/ V BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY VOL. 17 1968 — 1969 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) LONDON: 1973 DATES OF PUBLICATION OF THE PARTS No. i ...... 19 November 1968 No. 2 ...... 19 November 1968 No. 3 . • . . . . 19 November 1968 No. 4 ...... 4 February 1969 No. 5 ...... 4 February 1969 No. 6 ...... 7 February 1969 No. 7 ...... 14 March 1969 No. 8 29 April 1969 No. 9 30 May 1969 Printed in England by Staples Printers Limited at their Kettering, Northunts, establishment 2 9 CONTENTS ZOOLOGY VOLUME 17 PAGE No. i. Notes on woodpeckers (Picidae). By D. GOODWIN I No. 2. A review of the iguanid lizard genera Uracentron and Strobilurtis. By R. ETHERIDGE 45 No. 3. Nigerian lizards of the genus Agama (Sauria: Agamidae). By ALICE G. C. GRANDISON (Pis. 1-6) 65 No. 4. A revision of the amphipod genus Microdeutopus Costa (Gammaridea: Aoridae). By A. A. MYERS (PI. i) 91 No. 5. Convergence in the structure of the head and cuticle of Euchromadora species and apparently similar nematodes. By W. GRANT INGLIS 149 No. 6. A nomenclatural index to "A history of the British marine Polyzoa" by T. Hincks (1880). By J. S. RYLAND 205 No. 7. The clupeoid fishes described by Bloch and Schneider. By P. J. P. WHITEHEAD (Pis. 1-3) 261 No. 8. Cyclophyllidean cestodes from birds in Borneo. By MICHAEL D. B. BURT 281 No. 9. Type material of the families Lysianassidae, Stegocephalidae, Ampeliscidae and Haustoriidae (Crustacea: Amphipoda) in the collections of the British Museum (Natural History). By M. H. THURSTON & ELIZABETH ALLEN 347 Index to Volume 17 389 NOTES ON WOODPECKERS (PICIDAE) D. GOODWIN _ BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 17 No. i LONDON: 1968 NOTES ON WOODPECKERS (PICIDAE) BY D. GOODWIN Pp. 1-44: 10 text-figures. BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 17 No. i 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. ij, No. i of the Zoology series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). Trustees of the British Museum (Natural History) 1968 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 19 November, 1968 Price Sixteen Shillings NOTES ON WOODPECKERS (PICIDAE) By D. GOODWIN THESE notes are based on examination of the woodpeckers, Picidae, in the British Museum (Natural History) during revision and re-arrangement of the collection. The only species I know in life are the Green Woodpecker Picus viridis, the Greater Spotted Woodpecker Dendrocopos major and the Lesser Spotted Woodpecker D. minor; I have also briefly observed the Grey-headed Woodpecker Picus canus, the Black Woodpecker Dryocopus martius, the Syrian Woodpecker Dendrocopos syriacus, the Middle Spotted Woodpecker D. medius, the Hairy Woodpecker D. villosus, the Downy Woodpecker D. pubescens, the Sapsucker Sphyrapicus varius, the Red-bellied Woodpecker Melanerpes carolinus and the Golden-shafted Flicker Colaptes auratus during visits to Austria and North America. Suggestions or conclusions presented here are, therefore, based mainly on external morphological characters. For this reason I have not suggested alterations from the most recent comprehensive check list (Peters, 1948) except when the evidence in favour of such a course appears overwhelming. Section i : General notes on the family REMARKS ON AFFINITIES, CONVERGENCE AND COLORATION Peters (1948) divides the true woodpeckers (Picinae) into two major groups on the basis of correlated foot and bill characters : those in which the outer hind toe is not longer than the outer front toe, the nasal shelf narrower and the nostril near the culmen ; and those which have the outer hind toe longer than the outer front toe, a wider nasal shelf, and the nostril mid-way between the culmen and the edge of the upper mandible or nearer the latter. When the woodpeckers are thus divided into two groups two significant correlations emerge. First, most, if not all, genera, many or all of whose members are known to be partly or entirely ground feeders, or to feed largely by picking insects from the surface of trunks, branches or leaves or by fly-catching, come into the first group. This is the case with Nesoceleus, Picus, Dryocopus, Melanerpes, Asyndesmus, and Micropternus. Second, some " pairs " of genera whose geographical distributions, colour patterns and general similarity suggest close relationships between them — Geocolaptes and Mesopicos, Micropternus and Blythipicus, and Dinopium and Chrysocolaptes — are separated. Thus it seems likely that these bill and foot differences may be rather labile adaptive characters and that the toe formation may have become differentiated in closely related forms in the course of adaptive radiation. In Europe woodpeckers divide easily into four taxonomically distinct genera — Picus, Dryocopus, Dendrocopus and Pico'ides — with no troublesome intermediate forms. The same does not hold true for most other geographical regions. Even such visually striking differences in coloration as those between " pied " woodpeckers and " green " woodpeckers are bridged by " pied " woodpeckers that show consider- able yellow or green suffusion, such as Dendrocopos temminckii and Melanerpes ZOOL. iy, I. J§ 4 D. GOODWIN striatus; and by "green woodpeckers, such as Veniliornis spilogaster, which, show an underlying pattern virtually identical to that of some of the black and white forms. As in other groups of birds there is evidence that relatively concolorous forms with a simplified colour pattern can derive from forms with a more complex pattern, although in the woodpeckers most of these are still at a stage where their affinities are obvious; for example Melanerpes herminieri and Sapheopipo noguchii. The following general trends in coloration and colour pattern are evident. Forest forms tend to have boldly patterned black and white colour patterns, usually with red or yellow signal or display markings. Forms inhabiting more open woodland are usually more cryptic ; either through a predominantly olive-green colour, at least on the upper parts; or, when predominantly black and white in coloration, through a barred (" ladder-backed ") or profusely mottled pattern. Forms that have become largely terrestial tend to be more fully cryptically coloured, and show reduction or absence of the red or yellow markings on crown or nape that are otherwise so nearly universal in the woodpeckers. In view of the number of species whose behaviour and ecology are little known the above remarks are, of course, tentative. On the other hand, apparent exceptions to those general rules may prove not to be so when we have further information. For example, the Syrian Woodpecker D. syriacus which is black and white with a red nuchal patch in the male inhabits, at least in its European range, rather open cultivated and settled regions, but this choice of open habitat may be, in an evolutionary sense, a recent development. Again, the Green Woodpecker P. viridis feeds largely on the ground through it appears colourful enough at close quarters. When, however, it is on the ground in typical feeding sites, such as grassy slopes, parkland, and woodland openings, its plumage is highly procryptic and even the red on its head, which is darker and less " fiery " in tone than that of many more arboreal species, does not show up very conspicuously. It must be mentioned that conspicuous red, yellow or white rumps can be concealed by the folded wings, and they usually are when their owners are alarmed, so that their possession by such species as the Ground Woodpecker Geocolaptes olivaceus and the Andean Flicker Colaptes rupicola, does not significantly affect the cryptic character of the rest of their plumage. There are widely separated allopatric forms that show considerable general resemblance to each other. In some cases the details of their respective colour patterns make it fairly certain that the resemblance is due to convergence. This is so, for example, with the two ground-living woodpeckers, Colaptes rupicola of South America and Geocolaptes olivaceus of South Africa, whose affinities are clearly with other American and African forms respectively, not with each other. The Chestnut-coloured Woodpecker Celeus castaneus of Central America is related most closely to other Celeus species, and thence to other American genera, and is, obviously, not at all closely allied to the Asiatic Rufous Woodpecker Micropternus brachyurus, to which its rufous, black-barred plumage gives it considerable superficial resem- blance. The small green woodpeckers of Central and South America, of the genus Veniliornis, show such close resemblance to the African genera Dendropicos and Campethera that I feel uncertain whether this is due to genuine close affinity or to convergence, although I think the former less likely. NOTES ON WOODPECKERS 5 The Ethiopian region has no arboreal woodpecker that is very large in size. Although the smallest woodpeckers in the Nearctic, Neotropical and Oriental regions are all about the same size, the largest arboreal African woodpeckers (Thripias) are much smaller than the largest arboreal species in all the other woodpecker- inhabited regions. The largest African species, the terrestial Geocolaptes olivaceus, is, however, about the same size as, or only a little smaller than species that are equally (Colaptes rupicola) or to some considerable extent (other Colaptes species, some Picus species) terrestial in the Nearctic, Palearctic and Neotropical regions. Also no African woodpecker is boldly black and white in colour although the African barbets have produced some predominantly black and white species. Another interesting fact is that although all other continents inhabited by wood- peckers have several very similarly coloured " green " woodpeckers, there are no green species in North America, where their ecological equivalents would seem to be the brownish and barred flickers, Colaptes, and the " ladder-backed " species of Melanerpes (Centurus). FIG. i. (a to e) Diagrammatic outline sketches of the largest species of arboreal wood- pecker in, respectively, the Nearctic, Palearctic, Oriental, Neotropical and Ethiopian regions ; f shows comparative size of the smallest woodpecker species (piculets excluded) in all of the above regions. It should be noted that the largest Nearctic species, the Imperial Woodpecker Campephilus imperialis, is rare and restricted in range; the largest widespread species in the Nearctic, the Pileated Woodpecker, Phloeoceastes pileatus, is approximately the same size as the largest species in each other region, the Ethiopian excepted. SEXUAL DIMORPHISM The great majority of woodpeckers are sexually dichromatic. In nearly all the difference consists of the male having red or, less often, yellow areas on the head that are absent in the female. This may involve only the male having red on the head or both sexes may have heads that are predominantly red in colour but still with the 6 D. GOODWIN female having a little less than the male. Even in species such as Colaptes rupicola and Micropternus brachyurus, in which the male has only a little rather dull red on the malar region, the sexual distinction is clear cut and not just a matter of the male being slightly brighter or having his bright colours slightly more extensive than the female's. This latter form of sexual dichromatism, common in so many birds, is found in only a very few woodpeckers. Only in five species are the sexes alike or nearly so. These are the Middle-spotted Woodpecker Dendrocopos medius, the Sapsucker Sphympicus varius (the predomi- nantly red-headed races daggetti and nuchalis) Lewis's Woodpecker Asyndesmus lewis, the Red-headed Woodpecker, Melanerpes erythrocephalus and the Porto Rican Woodpecker M. portoricensis . In D. medius, both sexes have the crown red, a very common feature of males of allied Dendrocopos species. There is, however, some sexual difference in that the red on the female is not quite so bright as on the male and does not usually reach quite so far back on the nape where it is tinged with yellow. This is the kind of sexual dimorphism common to most bird species in which the sexes are usually said to be alike. It would, presumably, not give so immediate or positive a clue to sex as the type of difference usual in woodpeckers. In the red- headed form of 5. varius there is also a marginal tendency for the male to be more intensely coloured, but in M. erythrocephalus there is no constant difference in brilliance between the sexes. There does not seem any obvious or likely reason why any of these three forms should be able to dispense with the usual type of sexual dimorphism which, since it is nearly universal, must have, in most species, strong selective value. D. medius is, however, sympatric with congeneric and rather similar species, and in its case selection for specific distinctness may have operated at the expense of sexual differentiation. One species that is in many places sympatric with it, D. leucotos, is extremely similar in coloration and colour pattern except that the female has a black instead of a red crown ; it is also larger in size. Asyndesmus lewis is a rather aberrant species which differs much in feeding habits from the more typical woodpeckers (see Bent, 1939). Its colour pattern, although showing some connections with those of some Melanerpes species, to which it is clearly allied, is unusual and it has no red on crown or nape. Typically the female is marginally duller than the male. M. portoricensis shows in extreme degree the loss of distinctive markings common in isolated island forms, being all black except for a dark reddish tinge on the underparts. The sexes are alike. Colaptes auratus is the only species which shows both conspicuous red markings (a red nuchal band) and clear cut sexual dimorphism that does not involve a lack of red in the female, the male alone having black malar stripes. Other flickers, leaving aside for the moment the aberrant Fernandina's Flicker Nesoceleus, differ sexually in the male having red or partially red malar stripes and the female not. In C. pitius the red on the malar stripes of the male is reduced to a pale, dull pink on the tips of the feathers, whose bases are blackish, so that the sexual difference in this species is somewhat intermediate in character between that of auratus and other flickers although less conspicuous than in either. The only woodpeckers in which neither sex shows a trace of red or yellow in the plumage are Nesoceleus fernandinus and the Heart-spotted Woodpecker Hemicircus NOTES ON WOODPECKERS 7 canente. In Nesoceleus the sexes differ in the male having a black and the female a speckled malar stripe; in H. canente the male has the forehead and crown white, the female has a spotted forehead and crown. Besides the colour differences described above female woodpeckers, like the females of most other birds, often average slightly smaller in bill size than the males. In some island forms this difference and the correlated difference in body size is consider- able. In a recent important paper on this subject (Selander, 1966) it has been shown that such differences are correlated with sexual differences in feeding ecology and serve to lessen or prevent feeding competition between male and female. The piculets, sub-family Picumninae, show " typical " woodpecker sexual dimorphism except that in two species the " male's " colour is chestnut, not pure red or yellow. JUVENILE PLUMAGES Juvenile woodpeckers, like juvenile barbets, Capitonidae, but unlike most other birds, usually show similar bright red or yellow pigments to those of the adults. In some species, of which the Greater Spotted Woodpecker Dendrocopos major is an example, the juvenile male may have differently placed and more extensive red markings than his father, and the juvenile female show red areas that are lacking in the adult female. Many juvenile woodpeckers have a greater number of attenuated tail feathers similar in character to the stiff, central feathers used to prop the bird when resting or working on a tree trunk than have the adults. The juvenile's wing may show a reduction in size of the two innermost primaries and consequently rather more pointed wing. These facts prompted Kipp (1956) to suggest that the juvenile plumages of woodpeckers represented a probable future stage in their evolution as they could, he thought, neither represent a recapitulation of a prior evolutionary stage nor be of value to the juvenile as distinct from the adult. Verheyen (1957) disagreed with this conclusion; pointing out that, in many other birds, the juvenile rectrices are to some degree narrower and more pointed than the adult's and that the reduction in size of the inner primaries was probably connected with the need for quick growth to enable the early onset of the first moult. This was also discussed by Sibley (1957), who pointed out that this reduction of the innermost primaries occurs in some species only, and is almost certainly adaptive. The Stresemanns (1966) in their comprehensive study on moult in birds agree with this conclusion. Verheyen also emphasized, correctly, that the juvenile plumage of woodpeckers is, like that of most other birds, usually of a weaker and more downy character than the adult plumage. He suggests that this might be of particular disadvantage to a woodpecker and hence one reason for the quick onset of the first moult. This, in some species, begins before fledging, at which time the two reduced inner primaries above mentioned, may have been replaced by adult feathers. Verheyen suggested that the bright red or yellow pigments of juvenile woodpeckers, and the fact that in many woodpeckers the juveniles show the same sexual dimorphism as the adults, might be due to an original (presumably duller and less dimorphic) juvenile plumage ZOOL. 17, I. I§§ 8 D. GOODWIN having been completely suppressed in the course of evolution. He considered that the present juvenile plumage was originally the first post- Juvenal dress. I think this is an unlikely hypothesis. Except for the bright red or yellow pigments often present, this plumage has the typical characteristics commonly found in juvenile plumages in other groups: a looser and more downy or woolly texture of the contour feathers; weaker shafts of the quill feathers; a tendency for transverse barring, dark or light shaft streaks or pale feather tips (not present or not so pronounced in adult) to be present or more prominent and to give a more spotted or barred general appearance and a tendency for black areas to be browner and less glossy. All these characters are not, of course, usually found in the juvenile plumage of any one species. The red or yellow pigments themselves may be less fully developed in the juvenile plumage. The red is often lighter or more orange-red than that of the adult; and in many species the red tips of the feathers are less extensive, so that the juvenile appears to have a scattering of red speckles where the adult, if in unworn plumage, appears uniform red. In a few cases the lighter red of the juvenile may make it appear more brilliant. Similarly, in those species which have golden yellow instead of red on the head, this is often of a darker and more orange tint in the juvenile. It is possible, however, that in both these instances the juveniles may not appear more impressive to avian as they do to human eyes. Verheyen divided the African and Eurasian species he studied into four groups: those in which the juveniles resembled the respective adults in their degree of sexual dimorphism; those in which the juveniles resembled neither parent; those in which both sexes resembled the adult male, and those in which both sexes resembled the adult female. Most other woodpeckers fall into one or other of Verheyen's categories, which do not, however, cover the whole range of juvenile plumages. It seems worthwhile to describe briefly the likenesses and differences (vis a vis the adult) found in the coloration and sexual dimorphism of the juvenile plumages, and their generic distribution. In so-doing I shall not take account of such minor differences as are generally characteristic of juvenile plumage. Thus a juvenile may be said to be " like the adult " if it has the same colour pattern and general coloration, even though it may, for example, have an orange-red instead of a scarlet cap. It seems pertinent to digress here to mention the hazards when ascertaining the juvenile plumage in species of which few reliably sexed juvenile specimens are available. It has become very evident in the course of this study, that in the past many collectors have labelled young woodpeckers as males or females according to whether their head coloration most resembled that of the male or female adult. With well-known European and American species this guess-sexing is immediately obvious, but is is much less easy to detect with species represented only by one or two specimens. Another hazard is that in some species, of which the Three-toed Wood- pecker, Pico'ides tridactylus is an example, the feathers of the forehead and f orecrown (sexually differentiated in the adult) are moulted very soon indeed after fledging, so that specimens in almost complete juvenile plumage, but with adult plumage on front and top of the head, may be much commoner in collections than birds in complete juvenile plumage. In general colour pattern and the (usually sexually dimorphic) head markings, NOTES ON WOODPECKERS 9 juvenile plumages, come into one or more of the following categories. Categories 2 to 6 involve coloration of male juveniles, 7 to 9 of females, and 10 to 12 of both sexes. (1) The juvenile male and female are like respective adults in the extent of red or yellow on the head. In this category are some species of Colaptes, some (possibly all) species of Piculus and Chrysoptilus , some species of Melanerpes (cruentatus, formicivorus , candidus and possibly some others), Trichopicus (although in this species the yellow on the throat, which is not a sexual character, is absent in the juvenile), Celeus flavescens (most probably other species of Celeus also), some species of Phloeoceastes (lineatus, pileatus and, probably, schulzi and galeatus), Dryocopus, Picus (with two possible exceptions and a tendency for restriction of lipochrome pigment in the juveniles of the yellow-naped species), Gecinulus (females of both species and males of one), Dinopium benghalense and D. javanense (but in the latter the front of juvenile's forehead is brownish so that it is intermediate in this character between benghalense and the other Dinopium species) , Chrysocolaptes (festivus only ; although the male of validus is very similar to the adult but has yellowish brown throat and malar region), Dendrocopos (males of all the red-crowned species and females of a very few species) , Meiglyptes (but juvenile males often show a reddish tinge on forehead, which is rarer in adult males), Blythipicus (Pyrrhotis], Micropternus (but a few juveniles show a trace of red on the nape). (2) The juvenile male has less extensive red or yellow areas on head than has the adult male. This broad category is best subdivided into the following groups. (a) Juvenile male has red (or yellow] only on the hind crown and lor nuchal region, adult male has entire top of head, and sometimes forehead also, red or yellow. In this group belong the Campethera species. In nubica, punctuligera and bennettii juveniles resemble the adult female in having red only on the hind crown and nape, the forehead and forecrown is, however, more or less dull grey or blackish, not spotted or streaked with white as in adult female. In maculosa and permista the differences between adult and juvenile males are similar. Our few specimens of juvenile females of these species show less extensive red on the nuchal region than males. In caroli and nivosa, in which the adult females has no red and the adult male has red only on the nuchal region, the juvenile males appear to have no red and thus resemble the female. This statement is, however, based on only one juvenile male specimen of each species in our collection (which I have reason to believe were reliably sexed) and on Verheyen's description of caroli. In Dinopium rafflesii the juvenile male has the forehead brownish, shading to blackish on the crown with a few red flecks, narrow greenish gold fringes to some of the feathers and only the hind crown and long nuchal crest red; in D. shorii the juvenile male has the forehead streaked dark brown and dull cream; in both these species the adult males have the top of the head entirely red. In Chrysocolaptes lucidus (guttacristatus and allied mainland races) the young male also lacks red on the forehead. This species shows, however, an interesting situation. The juvenile male at first has a brownish black forehead, unspotted or with only obscure spots. Before the juvenile plumage as a whole is shed some of the D. GOODWIN forehead feathers are replaced by boldly spotted ones similar to those on the head of the adult female. These spotted feathers may extend well onto the crown where they have, presumably, replaced juvenile red feathers. Later these spotted feathers are replaced by red ones but some birds show feathers intermediate between the adult red feathers and the black and white spotted feathers, that is red feathers somewhat intermediate in form and with a white spot. These spotted feathers on the forehead do not seem to represent an intermediate plumage in any true sense but they do give the owner's head some " female " characteristics. From the available specimens it is difficult to judge whether differences are due to age or individual variation and observations on living birds are needed. FIG. 2. Diagrammatic sketches of heads of Hemicircus concretus to show differences in colour pattern between adults (left) and juveniles (right) of both sexes; males above, females below. Shaded areas are dark grey, stippled areas bright red or orange-red, unshaded areas buff; the scale-like markings on the crest of the juvenile female represent grey feather edges. Hemicircus concretus shows a striking difference between the juvenile and adult male. The latter has the entire top of the head, from forehead to nuchal crest, bright red, the rest of its head is dark grey. The juvenile male has the forehead and crown buff and only the nuchal crest (ends of the long head feathers) red. In the race sordidus the adult has the nuchal area largely grey and the juvenile male has much less red than in the juvenile of the nominate form. In both the juvenile shows a suggestion of a buff malar stripe which is lacking in the adult. It may be mentioned here that although the adult female's head and crest are grey the juvenile female is like the juvenile male except that her nuchal region is buff like her crown, not red. NOTES ON WOODPECKERS II (b) Red on juvenile male does not extend to, or so far down, nape as in adult male. In this group belong only the species of Mesopicos. (c) Adult male has head largely or entirely red; juvenile male has less red on head. Phloeoceastes : in guatemalensis and melanoleucos , juveniles are like the adult female, thus having black forehead and forecrown and much less extensive red areas that adult males. In robustus (but this is surmised from only one specimen) the juveniles also, probably, resemble the adult female. In haematogaster , however, in which, as in robustus, the adult female has the forecrown red, the juveniles of both sexes have the forehead and forecrown blackish. They also have buff instead of red rumps although both on rump and head the juvenile feathers are soon replaced. In leucopogon the juveniles of both sexes are said (Laubmann, 1930) to have entirely black heads but this statement was based on one moulting juvenile. In pollens, the only form in which the female has no red on the head, an unsexed fledgling is like the adult female. FIG. 3. Diagrammatic sketches to show difference in colour patterns of heads of adults (right) and juvenile male (left centre) of Ipocrantor magellan icus ; the distribution of red and black on juvenile female is similar to adult female's. In Ipocrantor (magellanicus] the male has the head entirely red; the juvenile male has a black head with forehead, crown, nape, malar stripes and chin red. Both juvenile and adult females have only the chin and loral region red. In the red-headed forms of Sphyrapicus varius (ruber and daggetti] the generally smoky-brown juvenile has the head extensively tinged with red but this red is not so extensive and not nearly so bright as that of the adult. So far as adult- juvenile 12 D. GOODWIN differences are concerned the situation is the same with those races of varius in which the adults have less red on the head. (d) Adult male has some red on head, juvenile male has none. In Campethera nivosa and C. caroli the adult males (but not the females) have red on the nape, although in carola this is reduced to a scattering of red-tipped feathers. If a male and a female juvenile of carola and a male juvenile of nivosa are r rrrectly sexed, as I believe to be the case, then in neither of these species has the juvenile male any red on the head. The juvenile of caroli (both sexes) also differs in having less yellow pigment, in the top of its head being blackish brown instead of dark green and the yellow, dark-speckled throat and eye-stripe of the adult being replaced by pale brownish. In the nominate form of Sphyrapicus varius the adult male has red crown and throat, the mainly brownish (instead of black and white) juvenile male has no red but it usually moults some or all of the feathers on its forehead and crown before it attains other parts of the adult dress. The juvenile male of S. thyroideus has the same black and white colour pattern as the adult male but has a white instead of a red throat. There appears to be only one published description of a juvenile male of Campephilus principalis (Tanner, in Bent, 1939) . This bird resembled in colour the adult female and lacked the red nuchal crest of the male (which extends forwards on sides of head). It did not begin to show red feathers until it was about 3§ months old. It is highly likely this is the normal situation and possible that some of the few other juveniles seen or collected, and sexed as females were, in fact, males also. It is also probable that the closely similar and possibly conspecific C. imperialis agrees with principalis in this. (3) Juvenile male has red (or yellow) on forehead and /or crown, adult male has this colour only on nuchal region. In this category are many species of Dendrocopos, one species (rubiginosus) of Blythipicus and, but to a marginal degree only, Dendropicosfuscescens. In this latter the red on the adult male's nape extends forward nearly to the crown. (4) Juvenile male has red on crown extending a little further towards forehead than red of adult male. In this category come all species of Mesopicos (3), both species of Gecinulus and, marginally, Colaptes auratus in which the juvenile male has a strong tinge of red on the crown. (5) Juvenile male and adult male both have red (or yellow) crown, this colour sometimes extending further over the nuchal region in the adult. This includes those species (many) of Dendrocopos in which the adult male has a red crown; Mesopicos (these come into category 4 also), Thripias, Pico'ides, Dryocopus, Chrysocolaptes; most species of Dinopium and Picus; some species of Dendropicos and some (possibly all?) species of Veniliornis. (6) Adult male has red only on malar region; juvenile male has red on forehead and crown also. Mulleripicus pulverulentus. I think it highly likely that the case is the same in M. fuliginosus, of which I have been unable to find either juveniles or descriptions NOTES ON WOODPECKERS 13 of them. Meiglyptes may show a suggestion of this condition, some male juveniles being tinged with red on the forehead. FIG. 4. Diagrammatic sketches to show colour patterns of heads of adult males (left) and juvenile males (right) of Mulleripicus pulverulentus and M. julvus. (7) Juvenile female resembles adult female in having no red on head, in species in which males have red on head. Those species of Colaptes in which the males have red malar stripes ; a few species of Dendrocopos, including the aberrant D. albolarvatus ; some species of Picus (squamatus and its allies), Meiglyptes, Chrysocolaptes and Dinopium and in two related and rather aberrant species of Campethera (nivosa and caroli) . (8) Juvenile female and adult female differ appreciably in colour of head but neither has any red although the male has. One species only, Hemicircus concretus, in which the adult female has the head (and most of the body) a concolorous olivaceous grey but the juvenile has the top of head and crest buff. (9) Juvenile female has some red (or yellow) on head; none present in adult female. Most species of Dendrocopos, Pico'ides, Mesopicos, Thripias, most species of Dendropicos, Veniliornis callonotus and, in all probability, other species of Veniliornis also. (TO) Juveniles of both sexes have strikingly different coloration and colour pattern from adults. 14 D. GOODWIN Melanerpes erythrocephalus. The juvenile appears predominantly greyish brown except on wings and tail owing to extensive pale brownish to brownish white fringes, central streaks and/or cross bars on the blackish cover feathers. There is only a little red around the eye, in contrast to the entirely red head of both sexes of the adult. The juveniles of the nominate form of Spyrapicus varius (and to a lesser extent those of other forms of the species) are very similar in general coloration (and in the plumage patterns by which this is achieved) to the juvenile of M. erythrocephalus described above. Hemicircus concretus comes less definitely into this category. The colour pattern of the juveniles (but not their coloration) is very close to the adult male's (see Text-fig. 2) but they also show some indication of a buff malar stripe which is lacking in the adults. (n) Adults differ in coloration and colour pattern, juveniles very like respective adults but juvenile male lacks the red throat of the adult male. Sphyrapicus thyroideus. This species is of interest in that the differences between the male and female (at all ages) closely parallel the differences between adults and juveniles in the related S. varius. (12) Neither adults nor juveniles have red or yellow on head; juveniles resemble respective adults (allowing for very minor differences) . Hemicircus canente. It seems probable that Nesoceleus comes into this category but I can find neither juveniles of this species nor descriptions of them. The aberrantly coloured Asyndesmus lewis does not fit readily into any of the above categories. Its juveniles are similar to but generally duller than the adults. They lack the silver collar and upper breast, have only traces of red on the face and have white sub-terminal spots (not present in the adult) on the feathers of the hind neck. DISCUSSION. Thus, taking the woodpeckers as a whole, juveniles tend to be less, although often only a little less, conspicuously coloured than the adults. Sexual dimorphism is usually present in the juvenile plumage but is often much less marked than in the adult plumage. In species in which the adult males or adults of both sexes have the head almost entirely red the juveniles (where known) have appreciably less red on the head. Juveniles, taking all species into consideration, approximate more closely to the familial mean in the amount of red on their heads than do adults. This is well exemplified, within a small genus, by Mulleripicus pulverulentus and M. fulvus (see Text-fig. 4) and certainly suggests that the juvenile colour pattern resembles a previous stage in the species' evolutionary history, as Voous (1947) claimed for the red crowns of those species of Dendrocopos which show this feature only in their juvenile plumage. Certainly speciation in woodpeckers has involved both reductions and increases in the red areas of the adult plumages. It is likely, however, that many features of the juvenile plumages may be functional and hence of selective value. This seems obvious in Melanerpes erythrocephalus and Sphyrapicus varius, whose much less conspicuous juvenile plumage must give some protection against aerial predators ; both these species are partly migratory and often inhabit relatively open, although wooded, country. Under such conditions it may be of great advantage to the inexperienced juveniles to be less conspicuous. The NOTES ON WOODPECKERS 15 same is probably true for some other species in which the differences between young and adults are much less marked. In Picus viridis, for example, the more or less speckled and barred (but otherwise similarly coloured) juveniles are even less conspicuous when feeding on the ground than are the adults. In the many species in which the juveniles are like the adults it is evident that here any advantages of a less conspicuous juvenile plumage are outweighed by others. Probably adult coloration is advantageous in the acquisition of territory or use of feeding areas in competition with adults. On the other hand the tendency in so many species, some of them congeneric with those showing strong sexual dimor- phism, for the male and female juveniles to be almost alike, differing only in the female showing less extensive red or yellow areas than those of the male, may perhaps give other advantages which are more important for these species. It might, perhaps, lessen any tendency that might otherwise exist for either parent to " favour " young of one sex or it might better enable the female juvenile to hold her own in competitive strife with her brothers. Young Dendrocopos major, for example, may fight even in the nest hole (Sielmann, 1959). Where they are numerous, one sees much aggression and frequent fights between fledged young before (as well as after) they have moulted their juvenile plumage. Where the juvenile lacks red on the head or has it more restricted than the adult it is tempting to suggest that this might function to reduce or inhibit aggressiveness on the adult's part, particularly in those species in which the juvenile lacks the red precisely on those parts, forehead and forecrown, which are presented to the adult when soliciting or taking food from it. This may be so but against this hypothesis is the fact that in some other species, including a large number of Dendrocopos, it is precisely these areas (forehead and forecrown) which are red in juveniles and not in the adult. It is often implied that the juveniles of those species of Dendrocopos that have a red nuchal patch have the entire top of the head red but in fact the young male, although having a red forehead and forecrown, is black on the nape where the adult male is red. The same is of course, true of the young female who has less red on her head (as a rule) than the young male. In these species of Dendrocopos the " replacement " of the red on the napes of adults by red on foreheads and crowns of juveniles reaches an almost bizarre state in the Red-cockaded Woodpecker D. borealis in which the adult male has a tiny red mark (the " cockade ") on the sides on the hindcrown and the juvenile male an equally small red mark in the middle of his forecrown. It is, of course, feasible that in some species some inhibition of adult agression could best be achieved by the young not presenting red markings to the feeding parent and in other species, in which the relative balance of fear and aggression differed, in their doing precisely that! As Dendrocopos major is often instanced as a supposed example of a species in which the young are more brightly coloured that the adult it seems pertinent to digress here to say it is, in my opinion, very doubtful if the red foreheads of the juveniles make them more conspicuous to predators than are their parents. The same is probably true for the other species showing this character. All the boldly-marked black and white woodpeckers are, at least to human eyes, relatively conspicuous as compared with olivaceous or " ladder backed " species. ZOOL. 17, I. !§§§ 16 D. GOODWIN Section 2 : Taxonomic notes on genera and species MELANERPES, CENTURUS, TRIPSURUS AND LEUCONERPES Peters (1948) included all the above genera in Melanerpes with the exception of Leuconerpes, which he retained as a monotypic genus for the White Woodpecker candidus. Hargitt (1890) had used a similarly broad conception of Melanerpes but had also included candidus therein. De Schauensee (1966) retains the genus Leuconerpes but notes that it "is perhaps not separable from Melanerpes. " Selander & Giller (1963) give cogent reasons for treating Tripsurus as a synonym of Centurus but say that they do not feel justified in including both these genera in Melanerpes. However, I think that that is the most reasonable decision, unless all three genera are to be maintained. Both in its morphological characters and what is recorded of its ecology (see esp. Selander & Giller) Tripsurus seems perfectly to bridge the gap (such as it is) between Centurus and Melanerpes (sensu strictu). I provisionally maintain the genus Melanerpes as used by Peters, except as hereunder discussed. The species candidus does not seem to me to be sufficiently distinct to warrent upholding the monotypic genus Leuconerpes. Its very distinct coloration, with entirely white head, parallels that of the few white-headed African barbets of the genus Lybius, the differences between most Lybius species and the white-headed forms being comparable to those between candidus and related species. Wetmore (1926) gave information on field habits of candidus, including its gregariousness (a feature also found in some Melanerpes species) and added that the genus Leuconerpes " in external characters . . . seems to be only slightly differentiated from Tripsurus. " Sharncke (1931) concluded from studies of its tongue musculature that candidus was a primitive form and closely linked with Sphyrapicus. He found, however, several similarities between its tongue and that of Melanerpes aurifrons. Pending further and more detailed studies it is, I think, best to include candidus in the genus Melanerpes. Melanerpes striatus The Hispaniolan Woodpecker is a very distinct species which stands out among other species of Melanerpes by reason of its red rump, greenish yellow and black barred upperparts, yellow-green belly and, on closer investigation, its rather long tail and the marked sexual dimorphism in length of bill. For these and other reasons Selander & Giller (1963) argued that the monotypic genus Chryserpes should be used for this species. Later, however, after a detailed study of the species in the field in Hispaniola, the senior author (Selander, 1966) changed his opinion and was in favour of including it in Centurus (Melanerpes} . I do not know this species in life but on its museum characters I am entirely in favour of Selander's second decision. The barred pattern of its upperparts only differs essentially from other barred Melanerpes (Centurus) species in the greenish yellow instead of white on the paler parts of the feathers ; this would seem to involve only the spread of lipochrome pigments to these areas. In this connection it may NOTES ON WOODPECKERS 17 be significant that M. superciliaris from Cuba has the white parts of the mantle suffused with yellow and M. radiolatus from Jamaica has much dark yellow on the lower breast and belly. The head colour pattern of striatus is rather like that of superciliaris. Its largely yellowish green coloration gives striatus a certain resem- blance to some species of Chrysoptilus but it shows no trace of the conspicuously barred and spotted underparts of these. The red patch on the rump is shared by no other New World woodpecker except Veniliornis kirkii but I do not think any of the Chrysoptilus species or V. kirkii are very close relatives of striatus. M. flavifrons, M.formicivorus & M. chrysauchen The Yellow-fronted Woodpecker M. flavifrons and the Golden-naped Woodpecker M. chrysauchen are allopatric and seem best given specific rank although they certainly comprise a superspecies. The differences of the colour patterns of the heads of these two are very similar to those between the North and Central American forms of the Acorn Woodpecker M. formicivorus and the South American form (flavigula) of that species. The differences between nominate chrysauchen from Costa Rica and the Pacific slope of western Panama and the race pulcher from northern Colombia may be in part or whole due to character displacement in reference to the different races of formicivorus with which they are sympatric (see Text-fig. 5). Melanerpes rubrifrons Peters (1948) gives the Red-crowned Woodpecker rubrifrons specific rank but others (Stresemann, 1924, Greenway & Griscom, 1941, Todd, 1946 & Haverschmidt, 1956) have concluded that is a colour phase (morph) of M. cruentatus. I fully agree with this latter opinion which is substantiated by the specimens in the B.M. collection. Besides phenotypically " pure " specimens of each form there are several that are intermediate, to varying degrees. The African Woodpeckers All the sub-Saharan African woodpeckers seem to be more closely allied to one another than to any non- African species or genus. They are, I think, representatives of the stock ancestral also to Picus and Dendrocopos. Adaptive radiation has, however, resulted in similar but less marked divergences to those shown by wood- peckers in Eurasia and America. The second possibility: that the African forms are polyphyletic, with Thripias and Dendropicos closer to Dendrocopos, and Mesopicos and Campethera to Picus, is, I think, rather less likely. In spite of some remarkable general resemblances in coloration I think that the South and Central American green woodpeckers are more closely related to other New World forms, in the genera Colaptes and Melanerpes, than they are to African forms. When comparing, for example, species of Dendropicos and Veniliornis that look much alike, one finds that some details of their colour patterns differ from each other and are, respectively, closer to those of other African and South American species. There have been divergencies of opinion as to how many genera of African wood- i8 D. GOODWIN FIG. 5. Sketches to show colour patterns of heads of males (left) and females (right) of Melanerpes formicivorus (top) which overlaps in range with M. chrysauchen (second row) in Central America and of the form of M. formicivorus (third row) which overlaps with M. chrysauchen in northern South America. (See text). peckers to recognize. Hargitt (1890) recognized the genera Campethera, Dendropicos, Mesopicos, Thripias and Geocolaptes, mainly on structural characters with emphasis on shape and size of bill. Other authors, notably Chapin (1939) recognized further genera but Mackworth-Praed & Grant (1957) followed Hargitt except for resurrecting the genus Ipophilus for the two species obsoletus and stierlingi. Peters (1948) put elliotii and johnstoni (now reckoned as a race of elliotii) in the genus Polipicos (following Chapin) but otherwise recognized the same genera as Hargitt and Praed and Grant although not with precisely the same allocation of species within them. I have followed Peters' nomenclature except where otherwise stated. NOTES ON WOODPECKERS 19 DENDROPICOS Peters includes in this genus fuscescens, stierlingi, elachus, abyssinicus, poecilolaemus , gabonensis and lugubris. I concur with him in thinking that these species are probably more closely allied to each other than any is to species in another genus. This assumes, as I think is likely, that some differences of general coloration such as a tendency towards light, barred or spotted plumage in dry savannahs; darker, greener coloration in forests and the acquisition of a red rump have arisen indepen- dently in different (but related) stocks. Their head markings suggest the Dendropicos species are nearer to Thripias and Mesopicos than to Campethera. The extreme similarity between Dendropicos lugubris and Campethera nivosa is, I think, due to parallel development but it may indicate true affinity in spite of differences in bill shape. " Dendrocopos " obsoletus Most writers on African woodpeckers (Bates, 1937, Chapin, 1939, Hargitt, 1890, Peters, 1948, Voous, 1947) put the Brown-backed Woodpecker, obseletus, in the genus Dendrocopos (or in Yungipicus which is now usually regarded as a synonym of Dendrocopos) but Grote (1928) and Bannerman (1933) placed it in Dendropicos. Voous in his monograph on Dendrocopos says that Grote and Bannerman had " abusively referred (it) to Dendropicos " and adds that this latter genus is quite distinct in having reddish or yellowish shafts to rectrices or remiges and usually in addition a distinct greenish tinge of the plumage. He considered, however, that stierlingi was correctly referred to Dendropicos. Mackworth-Praed & Grant, 1957, place obsoletus and stierlingi together in the genus Ipophilus. I think that Bannerman and Grote were right to include obsoletus in Dendropicos. Among them the Dendropicos species show varying amounts of green coloration and/ or yellow or reddish quill shafts. In two of them, stierlingi and elachus, this is reduced to a not very pronounced yellow tinge in the shafts of the wing and tail quills (stierlingi) or tail quills only (elachus). If either of these two species are com- pared with any other Dendropicos species it will be seen that the " gulf " between them, so far as yellow pigmentation is concerned, is greater than that between either one of them and obsoletus. Voous links obsoletus with the pygmy woodpeckers of south Asia, Dendrocopos nanus and its allies, but in plumage pattern obsoletus is closer to Dendropicos species than it is to any of the Asian forms. In its coloration it is nearest to elachus although it lacks the latter's scarlet rump which is, in all probability, a species recognition mark serving to isolate obsoletus and elachus in the areas where their ranges overlap. Such species as obsoletus and elachus undoubtedly indicate the close relationship between " green " and " pied " woodpeckers or the potentiality for " green " and " pied " forms to evolve, in different geographical regions, from a common stock. I think, however, there is little doubt but that the true affinities of obsoletus are, as Bannerman and Grote decided, with Dendropicos. 20 D.GOODWIN Campethera bennettii, C. scriptoricauda & C. nubica Benson (1952) gives evidence for the conspecifity of Bennett's Woodpecker C. bennettii and the Tanganyika Woodpecker C. scriptoricauda. In C. bennettii vincenti (Grant & Mackworth-Praed 1953) the throat colour of females is intermediate between the chocolate brown of bennettii and the creamy white of scriptoricauda, although it does not have the black spots of the latter. Some males of vincenti also have the ear coverts a less pure white than those of other forms of bennettii. Also a female scriptoricauda, from Blantyre, shows some approach to bennettii in colour, having fewer and smaller spots than usual, on a beige-tinted throat. It is unlikely that these specimens merely indicate occasional hybridization between two " good " species as there seem to have been no " pure " specimens of either bennettii or scriptoricauda collected within the range of vincenti and, more importantly, Mr Benson informs me (pers. comm.) that the calls and ecology of bennettii and scriptoricauda do not differ. I therefore agree with him that, on present evidence, scriptoricauda is best considered as a race of bennettii. The very distinctive chocolate and white markings on the face and throat of the females in most forms of bennettii and, possibly, also the very white cheeks and intense red malar stripes of the male, probably function as isolating mechanisms in reference to C. abingoni, with which the chocolate-throated forms of bennettii overlap widely in range whereas scriptoricauda (and C. nubica} overlap only marginally with abingoni. The Nubian Woodpecker C. nubica does not seem to differ in any significant plumage character from C. bennettii scriptoricauda; they differ only in the unspotted throat of nubica. Localities of specimens in the British Museum collections do not suggest any actual overlap of distribution. I provisionally maintain them as separate species, however, because Mr. C. W. Benson informs me that their calls differ, or at any rate those calls most usually uttered, and they differ in ecology. Campethera nubica & P. punctuligera I provisionally treat the Nubian and the Fine-spotted Woodpeckers, nubica and puctuligera, as members of a superspecies but I think it likely that they may prove to be conspecific. Although their respective ranges come close together in the southern Sudan and the eastern Congo I can find no evidence of both having been taken or identified in the same area (see also Chapin, 1939). The form of punctuligera, C. p. balia, from the south-eastern Sudan and eastern Congo is largely intermediate in colour and colour pattern. The differences between the two forms are, chiefly, the size and formation of the spots on the breast and the presence (in punctuligera} or absence (in nubica} of spotting on the throat. One female of balia from Baginzi, Bahr el Gazal (B.M. No. 1919. 10.7.23.) comes closer to nubica than most, having no spots on the throat, very dull underparts (less yellow tinge than usual in punctuligera} with the breast spots closely approaching those of nubica in pattern and general appearance although still much smaller than those of nubica. A juvenile from Angba, on the river Welle, also much resembles nubica in its breast spots. NOTES ON WOODPECKERS 21 Chapin (1939) says that the calls and habits of punctuligera and nubica are extremely similar and that " adherents of the Formenkreise theory " would surely consider them as forms of one species. A main difficulty and the one that chiefly prompts me still to treat them as species is that in appearance punctuligera could equally well be a form of bennettii, with which it is also allopatric. In fact there is an even closer resemblance in colour pattern between punctuligera and bennettii scriptoricauda than between the former and nubica. Present evidence suggests differences between the calls of bennettii and those of scriptoricauda but this negative evidence is of no great significance until the vocalizations of all three forms have been recorded in more detail. I suspect they may prove all to be members of a superspecies or even conspecific. Campethera notata I think Clancey (1958) is, right in suggesting that the Knysna Woodpecker notata and the Golden-tailed Woodpecker abingoni are possibly conspecific and certainly members of a superspecies. In spite of their great similarities, however, the pattern of the breast spots in notata is closer to those of nubica, bennettii and punctuligera, with none of which is notata sympatric, than to abingoni although such breast spots as those of notata could easily drive from the streaks of abingoni or vice versa. Campethera cailliautii, C. taeniolaema & C. maculosa These three species are all very closely allied, the Little Spotted Woodpecker, cailliautii and the Fine-banded Woodpecker taeniolaema being mainly allopatric. Their ranges overlap in the eastern Congo and adjacent areas where they appear, however, to be ecologically isolated, taeniolaema inhabiting the highlands and cailliautii the lowlands. They are very similar in coloration but cailliautii has the throat and face buffish with blackish, speckly-looking, barring whereas taeniolaema has the face and throat greenish white very finely barred with greyish olive. From the Golden-backed Woodpecker maculosa, with which it is sympatric, cailliautii differs in the male having a slightly brighter red crown, greener and les gold-tinged upperparts and less boldly spotted on the throat. The females differ rather more markedly in maculosa lacking the red nape of cailliautii; she also has her entirely blackish crown spotted with deep buff, the spots on the head of cailliautii being pale buffish to white. Campethera tullbergi The rather distinct Tullberg's Woodpecker appears to be closest to cailliautii (permista) with which it is sympatric. The carmine patch on the carpel area of the wing probably serves as a species-specific recognition mark in reference to cailliautii. I fully concur with Peters (1948) that C. wellsi should be considered a race of tullbergi. Campethera nivosa The Efulen race of the Buff-spotted Woodpecker, efulensis Chubb, appears to be inseparable from nominate nivosa. One of three specimens from the type locality 22 D. GOODWIN (Efulen, Cameroon) can be matched by a specimen of nivosa from Portuguese Guinea. The large series from Kumba, Cameroons, vary from typical " nivosa " to typical " efulensis " in colour. The races poensis and herberti are slightly differentiated but recognizable. MESOPICOS The Grey Woodpecker M. goertae and the Olive Woodpecker M. griseocephalus are very closely related and could certainly be considered as members of a superspecies or even as conspecific but for the fact that their ranges overlap in parts of Kenya and north-eastern Tanzania. We have, however, no records of both species from precisely the same place although there are specimens of both from north-eastern Tanzania, goertae from Nabara and griseocephalus from Same. In and near the area of apparent overlap they appear to be ecologically separated, griseocephalus being confined to highland forest. The only absolute colour differences between the adults are the darker grey head and golden-olive breast of griseocephalus. In goertae juveniles of both sexes have red on the head, the male usually rather more than the female. Some works on African birds imply that in griseocephalus the young are, except for being duller, like the respective adults. There are only six juvenile griseocephalus in the British Museum collection, some unsexed, others sexed as males. AU of them have red on the head ; in some of them the red is less extensive than in others and I suspect these may be females. Juveniles of griseocephalus are, incidently, paler and greyer in colour than adults and thus come even closer to goertae in appearance than do the adults. A few remarks on geographical variation in both species: From West Africa (Senegal, Gambia, etc.) across to Uganda and the southern Sudan M . goertae shows no striking differentiation, if allowance is made for individual differences (from the same area some birds may have an orange-red belly patch and others not) and those due to wear (which are often considerable). Within this region there is a tendency for birds from the more northern arid areas to be palest and to have more distinct barring on the wings and upperparts, those from the southern areas to be darker and duller. Specimens from Gambia to Uganda tend to have little or no orange-red, only yellowish, on the belly. Those from the extreme north-east of this area (parts of Southern Sudan) are more like those of Senegal in coloration than are the latter to Gambian birds! I think it best to recognize three races within the above areas : nominate goertae from Senegal, the paler konigi (Neumann, 1903) from the Lake Chad regions, parts of French Niger and French Sudan and centralis (Reichenow 1900) and agree with Mackworth-Praed & Grant (1957) in thinking agmen (Bates, 1932) and oreites (Grote, 1923) best treated as synonyms of centralis. This is an admittedly wide but I think workable interpretation of centralis but it must be borne in mind that no hard and fast line can be drawn between any of the above-mentioned forms of goertae. To the east of its range centralis intergrades with abessinicus (Reichenow, 1900). M. griseocephalus persimilis (Neumann, 1933), described from the Highlands of NOTES ON WOODPECKERS 23 Angola (Benguella) was recognized by Peters but has since been commonly treated as a synonym of ruwenzori (Sharpe, 1902). In the original description persimilis was separated on its smaller size. Besides the rather slight size difference our specimens from Angola, southern Congo and Tanzania mostly show quite conspicuous paler barring on the flanks which is absent from specimens of ruwenzori from Ruwenzori and Lake Kivu. I have, therefore, recognized persimilis. Mesopicos elliotii Chapin (1939) & Peters (1948) both use the genus Polipicus Cassin for Elliot's Woodpecker which was previously (Hargitt, 1890) and subsequently (Mackworth-Praed & Grant, 1957) included in Mesopicos. In structure of bill elliotii (with which johnstoni, formerly given specific rank, is now reckoned conspecific) is nearer to Mesopicos than to other African genera but in its other external characters it seems to be as close to Campethem. The two genera are, however, closely allied. It seems preferable to leave elliotii, at least provisionally, in Mesopicos rather than to place it in a monotypic genus. Mesopicos elliotii sordidatus (Bates, 1928) was described from a juvenile from Oku, west of Kumbo, Cameroons. Its supposed racial characters are merely those of the juvenile plumage. There are now two adults from Oku in the British Museum collection, these do not differe from examples of M. e. johnstoni of which I therefore consider M. e. sordidatus a synonym. THRIPIAS Peters (1948) includes the Bearded Woodpecker namaquus, the Fire-bellied Woodpecker pyrrhogaster and the Golden-crowned Woodpecker xanthocephalus in the genus Thripias. Many other authors, e.g. Hargitt (1890), Chapin (1939) and Grant & Mackworth-Praed (1957) have put Pyrrhogaster and xanthocephalus in Mesopicos, presumably mainly on account of their proportionately shorter wings (than those of namaquus) and maintained Thripias as a monotypic genus. I think Peter's arrangement is best. These three species all agree in having a relatively long and straight bill, black nuchal region in both sexes and similar characteristic facial markings. Against these I do not think that the proportionately longer bill and tail of namaquus warrants its generic separation. T. namaquus seems to link the other two Thripias species (as reckoned here) with Campethera rather than with Mesopicos. In this connection it may be mentioned that the female of Campethera bennettii has facial markings rather similar to, although not identical with, those of the Thripias species. Geocolaptes olivaceus I think the Ground Woodpecker is most closely related to other African forms and that its resemblance to some of the American flickers, Colaptes, (near which it is usually placed in systematic lists,) is due to convergent adaptations for terrestial or semi-terrestial life. It differs from the Colaptes species but very closely resembles 24 D. GOODWIN Mesopicos goertae and M. griseocephalus in colour pattern except for the barred tail and the vestigial red malar stripes of the male. Both these are, however, very widespread characters in " green " woodpeckers and occur, inter alia, in some Campethera species. It seems likely that olivaceus evolved from some form ancestral both to itself and to goertae and griseocephalus. It may well have retained (or developed) red malar stripes as a species-specific recognization mark in reference to griseocephalus (or its ancestor). The present apparently obsolescent state of the malar markings might indicate that the development of greater divergence in size and ecology has now obviated the need for such distinguishing markings between these two forms. Notes on Picus Peters (1948) includes in Picus the former genera Chrysophlegma — species mentalis andflavinucha, and the monotypic Callolophus — species mineaceus. Hargitt (1890) included the latter in Chrysophlegma also although he placed the smaller yellow- crested forms, puniceus, chlorolophus and chlorigaster with the " typical " green woodpeckers in the genus Gecinus. Within Peter's broad and inclusive conception of Picus the species fall into three rather distinct groups which are not precisely the same as those suggested by former allocations of species between Gecinus (or Picus} and Chrysophlegma. The latter genus was defined as having " wing rounded . . . nasal ridge almost obsolete, culmen ridge very blunt, tail two thirds length of wing " and later by Stuart Baker as having "... bill more curved and nasal ridge almost obsolete, well-developed nuchal crest. " These definitions have, apparently, restrained those who recognized Chrysophlegma from including in it the species chlorolophus chlorigaster and puniceus, which have rather straighter culmens that the other yellow-crested species. The group of most typical Picus species comprises viridis, canus, vaillantii, awokera, viridanus, vittatus, squamatus, xanthopygaeus , erythropygasus and rabieri. All these are relatively long-billed forms with fairly straight culman, nuchal crest absent or slight, primaries marked whitish and black or grey, and except in one race of canus, no red or chestnut colour on wings. P. rabieri has a more rounded wing and rather shorter bill than the others and approaches the next group in these features. The species in the second group are characterized by having yellow nuchal crests, shorter bills