorspringenden Partie " — as remarked by Nilsson- Cantell (1921 1284). Mouth parts have been figured previously by Nilsson-Cantell (1921). There is a long series of small teeth and a row of hairs on the labrum (Text-fig. 90). In some Nigerian specimens the labral teeth are less developed and less numerous. The palp (Text-fig. 96) is broadly oval with the inner margin straight or slightly concave. The mandible (Text-fig, gc) is close to that shown by Nilsson-Cantell (1921, fig. 52c, d). There have been divergent reports on the condition of the lower angle. Darwin (464) described it as " coarsely pectinated " but Nilsson-Cantell found most specimens to have only fine bristles here, a view concurred in by Barnard (1924). However, Nilsson-Cantell (1921) found an individual in which spines occurred among the hairs on the lower angle (his fig. 52^) . The specimen here figured has fine spines on the lower angle. THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 255 Maxilla I (Text-fig, gd) has a broad notch with small spines. Below it, the median spines are but little larger than the lower ones into which they grade smoothly. Maxilla II (Text-fig, ge) is broad with a deeply concave anterior margin, the median concavity being devoid of bristles as noted by Nilsson-Cantell. The number of segments in the cirri of two specimens, presumably from Madagas- car, were given by Nilsson-Cantell (1921). These compare favourably with counts FIG. g. Chthamalus dentatus Krauss : a, labrum ; b, palp ; c, mandible ; d, maxilla I ; e, maxilla II (all x 170) of a specimen from rubber tyre fender, Accra : /, compound spine from tip of cirrus II ( x 350) of a specimen from a buoy at Bonny, Nigeria ; g, tip of penis (X8o). TABLE 4 Scutum I II III IV V VI mm. L 7-6 6-5 I5-I5 16 •16 18 •18 18 •19 i •85 R 7-6 6-5 I4-I5 15 •17 18 •17 19 •18 L 7-8 7'5 14-14 17 •16 16 •16 18 •18 2 •ii R 8'7 7-6 13-12 15 • 17 18 •18 19 •18 L 8-7 7'7 20-19 23 •23 24 •24 24 •24 3 •5i R 9-7 7-6 2O-2I 23 •24 24 •24 25 •24 L 7-6 6-5 I8-I9 22 •23 24 •25 27 •24 3 '44 R 7-6 6-5 I7-I8 22 •23 25 •24 25 •25 256 H. G. STUBBINGS made on specimens from Christiansborg, Accra and from Nigeria collected by the author. The number of segments in the right and left cirri of two specimens from each locality are given in Table 4. Specimen Accra .... Accra .... Nigeria .... Nigeria .... TABLE 4. Number of segments in left and right cirri of four C. dentatus, with scutal lengths as an index of barnacle size. The number of segments in the longer cirri in the Nigerian specimens but not in those from Accra is thus rather higher than previously recorded. Because of the distortion of some of the shells due to crowding the relative size of the above specimens is indicated by reference to the length of the base of the scutum. The Nigerian specimens were considerably larger than those from Accra, probably attributable to a more favourable environment on the navigation buoys. This size variation could account for the observed difference in segment counts. Pectinate spines (Text-fig, o,/) are present on the terminal segments of cirrus II. The penis is closely annulated, the annuli broadening towards the distal end until about half as long as broad (Text-fig, gg). The distal segments are sparsely clothed with long hair-like setae. A ring of similar setae surrounds the tip. C. dentatus is found on the whole west coast of Africa, south of Cape Verde. There seems little doubt of its northern limit. There are few specimens in the IFAN collections at Dakar, suggesting that it is not common there. Furthermore, Sourie (1954) who investigated the coastline of Mauretania and Senegal rather thoroughly does not record it at all. Hoek (1883) records C. dentatus from the Cape Verde Is., but from his account the identification is uncertain. The species was not found in the archipelago by the " Calypso " expedition in 1959. In the Gulf of Guinea the species is known from the islands of Principe and Sao Tome (Stubbings, 19616). South of the River Congo there are few records, largely due to the lack of collecting on this coastline. At the Cape it is common from False Bay to Durban (Sandison, 1954). According to Miss Sandison it is uncommon in the cold water current on the west side of the Cape Peninsula and this, presumably, will apply to the Atlantic coast of South Africa and South- West Africa where in- fluenced by the cold Benguela current. As at Capetown, however, it is not entirely absent from this section of the coast, as witness the record for Walvis Bay (Stubbings, 1963)- Four isolated records, two in Madagascar (Nilsson-Cantell, 1921), one each in Mauritius (Weltner, 1897) and Aden (cited by Nilsson-Cantell, 1921, without author- THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 257 ity) extend the distribution of C. dentatus into the Indian Ocean and a fifth (Weltner, 1897) to Singapore. Chthamalus aestuarii Stubbings Chthamalus aestuarii Stubbings, 1963, 7, text-figs. 2, 3. Chthamalus stellatus depresses: Nilsson-Cantell, 19386, 177, text-fig. 2. Chthamalus cirratus: Kolosvary, 19416, 70. Chthamalus cirratus: Kolosvary, 1 943*3, 75. Chthamalus stellatus f. depressus: Gauld, 1957, 10. Chthamalus rhizophorae: Longhurst, 1958, 32, 59, 85. Chthamalus withersi: Longhurst, 1958, 59, 85. LOCALITIES : Sierra Leone : (i) Bunce I. Sierra Leone River, on mangrove, 1955, coll. A. R. Longhurst, B.M. 1956 .1.7.3 (as C. rhizophorae Oliveira) ; (2) Sierra Leone River on mangrove, July 1955, coll. A. R. Longhurst B.M. 1957.6.3.3 (as C. withersi Pilsbry); (3) Reef at Wellington, Sierra Leone River on mangrove, 19.7.55, coll. A. R. Longhurst, B.M. 1956.7.1.5 (as C. withersi). Ghana: (i) R. Densu on mangrove, 3.4.49.; (2) Prince's Town, on mollusc shell fragment, 15.4.49 (*• spec.); (3) Ada, R. Volta, at H.W.M. on pier, 15.3.49 (numerous spec.). Nigeria: (i) Port Harcourt on concrete " dolphin ", 19.12.57 (37 spec.) (H.G.S. 350); (2) Bonny R. estuary, on concrete pier at Bonny, 28.11.57 (6 spec.) (H.G.S. 323)5 (3) Opobo, Imo River, E. Nigeria, on pier and on B. pallidus, 11.1.58 (12 spec.) (H.G.S. 373). In addition I have seen specimens collected by Miss Sandison from Badagri Creek and Kuramo Creek, near Lagos, from Porto Novo lagoon, Dahomey and the R. Baka, near Elmina, Ghana, all on bark, probably of mangroves. This species was described fully from material from the Congo estuary by Stub- bings (1963). The present material establishes its presence over the whole tropical West African coastline where conditions are favourable. In examining further material from a greater geographical area some variations in form have been observed and are illustrated here. In my original illustration a slightly eroded specimen in which growth lines were not apparent was figured. This specimen had radiating dark brown or black lines on the parieties. The young and uneroded specimen now figured (Text-fig. ioa) from the Sierra Leone river lacks these black lines and the incremental lines are quite distinct. The scutum (Text-fig. lob, c) has a much more oblique basi-tergal angle and the basal margin is less strongly bowed. In other respects the scutum resembles my earlier figures, in particular in the protuberant apical region of the articular ridge. Internally the tergum shows some variation. The articular furrow is always well developed and rather broad. Text-figs. iod, e are external and internal views of the terga of the specimen from which the scuta came. The articular furrow here was of average size. The depressor muscle crests on the other hand were reduced to one, instead of the normal three or four as in Text-fig. io/. This latter specimen had a very broad open articular furrow with a rather uneven surface. A specimen from Port Harcourt (Text-fig. log), showed a similar rough broad articular furrow. This 258 H. G. STUBBINGS FIG. 10. Chthamalus aestuarii Stubbings : a, uneroded specimen from Sierra Leone River showing distinct growth lines and no pigment bands ( x 17) ; b, c, external and internal views of the scuta, and d, e, of the terga of another Sierra Leone specimen ; /, internal view of tergum of a third specimen (b-f, all x&5) (B.M. 1957.6.3.3) ; g, tergum of a specimen from Port Harcourt, Nigeria, with broad articular furrow and roughened internal surface ( X28). rather massive valve was noteworthy for the very rough interior and for the apparent differentiation of a tiny spur from the basi-scutal angle. The mouth appendages with the exception of Maxilla II were illustrated by Stubbings (1963). Minor variations are shown by the appendages illustrated in Text-fig, ii. In a Port Harcourt specimen (Text-fig, na) the row of teeth on the labrum is rather shorter. In a specimen from Sierra Leone (B.M. 1957.6.3.3) (Text-fig, iib-d] the mandible has 3 simple teeth without subsidiary cusps. In the maxilla the central group of spines is demarcated by a very weak notch below as well THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 259 as by one above. In this it approaches C. rhizophorae Oliveira but the frontal margin of the maxilla is still straight and not stepped as in C. rhizophorae. Maxilla II has a concave anterior margin and a very strongly convex posterior margin. The setae are all simple. The cirri are as previously described. The number of segments in the right and left cirri of a Sierra Leone specimen are given in Table 5. TABLE 5 I II III IV V VI 8-6 8-6 8-8 9.9 17-20 24-24 25.?* I9-22 24-?* 20-23 j. 2I* + -2 right ...... left ...... * cirri IV and V of the right side lacked one ramus : cirri V and VI of the left side lacked the last few segments. TABLE 5. Number of segments in right and left cirri of a specimen of C. aestuarii. These figures are in good agreement with the segmentation of the cirri of a Congo specimen (Stubbings, 1963 : 9). The above-sited synonymy was reviewed at some length by Stubbings (1963) and that discussion need not be repeated here. With the exception of C. wither si and C. rhizophorae the species confounded with C. aestuarii have in general been described and figured adequately in the past. The appearance of full-grown C. wither si is now revealed through the work of Pope (1965) but no further figures of C. rhizophorae have appeared since the original descriptions by Oliveira (1940, 1941). As Miss Pope has cast doubt on the status of C. rhizophorae as a separate species — and by implication of C. aestuarii (1965 : 40) a further series of figures of scuta and terga of C. rhizophorae from the same general area as the type are given here (Text-fig. 12). Comparison of these valves with those of C. aestuarii (Text-fig. 10 and Stubbings, 1963, fig. 2) shows at once the differences and resemblances in the opercular valves of the two species. The shape of the articular ridge of the scutum alone (Text-figs. io&, c and iza, c) separates the two species immediately. The presence of pits in the inner surface of the scutum and to a lesser extent of the tergum also seems charac- teristic of C. rhizophorae. These pits are visible in Oliveira's (1940) photographs, but were not indicated in his line drawings of the valves (1941). Very heavy and thick valves may occur in C. rhizophorae (Text-fig. 120, d) and in terga of this type the articular surface is even broader than in C. aestuarii. The three ' species ' have been compared using material from Queensland (C. wither si), Brazil (C. rhizophorae} and West Africa. The chief points distinguishing C. aestuarii from C. wither si are described below. The psi (tft] shaped articulation of the opercular valves is shown by all except young uneroded aestuarii but not by withersi. The articular ridge and furrow of the scutum form only " a shallow wavy fold and trough " (Pope, 1965 : 42) in withersi. There are scarcely any interlocking surfaces. It is, in fact, one of the most poorly developed examples of this structure in Chthamalus. In aestuarii the ridge is much stronger and interlocks with the tergum. The area of the ridge towards the valve apex projects beyond the tergal margin of the scutum. In the tergum the chief ZOOL. 15, 6. 22 260 H. G. STUBBINGS FIG. ii. Chthamalus aestuarii Stubbings : a, labrum of a specimen from Port Harcourt ; b, mandible, c, maxilla I, d, maxilla II of the same Sierra Leone specimen as the opercular valves in Fig. lob-e (all xiyo). difference lies in the articular margin. As in wither si it is inturned at right angles to the plane of the main valve surface. But whereas in wither si there is only a very narrow articular surface in aestuarii it is much broader. To accommodate this greater articular surface the scutal margin is turned further back on itself so that much of the articular surface lies in a plane parallel to that of the main part of the valve. In young rhizophorae the articular ridge of the scutum is only modestly developed but its middle portion projects forward over the articular groove. In old and massive valves which have only suffered slight erosion, the interlocking surfaces of scutum and tergum are very broad and flat, being again roughly parallel to the external surface of the valves. On these characters it is quite possible to distinguish the three species. Differences in the mouth appendages are rather variable. The redescription of the labrum by Miss Pope removes the earlier distinction of a short line of teeth on its edge in wither si. The wither si mandible evidently varies in the number of major THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 261 teeth from three to occasionally four. The curiously " hard " outline of the teeth shown by Pilsbry and present in my material — seemingly due to the straight margins of the teeth in profile — is not seen in aestuarii or rhizophorae. The lower angle terminates in a few short spines in my withersi but in aestuarii and rhizophorae the number of spines is greater and they form a " comb " below the third tooth. There is some variability in these spines. Those shown in Text-fig. lib are appreciably shorter than those in my original description (S tubbings, 1963, fig. 30). In withersi, and rhizophorae the edge of maxilla I has two notches and is " stepped ", the lower sections protruding well in front of that or those above. In aestuarii the edge is almost straight with a small notch below the upper group of spines. In the specimen originally figured there was no lower notch. In that now shown there is a slight indication of this lower notch. It is more apparent than real, its presence being emphasized by the disposition of the spines and the thickening of the cuticle here. In view of the above differences it is considered that C. aestuarii andC. withersi are distinct species albeit occupying very similar habitats. Though less material has been available, it would seem that C. rhizophorae also is a distinct species. FIG. 12. Chthamalus rhizophorae Oliveira : internal views of a, b, right scutum and tergum of one specimen, c, d, right scutum and left tergum of a second, and e, left tergum of a third specimen (all 262 H. G. STUBBINGS Chthamalus fragilis Darwin LOCALITY: Bota, Cameroon: on Ostrea sp. from a lighter, 21.2.58 (22 spec.) (H.G.S.426). Chthamalus fragilis was recorded by Broch (19270) from Tangier and is grouped as an Atlantic-Mediterranean species by Kolosvary (19416), presumably on this record. There are no other records for north or west African coasts. The species is common apparently on the warmer Atlantic coastline of North America and in the West Indies (Darwin, 1854 ; Pilsbry, 1916 ; Nilsson-Cantell, 1933, 1939^ ; Kolosvary, 19430 ; Weiss, 1948). Broch (19270) did not figure the Tangier specimens. Figures have been given of American (Pilsbry, 1916, pi. 70 and text-fig. 83) and West Indian specimens (Nilsson-Cantell, 1928, text-fig. 14 ; 1933, text-fig, i). The walls of the present specimens are smooth and white with a thin pale yellow persistent epidermis. Growth lines are present and easily traced but are very low. Radii are present but small, and the interparietal areas are very narrow. The tergum (Text-fig. 130) agrees closely with that figured by Pilsbry (1916, text-fig. 83A) . The spur is distinct from the basi-scutal angle but is rounded, not pointed as in some of Pilsbry's examples and in the two shown by Nilsson-Cantell (1933, text-fig, i). The scuta are shown by Pilsbry only. That figured here (Text-fig. I3&), is in good agreement with Pilsbry's text-fig. 830 and his pi. 70, fig. 3. The depressor muscle pit is not appreciably deeper than the shallow depression behind the articular ridge. The adductor pit lies in a very pronounced furrow parallel to the occludent margin of the valve. The mouth parts are, as stated by Pilsbry, very like those of C. stellatus stellatus and agree with this author's figures for the latter species. FIG. 13. Chthamalus fragilis Darwin : internal views of a, tergum and b, scutum of a specimen from Bota, Cameroon ( x8o). THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 263 Pachylasma giganteum (Philippi) This Mediterranean species has been recorded twice only from other waters. Gruvel (19076, 1910) records it from Simonstown, South Africa in the collections of the " Gauss " and Kolosvary (19430) from Sette Cama, Gabon. The latter is the only record for West Africa. Darwin (1854) remarks that P. giganteum is often associated with B. tulipiformis, but although the latter species has been collected frequently no Pachylasma have been recorded. Other species of Pachylasma have been described from the Indian Ocean, South-east Asia and Australia. Specimens of all species of Pachylasma are rare and the degree of variation within any one species is uncertain. As Nilsson-Cantell (19326) has pointed out, it is probable that some at least of these described species are no more than sub-species of P. giganteum. Until more material is available especially of P. giganteum from the Mediterranean and Atlantic the validity of these species, and therefore, the geographical distribution of each, must remain problematical. BALANUS Balanus tintinnabulum (Linn.) Only one variety or subspecies of B. tintinnabulum is common on the west coast of Africa, namely B. tintinnabulum tintinnabulum (L.). The occurrence of two other varieties has been fairly definitely established but the presence there of several others mentioned in the literature is highly doubtful. Balanus tintinnabulum tintinnabulum (L.) Balanus tintinnabulum tintinnabulum Pilsbry, 1916, 55-57, pi. 10, figs. i-ie. LOCALITIES: Gambia: (i) off Gunjur, on hull of shark fishing boat, 3.3.51, coll. M. H. Routh, B.M. 1952.10.2.3 (14 half grown and n very small spec. ^-S5- 7a~f- LOCALITIES: Gambia: (i) off Gunjur from hull of shark-fishing boat, on B. t. tintinnabulum, 3.3.51, coll. M. H. Routh, B.M. 1952.10.2.3 (few juv. spec.); (2) from seawater intake of "African Queen", on B. t. tintinnabulum, 2.10.51, coll. M. H. Routh, B.M. 1952.10.2.5 (2 juv. spec.). Sierra Leone, (i) 12.1.52, coll. A. R. Longhurst, B.M. 1957.6.3.5 (19 dead spec.); (2) coll. on board S/T " Cape St. Mary " P. H. Creutzberg, pres. A. R. Longhurst, B.M. 1956.1.7.11 (9 dead and i live spec.). Balanus trigonus is of world- wide distribution in tropical and subtropical waters. In the Atlantic it occurs from the Azores and Madeira to the Cape of Good Hope in shallow water. It is present in the Mediterranean. On the African Mediterranean Coast there are records from Oran (Kolosvary, 19430). It was not reported by Broch (1927) from the Atlantic Coast of Morocco but Gruvel (1912) states that it occurs on all the Atlantic Coast of the Sahara (i.e. Mauretania). Sourie (1954), also reports its abundance at Cap Blanc, Mauretania and at Dakar, Senegal. It is very common on the Senegal Coast (Stubbings, 1965) and was present in almost all the " Calypso " collections made in the Cape Verde Archipelago (Stubbings, 1963). From south and east of Senegal there are very few records. Longhurst (1958) obtained it twice only in his survey of the benthos off Sierra Leone. There are no records from Liberia, the Ivory Coast or Ghana and only one from Lagos, Nigeria (Stubbings, 19610). From Liberia this lack of records may be due to lack of investi- gations but the absence of the species from Ghana must be real in view of the exten- sive collecting by Bassindale. It was not found among the islands of the Gulf of Guinea. 268 H. G. STUBBINGS South of the Bight of Biafra records are again more frequent and B. trigonus is recorded down to Tiger Bay on the southern border of Angola (Gruvel, 1912). There is a single record from Great Fish Bay, South-west Africa (Weltner, 1922). In this paper Weltner reports the species from various localities in the Cape and Barnard (1924) states that it is found all round the South African coast from False Bay to Zululand. It would appear, therefore, on present information that B. trigonus is common on the N.W. Atlantic coast of Africa, it being most abundant in the Senegal-Cape Verde area. From about 10° N. to 5° S. it is evidently uncommon. Collecting on the coasts of Angola, South-west Africa and the western coast of the Union has not been inten- sive, largely due to difficulties of access, but such records as we have would indicate that B. trigonus is present throughout this section of the coast. It is evidently common on the southern coast of the Union and well into the Indian Ocean. Balanus spongicola Brown B. spongicola is not contained in the present collections. It is widespread in the eastern Atlantic and Western Indian Ocean. In West Africa there are records from French Guinea (S tubbings, 19610;), Rio de Oro and Ghana (Stubbings, 19616) and Angola (Stubbings, 1963). It is known from the western Mediterranean, Algiers (Darwin, 1854) and Catania (Kolosvary, 19430). Northwards it extends to Portugal (Kolosvary, 19430), Madeira (Darwin, 1854), the Azores (Gruvel, 1920) and the south-west coast of England (Darwin, 1854 '> Norman & Scott, 1906). The identity of B. dollfusi Broch (19270) with B. spongicola has been discussed by the author (Stubbings, 1963) elsewhere. All records of this species in the literature refer to localities whence B. spongicola has been collected or are within its known geographical range, namely the Atlantic coast of Morocco (Broch, 19270) Rio de Oro (Nilsson-Cantell, 19396) and off the Congo estuary (Nilsson-Cantell, 19386). Gruvel (19076, 1910) records it from Simonstown, South Africa and Barnard (1924) from numerous places from False Bay to Zululand. In the Indian Ocean it is found in the Chagos Archipelago and Seychelles (Gruvel, 19070). Kolosvary (19430) cites specimens in Budapesth from the East Indies, but there are no other records from this region and confirmation by new material is desirable. Darwin (1854) placed some specimens from the West Indies in an unnamed variety, but was uncertain as to their true status. Pilsbry (1916) elevated these to specific status as B. calidus. It is probable that the W. Indian and tropical American material recorded as B. spongicola by Weltner (1897), Nilsson-Cantell (1927) and Kolosvary (19430) should be assigned to Pilsbry's species, though in neither of the two last cited works does the author mention B. calidus. Balanus perforatus Brug. This species is not represented in the Ghanaian or Nigerian materials here reported. Recent records of the species in West Africa are from the southern portion of its THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 269 range, south of the Bight of Biafra : FaussePointe Noire (Congo) (Stubbings, 19646) and Lobito and Moita Seca, Angola (Stubbings, 1963). B. perforatus exhibits the same divided distribution in West Africa as does B. trigonus. There are a number of records from Morocco and Mauretania (Broch, 19270; Gruvel, 1912) and the Canary Islands (Nilsson-Cantell, 1932*2). Darwin names the Gambia as a locality and Bruguiere (1789) cites Senegal in his original description, but there are no other localities recorded to the southward until Pointe Noire in Congo (formerly French Equatorial Africa) is reached. The species is unknown from south of Senegal to the southern border of the Gulf of Guinea. There are several records from the Congo and Angola but it has not been found on the south-west coast south of Lobito, Angola. Gruvel (19076, 1910) records the species from rocks at Simonstown, South Africa. The cited records include references to the vars. (or formae) angustus andfistulosus, both being growth forms, the former occurring where erosion is slight, the latter due to overcrowding. The collected and recorded material of B. perforatus from West Africa is unsatisfactory in that much of it consists of empty shells, dead when collected. There is, therefore, an element of doubt as to the true status of the species in some of the recorded localities. Dead shells may be the remains of an indigenous species, be derived from imported specimens that have bred but not been able to maintain themselves, or be merely the remains of imported shipborne specimens. It is probable that many of the present records represent shipborne introductions that have failed to maintain themselves for more than a short period. Until much more living material, preferably in clusters comprising individuals of various ages, is available, there must remain an element of doubt regarding the true status of B. perforatus in West Africa. In any case, it seems safe to presume its absence from about Lat. 10° N. to 5° S. The geographical limits of B. perforatus beyond south-west Europe and the west coast of Africa are somewhat debatable. Its general distribution in the Mediter- ranean is undisputed and Zevina & Tarasov (1954) have recorded it, as a new record for Russian waters, in the Black Sea. In the eastern North Atlantic it extends to south-west England and South Wales (Norris & Crisp, 1953), though it is absent from the Scilly Isles (Fischer-Piette, 1936). It was not found by Southward & Crisp (1954) in their survey of the intertidal fauna of the Irish coastline but Williams (1954) records it from about 20 m. in Strangford Lough. Occasionally, therefore, it may penetrate far into St. George's Channel. The Scottish record from the Jef- frey's collection (Pilsbry, 1916) is of doubtful validity as is even more so that from Kirkeness in Gruvel (1903). Darwin hesitated to accept Western Atlantic records for B. perforatus and in this he was followed by Pilsbry (1916) on the grounds that they had not been con- firmed by later records although Weltner (1897) cites specimens of the var. angustus on " Pur pur a haemastoma " from Brazil in the Berlin Museum. Nevertheless there are no recent records of western B. perforatus and its occurrence on both sides of the Atlantic must still be regarded as equivocal. The Andaman Islands record in Gruvel (1907^ must be in error. The author himself admits that the specimen was scarcely recognisable. ayo H. G. STUBBINGS Balanus eburneus Gould 1841 LOCALITY : Accra, on boat buoy, 14.1.51 (2 spec., i damaged). This is the only example of this species in the collection. The intact specimen measures 11-5 mm. carino-rostral diameter as compared with 14-15-5 mm. for material from Woods Hole, Mass., and is therefore not quite full-grown. The specimens are quite typical of B. eburneus in shell and valve structure. This American species has been recorded in the Old World on many occasions since first being reported by Ostroumoff (1892) from Sevastopol, Black Sea. Fischer- Piette & Prenant (1956) have summarized records of B. eburneus in the Black Sea and at Istanbul and give many localities on the Atlantic coast of North Spain. Its occurrence in the Adriatic has been reported by Kolosvary (19410, 19440, b) and in the Mediterranean by Bishop (1951) and Nilsson-Cantell (19380). Fischer-Piette & Prenant (1957) were surprised not to find B. eburneus in Portugal, south-west Spain or Morocco. There are no records for the West African coastline other than the above record from Accra. Its rarity suggests that it is not yet established on the West African coast but is occasionally imported in a breeding condition on ships. Balanus improvisus Darwin Balanus improvisus has a vast geographical distribution. Its presence on West European coasts and in the Mediterranean and Black Sea is amply documented, as is its presence on the Atlantic seaboard of North America. In the Tropical and South Atlantic there are a number of records from the West Indies (Darwin, 1854 ; Pilsbry, 1916 ; Henry, 1959) but fewer from the coast of South America, (Darwin, 1854 ; Weltner, 1897 ; Borradaile, 1916) and from West Africa. Broch (19270) recorded B. improvisus from Morocco and Gruvel (19076) from the Cape of Good Hope and (1912) from the Congo estuary. It was not present in collections reported on by the writer (19610, b, 1963, 19640, b}. It would appear, therefore, that B. improvisus is not common on the west coast of Africa. It is commonly transported by ships and records from estuarine areas such as the Congo, where not collected from ships' hulls, may represent shipborne introductions that have perished, or at least not multiplied greatly. It is evident that B. improvisus has not established itself in quantity as it has for instance in the Bosphorus (Neu, 1935, 1939) and in Japan where it has appeared in recent years (Kawahara, 1961, 1963). Balanus improvisus var. assimilis Darwin This variety of B. improvisus occurs in warm water and does not appear north of the Mediterranean whence it has been recorded by Gruvel (1903). It is known from the West Indies (Gruvel, 1903 ; Darwin, 1854 '> Nilsson-Cantell, 1928) and from Brazil (Weltner 1897). In the eastern Atlantic there are records from Mauretania and the Congo (Gruvel, 1912) and Bishop (1951) records specimens on ships coming from West African ports under circumstances suggesting that settlement took place there. THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 271 The possibility that white forms of B. amphitrite might be confused with B. im- provisus var. assimilis prompted a re-examination of much of the white " amphitrite " material to hand. No improvisus were found, the white " amphitrite " examined all proving to be thin-walled specimens of B. pallidus Darwin (= B. a. pallidus Darwin). These thin white forms all had opaque white lines opposite the internal ribs of the paries with thin, clear areas between, whereas Darwin expressly states that the translucent areas in B. improvisus var. assimilis correspond to the longitudinal parietal septa. The identity of the specimens was confirmed by the very long pallidus-type spur to the tergum. There can be no doubt, therefore, that this variety of B. improvisus is far from common in West Africa. Balanus amphitrite Darwin In considering B. amphitrite I have followed Harding (1962) in nomenclature. Hence the Darwinian varieties venustus, pallidus and stutsburi are dealt with separ- ately, the first as B. venustus and the other two as B. pallidus. The varieties com- munis Darwin, hawaiiensis Broch and denticulata Broch are combined as B. amphitrite amphitrite Darwin. Balanus amphitrite amphitrite Darwin Balanus amphitrite Darwin, 1854, 240, pi. 5, figs, ze, h, I. Balanus amphitrite hawaiiensis: Utinomi, igdoa, 43, text-figs. 1-3. Balanus amphitrite amphitrite: Harding, 1962, 274, pis. i, 2. LOCALITIES: Gambia: from seawater intakes of "African Queen", with B. pallidus and B. trigonus on B. t. tintinnabulum, 2.10.51, coll. M. H. Routh, B.M. 1952 . 10 . 2 . 5 (several spec.) . Sierra Leone: Kissy, Sierra Leone River (i) infra-littoral fringe on rocks, 19.7.55, coll. A. R. Longhurst, B.M. 1956.1.7.9 (19 spec.): (2) on bakelite panel exposed 22.3.54-14.6.54, a heavy encrustation (H.G.S.). Ghana: (i) Ankobra ferry, on logs with C. dentatus, 19.2.49 (many spec.); (2) the same, on copper sheathing of boat with epizoic juv. B. tintinnabulum (i spec.); (3) Accra, on boat buoy, 14.1.51 (15 spec.); (4) Labadi, Accra, on gastropod shell ; 22 -3 -58 (9 dead spec.) (H.G.S.); (5) Kpeshie Lagoon, Accra, on oyster shell, coll. D. T. Gauld (38 spec.). Nigeria: (i) Port Harcourt on hull of launch, (52 juv. spec.) (H.G.S.); (2) Opobo, Imo River, from concrete boat slipway (8 spec.) (H.G.S.). Considerable confusion has arisen over the identity of the forms variously regarded as varieties or subspecies, B. amphitrite communis Darwin (1854), hawaiiensis Broch (1922) and denticulata Broch (19276) doubtless stemming from the sketchy description of the earliest described. Utinomi (19600) advanced reasons for regarding hawaiien- sis and denticulata as identical, the former name having priority. At the same time Utinomi listed the criteria for distinguishing B. a. hawaiiensis from Darwin's var. communis. 