and, one species excepted, red coloration or at least a reddish tinge, on the wings. They comprise flavinucha and mentalis, which in addition to the characters listed above also have black and chestnut primaries and a spotted patch on the throat, and puniceus, chlorolophus and chlorigaster which have rather straighter culmens, no chestnut on their primaries which are dark and whitish, varying amounts of red on the head, and are smaller in size. To some degree these three forms link the original " Chrysophlegma " species to the first group but they are, I think, much more closely allied to the former. The third group consists of the single species mineaceus. This has a long and broad occipital crest, relatively short, broad bill, barred plumage pattern both above and below and rather harsh, coarse-feeling plumage. Although it may be justifiable NOTES ON WOODPECKERS 25 to include mineaceus in the present enlarged conception of Picus, (and certainly this is more reasonable than allying it with flavinucha and mentalis and yet not putting puniceus, chlorolophus and chlorigaster in the same genus), it is a very distinct species. It appears to be a link between Picus, via Picus puniceus, and the Rufous Woodpecker Micropternus to which it shows some approach in proportions, coloration, plumage pattern and plumage texture. I think it possible that more comprehensive studies might indicate that the species in this second group are less closely allied to those in the first group than are the latter to such Ethiopian genera as Campethera and Mesopicos. Picus canus In the eastern part of its range, where it and viridis do not overlap, the Grey-headed Woodpecker canus much more nearly approaches the Green Woodpecker viridis in size and proportionate length of bill. The only similarly-sized green woodpeckers which it overlaps in eastern Asia are P. erythropygius and P. rabieri, which are very distinct from canus, with striking colour differences. Picus dedemi of Sumatra seems rightly put as a form of canus in spite of its different colouring, with bronzy carmine, with a hint of green, replacing the green and olive of canus. The racial variation within the Paleartic populations of canus has been very fully discussed by Vaurie (1959). Picus vaillantii Vaurie (1965) put North African Green Woodpecker vaillantii, as a race of viridis and earlier (1959) anticipated this decision on the grounds that in everything except the black malar stripe of the male, which he considered an " alternate character " to the red and black malar stripe of the male viridis, P. viridis sharpei of the Iberian peninsula " bridges the gap " between viridis and vaillanti. He pointed out vaillantii had already been listed as conspecific with viridis by Goutenoire (1955) and that Dr. David Snow, who has observed this species in the wild, was also in favour of this decision. I cannot, however, see that sharpei is intermediate between [other races of] viridis and vaillantii except in bill shape and I do not think that minor details of size, size of bill or tone of general colour, in all of which vaillantii comes nearer to viridis than it does to European forms of canus, are of much significance when dealing with a geographically isolated " green woodpecker ". Especially in view of the amount of geographic variation shown by both viridis and canus and the obvious close relationship of the above three forms and the Japanese Green Woodpecker P. awokera. The female of P. viridis sharpei has the feathers of the forehead and crown red (or, to be more precise, these areas all tipped with red) as in other races of viridis, whereas these areas are without any red in females of vaillantii. Incidently, the red tips on the nape feathers of vaillantii, appear longer than and somewhat different in texture from those of viridis or canus. While vaillantii may be more closely allied to viridis than it is to canus or awokera, the differences in colour patterns of their heads, lack of red on malar stripe of male vaillantii and black and grey instead of red forehead and crown of female, might be 26 D. GOODWIN FIG. 6. Diagrammatic sketches to show colour patterns of heads of males (left) and females (right) of Picus viridis (top, Picus vaillantii (middle) and P. canus (bottom). pre-adapted to act as isolating mechanisms should the two ever overlap in range. Of its more striking plumage characters vaillantii shares one (crown and nape of male red) with viridis, two (no red on malar stripe of male, forehead and crown of female grey or grey and black) with canus and two (red on nape only of female, forehead and crown of female grey or black and grey) with the more geographically distinct awokera. I think it best, therefore, to give vaillantii specific status although it can, at least provisionally, be put in the same superspecies as viridis. Picus rabieri This is a very distinct species but, in spite of its rounded wings and somewhat different colour pattern, I think it is an offshoot of the viridis group although in wing shape and in its rather short bill it shows some approach (convergence) towards the flavinucha group. NOTES ON WOODPECKERS 27 Picus squamatus, P. viridanus, P. vittatus & P. xanthopygaeus These species form a closely related group and are essentially alike in coloration and colour pattern. The Red-rumped Woodpecker P. erythropygius is related to this group, as its colour pattern especially in the eye-striped morph, clearly indicates. The Scaly-bellied Green Woodpecker P. squamatus overlaps in part of its range with the Little Scaly-bellied Woodpecker P. xanthopygaeus from which it differs in being much larger, having very conspicuous black and white malar stripes, the upper breast plain greyish green without squamate markings and conspicuously barred central (and other) tail feathers. P. xanthopygaeus overlaps with the Burmese Scaly-bellied Woodpecker P. viridanus and marginally with the Laced Green Woodpecker P. vittatus from both of which it differs in having inconspicuous (almost obsolescent) malar stripes, more streaked (longitudinal squamate markings) throat and yellow or greenish yellow instead of yellowish green rump and upper tail coverts. P. vittatus and P. viridanus are extremely similar, differing in appearance only in viridanus having the streaky-looking squamate markings of the underparts extending forward onto the upper breast and throat where they are, however, less clearly denned than lower down. As given in many standard works the ranges of the two would appear to overlap in parts of Eastern Burma and south-western Siam. In the fairly extensive series of both in the British Museum (Natural History) there are no specimens of both species from any one area with the exception of one specimen of viridanus, allegedly from Tenasserim, where only vittatus normally occurs. I think this probably represents some error in labelling. Robinson & Kloss (1923) say that " where the ranges of the two species touch or approach there is not the slightest sign of intergradation ". In view of the extremely slight differences between them I do not find this statement very convincing. There are two specimens in the British Museum, No. 1887.8.10.1556 from Mergui and No. 1887.11.1.95 from " Burmah ", which are absolutely intermediate between " typical " vittatus and viridanus at first appearance. On closer examination they are seen to have squamate markings on the upper breast although these are fainter than usual in viridanus. It is significant, perhaps, that in all specimens of viridanus these squamate markings are least developed on the upper breast and throat, contrary to what one might expect if they functioned as a species recognition mark in reference to vittatus. Deignan (1963) puts viridanus as a race of vittatus and, although he does not there give his reasons for so-doing, all the evidence I have been able to find suggests that he is quite right in this decision. Picus chlorolophus & P. chlorigaster The number of races of the Small Yellow-naped Woodpecker chorolophus that has been described tends to obscure the main difference which is between the popu- lations in peninsular India and Ceylon and those from elsewhere. The former, at one time given specific status as P. chlorigaster, are not only geographically isolated but in some characters come closer to the related Crimson-winged Woodpecker P. puniceus 28 D. GOODWIN resembling it and differing from typical chlorolophus in having blackish lores, instead of whitish as in chlorolophus, and the top of the head entirely red. In the amount of red on their wings they are intermediate. The three forms, chlorigaster , chlorolophus and puniceus are allopatric except that an isolated race of chlorolophus (rodgeri) occurs in the mountains of Perak. There is thus a very limited geographical overlap of this form with puniceus although there is apparently no ecological or altitudinal overlap (Robinson, 1928). This montane race is rather dark in colour and thus superficially somewhat intermediate between chlorolophus and chlorigaster but in its main features it agrees with the former, with which its affinities clearly lie. I think it is preferable to treat chlorigaster as a member of a superspecies, together with chlorolophus and puniceus, rather than as a race of chlorolophus. DENDROCOPOS Voous (1947) shows that the American and Asiatic " ladder-backed " woodpeckers in this genus all inhabit areas that were not covered with Pleicestocene land-ice and which, in North America, represent the glacial forest refuges. He considers all these ladder-backed species to be (relatively) closely related in spite of their present discontinuous ranges. FIG. 7. Diagrammatic sketches to show colour patterns of heads of a typical Old World Dendrocopos (major], a typical New World ditto (villosus) and Pico'ides trydactylus (left to right) (In all species the heads shown are of females). All American ladder-backs agree with other American Dendrocopos species, and differ from most Old World Dendrocopos species, ladder-backed and otherwise, in lacking red on the ventral regions or under tail coverts. They also have a similar head pattern, with a dark post-ocular band, to other American Dendrocopos species. I think these facts indicate that all the New World Dendrocopos species are more closely related to each other phylogenetically than any one of them is to any Old World form. It seems more likely that the " ladder-back " pattern (so common in woodpeckers generally) has developed, or been retained if it was a primitive feature, independently in both Old and New Worlds, as an adaption to relatively open or well-lighted habitats in which it probably has protective value, than that some factor in the American environment has caused two or more phylogenetically distinct Dendrocopos stocks to lose all trace of red on the underparts and to acquire similar facial markings subsequent to their arrival in America. In this connection it is pertinent that in some species of Melanerpes the adults are plain backed and the juveniles ladder-backed NOTES ON WOODPECKERS 29 and in the Three-toed Woodpecker Picoides tridactylus some races are ladder-backed and others not (see Text-fig. 8). Of Old World species the " dwarf " woodpeckers, D. minor, D. kizuki, D. nanus et al., come nearest to the American forms in their colour patterns and are, I think, closest to them phylogenetically. FIG. 8. Diagrammatic sketches to show variation in back pattern within a species: Females of Pico'ides trydactylus from Sweden (left), South-eastern Tibet (middle) and eastern North America (right). The Hairy Woodpecker D. villosus and the Downy Woodpecker D. pubescens are now very largely sympatric. They presumably derived from a common ancestor at a relatively recent date as their colour patterns are virtually identical. They also show similar geographical variation although this, by itself, is of course, no argument for close relationship. Nuttall's Woodpecker D. nuttallii and the Cactus Woodpecker D. scalaris are also close relatives. They appear not to be sympatric and descriptions of their calls do not suggest any great difference of voice. They may prove to be conspecific but provisionally are, perhaps, best treated as members of a superspecies. The Red- cockaded Woodpecker D. borealis is, I think, closely allied to them. Its strikingly different facial pattern seems to have been achieved by the extreme reduction of the usual (in American Dendrocopos) black post-ocular stripe so that only a vestige of this marking remains. I think, however, that the great similarity in position and extent of the red on the head in this species and the Old World D. kizuki is due to convergence, within related stocks, and does not indicate that borealis is closer to 30 D. GOODWIN kizuki than are other American species. The juveniles of borealis and kizuki do not, apparently, resemble one another in position of red on the head as kizuki is said to resemble the adult in this respect. The Arizona Woodpecker D. arizonae and Strickland's Woodpecker D. stricklandi are allopatric, they are close to one another in appearance, voice, behaviour and ecology (Davis, 1965). On its upperparts stricklandi has a colour pattern intermediate between that of completely ladder- backed species and those, such as arizonae, with uniform dark back and rump. Individuals of arizonae which show traces of a barred dorsal pattern do, however, occur (Davis, 1965). The two are otherwise similar in colour but differ in stricklandi having dark streaks on the underparts with some indication of barring on the flanks, whereas arizonae is spotted, but the differences are less impressive when the markings on the individual feathers are compared. I think arizonae and stricklandi are best considered members of a superspecies. The two South American forms the Striped Woodpecker D. lignarius and the Chequered Woodpecker D. mixtus differ from all North American Dendrocopos species in having their central tail feathers strongly barred with white and they are generally lighter and more " spotty " in appearance. They show much resemblance in plumage pattern to Veniliornis spilogaster and I think this may indicate close relationship between Veniliornis and Dendrocopos. They agree, however, with the northern forms in general overall colour pattern. From their visual characters one would certainly suspect them of being conspecific but they apparently overlap in range, both being found in Nequen and the eastern Rio Negro in Argentina (Peters, 1948, Olrog, 1963) . Two specimens in the British Museum from Cordova, Argentina, (B.M. Nos. 1889.2.26.112 and 113) appear to be intermediate between D. lignarius and D. mixtus berlepschi. The male of the two agrees with mixtus in the extent of the red on its head but both are closer to lignarius in plumage pattern, which is, however, extremely similar in the two species (?) in any case. These two have longer bills than our specimens of berlepschi but we have only two of the latter and, at least in lignarius, this feature seems prone to individual variation. The White-headed Woodpecker Dendrocopos albolarvatus is rather aberrant in colour pattern and its tongue is said (Voous, 1947) to be less extensible than in other (all other?) Dendrocopos species. Its external features, particularly the typically dendrocopine correlation of a red nape in the adult adult male with a red crown and black nape in the juvenile indicate that Peters was right to include it in Dendrocopos. Dendrocopos darjellensis Measurements of our specimens seem to indicate a less clear-cut difference between nominate specimens of the Darjeeling Pied Woodpecker, darjellensis, and D. darjellensis desmursi (]. Verreaux) than did those measured by Vaurie (1959) We have, however, no topotypical specimens of desmursi from Sikang and only a few from Yunnan. Our only specimen of fumidus, from the Naga Hills, does not show the characters claimed for this race (Ripley, 1951) and thus tends to confirm Vaurie 's opinion that fumidus should be considered a synonym of darjellensis. This species overlaps widely with the Crimson-breasted Pied Woodpecker NOTES ON WOODPECKERS 31 D. cathpharius. The two are nearly identical in coloration but differ much in size. Their yellow-tinged cheeks and underparts, red breast bands and heavy streaking on the underparts are considered by Voous to be primitive characters. Whether this is so or not these characters might function now as species-specific signal markings. Dendrocopos leucopterus I concur with Voous (1947) and Vaurie (1959) in thinking that it is better to give leucopterus, provisionally, specific rank. This, however, is mainly on grounds of convenience. I think Vaurie has rather over-emphasised the differences between leucopterus and major. I have seen no juvenile specimens of leucopterus so do not know if I should think them so distinct from juvenile major as they are said to be. It seems, however, pertinent to remark that although, as Vaurie says, juvenile major of both sexes have red on the crown, that on the female is normally less extensive than the male's. So the fact of the juvenile female of leucopterus having (always?) an entirely black crown may not be of specific significance. Dendrocopos dorae Although at first glance the Arabian Woodpecker is rather like some of the species in the African genus Dendropicos it seems most closely allied to the Eurasian forms of Dendrocopos and is now rightly included in that genus. I think that dorae may be closest to medius ; to which it bears much resemblance when allowance is made for " fading " of the black areas to darkish brown (already often shown to some extent in some individuals of medius) and for reduction of the white areas. In size, texture and colour of the red crown feathers, and the streaking on its flanks it is very like medius. Indeed, the differences between these two woodpeckers are very similar in kind, although greater in degree, to those between the Arabian form of the Magpie Pica pica and its more northerly representatives. In dorae, like most Dendrocopos species but unlike medius, the female lacks red on the head but I do not think this feature of medius, which is unique within the genus, argues against their close relationship. D. medius and D. leucotos are unquestionably close relatives, but differ in this feature as they do in size and bill shape. Dendrocopos analis, D. atratus & D. macei I provisionally follow Peters & Deignan (1945) in treating analis as a form of the Fulvous-breasted Pied Woodpecker macei. They differ slightly in size, in analis having a spotted rather than a streaked breast, white ground colour on underparts and central tail feathers and upper tail coverts barred with white instead of uniformly black. They do not, however, differ in colour pattern of the head. Intermediates occur (fide Deignan, 1945) in the Chiang Mai area of Siam. Voous (1947) suggests that macei and the Stripe-breasted Pied Woodpecker atratus may be conspecific, quoting Standford & Ticehurst (1939) that where they appear geographically to overlap in range in northern Burma macei is a lowland and 32 D. GOODWIN atratus a highland form. He mentions specimens of macei obtained by Hume in Manipur, Assam and implies that these showed some intergradation with atratus. At least some of these Manipur specimens are in the British Museum collection and do not seem to differ from other specimens of macei, collected elsewhere. When they are compared with specimens of atratus, also collected in Manipur, the differences between them are at least as great as between specimens of both from elsewhere. That is, macei differs in being smaller, having an unstreaked lower throat, the underparts creamy rather than dusky yellowish and with not very prominent dark streaks, as against very prominent dark streaks in atratus. The red on the ventral region does not usually extend so far forwards, the red of the head is a little less bright and the red feathers are a little dissimilar in texture. These differences also hold for analis, which has, in addition, tail and rump feathers barred with white. D. analis and D. atratus overlap widely, geographically, but may be ecologically isolated. Thus some of the differences between analis and macei might have originated as species-specific recognition marks in reference to atratus. Dendrocopos major Vaurie (19590) puts the race parroti (Hartert) from Corsica as a synonym of the Sardinian form harterti (Arrigoni) . I prefer to recognize parroti as there appears to be very little overlap in bill length and the bills of all our specimens of parroti are noticeably more massive than those of specimens of harterti of the same sex. In view of the only slightly differentiated and intergrading continental races of this woodpecker that are currently recognized it seems better not to lump these two island forms. Vaurie considers it most probable that D. m. tianshanicus represents inter-specific hybridization between major and leucopterus. With one exception our specimens all seem nearer to major and I provisionally treat them as a form of major and not as hybrids. The exception to this, mentioned above, is a female from the Tekkes Plain (B.M. No. 1931.7.8.259) which was originally identified as leucopterus and later re-identified by Vaurie as tianshanicus, because it had a wing measurement of 137 mm. as against 120-129 mm. for 23 females of leucopterus measured by Vaurie and because he considered the white spots on its wings too small for leucopterus. I think this bird was correctly identified originally. The very large measurement of its right wing appears to be due to the make-up of the skin. The left wing measures only 130 mm. and n other females of leucopterus, picked at random, have wing measurements ranging from 125 to 131 mm. The white spots on its wings do not appear to be significantly smaller than those of some other specimens. PICOIDES I agree with Delacour that this genus is very close to Dendrocopos. Its lack of red pigments, or rather their replacement by yellow; its flatter bill and its lack of a fourth toe are, in combination, perhaps just sufficient grounds for keeping it in a separate genus. It is certainly convenient to do so and the two Pico'ides species are, unquestionably, much more closely related than either is to any other. NOTES ON WOODPECKERS 33 However, although Pico'ides seems to have diverged a little further from the Dendrocopos species in the course of evolution than these have from one another it is, I think, likely that it may be phylogenetically closer to the American Dendrocopos species than these are to Old World forms. The colour pattern of Pico'ides tridactylus is very close to that of Dendrocopos villosus. This is especially so with the northern Old World forms of tridactylus. The relatively greater amount of difference between the New World forms of tridactylus and villosus may be due to character displacement although in some other Holarctic species or superspecies there is a tendency for the New World representatives to show a greater amount of melanin pigment than the Old World forms (e.g. Numenius phaeopus, Branta bernicla). In this connection it is of interest that nominate P. t. trydactylus has a broad white stripe down its back like D. villosus whereas the American forms, e.g. P. t. fasciatus, show white cross-barring very like that of the " ladder-backed " Dendrocopos species and similar in position and extent to that of D. stricklandi. The Arctic Three-toed Woodpecker P. arcticus, which is entirely American in distribution, has a wholely black back with, sometimes, a hint of barring indicated by white tips to one or two feathers. In the large amount of black in its plumage generally arcticus stands in about the same relation to American forms of tridactylus as these latter do to most of the Old World forms of their species. Although now largely sympatric with the smaller tridactylus, arcticus must, presumably, have evolved in isolation from tridactylus or proto-tridactylus . The very dark, and now apparently geographically isolated, Tibetan form of tridactylus, P. t. funebris, is suggestive in this connection as it shows similar, and nearly as great, contrast to the northern Old World forms of its species as does arcticus to American forms of tridactylus. Sapheopipo noguchii The Okinawa Woodpecker, originally included in the genus Picus (Seebohm, 1887), was placed by Hargitt (1890) in the monotypic genus Sapheopipo, which he having: " wing very long and pointed; tail only equal to the second primary and falling short of the longest by about an inch ". It is found only in Okinawa, in the central Ryukyu Islands, south of Japan. The general coloration of noguchii is dark olive, more or less suffused with red, especially on the tips of the feathers, on back, rump and underparts; with dark brownish-black wings and tail. It is thus, superficially, most like the dark red Sumatran form of Picus canus, P. c. dedemi. It has, however, very clear indications (see Text-fig. 9) of head and breast markings nearly identical to those of many Dendrocopos species and quite unlike those of any species of Picus or Dinopium. Also those parts of the primaries that are not visible, or at any rate would not be conspicu- ous, when the wings are closed have white markings similar to and I think homologous with those of D. leucotos owstoni, the very dark race of D. leucotos from the northern Ryukyu islands. The legs and irides of noguchii and owstoni agree in colour although their bills do not (Kuroda, 1925). 34 D. GOODWIN The difference of coloration between leucotos and noguchii is very similar to what one finds between many other species and their island derivatives, for example the pigeons Columba arquatrix and C. pollenii. They seem to have involved, assuming noguchii to be derived from Dendrocopos stock, simply a spread of dark pigment together with some lessening of its intensity and the spread into some of these dark areas of red or yellow pigments; or, perhaps, their revelation through their no longer being completely masked by melanin pigments. The pale colour of the bill might be similarly caused. In proportion and in the shape and structure of its bill noguchii is quite close to leucotos although its bill is proportionately longer, a common feature with island forms. Its type of sexual dimophism is identical with that of leucotos and several other Dendrocopos species. D. leucotos owstoni is, presumably, a more recent invader of the Ryukyu Islands that is already beginning to evolve in the same direction as has noguchii. FIG. 9. Diagrammatic sketches to show resemblances and differences of colour patterns of Dendrocopos leucotos from northern Europe (left), same species from northern Ryu-kyu Islands (middle) and Sapheopipo noguchii (right). I think that noguchii is an offshoot of Dendrocopos stock and is rather more likely to have derived from Dendrocopos leucotos subsequent to the speciation of the latter than from a more primitive common ancestor to them both. It is said (Kuroda, 1925) to be found on or near the ground in damp woods or bamboo jungle and never in tall trees during the day (he does not say whether it roosts in tall trees but this is, perhaps, implied). This suggests that noguchii' s ecology may have already diverged considerably from that of typical Dendrocopos species and this, together with its unusual coloration, may, perhaps, justify retention of the monotypic genus Sapheopipo. NOTES ON WOODPECKERS 35 If, however, further studies should show that it does not differ so markedly in habits as Kuroda's notes suggest and that there are no striking anatomical differences I would be in favour of treating Sapheopipo as a synonym of Dendrocopos. CHRYSOCOLAPTES & DINOPIUM Delacour (1951) suggested that the absence of the fourth toe is not of great phylo- genetic significance in woodpeckers and that in this character Brachypternus with its greatly reduced fourth toe, placed by Peters (1948) in the synonymy of Dinopium, forms a link between the three-toed Dinopium species and Chrysocolaptes with four well-developed toes. I agree with his opinion on both points; there can be no doubt that the species comprising the above genera are all closely allied. I provisionally maintain the genus Chrysocolaptes mainly on grounds of taxonomic convenience. This can, perhaps, be further justified by the fact that the three species involved — lucidus , festivus and validus seem very close to each other. All, apart from having four toes, agree in having some white on hind neck and mantle, admittedly vestigial in lucidus, almost identical facial markings which are, however, approaching obsolescence in validus, and bills that are longer and more massive than those of Dinopium. Except in the case of Dinopium javanense and Chrysocolaptes lucidus, wherever two species of either (or both) of these two genera overlap widely in range they show fairly striking differences of colour pattern. The very similarly-coloured Golden-backed Three-toed Woodpecker, D. javanense and the Golden-backed Woodpecker C. lucidus are extensively sympatric and, in general, show similar geographic variation. They are most alike in western India, Burma and much of Malaysia where, except for the not very striking difference in the pattern on face and throat, colour differences between them are slight or non-existant. C. lucidus is, however, a larger bird with proportionately much larger bill. In other areas where they overlap the two show greater differences in coloration. On Java lucidus (race strictus) has a female with the top of the head yellow and rather obscurely spotted, and both sexes have only a tinge of red on the rump, whereas the javanense female has a black and white streaked crown and both sexes have a conspicuous red rump. On Palawan lucidus has a very distinct race (erythrocephalus) in which both sexes have a red face and pink throat, with the usual dark striations rather reduced, the female has the top parts of her head a mixture of red and yellow-olive with light dull yellow spots, the individual feathers being red near the base, then olive with a yellow terminal spot. The Palawan form of D. javanense (everetti) is very distinct, having a boldy striped buffish and black face and the female a black forehead and crown and red nape. It is noteworthy, however, that, compared with other forms of javanense, this race shows parallel development to lucidus on Palawan, in having more red on the head than is usual in the species : females of other forms of javanense have no red on the nape and the males no red on the malar region as has everetti. They also agree in having the dark and light facial markings less pronounced than in most other forms of their respective species. 36 D. GOODWIN Chrysoclaptes lucidus I provisionally follow Peters (1948) in his wide conception of this species because, on external characters, it is not possible to make any logical division among the many races or presumed races of lucidus. It is, however, quite possible that some of the island forms, especially in the Philippines, may have already reached specific level. Like previous workers, I cannot separate specimens from South India from those from southern Malaya although the latter are, on average, a slightly lighter golden- yellow on the upperparts. There seems no alternative, therefore, but to continue to employ the name chersonesus for both these populations although it is likely that, in a phylogenetic sense, chersonesus from southern India are more closely allied to guttacristatus from Bengal than they are to chersonesus from the Malay Peninsula. On the different Philippine islands the races (?) of lucidus show great variation of head colour (besides other differences). These are comparable to the differences of head colour between undoubted species of the Chrysocolaptes-Dinopium group elsewhere. This is no doubt due at least partly to the lack of any closely related sympatric species on most of these islands. Nominate lucidus from Mindanao is, perhaps rather unfortunately, a rather nondescript or intermediate form and as it has indistinct facial markings, mixed red and yellow upperparts and the top of the female's head yellowish and spotted it is difficult to ally it with any one, or even any one of the main groups of Philippine forms rather than another. The interesting sequence of the immature plumages of guttacristatus (and other races of lucidus!} is described in the section on juvenile plumages. Dinopium shorii & D.javanense The Himalayan Golden-backed Three-toed Woodpecker and D. javanense are very closely allied and must have derived relatively recently from a common ancestor but they appear now to overlap without interbreeding in northern Burma. There are specimens of both from Thayetmayo and the Arrakan Hills, in the British Museum. They appear to keep or even slightly to accentuate their minimal plumage differences — pale bases to the red head feathers in shorii, slightly different pattern of streaks on the throat — where they overlap. Admittedly hybrids between the two would be difficult to recognize. There is a slight difference of size and bill length between them and, no doubt, this correlates with differences of ecology. Dinopium rafflesii I concur with Peters in thinking it preferable to put the monotypic genus Gauropicoides (species rafflesii) into the synonomy of Dinopium. The Olive-backed Three-toed Woodpecker, rafflesii, seems to link Dinopium with Gecinulus, agreeing with the former in most characters but with Gecinulus to some extent in its softer feather texture and generally dark and rather concolorous body plumage. The wing markings are similar in both genera. NOTES ON WOODPECKERS The genus GECINULUS 37 I maintain this genus for the Pale-headed Woodpecker grantia and the Green Bamboo Woodpecker viridis. These two closely allied forms differ from the green woodpeckers of the genus Picus in having proportionately small bills and only three toes ; also in some plumage characters in which they are closer to Dinopium. Although rather discrete they appear, as has been mentioned above, to be linked with Dinopium by rafflesii and probably represent a link between Picus and the Dinopium- Chrysocolaptes group. DRYOCOPUS & PHLOEOCEASTES Peters (1948) placed the American species pileams, lineatus, erythrops and galeatus in the genus Dryocopus together with the old world forms martius and javensis. Presumably he did this because their outer hind toes are not longer than their outer front toes (Peters, 1948, introduction, p. 6). The characters of a narrower nasal shelf and nostrils nearer the culmen, which he says are correlated with the shorter outer hind toe, do not appear to be valid as there is much variation between species; on these characters lineatus, at least, seems to fit better with those species which have a longer outer hind toe. I do not think this relative length of the outer hind toe is, FIG. 10. Diagrammatic sketches to show colour patterns of heads of Phloeoceastes lineatus (top left), Dryocopus javensis (bottom left), Picus viridis (bottom middle), Phloeoceastes haematogaster (top right) and P. melanoleucos (bottom right). 38 D. GOODWIN by itself, a very important character. Their distribution and their plumage characters, especially the colour patterns of their heads all suggest that the American species placed by Peters in Dryocopus are closer to the species included by him in the genus Phloeoceastes than they are to Old World forms. I therefore think it best to place them in Phloeoceastes and restrict Dryocopus to the two Old World species martins and javensis. I think it possible that Dryocopus, as above restricted, might prove to be more closely related to other Old World genera, such as Mulleripicus and Picus. Kilham (1959) was impressed by the great similarity between some of the behaviour patterns he had observed in pileatus and some described for martins (Blume, 1956) but comparisons of these two with both (other) Phloeoceastes species and Mulleripicus are needed. Most of sketches and descriptions in Blume's comparative study of European woodpeckers (Blume, 1961) suggest to me that the displays, and possibly even the calls, of martins bear a more close resemblance to those of Picus viridis than to those described for pileatus by Kilham. Phloeoceastes divides readily into the lineatus species-group comprising lineatus (syn. erythrops), pileatus, schulzi and galeatus, all of which have rather softer, denser plumage with consequently longer nuchal crests and tend to have proportionately smaller bills, and the melanoleucos species-group, comprising melanoleucos , guatemalensis , pollens, haematogaster , rubricollis, robustus and leucopogon. Within the lineatus group pileatus, lineatus and schulzi are closer to each other in colour patterns than any of them is to galeatus. This may give a false impression, as the very differently coloured (largely buffish brown) head and underwing of galeatus may be the result of selection for species-specific markings in reference to lineatus which it overlaps widely in range. P. pileatus does not overlap any congener and schulzi would appear to have only a marginal overlap with galeatus and lineatus at the northern edge of its range. Both pileatus and schulzi have concolorous black underparts unlike any other close relatives. This might, however, be due to some environmental factor as these two are in predominently temperate climates north and south, respectively, of the ranges of other Phloeoceastes species. The species in the melanoleucos group all appear to be very closely related to each other. Within this group it is possible that relatively greater differences in colour pattern between any two species may not indicate comparable phylogenetic disparity as such differences may have evolved as isolating mechanisms in reference to each other. Although all species show differences in colour pattern these are, on the whole, most marked, and often correlated with differences of body proportions, in species that overlap widely in range. It is notable that both this and the lineatus group have evolved some forms predominantly white and others predominantly buff or chestnut on the under wing. I follow Peters in thinking it best to give specific rank to both guatemalensis and melanoleucos, although they can certainly be considered as members of a superspecies. Further studies may, however, show that they are in fact conspecific and if so the rather striking differences in colour patterns of their heads may be a case of character displacement as melanoleucos overlaps in range with robustus whose male has an almost entirely red head like that of guatemalensis. NOTES ON WOODPECKERS 39 Pholeoceastes lineatus & P. erythrops Peters (1948) says that it has been suggested that erythrops is a subspecies of the Lineated Woodpecker lineatus but that, if specimens have been correctly identified, they overlap over a wide area. He treats erythrops as a good species but remarks that " The inter-relationships of the lineatus-erythrops-schulzi group are quite involved and require additional field study ". Van Rossem (1934) and De Schauensee (1966) have both presented evidence that erythrops and lineatus are conspecific. I have compared specimens of both from Sapucay, Paraguay and they differ in no character except the presence (in lineatus) or absence (in erythrops) of white on the scapular region. Van Rossem states that some specimens of erythrops show white markings on the shoulder region and one of our specimens has one white-edged feather in this region. I think the evidence suggests that lineatus is dimorphic over the southern-most part of its range and that erythrops is merely a black-shouldered morph of this species. CAMPEPHILUS & IPOCRANTOR Hargitt (1890) followed Cabanis & Heine (1863) in placing the Magellan Woodpecker in the monotypic genus Ipocrantor; Peters (1948) placed it with the ivory-billed woodpeckers in the genus Campephilus. Peters was, presumably, led to this because its rather large size (in comparison with Phloeoceastes species), soft, glossy black plumage, large white wing patches and the long crest of the female combine to give it a very similar general appearance to an ivory-bill. Although it is possible that magellanicus does represent a divergence from the same stock as produced the ivory- bills, subsequent to the latter's splitting off from ancestral Phloeoceastes, I think this rather unlikely. The coloration and colour-patterns of the heads of both magellanicus and the ivory- bills are closer to those of Phloeoceastes than they are to each other. In both magellanicus and the ivory-bills the colour pattern of the head seems to represent a modification and some degree (considerable in magellanicus) of simplification of a pattern typical of Phloeoceastes. This simplification has involved a reduction in the amount of red in the male ivory-bills and its increase in the male of magellanicus. The white wing patch of magellanicus might represent a restricted version of that found in the ivory-bills but could equally (in both) be a development from the white or light bars that some Phloeoceastes species show on (some of) the inner webs of the primaries and secondaries. The under wing pattern of magellanicus is closer to that of Phloeoceastes melanoleucos and P. guatemalensis than it is to that of Campephilus. The bill of magellanicus is dark, not white or cream-coloured. Some species of Phloeoceastes have pale bills (others have dark bills). It is, I think, likely that the features in which magellanicus agrees more closely with the ivory-bills than it does with Phloeoceastes species: length of crest of female, relatively lax, glossy plumage and rather long tail (all, probably, correlated charac- ters) are due to convergence in related stocks but that each is, phylogenetically, at least as closely related to Phloeoceastes as to the other. I think, therefore, that it is preferable to retain the monotypic genus Ipocrantor for magellanicus. The only 40 D. GOODWIN reasonable alternative, and one that seems to have much to recommend it, would be to place all these large, related, predominantly black, white and red woodpeckers in the single genus Campephilus. I feel, however, that it would be premature for me to do so now in view of the work currently being done in the U.S.A. on the habits and ecologies of the American woodpeckers which will, doubtless, soon provide other and perhaps firmer grounds for taking or refraining from this course. Variant individuals The commonest plumage variation in woodpeckers, occurring in individuals of many green-backed and golden-backed species, is for the green or golden parts of the plumage, especially on the mantle, to be more or less suffused or mottled, with red. This correlates with the existence of several red-backed forms closely related to or even conspecific with green-backed and golden-backed forms. Other variants in the British Museum (Nat. Hist.) collection are listed and described below. Picus vittatus A male from Pulau Langkawi, Malay Peninsula (B.M. No. 1936.4.12.586) shows an apparent reduction of red and yellow pigments. Its breast, underparts and sides of face are paler and more buffy than in normal specimens. The green of its upper- parts is a colder hue, less tinged with yellow, and the red parts of its head a light flame-orange instead of the deep scarlet-crimson of normal birds. Dendrocopos assimilis A male from Khipri, Sind (B.M. No. 1898. 12. 12. 331) has the red on the head replaced by straw yellow except for the nuchal region and a line extending thence above each eye, where it is orange-red, not the normal dark scarlet. Dendrocopos himalayensis A supposed male from near Simla (B.M. No . 1887 .8.10.7) has the belly, flanks and much of the lower breast red, instead of this colour being confined to the under tail coverts and post-cloacal area as in normal specimens. Surprisingly, if it is correctly sexed, it shows much less red on the head than normal, this being confined to the tips of only about a third of the crown feathers. It would seem strange if the bird exhibited an excess of red pigment in one area yet a decrease of it elsewhere and I am inclined to think that it may be a female, in which case (the female having normally no red on the head) its head coloration would, like that of its underparts, represent an increase of red. Dendrocopos syriacus An adult female (B.M. No. 1888. 11.1.564) shows partial albinism. It has many white or partially white feathers on normally black areas of head, back and wings. The distribution of red is normal. NOTES ON WOODPECKERS 41 Dendrocopos atratus A presumed female from the Southern Shan States of Burma (B.M. No. 1903. 12. 24.53) shows an interesting coloration as a result of partial loss of melanin pigment. It gives a general impression of having most of the normally black areas replaced with light grey. More closely examined it shows, on its upperparts, a narrow edging of more intensely pigmented (darkish grey) plumage posterior to every white marking (except where this comes to the end of the feather) and a similar but broader dark edge immediately anterior to it. This is most clearly seen on the secondaries but holds good for all other feathers. On the outer webs of the outer primaries the dark grey areas are more extensive. The tail shows ill-defined light and darker grey transverse barring but the modified, supporting, central feathers have their tips and the edges of both webs more heavily pigmented, a dark brownish grey. The top of the head is noticeably darker than most of the back, being, at least so far as the tips of most feathers are concerned, a very brownish grey, almost blackish. Due, presumably, to the reduction of melanin the underparts look much paler and more yellowish than normal, the dark streaks are greyish brown, not brownish black, and the red under tail coverts lighter and brighter. It would be interesting to know whether the areas (tail tips, primary edges] etc.) which show stronger pigmentation in this individual also have in fact more melann in normal birds where it is not visually evident. As they all seem to be areas that are likely to need " extra " strengthening, this would seem probable. Dinopium benghalense A male from southern Konkan (B.M. No. 1887. 8. 10.1957) and a male from Khandeish (B.M. No. 1880. 1.1.225) both have many red-tipped feathers in the malar region, giving much the impression of the type of black-enclosed red malar stripe found in males of Picus viridis. Normally D. benghalense has no red in the malar region in any plumage phase. A female from Sind (B.M. No . 1941 . 5 . 30 . 1391) shows reduction of red and yellow pigments. The normally golden parts of the mantle are a pale straw yellow, the normally scarlet nuchal crest more or less cream, intermixed with some partially orange-pink feathers. A female from Lucknow (B.M. No . 1887 . 8 . 10 . 1932) although normal in coloration, seems worth mentioning here. It has an abnormally long bill with decurved culmen which looks more like a Hoopoe's bill than a woodpecker's. The culmen measures 64 mm., as against 36 to 38 in normal females, and overlaps the under mandible by 7 mm. The bill tips are narrow and flexible and it is impossible to believe that the bill could have been used at all for wood-pecking. The bird had evidently been able to obtain food as is adult and in good plumage. Phloeoceastes lineatus Male from eastern Ecuador in post-juvenal moult (B.M. No . 1940 . 12 . 5 .93). Has the central part of the crown and nuchal region bright straw yellow, tinged orange, 42 D. GOODWIN instead of red. The red on the forehead and sides of head is a little lighter in hue than usual. Dinopium javanense Three females of the race intermedium show some trace of red in the head plumage. No. 1921.12.31.92, from western Siam has several male-type red feathers, two of them long nuchal feathers, on the left side of its head. No. 1927.6.5.419, from Tonkin, has one red-tipped but otherwise female-type feather above its left eye and No. 1927 .6.5. 1587, from Cochin-China, has several red-tipped but otherwise sexually intermediate-looking feathers. Many of the above variants involve partial loss of pigmentation and are comparable in appearance, and probably in origin, to relatively common varieties in other groups. It is, however, of interest that where this involves only the partial substitution of red by yellow, as in the Dendrocopos assimilis and the Phloeoceastes lineatus, it is comparable to one of the more conspicuous differences in coloration often found between related species of woodpeckers. This, of course, is also true of the second type of variant included above, where individuals show red on areas not normally so-coloured in their sex or species. SUMMARY Some aspects of taxonomy and relationships of the woodpeckers are described and discussed. Certain close resemblances in coloration and ecology are due to conver- gence. In all geographic regions inhabited by woodpeckers the smallest species are about the same size ; the same is true for the largest species except that there are no large arboreal species in the Ethiopian region. The typical forms of sexual dimor- phism and the exceptions to them are described. Juvenile woodpeckers usually have bright pigments similar to, but not always identical with, those of the adults. When all species are considered, the juveniles approximate more closely to the family average in the amount of red or yellow on the head than do the adults. In no instance, however, is the juvenile plumage considered to be more conspicuous than the adult male's. Leuconerpes is treated as a synonym of Melanerpes. The species obsoletus is considered to be referable to Dendropicos not Dendrocopos. Geocolaptes is related to other African genera, not to Colaptes. It is thought that all the New World species of Dendrocopos are more closely related to each other than any one of them is to any Old World species. Sapheopipo is a recent derivative of Dendrocopos. Some of the Philippine Island races of Chrysocolaptes lucidus show colour differences comparable to those shown by different species of the Chrysocolaptes- Dinopium group elsewhere. The New World forms currently placed in Dryocopus seem better placed in Phloeoceastes. Unless all the large black, white and red American wood- peckers are to be treated as congeneric, it is best to retain the monotypic genus Ipocrantor for the species magellanicus. NOTES ON WOODPECKERS 43 REFERENCES BANNERMAN, D. A. 1933. The birds of tropical West Africa. BATES, G. L. 1928. Bull. Brit. Orn. Cl. 49 : 33. — 1932. Bull. Brit. Orn. Cl. 53 : 74. 1937. On certain genera of woodpeckers. Bull. Brit. Orn. Cl. 58 : 15-17. BENSON, C. W. 1952. Notes from Nyasaland, Ostrich 23 : 144-159. BENT, A. C. 1939. Life histories of North American woodpeckers. Smith. Inst. U.S. Nat. Mus. Bull. 174. BLUME, D. 1956. Verhaltensstudien an Schwarzspechten (Dryocopus martins). Vogelwelt 77 : 129-151. 1961. tiber die Lebensweise einiger Spechtarten. /. Orn., 102, Sonderheft 1-115. CABANIS, J. & HEINE, F. 1863. Mus. Hein., 4 : 99. CHAPIN, J. 1939. Bds. Belgian Congo 2, Bull. Amer. Mus. Nat. Hist. LXXV. DAVIS, J. 1965. Natural history, variation and distribution of the Strickland's Woodpecker. Auk. 82 : 537-590. DEIGNAN, H. G. 1963. Checklist of the birds of Thailand, U.S. Nat. Mus. Bull. 226. DE SCHAUENSEE, R. M. 1966. The species of birds of South America with their distribution. Pennsylvania. DELACOUR, J. 1951. The significance of the number of toes in some woodpeckers and kingfishers, Auk 68 : 49-51. GOUTTENOIRE, G. 1955- Inventaire des Oiseaux de Tunisie. Alauda 23 : 1—64. GRANT, C. H. B. & MACKWORTH-PRAED, C. W. 1953. A new race of woodpecker from Portuguese East Africa. Bull. Brit. Orn. Cl. 73 : 55. GRISCOM, J. C. & GREENWAY L. 1941. Birds of lower Amazonia. Bull. Mus. Comp. Zool. Harvard 88 : 83-344. GROTE, H. 1928. Ubersicht iiber die Vogelfauna des Tschadgebiets. /. Orn. 76 : 739-783. HARGITT, E. 1890. Cat. Bds. Brit. Mus., 18. HAVERSCHMIDT, F. 1956. Was ist Melanerpes hargitti Dubois? /. Orn. 97 : 411-414. KILHAM, L. 1959. Behaviour and methods of communication of Pileated Woodpeckers. Condor 61 : 377-187. KIPP, F. A. 1956. Progressive merkmale des Jugendkleides bei den Spechten. /. Orn. 97 : 403-410. KURODA, N. 1925. Avifauna of the Riu Kiu Islands and the vicinity, Tokyo. LAUBMANN, A. 1930. Vogel in: Wissenschaftliche Ergebnisse der Deutschen Gran Chaco- Expedition. Stuttgart. MACKWORTH-PRAED, C. W. & GRANT, C. H. B. 1957. African handbook of birds, Ser. i, Vol. i. (sec. ed.). NEUMANN, O. 1903. Neue africanische Species und Subspecies. Orn. Monatsber. 11 : 180- 185. OLROG, C. C. 1963. Lista y Distribucion de las Aves Argentinas. Op. Lill. 9 Universidad Nacional de Tucuman. PETERS, J. L. 1948. Check-list of birds of the world, 6. London. REICHENOW, A. 1900. Uber Mesopicos goertae und Verwandte. Orn. Monatsber. 8 : 58-59. ROBINSON, H. C. & KLOSS, C. B. 1923. The birds of South-West and Peninsular Siam. Journ. Nat. Hist. Soc. Siam 5 : 171-173. SCHARNKE, H. 1931. Beitrage zur Morphologic und Entwicklungsgeschichte der Zunge der Trochilidae, Meliphagidae und Picidae. /. Orn. 79 : 425-491. SEEBOHM, H. 1887. Notes on the birds of the Loo-choo Islands. Ibis 5, 5th Scr., : 173-182. SELANDER, R. K. 1966. Sexual dimorphism and differential niche utilization in birds. Condor 68 : 113-151. SELANDER, R. K. & GILLER, D. R. 1963. Species limits in the woodpecker genus Centurus (Aves). Bull. Amer. Mus. Nat. Hist. 124 : 213-274. SIELMANN, H. 1959. My year with the woodpeckers. London. 44 D. GOODWIN SIBLEY, S. G. 1953. The abbreviated inner primaries of nestling woodpeckers. Auk 74 : 102- 103. STANFORD, J. K. & TICEHURST, C. B. 1939. On the birds of northern Burma. Ibis 3, i4th set. : 1-45. STRESEMANN, E. 1924. Mutationstudien 16 : Melanerpes rubrifrons (Spix). /. Orn. 72 : 422- 423- STRESEMANN, E. & V. 1966. Die Mauser der Vogel. /. Orn. 107. Sonderheft. VAN ROSSEM, A. J. 1934. Notes on some races of Ceophloeus lineatus. Trans. San Diego Soc. Nat. Hist. 8, 1934 : 9~12- VAURIE, C. 1959. Systematic notes on Palearctic birds. No. 34 Picidae: The genera Picus and Dryocopos. Amer. Mus. Novit. No. 1945. - 19590. Systematic notes on Palearctic birds. No. 35 Picidae : The genus Dendrocopos (Part i) Amer. Mus. Novit. No. 1946. — 1965. The Birds of the Palearctic fauna. Non-Passeriformes. London. VERHEYEN, R. 1957. Bijzonderheden over het jongkleed der speckten (Picidae). Gerfaut 47 : 177-182. WETMORE, A. 1926. Observations on the birds of Argentina, Paraguay, Uruguay, and Chile. U.S. Nat. Mus. Bull. No. 133. ,rx. *SK<*»£ PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING IB A REVIEW OF THE IGUANID LIZARD GENERA URACENTRON AND STROBILURUS R. ETHERIDGE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 17 No. 2 LONDON: 1968 A REVIEW OF THE IGUANID LIZARD GENER URACENTRON AND STROBILURUS BY R. ETHERIDGE Department of Zoology, San Diego State College, San Diego, California, U.S.A. Pp. 45-64 : 2 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 17 No. 2 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. 17, No. 2 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) 1968 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 19 November, 1968 Price Eight Shillings A REVIEW OF THE IGUANID LIZARD GENERA URACENTRON AND STROBILURUS By RICHARD ETHERIDGE SYNOPSIS The external morphology, distribution, and systematics of the iguanid lizard genera Uracentron and Strobilums are reviewed. Four species of Uracentron and one species of Strobilurus are recognized, and a key is given to the species of Uracentron. Two species erroneously referred to Uracentron are discussed . INTRODUCTION Uracentron and Strobilurus are South American iguanid lizards belonging to a group of genera that I have called the " tropidurines ", (Etheridge, 1964 : 629). Within this group Uracentron is easily identified by its very large interparietal scale, and short, spiny tail, and by the absence of a row of scales aligned mid-dorsally. Strobil- urus is closely allied to it, and like Uracentron has a very large interparietal scale and a spiny tail ; it differs in having a row of scales aligned middorsally. Also, the tail of Uracentron is flat, nonautotomic, and scarcely more than half as long as the head and body, whereas the tail of Strobilurus is nearly cylindrical, autotomic, and about as long as the head and body. Of the species we now know under the genus Uracentron the first was described as Lacerta azurea by Linnaeus (1758 : 202). Prior to this the species had been figured by Seba (1734 : 2 : 62 : 6), and Linnaeus (1754 : 42) had described in some detail specimens then in the Museum of King Adolf Fredrik of Sweden. Lacerta azurea was subsequently combined with diverse other lizards that also have a spiny tail under the genera Stellio (Latreille, 1802 : 29 and 34 ; Daudin, 1802 : 36 and 40 ; Fitzinger, 1826 : 49), and Uromastyx (Merrem, 1820 : 56-7). The generic name Ura- centron was introduced by Kaup (1826 : 88), and in the following year he included Uracentron in his list of the New World lizards (Kaup, 1827 : 612). The latter publication was cited as the original description of the genus by Burt & Burt (1933 : 48). Wagler (1830 : 145) changed the spelling to Urocentron, and with few exceptions (Mertens, 1925 : 75 ; Dunn, 1944 : 89 ; Valdivieso & Tamsitt, 1963 : 31 ; Peters, 1967 : 36) all later authors either adopted the change, or altered it still again to Uranocentron (Gray, 1831 : 42, 1845 : 225; O'Shaughnessy, 1881 : 245), or to Urocentrum (Boulenger, 1894 : 729; Werner, 1900 : 4). Cuvier (1829 : 34) proposed Doryphorus as a substitute name, and for a time it was also in use (Schinz, 1835 : 92 i Guerin-Meneville, 1829-1838 : 8 ; Dumeril & Bibron, 1837 : 369 ; Guichenot, 1855 : 26 ; Dumeril, 1856 : 559 ; Cope, 1870 : 556). Boulenger (1885 : 182-184) recognized three species of Uracentron : azureum (Linnaeus, 1758), flaviceps (Guichenot, 1855), and castor (Cope, 1870), and later he described a fourth species, U. guentheri (Boulenger, 1894 : 729). A fifth species was described by Mertens (1925 : 75), U. werneri. Burt & Burt (1933 : 48-9) recognized five species of Uracentron in their Checklist of South American Lizards. They included U. meyeri (Werner, 1900 : 4), and U. palluma (Tschudi, 1845 : 35), ZOOL. iy, 2. 2§ 48 R. ETHERIDGE neither of which is correctly referred to the genus Uracentron, and they failed to include U. flaviceps and U. werneri. In the present study U. meyeri and U. palluma are removed from the genus Uracen- tron, U. castor is placed in the synonomy of U. flaviceps, and this species together with U. azureum, U. guentheri, and U. werneri are considered valid species. Strobilurus is, and has always been, considered monotypic, with the single species 5. torquatus (Wiegmann, 1834 : I8). ACKNOWLEDGEMENTS During the course of this study I have had the good fortune to work in a number of museums in Europe and in the United States. I should like to express my appre- ciation to those curators who have allowed me to examine the collections in their care : Mr. C. M. Bogert, the American Museum of Natural History, New York ; Dr. E. E. Williams, the Museum of Comparative Zoology, Harvard ; Dr. R. F. Inger, the Field Museum of Natural History, Chicago ; Dr. A. Leviton, the California Academy of Sciences ; Dr. J. A. Dixon, the Los Angeles County Museum ; Miss A. G. C. Grandison, British Museum (Natural History), London ; Monsieur J. Guibe, Museum National d'Histoire Naturelle, Paris; Dr. K. Klemmer, Natur- Museum und Forschungs-Institut Senckenberg, Frankfurt ; Dr. J. Eiselt, Natur- historischen Museum, Vienna ; Dr. C. Edelstam, Naturhistoriska Riksmuseet, Stockholm ; Dr. H. Wermuth, Staatliches Museum fur Naturkunde, Ludwigsburg ; Dr. J. A. Peters, United States National Museum, Washington ; Dr. G. Peters, Institut fur Spezielle Zoologie und Zoologisches Museum der Humbolt Universitat, Berlin ; Dr. W. Ladiges, Zoologisches Staatsinstitut und Zoologisches Museum, Hamburg ; Dr. F. W. Braestrup, Zoologiske Museum, K0benhavn ; Dr. W. Hellmich, Zoologisches Sammlung des Bayerischen Staates, Munich. I should also like to thank Mr. E. V. Malnate, Academy of Natural Sciences of Philadelphia, for providing me with information on the holotype of U. castor. I am especially grateful to Miss A. G. C. Grandison and her staff at the British Museum (Natural History) where most of this work, including the preparation of this manuscript, was undertaken. The following abbreviations are used : A.M.N.H. American Museum of Natural History, New York. A.N.S.P. Academy of Natural Sciences of Philadelphia. B.M.N.H. British Museum (Natural History), London. C.A.S. California Academy of Sciences, San Francisco. F.M.N.H. Field Museum of Natural History, Chicago. L.A.C.M. Los Angeles County Museum, Los Angeles. M.C.Z. Museum of Comparative Zoology, Harvard University, Cambridge. M.H.N.P. Museum National d'Histoire Naturelle, Paris. N.M.W. Naturhistorischen Museum, Wien (Vienna). N.R.M.S. Naturhistoriska Riksmuseet, Stockholm. S.M.F. Natur-Museum und Forschungs-Institut Senckenberg, Frank- furt. IGUANID LIZARD URACENTRON AND STROBILURUS 49 S.M.N.L. Staatliches Museum fur Naturkunde, Ludwigsburg. U.S.N.M. United States National Museum, Washington. Z . M . B . Zoologisches Museum der Humboldt Universitat , Berlin . Z.M.H. Zoologisches Staatsinstitut und Zoologisches Museum, Ham- burg. Z.M.K. Zoologiske Museum, K0benhavn (Copenhagen). Z.S.B.S. Zoologisches Sammlung des Bayerischen Staates, Miinchen. URAGENTRON Kaup, 1826, Isis (von Oken), 19 : 88. Type species Lacerta azurea Linnaeus. DIAGNOSIS. Uracentron belongs to that group of South American- West Indian iguanids that is distinguished by the presence of a large sternal fontanelle and the absence of femoral pores, the " tropidurines " (Etheridge, 1964 : 629). Within the tropidurines Uracentron is probably most closely allied to Strobilurus, Tropidurus, Plica, and Platynotus, all of which have a very large interparietal scale. Uracentron differs from these genera in having a short, spiny, nonautotomic tail that is about one half as long as the head and body ; in the others the tail is at least as long as the head and body, and is autotomic. Among South American iguanids only the unrelated Hoplocercus has a tail similar to Uracentron's, but Hoplocercus may be easily distinguished by its small interparietal and heterogeneous dorsal squamation. CHARACTERISTICS. Upper head scales polygonal, juxtaposed, slightly to moder- ately convex, with a distinctly granular surface. Supraorbital semicircles poorly differentiated ; scales in frontal and prefrontal regions similar to other snout scales. Three or four pair of frontals, of which one to three may be in contact medially between the orbits, or separated by a median row of small scales. One to three rows of enlarged supraoculars, at least some of which are transversely expanded. Enlarged supraoculars usually separated from frontals and frontoparietals by two rows of small scales. Interparietal scale very large, one third to one fourth as wide as the head, narrower in front, usually longer than wide, with a central " eye ". Nasal large, dorsal, separated from rostral and upper labials by a row of scales ; opening large, directed dorsally in the posterior part of the scale. One or two large, convex, overlapping canthals. Canthals followed by six elongate superciliaries, each overlapping the one behind, followed by three shorter scales that overlap in the opposite direction. Two or three large scales in loreal region. A wide, elongate subocular scale with a keel along its upper margin, preceded by two or three shorter but similar scales that border the orbit anteriorly. A single row of rather large loreolabials, ending below the subocular, which posteriorly forms a short suture with the last upper labial. Upper labials large, more or less rectangular. Temporals polygonal, juxtaposed, smooth or keeled. Lower labials similar to upper labials but larger still. No enlarged postmental scales. Gulars small, mostly hexagonal, convex, juxtaposed or subimbricate, reduced in front of the anterior transverse gular fold, enlarged and imbricate in front of the posterior transverse gular fold. Tympanum about as large as the eye, without projecting scales along its anterior margin. 5° R. ETHERIDGE No scale row aligned mid-dorsally. Dorsal and lateral scales of neck and anterior body small, convex and juxtaposed, or keeled and imbricate. Remaining dorsal and lateral scales of body larger, imbricate, smooth or keeled. Ventral body scales imbricate, smooth or faintly keeled, larger than dorsals. Tail short, not autotomic, about one half as long as the head and body, moderately or strongly depressed, with equal whorls of large, spinose scales. Limbs covered with rhomboidal, imbricate scales, smooth or keeled, those on the posterior surface of the thighs reduced. Scales of palms and soles imbricate, denti- culate, smooth or faintly keeled. Subdigital lamellae mostly tricarinate and tridenticulate, the keels becoming faint on the distal lamellae. Two transverse gular folds present, the anterior one just posterior to a line even with the lower border of the tympanum, and with normal gular scales ; the posteror fold formed by ventromedial extensions of the antibrachial folds, and enclosing reduced but not granular scales. Antibrachial folds extend up and back over the forelimbs, then slant back and downward along the side of the body, fading out about midway to the hind limbs. Sides of neck with well developed, irregular folds. A ventrolateral longitudinal fold present between the forelimb and hind limb insertions. Scales on top and sides of head and on chin with numerous, closely set scale organs. Body scales with a single scale organ or none at all. Caudal scales with several prominent scale organs along posterior edge on each side of keel. No femoral or preanal pores. Uracentron azureum (Linnaeus) Lacerta azurea Linnaeus, 1758, p. 202 (type locality, Africa; here restricted to the vicinity of Paramaribo, Surinam). Stellio brevicaudata Latreille, 1802, p. 29 (type locality, interior of Guiana and Surinam). Stellio azureus Latreille, 1802, p. 34. Stellio brevicaudatus Daudin, 1802, p. 40. Stellio azureus Daudin, 1802, p. 36. Uromastyx caeruleus Merrem, 1820, p. 56 (substitute name for Stellio azureus}. Uromastyx azureus Merrem, 1820, p. 57. Uracentron azureum Kaup, 1826, c. 88. Uracentron coeruleus Kaup, 1826, c. 88. Doryphorus brevicaudatus Cuvier, 1829, p. 34. Doryphorus azureus Cuvier, 1829, p. 34. Urocentron azurea Wagler, 1830, p. 45. Ophessa (Uranocentron) brevicaudatus Gray, 1831, p. 42. Ophessa (Uranocentron) azureus Gray, 1831, p. 42. Urocentron brevicaudatum^Wiegma.n-n, 1834, p. 48. Urocentron azureum Fitzinger, 1843, p. 77. Urocentron azureum Boulenger, 1885, p. 182. SYNTYPES. No. KaF 1900 : 113 (2 exs) in the Naturhistoriska Riksmuseet, Stockholm, Sweden. TYPE LOCALITY. In listing Africa as the type locality of Lacerta azurea, Linnaeus (1758 : 202) probably was misled by Seba (1734 : 2 : 62 : 6), who had figured this IGUANID LIZARD URACENTRON AND STROBILURUS 51 species and given Africa as its origin. Nevertheless, Linnaeus did have before him two specimens upon which his description was based, and there is reasonably good evidence that they had come from the neighbourhood of Paramaribo, Surinam. Many of the species of South American plants and animals described by Linnaeus were collected by Carl Gustaf Dahlberg, a man of Swedish nationality, who arrived in Surinam in 1746, where he acquired plantations on Peruca Creek, and Cottica River, within 100 km of Paramaribo. In 1754, during one of his return trips to Sweden, Dahlberg presented his collections to King Adolf Fredrik and in the same year Linnaeus (1754 : 42) described specimens of Lacerta azurea in the museum of the King. When Dahlberg returned to Surinam in 1755 he took with him Daniel Rolander at the request of Linnaeus, who at least partly financed Rolander's trip. Rolander was a former student of Linnaeus, and private tutor to his son. Rolander is known to have collected in the neighbourhood of Paramaribo, and also went up the Commewijne River. The unrest caused by the revolt of escaped negro slaves prevented him from penetrating deeper into the interior of Surinam, and in 1756 he returned to Sweden. Part of Rolander's collection was purchased by the Swedish Baron de Greer, who in turn presented the specimens to Linnaeus (Holthuis, 1959 : 17-21). Most of the South American species of lizards described by Linnaeus, including Uracentron azureum, are widely distributed in the northern part of the continent, and are known to occur at Paramaribo. It seems probable that many, if not all, of Linnaeus' specimens were from the collections of Dahlberg and Rolander, made in the vicinity of Paramaribo. I therefore propose the restriction of the type locality of Uracentron azureum to the vicinity of Paramaribo, Surinam. CHARACTERISTICS. The supraoculars are variable in shape ; one, two, or two and a half rows of enlarged scales may be present. The scales of the inner row are always to some extent transversely widened, but never strap-like. The enlarged supra- oculars are always separated from the superciliaries by two rows of small scales. The dorsal and lateral scales of the neck and anterior part of the body are very small, smooth, convex, and juxtaposed. Posteriorly the dorsal and lateral scales of the body become larger, imbricate, and obtusely keeled, largest in the sacrolumbar region. The ventral scales are smooth. Scale counts are given in Table I. The tail is moderately depressed, with about 21 whorls of large, spinose scales. A whorl halfway between the vent and the tip of the tail contains about 13 scales. The tail is 0.48 to 0.65 (mean, 0.58) times as long as the head and body. The snout-vent length of the largest male examined is 87 mm, of the largest female 86 mm. Colour in preservative : the upper surfaces are bluish-grey, with bold, black mark- ings. The medial half of the supraocular region is black, and there is usually a median, V-shaped mark in front of the frontal region. A W-shaped mark on the back of the head has its median apex on the interparietal scale, and its horns extend down to the suboculars. Two crescentric bands cross the neck, the extremities of the anterior one curving down through the tympani to the angle of the mouth. R. ETHERIDGE IGUANID LIZARD URACENTRON AND STROBILURUS 53 A series of similar crescentric bands cross the back, the extremities of the first curving down into the antihumeral folds. Five or six bands may be present between the shoulders and the sacrum, or the posterior one, two, or three bands may be broken and off-set at the dorsal midline, or may be broken up into a bold reticulum. A black reticulum covers the upper surfaces of the limbs, and is most intense on the hind limbs. The tail has irregular black marks above, and about one half of the spines along the sides of the tail are tipped with yellow. Below, the throat is dark bluish-grey. The belly is a lighter grey, and the ventral surface of the tail is dark bluish-grey along the sides, with a yellowish area down the middle. Mr. Marinus S. Hoogmoed has furnished me with colour notes on a living individual captured along Feticreek, Litani River, in Surinam. The colour is described as " head and back yellow-green mottled, tail with yellow points, throat yellow-green ; belly, underside of legs grey-blue ". Cott (1926) has published an excellent coloured illustration of this species. RANGE. (Text-fig, i). Records of Uracentron azureum are from British Guiana, French Guiana, and Surinam, and from along, or near the Amazon River in north- eastern Brazil, as far west as Manaus. U. azureum and U. flaviceps occur together at Manaus. MATERIAL EXAMINED. British Guiana : no specific locality B.M.N.H. 1905.10.21.1. Surinam : Bergenaal N.M.W. 13926 (2 exs) ; no specific locality. M.H.N.P. 2515, Z.S.B.S. 534/0, M.C.Z. 4500. French Guiana : Cayenne M.H.N.P. 196, 2514, 2396 ; Saint Laurent M.H.N.P. 24.121 ; Mornes du bas Mahurg M.H.N.P. 03.20 ; no specific locality M.H.N.P. 8211, 2516, B.M.N.H. 1920.1.20.1334, Z.S.B.S. 534/0. Brazil : Amazonas, Manuas B.M.N.H. 1922.6.15.1, C.A.S. 84247 ; Amazonas, Manjuru River A.M.N.H. 101945 ; Para, Ilha Marajo, Soute S.M.F. 24930 ; Para, Ilha Marajo, Dist. Calderao Z.S.B.S. 37/1924 (5 exs), Z.M.H. 4421 (5 exs), 4496 ; Para, Ilha Marajo B.M.N.H. 1923.11.9.68-72, 1926.5.5.5, S.M.F. 24931 ; Para, within 50 miles of Belem M.C.Z. 53216 ; Para, Amnuathena, Rio Tocantins Z.S.B.S. 674/1920 ; Amapa, Serra do Navio S.M.F. 59580 ; Maranhao S.M.F. 11202 ; no specific locality M.H.N.P. 92.290, 99.215, N.M.W. 13928.1, B.M.N.H. 51.7.17.45, A.M.N.H. 58277-9, 60330-2. South America : no specific locality B.M.N.H. 47.2.19.6, 59.12.28.2, 66.8.14.302, N.M.W. 13927 (2 exs), N.R.M.S. Kaf 1900 : 113 (syntypes), S.M.N.L. unnumbered. Additional records from Surinam have been provided to me (in litt. )by Mr. Marinus S. Hoogmoed : Litani Fetikreek, Sipaliwini, Paloemeu, Berg en Dal, and Moengo. Uracentron guentheri Boulenger Urocentrum gttentheri Boulenger, 1894, P- 729 (type locality : Iquitos, Peru). Urocentr on guentheri Burt and Burt, 1933, p. 49. HOLOTYPE. No. 93 . 7 . 10 . 13 (RR 1946 . 8 . 29 . 85) in the British Museum (Natural History), London. Collector, A. E. Pratt. CHARACTERISTICS. Four or five transversely widened, band-like supraocular scales are present in a single row, all in contact laterally with the superciliaries, or 54 R. ETHERIDGE the first two or three may be separated from the superciliaries by a row of very small scales. The dorsal and lateral scales of the neck and anterior part of the body are very small, smooth, convex, and juxtaposed. Posteriorly the dorsal and lateral scales of the body become larger, imbricate, and weakly, obtusely keeled, largest in the sacrolumbar region. The ventral scales are smooth. Scale counts are given in Table I. The tail is moderately depressed, with about 21 whorls of large, spinose scales. A whorl halfway between the vent and the tip of the tail contains about 13 scales. The tail is 0.52 to 0.57 (mean 0.54) times as long as the head and body. The snout-vent length of the largest male examined is 75 mm., of the largest female 75 mm. Colour in preservative : the upper surfaces are bluish-grey with black markings. Many of the head scale sutures are edged in black, and the supraocular region on each side is outlined in black, with a black bar across the middle. On the frontals is a median, longitudinal black bar. Five or six narrow, somewhat irregular, crescentric bands cross the posterior part of the head and neck, with the horns of each crescent pointing downward and forward. Across the shoulders is a similar, narrow crescent that descends on each side into the antihumeral folds. The upper surface of the limbs and most of the back are covered by a bold reticulum of black lines. In the middle of the anterior part of the back the reticulum tends to form short, transverse bars. On the hind leg the reticulum extends onto the posterior surface of the thigh. The caudal scales and the ends of their spines are outlined in yellow. The ventral surface of the head and body is light bluish-grey, becoming yellowish in the preanal area and under the tail. There are no records of the colour in life of this species, but is seems probable that the bluish-grey background of preserved animals is green in life. A black-and-white illustration of the pattern may be found in Boulenger, 1894 : pi. 47, fig. 3. RANGE. (Text-fig, i). Records of Uracentron guentheri are from the Rio Ucayali system in the Department of Loreto in eastern Peru, and from the western Amazonian basin in northwestern Brazil. The range of U. guentheri apparently overlaps that of U. flaviceps in eastern Peru and Brazil, and the two are definitely known to occur together at Iquitos, Peru. REMARKS. Boulenger (1894 : pi. 47, fig. 3) shows the tip of the tail of the holo- type of Uracentron guentheri blunt. Mertens (1925 : 76) used this as a character to distinguish it from U. werneri. However, the blunt tail of the holotype of U. guentheri is due to injury; the tip is missing and the end of the tail is healed over with scar tissue. The tail has i6| rows of spinose scales ; about 5^ rows are missing. MATERIAL EXAMINED. Peru : Dept. Loreto, lower Rio Cushdbatay A.M.N.H. 56410 ; Dept. Loreto, Roaboya, Rio Ucayali A.M.N.H. 57204 ; Dept. Loreto, Rian Rian, Contamana region, Rio Sahnaya Valley A.M.N.H. 57205 ; Dept. Loreto, Iquitos B.M.N.H. 93.7.10.3 (RR 1946.8.29.85) (Holotype). Brazil : Amazonas, Rio Jutai Z.M.B. 30974 ; Amazonas, Rio Solimoes, Lago Calado S.M.F. 30304/5.2. IGUANID LIZARD URACENTRON AND STROBILURUS 55 Uracentron werneri Mertens Uracentron werneri Mertens, 1925, p. 75 (type locality : upper Orinoco, Venezuela). HOLOTYPE. No. 11203 in the Natur-Museums und Forschungs-Institutes, Frank- furt am Main. Collector, G. Hubner. CHARACTERISTICS. The shape and size of the supraoculars are variable ; two or two and a half rows of enlarged scales with four or five scales in each, those of the inner row somewhat transversely widened, and those of the outer row separated from the superciliaries by two rows of small scales. The dorsal and lateral scales of the neck and anterior part of the body are very small, smooth, convex, and juxtaposed. Posteriorly the dorsal and lateral scales of the body become larger, imbricate, and remain smooth. The ventral scales are smooth. Scale counts are given in Table I. The tail is moderately depressed, with about 21 whorls of large, spinose scales. A whorl halfway between the vent and the tip of the tail contains about 13 scales. The tail is 0-56 to 0-60 (mean, 0-57) times as long as the head and body. The largest male examined is 60 mm. snout-vent length, the largest female is 58 mm. Colour in preservative : all upper surfaces are dark greyish-brown or bluish-black, somewhat lighter on the snout and sides of the head. A pattern of light spots and a dark nuchal collar is very faintly indicated in some specimens, including the holo- type. The holotype has a bold, yellowish, asymetrical spot on the top of the head that covers the interparietal and adjacent scales on both sides and to the rear, and extends forward on the right side only, narrowing towards the superciliary ridge. This mark appears to be anomalous, for it is absent in all specimens other than the holotype. According to Valdivieso and Tamsitt (1963:31) living animals of this species are bright green. RANGE. (Text-fig, i). Records of Uracentron werneri are from the upper Rio Orinoco valley in Venezuela, and from near the western tributaries of the Orinoco in Colombia. MATERIAL EXAMINED. Venezuela : Amazonas, Alto Orinoco S.M.F. 11203 (holotype). Colombia : Meta, near Macarena, Rio Guayabero A.M.N.H. 91755 ; Vaupes, Cerro Yapopoda M.C.Z. 67982. Published records from Colombia are Caqueta (Dunn, 1944 : 89), and near the Macarena Mountains, Meta (Valdivieso and Tamsitt, 1963 : 31). Uracentron flaviceps (Guichenot) Doryphorus Azureus, variety Dumeril, 1851, p. 85. Doryphorus flaviceps Guichenot, 1855, p. 26 (type locality : Sarayacu, Peru). Doryphorus castor Cope, 1870, p. 556 (type locality : Pebas, " Ecuador ", now Peru). Uranocentron flaviceps O'Shaughnessy, 1881, p. 245. Urocentr on flaviceps Boulenger, 1885, p. 183. Urocentron castor Boulenger, 1885, p. 184. HOLOTYPE. No. 6682 in the Museum National d'Histoire Naturelle, Paris. Collectors, Castelnau and DeVille. CHARACTERISTICS. The supraocular scales are variable ; usually there is an inner 56 R. ETHERIDGE row of four or five transversely widened scales, and an outer row of three or four scales that are about half as large as those of the inner row, with a single row of scales between the outer row and the superciliaries. The dorsal nuchal scales are somewhat conical and keeled, with tiny granules in between. These grade laterally into smaller, but otherwise similar scales on the sides of the neck. The dorsal body scales are larger, imbricate, and distinctly keeled, largest in the sacrolumbar region. The lateral body scales are a little smaller, but otherwise similar to the dorsals. The ventrals are smooth, or faintly keeled, especially in the pectoral region. Scale counts are given in Table I. The tail is very flat and wide, almost leaf-like, with 30 to 38 whorls of moderately large, spinose scales. A whorl halfway between the vent and the tip of the tail contains 9 to n scales. The tail is 0-48 to 0-63 (mean 0-50) times as long as the head and body. The largest male examined is 128 mm. snout-vent length, the largest female is 87 mm. Colour in preservative : juveniles and females are a rich, dark brown above, with numerous small, light spots; the spots are light bluish on the head, forelimbs, and anterior part of the body, becoming yellowish on the posterior part of the body, hind limbs, and tail. In juvenile males the light spots tend to be absent from a crescentric band across the nape, leaving a solid, dark band. The throat in females and juvenile males is light bluish with brown spots that may be arranged in irregular, oblique rows ; other ventral surfaces are light grey. In adult males the light bluish spots of the head and neck increase in extent, and join one another until the head and neck have small brown spots on a bluish background. The dark brown nuchal band becomes conspicuous, and is bordered behind by a light, bluish band. On the remainder of the body, limbs, and tail the light spots become obscure or disappear entirely, leaving a uniform dark brown surface. The ventral surfaces become brown as well, except for the chin and throat, and the ventral surface of the tail where some yellow spotting usually remains. According to my field notes of yth August, 1961, an adult male of 88 mm. snout- vent length, obtained 39 km. NNE of Oxapampa, Dept. Pasco, Peru (now L.A.C.M. No. 39161) had the following colour in life : dorsum of body, limbs, and tail very dark brown, finely punctate with yellow on limbs and sides of body ; head reddish- brown mottled ; collar a wide, dark brown band bordered behind by a yellow band ; chest and venter of limbs iridescent blue-green ; tail reddish below ; throat dirty yellow. Another male, 114 mm. snout- vent length, from the same locality, had the same pattern, but with colours more intense, and the head was bright reddish. After six years in preservative the reddish and yellow colours have become light bluish-green. Guichenot (1855, pi. 3, fig. 2) gives a coloured illustration of the holotype. RANGE. (Text-fig. 2). Records of Uracentron flaviceps are from the western part of the Amazonian Basin in northwestern Brazil, southeastern Colombia, eastern Ecuador, and eastern Peru. The species occurs together with U. guentheri in Peru, and with U. azureum in Brazil. 57 WEST LONGITUOC FIG. 2. Map of northern South America showing localities for Uracentron flaviceps. REMARKS. In his description of Doryphorus castor, Cope (1870 : 55) used as the principle diagnostic character the presence of the nostril between two scales rather than within a single scale. Boulenger (1885 : 184) recognized U. castor as a valid species, but called attention to its close similarity with U. flaviceps. Mr. E. V. Mal- nate has examined the holotype of U. castor (A.N.S.P. 11303), and compared it with specimens of U. flaviceps from eastern Peru. He has informed me (in litt.} that contrary to Cope's description each nasal opening is within a single scale, and in this and in all other details of scalation and colour pattern the specimen is indistinguish- able from U .flaviceps. In his list of the type specimens of lizards in the Paris Museum, Guibe (1954 : 42) incorrectly considered Uracentron flaviceps to be a synonym of U. azureum. 58 R. ETHERIDGE MATERIAL EXAMINED. Colombia : Amazonas, Leticia F.M.N.H. 83038, 78384-5 (2 exs) ; Putumayo, Santa Rosa de los Kofanes, upper Rio Guamues F.M.N.H. 165828. Ecuador : Sarayacu M.H.N.P. 6882 (holotype), B.M.N.H. 80.12.8.56 ; Canelos B.M.N.H. 80.12.8.57-9 ; Pastaza River M.C.Z. 37270 ; Napo, Santa Cecilia M.C.Z. 92519 ; no specific locality Z.M.B. 9988. Brazil : Amazonas, Rio Madiera A.M.N.H. 57207 ; Amazonas, Manaus N.H.M.P. 92.291 ; Amazonas, Rio Jutai Z.M.B. 30973 ; Amazonas, Rio Solimoes N.R.M.S. 3342 ; Amazonas, Lago Calado S.M.F. 30304/5.1. Peru : Loreto, Iquitos A.M.N.H. 56408, N.M.W. 13929 (2 exs), 13930, F.M.N.H. 45487 ; Loreto, Prov. Ucayali, Yarinacocha F.M.N.H. 45484, 56064 ; Loreto, Prov. Ucayali, Pucalpa F.M.N.H. 56061-3 ; Loreto, Prov. Bajo Amazonas, Quidtococha F.M.N.H. 45485-6 ; Loreto, upper Ucayali A.M.N.H. 71101-2 ; Loreto, Pebas A.N.S.P. 11303 (holotype Doryphorus castor) ; Pasco, 39 km NNE of Oxapampa L.A.C.M. 39161 ; Loreto, Pongo Manseriche, Marafion valley A.M.N.H. 56409 ; between Apaga and Nieva A.M.N.H. 57206. No specific locality B.M.N.H. 69 . 5 . 21 . 57. Species Referred Erroneously to Uracentron Urocentrum meyeri was described by Werner (1900 : 4) from Lima, Peru, and included by Burt & Burt (1933 : 49) in their list of South American lizards. The holotype and only specimen was destroyed in the Dresden Museum (No. 1764) during the last days of World War II. However, the description contains details that are sufficient to identify the specimen as belonging to the genus Stenocercus rather than Uracentron. The upper head scales are smooth, the interparietal scale small, and there is a projecting auricular scale. The tail is flattened, but scarcely wider than the sacral region, with spiny scales in rings, and is a little longer than the head and body length : total length 131 mm., tail length 77 mm. These are all characteristics of the spiny-tailed species of Stenocercus: atrigularis, roseiventris, marmoratus , crassi- caudatus, carrioni, and simonsi. Additional details in the description, and the type locality indicate that the specimen may have been one of Stenocercus crassi- caudatus. The sides of the neck have two converging folds, a postympanic and an antihumeral fold, with a horizontal fold between them ; the dorsal scales are small and granular. A median dorsal crest is not mentioned, and presumably was lacking. The ventral scales are said to be similar to the dorsals, but faintly keeled, almost smooth. Only the faintly keeled ventrals are not characteristic of 5. crassicaudatus. Although there are no records of S. crassicaudatus from Lima, the species does occur inland from Lima at higher elevations. The exact identity of Werner's species may never be known, but in any event it is clearly not referrable to Uracentron. Burt & Burt (1933 : 49) also included in their checklist of South American lizards the name Urocentronpalluma, which they credited to Tschudi (1846 : 35). However, it is clear from Tschudi's own treatment in his Fauna Peruana (incorrectly cited by Burt & Burt as 1845), and in an earlier work (Tschudi, 1845 : 157-8) that he is referring to Phymaturus palluma, which Tschudi correctly attributes to Molina (1782 : 217). Tschudi listed Phymaturus Gravenhorst as a subgenus of Urocentron, IGUANID LIZARD URACENTRON AND STROBILURUS 59 and gave as its only species in Peru " U. palluma Tschudi ". Although he followed the species with his own name, he listed Lacerta palluma Molina in its synonomy. Burt & Burt's error is clearly that of accepting Tschudi's placement of Phymaturus palluma in the genus Urocentron, and, apparently because Tschudi followed the species name with his own, considering Tschudi to be its author. Phymaturus is a valid, monotypic genus, containing only the species palluma, and quite unrelated to Uracentron. It differs from the latter in many ways, including the presence of tricuspid premaxillary teeth, preanal pores in males, a small interparietal scale, and a rather slender, somewhat spiny, autotomic tail. Relationships Within Uracentron Uracentron flaviceps stands apart from the other three forms of Uracentron as the most distinctive species. Compared with the others its tail is much flatter and wider, with smaller, less distinctly spinose scales. Its body scales are larger and consequently fewer in number around the middle of the body, and more distinctly keeled. U. flaviceps also attains a greater maximum size. Uracentron azureum, U. guentheri, and U. werneri are very similar in the form of their body and tail, and in scalation. U. guentheri differs from azureum and werneri in having greatly expanded, band-like supraoculars, and U. werneri differs from azureum and guentheri in having smooth dorsal scales in the sacrolumbar region. The most obvious differences among the three are their colour patterns : bold black cross-bands usually followed by a bold reticulum in azureum, a finer reticulum with narrow cross-bands on the neck in guentheri, and solid or faintly spotted in werneri. The most extreme example of breaking up of the posterior cross-bands into a reti- culum in azureum does not approach the pattern of guentheri, but transitional stages between the two are not difficult to imagine. The pattern of werneri might result from an overall darkening of the pattern of guentheri. The ranges of U. azureum, U. guentheri, and U. werneri are not known to contact one another, or to overlap. With the acquisition of more specimens from inter- mediate areas it may become possible to determine whether or not any two, or all three of these species intergrade. Until that time the best course would seem to be recognition of three separate species. 1 Tail very strongly depressed. More than 25 whorls of caudal scales from base to tip of tail. Dorsal body scales, including nuchals, distinctly keeled . . flaviceps - Tail moderately depressed. Not more than 25 whorls of caudal scales from base to tip of tail. Dorsal scales of neck and anterior part of body smooth ... 2 2 Four or five band-like supraoculars separated from superciliaries by one row of small scales, or in contact with superciliaries ....... guentheri - Supraoculars variable, but those enlarged separated from superciliaries by at least two rows of small scales .......... 3 3 Dorsal body scales in sacrolumbar region obtusely keeled. A bold dorsal pattern of black cross-banks or anterior cross-bands and posterior reticulum . . azureum - Dorsal body scales of sacrolumbar region smooth. Dorsum solid gray-brown, or faintly spotted werneri 60 R. ETHERIDGE STROBILURUS Wiegmann, 1834, Herp. Hex., p. 18 Type species Strobilurus torquatus Wiegmann. DIAGNOSIS. Strobilurus is a member of the tropidurine group of South American- West Indian iguanid lizards. It was not initially included in that group because I had no data on the structure of the skeleton (Etheridge, 1964 : 629). I have since determined, however, that a large sternal fontanelle is present, and on the basis of this and other osteological and integumentary characteristics, Strobilurus is clearly a member of the tropidurine group. Within the tropidurines Strobilurus is probably most closely allied to Uracentron, Tropidurus, Plica, and Platynotus, all of which have a very large interparietal sacle. The tail of Strobilurus is auto- tomic, about as long as the head and body, and provided with unequal whorls of spinose scales. In Uracentron the tail is about half as long as the head and body, and is not autotomic. In the other genera the tail is considerably longer than the head and body, and autotomic. Some species of Tropidurus have a moderately spiny tail due to sharp, projecting mucrons, but the spines are not nearly as well developed as they are in Strobilurus. Among other South American iguanids some species of Stenocercus also possess a tail about equal to the head and body length, and provided with unequal whorls of stout, spinose scales. However, the inter- parietal scale of Stenocercus is very small, or absent entirely. CHARACTERISTICS. Upper head scales polygonal, juxtaposed, with a distinctly granular surface. Supraorbital semicircles not well differentiated as such. Three pairs of frontals in contact between the orbits, preceded by two or three larger prefrontals and frontonasals on each side. Two rows of enlarged supraoculars, four or five scales in each row. Those on the inner row a little wider than long and separated from the supraorbital semicircles by two rows of smaller scales ; those on the outer row separated from the superciliaries by two rows anteriorly and one row posteriorly. Interparietal scale very large, about one third as wide as the head, narrowing anteriorly, and as wide as to a little wider than long. A median " eye " present anteriorly in the interparietal scale, and often a median pit followed by a median groove posteriorly. Nasal scale large, dorsolateral in position, with a small, dorsally directed nostril in the posterior part of the scale. Nasal separated from rostral and upper labials by a single row of scales. Two rather small but strongly convex canthals. Canthals followed by five or six elongate, curved superciliaries, each of which overlaps the one behind, these in turn followed by three shorter super- ciliaries that overlap in the opposite direction. Several polygonal scales in the postnasal-loreal-preocular region. An elongate subocular narrowing anteriorly, with a keel along its upper margin. Loreolabials in a single row below the nasals, two rows below the loreal region, and one row between the subocular and upper labials. Temporals imbricate, obtusely keeled. Tympanum large, about equal to eye opening, with two flat, projecting scales in front. Upper labials more or less rectangular. Lower labials about one and a half times wider than upper labials. No enlarged postmentals. Gulars smooth, imbricate, reduced in size medially and posteriorly. A row of scales aligned mid-dorsally from occiput to base of tail, its scales slightly IGUANID LIZARD URACENTRON AND STROBILURUS 61 larger than, but otherwise similar to adjacent scales, forming a low denticulation in individuals over 100 mm. snout-vent length. Dorsal nuchals rhomboidal, imbricate, sharply keeled, the keels rising to sharp mucrons. Lateral nuchals smaller, spinose along the lateral nuchal folds. Dorsal scales of body rhomboidal, keeled, imbricate, shortly mucronate, the keels forming oblique lines that converge posteriorly toward the dorsal midline. Lateral body scales slightly smaller, but otherwise similar to the dorsals. Ventral scales smooth, imbricate, about one and a half times as large as the dorsals. Tail about as long as the head and body, and autotomic. Base of tail slightly depressed and a little wider than the sacral region, with eight equal whorls of large, sharply keeled, and strongly spinose scales. Remainder of tail cylindrical and much narrower than the base, with scales of unequal whorls that correspond to the autotomy segments of the vertebral column. Anteriorly each segment with a ventrally incomplete anterior ring of small, smooth scales, and posteriorly an additional ventrally incomplete ring of small scales added to the anterior border of each segment. Dorsal scales of forelimb rhomboidal, keeled, imbricate, about as large as dorsal body scales. Ventral scales of forelimb smaller, smooth, imbricate. Dorsal scales of hind limb sharply keeled and spinose, similar to but much smaller than spinose scales of tail. Postfemoral scales reduced, keeled, imbricate. Scales of palms and soles sharply keeled and mucronate, some of them tricarinate and tridenticulate. Subdigital lamellae tricarinate and tridenticulate. A transverse fold extends across the throat just posterior to a line even with the tympani, but scales within fold not differentiated from adjacent gulars. A pair of short, deep, antihumeral folds enclosing very small scales, widely separated vent- rally by about 12 anterior pectoral scales. A dorsolateral fold extends posteriorly from upper border of tympanum about halfway back to shoulder ; a ventrolateral fold begins as a pair of converging folds that meet just posterior to tympanum, and extend back to intercept antihumeral fold. Scales on top and sides of head with numerous scale organs. Dorsal scales of body and limbs with one to three scale organs on the free edge, or none at all. Caudal scales with three to eight scale organs on each side of the keel, along the free edge. No femoral pores or preanal pores. Strobilurus torquatus Wiegmann Strobilurus torquatus Wiegmann, 1834, p. 18 (type locality, Brazil). Steironotus (Strobilurus) torquatus Fitzinger, 1843, p. 71. Doryphorus spinosus Guichenot, 1855, p. 27 (type locality, Bahia, Brazil). Strobilurus torquatus Boulenger, 1885, p. 181. SYNTYPES. Nos. 672-3 and 9215 in the Zoologisches Museum der Humboldt Universitat, Berlin. CHARACTERISTICS. Since Strobilurus is a monotypic genus the characteristics of torquatus are the same as those of the genus. The vertebral scales counted from the 62 R. ETHERIDGE occiput to a line even with the anterior surface of the thighs when the limbs are extended at right angles to the body number 39 to 57 (mean, 55-6) ; the paravertebral scales counted in the same way are 49-72 (mean, 67-6). There are 19 to 29 (mean, 24-1) lamellae under the fourth finger, and 23 to 34 (mean 28-3) under the fourth toe. The tail is 0-93 to 0-95 (mean, 0-94) times as long as the snout-vent length. The largest male examined is 97 mm. snout-vent length, the largest female is 106 mm. Colour in preservative : the head and neck above are bluish-grey, with bold, black markings. There is a narrow, black crescent on each side where the supraoculars contact the frontals and frontoparietals. A wide crescent extends across the anterior part of the neck from ear to ear, and from this crescent there extends anteriorly a medial bar. On each side a black crescent extends from the interparietal scale down to the angle of the mouth. A wide, black, crescentric collar about six or seven scales wide middorsally, extends down on each side into the antihumeral fold. Behind the black collar the dorsal surface of the body, limbs, and tail is somewhat darker greyish-green, with obscure lighter cross-bands. The chin and throat are marbled light and dark grey, and the belly is light grey. A coloured illustration may be found in Guichenot (1855, pi. 7, fig. la), labelled Doryphoms spinosus. RANGE. Strobilurus torquatus is known from the states of Pernambuco, Alagoas, and Bahia, Brazil. REMARKS. Wiegmann described Strobilurus torquatus in a footnote (1834 : ~&] and the subsequent taxonomic history of the genus and species has been relatively uncomplicated. The species has remained rare in collections. MATERIAL EXAMINED. Brazil : Alagoas, Sao Miguel M.C.Z. 59275 ; Bahia, no specific locality Z.M.B. 9215, 672-3 (syntypes), 8272, M.H.N.P. 5085 (2 exs, syntypes of Doryphorus spinosus}, N.M.W. 13908 (2 exs),B.M.N.H. 62.11.23.50, 1903.10.16.23; Pernambuco, no specific locality B.M.N.H. 88.4.18.6. South America : no specific locality M.H.N.P. 6880, N.M.W. 13909 (2 exs), B.M.N.H. xxiii.io4a. Scales along middle of black Scales around middle of body Subdigital lamellae of fourth finger Subdigital lamellae of fourth toe azureum 92-(lO7'O)-I2O TABLE I guentheri N = 6 werneri N = 3 !27-(i23-3)-i4i 98-(iio-4)-i28 24-(27-4)-30 3i-(3i-6)-34 flaviceps N = 29 74-(8o-o)-88 66-(78-2)-88 25-(29'7)-33 26-(30-3)-34 TABLE i . Some scale counts of Uracentron. Scales along middle of back counted from occi- put to line even with anterior margin of hind limb at right angles to body ; scales around middle of body counted halfway between limb insertions. Mean figures in parentheses. N = number of specimens examined. IGUANID LIZARD URACENTRON AND STROBILURUS 63 REFERENCES BOULENGER, G. A. 1885. Catalogue of the lizards in the British Museum (Natural History], vol. 2, ed. 2, London, xiii+497 pp., 24 pis. — 1894. Second report on additions to the lizard collections in the Natural History Museum. Proc. zool. Soc. Land. (1894) : 722-736, 3 pis. BURT, C. E. & BURT, M. D. 1933. A preliminary checklist of the lizards of South America. Trans. Acad. Sci. St. Louis 28 (i) : i-v-f 1-104. COPE, E. D. 1870. Eighth contribution to the herpetology of Tropical America. Proc. amer. phil. Soc., 553-559. COTT, H. B. 1926. Observations on the life-habits of some batrachians and reptiles from the lower Amazon : and a note on some mammals from Marajo Island. Proc. zool. Soc. Lond., 4 : 1161-1168, 6 pis. CUVIER, G. F. 1829. Le regne animal distribue d'apres son organisation, vol. 2, ed. 2, Paris. 406 pp. DAUDIN, F. M. 1802. 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Neue Classification der Reptilien nach ihren naturlichen Verwandtschaften. Vienna, viii + 66 pp. — 1843. Systema Reptilium. Fasciculus Primus. Amblyglossae. Vienna. vi+io6 pp. GRAY, J. E. 1831. in CUVIER, edit. GRIFFITH, The Animal Kingdom. Reptilia. London. 481 pp. 1845. Catalogue of the specimens of lizards in the collection of the British Museum. London xxviii-f-289 pp. GUERIN-MENEVILLE, M. F. E. 1829-1838. Iconographie du Regne Animal de G. Cuvier. Reptiles. Paris, 23 pp., 30 pis. GUIBE, J. 1954. Catalogue des types de lezards du Museum National d'Histoire Naturelle. Paris. 1 19 pp. GUICHENOT, A. 1855. in CASTELNAU, Animaux nouveaux ou raves recueillis pendant I' expedi- tion dans les parties centrales de I'Atnerique du Sud, de Rio Janeiro a Lima, et de Lima au Para. Reptiles. Paris. 96pp., 1 8 pis. HOLTHUIS, L. B. 1959. The Crustacea decapoda of Suriname (Dutch Guiana). Zool. Verh. 44 : 1-295, 68 figs., i6pls. KAUP, J. 1826. Behtrage zu amphibiologie und ichthinologgy. 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Miinchen. vi + 354 pp., 9 pis. WERNER, F. 1900. Reptilien und Batrachier aus Peru und Bolivien. Abh. Ber. K. zool. anthrop.-ethn. Mus. Dresden 9 : 1-4. WIEGMANN, A. F. A. 1834. Herpetologica Mexicana. I. Saurorum Species. Berlin, vi+54 pp., 10 pis. PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING NIGERIAN LIZARDS OF THE GENUS AGAMA (SAURIA : AGAMIDAE) HIS? ALICE G. C. GRANDISON BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 17 No. 3 LONDON: 1968 18 NIGERIAN LIZARDS OF THE GENUS AGAMA (SAURIA : AGAMIDAE) BY ALICE G. C. GRANDISON *"• — " " "*"/ t%',A • British Museum (Natural History) Pp. 65-90; 6 Plates: 2 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 17 No. 3 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. 17, No. 3 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 19 November, 1968 Price Eighteen Shillings NIGERIAN LIZARDS OF THE GENUS AGAMA (SAURIA : AGAMIDAE) By ALICE G. C. GRANDISON SYNOPSIS The external morphology, distribution and systematics of the species of Agama occurring in Nigeria are reviewed. The number and position of the dermal sense organs on the trunk scales are used to help clarify the status and relationships of the five species recognized. Agama agama benueensis Monard is shown to be a complex comprising two distinct species, one of which is described as new, the other sympatric with but unrelated to Agama agama (Linnaeus). Agama boensis Monard is not a valid form; the types are a mixed series of Agama sankaranica Chabanaud and Agama weidholzi Wettstein. INTRODUCTION UNTIL a few years ago it was believed that only three species of Agama occurred in Nigeria: the ubiquitous Agama agama (Linnaeus) and the two smaller and less common species A . sankaranica Chabanaud and A . gracilimembris Chabanaud. But in 1963 Dr. Hilary Fry of the Ahmadu Bello University, Zaria drew my attention to the existence of a form which he described as being of similar adult size to Agama agama and as occurring on the smaller inselbergs in the Zaria region of northern Nigeria. The adult male was said to be distinguished by four indigo or black, clearly defined blotches on the throat. It was immediately recognized from an examination of an adult male which Fry submitted that this Agama was not conspecific with any of the three known species. With the co-operation of members of the staff of his depart- ment intensive efforts were made to obtain further examples as well as ecological data. The material resulting from these field studies contained more adult male " blue throats " but not adult females belonging to the same form, instead examples of Agama sankaranica, which in the field had been confused with the apparently undescribed form, and gravid females of yet another apparently undescribed agama. The absence in collections of females and immature individuals of this " blue throat " continued until the following year when a Danish zoologist, Dr. Arne Schi0tz, who was working in the British Museum recognized the male blue-throated agama as being similar to ones he had collected in the Western Region. His large series, obtained on inselbergs near Idanre and Igbetti, included both sexes and juveniles and were in the collections of the Zoologiske Museum Copenhagen. A detailed study there of his material enabled me to appreciate the difficulty that other collectors had encountered in the field in distinguishing the females and immature individuals from the superficially similar A. sankaranica and to elucidate the intraspecific variation of all the Nigerian forms of Agama. The study was extended by referring to type and other material in several institutions and to the British Museum's collec- tions including those acquired during the 1962 Northern Nigeria expedition. Finally the specimens on which Monard based the name Agama agama benueensis were borrowed. ZOOL. 17, 3. 3 68 A. G. C. GRAND I SON The identity and relationships of Monard's Agama agama benueensis had not been questioned until Wermuth (1967) in a footnote pointed out that benueensis is probably no more than a local variety of Agama agama agama differing from it only in minute details of toe length and pattern. The original description of benueensis, extremely brief and inadequate, turned out to be based on a mixed series, some of the individuals being conspecific with the Nigerian " blue-throated " form, while others, strikingly different in their pattern and in the position of the nostril and with a lower average midbody count are closely related to Agama agama and apparently conspecific with a Nigerian population. The name benueensis is here applied to the " blue-throat " form and a lectotype is designated. The information now available on the ecology, morphology, distribution and variation of Agama agama, A . benue- ensis, A . sankaranica and A . gracilimembris is given and a new species is described. Descriptions are based on the observed variation in the examined material. Throughout this paper the following abbreviations are used to refer to specimens or collections. B.M. British Museum (Natural History) M.H.N.C.F. Museum d'Histoire Naturelle, La Chaux de Fonds M.N.H.N. Museum National d'Histoire Naturelle, Paris Z . M . C . Uni versitets Zoologiske Museum Copenhagen METHODS AND TERMINOLOGY The characters used in this study are defined as follows: Body scales. The shape, size and direction of imbrication of the trunk and flank scales, the extent of the keel relative to the axial length of each scale, from its anterior border to its apex. Microscopic structure of body scales. The number and position of the " hair- bearing " scale organs on the scales of the mid trunk region were studied by removing a series from the vertebral to the mid ventral row and mounting them according to the method described by Underwood (1957). Midbody scale count. The number of longitudinal scale rows around the middle of the trunk. Vertebral count. The number of transverse scale rows along the vertebral line from forelimbs (on a level with the axillae) to hind limbs (groin). Direction of imbrication of temporal and supratemporal scales. Direction of overlap of the scales in these regions, the regions being defined as : temporal, the area between the anterior rim of the ear and the postocular scales, overlying the temporal fossa and the squamosal and the posterior half of the jugal bones; supratemporals, the scales overlying the supratemporal arcade and the postorbital bone. Body length. The distance from the tip of the snout to the anterior margin of the vent, measured with dial calipers. Head length. The distance measured with dial calipers from the tip of the snout to the angle of the lower jaw at the posterior extremity of the articular. Upper labials. The number of scales bordering the upper lip, counted from the scale adjacent to the rostral to the corner of the mouth. NIGERIAN AGAMA LIZARDS 69 Lower labials. The number of scales bordering the lower lip, counted from the scale adjacent to the mental to the corner of the mouth. Lamellae under fourth toe. The number of transversely enlarged scales under the fourth toe from the base of the claw to the junction of the fourth toe with the third toe. Relative lengths of the toes. Comparisons of the first and fifth and of the third and fourth are expressed as the number of lamellae contained in the difference in lengths when the toes are laid flat and parallel to each other. C Q FIG. i. Supratemporal and temporal areas of Agama; underlying bones indicated by broken lines. E — Eye J— Jugal M — Maxilla P — Parietal Po— Postorbital bone Q — Quadrate S — Squamosal St — Supratemporal arcade T — Temporal fossa Ty — Tympanum DERMAL SENSE ORGANS Cohn (1914) was the first person to report on the occurrence of dermal sense organs in Agama and he described in detail their histology as well as their external appearance and distribution on various parts of the body. His study was restricted to Agama colonorum (= A. agama). Subsequently Schmidt (1920) and Preiss (1922) examined the scale organs of other species and genera of lizard, Schmidt extending his studies on reptile receptors to the Gekkonidae, Iguanidae and Agamidae but limiting his investigation of Agama to an Asian species A. sanguinolenta, which however exhibited several differences from A . agama, as described by Cohn, in the number and position of the organs on each trunk scale. It was not until 1937 and 1941 when Scortecci made an extensive survey of the lenticular and " hair-bearing " sense organs in the Agamidae that the density of these receptors and their positions on each scale were shown to have some value as a species character. (The agamid material which yo A. G. C. GRANDISON Scortecci used belonged to nineteen genera and eighteen species, his nine species of Agama all being African forms but none occurring in West Africa.) But despite the breakthrough afforded by Scortecci's meticulous comparative studies the potential of the receptor character for helping to distinguish agamid species was passed by, yet Underwood (1957) in surveying the scale organs in pygopod lizards had stressed that they would repay further study and Wayne King (1962) had effectively demon- strated their usefulness in sorting out the taxonomy of Caribbean sphaerodactyl geckos. The only attention recently given to these scale organs is by Miller and Kasahara (1967) who have studied their innervation and who remark on the tactile function of the " hair-bearing " receptor. In species whose mode of life is known correlations might be established between habits and the frequency with which the " hair " bearing organs occur, their position on the scales, their distribution on the body surface and the extent to which the " bristles " project beyond the edge of the scale. An investigation along these lines seems merited but unfortunately through lack of adequate field observations this could not be carried out on West African species of Agama. It has been found in the present study that the species number and the position of the " hair-bearing " organs on the trunk scale do not vary ontogeneticaly, nor sexually, but that there is some individual variation which is discussed under the species. Nor does this character vary geographically, at least not in four of the Nigerian species where material from throughout their known ranges has been examined. Whether there is geographic variation in Agama agama receptors has still to be established but none is evident in the Nigerian specimens. Another apparently reliable taxonomic character not previously employed in keying out agamas is introduced into this study, namely the direction of imbrication of the temporal and supratemporal scales. These two characters, receptors and direction of imbrication, both of which are probably related to the animal's ecology, seem to have greater significance in classification than some of the characters so rigidly adhered to in the past in keys and descriptions of species of agama ; certainly the results of studying them in the Nigerian species are encouraging and an extension of study to other groups of agamid lizards may help elucidate the status and relation- ships of the many forms that have been described. Agama agama (Linnaeus) (Map, PI. i, figs, c, d; Pis. 4, 5) Lacerta agama Linnaeus, 1758 : 207. MATERIAL EXAMINED. NIGERIA, Sokoto Prov., i mile north of Kware, 15 miles north of Sokoto B.M. 1962.1609; Katsina Prov., Rimi village, 13 miles South-east of Katsina B.M. 1962.1608; Kano Prov., Birnin Kudu B.M. 1962.1610; BornuProv., Maiduguri B.M. 1962.1611, Galtimare village, South of Maiduguri B.M. 1962.1612; Zaria Prov., Kawo, 4 miles north of Kaduna B.M. 1962.1596-98, Kujama, 25 miles South-east of Kaduna B.M. 1962.1599-1603, Zonkwa B.M. 1962.1604 and 1606, Birnin Gwarri B.M. 1967.2207, Galma Fadama B.M. 1967.2208, Old City Zaria NIGERIAN AGAMA LIZARDS 71 B.M. 1967.2209-2211, River Amethyst, Kaduna Road, Zaria B.M. 1967.2212; Plateau Prov., Jos B.M. 1962.1613-54, 10 miles south of Jos on Miango Road B.M. 1962.1593-95, Shendam B.M. 1962.1574-92; Oyo Prov., Igbetti Z.M.C. R3656i, 36567, 36569, 36573, 36565, 36644, 36655, 36745, 36286-9; Ondo Prov., Idanre Z.M.C. R36577-8, 36584, 36586. DESCRIPTION. Body subcylindrical, little depressed above, between 3-3 and 3-8 times the head length. The body of adult males rather triangular in section, the apex mid dorsal. Nostril on a line connecting the canthal supraciliaries with the rostral/first labial suture, that is on the canthal ridge ; situated in the posterior half or third of a strongly swollen, pear-shaped nasal shield and directed upwards, or upwards and outwards. A keel usually present between the nostril and the anterior edge of the nasal. On its posterior margin the nasal forms a suture, generally a broad suture, with the first canthal supraciliary. The supraorbital scales and the scales overlying the frontal and anterior portion of the parietal quite smooth, the other scales on the head keeled, particularly so in juvenile specimens. The direction of imbrication of the scales overlying the supratemporal vacuity is forwards but the direction of the scales covering the postorbital bone alters to downwards and slightly backwards. The scales overlying the temporal fossa, imbricate in the reverse direction and between the two areas and along an oblique line, which marks the position of the squamosal and the posterior limb of the jugal, is a single row of quite strongly keeled juxtaposed scales (PI. I, fig. c and Text-fig i). Occipital enlarged, of variable size, its greatest width generally half to three quarters the diameter of the tympanum. Seven to twelve upper labials; a similar number of lower labials. A long, low nuchal crest of approximately fourteen scales but no dorsal crest. A poorly developed caudal crest present in adult males. The shape of the ear roughly a right- angled triangle, the tympanic membrane rather superficial. No conspicuous fringe of long, pointed scales along the anterior border of the tympanum but a few erect scales which in juveniles may be only slightly raised. A row of erect or somewhat spinous scales from the commissure of the lips to immediately behind and below the tympanum where it joins with one or two (in adults invariably two) round groups of spinose scales on the swollen area at the angle of the jaws. At the upper posterior corner of the tympanum a less elevated and more elongated group of erect scales is present. Midway between a point half way along the nuchal crest and the lower posterior corner of the tympanum lies a small oval group of spinose scales; in front of and also behind this group a very small group of erect scales usually insignificant but if prominent the longest scales less than half the width of the tympanum. All these groups of erect scales are considerably more pronounced and the scales longer and more spinous in adult specimens, particularly in adult males, and in the juveniles the groups may be quite inconspicuous. A strong gular fold present in both sexes. Body scales homogeneous, those in the mid dorsal area flattened dorsoventrally, their distal margins rounded, often only feebly keeled and with short mucrones. Scales on the flanks more rhomboidal, particularly in adult males, the keels more pronounced and terminating in upward turning short mucrones ; distal margins some- what denticulate. Scales on either side of the nuchal crest small, barely mucronate, approximately equal in size to those between the neck pit and ear. 72 A. G. C. GRANDISON The sense organs of each trunk scale are situated at the distal margin or just below the dorsodistal edge of the scale. They are not confined to the area around the base of the mucrone but may extend along the entire distal aspect of the scale. No scale organ on a trunk scale has more nor less than one " bristle ". The number of receptors per scale varies from one to eight and while there is a tendency for the number to increase on the flanks and then diminish towards the venter no definite correlation could be established between the number of organs per scale and the longitudinal scale row although the body scale rows bearing the highest density are invariably between rows ten and sixteen. It is in this region of the trunk that the scales often have a rhomboidal shape with a somewhat denticulated distal margin and each organ is usually situated in an identation. Harris' (1963) statement that sense organs are absent from the ventral body scales is incorrect. Each ventral scale has one but no more than one " hair-bearing " sensory organ at the apex. The smoother scales of the middorsum generally have one to four sensory organs per scale. Cohn (1914) found only three organs on each of the scales of the back and describes the central one as placed somewhat to one side of the projecting tip of the keel. He reports a similar number of receptors on flank scales and says that the scales on the vertebral line may lack one or both lateral organs. He gives no indication of the origin of his material which he identifies as Agama colonorum, and there seems to be a strong possibility that a misidentification accounts for the much lower density of trunk scale organs in his specimens. Midbody scale counts of sixty-four Nigerian examples 59-77, the mean for thirty- five females being 68-2 and for thirty males 66-2; vertebral count 38-55. Mature males with eight to twelve, usually ten preanal pores. Eighteen lamellae under the fourth toe. First toe shorter than the fifth by two to seven (usually three to six) lamellae. Third toe shorter than fourth toe by one to two lamellae, very exceptionally subequal. Tail approximately twice the body length, depressed at the base and particularly so in adult males, its scales strongly keeled and mucronate and arranged in whorls. INTRASPECIFIC VARIATION. Loveridge (1936) in discussing intraspecific variation in Agama agama remarked on the extraordinary wide range of midbody count, 60-80, quoted by Boulenger (1885). Presumably Boulenger's counts were based solely on the material that was available in the British Museum at that time, namely twenty-one West African examples many of which bore no more precise locality than " West Africa ", and eight Angola and South West Africa examples, in other words a small sample from only a very limited portion of the range of the species. In the 1936 publication Loveridge demonstrated without regard to any sex difference a gradual reduction in the same count from east to west (Uganda to Senegal) and in the belief that populations occurring west of Ghana have a considerably lower average count he suggested recognizing these individuals as belonging to a distinct race, savatieri Rochebrune. The recognition of a western race based solely on this character was ill conceived and hasty, particularly in view of Loveridge having at his disposal only one individual from Ghana and none from the 1,000 mile stretch between Ghana and the Cameroun. In 1956 Grandison in reporting on over one NIGERIAN AGAMA LIZARDS 73 hundred Agama agama obtained in West Africa west of Ghana remarked on a south- west to northwest reduction from 68 to 60 in their average midbody count (range of the entire series 56-76). She assigned the individuals to A. agama savatieri, having overlooked the statement in Loveridge's later paper (1941) that Hallowell's name africana has priority over savatieri. Daniel's (1961) counts (M 61-6 <$ : 65-1 $) on twenty adult Liberian individuals which he identified as A. agama africana agree favourably with those given by Loveridge (1936, 1941) for Liberian material but suggest that Loveridge's samples in which the average number of scales round the body was sixty-two probably consisted mainly of males. The range of variation in midbody count in the sixty-five Nigerian examples studied here runs almost the whole gamut of the variation said to occur in africana but the average number of scales in males and females are intermediate between those quoted by Loveridge for Cameroun and Uganda individuals of the nominate form and those given by Loveridge, Daniel and Grandison for examples identified as africana and obtained west of Ghana. The Nigerian, Ghanaian and eighteen of the Senegal counts shown in the map are for adult specimens in the British Museum collection. U<^ 68-74( m, 21, m 45) ( 57-68(n 10 o; m 6/6) »»-72(n 35, m 66) S8-72(n 10?, \ 62-80(n 130*, m 70V-4) / <~ 68-74(n 7? 'm 71-41 [ n21, m65) 7 \ I !•»**»•*• FIG. ^ Geographic varation in midbody count in Agama agama. Thys van den Audenaerde (1963) in his study of Congolese species of Agama examined large samples of A . agama from the Congo and compared his findings with those of other authors for specimens from all regions in the range of the species. His thorough review of the pertinent literature available on Agama agama emphasizes the clinal sequence in midbody count from east to west although his argument is to some extent weakened by blind acceptance of identifications by others, to unequal sampling and to his not distinguishing between the counts for males and for females. He includes in his range of variation in A . agama a specimen from the Bauchi Plateau with ninety-two midbody scales that Grandison (1956) identified as A. sankaranica. While the individual is not available for re-examination it seems more than likely that it was incorrectly identified and is actually an example of A. benueensis, a species closely related to sankaranica and in its juvenile livery resembling sank- aranica but with a considerably higher midbody count. An A'ir specimen with a 74 A. G. C. GRANDISON low count of 58 was also mentioned by Grandison (1956) but excluded from the table of means; it too was probably incorrectly identified and may be an example of the large scaled species Agama boueti Chabanaud. This species was described from Gao on the Niger. Examination of type material shows that its trunk receptors are similar in number and position to those of A. bibronii Dumeril of North West Africa and are quite unlike any other West African form. Thys van den Audenaerde maintains that development of the keel and mucrone is correlated with body count, the lower the count the stronger the keeling, but I can establish no such link in the Nigerian material where variation in this character seems to be related to the proximity of sloughing. He further comments on the geographical variation that occurs in A . agama colouration, in particular of the head, throat and tail, which is perhaps correlated with the clinal gradient in scaling and he draws attention to Angolan intermediates between the nominate race and the Angolan race mucosoensis. A similar cline in the development of the nuchal crest which both Stejneger (1893) and Thys van den Audenaerde (1963) have suggested exists, has not yet been effectively demonstrated. Much more quantitative data on large samples from throughout the range of the species would have to be compared, sex with sex, if a true picture is to emerge. Meanwhile the only East African figures available are those of Stejneger based on only six Kenya examples, and of Barbour and Loveridge (1928) for eighteen adult Usambara specimens and in neither case is the sex or the proportion of the sexes stated. Since in Nigeria at least, there is a sex difference in the length and height of the crest, the spines being fewer and shorter in the females, the East African figures are not necessarily comparable with those for twenty adult male Nigerian individuals (13-17 : M 14-9). Thus the abundance of races of A. agama that have been described reflects not the occurrence of legitimate geographically localized subdivisions but accidents of sampling or more likely the work of splitters. There would appear to be no evidence of geographic isolates but only of marked individual variation within a population and Thys van den Audenaerde's conclusion that Agama agama is a large polymorphic species rather than a polytypic species would seem to be the logical one. COLOUR. Immature individuals and adult females are similar in colouration. In their reproductive colour phase the head is a medium brown or brown with green tinges, the back fawn or brown with darker brown marks of variable shape and position but often in the form of rather indistinct diamond-shaped outlines along the vertebral region. The upper part of the head, the temporal region and the neck have pale green spots and short longitudinal stripes of the same colour. The undersurfaces are a grey-white. Mature males that are dominant over territories are characterized in their reproductive colour phase by an orange head and neck, indigo blue body and by a median orange segment on a black-tipped tail. Other colour phases occur and these as well as the reproductive colour phase are admirably and fully described by Harris (1964). ECOLOGY. In Nigeria wherever the savanna and forest have been disturbed by human agency, bridges built, houses and shacks erected, trees felled, rubbish dumped, Agama agama seems to have established territories. It is very much a commensal NIGERIAN AGAMA LIZARDS 75 of man and is common around native houses, farm buildings, litter, wood piles and along paths. The species is also common in the boulder strewn country flanking the road from Jos to Miango. Schi0tz (personal communication) says that its habitat preferences do not include flat uniform savanna, nor large rocks, nor dense forest but as soon as such areas acquire human habitations or are in other ways disturbed by man Agama agama takes up residence. While this is largely true, the rocky mass of Kujama Hill south of Kaduna supports a number of Agama agama yet on the bare rock surfaces of the vast gneiss domes 1,000-3,000 ft. high at Igbetti and Idanre Agama agama is absent, its place being taken by Agama benueensis. RANGE. Senegal to Ethiopia, southwards to Tanzania and Angola. Agama paragama sp. n. (PI. i, figs, a, b; Pis. 4, 5) Agama agama benueensis Monard 1951 : 131 (part). MATERIAL EXAMINED. HOLOTYPE: NIGERIA, Zaria Prov., •£ mile Southwest of Ahmadu Bello University, Zaria City, collected by C. H. Fry from Parkia tree on 17.11.1964, B.M. 1967.2215 $. PARATYPES: NIGERIA, Zaria Prov., same locality as holotype B.M. 1967.2216, Mile 7, Zaria Road B.M. 1967.2214, Mile 5, Zaria Road B.M. 1967.2213, Ahmadu Bello University campus B.M. 1967.2217, no information, probably in vicinity of Zaria City B.M. 1967.2218, Birnin Gwarri, 62 miles Southwest of Funtua B.M. 1968. 497, Zaria City B.M. 1968.482-483, Veterinary Unit, Zonkwa B.M. 1962.1605, B.M. 1962.1607; BornuProv., Potiskum B.M. 1968.498, Yo B.M. 1968.496; Plateau Prov., Naraguta, Jos B.M. 1968.484-485. CAMEROUN, Bangouve M.H.N.C.F. 1162, Rei Bouba M.H.N.C.F. 1432, 1440, 1459, Mayo Sala M.H.N.C.F. 1374, Ngaouyanga M.H.N.C.F. 965 (six of the paralectotypes of Agama agama benueensis.