272 H. G. STUBBINGS FIG. 14. Balanus amphitrite amphitrite Darwin : internal views of terga of six specimens ; a, removed from hull of " Atlantide " at Le Havre ; b, Kpeshie Lagoon, Accra ; c, Labadi, Accra ; d, Freetown, Sierra Leone ; e, f, Lagos, Nigeria (" Atlantide " Sta. 93) ; (a, c X28 ; b, d-f, Xi4). Harding (1962) has given a much-needed re-description of the original Darwin specimens. From this it is evident that the Darwinian var. communis, bearing the name in his handwriting, has the characters usually ascribed to denticulata or hawaiiensis by later authors, and just those listed by Utinomi as typical of hawaiiensis and distinguishing it from communis. The Darwinian name has clearly become associated with the wrong set of characters, through lack of sufficiently precise information on the original material. Hence, presupposing communis and hawaiiensis to be distinct, specimens conforming to Utinomi's summary of hawaiiensis characters should be termed B. a. communis Darwin, or in modern terminology B. amphitrite var. or ssp. amphitrite Darwin. Those with the characters listed by Utinomi for B. THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 273 amphitrite communis, if distinct, cannot bear this name but must receive a new varietal or subspecific epithet. Before taking such action it is prudent to consider whether the apparently clear-cut distinctions between " communis " and " hawaiiensis " in Utinomi (19600) can be maintained in practice. Material referable to these forms has been gathered from the thirteen localities named below and examined for these supposedly diagnostic characters : Bathurst, Gambia Inhaca Is., Mozambique Freetown, Sierra Leone Izmir, Turkey Labadi ~| Haifa, Israel Kpeshie Lagoon > Ghana Kuwait Ankobra Ferry J Le Havre, France Lagos, Nigeria Devonport, England Zwartkops R., S. Africa The most clear-cut character, after dissection, is the number of teeth on the labrum. In the 28 specimens examined the smaller numbers found varied from as few as two /three (on the two halves) in a specimen from Lagos to 4 or 5, all widely spread and all on the anterior border close to the edge of the median notch. Only the tooth nearest the mid-line was seated in the notch itself. There was then a jump to 8 or 9, or more teeth, of which many were situated in the notch in the typical close-set " denticulata " manner. A clear distinction is evident between a labrum with a few teeth (2 to 5) and another with numerous teeth (8 or 9 or more). According to Utinomi (19600) the number of parietal tubes in the rostrum is significant, there being about 16 in denticulata and only about 9 in communis. Using the same specimens, the number of tubes varied from n to 21, never as low as nine. The two with only n rostral parietal pores had numerous teeth on the labrum. Specimens from all " tube No. categories " had numerous teeth on the labrum. Only " three " (2-5) such teeth occurred in specimens with 12-17 rostral pores. There is no relation between number of rostral pores and number of labral teeth. The tergal features of spur shape and proportions and number of depressor crests are easy to observe. According to Utinomi in denticulata the spur is broad, over one-third the width of the basal margin of the valve, short and rounded or truncate distally, less than its own width from the basiscutal angle. In communis it is said to be longer than wide, and separated by its own width from the angle. Utinomi figures both forms (19600, text-figs. 20, c) though here the differences are not so marked as some that occur. Forms of " denticulata " type tergum occur with a much broader spur (see for instance the lectotype of B. amphitrite amphitrite in Harding (1962) pi. I, figs. e,f). The length of the spur also is variable. According to Harding (1962) a few of the specimens labelled " communis " by Darwin have the tergal spur narrow as figured by him (1854, pi. 5, fig. 21) but the majority of Darwin's specimens have a broad spur. A series of six terga of specimens from various localities (Text-fig. 14) shows this range of spur form. The first two (Text-fig. 140, b) 274 H. G. STUBBINGS show the broad tergal spur, comparable to that of the lectotype ; in c it is reduced in width and in d and e is so narrow as to be longer than broad. The specimen / has a similar narrow spur. Here the margins are roughly parallel and the spur is more sharply marked off from the basal margin of the valve. Occasionally terga (and scuta) are thin, so that the growth lines are faintly visible internally. More often they are more heavily calcified and are opaque with a smooth internal surface. Roughened areas occur on some terga and rarely may cover the whole internal surface (Text-fig. 14^). The depressor muscle crests are very variable in number and development. Five or six well-developed crests appears to be the normal number with one or two very short and ill-defined crests nearer the spur. The outer crest or crests towards the carinal margin of the valve are often less sharply defined (e.g. Text-fig. 14^, e). Additional small ill-defined crests may be present on the basal margin, e.g. Text-fig. 146 and/. Text-fig. 140 shows a tergum with very poorly developed crests, only two rather low crests being present with very faint traces of a further two inside these. By plotting the distribution of pairs of characters it is possible to see how far they are linked, even when the separate characters are not expressed numerically. Thus radii with horizontal tops are associated almost exclusively with numerous teeth on the labrum (Text-fig. 15^), but oblique radii may be associated about equally with few or numerous labral teeth. There is better correlation between form of radii and form of spur (Text-fig. 156) : a narrow radius with oblique summit appears to be associated exclusively with the longer and narrow tergal spur. A broad spur was not found with this type of radius. But the broad radius with horizontal top is not associated only with a broad spur. In one-third of the admittedly small sample of 12 specimens, the spur was definitely longer than broad. The number of well- developed depressor crests on the tergum (Text-fig. 150) is on average higher (6-43) on forms with narrow oblique radii than on forms with broader radii with horizontal summits (4-9). Similarly the average number of crests is higher where the spur is narrow (e) or there are few labral teeth (/,. A broad spur is almost always associated with numerous labral teeth (d) but the converse does not hold (cf. radii and labral teeth in (a)). Forms with a narrow-spurred tergum and few or many labral teeth occur with equal frequency. We can thus pick out a set of features characterizing a " well developed " type of animal, viz : broad radii with horizontal summits, broad tergal spur, numerous labral teeth and 5 depressor crests (or occasionally less) on the tergum. This is, in fact, the result arrived at by Utinomi (19600) and the above features characterize his interpretation of B. a. hawaiiensis Broch (= denticulata Broch). The opposite characters — narrow radii with oblique summits, a narrow and longer tergal spur, and few teeth on the labrum and more than 5 depressor crests on the tergum also occur in association. These are the characters, again among others, listed by Utinomi for B. a. communis Darwin. But, as the diagrams in Text-fig. 15 indicate these associations are not constant — a narrow tergal spur, or narrow radii occur as frequently associated with numerous labral teeth. Hence, specimens occur with a " mixed " assemblage of characters and the question arises as to which of these must be regarded as determining the name to be assigned to the specimen. Of the THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 275 (a) (b) (c) CO ••• 2j.g sh — 55 7 Lab. teeth <5 "3K • Spur 1 t » 9 Crests 5 ::: # , , i , 4 • ••• ob. h. ob. h. ob. h. radii radii radii oO Lab. teeth <5 • •• h ••• (d) :*: Crests K &• (e) Crests I sh Spur :•: (0 on Turritella sp. coll. R. Bassindale, B.M. 1953.2.5.2 (several spec.); (59) trawled off Accra, with C. patula on Cymbium porcinum pres. E. Salzen, B.M. 1957.6.3.1 (46 + 2gjuv. spec.). Nigeria: Lighthouse Beach, Lagos: (i) on cork cast up on beach, with B. t. tintinnabulum and C. patula (several dead spec.) ; (2) on a stone cast up on beach, with a Gorgonian (i spec.). To avoid excessive repetition in listing the records of this very abundant species the stations off Accra made by Mr. Bassindale have been cited above by number only. Fuller station details are given in Tebble (1955) and Bassindale (1961). Specimens have been collected from shells of the following Mollusca and Crustacea : Gastropoda : Turritella annulata Kiener, Solarium granulatum Lam., Crepidula porcellana Lam., C. porcellana var. " Garnot " Adanson, Natica collaria Lam., N. fulminea Gmelin, Murex hoplites P. Fischer, M. bourgeoisi Tournouer, Phos grateloupianus Petit de la Saussaye, Nassa tritoniformis Kiener, N. cf. plicatella A. Adams, Fusus boettgeri v. Maltzan, Olivancillaria (Agaronia) hiatula Gmelin, Cymbium porcinum Lam., Cancellaria cancellata Linn., Drillia rosacea Reeve, Clava- tula muricata Lam., C. (Pusionella] nifat Adanson, Surcula coerulea Weinkauft, Turris pluteata Reeve, T. (Crassispina] carbonaria Reeve, Genota mitraeformis Wood, THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 281 Terebra senegalensis Lam. Lamellibranchiata : Cardium costatum Linn., C. ringens Gmelin, Dosinia africana, Gray. Crustacea : Balanus tintinnabulum var. tintinna- bulum (Linn.), Apiomithrax bocagei (Osorio), Calappa rubroguttata Herklots. The references in the literature to B. amphitrite var. venustus, Darwin by name are listed above. That they all refer to this variety is doubtful as discussed below. Weltner's record (1897) from Nagasaki was questioned by Nilsson-Cantell (19320) presumably on account of its geographical separation from other records, as no morphological questions were raised. There are only five illustrations of the variety in the literature. One, (Tarasov & Zevina, 1957) is a copy from Nilsson-Cantell (1925) which is not here considered to be var. venustus. There are thus only three valid illustrations ; of the whole shell (Darwin, 1854, pi. 5, fig. 20), of a scutum and tergum (Nilsson-Cantell, 19380, text-fig. 8) and of shell, opercular valves and appendages (Harding, 1962). Darwin's illustration of the hard parts of his varieties of B. amphitrite were not complete, which has undoubtedly led to some difficulty and misunderstanding of the varieties he proposed. Opercular valves of venustus are present in the Darwin collection of Cirripedia in the British Mudeum (Natural History) and these have been redescribed and figured by Harding (1962). The spur of the tergum is short and broad. This with the characteristic pink colour of the shell will in general distinguish the variety. From the very extensive Ghanaian material it is possible to give an extended description of the variety. The shell is typically pink, sometimes bright but more often rather dull or " washed out ". Pale pink specimens also occur, but the shell is never red and only rarely approaches the purplish pink, of deeply coloured B. pallidus. Narrow vertical stripes of a deeper pink are present on the paries but may be absent or difficult to distinguish from the ground colour. Rarely the pink stripes are the only colour present (cf. some var. stutsburi). Shells with an orange tint occur rarely among normal pink ones. From the many young specimens present it is evident that for some time after metamorphosis the shell of B. venustus is white. Pink pigment is developed quite suddenly as there is a rapid transition from white to pink shell. Young specimens thus have shells pink basally and with white summits to the compartments. The age, or size, at which pigment is first produced is very variable and a few never do so and remain pure white. When pigment is produced early in life, older specimens may be completely pink-shelled due to the disintegration of the tops of the com- partments. Rarely, specimens occur with pigment on the margins of the paries only. When rather elongated these tend to resemble the B. amphitrite var. pallidus of Darwin. Such a specimen was found in the R. Densu collection listed above. It had a typical B. venustus tergum as here described. The compartments are smooth or irregularly folded. Smooth specimens tend to be less heavily calcified and somewhat translucent so that the septate parietal canals are visible externally. The radii are moderately developed or narrow with very oblique summits when intact, white above and pink below. When intact the orifice is highly dentate. The epidermis is thin and not persistent. It bears hori- zontal rows of short fine hairs corresponding to the growth increments. Alae are well developed with less oblique summits than the radii. Internally the growth lines 282 H. G. STUBBINGS are prominent on alae and sheath. The sheath is overhanging, about one-third to one-half the length of the compartment. The compartments are prominently ribbed internally. The scuta of three West African specimens are illustrated (Text-fig, ija, c, e). The tergal and basal margins are of equal length, the basal angle being rounded off so that the slightly convex basal margin curves into the tergal. Growth lines are distinct but not prominent, the epidermis persistent in part and with hairs along the growth lines. The apex is acute when uneroded. There is a purple patch in the upper part of the valve but the apex is usually colourless. Rarely the whole scutum FIG. 17. Balanus venustus venustus Darwin. Internal views of the paired scutum and tergum of three specimens from off Accra : a, b, from Sta. 17 ; c, d, from sta. 31 ; e, f, from sta. 87 (a-d xi? ; e, f Xiq), THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 283 is colourless. There is no direct relationship between shell pigmentation and that of the opercular valves ; of two shells with virtually identical pigment development, one may have a very small pale purple patch on the scutum and the other a deep purple patch covering the apical third of the valve. Very young colourless shells have colourless scuta. The outer surface is faintly striated from the apex. Inter- nally the valve is thick with a prominent adductor ridge. The deep groove below the ridge and the deep depressor muscle pit between them delimit a raised area that in some specimens has the appearance of a low broad ridge. There is not a true second ridge. The adductor muscle pit is large and distinctly marked. The articular ridge is large, straight and reflexed over the articular groove. The inner surface of the valve is faintly striate in the area above the adductor ridge. The striations extend on to the articular ridge. The terga corresponding to these scuta are illustrated in Text-fig. 176, d, f. The valve is flat, triangular with a short broad spur about one third the width of the basal margin. Growth increments are marked by distinct grooves externally. The spur area also is demarcated by grooves only. The articular margin is slightly concave, the carinal margin convex, the basal margin on the carinal side straight or more usually slightly concave. Internally the articular ridge is long but not very prominent, the articular groove long and deep. Crests for the depressor muscle are only feebly developed and about four in number. The internal surface is more or less striated longitudinally, the striations being finely tuberculated. The above descriptions of scutum and tergum are in close agreement with the figures given by Harding (1962, pi. 6, figs, dr-g) for the lectotype of B. venustus venus- tus. Opercular valves of " venustus " have been figured by Nilsson-Cantell (1925, 1938^), the two figures representing very differently shaped valves. The Indian specimens (Nilsson-Cantell, 1938^), have valves similar to those shown here though there are some differences. Thus the scutum has a basi-tergal angle of < 90° and the corner of the valve is not rounded off : also, the articular ridge projects a little in front of the valve margin. The tergum is more angularly triangular mainly due to the very straight carinal margin. In other respects these valves appear to agree reasonably with the West African venustus. The variety figured by Nilsson-Cantell (1925) has very different valves. The tergum has a long narrow spur and many crests for the depressor muscles. The shell is said to be red (" rot ") and the colour to agree best with that of venustus. But no venustus in the present collection from West Africa is red. The statement that the stripes are not confluent — a character attributed to stutsburi — is not valid as stutsburi has often clear distinct stripes. In the scutum the depressor muscle pit is said to be indistinct (" unbedeutend "), which is not the case for venustus as here considered. It is considered that the specimens are referable to Darwin's var. Stutsburi, i.e. B. pallidus Darwin of this work. Excluding the rejected figures given by Nilsson-Cantell (1925) and copied by Tarasov & Zevina (1957) there were no illustrations or valid descriptions of the appendages of B. venustus until Harding (1962) published photographs of a lectotype from Natal, The mouth-parts of a specimen from Accra are described and figured 284 H. G. STUBBINGS here. The labrum (Text-fig. i8a) has the two halves of the margin set at an angle so that the median toothed area is prominent. There are two or three teeth on either side of the wide V-shaped notch. Sparse short hairs are present round the teeth and in the upper part of the notch. Hairs are absent in Harding's photograph (1962, pi. 6c). The palp (Text-fig. iSb) is rectangular, tapering slightly towards the rounded extremity. The anterior margin is slightly concave, the posterior convex. The hairs on the concave margin are short. A band of similar short hairs on the inner face of the palp merges into the long hairs of the extremity. A row of long hairs begins on the face, runs to the posterior margin of the palp and follows this margin to merge with the apical hairs. The mandible (Text-fig. i8c) has five teeth, the third to fifth lying close together. The first is normally simple but occasionally bifid at the tip. The second is simple or with a small cusp which when close to the tip renders this bifid. The third to fifth teeth are usually blunt from use but in the developing cuticle are sharp. The third tooth has one or two secondary cusps but occasionally may be simple. The FIG. 18. Balanus venustus venustus Darwin. Mouth parts of a specimen from Sta. 87 off Accra, a, labrum (X58) ; b, palp (X45) ; c, mandible; d, maxilla I (both x 100) ; e, maxilla II (X45). THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 285 fourth is double with two sharp equal points when new but usually worn down in use. The fifth is very close to the lower angle and when worn apparently confluent with it. In the developing cuticle it is distinct and there are two sharp stout spines below it on the lower angle. Harding's illustration (pi. 6n) would appear to be of a worn mandible as the lower angle is blunted. Maxilla I (Text-fig. i8d) has a straight biting edge. The lower pair of spines is about as large and stout as the upper pair. There are five intermediate spines, though occasionally four or even only three. There is a small tuft of short stout hairs or fine spines on the lower angle. Of six pairs of maxillae examined, in two there was a difference in number of intermediate spines in the left and right maxilla. Maxilla II (Text-fig. i8e) is broadly oval tapering slightly to the tip, which is densely clothed with long fine setae. There is a very small lower lobe. Cypris larvae were present on some mollusc shells. Two from a valve of Cardium ringens measured 0-75 mm. long x 0-31 mm. high x 0-27 mm. broad 0-73 X 0-30 X 0-26 and a third unattached specimen was 0-676 mm. long. The present material comes entirely from below low-tide level. According to Buchanan (1958) B. venustus is a common member of the sessile epifauna of the " inshore fine sand " and " sandy silt " communities off Accra, i.e. from 3-20 fathoms (5-5-36-5 m.) depth. The present collections show it to be abundant from 10-20 m. and rather less common from 20-25 m., that is over the deeper part of the fine sand community and the shallow part of the sandy silt community. Here it is abundant on shells of Cardium ringens Brug. and especially Turritella annulata Kiener, both living and dead. As many as 74 specimens, all but three alive, were found on a single Turritella shell. It is by no means confined to these molluscs as the extensive list above testifies. The two records of material cast on shore at Lagos suggest that venustus occupies a similar habitat there. Shallow water dredgings in the Bonny estuary, Eastern Nigeria, failed to obtain this variety. Records from the " Atlan- tide " and " Calypso " Gulf of Guinea expeditions (Stubbings, 19610., b) indicate that B. venustus is distributed from Gambia to the R. Congo. The fact that most speci- mens from the latter locality came from 25 m., or deeper and were dead, suggests that the normal habitat of B. venustus is in the 10-20 m. depth zone. A number of specimens from Accra Sta. 2 were attached in the grooves on the convex side of Cardium ringens valves. All were orientated along the grooves and elongated along the rostro-carinal axis, with the carinal end directed towards the highest part of the shell. Thus the few barnacles in the shallower part of the grooves near the umbo were orientated at 180° to the majority in the deeper furrows nearer the shell lip. B. venustus was not obtained from the intertidal zone by either of the above ex- peditions, nor by Bassindale or the author. Neither was it found on mangrove roots, a common habitat of estuarine species in West Africa. It is evidently confined to the open sea. This is of significance in that it supports the view that the form found by Gruvel (1912) on mangroves in the lagoon at Grand Bassam, Ivory Coast, 286 H. G. STUBBINGS and that reported by Nilsson-Cantell (1925) on mangroves from the Cameroons, are not venustus. The mangrove fringed estuarine creeks of tropical West Africa are the habitat of B. pallidus Darwin. B. venustus appears to be very widely distributed in tropical waters. Whereas now it is perhaps best known from West Africa it is also found in S. Africa (Darwin), Ceylon (Annandale, 1906) India (Nilsson-Cantell, 19380;). Weltner's record from Nagasaki (Japan) has already been noted as doubtful (Nilsson-Cantell, 19320). That it has not been recorded by Hiro (Utinomi) from Japan confirms this doubt. This absence from the literature on Far Eastern barnacles casts further doubt on the record by Tarasov & Zevina (1957). Balanus venustus niveus Darwin Balanus venustus niveus: Harding, 1962, 286, pi. 7, figs. a-r. Balanus venustus niveus: Stubbings, 19646, 340. A number of more or less translucent white " B. amphitrite " from the Sierra Leone river, Port Harcourt and Bonny, Nigeria, were at first assigned to this variety. A further examination cast doubts upon this determination. In the first place all the specimens were associated with more or less pigmented B. pallidus. A more detailed examination after removal of epizoic growth and debris from the compart- ments and opercular valves showed that an appreciable number had slight traces of pinkish-purple pigment, either as small streaks or spots on the body of the paries or as small marginal lines. Patches of pigment were present also on the scuta, though smaller than on well-pigmented individuals. As no pigment is recorded for the Darwin type of var. niveus, it seemed probable that these specimens were really very poorly coloured pallidus. In typical specimens, of course, the tergum of pallidus is very distinctive having a long spur and much emarginated basal margin whereas that of niveus is short and broad and obliquely truncated In the terga of the specimens in question the spur is not typical of either form, being shorter than in pallidus but less broad in proportion than in niveus. They could be regarded as rather atypical specimens of either variety. As, however, pigmented specimens in the same collections had similar terga it is considered that the whole sample in each case belonged to one variety and the material is all regarded as B. pallidus. There is not, therefore, any material in the collections under review that can be assigned with certainty to B. venustus niveus. In all the West African collections examined by the author with one exception, B. v. niveus has been absent. In material from the Congo estuary (Stubbings, 19646) a few specimens were found which appeared to belong to this variety. As concluded in that report, B. v. niveus is very rare in West African waters and may well occur sporadically as a shipborne immigrant. The above record from the R. Congo is the only one from the eastern Atlantic though Kolosvary (1951) cites the Mediterranean and Darwin (1854) South Africa. Neither record has been confirmed. Further afield there are a number of scattered records of niveus from the Indian Ocean and the Malay Peninsula. As the variety is THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 287 common on the American shores from Massachusetts to Southern Brazil and according to Kolosvary (1943*2) occurs at Montevideo, Uruguay and even in Tierra del Fuego, there is ample scope for shipborne transport of this variety as with other forms of " B. amphitrite " and some at least of these scattered records may represent such local introductions. ltd luti us fallax Broch Balanus (Hesperibalanus) fallax Broch, 1927, 26, pi. 2, figs. 12-17 » pi- 3. figs- 18, 19 ; text-figs. VII-IX. Balanus (Hesperibalanus) fallax: Utinomi, 19596, 402, text-fig, i. Balanus occidentalis Stubbings, igdia, 34, text-figs. 8-n ; 19616, 189. Balanus fallax: Stubbings, 1963, 30, text-figs. 10, n. LOCALITIES: Ghana: Accra, (i) Sta. 133, 51 m. (i spec.); (2) 14 m., in sponge, 27.4.51 (i spec.); (3) Osu Fisheries Sta., 19.5.51 (4 spec.); (4) near R. Densu, 15 miles from Accra, 16.5.51 (i spec.). The above material has been compared with the original material designated as " types " by Professor Broch. The shell and appendages have been redescribed in some detail (Stubbings, 1963) and it is only necessary here to comment on points of interest in the Ghanaian material. The specimens exhibit in general a more strongly developed shell. That from 14 m. in a sponge is mottled pink like the type material but the others have regular lines of pigment except towards the margins of the compartments where the lines are broken up into blotches or spots of a deep pink colour. None of the specimens shows longitudinal striations on the scuta. The internal sculpturing of the scuta is strongly developed (Text-fig, iga, c, e). The adductor scar is a distinct pit deepest on its apical border. There is no adductor ridge but the valve is appreciably thickened below the adductor pit. In the specimen from the R. Densu (Text-fig, ige) there is a faint suggestion of a short ridge parallel to the basal margin immediately below the adductor pit. The pit for the lateral depressor muscle is more marked in these somewhat heavier valves. The basitergal angle according to Broch (19270:, figs. 16, 19) and Stubbings (1963, fig. 10) is 90° or more. In one specimen (Text-fig, igc) this angle is less than 90° though in another specimen (Text-fig. 190) the angle is distinctly obtuse. A point not brought out in Broch' s figures is the scroll-like form of the upper part of the articular ridge of the scutum. This is well marked in the specimens here illustrated. The degree of development of this scroll affects the shape of the lower end of the articular ridge. If the scroll is weak, as in Broch's illustration and Text- fig, ige, then the ridge ends abruptly. But when the scroll is strongly marked (Text-fig, iga, c) then it extends to the lower part of the articular ridge and the ridge ends less abruptly, curving into the plane of the valve. Terga from two barnacles from Osu are shown in Text-fig. 196, d. The chief points of interest are the presence of fine hairs on the epidermis of the growth ridges, the broad spur and the strongly developed crests for the depressor muscles. There is a distinct tendency for small incipient crests to develop near the base of the spur. The specimen from near the R. Densu (Text-fig, igf, g) has a less robust tergum. 288 H. G. STUBBINGS FIG. 19. Balanus fallax Broch. a, b, internal views of scutum and tergum of a specimen from the Osu Fisheries Station, Accra, Ghana ; c, d, similar views of valves of another specimen from Osu : e, f, g, internal view of scutum and external and internal views of a tergum of a specimen from the R. Densu, Ghana (all X28). THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 289 The spur is much narrower and is pointed at its extremity instead of being rounded as in the heavier Osu specimens. There are no incipient crests in the concave basal margin of the valve. In both these points it agrees with the specimen figured by Utinomi (19596). Illustrations of the mouth appendages have been given by Broch (19270) and Stubbings (19610, 1963). A few notes on minor variations may be given. The teeth on either side of the labrum (Text-fig. 200) may be directed obliquely away from the mid-line instead of standing perpendicular to the border of the labrum. The palp (Text-fig. 2O&) agrees with that figured by Stubbings (19610). It is oval or somewhat rectangular, the anterior margin being almost straight, the posterior and distal margins curved. There is a double row of pectinate setae along the anterior margin. The mandible (Text-fig. 2oc) has been fully described previously. The third and fourth teeth are somewhat bifid and the lower angle has 3 more or less complex teeth, sharply pointed in some specimens but blunt in others. This is probably a matter of wear only. The setae on the lower margin may be described as stout. Whether they should be regarded as " spines " depends on the degree of individual develop- ment and a somewhat arbitrary interpretation of the point where a stout seta becomes a spine. The setae on the R. Densu mandible figured were appreciably stouter than those on an Osu specimen. The first maxilla of the Osu specimen (Text-fig. 20^) had the small spine lateral to the two large upper spines, as described by Broch. In the R. Densu specimen this spine was absent. The second maxilla is oval and tapers distally to a rounded point. The lower lobe is semicircular. In the R. Densu specimen the second maxilla was much narrower than in the Osu specimens. The segmentation of the cirri was given by Broch (19270) and in a paratype by Stubbings (1963). The number of segments in cirri I and VI was appreciably lower in the paratype. The segmentation in the cirri of the paratype agreed closely with that of B. occidentalis (= B. fallax) in Stubbings (19610). Three new counts of segments are now available (Table 6). They agree substantially with earlier counts. It may be noted that in the Accra specimen the number of segments in cirrus VI equalled that given by Broch. In no specimen so far examined has there been 15 segments in the anterior ramus of cirrus I as described by Broch. TABLE 6 Specimen I II III IV V VI Accra, 14 m. . . . . . 12-9 9-7 n-io 18-18 21-22 22-22 R. Densu ....... 13*7 10-10 10-9 18-14* I7'23 17*18 Osu ........ 11-7 9-8 9-8 n*-i4* 19-18 20-21 Mean ....... 12-7 9-8 10-8 17*17 19-21 19-20 TABLE 6. Number of segments in the cirri of three specimens of B. fallax Broch from three localities in Ghana. * tip of cirrus missing. The presence of a dorsal process at the base of the penis (Stubbings, 1963) has been confirmed. It is sharply conical, slightly curved and bears a single long seta (Text-fig. 20/). B. fallax is known only from the African coast, from Algiers in the Western H. G. STUBBINGS FIG. 20. Balanus fallax Broch a, labrum ; b, palp ; c, mandible ; d, maxilla I ; e, maxilla II ; /, base of penis with dorsal spine (or-c, e, f, X95 ; d x 180). Mediterranean, down the west coast as far as Cape Ledo in Angola. Where more intensive collecting has been done, e.g. Morocco, French Guinea and Angola, there are a number of records, suggesting that the species is common. But there are few specimens in the Ghana collections and none at all from the Gulf of Guinea, suggesting that B. fallax is not uniformly distributed over the West African coastline. Balanus (Conopea) calceolus Darwin LOCALITIES: Ghana: Accra, (i) Chorkor, on the horny stem of an alcyonarian, - . 