} These individuals differ from A . agama not only in body size, proportions, greater spinosity and in lower average midbody and vertebral counts but in an absence of sexual dimorphism in size. They resemble A . agama in the position of the nostril being on the canthal ridge and in having multiple sense organs on the distal edges of the scales of the flanks and the two forms are undoubtedly closely related but being sympatric in Nigeria specific status is given to paragama. Relatively few examples of paragama have been collected and the form is so far known only from northern Cameroun and from the Northern Region of Nigeria, although one specimen obtained by C. H. Fry in a crevice 10 ft. up a Terminalia tree in the Botanical Gardens, University College, Ibadan was thought by him to belong to this form; the specimen has not been seen by me. Although situated in the high forest belt Ibadan has many clearings that have the character of cultivated savanna woodland and support typical savanna species. Schi0tz (1963) describes the vegetation of the University College campus as " dry forest and culture savanna ". It is possible that where suitable habitats are available this species, like Agama benueensis, penetrates the Western Region's forest belt. 76 A. G. C. GRANDISON DESCRIPTION OF HOLOTYPE. Body subcylindrical, depressed above, 3-6 times the head length. Nostril on the canthal ridge, situated in the posterior half of a swollen, pear-shaped nasal shield and directed laterosuperiorly. Nasal keeled from its anterior tip to the upper rim of the nostril, forming a suture on its posterior margin with the first canthal supraciliary. The supraorbital scales and the scales overlying the frontal and the anterior portion of the parietal quite smooth, remaining head scales keeled. The direction of imbrication of the supratemporal scales is forwards and slightly upwards; anteroventrally the scales overlying the postorbital bone face downwards and backwards. The scales immediately in front of the ear, that is those overlying the temporal fossa, imbricate in the reverse direction to those covering the lower postorbital and between the two areas and along an oblique line marking the position of the squamosal and posterior limb of the jugal is a single row of large, keeled, juxtaposed scales. Occipital enlarged, its greatest width equal to three-quarters the horizontal diameter of the tympanum. Upper labials 8 : 9 (one divided) ; lower labials 9 : 10. A short but conspicuous nuchal crest formed from nine, long, conical scales, the longest equal to three-quarters the width of the tympanum; no dorsal nor caudal crest. Ear triangular. On anterior margin of ear a fringe of two to three long, pointed scales, on upper posterior border one long, pointed scale surrounded by several erect scales (this group barely developed on left side of head). At the posterior extremity of the lower jaw, that is behind and below the ear, a small, round group of spinose scales of which one scale is three to four times longer than the others. A similar group immediately above. Between a point midway along the nuchal crest and the lower, posterior corner of the tymp- anum lies a prominent round tuft of long spinose scales the longest scale three- quarters the tympanic diameter; in front (right side only), and also behind, a small, round but insignificant group of erect scales. (The lengths of the scales comprising the nuchal crest and the groups of spinose neck and ear scales are considerably longer and more pronounced in male specimens : see note on variation.) A well developed gular fold. Body scales homogeneous, the longitudinal rows converging towards the midline, scales on the dorsum and flanks equally strongly keeled and mucronate ; the mucrones of the flank scales rather broad at their base, tapering rapidly and rising at an angle to the scale. Distal margins of the body scales rounded on each side of the mucrones. Ventral scales smooth. Scales on either side of the nuchal crest with strong, compressed keels and long, almost erect, mucrones, the scales at least twice the size of those between the neck pit and ear. The maximum number of sense organs on the middorsal and flank scales is nine, the minimum three. They are situated just under the dorsodistal margin and may extend as far as the " shoulders " of the scale. No scale organ on a trunk scale has more nor less than one " bristle ". The highest density of receptors on midtrunk scales occurs on longitudinal rows 3-5. Each ventral scale bears a receptor at its apex. Midbody scale count 60, vertebral count 30, eighteen lamellae under the fourth toe. First toe four or five lamellae shorter than the fifth toe, third toe half to one lamella shorter than the fourth. Tail one and a quarter times body length (tip of tail missing), depressed at base, NIGERIAN AGAMA LIZARDS 77 only very slightly compressed distally, its scales keeled and strongly mucronate and arranged in whorls. Length of body 94-0 mm. VARIATION. The following variation in meristic characters in the paratype series was noted. Nigerian individuals: midbody scale count 50-66 (M 58-5), vertebral count 26-34 (M 30-1), number of lamellae under the fourth toe seventeen to twenty- one, first toe shorter than fifth toe by between three and five lamellae, third toe half to two, usually one to two lamellae shorter than fourth toe. Differences in the develop- ment of the rosettes of neck and ear spines and in the lengths of the scales forming the nuchal crest are attributable to sex and maturity, in adult males the longest of the pointed nuchal crest scales being as much as equal to the width of the tympanum, likewise the most prominent scales in the neck tufts, whereas in females they are less than three-quarters the diameter of the tympanum. The number of scales form- ing the nuchal crest varies from eight to ten, the usual number being eight. The density and distribution of " hair-bearing " receptors on the midbody scales of the paratypes is similar to the holotype. Sex dichromatism is discussed below. The tail of the adult male is considerably more compressed than that of the female and has a slight crest ; the base of the tail is depressed but not to the same extent as in Agama agama. Juvenile A. par agama in which the neck and ear rosettes and nuchal crest are insignificant can be readily distinguished from young individuals of A. agama by the relative size of the scales above and below the central neck rosette and by the roundness of the rosette. Further distinguishing features are the pattern, lower vertebral count, gradual tapering of the tail and less depressed base to the tail. All the Cameroun examples, except an adult male (Mayo Sala M.N.H.C.F. 1374), fall within the range of variation in midbody count of the Nigerian individuals but they have a slightly higher number of transverse rows on the trunk, their vertebral count being 36-38. The Mayo Sala example with seventy-two midbody scale rows is well outside the normal range of variation in the species yet in its other morpho- logical characters it agrees with par agama. COLOUR. Male. In life the upper surfaces of the body and limbs of the adult <$ are a metallic blue with white flecks. On the back the flecks tend to be arranged in narrow transverse lines, no more than one scale in width, with approximately eight lines between fore and hind limbs. The vertebral zone is whitish, the head chalk white, the proximal two-thirds of the tail whitish, the tail tip black. Except for the whitish areas the general appearance of the $ is rather similar to the reproduc- tive colour phase of a male Agama agama. Harris (1964) describes in detail the various colour phases of both <$ and $ Agama agama and while he refers to a geograph- ical variation in the colour of the head of adult males, northern individuals (Kano) having a deep yellow while more southern populations have either orange or vermilion, he makes no mention of any adult male agama in which the orange or yellow tones are replaced by white. FEMALE. None of the colour phases described by Harris for female A. agama occurring in Nigeria corresponds to, or in any way approaches the colour pattern 78 A. G. C. GRANDISON of female A. paragama which is characterized by four or five pairs of more or less clearly defined, large, round, brown spots on a lighter brown (orange in life) background on the dorsum between fore and hind limbs. In those specimens in which the spots are ill-defined the lateral halves of the spots are indicated by narrow, dark brown outlines. The flanks and the upper surfaces of the limbs and tail are of a similar shade of brown to the spots; the head is mottled brown. In both sexes the throat, and in some specimens also the sides of the neck and chest, has a dark network on a cream ground which takes the form of isolated, round, cream spots and is dissimilar to the gular pattern of A. agama which tends rather to consist of a longitudinal arrangement of dark lines or blotches (Harris, 1964). The distinctness of the gular spotting does not appear to bear any relation to the maturity or sex of the individual, for example one gravid female has a pronounced pattern while in another a pattern is barely discernible. No pattern whatsoever is present on the throat of one adult male but a distinct network is present in another mature male Adult female paragama are larger and considerably more robust than female agama. Three gravid female paragama vary in body size from 96-107 mm. while the gravid female agama examined in this study vary from 86-91 mm. Harris (1964) records 97 mm. as the average body length of sixty-eight Nigerian adult female agama and Chapman & Chapman (1964) regard all Ghanaian female A . agama of about 90 mm. as mature. Adult male paragama are considerably smaller (99-108 mm.) than mature male agama obtained in the same general area (107-137 mm.). ECOLOGY. Little information on the ecology of paragama is available. Fry (in litt.) states that the species is to be found on the trunks and lower limbs of shade trees and collectors have suggested that it occupies a different ecological niche from A. agama but only the specimens collected at Mile 7 Zaria, Potiskum and Birnin Gwarri bear information that suggest an arboreal existence (Mango and Parkia trees) ; the Yo specimen, a juvenile, was taken on the walls of a house and the rest of the examined material have no biotope data. RANGE. West Africa: Northern Region of Nigeria to northern Cameroun. Agama benueensis Monard (PL 2, figs, a-d; Pis. 4, 5) Agama agama benueensis Monard, 1951 : 131 (part). Agama agama boensis Monard, 1951 : 130 (part: males). ? Agama sankaranica: Grandison, 1956 : 231 (part: Bauchi specimen). MATERIAL EXAMINED. CAMEROUN, Upper Benue, Ngaouyanga M.H.N.C.F. 956-7, Sakdje M.H.N.C.F. 1269, Mayo Sala M.H.N.C.F. 1373, Rei Bouba M.H.N.C.F. 1442, 1460, 1517 (seven of the paralectotypes of A. a. benueensis), Ngaouyanga M.H.N.C.F. 961 (designated lectotype of A. a. benueensis). NIGERIA, Oyo Prov., Igbetti Z.M.C. R 36342, 36345, 36347-51. 36353, 36389, 36392> 36394, 36397, 36399* 36401, 36599, 36601-12, 36614-20, 36661-2, 36702-3; Ondo Prov., Orosuta, Idanre Z.M.C. no number, Idanre Z.M.C. R 36576, 36579, 36580-2, 36585, 36588-9; Zaria NIGERIAN AGAMA LIZARDS 79 Prov., R. Amethyst, Kaduna Road B.M. 1967.2234-8, B.M. 1967.2263, Ahmadu Bello University campus, Zaria B.M. 1967.2239-44, Siberia, A.B. University campus Zaria B.M. 1967.2254-7, Old City, Zaria B.M. 1967.2245-7, B.M. 1967.2258-61, Mile 5, Zaria Road B.M. 1967.2248-52, B.M. 1967.2253, Zaria B.M. 1967.2262, Near Oomawa, at foot of Kwatarakwashi rock B.M. 1968.490-495; Plateau Prov., Jos B.M. 1968.486-487, Hillcrest School, Jos B.M. 1968.488. REMARKS. The whereabouts of four of the type specimens of Agama a. benueensis (959, 1372, 1441 and 955) listed by Monard (1951) are not known at present. Thys van den Audenaerde tells me he believes that part of the Monard material originally lodged in the herpetological collection at La Chaux de Fonds was transferred to other European institutions and that the missing Cameroun agamas may have been among it. The thirteen individuals from the benueensis type series that are available and have been re-examined belong to two morphologically very different forms, and as referred to in the introduction to this paper, the name benueensis is applied to the blue-throated form. The other form is described as A. paragama sp. n. The Monard specimens here assigned to A. benueensis are those males from the Upper Benue that are described by him (1951) as having " une tache noiratre a la gorge ", " une tache foncee a la gorge " and " une raie vertebrale claire ", as well as four females and a juvenile male that have a feebly developed pattern characteristic of immature and female Nigerian benueensis, and two males from Tibati, Adamaoua Plateau which he assigned to A, agama boensis. The Tibati males, M.H.N.C.F. 673, 784, have been examined by Dr. Braestrup who reports (in litt.) that both individuals have chin as well as gular spots. The presence of the gular spots, even if faint, identifies them as benueensis for this species is the only West African agama that has a bilobed blotch in this region in the adult male. In the Tibati males the midbody count is about 74, the vertebral counts 45 and 47 which accord with Cameroun examples of benueensis. Neither I nor Dr. Braestrup has seen the female, M.H.N.C.F. 726, which Monard also referred to A. a. boensis and its identification remains in doubt. DIAGNOSIS. A large sized Agama (body length of $ up to 113-0 mm., of $ to 74-0 mm.) related to A. sankaranica and characterized by a high average number of midbody scale rows, 74-98 (M 88 : N 71) in Nigerian populations, ventrally directed temporal scales and marked sexual dimorphism. Dorsal trunk scales each with one to three single " hair-bearing " scale organs close to base of mucrone, nostril below canthal ridge; in these respects similar to A. sankaranica. DESCRIPTION OF LECTOTYPE M.H.N.C.F. 961, <$. Body subcylindrical, little depressed above, 3-7 times the head length. Nostril below a line connecting the canthal-supraciliaries with the rostral-labial suture, its aperture round, situated in the posterior third of an elongated, pear-shaped nasal, directed superior-laterally. From the upper rim of the nostril to the front of the nasal shield is a distinct ridge. Canthal-supraciliaries 8:9, the anterior forming broad sutures with the nasals at their posterior borders. Supraorbitals quite smooth, supratemporals distinctly keeled, temporals only very slightly keeled. The scales overlying the supratemporal arcade and the postorbital bone directed forwards and slightly upwards; ventrally 8o A. G. C. GRANDISON the imbrication of the scales covering the posterior part of the jugal, the broad portion behind and below the eye, as well as those over the temporal fossa directed downwards and slightly backwards; no row of keeled juxtaposed scales. Occipital plate enlarged, broadly oval, its length equivalent to the horizontal diameter of the tympanum. Seven to ten upper labials, eight to twelve lower labials. Tufts of enlarged, conical, rather flat scales around the ear, one, consisting of three scales, on the anterior border, one on the upper posterior border, on the lower posterior border a group of about four to five more prominent conical scales and below this group a curved row of about five flattish conical scales. At the side of the neck midway between the tympanum and a shallow neck pit, but a little above, a rosette of spines, the longest scale in the rosette two-thirds the length of the nasal. Behind and a little below this rosette and situated above the neck pit an oblique or vertical line of conical scales. Above and slightly behind the tympanum a prominent erect scale surrounded by a few lower but erect scales. Ear subtriangular, its horizontal diameter equal to the distance from the tip of the snout to the middle of the nasal. A gular pouch present. Nuchal crest low, formed from about ten conical scales; no dorsal nor caudal crest. Dorsal scales imbricate, homogeneous, strongly converging on the vertebral line, strongly keeled but the keel not markedly compressed, mucrone finely tapering, ventrals and subcaudals smooth. A maximum of two scale organs on each body scale but usually only one and situated under the dorsodistal edge of the scale close to the base of the mucrone. Each scale organ bears a single " hair ". Midbody scale count 78; vertebral count 46; preanal pores in two transverse rows, 13 anteriorly, 15 posteriorly. The adpressed hind limb reaches to posterior border of the tympanum. Nineteen lamellae under the fourth toe. First toe three or four lamellae shorter than fifth toe, third toe one lamella shorter than fourth toe. Tail slightly depressed at the base; broken. Caudal scales keeled and mucronate, arranged in distinct whorls. Body size 94-0 mm. COLOUR IN ALCOHOL. Apart from a conspicuous whitish vertebral line approxi- mately three scales wide and extending from the nape to the base of the tail there is little colour pattern discernible. Any pattern that may have been present on the flanks of the living animal has become obscured by preservation and sloughing and only a slight mottling of cream and dark brown is now evident. The throat bears a large, blackish, bilobed patch at the level of the gular fold ; in front of this patch and extending on to the chin are several small, dark spots on a cream background. The rest of the undersurfaces are a grey-cream. COLOUR IN LIFE OF NIGERIAN EXAMPLES. Dorsum grey or brown with whitish dark-edged ocelli on the flanks which are arranged in two or three longitudinal rows, the spots in the outer, or middle in the case of individuals with three rows on each side, generally larger than the others. Across the back from the shoulders to the tail a series of dark chocolate brown bands which, particularly in juveniles, are considerably broader and often diamond-shaped on the vertebral line and enclose a narrow, longitudinal, whitish streak. The streaks become confluent in some adult NIGERIAN AGAMA LIZARDS 81 males and females (see sex dimorphism) and form a diffuse yellowish vertebral stripe. Schi0tz (personal communication) describes the entire dorsum and belly of adult males as having a distinct bluish tinge, but not of such an intense blue as the Agama agama dominant male, while the chin and throat are yellow with two to four dark blue or black patches. The upper surface of the male's head is irregularly mottled green; the tail is greenish with indistinct yellow bands. The ground colour of the female varies according to the colour of the rocks or soil in its habitat. At maturity the transverse bands on the trunk are reddish brown and the sides of the head are greenish. The ventral surface of both the female and the juvenile is white but the chin and gular region of most individuals has a network of dark lines forming irregular spots. INTRASPECIFIC VARIATION. An analysis of meristic variation in the material examined shows some geographical differences. The samples from the western region (Igbetti and Idanre) have the highest midbody and vertebral counts, midbody 84-98 (M 91-3 : N 34), those from the Jos Plateau and the Cameroun the lowest, 68-82 (M 76 : N n) while the gradation in the Zaria Province population almost spans these two extremes, 74-96 (M 85-8 : N 34). The variation in these characters for the entire series is : midbody 68-98 (M 87-0 : N 79), vertebral 41-56 (¥486 : N 76). The correctness of the identification of the individuals at the extremes of the variation is not in doubt, a combination of other characters such as throat mark- ings, position of nostril, trunk scale organs and direction of imbrication of the post- orbital/temporal scales assigning them unquestionably to benueensis. Variation in the number of fourth toe lamellae (nineteen to twenty-three) and in the relative lengths of the first and fifth, and third and fourth toes is independent of the variation observed in midbody and vertebral counts and appears to have no geographical nor ontogenetic significance. The third toe may equal the fourth but generally it is shorter by half to one lamella. Variation in the relative lengths of the first and fifth toes is greater; the first is always shorter than the fifth but the difference may be as many as four lamellae. In the majority of the examples studied the lateral head scales between the eye and the ear all imbricate downwards or downwards and slightly backwards but in a few individuals, juveniles as well as adults, the scales in the lower posterior quarter of this area, that is those overlying the temporal fossa are directed forwards and upwards, in other words in a similar direction to those of Agama agama; however they are not as strongly keeled as in agama nor are they separated from the post- orbital /supratemporal scales by an oblique row of juxtaposed scales. Scales on either side of the nuchal crest are not or only slightly mucronate and are only slightly keeled. They are considerably larger than the scales below the central group of spines on the side of the neck. SEXUAL DIMORPHISM AND DICHROMATISM. Some sexual dichromatism is evident, notably in the conspicuous gular and chin spots which are a characteristic feature of the adult male. The twin gular spots develop before the chin spots and first appear as a darkening of the median gular region which later becomes an intense black or navy bilobed blotch; occasionally the lobes are separated on the midline. ZOOL. 17, 3. 4 82 A. G. C. GRANDISON The gular spots do not develop until the male reaches a body size of over 73 mm., although there is one exception among the fifty-six males examined, a 69-7 mm. individual collected at Igbetti. By 80 mm. most of the males have developed two additional dark spots; these are on the chin and somewhat oval and parallel to each other on either side of the median line. The development of these gular and chin spots in adult male benueensis does not seem to be seasonal; they are present in Nigerian individuals collected in January, February, March, June, July and August. The three largest males in the Cameroun series (72-5, 78-3 and 74-0 mm.) which were obtained in July and in September have only gular spots. Adult males retain some evidence of the juvenile pattern; generally the two longi- tudinal rows of ocelli along each flank are discernible, although feebly, as well as remnants of the five dark transverse lines in the mid dorsal region on each side of the midline from shoulders to groin but the light coloured vertebral streaks, a character- istic feature of the juvenile livery, invariably coalesce and form an ill defined light vertebral zone extending from the nape to the tail. The female sample is insufficient to be certain whether a similar light vertebral zone always develops at maturity. Only six of the fifteen females are gravid and of these six only the two Cameroun examples (M.N.H.C.F. 1460, 1269) have a clearly defined continuous light line. In the other examples the vertebral pattern is obscure in all but one of the Igbetti examples (Z.M.C. 36351). In both sexes the dorsolateral series of ocelli becomes progressively less distinct as the individuals mature. Egg bearing females were obtained in July, August and September. Preanal pores are usually ten to twelve in a single series but in three Idanre males, the only male benueensis collected there, and in the lectotype the pores are in double series 13 + 14, 10 + 13, 8 + 13, 13 + 15 respectively. ECOLOGY. In the western region of Nigeria Schi0tz (personal communication) says that A. benueensis is strictly confined to large rocky outcrops such as the vast gneiss inselbergs where it is common both in the savanna (Igbetti) and in the forest (Idanre). Other ecological niches that he searched in southern Nigeria, such as mounds of boulders, small rocky areas, produced no examples. He noted that benueensis hides in rock crevices and among vegetation at the perimeter of an insel- berg but conducts its feeding and sexual activities on the actual surface of the rock. Farther north on the Jos Plateau the species is much less abundant but it is again quite common in Zaria Province on the inselbergs around Zaria City as well as in a variety of other habitats such as on mounds of laterite on open farmland in the Galma Fadama, a locality some miles from the nearest inselbergs, on sandy river banks (Amethyst River) and in sandy stream beds (R. B. Walker, personal communi- cation) . Near Sabon Gari, in an area of scrap cars and garbage tips, both A . benueensis and A. agama were taken and no specific habitat preference was noted but Schi0tz maintains that at Igbetti A. benueensis, A. sankaranica and A. agama can be seen virtually only a few feet from each other yet each still adhering to its preferred habitat. A careful search by Schi0tz of the Shai Hills in Ghana and suitable localities in the Eastern Region of Nigeria failed to produce any benueensis and he believes the species NIGERIAN AGAMA LIZARDS 83 may also be absent from the Bamenda region of the Cameroun. Agama benueensis seems to be endemic to the Western and Northern regions of Nigeria and the upper Benue valley in the Cameroun. Monard (1951) does not record the biotopes for the type series of benueensis and he describes only the general areas around the villages. The I : 1,000,000 maps of the Upper Benue show the villages where benueensis were taken as being in an area deeply dissected by the Benue and its tributaries and at altitudes varying from 740 ft. to 1,400 ft.; only Ngaouyanga is on a high steep-sided wooded platform, the other localities are in or at the sides of valleys in woodland savannah. RANGE. West Africa: Western and Northern Regions of Nigeria and northern Cameroun. Agama sankaranica Chabanaud (PI. 3, figs, a, b; Pis. 4, 5) Agama sankaranica Chabanaud 1918 : 105. Agama boensis Monard 1940 : 155 (part: adults only). MATERIAL EXAMINED. PORTUGUESE GUINEA, Madina Boe M.H.N.C.F. 865, 867-8 (three of the syntypes of Agama boensis). REPUBLIC OF GUINEA, Beyla M.N.H.N. 21.309-316, Kankan M.N.H.N. 21.297-298, Kerouane M.N.H.N. 21.299-308, B.M. 1921.11.12.1-5. MALI, Moussaia, Sankaran M.N.H.N. 1901.395 (holotype of Agama sankaranica) . GHANA, B.M. 1930 .6.9.8 Korley Bu, Accra B.M. 1931 .5.6.3, B.M. 1932.6.1.5-7, Tamale B.M. 1927.9.27.199, Tafo B.M. 1962.910. NIGERIA, Benue Prov., Makurdi B.M. 1937.12.4.14; Plateau Prov ., Shendam B.M. 1962.1573; Kano Prov., Kano B.M. 1961.2069, B.M. 1962.567 (no. 567 later skeletonized); Zaria Prov., B.M. 1967.2206, Rimi, South-east of Kaduna B.M. 1962.1572, Zonkwa B.M. 1962.1571, Ahmadu Bello University campus, Zaria B.M. 1967.2205, 14 miles South of Zaria B.M. 1967.2204; Lagos Prov., Lagos B.M. 1960.1.7.28; Oyo Prov., Igbetti Z.M.C. R 36386, 36388, 3639°. 36393, 36395-6, 36398, 36403, 36405, 36407, 36597, 36607, 36697-8, 36691, 36758. DIAGNOSIS. A medium sized Agama (body length of adult males averaging 66 mm., of mature females 76 mm.) belonging to the group of agamas that have the nostril below the canthal ridge, homogeneous scaling and dermal sense organs not exceeding three in number on each trunk scale. Related to Agama benueensis Monard but distinguished by a lower average midbody count (69-6) and by dorsally directed temporal scales. DESCRIPTION. Body subcylindrical, scarcely depressed above, 3-3-7 times the head length. Nostril below the canthal edge, situated in the posterior half of a slightly swollen, oval nasal, directed laterally. No post nasal separating the first canthal supraciliary from the nasal. Scales of the head keeled; the interorbital scales considerably smaller than the supraorbitals ; imbrication of temporals directed upwards and of supratemporals directed upwards and forwards (PI. 3 fig. b) Occipital large, its greatest width equivalent to or as much as one and a half times greater than the diameter of the tympanum. Eight to twelve upper labials; a ZOOL. 17, 3. 4§ 84 A. G. C. GRANDISON similar number of lower labials. Usually three tufts of long, conical spines on pos- terior border of ear ; behind and above these a single group of spines and yet farther back and just above a shallow neck pit another group of somewhat shorter spines. A weakly developed gular pouch. A low nuchal but no dorsal or caudal crest. Dorsal scales homogeneous, broadly rounded, strongly keeled and mucronate, the sharp high keel extending the length of each scale, the mucrone very slender. Scales on the flanks slightly smaller than dorsals, similarly keeled and mucronate. Ventrals keeled only in juveniles. " Hair-bearing " sensory pits on the dorsal midtrunk scales confined to around the base of the mucrones, at or just under the dorsodistal edges of the scales. The maximum number of scale organs on each scale is three but invariably no more than two are present. In the vertebral region at midbody there are generally two " hair-bearing " pits, each with a single hair-like projection, the one pit close to the base of the mucrone or under the mucrone and concealed in dorsal view, the other on the opposite side of the mucrone but near the apex of the scale. On more lateral scales there is a tendency for only the more lateral, that is lateral with respect to the mucrone, to develop. The number and position of the scale organs on the dorsal scales bear a close similarity to the condition obtaining in East African rueppelli and described and figured by Scortecci (1937). Midbody scale count 64-78 (M 69-6 : N 61); vertebral count 32-46 (M 38-8 : N 61). Mature males with eight to twelve usually ten preanal pores in a single transverse row. Fifteen to twenty-one lamellae under the fourth toe. First and fifth toes subequal or first toe one lamella shorter. Third toe usually subequal to fourth toe, rarely half to one lamella shorter or longer than fourth. Tail one and three quarter times the body length, depressed at the base, slender and cylindrical distally, its scales strongly keeled and very distinctly mucronate, not arranged in distinct whorls. COLOUR IN ALCOHOL. Upper parts reddish brown. Invariably a conspicuous yellowish or russet vertebral stripe from behind the head to the proximal third of the tail, much broader and always prominent in the cervical region but narrowing rapidly to a width of approximately two scales on the rest of the body. On each side of this stripe and from nape to tail broad, dark brown, transverse bands, six between the nape and hind limbs. At the lateral extremities of these bands light-coloured ocelli or elongated blotches which in some specimens coalesce to form a light dorsolateral stripe. In some examples the vertebral stripe widens at the level of each transverse band to form light diamond-shaped areas. Halfgrown individuals occasionally have indistinct marbling on the flanks below the ocelli or dorsolateral stripe. In one mature male, collected at Zaria, the transverse bands are broken up into irregu- larly shaped blotches. Between the eyes a thin dark transverse line is present; four similar lines radiating from the eyes to the lips are usually evident. Limbs indistinctly cross-barred with narrow dark lines. SEXUAL DIMORPHISM. There are no obvious external differences between the females and immature males. Adult males have eight to twelve preanal pores and also broken, longitudinal, blue stripes on the chin and throat which become more prominent with maturity and often enclose a large, dark blue, median gular spot NIGERIAN AGAMA LIZARDS 85 and a lozenge-shaped chin blotch. The rosettes of neck spines and the lips may also be blue in mature males. Adult females are on average rather larger than adult males. The body length of mature females varies from 68-85 mm- (average 76-2 mm.) while males vary from 62-76 mm. (average 66-4 mm.). REMARKS. The examined material consists of examples from throughout the known range of the species. No evidence of geographical variation can be demon- strated. The examples from the extreme west of the geographical range of sank- aranica are the three adults, two males and one female, in the series of syntypes of Agama boensis Monard. These individuals unquestionably belong to sankaranica. The throats of the males have the blue blotches and stripes that are typical of mature male sankaranica, furthermore the scale counts and other morphological characters including the density and distribution of trunk receptors agree well with sankaranica although one male with a midbody count of 64 is exceptional in having 20 : 21 lamellae under the fourth toe, which is outside the range for all other specimens examined (fifteen to nineteen). The fourth syntype of boensis, a juvenile, lacks a nuchal crest and clusters of neck spines and has a single apical scale organ on each body scale; it is conspecific with Agama weidholzi Wettstein. (Grandison, in press.) The Cameroun individuals assigned by Monard (1951) to A. a. boensis are discussed in the section on A. benueensis. ECOLOGY. In the field this species has often been confused with female and imma- ture Agama benueensis and as a result some ecological information claimed to be on A. sankaranica has had to be disregarded or treated with suspicion in this study. Dr. Arne Schi0tz' reliable field notes and comments make it clear that he readily distinguishes sankaranica from benueensis and his notes correspond with my own field observations in stating that unlike all other Nigerian species of Agama, with the exception of gracilimembris, sankaranica does not form family groups but is a solitary creature. The species has been observed and captured in ploughed farmland and in maize and cassava plantations when running along the rutted ground, also on paths in grass covered savanna. Schi0tz' notes state that he saw the species on the ground only in grass covered savanna or in rather dense tree savanna and never in forest nor on rocks. The localities at which examples of A. sankaranica have been collected are, with one exception, in the Doka and True Guinea woodland belts from Nigeria westwards to Portuguese Guinea; the exception is an individual taken at Kano which is towards the southern limit of Sudan woodland. RANGE. West Africa: Portuguese Guinea to Nigeria. Agama gracilimembris Chabanaud (PI. 3, ngs. c, d; PI. 6) Agama gracilimembris Chabanaud, 1918 : 106. MATERIAL EXAMINED. DAHOMEY, M.N.H.N. 04.114-5 (syntypes). NIGERIA, Benue Prov., Lafia B.M. 1938.3.1.47, Wukari B.M. 1938.3.1.48-9; Plateau Prov., Shendam B.M. 1962.1570; Zaria Prov., Zonkwa B.M. 1961.949, B.M. 1962.1569, Samara Bush, Zaria (on exchange to Vienna Mus.) ; OyoProv., Igbetti Z.M.C. R 36654, R 36701, R 36263; Kano Prov., Kano B.M. 1961.2067-8, B.M. 1962.566, CENTRAL 86 A. G. C. GRANDISON AFRICAN REP., " Pays des Senoussi " M.N.H.N. 17.191 (according to information received from Dr. J. Guibe this specimen was collected in the neighbourhood of Ndelle, 8° 25' N : 20°36'E). DIAGNOSIS. A small sized Agama (body length of males up to 47 mm., of females to 57 mm.) lacking tufts of erect spiny scales behind the ear and bearing only one receptor on each body scale, related to Agama weidholzi Wettstein but distinguished by heterogeneous body scales, strongly keeled head scales and a low fourth toe lamellar count (13-14). DESCRIPTION. Body slightly depressed, 3-3-6 times the head length. Nostril below the canthal edge, situated in the posterior half of a convex, often keeled, oval or pear-shaped nasal; directed laterally. Interorbital scales as large or larger than the supraorbitals. Above the nasal a continuous series of three clearly defined, somewhat elongated scales runs from the rostral to the canthal-supraciliaries. The first canthal-supraciliary is separated from the nasal by a small postnasal. Scales of the head strongly keeled, somewhat rugose ; imbrication of temporal scales directed downwards and slightly backwards. Occipital large, its greatest width equivalent to the diameter of the tympanum. Nine to twelve, usually ten upper labials; eight to eleven, usually ten lower labials. No tufts of spines around the ear, instead single, short, conical scales close to the border of the ear and three to four tubercles on the side of the neck in a slightly curved row extending from the upper edge of the ear to above a point midway between the neck pit and the arm insertion ; addi- tional tubercles irregularly scattered below this row and between the ear and the neck pit, the lowest at the jaw angle. No gular fold. No trace of a nuchal or dorsal crest. Dorsal scales heterogeneous, those in the vertebral region a little larger than the lateral series; on the flanks, and irregularly disposed, some large scales which are as big and occasionally bigger than the vertebrals. The scales of the midtrunk region broadly oval or pear-shaped with pronounced " shoulders ", dorso ventrally flattened on each side of a high keel, the edges of which rise abruptly from the scale. The keel extends half to two-thirds the length of the scale but does not project beyond the scale apex to form a mucrone. A single sensory organ is present on the dorsodistal surface of each of the midbody scales just under the keel tip or very slightly to one side and sometimes evident only when the scale is viewed from behind. Generally each of these organs bears one but never more than one " hair ", which projects slightly beyond the apex of the scale and by macro- scopic inspection may be mistaken for a mucrone. Gular and ventral scales strongly keeled. Midbody scale count 70-85 (M 74-8 : N 16) vertebral count 30-46 (M 37 : N 15). Mature males with eight to twelve preanal pores in a single transverse row. The adpressed hind limb reaches to the tympanum. Thirteen to fourteen lamellae under the fourth toe. The first toe is usually either one lamella longer than the fifth toe or equal to it; rarely (two examples) is it shorter (half lamella). The third toe exceeds the fourth toe in length by half to two lamella. Tail one and a half times the body length, depressed at the base, slender and cylindrical distally its scales strongly keeled and slightly mucronate, and not arranged in whorls. Body length of gravid females 49-0-56-7 mm., of mature males 44-0-47-0 mm. NIGERIAN AGAMA LIZARDS COMPARISON OF CHARACTERS OF NIGERIAN SPECIES OF Agama agama* paragama* benueensis* sankaranica gracilimembris Dorsal body Homogeneous scales Homogeneous Homogeneous Homogeneous Heterogeneous Position of On canthus On canthus Below canthus Below canthus Below canthus nostril Dorsal body + scales keeled + (strongly) + + (strongly) Keeled but not mucronate and mucronate Keeled ventrals — — + (juveniles only) + Midbody 59~77 count (M68 • 2 ? : M66 • 2 (i) lamellae) Third to fourth toe = or<(i-3) <(i-2) = or <(i-i) — or<(i) or >(I~2) Body length 84-91 mm. of gravid °. 96-107 mm. 65-74 mm. 68-85 rnm. 49-57 mm. Body length 107-137 mm. of adult c? 99-108 mm. 80-113 mm. 62-76 mm. 44-47 mm. Tufts of spines + behind ear and + + + — on side of neck * Based on Nigerian individuals only. 88 A. G. C. GRANDISON COLOUR IN ALCOHOL. <$ Upper surfaces of body and limbs greyish brown ; usually a clearly defined light vertebral stripe which extends from behind the occiput to the base of the tail. Straddling the vertebral stripe from nape to base of tail are nine < > shaped, dark chocolate brown marks which in some specimens are rather obscure. lu the same region and alternating with these marks, diamond-shaped brick orange zones occasionally present. The lips, snout, supraorbital and tympanic and parietal regions invariably dark brown or blue black but a lighter area over the occipital is always present. Dark, irregular, longitudinal lines are usually present on the throat, chest and belly but they may extend no farther back- wards than the gular region. In a sexually mature male these lines are dark brown and closely set to give a brownish appearance to the entire belly; the throat has a dark median area. 9- The entire dorsal surfaces of the body, tail and limbs brown with obscure, irreg- ularly placed lighter areas. An ill defined light vertebral stripe occasionally present. Snout, supra-orbital area, cheeks and olbique stripe from below eye to commissure of lips dark brown; occipital, temporal and nuchal regions light fawn. Above and behind each ear a large ill defined blackish patch in which the small pointed scales stand out clearly as light bluish or greyish spots. Lower surfaces greyish white with faint longitudinal dark lines from throat to belly. In some specimens these lines do not extend farther than the chest. ECOLOGY. A. gracilimembris is the rarest of the West African savannah species and has been recorded from only a few localities all of which are in the Doka and True Guinea woodland vegetation belts and on the extreme southern limit of Sudan woodland (at Kano). Its biotope has not been described. RANGE. West Africa : Dahomey to Ubangi Chari. ACKNOWLEDGEMENTS The stimulating enthusiasm, interest and collecting ability of R. B. Walker and Dr. C. H. Fry of the Ahmadu Bello University is gratefully acknowledged and thanks are extended to Prof. A. P. Mead for his cooperation. Thanks are also due to Dr. G. T. Dunger who co-operated in obtaining samples from the Jos area. I am especially indebted to Dr. A. Schi0tz, Danmarks Akvarium for graciously making available his field notes on his extensive collections and for valuable discussions and hospitality during my Copenhagen visit. Dr. F. W. Braestrup, Copenhagen Museum has not only taken a keen interest in this study but has been a source of encourage- ment and assistance. To Professor J. Guibe, Paris Museum, Dr. V. Aellen, Geneva and Dr. R. Matthey, La Chaux de Fonds I am also indebted for the loan of speci- mens, for supplying information on specimens of Agama in their care or for museum facilities. I wish to thank too A. F. Stimson, Paula D. Jenkins and Belinda A. Brindley for their help in scale counting and in preparing slides, skeletons and line drawings. Summary oftaxonomic changes : Agama boensis Monard part = A . sankaranica Chabanaud Agama boensis Monard part (juvenile) = A. weidholzi Wettstein Agama a. benueensis Monard part = A . benueensis Monard Agama a. benueensis Monard part = A . paragama sp. n. NIGERIAN AGAMA LIZARDS 89 REFERENCES BARBOUR, T. & LOVERIDGE, A. 1928. A comparative study of the herpetological faunae of the Uluguru and Usambara Mountains, Tanganyika Territory, with descriptions of new species. Mem. Mus. Comp. Zool. Harv. 50 (2) : 87-265, 4 pis. CHABANAUD, P. 1917. Enumeration des reptiles non encore etudies de 1'Afrique occidentale, appartenant aux Collections du Museum, avec la description des especes nouvelles. Bull. Mus. Hist. nat. Paris, 23 : 83-105, 13 text figs. 1918. Etude complementaire de deux Agama de 1'Afrique occidentale et description de quatre especes nouvelles de reptiles de la meme region. Bull. Mus. Hist. nat. Paris, 24 : 104-112. CHAPMAN, B. M. & CHAPMAN, R. F. 1964. Observations on the biology of the lizard Agama agama in Ghana. /. Zool. Lond. 143 : 121-132, 6 figs. COHN, L. 1914. Die Hautsinnesorgane von Agama colonorum. Zool. Anz. 44 : 145-155, 7 text-figs. DANIEL, P. M. 1961. Notes on the life history of Agama agama africana (Hallowell) in Liberia. Spec. Publs. Ohio herpet. Soc. no. 3 : 1-5, 2 tables. GRANDISON, A. G. C. 1956. On a collection of lizards from West Africa. Bull. Inst. fr. Afr. noire (A) 18 : 224-245, 4 text-figs, 2 maps. 1969. Agama weidholzi (Sauria: Agamidae) of West Africa and its relationship to Agama gracilimembris. Ibid, (in press). HARRIS, V. A. 1963. The anatomy of the Rainbow Lizard. Hutchinson Tropical Monographs, London: 1-104, 39 text-figs. 1964. The life of the Rainbow Lizard. Hutchinson Tropical Monographs, London: 1-174, 39 text-figs. KEAY, R. W. J. 1959. Vegetation map of Africa south of the Tropic of Cancer. Oxford University Press, London. KING, W. 1962. Systematics of lesser Antillean lizards of the genus Sphaerodactylus. Bull. Fla. St. Mus. biol. Sci. 7 : 1-52, 17 text-figs. LINNAEUS, C. 1758. Systema Naturae. Tenth Edition: 824 pp. LOVERIDGE, A. 1936. African reptiles and amphibians in Field Museum of Natural History. Publs. Field Mus. nat. Hist. Zool. ser. 22 : i-m. 1941. Report on the Smithsonian-Firestone Expedition's collection of reptiles and amphibians from Liberia. Proc. U.S. natn. Mus. 91 : 113-140, i text-fig. 1952. Mission A. Villiers au Togo et au Dahomey (1950) XII. Tortoises and lizards. Bull. Inst. fr. Afr. noire (A) 14 : 229-242. MILLER, M. R. & KASAHARA, M. 1967. Studies on the cutaneous innervation of lizards. Proc. Cal. A cad. Sci. 34, 16 : 549-568, 20 text-figs. MONARD, A. 1940. R6sultats de la Mission scientifique du Dr. Monard en Guinee Portugaise (I937-I93^)- VIII Reptiles. Archos. Mus. Bocage, 11 : 147-180, 2 pis., 5 text-figs. 1951. Resultats de la mission zoologique suisse au Cameroun. Mem. Inst.fr. Afr. noire, 1 : 1-244, ! 3 figs. PREISS, F. 1922. Uber Sinnesorgane in der Haut einiger Agamiden. Jena Z. Naturw. 58 : 25-76, 9 text-figs. ROCHEBRUNE, A-T, DE, 1884. Faune de la Senegambie. Reptiles. Octave Doin, Paris, 89 : 1-22 1, 20 pis. SCHIOTZ, A. 1963. The Amphibians of Nigeria. Vidensk. Medd. dansk. naturh. Foren. 125 : 1-92, 28 text-figs., 4 pis. SCHMIDT, K. P. 1919. Contributions to the herpetology of the Belgian Congo based on the collection of the American Congo Expedition, 1909-1915. Bull. Am. Mus. nat. Hist. 39 : 385-624, 27 figs., 32 pis. SCHMIDT, W. J. 1920. Einiges uber die Hautsinnesorgane der Agamiden insbesondere von Calotes, nebst Bemerkungen uber diese Organe bie Geckoniden und Iguaniden. Anat. Anz. 53 : 113-139, 1 6 text-figs. go A. G. C. GRANDISON SCORTECCI, G. 1937. G"li organ! di senso della pelle degli Agamidi. Memorie Soc. ital. Sci. nat. 10 (u) : 159-206, 39 text-figs., 2 pis. 1941. I recettori degli agamidi. Memorie Soc. ital. Sci. nat. 10 (m) : 209-326, 80 text- figs. STEJNEGER, L. 1893. On some collections of reptiles and batrachians from East Africa and the adjacent islands, recently received from Dr. W. L. Abbott and Mr. William Astor Chanler, with descriptions of new species. Proc. U.S. natn. Mus. 16 : 711-741. THYS VAN DEN AUDENAERDE, D. F. E. 1963. Les Agamidae du Congo: les especes et leur distribution geographique. Rev. Zool. Bot. afr. 68 : 203-250, 4 text-figs. UNDERWOOD, G. 1957. On lizards of the family Pygopodidae. A contribution to the morph- ology and phylogeny of the Squamata. /. Morph. 100 (2) : 207-268, n text-figs. WERMUTH, H. 1967. Liste der rezenten Amphibien und Reptilien: Agamidae. Tierreich, 86 : 1-127. WETTSTEIN, O. 1932. Eine neue Eidechse aus Senegambien. Zool. Anz. 99 : 303-305, i text-fig. PLATE I a. Agama paragama sp. n. Holotype B.M. 1967.2215. b. Agama paragama sp. n. Lateral view of the head of the holotype. c. Agama agama L. Lateral view of the head of a juvenile female B.M. 1967 .2212. d. Agama agama L. Dorsal view of same individual. Bull. BY. Mus. nat. Hist. (Zool.) 17, 3 PLATE i ZOOL. 17, 3. PLATE 2 a. Agama benueensis Monard. Dorsal view of an immature male B.M. 1967.2239. b. Agama benueensis Monard. Dorsal view of an adult male B.M. 1967.2234. c. Agama benueensis Monard. Throat of a sexually mature male B.M. 1967.2253. d. Agama benueensis Monard. Laterial view of the head of an immature male B.M. 1967 . 2239. Note developing dark gular patch. Bull. Br. Mus. nat. Hist. (Zool.) 17, 3 PLATE 2 ZOOL. 17, 3. 5§ PLATE 3 a. Agama sankaranica Chabanaud. Dorsal view of an adult female B.M. 1967.2204. b. Agama sankaranica Chabanaud. Lateral view of the head of the same individual. c. Agama gracilimembris Chabanaud. Lateral view of the head of an adult male B.M. 1961 . 949. d. Agama gracilimembris Chabanaud. Dorsal view of same individual. Bull. Br. Mus. nat. Hist. (Zool.) 17, 3 PLATE 3 PLATE 4 Stereoscan electron microscope photographs of dorsal midbody scales. Arrows point to the sense organs. Upper row left to right. Agama agama B.M. 1962.1595 x 48 Agama par agama sp. n. Holotype X 28 Lower row left to right. Agama benueensis B.M. 1967.49 x 52 Agama sankaranica B.M. 1967.2204 x 47 Bull. Br. Mus. nat. Hist. (Zool.) 17, 3 PLATE 4 PLATE 5 Stereoscan electron microscope photographs of the apices of the same trunk scales as illustrated in Plate 4 showing the positions of the sense organs relative to the keel and dorsodistal margin and the single " hair " protruding from each receptor. Upper row left to right. Agama agama X 70 Agama paragama X 70 Lower row left to right. Agama benueensis X 131 Agama sankaranica X 260 Bull. BY. Mus. nat. Hist. (Zool.) 17, 3 PLATE 5 PLATE 6 Stereoscan electron microscope photographs of a dorsal midbody scale of Agama gracili- membris B.M. 1961 .949 showing the position of the single " hair-bearing " sense organ directly below the termination of the keel. Left X no. Right X 580. Bull. BY. Mus. nat. Hist. (Zool.) 17, 3 PLATE 6 PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING A REVISION OF THE AMPHIPO: GENUS MICRODEUTOPUS COSTA (GAMMARIDEA : AORIDAE) A. A. MYERS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 17 No. 4 LONDON: 1969 A REVISION OF THE AMPHIPOD GENUS MICRODEUTOPUS COSTA (GAMMARIDEA : AORIDAE) BY A. A. MYERS Department of Zoology University College of Swansea Pp. 91-148; i Plate, 22 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 17 No. 4 LONDON: 1969 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. 17, 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. nat. Hist. (ZooL). Trustees of the British Museum (Natural History) 1969 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 4 February, 1969 Price £i 55. A REVISION OF THE AMPHIPOD GENUS MICRODEUTOPUS COSTA (GAMMARIDEA : AORIDAE) A. A. MYERS SYNOPSIS The systematic position of the Aoridae and the phylogenetic relationships of its members are discussed. All the known species of Microdeutopus and one new species are described and figured, with keys for their identification. The synonymy of each species of Microdeutopus is reviewed, and details of their ecology and distribution are given. A catalogue of material studied in the present work, and the location of type material, is given in the appendices. INTRODUCTION THE need for a revision of a number of genera in the Aoridae has long been indicated. Enequist (1949) pointed out that there is uncertainty about the delimitation of the species of the genera Microdeutopus and Lembos. Andersson (1954) elucidated the relationships of Microdeutopus gryllotalpa Costa, M. anomalus (Rathke) and M. propinquus Sars, using material collected in the Gullmarfjord and collections in the Swedish State Museum. His findings have been substantiated in the present work, which included studies on laboratory-reared material. Generally, however, field workers have experienced difficulties in the identification of aorid material, as is clear from the numbers of wrongly identified specimens that exist in many institu- tions. The literature is so widely scattered, and the synonymy, particularly of the European species, so confused, that a thorough revision of the synonymy and review of the literature are needed. During the present investigations, fresh material was examined wherever possible together with preserved material from the British Museum and a number of other institutions and specimens from private donors (see Appendix i). Laboratory reared material of M. gryllotalpa and M. anomalus was also studied. Identification of Aoridae is frequently hindered by their fragility, specimens collec- ted in dredge samples often being devoid of all appendages except gnathopoda, pleopoda and uropoda. Fortunately male Microdeutopus species can readily be identified by the structure of the gnathopoda, uropoda and telson, and by the form of the processes which arise from the peraeon segments in the mid- ventral line. These ventral processes have been shown to occur in all species of Microdeutopus which have been examined for this character. The only species not examined is M. haswelli Stebbing, which at this stage seems better not dissected since it is repre- sented only by the single holotype specimen obtained on loan from the Australian Museum, Sydney. In general, the structure of the mouthparts is not a valuable taxonomic criterion, except at a generic level and above, though in some species, certain minor differences ZOOL. 17, 4 6 94 A. A. MYERS can be recognized as in the structure of the mandibular processes of the paragnath in M. schmitti Shoemaker and M . hancocki Myers. Kinne (ig63a, 1963^ 1964) has shown that temperature, salinity, food, and com- binations of these factors affect maximum size in some organisms. Personal observations in the laboratory (Myers, J_968d) indicate that some, if not all these factors, also affect growth rates and maximum size in M. gryllotalpa and presumably, therefore, in other Microdeutopus species. Field observations also suggest that the various species reach differing maximum sizes, in different localities, and it is therefore not possible to give an overall indication of the size at which the male of any Microdeutopus species reaches maturity and is hence identifiable. In particular it should be noted that high latitude specimens are generally much larger than low latitude specimens of the same species. In the systematic section the maximum recorded size, measured from the anterior margin of the cephalosome to the tip of the telson, is given for each species and each sex. Male Microdeutopus are readily identifiable, and descriptions of all known species are given below, together with keys for their identification. The identification of females presents considerable difficulty, however, since aorid genera are based almost entirely upon adult male characters, of forms which show extensive sexual dimor- phism. There are no characters exhibited by females which are diagnostic of the genus Microdeutopus. Many resemble the male in the structure of the antennules, antennae, uropoda and telson, thus exhibiting features which differ in one way or another from those of other Microdeutopus females, and probably from those of other female Aoridae. However, until the females of all the known Aoridae are studied, identification of solitary females within this family must remain questionable. Full descriptions and figures of ovigerous females, are given here, to aid provisional identification and also to prevent the confusion of females in samples containing more than one Microdeutopus species. SYSTEMATIC POSITION OF THE AORIDAE The Aoridae are closely related to the Photidae, from which they differ in having the first pair of gnathopoda more markedly developed than the second. Aorcho Barnard is intermediate in having the first and second gnathopoda of equal size and can be assigned to either family (Barnard, 1964). Also intimately related to the Aoridae are the Corophiidae, which differ from the Aoridae only in the degree of depression of the pleon. The loss of the accessory flagellum of the antennule has been described as an advanced character in Amphipoda (Barnard, 1962), and it is suggested here that the corophiid tendency towards the loss of a uropod 3 ramus is also advanced, so does not necessarily indicate close relationship between the posses- sors of this character. Barnard (1962) has pointed out that it has become increas- ingly difficult to assign, between the two families, various members of the Photidae and Corophiidae, because here too the differences involve quantitative aspects of depression or compression of the pleon. It seems probable that the family Coro- phiidae is polyphyletic and includes, within its framework, advanced members of both the Aoridae and Photidae. It is therefore not possible to indicate any circum- REVISION OF MICRODEUTOPUS 95 scribed diagnosis for the family Aoridae, which can only be described as those mem- bers of the aorid/photid/ischyrocerid/corophiid complex which exhibit greater development of gnathopod I than gnathopod 2, coupled with a tendency towards retention of such primitive characters as an accessory flagellum on the antennule, uropod 3 with two rami, contiguous coxae, and no marked depression of the pleon. The subdivision of the family Aoridae is also problematical since the complex permutations of characters among the species makes a systematic classification, based on possible phylogenetic relationships, exceedingly difficult. The accessory flagellum of the antennule exhibits a complete gradation of stages, from (a) primitively multi-articulate as in Aora typica Kroyer, Microdeutopus anomalus (Rathke), Lembopsis spinicarpus Pearse, Lembos viguieri Chevreux and others, to (b) reduced to two or three articles of which the terminal one is usually vestigial, e.g. in Micro- deutopus armatus Chevreux, Lemboides afer Stebbing, and Neomegamphopus roose- velti Shoemaker, or microscopic as in Aoroides columbiae Walker. The rami of uropod 3 may be (a) more or less equally developed as in Amphideutopus oculatus Barnard, Microdeutopus versiculatus (Bate), Lembos longipes Lilljeborg, and Aorcho delgadus Barnard; (b) the endopodite may be greatly reduced as in Acuminodeutopus heter- uropus Barnard ; (c) both rami may be minute, as in Paradryope orguion Stebbing and Dryopoides westwoodi Stebbing; or (d) there may be a single ramus only as in Neomicrodeutopus cabindae Schellenberg. The antennae exhibit considerable variation in form. They may be (a) shorter than the antennules stout, sub-pediform, with few flagellar articles, resembling those of the Corophiidae, as in Microdeutopus haswelli Stebbing; (b) less stout, as in M. gryllotalpa Costa, Aora typica Kroyer, Lembos websteri Bate and Lemboides afer Stebbing; (c) slender though still shorter than the antennules, as in Microdeutopus stationis Delia Valle, Lembopsis spinicarpus Pearse and Lembos macromanus (Shoemaker) ; or (d) very long and slender, exceeoling the length of the antennules, with multiarticulate flagella, as in Paradryope orguion Stebbing and Amphideutopus dolichocephalus Myers. The development of median spines on the ventral surface of the peraeon segments is of frequent occurrence in the Aoridae, but such spines are not restricted to the Aoridae, being also exhibited by various members of the related family Corophiidae, including certain species of Unciola Say and Grandidierella Coutiere. Most classifications of the Aoridae have been based upon the structure of the male gnathopod i, the gross development of which is so characteristic a feature of the Aoridae, though it is not diagnostic. Two main tendencies are discernible in this appendage: first, the great development of one or more of the articles, frequently accompanied by reduction of some of the others, and secondly, the production of toothlike processes on one or more of the articles. Classifications based on either of these characters are arbitrary, since it would be unreasonable to assume that either of these tendencies has occurred only once in the evolution of the family. Never- theless, a good workable classification is that of Barnard (1958, 1962, 1964) which is adopted here with two important modifications suggested by present work. Firstly Microdeutopus tridens Schellenberg and M. kraemmeri Reid are transferred to the genus Lembopsis Pearse (see Myers I968c); and secondly, the genus Coremapus Norman is incorporated in the genus Microdeutopus (see p. 102). 96 A. A. MYERS Genus MICRODEUTOPUS Costa 1853 Microdeutopus Costa, 1853 : 178. Autonoe Bruzelius, 1859 (pro-parte) : 23. Stimpsonia Bate, 1862 : 162. Microdeuteropus Norman, 1868 : 281. Stimpsonella Delia Valle, 1893 : 421. Coremapus Norman, 1905 : 78. TYPE SPECIES : Microdeutopus gryllotalpa Costa. DIAGNOSIS. Head lobes moderately produced, obtuse; antennules longer than the antennae (with the exception of male M. chelifer (Bate)) ; article i of mandibular palp the shortest, article 3 the longest, article 3 broad centrally, narrowing terminally; mandibular process of paragnath acute ; gnathopod i of male larger than gnathopod 2, complexly chelate, article 5 considerably larger than article 6 ; females with the first and second gnathopods dissimilar; uropod 3 with two more or less subequal rami; telson simple. HISTORICAL. The genus was established by Costa (1853) to include the single species M. gryllotalpa Costa, independently described by Bruzelius under the name Autonoe grandimana. Bate (1856) erected the genus Lembos to include the species L. cambriensis Bate, L. damnoniensis Bate, L. versiculatus Bate and L. websterii Bate, but later (1862) relegated all these species to the genus Microdeutopus Costa (corrected from Microdentopus in the text, to Microdeutopus in the appendix). In the same year he erected the genus Stimpsonia, to include as monotype chelifera Bate, while admitting that " the genus probably bears too close a resemblance to Microdeutopus to be retained as generically distinct ". Delia Valle (1893) noted the preoccupation of Stimpsonia among the Nemertina (Stimpsonia Girard, 1853) and erected Stimpsonella, including in the genus, armatus Chevreux with chelifera Bate. Norman (1868) altered Microdeutopus to Microdeuteropus for grammatical reasons but later (Norman, igo5a) reverted to the original spelling. Norman (i905b) erected the monotypic genus Coremapus for versiculatus Bate. LITERATURE. The best general account is that of Delia Valle (1893) who gives ten species. Of these, however, M. gryllotalpa Costa and M. minax Smith are synonymous, M. tennis Dana is not a species of Microdeutopus and M. titii Heller is a dubious species, while the distinct species M . versiculatus is treated as a synonym of M. anomalus Rathke. The keys, which apply to males only, are adequate. Stebbing (1906) gives a good account of eight species, and not eleven as it would seem, for M. anomalus (Rathke) and M. propinquus Sars are synonymous, M. megnae Giles is attributable to Grandidierella Coutiere, and the doubtful M. titii Heller is listed with reservation. Descriptions of the females are inadequate, as in most literature, but the keys which apply to males only are adequate. Chevreux and Fage (1925) give seven species and their keys are modified after Stebbing (1906). Barnard (1958) gives a catalogue of species, and other useful taxonomic works are those of Haswell (1882) describing i sp. from Australia, Sars (1894) 2 spp. from Norway, Stephenson (i92ga) 2 spp. North Sea and Baltic, Shoemaker (1938) i sp. Mexico, Schellenberg (1942) 2 spp. Germany, Gurjanova (1951) 3 spp. U.S.S.R. and Myers (i968b) 3 spp. Central America. REVISION OF MICRODEUTOPUS 97 KEY TO THE MALE MICRODEUTOPUS OF THE WORLD N.B. in most figures of limbs, article i is omitted so article 2 is the most proximal shown. 1 a. Gnathopod 2 chelate ........... 2 b. Gnathopod 2 not chelate .......... 3 2 a. Gnathopod 2 with articles 5 and 6 subequal in length and breadth, article 2 expanded, crenulated on anterior margin (Text-fig. i8d) . M. armatus (p. 129) b. Gnathopod 2 with article 6 much shorter and broader than article 5, article 2 unexpanded, anterior margin smooth, (Text-fig. IQC) . . M. chelifer (p. 132) 3 a. Gnathopod i with the anterior margin of article 5 produced into a single tooth. Accessory tooth if present, arising within the posterior margin (Text-fig. 8b) . 4 b. Gnathopod i with article 5*8 anterior margin produced into 2 or more teeth (Text- fig. 2a) . 7 4 a. Gnathopod 2 densely setose, the setae finely pectinate; article 4 extending over greater length of article 5 (Text-fig. 31) . . . M. versiculatus (p. 101) b. Gnathopod 2 moderately setose, the setae not pectinate; article 4 relatively short (Text-fig, gc) (" anomalus " group) 5 5 a. Gnathopod 2 with articles 5 and 6 short and broad, palmar angle of article 6 almost transverse (Text-fig, ga) ; uropod 3 with rami relatively short and broad M. damnoniensis (p. 114) b. Gnathopod 2 with articles 5 and 6 slender, elongate, palmar angle of article 6 oblique (Text-fig, gc) ; uropod 3 with rami relatively elongate and slender (Text-fig, ge) 6 6 a. Gnathopod i with the anterior distal margin of article 5 produced into a slender, straight or inwardly curved tooth, generally with an accessory tooth arising near its base within the posterior margin (Text-fig. 8b) . M. anomalus (p. no) b. Gnathopod i with article 5*3 anterior margin produced into a broad based stout tooth, never with accessory tooth (Text-fig. 8h) . . . M. algicola (p. 117) 7 a. Gnathopod i with article 5 having the proximal (outer) of two teeth longer than the other (Text-figs. 2e, 22a) ......... 8 b. Gnathopod i 's article 5 not as above ........ 9 8 a. Antenna stout, subpediform, with the first flagellar article grossly developed (Text-fig. 4b); gnathopod 2 with article 2 elongate and slender (Text-fig. 3h) M. haswelli (p. 107) b. Antenna relatively slender, the first flagellar article not as above (Text-fig. 21), gnathopod 2 with article 2 broad and short (Text-fig. 22g) . M. sporadhi (p. 136). 9 a. Gnathopod i with the distal tooth on article 5 the longest (Text-fig. 2b) . . 10 b. Gnathopod i with the central tooth on article 5 the longest (Text-fig. 2d) M. stationis (p. 104) 10 a. Gnathopod 2 with article 2 convex anteriorly, crenated on the anterior margin (Text-fig. 3g) M. gryllotalpa (p. 98) b. Gnathopod 2 with article 2 concave anteriorly and its anterior margin smooth (Text-fig. 14!) (" schmitti " group) n n a. Gnathopod i having article 5 with a tooth on the anterior margin (Text-fig. I3a) 12 b. Gnathopod i having article 5 without a tooth on the anterior margin (Text-fig. I3b) M. schmitti (p. 120) 12 a. Gnathopod i having article 6 with the palmar margin produced into a forward directed tooth opposable to article 7; posterior margins of articles 4, 5 and 6 with very long setae (Text-fig. 136) ..... M. trichopus (p. 124) b. Gnathopod i having article 6 with the palmar margin at most produced into a small lobe, which is never directed forward; posterior margin of article 4 only having very long setae, those of articles 5 and 6 being relatively short (Text-fig. i3a) .......... M. hancocki (p. 124) 98 A. A. MYERS Microdeutopus gryllotalpa Costa (Text-figs. la, 2a-c, 3a-c, g, m, 5b, 6c-d, 2oa) Microdeutopus gryllotalpa Costa, 1853 : 178; Costa, 1857 : 231, pi. 4, fig. 10; Boeck, 1870 : 156; Boeck, 1876 : 565, pi. 29, fig. 6; Blanc, 1884 : 75, pi. 4, figs. 82-90; Hoek, 1889 : 226; Delia Valle 1893 : 411, pi. i, fig. 12, pi. n, figs. 25-43; Sars 1894 : 543. P^ I92> ng- 2> Sowinsky !895 : 237, pi. 4, fig. 6; Sowinsky, 1898 : 480; Stebbing, 1906 : 590; Norman, 1907 : 368, pi. 16, fig. 3, pi. 17, figs. 6-7; Chevreux & Fage, 1925 : 299, fig. 310; Stephenson, 1927: 124; Stephensen. igzqa : 151, fig. 273; Oldevig, 1933 : 213, fig. 95; Cecchini & Parenzan, 1934 : 213, fig. 43; Loven, 1934: 6—8, fig. 2; Miloslavskaia, I939a : 121; Schellenberg, 1942 : 186, fig. 154; Soika, 1949 : 199; Gurjanova, 1951 : 831, fig. 580; Barnard, 1958 : 29. [Nori] Microdeutopus gryllotalpa Bate, 1862 : 163, pi. 30, fig. i. [Nori] Microdeutopus gryllotalpa Nebeski, 1880 : 45, fig. 41. [Nori] Microdeutopus grillotalpa [sic] Sowinsky, 1880 : 125, pi. 5, fig. 17 a-d. Autonoe grandimana Bruzelius, 1859 : 26, pi. i, fig. 5. Microdeutopus grandimanus (Bruzelius) Bate, 1862 : 378. Microdeutopus minax Smith, 1874 : 562. Microdeutopus bidens Sowinsky, 1880 : 129, pi. 5, fig. 18. [?]Amphithoe salenskiiCarus, 1885 : 396. Microdeuteropus gryllotalpa (Costa) Norman, 1886 : 16; Chevreux & Bouvier, 1893 : 133. Microdeuteropus minax (Smith) Norman, 1886 : 17. TYPE LOCALITY. Lago di Fusaro, Italy. DIAGNOSIS OF MALE. Antennule about half the body length, the first peduncular article slightly shorter than the second, the third about one third the length of the second ; flagellum longer than the peduncle normally with up to twenty- two articles (maximum recorded twenty-six) ; accessory flagellum with two articles, the second article rudimentary. Antenna slightly under two thirds the length of the antennule, the fourth and fifth peduncular articles subequal; flagellum shorter than the fifth peduncular article, normally with up to nine articles (maximum recorded eleven). Peraeon segments 2-7 each with a process in the mid-ventral line, those of segments 2 and 3 short, straight, those of segments 4-7 short, robust, posteriorly directed; processes 6 and 7 with short setae arising from their posterior margins. Gnathopod 1 with article i moderately produced anteriorly, acute ; article 2 short and expanded distally; article 5 immensely expanded, in some specimens as broad as long, the posterior distal margin produced into two to four teeth, of which the most distal is always the longest ; article 6 short, the posterior margin irregularly lobed ; article 7 sometimes short and robust, sometimes relatively long and slender. Gnathopod 2 with article 2 greatly expanded, the anterior margin convex and crenated; articles 5 and 6 subequal in length, 5 slightly the broader, articles 4, 5 and 6 with long pectin- ate setae on the anterior margins. Uropod 3 with rami equal in length to the peduncle exopodite slightly the longer of the two rami; each ramus with a transverse row of three closely associated spines on the outer dorsal margin, together with a number of paired and solitary spines of variable distribution, terminal setae relatively long. Telson with each terminal crest bearing on its distal dorsal margin, a group of up to six setae. Maximum size of male : 8 mm. REVISION OF MICRODEUTOPUS 99 FEMALE. As the male except for the absence of ventral peraeon processes and the structure of the sexually dimorphic gnathopoda. Gnathopod i with article 2 moderately expanded; articles 5 and 6 subequal in length, 5 slightly the broader, 6 with the anterior and posterior margins roughly parallel; article 7 equal in length to the angular palmar region. Gnathopod 2 with FIG. i. Microdeutopus gryllotalpa Costa, (a) Male head, Swansea, Wales. Microdeutopus versiculatus (Bate), (b) Male head, Falmouth Harbour, England. ioo A. A. MYERS article 2 slightly expanded, crenulated on the anterior margin; articles 5 and 6 elongate, subequal; articles 4, 5 and 6 with pectinate setae on their anterior margins. Maximum size of female: 10 mm. DISCUSSION. M. gryllotalpa is closely related to M. versiculatus but differs in both sexes by the second gnathopod (Text-figs. 3g, 5b) not having the fourth article greatly produced along the posterior border of the fifth, nor the fifth and sixth FIG. 2. Microdeutopus gryllotalpa Costa, (a) Male gnathopod i, Swansea, Wales, (b) Male gnathopod i, Swansea, Wales, (c) Male gnathopod i, Isefjord, Denmark. Microdeuto- pus stationis Delia Valle. (d) Male gnathopod i, St. Peter's Port, Guernsey, (f) Male gnathopod i, Naples, Italy. Microdeutopus haswelli Stebbing. (e) Male gnathopod i, Port Jackson, Australia (Holotype). REVISION OF MICRODEUTOPUS 101 articles so markedly elongate and slender. Females of M. gryllotalpa are readily identifiable by the structure of the fifth article of the second gnathopod (Text-fig. 5b) , which has long pectinate setae arising from the entire length of the anterior margin, whereas in all other Microdeutopus species (except M. versiculatus) the long setae arise from the anterior corner only of the fifth article. ECOLOGY. In the intertidal zone in rock pools amongst Chaetomorpha and other algae; amongst Zoster a spp.; and in salt marshes (Chevreux & Page, 1925), often associated with Ruppia maritima as at Roscoff (Truchot, 1963). Also occurring sub-littorally to a depth of about 150 metres on oyster beds, and amongst algae, tunicates, sponges, polyzoans, Mytilus etc., always in areas of high detritus accumu- lation. It particularly favours docks and other man-made installations, and is frequently associated with Corophium acherusicum Costa in both the Palearctic and Nearctic regions. DISTRIBUTION. From the Lofoten Islands southwards along the coasts of Norway, Sweden, Denmark, including the Baltic, and Holland, around the coasts of the British Isles, chiefly in the south-west, along the Atlantic coasts of France, Spain and Portugal, and in the Mediterranean Ligurian, Tyrrhenean, Adriatic, Aegean and Black Seas. Also, on the north-eastern seaboard of the United States of America (Massachusetts, Rhode Island and Connecticut) . Microdeutopus versiculatus (Bate) (Text-figs, ib, 3i, k, 5a, 6a-b, 2oc) Lembos versiculatus Bate, 1856 : 58; Bate, 1857 : 142. Microdeutopus versiculatus Bate, 1862 : 165, pi. 30, fig. 5; Bate & Westwood 1863 : 295; Walker, 1895 : 469; Chevreux 1900 : 89; Stebbing, 1906 : 593. Microdeuteropus versiculatus (Bate) Norman, i86ga : 282; Stebbing, 1874 : 12, pi. i, figs. 2, 2a-f. Chevreux & Bouvier, 1893 : 134. Autonoe longipes (Lilljeborg) Boeck, 1876 (pro-parte) : 574. Autonoe versiculata (Bate) Norman, 1886 : 17. Microdeutopus anomalus (Rathke) Delia Valle, 1893 (pro-parte) : 418. Coremapus versiculatus (Bate) Norman, igo5b : 78; Chevreux & Page, 1925 : 301, fig. 312; Stephensen, i92ga : 152, fig. 276; Cecchini & Parenzan, 1934 : 217, fig. 45; Miloslavskaia, I939a : 125, fig. 25; Soika, 1949 : 200; Gurjanova, 1951 : 835, fig. 585; Barnard, 1958 : 29. TYPE LOCALITY. Plymouth, Devon, England. DIAGNOSIS OF MALE. Antennule about equal to the body length, the first pedun- cular article slightly shorter than the second, the third about one third the length of the second; flagellum almost twice the length of the peduncle, normally with up to twenty articles (maximum recorded twenty-two); accessory flagellum with two articles, the second article rudimentary. Antenna little more than half the length of the antennule, the fourth peduncular article slightly longer than the fifth ; flagel- lum shorter than the fifth peduncular article, normally with up to eight articles (maximum recorded nine). Per aeon segments 2-7 each with a process in the mid- ventral line; that of segment 2 slender, spiniform, directed anteriorly; those of segments 3 and 4 slender, spiniform, straight; those of 5-7 robust, decreasing in size antero-posteriorly, with processes 6 and 7 having setae arising from their posterior loa A. A. MYERS margins. Gnathopod i with article I moderately produced anteriorly, acute; article 2 considerably expanded; article 5 much enlarged, longer than broad, the posterior distal angle produced into a single, robust tooth; article 6 short, the post- erior margin irregularly lobed; article 7 short and robust. Gnathopod 2 with article 2 somewhat expanded ; article 4 well developed, extending over the greater length of the posterior margin of article 5 ; article 5 longer than article 6, elongate and slender, about three times as long as broad; article 6 also very long and slender, about three and a half times as long as broad; article 7 very short, long pectinate setae arising from the anterior margins of articles 4, 5 and 6. Uropod 3 with rami sub-equal, equal in length to the peduncle ; exopodite with a transverse row of three closely associated spines on the outer-dorsal margin, basal to which is a solitary spine ; endopodite with a similar transverse row of closely associated spines on the outer-dorsal margin, and in addition on the inner margin, a series of well separated spines; terminal setae of both rami relatively long. Telson with each terminal crest bearing on its distal dorsal margin a group of two to three setae. Maximum size of male: 7 mm. FEMALE. As the male except for the structure of the sexually dimorphic gnatho- poda and absence of ventral peraeon processes. Gnathopod i with article 2 short and considerably expanded, about two thirds as broad as long; article 5 scarcely longer than article 6, with a dense clothing of comb setae along the entire length of the posterior border; article 6 almost as broad as 5, with the anterior and posterior margins roughly parallel; article 7 moderately long, equal in length to the palmar region. Gnathopod 2 with article 2 elongated and slender, otherwise identical with the corresponding appendage of the male. Maximum size of female : 8 mm. DISCUSSION. First designated as a nomen nudum by Bate (1856) the species was later described from the female (Bate, 1857, 1862). The male was first described by Norman (1868) and later by Stebbing (1874) who also gave figures. The sugges- tion that a monotypic genus should be erected for this species (Norman, i905b) is not accepted. The present species bears a close relationship with M. gryllotalpa Costa in the similarity of the male first gnathopod of the two species. That of M. versicu- latus (Text-fig. 3k) differs in having a single tooth on the posterior distal angle of article 5, whereas in M. gryllotalpa (Text-figs. 2a-c) there are two to four. The two species are also similar in the structure of the male second gnathopod. Article 2 of that appendage is expanded, though less markedly in M. versiculatus (Text-fig. 3i) than in M. gryllotalpa (Text-fig. 3g) and article 5 has pectinate setae arising from the entire anterior margin (also a character of the female). Finally, males of both species have ventral processes on all peraeon segments except the first. The remarkable slender, setose second gnathopod of M. versiculatus (Text-fig. 5a), which led Normon (iQ05b) to erect a monotypic genus for the species, is an extreme modification of the type found in M. gryllotalpa and serves to distinguish M. versiculatus in both sexes from all other Microdeutopus species. ECOLOGY. In the intertidal zone (Oldany, Sutherland) , but much more frequently in the sub-littoral to a depth of about 150 metres (off the Shetland Islands) amongst REVISION OF MICRODEUTOPUS 103 Fig. 3. Microdeutopus gryllotalpa Costa, (a) Male gnathopod i, 1-8 mm. <$, Swansea, Wales, (b) Male gnathopod i, 2-9 mm. <$, Swansea, Wales, (c) Male gnathopod i, 3-9 mm. cJ, Swansea, Wales, (g) Male gnathopod 2, Swansea, Wales, (m) Male telson, Swansea, Wales. Microdeutopus stationis Delia Valle. (d) Male gnathopod i, 1-9 mm. fig- 44>' Miloslavskaia, i939a : 122; Gurjanova, 1951 : 832, fig. 581 ; Barnard, 1958 : 29. [Wow] Microdeutopus stationis Sowinsky, 1895 : 237, pi. 4, figs. 1-5; Sowinsky, 1898 : 480. TYPE OF LOCALITY. Naples, Italy. DIAGNOSIS OF MALE. Antennule about equal to the body length, the first pedun- cular article slightly shorter than the second, the third less than one third the length of the second; flagellum twice the length of the peduncle, normally with up to twenty-six articles (maximum recorded twenty-nine) ; accessory flagellum with two to five articles, the terminal article rudimentary. Antenna two thirds the length of the antennule, the fourth and fifth peduncular articles subequal; flagellum equal in length to the fifth peduncular article, normally with up to twelve articles (maximum recorded fourteen). Per aeon segments 2-4 each with a process in the mid-ventral line, that of segment 2 slender spiniform, straight, those of segments 3 and 4 becoming progressively shorter and more robust. Gnathopod ~L with article I strongly produced anteriorly, very attenuated; article 2 scarcely expanded; article 5 very enlarged, longer than broad, the posterior distal angle produced into three teeth of which the central is the longest, the inner two slightly recurved; article 6 short, the posterior margin convex distally, concave basally; article 7 long and slender. Gnathopod 2 with article 2 not expanded; article 5 longer and slightly broader distally than article 6. Uropod 3 with the rami slightly shorter than the peduncle, the expedite slightly the longer of the two rami, bearing on its outer distal margin a group of three closely associated elongate spines, with a pair of spines (of which one is elongate) basal to these and not infrequently a further solitary spine basal to the pair; inner margin of exopodite with one or two solitary spines; endopodite outer margin with a pair of spines, basal to which is a solitary spine; inner margin with up to two pairs of spines and a solitary spine; terminal spines and setae of both rami very short. Telson with each terminal crest bearing on its distal dorsal margin, a group of three short stout spines and a single fine seta. Maximum size of male : 10 mm. FEMALE. As the male, except for the structure of the sexually dimorphic gnatho- poda, and absence of ventral peraeon processes. Gnathopod i with article 2 scarcely REVISION OF MICRODEUTOPUS 105 FIG. 4. Microdeutopus statlonis Delia Valle. (a) Male head, St. Peter's Port, Guernsey. Microdeutopus haswelli Stebbing. (b) Male head, Port Jackson, Australia (Holotype). io6 A. A. MYERS expanded, articles 5 and 6 subequal in length, the sixth distinctly broader distally, narrowing at the junction with the fifth article, the palmar angle smoothly rounded; article 7 elongate and slender. Gnathopod 2 with article 2 relatively slender, article 5 slightly shorter than article 6, setae not pectinate. Maximum size of female : 12 mm. DISCUSSION. The species was first described by Nebeski (1880) from the Adriatic, and erroneously ascribed to M. gryllotalpa Costa. That author's illustrations (Nebeski, 1880, fig. 41) of the growth stages of the male gnathopod i clearly indicate FIG. 5. Microdeutopus versiculatus (Bate), (a) Female gnathopod 2, Plymouth, Devon. Microdeutopus gryllotalpa Costa, (b) Female gnathopod 2, Swansea, Wales. Micro- deutopos stationis Delia Valle. (c) Female gnathopod 2, Naples, Italy. that he was dealing with the species later described by Delia Valle under the name M. stationis. M. stationis is easily distinguishable in the male from all other described species in the genus by having the posterior distal angle of article 5 of the first gnathopod (Text-figs. 2d, f) produced into three teeth, of which the central is the longest. Females have few diagnostic characters, but important characters taken in combina- tion are the telson (Text-fig. 31) bearing both spines and setae, uropod 3 (Text-fig. 6f) with groups of spines on the rami, accessory flagellum with two to four articles and long slender antenna with multi-articulate flagellum. ECOLOGY. Recorded in Sacchoriza holdfasts, among Posidonia, on oyster beds and over fine sands, in regions of relatively low detritus contamination, in the sub- littoral to a depth of 50 metres. REVISION OF MICRODEUTOPUS 107 DISTRIBUTION. Guernsey (St. Peter's Port), France (Arcachon, Roscoff, Perros Guirec), Corsica (Ajaccio, Bonifacio), Algeria (Bone, La Calle), Tunisia (Golfe de Gabes), Italy (Naples, Trieste), Greece (Khios). Microdeutopus haswelli Stebbing (Text-figs. 2e, 3h, 4b) Microdeuteropus chelifer Hasell, 1879 : 340, pi. 22, fig. 3; Delia Valle, 1893 : 421. Microdeutopus chelifer Haswell, 1882 : 265. Microdeutopus haswelli Stebbing, 1899 : 350; Stebbing, 1906 : 591; Stebbing, 1910 : 647; Sheard, 1937 : 2^; Barnard, 1958 : 29. TYPE LOCALITY. Clark Island, Port Jackson, Australia. The present species is represented by a single male specimen in Haswell's collec- tions, at the Australian Museum, Sydney. Haswell did not designate types for any of his material, but this specimen is almost certainly the holotype, agreeing in the main with Haswell's description, and is here considered as such. An expanded description of the more relevant features of the holotype is given. The range of variation of this species is not known, as further material has not been forthcoming. DESCRIPTION OF MALE HOLOTYPE. Head lateral lobes slightly produced, obtuse; eyes round. Antennule slightly less than half the body length, the first and second peduncular articles subequal, the third slightly over a third the length of the second ; flagellum longer than the peduncle, with seventeen articles; accessory flagellum with five articles, the fifth article rudimentary. Antenna shorter than the antennule, sub-pedif orm, the fifth peduncular article slightly longer than the fourth ; flagellum with four articles, the first article long and robust, over half the length of the fifth peduncular article, the second short, the third and fourth articles progressively shorter, obscurely articulate. Gnathopod I with article I moderately produced anteriorly, obtuse; article 2 short and expanded, the anterior margin excavate; article 5 much enlarged, longer than broad, the posterior distal angle produced into two teeth, of which the outer is the longer and recurved; article 6 short, the posterior margin concave centrally; article 7 very stout. Gnathopod 2 with article 2 very elongated and slender, concave anteriorly, articles 5 and 6 slender with article 5 slightly the longer. Epimeral plate 3 with the posterior free corner produced into a tooth, above which is inserted a short seta. Uropod 3 with the peduncle exceeding the length of the very short and stout rami, of which the exopodite is very slightly the longer; each ramus with a terminal group of setae which exceed the length of the ramus. Telson with the terminal crests moderately well developed, each bearing on its distal, dorsal margin, a group of three setae, of which the central is the longest. Length : 4-5 mm. DISCUSSION. The very robust, sub-pediform antenna, (Text-fig. 4b) with the flagellum very reduced and the first flagellar article robust, approaches the condition found in many Corophiidae. However, the lateral compression of the urosome, biramous third uropoda, and moderately well developed coxae, are characteristic of aorid/photid stock. M. haswelli differs from all other Microdeutopus species, ZOOL. 17, 4 7 io8 A. A. MYERS except M. sporadhi sp. nov. (p. 136), in having the posterior distal angle of article 5 of the male first gnathopod (Text-fig. 2e) produced into two teeth, of which the outer is the longer. The elongated, slender article 2 of the male second gnathopod (Text- fig. 3h) is also a distinctive feature. FIG. 6. Microdeutopus versiculatus (Bate), (a) Female gnathopod i, Plymouth, Devon, (b) Male uropod 3, Falmouth, Cornwall. Microdeutopus gryllotalpa Costa, (c) Female gnathopod i, Swansea, Wales, (d) Male uropod 3, Arcachon, France. Microdeutopus stationis Delia Valle. (e) Female gnathopod i, Naples, Italy, (f) Male uropod 3, Naples, Italy. REVISION OF MICRODEUTOPUS 109 ECOLOGY. Unknown. DISTRIBUTION. Not yet recorded from other than the type locality. THE " ANOMALUS " GROUP OF SPECIES The systematics of the anomalus group of species has long been under dispute. In 1856 Bate erected two species, Lembos damnoniensis and L. cambriensis, but gave no descriptions and only in the following year attributed to the former species " first hand with a thumb on carpus " to the latter " first hand without thumb ". White (1857), no doubt in error, transposed the " mn " in L. damnoniensis, to " nm " in L. danmoniensis , and later Bate (1862) and Bate and Westwood (1863) relegated L. cambriensis to a synonym of M. anomalus (Rathke) and L. damnoniensis to a synonym of M. gryllotalpa Costa. Norman (1868) considered that M. gryllo- talpa Bate represented a young male M. anomalus, but later (Norman iQOSb) decided that M. damnoniensis Bate, was a distinct species, and that M. gryllotalpa Bate (non M. gryllotalpa Costa) represented that species. Sars (1894) figures a species under the name M. propinquus Sars, but in the text relegates it to a synonym of M . danmoniensis (Bate) (altered presumably in error, or following White, 1857, from M. damnoniensis}. St ebbing (1906) considered that M. propinquus Sars was not synonymous with M. damnoniensis (Bate) and reinstated it as a distinct species with synonym M. danmoniensis Sars (non M. damnoniensis (Bate)). Andersson (1954) considered M. propinquus Sars to be a juvenile form of M. anomalus (Rathke) and in the synonymy also listed L. damnoniensis Bate. Delia Valle (1893) described a new species M. algicola, and placed M. damnoniensis in the synonymy of M. anomalus (Rathke). Present investigations suggest that there are three distinct species in the "anomalus" group; M. anomalus (Rathke), M. damnoniensis (Bate) and M. algicola Delia Valle. Males of the " anomalus " group of species are distinguished from those of all other Microdeutopus species, except M. versiculatus (Bate) M. chelifer (Bate) and M. armatus Chevreux by having the posterior margin of gnathopod I produced into a single marginal tooth. In M. anomalus an accessory tooth is usually present, but this always arises within the posterior margin. They differ from the above three species in the structure of gnathopod 2, which is neither chelate as in M. chelifer and M. armatus not densely setose as in M . versiculatus. Females of the " anomalus " group of species are notable for their lack of diagnostic features, and are therefore identifiable only by negative characters. They are distinguishable from M. versiculatus, M. gryllotalpa and M. stationis by the absence of paired spines from the rami of the third uropod, from the first two species by the absence of pectinate setae on gnathopod 2. They differ also from M. stationis and from the " schmitti " group of species by the lack of any spines as distinct from setae on the telson terminal crests, and from M. armatus by the lack of any pronounced projection at the antero- distal corner of article 2 or gnathopod 2. Within the " anomalus " group of species the identification of females is even more difficult, particularly in the absence of males. In practice, when two or more of the " anomalus " group occur together no A. A. MYERS in a sample, the females have been separated with some degree of accuracy and grouped together with the corresponding males, according to characters summarized in Table i. Owing to confusion between the three species, records are unreliable and so the ecology and distribution of the " anomalus " group of species is poorly known. Character 1. Maximum size 2. Antennule a. Ratio of peduncular articles b. Average number of flagellar articles c. Number of articles to accessory flagellum excluding vestigial terminal article 3. Uropod 3 a. Peduncle b. rami M. anomalus 10 mm. 23 3-4 Flanges poorly developed, dorsal and inner longest slender, elongate M. damnoniensis 4 • 5 mm. 14 Flanges well developed inner much the longest broad, robust M. algicola 4-5 mm. i* 3 as damnoniensis as anomalus TABLE i. The main identification characters for females of the "anomalus" group of species. * = From Delia Valle (1893). Microdeutopus anomalus (Rathke) (Text-figs. 7a, 8a-e, QC, e, loc, f, 2ob, d) Gammarus anomalus Rathke, 1843 : 63, pi. 4, fig. 7; Lilljeborg, 1855 : 457. Autonoe anomala (Rathke) Bruzelius, 1859 : 25, fig. 4. Autonoe sp. Smith, 1874 : 562. Microdeutopus anomalus (Rathke) Boeck, 1870 : 157; Catta, 1875 : 167; Boeck, 1876 : 567, pi. 25, fig. 5; Delia Valle, 1893 : 417, pi. 56, fig. 41 ; Sars, 1894 : 540, pi. 191 ; Sowinsky, 1898 : 480, pi. 10, figs. 20-24; Stebbing, 1906 : 591; Kunkel, 1910 : 76, fig. 29; Chevreux & Fage, 1925 : 298, fig. 309; Stephensen, 1927 : 124; Stephensen, i92ga : 152, fig. 275; Oldevig, 1933 : 212, fig. 94; Miloslavskaia, i939a : 123; Carausu & Carausu, 1942 : 74, fig. 5; Schellen- berg, 1942 : 188, fig. 155; Soika, 1949 : 198; Gurjanova, 1951 : 833, fig. 582; Andersson, 1954 : 252, figs. 1-4; Barnard, 1958 : 29. [Nori] Microdeutopus anomalus Bate, 1862 : 164, pi. 30, fig. 3. Microdeuteropus anomalus (Rathke) Norman, i86ga : 281; Norman, 1886 : 16. Microdeutopus propinquus Sars, 1894 : 542, pi. 192; Stebbing, 1906 : 592; Stephensen, 1927 : 125; Stephensen, i92ga : 152, fig. 274; Oedevig, 1933 : 212, fig. 95; Schellenberg, 1942 : 189, fig. 156; Dahl, 1946 : 5; Gurjanova, 1951 : 834, fig. 583. Microdeutopus danmoniensis [sic] Sars, 1894 : 542, pi. 192. Microdeutopus stationis Sowinsky, 1895 : 237, pi. 4, figs. 1-6. TYPE LOCALITY. Christiansund, Norway. REVISION OF MICRODEUTOPUS in DIAGNOSIS OF MALE. Antennule about two thirds the body length, the first peduncular article shorter than the second, the third about one third the length of the second; flagellum slightly less than twice the length of the peduncle, normally with up to twenty-three articles (maximum recorded twenty-seven) ; accessory flagellum with four to five articles, the terminal article of which is rudimentary. Antenna two thirds the length of the antennule, the fourth and fifth peduncular articles subequal; flagellum about equal to the fifth peduncular article, normally with up to six articles (maximum recorded eight) . Per aeon segments 2-4 each with a spine in the mid-ventral line, those of segments 2 and 3 the longest and most slender, that of segment 4 the shortest and most robust ; segment 5 in some specimens with a vestigial spine. Gnathopod i with article i strongly produced anteriorly, very attenuated; article 2 elongate and relatively slender; article 5 very enlarged, longer than broad, the posterior distal angle produced into a straight or slightly recurved tooth, at the base of which in mature specimens is characteristically an accessory tooth, which arises within the posterior margin and is not an extension of it as in other Microdeutopus species ; article 6 with the posterior margin concave, excepting the palmar angle; article 7 variable, usually relatively long and slender. The accessory tooth on article 5 varies considerably in size, sometimes being so redu- ced that it does not project over the posterior margin and appears obsolete; occasion- ally in adults and frequently in juveniles it is absent. The aberrant form of this species described and figured by Chevreux and Fage (1925) with two accessory teeth, has not been observed in the present investigations. Gnathopod 2 with article 2 unexpanded; articles 5 and 6 elongate, 5 slightly the broader; article 6 almost three times as long as broad, with the palmar angle oblique ; article 7 relatively long. Uropod 3 with the peduncle slender, its outer, inner and dorsal margins each produced into a flange of which the dorsal and inner are the most pronounced ; rami equal to or slightly exceeding the length of the peduncle ; endopodite slightly shorter than the exopodite, with four spines on the inner margin and two on the outer, none of which are closely associated; exopodite with two spines on the outer margin only. Telson, with each terminal crest bearing on its distal dorsal margin, a pair of unequal setae. Maximum size of male : 8 mm. FEMALE, As the male, except for the structure of the sexually dimorphic gnatho- poda, and absence of ventral peraeon spines. Gnathopod i with article i somewhat produced anteriorly, rounded; article 2 unexpanded; article 6 longer than article 5, distinctly broader distally, and constricted at the junction with article 5 ; article 7 longer than the palmar region. Gnathopod 2 with article 2 elongate and slender; article 6 considerably longer and more slender than article 5. Maximum size of female : 10 mm. DISCUSSION. M. anomalus is distinguishable in the male, in its typical form, from all other Microdeutopus spp. by the presence of an accessory tooth, arising within the posterior margin of article 5 of gnathopod i (Text-figs. 8a, b, d, e). Past workers have described M . anomalus as having this accessory tooth (or teeth) always present, although this is by no means the case, particularly in immature material, and have frequently utilized this character as a basis for a key (Stebbing, 1906, Chevreux & A. A. MYERS FIG. 7. Microdeutopus anomalus (Rathke). (a) Male head, Plymouth, Devon. Micro- deutopus damnoniensis (Bate), (b) Male head, Plymouth, Devon. REVISION OF MICRODEUTOPUS FIG. 8. Microdeutopus anomalus (Rathke). (a) Male gnathopod i, Falmouth, Cornwall, (b) Male gnathopod i, Bonifacio, Corsica, (c) Male gnathopod i, Falmouth, Cornwall, (d) Male gnathopod i, Isle of Mull, Scotland. (e) Male gnathopod i, Isle of Mull, Scot- land. Microdeutopus damnoniensis (Bate), (f) Male gnathopod i, Plymouth, Devon. Microdeutopus algicola Delia Valle. (g) Male gnathopod i, Naples, Italy, (h) Male gnathopod i, Bone, Algeria. H4 A. A. MYERS Fage, 1925) . This has often led to the attributing of immature material of the present species to M. damnoniensis (Bate), the separation of which must be made with great care. In its immature form, or in mature specimens lacking an accessory tooth, M. anomalus is distinguishable from M. algicola Delia Valle by the form of the primary tooth which arises at the posterior distal angle of article 5 and is relatively slender, though never as slender as in M. damnoniensis. In M. algicola (Text-figs, 8g-h) a considerable area of the posterior margin of article 5 contributes to the development of the tooth, which is hence very robust. M . anomalus differs from M . damnoniensis also in the form of the tooth on article 5 of gnathopod I, but is more easily distinguished from that species by the structure of the second gnathopod, which is slender and elongate with an oblique palm in M. anomalus (Text-fig. QC), relatively short and broad, with an almost transverse palm in M. damnoniensis (Text-fig, ga). In addition, the rami of uropod 3 are long and slender in M. anomalus (Text-fig. 96) ,whereas they are short and broad in M. damnoniensis (Text-fig. 9g). ECOLOGY. A sub-littoral species in depths ranging down to 200 metres, but occasionally extending into the intertidal zone. In areas influenced by detritus, often among arborescent weeds, but also in Zostera beds, and among shells, poly- zoans, sponges, tunicates and Mytilus. In the Black Sea, associated with Modiola phaseolina and Amphiuraflorifera. DISTRIBUTION. Norway south of the Lofoten Islands, Sweden, Denmark, including the Baltic, the British Isles, particularly the South and West coasts, the channel coasts of France, including the Channel Islands, along the Atlantic coasts of France, Spain, Portugal and North West Africa to the Canary Isles. Throughout the Mediterranean, Adriatic, Aegean, Ionian and Black seas, in the Bermudas and on the north-eastern seaboard of the United States of America (Rhode Island and Massachusetts). Microdeutopus damnoniensis (Bate) (Text-figs, yb, 8f, ga, g, zoa-b, 2oh, j) Lembos damnoniensis Bate, 1856 (nomen nudum): 58; Bate, 1857 : 142. Lembos danmoniensis [sic] (Bate) White, 1857 : 180. Microdeutopus gryllotalpa Bate, 1862 : 163, pi. 30, fig. i; Bate & Westwood, 1863 : 289. Microdeutopus damnoniensis (Bate) Norman, i9O5a : 24; Stebbing, 1906 (pro-parte) : 593; Chevreux & Fage, 1925 (pro-parte): 297, fig. 308; Barnard, 1958 : 29. Microdeutopus danmoniensis [sic] (Bate) Norman & Scott, 1906: 83. [Non] Microdeutopus danmoniensis [sic] Sars, 1894:542, pi. 192. TYPE LOCALITY. Plymouth, Devon, England. DIAGNOSIS OF MALE. Antennule slightly over one half the body length, the first and second peduncular articles subequal, the third almost one half the length of the second; flagellum a little longer than the peduncle, normally with up to fourteen articles (maximum recorded fifteen) ; accessory flagellum with two articles the terminal article rudimentary. Antenna shorter than the antennule, the fourth and fifth peduncular articles subequal ; flagellum about equal to the fifth peduncular article normally with up to seven articles (maximum recorded eight). Per aeon REVISION OF MICRODEUTOPUS 115 segments 2-4 each with an anteriorly directed spine in the mid-ventral line, that of segment 2 the longest. Gnathopod i with article i moderately produced anteriorly, less acute than in M. anomalus or M. algicola ; article 2 expanded somewhat distally ; article 5 very enlarged, longer than broad, the posterior distal angle produced into a single, long, slender, inward curved tooth which in fully mature specimens consider- ably exceeds half the length of article 6 ; article 6 with the posterior margin smoothly rounded; article 7 relatively long with accessory teeth on the posterior margin. Gnathopod 2 with article 2 expanded; articles 5 and 6 short, 5 slightly the longer, but scarcely broader than article 6, which is scarcely twice as long as broad, and has the palmar angle almost transverse; article 7 short and robust. Uropod 3 having the peduncle with its lateral and dorsal flanges very distinct, as in M. algicola; rami short and very stout, not exceeding the length of the peduncle, the exopodite the longer of the two rami, with two separated spines on the outer margin, the more distal of which is much the longer, endopodite with a single median spine on the outer margin, and a single more distal spine on the inner, each ramus terminating in a group of spines of which one is relatively long. Telson as M. anomalus. Maximum size of male : 4-5 mm. FEMALE. As the male, except for the structure of the sexually dimorphic gnathopoda, and the absence of ventral peraeon spines. Gnathopod i with article i not markedly produced anteriorly, very obtuse; article 2 somewhat expanded; article 6 slightly longer than article 5 and narrowing somewhat at the junction with it; article 7 longer than the palmar region. Gnathopod 2 with article 2 unexpanded and article 6 somewhat longer and more slender than article 5, with the palmar margin angular. Maximum size of female : 4-5 mm. DISCUSSION. Andersson (1954) using material of Swedish origin concluded that M. propinquus Sars and M. damnoniensis (Bate) were both synonyms of M. anomalus (Rathke). Present investigations, whilst supporting the contention that M. propinquus is synonymous with M. anomalus, indicate that M. damnoniensis certainly is not. Since there are no substantiated records of M. damnoniensis north of latitude 51° N. it is probable that Andersson was working entirely with a population of M. anomalus. The distinction between M. anomalus and M. damnoni- ensis is not simply one of presence or absence of an accessory tooth, and most so- called M. damnoniensis in the literature are in fact immature M. anomalus. Never- theless, true M. damnoniensis does exist and males are distinguishable from those of M. anomalus and M. algicola by a number of important characters, including the very slender, inward curved tooth on article 5 of gnathopod i (Text-fig. 8f), the short, broad articles 5 and 6 of gnathopod 2 (Text-fig, ga), and the short, robust uropod 3 rami (Text-fig, gg). In addition, the number of articles to the accessory flagellum of the antennule remains constant from eclosion to maturity, in contrast with the condition in M. anomalus where the number of articles increases with age. ECOLOGY. Unlike M. anomalus, the present species occurs most commonly in the intertidal zone, among sponges, polyzoans and corallines, especially in rock pools, or in the shallow sub-littoral. u6 A. A. MYERS FIG. 9. Microdeutopus damnoniensis (Bate), (a) Male gnathopod 2, Plymouth, Devon, (g) Male uropod 3, with transverse section across peduncle at z, Plymouth, Devon. Microdeutopus algicola Delia Valle. (b) Male gnathopod 2, Naples, Italy, (d) Male telson, Naples, Italy, (f) Male uropod 3, with transverse sections across peduncle at x and y, Naples, Italy. Microdeutopus anomalus (Rathke). (c) Male gnathopod 2, Plymouth, Devon, (e) Male uropod 3, with transverse section across peduncle at w, Plymouth, Devon. REVISION OF MICRODEUTOPUS 117 DISTRIBUTION. Very difficult to ascertain, due to confusion in the literature with immature M. anomalus. Present investigations have confirmed material from Drake's Island and Wembury Bay, Plymouth, Jersey and Guernsey, and Bandol, France. TYPE MATERIAL. A lectotype has been erected from Bate's syntypic series (see Appendix 2) . Microdeutopus algicola Delia Valle (Text-figs. 8g-h, gb, d, f, lod-e, 2of) [?] Microdeutopus grillotalpa Sowinsky, 1880 : 125, pi. 5, fig. lya-d. Microdeutopus algicola Delia Valle, 1893 : 41**, pi. i, fig- 3. pi- n, figs. 1-12; Norman, igc^a : 25- Microdeutopus damnoniensis (Bate) Stebbing, 1906 (pro-parte) : 593; Chevreux & Page, 1925 (pro-parte): 297, fig. 308; [?] Miloslavskaia, I939a : 124, fig. 24; [?] Soika, 1949 : 199, [?] Gur- janova, 1951 : 834, fig. 584. TYPE LOCALITY. Mergellina, Naples, Italy. DIAGNOSIS OF MALE. Antennules and antennae have been missing in material examined in present investigations, and the description of these appendages here is extracted from the description and figures of Delia Valle (1893). Antennules with the first and second peduncular articles sub-equal, the third about one third the length of the second; flagellum longer than the peduncle with twenty articles; accessory flagellum possibly with two articles, but almost certainly three, of which the terminal is rudimentary as figured (for M. grillotalpa) by Sowinsky (1880). Antenna shorter than the antennule, the fourth peduncular article slightly longer and more robust than the fifth, flagellum about equal to the fifth peduncular article, with eight articles. Per aeon segments 2-4 each with an anteriorly directed spine in the mid- ventral line, that of segment 2 the longest; segment 5 with a vestigial spine. Gnathopod I with article I moderately produced anteriorly, acute, though less attenuated than in M. anomalus; article 2 relatively robust, expanded somewhat distally; article 5 very enlarged, the posterior margin produced into a single, robust, broad based tooth not exceeding half the length of article 6 ; article 6 with the posterior margin convex distally, straight or slightly concave basally; article 7 robust, with accessory teeth on the posterior margin. Gnathopod 2 with article 2 having the anterior margin produced into a flange which terminates distally in a blunt process ; articles 5 and 6 slender, subequal in length, article 5 slightly the broader; article 6 over twice as long as broad, with the palmar region oblique ; article 7 short and robust. Uropod 3 with the peduncle broad, due to the marked development of the lateral flanges, particularly the inner, which exceeds in length both the outer flange and the dorsal crest; rami long and slender, about equal in length to the peduncle, the exopodite slightly the longer, not differing in spination from M. anomalus of a similar size. Telson as in M. anomalus. Maximum size of male : 4-5 mm. FEMALE. As the male, except for the structure of the sexually dimorphic gnatho- poda, and absence of ventral peraeon spines. Gnathopod I with article I not markedly produced anteriorly, obtuse ; article 2 robust, otherwise scarcely differing n8 A. A. MYERS from that of M. damnoniensis. Gnathopod 2 with article 6 having the palmar angle rounded, otherwise as that of M. damnoniensis. Maximum size of female 4-5 mm. DISCUSSION. M. algicola is the most robust member of the " anomalus " group of species. In facies it more closely resembles M. stationis, than it does M. anomalus FIG. 10. Microdeutopus damnoniensis (Bate), (a) Female gnathopod i, Plymouth, Devon. (b) Female gnathopod 2, Plymouth, Devon. Microdeutopus anomalus (Rathke). (c) Female gnathopod i, Plymouth, Devon, (f) Female gnathopod 2, Plymouth, Devon. Microdeutopus algicola Delia Valle. (d) Female gnathopod 2, Naples, Italy, (e) Female gnathopod i, Naples, Italy. or M. damnoniensis, and it is noteworthy that a male M. algicola was discovered during present re-examinations of M. stationis material in the Chevreux collections. Male M. algicola are distinguishable from those of M. damnoniensis, in having the tooth on article 5 of gnathopod i (Text-figs. 8g-h) broad-based and triangular, whereas in M. damnoniensis (Text-fig. 8f) it is slender throughout its length, arising REVISION OF MICRODEUTOPUS 119 at the posterior distal angle only. The present species also differs from M. damnon- iensis in the structure of gnathopod 2 (Text-fig, gb), which has articles 5 and 6 slender, as in M. anomalus and in the form of uropod 3 (Text-fig, gf), which has slender elongate rami. In lacking an accessory tooth on article 5 of the male gnathopod i, M. algicola differs from typical M. anomalus which usually possesses such a tooth. In specimens of the latter species lacking an accessory tooth (including immature specimens) the primary tooth is structurally very different, being longer and more slender, with the posterior margin more smoothly rounded, and with a narrower base, than it is in M. algicola. In addition, the flange on the anterior margin of article 2 of gnathopod 2 terminates distally in a more markedly developed process in M. algicola than in M . anomalus. ECOLOGY. At Mergellina (Delia Valle, 1893) among attached algae on the shore, and at Bone in 15-20 metre depth. DISTRIBUTION. Difficult to ascertain, due to the scarcity of reliable records. The present author has seen material from Naples, Italy and Bone, Algeria, and considers that the M. grillotalpa Sowinsky from Sevastopol in the Black Sea, refers to this species. The species may prove to be widely distributed in the Mediterranean region, and Black Sea. THE " SCHMITTI " GROUP OF SPECIES The " Schmitti " group of species occurs over a wide geographical area, around both the Pacific and Atlantic coasts of the Americas. So far material has been examined from a number of isolated geographical localities, and regional morpholo- gical differences are difficult to assign to varietal, subspecific and specific status. Nevertheless there seems at present to be justification for grouping these forms into at least three species, of which M. schmitti Shoemaker and M. hancocki Myers are sympatric, overlapping in their distribution and maintaining their separate identities. It is clear that ecological studies on the " schmitti " group of species would be of considerable value. Males of the " schmitti " group, with the exception of M. schmitti itself, differ from those of all other species of Microdeutopus by the development of a tooth on the anterior margin of article 5 of gnathopod i. M. schmitti differs from M. gryl- lotalpa Costa (the only species with which it could be confused) by having article 2 of gnathopod 2 concave and smooth on the anterior margin, whereas in M. gryllo- talpa the anterior margin is convex and crenated. Females of M. schmitti differ from those of all other known species in the genus (but see also Hansenella longi- cornis Chevreux, p. 138) by the development of a tooth or teeth on the posterior margin of article 5 of gnathopod i. Females of other species in the " schmitti " group are without recognizable diagnostic features and their identification in the absence of males cannot be attempted with certainty. Most apparent differences in structure between M. schmitti and the smaller species M. hancocki and M. trichopus are due to neoteny. The rami of the third uropod of mature female M. hancocki and M. trichopus, for example, correspond to those of immature specimens of M. schmitti of a similar size. I2O A. A. MYERS Microdeutopus schmitti Shoemaker (Text-figs, n, i3b, d, i4a-c, f, g, 1, 15, i6a-b, d, 2om, pi. ib) Microdeutopus schmitti Shoemaker, 1942 : 18, fig. 6; Barnard, 1958 : 29; Barnard, 1959 : 32, pi. 9; Myers, 19685 : in press. TYPE LOCALITY. Bahia de Magdalena, Baja California, Mexico. DIAGNOSIS OF MALE. Pamgnaths with the mandibular processes relatively short and stout. Antennule slightly over one third the body length, the first and second peduncular articles subequal, the third a little less than one half the length of the FIG. ii. Microdeutopus schmitti Shoemaker. Male head, Bahia de Salinas, Costa Rica. second; flagellum about equal in length to the peduncle, normally with up to ten articles (maximum recorded twelve); accessory flagellum with three articles, the third article rudimentary. Antenna about two thirds the length of the antennule, the fourth and fifth peduncular articles subequal; flagellum shorter than the fifth peduncular article, with four articles, of which the first is equal to the combined length of the terminal three. Peraeon segments 3-6 each with a short, robust, REVISION OF MICRODEUTOPUS FIG. 12. Microdeutopus hancocki Myers, (a) Male head, Bahia de Salinas, Costa Rica (Paratype). Microdeutopus trichopus Myers, (b) Male head, Isabela Island, Galapagos (Paratype). 122 A. A. MYERS anteriorly directed spine in the mid- ventral line. Gnathopod i with article i moderately produced anteriorly, rounded ; article 2 expanded, oval, article 4 roughly triangular, with relatively short setae on the posterior margin ; article 5 longer than broad, with- out teeth on the anterior margin, the posterior distal angle produced into a large, stout tooth, at the base of which are from one to four small forward projecting teeth; article 6 short, the posterior margin with a distal and proximal lobe; article 7 relatively long, with accessory teeth on its posterior edge. Gnathopod 2 with article 2 somewhat concave on its anterior margin, the anterior distal corner produced into a rounded lobe; article 5 slightly longer than article 6 and expanded distally, where it considerably exceeds the width of article 6; articles 4, and 5 with numerous long, finely pectinate setae arising from the anterior and posterior margins. Uropod 3 with the rami subequal, slightly longer than the peduncle; endopodite with two to four equally spaced spines on the inner margin and one or two on the outer; exopodite with two well separated spines on the inner margin, and on the outer a solitary spine, basal to which in fully mature adults is a pair of closely associated spines, the outer of which is the longer. Telson with each terminal crest bearing on its distal dorsal margin a group of three or four setae, two or three of which are short and spiniform. Additional setae may appear spiniform during growth, which is in contrast with the condition in M. stationis where the spines are always quite distinct from the setae. Maximum size of male : 5 mm. FEMALE. As the male except for the absence of ventral peraeon spines, and the sexually dimorphic gnathopoda. Gnathopod i with article 2 unexpanded; articles 5 and 6 subequal in length, article 5 distinctly the broader, with the posterior distal angle produced into one or two small teeth, article 6 with the anterior margins roughly parallel; article 7 slender, considerably longer than the palmar region with accessory teeth on the posterior edge. Gnathopod 2 with article 2 unexpanded; articles 5 and 6 subequal in length, article 5 roughly pentagonal with the anterior margin almost straight, broadening distally, where it considerably exceeds the breadth of article 6 ; anterior and posterior margins of articles 4 and 5 bearing long pectinate setae. Maximum size of female : 6 mm. DISCUSSION. M. schmitti is closely related to M. hancocki and M. trichopus but differs from these in the male, by having the anterior margin of article 5 of gnathopod i (Text-figs. I3b, d) devoid of any toothlike processes and in the female by the pres- ence of teeth on the posterior distal angle of article 5 of gnathopod i (Text-fig. i6b,d). ECOLOGY. In the sub-littoral to a recorded depth of 42 metres (off Point Loma Light, California). Tolerant of a wide range of conditions, on mud, fine or course sand, and rock substrata, among various vegetation, shells, and coral. DISTRIBUTION (Text-fig. 15). Along the coasts of California (Cayucos, Point Conception, Point Loma, San Clemente, and Newport), Baja California (El Coyote, Bahia de San Quintin, Bahia de Magdalena and Cape San Lucas) and Costa Rica (South Viradores Islands, Playa Blanca and Bahia de Salinas). REVISION OF MICRODEUTOPUS 123 FIG. 13. Microdeutopus hancocki Myers, (a) Male gnathopod i, Bahia de Salinas, Costa Rica (Holotype). (c) Male gnathopod i, Salango Island, Equador. Microdeutopus schmitti Shoemaker, (b) Male gnathopod i, Bahia de San Quintin, Baja California. (d) Male gnathopod i, Bahia de Salinas, Costa Rica. Microdeutopus trichopus Myers. (e) Male gnathopod i, Isabela Island, Galapagos (Holotype). Microdeutopus sp. nov.? (f) Male gnathopod i, Tortugas, Florida, U.S.A. (g) Male gnathopod i, Tortugas, Florida, U.S.A. ZOOL. 17, 4 8 124 A. A. MYERS Microdeutopus hancocki Myers (Text-figs. I2a, I3a, c, I4d, h, m, 15, i6c, e, 2ok, pi. la) Microdeutopus hancocki Myers, ig68b : In press. TYPE LOCALITY. Bahia de Salinas, Costa Rica. DIAGNOSIS OF MALE. Antennules and antennae not noticably different from those of M. schmitti. Paragnaths with the mandibular processes longer and more slender than in M. schmitti. Per aeon segments 3-5 each with a short robust, anteriorly directed spine in the mid- ventral line, segment 6 with a small or vestigial spine. Gnathopod I with article I moderately produced anteriorly, rounded; article 2 markedly expanded anteroproximally ; article 4 roughly triangular, with very long setae on the posterior margin ; article 5 longer than broad, oval, with a median dentiform process on the anterior margin, and at the posterior distal angle, three stout teeth, the most distal of which is the longest; article 6 short, with a single lobe on the posterior margin ; article 7 relatively long, with accessory teeth on the posterior edge. Gnathopod 2 with article 2 concave on the anterior margin ; article 5 consider- ably longer than article 6 but scarcely broader; articles 4 and 5 with numerous long, finely pectinate setae arising from the anterior and posterior margins. Uropod 3 and telson scarcely differing from those of M. schmitti of a comparable size. Maximum size of male 3-8 mm. FEMALE. As the male except for the absence of ventral peraeon spines, and the sexually dimorphic gnathopoda. Gnathopod i similar to that of M . schmitti but with the posterior margin of article 5 without teeth at the posterior distal angle. Gnathopod 2 scarcely differing from that of M. schmitti, but with the pentagonal shape of article 5 less well marked. Maximum size of female 4-1 mm. DISCUSSION. M. hancocki is very closely related to M. schmitti, from which it differs in the male having a tooth-like process on the anterior margin of article 5 of gnathopod i (Text-fig. I3a, c), and article 4 of this appendage bearing very long setae. In addition the structure of the ventral peraeon spines (Text-fig. 2ok) and paragnath (PI. la) and the shape of the second gnathopod (Text-fig. I4m) distinguishes the present species from M. schmitti. It differs from M. trichopus by having article 6 of gnathopod i devoid of any forward projecting teeth on its posterior margin. ECOLOGY. Apparently more habitat-specific than M. schmitti, having been recorded at present only from sandy bottoms to a depth of about 18 metres. DISTRIBUTION (Text-fig. 15). Costa Rica (Bahia de Salinas), Panama (Bahia Honda), Equador (Salango Island), and Galapagos (Isabela Island). crest of male telson, Bahia de Salinas, Costa Rica (Paratype). (m) Male gnathopod 2, Bahia de Salinas, Costa Rica (Paratype). Microdeutopus trichopus Myers. (e) Male uropod 3, Isabela Island, Galapagos (Holotype). (i) Dorsal view of left terminal crest of male telson, Isabela Island, Galapagos (Holotype). (j) Male gnathopod 2, Isabela Island, Galapagos (Paratype). Microdeutopus sp. nov.? (k) Male gnathopod 2, Tortugas, Florida. REVISION OF MICRODEUTOPUS 125 FIG. 14. Microdeutopus schmitti Shoemaker, (a)-(c) Male uropod 3, all figured the same size (and therefore each at a different scale) for easy comparison, (a) 3-0 mm. $ (b) 4-5 mm $ (c) 5-0 mm. $, Bahia de San Quintin, Baja California, (f) (g) Dorsal view of left terminal crest of male telson, Bahia de San Quintin, Baja California. (1) Male gnatho- pod 2, Bahia de San Quintin, Baja California. Microdeutopus hancocki Myers, (d) Male uropod 3, Bahia de Salinas, Costa Rica (Holotype). (h) Dorsal view of left terminal 126 A. A. MYERS Microdeutopus trichopus Myers (Text-figs. I2b, 136, 146, i, j, 15, i6f-g, 20g) Microdeutopus trichopus Myers, ig68b : In press. TYPE LOCALITY. East of south end of Isabela Island, Galapagos. DIAGNOSIS OF MALE. Antennule slightly over one half the body length, the first peduncular article slightly shorter than the very slender second article, the third about a third the length of the second article; flagellum somewhat longer than the peduncle, normally with up to twelve articles (maximum recorded thirteen) ; accessory flagellum with three articles, the third rudimentary. Antenna about two thirds the length of the antennule, the fourth and fifth peduncular articles elongate, slender subequal; flagellum shorter than the fifth peduncular article with four, rarely five articles, the first article long, about equal to the combined length of the terminal three or four. Peraeon segments 3-6 each with a long and slender spine in the mid- ventral line, those of segments 3 and 4 directed anteriorly those of segments 5 and 6 straight or slightly recurved. Gnathopod I with article I moderately produced anteriorly, rounded ; article 2 expanded anteroproximally ; article 4 short and bulky ; article 5 oval, longer than broad, with a median dentiform process on the anterior margin and at the posterior distal angle, a short stout tooth, basal to which on the posterior margin is a further small tooth ; article 6 over one half the length of article 5, the palmar angle produced into a short, blunt ending, forwardly projecting tooth, opposable to article 7; article 7 of moderate length, with accessory teeth on the posterior margin ; the posterior margins of articles 4, 5 and 6 densely setose, the setae exceptionally long, in the largest males measuring 0-5 mm., 0-4 mm. and 0-3 mm. on each article respectively. Gnathopod 2 with article 2 somewhat expanded distally, the anterior margin slightly concave; articles 5 and 6 subequal in length, article 5 slightly the broader. Uropod 3 with the rami long and slender, longer than the peduncle, the exopodite slightly the longer of the two rami; each ramus with a single spine on each of the inner and outer margins. Telson with each terminal crest bearing on its distal dorsal margin a long seta and a short stout spine. Maximum size of male : 3-8 mm. FEMALE. As the male except for the absence of ventral peraeon spines, and sexually dimorphic gnathopoda. Gnathopod I with article 2 unexpanded, articles 5 and 6 slender, subequal in length, article 6 somewhat broader distally than at the junction with article 5; article 5 without teeth at the posterior distal angle; article 7 longer than the palmar region, with accessory teeth on the posterior edge. Gnatho- pod 2 with article 2 unexpanded; articles 5 and 6 subequal in breadth, article 6 slightly the longer, broadening distally; article 7 short and stout. Maximum size of female : 4-0 mm. DISCUSSION. M. trichopus differs in the male from all other known Microdeutopus species (but see following form and Lembopsis Pearse, (p. 139), in having the posterior margin of article 6 of gnathopod I (Text-fig. 130) produced into a forward projecting tooth opposable to article 7. ECOLOGY. M. trichopus appears to be a relatively deep water species having REVISION OF MICRODEUTOPUS 127 been absent from samples of M. hancocki taken in shallow waters in geographical areas where the present species is known to occur. Recorded from 58-110 metres over mud, and also among nullipores on sandy bottoms. DISTRIBUTION. Galapagos (two localities off Isabella Island). Microdeutopus sp. nov.? (Text-figs. i3f-g. i4k, 15, 20) Two males and a single female of a species of Microdeutopus of the " schmitti " group, were examined, from Tortugas, Florida. This material most closely resembles M. trichopus from the Galapagos archipelago but it differs from that species particu- larly in the structure of the male gnathopod i. This has article 4 elongate and M. schmitti Shoe. M. hancocki Myers M.trichopus Myers ] Microdeutopus sp. nov. ? FIG. 15. The recorded distribution of the " schmitti " group of species in the Americas. rectangular in the present material, whereas it is short and triangular in M. trichopus, in addition article 6 is short and article 7 opposable to the teeth of both articles 5 and 6 in present material, whereas in M. trichopus article 6 is over half the length of the article 5, with article 7 opposable to the teeth of article 6 only. Lastly, the setation of gnathopod i is considerably more sparse in present material than in M . trichopus. On gnathopod 2 article 5 is slightly longer than article 6 in present material, whilst the same two articles are subequal in M. trichopus. 128 A. A. MYERS FIG. 16. Microdeutopus schmitti Shoemaker, (a) Female gnathopod 2, Bahia de San Quin- tin, Baja California, (b) Female gnathopod i, Bahia de San Quintin, Baja California, (d) Female gnathopod i, Bahia de Salinas, Costa Rica. Microdeutopus hancocki Myers, (c) Female gnathopod 2, Bahia Honda, Panama (Paratype). (e) Female gnathopod i, Bahia Honda, Panama (Paratype). Microdeutopus trichopus Myers, (f) Female gnatho- pod 2, Isabela Island, Galapagos (Paratype). (g) Female gnathopod i, Isabela Island, Galapagos (Paratype). REVISION OF MICRODEUTOPUS 129 The present form is separated geographically from M. trichopus by the Central American isthmus, which has not been severed since the late Miocene, so one might expect the two forms to be specifically distinct. This possibility is strengthened by what is known so far of the ecology of the two forms, M. trichopus occurring in, and apparently restricted to, muddy and sandy bottoms in over 50 metres depth, while present material is found in relatively shallow waters amongst weeds. More abundant material from a wider range of localities is required before the true taxonomic status of the present form can be elucidated. DESCRIPTION OF MALE. Antennules and antennae missing. Per aeon segments 3-6 each with a spine on the mid- ventral line, those of segments 3 and 4 directed anteriorly, those of segments 5 and 6 straight or slightly recurved. Gnathopod I with article i moderately produced anteriorly, rounded ; article 2 markedly expanded anteroproximally; article 4 very elongate, roughly rectangular; article 5 oval, longer than broad, with a median dentiform process on the anterior margin, and at the posterior distal angle, a short stout tooth, basal to which, on the posterior margin, are two further small teeth; article 6 about one third the length of article 5, the palmar angle produced into a short, blunt ending, irregular, forward projecting tooth, basal to which is a small lateral projection; article 7 relatively long, opposable to the teeth of both articles 5 and 6; the posterior margins of articles 4, 5, and 6 bearing long setae. Gnathopod 2 with article 2 very slender basally, markedly expanded proximally ; article 5 somewhat longer and broader than article 6. Uropod 3 with the rami long and slender, longer than the peduncle, the exopodite the longer of the two rami, each ramus with a single spine on its inner margin. Telson with each terminal crest bearing on its distal dorsal margin, a long seta, and a short spine. Length of males : 3-4 mm. and 3-5 mm. FEMALE. Antennules missing. Antennae about one third the body length, the fourth and fifth peduncular articles elongate, slender, subequal; flagellum shorter than the fifth peduncular article, with four articles, the first article long, slightly shorter than the combined length of the terminal three. Gnathopods i and 2 not differing from those of M. trichopus. Length of female : 3-3 mm. MATERIAL, i $ and i $; station 48-30, Qth August, 1930, east and south of Loggerhead Light, Dry Tortugas, Monroe Co., Florida, in 20-20 metres, i <$; station 12-31, 26th June, 1931, Tortugas, Florida (24° 36' N. 82° 56' W. approx. from chart) in 18-20 metres (low water) net tow-weedy haul. Microdeutopus armatus Chevreux (Text-figs, ryd, 18, 20!) Microdeutopus armatus Chevreux, 1886 : XLI; Chevreux, i88jb : 312, pi. 5, figs. 11-12; Chevreux, iSSya : 92; Stebbing, 1906 : 589; Chevreux & Page, 1925 : 296, figs. 303, 307; Barnard, 1958 : 29. Stimpsonella armata (Chevreux) Delia Valle, 1893 : 422, pi. 4, fig. 8, pi. n figs. 13-24; Chevreux, 1900 : 89; Norman i905a : 25. 130 A. A. MYERS TYPE LOCALITY. North East of Basse-Kikerie, France. DIAGNOSIS OF MALE. Antennule between one half and two thirds the body length, the second peduncular article longer than the first, the third a little over one third the length of the second; flagellum longer than the peduncle, normally with up to fourteen articles (maximum recorded sixteen) ; accessory flagellum with two articles, FIG. 17. Microdeutopus chelifer (Bate), (a) Male head, with (b) a transverse section and (c) a lateral view of the third peduncular article of the antenna, Jersey, Channel Islands. Microdeutopus armatus Chevreaux. (d) Male head, Villefranche, France. REVISION OF MICRODEUTOPUS 131 the second article rudimentary. Antenna about two thirds the length of the antennule, the fourth and fifth peduncular articles subequal ; flagellum slightly shorter than the fifth peduncular article with five articles. Peraeon segments 2-5 each with a spine in the mid- ventral line, that of segment 2 slender hook-shaped, directed anteriorly, those of segments 3-5 robust anteriorly directed, that of segment 5 the shortest. Gnathopod 1 with article I moderately produced anteriorly, acute; article 2 short and greatly expanded ; article 5 almost as broad as long, the posterior distal angle produced into a large, broad, blunt ended tooth; article 6 short, the posterior margin irregularly lobed; article 7 relatively long. Gnathopod 2 with article 2 expanded, the anterior margin produced into a crenated flange; articles 5 and 6 roughly rectangular, subequal in length and breadth, article 6 with the palmar angle produced into a short tooth. Uropod 3 with the rami subequal, slightly shorter than the peduncle; endopodite with a pair of spines on the inner dorsal margin near the apex, and one long and one short terminal seta, exopodite with a similar pair of terminal setae. Telson with each terminal crest bearing, on its distal dorsal margin, a long seta. Maximum size of male : 3 mm. FEMALE. As the male except for the structure of the sexually dimorphic gnatho- poda, and absence of ventral peraeon spines. Gnathopod i with article 2 somewhat dilated; article 6 longer than article 5, with the anterior and posterior margins roughly parallel, article 7 very elongate, exceeding in length the palmar region. Gnathopod 2 with article 2 expanded, the antero-distal corner produced into a lobe; article 6 considerably longer than article 5, with the palmar angle produced into a broad- based triangular process. Maximum size of female : 4 mm. DISCUSSION. M. armatus differs in the male from all other species in the genus, except M. chelifer, in having the second gnathopod chelate. It differs from the latter species by having the fifth and sixth articles of gnathopod 2 (Text-fig. i8d), subequal in length and breadth, whereas in M. chelifer (Text-fig. IQC), article 6 is much shorter and much broader than article 5. There is no justification for placing these two species together in a distinct genus (Stimpsonella Delia Valle) as suggested by Delia Valle (1893) on the basis of the chelate male second gnathopoda, since in other respects they exhibit no greater affinity with each other than they do with other Microdeutopus species. The most generally useful character enabling the identi- fication of females of M. armatus is the structure of the second gnathopod (Text-fig. i8e), which has a pronounced projection at the antero-distal corner of article 2, and a triangular process at the palmar angle of article 6. In addition article 2 of the first gnathopod (Text-fig. i8a) is somewhat dilated. ECOLOGY. On soft grey mud (type locality), fine or coarse sands, in the sub- littoral, in 10-90 metres. At Naples (Delia Valle, 1893), associated with Urothoe and Ampelisca. DISTRIBUTION. France, from the type locality, southwards along the coasts of the Bay of Biscay (Le Croisic, Gulf of Gascony), Portugal, and in the Mediterranean off the coasts of France (Antibes, Villefranche) Corsica (Bonifacio) and Italy (Naples). A. A. MYERS Microdeutopus chelifer (Bate) (Text-figs. I7a-c, 19, 2on) Stimpsonia chelifera Bate, 1862 : 162, pi. 29, fig. 9; Bate & Westwood, 1863 : 285; Stebbing, 1878 : 34, pi. 5, figs. 2-3; Norman, 1886 : 17. Microdeutopus chelifer (Bate) Stebbing, 1888 : 334; Stebbing, 1906 : 589; Chevreux & Page, 1925 : 295, fig. 306; Barnard, 1958 : 29. [Nori\ Microdeutopus chelifer Haswell, 1879 : 340, pi. 22, fig. 3. Stimpsonella chelifera (Bate) Delia Valle, 1893 : 424, pi. 56, figs. 42-45; Norman, igosa : 25; Norman, I905b : 82; Norman & Scott, 1906 : 84; Norman, 1907 : 368. TYPE LOCALITY. Salcombe, Devon, England. DIAGNOSIS OF MALE. Antennule slightly over one half the body length, the second peduncular article a little longer than the first, the third over one third the length of the second; flagellum longer than the peduncle, normally with up to fifteen articles (maximum recorded seventeen); accessory flagellum with three articles, the third FIG. 1 8. Microdeutopus armatus Chevreux. (a) Female gnathopod i, Le Croisic, France, (b) Male gnathopod i, Le Croisic, France, (c) Male uropod 3, Le Croisic, France, (d) Male gnathopod 2, Le Croisic, France, (e) Female gnathopod 2, Le Croisic, France, (f) Male telson, Le Croisic, France. REVISION OF MICRODEUTOPUS 133 article rudimentary. Antenna longer than the antennule, the third peduncular article with its lower outer margin produced into a flattened lobe, the fourth pedun- cular article very slightly shorter than the fifth; flagellum longer than the fifth peduncular article, normally with up to ten articles (maximum recorded eleven). FIG. 19. Microdeutopus chelifer (Bate), (a) Male gnathopod i, Paignton, Devon, England, (b) Male gnathopod i, articles 4 and 5, Khios, Greece, (c) Male gnathopod 2, Paignton, Devon, England, (d) Male (Immature) gnathopod 2, articles 6 and 7, Khios, Greece, (e) Male uropod 3, Brittany, France, (f) Female gnathopod i, Guernsey, Channel Islands, (g) Female gnathopod 2, Guernsey, Channel Islands. 