3 . 49 (2 dead and without opercular valves) ; (2) Chorkor, on a violet and yellow alcyonarian and other alcyonarians, 1.2.50 (n + 13 dead spec.); (3) Chorkor, on THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 291 the same violet and yellow alcyonarian, 12.5.50 (i spec.) ; (4) Christiansborg shore, on the same alcyonarian sp., 19.11.49 (4+3 juv. spec.); (5) 10 miles off-shore, on a white alcyonarian, 8.5.49 (3 + J dead spec.) ; (6) off Accra, on the violet and yellow alcyonarian, 13 .11.51 (i spec.) ; (7) 15 m. from Accra beyond Densu 16 . 5 . 51 (6 spec.). Specimens from locality (7) above contained developing eggs of average size 222 X 109 (i. B. calceolus is known from the Mediterranean and the Atlantic off the Straits of Gibraltar (Darwin, 1854 ; Caziot, 1921 ; Utinomi, 19596). On the West coast of Africa it is recorded from Rabat, Morocco (Broch, 19270), Cap Blanc, Mauretania (Broch, 19246), Sierra Leone (Nilsson-Cantell, 1928), Ghana (Gauld, 1957), the Congo and Angola (Nilsson-Cantell, 19386 : Stubbings, 1963, 19646). From South Africa Stebbing (1910) records it from Bird Island, Algoa Bay. B. calceolus has therefore been found at localities spaced along the whole west coast of Africa. There are some considerable gaps between known localities. It is surprising that there are no specimens in the collections of the IFAN, Dakar, in view of the extensive collecting done by that Institution. The absence of records in the Gulf of Guinea may be due to less intensive collecting, or may represent a dis- continuity in distribution clearly seen for several other species. Proceeding eastward it is known from Wasin, E. Africa (Gruvel, 19070:), Madras (Darwin, 1854 ; Nilsson-Cantell, 19380), the E. Indies (Broch, 1922, 1931 ; Pilsbry, 1916), Japan (Weltner, 1897 ; Hiro, 1937 ; Utinomi, 1949, 1958) and from Western Australia (Weltner, 1897). B. calceolus has therefore an " old world " distribution in warm to tropical seas, extending from the eastern Atlantic eastward to the Indian Ocean and the seas bordering S.E. Asia. It has not been recorded from the remainder of the Pacific and is unknown from the western Atlantic Ocean. TETRACLITA Schumacher Very few specimens of Tetradita have been collected in West Africa and the several groups have been assigned to different species by the respective authors. Pilsbry (1916) and Gauld (1957) each record one species from the coast. I disagree with Gauld as to some of his material and consider it belongs to a third species T. divisa Nilsson-Cantell (1921). It is questionable whether there are indeed three species in the area but much more material is required from the old collecting sites and elsewhere before an authoritative pronouncement can be made as to how many and which species occur there. The three species in question are discussed below. Tetradita divisa Nilsson-Cantell 1921 Tetradita divisa Nilsson-Cantell, 1921, 362, pi. 3,* fig. n, text-fig. 83. Tetradita divisa: Hiro, 1939, 275, text-fig. 15. LOCALITY: Ghana: Hospital Reef, Axim, 13.4.49 (2 sPec- one without soft parts or opercular valves) . This species is closely related to T. purpumscens (Wood) with which it agrees in having a transversely elongated scutum. Whereas T. purpurascens has the spur of ZOOL. 15, 6. 24 2Q2 H. G. STUBBINGS the tergum united with the basiscutal angle and the valve elongated, T. divisa has a triangular tergum with distinct short and broad spur, separated from the basiscutal angle by about half its width (Text-fig. 2ia). Nilsson-Cantell (1921) states that the scutum has no adductor ridge or crests for depressor muscles, but Hiro (1939) finds the adductor ridge may or may not be present and figures (1939, fig. 146) a scutum with a ridge. Zevina & Tarasov (1963) state that the ridge is little developed and do not show it in their figure. The present complete specimen has an adductor ridge (Text-fig. 216). There are no crests for the lateral depressor muscles but a series of about 6 shallow pits for the muscle insertions. The compartments have well developed radii with horizontal summits. In both specimens the walls are worn, in one so much so that only faint traces of the radii remain. The growth lines on uneroded compartments are fringed with hairs. There are up to six rows of pores in the paries, rather more than described by Hiro. No FIG. 21. Tetraclita divisa Nilsson-Cantell. a, internal view of tergum, b, of scutum (both X4o) ; c, labrum, d, palp, e, mandible,/, maxilla I, g, maxilla II (c, e-g, x 160 ; d, x 120). THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 293 trace of basis remains in one specimen. In the other there are fragmentary remains of a rather thin calcareous basis near the walls. The central area of the basis is wanting. Hiro states that the basis in T. divisa is calcined peripherally only. The mouth parts agree substantially with the description of the species by the above authors. The labrum (Text-fig. 2ic) is even more weakly bowed than in Nilsson-Cantell's figure, being almost straight from side to side with minor irregu- larities. There are three or four slight prominences, scarcely teeth, on either side of the middle line, as against only two in Hiro's specimen. The palp (Text-fig, zid) is very similar to Nilsson-Cantell's figure even to the rather brushlike plumose setae on the anterior edge. The mandible (Text-fig, zie) has a subsidiary cusp or cusps on all but the first tooth as in Nilsson-Cantell's and Zevina & Tarasov's figures. The fourth tooth has several sharp-pointed cusps as figured by the above authors. Maxilla I (Text-fig. 2i/) agrees very closely with those figured by Hiro and Nilsson- Cantell. There are minor differences in the arrangement of the central spines rela- tive to the notch in Zevina & Tarasov's figure. Maxilla II (Text-fig. 2ig) has been figured by Zevina & Tarasov (1963), who show no plumose setae. Plumose setae in the present specimen are restricted to a group at the tip of the appendage and to a second group on the rather protuberant lower part of the anterior border. The number of segments in the cirri is given in Table 7. TABLE 7 I II ill IV v VI Ghana . . . . . . . . 10-5 6-5 5-5 ii-n 12-13 I4'I4 Nilsson-Cantell 8-4 6-5 6-5 12-12 12-12 12-12 Hiro ......... 10-6 8-7 7-6 11-12 14-14 14-14 r, . , T, f 8-5 6-6 6-4 --8 8-7 9-8 Zevina and Tarasov . . . . . . < J \ — 6-5 6-4 7-6 11-9 10-9 TABLE 7. Comparison of number of segments in the cirri of a specimen of T. divisa from Ghana with the number given by Nilsson-Cantell (1921), Hiro (1939) and Zevina & Tarasov (1963). There is close agreement between the number of segments given by the earlier authors and that found in the present specimen. The figures quoted by Zevina & Tarasov (1963) for cirri IV to VI are appreciably lower. T. divisa was originally described (Nilsson-Cantell, 1921) from Sumatra and Java from material collected by C. W. A. Aurivillius. The only subsequent records are by Hiro (1939) from the east coast of Formosa and by Zevina & Tarasov (1963) from the South China Sea. Its occurrence on the west coast of Africa extends enormously the known distribution of this species and suggests a probable distribution over much of the tropics of the Old World, as it is unlikely this will prove to be an isolated occurrence. Tetraclita purpurascens Wood T. purpurascens is included here on the authority of Gauld (1957) who recorded specimens from Victoria, Cameroon and Axim, Ghana. There are no other records of this species from West Africa. It is stated that the specimens corresponded 294 H. G. STUBBINGS " exactly " with Darwin's (1954) description. Dr. Gauld informs me (in litt.) that this statement referred specifically to a group of specimens from Victoria and might not be true for the material from Axim. It is possible that this Axim material was originally part of that here described as T. divisa Nilsson-Cantell. Unfortunately the Victoria material is no longer available for re-examination. Tetraclita squamosa squamosa (Bruguiere) Tetraclita squamosa squamosa: Pilsbry, 1916, 251. Pilsbry (1916, 252) cited two groups of specimens from " West Africa " and " Cape Palmas " (Liberia) which he assigned to the typical subspecies of T. squamosa at the same time noting certain differences. It has not been recorded subsequently from West Africa. PYRGOMA Leach 1817 Pyrgorna anglicum G. B. Sowerby LOCALITIES: Ghana: (1-5) Accra: (i) Sta. 45, 35m., on dead coral (i spec.); (2) Sta. 131, 37 m., attached on or near base of the living solitary coral Balanophyllia formosa Gravier, 2.5.51 (5 + i dead spec.); (3) Sta. 133, 51 m., on a single dead solitary coral (6 dead spec., also remains of the bases of 12 others) ; (4) no locality, on Oculina (Schizoculina) fissipora Milne-Edwards & Haine received from Dr. D. T. Gauld, University of Ghana (4 dead spec.); (5) on Caryophyllia sp. -.2.55, coll. J. B. Buchanan (7 spec.); (6) Axim. "from coral Astrangia sp. " -.6.55, coll. J. B. Buchanan (3 spec, without basis or soft parts). FIG. 22. Pyrgoma anglicum Sowerby from Accra, a, labrum c, maxilla I, d, maxilla II (all XI35). , b, mandible, THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 295 Nigeria : Bonny River Estuary, off Bonny, from a coral dredged in shallow water, 28.1.58 (21 + 3 dead spec.) (H.G.S.). Dr. Buchanan states (in litt.) that he collected Pyrgoma in 1954 from 20 m. off Takoradi, Ghana on Oculina sp. and in 1955 from the shore at Dixcove, Ghana on Porites sp. and Siderastrea radians. Illustrations of some of the appendages have been given by Broch (19270). The labrum is of the high conical form described by Broch but has three distinct teeth, not one, on either side (Text-fig. 220) , thus agreeing with Darwin's description of the labrum of this species. The outermost of the three labral teeth is smaller than the other two. The palp is oval and of the typical form. The mandible (Text-fig. 226) has a rather straight edge, the teeth being rather broad and blunt. The third tooth is slightly bifid. In Broch's specimen the second tooth was slightly bifid. The lower angle is composed of two blunt rounded teeth. There are very few hairs on the sur- face of the appendage. The first maxilla (Text-fig. 22c) agrees closely with that figured by Broch. The edge is straight and without a notch. The large upper spine is followed by eight others of which the first, fifth and last two are rather larger than the others. The surfaces of the appendage are hairy. The second maxilla (Text-fig. 22d) is narrowly oval with a small oval lower lobe. It is densely setose along the inner margin and apically. The segmentation of the cirri is shown in Table 8, agreeing reasonably well with that given by Broch (19270) : TABLE 8 I II ill IV V VI 16-9 7.9 8-9 15-16 18-18 19-19 . 17-8 7-9 8-10 17-19 18-22 20-23 Accra ..... Broch TABLE 8. Number of segments in the cirri of a specimen of P. anglicum from Accra, compared with Broch's (1927) figures. The segments of the inner ramus of cirrus I are moderately protuberant, those of the outer ramus scarcely so. Cirrus II is very small. Hooked spines occur on segments 2-6 of the inner ramus of cirrus III. Cirri IV-VI have rather fewer segments than in Broch's specimen. In cirrus IV the first four segments are dis- tinguishable though probably fused. The number of hooked spines on each segment of the inner ramus of cirrus IV, compared with the number given by Broch (19270) , is shown in Table 9. TABLE 9 Segment .....1234567 8 910 Accra . . . . .12222221 +r i o Broch 1221222 2 i r fused segments r = rudimentary spine TABLE 9. Number of hooked spines on the lower segments of the inner ramus of cirrus IV in P. anglicum. 2g6 H. G. STUBBINGS The penis is long and closely annulated and has a basidorsal point. The coral host of the Bonny specimens eventually grows over the barnacle and entombs it. From the nodular form of the coral specimen and the fact that several nodules revealed enclosed Pyrgoma when broken open it is possible that many Pyrgoma are so buried. Scuta and terga are present in many of these buried shells so overgrowth of the coral presumably occurs whilst the barnacle is still alive. Rees (1962, 1966) has discussed the distribution of P. anglicum with particular reference to its coincidence with that of the solitary coral Caryophyllia smithii in British waters. Rees also indicates the wider distribution of the species southwards to Spain, the Mediterranean and West Africa. To Rees's references may be added Sicily (Kolosvary, 1951) and the Cape Verde Archipelago (St. lago and Brava) (Stubbings, 1963). These localities confirm Darwin's (1854) earlier records. Gauld (1957) cites Astrangia and Caryophyllia as host corals off Accra, Ghana, but these coral identifications need confirmation. Gruvel (1903), not quoted by Rees, reported specimens in the Paris Museum from Algeria on Coenocyathus anthophyllites and others from " Travailleur " St. 34 on Dendrophyllia cornigera, the same host as recorded by Broch (19270) from the coast of Morocco. The present records from Bonny, E. Nigeria represent the furthest kown southerly limit of the species in the Eastern Atlantic. The Far Eastern records cited by Rees may be extended by adding Madras (Nilsson-Cantell, 19380) and Seto, Japan (Hiro, 1937). Hiro (1935) also apparently accepts Broch's (1931) record of " Pyrgoma sp. afL anglicum " from Sagami Bay, Japan as a true P. anglicum. CHELONIBIA Leach 1817 Four species of Chelonibia have been recorded from West Africa. Chelonibia testudinaria (Linnaeus) LOCALITIES : Senegal : from scutes of a turtle, coll. Captain Moloney, B.M. 85.5 (5 large and 2 very small spec.). Ghana, Accra, 8, pres. Dr. D. T. Gauld (no further data). The Ghanaian specimens range in size up to 30 mm. carino-rostral diameter. All the shells are disarticulated and the soft parts wanting. The " teeth " on the radii are poorly developed or wanting in these small specimens. The species is found in Ghana on turtles coming ashore on the sandy beaches and killed by the villagers, but few specimens seem to have been collected. It has been recorded previously from " equatorial Africa " (Gruvel, 1905), Cameroons (Nilsson- Cantell, 1921), Ghana (Gauld, 1957) and Senegal (Stubbings, 1965). It has not been recorded from the more temperate waters of north-west Africa. A number of records for the Mediterranean including the Adriatic are summarized by Kolosvary (1943*2, 1951). Barnard (1924) records it from Table Bay, South Africa. Further afield C. testudinaria is generally distributed throughout the tropical and warmer temperate seas. THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 297 Chelonibia caretta (Spengler) Records of this species on the West African coast are few and widespread. Darwin (1854) recorded it from the " West Coast of Africa " without further indication of locality. Broch (19246) described it from the Baie du Levrier, Mauretania and in the same year Barnard (1924) from Table Bay, South Africa. Pilsbry (1916) also mentions specimens from the Cape of Good Hope. There appears to be no other record from the west coast of Africa. It is not represented in any of the recent expedition collections. Neither is it present in the collections of the Institut Fran£ais d'Afrique Noire. We may conclude, therefore, that C. caretta is rare on the west coast of Africa. Chelonibia manati Gruvel Chelonibia manati Gruvel, 1903, 116, pi. 2, figs. 14, 17, 18; pi. 4, figs. 15, 16. Chelonibia manati: Stubbings, 1965, 894, text-figs. 5, 6. Typical C. manati had not been reported since the original description by Gruvel based on material brought from the Congo by Dybowski in 1896 until redescribed by the present author from Senegalese material (Stubbings, 1965). Pilsbry (1916) erected two subspecies lobatibasis and crenatibasis for material that he assigned only tentatively to C. manati. Insofar as could be determined these variant specimens probably came from turtles. The greater resistance to penetration of the turtle shell as compared to the skin of the manatee would account for the relatively intact basal margin of the barnacle shell compartments. This has already been shown to happen on the manatee when adjacent individuals overlap (Stubbings, 1965). It is possible, therefore, that C. manati occurs on turtles as well as on the type-host but specimens would then be atypical in having a relatively intact basal margin to the compartments. On the other hand a specimen growing by chance on a soft area of the turtle shell or skin would develop branching parietal ribs approximating to the normal type. This is probably what happened in the case of the incompletely known C. ramosa described by Korschelt (1933), which may well be a growth form of C. manati. Distribution : embedded in the skin of the manatee Trichechus senegalensis in West African estuaries. It may occur also on turtles in W. African waters and if so, owing to the possibility of distribution by currents and wind, it is liable to occur rarely over a much wider range. Chelonibia patula (Ranzani) LOCALITIES: Senegal, off Goree, 99 m. on anchor, coll. Capt. Moloney, B.M. 85.5 (2 shells without soft parts). Gambia: (i) off Langiang Point on Callinectes ? marginatus with B. venustus, 10.6.51, coll. M. H. Routh, B.M. 1952.10.2.8 (3 large and 14 smaller spec.); (2) River Gambia, coll. Capt. Moloney, B.M. 01.4 (3 dry shells); (3) off Gunjur, with B. t. tintinnabulum from hull of shark-fishing boat, 3.3.52 coll. M. H. Routh, B.M. 1952.10.2.3 (2 spec). 2Q8 H. G. STUBBINGS Sierra Leone: (i) 30 m., trawl, from carapace of Neptunus validus, 20.9.54, coll. A. R. Longhurst, B.M. 1957.6.3.6 (3 + 7 juv. spec.); (2) 30 m., trawl, from carapace of Neptunus validus, 28.9.54, coll. A. R. Longhurst, B.M. 1956.1.7.12 (4 + 3 attached juv. spec.). Ghana: off Accra: (i) 14 m., with B. venustus on shells, 27.4.51 (9 + 8 juv. spec.); (2) 14 m., with B. venustus on Cymbium shell occupied by a Pagurid, 30.4.51 (3 + 10 juv. spec.); (3) 15 m., on empty Cymbium shell, n.d., (i spec.); (4) on carapace of Calappa rubroguttata Herklots, 9.4.51 (2 spec.); (5) 10 m., with B. venustus on Cymbium fragment, 9.1.52 (2 -f- 10 juv. spec.); (6) off Chorkor, Accra, 13 m., on C. rubroguttata, n.d., (8+4 juv. spec.); (7) trawl, on Cymbium porcinum with B. venustus, coll. E. Salzen, B.M. 1957.6.3.1 (9 -j- 21 juv. spec.). Nigeria : (i) Lagos, attached to cork thrown up on beach (dead shells only) (H.G.S.) ; (2) Bonny River, E. Nigeria, from No. 2 fairway buoy about 3 ft. below water line (13 spec.). The scutum and tergum have been inadequately figured previously and those of a specimen from Bonny are shown in Text-fig. 23. The scutum (Text-fig. 236, d) shows no trace of longitudinal furrows but growth lines are deeply sculptured. On the tergum (Text-fig. 230, c) the successive growth increments are demarcated by furrows, the incremental areas being convex in section and ornamented by fine grooves. The mouthparts, likewise inadequately figured, agree with Darwin's description. The FIG. 23. Chelonibia patula (Ranzani). Scuta and terga of a specimen from the Bonny River, Nigeria : a, b, external surface of left tergum and scutum, c, d, internal surface of right tergum and scutum, (all X 14). THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 299 labrum (Text-fig. 240) is deeply notched, and armed on each side with 20-24 promi- nent teeth diminishing in size from the midline outwards. The palp (Text-fig. 246) is club-shaped slightly concave on the anterior margin and only sparsely clothed with setae at the distal extremity. The mandibles are 5-toothed. On the specimen dissected (Text-fig. 240) only the 2nd and 3rd teeth bear subsidiary points and not the second and fifth as described by Darwin. The lower angle bears short stout spinules. The first maxilla (Text-fig. 24^) is not notched. There are two stout upper spines and a row of lesser spines on the edge. The lower angle is setose. The second maxilla (Text-fig. 240) bears a small oval secondary lobe. Cirri I and II are short and stout with densely setose and moderately protuberant segments. Cirrus III is intermediate between the first two and cirri IV-VI which FIG. 24. Chelonibia patula (Ranzani) : mouth parts of the same specimen as in Fig. 23 a, labrum, b, palp (both x 50) ; c, mandible ; d, maxilla I (both x 80) ; e, maxilla II 300 H. G. STUBBINGS are similar. The inward direction of the setae on cirri I-III which lie more on the inner face of the segments and not on their anterior faces is marked. The segmenta- tion of the cirri in the specimen dissected is as follows : TABLE 10 I II III IV V VI 16-15 14-14 20-21 32-33 41-35 41-39 TABLE 10. Segmentation of the cirri of a specimen of C. patula from the Bonny River, E. Nigeria. C. patula is known from French Guinea, Sierra Leone, Ivory Coast, Ghana and Nigeria and presumably therefore occurs throughout subtropical and tropical West Africa. It is commonly found on the larger crabs and Gastropoda, notably old Cymbium shells occupied by Pagurids (Gauld, 1957 ; personal observations) and on artifacts such as navigation buoys. It is a sub-littoral shallow-water species, only occasionally cast ashore on floating or sunken debris. Beyond the west coast of Africa it is known from the Mediterranean and from most tropical and subtropical regions. Coronula complanata (Morch) The authority for including C. complanata in the West African fauna would appear to be Pilsbry (1916) who records the presence of some old material labelled West African without name of collector in the U.S. National Museum. But from the tone of the passage the author did not place much reliance on this evidence of provenance. There are no other records for the coast, the nearest being from Table Bay and Simonstown (Barnard, 1924) and the Cape of Good Hope (Nilsson-Cantell, 19320). In the absence of commercial whaling off the coast of West Africa and the very slender chance of a stranded whale with this barnacle being recorded on that coast it is unlikely that confirmatory material will ever be forthcoming. Platylepas hexastylos (O. Fabr.) P. hexastylos is not represented in the Ghanaian and Nigerian collections. Records from West Africa are surprisingly few, doubtless due to the unpredictable occurrence and chance collection of the host animals. Specimens are recorded from the Gambia (Darwin, 1854), Mauretania and Morocco (Broch, 19246, 1927 respectively), Sierra Leone and Senegal (Stubbings, 1965). There are more records from the Mediter- ranean, from turtle hosts. Elsewhere, P. hexastylos is known from the tropical and sub-tropical west Atlantic, from S.E. Asia and from Eastern Australia. Stomatolepas elegans (O. G. Costa) Stomatolepas elegans: Pilsbry, 1916, 289, pi. 68, figs. 2, 2«. Stomatolepas elegans: Hiro, 1936, 312, text-figs. 1-5. The West African status of this species is based on a single specimen in the collections of the Institut Fran9ais d'Afrique Noire from Caretta caretta (Stubbings, THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 301 1965) on which host it was associated with Chelonibia testudinaria and Platylepas hexastylos. Hiro (1936) considered that there was only one species, Stomatolepas elegans (Costa) and that 5. praegustator Pilsbry (1910) and 5. transversa Nilsson-Cantell (19306) were synonymous. The scattered localities recorded by these authors in- dicate that Stomatolepas is widespread in tropical and sub-tropical waters. On the basis of known records it is nowhere common, though this may be a result of the inconspicuous character of the species when embedded in the rough skin of turtles. Xenobalanus globicipitis Steenstrup X. globicipitis is of world- wide distribution as might be expected of a species asso- ciated with Cetacea. Records for West Africa come solely from the collections of the Institut Fran£ais d'Afrique Noire and are all from Senegal (Stubbings, 1965). The nearest other records are those made by the Prince of Monaco in the Atlantic off the mouth of the Mediterranean and in the Azores (Gruvel, 1920). The cetacean hosts of X. globicipitis were listed by Nilsson-Cantell (1930^). The IFAN specimens were found on Delphinus delphis, Phocoena phocoena and Feresa attenuata, all except the first being new hosts for this barnacle. ACROTHORACICA No species of Acrothoracica were known from West Africa prior to 1957. In that year Gauld reported the occurrence of Kochlorine hamata in old shells from the Ghana coast. Subsequently, with increased interest in the Cirriped fauna of West Africa, a further three new species have been described and it would seem highly probable that yet others will be found eventually. CRYPTOPHIALIDAE CRYPTOPHIALUS Darwin 1854 Cryptophialus coronatus Tomlinson Cryptophialus coronatus Tomlinson, 1960, 404, figs. 1-3. This species was found in the shell of Haliotis tuberculata L. from Goree Island, Senegal. It has not yet been recorded from any other locality. Cryptophialus variabilis Stubbings Cryptophialus variabilis Stubbings, 19616, 189, fig. 6. This species was found in molluscan and Balanus tintinnabulum shell rubble from the Island of Principe, Gulf of Guinea. KOCHLORINIDAE Kochlorine hamata Noll LOCALITIES: Ghana: (1-5) Accra: Sta. 10 (5 spec.); Sta. 81 (5 spec.); Sta. 106 (210 spec.); Sta. 107 (266 and 25 spec.); Sta. unknown (249 spec.); (6) Prince's 302 H. G. STUBBINGS Town shore (c. 12 spec.) ; (7) 2 miles west of River Densu, 7-3 m., in shell encrusted with the Polyzoan Membranipora arborescens (Canu & Bassler) (3 spec.). Gulf of Guinea off Annobon : "Calypso" Sta. 107, 1° 26' 15" S. 5° 35' 40" E., 60 m. in nodule of shell material, 4.7.56 (2, possibly more spec.). All the Ghanaian specimens except those from Prince's Town and the R. Densu locality were in shells of Murex bourgeoisi Tournouer. The number of specimens infesting a shell may be very high, each of the figures (210), (266) and (249) above representing the total number of Kochlorine recovered from two infested Gastropod shells, broken up and digested with acid. There may therefore be appreciably more than 100 specimens of K. hamata in an infested Murex bourgeoisi shell. Living Murex, shells occupied by hermit crabs and empty shells are colonized. Murex appears to be infested more readily than other gastropod shells. The Prince's Town specimens were recovered from an unidentified fragment of a large thick- walled gastropod shell,, bored also by the mollusc Lithophaga aristata (Solander). LITHOGLYPTIDAE Kochlorine inermis Stubbings Kochlorine inermis Stubbings, 19646, 343, fig. 5. This species was described from a single specimen found in the tergum of a small B. tintinnabulum taken at Malembe, Angola. As yet there has been no further record of this species. Kochlorinopsis discoporellae gen. et sp. n. LOCALITIES: Senegal, coll. I. Marche-Marchad (i) South of Goree I., 38-42 m. in Cupuladria canariensis (Busk), 27.10.53: (2) the same, 40-41 m., in C. canariensis, 24 . 2 . 53 ; (3) South of Cape Verde Peninsula, 95 m., in C. canariensis and Discoporella umbellata (Defrance), 18.2.54; (4) Baie de Seminoles, Goree, 38m., in Cupuladria multispinata (Canu & Bassler) and D. umbellata, 8.12.53; (5) SW of Madeleines Light, 48 m., in D. umbellata, 15.9.53; (6) " Alignment Goree-Cap Manuel, lantern de Madeleine, 35-42 m., in D. umbellata, 19.10.56; (7) 20-25 miles off Saloum, 35-37 m., in D. umbellata, 8.3.55. Guinea : Parages des Fles de Los, Conakry, 19 m., in C. multispinata and Cupuladria owenii (Gray), 21.1.53, coll. I. Marche-Marchad; (2) "Calypso" Gulf of Guinea Cruise, Sta. 7, 9° 40' N., 13° 53' W., 18 m., in C. owenii; (3) Atlantide Sta. 146, 9° 27' N., 14° 48' W., 51 m., in C. canariensis and D. umbellata. Ivory Coast: " Calypso " Gulf of Guinea Cruise Sta. 17, 5° N., 5° 28' 30" W., 27 m., in C. canariensis and D. umbellata. Ghana: Accra (i) Sta. 132, 44 m., in D. umbellata, 2.5.51; (2) Sta. 133, 51 m. in D. umbellata, 2.5.51 (9 spec.). Gabon: (i) " Calypso " Gulf of Guinea Cruise Sta. 45, o° 25' N., 9° E., 73 m., in Discoporella reussiana (Manzoni); (2) " Discovery " Sta. 279 off Cape Lopez, 58-67 m., in C. canariensis, 10.7.27. The original specimens from which this new species is described, and the type selected, came from Mr. Bassindale's Sta. 133 off Accra, Ghana, (Bassindale, 1961, text-fig. 2). This is therefore the type locality. Miss P. L. Cook in her study of THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 303 West African Cupuladriid Polyzoa (1965(2) found Acrothoracica in a number of species from several localities. Through her courtesy in passing these to me for study I am able to give the other records listed above. I have to thank Dr. Torben Wolff of the University Zoological Museum, Copenhagen for the use of material from " Atlantide " Sta. 146 off Guinea, which also was recognised by Miss Cook in her examination of the " Atlantide " Cupuladriidae. Diagnosis: Mantle extremely flattened. No flattened attachment "pad" but the mantle adjacent to the dorsal lip of the orifice is thickened somewhat and protuberant. Mantle with longitudinal and transverse muscles. Orifice slit-like with inner lip armed with bifid or more complex spines. Mouth cirri well developed on long two jointed pedicels with rami of five and four segments. Three pairs of terminal cirri. Caudal appendages present, two-segmented. TYPES: Holotype BM 1966.12.5.1. paratypes BM 1966.12.5.2-8 from Accra Sta. 133. The appearance of the orifices of the cavities inhabited by K. discoporellae in a zoarium of D. umbellata is shown in PI. la. The location of the cirripedes within the thickness of the zoarium is revealed by decalcification (PL 16). The mantle is oval, light brown (in spirit) about 3 mm. long by 2 mm. broad and extremely flattened laterally so that the whole animal is somewhat transparent viewed by transmitted light. The orifice is elongated, slit-like and not on a pro- longation of the mantle. The margins of the orifice are bordered by setae and short stout" stellate " spines with two or three or more recurved teeth (Text-fig. 250, 260). Towards the ventral side the lips of the orifice are prolonged into short triangular processes. Beyond these the margins carry a row of fine stout spines forming a " comb " collar. The ventral margin of the mantle is crenated. It bears four or five recurved hook-like spines reducing in size posteriorly where several of the typical stellate spines finish the row. In young specimens the antennules can be seen lying close against the dorsal side of the mantle (pi. ic). The mantle has longitudinal muscle bands running from a little below the orifice toward the opposite end of the mantle parallelling the ventral margin. There are no such muscles over the dorsal half of the mantle which is thin and transparent. The muscles are attached to a thickened band of the mantle delimiting the non- muscular area. The circular muscle system is less developed. Fine muscle bands attached to the thickened attachment process radiate towards the ventral margin of the mantle. In the posterior part of the mantle these muscles run from the thickened band to the ventral margin. Retractor muscles from the orifice run to the body and to the thickened attachment area. These correspond to the retractor corporis and retractor orificii muscles of Utinomi (1950). There is no " retractor palii rostralis " (Utinomi 1963). Some of the anatomy as elucidated from a whole mount is shown in Text-fig. 256. There is a single pair of lobe-like processes on the ventral side of the body, probably comparable to the two pairs described by Utinomi (1950) in Berndtia purpurea. Mouth cirri, three pairs of terminal cirri and caudal appendages are present. The mouth cirri (Text-fig. 260) have a long two-segmented pedicel, the first segment being greatly elongated. The rami are composed of 5 and 4 segments respectively. A mouth cirrus of another specimen had rami of 7 and 5 304 H. G. STUBBINGS r.c r.o FIG. 25. Kochlorinopsis discoporellae gen. et sp. n., a, whole animal to show mantle muscula- ture, ovary and egg mass. The broken line within the mantle indicates the ventral surface of the body : the cirri have been omitted ( x 25) ; b, animal with mantle removed ( x 38). Setae omitted from the cirri. C.2 mouth cirrus ; i.g. infra-oesophageal ganglion ; r.c. retractor corporis muscle ; r.o. retractor orificii muscle ; s.g. supra-oesophageal ganglion. segments. The terminal cirri are biramous and multi-articulated. The first pair is slightly shorter than the succeeding two pairs. The segments are about twice as long as broad proximally, increasing to about four times as long as broad distally. Each segment bears two pairs of setae anteriorly, the upper pair long and stout, the lower short and fine. Posteriorly each bears a single seta or pair of setae. The caudal appendages are short and two-segmented. The terminal segment bears three short setae. The labrum is strongly folded so as to embrace laterally the mouth appendages. Its margin is entire and convex and without teeth or hairs. There is a slight sugges- tion of serration of the margin close to the middle line but there are no teeth. Behind the margin two single lines of long setae run backward parallel to the midline. In a flattened preparation these point towards the middle line but in life they stand more or less upright. There are a few shorter hairs scattered between these lines. The palps (Text-fig. 266) are very small and conical with a long base of attachment. THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 305 They bear a few setae on the outer margin and three or four longer ones at the tip. The mandible (Text-fig. 260) has three simple teeth and a dentate lower angle with about 6 fine teeth. The lower margin has a short row of fine setae. The maxilla (Text-fig. 26d) has a long straight upper margin and short lower mar- gin, the articulation thus being rather long. The anterior margin has a very marked broad notch bearing two stout but short spines. There are two large upper spines strongly curved towards the notch. At their base laterally is a short stout spine similarly curved. The lower and larger part of the anterior edge bears two pairs of short stout spines separated by a gap with short setae. The lower angle is receding and carries a few short spines grading into a few slender setae on the lower margin. There are a few small spines on the surface of the appendages. The second maxilla is typical of the Acrothoracica, tapering to a point distally and with a very thick outer margin so that the appendage is wedge-shape. The inner margin is concave distally and convex proximally giving the impression of a greatly enlarged lower lobe here. The apical portion is thickly covered with long setae. DWARF MALE : dwarf males are found in association with most females. One is FIG. 26. Kochlorinopsis discoporellae gen. et. sp. n., a, mantle edge showing spines and comb collar (X93) ; b, palp (xiyo) ; c, mandible ; d, maxilla I (both X38o) ; e, mouth cirrus (xiyo); /, dwarf male (xi6o). 