134 A. A. MYERS Peraeon segments 2-4 each with an anteriorly directed process in the mid-ventral line, those of segments 2 and 3 long, slender, spiniform, that of segment 4 short, robust. Gnathopod i with article I strongly produced anteriorly, very acute; article 2 scarcely expanded; article 5 very enlarged, but longer than broad, the posterior margin produced sub-distally into a long, narrow, inward curving tooth, distal to which in some, but not all specimens, is a further small tooth; article 6 relatively long, the posterior margin undulate; article 7 robust. Gnathopod 2 with article 2 narrow and elongate, with a small projection at the antero-distal corner; article 6 shorter and broader than article 5, with the palmar region produced into a process, which in young males is short and broad, bearing a stout spine at its tip, but in mature males is drawn out into a long inward curving tooth, when the spine is usually obsolete; article 7 very long, opposable to the palmar process. Uropod 3 with the rami elongate and slender, subequal in length with the peduncle ; FIG. 20. Ventral peraeon processes in Microdeutopus species, (a) M. gryllotalpa Costa Isefjord, Denmark, (b) M. anomalus (Rathke) Falmouth, Cornwall, England, (c) M. versiculatus (Bate) Falmouth, Cornwall, England, (d) M. anomalus (Rathke) Plymouth, Devon, England, (e) M. stationis Delia Valle Guernsey, Channel Islands, (f) M. algi- cola Delia Valle Bone, Algeria, (g) M. trichopus Myers Isabela Island, Galapagos, (h) M. damnoniensis (Bate) Bandol, France, (i) M. sp. nov. ? Tortugas, Florida, U.S.A. (j) M. damnoniensis (Bate) Plymouth, Devon, England, (k) M. hancocki Myers Bahia de Salinas, Costa Rica. (1) M. armatus Chevreux Villefranche, France, (m) M. schmitti Shoemaker Bahia de Salinas, Costa Rica, (n) M. chelifer (Bate) Jersey, Channel Islands. REVISION OF MICRODEUTOPUS 135 exopodite slightly the longer of the two rami, with two isolated spines on the outer margin, of which the more distal is particularly long; endopodite with two well separated spines on the inner margin and one on the outer; each ramus with a term- inal cluster of spines, of which one is greatly elongated. Telson with each terminal crest bearing on its distal dorsal margin, a group of up to 3 setae. Maximum size of male : 8 mm. FEMALE. As the male except for the absence of ventral peraeon spines and the sexually dimorphic antennae and gnathopoda. Antenna with the third peduncular article unmodified, and the entire appendage less well developed than that of the male, such that it does not exceed the length of the antennule. Gnathopod i with article 2 moderately expanded; article 6 longer than article 5, about twice as long as broad, with the anterior and posterior margins roughly parallel; article 7 robust, equal in length to the palmar region. Gnathopod 2 similar to gnathopod I out more slender, the palmar region more transverse. Maximum size of female : 8 mm. DISCUSSION. As in the preceding species, the second gnathopoda of the male are chelate. In the present species, however, article 6 of this appendage (Text-fig, igc) is much shorter and broader than article 5, whereas in M. armatus (Text-fig. i8d) the fifth and sixth articles are more or less subequal in length and breadth. M. chelifer is unique among the known Microdeutopus species in having sexually dimorphic antennae, but this feature is not considered to be a sufficient reason for separating the present species from the genus Microdeutopus (see also Stebbing, 1888). Females have few diagnostic characters, but important identification features when taken in combination are : the accessory flagellum of the antennule having three articles (Text-fig. lya), the third uropod rami having their sparse spination of solitary spines, the more distal of two spines on the outer margin of the exopodite being exceptionally long (Text-fig. IQC) and the telson bearing setae, but no spines. ECOLOGY. In the scarcely tidal Mediterranean the present species occurs in shallow waters, among Cystoseira, Chondrus, Saccorhiza etc. On the Atlantic and English Channel coasts, the species is also found in the shallow sub-littoral among Laminaria holdfasts and other weeds, but it most frequently inhabits the littoral zone, among algae in rock pools often high up on the shore, particularly in sheltered places with a high accumulation of silt, often where there is some brackish water influence. DISTRIBUTION. Along the south-western coasts of the British Isles (Studland, Torbay, Wembury Bay, Salcombe, Port wrinkle) , the Channel Islands and the adjoin- ing French coast, and thence at a number of localities along the coasts of the Bay of Biscay to Guethery in the extreme south. Recorded from Cadiz, and probably occurs in a number of favourable localities around the coasts of Spain and Portugal. In the Mediterranean it has been recorded from the coast of France (Sete, Monaco, Antibes, Villefranche) and in the Aegean from the Greek Island of Khios, off the Turkish coast. The IStimpsonia sp. of Sowinsky (1898) from the Black sea may refer to this species. 136 A. A. MYERS Microdeutopus sporadhi sp. nov. (Text-figs. 21, 22) DESCRIPTION OF MALE HOLOTYPE. Mouthparts not differing significantly from those of other members of the genus. Head lateral-lobes moderately well developed, obtuse; eye reniform. Antennule about two thirds the body length, the first pedun- cular article shorter than the second, the third less than one half the length of the FIG. 21. Microdeutopus sporadhi sp. nov. Anterior end of male holotype, Emborios Bay, Khios, Greece. second; flagellum about one and one half the length of the peduncle, with thirteen articles; accessory flagellum with two articles, the second article minute, shorter than the first article of the primary flagellum. Antenna shorter than the antennule, the fourth and fifth peduncular articles subequal ; flagellum scarcely longer than the fifth peduncular article, with seven articles. Peraeon segments 2 and 3 each with a robust, anteriorly directed spine in the mid-ventral line, that of segment 2 the longer. Gnathopod i with article I strongly produced anteriorly, somewhat attenuated, article 2 elongate, relatively slender, but broadening distally, its anterior margin excavate for the reception of the carpus when folded, article 4 slender, scarcely over lapping the posterior margin of article 5, article 5 grossly expanded, longer than broad, REVISION OF MICRODEUTOPUS 137 slender basally, broad distally, with groups of long fine setae arising from within the distal and postero-distal margins, and with the posterior distal angle produced into two distinct teeth, the proximal (outer) tooth curved inward, relatively long and slender, terminating acutely, the distal (inner) short, robust, obtuse, article 6 elongate and slender, broadening somewhat distally, but with the palmar region almost obsolete, the posterior margin almost straight, article 7 very long, though slightly shorter than article 6, with accessory teeth on the posterior margin. Gnatho- pod 2 with article 2 relatively short and broad, the anterior margin almost straight, articles 5 and 6 subequal in length and breadth, article 7 relatively short. Peraeo- pods 3-7 of the typical Microdeutopus structure. Uropod 3 with the rami short, not exceeding the length of the peduncle, exopodite slightly the longer of the two rami, devoid of spines on the lateral margins, terminating distally in a group of five long setae, of which the two most distal arise from a vestigial second article; endopodite with a single median spine on the inner margin and a group of four FIG. 22. Microdeutopus sporadhi sp. nov. Male paratypes. (a) Gnathopod i. (b) Telson. (c) Uropod 3. (d) Paragnath. (e) Terminal article of maxillule palp, (f) Maxillule. (g) Gnathopod 2. (h) Ventral peraeon processes, (i) Mandible. 138 A. A. MYERS terminal setae. Telson of the usual Microdeutopus form, each terminal crest bearing on its distal dorsal margin a pair of long setae of unequal length. Length : 3-0 mm. MALE PARATYPES. Agree closely with the holotype in all diagnostic characters. Antennal flagellum of one paratype with 8 articles, otherwise this appendage also invariable. Length of paratypes 2-8-3-2 mm. FEMALE. Unknown. TYPES, Deep-sea trawling (Kirkpatrick, 1889). i : approx. 50° 50' N, 11° 26' W; 576 m. ^ " L'Hirondelle " (Jullien & Calvet, 1903). (Longitudes refer to Greenwich, see p. 238.) 42 : 46° 47' N, 3° 52' 30 " W; 136 m. 44 : 46° 27' N, 4° 10' W; 166 m. 46 : 46° 25' N, 3° 35' 30" W; 155 m. 85 : 46° 31' N, 4° 32' W; 180 m. V " Princess Alice II ", " L'Hirondelle II " (Calvet, 1931). 1043 : 59° 03' N, i° 48' W; 88 m. 1535 : 47° 46' N, 5° 40' W; 132 m. 2974 : off Belle-He; 85 m. ^ " Vienne " on La Chapelle Bank (Gu6rin-Ganivet, 14 : 47° 37' N, 6° 30' W; 200 m. J5 : 47° 3°' N, 6° 50' W; 400 m. ZOOL. 17, 6. 16 238 J. S. RYLAND years, collections of Polyzoa have been made by R. V. " Sarsia " (Ryland, in preparation) . The northern boundary at 62 ° N passes clear of the Shetland Islands to the Norwe- gian coast at Nordfjorden. The well-studied area around Bergen is on the same latitude as Shetland. The Faroe Islands, which straddle latitude 62° N, have also been included. The natural western boundary follows the continental slope (the 1500 m. isobath approximately sets the limit at the present time) from Faroe, west of Faroe Bank, Rockall Bank and Porcupine Bank, then across the Western Approaches into the Bay of Biscay. The eastern boundary lies at 14° E, which includes the Oresund and Kiel Bay within the area, but excludes the Baltic Sea. A small number of High Arctic species, that do not occur in Norway south of the Lofoten Islands, have been recorded from the Faroe Islands (Kramp, 1934)— apparently colonized from Greenland and Iceland via the North Atlantic Trans- versal Ridge. Further southward spread by way of the Wyville Thomson Ridge to northern Scotland might also be possible, and one species (Celleporina surcularis) may perhaps have reached the Shetland Islands by this route (see comment on p. 242). Many of the records, particularly from deep water, come from the reports of research ships, and the positions of relevant stations are indicated in Text-fig. 4, together with other localities mentioned in this list. (It should be noted that Jullien & Calvet, 1903, like Calvet, 1907, and Guerin-Ganivet, 191 la, measure longitude with reference to the Paris meridian, 2° 20' 14" east of Greenwich).1 Some of the species listed are very imperfectly known, and the generic placing may be rather conjectural. In square brackets one or more references to descrip- tions follows the name, and then come details of the occurrence of the species. POLYZOA CHEILOSTOMATA (ANASCA) Membranipora tenuis Desor, 1848 [Marcus, 1940 : 126; Prenant & Bobin, 1966 : 120], Present in the Oresund (Levinsen, 1894; Marcus, 1940). Membranipora tuberculata (Bosc, 1802) [Prenant & Bobin, 1966 : 115]. Washed up on Sargassum weed at Brest (Guerin-Ganivet, 19116, as M. tehuelcha) . Conopeum seurati (Canu, 1928) [Prenant & Bobin, 1966 : 1207]. In brackish water, east, south and west coasts of Britain (Markowski, 1960, as C. spiculatum; Cook, 1962). Sarsiflustra abyssicola (M. Sars in G. O. Sars, 1872) [Kluge, 1962 : 319; Prenant & Bobin, 1966 : 173]. Taken by the " Michael Sars " off the north-west of Scotland, St. 76 (Nordgaard, 1907). Type-species by monotypy of Sarsiflustra Jullien (Jullien & Calvet, 1903 : 43)- i My attention was drawn to this by Miss P. L. Cook. "BRITISH MARINE POLYZOA" INDEX 239 Chartella calveti (Guerin-Ganivet, 1911) [Prenant & Bobin, 1966 : 193]. Occurs in shallow water in the Brest-Roscoff area. Placed by Prenant & Bobin in Terminoflustra, but see the comment on this genus under 122 (p. 212). (?) Crassimarginatella tensa (Norman, 1903 : 595). Dredged from fjords around Bergen and in Hardangerfjorden, and described as Oochilina tensa. The correct generic placing is not clear, but Oochilina is a junior objective synonym of Crassimarginatella Canu (1900). Alderina aquitanica (Jullien, 1903) [Jullien & Calvet, 1903 : 45; Prenant & Bobin, 1966 : 214]. Off Brittany, St. 42. Described as Biflustra aquitanica, but Prenant & Bobin appear correct in referring the species to Alderina Norman. Callopora rylandi Bobin & Prenant, 1965 [Prenant & Bobin, 1966 : 231]. Description based on specimens from Roscoff. Material from the Channel Islands is present in the British Museum (Natural History) . Recently discovered in south-west Wales. Tegella arctica (d'Orbigny, 1853) [Kluge, 1962 : 280]. Faroe Islands (Kramp, 1934). Tegella armifera (Hincks, 1880) [Kluge, 1962 : 276; Prenant & Bobin, 1966 : 242]. Dredged to the east of the Orkney Islands, St. 1043 (Calvet, 1931). Cauloramphus cymbaeformis (Hincks, 1877) [Kluge, 1962 : 298]. Faroe Islands (Kramp, 1934). Chlidonia pyriformis (Bertoloni, 1810) [Levinsen, 1909 : 197, as C. cordieri]. Calvados (Normandy) (Busk, 1884 : 8, as C. cordieri). Doryporella reticulata Ryland, 1963 [Ryland, 19636 : 9]. Bergen, where Korsfjorden meets the Atlantic, 75-180 m. Calpensia nobilis (Esper, 1796) [Prenant & Bobin, 1966 : 337]. Guernsey (Norman, 1907, as Micropora impressa} ; La Chapelle Bank (Guerin- Ganivet, igna, as M. impressa). For taxonomic discussion see under 174 (p. 216). Scrupocellaria elongata (Smitt, 1868) [Kluge, 1962 : 382, as S. arctica; Prenant & Bobin, 1966 : 409]. The var. congesta Norman (1903) was found in shallow water near Bergen (Ryland 19636 : 12, with a note on the name of the species). Scrupocellaria grimaldii Jullien, 1903 [Jullien & Calvet, 1903 : 34; Prenant & Bobin, 1966 : 405]. Off Brittany, St. 42 and 85. It seems almost certain that this is actually 5. inermis Norman (Hincks, 1880 : 46), but slight doubt remains as Jullien's illustration appears to indicate the presence of a single axillary vibraculum. Scrupocellaria intermedia Norman, 1893 [Kluge, 1962 : 378 ; Prenant & Bobin, 1966 : The southernmost Norwegian record is from Trondheimsfjorden (Norman, 1893), but the species was identified by Nichols (1911 : 9) from St. 151, 353 and 477. Notoplites harmeri Ryland, 1963 [Ryland, 19636 : 15]. Bergen, where Korsfjorden meets the Atlantic, 75-180 m. Notoplites smitti (Norman, 1868) [Kluge, 1962 : 371]. 240 J. S. RYLAND A High Arctic species recorded, presumably in error, by Nichols (1911). How- ever, Prenant & Bobin (1966 : 446) can hardly be correct in stating that Notoplites jeffreysii is the species intended, since Nichols observed a fork-shaped scutum. Nordgaardia pusilla (Nordgaard, 1907). " Michael Sars " St. 76 (Nordgaard, 1907 : 5). Described as Synnotumpusillum, but Kluge (1962 : 362) created for it the new genus Nordgaardia. Dendrobeania elongata (Nordgaard, 1906) [Kluge, 1962 : 345, as Bugula; Prenant & Bobin, 1966 : 479]. (See addendum.) The original specimen came from Boknfjorden, Stavanger, 140-343 m., which is still the most southerly known locality. Dendrobeania fruticosa (Packard, 1863) [Kluge, 1962 : 331 ; Prenant & Bobin, 1966 : 476]. No British records were included by Hincks (1880 : 92), but the species has been recorded off Cullercoats, Northumberland, by Roper (1913 : 38), and further east in the northern North Sea (" Michael Sars " St. 278, Nordgaard, 1907). Also Faroe Islands (Kramp, 1934). Var. quadridentata Loven in Smitt, 1868 [Kluge, 1962 : 333] was recorded earlier by Nordgaard (1900) off the west coast of Norway, St. 10 of the North-Atlantic Expedition, but the synonymy of this var. has become so confused with that of D. elongata (Nordgaard, 1906, q.v.), that it is impossible to be sure to which taxon this refers. Bugula fulva Ryland, 1960 [Ryland, 1960 : 78]. Widely distributed around the British Isles. The range of variation of B. fulva is now known (Ryland, 1962) to include the form originally described as B. aquilirostris. Bugula neritina (Linnaeus, 1758) [Ryland, 1960 : 74]. Widely distributed in ports along the south coast of Britain; presumably introduced by shipping. Bugula simplex Hincks, 1886 [Ryland, 1960 : 91]. Various localities in southern Britain (Ryland, 19580) . Bugula stolonifera Ryland, 1960 [Ryland, 1960 : 78]. Ports in south-west Britain. Kinetoskias smitti (Danielssen, 1868) [Marcus, 1940 : 194; Prenant & Bobin, 1966 : 545]- Recorded from the fjords of southern Norway (Nordgaard, 1906), the Skagerrak (Silen, 1936) and off the south-west of Ireland (Nichols, 1911, St. 506). Kinetoskias arborescens (Danielssen, 1868) [Prenant & Bobin, 1966 : 548; Smitt, iS68a : 292, 353, PI. XIX, 28-31, as Bugula umbella]. Taken by the Norwegian North-Atlantic Expedition, St. 2, from 1229 m. in Sognefjorden (Nordgaard, 1900). This is the most southerly record. Erymophora gracilis (Nichols, 1911) [Prenant & Bobin, 1966 : 563]. Recorded by Nichols (1911 17; St. 477) off the south-west of Ireland, as Brettia pellucida var. gracilis nov., later raised to specific level as the type of Erymo- phora gen. n. by Hastings (1943). (See addendum.) Gephyrotes nitidopunctata (Smitt, 1868) [Kluge, 1962 : 388; Prenant & Bobin, 1966 : 585]. "BRITISH MARINE POLYZOA" INDEX 241 Recorded from various fjords in the Bergen area (Nordgaard, 1906; Ryland, 19636). CHEILOSTOMATA (ASCOPHORA) Haplopoma planum Ryland, 1963 [Ryland, 19630 : 14]. This is the northernmost species of Haplopoma, until recently confused with H. impressum auctt., which has its southern limit in the Bergen area, Hardanger- fjorden, and the Shetland Islands. The " Michael Sars " record, St. 82 (Nordgaard, 1907), presumably belongs to this species. Hippothoa expansa Dawson [see 291, p. 224] var. transparens Roper, 1913. Northumberland. Almost certainly a growth form of Celleporella hyalina (Linnaeus) [271, p. 223], and not of H. expansa. Myriozoella plana (Dawson, 1859) [Kluge, 1962 : 478, as Schizoporella Crustacea (Smitt)]. (See addendum.) Myriozoum crustaceum Smitt is type-species by monotypy of Myriozoella Levinsen (1909). Faroe Islands (Kramp, 1934, as Myriozoum crustaceum). Schizomavella glebula (Jullien, 1903) [Jullien & Calvet, 1903 : 81]. St. 46. The suboral avicularium and type of ancestrula are characteristic of Schizomavella and not Schizoporella. Schizoporella longirostris Hincks, 1886. Collected recently at several localities among the Isles of Scilly, from 12-51 m. (P. J. Hayward, personal communication and in preparation). Schizoporella magnifica Hincks, 1886 [Ryland, 19686]. Recorded from two localities in southern Britain. (?) Schizoporella nomismata Jullien, 1903 [Jullien & Calvet, 1903 : 80]. Off Brittany, St. 42. Correct generic placing not clear. Smittina cervicornis (Pallas, 1766) [Gautier, 1962 : 204]. St. 15, La Chapelle Bank (Guerin-Ganivet, 19110). I agree with Cook (1968) that Smittina, rather than Porella, is the correct genus for this species. (?) Smittina decipiens Jullien, 1903 [Jullien & Calvet, 1903 : 101, 149]. Taken several times in the Bay of Biscay (Jullien & Calvet, 1903; Calvet, 1931), twice close to the coast of Brittany (Calvet, 1931 : St. 1535 and St. 2974 off Belle- He) and also from near the Orkney Islands (St. 1043). Correct generic placing not clear. Smittoidea grimaldii (Jullien, 1903) [Jullien & Calvet, 1903 : 99, as Smittia]. Off Brittany, St. 85. Referable to Smittoidea Osburn (1952). Porella patula (M. Sars, 1851) [Marcus, 1940 : 269]. Occurs in the Bergen area (Ryland, 19636) and extends into the Skagerrak and Kattegat (Silen, 1951). Nordgaard (1912) showed that Lepralia aperta Boeck was synonymous with Sars' species. Kluge (1962 : 452), however, employs Boeck's name. Porella proboscidea Hincks, 1888. Obtained in the North Sea by the " Michael Sars ", St. 277 (Nordgaard, 1907). There is some doubt as to which species this record actually refers. Kluge (1962) considered that Nordgaard's records of P. proboscidea belonged to P. smitti Kluge ZOOL. 17, 6 i6§ 242 J. S. RYLAND (1907), though Nordgaard (1918) maintained that he had identified the species correctly. Phoceana columnaris Jullien, 1903 [Jullien & Calvet, 1903 : 107, 154; Gautier, 1962 : 212]. Recorded by Jullien & Calvet from St. 44. Escharella ventricosa Hassall, 1842, var. peristomata Kluge, 1962 [Ryland, 19636 : 26]. Shallow fjords, near Bergen. The status of this taxon requires investigation. Hemicyclopora multispinata (Busk, 1861) . Recorded by Nichols (1911 : 21) from St. 151, 194 and 504. There is some doubt, however, as to whether this species (and not H. polita only) was obtained (Ryland, 19636 : 27, footnote). Mucronella canalifera Busk (1884) is placed by Cook (1968) under the synonymy of H. multispinata. Marguetta pulchra Jullien, 1903 [Jullien & Calvet, 1903 : 73]. St. 14, La Chapelle Bank (Guerin-Ganivet, 19110). Bryocryptella torquata (Jullien, 1903) [Jullien & Calvet, 1903 : 77, as Cryptella], St. 14, La Chapelle Bank (Guerin-Ganivet, 19110). Rhamphostomdla ovata (Smitt, 1868) [Kluge, 1962 : 540]. Faroe Islands (Kramp, 1934). Rhamphostomella plicata (Smitt, 1868) [Kluge, 1962 : 544]. Faroe Islands (Kramp, 1934). Phidolopora elongata (Smitt, 1868) [Kluge, 1962 : 527, as Retepora]. Radosund near Bergen (Nordgaard, 1906). Referred to Phidolopora Gabb & Horn by Hansen (1962). Sertella aquitanica Jullien, 1903 [Jullien & Calvet, 1903 : 60], St. 15, La Chapelle Bank (Guerin-Ganivet, 19110) . The same author records it from the Concarneau region, " ou cette espece n'est pas absolument rare ". Sertella grimaldii (Jullien, 1903) [Jullien & Calvet, 1903 : 62]. St. 44. Although Jullien (p. 56) did not place this species in Sertella gen. n., he stated that it was congeneric with Retepora cellulosa Smitt, which Harmer (1933) referred to Sertella (see next entry) . Sertella septentrionalis Harmer, 1933 [Kluge, 1962 : 526, as Retepora cellulosa; Gautier, 1962 : 233] . St. Kilda (Ryland, 19650). Name proposed by Harmer (1933 : 620) for Retepora cellulosa Smitt (19686) non Linnaeus (1758). Adeonellopsis distoma (Busk, 1859). St. 14, La Chapelle Bank (Guerin-Ganivet, 19110). Celleporina surcularis (Packard, 1863) [Kluge, 1962 : 553]. Norman (1907) may have been correct in supposing that his specimen came from the Shetland Islands, as C. surcularis has been recorded from the Faroe Islands (Kramp, 1934), though in Norway it is not known to occur south of Tromso (Nordgaard, 1918). TurUcellepora coronopus (Wood, 1844) [Gautier, 1962: 260, as Schismopora}. Recorded by Jullien & Calvet (1903) from St. 46 and 85. It should be noted that T. coronopus was not then being distinguished from T. tubigera Busk, and it is uncer- "BRITISH MARINE POLYZOA" INDEX 243 tain which species was intended (Lagaaij, 1952; Gautier, 1962). Dinard (Balavoine, 1958,1959). Turbicellepora coronopusoida (Calvet, 1931) [Calvet, 1931 : 119; Gautier, 1962 : 261, as Schismopom] . Included in species list of St. 1043 from near the Orkney Islands, though this station is not mentioned on p. 119. Possibly a mistake, since the species is other- wise only known from the Mediterranean (Gautier, 1962). Turbicellepora smitti (Kluge, 1962) [Ryland, 19636 : 34]. Recorded from the Bergen area (Ryland) and the Kattegat (Smitt, 18686), and from the Faroe-Shetland Channel, "Michael Sars" St. 63 and 64 (Nordgaard, 1907). Gemellipora eburnea Smitt, 1873 [Busk, 1884 : 5, PL XXXIV, i, as Pasythea; Osburn 1940 : 463, PL 9, fig. 73-4]. Recorded by Nichols (1911 : 23) St. 151 and 479. CTENOSTOMATA Alcyonidium candidum Ryland, 1963 [Ryland, 19636 : 36]. Found on stones and on dead Lophelia coral from depths exceeding 50 m. in fjords in the vicinity of Bergen. Alcyonidium proliferans Lacourt, 1951 [Prenant & Bobin, 1956 : 208]. Described from the Dutch coast, but possibly only a form of A. gelatinosum (L.) (Hincks, i88o« : 491). Alcyonidium variegatum Prouho, 1892 [Prenant & Bobin, 1956 : 211]. For long only known from a single locality in the Mediterranean (Banyuls), but recently recorded from the Isle of Man (Eggleston, 1963). Arachnidium simplex Hincks, 1880 [Hincks, i88oc : 284; Prenant & Bobin, 1956: 26]. " 71 miles W by S from Fastnet, 315 fath." (Kirkpatrick, 1889 : 446). Immergentia suecica Silen, 1947 [Prenant & Bobin, 1956 : 230]. In dead Pecten shells, Gullmarfjorden, Sweden. (?) Immergentia sp. Borings in a shell bearing Hippothoa divaricata from Guernsey, 18-27 m. (Brit. Mus. 1897 .5.1. 792) appear to belong to a species of Immergentia (Hastings, personal communication). Tanganella muelleri (Kraepelin, 1887) [Braem, 1951 : 22]. Braem considered this controversial species sufficiently distinct from Victorella pavida Kent to create for it the new genus Tanganella. It was found in brackish waters near the Baltic coast of Germany. Bulbella abscondita Braem, 1951 [Braem, 1951 : 34]. Found by Braem (p. 44) amongst Victorella pavida Kent, from London. Hypophorella expansa Ehlers, 1876 [Prenant & Bobin, 1956 : 272]. Found in the tube lining of the polychaete Chaetopterus. First recorded in Britain by Harmer (1897), but now known from a number of localities in the British Isles, and as far north as Trondheim in Norway (Dons, 1939). Amathia pruvoti Calvet, 1911 [Prenant & Bobin, 1956 : 287]. Found in Chichester Harbour (Stubbings & Houghton, 1964). 244 J- s- RYLAND Amathia vidovici (Heller, 1867) [Prenant & Bobin, 1956 : 283]. Roscoff (Echalier & Prenant, 1951). (?) Bowerbankia pusilla Jullien, 1903 [Prenant & Bobin, 1956 : 308]. St. 85 (Jullien & Calvet, 1903 : 31). Penetrantia ditrupae (Norman, 1907) [Prenant & Bobin, 1956 : 324]. Found in the calcified tubes of the serpulid polychaete Ditrupa arietina dredged in the Shetland Islands. On the advice of Dr. Anna Hastings I have followed Silen's suggestion (1947 : 38) that Norman's Terebripora ditrupae may be a species of Penetrantia, though Marcus (1938 : 291) and Prenant & Bobin (1956 : 324) refer it to Spathipora. Obviously no placing can be final when based on borings only. Penetrantia concharum Silen, 1946 [Prenant & Bobin, 1956 : 330]. Living in the dead shells of many species of mollusc, from Gullmarfjorden, Sweden. Recorded from the Isle of Man (Bruce, Colman & Jones, 1963). Mimosella verticillata (Heller, 1867) [Prenant & Bobin, 1956, 264]. Roscoff (Echalier & Prenant, 1951). Mentioned by Hincks, p. 548, as Lagenella nutans Joliet. Monobryozoon ambulans Remane, 1936 [Prenant & Bobin, 1956 : 335]. Discovered in " Amphioxus sand " off Heligoland. Monobryozoon limicola Franzen, 1960. Described from a muddy substratum in Gullmarfjorden, Sweden. CYCLOSTOMATA Tubulipora aperta Harmer, 1898 [Harmer, 1898 : 101; Marcus, 1940 : 66; Kluge, 1962 : 96]. Bergen area and the Skagerrak. Tubulipora flabellaris (Fabricius, 1780) [Harmer, 1898 : 99; Marcus, 1940 : 60; Kluge, 1962 : 91]. The west of Scotland record (O'Donoghue, 1935) may or may not be an error, but there are records from the Skagerrak, Kattegat, Oresund, and from the Heligoland area. Tervia irregularis (Meneghini, 1847) [Busk, 1875 : 13, PI. 12, as Idmonea]. Appears to be common in the Bay of Biscay below 45° N, and was also recorded by Jullien & Calvet (1903) from St. 44, and by Nichols (1911 : 4) from St. 504. Bicrisia abyssicola Kluge, 1962 [Ryland, 19676 : 270]. St. Kilda (Ryland, 19650); fjords in the Bergen area (Ryland, 19676). Crisia constans Kluge, 1946 [Kluge, 1962 : 155; Ryland, 19676 : 273]. Recorded by Ryland from near Bergen, where Korsfjorden meets the Atlantic, 100-180 m. Crisia klugei Ryland, 1967 [Kluge, 1962 : 157, as C. denticulata var. borgi; Ryland, 19676 : 275]. The C. denticulata of writers on sub-arctic Polyzoa. St. 10 of the Norwegian North- Atlantic Expedition (Nordgaard, 1900) ; Bergen, where Korsfjorden meets the Atlantic (Ryland, 19676); the Skagerrak (Smitt, 1865). "BRITISH MARINE POLYZOA" INDEX 245 Crisia ramosa Harmer, 1891 [Harmer, 1891 : 134, 163]. This widely distributed British species was probably the C. denticulata var. a of Hincks (18800 : 422), but the diagnostic characters, especially of the gonozooid, were not described. Disporella venabulum Jullien, 1888 [Busk, 1859 : ri6> as Discoporella grignonensis ', Lagaaij, 1952 : 182, as Lichenopora]. Recorded (without further discussion) from the region of Dinard by Balavoine (1959 : 13), although his was the first published non-fossil record for this species. The identification was evidently made using Lagaaij (1952), who suspected (p. 182) that the Recent material of D. hispida in the British Museum (Natural History) might include specimens of the present species. (?) Lichenopora prolifica Jullien, 1903 [Jullien & Calvet, 1903 : 119]. St. 85. " Both from the description and the figure it seems clear to me that this is no Calyptrostegous species at all, but I am not prepared to say where it ought to be placed " (Borg, 1944 : 234). ENTOPROCTA Pedicellina hispida Ryland, 1965 [Ryland, 19656 : 200]. Plymouth and Finistere. Barentsia benedeni (Foettinger, 1886) [Prenant & Bobin, 1956 : 112]. Known from Hull (Ritchie, 1911) and Ostend. Possibly only a form of B. gracilis (M. Sars). Barentsia laxa Kirkpatrick, 1890 [Nielsen, 19640 : 57]. Kattegat and Oresund (Nielsen, 19640), Bergen fjords (Nielsen, 19646); Cullercoats (Eggleston & Bull, 1966). Barentsia mutabilis (Toriumi, 1951) [Ryland, 19610, as Pseudopedicellina]. Two localities in Pembrokeshire. Pseudopedicellina Toriumi was considered a synonym of Barentsia by Ryland (19656). Loxosoma agile Nielsen, 1964 [Nielsen, 19640 : 13]. On Nicomache trispinata, Kattegat and Oresund (Nielsen, 19640), and Bergen fjords (Nielsen, 19646). Loxosoma annelidicola (van Beneden & Hesse, 1864) [Prenant & Bobin, 1956 : 34]- On maldanid polychaetes from the French side of the English Channel; also from Holyhead (Ryland & Austin, 1960). Loxosoma claparedei Bobin & Prenant, 1953 [Prenant & Bobin, 1956 : 32]. On Notomastus latericeus, from Roscoff. Loxosoma loricatum Harmer, 1915 [Prenant & Bobin, 1956 : 39]. On Hermione hystrix, Roscoff. Loxosoma loxalina Assheton, 1912 [Prenant & Bobin, 1956 : 36]. In the tube of an unidentified maldanid polychaete, Sound of Mull, Scotland, Loxosoma monensis Eggleston, 1965. On unidentified worm tube, Isle of Man. 246 J. S. RYLAND Loxosoma pectinaricola Franzen, 1962. On Pectinaria belgica, Skagerrak to Oresund (Franzen, 1962; Nielsen, 19640) and Bergen fjords (Nielsen, 19646) ; Cullercoats (Eggleston & Bull, 1966). Loxosoma rhodinicola Franzen, 1962. On Rhodine loveni, Gullmarfjorden, Sweden. Recorded as Loxosoma sp. by Mclntyre (1961) on the same host in Loch Nevis and on the Fladen Ground (North Sea, off Aberdeen); also by Nielsen from the Skagerrak and Oresund (19640) and Bergen fjords (19646). Loxosoma saltans Assheton, 1912 [Prenant & Bobin, 1956 : 36]. In tube of the maldanid polychaete Leiochone clypeata (= Clymene ebiensis], Sound of Sleat, Skye. Loxosoma significant Nielsen, 1964 [Nielsen, 19640 : ii]. On the maldanid Asychis significant, Skagerrak to Oresund (Nielsen, 19640), and Bergen fjords (Nielsen, 19646). Loxosoma singulare Keferstein, 1862 (non Hincks, 18800) [Prenant & Bobin, 1956 : 29]. On Notomastus latericeus, on the French side of the English Channel. Loxomespilon perezi Bobin & Prenant, 1953 [Prenant & Bobin, 1956 : 40]. On Sthenelais boa, Roscoff ; Plymouth (Marine Biological Association, 1957) . Loxosomella alata (Barrois, 1877) [Prenant & Bobin, 1956 : 57, as Loxocalyx]. On sponge Dysidea, Barfleur. I follow Nielsen (19640) in uniting Loxocalyx with Loxosomella. Loxosomella antedonis Mortensen, 1911. Swansea docks (Ryland & Austin, 1960) . Loxosomella aripes Nielsen, 1964 [Nielsen, 19646 : 4]. On Laonice cinata, Bergen. Loxosomella arvyae Bobin & Prenant, 1953 [Prenant & Bobin, 1956 : 81]. On Phascolion strombi, Roscoff; Isle of Man (Eggleston, 1965). Loxosomella atkinsae Bobin & Prenant, 1953 [Prenant & Bobin, 1956 : 92]. On Phascolion strombi, Concarneau, Roscoff; Isle of Man (Eggleston, 1965); Kattegat (Nielsen, 19640). Nielsen found that L. bouxini was based on the contrac- ted state of L. atkinsae. Loxosomella bouxini Bobin & Prenant, 1953. See L. atkinsae. Loxosomella brucei Eggleston, 1965. Attached to the lining of an unidentified worm-tube, Isle of Man. Loxosomella compressa Nielsen & Ryland, 1961. On the setae of Lagisca extenuata, Bergen; on L. extenuata and Gattyana cirrosa, Kattegat (Nielsen, 19640) ; on the former, Isle of Man (Eggleston, 1965) and on both, Cullercoats (Eggleston & Bull, 1966). Loxosomella crassicauda (Salensky, 1877) [Prenant & Bobin, 1956 : 70]. Type-species by original designation of Loxosomella Mortensen, 1911. Plymouth (Atkins, 1932). Loxosomella cuenoti Bobin & Prenant, 1953 [Prenant & Bobin, 1956 : 94]. On Phascolion strombi, Concarneau. "BRITISH MARINE POLYZOA" INDEX 247 Loxosomella discopoda Nielsen & Ryland, 1961. On the ophiuroid Amphilepis norvegica, Bergen area; Skagerrak (Nielsen, 19640). Loxosomella elgans Nielsen, 1964 [Nielsen, 19640 : 18]. In tubes of Praxillella praetermissa and Rhodine gracilior from the Kattegat; common in the Bergen area, in tubes of Praxillella and Heterodymene robusta (Nielsen, 19646). Loxosomella fagei Bobin & Prenant, 1953 [Prenant & Bobin, 1956 : 95]. On Hermione hystrix, Roscoff . Loxosomella fungiformis Bobin & Prenant, 1953 [Prenant & Bobin, 1956 : 87]. On Phascolion strombi, Concarneau; Isle of Man (Eggleston, 1965). Doubtfully distinct from L. atkinsae Bobin & Prenant, of which it probably represents a highly contracted state. Loxosomella glandulif era Franzen, 1962. In tubes of the polychaete Panthalis oerstedi from the Kattegat (Franzen, 1962 ; Nielsen, 19640) . Loxosomella harmeri (Schultz, 1895) [Nielsen, 19640 : 28]. On Gattyana cirrosa, from the Kattegat and Oresund; on Gattyana and from Chaetopterus tubes, Cullercoats (Eggleston & Bull, 1966). Loxosomella kefersteinii (Claparede, 1867) [Prenant & Bobin, 1956 : 68]. Swansea docks (Ryland & Austin, 1960). Loxosomella marsypos Nielsen & Ryland, 1961. In the tube lining of Eunice pennata, Bergen ; Kattegat (Nielsen, 19640) ; in tubes of Chaetopterus variopedatus, Isle of Man (Eggleston, 1965). Loxosomella murmanica (Nilus, 1909) [Prenant & Bobin, 1956 : 72]. Nielsen (1967) considers that L. nitschei Prenant & Bobin (non Vigelius, see next entry) and L. murmanica, which both occur on Phascolion strombi, are identical. Widely distributed from the English Channel northwards. Loxosomella nitschei (Vigelius, 1882) [Ryland, 19616 : 35]. Cullercoats (Roper, 1913; Eggleston & Bull, 1966); also Bergen (Ryland, 19616). On various Polyzoa. This is not L. nitschei Arvy & Prenant (1952), Prenant & Bobin (1956) or Nielsen (19640) — see L. murmanica (above). Loxosomella nordgaardi Ryland, 1961 [Ryland, 19616 : 31]. Described from fjords in the neighbourhood of Bergen; Kattegat (Nielsen, 19640); Cullercoats (Eggleston & Bull, 1966). On various species of polyzoa. Loxosomella obesa (Atkins, 1932) [Prenant & Bobin, 1956 : 66]. Described from Plymouth; also recorded from Anglesey; the Isle of Man (Eggle- ston, 1965); and Bergen (Nielsen, 19646). On Aphrodita aculeata. Loxosomella ornata Nielsen, 1964 [Nielsen, 19640 : 24]. In the tubes of Thelepus cincinatus, from the Skagerrak to the Oresund. Loxosomella polita Nielsen, 1964 [Nielsen, 19640 : 22]. In tubes of Petaloproctus tennis and Nicomache lumbricalis from the Kattegat and Oresund; Bergen (Nielsen, 19646). Loxosomella scaur a Nielsen, 1964 [Nielsen, 19640 : 47]. On the setae of Nephtys spp., Skagerrak and Kattegat. 248 J. S. RYLAND Loxosomella similis Nielsen, 1964 [Nielsen, 19640 : 20]. In tubes of Nicomache trispinata from the Kattegat; Bergen (Nielsen, 19646). Loxosomella teissieri (Bobin & Prenant, 1953) [Prenant & Bobin, 1956 : 53, as Loxocalyx] . On the sponge Dysidea fragilis, Roscoff. I follow Nielsen (19640) in uniting Loxocalyx with Loxosomella. Loxosomella varians Nielsen 1964 [Nielsen, 19640 : 49], On Nephtys spp., Skagerrak to Oresund, very abundant; Bergen (Nielsen, 19646). 4. SUMMARY Each species described by Hincks in the " British Marine Polyzoa ", together with any varieties, is listed beside its appropriate page number. The name now recommended or generally accepted for each is then given, with author and date of introduction. If Hincks' name embraced more than one species, this is indicated. Where the present nomenclature differs from that of Hincks, brief documentation is given. In addition, 127 species which have been recorded since 1880 in European waters between latitudes 46° and 62° N are listed, together with the references docu- menting their discovery in the British area and providing an illustrated description. These supplementary records comprise fifty-seven cheilostomes, eighteen ctenostomes, nine cyclostomes and forty-three entoprocts. 5. REFERENCES ABILDGAARD, P. C. 1806. In O. F. Miiller: Zoologia Danica, 4 : 1-46. ADAMS, J. 1800. Descriptions of some marine animals found on the coast of Wales. Trans. Linn. Soc. Land. 5 : 7-13. AGASSIZ, L. 1846. Nomenclatoris Zoologici. Index universalis. Soloduri, 393 pp. ALDER, J. 1856. A catalogue of the zoophytes of Northumberland and Durham. Trans. Tyneside Nat. Fid Club, 3 : 93-162. (Reprinted with altered pagination 1857, F. & W. Dodsworth, Newcastle-on-Tyne, 72 pp.). — 1857. On a new species of Bugula. Q. Jl microsc. Sci. 5 : 174-175. 1864. Descriptions of new British Polyzoa, with remarks on some imperfectly known species. Q. Jl microsc. Sci., N.S. 4 : 95-109. ALLMAN, G. J. 1869. On Rhabdopleura, a new form of Polyzoa, from deep-sea dredging in Shetland. Q. Jl microsc. Sci., N.S. 9 : 57-63. ARVY, L. & PRENANT, M. 1952. Loxosoma nitschei Vigelius sur les Phascolions de Dinard. Bull. Lab. marit. Dinard, 36 : 2-5. ASSHETON, R. 1912. Loxosoma loxalina and Loxosoma saltans — Two new species. Q. Jl microsc. Sci. 58 : 117-143. ATKINS, D. 1932. The Loxosomatidae of the Plymouth area, including L. obesum sp. n. Q. Jl microsc. Sci. 75 : 322-391. AUDOUIN, V. 1826. Explication sommaire des Planches de Polypes de 1'figypte et de la Syrie. Description de I'Egypte, Histoire Naturelle, 1 : 225-244. (Planches 1-14 par J. C. Savigny) . 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Archs Zool. exp. gen. 6 : 305-357. WATERS, A. W. 1878. Bryozoa (Polyzoa) from the Pliocene of Bruccoli (Sicily) Manchr geol. Min. Soc. Trans. 14 : 465-488. 1879. On the Bryozoa (Polyzoa) of the Bay of Naples. Ann. Mag. nat. Hist. Ser. 5, 3 : 27-43, 114-126, 192-202, 267-281. — 1897. Notes on Bryozoa from Rapallo and other Mediterranean localities : chiefly Cellu- lariidae. /. Linn. Soc. (Zool.) 26 : 1-21. 1899. Bryozoa from Madeira. // R. microsc. Soc. 1899 : 6-16. 1900. Bryozoa from Franz-Josef Land, collected by the Jackson-Harms worth Expedition, 1896-1897. Pt. I. Cheilostomata. /. Linn. Soc. (Zool.} 28 : 43-105. 1909. Reports on the marine biology of the Sudanese Red Sea — XII. The Bryozoa. Part I. Cheilostomata. /. Linn. Soc. (Zool.) 31 : 123-181. 1910. Reports on the marine biology of the Sudanese Red Sea — XV. The Bryozoa. Part II. Cyclostomata, Ctenostomata, and Endoprocta. /. Linn. Soc. (Zool.) 31 : 231- 256. 1918. Some Mediterranean Bryozoa. Ann. Mag. nat. Hist. Ser. 9, 2 : 96-102. WOOD, S. V. 1844. Descriptive catalogue of the z lophytes of the Crag. Ann. Mag. nat. Hist. 13 : 10-21. 260 J.S.RYLAND 6. ADDENDUM Mature & Schopf (1968), in a publication received since the type of the present paper was set, have given details of type material of American Polyzoa described by Verrill & Dawson. The new information about four of these species affects the nomenclature adopted in this list, and is summarized below : (1) 291 Hippothoa expansa Dawson, 1859 [?• 224]- The species described by Hincks differs in a number of ways from Dawson's syntypes, and is to be regarded as a separate species needing a new name. (2) Dendrobeania elongata (Nordgaard, 1906). [Additional species, p. 240.] Nordgaard's species is recognized as being identical with Bugula decorata Verrill, so that the correct name is Dendrobeania decorata (Verrill, 1879). (3) Erymophora gracilis (Nichols, 1911). [Additional species, p. 240.] This species is recognized as identical with Bugulella fragilis Verrill (1879). As B. fragilis is the only valid genosyntype, it follows that Erymophora Hastings (1943) is a junior synonym of Bugulella Verrill. The correct name of the species is Bugulella fragilis Verrill, 1879. (4) Myriozoella plana (Dawson, 1859). [Additional species, p. 241.] Myriozoum crustaceum Smitt (18686) differs in several ways from Lepralia plana Dawson, and is to be regarded as a distinct species. The correct name for the present species, therefore, is Myriozoella Crustacea (Smitt, 1868). PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING THE CLUPEOID FISHES BY BLOCH AND SCHNEIDER P. J. P. WHITEHEAD BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 17 No. 7 LONDON: 1969 THE CLUPEOID FISHES DESCRIBED BY BLOCH AND SCHNEIDER BY P. J. P. WHITEHEAD Pp. 261-279; 3 Plates, 2 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 17 No. 7 LONDON: 1969 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in Jive 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. 17, 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) 1969 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 14 March, 1969 Price Eighteen Shillings THE CLUPEOID FISHES DESCRIBED BY BLOCH AND SCHNEIDER By P. J. P. WHITEHEAD INTRODUCTION THIS is the fourth in a series of studies (Whitehead, Boeseman & Wheeler, 1966; Whitehead, 1966, 1967) dealing with a fundamental problem in the systematics of clupeoid fishes — the problem of poor original descriptions relating to material that has rarely been re-examined. The clupeoid species described by Bloch and by Schneider are particularly important because they provide some of the oldest names still in use and because the types have not been redescribed in the modern era of clupeoid systematics, that is to say since Regan's pioneering studies of some 40 years ago. The Bloch & Schneider types have been in the Zoologisches Museum in Berlin for nearly 170 years, but damage during the 1939-45 war and subsequent curatorial difficulties in restoring the large collection to a workable state has led to the loss or misplacement of specimens. Thorough search produced 4 clupeoid types but one could not be found (Clupea malabaricus} and a neotype is recommended. Certain other specimens are commented on here since misidentifications have led to errors in the literature. Marcus Elieser Bloch (1723-99) published about a dozen ichthyological papers, but his major work was a i2-volume compilation entitled Allgemeine Naturgeschichte der Fische, 1782-95 (which was made up of the 3 parts of his Oeconomische Naturgeschichte der Fische Deutschlands, 1782-85 and the 9 parts of his Naturges- chichte der ausldndischen Fische, 1783-95; the 432 plates were numbered as a continuous series). A French translation of this work (by Laveau) appeared in 12 parts in 1785-88 and 1797, and a cheap edition by Castell in 10 volumes was published in 1801 in " Suites a Buffon " (volumes 32-41), with Bloch's plates copied and reduced by de Seve. After Bloch's death, Johann Gottlob Schneider (1750-1822) completed his colleague's work by compiling a synopsis of all described species under the title M. E. Blochii, . . . Sy sterna Ichthyologiae, iconibus ex ittustratum, published in 1801. In this work Schneider marked with an asterisk those species represented by specimens in the Museum Blochiana. Those not marked were thus unavailable to Bloch and the description either follows earlier authors or must be attributed to Schneider (who often wrote " Schn. " after his additions to Bloch descriptions). Bloch listed 12 species under the Linnaean genus Clupea (see Table i). All but one are clupeoids (8 Clupeidae, 3 Engraulidae but r Megalopidae) and three were new species (C. africana, C. nasus and C. malabaricus}. In addition to Bloch's 12 species, Schneider listed a further 10 and proposed a new clupeoid genus, Gnathobolus ( = Odontognathus Lacepede). Of Schneider's additional species, 4 are not clupeoids ZOOL. I7? 7. I7§ 264 P. J. P. WHITEHEAD TABLE i THE CLUPEOID FISHES OF BLOCK AND SCHNEIDER Bl. Page and volume number in Oecon. Nat. Fische Deutsch. or Nat. ausl. Fische. Schn. Page number in Syst. Ichth. Blochii. Plate. Plate numbers in Bloch's works (which were numbered consecutively). (new names in bold face) CHIROCENTRIDAE i. Clupea dentex Forssk. A. CLUPEOID SPECIES Bl. Schn. Plate — 428 — Identification Chirocentrus dorab (Forsskal, 1775) CLUPEIDAE 2. Clupea harengus L. 1 186 422 29 (i) 3. Clupea sprattus L. 1 206 423 29 (2) 4. Clupea pilchardus IX 40 425 406 5. Clupea trissa (i.e. thrissa L.) IX 35 — 424 4°4 6. Clupea alosa L. 1 209 432 30 (2) 7. Clupea sinensis L. IX 38 424 405 8. Glupea africana Bloch IX 45 425 407 9. Glupea melastoma Schn. 427 10. Gnathobolus [mucronatus] Lac. — 556 1 1 . Glupea nasus Bloch IX 116 426 429 (i) Clupea harengus Linnaeus, 1758 Sprattus sprattus (Linnaeus, 1758) Sardina pilchardus (Walbaum, 1792) Figure — Opisthonema oglinum (Le Sueur, 1817), text compo- site (see p. 266) Alosa alosa (Linnaeus, 1758) Hilsa kelee (Cuvier, 1829) Ilisha africana (Bloch, 1795) Ilisha melastoma (Schneider, 1801) Odontognathus mucronatus (Lacepede, 1801) Nematalosa nasus (Bloch, 1795) ENGRAULIDAE 12. Clupea encrasicolus L. I 212 13. Clupea atherinoides L. IX 46 14. Clupea baelama Forssk. 423 1 8. Clupea mystus L. 30 (2) 427 408 (i) — 429 — 15. Clupea malabaricus Bloch 1X115 425 432 16. Glupea mystax Schn. 426 (pi. 83) 17. Clupea setirostris Brouss. 428 428 Engraulis encrasicolus (Linnaeus, 1758) Pterengraulis atherinoides (Linnaeus, 1766) Thrissina baelama (Forsskal, 1775) Thryssa malabarica (Bloch, 1795) Thryssa mystax (Schneider, 1801) Thryssa setirostris (Broussonet, 1782) Coilia mystus (Linnaeus, 1758) B. NON-CLUPEOID SPECIES ALBULIDAE 19. Glupea brasiliensis Marc. — 427 — Albula vulpes (Linnaeus, 1758) CLUPEOID FISHES OF BLOCH AND SCHNEIDER 265 Bl. Schn. Plate MEGALOPIDAE 20. Clupea thrissoides Schn. 424 21. Clupea cyprinoides Brouss. IX 32 403 NOTOPTERIDAE 22. Clupea synura Schn. DOUBTFUL 23. Clupea brunnichii Schn. 426 424 — Identification Megalops cyprinoid&s (Broussonet 1782) M. cyprinoides Notopterus notopterus (Pallas, 1769) ? (see p. 274) (Table i) and of the remainder (i Chirocentridae, 2 Clupeidae and 4 Engraulidae) 2 were described as new (C. melastoma and C. mystax). The older jars in Berlin usually bear small rectangular labels about 50 mm. long. The labels fall into 3 groups whose relative ages can be inferred from instances where a second or third label was pasted over an older one (as in the case of Clupea melastoma — see p. 270). Type a label. Plain white paper, less than 25 mm. broad, without border, writing in flowing script. It is not certain if these date from Bloch (see under Clupea melastoma, p. 270). Type b label. Thicker paper, about 25 X 50 mm., with characteristic printed border of 2 black lines thickened along the lower and right edges of the outer border (and reverse for inner border) to give inset effect. Species name, locality and " Bl. " are printed by hand in heavy Gothic script. Register numbers appear to have been added later. The labels are sometimes coloured green, blue, yellow, etc., possibly in accordance with geographical areas. Type c label. Similar to the above but the outer border thick on all 4 sides and the inner border thin. Information on label in fine flowing script. A label bearing the name Chatoessus shows that these were in use after Cuvier's proposal of that name in 1829. Register numbers on these labels appear to be contem- porary with the label. The labels are sometimes coloured. Some of the jars also bear a small, square label (about 10 X 10 mm.) with a number written in heavy strokes as in type b labels. The numbers occasionally match the species numbers used by Bloch (or by Schneider) but this appears to be coincidence. The specimens preserved in alcohol are in good condition although often rather flaccid. The dry specimens, of which many seem to have been lost, are in poor condition. The specimens are listed in a catalogue dated 1860; the entries were probably copied from an older catalogue which no longer exists. ACKNOWLEDGEMENTS I am indebted to Dr. Kurt Deckert for permission to examine the collection in Berlin and for facilities provided; Dr. Christine Karrer greatly assisted in the search for specimens and literature. 266 P. J. P. WHITEHEAD THE CLUPEOID FISHES OF BLOCK AND SCHNEIDER GHIROCENTRIDAE i. Clupea dentex = Chirocentrus dorab (Forsskal, 1775) Clupea dorab Forsskal, 1775, Descript. Animal. : xii, 72. Clupea dentex Schneider, 1801, Syst. Ichth. Block. : 428 (Red Sea). Schneider merely proposed a new name for Forsskal's species. CLUPEIDAE 2. Clupea harengus = Clupea harengus Linnaeus, 1758 Clupea harengus Linnaeus, 1758, Syst. Nat., loth ed. : 317; Bloch, 1782, Oecon. Naturg. Fische Deutsch., 1 : 186, pi. 29 (i); Schneider, 1801, Syst. Ichth. Bloch. : 422. 3. Clupea sprattus = Sprattus sprattus (Linnaeus, 1758) Clupea sprattus Linnaeus, 1758, Syst. Nat., loth edition : 318; Bloch, 1782, Oecon. Naturg. Fische Deutsch., 1 : 206, pi. 29 (2); Schneider, 1801, Syst. Ichth. Bloch. : 423. 4. Clupea pilchardus — Sardina pilchardus (Walbaum, 1792) Clupea pilchardus Walbaum, 1792, in Artedi, Gen. Pise. : 38; Bloch, 1795, Naturg. ausl. Fische, 9 : 40, pi. 406; Schneider, 1801, Syst. Ichth. Bloch. : 425. 5. Clupea trissa (or thrissa) (PI. la) = Opisthonema oglinum (Le Sueur, 1817) [Halex festucosus — Plumier MS name on drawing, see below] . [Clupea minor, radio ultimo pinnae dorsalis longissimo — Brown, 1756, Civil Nat. Hist. Jamaica : 4431- [Cailleu-Tassart des Antilles Duhamel, 1772, Traite General Pesches., 3 (2) : 548, pi. 31 (3) — on Barbotteau], Clupea trissa Bloch, 1795, Naturg. ausl. Fische, 9 : 35 (emendation of thrissa). Clupea thrissa (non Linnaeus, 1758) Bloch, idem, 1795, pi. 404; Schneider, 1801, Syst. Ichth. Bloch. : 424. Megalops oglina Le Sueur, 1817, /. Acad. nat. Sci. Philad., 1 : 359. Bloch's plate 404 is a copy of a drawing by Father Plumier (see Plate ib), almost certainly the one entitled Halex festucosus now in the Bibliotheque Centrale in Paris (MS. 24, pi. 4). Bloch based his description entirely on the drawing, which shows the Western Atlantic Opisthonema oglinum. The name thrissa (trissa, triza, etc.) stems from the description of a Chinese gizzard shad (i.e. Clupanodon thrissa) by Osbeck (1757) which was first published as a valid name by Linnaeus (1758). Unfortunately Linnaeus and many subsequent authors CLUPEOID FISHES OF BLOCK AND SCHNEIDER 267 included the Western Atlantic Opisthonema in their descriptions or synonymies of the Chinese Clupanodon. Thus Linnaeus (1758) not only made reference to Brown's Clupea minor from Jamaica, but also identified a specimen of Opisthonema oglinum, sent by Charles Blagden from Carolina, as Clupea thrissa (Giinther, 1899 — specimen in Linnean Society collection, London). Considerable confusion arose because " thrissa " was claimed as type of both the Chinese Clupanodon of Lacepede and the American Opisthonema of Gill, the filamentous last dorsal ray being a feature common to the two genera. Although Bloch's description of Clupea trissa clearly refers to the American Opisthonema oglinum, Bloch mentioned the Chinese records of Osbeck and Linnaeus as well as the excellent description and figure of Clupanodon thrissa by Broussonet (1782). 6. Clupea alosa = Alosa alosa (Linnaeus, 1758) Clupea alosa Linnaeus, 1758, Syst. Nat., loth ed. : 318; Bloch, 1782, Oecon. Naturg. Fische Deutsch., 1 : 209, pi. 30 (i); Schneider, 1801, Syst. Ichth. Bloch. : 423. 7. Clupea sinensis (PI. ic) = Hilsa kelee (Cuvier, 1829) ? Clupea sinensis Linnaeus, 1758, Syst. Nat., loth ed. : 319; Bloch, 1795, Naturg. ausl. Fische, 9 : 38, pi. 405; Schneider, 1801, Syst. Ichth. Bloch. : 424. Clupea chinensis Cuvier, 1817, Regne Animal, ist ed. : 174 (on Bloch — omitted from 2nd ed.). Clupea kelee Cuvier, 1829, Regne Animal, 2nd ed., 2 : 320 (on Keelee of Russell). Clupeonia blochii Valenciennes, 1847, Hist. Nat. Poiss., 20 : 353 (on Bloch's Clupea sinensis). SPECIMENS. A skin, right side only, with label on reverse " Clupea sinensis ", mounted on stand on base of which is a label (without border) " Clupea sinensis Lin. M. Bl. Bl. t. 405 " and a smaller label " 98. " pasted below. The specimen has the catalogue number 21893. It is probably the one sent from Tranquebar by the Rev. John. Description. The specimen is in poor condition, but the following measurements were possible. S.L. 161 mm. ; tot. 1.205 nun.; body depth 69-3 mm. ; head length 51-9 mm. ; pectoral length 30-3 mm. ; pre-dorsal distance 80-2 mm. ; pre-pelvic distance 94-4 mm. IDENTIFICATION. The notched upper jaw, striated maxilla, presence of fronto- parietal striae and deep body identify the fish as Hilsa kelee (diagnosis in Whitehead, 1965 : 132). The characteristic black spots along the flanks, retained in some preserved specimens of H. kelee, are absent and all black markings appear to be dirt. The specimen was twice examined by Prof. Peters, first for Giinther and then for Day. Day (1878 : 641) believed the specimen to be Hilsa toli (Valenciennes), but Giinther (1868 : 446) identified it as Hilsa ilisha (Ham. Buch.) ; most of Giinther's ilisha material was, however, H. kelee (Whitehead, 1965 : 145). 268 P. J. p. WHITEHEAD There is a second and smaller specimen of Clupea sinensis in the collections in Berlin. This fish, 93 mm. S.L. and 125 mm. tot. 1., is preserved in alcohol. The jar bears a type c label (see Introduction, p. 265) " Clupea sinensis Bl.* Indischer Oz. 3846 A ^ 1.1. 50/0 [?] " and beneath this a type b label " C. sinensis Lin. 3.4.15 Ind. Oc. 3846 Bl." The significance of the asterisk is not now known. This juvenile fish has no striae along the maxilla, a barely developed cleithral lobe, few fronto-parietal striae, 18 + 13 scutes, pectoral fin tips just reaching to pelvic base, caudal lobes less than head length, the pseudobranch attenuated and with a groove below it, and an operculum twice as deep as broad. This clearly identifies it as Hilsa ilisha. Bloch (in the Castel edition of 1801) stated that he received Clupea sinensis from the Rev. John of Tranquebar under the name poiken or nannalai, a fact not given in the original edition of 1795 (in which Bloch records Japan and Tranquebar, as well as American coasts, as localities.) Although the juvenile specimen mentioned above bears a type b label, it seems unlikely that such a small fish was used for Bloch's figure or description. Bloch's figure of Clupea sinensis (pi. 405) formed the basis for Clupeonia blochii Valenciennes. Although the standard length of the larger specimen is 64mm. shorter than that of the figure, Bloch's drawings were not necessarily life size (the pilchard is 12 inches long, for example) and the present specimen is quite likely model for the drawing and can be considered the type of Clupeonia blochii. Accord- ing to Day (1878), Prof. Peters also believed this fish to have been the basis of the drawing. Clupea sinensis Linnaeus has been discussed as a possible senior synonym of Hilsa toll, but no conclusion was reached (Whitehead, 1965 : 144). The Linnaean species might have been the same as Bloch's (i.e. Hilsa kelee) although Hong Kong appears to be at the limit of the range of that species (no record in the literature but four juveniles in British Museum collection). The name sinensis suffered various interpretations (e.g. Lacepede's Clupanodon sinensis — see Whitehead, 1967 : 57) and was finally abandoned until revived by Fowler (1941 : 631) as a senior synonym of Hilsa toll. The type is not in Uppsala (Lonnberg, 1896; Holm, 1957) nor in London (Gunther, 1899) and in the absence of better evidence of its identity than the original description, the name Clupea sinensis Linnaeus is best rejected as a nomen dubium. 8. Clupea africana (PI. 2a) = Ilisha africana (Bloch, 1795) Clupea africana Bloch, 1795, Naturg. ausl. Fische, 9 : 45, pi. 407 (Accra; Br. St. 5, D 18, P 16, V6, A 46); Schneider, 1801, Syst. Ichth. Bloch. : 425. TYPE. A specimen in alcohol in a jar bearing a green type b label " C. africana. Bl. Aethiop. M. 3874 Bl. " to which the name Pellona Iserti V. has been added at a later date. CLUPEOID FISHES OF BLOCK AND SCHNEIDER 269 DESCRIPTION. Holotype, a fish of 154-211101. S.L. (193 mm. tot. 1.), in fair con- dition but snout damaged, belly split below pectoral fin, back damaged behind head and scales shed, ZMB. 3874. Br. St. 6, Diii 13, P i 12, V i 5 (or i 6), A iii 43, g.r. 23, scutes 26+7 (or 8). In percentages of standard length: body depth 33-1, body width 5-2, head length 26-8; snout length 7-9, eye diameter 8-1, upper jaw length 13-2, lower jaw length 12-5; pectoral fin length 17-8 (damaged, estimated 19-1), pelvic fin length 3-0 (dam- aged, estimated 3-9), length of anal fin base 41-3; pre-dorsal distance 53-8, pre-pelvic distance 46-0, pre-anal distance 57-8. Body strongly compressed, its width 5 times in depth, the latter greater than head length; belly keeled, scutes trenchant, especially behind pelvic fins. Snout a little shorter than eye diameter. Lower jaw strongly projecting with 3 (right) and 5 (left) conical teeth on either side of the symphysis. About 10 small teeth on each pre-maxilla. Two supra-maxillae (anterior one missing on right side), the posterior with ventral bulge and tapering posterior tip which just fails to reach tip of maxilla; the latter blunt posteriorly and with a series of fine teeth along ventral margin. No hypo-maxilla present. Pseudobranch present, exposed, with 18 filaments the longest being | eye diameter. Gillrakers moderate, the longest £ eye diameter, the corresponding gill filaments half that length. Dorsal surface of head with 2 prominent longitudinal ridges on each side, the outer one interrupted behind eye, and a third short ridge above the temporal foramen. Dorsal fin origin nearer to snout than to caudal base by ^ eye diameter, in advance of vertical from anal origin by £ eye diameter. Pectoral tips reaching to beyond vertical from pelvic base ; base of first ray on a level with lower border of eye. Pelvic base 1 1 eye diameters in front of anal origin. The latter equidistant between eye centre and caudal base and below vertical from sixth branched dorsal ray. COLOUR. Upper i of body brown, remainder silvery; no marks on body or fins. IDENTIFICATION. This is the only member of the Pristigasterinae known from West Africa and the long anal fin and projecting lower jaw make it unmistakable. The types of the following nominal species have been examined and found to be conspecific (Whitehead, 1967 : 112). Pellona gabonica Dumeril, 1858 Pristigaster dolloi Boulenger, 1902 Ilisha melanota Derscheid, 1924 TYPE STATUS. The label on the jar (type b) suggests that this specimen was from the Museum Blochiana, although Schneider (1801 : 425) did not place an asterisk against it, possibly in error. Again, the locality given (Aethiopian Sea) is curious because Bloch (1795 : 45) specifically states Acara (i.e. Accra). But it seems preferable to regard these as errors and to recognize the specimen as the holotype of Clupea africana Bloch. Valenciennes (1847 : 307) may have based his description of Pellona iserti on specimens but this is not clear from the description ; there are no such specimens in 270 P. J. P. WHITEHEAD Paris (Whitehead, 1967 : 112). He cited Block's name and quoted Bloch's meristic figures but proposed a new name because of differences between his material (or drawing or notes by Dr. Isert) and Bloch's figure (profile of the back more convex but belly less so, etc.). Valenciennes' species is clearly the same as Bloch's, and since only a single West African species of Ilisha is known, no point would be served in providing a type for Pellona iserti. 9. Clupea melastoma = Ilisha melastoma (Schneider, 1801) Clupea melastoma Schneider, 1801, Syst. Ichth. Block. : 427 (Indian Ocean, near Coromandel; Br. St. 6, Dis, P 14, V6, A 34). Pellona micropus Valenciennes, 1847, Hist. Nat. Poiss., 20 : 320. Ilisha brachysoma Bleeker, 1852, Verh. Bat. Gen., 24 : 22. TYPE. A specimen in alcohol in a jar with a yellow type c label " Pellona mela- stoma Bl. Schn. Indisch. Oz 3842 Bloch ", beneath which is a type b label " C. melastoma Bl. S Ind. Oc. Bl. ", beneath which is yet a third label (type a] " Clupea melastoma (Kutta Wai Bl) Bl. " A very small square label on the jar is inscribed " 10 ". DESCRIPTION. Holotype, a fish of 118-9 mm. S.L. (154-9 mm. tc)t- !•)> caudal tips complete but tips of pelvics, pectorals and dorsal damaged, as also belly below pectorals, ZMB. 3842. Br. St. 6, D iv 13 (last ray double), P i 14, V i 5, A ii 35, g.r. 23, scutes 8 [+ 2] 8 + 9. In percentages of standard length: body depth 37-8, head length 28-2; snout length 6-8, eye diameter 10-0, length of upper jaw 13-6, length of lower jaw 14-0; pectoral fin length 18-7, pelvic fin length (tip broken) 4-9, length of anal base 33-8 ; pre-dorsal distance 49-2, pre-pelvic distance 48-1, pre-anal distance 65-1. Body strongly compressed, deeper than head length, belly sharply keeled. Snout a little shorter than eye diameter. Lower jaw strongly projecting, about 5 small conical teeth on either side of symphysis. Small granular teeth present on tongue. Pre-maxilla with a single row of small conical teeth giving way towards the centre of the upper jaw to a row of small papillae. Two supra-maxillae, the anterior 57 mm. long with narrow anterior shaft, the posterior with expanded portion 6-3 mm. long, with ventral bulge but tapering posteriorly to reach tip of maxilla (which extends to vertical from anterior margin of pupil) ; maxilla with minute teeth along lower edge. No hypo-maxilla. Pseudobranch present, exposed, with 21 filaments the organ about | eye diameter in both length and breadth. Dorsal surface of head with a continuous ridge on each side flanked by a second ridge (discontinuous behind eyes) and two smaller ridges on parietal region. Dorsal origin nearer to snout than to caudal base by | eye diameter, behind vertical from pelvic base by f eye diameter. Pectoral tips just reaching to pelvic base. The latter slightly nearer to pectoral base than to anal origin; no axillary scale. Anal origin under last dorsal ray. CLUPEOID FISHES OF BLOCH AND SCHNEIDER 271 Scales with 2 (anterior) to 4 (posterior) continuous vertical striae and 6 striae interrupted (anterior) or overlapping (posterior) in front of this ; posterior margin of scale slightly crenulated and bearing fine, short horizontal striae. IDENTIFICATION. Norman (1923), Fowler (1941) and Whitehead, Boeseman & Wheeler (1966 : 92) believed that Schneider's Clupea melastoma was a species of Pellona (i.e. Pellona ditchela Valenciennes), chiefly because of the low anal count (34 fide Schneider; 33-37 in Pellona and 37-52 in species of Ilisha). Schneider's statement that " margine ossium maxillarum ensiformium anterior e toto crenulato " was taken as further evidence for Pellona since it suggested the toothed hypo- maxilla present in Pellona but absent in Ilisha. In the list of corrigenda, however, Schneider substituted labialum for maxillarum, and the resemblance to Pellona is further diminished by the higher anal count found in the specimen (37 not 34). The identity of Clupea melastoma is problematical. In the key given by White- head et alii (1966 : 93) it lies between the deep-bodied Ilisha brachysoma (Bleeker) and the more slender /. megaloptera (Swainson). So also does the holotype of Ilisha micropus (Valenciennes) (Whitehead, 1967 : 116). These 3 nominal species may merely represent growth forms of a single widespread and variable species, in which case Schneider's Clupea melastoma should be included. The name melastoma has not been used for over 50 years as a senior synonym and under Rule 23 (b) it should be rejected as a nomen oblitum. This is unfortunate because the next oldest name, Swainson's megaloptera, lacks a type and the descrip- tion is poor. Use of the name melastoma would not conflict greatly with usuage and application will be made to retain it. 10. Gnathobolus [mucronatus] (PL 2b) = Odontognathus mucronatus (Lacepede, 1800) Odontognathus mucronatus Lacepede, 1800, Hist. Nat. Poiss., 2 : 220, 221, pi. 7 (2). Gnathobolus [mucronatus inferred] Schneider, 1801, Syst. Ichth. Block. : 556. Valenciennes (1848 : 87), in a particularly uncharitable passage, pointed out that had Lacepede not been so dominated by his own ideas of systematics and had he consulted his material instead of following Gmelin, he would have discovered that the curious toothed " horns " curving forward on either side of the mouth of his Odontognathus mucronatus were nothing else but the maxillae bent forwards. Schneider (1801), with only Lacepede's description and bizarre figure by de Seve (see PI. 2b), recognized the " horns " as pieces of the upper jaw and proposed what he felt to be a more appropriate name (Gnathobolus}. Cuvier (1829 : 321) acknow- ledged that the jaw position was an artifact, but continued to use the name Gnatho- bolus, as also did Valenciennes (loc. cit.). 