306 H. G. STUBBINGS shown in Text-fig. 26/. It is triangular in side view, flattened and reminiscent in form of that of K. hamata Noll though not so markedly concave on the side opposite the antennae. One two- jointed antenna was present on the specimen. The posterior end is slightly drawn out into a blunt process, presumably the penis sheath. This end of the animal is deeply staining and appears to contain a muscular tube, the penis in the contracted state. The internal anatomy is only partially visible. The penis appears to swell out and fill the posterior part of the organism. There appears to be a wide tubular structure lying near the surface close to the margin between penis and antenna, probably the vesicula seminalis. A number of deeply staining irregular or rounded bodies in the opposite half of the animal are presumably lobes of the testis. The dimensions of this specimen were length 0-33 mm. depth 0-20 mm. In having well-developed mouth cirri, an attachment disc, short labrum and no lateral bar this species resembles both Lithoglyptes and Kochlorine. Lithoglyptes has four pairs of terminal cirri as against three pairs in Kochlorine. On this basis the present species should be referred to Kochlorine. The absence of the long retrac- tor muscle along the rostral side of the mantle (" retractor pallii rostralis " of Utinomi, 1963) is against this. The genera Kochlorine and Balanodytes both have this prominent paired muscle band which Utinomi (1963) and Tomlinson (1963) consider as diagnostic of the family Kochlorinidae. Utinomi considers (1963 : 70) that the number of pairs of cirri and the presence or absence of caudal appendages should be regarded as of generic value only and that the musculature is a better criterion of family relationships. On this basis K. discoporellae should be assigned to the Litho- glyptidae (Aurivillius, 1894 ; emend. Tomlinson & Newman, 1960) as " retractor corporis " and " retractor orificii " muscles are present. This procedure demands, however, a further amendment of the family diagnosis to include forms with only three pairs of terminal cirri. Furthermore inclusion of this species in Lithoglyptes would entail redefining this genus. Hence it is proposed to erect a new genus within the Lithoglyptidae to contain species with only three pairs of terminal cirri. The species is accordingly named Kochlorinopsis discoporellae g. et sp. n., recording its cirral affinity with Kochlorine and its association with the Polyzoan Discoporella umbellata. The diagnoses of family and genus follow : Family LITHOGLYPTIDAE emend. Mouth cirri well-developed, the pedical two-jointed. Three to five pairs of terminal cirri, if less than five pairs, then caudal appendages present. No gut teeth or gizzard. No lateral bar. Attachment disc present, retractor orificii and r. corporis muscles present. Genus KOCHLORINOPSIS nov. Lithoglyptidae with three pairs of terminal cirri : caudal appendages present, two- jointed : mouth cirri with number of segments in rami varying from five and four to seven and five. TYPE, K. discoporellae nov. THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 307 3. THE WEST AFRICAN CIRRIPED FAUNA From geographical plots of the distribution of the various species it is possible to distinguish several groups of species in which the individual members have approxi- mately the same limited geographical range. Other species are apparently cosmo- politan. A considerable number are still insufficiently known for their geographical range to be determined. The following groups of species can be distinguished : (1) Northern species (Text-fig. 27A) Species with a wide range in latitude extending from high latitudes in the northern hemisphere into sub-tropical and tropical waters. Two species only have this distribution, Scalpellum scalpellum (L.) and Chthamalus stellatus stellatus (Poli). S. scalpellum ranges south to the Congo and Angola. The main distribution of C. stellatus appears to end in the latitude of Dakar (Cape Verde) though small numbers have been recorded from as far south as Angola. It is probable, however, that the records of this species from south of the Bight of Biafra, as well as those from the Eastern Mediterranean are erroneous and due to misidentifi cations. (2) Western English Channel to the Gulf of Guinea (Text-fig. 27B-D) In this series are included species with a northern limit of distribution in the south-west English Channel, on the Atlantic coast of Europe or in the Western Mediterranean and which extend southwards through the Gulf of Guinea. In this category, in order of diminishing northward distribution fall : Pyrgoma anglicum Sowerby Balanus perforatus Bruguiere B. spongicola Brown B. tulipiformis Darwin B. fallax Broch Much of the geographical range of P. anglicum coincides with that of B. perforatus, the major differences being the extension of the range of B. perforatus into the Eastern Mediterranean and south of the Gulf of Guinea and the presence of P. anglicum in the Cape Verde Archipelago where B. perforatus is hitherto unknown. (3) South-west Europe to Cape Verde (Text-figs. 270, 28A) Species distributed from the Atlantic coast of Europe or the Western Mediterranean to the region of Cape Verde only. Mitella pollicipes (Gmelin) Heteralepas cornuta (Darwin) Paralepas minuta (Philippi) H. cornuta is still further restricted in distribution in that it is known only from north-west Africa, from Morocco to the Cape Verde Islands. ZOOL. 15, 6. 25 H. G. STUBBINGS C D FIG. 27. Distribution of some West African barnacles. A. The northern components Sc. scalpellum (L.) and C. stellatus (Poli). The far northern distribution of Sc. scalpellum is omitted. The eastern Mediterranean distribution and that south of the Gulf of Guinea for C. stellatus may be erroneous due to mis-identifications. B, C, D, South-west Europe, Western Mediterranean, North-west and West African species. B. perforatus extends also to the Eastern Mediterranean. THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 309 (4) Endemic West African species (Text-fig. 28s, c) There are only five species that can be regarded as possibly endemic in tropical West Africa : Smilium renei (Gmelin) Ibla atlantica sp. n, Chelonibia manati Gravel Balanus pallidus Darwin Chthamalus aestuarii Stubbings 5. renei, C. manati and B. pallidus extend northwards to Dakar ; C. aestuarii is not known as yet north of Sierra Leone and Ibla atlantica only from off Sierra Leone. The known southern limit of all except the Ibla is northern Angola. B. pallidus is an abundant intertidal species in West Africa. There are a few records from other parts of the old world (" P " in Text-fig. 28c), but these could be instances of ship- borne introductions or even misidentifications. Darwin (1854) mentioned seeing specimens of B. pallidus " from the West Indies ". Specimens from Cananeia, S. Paulo, Brasil received through the kindness of Dr. V. Zullo are of this species, which may, therefore, occur naturally on both sides of the tropical Atlantic. It is possible that B. tintinnabulum zebra is a West African variety but here again it may well be that the pattern of its natural distribution, which is very imperfectly known, is blurred by records of ship-borne specimens. (5) South African species (Text-fig. 280) Only two species can be considered as South African with an extended distribution northward in the Atlantic namely Chthamalus dentatus and Balanus maxillaris but the evidence is not clear cut. C. dentatus is a common species at the Cape and on exposed coasts in West Africa as far as Dakar. It is recorded from several widely spread Indian Ocean localities so should perhaps be regarded as an Indian Ocean form that has penetrated to the Atlantic. Until records for the Indian Ocean are much more numerous, however, its true status is debatable. It is questionable whether B. maxillaris is native to West Africa. It is well- known at the Cape but the sole record north of Angola is for Port Etienne, Maure- tania, which suggests that this may be a chance introduction. Further records from Gough Island and between South America and the Falkland Islands in the South Atlantic (Nilsson-Cantell, 19390) suggest that B. maxillaris may be really a southern temperate species widespread in those latitudes as suggested by Kolosvary (i943&). (6) Circumtropical species Several species are known from the tropics of both the old and new world : Trilasmis (Poecilasma] kaempferi (Darwin) Octolasmis lowei (Darwin) B. amphitrite amphitrite Darwin B. calceolus Darwin (old world tropics only) B. tintinnabulum tintinnabulum (L.) B. trigonus Darwin B. venustus Darwin 3io H. G. STUBBINGS FIG. 28. Distributions of African species : A, temperate-subtropical Western Mediter- ranean and N.W. African : B, C, West African-Gulf of Guinea tropical species. Three isolated reported occurrences of B. pallidus in North-west Africa, the Red Sea and Gulf of Aden are ringed and marked with a " P " : D, the temperate South- African element — C. dentatus Krauss extending throughout tropical West Africa and the southern B. maxillaris with a single record from the Cap Blanc region. THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 311 The inclusion of 0. lowei in this group is dependent upon the interpretation of the synonymy of this species. If the Western Atlantic and Pacific specimens are con- specific with those of the Eastern Atlantic then the species is certainly circum- tropical. B. venustus Darwin is present on the whole West and South African coast from Port Etienne, Mauritania to Natal, with an outlying northern record from Cadiz Bay, southern Spain. It is particularly abundant off Ghana and this might well be the centre of distribution of the species. The picture is complicated, however, by an isolated series of records in the Bay of Bengal from Ceylon, to Akyab in Burma. There are no East African or Arabian Sea records linking these two areas. (7) Deepwater elements The few species so far collected from beyond the continental shelf indicate the presence of a deep-water barnacle fauna here as elsewhere. The following five species are the only ones so far recorded : Smilium longirostrum (Gruvel) Scalpellum imperfect/urn Pilsbry 5. trapezoideum Hoek 5. velutinum Hoek Verruca striata Gruvel By comparison with the known fauna of the Azores region it may be expected that many others await discovery. (8) Introduced species Several species recorded sporadically are well-known to have spread from their original homes by the unwitting agency of man. Others may have spread in the same way but have not been detected. The following species, found sporadically in West Africa, may have this doubtful status. Octolasmis hoeki (Stebbing) B. venustus niveus Darwin Chthamalus fragilis Darwin B. nigrescens Lamarck C. stellatus bisinuatus Pilsbry B. tintinndbulum concinnus Darwin Balanus eburneus Gould The first five have representatives in North and Central American waters, the two Balani most probably having been spread from there by man. B. nigrescens was probably introduced from the western Pacific. B. tintinndbulum may also have been introduced on ships from that ocean. Two further varieties, B. t. azoricus Pilsbry and B. t. spinosus (Gmelin) may also be incomers, the first from the Azores presumably being carried by ships. B. t. spinosus, as suggested elsewhere (Stubbings, 19616) is perhaps another island race that occasionally reaches the African mainland. There is left a number of species recorded from West Africa for the distribution of which there is wholly inadequate knowledge. The very few records from African waters (in many instances only a single record) are inadequate to establish these as 312 H. G. STUBBINGS W. African species. The following all require confirmation from new finds : Trilasmis (Poecilasma] crassum (J. E. Gray) B. amphitrite albicostatus Pilsbry Octolasmis tridens (Aurivillius) Tetradita divisa Nilsson-Cantell Chthamalus stellatus bisinuatus Pilsbry T. purpurascens Wood Pachylasma gigantea (Philippi) T. squamosa squamosa (Brag.) Balanus tintinnabulum maroccana Broch In considering the above groupings of species the pelagic species of Lepas and Conchoderma and those species found on turtles or cetaceans have been excluded. The wide-ranging habits of the host animals or the drift of floating debris used as a support render specific zoogeography of these barnacles valueless. The one exception has been Chelonibia manati ; the estuarine habitat and sluggish habits of the sirenian preclude long migrations. After removal of these nine species, the deep water species, the introduced species and those with imperfectly known distribution, some 24 species are left for which the zoogeographical information is more or less adequate. Three of these species, T. kaempferi, B. t. tintinnabulum and B. trigonus are circumtropical. Two, B. v. venustus and B. calceolus and possibly a third B. a. amphitrite are old world tropical species but have a restricted Atlantic distribution. They have been placed in Table ii with species of similar distribution. Where a species is apparently at the limit TABLE n W. N.W. W. S.W. S. Species S.W. Europe Med. Afr. Afr. Afr. Afr. Sc. scalpellum -f + + -f- -f B. perforatus . . + + + -f- (+) — Ch. stellatus + + + (+) — — B. tulipiformis + + + -j- B. spongicola + -f- + -f- — — P. anglicum. + + + + — M. pollicipes . . + + -f- — — — P. minuta ... -f- + (+) — — B. a. amphitrite . . + + + — — B. calceolus ... + + + — — B.fallax ... + + + (+) B. v. venustus . . ( + )( + ) + + H. cornuta ... + (+) — — O. lowei ... + -f — — B. pallidus . ( -f ) + — — Ibla atlantica . . — -f- — — 5. renei ... — + — — Ch. aestuarii . . — — -{- — — C. manati ... — — -f — — Ch. dentatus . . -f -f- -f B. maxillaris . . (-f-) — _)_ _|_ Totals ... 7 12 16 19 5 3 % ... 37 63 84 100 26 16 TABLE n. The northward and southward Atlantic distribution of 21 tropical West African barnacles. THE CIRRIPED FAUNA OF TROPICAL WEST AFRICA 313 of its range in an area the " plus " symbol for presence is placed in brackets thus ( +). The northern affinities of the West African cirriped fauna are obvious. Of the remaining 19 species occurring between Dakar and Lobito, Angola, and listed in the table below, 16 (84%) occur to the north of Dakar and 12 of these (63 %) extend into the western Mediterranean. Only one species, B. perforatus, is known reliably from the Eastern Mediterranean. Although a second, C. stellatus, is recorded from there this may well be an error of identification. Seven species (37%) extend into S.W. Europe, with a few reaching the English Channel. On the other hand only 5 (26 %) are known from S.W. Africa and 3 (16 %) from the Cape. ACKNOWLEDGMENTS In completion of this final and at the present time comprehensive report on the barnacles of the West Coast of Africa it is a pleasure to repeat my thanks to those Directors of Museums or Institutions who have entrusted to me the cirripede collec- tions in their charge : also to the many individuals who have assisted me, with specimens, identifications and otherwise. My especial thanks are due to the Director of the British Museum (Natural History) for continued access to the collections and library of the Museum and for agreeing to publish this present work. To the Keeper of Zoology of the Museum and members of his staff I tender my thanks for many kindnesses in identifying, or checking my identifications of, the hosts of many epizoic species, and otherwise. To Dr. J. P. Harding I owe personal thanks for permission to use his photographs of K. discoporellae reproduced in PI. i. SUMMARY 1. The distribution of seventy species and varieties of barnacles, including five species of Acrothoracica in West Africa is recorded. 2. Two new species are described. Ibla atlantica sp. n. from the continental shelf off Sierra Leone represents the first record of this genus in the Atlantic. A new Acrothoracican, Kochlorinopsis discoporellae gen. et sp. n. has characters between Kochlorine and the Lithoglyptidae. It is referred to the latter family, for which a redefinition is given. 3. The type material of Dichelaspis hoeki Stebbing and D. antiguae Stebbing has been re-examined and compared with new material. The union of the two species as D. hoeki by Annandale (1910), followed by Nilsson-Cantell (1927) (as 0. hoeki (Stebbing)) is rejected and D. antiguae is reinstated as a species, 0. antiguae (Stebbing) . It is not, as far as is known, found in the eastern Atlantic. 4. The forms Balanus amphitrite communis, B. a. hawaiiensis and B. a. denticulata as found in W. Africa, have been re-examined in the light of Harding's (1962) redescription of Darwin's type material. No constant differences could be detected on which to separate the varieties and they are united as B. amphitrite amphitrite Darwin. 5. Harding (1962) could not find constant differences for separating the two " amphitrite " varieties called by him B. pattidus pattidus Darwin and B. pattidus stutsburi Darwin. The latter name is reduced to a synonym and both white and coloured shells are referred to the species B. pattidus Darwin. 3H H. G. STUBBINGS 6. Amplified descriptions have been given of the following : Chthamalus dentatus Krauss, C. aestuarii Stubbings, Balanus perforatus Bruguiere, B. venustus venustus Darwin, B. fallax Broch, Tetradita divisa Nilsson-Cantell and Chelonibia patula (Ranzani) . 7. 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The seasonal occurrence of sedentary marine organisms in Biscayne Bay, Florida. Ecology 29 : 153-172. WELTNER, W. 1887. Die von Dr, Sander 1883-85 gesammelten Cirripedien (Acasta scuticosta sp. n.). Arch. Naturgesch. Jahrg. 53, Bd. i : 98-117, pis. 3, 4. 1897. Verzeichnis der bisher beschriebenen recenten Cirripedienarten. Arch. Naturgesch. Jahrg. 63 Bd. 1 : 227-280. 1922. Cirripedia der Deutschen Tief see-Expedition. Wiss. Ergebn. Deutsche Tiefsee- Exped. 23 (2) : 59-112, pis. 2-4. WILLIAMS, G. 1954. Fauna of Strangford Lough and neighbouring coasts. Proc. R. Irish A cad. 56B : 29-133, pis. 2-4. WOOD, E. J. F. & ALLEN, F. E. 1958. Common marine fouling organisms of Australian waters. Dept. of the Navy, Melbourne, Australia. ZEVINA, G. B. & TARASOV, N. I. 1954. [Acorn barnacles new to Soviet Waters of the Black Sea.] Trudy Sevastopol', biol. Sta. 8 : 341-346. 1963. [Cirripedia Thoracica of the mainland coasts of south-east Asia.] Trudy Inst. Okeanol. 70 : 76-100. PLATE i Kochlorinopsis discoporellae g. et sp. n. a. Frontal view of a specimen of a host Polyzoan Discoporella umbellata (Defrance) with a number of embedded K. discoporellae X4- b. Basal view of the same after decalcifying with diluted hydrochloric acid so that the embedded cirripedes of various sizes can be seen X 4. c. A recently settled specimen of K. discoporellae photographed by transmitted light showing the paired antennules lying against the dorsal side of the mantle X54- Bull. Br. Mus. nat. Hist. (Zool.) 15, 6 PLATE i PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING POLYZOA (BRYOZOA) FROM WEST AFRICA THE PSEUDOSTEGA, THE CRIBRIMORPHA AND SOME ASCOPHORA IMPERFECTA i?f>7 r* PATRICIA L. COOK BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 15 No. 7 LONDON: 1967 POLYZOA (BRYOZOA) FROM WEST AFRICA THE PSEUDOSTEGA, THE CRIBRIMORPHA AND SOME ASCOPHORA IMPERFECTA BY PATRICIA L. COOK Pp. 321-351 ; 2 Plates, 14 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 15 No. 7 LONDON: 1967 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the 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. 15, No. 7 of the Zoological 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.). Trustees of the British Museum (Natural History) 1967 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 14 November 1967 Price Thirteen Shillings POLYZOA (BRYOZOA) FROM WEST AFRICA THE PSEUDOSTEGA, THE CRIBRIMORPHA AND SOME ASCOPHORA IMPERFECTA By PATRICIA L. COOK CONTENTS Page INTRODUCTION ........... 323 PSEUDOSTEGA LEVINSEN ......... 324 Cellaria Ellis & Solander ........ 324 C. fistulosa auctt. (non Linnaeus) ....... 325 C. sinuosa (Hassall) ......... 325 C. salicornioides "var. normani" Hastings ..... 326 C. elongatoides Bassler. ........ 326 C. atlantida n. sp. ......... 327 CRIBRIMORPHA LANG ......... 328 Membraniporella Smitt . . . . . . . .328 M . nitida (Johnston) ......... 330 M. marcusi n. sp. ......... 331 Cribrilaria Canu & Bassler ........ 333 C. radiata (Moll) .......... 333 ASCOPHORA IMPERFECTA HARMER ....... 334 Tremogasterina Canu ......... 335 T. perplexa n. sp. ......... 336 Exechonella Canu & Bassler . . . . . . . . 337 E. antillea (Osburn) ......... 337 E. gigantea n. sp. ......... 339 Triporula Canu & Bassler ........ 340 T. stellata (Smitt) ......... 343 Hippopetraliella Stach. ........ 345 H. africana n. sp. ......... 345 Metrarabdotos Canu ......... 346 M. unguiculatum Canu & Bassler ....... 346 SUMMARY ........... 348 ACKNOWLEDGMENTS .......... 349 REFERENCES ........... 349 INTRODUCTION THE Collections from which specimens are described here comprise the " Calypso " Collection I, from Senegal and the Bay of Biafra, and Collection II, from the Cape Verde Islands; the Marche-Marchad Collections, from Senegal and the Achimota Collections, from Ghana, see Cook (ig6^a : 44). Some additional specimens from the west African Expeditions of the " Atlantide ", and " Galathea ", and from west African Stations from the Mortensen Java-S. Africa Expedition, all of which are stored at the Universitetets Zoologisk Museum, Copenhagen, are also included. The holotypes of new species, together with representative specimens of all the material ZOOL. 15, 7. 26 324 PATRICIA L. COOK described are deposited in the British Museum (Natural History), unless otherwise stated. Specimens were treated with eau de javel for examination of calcareous parts, and decalcified to show chitinous parts. The following measurements were made where possible : Lz Length of zooid Iz Width of zooid Lor Length of orifice lor Width of orifice Lfo Length of fertile orifice Ifo Width of fertile orifice Lop Length of operculum lop Width of operculum Lov Length of ovicell lov Width of ovicell Le Length of embryo le Width of embryo Lan Length of ancestrula lovor Width of ovicellular orifice Lav Length of avicularium Lm Length of mandible Specimens in the Collection of the British Museum (Natural History) are referred to by their registered numbers, thus: 1964. 3. 20. 15. The Pseudostega are represented by 5 species, 3 of which occur south of the Canary Islands; the number of species of Cribrimorpha is also small, only 3 being represented. The Ascophora Imperfecta includes those genera in which the oper- culum and contiguous frontal membrane are covered by an overarching frontal shield (see Harmer, 1957 : 645). A true compensation-sac is therefore absent. There is evidence that at least 2 distinct types of development of the frontal shield exist, and examples of both have been found in the genera in these Collections. The first type resembles that seen in Tremogasterina (see Powell & Cook 1967), which appears to indicate affinities with some Cribrimorpha (see pp. 337, 342). The second type is that originally described in Umbonula by Harmer (1957) for the Ascophora Imperfecta (see pp. 345, 347). Division PSEUDOSTEGA Levinsen CELLARIA Ellis & Solander Cellaria Ellis & Solander, Harmer, 1923 : 301; Hastings, 1947; Lagaaij, 1952 : 47, Galopim de Carvalho, 1963. Zoarium free, erect, cylindrical, typically jointed, with chitinous rootlets. Opesiae reduced, scarcely larger than the orifice. Avicularia vicarious. Ovicells entotoichal. The characters used to define species in this genus need further study, and a complete revision of the Atlantic and Mediterranean " species ", at least, is urgently required1. Two species of Cellaria were recorded by Calvet (1907 : 400-401) from the north- west African region. These were C. salicornioides (Lamouroux), from north-west Morocco (from 636 m.), and C. fistulosa (Linnaeus) from the Canary Islands (from 3700 m.). Norman (1909) recorded C. fistulosa from Madeira, from " deep water ", 1 A discussion of Cellaria, with descriptions of several species has recently been given by Prenant & Bobin (1966 1372). POLYZOA FROM WEST AFRICA 325 and also listed C. johnsoni (Busk). His specimen labelled with this name is referable to C. salicornioides " var. normani" (see below). C. salicornioides and C. fistulosa were recorded by Canu & Bassler (19286 : 25) from Morocco. C. fistulosa has not been found to occur in these west African Collections, its most southerly limit appears to be the Canary Islands. C. sinuosa (see below) does not appear to occur south of Tangier. Cellaria fistulosa auctt. (non Linnaeus) (Text-fig, i) Cellularia fistulosa (Linnaeus) Norman, 1909 : 292, Madeira. Canu & Bassler, 19286 : 25, Morocco, 28-155 m., sandy bottom. Cellaria fistulosa auctt. (non Linnaeus) Harmer, 1923 : 302. MATERIAL EXAMINED. B. M. Coll. Madeira, 1911. 10. i. 772, 3 worn fragments, Norman Coll. Pedra da Malha, off Porto Santo, 55 fath. 1910. 10. 30. 21, Kirkpatrick Coll. Off Tangier, 1903. 4. 29. roA and 16, 30-40 fath. DIMENSIONS. (African material) . Lz 0-50-0-55 mm., Iz 0-24-0-26 mm., Lor 0-12- 0-14 mm., lor 0-07-0-09 mm. (Ovicelled zooids, av. Lz 0-50 mm., Iz 0-29 mm., Lovor o-io mm., lovor 0-06 mm.) Lav 0-12-0-14 mm., lav 0-14-0-15 mm., Lm 0-04-0-05 mm., 1m 0-08-0-09 mm. Av. L internode, 9 mm. Zoarium with short internodes, chitinous joints tubular. Orifice near the middle of the zooid, with small, paired, proximal denticles. Ovicellular orifices oval. Avicularia with small rounded mandibles directed distally, or slightly distal-laterally. Cellaria fistulosa auctt. is not the same species as that introduced by Linnaeus. I here follow Lagaaij (1952) in using the name C. fistulosa auctt. in the sense res- tricted by Harmer (1923), pending a full revision of the genus. Cellaria sinuosa (Hassall) (Text-fig. 2) Cellaria fistulosa (Linnaeus, non auctt.), Harmer, 1923 : 303. Cellaria sinuosa (Hassall), Lagaaij, 1952 : 48, pi. 4, figs. 4, 5, Pliocene, Europe. MATERIAL EXAMINED. B. M. Coll. Off Tangier, 1903. 4. 29. loB, 30-40 fath. DIMENSIONS. Lz 0-45-0-56 mm., Iz 0-23-0-26 mm., Lor 0-06-0-08 mm., lor o-io- 0-12 mm. (Ovicelled zooids, av. Lz 0-50 mm., Iz 0-24-0-26 mm. Lovor 0-02-0-04, lovor 0-07-0-09), Lav 0-12-0-17 mm., lav 0-16-0-20 mm., Lm 0-07-0-09 mm., 1m 0-13-0-15 mm., Av. L internode, 15 mm. Zoarium robust, with very long internodes, chitinous joints tubular. Orifice in the distal half of the zooid, with a prominent proximal lip. Minute distal and proximal denticles present. Ovicellular orifice transverse, frequently slit-like. Avicularia small, mandibles rounded, directed distally. C. sinuosa has long internodes, and a robust zoarium, like that of C. elongatoides (see below). It does not seem to occur south of Tangier, and, like C. fistulosa, has not been reported from the more southerly parts of the west African coast. 326 PATRICIA L. COOK Cellaria salicornioides " var. normani " Hastings (Text-fig. 3) Cellularia nodosa (d'Orbigny) Norman, 1909 : 293, pi. 42, figs. 4, 5, Madeira (not C. nodosa d'Orbigny, 1851 : 31). Cellularia johnsoni (Busk), Norman, 1909 : 293, Madeira. Cellaria salicornioides var. normani Hastings, 1947 : 221, pi. 2, fig. B. MATERIAL EXAMINED. Marche-Marchad Coll. II. 3oD, Sud de Bale de Goree, i8.ii.54, 95 m. Coll. III. 28 J, dragage 4, " Gerard Freca " i8.ii.54, 97-98 m. Achimota Coll. II, Stn. 117, Trawl 2, 5-iv.5i, 64 m, I3E. Copenhagen Museum. " Atlantide " Coll, Stn 146, 9° 24' N., 14° 48' W., 13 . iv .46, 50-51 m., 45R. British Museum. Holotype of C. nodosa Norman, 1911 . 10 . i . 777, Madeira. Para- type of C. nodosa, 1911. 10. I. 778, Madeira, Norman Coll. As C. nodosa, Porto Santo Is., 65 fath., 1945. 7. 10. 4 and 1910. 10. 30. 18, Kirkpatrick Coll. As C. john- soni, Madeira 30-70 fath., 1911. 10. i. 733, Norman Coll. Gulf of Guinea, 1964. Redier Coll. DIMENSIONS. Lz 0-23-0-40 mm., Iz 0-14-0-21 mm., Lor 0-05-0-06 mm., lor 0-06- 0-09 mm., (Ovicelled zooids av. Lz 0-41 mm., Iz 0-25 mm., lovor 0-07 mm.), Lav 0-46-0-50 mm., lav 0-21-0-23 mm., Lm 0-14-0-16 mm., 1m 0-17-0-19 mm. Av. L internode, 6 mm. Zoarium slender, fertile parts wide and conspicuous, chitinous joints tubular. Branches frequently formed by lateral offsets, rather than by dichotomy. Zooids elongated. Orifice curved proximally, without denticles. Ovicellular orifice cresentic. Avicularia large, rostrum rounded distally, raised. Mandible directed distally. C. salicornioides "var. normani" has been fully discussed by Hastings (1947). Approximately half the branches in these specimens are formed by lateral offsets, the remainder being the result of dichotomous branching. The internodes are all more slender than those of typical C. salicornioides, and the west African material has therefore been referred to the variety, as has Norman's material labelled " C. johnsoni ", see above. It seems doubtful, however, that the form can rank as a subspecies. A large number of colonies, collected by Kirkpatrick from Madeira (Agua de Baixa, off Porto Santo Is., 65 fath., 1910. 10. 30. 18), illustrate the method of growth. They originate on small Gorgonid stems, and grow in profusion, accompanied by Ascidians, worm tubes, Foraminifera, Sertularian Hydroids and small Lamellibranch Molluscs. The Cellaria internodes are themselves encrusted at the base by a Sponge. Cellaria elongatoides Bassler (Text-fig. 4) Cellaria elongata Canu & Bassler, 19286 : 24, pi. 2, figs. 7-9, Morocco, 170 m. (preoccupied by C. elongata Canu, 1908, a south American fossil species). Cellaria elongatoides Bassler, 1936 : 161 (for C. elongata Canu & Bassler, 1928). POLYZOA FROM WEST AFRICA 327 MATERIAL EXAMINED. " Calypso " Coll. I, Stn. i, 21° 05' N., 17° 14' W., io.v.56, 43-45 m., C52A. Marche-Marchad Coll. II. 4A, 8A, I2E, 27 A, Large de Goree, 5-vii.55, 50 m., 7L, Bale de Goree, 50-100 m., 32 J, Sud de presque 1'Ile de Cap Vert, iS.ii. 54, 95 m. Coll. III. 28D, dragage 4, " Gerard Freca ", iS.ii. 54, 97-98 m. Achimota Coll. II. Stn. 117, Trawl 2, 5 .iv.5i, 64 m., I3D. Copenhagen Museum, Grand Canary, 24.111.30, 15-20 fath., 8gM, Mortensen Coll. " Atlantide " Coll., Stn. 60, 5° 06' N., 9° 34' W., 9.1.46. 54A. Stn. 146, 9° 24' N., I4°48'W., I3.iv.46, 50-51 m. 45R. Stn. 161, off Bathurst, Gambia, 24.^.46, 18 m. 58C. Stn. 163, 13° 43' N., 17° 23' W., 25 .iv.46, 65-89 m. 6oC. DIMENSIONS. Lz 0-40-0-45 mm., Iz 0-29-0-32 mm., Lor 0-07-0-09 mm., lor 0-15- 0-18 mm., (Ovicelled zooids av. Lz 0-55 mm., Iz 0-32 mm., lovor o-io-o-ir mm.), Lav 0-09-0-10 mm., lav 0-11-0-12 mm., Lm 0-03-0-04 mm., 1m 0-11-0-12 mm. Av. L internode n mm. Zoarial internodes frequently very long (longest observed 20 mm.), and curved, chitinous joints tubular. Zooids wide, their boundaries well-marked. Orifices with large, paired proximal denticles. Ovicellular orifices with a large proximal denticle. Avicularia small, not common, rostrum sub-triangular, very slightly raised distally. Mandible semi-circular, directed distally. The most striking characters of this species are the large oral denticles and the small avicularia. C. elongatoides is a robust species, and the zoaria can easily be distinguished from the more delicate C. salicornioides " var. normani " by eye. Much of the material consists of isolated internodes, but some complete colonies are present in the specimens from Senegal in the Marche-Marchad Collection. The early stages arise from Hydroid stems, or are attached by rootlets to small conglome- rate masses of sand grains, shell fragments and Foraminifera. Frequently the older, dead parts of the colonies have been colonized by species of Foraminifera. Cellaria appears to be a genus capable of directly colonizing unstable substrates, although it is not exclusively associated with them. C. forceps Lagaaij (1952 : 53, pi. 4, fig. 6), from the Coralline Crag (Pliocene, Britain), resembles C. elongatoides, but has larger avicularia, and a bifid denticle on the proximal side of the ovicellular orifice. Cellaria atlantida n. sp.2 (Text-fig. 5) HOLOTYPE. Zoologisk Museum, Copenhagen, marked slide, " Atlantide " Stn. 123, 57Ept. MATERIAL EXAMINED. Copenhagen Museum. " Atlantide ", Stn. 123, 2° 03' S., 9° 05' E., 5.iii.46, 50 m., 57E, paratypes. DIMENSIONS. Lz o-5o-o-58mm, Iz 0-23-0-27111111, Lor o-oS-o-ogmm, lor o-i-o-i2mm (Ovicelled zooids Lz 0-49-0-53, Iz 0-25-0-29, lov o-og-o-iomm), Lav o-55-o-6omm, lav o-28-o-3omm, Lm o-23-o-27mm, 1m o-i8-o-2omm, Av. L. internode 9 mm. 2 Named after the yacht " Atlantide ", in which the specimens were collected. 328 PATRICIA L. COOK Zoarial internodes occasionally long (longest observed, 18 mm.), but not curved. Chitinous joints tubular. Orifices curved proximally, with paired denticles. Ovicell- ular orifices with a very large proximal denticle. Avicularia as large as the zooids, occasionally larger. Rostrum elongated, rounded distally, not raised. Avicularian cryptocyst extensive. Mandible rounded distally, directed distally. C. atlantida resembles C. salicornioides in its large avicularia, but differs in its denticulate orifice and ovicellular orifice, which are somewhat similar to those of C. elongatoides. Like C. elongatoides , C. atlantida is a robust species, the fertile parts of the internodes reaching i mm. in diameter. C. elongata Canu (1908 : 267, pi. 4, fig. 5, Patagonian, Argentina), resembles C. atlantida, but differs from it in its lack of avicularian cryptocyst and its smaller dimensions (Lz 0-38-0-42 mm., Iz 0-19-0-21 mm., Lor 0-06 mm., lor 0-09 mm.). FIGS. 1-5. Cellaria. Scale = 0-5 mm. i. C. fistulosa auctt. (non Linnaeus). Off Tangier, 1903. 4. 29. i6pt. 2 zooids with ovicells, and i avicularium. 2. C. sinuosa (Hassall). Off Tangier, 1903. 4. 29. loB. 2 zooids, i with an ovicell, and i avicularium. 3. C. salicornioides " var. normani " Hastings. Gulf of Guinea, 1964, Redier Coll. i zooid with ovicell and i avicularium. 4. C. elongatoides Bassler. Marche-Marchad Coll. II. Senegal, 4A. 2 zooids, one with an ovicell, and i avicularium. 5. C. atlantida n. sp. i zooid with an ovicell, and i avicularium. The avicularian cryptocyst appears to become secondarily calcified in some cases, extending almost to the distal end of the rostrum. The mandible is still present. Most of the colonies from Stn. 123 were young, but nearly all had ovicelled zooids. One zoarium arose from rootlets attached to a lamellibranch shell. Division CRIBRIMORPHA Lang MEMBRANIPORELLA Smitt, 1873 Zooids with a frontal shield formed of overarching spines which fuse leaving lacunae between them. Ovicell hyperstomial, closed by the operculum. Membraniporetta is here included in the Cribrimorpha purely for convenience. POLYZOA FROM WEST AFRICA 329 A summary of the discussions relating to its systematic position has recently been given by Bobin & Prenant (1965 : 289), see also Prenant & Bobin (1966 : 568). The range of variation of species assigned to this genus is large. It is possible that the similarities of character of the species included are only superficial; but they are here retained in the genus until work can be done on the early astogeny, and on the development of the spines, ovicells and avicularia. The type-species, M. nitida, possesses diatellae ; all other species listed here have septulae. It is not known at present whether this is a significant character, as, for example, forms with and without diatellae are known in the Ascophoran genus Fenestrulina (see Hastings, 1932 : 427). Avicularia are present in M, nitida and M. agassizii, but they are not found in the other species. The frontal shield of spines may show great inter- and intraspecific variation. In M. nitida (notably in some zooids of the type-specimens), the spines may be few in number, with large lacunae. The appearance of these zooids is then similar to those of M. aragoi s.s., other zooids are almost indistinguishable from those of M. distans. In both these cases, however, specimens of these species may be distinguished from M. nitida by the possession of septulae, and by the form of the ovicell. The following species may be assigned to Membraniporella : — Membranipora agassizii Smitt, 1873 : n. pi. 5, figs. 103-106, Cuba. Erect, avicularia present, spines little branched. Membraniporella antarctica Kluge, 1914 : 677, pi. 33, fig. 7, Antarctic. Encrust- ing, septulae present, avicularia absent, spines little branched. Flustra aragoi Audouin, 1826 : 240, pi. 10, fig. i, ?Red Sea, East Indies, Florida. Encrusting, septulae present, avicularia absent, spines little branched. Membraniporella aragoi var. pacifica Osburn, 1950 : 174, pi. 27, figs. 3, 4, Gulf of California to Galapagos. Avicularia absent, spines unbranched. Membraniporella distans MacGillivray, 1883 : 130, pi. i, figs. 5, 5a, 5b. Port Phillip Heads, Australia. Encrusting, septulae present, avicularia absent, spines little branched. Membraniporella magnifica Thornely, 1912 : 144, pi. 8, fig. 5, Cargados, Indian Ocean. Encrusting, opercula thickened and elongated, septulae present, avicularia absent, spines well-branched. Lepralia nitida Johnston, 1838 : 277, pi. 34, fig. 7, S.W. coasts of Britain and France, Mediterranean, west Africa. Encrusting, diatellae present, avicularia present, spines little branched. Membraniporella petasus Canu & Bassler, 19280 : 36, pi. 4, figs, i, 2, Cuba. Encrusting, avicularia absent, spines branched and fused so that the frontal shield appears porous in the centre. Membraniporella pulchra Osburn, 1950 : 176, pi. 27, figs. 5, 6, west Mexico. Encrusting, avicularia absent, spines numerous, unbranched, central area with irregular lacunae. ZOOL. 15, 7 26§ 330 PATRICIA L. COOK The following species have in the past been assigned to Membraniporella, but are probably referable to other genera :— Cribrilina alcicornis Jullien, 1883 : 12, pi. 14, figs. 23-25. Membraniporella baueri Soule, 1959 : 45, text-fig. 5. Membraniporella corbicula O'Donoghue & O'Donoghue, 1923 : 172, pi. 3, fig. 20. Membraniporella crassicosta Hincks 1888 : 216, pi. 14, fig. 5. Lepralia sceletos Busk, 1858 : 262, pi. 20, fig. 3. Membraniporella nitida (Johnston) Membraniporella nitida var. intermedia Norman, 1909 : 288, pi. 36, fig. 7, Madeira. Membraniporella nitida (Johnston) Bobin & Prenant, 1965 : 277, Text-figs, i, I, II; 2, I, I1; 3, I-III; 4, I, II; 5, I-III; 6, I, II; 7, I, II, VI, Roscoff, France. MATERIAL EXAMINED. British Museum. Great Britain, Berwick Bay, 1847. 9. 16. 66 pt. (type-material), 1847. 9. 16. 51 (fig'd Busk, 1854, pi- 76, fig- i), Johnston Coll. Channel Islands, 1963. 3. 30. 132, Harmer Coll. Madeira, 1911. 10. i. 611 and 660 (type-material of M. nitida var. intermedia), 1879. 5- 2^- 16, Watson Coll. Guernsey, 1919. 6. 24.24, Norman Coll., and many other specimens. M. nitida does not occur in the west African Collections. DIMENSIONS. Lz 0-32-0-51 mm., Iz 0-29-0-37 mm., Lav 0-13-0-16 mm., Lm 0-08- o-io mm., Lov 0-15-0-20 mm., lov 0-19-0-25 mm. Membraniporella with the frontal shield formed by 8-20 spines, each slightly branched and fused at the tips. Diatellae present. Two pairs of oral spines, present, unbranched. Avicularia present, mandible acute. The ancestrula (seen in 1963. 3. 30. 132) is membraniporan, with 10-12 long, slightly curved spines, which are not fused at the tip. The avicularian chambers do not reach the basal lamina, although frontally they appear to occur between the zooids. The chambers are not associated with the dietellae in position (seen in Guernsey, 1919. 6. 24.24, Norman Coll.). The type-specimen (encrusting shell) has 10-12 wide frontal spines, little branched at the tip, with a wide central area of fusion. Avicularia are rare and small, ovicells plentiful. Busk's figured specimen has longer zooids, but is otherwise very similar. The characters of M. nitida var. intermedia are not sufficiently distinct or consistent even to rank as varietal. Avicularia are more common, and slightly larger than in the type specimen. In some zooids, as in the type, the number of frontal spines is reduced, and the lacunae between them is correspondingly wider; but in the same colony zooids may be found with numerous frontal spines. Bobin & Prenant (1965) described an interesting convergence of characters in Membraniporella and Callopora, an Anascan genus. The species they defined and distinguished, M. nitida and C. rylandi, both had similar frontal spines, diatellae and avicularia, but differed in the relationships of the ovicell and operculum. C. rylandi was described from Roscoff, and there are specimens of the species, from the Channel Islands, which had previously been assigned by Hincks and Norman to M. nitida, in the British Museum. All the material from Madeira is referable to M. nitida, and the distribution of C. rylandi does not apparently extend as far to POLYZOA FROM WEST AFRICA 331 the south as west Africa. Another species of Callopom, which also resembles Membraniporella, does, however, occur from this area (see Cook, 1968, in press). FIGS. 6, 7. Membraniporella. Scale = 0-5 mm. 6. M. marcusi, n. sp. Achimota Coll., Ghana, 5QH. Zooid with ovicell, showing the formation of the frontal shield by the paired distal, and 2 lateral and i proximal branched spines. 7. M. aragoi (Audouin), Maturo Coll., Florida, 1965. 8. 6. 2. Zooid, showing the formation of the frontal shield by 8 little-branched spines. Membraniporella marcusi3 n. sp. (Plate i, fig B, Text. fig. 6) Membraniporella aragoi (Audouin) Marcus, 1938 : 30, pi. 6, fig. 16, Santos, 20 m., Brazil (not Flustra aragoi Audouin 1826 : 240, see below). HOLOTYPE: B. M. Achimota Coll., 59H, figured specimen. MATERIAL EXAMINED. " Calypso " Coll. II. Stn. 75, 16° 04' 20" N., 22° 58' 10" W., 25.xi.59, 45 m., Cio6B, on gorgonid stem. Achimota Coll. Stn. 35, Dredge i, 2i.xii.5o, 37m., 59!!; Stn. 45, Dredge i, 31. xii. 50111., 62A; Stn. 47, Dredge i, 4.i.5i, 44m., 14!.; Stn. 48, Dredge 2, 4.1.51, 44m., 466; Stn. 72, Dredge 3, 24.1.51, 38m., 6iA; Stn. no, Trawl i, 4.iv.5i, nom., 486; Stn. in, Trawl 2, 4.iv.5i, 43m., 49!.; Stn. 112, Trawl 3, 4.iv.5i, 43 m., 6oD; Stn. 131, Trawl i, 2. ¥.51, 37 m., 4iC, 43C; Stn. 132, Trawl 2, 2. ¥.51, 44m., 4oN, 426; Stn. 133, Trawl 2, 2. ¥.51, 51 m., 456; all on Jullienella foetida. " Atlantide " Coll. Copenhagen Museum. Stn. 44, 10° 22' N., 16° 22' W., 17. xii. 45, 41-55 m., sand and shell, 63C, on Cupuladria canariensis. Stn. 146, 9° 24' N., 14° 48' W., I3.iv.46, 50-51 m., stones, 45P, on Discoporella umbellata. M. marcusi is not present in Marche-Marchad Collection. DIMENSIONS. Lz 0-45-0-67 mm., Iz 0-40-0-52 mm., Lov 0-10-0-17 mm. Zoarium encrusting. Zooids with cryptocyst and gymnocyst small. Frontal 3 Named after Professor Ernst Marcus, who first described this species. 332 PATRICIA L. COOK shield formed by the fusion of the tips part of the paired oral spines and of 3-5 irregularly and profusely branched spines occurring round the opesia. One pair of branched oral spines, part of which are fused with the lateral walls of the ovicell, when present. Ovicell small, shallow, with a raised frontal protuberance. The ancestrula, like that figured by Savigny for M . aragoi (see below) , is membrani- poran, with 4-6 simple marginal spines, which are not fused at the tip. The average Lan is 0-25 mm. Membraniporella marcusi has many features in common with M. aragoi, but differs entirely in the number and form of the spines. Although Savigny's excellent figures makes recognition of M. aragoi easy, no material from the type-locality is available, and, in view of the variation of character within Membraniporella, and the apparent close convergence of the genus with Callopora, it is unfortunate that certain details of the structure of M. aragoi are unknown. For example, specimens assigned to M. aragoi from other areas (see below) have septulae, but, as noted above, zooids of M. nitida and M. distans may appear identical, in the absence of other distinguish- ing characters, yet one has dietellae, the other, septulae. Waters (1909 : 167) recorded M. aragoi from the Red Sea, and it may be presumed that this is the type-locality. Recent collections made there by the author, and by others, have unfortunately failed to provide any specimens. Harmer (1926 : 473) reported M. aragoi from the East Indies, and his figure is very similar to Savigny's. Kluge (1955 : 106) recorded it from Japan. It must be assumed that these forms are the same as that mentioned as occurring in the Caribbean by Osburn (1950 : 175) . The British Museum does possess a specimen from Florida (1965. 8. 6. 2, Mature Coll)., which is also very similar in character to Savigny's figure. The zooids have 6-8 regular, little-branched spines round the opesia, and the branched oral spines are fused with the proximal part of the lateral walls of the ovicell as in Savigny's figure i3. This type of fusion also occurs in M. marcusi and M. magnifica. Septulae are present in the Floridan specimen. The zooids are, on average slightly smaller than those of M. marcusi. Both the west African specimens of M. marcusi and Marcus's figure differ from M . aragoi in the smaller number of spines round the opesia. There are usually 4, but occasionally 5 are present. These spines are profusely branched, spines of the 3rd and occasionally the 4th order occurring, (compare Text-figs. 6, 7). In all the plentiful material from west Africa, there is no tendency towards production of a frontal shield of simple little-branched spines, as in M. aragoi. Some scattered zooids do, however, have only 3-4 short, simple, curved spines, which do not fuse at the tip, which appears to be a reversion to the ancestrular condition. The west African material differs from the Brazilian only in the rarity of occurrence of the single distal spine figured by Marcus. The development of the spinous processes across the frontal membrane in M. marcusi shows some similarities to that seen in Cribrilina punctata (see below). The distal pair of spines appears to grow at the same rate as the more proximal spines, and does not fuse first (cf. Cribrilaria radiata, p. 333). The distal ist and 2nd order branches of the oral spines are free, and either directed upward, or fused with the lateral walls of the ovicell, if present. The most proximal 2nd order branches POLYZOA FROM WEST AFRICA 333 of the oral spines fuse first with their most distal counterparts of the nearest opesial spines. They then continue to grow across the frontal membrane and fuse centrally, forming the equivalent of an apertural bar. In M. aragoi the bar is formed entirely by the distal pair of opesial spines, not by part of the oral spines. The Achimota Collection specimens of M. marcusi all encrust the arenaceous test of the Foraminiferan, Jullienella foetida Schlumberger (see N0rvang, 1961, and p. 348). This appears to be one of the principal substrates available for encrusting Polyzoa, which are otherwise unable to settle on the muddy sea-bottom of the Gulf of Guinea (see Buchanan, 1958 and Bassindale 1961). It is interesting that the specimens from further North, where Jullienella may be less common (see Cook, 19646 : 73), have colonized specimens of species of Cupuladriidae, which are almost exclusively associated with sandy and muddy sea-bottoms. Osburn (1950 : 176) noted that his specimens of M. pulchra exclusively encrusted the basal side of specimens of Cupuladria. CRIBRILARIA Canu & Bassler Cribrilaria Canu & Bassler, 19286 : 27, footnote. The genus has been discussed by Canu & Bassler (19290 : 33) and by Lagaaij (1952 : 57). Cribrilaria radiata (Moll) Colletosia radiata (Moll) Harmer, 1926 : 475, pi. 34, figs. 15-18, East Indies. Cribrilaria radiata (Moll) Lagaaij, 1952 : 57, pi. 4, fig. 9, Tertiary, Europe. MATERIAL EXAMINED. " Calypso " Coll. I. Stn. 17, 5° N., 5° 28' 30" W.,2i.v.56, 27 m., C56D, on Jullienella. Stn. no, Grand Frere N.E., i° 2' 45" N., 7° 17' 37" E., 7.vii.56, 25-40 m., CgE, on stone with Callopora sp.; Coll. II. Stn. 14, 14° 53' 43" N., 23° 31' 24" W., I7.xi.59, 25-30 m., C68H; Stn. 31, 14° 53' 55" N, 23° 29' 58" W., I9.xi.59, 75-170 m., C66J, with Crepidacantha setigera; Stn. 42, Punto do Anciao, 20.xi.59, I5-~3° and 60 m., Cio4B, on shell; Stn. 75, 16° 04' 20" N., 22° 58' 10" W., 25.xi.59, 45 m., Cii7B. Marche-Marchad Coll. I. 4E, Cap Matakong, Guinee Ise, on shell with many other species. Coll. II. 3iE., S.W. Ise de Madeleines, 9.1.54, 47-5 m., on bivalve shell. Achimota Coll. Stn. 126, Trawl 3, I2.iv.5i, 20 m., 37 I. Copenhagen Museum. Las Palmas, 150-160 m., 746. La Luz, Gran Canaria, 15-20 fath.,88E; 100 fath., 8gG. Mortensen Coll. " Galathea " Stn. 4, 22° 19' N., 17° 05' W., 2.xi.5o, 62 m., 8oE. " Atlantide " Stn. 44, 10° 22' N., 16° 22' W., i7.xii.45, 41-55 m., 63E. Stn. 45, 9°23'N., I5°07'W., i8.xii.45, 30-36 m., 78E. Stn. 145, 9° 20' N., 14° 15' W., I3.iv.46, 32m., 76, noN. Stn. 146, 9° 24' N., i4°48'W., 13.^.46, 50-51 m., 45E, I07H, n6A. British Museum. Madeira, 1899. 7. I. 1348, 1354, 1355, 4725, Busk Coll.; 1899. 5. i. 721, Hincks Coll; Port of Oratava, Canary Is., 50 fath., 1899. 7. I. 1350, Busk Coll.; Tangier, 1899. 7. i. 1356, 1357, 1360, Busk Coll.; Gulf of Mexico, 28° 58' N., 89° 9' W., South Pass of Mississippi River, 1961. n. 2. 39, Cheetham Coll. 334 PATRICIA L. COOK DIMENSIONS. Lz 0-49-0-60 mm., Iz 0-23-0-45, Lor 0-05-0-07 mm., lor 0-07-0-09 mm., Lov 0-I3-O-I9 mm., lov 0-17-0-20 mm., Lav 0-21-0-32 mm., Lm 0-17-0-27 mm. Zoarium encrusting. Frontal shield of zooids formed by 10-24 costae. 5-7 oral spines. Avicularia vicarious, between the zooids, chambers reaching the basal lamina, mandibles acute, often elongated. Ovicells small, imperforate, slightly raised in the centre. Diatellae present. Ancestrula membraniporan. The range of variation of this wide-spread form has been fully discussed by Harmer (1926 : 475-8). It has been reported from Madeira by several authors. The west African form shows the small, raised bosses or papillae at the bases of the costae, and the avicularia frequently have greatly elongated mandibles, and raised rostra. C. radiata occurs on shell, stone, and Jullienella, The development of the frontal shield shows interesting similarities with that of some specimens of Triporula (see below). Division ASCOPHORA IMPERFECTA Harmer Ascophora Imperfecta Harmer, 1957 : 645. Harmer defined the Ascophora Imperfecta to include those genera in which the frontal membrane and contiguous operculum were covered by an overarching frontal shield. A true compensation sac, developing from a group of cells at the proximal edge of the operculum, is therefore not developed. Two distinct methods of formation of the frontal shield appear to be present in the species referred here to the Ascophora Imperfecta. One type is that already des- cribed by Harmer. The frontal shield develops as a curved lamina, with its concavity directed proximally, which grows over the frontal membrane in a distal direction. Genera with this type of frontal shield are here represented by species of Hippo- petraUella and Metrambdotos. In the second type of development a pair of distal processes first fuses centrally to form a bar which delineates the orifice proximally. An irregular series of flattened, rounded processes then develops across the frontal from the lateral and proximal walls of the zooid. These processes later fuse, leaving large foramina which then calcify further. This type of development has only recently been observed. It is known to occur at present in three genera; Tremo- gasterina (see Powell & Cook, 1967), Exechonella and Triporula (see below). Further observations will almost certainly show that other genera have a similar type of development, and probably that still other, differing, types of frontal structure are present in the Ascophora Imperfecta. The second type of development resembles that seen in two species of Cribimorph, Cribrilaria radiata and Cribrilina punctata. In C. radiata the distal pair of costae fuse first, forming the apertural bar which delimits the proximal edge of the orifice. On the proximal side of these costae, lacunae are left uncalcified where the fusion is incomplete (see Harmer, 1926 : 477). The remaining costae advance across the frontal membrane as a series of hollow, flattened spinous processes, which form cross connections, producing the rows of intercostal lacunae. This development is superficially similar to that seen in some specimens of Triporula stellata (see p. 342). In Cribrilina punctata the development is of a slightly different nature, A pair of POLYZOA FROM WEST AFRICA 335 distal spines develop, but does not always fuse first. Simultaneously other irregular spinous processes arise round the gymnocyst. In the material examined, the order and position of the fusions of these processes seem to have a fairly constant pattern (see Text-fig, n). The resulting foramina are large, and at first, irregular. Later calcification produces the regularly porous frontal shield characteristic of the species. This type of development resembles both that of M. marcusi (see above), and that of Exechonella tuberculata (see below). Some of the developmental stages described by Larwood (1962 : 25-32) in the Cribrimorpha differ from those described above . Before any analogies and homologies can be established, much further work must be done on the development of living colonies of a large number of genera (see Osburn, 1940 : 366, 368). It is interesting to note, however, that Waters (1923 : 548-9) described apparent lateral processes involved in the calcification of the frontal wall of the Ascophoran, Lepralia otto-mulleriana Moll (see also Larwood, 1962 : 28). A specimen, labelled L. otto-mulleriana by Waters, and sent by him to O'Donoghue, has recently been incorporated into the British Museum from the O'Donoghue Collection. It is from Santa Margharita; the material described by Waters in 1923, was from Mentone. Unfortunately, the specimen was preserved dry, but restoration with 10 % solution of trisodium phosphate revealed the following features. The primary frontal wall is very thinly calcified. It appears to be developed from a series of broad plates, a distinct central suture, with 3-4 lateral sutures extending from the margins of the zooids, and alternating with the marginal pores, being visible. Occasional small pores occur on the suture lines, where the plates have joined incompletely. The secondary calcification extends from around the marginal pores, and does not obscure the central pores. If a suboral avicularium is developed, its chamber is formed by secondary calcification. The specimen is now preserved in alcohol (registration number 1966. 9. 2. 3.). It is not yet known how much importance may be attached to the type of ancestrula found in Polyzoa, or how much inter- and intraspecific variation may occur. The ancestrulae of Membraniporella, Cribrilaria radiata and Cribrilina punctata are membraniporan (see above), as is that of Tremogasterina. That of Hiantopora bidenticulata Canu & Bassler, which is here provisionally associated with Triporula, is cribriform (see below). A similar ancestrula was described for the otherwise apparently Ascophoran species, Temachia opulenta, by Jullien (1883 : 509, pi. 14, figs. 26-29), and for Schizoporella kuensis Okada & Mawatari, by Mawatari (1952 : 279, text-fig. 13). The ancestrulae of Exechonella and Triporula however, are smaller than, but are essentially of the same character as later developed zooids. TREMOGASTERINA Canu Tremogasterina Canu, Powell & Cook, 1967. Zoarium encrusting. Zooids with a frontal shield formed by the coalescence of marginal processes, leaving large foramina. Orifice contracted laterally, with oral spines. Ovicells hyperstomial, Avicularia usually present. Diatellae and septulae present. 336 PATRICIA L. COOK FIG. 8. Tremogasterina. Scale = 0-5 mm. T. perplexa n. sp. " Atlantide " Coll., Stn. 146, off French Guinea, loyF. 4 zooids, i with an ovicell. Note the large, tubular frontal foramina, and the irregular growth of the tuberculate frontal processes. Tremogasterina perplexa* n. sp. (Text-fig. 8) HOLOTYPE (and only specimen) : — " Atlantide ", Stn. 146, 9° 24' N., 14° 48' W., I3.iv.46, 50-51 m., I07F. Copenhagen Museum. DIMENSIONS. L z 0-42-0-65 mm., 1 z 0-35-0-42 mm., Lor 0-12-0-14 mm., lor 0-11-0-13 mm. LSpine 0-25-0-42 mm. Tremogasterina with no avicularia. Zooids separated by deep grooves. Orifices elongated, with the proximal border curved distally. Six large oral spines present. Frontal shield smooth, with a central area perforated by 5-9 lunate foramina. Two distal diatellae, and 2-4 lateral septulae. Ovicells with a frontal area. In spite of the complete absence of avicularia, the affinities of this species are with Tremogasterina. The zooids resemble the periancestrular zooids of T. robusta (Hincks), and the zooids of T. spathulata (Canu & Bassler) (see Powell & Cook, 1967), 4 Latin, perplexus, involved, referring to the difficulties in assigning secondarily calcined zooids to the genus Tremogasterina. POLYZOA FROM WEST AFRICA 337 in the small cribriform central area of the frontal shield. The form of the orifice, the oral spines, the ovicell, diatellae and septulae are all characteristic of the genus. The colony comprises approximately 75 zooids, and encrusts the surface of a dead colony of Membranipora arborescens (Canu & Bassler), which itself grows over an echinoderm test. At the growing edge a few zooids show that the frontal shield develops from a series of coalescing, spinous processes, as in the other species of the genus. The oral spines are large, and very long; the most proximal pair frequently being the stoutest. A pair of small distal diatellae, and 2-4 lateral septulae are present. The orifice is elongate like that of T. robusta', the opercula are dark brown, and appear to have a marginal sclerite. The proximal border of the orifice is curved distally. Secondary calcification greatly alters the appearance of the zooids. The foramina become surrounded by short tubular extensions, similar to those found in Exechonella tuberculata (MacGillivray) . The distal part of the frontal shield, or apertural bar, becomes enormously thickened; a high, ridged mucro sometimes developing, which may obscure the orifice and eventually the frontal foramina. The spines are present in nearly all the zooids. One ovicell is present, it has a small frontal area. EXECHONELLA Canu & Bassler Exechonella Canu & Bassler, 1927 : 4. Harmer, 1957 : 652. Zoarium encrusting. Zooids large. Frontal shield with numerous large foramina. Avicularia present or absent. Avicularia have been recorded in E. magna, E. antillea and E. pumicosa, but their occurrence in the last two forms is apparently sporadic (see below). Ovicells have been reported in one species, E. discoidea Canu & Bassler (19296 : 123). Lagenipora tubulosa d'Orbigny, described by Canu (1909 : 450, pi. 12, fig. n), from the Burdigalian of Dax, France, belongs to Exechonella. The zoaria of E. antillea in these Collections are nearly all slightly worn, and well- preserved growing edges are not present. A few zooids, particularly one in specimen Achimota 65 C, show evidence that the frontal shield has been formed by the fusion of irregular processes. The development of the frontal shield in Exechonella has, however, been seen particularly clearly in specimens of E. tuberculata (MacGillivray) from Port Phillip Heads, Australia (1897. 5. i. 930, 931, J. Bracebridge Wilson Coll.). The proximal part of the orifice is formed by the central fusion of the 2 most distal of a series of irregular lateral and proximal processes (see Text-figs. 9, 10). The processes fuse, forming large foramina, which then calcify further, forming tubular processes in this species. Exechonella antillea (Osburn) (Plate i, fig. E) Lepralia antillea Osburn, 1927 : 6, text-fig. 6, Cura9ao on shell, pottery. Exechonella pumicosa Canu & Bassler, 19280 : 70, pi. 14, fig. i, text-fig, na, South of Florida, 40 fath., on shell. 338 PATRICIA L. COOK Exechonella antillea (Osburn) Osburn, 1940 : 366, Porto Rico, on rock and shell. 5-20 fath; I()5O ' 95. pi- IO» ngs- 9> IO> Gulf of California; Shier, 1964 : 616, N.W. Florida, on rock and shell. lExechonella brasiliensis Canu & Bassler, igaSc : 72, pi. 3, fig. 5, Brazil. Exechonella sp. (i8.A from Ghardaqa); Harmer, 1957 : 653. MATERIAL EXAMINED. " Calypso " Coll. I. Stn. P2i, Entre Pt2 da Mina et Ise S. Ana, 4.vii.56, 8-iom., C8E; Stn. no, Grand Frere, N.E., i°2o'45"N., 7° 17' 37" E., 7.viii.56, 5-46 m. Achimota Coll. Stn. V, Tenpobo | m. square section on rock, 5.11.50, 8B, C, 65C, Coll. II, 176; Stn. 69, Dredge 5, 22.1.51, 22 m., goG. Copenhagen Museum " Atlantide " Coll. Stn. 146, 9° 24' N., 14° 48' W., 13.^.46, 50-51 m., I07V. British Museum, Ghardaqa, Red Sea (i8A), 1961.10.30.3. Aden, Khormaksar sands, on shell, 1965. 9. 4. 2, Sgt. Cambridge Coll. St. Thomas, W. Indies, 1887. 