272 P. J. P. WHITEHEAD ii. Clupea nasus (PI. 2C) = Nematalosa nasus (Bloch, 1795) Clupea nasus Bloch, 1795, Naturg. ausl. Fische, 9 : 116, pi. 429 (i) (Tranquebar ; Br. St. 4, Di6, Pi3, V6, A 20); Schneider, 1801, Syst. Ichth. Bloch. : 426. TYPE. A specimen in alcohol in a jar with a type c label " Chatoessus nasus Bl. Indisch. Oc. 3898 Bloch." and a small square label " 12 ". DESCRIPTION. Holotype, a fish of no-o mm. S.L. (147-1 mm. tot. 1.), in fair condition, pelvic tips broken and tenth post-pelvic scute damaged, ZMB. 3898. Probably sent by the Rev. John of Tranquebar. Br. St. 6, D iv 14, P i 15, V i 7, A iii 20, g.r. (numerous), scutes 17 + 12. In percentages of standard length: body depth 38-3, head length 26-8; snout length 6-3, eye diameter 8-0, upper jaw length 8-3, lower jaw length 9-7; pectoral fin length 22-4, pelvic fin length 10-3 (damaged, 10-9 estimated), dorsal filament length 36-6, length of anal base 22-1; pre-dorsal distance 46-4, pre-pelvic distance 46-6, pre-anal distance 72-0. Body compressed, belly keeled, especially behind pelvic base, body depth greater than head length. Snout shorter than eye diameter. Lower jaw inferior, edge of dentary flared outwards. Upper jaw short, maxilla reaching to vertical from anterior pupil border, hind end of maxilla rounded; a single (posterior) supra- maxilla with short anterior shaft, curving downwards posteriorly. No teeth in jaws. Pseudobranch present, exposed, with 22 filaments, the base of the organ ridged with a small groove below. Gillrakers fine, short, about | diameter of pupil, cor- responding gill filaments twice as long, the anterior and posterior hemibranchs equal. Cleithral lobe present but small. Lower edge of second suborbital in contact with upper margin of pre-operculum and reaching forward to articulation of lower jaw before rising at an angle of about 20° to the vertical to reach a point beneath the anterior pupil border. Dorsal fin with last ray filamentous, its length almost equal to body depth, dorsal origin nearer to snout than to caudal base by if eye diameter and before vertical from pelvic base by ^ eye diameter. Pectoral tips just reaching beyond pelvic base. Pelvic base below 2nd unbranched dorsal ray, nearer to pectoral base than to anal origin by f eye diameter. Anal origin nearer to pelvic base than to caudal base by f eye diameter. Scales (about 45 in lateral series) with one continuous and o (anterior) to 3 (pos- terior) interrupted striae, the scale border slightly crenulated and bearing short horizontal striae. IDENTIFICATION. Nematalosa nasus is distinguished from other members of this genus by the shape of the second suborbital (Whitehead, 1962). In other species the lower edge of this bone does not reach forward as far as the lower jaw articulation before rising at an angle of 45°, leaving an exposed trapezoidal space on the cheek (Text-fig. la). In N. nasus the lower edge of the 2nd suborbital reaches the lower CLUPEOID FISHES OF BLOCH AND SCHNEIDER 273 jaw articulation and then rises quite steeply (20° in the type, but often almost vertical), leaving little of the cheek exposed (Text-fig, ib). The species is otherwise very similar to TV. come (Richardson). FIG. i . Shape of 2nd sub-orbital (stippled) in species of Nematalosa. a. Nematalosa come, 128 mm. S.L., BMNH. 1844.2.21.69. b. Nematalosa nasus, 135 mm. S.L., BMNH. 1881.2.7.1880. ENGRAULIDAE 12. Clupea encrasicolus — Engraulis encrasicolus (Linnaeus, 1758) Clupea encrasicolus Linnaeus, 1758, Syst. Nat., loth ed. : 318; Bloch, 1782, Oeconom. Naturg. Fische Deutsch., 1 : 212, pi. 30 (2). Clupea encrasicholus : Schneider, 1801, Syst. Ichth. Bloch. : 423. Variations in the spelling ot the species name still persist. Linnaeus (1735) first used encrasicholus, but the spelling in the loth edition is definitive. 13. Clupea atherinoides (PI- 3a) = Pterengraulis atherinoides (Linnaeus, 1766) Clupea atherinoides Linnaeus, 1766, Syst. Nat., i2th ed. : 523 (Surinam); Bloch, 1795, Naturg. ausl. Fische, 9 : 46, pi. 408 (i) (Surinam; Br. St. 12, Dn, Pi4, V 8, A 35); Schneider, 1801, Syst. Ichth. Bloch. : 427. SPECIMEN. A specimen in alcohol, 141-1 mm. S.L. (174-5 mm. tot. 1.), in a jar bearing a green label " Engraulis atherinoides L. Surinam 3838 Bloch ". The fish is in fair condition, rather soft, the silver lateral stripe visible but the body darkened and the scales black. 274 P. J. P. WHITEHEAD IDENTIFICATION. The specimen is clearly the South American P. atherinoides as described and figured by Hildebrand (1964); Bloch's description and figure may well have been based on the present specimen. Bloch, however, believed the Medi- terranean Clupea maxilla superiore longiore, fascia laterali argentea of Brunnich (1768 : 101) to be this species and also identified a Tranquebar specimen as Clupea atherinoides. The latter was most likely Stolephorus indicus (van Hasselt), while the former resembles Engraulis encrasicolus (the only Mediterranean anchovy species) except in its high anal count of 24 (about 17-19 in E. encrasicolus, including the three small unbranched rays) and low branchiostegal count (8, cf. 12). For this Brunnich species Schneider (1801 : 424) proposed the name Clupea brunnichii, but it cannot be positively identified with any known Adriatic clupeid (Alosa, Sprattus, Sardina and Sardinella have rather short maxillae quite unlike Engraulis, while an anal count of 24 would be exceptional even in Alosa). 14. Clupea baelama = Thrissina baelama (Forsskal, 1775) Clupea baelama Forsskal, 1775, Descript. Animal. : 72 (Djidda) ; Schneider, 1801, Syst. Ichth. Bloch. : 429. Bloch ignored this Forsskal species although he was well aware of Forsskal's work. 15. Clupea malabaricus (PI. 3b) = Thryssa malabarica (Bloch, 1795) Clupea malabaricus Bloch, 1795, Naturg. ausl. Fische, 9 : 115, pi. 432 (Tranquebar; Br. St. 8, D8, Pi4, Vy, A 38); Schneider, 1801, Syst. Ichth. Bloch. : 425. TYPE. Despite intensive search, no Bloch specimen of this species could be found, either in alcohol or in the dry collection. The genus Thryssa requires revision and the provision of types for those nominal species that have hitherto been known only by poor original descriptions would help to remove a great deal of existing confusion. A specimen has, therefore, been chosen which can stand as putative neotype pending a full revision. DESCRIPTION. Putative neotype, a ripe female of 158-2 mm. S.L. (197 mm. tot. 1.) in good condition except belly slit on left side, ex Madras (Day collection), BMNH. 1868.10.25.27. Br. St. 12, D I iii n, P i 12, V i 6, A iii 36, g.r. 14 + 17, scutes 15 + 9. In percentages of standard length: body depth 36-5, width of body 10-0, head length 24-9; snout length 5-1, eye diameter 6-0, length of upper jaw 24-7, length of lower jaw 17-6; dorsal fin height 21-3, pectoral fin length 18-8, pelvic fin length 9-2, height of anal fin (greatest) 12-2, length of anal fin base 33-0; pre-dorsal distance 51-7, pre-pelvic distance 43-9, pre-anal distance 65-8; depth of caudal peduncle 10-7. Body compressed, its width 3-6 times in its depth, head length less than body depth. Snout a little smaller than eye diameter. Mouth at an angle of about 15° CLUPEOID FISHES OF BLOCK AND SCHNEIDER 275 to the horizontal. Lower jaw with a single series of close-set teeth. Upper jaw with a single series of small teeth along entire edge; maxilla reaching gill opening and projecting 0-9 mm. beyond (10-3 mm. behind end of posterior supra-maxilla) ; the latter with slender anterior shaft expanding posteriorly to form a rounded ventral lobe 3-1 mm. long; anterior supra-maxilla round, minute (1-15 mm. long) (Text-fig. 2). FIG. 2. Thryssa malabarica, 158-2 mm. S.L., putative neotype, BMNH. 1868.10.25.27. Maxilla extends 0-9 mm. beyond gill opening. Pseudobranch present, concealed. Gillrakers short, the longest just over eye diameter, the corresponding gill filaments a little shorter (5-3 mm.) ; gillraker serrae not clumped ; 6 short gillrakers present on posterior face of 3rd epibranchial. Vomer with two clumps of teeth (left I, right 3) ; a line of granular teeth on edges of palatine and endo-pterygoid and patch of granular teeth on ecto-pterygoid. Sterno-hyoideus muscle reaching forward to beyond hind border of branchiostegal membrane. Posterior frontal fontanelles present, triangular. Venulose flap of skin behind operculum bearing about sixteen horizontal rows of dark brown dots. Dorsal preceded by scute-like plate bearing a small spine, base of fin invested in low scaly sheath; dorsal origin about equidistant between snout and caudal base and above vertical from pelvic tips. Pectoral tips reaching to pelvic base; axillary scale present, just over £ length of fin. Pelvic base nearer to pectoral base than to anal origin by just over i pupil diameter; axillary scale present, a little more than half length of fin. Anal origin f pupil diameter behind vertical from last dorsal ray, nearer to caudal base than to pectoral base by f pupil diameter; fin covered to half its height by scaly sheath. 276 P. J. P. WHITEHEAD Scales with exposed portion bearing reticulated striae; unexposed portion with up to 8 vertical striae, the anterior striae becoming interrupted in the centre of the scale in the more posterior scales. COLOUR. Upper ^ of body brown, remainder silver, fins yellowish. Dark brown speckled pigmentation on pectoral fins, maxilla (except second supra-maxilla, which shows up white) , margin of anal fin and venulose humeral region (a similar but much more intense pattern of pigmentation was described for Arabian Sea specimens (Whitehead, 1968) but in addition to the areas mentioned the dark pigments also occurred on the last unbranched dorsal and anal rays and on the margins of these fins and also on the caudal margin). IDENTIFICATION. The most recent key to the species of Thryssa (Whitehead, 1968) makes a major separation of species on the length of the maxilla, i.e. reaching to gill opening, to just beyond, to pectoral base, to pectoral tip or beyond. But species with a short maxilla failing to reach the pectoral base may yet prove to be mere ontogenetic forms of the long maxilla group (see below under Clupea mystax}. For Clupea malabaricus, Bloch's figure shows the maxilla just reaching the gill opening (see PL 3b) whereas in the specimen selected here the maxilla projects 0-9 mm. beyond. The difference is small and in fact is very much less than is found between individuals of other species (e.g. T. dussumieri and T. vitrirostris — White- head, 1968). Closely related to T. malabarica is T. kempii (Chaudhuri) of Lake Chilka in which the maxilla projects beyond the gill opening and about half-way to the pectoral base (photograph of holotype No. F. 8782/1 in the collection of the Zoological Survey of India kindly sent by Mr. M. Babu Rao). This species may prove to be T. malabarica. 16. Clupea mystax (PI. 3c) = Thryssa mystax (Schneider, 1801) Clupea mystax Schneider, 1801, Syst. Ichth. Block. : 426, pi. 83 (Malabar: Br. St. 5, 014, Pi4, V5, A 34 — the Br. St. clearly an error, as also the pelvic count). TYPE. A fish preserved in alcohol in a jar bearing a type b label " C. mystus Lin. Ind. Oc. 3884 Bl. " and a small square label " 16 ", matching an entry in the register " Engraulis mystus L. Ind. Ocean Bloch. " Schneider (1801 : 426) did not indicate a Bloch specimen (by means of an asterisk) for this species although one is noted for Clupea mystus (a Linnaean species of Coilia — see Whitehead, 1967 : 141). The present specimen so closely matches Schneider's figure, however, that it must be presumed the type. DESCRIPTION. Holotype, a fish of 134-7 mm- S.L. (160-7 mm- tot. 1. but caudal tips damaged, estimated ca. 165 mm.), in poor condition, especially broken caudal tips, belly (split behind pectoral), right maxilla (tip loose) and loss of scales, ex Malabar, ZMB. 3884. Br. St. 13, D I iii 12, P i 12, V i 6, A iii 33, g.r. n + 14, scutes 17 + 10. CLUPEOID FISHES OF BLOCK AND SCHNEIDER 277 In percentages of standard length : body depth 26-4, width of body 8-1, head length 25-7; snout length 4-9, eye diameter 5-9, length of upper jaw 30-4, length of lower jaw 19-6; length of pectoral fin 17-7, length of pelvic fin 8-8, length of anal base 30-2 ; pre-dorsal distance 53-0, pre-pectoral distance 27-2, pre-pelvic distance 41-8, pre-anal distance 63-4. Body compressed, its width 3^ times in its depth, belly sharply keeled, head length about equal to depth. Snout a little shorter than eye diameter. Lower jaw inferior, with a single series of fine conical teeth. Mouth slightly oblique, set at 15-20° from the horizontal. Upper jaw long and entirely toothed, maxilla projecting 3-7 mm. beyond pectoral base and 14-9 mm. beyond second supra-maxilla ; the latter with slender anterior shaft followed by an expanded portion 2-8 mm. long and about as deep; anterior supra-maxilla present, I mm. long. Pseudobranch present, concealed. Gillrakers short, the longest f eye diameter, the corresponding gill filaments | eye diameter ; gillraker serrae not clumped ; 7 small gillrakers on posterior face of 3rd epibranchial. Sterno-hyoideus muscle reaching forward to beyond hind border of branchiostegal membrane. Posterior frontal fontanelles present, triangular, 3-2 mm. long. Dorsal preceded by small scute-like plate bearing a spine, base of fin invested in low scaly sheath ; dorsal origin nearer to snout than to caudal base by | eye diameter, behind vertical from pelvic base by if eye diameters. Pectoral tip projecting 3-8 mm. beyond pelvic base; axillary scale present, half length of fin. Pelvic base nearer to pectoral base than to anal origin by if eye diameters; axillary scale present, almost equal to fin length. Anal origin nearer to pectoral base than to caudal base by £ eye diameter; base of fin invested in low scaly sheath. Scales with unexposed portion bearing i large and about 6 finer and more irregular striae the most anterior of which sometimes fail to meet in the centre of the scale; exposed portion becoming reticulated posteriorly. IDENTIFICATION. According to the most recent key (Whitehead, 1968), Thryssa mystax most closely resembles T. purava (Ham. Buch.) but differs in having a lower anal count (35-41; cf. 43-49), a higher number of pre-pelvic scutes (17-19; cf. 14-17) and a less oblique mouth. Schneider's description is poor, but his figure (shown here, PI. 3c) is adequate except that the maxilla is shown projecting too far beyond the pectoral base, the anal base is slightly too long and the origins of the dorsal, pelvic and anal fins are too advanced. Elsewhere (Whitehead, 1966 : 43) it was suggested that maxilla length may vary with size of fish and that larger fishes with shorter maxillae (C. hamiltonii of authors) may be C. mystax. 17. Clupea setirostris = Thryssa setirostris (Broussonet, 1782) Clupea setirostris Broussonet, 1782, Ichthyologia, i : no pagination (Tanna I., Pacific). Clupea setirostris: Schneider, 1801, Syst. Ichth. Block. : 428. This Broussonet species is not mentioned by Bloch, but Schneider cited the original description and quoted the then MS name of J. R. Forster, Clupea mystacina 278 P. J. P. WHITEHEAD (published some 40 years after Forster's death by Lichtenstein, 1844). The very long maxilla in this species, to beyond pelvic base, makes T. setirostris unmistakable. 18. Clupea mystus = Coilia mystus (Linnaeus, 1758) Clupea mystus Linnaeus, 1758, Syst. Nat., loth ed. : 319; Schneider, 1801, Syst. Ichth. Block. : 428. This was another species not included by Bloch. The synonymy of C. mystus has been discussed recently (Whitehead, 1967 : 149). REFERENCES BLOCK, M. E. 1782—85. Oeconomische Naturgeschichte der Fische Deutschlands. Berlin, 3 parts in i & Atlas (also issued as parts 1-3 of his Allgemeine Naturgeschichte der Fische}. J 783-95. Naturgeschichte der ausldndischen Fische. Berlin, 9 parts in 2 & Atlas (parts 4-12 of his Allgem. Naturg. Fische). 1785-88 and 1797. [Laveau translation] Ichthyologie, ou Histoire naturelle, general et particuliere des poissons. Berlin, 2 series. 1801. [Rene-Richard Castel translations] Histoire naturelles des poissons, avec les figures dessinees d'apres nature par Bloch, ouvrages classe par ordres, genres et especes, d'apres le systeme de Linne. Paris, 10 vols, an X (23 Nov. 1801-22 Sept. 1802). BROUSSONET, P. M. A. 1782. Ichthyologia, sistens piscium descriptiones et icones. Decas i, London 41 pp., n plates. BRUNNICH, M. T. 1768. Ichthyologia Massiliensis, sistens piscium descriptiones eorumque apud incolas nomina. Accedunt Spolia Maris Adriatici. Hafniae et Lipsiae, no pp. CUVIER, G. L. C. F. G. 1829. Regne Animal, 2nd ed., 2, Paris, 406 pp. DAY, F. 1878. The fishes of India, Williams & Norgate, London, 778 pp. FOWLER, H. W. 1941 . Contributions to the biology of the Philippine Archipelago and adjacent regions. Bull. U.S. natn. Mus., 13 (100) : 1-879. GUNTHER, A. K. L. G. 1868. Catalogue of the fishes in the British Museum, 7, London, 512 pp. 1899. President's anniversary address (on the Linnaean fishes in the Linnean Society collection). Proc. Linn. Soc. London, May 1899 : 15-38. HILDEBRAND, S. F. 1964. Engraulidae in Fishes of the Western North Atlantic, part 3. Sears Foundation for Marine Research, New Haven, Memoir i, 630 pp. HOLM, A. 1957. Specimina Linnaeana i Uppsala beverade Zoologiska Samlingar fran Linn6s tid. Uppsala Univ. Arsshrift (1957), No. 6 : 1-68 (English summary). LICHTENSTEIN, M. H. C. 1844. Descriptiones Animalium quae in itinere ad mavis australis terras per annos 1772, 1773 et 1774 suscepto collegit observavit et delineavit Joannes Reinoldus Forster, Berlin (on J. R. Forster's original MS from Capt. Cook's 2nd voyage — now in Staatsbibliothek, Berlin). LINNEAUS, C. 1735. Sy sterna naturae, sive, regna tria naturae systematice proposita per classes, ordines, genera et species, ist ed. Lugduni Batavorum, 7 pp. 1758. Systema naturae, sive, regna tria naturae systematice proposita per classes, ordines, genera et species. loth ed., 1, Holmiae, 824 pp. LONNBERG, E. 1896. Linnean type specimens of ... fishes in the R. University of Uppsala. Bihang. K. svenska Vet. Acad. Handlingar., 22 (4) : 3-45. NORMAN, J. R. 1923. A revision of the genus Ilisha and allied genera. Ann. Mag. nat. Hist. (9) 11 : 1-22. OSBECK, P. 1757. Dagbok ofver en Ostindisk Resa aren 1750-52. Stockholm, 376 pp. CLUPEOID FISHES OF BLOCH AND SCHNEIDER 279 SCHNEIDER, J. G. 1801. M. E. Blochii . . . Systema Ichthyologiae iconibus ex illustratum. Post obitum auctoris opus inchoatum absolvit, correxit, interpolavit Jo. Gottlob Schneider, saxo, 2, Berolini, 584 pp. VALENCIENNES, A. 1847. Histoire naturelles des poissons, 20, Paris, 472 pp. • 1848. Histoire naturelles des poissons, 21, Paris, 536 pp. WHITEHEAD, P. J. P. 1962. A review of the Indo-Pacific gizzard shad genera Nematalosa, Clupandon and Konosirus. Bull. Br. Mus. nat. Hist. (Zool.), 9 (2) : 87-102. — 1965. A preliminary revision of the Indo-Pacific Alosinae (Pisces : Clupeidae). Bull. Br. Mus. nat. Hist. (Zool.), 12 (4) : 115-156. — 1966. The elopoid and clupeoid fishes of Richardson's " Ichthyology of the seas of China and Japan " 1846. Bull. Br. Mus. nat. Hist. (Zool.), 14 (2) : 15-44. 1967. The clupeoid fishes described by Lacepede, Cuvier and Valenciennes. Idem, Suppl. 2 : 1-180. 1968. Indian Ocean anchovies collected by the Anton Bruun and Te Vega, 1963-4. /. mar. biol. Assn. India, 9 : 13-37. BOESEMAN, M. & WHEELER, A. C. 1966. The types of Bleeker's Indo-Pacific elopoid and clupeoid fishes. Zool. Verhandl. Leiden, No. 84 : 1-152. PLATE I a. Clupea thrissa from Bloch, pi. 404 (— Opisthonema oglinum — see p. 266). b. " Halex festucosus ". Plumier drawing on which the Bloch figure was based — see p. 266 (z= Opisthonema oglinum). c. Clupea sinensis from Bloch, pi. 405 (= Hilsa kelee — see p. 267). Bull. Br. Mus. nat. Hist. (Zool.) 17, 7 PLATE i , ZOOL. 17, 7. 18 PLATE 2 a. Clupea africana from Bloch, pi. 407 (= Ilisha africana — see p. 268). b. " Odontognathe aiguillonne " from Lacepede, vol. 2, pi. 7 (2), opp. p. 182 (= Odontognathus mucronatus — see p. 271). c. Clupea nasus from Bloch, pi. 429 (= Nematalosa nasus — see p. 272). Bull. Br. Mus. nat. Hist. (Zool.) 17, 7 PLATE 2 PLATE 3 a. Clupea atherinoides from Bloch, pi. 408 (i) (= Pterengraulis atherinoides — see p. 273). b. Clupea malabaricus from Bloch, pi. 432 (= Thryssa malabaricus — see p. 274). c. Clupea mystax from Schneider, pi. 83 (= Thryssa mystax — see p. 276). 12 MAR 1969 V, Bull. Br. Mus. nat. Hist. (Zool.) 17, 7 PLATE 3 a PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO MICHAEL D. B. BURT BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 17. No. 8 LONDON : 1969 CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO BY MICHAEL D. B. BURT Pp. 281-346 ; 60 Text- figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 17 No. 8 LONDON: 1969 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in Jive 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. 17, 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) 1969 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 29 April, 1969 Price £i. 125. - CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO By MICHAEL D. B. BURT CONTENTS ABSTRACT ..... INTRODUCTION ..... HOST-LIST ..... MORPHOLOGICAL DESCRIPTIONS . Family Anoplocephalidae Genus Paronia Diamare Family Davaineidae Genus Raillietina Fuhrmann Family Hymenolepididae Genus Hymenolepis Weinland Genus Fimbriaria Frolich . Family Dilepididae Genus Paricterotaenia Fuhrmann Genus Dilepis Weinland . Genus Liga Weinland Genus A nomotaenia Cohn . Genus Parvitaenia Burt Genus Vitta Burt Genus Kowalewskiella Baczynska Genus Ascometra Cholodkovsky Genus Notopentorchis Burt Family Acoleidae Fuhrmann Genus Gyrocoelia Fuhrmann ACKNOWLEDGMENTS REFERENCES Page 283 284 284 285 285 285 287 287 310 310 3i5 315 315 316 317 3i9 324 325 328 331 333 336 336 341 342 ABSTRACT Twenty seven species of cestodes, collected from 20 species of birds in Borneo, are identified and described in the following order : Paronia bocki Schmelz; Raillietina echinobothriaa (Megnin); R. johri Ortlepp; R. parviun- cinata Meggitt et Saw; R. sequens Tub. et Masil. ; R. allomyodes K6tlan; Raillietina sp. ; R. siamensis Schmelz; Hymenolepis mahonaenom. nov. ; Hymenolepis sp.; Fimbriaria fasciolaris (Pallas) ; Paricterotaenia burti Sandeman ; Dilepis ardeolae Singh ; Liga facile (Meggitt) ; A nomo- taenia depressa (Siebold) ; A . nymphaea (Schrank) ; A . tringae (Burt) ; Parvitaenia sp. ; Vitta rustica (TSTeslobinsky) ; Dilepid sp. ; Kowalewskiella susanae sp. nov.; Ascometra sp. ; Notopen- torchis collocaliae Burt; Gyrocoelia perversa Fuhrmann; Acoleid spp. (3). Of these, two are new species, one is recorded from Asia for the first time, and the synonymy and taxonomy of each species is discussed. The descriptions are as complete as the material allowed and are supported by 60 text figures and 12 tables. ZOOL. 17, 8. 19 284 M. D. B. BURT INTRODUCTION THE following cestodes form part of a collection of parasites collected from verte- brates by Dr. Robert E. Kuntz, United States Navy Medical Corps, in Borneo and sent to Dr. T. W. M. Cameron, Institute of Parasitology, Macdonald College, for identification. The species described are listed in systematic order. There are several new host-records and two new species are described. All drawings were made with the aid of a Wild drawing tube. Whole mounts were stained either in acetic acid alum-carmine or in Ehrlich's haematoxylin. Horizontal sections cut at 15 /*, and transverse sections cut at 5-10 /i, were stained in Ehrlich's haematoxylin and counterstained in eosin. Examination of rostellar hooks, sucker spines, and onco- sphere hooks was facilitated by using Berlese fluid. Material prepared as whole mounts has been deposited in the collections of the British Museum (Natural History) . HOST LIST APODIFORMES A pus affinis (J. E. Gray) ANSERIFORMES Anas boschas domestica L. CH A RA DRIIFORMES Actitis hypoleucos (L.) Charadrius leschenaultii Lesson Numenius phaeopus (L.) Tringa glareola L. CICONIIFORMES Butorides striatus (L.) COL UMBIFORMES Columba livia domestica L. Treron curvirostra (Gmelin) Treron vernans (L.) Streptopelia chinensis (Scopoli) CUCULIFORMES Cacomantis merulinus (Scopoli) *Centropus toulou (P.L.S. Miiller) Anomotaenia depressa (Siebold, 1836) Notopentorchis collocaliae Burt, 1938 Fimbriaria fasciolaris (Pallas, 1781) Liga facile (Meggitt, 1927) Gyrocoelia perversa Fuhrmann, 1899 Paricterotaenia burti Sandeman, 1959 Raillietina (Paroniella) siamensis Schmelz, 1941 (?) Species i Species 2 Species 3 Anomotaenia nymphaea (Schrank, 1790) Anomotaenia tringae (Burt, 1940) Kowalewskiella susanae n. sp. Raillietina (Raillietina) johri Ortlepp, 1938 (?) Dilepis ardeolae Singh, 1952 (?) Parvitaenia sp. Raillietina (Raillietina} sequens Tubangui et Masilufigan, 1937 Raillietina (Raillietina) sp. Raillietina (Raillietina) allomyodes K6tlan, 1921 Raillietina (Raillietina) johri Ortlepp, 1938 Raillietina (Raillietina) sequens Tubangui et Masilufigan, 1937 Dilepididae Ascometra sp. CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 285 GALLIFORMES Gallus gallus (L.) dom. Raillietina (Raillietina) echinobothrida (Megnin, 1881) Raillietina (Raillietina) parviuncinata Meggitt et Saw, 1924 PASSERIFORMES Aegithina tiphia (L.) Hymenolepis mahonae nom. nov. Raillietina (Raillietina] sequens Tubangui et Masilungan, 1937 Anthreptes malacensis (Scopoli) Hymenolepis sp. Hirundo rustica L. Vitta rustica (Neslobinsky, 1911) Nectarinia calcostetha Jardine Hymenolepis sp. PICIFORMES Megalaima chrysopogon (Temminck) Paronia bocki Schmelz, 1941 Raillietina (Paroniella) siamensis Schmelz, 1941 Meiglyptes tukki (Lesson) Raillietina (Paroniella) siamensis Schmelz, 1941 * Centropus toulou (P. L. S. Miiller) is not recorded from Borneo (Smythies: The Birds of Borneo, 1960) C. t. toulou (P. L. S. Miiller) is found in Madagascar, C. t. insularis Ridgway is confined to Aldabra Is. and C. t. assumptionis confined to Assumption Is. The Bornean Coucals are the Common Coucal, Centropus sinensis (Stephens), the Lesser Coucal C. bengalensis (Gmelin) which is also common, and the rare Short-toed Coucal, C. rectunguis Strickland, a rare bird known from four specimens. MORPHOLOGICAL DESCRIPTIONS Family ANOPLOCEPHALIDAE Cholodkovsky, 1902 Genus PARONIA Diamare, 1900 Paronia bocki Schmelz, 1941 . (Text-figs. 1-3) HOST: Megalaima chrysopogon 8891*. Only fragments of what appears to be one, incomplete worm are present and these total about 80 mm. in length. The maximum breadth is 2 mm. The proglottides are all broader than long with a tendency, in the more gravid proglottides, to become square. The scolex is missing. The testes are numerous (Text-fig, i) and develop before the ovaries. Over 150 testes were seen in one proglottis, but they are difficult to count accurately owing to the displacement of the genitalia by the unnaturally swollen excretory vessels. In some proglottides the testes attain a maximum diameter of about 70 /*, but are generally smaller averaging about 55 n in diameter. The cirrus-sac measures 400-600 /i in length, but may be longer as, in many instances, there are pronounced twists which tend to foreshorten it. The diameter of the cirrus-sac remains fairly constant at 40-55 /* and in some proglottides the cirrus can be seen protruding as a short nipple-like projection from the genital atrium (Text-fig. 2). *Host name and number according to collection records. 286 M. D. B. BURT The deeply-lobed ovaries are 380-520 fi by 200-240 {i and are fan-shaped, The vitelline glands, which measure 150-170 ju, by 110-125 /*» tend to be surrounded by the ovarian lobes (Text-fig. 3). The uterus breaks down into uterine capsules, each of which contains a single egg. Although initially the two uteri are quite separate, the capsules eventually fill the whole proglottis. The outer membrane of the egg measures 50-60 fi by 38- 48 fi and the inner membrane measures 28-34 /* m diameter. The hooks of the oncospheres were not seen sufficiently clearly to be measured. lOOjj FIGS. 1-3. Paronia bocki Schmelz, 1941. Early proglottis showing distribution of testes (Fig. i), opening of cirrus-sac (Fig. 2), and part of a later proglottis showing one of the two ovaries and the associated vitelline gland (Fig. 3). The excretory vessels appear to be unnaturally swollen and have the effect of crowding the genitalia into isolated regions in most proglottides. The transverse excretory vessel, which joins the two ventral excretory vessels posteriorly in each proglottis, is particularly prominent and swollen in fully-mature and gravid proglottides. DISCUSSION. Schmelz (1941) differentiates his species from Paronia carrinoi Diamare, 1900, on the formation of the uterus in gravid proglottides. In P. bocki, the two uteri eventually join to form a continuous field of uterine capsules, whereas in P. carrinoi both uteri remain separate. This feature, together with the difference in the size of the cirrus-sac, also serves to differentiate P. bocki from P. pycnonoti CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 287 Yamaguti, 1935. Although the number of testes in the present material may be fewer than the number described by Schmelz, and although there is a slight discrep- ancy in the sizes of the ovaries (380-520 /JL by 200-240 /* in the present material and 700 by 290 /* in Schmelz's material) in view of the close agreement of other characters, there seems little doubt that the present material should be ascribed to P. bocki Schmelz, 1941. Further differences between the known species of Paronia can be seen readily in Table I. P. carrinoi is listed by Yamaguti (1959) as P. carrinii and is included as a synonym of P. trichoglossi (Linstow, 1888). As indicated by Spasski (1951), Linstow, in his original description, mentions that the material is fragmented, without a scolex, and gives only the length and breadth of the fragments, the sizes of some proglottides, and the size of the eggs and oncospheres. Furthermore, Linstow indicates that " The specimens possibly belong to Taenia leptosoma, Diesing, found in various parrots ", and this worm, T. leptosoma, is now considered as belonging to the genus Raillietina. It would appear then that the grounds for including Diamare's species as a synonym of such an inadequately-described worm, are not really sufficient, and it is here proposed that the specific name carrinoi be retained and that it should not become a synonym of P. (?) trichoglossi. Family DAVAINEIDAE Fuhrmann, 1907 Genus RAILLIETINA Fuhrmann, 1920 Raillietina (Raillietina} echinobothrida (Megnin, 1881) (Text-figs. 4-8) Taenia bothrioplites Piana, Davainea paraechinobothrida Magalhaes, 1898. Davainea volzi Fuhrmann, 1905. Davainea penetrans Baczynska, 1914. Raillietina grobbeni Bohm, 1925. Raillietina pseudoechinobothrida Meggitt, 1926 HOST. Domestic fowl (Gallus gallus (L.) dom.) 8696. The longest specimen is 105 mm. long with a maximum breadth of about 3 mm. The proglottides are all much broader than long, the ratio of breadth to length tend- ing to increase from immature to mature proglottides and tending to decrease from mature to gravid proglottides. The genital apertures are unilateral and are situated laterally and slightly posteriorly to the middle of each proglottis. The genital ducts pass between the dorsal and ventral excretory vessels. The size of the scolex (Text-fig. 4) shows considerable variation, measuring 170- 265 (i long by 240-330 IJL broad. The four suckers are circular to oval in outline and measure 90-155 /* by 52-90 /*. They are profusely armed with spines (Text-fig. 5) which are 9-17 fi long. The rostellum, when everted, is roughly spherical with a diameter of about 100 fi. It is armed with a double circlet of about 200 hooks (Text-fig. 6) which are 12-14 fi long. ! .;: 'N £ New Guin £ 1 CO § IT 0 Australia; New Guin< Sumatra ; Bengal C T>, Formosa 1 rt ? ^ .9 3 5 S s g e * U co < I r^ttuopia Borneo 1 H •C • x oo 4J ^O 2 (j o 1 T o CO o o M f ' 3 3 CO t" O rj- B '3 CO 1 1 1 CO CO 1 N ^ *-& * v . ^? 1 I -a .g CM M CM 1 3 0 - •' •O O co fli ^ § 4) n> ' s ^ 1 _0 11 I bo .3 co 3 i be 2, 1 o "bb i _g 0 f ^ i • ^ 3 .3 5 en CO K g .S H-> O o o hH ^ || S 1 oo CO £ 00 X 10 ! 00 vO I o J w '§ Is S .3 * m si '» h O O o •n PQ *o CM CM >o O CM ^ rrt o o O t*N» CM M M Q OO o fe o H S S5 o M en 1 i CM O i ° i i 00 o M IS CO CM M M CD O « a CO ,£! g CM o o o o CD o J> O CM o Q in o O ( 5 v§ rtl S 5P -*H N 1 o j* N 1 0. 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' in t t !- ff . r i I 1 M £ " A N M M 3 w echino- bothrida rt •Q , rt 10 jj 10 120-250x4 250-550 100—140 CO o 7 o 4 I3O-2OO CO O i 1 M in r »n M in vO H A m M 1 ? co c t 2 t ;!M -1 l alternating &•) bo CO X CO t»» 5; ° 0. „ O N IO >n t 0 ^ o 0 at > « i Qi O SI o o vO VQ OO Oi & 1 A N oi, ^ r « in o CM f 1 1 o 10 7 I P M JS Ei| *Q in CO M M M M s oo >> bO pseudo- echino- bothrida bo M j|S> M X o 1 1 o 2 2 " N co 1 1 1 o 1 ! J "S-3 1*3 ^ Sofi 4) V fcJ ^ 00 HH «g 9. V CO OO ^ ( ^ "3 pQ I S to f^H N :O O^ X 1 ? % O M 10 M ? Tj- M *• H *> i*i sj O"» X O 00 A ^ A A 5 A J A oi 1 •<-> 10^ 8 M CO in oo o o M N •O CO < ^ *"* a ir a N N H M P rt f>. bo echino- bothrida Is N & 4) H S Tf X o in M 250-450 88-150 A * 0 0 N M 90-200 10 10 M M A ^ M rx. A O ro O •*• N A^ A m co M N H 00 ^5-4° 10-14 Irregularl; alternatin ,0 £ 1 •a o a ¥ — *» 0. 's: 1 'o V & C/j Describ •tH fl leter in /j) diameter in ^u) (number) length) in (i .meter in ft) (number of ro^ (length in p) (number) a _ n S. 2 a U *" 1 ^ i 1° -±- 0) number per ca diameter in p) s (diameter in rtures (arrang ^ £> OQ Scolex (diari Rostellum ( a ^ "a3 o Suckers (dia CO II en Sr CO 4) H 0 ^ w W " i 1 •% j 11 o c <§ (S 294 M.D.B.BURT 36-60 11. The cirrus-sac, 85-115 jn long by 40-60 fi in diameter, is roughly flask- shaped and has a thick muscular wall. In a few proglottides the profusely armed cirrus could be seen as a small bulbous projection from the genital atrium. There is an internal seminal vesicle with a diameter of 15-30 /*. The vas deferens is much coiled and lies parallel to the anterior margin of each proglottis. The bilobed ovary lies in the centre of the proglottis and measures 115-135 /^ across both lobes. The vitelline gland, situated posteriorly to the isthmus of the ovary, is compact and measures 45-75 /* by 30-55 /*. The shell-gland, 20-25 /* m diameter, lies more or less centrally over the isthmus of the ovary, occasionally being slightly displaced so that it lies partly over one of the lobes of the ovary. The vagina expands before reaching the centre of the proglottis to form a receptaculum seminis, 25-45 /i long by 15-25 fi in diameter. The opening of the vagina is posterior to the opening of the cirrus-sac. The uterus breaks down to form 18-30 capsules per proglottis and occasionally more, the size of the capsules varying from 50-80 /* in long diameter by 40-65 fi in short diameter. In some few capsules, which contained more than the normal number of eggs, the long diameter reached as much as 100 /* or more. There are 8-12 eggs per capsule usually, while some capsules contained as few as 6 and others as many as 16 eggs. The eggs measure 35-40 /i by 27-33 /* and the contained oncospheres are about 15 /* in diameter. DISCUSSION. Raillietina (Raillietina) polychalix Kdtlan, 1921 , was described by Johri (1934) irom. Psittacula krameri manillensis and from Columba lima domestica, although the worms from Columba differed quite markedly from those found in Psittacula. According to Johri, however, these differences are not sufficiently great to warrant separation of the worms and the erection of a new species, and accordingly the worms from both hosts were identified as polychalix. Ortlepp (1938), however, feels that the differences between the worms from Columba and those from Psittacula are too great to allow the inclusion of both groups of worms in the species polychalix and erects a new species, Raillietina (Raillietina) johri, to contain those worms which were found in Columba livia domestica. The two differences that Ortlepp considers to be most significant are the different sizes of the cirrus-sac and the different number of rostellar hooks (See Table III). TABLE III Raillietina (Raillietina} polychalix Kotlan, 1921, as described by Johri (1934). Number of Length of Host rostellar hooks cirrus sac Columba 324 125-1 30^ Psittacula 190 6i/j, Perhaps it should also be pointed out that the worms described by Johri that came from Psittacula show even less similarity to Kotlan's species polychalix than did the worms from Columba, and accordingly it would appear that neither of the worms which Johri described is in fact polychalix Kotlan. Proper identification of the worms from Psittacula will have to await a full re-examination of the material and comparison with known species. CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 295 The present material agrees well with the few characters that are given for Raillietina (Raillietina} johri and also seems to differ in one or more characters from all other species of Raillietina from birds, where the size of rostellar hooks is within the same range of 12-15 fi, and where the number of testes also falls within the same range of 6-12 testes per proglottis (See Table IV). As can be seen, the only species with which the present material does not differ significantly, apart from johri, is Raillietina (R.) circumcincta (Krabbe, 1869) but this worm is very poorly described and furthermore comes from a well-separated order of birds, the Ciconiiformes. According to the label in one of the tubes containing worms of the present species the host is Tringa glareola, a charadriiform bird, but as there does not appear to be any substantiated record of a Raillietina from any member of the Charadriiformes (see discussion following description of Kowalewskiella susanae n. sp.), it is not unlikely that there may have been a mix-up in the labels. All measurements of the present material from Tringa glareola are recorded separately, as follows, to facilitate comparison in the event of there being any further recorded Raillietina from a Charadriiform bird. Strobila — 41 X 1-15 mm.; scolex — 165 x 210 p; rostellum — 100 JLL diam. ; rostellar hooks — 200-250 hooks, in double circle, about 12 fi long (size measured from whole mount); suckers — armed, 65-70 /i in diameter; genital apertures — unilateral; testes — 6-1 1, 37-52 fi in diameter, on both sides of the female genitalia; cirrus-sac — 95-130 /* by 50-75 /* ; internal seminal vesicle present ; ovary-bilobed ; vitelline gland — median, posterior to ovary, ovoid; receptaculum seminis — 59;* x 45 /^; egg capsules — not extending beyond longitudinal excretory vessels, containing about seven eggs per capsule. Raillietina (Raillietina} parviuncinata Meggitt et Saw, 1924 (Text-figs. 11-13) HOST. Domestic fowl (Callus gallus (L.) dom.) 8696. The mature worm measures 35 mm. long by 0-9 mm. in maximum breadth. The proglottides are broader than long, but only immature and early mature ones are present. The genital apertures are unilateral and open in the anterior half of the margin of each proglottis. The scolex (Text-fig, n) measures 170/4 long by 240 /* in diameter and bears four, heavily armed suckers. The suckers are oval measuring 92-100 X 52-66 fi and the sucker spines, while being difficult to measure accurately, appear to reach up to 12/4 long. The rostellum is not evaginated and bears a complete double row of about 200 hooks which are 8-10 /* long. In the early mature proglottides present, the testes number 20 to 30 and are 38-46 x 34-43 /*. The cirrus-sac was not clearly seen, but appears to be about 70-80 jn long by about 40-50 fi in diameter. DISCUSSION. Although there are several discrepancies between the present material and those worms described by Meggitt and Po Saw (1924), it is apparent OJ -|J N .0 a.n, Ortlepp, Borneo I 1938 material OH OH 00 ^ ro JJ ON X in in o ro o in v X , rooo M""* w ^oinOMv M\O OO rtw OOwOwi °y i i ^ivl^ OO 0 vo IvOIOOOv O\O ^ oo * a ^ 1 ct * 1 5T f I To N i 'ro M ' « oo >n M M M ON M ro ro Ii O cM i 1 M i i-i 1 *o 2 0 0 i — > 0 O M 1 1* 1 1 AN 1 Z 1 -l-> CO M '$ .H -d £H a f^ o S rt tuO "e r3 1 JS +j -0 M N ON M M X in N OOOlN^i I1"1 O T ?OMOM' 'o "*" i-i N in ^" M O •&, in M a. yp '•S - C/} C4 N N M d) '§ « | % m ON M ON M S ON X o 0^&|u-,00 ^M" ^ x «n fli-ilMO0 MN ^ O ^ w O N 00 PI .a •** 1 '«• Baczynska, Tl- M ON M Baczynska Tt- — ON H 00 H X o I ^! ^ cT&g^" i^lloN 2 ° S^^^-o Io'£>tflf0 ° * o ."> ro 1 [e 0 ^_> ON | 0^ 00 in N 00 "tfl o i o ^ o i i-1 ^ i !r! s S •«* m ON fl ON X 00 0 1. ro T I N ' in * i-i o t^ f. -£> ^ •^ M ''j M o i-1 «O M *, $ 1 icracantha uhrmann, ON 0 ON M N ON M N M X o oo C^ V O «O • v ,^ OOO T(-AOOOO OO NMMininMM TfvOt^ i ^, | A.ti^ 4 g VAV 00 M M M 6 ^ ' M gL OT}- H n in in M •" S fc "9 M M s 1 « ^e c" g 00 ) species : micracar, Fuhrma: ON M Fuhrma g C^ M OXOOI "I 1 g 1 I M 1 1 1 M •?» •~» <3 1 *«* i ,D cfl cf 1 1 X o Mill M 1 1 1 Pn «n g S M % g (J H5 olumb ormes 1 ^- H M rj rj l|| *> - oo s s 3 t; c3* ken from : 1 £ a. d 5. •«H 1 cu S _c I-I (fl +•» ^ *. .5 t? -B a '^ *\*~ « fl l_l c^ Jn -+J ! "So j fi rt 2 fl OJ -M 1 || meter ii -4-> bo rt "a 3 _fl_ § .2 2- X'c3 (D fj ! 1 5, S, I .2 2. i tn 'S E ^ C/3 o ::: B ^i t^ ^3 rt ~z i > C!LU3 *^ ^ — •* co O> Q Strobila (1 n Scolex (dia Rostellum Rostellar j hooks Suckers (di Sucker spii 1 cn H Cirrus sac i C V bO w Oncospher Avian hosi CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 29? that in their description there are several typographical errors. The maximum breadth of the worm, for instance, is quoted as being 0-2 mm. and yet in the text- figure on page 325 a mature proglottis is drawn which, according to the scale given, measures over i-o mm. in breadth. The cirrus-sac in their description is described as being 0-58 to 0-84 mm. in length, whereas it is again clear from the drawing on page 325 that the length should be 0-058 to 0-084 nm]- FIGS. 11-13. 13 Raillietina (R.) parviuncinata Meggitt et Saw, 1924. Scolex (Fig. 11), mature proglottis (Fig. 12), and cirrus-sac (Fig. 13). In view of the close agreement between the length of rostellar hooks (7-9 fi — Meggitt and Po Saw; 8-10 fi — present material); the number of testes (24-39 — Meggitt and Po Saw ; 20-30 — present material) ; and the length of the cirrus-sac (58-84 ft — Meggitt and Po Saw ; 70-80 /* — present material) the present material is tentatively identified as Raillietina parviuncinata. This identification should remain tentative owing to the fact that parviuncinata was recorded initially from an anseriform bird, whereas the present material comes from a galliform bird. So far as can be ascertained, however, the present worms do not resemble any kndwn species from a galliform bird well enough to warrant identification with them. \ (, Raillietina (Raillietina) sequens Tubangui et Masilungan, 1937 > J (Text-figs. 14-18) HOSTS. Domestic pigeon (Columba lima domestica L.) 9019. Streptopelia chinensis 9409, 9151. Aegithina tiphia 9047. The largest worm measures 100 mm. in length and the maximum breadth is 1-5 mm. The proglottides are all broader than long, with the ratio of breadth 298 M. D. B. BURT to length generally increasing towards the posterior end of the worm. The genital apertures are unilateral and the genital ducts pass between the excretory vessels. The scolex (Text-fig. 14) varies in diameter from 80 to 122 /* and bears a rostellum which measures 45-90 /* (everted and withdrawn) in diameter by 30-53 /* long. The rostellar hooks (Text-fig. 15) are arranged in two separate rows ; those of the anterior row being slightly larger, measuring 7-5-8 /* in length while those of the posterior row measure only 6-7 /* in length. The four suckers measure 30-51 /* X 20-45 fi and are armed with spines (Text-fig. 16) which are 2-10 p long. 15 IQOM FIGS. 14-16. Raillietina (R.) sequens Tubangui et Masilungan, 1937. rostellar hooks (Fig. 15), and sucker spines (Fig. 16). 16 Scolex (Fig. 14), There are 6-10 testes (Text-fig. 17) one to three usually lying porally, which measure 45-60 /* x 40-55 fi. The cirrus-sac (Text-fig. 18) is 80-115 fi long x 40-60 fi in diameter and contains a slightly coiled cirrus, which is armed with hair- like, cuticular spines, and an internal seminal vesicle of 35-40 fi by about 30 fi. The vas deferens becomes greatly swollen with sperm and highly twisted. The ovary is bilobed with each of the two lobes tending to subdivide further into smaller lobules. It is situated medially and ventrally being contained within the confines of the ventral excretory vessels. The vitelline gland, lying immediately 18 FIGS. 17 and 18. Raillietina (R.) sequens Tubangui et Masilungan, 1937. Mature proglottis (dorsal view) (Fig. 17) and cirrus-sac (ventral view) (Fig. 18). CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 299 posteriorly to the ovary, is irregularly ovoid and measures 30-40 fi in diameter. Immediately in front of the vitelline gland, but posterior to the ovary, is the shell- gland of 20-25 /i in diameter. The receptaculum seminis is just posterior to the cirrus-sac and tends, in many cases, to lie slightly ventrally to it. It is highly variable in size, measuring up to 50 (i x 25 fi. The uterus, which arises immediately dorsally to the shell-gland, is initially a sac-like structure, but eventually occupies the whole of the medulla before breaking down into uterine capsules. The number of capsules per proglottis is variable, but generally falls between 50-70, each capsule measuring 50-100 fi x 35~55 /*. There are 2-8 oncospheres per capsule and these measure 25-40 p x 15-35 /*. The hooks of the oncospheres are small, ranging from 5-5 /i to about 7 fi in length. DISCUSSION. As can be seen in Table V there are several worms which have only a few testes and which also possess small rostellar hooks. In many instances the only difference between separate species is that one species may have one or two testes more, or fewer, than another. This hardly seems to be a justifiable criterion for the erection of new species, as it is abundantly clear in the present material that the range in number of testes is fairly wide. However, it should be noted that this range of 6-10 testes has not been seen in any single worm, but represents the total range in all the worms which otherwise are more or less identical from the different hosts mentioned. Thus, some individual worms show a range of 6-7 testes ; others a range of 6-8; others a range of 8-10 and so on. While this observation could be interpreted as indicating that there are two or more separate species present, in view of the extremely close similarity of other features, and the fact that there is no other constant difference manifest between worms showing differences in number of testes, it is here proposed that the range of 6-10 testes represents an intraspecific variation in the species R. sequens. Furthermore a re-examination of the species listed in Table V, and also many other species in the genus Raillietina, may very well show that the intraspecific variation is so great that several species which at present are considered as distinct should in fact be united into a single species. In view of the fact that it has not been possible, as yet, to see sufficient type-material, redescriptions from the suggested re-examination will not be presented at this time. Except in the number of testes, as has just been discussed, the present material appears to agree in all respects with worms described by Tubangui and Masilungan (1937) as Raillietina (R.) sequens and is, accordingly, here identified as that species. Raillietina (Raillietina) allomyodes Kdtlan, 1921 (Text-fig. 19) HOST. TreronvernansSg^i. One mature, but non-gravid, worm was present. It measures 15 mm. long and has a maximum breadth of 0-8 mm. The worm is highly contracted and accordingly the shape of the proglottides, which ranges from 24 times broader than long in imma- ture proglottides to n times broader than long in the last mature proglottides, is of little significance. The genital apertures are unilateral. X Til o o £ I 35 C •3 &_ M ^| M s~ I Jf ^ 4J r§ -r< M 'Sb ro 1 1 * ' I SI f I : w 111 **< •» on M •is1 III «^ ON N §11 u | g i II t^ M X o 1 1 2 I 1 1 \ N M 1 Unilateral Unilateral Unilateri Columbiformes Columbiformes Piciforme 0> CO 5 fa fa °efl -d Pn W CO « 8 Q g 0 0 X 8 rt g H -4-> en CD O §L ^ M O \J( H o X ON 00 1 O OO N 7 00 \ M 1 O M o >n I V" I 1 , bo P en oligocanth Fuhrman] 1909 Fuhrman (1909) X 0 1 in O ro O M ro Tf 7 O 00 M \ 1 1 x o 00 " I ro Irregularl alternatin rypturifon 'o O en 1 •!3 "^ ^ CO VD • bo M |l ^ ON rt ^ o X •^i- 1 1 90-100 2O-2I 1 \ IO-I2 1 - 1 1 ailateral mbiformes £* " in £ 2, *«* 1 6 CO g O E f ^ C fl 'a? •* 1 O I p Strobila (length X max. breadth in mi Scolex (diameter in ft) Rostellum (diameter in ft) IT '~ g to u |J3 1 § Suckers (diameter in ft) Sucker spines (length in ft) / (number) Testes < L (diameter in ft) Cirrus sac (length x diameter in ft) f (number/capsul \ (diameter in ft) Oncospheres (diameter in f Genital apertures (arrangement) Avian host (order) weissi weissi Borneo Joyeux, var. material 1923 valliclusa Joyeux & Baer, 1936 Joyeux Sawada (1923) (1965) 00 o X HI N X o u-> HI A •o HI 'o A ° u-) p. •o 0 >o HI HI O HI 1 150—300 200-250 210 i6&ig 20-25 i8&2i X t^ 1 OO >o o o HI ||? 11 C HI 00 HI X o O oo 7 o O 00 HI iO 8 ON S w •* o HI *0 M 4 1 M 0 « ° r °? r " oo PO o IN PO 1 u I-t-> i olumbi formes '$> £^" w N HI 0 M C P O Tf « M _ ON HI HI I 8 11 |.~-tfc "Q ON 1 PO o 1 o V 1 1 AN 1 x 1 1 1 5 % % ^ M ri j3 *->£- HI PO 1 1 W' 1 vO ^ t— * g-a •&,)—) fj 5 HI P S-" 'SB •** ON •B *rt to 4> « ;zj PO 11 tf lj! M X o Tl- , vO CO PO HI CO M 1 o N O TO I-1 I-1 \O I S V 1 1 1 I -4-> ""^ ' — o 3 O "^ O ^ CO M 3 O CO PO 1 P fi <£ PO 8 ON IIs 1 *'J3 If >— >-ii. o X CO o »o N 1 Tf 00 ON IS 1 1 HI M X 1 1 £ "° "rt 10 IH •P -2 f* -a ^ '3 Passeri- formes •&, £ TT M M N M H^ r" n ON o> t HI Tt- X TO • •4 S of £ .« >0 So°^ |l~ a 250-274 X 00 I PO PO HI N 8 7 M 142 XII. t- N OO ON N HI f^ » $• oo * M 1 1 X A & *° ?, Oi 17x14 Unilater Columb formes a •a $ rt rt "» HI •ON 0 B 9) a •«* rt M •S o o go X o O 8 T o HI 1 | | 1 S 1 g 5$ j-j W o Tf M 00 HI 1 1 1 | vl 1 M 7^3 JJ3 M s O 3 P 8 S 'o i & CO 1 ? Description ta Strobila (length x max. breadth in mi Scolex (diameter in fi) Rostellum (diameter in fi) ^ -^ PI (-1 "* 11 1^ 'v o P Suckers (diameter in fi) Sucker spines (length in //) / (number) Testes < V. (diameter in fi) Cirrus sac (length X diameter in fi) C /number/capsule \ (diameter in fi) Oncospheres (diameter in fi Genital apertures (arrangement) Avian host (order) 304 M.D.B.BURT the latter possesses only 12 testes. Looking at the variety weissi valliclusa, however, it is clear that the range quoted for this variety, of 12-15 testes, falls exactly between the ranges for the two species just mentioned. Furthermore, in view of the evidence presented in the discussion on Raillietina (R.} sequens it would appear that the number of testes can show a relatively wide range within a single species and accor- dingly not only would taiwanensis and weissi be synonymous, but, further, these would simply represent variations of Kotlan's species, allomyodes. Although it is not intended that these, or any others, be united into a single species at the present time, the possibility that a difference in host-species may affect differently the morpho- logical development of the infesting worms is obviously a strong possibility which warrants experimental verification. The present material differs in at least one respect from all the other species des- cribed, but in view of the close similarity of allomyodes, taiwanensis and weissi to each other and to the present worms, these are identified as belonging to that group all members of which should, on grounds of priority, be referred to the species allomyodes, Raillietina (Raillietina) sp. HOST. Treron curvirostra 9265. The strobila measures 50 mm. long by 1-15 mm. in maximum breadth. Nearly all the segments are much broader than long although the last few segments have a tendency to become square or slightly longer than broad. The genital apertures are unilateral and situated in the anterior half of each proglottis. The scolex is missing. There are 20-24 testes which measure 45-60 fi X 23-37 /* and which are situated in two groups on either side of the female genitalia. The group on the aporal side contains more than that on the poral side. The cirrus-sac is long and unusually thin, measuring 110-125 /* l°ng by only 15-25 fi in diameter. No seminal vesicle was seen and in none of the proglottides was the cirrus extruded. The vas deferens lies in large loose coils in the anterior, poral moiety of each proglottis. The ovary is bilobed, each of the two lobes being digitate, and is situated centrally in the proglottis. The vitelline gland, 37-45 /* X 15-23 /i, is compact and irregu- larly lobed, lying immediately behind the ovary. The uterus breaks down to form about 25 capsules per proglottis which measure 115-150 fi x 85-100 fi. In each capsule there are 6-9 eggs which measure 28-32 fi X 20-25 fi, while the contained oncospheres are about 14 p in diameter. DISCUSSION. Table VII contains those species of Raillietina (Raillietina) which : (a) parasitize birds; (b) possess a number of testes which falls in the range 20-24; and (c) possess a cirrus-sac the length of which falls in the range 100-150 /*. As can be seen from the table, the present material agrees reasonably well with several species, but does not agree in every respect with any single species. However, in view of the fact that the scolex is missing and in view of the reasonable similarity to several other species this worm has neither been given the status of a new species, nor has it been identified with any existing species. CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 305 Raillietina (Paroniella) siamensis Schmelz, 1941 (Text-figs. 20-24) HOSTS. Megalaema chrysopogon 8750, 8891, 9418; Meiglyptes tukki 9274; Charadrius leschenaultii 9486. The longest specimen is 35 mm. and the maximum breadth from any of the worms is 1-15 mm. The proglottides are all broader than long and in mature proglottides the breadth varies from two to four times the length. The genital apertures are unilateral and the genital ducts pass between the dorsal and ventral excretory canals. FIGS. 20-22. 20 Raillietina (Paroniella) siamensis Schmelz, 1941. Scolex (Fig. 20), rostellar hooks (Fig. 21), and sucker spines (Fig. 22). The scolex (Text-fig. 20) measures 490-525 fi across the region of the suckers and has a length of about 350 ju,. The rostellum is 174-215 JLI in diameter by 140-178 jn in length and bears a double crown of about 200-300 hammer-shaped hooks (Text-fig. 21). The hooks in the anterior row are larger than those of the posterior row; the former being about 28 fi long while the latter are only 19-23 fi long. There are four well-developed suckers, armed with spines (Text-fig. 22) of 8-18 /* in length. In some specimens the anterior part of the scolex could be seen to be covered with hair-like spines less than 3 /* long. This was easily seen where the rostellum was invaginated. There are 20-40 testes (Text-fig. 23) which lie in two separate fields ; the larger group lying aporally comprising 12-30 testes. The testes measure 40-90 fi X 37-68 /*. The cirrus-sac has a length of 90-130 /* and a maximum diameter of 60-74 /*• It has a thick wall of about 8 fi and opens into the genital atrium anteriorly to the opening of the vagina in the anterior half of the lateral margin. There is neither 1 .g *=a^- e . o O 4-> w S 'o % N ca ^_^ rt _M_ CO o o 240-280 •4- 10 M ro ro M "O r^ O •0 oo (O o 100-130 X 60-70 I 1 S T Unilateral .es Galliformes o •8* *" pH *" * N 0 ro g 0 0 o * M "O X Tf 1 g o M ^ ll 3 CD CO CO ^ C7» X •HT o o i— i A A M CO f H 1/1 IO o M {Q ^ | O i ig 1 o "* O T3 o o ~~^ vO CM (N M <0 * •S T 00 'S G M i— > 3 '"'K 3 c5 o M ,3 So ffi (H o o cij 00 t/5 CD 2 rf rt ^ O o OO "c3 i 1 3 fli B ^ C/5 *" xogi-ogi o CM 0 H o CM A OO M 2O-22 1C ir» * ll X O M M 1 \ I Unilater Columbifor $ § a" 10 S -S N B *o o Tf r Tt- 0 0 CM A O ir> £• CM T£ X t""1* ^" M 00 H f I ON ° 1 1 % brmes "I |,M H « i. r^ M CO 0 M ro M 5? N M 3 a G •M-H "2 M P 0 1-H 1 r™^ . tn PABLE \ N i CM CD be c" r* a ? rt O\ i§J *D£j M 3 G if IH O* •3 -^ 3 M M X o I8O-20O % M 8 ro • O »o ! O 7 0 00 v£> I2O-I4O o M ol, 1 I nilateral mbiforme r^ 9 1 & M P ^3 IH "o CD u ^ • VI CD -4-J ill •£ 'JK CO X O o O CO rmous mber CO j N o ro 1 1 O X 1 1 ^ <->.. liforme W "^ W M W ^' _j_ O 3 IO c CD G C CM 0 6 cj M o •^ O 1 en 1 a CO zruensis ihrmann, 1911 uhrmann (1911) N X O O co'o' 1 oo O ro O M CO II o 0 0 CM A 00 M CO M 001-06 1 O ^ si 0 IH 1 1 1 nilateral :taciforme Q PH PH P 'tn '•a fc Raillietina (Raillie 1 S. C/2 Description taken from : Strobila (length X max. breadth in mm. Scolex (diameter in ft) Rostellum (diameter in fi) "5 I 1 W) 1 i I* CD O -4-> O en ^ 0 -f K Suckers (diameter in ft) •*5 G 1 G _0 ¥ 1 c/) If *?<*> ^t* rt P IS a J3 "J "3 O 8S ^} O OO N O oo" OH ON rt M « 10 iO 10 u-> X 10 Tl- 00 O4 O ^" % "•> 8 1 ' 1 °° 1 2 r-- 2 o t^ ON ^ CD 'ON in o 04 si )-> § 'i J M. ^ i^ CO JH rt ON M X S- 1 $ <2 & 1 ^ | M £j ^ 1 1 1 •S to "3 V| ^ O rt -g M ^ O o N ' M N' ^ a fl § CO i-J C/2 t in iripendra Sharma, 1943 a ^ J3 ON N M X o in i o oo M Cl r^ O i N (M I i i 00 M 0 M VO » M IH in M M in 10 ON 8 ^ ' Jnilateral umbiforme n ON 17x14 Unilateral lumbiforme s 04 o O N en 1 «s s £ d E 3_ 5 ^ CO 5 3. C 1 _G C 5 3 _fl fl 'S; 1 .a X. Description ta Strobila (length X max. breadth Scolex (diameter in p\ Rostellum (diameter i Rostellar / (number) .2 c ^ en | Suckers (diameter in , Sucker spines (length f (number) Testes -I (diameter: Cirrus sac (length x diameter in r (number/c u u 1 .2 2- -v— ' en bo bx> W Oncospheres (diamett Genital apertures (arrangement) Avian host (order) 308 M. D. B. BURT an internal nor an external seminal vesicle, although in fully-mature and gravid proglottides, the vas deferens becomes greatly swollen with sperm and may act as an external seminal vesicle. In early proglottides the vas deferens tends to lie along the anterior edge of the segment, but, as it becomes filled with sperm, it comes to occupy most of the anterior poral quarter of each proglottis and lies in large loose coils. The ovary is fanlike and deeply lobed with a maximum breadth of about 300 /* and lies ventrally in a median position. The vitelline gland, lying immediately posteriorly and slightly porally to the ovary, is irregularly ovoid and measures 104-180 fi X 65-100 fi. The shell-gland, with a diameter of about 15 fi, lies between the ovary and the vitelline gland. On leaving the genital atrium, the vagina lies parallel to the anterior margin and widens out, before reaching the centre of each proglottis, to form a receptaculum seminis. 23 FIGS. 23 and 24. Raillietina (Paroniella) siamensis Schmelz, 1941. Mature proglottis (Fig. 23), and egg with contained oncosphere (Fig. 24). The uterus can first be seen as a transverse band across the anterior region of the proglottis, and it eventually breaks down to form uterine capsules, each of which contains but one egg. The size of the capsules is 34-38 /* X 30-32 fi ; the diameter of the contained eggs (Text-fig. 24) is 25-27 ju, x 22-24 P'> and the diameter of the embryos is 9-15 /*. The oncosphere hooks are small, measuring 7-9 ju, in length. DISCUSSION. As can be seen from Table VIII, the present material agrees closely with that described by Schmelz (1941) as Raillietina (Paroniella) siamensis and also with material described by Johnston (1914) as Davainea sphecotheridis. Schmelz separated his species from that of Johnston on the following grounds : (a) sphecotheridis is a parasite of Passeriformes whereas siamensis is found in Capitoniform.es ; (b) The scolex of sphecotheridis bears a great number of minute spines, particularly at the base of the rostellum, whereas siamensis does not have these; (c) there are fewer and smaller testes in sphecotheridis than in siamensis ; (d) the rostellar hooks are smaller in sphecotheridis, and there appear also to be differences in the length and breadth of the strobila, in the length of the cirrus- sac and in the diameter of the embryo. CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 3°9 TABLE VIII Comparison of Borneo material with Davainea sphecotheridis and Raillietina siamensis Species : Strobila: (length x max. breadth in mm.) Scolex (diameter x length in /*) Rostellum (diameter x length in /*) f~ (number) Rostellar I hooks | (length in ft) Suckers (diameter in n) Sucker spines (length in fj,) C (number) \ (diameter in /i) Cirrus sac (length X diameter in p) Ovary (shape and size in p) Vitelline gland (shape and size in ft) Egg capsule (diameter in p) Embryo (diameter in p) Testes Davainea sphecotheridis IOOX2 360 very great number ant. row : 20 post, row: 15 140 up to 10 ca. 30 25-30 Xi 5-20 100 X4O digitiform lobes solid and rounded 27x20 (egg) Raillietina siamensis 45X4-8-4-9 400-440 X 240 Borneo material 35XI-I5 490-525 X 360 130-150 174-215x85-150 240 200-300 ant. row : 24 ant. row: 28 post, row : 20 post, row: 19-23 200 174-215 x 140-178 — 8-1 8 50-60 20-40 80x60 40-90 x 37-68 1 50-1 60 x 66-68 90-1 30 x 60-74 495 145 30-36 12-14 fanlike, deeply lobed ; 300 irregularly ovoid 104-180 X 65-100 34-38 X 30-32 9-15 However, it is clear from the present material that some of these characters which purportedly separate these species are, in fact, common to both. The minute hooks described by Johnston on the scolex of sphecotheridis are abundantly clear in the present material but only when the rostellum is not evaginated, which would suggest that they are extremely caducous and which might well explain why Schmelz did not see any such spines, although he was looking for them, in his material. The number of testes in the present material seems to fall half-way between the number described for sphecotheridis and the number described for siamensis, and it is quite possible that the number of testes is a variable character within the limits quoted. The difference in the sizes of rostellar hooks seems hardly sufficient for this not to be due to measuring technique, particularly in view of the fact that when the hooks of the present material were measured from whole mounts in Canada balsam the lengths were considerably lower than those actually recorded from squash preparations in Berlese fluid. In view of the number of small discrepancies present between the two species, it may be that they are in fact separate species, but the present material suggests that there is a range of variation which may well encompass both described species. However, without examining the type-material of sphecotheridis and that of siamensis, it would not be wise to make these two species synonymous and accordingly the 310 M.D.B.BURT present material is tentatively identified as Raillietina (Paroniella) siamensis, as it seems to fit Schmelz's description slightly better than that of Johnston, but it is strongly suggested here that siamensis is a synonym of sphecotheridis. It is probable that there is a mistake in labelling in some of the present material as a Charadriiform bird, namely Charadrius leschenaultii, is apparently a host for this species, whereas this order of birds has not previously been shown to carry species of Raillietina (see discussions following descriptions of Raillietina (R.) johri Ortlepp, 1938 and Kowalewskiella susanae n. sp.). Furthermore, no differences could be found between those worms supposedly from Charadrius leschenaultii and those worms from the piciform hosts. Family HYMENOLEPIDIDAE Fuhrmann, 1907 Genus HYMENOLEPIS Weinland, 1858 Hymenolepis mahonae nom. nov. (Text-figs. 25-28) Hymenolepis fringillarum of Mahon, 1958, nee Rudolphi, 1810. HOST. A egithina tiphia 9045 . There were several worms present only two of which, however, possessed scolices. Of these two, only one (Text-fig. 25) bore rostellar hooks. The longest complete worm measures 7 mm. and has a maximum diameter of 0-5 mm. There are fragments present, however, where the maximum breadth is almost 0-8 mm. The proglottides are all broader than long, the ratio of breadth to length varying from 3 : i to 8 : i depending on the part of the worm. The genital apertures are unilateral and the genital ducts pass dorsally to both the ventral and the dorsal excretory vessels. The scolex is 96 /* long and 140 /* in maximum breadth across the widest part. There are four, apparently unarmed, suckers which measure 62-74 fi X 50-56 /*. The rostellum, 82 fi long by 64 p in diameter, bears 10 hooks which are 23-28 /* long (Text-fig. 26). The three testes are arranged in a triangle (Text-fig. 27) with one poral and posterior, one aporal and posterior, and one aporal and anterior. The testes measure 60-65 /* X 49-6i /*• There are both external and internal seminal vesicles. The external seminal vesicle is 25-35 fi in diameter and lies anteriorly about the centre of the proglottis. The internal seminal vesicle is larger and measures 40-50 /* in diameter. The cirrus-sac was not clearly seen, but appears to be about 65-75 ju, long by about 40 /* in diameter. The ovary, bilobed and central, measures up to about 200 /* across its total width, by 50-80 fi. Posterior to the ovary, and lying slightly aporally is the compact vitelline gland which measures 50-55 fi X 36^-41 /£. The receptaculum seminis lies prominently in the anterior, median portion of each proglottis and, when full of sperm, reaches a size of 52-70 //. x 48-55 /*. CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO The present specimens are not sufficiently well preserved to allow the elucidation of details of uterus development. The eggs, however, are 42-48 fi in diameter (Text-fig. 28) while the contained oncospheres are 26-32 fi X 20-30 /*. The hooks of the oncosphere measure 18-20 fi in length. 