8. i. 1-3, " Challenger " Coll. E. antillea is not present in the Marche-Marchad Collection. DIMENSIONS. Lz 0-70-0-95 mm., Iz 0-60-0-80 mm., Lor 0-20-0-23 mm., l°r 0-24-0-26 mm. Lav 0-07-0-10 mm., Lm 0-03-0-06 mm. 15-40 foramina, av. width 0-025 mm. Zooids large, very irregular in shape, separated by deep grooves. Operculum dark, with marginal and proximal sclerites. Orifice with raised, flaring peristome, sometimes with spinous processes. Frontal foramina round, regular, funnel-shaped, surrounded by a salient rim when fully developed. Avicularia very rare, minute, developed from a foramen, mandible rounded. The range of variation of all characters in this species is large. The zooids vary greatly in size, those near the ancestrula being very small. The later-developed zooids are large, and very irregular in shape. The shape of the orifice may be wider than long, or longer than wide. The position of the oral condyles varies from half, to two thirds of the way down the orifice from its distal border. The size and direc- tion of the condyles also varies. In the " Atlantide " material they are large, and point inward across the orifice. In the Achimota specimens the condyles are smaller, and point downward into the cavity of the zooids. The frontal foramina, when fully developed, are surrounded by salient rims, which are funnel-shaped. The number of pores is directly related to the size of the zooids, and ranges from 15-40. The zoaria encrust shell and stone, and the older, dead parts of colonies of Stegano- porella magnilabris. The peristomes often show signs of wear, but some zooids, growing in the hollows of shell etc., notably those from Achimota Stn. 69, and " Atlantide " Stn. 146, have well raised distal and lateral flaring expansions. Four zooids in the Achimota specimen also have a short spike-like protuberance on the proximal side of the orifice. The " Atlantide " specimen has zooids with long spinous processes arising from the edge of the peristomes. Minute avicularia are present in the material from the West Indies and the " Atlantide " Collection. They appear to be derived from one of the lateral frontal foramina, and are usually found at the lateral borders of the zooids (see Osburn, 1927 and 1940). The subrostral chamber is raised, sometimes tubular. The avicularian POLYZOA FROM WEST AFRICA 339 mandible is short and rounded, and is variously orientated. Minute avicularia are also very occasionally present in the colonies from the Red Sea, which agree in all characters with those from West Africa. Osburn specified the presence of avicularia in E. antillea, and they were described by Canu and Bassler in E. pumicosa. Avicu- laria are not always present and are usually absent from large areas of the colony. Osburn (1950) did not mention their occurrence in his California material. Osburn considered that E. pumicosa (Canu & Bassler) was a synonym of E. antillea. E. brasiliensis (Canu & Bassler) was also described with small avicularia, but the number of frontal foramina was very small (13-15). In all other respects E. brasiliensis greatly resembles E. antillea and is certainly a closely related form; the discovery of further material may show that it is synonymous with E. antillea. FIGS. 9, 10. Development of the frontal shield in Exechonella. Scale = 0-5 mm. 9. E. tuberculata (MacGillivray) . Port Phillip Heads, 1897. 5. i. 930. Zooid at the growing edge, showing formation of the apertural bar and frontal foramina, which rapidly become calcined to form tubular processes. 10. E. tuberculata. Port Phillip Heads, 1897. 5. i. 931. Zooid at a later stage of development, showing closure of the apertural bar. Exechonella gigantea n. sp.5 (Plate 2, figs. C, D) HOLOTYPE (and only specimen), B. M. Sud de Goree, 24. u. 53, 4-41 m., nj, Marche-Marchad Coll. I. DIMENSIONS. Lz i-20-i-6o mm., Iz 0-90-1-40 mm., Lan i mm., Lor 0-20-0-26 mm., lor 0-20-0-26 mm. 60-80 frontal foramina, average width 0-02 mm. Zoarium encrusting. Zooids extremely large . Orifices with a thickened peristome, raised laterally and distally. Operculum with a marginal sclerite and thickened rim. Avicularia and oviceUs not seen. The zoarium completely covers one valve of a small lamellibranch shell 10 mm. in diameter, and comprises approximately 200 zooids. The ancestrula is smaller than, but has the same characters as the later developed zooids. The zooids on the 5 Latin, giganteus, large, referring to the size of the zooids. 340 PATRICIA L. COOK convex (and presumably exposed) side of the shell have low, thickened peristomes. The peristomes of those in the concavity have raised, flaring expansions, which tend to be bifid distally (see pi. 2, fig. D). Although the zooidal characters of E. gigantea are very similar to those of E. antillea, the size of the zooids, which shows no intergrading, appears to exclude it from that species. Cydicopora grandis Duvergier (1921 : 174, pi. 2, figs. 2, 3), from the Lower Miocene of Bordeaux, is very similar to E. gigantea in all characters, and is certainly referable to Exechonella. The zooids are slightly smaller, and the orifices have distinct oral condyles, which are absent in E. gigantea. Duvergier also recorded that avicularia were present in C. grandis but in view of the variability of their occurrence in E. antillea, this is probably not a difference of any specific value. The discovery of further material of E. gigantea, may show that it may be referred to E. antillea, but the present specimen appears to be distinct from that species. TRIPORULA Canu & Bassler Triporula Canu & Bassler, 1927 : 8. Enantiosula Canu & Bassler, 1930 : 23. TYPE-SPECIES, Escharipora stellata Smitt, 1873. Zoarium encrusting. Zooids with a frontal shield pierced by stellate pores formed from irregular foramina. Diatellae present. Paired lateral avicularia (not replacing diatellae) present, bar absent. Ovicells not known. Smitt (1873 : 24-6) included Escharipora (?) mucronata as the first species, with E. stellata, and also suggested that Eschar a lichenoides Busk, 1854 (= Adeonellopsis yarraensis (Waters) (seeHarmer 1957 : 799) and Lepralia distoma Busk, 1856 (= A. distoma, see Canu & Bassler, 1920 : 564), were congeneric. E. mucronata is referable to Tremogasterina (see Powell & Cook, 1967 : 9), and E. stellata was formally chosen as the type-species of Triporula by Canu & Bassler (1927). Enantiosula, with the type-species E. manica, was defined with the frontal " a tremocyst ". Examination of specimens shows that the frontal shield of E. manica is of the same nature as that of T. stellata and that the two species are not only congeneric but synonymous. Canu & Bassler (19296 : 385) suggested a close relationship between Triporula and Anarthropora Smitt. Brown (1952 : 357), who examined specimens of both A. monodon (Busk), the type-species, and Triporula biarmata (Waters) (see below), considered them to be congeneric. Both forms are similar, and the two genera may be placed in the same family. The structure of the frontal shield in A . monodon (seen in 1963. 3. 30. 9, Shetland, 80-170 fath., Harmer Coll., 1919. 6. 25. 128, Shetland, Norman Coll., and 1899. 7. I. 1452, Guernsey, Busk Coll.), shows that it belongs to the Ascophora Imperfecta. It resembles Triporula in possessing diatellae, numerous stellate pores, and in the absence of spines and ovicells. It differs in the following important characters, however. The small, distal avicularium is in series with, and replaces, one of the diatellae; there are no lateral avicularia, and the proximal avicularium develops from a fold in the peristome, which is a proximal upward extension of the apertural bar. It therefore seems advisable to maintain the two genera separately. POLYZOA FROM WEST AFRICA 341 A. horrida (Kirkpatrick, 1888 : 76, pi. 8, fig. 2, Mauritius) was pro- visionally referred to Teuchopora by Harmer (1957 : 654). Hiantopora bidenticulata Canu & Bassler (19296 : 115, pi. n, figs. 9-11, Philippines) shows several characters similar to those of Triporula. Comparison with specimens of the type-species of Hiantopora (H.ferox MacGillivray, 1899. 7. 1. 1371, Busk Coll., and 1897. 5. i. 620, Australia, see Harmer, 1926 : 237), shows that H. bidenticulata is not referable to Hiantopora. A specimen from Funafuti (60 fath., 1903. i. 29. 23) has slit-like diatellae, and paired avicularia arising in the same manner as those in Triporula (see below) . The frontal shield resembles that of Triporula, but the fora- mina are irregular and not stellate. H. bidenticulata differs from Triporula in possessing distal spinous processes (which are, however, not true spines), on the peristome. There is no distal avicularium or enlarged pore as in Triporula. The irregular frontal foramina resemble those of Tremogasterina spathulata, see Powell & Cook (1967 : n), but the avicularia have no bar. Generally, the characters of H. bidenticulata are very similar to those of Triporula, and until plentiful material allows the complete examination of the species, it may be associated with that genus. " Triporula " bidenticulata differs from other known species of Triporula in possess- ing a cribrimorph ancestrula. One is present on the slide from Funafuti. There are 7 costae on each side, and 5 oral spines. Triporula plana was described (as Enantiosula) by Osburn (1952 : 469, pi. 57, figs. 8-9) from Lower California, from 30 fath. It differed from E. manica in its smaller lateral avicularia, larger distal avicularium, and smaller zooids. Triporula biarmata (Waters) was placed in Anarthropora by Brown (1952 : 357, see above), who chose the specimen (B. M. Palaeontology Department, D 32974) from Mount Gambier, S. Australia, as lectotype. The zooids of this specimen are small (Lz 0-43-0-52 mm., Iz 0-22-0-26 mm.), whereas those of Recent material (1897. 5. i. 688-692, Port Phillip Heads, Victoria, Bracebridge Wilson Coll.) are considerably larger (Lz 0-75-0-90 mm., Iz 0-38-0-44 mm.). Some fossil specimens (e.g. D. 35460 and 34750) also have large zooids, however, as had Waters's figured specimens (see Brown, 1952). The Recent specimens from Port Phillip have ancestrulae; these are smaller than, but resemble the other zooids in all characters. Some Eocene species from North America are apparently closely related to Tri- porula ; all have large frontal pores and lateral oral avicularia. Cheilopora (?) labiosa Ulrich, 1901 : 220, pi. 60, figs. 15, 16, see Canu & Bassler, 1920 : 526, pi. 2, figs. 1-3, Lower Eocene, Maryland. Described with lateral avicu- laria, ovicells absent. Meniscopora subplana Ulrich, 1901 : 219, pi. 59, figs. 17-18, see Canu & Bassler, 1920 : 556, pi. 2, figs. 4-5, Lower Eocene, Maryland. Described with plurilaminar zoaria, frontal, marginal and basal pores (Pdiatellae), and paired avicularia. Ovicells absent, but enlarged " gonoecia " present. Cheilopora prelucidioides Canu & Bassler, 1920 : 527, pi. 68, figs, i, 2, Upper Eocene, S. Carolina. Described with lateral avicularia and fusiform diatellae. Anarthropora (?) verrucosa Canu & Bassler, 1920 : 430, pi. 7, fig. 20, Lower Eocene, Georgia. Described with three oral avicularia, ovicells absent. Phylactella magniporosa Canu (1918 : 306, pi. 9, fig. 7), from the Lutetian (Lower 342 PATRICIA L. COOK Eocene) of Aude, France, resembles Triporula in the shape of the orifice and porous frontal shield. Ovicells were stated to be present, however, and avicularia were apparently absent. The development of the frontal shield and avicularia in Triporula has been seen in T. stellata (in both the type specimen and in material from west Africa), and in T. biarmata (see below). It resembles that seen in Tremogasterina (see Powell and Cook, 1967) and differs from the " umbonuloid " development described for the Ascophora Imperfecta by Harmer (1957). The future shape of the zooids may be seen at the growing edge of the colony, delineated by the diatellae, which apparently form completely very early in the development of a new zooid. A thin calcified lamina then grows upward above the diatellae, until it is approximately twice their height, the zooids of Triporula being very deep. The lamina then curves inward slightly, and a series of ridges develops laterally and distally. These ridges bear no relationship in frequency or position to the diatellae. As the ridges deepen and lengthen, the spaces between them become partially roofed over by calcification above the level of the primary lamina. This produces a series of marginal pores round the entire zooid, which may at first sight be mistaken for diatellae. A lateral pair of these pores remains open, forming the subrostral chambers of the lateral avicularia. The most distal pore may remain open, or may form a distal avicularium. The more proximal pores appear to close, forming a continuous, ridged lamina, round the zooid. In describing E. manica, Canu & Bassler (1930 : 24) considered that the avicularia replaced the diatellae. Canu & Bassler's figure (pi. 3, fig. 10) purported to show the avicularia " visible among the diatellae ". The distal part of the photograph shows the young zooids with a row of diatellae, all of the same size; no avicularia are visible. The more proximal zooids show the next stage of development. The aperture of the zooids is reduced by the incurving of the lamina, which shows the marginal pores, and the larger, distal lateral pair which will become the subrostral chambers of the avicularia. Unfortunately, the apertures of the pores have been retouched in the photograph. Osburn (1952 : 469) mentioned that the avicularia in E. manica were not interzooecial. The next stage of development may differ slightly in T. biarmata from that in T. stellata. In T. biarmata two free distal processes arise from the lamina and fuse centrally, forming an apertural bar. The lateral and proximal parts of the lamina advance irregularly above the frontal membrane, leaving rounded foramina (see Text-fig. 14). In T. stellata, long, free, irregular processes coalesce over the frontal membrane, leaving irregular foramina in some specimens (see Text-fig. 12). In other specimens, the processes appear as flattened spines, and advance regularly across the frontal in a manner very similar to that seen in Cribrilaria radiata. In both T. biarmata and T. stellata, minute spinules growing inward from the edges of the foramina form the stellate pores characteristic of the genus. The avicularian chambers enlarge and usually obscure the lateral ridges. The aperture lengthens into a tubular peristome and in older zooids, there is a tend- ency for the frontal pores to become occluded by calcification. Ancestrulae have been seen in T. biarmata and T. stellata. They are smaller, and II POLYZOA FROM WEST AFRICA 12 ^. 13 343 FIGS. 11-14. Development of the frontal shield in Cribrilina and Triporula. Scale = 0-5 mm. ii. Cribrilina punctata (Hassall). Rasmussen Coll., Fredericksund, Denmark. Zooid from the growing edge, showing the spinous processes anastomosing to form rounded foramina. 12. Triporula stellata (Smitt). Redier Coll., Gulf of Guinea, 1964. Zooid showing formation of the apertural bar and anastomosing spinous processes. 13. T. stellata. " Calypso " Coll., Stn. 17, Cs6 I. 2 zooids showing the formation of the apertural bar (on right) and avicularian chambers (on left). 14. T. biarmata (Waters). Hincks Coll., Port Phillip Heads, 1899. 5. i. 690-692. Zooid from the growing edge, showing formation of the apertural bar and frontal foramina. have fewer frontal pores than the later developed zooids, but are otherwise similar in character. Ovicells have not been seen in Triporula. Enlarged apertures, which may have marked the position of fertile zooids were described in Adeonellopsis coccinella, but none have been seen in any of the specimens examined here. Canu & Bassler (1930 : 24) stressed that the zooids of the superposed layers of colonies of Enantiosula grew exactly above the zooids of the lower layer. The edges of the superposed layers are very regular, but the position and orientation of the zooids does not appear to bear any relationship to the underlying layers. The zooecia of the primary layer in T. stellata (e.g. " Calypso " €56!), which encrusts stone, appear to be slightly longer, and more irregular in shape, than those of other colonies, which are plurilaminar. Triporula stellata (Smitt) (Plate 2, figs. A, B, Text-figs. 12, 13) Escharipora stellata Smitt, 1873 : 26, pi. 6, figs. 130-133, Florida, 42-183 fath. (not Escharipora stellata MacGillivray = T. biarmata (Waters), see above). ? Adeonellopsis coccinella Canu & Bassler, 1923 : 161, pi. 24, figs. 5-8, Lower Miocene, Florida. Triporula stellata (Smitt) Canu & Bassler, 1927 : 8. Enantiosula manica Canu & Bassler, 1930 : 23, pi. 3, figs. 6-n, text figs. 5A-F, Galapagos. Osburn 1925 : 469, pi. 57, Gulf of California to Galapagos. MATERIAL EXAMINED: Holotype, Naturhistoriska Riksmuseet, Stockholm, No. 1805, La 19, 218. "Calypso" Coll. I, Stn. i, 21° 05' N., 17° 14' W., io.v.56, 43-45 m., C5iA, on coral. Stn. 17, 5° N., 5° 28' 30" W., 2I.V.56, 27m., C56I. Stn. 25, 4° 36' 5" N., 344 PATRICIA L. COOK i°3i"W., 24.V.56, 50 m., CioQ, on stone. Stn. 107, i° 26' 15" S., 5° 35' 40" E., 4.vii.56, 60 m., Ci3C, on coral. Marche-Marchad Coll. iB, Sud de presque 1'ile du Cap Vert, iS.ii. 54, 46-50 m. 4D, Cap Matakong, Guinee Ise, on shell, with many other species. i6P, either S.W. Madeleines, I5.ix.53, 48m., or Sud de Goree, I3.xi.53, 34-37 m., on Cleido- chasma oranense; 23M, Drag. I, iS.ii. 54, no other information. 3oF, Sud de presque 1'ile du Cap Vert, i8.ii.54, 95 m. Achimota Coll. Stn. O, Dredged shingle, 8 fath., 2 miles off Densu, Accra, 8.iv.4g, 64!!, on Cleidochasma oranense. Stn. 47, Dredge i, 4.1.51, 44 m., I4R (ancestrula) . Stn. 117, Trawl 2, v.4.51, 64m., 32? and Coll. II, I3K. Achimota, July, 1947, no other information, C, B, on Cleidochasma oranense. Copenhagen Museum. " Galathea " Stn. 4, 22° 19' N., 17° 05' W., 2.xi.5o, 62 m., 966. " Atlantide " Stn. 75, 4° 44' N., i° 36' W., 23^.46, 46 m., 5C. Stn. 145, 9° 20' N., 14° 15' W., 13 .iv.46, 32 m., 7H, noE. Stn. 146, 9° 24' N., 14° 48' W., 13.^.46, 50-51 m., 44L, 45H, I07C. Stn. 141, off Freetown, 9.^.46, 15 m., 1096. Also at Stn. 133. British Museum. Capo Blanco, west Africa, 11-35 fath., 1921 . 5 . 23 . 8. Galapagos, 33. 5 fath., 1933, 12.10.3. Canu & Bassler Coll. Gulf of Guinea, 1964, Redier Coll. The great majority of the specimens are plurilaminar, and many encrust large, erect, worn colonies of other Polyzoa, notably those of Cleidochasma oranense (Waters) . DIMENSIONS. Holotype. Lz 0-40-0-50 mm., Iz 0-30-0-35 mm., L or 0-05-0-07 mm., 1 or 0-15-0-17 mm., av. L distal av 0-08 mm., Lm 0-04 mm., av. L lateral av 0-15 mm., 1 m. 0-07 mm. West African specimens. Lz 0-35-0-60 mm., Iz 0-30-0-42 mm., L or 0-06-0-11 mm., 1 or 0-10-0-14 mm., av. L distal av 0-05 mm., Lm 0-03 mm., L lateral avo-io- 0-23 mm., Lm 0-05-0-17 mm. Zoarium encrusting, plurilaminar. Zooids with a horse-shoe shaped orifice, and a raised peristome. Frontal pores stellate. Large, paired lateral-oral avicularia present, with occasionally one small, distal avicularium. Mandibles acute, curved, directed proximally in the distal avicularium, and distally, round the sides of the peristome, or toward the orifice, in the lateral avicularia. Diatellae present. Ovicells not known. Triporula manica was described (as Enantiosula) by Canu & Bassler (1930 : 23) from the Galapagos Islands, from 33-5 fath. The species was denned as having two avicularia, but Osburn (1952 : 469) pointed out that a third, small, distal avicularium was often present, although it might be vestigial and replaced by a pore. The zooids of Osburn's specimens were larger than those of T. stellata, but those of a paratype specimen (Galapagos, Stn. 02815, 33-5 fath., 1933. 12. 10. 3, Canu & Bassler Coll.) are very similar (Lz 0-40-0-50 mm., Iz 0-30-0-40 mm., Lor 0-10-0-14 mm., lor 0-12-0-16 mm., L lat. avic. 0-10-0-18, Lm 0-08-0-15 mm.). Smitt (1873) described the differences in appearance produced by secondary calcification. At the growing edge, the operculum is just visible, the avicularia are distinct, and the pores large and stellate. As calcification proceeds the orifice becomes tubular, then immersed. The avicularian rostra also becomes less obvious, and the pores may be occluded. The distal avicularium, which was described by POLYZOA FROM WEST AFRICA 345 Smitt, is frequently replaced by a small pore, and even when present, is always small, and is soon immersed by secondary calcification. The position and size of the lateral avicularia is variable. Those figured by Smitt are small, and directed toward the proximal border of the orifice. They should be compared with those in the photograph of his specimen (pi. 2, fig. B), which show the effects of secondary calcification, and are frequently directed almost vertically. In the west African material, the avicularia are generally large, and directed towards the distal border of the orifice, as in " E. manica ". The distal ends of the rostra nearly meet in some specimens. It is noticeable that in hollows in the colony, and on the concave sides of shells, the avicularia are very large, and the rostra raised, obscuring the sides of the orifices. In other parts of the colony, and in some entire specimens, the avicularia are, however, much smaller (as in specimen Achimota C, B), and resemble Smitt's figures. Specimens from the Gulf of Mexico (" Cavalier" Stn. 311, Lagaaij Coll.), which I have examined are very similar in character to those from west Africa. Three avicularia are usually present, the rostra of the lateral pair extending at least halfway up the sides of the orifice. The frontal pores vary in development from irregular foramina to stellate pores. HIPPOPETRALIELLA Stach Hippopetraliella Stach, Harmer, 1957 : 7°°- Hippopetraliella africana n. sp.6 (Plate i, fig. C) HOLOTYPE, B. M. Achimota Coll., Stn. G, 38D, figured specimen. MATERIAL EXAMINED. Achimota Coll. Stn. E, Christiansborg shore, 15.1.49, 34D; Stn. F, as above, 14.^.49, nB; I3A; Stn. G, as above, I9.xi.49, 38D; Stn. U, Tenpbobo shore, 13.^.49, 67C; Stn. 47, Dredge i, 4.1.51, 44m., I4G; Stn. 69, Dredge 5, 22.1.50, 22 m., (dead). H. africana is not present in the " Calypso ", Marche-Marchad or " Atlantide " Collections. DIMENSIONS. Lz 0-90-1-20 mm., Iz 0-43-0-82 mm., Lo 0-23-0-27 mm., lo 0-22-0-25 mm., Lop 0-22-0-25 mm., lop 0-22-0-23 mm., Lav 0-10-0-27 nun., Lm 0-05-0-12 mm., Lov 0-41-0-50 mm., lov 0-40-0-45 mm. Zoarium unilamellar, loosely encrusting, with erect expansions. Orifices elongated, with a pair of oral condyles. Operculum with an incomplete proximal sclerite. Ovicells globose, prominent, minutely porous. Avicularia oral, frequently paired, with a complete bar, directed laterally. Basal surface with from one to five multi- porous rootlet plates per zooid, frequently very large, surrounded by a raised rim. The frontal shield develops in the typically umbonuloid manner described by Harmer for the Ascophora Imperfecta. 6 Named as the first representative of the genus from west African waters. 346 PATRICIA L. COOK H. africana has many characters similar to those of H. dorsipora (Busk). The zooids are slightly larger, but the small oral avicularia are of a similar size and shape. The mandible has, however, no teeth as are found in H. dorsipora. Occasionally, larger, more elongated avicularia, directed distally, are present ; they have a rounded mandible with a strong central sclerite. The ovicells are more prominent than those of H. dorsipora and have a raised proximal rim. H. africana also differs in that the orifice is longer than wide, and the operculum has an incomplete proximal sclerite. The basal rootlet plates are very large, some almost covering the entire basal wall. H. africana differs from both Petraliella bisinuata and P. marginata Canu & Bassler (19280 : 78, 80, Gulf of Mexico) in its smaller, rounder avicularia, and lack of any serration on the proximal rim of the orifice. H, africana is a shallow water species, distributed from the littoral region to 44 m. It is the only representative of the family so far recorded from west Africa. Harmer (1957 : 694) noted that no species belonging to the Petraliellidae had been described from the Atlantic. Utinga castanea (Busk) has been recorded from Brazil, see Marcus (1949 : 19) ; it differs from H. africana in the shape of the orifice, in the distribution of the avicularia, and in the size of the basal rootlet pores. METRARABDOTOS Canu Metrarabdotos Canu, 1914 : 472; Buge & Galopim de Carvalho, 1963 and 1964. Brown (1958 : 64) showed that Metrarabdotos is not a synonym of Trigonopora, as was stated by Canu & Bassler (1935 : 53). The genus has been discussed recently by Buge (1957, as Trigonopora}, and Buge and Galopim de Carvalho (1963 and 1964). The great majority of the species is known only from fossil deposits. Metrarabdotos unguiculatum Canu & Bassler7 (Plate i, fig. D) Metrarabdotos unguiculatum Canu & Bassler, I92ga : 128, pi. 23, figs. 6-9, text-fig. 26, Yucatan, Gulf of Mexico, Straits of Florida, coral and sand, 21-56 fath. ig28c : 92, pi. 8, fig. 9, Brazil, 49m. Calvet, 1931 : 112, Cape Verde Islands, 91 m. Redier, 1965 : 388, Gulf of Guinea, 70 m., mud, sand, shell, gorgonids. Trigonopora unguiculatum (Canu & Bassler) Marcus, 1955 : 304, pi. 6, fig. 70, Brazil, 35 m. Metrarabdotos unguiculatus (sic) Canu & Bassler, Buge & Galopim de Carvalho, 1963 : 181, figs. 23-24. 1964 : 656. MATERIAL. " Calypso " Coll. I. Stn. 25, 4° 36' 5" N., i° 31' W., 24^.56, 50 m., CioS, on stone. Coll. II. Stn. 75, 16° 04' 20" N., 22° 58' 10" W., 25.xi.5g, 45 m., Ci03A, on Echinoderm test. Marche-Marchad Coll., I. Sud de Goree, 24.xi.53, 40-41 m., nA; S.W. de Madeleines, 9-i.54, 45-46 m., 26A; as above, i5.ix.53, 48m., 24A. Col. II.; S.W. Madeleines, I5.ix.53, 48m., 406; Sud de Goree, 27-X.53, 38-42 m., 4iA; as above, I3.xi.53, 34-37 m., 43 A. 7 Dr. A. C. Cheetham, who is currently revising this genus, has examined specimens from west Africa and informs me (in lift i . iii . 67) that he considers them to be distinct from M. unguiculatum s s. POLYZOA FROM WEST AFRICA 347 Achimota Coll. Stn. L, dredge in 4 fath., 2 miles W. of Densu, i mile offshore, 2.iii.49, 276; Stn. O, off Densu River, 8 fath., 8.iv.49, 64!, 8416 + 156 (Coll. II); Stn. W, Apam shore, 3.iv.5o, 66A; Stn. 47, dredge i, 4-i.5i, 44m., I4f; Stn. 48; dredge 2, 4.i.5i, 44m., 22D, 46N., on /. foetida, 63C; Stn. 117, Trawl 2, 5.iv.5i, 64m., 32B. Coll. II, Chorkor shore, 22A, on stone. Copenhagen Museum. " Atlantide " Coll. Stn. 75, 4° 44' N., i° 36' W., 23.1.46, 46 m., 5A 4iA, 646. Stn. 85, 5° 37' N., o° 38' E., 30.1.46, 28-50 m., io8D. Stn. 145, 9°2o'N., i4°i5'W., 13.^.46, 32m., 44K, noC. Stn. 146, 9° 24' N., 14° 48' W., 13.^.46, 50-51 m., 45F, I07K. Stn. 147, 9° 28' N., 14° 58' W., 14.^.46, 45 m., 77G. British Museum. Gulf of Bahia, 1899. 5. i. 792, (as Lepralia areolata). Hincks Coll; John Adams Bank, off Brazil, 1899. 7. i. 529, 5298, Busk Coll; Victoria Bank, off Brazil, 1854. II- J5- 325-331 pt-, Brit. Mus. Cat. Coll. Gulf of Mexico, Albatross Stn. 2405, 30 fath., sand, 1932. 3. 7. 81, Canu & Bassler Coll; MBr/Ai C, Longhurst Coll. DIMENSIONS. Lz 0-76-0-93 mm., Iz 0-42-0-67 mm., Lo (secondary) o-22-o-25mm, lo 0-17-0-22 mm., Lav 0-21-0-41 mm., Lm 0-17-0-32 mm., Lfz 0-94-1-20 mm., Lov 0-41-0-59 mm., Ifo 0-45-0-56 mm. Zoarium erect, tubular, arising from an encrusting base, usually violet or purple in colour, even when preserved. Zooids with conspicuous marginal pores, bordered by ridges, and 5-7 lateral and 1-2 distal multiporous septulae. Orifice with a raised peristome, which has a proximal sinus and a channel running down inside to the operculum. Avicularia lateral-oral, replacing marginal pores. The rostrum acute, curved, directed distally, and the mandible running up the side of the peristome, curving into the secondary orifice. Ovicells enormous, with a porous frontal and large marginal pores; orifice very wide, slit-like, proximal lip everted and thickened. M. unguiculatum is one of the most easily recognizable of all the species from west Africa. The large, erect, tubular zoaria, and the purple colour, make it possible to distinguish it in samples with the naked eye. The colour is principally provided by the thick cuticle covering the zooid, and by the polypides and frontal membranes. The cuticle is also frequently covered by a layer of diatoms in specimens from both sides of the Atlantic (see also Marcus, 1955 : 304). The specimens Achimota Coll. II, 22A and " Calypso " Coll. C I03A, are encrusting and plurilaminar. The colour is pale pink, and the avicularia and zooids are slightly smaller (on average) than those of the erect specimens. The frontal shield develops in the umbonuloid manner (see p. 324). The large marginal pores are at first conspicuous, but as the shield develops they become bordered by ridges of calcification which raise the level of the margins of the zooids laterally and proximally. The funnel-shaped openings between the ridges then become tuberculate. The primary orifice is rounded and slightly elongated. As the peristome develops a pair of processes delineate a rounded sinus proximally at its base. The processes extend with the upward growth of the peristome, forming a channel, and also produce a proximal sinus in the secondary orifice. The relationships of the frontal membrane and operculum have been described, 348 PATRICIA L. COOK for M. gulo (see below), by Marcus (1955 : 304). Dissection of these specimens has shown that in M. unguiculatum there is no well defined operculum. A thickened, semicircular, flap-like extension of the frontal membrane protrudes into the base of the peristome. It is attached laterally by muscles inserted into the lateral walls of the base of the peristome. Although both Canu & Bassler and Buge & Galopim de Carvalho have described small avicularia at the edges of the orifice of the fertile zooids, none have been seen in any of the material listed above. An enlarged marginal pore is, however, frequently present. The embryos of M. unguiculatum are very large (average dimensions Le 0-45 mm., le 0-37 mm.). Apart from Buge & Galopim de Carvalho's mention of an unnamed Recent species from the Eastern Atlantic (1963 : 187), the only Recent record attributed to M. unguiculatum from the west African coast before that of Redier (1965) was that of Calvet (1931 : 112), who listed it from the Cape Verde Islands (91 m., on Litho- thamnion). This is remarkable, as M. unguiculatum is a robust and easily recogniz- able species, which occurs in these Collections in large quantities. Canu & Bassler 1925 and 19286) did not, however, find it in their Moroccan material, and its distri- bution appears not to extend further north than the Cape Verde area. Marcus (1955 : 304, pi. 6, figs. 71-72, pi. 7, figs. 73-77) described, as Trigonopora gulo, a Recent species from Brazil (Espirito Santo, 35 m.). The avicularia were either small, or large, with bifurcate mandibles, there were none present on fertile zooids. Buge & Galopim de Carvalho did not include M . gulo in their survey of the genus. It seems possible that M. tuberosum Canu & Bassler (19286 : 92, pi. 8, figs. 3, 4, off Brazil, 36 m.), which was figured with small avicularia, may be closely related to M. gulo. Canu & Bassler (19286 : 92, pi. 8, fig. 9, Brazil, 49m.), also recorded M. unguiculatum, and the majority of specimens in the British Museum Collection is from the same area (see above). M. unguiculatum appears to be associated with moderately deep water, and sandy sea bottoms. Lagaaij noted (1952 : 124, footnote), that the distribution of his Trigonopora nysti in sand and clay deposits, seemed " to indicate a rather deep-water facies ". The species described by Lagaaij under Trigonopora are all referable to Metrarabdotos (see Brown, 1958 : 64). The specimens from shallow water in the west African Collections are usually dead and worn, the remaining specimens are from 34-64 metres. Several of the Achimota Collection Stations at these depths are in or near the area of the " silty sand community " defined by Buchanan (1958). The majority of the specimens comprises broken fragments, but two show evidence of the mode of growth. " Calypso ", CioS encrusts a stone, but includes the base of an erect, tubular branch. Achimota 46N is another erect fragment growing from a base which encrusts a fragment of Jullienella foetida. J. foetida is a Foraminiferan with an argillaceous test which forms large foliaceous colonies. It is one of the dominant forms of the silty sand community off the coast of Ghana (see Buchanan, 1958, and also N0rvang, 1961). SUMMARY Five species of Cellar ia are described from the west African region, one of which POLYZOA FROM WEST AFRICA 349 is considered to be new. Only three species belonging to the Cribrimorpha have been found from the area. One, Membraniporella marcusi, from west Africa and Brazil, is here considered to be distinct from M. aragoi, a similar species which occurs in the Red Sea and the Gulf of Mexico. The Division Ascophora Imperfecta com- prises two distinct groups of genera, which differ in the method of development of the frontal shield. Both groups are represented from west Africa. The first, in which the shield is formed from the anastomosis of spinous processes, and is similar to that found in the Cribrimorpha, includes species of Tremogasterina, Exechonella and Triporula. The second, in which the shield develops in the umbonuloid manner described by Harmer (1957), includes species of Hippopetraliella and Metrarabdotos. ACKNOWLEDGEMENTS My thanks are due to Dr. A. Andersson (Naturhistoriska Riksmuseet, Stockholm), Dr. F. Mature (University of Florida) and Dr. E. Rasmussen (Copenhagen University), who kindly lent specimens for examination and comparison. Drs. J. Knudsen and Torben Wolff (Zoologisk Museum, Copenhagen), provided the opportunity for examination of the " Atlantide " Collection from west Africa. Drs. A. Cheetham (Smithsonian Institution, Washington) and R. Lagaaij (Shell Exploratie en Productie Laboratorium, The Netherlands) have given much helpful advice. Finally I should like to thank Messrs. H. V. Christensen (Zoologisk Museum, Copenhagen) and J. V. Brown (British Museum, Natural History) , for their care and patience in the prepara- tion of the photographs. REFERENCES AUDOUIN. J. V. 1826. Explication sommaire des Planches de Polypes de 1'Egypte et de la Syrie, in Description de 1'Egypte, Hist. Nat., i, 4. Savigny, J.C., ? date, pis. i-xiv, Paris. BASSINDALE, R. 1961. On the Marine Fauna of Ghana. Proc. zool. Soc. Lond., 137, 4 : 481- 510. BASSLER, R. S. 1936. Nomenclatorial notes on fossil and Recent Bryozoa. /. Wash. Acad. Sci., 26, 4 : 156-162. BOBIN, G. & PRENANT, M. 1965. Callopora rylandi, n. sp. (Bryozoaires Chilostomes, Alderi- nidae). Cah. biol. mar., 6 : 277-291. BROWN, D. A. 1952. The Tertiary Cheilostomatous Polyzoa of New Zealand. Brit. Mus. (N.H.) : i-xii, 1-405, London. 1958. Fossil Cheilostomatous Polyzoa from South-West Victoria. Mem. Geol. Surv. Victoria, 20 : 1-90. BUCHANAN, J. B. 1958. The bottom fauna communities across the continental shelf off Accra, Ghana (Gold Coast). Proc. zool. Soc. Lond., 130, i : 1-56. BUGE, E. & GALOPIM DE CARVALHO, A. M. 1963. Revision du genre Metrarabdotos Canu, 1914 (Bryozoa, Cheilostomata) . Rev. Fac. Ci&nc. Lisboa, 2C, u, 2 : 137-196. 1964. Nouvelles observations sur le genre Metrarabdotos Canu, 1914. Garcia de Orta (Lisboa), 12, 4 : 651-656. BUSK, G. 1854. Catalogue of the Marine Polyzoa in the . . . British Museum, part 2 : i-viii, 55-120, London. 1856. Zoophytology. Quart. J. Micr. Sci. 6 : 124-130. 1858. Op. cit., On some Madeiran Polyzoa. Ibid. : 261-263. CALVET, L. 1907. Bryozaires, Exped. Sci. Travailleur et Talisman 1880-1883, 8 : 355-495. 1931. Bryozoaires provenant des Campagnes Scientifiques du Prince Albert ler de Monaco. Res. Camp. Sci. Monaco, 83 : 1-152. 350 PATRICIA L. COOK CANU, F. 1909. Les Bryozoaires fossiles des Terrains du Sud-Ouest de la France, III, Burdigalien. Bull. Soc. geol. Fr., 4, 9 : 442-456. 1914. Op. cit., VIII, Rupelien de Gaas. Ibid., 14 : 465-474. — 1918. Les Bryozoaires fossiles de la region des Corbieres. Ibid., 18 : 294-314. CANU, F. & BASSLER, R. S. 1920. North American Early Tertiary Bryozoa. Bull. U.S. nat. Mus., 106 : 1-879. — 1923. North American Later Tertiary and Quarternary Bryozoa. Ibid., 125 : 1-302. — 1925. Les Bryozoaires du Maroc et de Mauritanie, ler Memoire. Mem. Soc. Sci. nat. Maroc, 10 : 5-79. — 1927. Classification of the Cheilostomatous Bryozoa. Proc. U.S. nat. Mus., 69, 14 : 1-42. — 1928^. Fossil and Recent Bryozoa of the Gulk of Mexico. Ibid., 72, 14 : 1-199. — 19286. Les Bryozoaires du Maroc et de Mauritanie, 2e Memoire. Mem. Soc. Sci. nat. Maroc., 18 : 7-85. — 1928^. Bryozoaires du Bresil. Bull. Soc. Sci. nat. med. Seine-et-Oise, 2, 9, 5 : 58-110. — I929». Bryozoaires Eocenes de la Belgique. Mem. Mus. roy. Hist. nat. beige., 39 : 3-68. — 19296. Bryozoa of the Philippine Region. Bull. U.S. nat. Mus., 100, 9 : 1-685. 1930. The Bryozoan Fauna of the Galapagos Islands. Proc. U.S. nat. Mus., 76, 13 : 1-78. COOK, P. L. i964«. Polyzoa from west Africa. I. Notes on the Steganoporellidae, Thala- moporellidae and Onychocellidae (Anasca, Coilostega). Ann. Inst. Oceanogr. (Calypso, VI), 41 : 43-78. — 19646. Polyzoa from west Africa. Notes on the genera Hippoporina Neviani, Hippo- porella Canu, Cleidochasma Harmer and Hippoporidra Canu & Bassler (Cheilostomata, Ascophora). Bull. Brit. Mus. (N.H.) Zool., 12, i : 1-35. 1968. Polyzoa from west Africa. The Malacostega, part i. Ibid, (in press). DUVERGIER, J. 1921. Note sur les Bryozoaires du Neogene de 1'Aquitaine. Act. Soc. Linn. Bordeaux, 72 : 5-41. GALOPIM DE CARVALHO, A. M. 1963. Les Cellaria (Bryozoa, Cheilostomata) du Tertiare du Portugal. Rev. Fac. Cienc. Lisboa, 2C, 11, i : 1-28. HARMER, S. F. 1923. On Cellularine and other Polyzoa. /. Linn. Soc. (Zool.), 35 : 293-361. — 1926. The Polyzoa of the Siboga Expedition, Part 2, Cheilostomata, Anasca. Rep. Siboga Exped., 28b: 181-501. I957- Op. cit., Part 4, Cheilostomata, Ascophora. Ibid., 28d : 641-1147. HASTINGS, A. B. 1932. The Polyzoa, with a note on an associated Hydroid. Rep. Great Barrier Reef Exp., 4, 12 : 399-458. — 1947. Notes on Polyzoa (Bryozoa), III. On some Species of Cellaria . . . Ann. Mag. nat. Hist, (n) 13 : 217-241. HINCKS, T. 1888. The Polyzoa of the St. Lawrence . . . Ibid. (6) 1 : 214-227. JOHNSTON, G. 1838. A History of the British Zoophytes. Ed. i. Edinburgh, London and Dublin. JULLIEN, J. 1883. Dragages du Travailleur, Bryozoaires. Bull. Soc. zool. Fr. 7 : 497-529. KIRKPATRICK, R. 1888. Polyzoa of Mauritius. Ann. Mag. nat. Hist. (16) i : 73-85. KLUGE, G. A. 1914. Die Bryozoen der Deutschen Siidpolar-Expedition, 1901-3. i. Deutsche Siidpolar-Exped., 15, Zool. 7 : 601-678. — 1955. Bryozoa, in Ushakov, P. V. Atlas of the invertebrates of the far eastern seas of the U.S.S.R. Acad. Imp. Sci. Leningrad : 99-111. LAGAAIJ, R. 1952. The Pliocene Bryozoa of the Low Countries. Meded. Geol. Sticht., C5, 5 = 1-233- LARWOOD, G. P. 1962. The Morphology and Systematics of some Cretaceous Cribrimorphs, Polyzoa (Pelmatoporinae). Bull. Brit. Mus. (N.H.) Geol., 6, i : 1-285. MACGILLIVRAY, P. H. 1883. Descriptions of New, or Little Known, Polyzoa, Part 2. Trans. Proc. roy. Soc. Viet., 19 : 130-138. MARCUS, E. 1938. Bryozoarios Marinhos Brasileiros, 2. Bol. Fac. Filos. Cienc. S. Paulo, Zool. 2 : 1-137. POLYZOA FROM WEST AFRICA 351 MARCUS, E. 1949. Some Bryozoa from the Brazilian Coast. Comun. Zool. Mus. Hist. Nat. Monte- video, 3, 53 : 1-33. 1955. Notas sobre Briozoos marinhos Brasileiros. Arq. Mus. nac., 42 : 273-324. MAWATARI, S. 1952. Bryozoa of the Kii Peninsula. Publ. Seto Mar. Lab., 2, 2 : 261-288. NORMAN, A. M. 1909. The Polyzoa of Madeira and neighbouring islands. /. Linn. Soc. (Zool.) 30 : 275-341. NORVANG, A. 1961. Schizamminidae, a New Family of Foraminifera. Atlantide Rep., 6 : 196-201. O'DoNOGHUE, C. H. & O'DoNOGHUE, E. 1923. A Preliminary list of Polyzoa (Bryozoa) from the Vancouver Island Region. Stud. Biol. Stas. Can., 1 : 143-201. d'ORBiGNY, A. 1851. Pal6ontologie Francaise, Terrains Cr6tac6s, 5, Bryozoaires, Paris. (Pages 1-188 were published in 1851, see Harmer, 1957 : m8). OSBURN, R. C. 1927. The Bryozoa of Cura9ao. Bijdr. Dierk., Amsterdam, 25 : 123-131. 1940. Bryozoa of Porto Rico. Scientific Survey of Porto Rico and the Virgin Islands, 16, 3 : 321-486. 1950. Bryozoa of the Pacific Coast of America, Part i, Cheilostomata Anasca. Rep. Allan Hancock Pacific Exped., 14, i : 1-269. 1952. Op, cit., Part 2, Cheilostomata-Ascophora. Ibid., 14, 2 : 271-611. POWELL, N. A. & COOK, P. L. 1967. Notes on Tremogasterina Canu and Tremogasterina robusta (Hincks). Cah. biol. Mar, 8: 7-20. PRENANT, M. & BOBIN, G. 1966. Bryozoaires Chilostomes Anasca. Faune de France, 68: i- 647, Paris. REDIER, L. 1965. Hydraires et Bryozoaires du Golfe de Guin6e. Bull. Mus. Hist. Nat. Paris, 2, 37 : 367-394. SHIER, D. E. 1964. Marine Bryozoa from Northwest Florida. Bull. mar. Sci. Gulf Caribbean, 14, 4 : 603-662. SMITT, F. A. 1873. Floridan Bryozoa, Part 2. K. svenska VetenskAkad. Handl., 11, 4 : 1-83. SOULE, J. D. 1959. Results of the Puritan- American . . . Expedition to Western Mexico, 6, Anascan Cheilostomata (Bryozoa) of the Gulf of California. Amer. Mus. Nov. No. 1969 : i- 54- THORNELY, L. R. 1912. The Marine Polyzoa of the Indian Ocean from H.M.S. ' Sealark '. Trans. Linn. Soc. Lond. (Zool.), 15 : 137-157. ULRICH, E. O. 1901. Bryozoa, in Maryland Geological Survey, Eocene. Md. geol. Surv. stratigr. Mem. : 205-222. WATERS, A. W. 1909. Reports on the Marine Biology of the Sudanese Red Sea ... 12, The Bryozoa, part i, Cheilostomata. /. Linn. Soc. (Zool.), 31 : 123-181. 1923. Mediterranean and other Cribrilinidae . . . Ann. Mag. nat. Hist., (9) 12 : 545-573. PLATE i Membraniporella, Hippopetraliella, Metrarabdotos and Exechonella. FIG. A. Membraniporella aragoi (Audouin). Florida, 1965. 8. 6. 2., Mature Coll. Zooids and ovicells. x 16. FIG. B. M. marcusi n. sp. 5QH, Achimota Coll., Ghana. Zooids and ovicells, note ances- trula. X 13. FIG. C. Hippopetraliella africana n. sp. 38D, Achimota Coll., Ghana. Zooids and ovicells. XI6. FIG. D. Metrarabdotos unguiculatum Canu & Bassler. 326, Achimota Coll., Ghana. Part of a tubular branch, showing zooids with avicularia, and two ovicells. Note the umbonuloid development of the frontal wall of the distal zooids, and, proximally, one partially developed, aborted ovicell. x 11.5. FIG. E. Exechonella antillea (Osburn). &5C, Achimota Coll., Ghana. Zooids showing slightly raised peristomes and large frontal foramina, x 20. Figures A-D photographed by Mr. H. V. Christensen (Zoologisk Museum, Copenhagen), figure E photographed by Mr. J. V. Brown (British Museum, Natural History). Bull. Br. Mus. nat. Hist. (Zool.) 15, 7 PLATE i PLATE 2 Triporula and Exechonella. FIG. A. Triporula stellata (Smitt). " Atlantide " Stn. 133, Angola, Zoologisk Museum, Copenhagen. Zooids and avicularia. Note developing frontal shield of the distal (left-hand) zooid. x 43. FIG. B. T. stellata. S.W. of Tortugas, 60 fath., Pourtales Coll., No. 218, No. 1805, Natur- historiska Riksmuseet, Stockholm. Zooids and avicularia. Note the difference in appearance of the zooids resulting from secondary calcification. x 55. FIG. C. Exechonella gigantea n. sp. Marche-Marchad Coll. I, nj, Senegal. Holotype Con- vex face, of zoarium. Note the short, thickened peristomes. x 8. FIGS. D, E. The same specimen. Concave face of zoarium. Note the, prominent peristomes at the centre of the colony. Composite picture taken at two levels of focus, x 8. Photographs by J. V. Brown. Bull. BY. Mus. nat. Hist (Zool.) 15, 7 PLATE 2 PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING I , CAUCUS LONGICAUDATUS \$ BRADY, 1899 (CALIGIDAE: COPEPODA) R. R. PARKER BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 15 No. 8 LONDON: 1968 -9 CALGIUS LONGICAUDATUS BRADY, 1899 ' (CALIGIDAE: COPEPODA) BY R. R. PARKER n Fisheries Research Board of Canada Biological Station, INanaimo, B.C. ^•353-368; 17 Text-figures. BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 15 No. 8 LONDON: 1968 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the 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. 15, No. 8 of the Zoological 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.) Trustees of the British Museum (Natural History) 1968 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 9 January, 1968 Price Eight Shillings. CAUCUS LONGICAUDATUS BRADY 1899 (CALIGIDAE: COPEPODA) By R. R. PARKER INTRODUCTION IN 1899 Brady described and illustrated a single specimen of Caligus obtained in a surface plankton-net tow at Port Chalmers, New Zealand. Both his description and figure were inadequate for identification of the species; the figure erroneously depicted the 2nd antennae and omitted the sternal furca and ist thoracopods, etc. One cannot be certain even of the sex although Hewitt (1963), who further described the species from Brady's figure, thought the specimen to be a male. Aside from a listing by Hutton (1904), the binomen had not been mentioned again in the literature until 1963. Cleugh (1966) disagreed with Hewitt's proposal for a new name and argued that the name C. longicaudatus Brady was valid; but he called attention to the facts that the species was founded on a single specimen, was inadequately described, and required validation. A letter to Dr. J. P. Harding quickly established that Brady's specimen existed in the British Museum (Natural History). Through the efforts of Miss Patricia Lofthouse, Entomostraca Section, the Trustees of the Museum kindly loaned Brady's specimen for examination. The specimen was mounted, ventral side up, on a glass slide. The cover slip was crazed, having been supported by the large maxillipeds while the mountant had shrunk. The specimen could not be examined in that condition. From the report by Stock (1960) it was learned that Brady's specimens were usually mounted in a glycerin-gelatin medium and could be recovered by soaking in water. (Dr. Stock's description of the state of the mounted specimens aptly applies in the present case.) Thus it was possible to recover the specimen, which was severely damaged, and by clearing with lactic acid, to redescribe it. The specimen was drawn in detail, using a camera lucida, by Miss Margaret Dean of this laboratory. I am also indebted to Dr. Z. Kabata and to Dr. L. Margolis for consultation in this project. The specimen has been returned to the British Museum (Natural History), as No. 1951.8.10.141 Type, in ethanol. The following redescription presents all of the detail which could be discerned from the type specimen. I have deliberately presented a very brief verbal account, relying upon the figures for anatomical detail. Caligus longicaudatus Brady, 1899 The specimen is a male. Exclusive of the terminal spines of the caudal lamellae the total length was 4-93 mm, (Brady gave 5 mm.). From the dorsal aspect (Text- fig, i) the carapace length including the frontal lobes is 58% the total length or approximately 2-84 mm. The maximum width of the carapace measured 2-93 mm., however the specimen was severely flattened in its former mount and did not regain its original shape. The dorsal surface of the carapace was marked with scattered ZOOL. 15, 8. 27 356 R. R. PARKER minute branched cilia, positioned in a bilaterally symmetrical pattern. Their func- tion is unknown to me. Similar cilia also were found on the dorsal surface of the 3rd thoracopods, the 4th and genital segments, and the abdomen. The abdomen is of 2 segments, the anterior being slightly wider but about half the length of the posterior segment. The caudal lamellae bear the usual 3 small and 3 large ciliated setae (cilia not shown) and the entire medial edge of each lamella is ciliated. From the ventral aspect (Fig. 2) the lunules appear semi-circular and prominent. FIGS. 1-2. Caligus longicaudatus. i. Dorsal view. 2. Ventral view. CALIGUS LONGICA UDATUS 357 The frontal plates are about 0-23 mm. long and are bordered by a hyaline fringe broken at the midline by the embryonic cement gland. Cephalic appendages are shown in larger detail in Text-figs. 3-9. Antennules (Text-fig. 3) are composed of 2 segments. The basal segment bears 25 spines ventrally and 2 spines dorsally which are arranged along the lateral edge. Twenty-three of the ventral spines are definitely ciliated (cilia not illustrated). The remaining 2, situated at about mid-length, are thought to be naked. The 2 dorsal spines are ciliated. The tip of the distal segment bears 13 naked spines, 2 of which appear to have a common base. A I4th naked spine originates from the medial edge at about mid-length of the segment. As is shown in Fig. 3, the spines of the basal segment are of variable lengths, but are relatively long for members of this genus; 2 spines originating from the distal edge project nearly to the tip of the distal segment. Antennae (Text-fig. 4) are modified as is usual for the male and are composed of at least 3 segments (a basal segment may be incorporated into the ventral cephalic surface). The first and second segments are sculptured with adhesion pads. The distal segment is modified into a small claw which bears a fine membrane along each edge. Originating with these membranes are 2 accessory spines. The post-antennary process (Text-fig. 5) is strongly developed, slightly hooked and pointed. Two papillae on the base give rise to hair-like setae and a third papilla near the base gives rise to an additional hair-like seta. The mouth cone (Text-fig. 6) is the usual tube-within-a-tube structure with the mandibles entering mandibular tubes (formed by involutions of the ectoderm) near the base of the mouth cone. The mandibular tubes penetrate the mouth cone distally, thus separating the cone into dorsal and ventral flaps. A circular flap, apparently separated from the ventral flap proper, extends the ventral region of the mouth cone and is bordered distally by a fringed membrane. A similar membrane is thought to border the dorsal flap. Mandibles (Text-fig. 7) are composed of 4 segments; the distal segment bears 12 teeth on the medial edge. The maxittules (Text-fig. 8) are largely incorporated into the ventral integument. Anteriorly a papilla bears 3 flattened spines of dissimilar lengths. Posteriorly the maxillule is represented by a prominent heavy spine. Faint striations were seen on the distal half along the ventro-lateral edge. Posterior to the maxillules are a pair of adhesion pads (Text-fig. 9) which may be of taxonomic significance. Maxillae (Text-fig. loa, b) are interpreted to be composed of 4 segments. The basal segment is short and heavy, followed by an equally thick but long second seg- ment. The 3rd segment is long and slender and bears the usual fringed lappet about two-thirds the segment length distally. In addition, a clearly defined fringe extends from this lappet distally to the base of the fourth segment. The laterodistal corner is extended into a prolonged pointed extension, fringed along both borders. The 4th segment is nearly one-third the length of the third segment, tapers to a point and bears 3 definite rows of cilia along its length, possibly a fourth row dorsally, which are so spaced to divide the circumference into 4 equal parts. The maxillipeds (Text-fig, na, b) are strongly developed, as is usual for males of the genus. The medial edge is developed into a protruding rough spine into which the 358 R. R. PARKER FIGS. 3-9. Caligus longicaudatus. 3. Right antennule, ventral view. 4. Right antenna, in situ, ventral view. 5. Right post-antennary process, in situ, ventral view. 6. Mouth cone, in situ, ventral view. 7. Left mandible, ventral view as seen in the mouth cone. 8. Maxillule, in situ, ventral view. 9. Adhesion pads, in situ, ventral view. FIGS. 4, 5, 6, 8 and 9 are depicted in positional relationship to the frontal membrane and and lunule show above them as seen from ventral view. CALIGUS LONGICAUDATUS 359 O.I mm O.I mm FIGS. ioa-i3b. Caligus longicaudatus . loa. Right maxilla, ventral view. lob. Enlarge- ment of apical end of left maxilla, ventral view. na. Right maxilliped, ventral view, lib. Portion of left maxilliped, ventral view. 12. Sternal furca, ventral view. i3a. Right ist thoracopod, ventral view. i3b. Enlargement of apical portion of right ist thoracopod. 360 R. R. PARKER tip of the maxilliped fits when closed. The 2 distal segments together form a claw. At the suture line, on the distal segment, are located 2 small spines. The distomedial edge of the apical segment is sharp, resembling the blade of a knife. The sternal furca (Text-fig. 12) of the specimen is damaged, however the right prong is flattened and slightly spatulate at the tip. The ist thoracopods (Text-fig. I3a, b) consist of a 2-segmented protopod and a 2-seg- mented exopod. The endopod is represented by a small papilla projecting into a minute spine. The 4 apical spines of the distal exopodal segment are unbranched and apparently naked. The 2nd thoracopods (Text-fig. 14) consist of a 2-segmented protopod, and 3-seg- mented exopod and endopod. The yd thoracopods (Text-fig. 15) are not unusual. The heavy spine on the ist exopodal segment is flattened, curved, with a knife edge laterally. Dorsally, a small, ciliated (cilia not shown Text-fig, i) spine originates from the protopod and projects over the base of the exopod. The 4th thoracopods (Text-fig. 16) are composed of 3 segments ; the lateral edge of the distal segment does not posess a spine. The apical spines of the 2nd and 3rd seg- ments are finely ciliated along both borders. Those of the 3rd segment are covered at their bases by finely pectinated lamellae. The $th thoracopods (Text-fig. 17) are represented by 2 papillae, the lateral one bears a single ciliated spine, the medial papilla bears 2 similar ciliated spines. The 6th thoracopods (Text-fig. 17) are found caudally on the ventral surface of the genital segment. They consist of a pair of caudally projecting lobes terminated by 2 ciliated spines. A small naked spine is located on the genital segment between the 5th and 6th thoracopods. DISCUSSION While Brady's name C. longicaudatus is available for his specimen it remains to be determined if the species was described prior to iSgg1 under another name. Ac- cording to our records there were 108 species named to the genus prior to that date. Of these 32, as listed in Table I, are recognized as belonging to other genera. In- cluded in this list is C. lessonius Risso, 1826, described as possessing foliaceous plates attached to the first 2 segments of the abdomen. A further 20 species, as listed in Table II, are recognized to be junior synonyms of previously described species. Included here is C. appendiculatus F. Miiller, 1852, described from a chalimus stage found on Cyprinus rutilus on the shore of the Baltic Sea. The binomen was last mentioned by Nordmann (1864), i.e. more than 100 years ago. Therefore, according to Article 23 (b) of the International Code of Zoological Nomen- clature (1961), it has no status. It is obvious, however, that Miiller described a juvenile male C. lacustris Steenstrup and Liitken, 1861. The male of this species was not figured nor adequately described until Redeke's 1939 paper. In Table III a list of 13 species names considered nomina nuda is presented. Only by examining the type material can the status of these species be established. All 1 Although 1901 is the commonly quoted date of publication of Brady's paper, it appeared as a separate in 1899 and was bound, with other sections, into a volume issued in 1901. CALIGUS LONGICAUDATUS 361 14 FIGS. 14-17. Caligus longicaudatus . 14. Right 2nd thoracopod, ventral view. 15. Right 3rd thoracopod, ventral view. 16. Right 4th thoracopod, ventral view. 17. Ventral view of left caudal portion of genital segment showing 5th and 6th thoracopods. 362 R.R.PARKER but two of the remaining species are grouped in Tables IV to VI according to ana- tomical features which readily distinguish them from C. longicaudatus. The two species remaining for consideration are C, lacustris Steenstrup and Liitken, 1861, and C. longipedis Bassett-Smith, iSQSc. C. lacustris has only been reported from fresh water or from freshwater fishes which have been found in estuarine waters of low salinity. The comparative lengths of the apical spines of the 4th thoracopod are completely dissimilar to those of C. longicaudatus. C. longipedis also can be separated from C. longicaudatus by the relative much longer length of the 4th thoracopods of the former, although the two species are in many ways similar. It is concluded that C. longicaudatus Brady, 1899, is a valid species although known only from a single male specimen taken in a surface townet. Of the 187 species named to the genus Caligus since 1899 there are 162 considered valid members of the genus today. I have compared the descriptions of these species with C. longicaudatus and cannot definitely assign any of the former to the latter species. C. pterois Kurian, 1949, and C. debueni Stuardo and Fagetti, 1960, are so incompletely described that no comparison is possible. C. brevicaudatus A. Scott, 1901, and C. calotomi Shiino, 1954, are known from females only. These species are similar to C. longicaudatus in many morphological details, i.e., thoracopod structure and the sternal furca, yet the gross body proportions may indicate dissimilarity. C. tetrodontis Barnard, 1948, is incompletely described. From the details provided by Barnard the species may be identical. These five species are given the status species inquirenda. It is not surprising that C. longicaudatus has not been reported since 1899. Rela- tively little work on the caligids has taken place in New Zealand waters. From the description provided herein it should now be possible to recognize C. longicaudatus Brady, 1899, and it should be looked for in the New Zealand fauna. TABLE I List of Caligus species named prior to 1899 transferred to other genera Caligus sp. Author Transferred by productus n. sp. Mtiller, 1785 Latreille, 1829 crassus n. sp. Abildgaard, 1794 Kr0yer, 1837-1839 oblongus n. sp. Abildgaard, 1794 Kr0yer, 1837-1839 imbricatus n. sp. Risso, 1816 Desmarest, 1825 piscinus n. comb. Lamarck, 1818 (in part) Kr0yer, 1837-1839 bicolor n. comb. Lamarck, 1818 Baird, 1850 smithii n. comb. Lamarck, 1818 Baird, 1850 heptapus n. sp. Otto, 1821 Kroyer, 1837-1839 molvae n. sp. Latreille, 1825! Wilson, 1905 lessonius n. sp. Risso, 1826 (see text) paradoxus n. sp. Otto, 1828 Kr0yer, 1837-1839 pharaonis n. sp. Nordmann, 1832 Humes, 1965 hippoglossi n. comb. Kr0yer, 1837-1839 Baird, 1850 pectoralis n. comb. Kr0yer, 1837-1839 Baird, 1850 CALIGUS LONGICAUDATUS 363 C aligns sp. salmonis n. sp. sturionis n. sp. nordmannii n. sp. ornatus n. sp. vespa n. sp. stromii n. nom. gracilis n. sp. bagri n. sp. branchialis n. sp. heckelii n. sp. macrurus n. sp. pacificus n. sp. brevipedis n. sp. obscurus n. comb. nautili prov. n. benedeni n. sp. hirsutus n. sp. parvus n. sp. TABLE I — contd. Author Kr0yer, 1837-1839 Kr0yer, 1837-1839 Milne-Edwards, 1840 Milne-Edwards, 1840 Milne-Edwards, 1840 Baird, 1848 Van Beneden, 1851 Dana, 1852 Malm in Steenstrup and Liitken, 1861 Kollar in Kroyer, 1863 HeUer, 1865 Gissler, 1883 Bassett-Smith, i896a, b Bassett-Smith, i8g6b Willey, 1896 Bassett-Smith, i8g8a Bassett-Smith, i8g8a Bassett-Smith, iSgSa Transferred by Smith, 1874 Wilson, 1905 Baird, 1850 Bassett-Smith, 1899 Baird, 1850 Baird, 1850 Carus, 1885 Dana, 1853 Bassett-Smith, 1899 Kroyer, 1863-1864 Yamaguti, 1963 Wilson, 1905 A. Scott, 1901 Bassett-Smith, 1899 Stebbings, 1900 Bere, 1936 Wilson, 1912 Rangnekar and Murti, 1964 1 1 have been unable to locate this reference. TABLE II List of species names junior in synonomy within the genus C aligns (prior to 1899) C aligns sp. mulleri n. sp. piscinus n. comb. bicuspidatus n. sp. americanus n. sp. minutus n. nov. leptochilus n. sp. appendiculatus n. sp. elegans n. sp. coryphaenae n. sp. abbreviatus n. sp. aeglefini n. sp. carangis n. sp. fallax n. sp. lumpi n. sp. monacanthi n. sp. nanus n. sp. borealis n. sp. bengoensis n. sp. scomberi n. sp. longicaudus n. sp. Leach, 1816 Lamarck, 1818 (in part) Nordmann, 1832 Pickering and Dana, 1838 Kr0yer, 1837 Frey and Leuckart, 1847 F. Miffler 1852 Van Beneden, 1851 Steenstrup and Liitken, 1861 Kr0yer, 1863-1864 Kr0yer, 1863-1864 Kr0yer, 1863-1864 Kr0yer, 1863-1864 Kr0yer, 1863-1864 Kr0yer, 1863-1864 Kroyer, 1863-1864 Olsson, 1877 T. Scott, 1894 Bassett-Smith, i8g6b Bassett-Smith, i8g8a Authority Kr0yer, 1837-1839 Kr0yer, 1837-1839 Nordmann, 1832 Steenstrup and Liitken, 1861 HeUer, 1866 Steenstrup and Liitken, 1861 (see text) Steenstrup and Liitken, 1861 Pillai, 1963 T. Scott, 1905 Wilson, 1905 Wilson, 1905 Dollfus, 1953 Olsson, 1869 Wilson, 1905 Parker, 1965 Markevich, 1956 Bassett-Smith, 1899 Wilson, 1905 Bassett-Smith, 1899 364 R. R. PARKER TABLE III List of Caligus species named prior to 1899 considered nomina nuda or species inquirenda Caligus sp. scutatus Milne-Edwards, iS^o1 cristata Gould, 1841 gayi Gay, 1849 gracilis Dana, 1852 scrombri J. V. Thompson in W. Thompson, 1856 platessae Van Beneden, 1871 lepidopi Richiardi, 1880 petersii Richiardi, 1880 serrani Richiardi, 1880 smaris Richiardi, 1880 trachini Richiardi, 1880 trachuri Richiardi, 1880 fissus Richiardi, 1880 1 The type material is not at the Museum National D'Histoire Naturelle, Paris. TABLE IV List of Caligus species named prior to 1899 characterized by absence of 3 ciliated spines on medial margin, distal segment of ist thoracopod (not included in Tables I-III) Caligus sp. Spines productus Dana, 1852 absent alalongae Kr0yer, 1863-1864 absent trichiuri Kr0yer, 1863-1864 present but naked haemulonis Kr0yer, 1863-1864 absent affinis Heller, 1866 absent dakari Van Beneden, 1892 absent CALIGUS LONGICA UDATUS 365 TABLE V List of Caligus species named prior to 1899 characterized by 4th thoracopods composed of 4 segments (not included in Tables I-IV) Caligus sp. diaphanus Nordmann, 1832 thymni Dana, 1852 isonyx Steenstrup and Liitken, 1861 chorinemi Kollar in Kr0yer, 1863-1864 pelamydis Kr0yer, 1863-1864 constrictus Heller, 1865 tenax Heller, 1865 torpedinis Heller, 1865* trachynoti Heller, 1865 vexator Heller, 1865 murrayanus T. Scott, 1894 arii Bassett-Smith, i8g8b cybii Bassett-Smith, i8g8a phipsoni Bassett-Smith, i8g8a platytarsis Bassett-Smith, i8g8b robustus Bassett-Smith, 18980 1 Brian, 1924, considered C. torpedinis a possible synonym of C. diaphanus Nordmann. TABLE VI List of Caligus species named prior to 1899 characterized by 4th thoracopod composed of 3 segments, with a spine present on lateral edge of the distal segment in addition to 3 terminal spines (not included in Tables I-IV) Caligus sp. elongatus Nordmann, I8321 kroyerii Milne-Edwards, 18402 rapax Milne-Edwards, i84O2 rissoanus Milne-Edwards, 18402 angustatus Kr0yer, 1863-1864 belones Kr0yer, 1863-1864 gurnardi Kr0yer, 1863-1864 stromatei Krayer, 1863-1864 trachypteri Kr0yer, 1863-1864 infestans Heller, 1865 irritans Heller, 1865 dubius T. Scott, 1894 cossackii Bassett-Smith, i8g8b 1 Type material was obtained from Institut fur Spezielle Zoologie und Zoologisches Museum, Berlin and examined by the author. This loan was arranged through the kindness of Dr. H. E. Gruner, Humboldt-Universitat zu Berlin. 2 Type material was obtained from the Paris Museum and examined by the author. This loan was arranged through the kindness of Dr. J. Forest, Museum National D'Histoire Naturelle, Paris. 366 R. R. PARKER TABLE VII List of Caligus species named prior to 1899 of which the male is characterized by an abdomen composed of a single segment (not included in Tables I-VI) Caligus sp. curtus O. Miiller, 1785 minimus Otto, 1821 centrodonti Baird, 1850 balistae Steenstrup and Liitken, 1861 cheilodactyli Kr0yer, 1863-1864 hyalinus Chernyavsky, 1868 REFERENCES ABILDGAARD, P. C. 1794. Beskrivelse af tvende nye insecter henhbrende under den Linnriske Slagt Monoculus, og den Miillerske Slagt Caligus. Skr. Naturh.-Selsk. Kj0benhavn, 3 (2) : 46-54. BAIRD, W. 1848. Description of the Caligus stromii. Ann. Mag. nat. Hist., N.S., 5 : 396-397. 1850. The natural history of the British Entomostraca. 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U.S. natn. Mus., 42 : 233-243. YAMAGUTI, S. 1963. Parasitic Copepoda and Branchiura of Fishes. Interscience Pub., 1104 pp. INDEX TO VOLUME 15 The page numbers of the principal references and the new taxonomic names are printed in bold type. Abalistes Acanthogobius Acanthuridae Acasta 146 143 140 229 acidens, Haplochromis 46, 70, 72, 78-77, IOQ, IIO, 112, 114, 115, 117, Adeonellopsis .... 340, 343 aegyptiaca, Nardoa . . . .169 aestuarii, Chthamalus . 229, 230, 231, 232, 251, 252, 257-26i, 309, 310, 312,314 afifinis, Stellicola . 201, 206-2og, 210, 211, 212, 220, 222-224, Pls- 4~6 Africana . . . 4-8, 11-13, 22> 24 africana, Heterakis . . . .22 africana, Hippopetraliella . 345-346, PI. i agassizii, Membraniporella . . .329 Agassizii, Nardoa . . . . .172 agassizii Smitt, Membranipora . . 329 ajax, Tamaria . . . 169, 194, 195 akaara, Epinephelus . . 136-137, PI. 2 albicostatus, Balanus amphitrite 231, 277, 312 albopunctata, Teuthis . . . .140 albo-rubrum, Holocentrum . . 132, 151 alcicornis, Cribrilina .... 330 Alectis ....... 134 Aleuteres . . . . . .147 altigenis, Haplochromis 59, 60-65, 68, 92, 109-115 Alutera ...... 147 alvarengai, Moaciria . .' . .22 Ambassidae . . . . . .136 Ambassis . . . . . .136 Amblyopus . . . . . .145 Amphacanthus . . . . .140 Amphiprion . . . . . .142 amphitrite, Balanus . . 271, 281, 286 amphitrite, Balanus amphitrite 230, 231, 271- 277. 3°9, 312, 313 amphitrite albicostatus, Balanus 231, 277, 312 amphitrite amphitrite, Balanus . 230, 231 271-277, 309, 312, 313 amphitrite communis, Balanus 271-276, 313 amphitrite denticulata, Balanus 230, 271, 272, 273, 274, 275, 276, 313 amphitrite hawaiiensis, Balanus 230, 271, 272, 273, 274, 275, 276, 313 amphitrite pallidus, Balanus 271, 277, 278, 281 amphitrite stutsburi, Balanus 203, 271, 277, 278 amphitrite venustus, Balanus . 271, 280, 281 Anarthropora .... 340, 341 anatifera, Lepas . . . .231, 237 andamanensis, Fromia . . . 188, 189 anglicum, Pyrgoma 232, 294-296, 307, 308, 312 Anguilla . . . 124, 129, 149-150 anguillaris, Amblyopus .... 145 anguillaris, Plotosus .... 131 anguillaris, Taenioides .... 