26 loop FIGS. 25 and 26. Hymenolepis mahonae nom. nov. Scolex (Fig. 25), and rostellar hooks (Fig. 26). FIGS. 27 and 28. Hymenolepis mahonae nom. nov. (Fig. 28). Mature proglottis (Fig. 27), and egg DISCUSSION. The present material agrees well with material described by Mahon (1958) as Hymenolepis fringillarum (Rudolphi, 1810). Mahon's description, however, does not agree sufficiently well with the description of Rudolphi (1810), nor with the description of Joyeux and Baer (1936), to allow for its inclusion in the specise fringillarum. For purposes of comparison, Mahon includes the figures quoted by Joyeux and Baer in her description, and although the scolex in her material has a diameter of less than half that of the worms described by Joyeux and Baer, and although the cirrus-sac in her material is only a little over half the size of the cirrus sac in the other material, she still identified her material as H. fringillarum (see Table IX). It is proposed here that the worms, identified by Mahon as Hymenolepis fringillarum, are, in fact, identical with the present material and represent a new 312 M. D. B. BURT species. The new species has been given the patronymic mahonae in honour of Dr. June Mahon who first described it. Table X shows all the species of Hymenolepis (sensu lato), which possess 10 rostellar hooks, so far recorded from Passeriformes and serves to illustrate the differences between the new species and the other existing species. In order to facilitate comparison of the species, it was thought better to retain the generic term Hymenolepis (sensu lato) despite the work of Spassky and Spasskaja (1954) and Yamaguti (1959) who have sub-divided this vast genus into many smaller genera employing, unfortunately, what in some cases may be regarded as questionable criteria for erecting new genera. TABLE IX Hymenolepis fringillarum (Rudolphi, 1810) compared with H. fringillarum of Mahon (1958) and present material from Borneo Species : Description taken from : Strobila (length x max. breadth in mm.) Scolex (diameter in p) Rostellar f (number) hooks \ (length in /*) Suckers (diameter in p) Testes (diameter in fi) Cirrus sac (length X diameter in p) Eggs (diameter in //) Oncospheres (diameter in fj,) Oncosphere hooks (length in Genital apertures (arrangement) fringillarum (Rudolphi, 1810) Joyeux & Baer (1936) Lopez- Neyra (1942) Mettrick (1958) 32-100 xo-8-i 210—300 10 26-28 90-100 x 100-120 150-170 95-110x40 57x34 u) 48 X 36 20 Unilateral fringillarum (Mahon, 1958, nee Rudolphi, 1810) Mahon (1958) 127-145 10 26 54-58 x 76-79 54X40 mahonae nom. nov. Unilateral 7x0-5(0-8) 140 10 23-28 50-56 X 62- 74 49-61 x 60-65 65-75X40 42-48 26-32 x 20-30 1 8-20 Unilateral It should be mentioned that Spassky and Spasskaja (1954) transferred H. fringil- larum to one of their new genera, namely Passerilepis, and Yamaguti (1959) lists fringillarum as a synonym of Passerilepis passeris (Gmelin, 1790). Mettrick (1958), however, redescribed fringillarum retaining both the older generic name of Hymenolepis and also the specific name of fringillarum, but he did not discuss any possible synonymy of this worm. Although there may be valid reasons for retaining both specific names, fringillarum (Rudolphi, 1810) is listed in Table X with passeris (Gmelin, 1790) in accordance with Ransom (1909) and Yamaguti (1959). * 00 x £. o 00 ^ O O r Jl V V o zosteropis (Fuhrmann, igi8) M Tj- ON H $ •i 3 o X N C4 0 N 1 ° T5 °° 1 w A °° CO 1 X o A H o 1 >o 5" isseriformes CO u K PH Passerifon i-3 2 «•"§ * ON £>;Q- g1 •^^ "e"^ 4) N rf O 80-110 X 1-1-8 032-002 o o oo o V ^ T5 *o 1 o | v O H OO A oo tx 10 OO O&I-O2I 00-270 x 70-140 1 X o 1 N O CO 1 8-20 seriformes , *"""** 3 *O ' -g N i 5 W ^ PH SH •H ri So" ^^ en bo | 3. 0 m Us II" "5 Hughes (194 Lopez-Neyr (19420/6) Mettrick (195 ? • 2-8 • I X 002-09 250-35° o O O N M •0 H | | oo 10 H t^. I5O-2OO 130-190 x 85-110 oo X 0 IH H n- X4 vO N ^A 5- "•s 20-22 Passeriforme X w u be M 1 i passeris (Gmelin, 1790) =fringillarum (Rudolphi, 1810 Joyeux & Baer , (1936) Lopez-Neyra (ig42o/6) Mettrick (igs8) X o V ooi CM ° CO 210-300 IO 26-28 IOO-I20X 90-100 150-170 95-100x40 57x34 CO X oo 20 Passeriformes 3 •H M P ri 1-*.. en C} /^j J2 •S a ps* •*• X V0 a | ||| N Tt- o X S o o o o >O M CO t^ | o M >o CO I * •d 1 ?£" 0 ON && o N A 00 HH 8 ° ? ? M 00 U-) M 00 go-ioo 80-300 X 45 -60 x 36-65 s* •ii CO w 2O sseriformes C •^ i_J V H oo 1 £1 Tf PH fc! S a • iH o *•""•* X^HV CJ 43 g 5S. ft CO a a 's; 3 ri a o I Q ibila (length X max. breadth in .ex (diameter in ft) tellum (diameter in ,« tellar f (number) 3oks \ (length in ,w) iers (diameter in /*) :es (diameter in /*) us sac (length x diameter in p s (diameter in p) ospheres (diameter in osphere hooks (length in host (order) 1 o (j en tn s 75 00-^3 4> be be g o .a a > c/5 PH PM CO H b W O 0 < ZOOL. 17, 8. 3i4 M.D.B.BURT Hymenolepis sp. (Text-figs. 29-32) HOSTS. Anthreptes malacensis 9318, 9358; Nectarinia calcostetha 9092. Mature worms measure up to 30 mm. long by 0-35-0-46 mm. in maximum breadth. The genital apertures are irregularly alternating and the genital ducts pass dorsally to both the longitudinal excretory vessels. FIGS. 29 and 30. Hymenolepis sp. Scolex (Fig. 29) and rostellar hooks (Fig. 30). The scolex (Text-fig. 29) measures 104-121 /* X 115-126,11 and bears an armed rostellum and four, apparently unarmed, suckers. There are 8 hooks (Text-fig. 30) on the rostellum and these measure 28-36 /n long. The suckers are 46-56 // X 33~49 /*• The three testes (Text-fig. 31) lie in a straight line and, when mature, measure 64-110 ju, x 48-67 /n. The cirrus-sac is 78-88 p long and has a diameter of 25-33 /^. Although the cirrus was not seen in an extruded position, it could be seen lying within the cirrus-sac and has a diameter of about 6 p. There is present both an external and an internal seminal vesicle, the latter measuring 21-25 /* X I5~23 /*• The vas deferens is slightly coiled and lies anteriorly to the cirrus-sac, roughly parallel to the anterior edge of the proglottis. FIGS. 31 and 32. Hymenolepis sp. Mature proglottis (Fig. 31), and egg with enclosed oncosphere (Fig. 32). 315 The ovary is distinctly trilobed and lies in the middle of the proglottis. The vitelline gland, lying immediately posteriorly to the ovary, is compact, measures 65-85 /A X 43-52 fi, and lies ventrally to the middle testis. The receptaculum seminis is large and increases to a maximum size of 163 JLL x 80 /A, although in most mature proglottides it measures about 140 /i by about 70 fi. There is a sudden transition between mature and gravid proglottides, the eggs being only just visible in the last mature proglottis and filling the whole of the medulla, extending beyond the excretory vessels, in the first gravid proglottis. The eggs (Text-fig. 32) measure 25-40 ft in diameter and the contained oncospheres are 10-14 fjL in diameter. The hooks of the oncosphere are 7-9 p long. DISCUSSION. The present material does not resemble any species listed by Fuhrmann (1932), Lopez-Neyra (19420, b), or by Yamaguti (1959) as coming from Passeriformes. However, in view of the fact that it has not been possible to examine all the references to Hymenolepis (sensu lato) it was considered advisable not to erect a new species to contain this worm at this time. Genus FIMBRIARIA Frolich, 1802 Fimbriaria fasciolaris (Pallas, 1781) HOST. Domestic duck (Anas boschas L. dom.) 9575. Specimens are greatly contracted, but appear to fit the description by Wolffhugel (1936), who also lists the full synonomy of this species. Webster (19430), in his review of the Fimbriariinae, mentions that he found a smaller range in the number of longitudinal muscle-bundles in F. fasciolaris than is quoted by Wolffhugel, the former finding only 110-120 in his material whereas Wolffhugel gives the range as 60-120. Although the present material is not well enough preserved for any des- cription, its similarity both to the worms described by Wolffhugel (1936) and to specimens in the Helminthological collection of the British Museum (Natural History) identified as Fimbriaria fasciolaris makes identification reasonably positive. Family DILEPIDIDAE Fuhrmann, 1907 Genus PARICTEROTAENIA Fuhrmann, 1932 Paricterotaenia burti Sandeman, 1959 (Text-fig. 33) HOST. Charadrius leschenaultii 9110. One, small immature worm was present which measured 0-35 mm. long by 0-12 mm. in maximum breadth. Only two immature proglottides were present, both being twice as broad as long. The scolex (Text-fig. 33) is 200 p X 240 p and possesses four unarmed suckers, 120-140 ju X loo-no p. The rostellum, 150 // long by 70 JLI in diameter, bears 16 hooks arranged in what appears to be a single row. The hooks are 50-52 fi long. No genitalia were seen at all. M. D. B. BURT 33 FIG. 33. Paricterotaenia burti Sandeman, 1959. Scolex. DISCUSSION. The present material appears to agree sufficiently well with Paricterotaenia burti Sandeman to warrant identification with that species. Sandeman (195 90) erected the species to contain worms he found in Lymnocryptes minimus and Numenius arquatus from the River Eden, Fife, and also to contain, in part, Paricterotaenia stellifera (Krabbe, 1869). In the original description of P. stellifera Krabbe gives two sets of hook characteristics : one with 10 hooks, 55 /* long; and the other with 14 hooks of length 46-51 (i. The former set of hook characters is that which has become ascribed to P. stellifera (Krabbe, 1869) while the latter, prior to Sandeman, had been ignored. Hooks of the present material agree closely with those drawn and described by Krabbe and also those described by Sandeman which pertain to P. burti. The measurements of the scolex and rostellum also agree reasonably closely with Sandeman's description and, further- more, the worm described by Krabbe, by Sandeman, and the present worm were all found in Charadriiiormes. This appears to be the first record of this worm in Asia. Genus DILEPIS Weinland, 1858 Dilepis ardeolae Singh, 1952 (?) (Text-fig. 34) HOST. Butorides striatus 9040. The worms are small, measuring up to about 3 mm. long by 0-25 mm. in maximum breadth. The genital apertures are unilateral. The scolices are all missing, although several worms are present. There are 7 testes (Text-fig. 34) lying mainly posteriorly and dorsally to the female genitalia and measuring 30-40 ju, in diameter. In some proglottides, the testes can be seen extending laterally and anteriorly to the ovary on the aporal side. Neither an internal nor an external seminal vesicle was seen. The cirrus-sac, CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 317 FIG. 34. Dilepis ardeolae Singh, 1952 (?) Mature proglottis. though not seen clearly, appears to measure about 80 /* long by about 25 fi in dia- meter, but this is probably a low figure as there were no gravid proglottides present for comparison. The cirrus is armed, but was not seen in the extruded position. In the anterior region of maturing proglottides, the vas deferens becomes profusely coiled and probably serves as an external seminal vesicle when it swells up with sperm. Lying approximately in the middle of the proglottis is the bilobed ovary, each lobe tending to be more or less spherical. The vitelline gland, lying immediately posteriorly to the ovary, is compact and measures about 30 /* x 20 /*. In the majority of cases, the vagina opens into the genital atrium posteriorly to the opening of the cirrus-sac, but this is not constant. Towards the centre of the proglottis the vagina opens into a receptaculum seminis which is broadly fusiform. DISCUSSION. The only feature in which the present material differs from that described by Singh (1952) as Dilepis ardeolae is the size of the cirrus-sac. In Singh's material the length of the cirrus-sac is given as 0-248-0-31 mm. and the diameter as 0-031-0-037 mm. The cirrus-sac in the present material, however, appears to be much smaller, being only about 80 /* long by 25 fi in diameter, but as there are no gravid proglottides present it is difficult to state whether the size of cirrus-sac meas- ured is the largest size in a complete worm. Despite the apparent discrepancy in size of cirrus-sac, the other characters are in such close agreement that the present worms are tentatively identified as Dilepis ardeolae Singh, 1952. Moreover, the fact that the host Ardeola grayi from which Singh described his species, is closely related both in habitat and phylogenetically to Butorides striatus, the host from which the present material comes, further suggests that the worms may be of the same species. Genus LIGA Weinland, 1857 Liga facile (Meggitt, 1927) Szpotanska, 1931 (Text-figs. 35-36) A nomotaenia facile Meggitt, 1927. Anomotaenia trivialis Meggitt, 1927. HOST. Actitis hypoleucos 9196. The worms are very small, not exceeding i mm. in length, and without exception are immature, no genitalia being seen at all. The maximum breadth in all the worms M. D. B. BURT present is the breadth of the scolex across the region of the suckers, as in no instance does the breadth of any of the proglottides exceed the breadth of the scolex. The scolex (Text-fig. 35), including the rostellum, measures 160-170 p in length and is 130-150 ju, broad. The suckers measure 79-82 /* x 72-76 /* and are unarmed. There are, however, some marks which may be the scars of attachment of acetabular spines along the posterior edge of the suckers. The rostellum, measuring 80-100 fi long by 40-60 /* in diameter, bears 20 hooks in two alternating rows. The hooks (Text-fig. 36) in each row are of different sizes: those lying posteriorly measure 41-43 fji long, while those lying anteriorly measure 46-51 // long. IQOM H FIGS. 35 and 36. Liga facile (Meggitt, 1927). Scolex (Fig. 35) and rostellar (hooks Fig. 36). DISCUSSION. The present material differs only slightly from that described by Meggitt (19276) in that the scolex of Meggitt's material is 270 fi x 290 /*, whereas the scolex in the present material is 130-150 /* X 160-170 ju. In view of the close correlation, however, which obtains in the number and size of the rostellar hooks, the size of the rostellum, the small size of the whole worm; and, furthermore, in the fact that both worms are parasites of wading birds, it seems reasonable to identify the present material as Liga facile (Meggitt, 1927). Szpotanska (1931) reviewed the genus Liga Weinland, 1857, and transferred Anomotaenia facile Meggitt, 1927, and A. trivialis Meggitt, 1927, to this genus. She regarded trivialis as a synonym of facile and redescribed the latter species using new material from Burhinus oedicnemus. Sandeman (1959^), while accepting that both species belong to the genus Liga, nevertheless regards facile and trivialis as distinct species, but has not yet published his evidence for this view. Williams (1962), however, having examined the type-material of both facile and trivialis, supports the view of Szpotanska that the two species are not distinct. On the basis CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 319 of his comparison of Meggitt's type-material with the descriptions of both Meggitt and Szpotanska, he states, in his detailed review of the genus Liga, that there seems to be " scant evidence for regarding A. facile and A. trivialis as distinct species ". The main feature of difference between facile and trivialis from Meggitt's description is the size of the rostellar hooks. In A . facile the hooks are 40-50 /* long, while in A . trivialis the hooks are 38-39 /* long. As can be seen in the present material, however, the hooks in the anterior row are longer than those in the posterior row. Accordingly, it is not improbable that Meggitt measured hooks from both rows in the material he described as A . facile, but measured only hooks in the posterior row in the material he described as A. trivialis. The fact that Williams (1962), on re-examining the type-specimens of A . trivialis, found hooks as long as 45 ju, supports this postulation. Therefore, there appears to be no justification for regarding facile and trivialis as distinct species. Genus ANOMOTAENIA Cohn, 1900 Anomotaenia depressa (Siebold, 1836) Fuhrmann, 1908 (Text-figs. 37-39) Taenia depressa Siebold, 1836. Ligafrigida (Meggitt, 1927). HOST. Apus affinis 8784. The present material comprises fragments, all of which probably come from a single worm. The total length of all the fragments is 8 mm. and the maximum breadth is 0-6 mm. The immature proglottides at first tend to be rather square in shape, but as they mature they become longer than broad. The genital apertures are irregularly alternating. The scolex is missing. There are 30-40 testes (Text-fig. 37) which surround the female genitalia laterally and posteriorly and which measure 40-70 /u in diameter. The cirrus-sac (Text-fig. 38) is large, with a length of 400-480 p and a diameter of 80-100 p. The cirrus appears to be fairly short when extruded, having a maximum observed length of 150 fi. The diameter of the cirrus base is about 30 /* and the diameter of the tip is 12 fi. The cirrus is covered for most of its length with fine spines about 4-5 /* long. There is no external seminal vesicle present, but the vas deferens, on entering the cirrus-sac, expands to form an internal seminal vesicle in the proximal part of that organ. The ductus ejaculatorius is quite narrow and lies convoluted in the distal part of the cirrus-sac before entering the cirrus. The deeply-lobed ovary is situated posteriorly and ventrally to the cirrus-sac which, in some proglottides, overlies part of the ovary. The vitelline gland, lying posteriorly to the ovary and anteriorly to the testes, is irregularly lobular. The vagina has its opening into the genital atrium posterior to that of the cirrus-sac and lies parallel to the cirrus-sac until it eventually leads into the receptaculum seminis through a peculiar, " chitinoid " structure (Text-fig. 39). This structure is roughly dumb-bell shaped and has a length of 55-57 /*. The diameter of the part 320 M. D. B. BURT 37 FIGS. 37-39. Anomotaenia depressa (Siebold, 1836). Mature proglottis showing male genitalia and vagina (Fig. 37), cirrus-sac and part of the vagina (Fig. 38), and chitinoid vaginal apparatus (Fig. 39). closest to the vagina is the same as the diameter of the part next to the receptaculum seminis and measures 27-29 /*. The diameter of the narrow constriction is 8-10 /*. The receptaculum seminis, situated immediately behind the cirrus-sac and dorsal to the vitelline gland, has a twist in it and when filled with sperm reaches up to 100 /* in overall length by about 40 fi in diameter. Gravid proglottides are not present. DISCUSSION. Although the scolex is missing and gravid proglottides are not present, this worm is almost certainly the same species as that described by Joyeux and Baer (1936) from Apus apus. Through the kindness of Professor J. G. Baer, I have had a chance to compare the above material with worms from his own collection which he had identified as Anomotaenia depressa and undoubtedly the worms were identical. The " chitinoid " structure present at the junction of the vagina and the recepta- culum seminis was described by Joyeux and Baer (1936) thus: " Le vagin est entoure, pres du receptacle seminal, d'un manchon cellulaire, auquel fait suite un appareil de fermeture chitineaux, a 1'entree de ce receptacle." In order to measure the structure in the present material accurately, two proglottides were mounted, unstained, in Berlese fluid which rendered most of the tissue transparent. Those structures which showed to advantage after this treatment were the cirrus-sac, the CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 321 narrow portion of the ductus ejaculatorius, the cirrus with its armature of spines, and the structure surrounding the vagina as it enters the receptaculum seminis. Dollfus (1958, p. 515), in a footnote, suggests that the structure may be made of a scleroprotein, but that its chemical nature is not known. In the same paper Dollfus reviews those species belonging to the genera Anomotaenia Cohn, 1900; Pseudangularia Burt, 1938 (a); Neoangularia Singh, 1952; and Neoliga Singh, 1952; which possess this structure which he describes as " un appareil occlusif entre le receptacle seminal et le vagin distal ". Dollfus (1958) discusses fully the complicated synonymy of Anomotaenia depressa (Siebold, 1836) and cites all the descriptions of note. Most of these descriptions are inadequate on their own, and accordingly Dollfus gives a compound description which takes into account those given by Krabbe (1869), von Linstow (1897), Fuhrmann (18990), Lopez-Neyra (1923), Joyeux and Timon-David (1934) and which agrees with data taken from his own material. In the resume of his paper Dollfus stresses the insufficiency of present knowledge of both Anomotaenia depressa (Siebold, 1836) and A. cyathiformis (Frolich, 1791), particularly as both these species have been described both from Passeriformes and Cypseliformes. Furthermore, several authors have used, for descriptions of cyathiformis, characters of worms taken from hosts belonging to both these orders. A more recent, though short, description of A. depressa, which agrees with that of Joyeux and Baer (1936), is given by Vojtechovska-Mayerova (1952). Anomotaenia nymphaea (Schrank, 1790) (Text-figs. 40-41) Taenia nymphaea Schrank, 1790 HOST. Numeniusphaeopusg^o. Two, small immature worms only were present. Both had complete scolices with full complements of rostellar hooks. A squash preparation was made of one of the scolices while the other was mounted whole in Canada-balsam. The longer of the two worms measures 0-75 mm. with a maximum breadth of o-ii mm. The scolex (Text-fig. 40) is 200 p long, including the length of the rostellar sac, and has a diameter of 250 ju. The suckers are ovoid and measure 90-110 /* X 80-90 fi. No sucker spines were seen. The rostellum is invaginated, measures 200 /* from its tip to the posterior extremity of the rostellar sac by no fi across the broadest part, and carries 20-22 hooks arranged in a double row. The rostellar hooks (Text-fig. 41) are of two sizes, the larger, lying anteriorly, are 75-80 p long, while those in the posterior row are 65-70 /a long. DISCUSSION. The present material agrees well with material described by Joyeux and Baer (1936) and by Mahon (1958), although Mahon's material was probably in a greater state of relaxation than the present, as the diameter of the suckers in her material is considerably greater. Sandeman (19590:) has so far been unable to deal with the synonymy of this species, but is at present working on this. 322 M. D. B. BURT 40 FIGS. 40 and 41. Anomotaenia nymphaea (Schrank, 1790). Scolex (Fig. 40) and rostellar hooks (Fig. 41). Anomotaenia tringae (Burt, 1940) Sandeman, 1959 (Text-figs. 42-43) Paricterotaenia tringae Burt, 1940. Anomotaenia par amicrorhyncha Dubinina, 1953. HOST. Tringa glareola 9255. The worms measure up to 16 mm. long by 0-73 mm. in maximum breadth. In immature proglottides the breadth is about twice the length, but as the proglottides mature and become gravid, so does the ratio of length to breadth increase until in the gravid proglottides the length is about equal to the breadth. The genital aper- tures are 89 % regularly alternating. The genital ducts pass between the dorsal and ventral excretory vessels. Scolices are not present. There are 9-11 testes (Text-fig. 42), most of which lie aporally, and they measure 45-70 IJL x 25-40 fi. There does not appear to be either an internal nor an external seminal vesicle. The cirrus-sac measures 87-100 fi long by 25-35 /* m diameter and has a typical constriction about halfway along its length. In many instances the unarmed cirrus could be seen projecting into the vagina. The ovary is irregularly digitate, lies in the anterior third of mature proglottides. and stretches from the aporal excretory vessels almost to the poral excretory vessels. The slightly lobular vitelline gland lies posteriorly to the ovary, but in front of the testes, and measures 45-75 ^ across. Lying to the poral side of the vitelline gland is the receptaculum seminis which is ovoid and measures 59-74 n X 46-64 ju, when not swollen with sperm, but when swollen reaches up to 130 fi x 70 fi. CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 323 FIG. 42. Anomotaenia tringae (Burt, 1940). Mature proglottis. FIG. 43. Anomotaenia tringae (Burt, 1940). Gravid proglottis. The uterus is sac-shaped initially, becomes increasingly lobular as the eggs ripen (Text-fig. 43), until in fully-gravid proglottides it tends to break down into pseudo uterine capsules. The eggs measure 34-39 ju X 31-34 // and the contained onco- spheres are 25-30 p X 24-25 fi. The hooks of the oncosphere were not fully formed. DISCUSSION. The present material agrees well with the type-material of Paricterotaenia tringae which Mr. D. R. R. Burt has so kindly placed at my disposal, and with material described by Baer (1959). Despite the fact that the scolex is 324 M. D. B. BURT wanting in the present material, such good correlation obtains through direct com- parison of the rest of the anatomy with the type-material, that there remains no doubt as to the identity of the present worms. Sandeman, (19590) transfers this species from the genus Par icier otaenia to the genus Anomotaenia. Burt (1940) states that owing to the fact that there does not exist a dilepid genus with a double crown of hooks and regularly alternating genital apertures " The choice lies therefore between Anomotaenia and Par icier otaenia ". Burt chose to place the species in the genus Par icier otaenia on the grounds that the scolex and rostellum are very similar to those seen in several species of Par icier otaenia. The transference of this species to Anomotaenia by Sandeman appears to be purely on the grounds of its possessing a double row of hooks and this is accepted by Baer (1959). Until such a time as Sandeman publishes his larger work on the dilepids of waders, and explains his justification for such a treatment, it is probably best to leave this species in the genus Anomotaenia. Genus PARVITAENIA Burt, 1940 Parvitaenia sp. (Text-fig. 44) HOST. Butorides striatm 9040. The worm is incomplete, lacking a scolex. It measures 14 mm. in length by 0-9 mm. in maximum breadth. The proglottides are all broader than long and the genital apertures are irregularly alternating. FIG. 44. Parvitaenia sp. Mature proglottis. There are 50-52 testes (Text-fig. 44), half lying anteriorly to the female genitalia and half lying posteriorly, which measure 30-50 p x 15-45 p. The cirrus-sac is not distinct, but appears to be about 115 /* long and 45 /i in diameter. It lies close to the anterior margin of the proglottis, opening into the genital atrium on the lateral margin. The cirrus is long and in many cases can be seen projecting into the vagina of the same proglottis and reaching almost to the receptaculum seminis. The ovary is distinctly bilobed, the combined length of both lobes measuring 345-395 fjb. The vitelline gland, lying more or less in the centre of the proglottis and posterior to the ovary, is irregularly bilobed having a width of 90-110 /«. Lying between the ovary and the vitelline gland is the roughly spherical shell-gland which has a diameter of about 45 /*. CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 325 There are no gravid proglottides, but in the most mature of those present eggs can be seen accumulating in the anterior poral part of each proglottis. The paren- chyma contains many calcareous corpuscles. DISCUSSION. It is not possible to identify this material as to species owing to the lack of a scolex. The internal anatomy, however, strongly suggests that it belongs to the genus Parvitaenia Burt, 1940. Baer and Bona (1960) slightly alter Burt's generic diagnosis and consider the genus to contain 13 valid species: P. ardeolae Burt, 1940; P. macropeos (Wedl, 1855); P. cochlearii Coil, 19550; P. purpurea Johri, 1959; P. magna Baer, 1959; P. macrophallica Baer and Bona, 1960; P. microphallica Baer and Bona, 1960; P. ambigua Baer and Bona, 1960; P. ardeae (Johnston, 1911) ; P. glandularis (Fuhrmann, 1905) ; P. aurita (Rudolphi, 1819) ; P. davipera Baer and Bona, 1960; and P. pseudocydorchida Baer and Bona, 1960. Of these, five are new species and four are new combinations. The full descriptions of the new species are to be given at a later date and, until such time as these descriptions appear, simply allocating the present material to the genus Parvitaenia should suffice. Genus VITTA Burt, 1938 Vitta rustica (Neslobinsky, 1911) Baer, 1959 (Text-figs. 45-46) Anomotaenia rustica Neslobinsky, 1911. Vitta magniurtcinata Burt, 1938. HOST. Hirundo rustica 9202. The strobila measures 25 mm. long by 2-5 mm. in maximum breadth. The prog- lottides are all broader than long, the ratio of breadth to length varying from 3 : i to 5 : i depending both on the state of contraction of the worm and on the site of the proglottis within the strobila. The genital apertures are irregularly alternating and situated laterally, close to the anterior margin of each proglottis, while the genital ducts pass dorsally to both the ventral and dorsal excretory vessels. The scolex (Text-fig. 45) has a diameter of 370 p and a length, including the length of the rostellum, of 330 p. The suckers are 73-81 p X 67-79 A* and are unarmed. The rostellum, 185 /i long by 165 fi in diameter, bears 42-45 hooks arranged in two rows in such a way that for every one hook in the anterior row, there are two hooks in the posterior row. The hooks are 50-60 fi long, those hooks in the posterior row being slightly longer than those of the anterior row. There are 60-90 testes (Text-fig. 46) which, when fully developed, measure 62- 77 ft x 60-67 /*» and which are arranged posteriorly and laterally to the vitelline gland and dorsally to the ovary. There is no external seminal vesicle as such, but the vas deferens expands in the region of the cirrus-sac and lies in large loose coils which function as an external seminal vesicle. On entering the cirrus-sac, the vas deferens forms an internal seminal vesicle. The cirrus-sac, lying parallel and close to the anterior margin of the proglottis, measures 320-360 ju, long by 65-85 fi in diameter. The ovary is bilobed, each lobe being fan-like and divided into a great number of smaller lobes which reach the excretory vessels on both sides of the proglottis. 326 M. D. B. BURT FIG. 45. Vitta rustica (Neslobinsky, 1911). Scolex. FIG. 46. Vitta rustica (Neslobinsky, 191 1). Mature proglottis. The vitelline gland is U-shaped and measures 90-110 /i x 50-70 /*. Situated dorsally and slightly anteriorly to the vitelline gland is a well-defined shell-gland, 45 ju, in diameter. In the more mature proglottides, the receptaculum becomes quite swollen, often exceeding the size of the cirrus-sac. CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 327 DISCUSSION. The above material agrees well with that described by Burt (19386) as Vitta magniuncinata and with that described by Baer (1959) as V. rustica. The genus Vitta was erected by Burt (19386) to contain two different worms taken from Hirundo rustica gutteralis Scop., 1786, namely magniuncinata and minutiuncinata. The genus closely resembles Anomotaenia Cohn, the main feature of difference being that in Anomotaenia the genital ducts pass between the excretory vessels, whereas in Vitta they pass dorsal to both. Baer (1959) transfers Anomotaenia rustica Neslobinsky, 1911, to the genus Vitta and places magniuncinata as a synonym of rustica. He also provides a key to the four species of Vitta which he recognizes, using only the characters of hook number and size, and number of testes. These four valid species are: V. parvirostris (Krabbe, 1869); V. minutiuncinata Burt, 1938; V. undulatoides (Fuhrmann, 19086); and V. rustica (Neslobinsky, 1911). Yamaguti (1959) includes Vitta as a synonym of Angular ella Strand, 1928, but gives no indication as to what grounds he has for this. The diagnostic characters of the genera differ essentially in the arrangement of the rostellar hooks. In Vitta there is a double row, whereas in Angular ella there is a single row. On this basis, it is proposed to retain, as valid, the genus Vitta Burt, 1938. Dilepidid sp. (Text-figs. 47-48) HOST. Cacomantismerulinusg^o^. The strobila measures 3-5 mm. long by 0-7 mm. in maximum breadth. The genital apertures are unilateral. The worm is in a poor state of preservation and did not stain well. The scolex (Text-fig. 47) is 160 p long, including the length of the rostellum, by 260 /j, broad across the suckers. The four suckers are circular in outline and measure loo-no n in diameter. Sucker spines were not seen. The rostellum, 65 fi long and 45 fi in diameter, bears a single row of 10 hooks which are not evenly spaced round the periphery therefore suggesting that there may have been some hooks lost. The length of the hooks is 18-19 p and their shape can best be seen in Text-fig. 48. FIGS. 47 and 48. Dilepidid. Scolex (Fig. 47), and rostellar hooks (Fig. 48). 328 M.D.B.BURT The only genitalia that can be made out with any certainty are the testes, which number 24-30, and which measure 45-75 /£ X 23-38 fi. In one segment there was a suggestion of double genitalia, but this clearly was not the normal arrangement. DISCUSSION. The worms show many features of the Dilepididae, but could belong to a wide range of genera. But for the shape of hooks, the data available from this worm agree very closely with those of Anomotaenia mutabilis (Rudolphi, 1819) Fuhrmann, 1907, even as to the order of host it infests. Owing to the lack of mature and gravid proglottides which can be described, identification further than the family is not possible. Genus KOWALEWSKIELLA Baczynska, 1914 Kowalewskiella susanae n. sp. (Text-fig. 49) HOST. Tringa glareola 9255 . Several worms are present in a good state of preservation. They measure up to 25 mm. long by 0-525 mm. in maximum breadth. The immature proglottides are slightly broader than long, but the mature proglottides become twice as long as they are broad. The genital apertures are irregularly alternating. The genital ducts pass between the dorsal and ventral excretory vessels. The scolex measures 90-100 ju, in diameter, but is bent and squashed so that the length cannot be measured accurately. The suckers are about 45 p in diameter, but these too are squashed. Only three hooks were seen on the rostellum, of which only one could be measured, its length being about 8 ft. There are 21-30 testes (Text-fig. 49) arranged in two groups, one group lying anteriorly to the female genitalia and the other lying posteriorly. The number of testes in each group is approximately equal and the size of the testes is subject to wide variation measuring 45-65 //. X 35-60 /i. There is neither an internal nor an external seminal vesicle present, the vas deferens, however, probably acts as an external seminal vesicle as it becomes greatly swollen with sperm in mature proglot- tides. The cirrus-sac is 92-98 fi long by 53-60 fi in diameter and is situated laterally, in about the middle of the proglottis. It opens into a well-developed genital atrium. In the mounted material, the cirrus is present only as a short papilla, but there is a convoluted ductus ejaculatorius still present in the cirrus-sac. The ovary is distinctly bilobed, each lobe being separated by a relatively long isthmus. The lobes are irregularly lobular and, in proglottides where the cirrus-sac is well developed, the total breadth of the ovary is 150-250 /*. The vitelline gland, lying posteriorly to the ovary, measures 55-90 /* X 34-52 /*. Just in front of the vitelline gland, but dorsal to it and behind the ovary, is the shell-gland which has a diameter of 20-25 /*• The receptaculum seminis lies between the cirrus-sac and the ovary and shows wide variation in size depending on the quantity of sperm present, the normal variation being 60-90 fi X 40-50 /*. Gravid proglottides were not found. British Museum (N.H.) Reg. no. 1968.4.25.47-48. CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 329 FIG. 49. Kowalewskiella susanae n. sp. Mature proglottis. DISCUSSION. The present material agrees well with material from the same host described by Baer (1959) as Kowalewskiella cingulifera (Krabbe, 1869) Sandeman, 1959. Sandeman (1959^) re-erects the genus Kowalewskiella Baczynska, 1914, to contain all those species in which the testes are divided into two groups, one anterior to the female genitalia, and the other posterior. In accordance with this, he transfers Choanotaenia glareolae Burt, 1940, Choanotaenia stagnatilidis Burt, 1940, and Choanotaenia hypoleucia [sic] Singh, 1952, all to the genus Kowalewskiella. Further- more despite the differences apparently inherent between these species and Krabbe's, species cingulifera, he regards them all, as well as Kowalewskiella longiannulata Bacyznska, 1914, as synonyms of cingulifera. Baer (1959) accepts this view and in a personal communication points out that " K. cingulifera forms a group within which variations appear to be considerable ". This assumption is presumably based on the fact that Baer (1959) described some worms from Tringa glareola, from the Belgian Congo, as K. cingulifera. As can be seen from the accompanying table (Table XI) if Baer's material is, in fact, cingulifera then there would be present very great variation within the species; more variation than can readily be accepted as existing in any one species. If the two species of Burt (1940), the species described by Singh (1952) and that of Baczynska (1914), are also considered as cingulifera, then this increases the already wide variation even more. Sandeman (1959^) does not outline his reasons for placing these various worms in synonymy with cingulifera, but it is doubtful if this sweeping treatment is justified in the present circumstances and, accordingly, it is here suggested that cingulifera, longiannulata, glareolae, stagnatilidis and hypoleuca are all separate species. To these five valid species should also be added Kowalewskiella bodkini (Vevers, 1923) n. comb. (— Raillietina (Skrjabinia) bodkini Vevers, 1923), K. buzzardia Tubangui and Masilungan, 1937, and K. susanae n. sp. It is unfortunate that Krabbe's original description is not more complete, but it can be differentiated from any of the others on the large size of the scolex, and on the large number of rostellar hooks. K. longiannulata can be distinguished from the rest by the fact that it has a 1* T 0 o CO 0 j?c X I . M fe S^ « £ •o o X H | CQ S 10 0 x o x o •, 00 $ ? o^ &r ^ M ] CO I o 01 (N 1O O •*" X bp ON rt x M .** -rt d 11 I 0 o bodkini B" S ^ 04 O* X O •o >o 10 8 to o fci C S "b/j 1» 42 bo N x T o_ o JP ^- -i C ON _G ON 1 -4-> ^ GO^ M 00 — U-N -t- 00 '^ _b ^ O * — X w -S v • ^* 4e _ti» . ^-^ o ^ 8 PQ !-0 S •S "5 s 3 CQ, Tf Oi 1 1 H T 1 ""> 0 « H 1 1 M Q on S-H O en SJ ~s 1 s " TT ON c « "5" i! a 'o *« """ o CD (X en ^. p. Q r3 cS iO O C A 1 s It m "/: C N' U ^ O^ ^ en o 1 o X 0 » ^ "o |- i PQ i n _2 • — o en Q 'C (0* o cfi ^ 5, •tr ON 2 ON X S o 1 | ^ 2 I O o M 0 •~ "1 = I! V ^O "** X ° '-S N o "•> -^ ^ IT T ^ § T A ^5 i 6 « a 6-2 IO X 75-108 e S Sf s .* ~ 'v o K. *H 0-2 4°- ^- is? II ^ Sp Totanu calidri «a .5 S o 4J O t/3 r/; CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 331 small scolex, small cirrus-sac, but many testes. The two species described by Burt (1940) are well separated on the constant difference in number of testes and in view of the fact that the cirrus-sac in glareolae appears to be significantly larger than that in stagnatilidis , even though the outside lower limit of the former overlaps with the upper limit of the latter. Singh (1952), describes the scolex of hypoleuca as 0-634- 0-672 mm. x 0-48-0-63 mm. These measurements, however, are at variance with the figure he draws and the difference does not seem to be merely a matter of a factor of 10. The suckers are described as 0-043-0-052 mm. in diameter, while the diameter of the scolex is, as stated above, 0-48-0-63 mm. In his drawing, Singh figures four suckers on the scolex and the sizes of the suckers drawn are such that it would be possible to place three suckers, side by side, and still not project beyond the side of the scolex. Thus, the scolex diameter must be about 130 ft or 0-130 mm. According to the scale given, the breadth of the scolex across the suckers is about no fi. Thus, it seems unlikely that the measurements given by Singh refer to the scolex of this species at all and should probably be completely ignored. However, the large size of the cirrus-sac and the large number of testes all serve to separate it from any of the other species. K. bodkini resembles both glareolae and stagnatilidis in many respects, but can be separated from both species on account of the size of the cirrus-sac. In bodkini, even the largest cirrus-sac present, which was measured personally from the holotype and paratype material, is still 40 /* smaller than the lower limit quoted by Burt (1940) for stagnatilidis. Finally, the small number of testes in susanae n. sp. serves to separate it from any of the other species. The small cirrus-sac also separates it from glareolae, stagnatilidis and hypoleuca. The new species described above has been named susanae in grateful recognition of the help received from Miss Susan Burt in the preparation of this manuscript. Genus ASCOMETRA Cholodkovsky, 1913 Ascometra (?) sp. (Text-figs. 50-52) HOST. Centropus toulou 8927. The strobila measures 55 mm. long by 0-9 mm. in maximum breadth. The imma- ture and mature proglottides are up to three times as broad as they are long, while the length of the gravid proglottides is up to twice the breadth. The genital apera- tures are almost unilateral, only 5 % of the apertures occurring on the other side. The apertures in this 5 % all occur singly. The scolex (Text-fig. 50) is 82 /j, long and 62 p in diameter. There may be four suckers present, but the only two seen clearly are 45-50 /JL in long diameter. Sucker spines are not present. For a description of the rostellum (?) see the discussion following this description. There are 6-13 testes (Text-fig. 51) lying mainly posteriorly to the ovary although they may come to lie laterally in some proglottides. The testes measure 37-52 fi X 30-40 /£. The cirrus-sac, 52-64 /i long by 30-50 /* in diameter, is situated laterally about one-third the length of the proglottis from the anterior margin. The cirrus, ZOOL. 17, 8. 22§ 332 M. D. B. HURT 50 FIGS. 50 and 51. Ascometra (?) sp. Scolex (Fig. 50) and mature proglottis (Fig. 51) 7 FIG. 52. Ascometra (?) sp. Gravid proglottis. apparently unarmed and short, could be seen in a few proglottides projecting from the cirrus-sac. The largest cirrus seen was 35 /i long by 4/4 in diameter. The bilobed ovary tends to be slightly displaced towards the aporal side of the proglottis but as it is not lying flat in the majority of proglottides, its full extent is difficult to determine accurately. Immediately behind the isthmus of the ovary is the ovoid vitelline gland which measures 50-75 fi X 45-60 fi. The vagina, which CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 333 opens into the genital atrium posteriorly to the opening of the cirrus-sac, is relatively short, opening into a small receptaculum seminis which lies almost immediately behind the cirrus-sac. The receptaculum seminis measures 35 fi x 20 /^. The uterus (Text-fig. 52) exists as a simple sac-like structure initially which lies in the posterior half of the proglottis within the medulla and not extending beyond the the excretory vessels. Just in front of the uterus lies the paruterine organ which gradually surrounds the uterus and rounds it off into an almost spherical structure. DISCUSSION. The structure of the scolex is very difficult to ascertain as it is lying in such a position that what at first sight looks like a rostellum may in fact be a third sucker superimposed on a fourth one. In the event that the scolex bears four suckers and no rostellum, it is almost certain that the worm belongs to the genus Ascometra Cholodkovsky, 1913, or perhaps to the genus Orthoskrjabinia Spassky, 1947. If the genital apertures are considered as being unilateral, thereby ignoring the 5% of genital apertures which appear on the " wrong " side, the worm should belong to the genus Ascometra; but if, on the other hand, the genital apertures are simply considered as irregularly alternating, this character would place the worm in Spassky's genus Orthoskrjabinia. In view of the fact that in neither of the two men- tioned genera does there exist a worm which agrees sufficiently well with the present worm to warrant identification with it, and in view of the fact that the worms are ton in good enough state of preservation to allow a full description, it has been thought best that the present material be left unidentified until such a time as more material from Centropus toulou becomes available. Genus NOTOPENTORCHIS Burt, 1938 Notopentorchis collocaliae Burt, 1938 (Text-figs. 53-55) HOST. Apus affinis 8784. The worm is 20 mm. long and 0-48 mm. in maximum breadth. The proglottides are all, except for some of the most gravid ones, broader than long, the ratio of breadth to length varying from 4 : I to 3 : 2. The genital apertures are irregularly alternating and the genital ducts pass ventrally to the excretory vessels. The scolex (Text-fig. 53) measures no p in diameter and has a length, including the length of the rostellum, of 120 /t. The suckers are 65-70 /* x 50-55 fi and are unarmed. The rostellum, 60 fi long by 70 /* in diameter, is also unarmed but it is presumed that the hooks of the rostellum have been lost. There are usually five testes (Text-fig. 54), but an occasional proglottis may contain four or six testes. These are situated posteriorly and laterally to the female genitalia with a tendency also to overlie parts of the ovary and the vitelline gland and they measure 25-30 JLL x 18-22 ju. The cirrus-sac is small and most easily seen in gravid proglottides where it measures 30-35 JLL in length by 16-20 ju, in diameter. The ovary, lying in the centre of the proglottis, is large and irregularly lobular, while the vitelline gland, which lies posteriorly to the ovary, is also well developed and measures 40-45 /* in diameter. Situated above the posterior part of the ovary is the shell-gland which has a diameter of 30-35 fi. The vagina, which opens into 334 M. D. B. BURT the genital atrium posteriorly to the opening of the cirrus sac, passes ventrally to the excretory vessels and opens into a receptaculum seminis which, in gravid proglottides where it reaches its maximum size, measures 24-33 /£ x 16-20 /*. The uterus is initially present as a simple sac which rounds off to form a sphere, 120-180 fjL in diameter (Text-fig. 55). There are 20-35 eggs Per proglottis and the con- tained oncospheres measure 30-38 fi x 24-30 /JL. The hooks of the oncosphere are of two different sizes : the two lateral pairs are smaller, measuring 13-15 /* in length while the medial pair is larger, measuring 16-20 /* in length. 200 y IOO p 55 FIGS. 53-55. Notopentorchis collocaliae Burt, 1938. Scolex (Fig. 53), mature proglottis (Fig. 54), and gravid proglottis (Fig. 55). DISCUSSION. The present material agrees very closely with the Ceylon material described by Burt (19380) as Notopentorchis collocaliae for which he created a new species and erected a new genus. Baer (1959), however, transfers to the genus Notopentorchis Burt, Paruterina javanica Hiibscher, 1937 and P. bovieni Hiibscher, 1937 and places N. collocaliae as a synonym of what is now Notopentorchis javanica (Hiibscher, 1937). In Burt's paper, the number of rostellar hooks is not given and Baer estimates the number, from the drawing of the scolex, to be about 50. Through the kindness of Mr. D. R. R. Burt, I have been able to examine the type material of Notopentorchis collocaliae and find there to be between 30-35 hooks, probably about 32. As there are at least 50 hooks in javanica, according to Baer who re- examined the type material of Hiibscher, this would seem to represent a valid differentiating character. This difference, taken with the significant difference in the size of the cirrus-sac, undoubtedly separates these two worms and, accordingly, it is proposed here that Notopentorchis collocaliae is a valid species and should not CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 335 be regarded as a synonym of N. javanica. Singh (1952) describes the species Notopentorchis micropus from Micropus affinis. Although his species differs from collocaliae " in the shape of the hooks, the size of the hooks of the two crowns, size of cirrus pouch, shape of ovary and development of uterus and paruterine organ ", it does not differ from Hiibscher's species, javanica, with which Singh did not compare it. The apparent difference in number of testes (5 in micropus and 8-10 in javanica} is not a valid difference as re-examination of Hiibscher's type material of javanica by Baer indicated that there were never 8-10 testes but only 5. Mokhehle (1951) creates the species Sphaeruterina caffrapi which appears to differ from Notopentorchis javanica only in the fact that the genital ducts pass dorsally to the excretory vessels instead of ventrally. Baer (1959), however, points out that while the genital ducts are described as passing dorsally to the excretory vessels, they are drawn as passing ventrally. Mokhehle (1951) described a second species of Sphaeruterina, namely S.dikeniensis,a.nd again describes the genital ducts as passing dorsally to the excretory vessels but draws them as passing ventrally. Baer assumes that Mokhehle is correct in his drawing but wrong in his description and believes that both worms should be in the genus Notopentorchis, the first one as a synonym of javanica and the second, Sphaeruterina dikeniensis, as a synonym of Notopentorchis vesiculigera (Krabbe, 1882). The accompanying table (Table XII) best illustrates the differences between the four valid species of Notopentorchis. TABLE XII Descripi Strobila (1< max. br Scolex (die Rostellum Rostellar hooks Suckers (d Testes (nu Cirrus sac diamete Oncosphei Valid species of Notopentorchis compared with material from Borneo. Species: vesiculigera bovieni javanica collocaliae Borneo Krabbe, Hiibscher, Hiibscher, Burt, material 1882 1937 1937 1938 tions taken from : Krabbe Hiibscher Hiibscher Baer Burt (1882) (1937) (1937) (1959) (i938a) mgthx 100X1-5 78x0-918 26-5x0-5 25x0-59 26x0-29 20x0-48 sadth in mm.) imeterin^) 408 228 183-260 150 no (diameter in yu) 252 135 90-100 82 70 '(number) 50 70 44-48 5° 3O~35 (personal "g count from "^ type material) ^ o (length in p) 37-46* 60 & 30 25-28 30-31* 27*24 A 20—26 25-26 iameter in p) 180-228 102 68-75 X 75 65-70 X 57-75 50-55 mber) 9-12 8-10 5-6 5 5 (4-6) (length X 150-170 X 110-120 x 57-68 X 35 X 3°-35 X r in /LI) 40 40-50 23 30 16-20 es (diameter in p) — 39x24 27-30 31-32 28 30-38 x 24-30 Oncosphere hooks 17-19 18 15 (length in p) 19 13-15 x 16-20 Host Hirundo Macropteryx Macropteryx rustica longipennis longipennis Cypselus apus Apus caffer Collocalia Apus unicolor affinis 336 M. D. B. BURT Family AGOLEIDAE Fuhrmann, 1907 Genus GYROCOELIA Fuhrmann, 1899 Gyrocoelia perversa Fuhrmann, 1899 (Text-figs. 56-59) Gyrocoelia paradoxa (von Linstow, 1906) Fuhrmann, 1908. Gyrocoelia milligani Linton, 1927. Gyrocoelia pagollae Cable and Myers, 1956. HOST. Charadrius leschenaultii 8999 ; 9334. One full mature " female " strobila was found in host 8999, and two small, barely mature worms — one male and one " female " — were found in host 9334. The scolex (Text-figs. 56 and 57) measures 290-325 ^ in diameter by 250-280 ft in length including the length of the rostellum. The suckers are oval in outline and measure 145-170 ju x 100-125 //.. No sucker spines were seen. The rostellum measures 150-250 /* from the top of the apical cushion to the bottom of the rostellar sac and the diameter of the apical cushion is 50-80 fi. No rostellar hooks were seen. The male strobila (Text-fig. 58) is not fully mature although, in the distal part of the strobila, several cirri can be seen in various stages of extrusion. The length of the strobila is 8 mm. and the maximum breadth is 0-9 mm. The testes are present in a compact group about the centre of the proglottis, but are so crowded together that it is not possible to make an accurate count of their number. There appear to be, FIGS. 56 and 57. Gyrocoelia perversa Fuhrmann, 1899. Scolex with everted rostellum (Fig. 56) and scolex with retracted rostellum (Fig. 57). CYCLOPHYLLIDEAN CESTODES FROM BIRDS IN BORNEO 337 however, about 30 testes, the maximum diameter of the most mature testes present being 30-40 fi. The cirrus-sac measures 240-280 /* X 120-130 fi and passes between the ventral and the dorsal excretory canals. The conical cirrus measures about 250 ju, in its extended form, has a diameter of 75 ju, at its base, and a diameter of about 35 fi at its apex. During the early part of its extrusion, the long spines present on the proximal quarter of the cirrus are easily seen and these measure 10-15 /* m length when measured from a whole mount. The smaller " female " strobila is very similar in external dimensions to the small male, while the larger " female " (Text-fig. 59) measures 53 mm. long by 2-5 mm. in maximum breadth. The ovary is a little over 500 fi across by 50-100 [i. The elongated vitelline gland measures 230-250 fi x 30-45 /*. The " receptaculum seminis " could be seen lying between the ovary and the vitelline gland. In the larger of the two worms, the cirrus-sac measures 420-500 fi x 110-150 fi\ while in the smaller worm the cirrus-sac is 300-320 ft x 120-140 /* . The eggs, present only in the larger worm, measure 68-73 fi x 35-40 fi while the contained oncospheres are 20-30 fi in diameter. The hooks of the oncosphere were not examined as squash preparations, but on a whole mount these appeared to be about 8 JLL long. The muscular system is very well developed and comprises 100-150 large bundles of longitudinal muscle fibres in two poorly separated layers. Each bundle contains up to 100 individual muscle fibres. The transverse muscles lie between the longi- tudinal muscle bundles and also round the central medulla. Close to the cuticle lie further individual longitudinal muscle fibres interspersed with large parenchyma cells. FIGS. 58 and 59. 59 Gyrocoelia perversa Fuhrmann, 1899. Part of a male strobila (Fig. 58) and part of a " female " strobila (Fig. 59). 338 M.D.B.BURT DISCUSSION. The present material agrees reasonably closely with that described by Baer (1959) ( ID N ^ ro O! HH HH I" g PQ 'H ocT 5 !£? to Q W 1 a -< pq H ,2 fe S, "2 oo" M i 2 u-j M ir> O en W r-l O^i O O^ C4 E ^ H CO 8 03 > AH CO S U 00 g CM o> t/3 M CO .s I 1 '0 Is O 4 ^ w 'C CO .8 « j 1^— ( rt 01 »— i W O 'o P ^ ^ « a; o3 3 „_ UO > T3 o 0 0 «T "o "o 'o S i cn n! .0 ,0 .0 ^ § ^ S * AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 353 The type-status of the specimen or specimens is given together with the registration number, locality data, number of specimens and the means of preservation (i.e. alcohol, microslides, or dry). There is doubt over the exact status of some of the type material from the Norman Collection. Norman marked as " cotype " specimens of any species which he received from the author of the species concerned. It has not been possible, in a few cases, to relate exactly the locality on the label with that given in the original description. Where there is doubt, this is indicated in the text. Local- ities of " Discovery " material determined by K. H. Barnard and marked thus * are those nominated as " Types " in his report (Barnard, 1932). This list includes type material incorporated into the collections up to and inclu- ding 31 March 1968. The authors acknowledge with thanks the help and encouragement given them by Dr. A. L. Rice and Mr. R. W. Ingle. Family LYSIANASSIDAE ACIDOSTOMA Liljeborg Acidostoma obesum (Bate) 1862 : 74 [Anonyx obesus], HOLOTYPE: Reg. no. 1952 15:7:2 (alcohol). LOCALITY : Moray Firth, Scotland. COLLECTION: Bate. ACONTIOSTOMA Stebbing [Acontiostoma magellanicum] Stebbing 1888 : 714 (transferred to A contiostoma marionis Stebbing by Barnard 1932). HOLOTYPE: Reg. no. 1889 : 5 : 15 : 34 (4 slides). LOCALITY: Sta. 313. Off Cape Virgins, Patagonia, 52° 20' S 67°39'W, 55 fm, 20 January 1876. COLLECTION: " Challenger ". Acontiostoma marionis Stebbing 1888 : 709. HOLOTYPE: Reg. no. 1889 : 5 : 15 : 33 (specimen in alcohol + 6 slides). LOCALITY: Sta. 145. Off Marion Island, South Indian Ocean, 50-75 fm, 27 December 1873. COLLECTION: " Challenger ". ALLOGAUSSIA Schellenberg Allogaussia lobata Barnard 1932 : 67. SYNTYPE : Reg. no. 1936 : n : 2 : 346* (i specimen, alcohol). LOCALITY: Sta. 170. Off Cape Bowles, Clarence Island, South Shetland Islands, 61° 25' S 53° 46' W, 342 m, 23 February 1927. SYNTYPES: Reg. no. 1936 : n : 2 : 347-348 (1+2 specimens, alcohol). ZOOL. 17, 9. 23§§ 354 M. H. THURSTON & ELIZABETH ALLEN LOCALITY: Sta. 142. East Cumberland Bay, South Georgia, 88-273 m, 30 Decem- ber 1926. Sta. 208. Off Livingston Island, South Shetland Islands, 8oo-(o) m, 7 April 1927. COLLECTION: " Discovery " Committee. Allogaussia navicula Barnard 1932 : 65. SYNTYPES: Reg. no. 1936 : n : 2 : 340-345* (4 specimens, alcohol). LOCALITY : Sta. 175. Bransfield Strait, South Shetland Islands, 63° 17' S 59° 48' W, 200 m, 2 March 1927. SYNTYPES: Reg. no. 1936 : n : 2 : 337-339 (7 specimens, alcohol). LOCALITY: Sta. 170. Off Cape Bowles, Clarence Island, South Shetland Islands, 61° 25' S 53° 46' W, 342 m, 23 February 1927. COLLECTION: " Discovery " Committee. AMARYLLIS Haswell Amaryllis bathycephala Stebbing 1888 : 699. HOLOTYPE: Reg. no. 1889 : 5 : 15 : 30 (specimen in alcohol + 4 slides). LOCALITY: Sta. 161. Off Melbourne, Australia, 33 fm, i April 1874. COLLECTION: " Challenger ". Amaryllis tenuipes (Walker) 1904 : 241 [Vijaya tenuipes]. SYNTYPES: Reg. no. 1905 : 2 : 18 : 10 (i specimen, alcohol). 1905 : 2 : 18 : 308-309 (2 slides). LOCALITY : Ceylon, shallow water. COLLECTION: Herdman. AMBASIOPSIS Barnard A mbasiopsis georgiensis Barnard 1931 : 425. SYNTYPES: Reg. no. 1936 : n : 2 : 136-137 (3 specimens, alcohol). LOCALITY: Sta. 159. 53° 52' S 36° 08' W, 320 m, 21 January 1927. COLLECTION: " Discovery " Committee. Ambasiopsis uncinata Barnard 1932 : 46. SYNTYPES: Reg. no. 1936 : u : 2 : 146-148 (4 specimens, alcohol). LOCALITY: Sta. 170. Off Cape Bowles, Clarence Island, South Shetland Islands, 61° 25' S 53° 46' W, 342 m, 23 February 1927. COLLECTION: " Discovery " Committee. ANONYX Kr0yer \Anonyx ampulloides] Bate 1862 : 78 (transferred to Anonyx nugax (Phipps) by Steele & Brunei, 1968). HOLOTYPE: Reg. no. 68 : 107 (15) (alcohol). LOCALITY: Japan. COLLECTION: Stimpson. AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 355 Anonyx annulatus Bate 1862 : 79 (removed from Anonyx by Steele & Brunei, 1968, but correct generic position not indicated). HOLOTYPE: Reg. no. 68 : 107 (13) (alcohol). LOCALITY: Japan. COLLECTION: Stimpson. Anonyx bispinosus Steele 1967 : 257. HOLOTYPE: Reg. no. 1966 : 3 : 28 : 1-2 (part) (i specimen in alcohol -f 12 slides). PARATYPE: Reg. no. 1966 : 3 : 28 : 1-2 (part) (i specimen, alcohol). LOCALITY : Off E coast of Kolguev Island, Barents Sea. COLLECTION: Stebbing. Anonyx exiguus Stimpson 1853 : 51. SYNTYPE: Reg. no. 68 : 107 (29) (i specimen, alcohol). LOCALITY: Off Cheney's Head, Grand Manan, Bay of Fundy, New Brunswick. COLLECTION: Stimpson. Anonyx indicus Giles 1890 : 69. SYNTYPE: Reg. no. 1968 : 129 (i slide). LOCALITY: Off the Seven Pagodas, Madras, India, 5-10 fm. COLLECTION: Giles. [Anonyx melanophthalmus] Norman 1867 : 201 (transferred to Orchomene hanseni Meinert by Sars 1895). SYNTYPES: Reg. no. 1911 : n : 8 : 13346 (i specimen, alcohol). 1911 : ii : 8 : M727-M729 (3 slides). LOCALITY : Sleat Sound, Isle of Skye, Scotland, 1866. COLLECTION: Norman. Anonyx punctatus Bate 1862 : 78 (removed from Anonyx by Steele & Brunei, 1968, but correct generic position not indicated). HOLOTYPE: Reg. no. 68 : 107 (14) (alcohol). LOCALITY: Behring Strait. COLLECTION: Stimpson. ARISTIAS Boeck Aristias antarcticus Walker 1906 : 454. SYNTYPES: Reg. no. 1907 : 6 : 6 : 92-93 (2 specimens, alcohol). LOCALITY : McMurdo Sound, Ross Sea, 20 fm, 28 February 1902. SYNTYPES: Reg. no. 1907 : 6 : 13 : 6-7 (2 slides). LOCALITY: no data, 5 June 1902. COLLECTION : National Antarctic Expedition. Aristias collinus Barnard 1932 : 44. SYNTPYE: Reg. no. 1936 : n : 2 : 134* (i specimen, alcohol). 356 M. H. THURSTON & ELIZABETH ALLEN LOCALITY: Sta. 170. Off Cape Bowles, Clarence Island, South Shetland Islands, 61° 25' S 53° 46' W, 342 m, 23 February 1927. SYNTYPE: Reg. no. 1936 : n : 2 : 135 (i specimen, alcohol). LOCALITY: Sta. 175. Bransfield Strait, South Shetland Islands, 63° if S 59° 48' W 200 m, 2 March 1927. COLLECTION: " Discovery " Committee. Aristias pacificus Schellenberg 1936 : 153. SYNTYPES: Reg. no. 1967 : I : 10 : 17-42 (26 specimens, alcohol). LOCALITY: NE corner of Union Bay, Vancouver, British Columbia, 16-20 fm, 4 July 1923 (In atrial cavity of Ascidiopsis paratropa). COLLECTOR: Prof. C. H. O'Donoghue. Aristias symbiotica Barnard 1916 : 121. SYNTYPES: Reg. no. 1928 : 4 : 13 : 16-20 (6 specimens, alcohol). LOCALITY: False Bay, South Africa. Cape Hangklip bearing NE ^E, distance 27 miles, 105 fm, 26 February 1902. (From branchial cavity of Ascidians). DONOR : South African Museum. Chevreux Aroui setosus Chevreux 1910 : 170. SYNTYPES: Reg. no. 1912 : 4 : 4 : 5-7 (3 specimens, alcohol). LOCALITY : NE of Cap de Garde, Golfe de Bone, Algeria, 65 m. On Spatangus sp. COLLECTOR: E. Chevreux. ARUGELLA Pirlot Arugella falklandica (Barnard) 1932 : 39 [Lysianassa falklandica], SYNTYPES: Reg. no. 1936 : n : 2 : 112* (i specimen, alcohol). LOCALITY: Sta. WS 85. 8 miles S66° E of Lively Island, East Falkland, Falkland Islands, 79 m, 25 March 1927. SYNTYPES: Reg. no. 1936 : n : 2 : 113-114 (2 specimens, alcohol). LOCALITY: Sta. WS 97. South Atlantic, 49°oo' S 61° 58' W, 146-145 m, 18 April 1927. COLLECTION: " Discovery " Committee. [AUSTRONISIMUS Barnard] [Austronisimus rhinoceros] Barnard 1931 : 425 (transferred to Paralysianopsis odhneri Schellen- berg by Barnard 1932). HOLOTYPE: Reg. no. 1936 : n : 2 : 108 (alcohol). LOCALITY: Sta. WS 25. Undine Harbour, South Georgia, 18-27 m, *7 December 1926. COLLECTION: " Discovery " Committee. AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 357 BATH? AMARYLLIS Pirlot Bathyamaryllis haswelli (Stebbing) 1888 : 703 {Amaryllis haswelli}. HOLOTYPE : Reg. no. 1889 : 5 : 15 : 31 (specimen in alcohol + 8 slides). LOCALITY: Sta. 78. Off the Azores, 37° 26' N 25° 13' W, 1000 fm, 10 July 1873. COLLECTION : " Challenger ". CHEIRIMEDON Stebbing Cheirimedon crenatipalmatus Stebbing 1888 : 638. HOLOTYPE: Reg. no. 1889 : 5 : 15 : 13 (4 slides). LOCALITY: Sta. 149!!. Off Cumberland Bay, Kerguelen, South Indian Ocean, 127 fm, 29 January 1874. COLLECTION: " Challenger ". Cheirimedon fougneri Walker 1903 : 41. SYNTYPES: Reg. no. 1903 : 10 : 15 : 13-15 (9 specimens in alcohol + n slides). LOCALITY: Ross Sea, 78° 35'S, surface, 18 February 1900. COLLECTION: " Southern Cross ". [Cheirimedon hanseni] Walker 1903 : 42 (transferred to Cheirimedon crenatipalmatus Stebbing by Schellenberg 1926). SYNTYPES: Reg. no. 1903 : 10 : 5 : 20-22 (8 specimens in alcohol + i slide). LOCALITY: Cape Adare, South Victoria Land, 7 fm, i November 1899. COLLECTION: " Southern Cross ". CHIRONESIMUS Sars Chironesimus rotundatus (Barnard) 1925 : 329 [Lakota rotundatus]. SYNTYPES: Reg. no. 1928 : 4 : 13 : 28-35 (10 specimens, alcohol). LOCALITY: South Africa. DONOR: South African Museum. CRYBELOCEPHALUS Tattersall Crybelocephalus megalurus Tattersall 1906 : 33. SYNTYPE: Reg. no. 1947 : n : 4 : 18 (i specimen, alcohol). LOCALITY: 50 miles N by W of Eagle Island, Co. Mayo, Eire, 1150 fm, May 1905. COLLECTION : Tattersall. CYCLOCARIS Stebbing [Cyclocaris faroensis'] Norman 1900 : 197 (transferred to Cyclocaris guilelmi Chevreux by Stebbing 1906). SYNTYPE: Reg. no. 1911 : n : 8 : 12755 (i specimen, alcohol). 1911 : ii : 8 : M686 (i slide). 358 M. H. THURSTON & ELIZABETH ALLEN LOCALITY: "Triton" Sta. 8. Faeroe Channel, 60° 18' N 06° 15' W, tow net at 640 fm, 1882. COLLECTION: Norman. Cyclocaris tahitensis Stebbing 1888 : 664. HOLOTYPE : Reg. no. 1889 : 5 : 15 : 20 (3 slides). LOCALITY: Off Tahiti, 2 October 1875. [This corresponds with Sta. 279. Off Papeete, Tahiti, 420-680 fm.] COLLECTION: " Challenger ". CYPHOCARIS Boeck Cyphocaris challengeri Stebbing 1888 : 661. HOLOTYPE: Reg. no. 1889 : 5 : 15 : 19 (4 slides). LOCALITY : 400 miles N of the Sandwich Islands. [Probably near Sta. 256.] COLLECTION