145 Anguillidae . . . . . .129 angustus, Balanus perforatus . . 231, 269 annularis, Mesoprion .... 137 anomala, Psenopsis . . . 134, PI. i anserifera, Lepas . . 231, 237-238, 239 Ansiruptodera . . . 4, 6, 7, 8, 23, 25 ansiruptodera, Aspidodera . . .23 antarctica, Membraniporella . . . 329 antiguae, Dichelaspis 242, 247, 249, 250, 313 antiguae, Octolasmis 242, 245, 246, 247-25O, 313 antillea, Exechonella . 337-339, 340, PI. i antillea, Lepralia ..... 337 Apistes. ...... 140 Apistus ...... 141 Aploactis ..... 124, 141 apogonoides, Haplochromis 105-io8, 117 Apolectis ...... 153 Apocryptes ..... 144, PI. 3 aragoi, Flustra .... 329, 331 aragoi, Membraniporella 329, 331, 332, 349, PI. i aragoi pacifica, Membraniporella . . 329 arborescens, Membranipora . . -337 areolata, Lepralia ..... 347 argenteus, Haplochromis 81, 82, 84-87, 109, III, 112, 113 argenteus, Otolithes .... 139 argenteus, Otolithus . . . 139 argenteus, Pomadasys .... 137 argus, Ophicephalus .... 132 argus, Scatophagus .... 140 armata, Citula .... 135, 154 armata, Fromia . . . . .191 armatus, Caranx .... 135, 154 armatus, Trichiurus . . . 133, 152 Ascaridia . . 4, 5, 6, 7, 13, 14, 17, 24 Ascaridiidae . . . . -9. io> 24 asper, Uranoscopus . . . .146 Aspidodera . . . . 4, 6, 7, 8, 23, 25 37° INDEX Aspidoderidae . . . 8-9, 10, 23 Aspidoderinae ..... 23 assimilis, Balanus improvisus . 231, 270-2yi Astatotilapia . . . . . 73, 117 Asterias . . . . . .172 asterinae, Stellicola .... 205 Asterope ...... 218 astridae, Ophidiaster . . 169, 195, 196 astropectinis, Stellicola . 205, 221, 223, 224 astylosterni, Africana .... 5 Atherina . . . . . 133 Atherinidae . . . . . 133 atlantida, Cellaria . . . 327-328 atlantica, Ibla 230, 231, 235-237, 309, 312, 313 attenuata, Hacelia . . . .196 Atule 134-135, PI. i aucupatorius, Boleophthalmus . 144 aurantiacum, Pelor . . . 155 aurantiacus, Pseudobragus . . . 130 auritum, Conchoderma . . . 231, 240 aurivillii, Dichelaspis .... 242 azoricus, Balanus tintinnabulum 265-266, 311 Bagridae ..... 130-131 Bagrus ..... 130-131 Balanodytes ...... 306 balansae, Fromia . . . . .191 Balanus 229-232, 263-291, 307, 308, 310-314 Balistes . . . . . 124, 146 Balistidae ..... 146-147 barbarae, Haplochromis 93-97, TI4> JI5> IJ7 bartoni, Haplochromis . 37, 42, 46, 72, 81, 82, 109, 112, 114 baueri, Membraniporella . . . 330 bayoni, Haplochromis 58, 60, 62, 109, in, 112, "3 Bellaplectana ..... 4 Belone ....... 131 Belonidae ...... 131 berda, Chrysophrys . . . .138 biaculeatus, Triacanthus . . 146, 156 biarmata, Triporula . 340, 341, 342, 343 bicinctus, Amphiprion .... 142 bidenticulata, Hiantopora . . 335, 341 bisinuata, Petraliella .... 346 bisinuatus, Chthamalus stellatus 231, 252, 311, 312 bleekeri, Atherina ..... 133 Blepharis . . . . . .134 boddaerti, Boleophthalmus . . . 144 Boleophthalmus . . . 126, 144, 155 boops, Haplochromis . 42, 46, 47-49, 50, 51, 97, 109, no, 116, 117 borneensis, Anguilla . . . .150 boro, Pisodonophis . . . .130 bouvieri, Linckia ..... 195 brasiliensis, Exechonella . . 338, 339 brevicaudata, Strongyluris ... 23 breviceps, Aploactis . . . 124, 141 breviceps, Synanceia . . . .141 brevirostris, Leiognathus . . . 135 brevirostris, Triacanthus . . 146, 156 brownae, Haplochromis 95, 96, 108, no, 113, 114 brunneus, Glossogobius . . . 143, PI. 3 caeruleus, Stellicola 201, 205, 219-221, 222-224, Pis. 15-17 calceolus, Balanus . . 290-291, 309, 312 calceolus, Conopea . . 232, 290-291 Caligus ..... 353-368 Callionymidae .... 145-146 Callionymus ..... 145-146 Callopora . . . 330, 331, 332 campylostomus, Boleophthalmus 126, 144, 155 cancrivorus, Ophichthys . . .130 cancroides, Caranx . . . 134, PI. i Carangidae ..... 134-135 Caranx . . . 134-135, 154, PI. i Carcharhinidae . . . . .128 Carcharhinus . . . . .128 Carcharias . . . . . .128 caretta, Chelonibia . . . 232, 297 carinata, Certonardoa . . . .186 carinata, Nardoa . . . . .186 carinifera, Asterope . . . .218 castanea, Utinga ..... 346 caudimaculata, Belone . . . .131 cavifrons, Haplochromis. 41, 55, 109, 113, 114 Celerina .... 193-194, PI. 6 Cellaria .... 324-328, 348 Cellularia ...... 325, 326 cephalotus, Mugil . . . . 133 cephalus, Mugil ..... 133 Certonardoa . . . . . .186 Channa ...... 132 Channidae . . . . . .132 Chatoesus . . . . . 128, PI. i Cheilopora . . . . . .341 Chelonibia 229, 232, 296-300, 309, 310, 312, 314 Cheloniketerakis ..... 4 chelonophilum, Conchoderma virgatum . 231, 240-241 Chetia 38 Chiloscyllium . . . . .128 chinensis, Leucosoma . . . .129 chinensis, Salanx . . . . .129 chinensis, Sphyraena . . . 133 Choriaster ..... 201, 218 chromogynos, Haplochromis . . 66, 94, 96 chrysargyrus, Amphiprion . . . 142 chrysogaster, Amphiprion . . .142 Chrysophrys . . . . . .138 Chthamalus . 229, 230, 232, 251-262, 307-314 ciliaris, Alectis ..... 134 ciliaris, Caranx .... 135, 154 cinereus, Haplochromis . . . .96 cinereus, Muraenesox .... 129 cirratus, Chthamalus . . . 230, 257 Citula . . .... 135, 154 clarkii, Amphiprion . . . .142 INDEX 371 clathrata, Anguilla clausi, Stellicola clavigera, Mithrodia Clupanodon . Clupea .... Clupeidae clupeoides, Coilia . coccinella, Adeonellopsis . Coilia .... Colletosia Collichthys . commersonii, Ambassis . 124, 129, 149 205, 221, 223, 224 209 . 128, PL i . 128 . 128 129 • 343 129, 149 • 333 • 139 136 communis, Balanus amphitrite 271, 272, 273, 274. 275, 276, 313 complanata, Coronula . . . 232, 300 Conchoderma . . 231, 240-241, 312 concinnus, Balanus tintinnabulum 231, 265, 266, 311 concinnus, Megabalanus tintinnabulum . 231 Congridae . . . . . .130 Conopea .... 232, 290-291 corbicula, Membraniporella . . . 330 Coronula ..... 232, 300 cornuta, Heteralepas 231, 239, 240, 307, 310, 312 coronatus, Cryptophialus . . 232, 301 crassicosta, Membraniporella . . 330 crassum, Poecilasma . . 231, 241, 312 crassum, Trilasmis . . 231, 241, 312 crenatibasis, Chelonibia manati . .297 Cribrilaria . . 332, 333~334- 335. 342 Cribrilina . . 330, 332, 334, 335, 343 cribrosus, Retaster .... 206 crispatus, Balanus tintinnabulum . . 265 crocodilus, Thysanophrys . . . 142 cronus, Haplochromis . . 96, 108, 115 cruciata, Tetractis . . . .171 Cryptocentrus . . . . .143 cryptodon, Haplochromis . 96, 114, 115 Cryptophialus .... 232, 301 crypturi, Heterakis .... 20 Ctenochromis . . . . .87 Ctenogobius . . . . . 143 Culcita . . . . .201, 208 cumingi, Ferdina ..... 192 cumingi, Ibla .... 236, 237 curticaudatus, Stellicola . 205, 222-223 curvicornis, Callionymus . . . 146 cuvieri, Pelor . . . . 155 Cyclicopora ...... 340 Cymodocea ...... 218 Cynoglossidae . . . . .132 Cynoglossus . . . . . .132 Dendrophysa . . . . 139 dentatus, Chthamalus 229, 231, 251, 252-257, 309, 3i°. 312, 314 dentex, Haplochromis . 58, 59, 60, 109, 112 denticulata, Balanus amphitrite 230, 271, 272, 273, 274, 275, 276, 313 depressus, Chthamalus stellatus 230, 251, 257 diacanthus, Serranus . . . .136 Dichelaspis . . . 242, 247, 249, 313 dichrourus, Haplochromis 64, 65-6g, 92, 109, 116 discoidea, Exechonella .... 337 discoporellae, Kochlorinopsis 232, 302-306, 313. PI- i distans, Membraniporella . . 329, 332 distoma, Adeonellopsis .... 340 distoma, Lepralia ..... 340 divisa, Tetraclita . 232, 291-293, 294, 312, 314 dollfusi, Balanus .... 230, 268 dorsipora, Hippopetraliella . . . 346 Drepane ...... 140 Drombus ...... 143 dussumieri, Platyglossus . . . . 142 dybowskii, Balanus . . 230, 277, 278 eburneus, Balanus . . 231, 270, 311 Echeneidae . . . . . .147 Echeneis ..... 147, 157 egeriae, Gomophia egyptiaca 169-171, 177-179, i8i.Pl. i egyptiaca, Gomophia 169, 172-179, 181, 192 egyptiaca, Gomophia egyptiaca 170-171, PL i egyptiaca egeriae, Gomophia 178, 179, 169-171, 181, PI. i egyptiaca egyptiaca, Gomophoria 170, 171, PL i ehrenbergii, Platax .... 140 elegans, Fromia . . 169, 188, 189, 191 elegans, Stomatolepas . . 232, 300-3OI elongata, Cellaria .... 326, 328 elongatoides, Cellaria . 325, 326-327, 328 elongatus, Polycaulus .... 141 empodisma, Haplochromis . . 108, 115 Enantiosula . . . 340, 341, 343, 344 Engraulidae . . . . . .129 Ephippidae ..... 139-140 Epinephelus .... 136-137, PL 2 Equula. ...... 135 equula, Genres ..... 136 erythropterus, Lutjanus . . . 137 erythrourus, Gerres .... 136 Eschara ...... 340 Escharipora ..... 340, 343 estor, Haplochromis . 59, 60, 44, 65, 109, 112 Eupleurogrammus . . . .152 Exechonella . 334, 335, 337-34°. 349, Pis. i, 2 exornatus, Julis .... 142, PL 3 fallax, Balanus . 230, 232, 287-290, 307, 308, 312, 314 fallax, Hesperibalanus .... 232 faouzii, Nardoa . . 169, 183, 186, 187 fario, Epinephelus ..... 136 fasciatomaculatus, Epinephelus . . 136 fasciato-punctatus, Gobius . . . 142 fasciatus, Blepharis . . . .134 fascicularis, Lepas . . . .231, 239 femineus, Stellicola 201, 213-215, 216-220, 222-224, PI- 8-1 1 372 INDEX Fenestrulina . . . . . .329 Ferdina . . 169, 189, 191-193, PI. 6 ferox, Hiantopora ..... 341 filifer, Cryptocentrus .... 143 Gobius ...... 143 fistulosa, Cellaria . . . 324, 325, 328 fistulosa, Cellularia .... 325 fistulosus, Balanus perforatus . . 231, 269 flavescens, Ferdina . 169, 191-192, PI. 6 flavipinnis, Haplochromis 73, 109, 113, 114 flaviventris, Chetia . . . .38 Flustra. ..... 329, 331 forceps, Cellaria . . . . .327 Formio. ...... 153 Formionidae . . . . .134 fragilis, Chthanalus . 231, 254, 262, 311 fragilis, Chthamalus stellatus . . .251 frequens, Stellicola . 205, 221, 223, 224 frianti, Nardoa 169, 171, 175-180, 187, PI. i Fromia. . . 169, 183, 188-193, PI- 6 Fugu 147, 157 fulvidraco, Pelteobagrus . . . 131 fuscescens, Teuthis .... 140 galatheae, Nardoa . . . 183, 184, 188 Gallichthys . . . . . 134 gallinarum, Ascaris . . . 19, 20 gallus, Caranx . . . . .134 gamophia, Nardoa . . . .172 Ganguleterakis ..... 4 Genres ....... 136 Gerridae ...... 136 ghardaqana, Fromia . . . .191 gigantea, Exechonella . 339-340, PI. 2 giganteum, Pachylasma . . 231, 263, 312 girardi, Gireterakis . . . .21 Gireterakis .... 4-8, n, 12, 25 giuris, Glossogobius .... 142 giuris, Gobius . . . . .142 globicipitis, Xenobalanus . . 232, 301 Glossogobius .... 142-143, PI. 3 Gobiidae ..... 142-144 Gobioididae . . . . . .145 Gobius . . . 142-143, 156, PI. 3 Gomophia . 169-179, 181, 192, 193, PI. i gomophia, Nardoa. . 169, 172-173, 175, 177, 179, 180, 182, 187, Pis. 2, 5 gomophia, Scytaster . . . 172, 1 80 gowersi, Haplochromis 37, 59, 64, 65, 109, in, 112 gracilis, Stellicola 205, 208, 209, 212, 223, 224 grammica, Plagiusa . . . .132 grammicus, Cynoglossus .... 132 grandis, Cyclicopora .... 340 granti, Haplochromis . . . 107, 108 granulatus, Choriaster . . . 201, 218 guiarti, Haplochromis . -35. 59. 60, 70, 71, 72, 75, 77, 81, 87, 89, 92, 95, 96, 109-114 gulo, Metrarabdotos .... 348 gulo, Trigonopora ..... 348 guttata, Inegocia . guttatus, Platycephalus . Hacelia hadracantha, Fromia Halichoeres . hamata, Kochlorine Haplochromis harancha, Ophisurus hasta, Chrysophrys Pristipoma Sparus . 142 . 142 169, 195, 196 . 191 • 142, PL 3 232, 301-302, 306 29-iig . 130 . 138 • 137 138 hawaiiensis, Balanus amphitrite 230, 271-276, 313 heffernani, Clerina . . 193-194, PI- 6 heffernani, Ferdina . . 169, 191-194 helichostichus, Ophidiaster . 169, 195-ig6 helicostichus, Hacelia . . . 195, 196 hemiopla, Fromia . . . . .191 hemprichi, Ophidiaster .... 196 hermannianus, Amblyopus . . .145 Hesperibalanus .... 232, 287 Heterakidae ..... 9, 10, 18 Heterakinae . . . . . .18 Heterakis . 4, 6, 8, n, 13, 14, 18, 19, 24 Heterakoidea ..... 1-28 Heterakoides . . . . . 4, 5 Heteralepas . 231, 239, 240, 307, 310, 312 Hexactis ...... 171 hexastylos, Platylepas . . . 232, 300 Hiantopora ....... 335, 341 hillii, Lepas . . . . .231, 238 hindsii, Callionymus .... 145 Hippopetraliella . 334, 345-346, 349, PI- i histophorus, Scartelaos . . 144-145, 155 hoeki, Dichelaspis . . . 242, 249, 313 hoeki, Octolasmis 231, 242-247, 248, 249, 250, 3", 313 Holocentridae . . . . .132 Holocentrum .... 132, 151 Holocentrus ..... 132, 151 holothuriae, Stellicola . . 205, 221-224 Hoplotilapia ...... 95 horrida, Anarthropora . . . .341 Ibla . . 230, 231, 235-237, 309, 312, 313 idiotica, Ibla .... 236, 237 imperfectum, Scalpellum . 231, 234, 311 improvisus, Balanus . . . 231, 270 improvisus assimilis, Balanus . . 270-271 inarmatus, Hacelia .... 196 indica, Alectis . . . . .- . 134 indica, Fromia . . 169, 188-189, 191 indicum, Chiloscyllium . . . .128 indicus, Platycephalus .... 141 Pseudupeneus ..... 138 Upeneus . . . . . .138 indicus, Scytaster . . • . . .188 Inegocia ...... 142 inermis, Kochlorine . . . 232, 302 Inimicus .... 141, 154, PI. 3 INDEX 373 insidiator, Platycephalus . . . 141 interlabiata, Heterakis .... 20 intermedia, Lauroia . . . .10 intermedia, Membraniporella nitida . 330 intermedius, Trichiurus . . . 133, 152 isolonche, Heterakis . . . .20 japonica, Anguilla . . . . 129, 149 Sillago 139 japonicum, Pelor .... 141, 154 japonicus, Callionymus . . . 145-146 Inimicus .... 141, 154, PI. 3 Mugil . . .133 Priacanthus . . . . .137 Uranosiopus . . . . .146 jarbug, Therapon . . . . .138 jnanendrae, Pseudaspidodera ... 20 johnsoni, Cellaria ..... 325 johnsoni, Cellularia . . . .326 Julis 142, PI. 3 kaempferi, Poecilasma . . 231, 241, 309 kaempferi, Trilasmis . . 231, 241, 312 kalla, Caranx . . . 134-135, PI. i knutteli, Gobius . . . . 143 Kochlovine . . . 232, 301-302, 313 Kochlorinopsis . 232, 302-306, 313, PI. i kokius, Gobius ..... 143 kossmanni, Stellicola . . 201, 202-2O5, 206-207, 210-212, 220-224, Pis. 1-4 kiiensis, Schizoporella .... 335 labiosa, Cheilopora .... 341 Labridae . . . . . .142 lacrimosus, Haplochromis ... 96 laevigata, Linckia .... 183, 216 laevis, Aleuteres ..... 147 Lagenipora . . . . . -337 lamprogenys, Haplochromis . . 78, 83 lankensis, Stellicola 205, 209, 212, 222-223 latus, Sparus. . . . . .138 Lauroia . . 4-7, 10, 13, 23, 24, 25 Lauroiinae ..... .23-24 lautus, Stellicola . . 205, 221, 223, 224 leachi, Leiaster . . . . .213 Leiaster ..... 213, 215 Leiognathidae . . . . 135 Leiognathus . . . . . 135 lemonnieri, Nardoa . 169, 182-186, 188 Lepas .... 231, 237-239, 312 Lepralia . 329, 330, 335, 337, 340, 347 Lepturacanthus . . . . 1 33, 152 lepturus, Congrus .... 124, 130 Trichiurus .... 133, 152 Uroconger . . . . . .130 Leucosoma . . . . . .129 lichenoides, Eschara . . . .340 Lichomolgus ..... 205 limbatus, Bagrus . . . . .131 lincki, Protoreaster . 201, 202, 218, 219 Linckia 172, 181, 183, 192, 194, 195, 216, 219 Linckiomolgus . . . . .219 lineatus, Plotosus . . . . .131 Lithoglyptes . . . . .306 lobatibasis, Chelonibia manati . . . 297 longecaudatus, Callionymus . . . 145 longicaudatus, Caligus . . . 353-368 longicaudatus, Stellicola . . 205, 222, 223 longimana, Drepane . . . .140 longirostris, Haplochromis 81, 85-87, 109, 112- 114 longirostris, Paratilapia . . . 60, 84 longirostrum, Smilium . 229, 231, 234, 311 longiseta, Stellicola 201, 209-213, 214, 220, 222-224, Pls- 6-8 lowei, Octolasmis . 231, 241-242, 309, 311, 312 lucida, Collichthys . . . . 139 Sciaena ...... 139 lucidus, Collichthys . . . .139 Luidia . . . . . . 206 luidiae, Stellicola . . 205, 221, 223, 224 Lutjanidae ...... 137 Lutjanus ...... 137 macrodon, Haplochromis . 78, 80, 83 macrognathus, Haplochromis . 55, 56, 77, 109, III, 112, 113 macrolepidotus, Mugil . . . 133 macrops, Haplochromis . . . .96 Maculaster ...... 206 maculata, Channa ..... 132 maculatus, Chatoessus . . . 128, PI. I Ophicephalus . . . . .132 Ophiocephalus . . . . .132 maculata, Luidia ..... 206 maculata, Maculaster maculata . . 206 maculata maculata, Maculaster . . 206 maculipinna, Haplochromis 37, 42, 43-47, 49> 51, 76, 109, no maculipinna, Paratilapia ... 43 magna, Exechonella . . . -337 magnifica, Membraniporella . . 329, 332 magniporosa, Phylactella . . . 341 major, Chrysophyrs .... 138 Gallichthys 134 Pagrus ...... 138 malabaricus, Caranx .... 135 Epinephelus . . . . 136 mammillatus, Pentaceraster . . 201, 218 mamillifera, Nardoa 169, 175, 177, 179, 180, 187 manati, Chelonibia 229, 232, 297, 309, 310, 312 manati crenatibasis, Chelonibia . . 297 manati lobatibasis, Chelonibia. . . 297 mandibularis, Haplochromis . 72, 81, 82, 109, 112, 113 manica, Enantiosula 340, 341, 342, 343, 344, 345 manica, Triporula ... . 344 marcusi, Membraniporella 331-333, 335, 349, PI- i margaritiferus, Amphacanthus . . 140 374 INDEX margariturus, Gobius Rhinogobius marginata, Petraliella marmorata, Anguilla marmoratus, Sebastes Sebasticus 143 143 346 150 141 141 maroccana, Balanus tintinnabulum 231, 266, 312 maroccana, Megabalanus tintinnabulum . 231 martini, Haplochromis . 65, 89, 91, 92, 113 maxillaris, Balanus 231, 267, 309, 310, 312 maxillaris, Haplochromis . . .115 maxillaris, Megabalanus . . .231 Megabalanus . . . . .231 megaloplax, Linckia ... . . 194 melanopterus, Carcharhinus . . .128 Carcharias . . . . . .128 (Prionodon) . . . . .128 melo, Trachinotus .... 134, PL i Membranipora .... 337, 329 Membraniporella . 328-333, 335, 349, PL i Meniscopora . . . . . .341 mento, Haplochromis 58, 59, 60, 62, 75, 77, 81, 109, in, 112 Mesoprion . . . . . 137 Meteterakinae . . . . .21 Meteterakis . . . 4-8, n, 12, 21, 25 Metrarabdotos . 334, 346-348, 349, PL i michaeli, Haplochromis 65, 89, 91, 92, 108, 113 Microbuglossus . . . . .131 microdon, Haplochromis . 96, 114, 115 milleporella, Fromia . . . 191, 193 Minons ...... 141 minons, Apistus . . . . .141 minuta, Paralepas . 231, 240, 307, 310, 312 Mitella . 231, 232-233, 307, 308, 312 Mithrodia .... 209, 210, 212 Moaciria . . . 4-7, 11-13, 22, 25 mollis, Nardoa . . 169, 183-186, 188 moluca, Pempheris .... 137 Monacanthus . . . . .147 monilis, Fromia .... 191, 193 monodactylus, Minous .... 141 monodon, Anarthropora . . . .340 mucronata, Escharipora . . . 340 Mugil ....... 133 Mugilidae ...... 133 Mugiloididae ..... 146 Mullidae ...... 138 Mullus ....... 138 Muraenesocidae . . . . .129 Muraenesox . . . . . .129 muticus, Trichiurus . . . 133, 152 mystus, Coilia . . . . .129 nageb, Pristipoma . . . . .137 Nardoa . . . 169-igS, Pis. 1-5 naucrates, Echeneis . . . 147, 157 nebulosa, Percis ..... 146 Neoferdina ..... 191, 192 niger, Parastromateus . . . 134, 153 Stromateus nigrescens, Astatotilapia . nigrescens, Balarus nigrescens, Halichoeres . nigrescens, Haplochromis nigrescens, Megabalanus nitida, Lepralia nitida, Membraniporella • 134 73. "7 231, 266-267, 311 . 142, PL 3 73, 77, 114 . 231 • 329 329, 330-331, 332 nitida intermedia, Membraniporella . 330 niveus, Balanus venustus 232, 286-287, 311 nodosa, Cellularia ..... 326 nodosa, Fromia . . . 189-I9I, PL 6 nodosa, Linckia . . . . 195 nodosus, Protoreaster . . . .218 Nomeidae . . . . . .134 novaecaledoniae, Nardoa 172, 173, 183-186, 188, PI- 5 normani, Cellaria salicornioides 325, 326, 327, 328 nuchalis, Equula . . . . 135 nyanzae, Haplochromis 37, 42, 46, 77, 82, 109 nysti, Trigonopora . . . -348 obesus, Haplochromis . . . 96, 115 obtusa, Nardoa . . 169, 176, 180, 181 obtusata, Sphyraena . . . 133 obtusus, Scytaster .... PL 4 occidentalis, Balanus . . 230, 287, 289 ocellatus, Fuga (Torafuga) . . 147, 157 Tetrodon ..... 147, 157 Octolasmis . 231, 241-250, 309, 311, 312, 313 Odontoterakis . 4, 6, 8, 11-14, J8» 20, 24 offreti, Ferdina . . . . 189, 191 offreti, Neoferdina . . . .191 ommatura, Solea .... 132, PL i Ophicephalus . . . . .132 Ophichthidae . . . . .130 Ophidiaster . 169, 175, 177, 181, 186, 194, 195-196 Ophisurus . . . . . .130 opulenta, Temachia .... 335 orbicularis, Platax .... 139-I4O oreastriphilus, Stellicola 201, 205, 218-219, 220, 222-224, PL 14 Orectolobidae . . . . .128 orthostoma, Haplochromis 100-102, 103, 105, 114 otaitensis, Pempheris .... 137 Otolithes . . . . . -139 Otolithus . . . . . -139 otto-mulleriana, Lepralia . . . 335 ovata, Solea ...... 131 ovatus, Microbuglossus .... 131 oxyrhynchus, Therapon . . . .138 oyena, Genres ..... 136 pachycephalus, Haplochromis . 39-42, 46, 51, 97, 109, no, 116, 117 Pachylasma .... 231, 263, 312 pacifica, Fromia . . V . .191 pacifica, Membraniporella aragoi . . 329 INDEX 375 Pagrus ...... 138 pallens, Ophichthys . . . .130 pallidus, Balanus 229, 230, 231, 271, 277-aSo, 281, 286, 309, 310, 312, 313 pallidus, Balanus amphitrite 271, 277, 278, 281 pallidus, Balanus pallidus . . 277, 313 pallidus pallidus, Balanus . . 277, 313 pallidus stutsburi, Balanus 230, 277, 281, 283, 313 paraguiarti, Haplochromis . 69-72, 76, 77, 81, 109, no, in, 112, 113, 117 Paralepas . . 231, 240, 307, 310, 312 Parapercis . . . . . .146 Paraspidodera . . 4, 6-8, 13, 24, 25 Parastromateus . . . . 134, 153 Paratilapia . 32, 43, 51, 60, 65, 73, 78, 84, 87 Pareterakis . . . . . • 4, 5 parorthostoma, Haplochromis 102, 108-105, 114, 117 parvidens, Haplochromis . 77, 114, 115 parvidens, Paratilapia ... 73, 76 patula, Chelonibia . 232, 297-3OO, 314 pauciforis, Nardoa . . 182, 183, 186, 188 pavonis, Pseudaspidodera ... 20 pectinata, Lepas . . . 231, 238, 239 pectinirostris, Boleophthalmus . . 144 pectoralis, Ctenochromis . ... 87 pectoralis, Paratilapia .... 87 pellegrini, Haplochromis 56, 59, 60, 68, 109, 112, 114 Pelmatochromis . . 32,38, 60,78,100 Pelor . . . .141, 154, 155, PL 3 Pelteobagrus . . . . -131 Pempheridae. . . . . .137 Pempheris . . . . . .137 Pentaceraster . . . . 201, 218 Pentadactylosaster . . . .171 Percis ....... 146 percoides, Haplochromis 73, 109, 113, 114 perforatus, Balanus 231, 268-269, 307, 308, 312, 313. 3H perforatus angustus, Balanus . . 231, 269 perforatus fistulosus, Balanus . . 231, 269 Periophthalmidae .... 144-145 perplexa, Tremogasterina . . 336-337 perrieri, Nardoa . . . . .172 petasus, Membraniporella . . . 329 Petraliella . . . . . 346 pharyngeodentata, Aplectana ... 4 Phylactella 341 phytophagus, Haplochromis ... 77 Pisodonophis . . . . .130 plagiostoma, Haplochromis . 55, 56, 113, 114 plagiosum, Chiloscyllium . . . 128 Plagiusa ...... 132 plana, Triporula . . . . .341 Platax .... 134, 139-140, 153 Platycephalidae .... 141-142 Platycephalus . . . . 141-142 platycephalus, Gobius . . . 143, PI. 3 Platyglossus . . . . . .142 Platylepas ..... 232, 300 playfairii, Coilia . . . . 129, 149 pleurobranchi, Stellicola . 205, 221-223 Plotosidae . . . . . .131 Plotosus ...... 131 Poecilasma . . . 231, 241, 309, 312 pollex, Stellicola . 201, 216-2i8, 220-224, Pis. 11-14 pollicipes, Mitella 231, 232-233, 307, 308, 312 Polycaulus . . . . . .141 Pomacentridae . . . . .142 Pomadasyidae . . . . .137 Pomadasys . . . . . 137 Poraster ..... 201, 218 praegustator, Stomatolepas . . . 301 prelucidioides, Cheilopora . . . 341 Preterakis . . . . . . 4, 5 Priacanthidae . . . . 137 Pricanthus 137, PI. 2 Prionodon . . . . . .128 Pristipoma . . . . . 137 prodromus, Haplochromis . . . 107 prognatha, Paratilapia . . . 43, 51, 78 prognathus, Haplochromis . 42, 58, 60, 70, 72, 73, 77, 78-83, 85, 87, 109-115 propetumescens, Tamaria . . 194, 195 Prosopodasys . . . . .140 Protoreaster . . . 201, 202, 218, 219 Psenopsis . . . . . 134, PI. i Pseudaspidodera . 4, 6, 8, 11-14, J8> 20 Pseudaspidoderina . . . . 4, 24 Pseudaspidoderoides . . . . 4, 5 Pseudobagrus . . . . 130, 131 pseudopellegrini, Haplochromis 56-6o, 62, 64 65, 109, no, in, 112, 117 Pseudupeneus . . . . .138 pulchella, Parapercis .... 146 Percis ...... 146 pulchra, Membraniporella . . 329, 333 pumicosa, Exechonella . . . 337, 339 punctata, Cribrilina . 332, 334, 335, 343 punctata, Drepane. . . . .140 purpurascens, Tetractita 232, 291, 293-294, 312 pygmaea, Ibla ..... 236 Pyrgoma . . 232, 294-296, 307, 312 quadrivalvis, Ibla ..... 236 quadrilineatus, Therapon . . . 138 radiata, Colletosia ..... 333 radiata, Cribilaria . 332, 333~334. 335. 342 Raillietakis ...... 4 rebeli, Haplochromis ... 78, 83 Reeves, J. 121-165, Pis. 1-3 reevesii, Callionymus .... 145 Leucosoma . . • • .129 renei, Scalpellum ..... 233 renei, Smilium 229, 230, 231, 232, 233, 309, 310, 312 Retaster 206 376 INDEX retrodens, Hoplotilapia .... 95 Rhinogobius . . . . . .143 rhizophorae, Chthamalus 230, 257, 259, 260, 261 Richardson, Sir John . . . .123 ripilepis, Drombus ..... 143 Gobius ...... 143 roberti, Astatotilapia 73 robusta, Tremogasterina . . . 336, 337 rosea, Nardoa 176, 177, 179, 181-183, 187, PI. 5 ruber, Holocentrus . . . 132, 151 rubrum, Holocentrum .... 132 rugosus, Amblyopus .... 145 russelii, Dendrophysa .... 139 Umbrina ...... 139 Upeneus ...... 138 russellii, Umbrina . . . . .139 rylandi, Callopora . . . . 330 Salangidae . . . . . .129 Salanx ....... 129 salicornioides, Cellaria . . 324, 325, 328 salicornioides normani, Cellaria 325, 326, 327, 328 sauvagei, Haplochromis . . . 107, 108 savala, Lepturacanthus . . . 133, 152 Trichiurus . . . . . 133 savignyi, Maculaster .... 206 Scalpellum 229, 230, 213, 233, 234, 307, 308, 311, 312 scalpellum, Scalpellum 229, 231, 234, 307, 308, 312 Scartelaos .... 144, 155, 156 Scatophagidae . . . . .140 Scatophagus . . . . . .140 sceletos, Lepralia ..... 330 Schizoporella ..... 335 schmideliana, Culcita . . . 201, 218 Sciaena ...... 139 Sciaenidae . . . . . .139 scoleciformes, Aspidodera ... 23 Scorpaenidae .... 140—141 scripta, Alutera ..... 147 scriptus, Monacanthus .... 147 scyllia, Triakis . . . . .128 scyllium, Triakis ..... 128 Scyris 134 Scytaster . . . 172, 186, 188, PI. 4 Sebastes .... 140, 141, PI. 2 Sebasticus . . . . . .141 Sebastoides .... 140-141, PI. 2 segmentata, Ibla .... 236, 237 semiregufaris, Scytaster . . . .186 serperaster, Apocryptes . . . 144, PI. 3 Serranidae ..... 135-136 Serranochromis . . . . .32 serranoides, Haplochromis . . 32, 33, 38 Serranus ..... 136, PI. 2 serranus, Haplochromis . . 32, 33, 35-37, 38, 42, 44, 46, 49, 51, 53, 54, 58, 70, 77, 81, 82, 89, 109, no, 113, 114 serranus, Paratilapia . . 32, 65, 73, 87 servus, Therapon . Seserinus Sexansodera . shihpan, Serranus . Sillaginidae . Sillago . sinense, Pelor sinuosa, Cellaria . 138 134. !39, 153 4 . 136, PL 2 • 139 • 139 • 155 . 325, 328 Smilium . 229, 231, 232, 233-234, 309-312 Smitt, Membranipora agassizii . .329 Solea ..... 131, 132, PI. i Soleidae ..... 131—132 Sparidae ...... 138 Sparus ....... 138 spathulata, Tremogasterina . . 336, 341 speciosus, Leiaster . . . 213, 215 spekii, Haplochromis 32-38, 42, 46, 51, 77, 109-111 spekii, Pelmatochromis . 32, 38, 60, 78, 100 Sphaeroides . . . . . .157 sphenisci, Nardoa . 173-175, 177, 178, 179, 181, 182, 187, PL 3 Sphyraena . . . . . -133 Sphyraenidae . . . . -133 Spinaspidodera ..... 4 Spinicauda . . . 4-8, 11-13, 2I> 22> 24 spinicauda, Ascaris . . . .22 Spinicaudimae . . . . .21 spinosa, Pseudaspidodera ... 13 spinosissimus, Holocentrus . . .151 spinosus, Balanus tintinnabulum 231, 265, 311 spinosus, Megabalanus tintinnabulum . 231 spongicola, Balanus 230, 231, 268, 307, 308, 312 spumosa, Heterakis .... 20 squamosa, Tetraclita squamosa 232, 294, 312 squamosa squamosa, Tetraclita 232, 294, 312 squamulatus, Haplochromis 64, 65, 87-93, JI3 squamulosa, Nardoa . . . .186 steindachneri, Haplochromis . . 78, 83 stellata, Escharipora . . . 340, 343 stellata, Triporula 334, 340, 342, 343-345, PL 2 stellatus, Abalistes. .... 146 Balistes . . . . . .146 stellatus, Chthamalus 229, 251,252, 307, 308, 3i2, 313 stellatus, Chthamalus stellatus 231, 251-252, 262, 307 stellatus bisinuatus, Chthamalus . 231, 251, 252, 311, 312 stellatus depressus, Chthamalus 230, 251, 257 stellatus fragilis, Chthamalus . . .251 stellatus stellatus, Chthamalus 231, 251-252, 262, 307 Stellicola . . . « - . 199-225 stigmothonus, Acanthogobius . . . 143 Gobius . ..... 143 Stomatolepas . . . 232, 300-301 striata, Verruca . . . 231, 251, 311 Stromateus ...... 134 INDEX 377 Strongylura . . . . . .131 Strongylura, Strongylura ... 131 Strongylurs . . -4, 6-8, 11-13, 23. 25 strongylurus, Belone . . . -131 stutsburi, Balanus amphitrite . . 230, 271, 277, 278 stutsburi, Balanus pallidus . 230,277,281, 283,313 subplana, Meniscopora . . . .341 subvittatus, Mullus .... 138 Upeneus ...... 138 Upenoides ...... 138 superbus, Poraster . . . 201, 218 Synanceia ...... 141 Synancejidae . . . . .141 Synaptura ...... 132 taeniatus, Haplochromis . . . 78, 80, 83 Taenioides . . . . . .145 Tamaria . . 169, 186, 194-195, 196 tayenus, Priacanthus . . . 137, PI. 2 teira, Platax . . . 134, 139, 153 Temachia ...... 335 testudinaria, Chelonibia . . . 232, 296 Tetraclita . . 232, 291-294, 312, 314 Tetractis ...... 171 Tetraodontidae . . . . 147 Tetrodon ..... 147, 157 Teuchopora . . . . . .341 Teuthis ...... 140 Therapon ...... 138 Theraponidae . . . . .138 theraps, Therapon . . . .138 thorelli, Stellicola . . . 205, 222, 223 thrissa, Clupanodon . . . 128, PI. i Clupea . . . . . .128 thuragnathus, Haplochromis 42, 46, 49-5 1, 109, no, 116, 117 Thysanophrys . . . . .142 tigrinum, Pelor . . . 141, 154, PI. 3 tintinnabulu, Balanus . . 263, 264, 265 tintinnabulum, Balanus tintinnabulum 23 1 , 263-264, 265, 309, 312 tintinnabulum, Megabalanus tintinnabulum 231 tintinnabulum azoricus, Balanus 265-266, 311 tintinnabulum concinnus, Balanus . 231, 265, 266, 311 tintinnabulum concinnus, Megabalanus . 231 tintinnabulum crispatus, Balanus . . 265 tintinnabulum maroccana, Balanus 231, 266, 312 tintinnabulum maroccana, Megabalanus . 231 tintinnabulum spinosus, Balanus 231, 265, 311 tintinnabulum spinosus, Megabalanus . 231 tintinnabulum tintinnabulum, Balanus . 231, 263, 264, 265, 309, 312 tintinnabulum tintinnabulum, Megabalanus 231 tintinnabulum zebra, Balanus 231, 264-265, 309 tintinnabulum zebra, Megabalanus . .231 Torafugu ..... 147, 157 Torquigener . . . . . .157 trachinoides, Apistes Prosopodasys Vespicula . Trachinotus . transversa, Stomatolepas trapezoideum, Scalpellum travassosi, Lauroia Tremogasterina 140 140 . 140 . 134. PI- i . 301 231, 234, 311 • 5. io, 24 324. 334.335-337. 34°, 341. 342, 349 Triacanthidae . . . . .146 Triacanthus .... 146, 156, 157 triaculeatus, Meteterakis . . 12, 21 triaculeatus, Spinicauda .... 5 Triakidae . . . . . .128 Triakis ....... 128 Trichiuridae . . . . . 133 Trichiurus ..... 133, 152 tricuspidatus, Congrus . . . .129 tridens, Haplochromis . . 97-ioo, 114 tridens, Octolasmis . . 231, 241, 312 trigonus, Balanus . 231, 267-268, 309, 312 Trigonopora ..... 346, 348 trigrammus, Cynoglossus . . . 132 Trilasmis . . . 231, 241, 309, 312 trimaculatus, Serranus .... 136 trinidadensis, Lauroia . . . .10 Triporula . 334, 335, 340-345, 349, PL 2 triseriata, Tamaria .... 195 Trixiphichthys . . . . .157 Trypauchen . . . . . -145 Trypauchenidae . . . . 145 tuberculata, Exechonella . 335, 337, 339 tuberculata, Linckia . . . .181 tuberculata, Nardoa 169, 175-177, 180-182, 186, 187, PI. 4 tuberculata, Ophidiaster. . . .181 tuberosum, Metrarabdotos . . . 348 tubulosa, Lagenipora . . . -337 tulipiformis, Balanus 231, 266, 307, 308, 312 tulipiformis, Megabalanus . . .231 tumescens, Ophidiaster .... 194 tumescens, Tamaria . . 169, 194-195 tumida, Fromia . . . 169, 188, 189 tumulosa, Nardoa . 169, 175, 177-180, 187 Umbonula ...... 324 Umbrina ...... 139 uncinata, Paraspinodera ... 24 unguiculatum, Metrarabdotos . 346-348, PI. i unguiculatum, Trigonopora . . . 346 unguiculatus, Metrarabdotos . . . 346 unicolor, Pagrus . . . . .138 Upeneus ...... 138 Upenoides ...... 138 Uranoscopidae . . . . .146 Uranoscopus . . . . . .146 uranoscopus, Polycaulus. . . . 141 Uroconger . . . . . .130 Utinga 346 Vachell, Rev. G. . . 121-165, Pis. 1-3 378 INDEX vachellii, Ambassis Bagrus .... Balistes .... vachellii, Pseudobagrus Sebastes .... Sebastodes Seserinus .... vagina, Trypauchen variabilis, Cryptophialus variolata, Asterias variolata, Hexactis variolata, Nardoa . 169, 171 variolata, Tetractis cruciata variolatus, Pentadactylosaster variolatus, Scytaster variolosa, Linkia . velutinum, Scalpellum . 229, venator, Haplochromis . venustus, Balanus . venustus, Balanus amphitrite . venustus, Balanus venustus . venustus niveus, Balanus 232, venustus venustus, Balanus . Verruca . . . 229, . 136 • 136 124, 146 . 130 140 140-141, PI. 2 134. 139, 153 • 145 232, 301 . 172 . 171 , 172, 182-188 . 171 . 171 172 172 , 231, 234, 311 81, 83 271, 3°9. 3" 271, 280, 281 232, 280-286, 312, 314 , 286-287, 311 232, 280-286, 312, 314 231, 251, 311 verrucosa, Anarthropora . . -341 versluysi, Haplochromis . . 78, 83 vespertilis, Platax . . . . .140 Vespicula . . . . . .140 victorianus, Haplochromis 37, 42, 46, 51, 53 54. 77. 99. ioo, 109, in virgatum, Conchoderma . . 231, 240 virgatum chelonophilum, Conchoderma . 231, 240-241 viridis, Boleophthalmus Gobius Scartelaos vulgaris, Anguilla . watsoni, Ophidiaster weberi, Triacanthus withersi, Chthamalus woora, Minous xanthurus, Amphiprion Xenobalanus . xenognathus, Haplochromis xenostoma, Haplochromis yarraensis, Adeonellopsis . 144 . 156 . 156 129, 150 169, 175-177 • 157 23°. 257, 259, 260, 261 . 141 . 142 232, 301 . 108 51-56, 81, 113 34° zebra, Balanus tintinnabulum 231, 264-265, 309 zebra, Megabalanus tintinnabulum . .231 zebra, Synaptura ..... 132 zebra, Zebrias . . . 132, PI. i Zebrias ..... 132, PI. i PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING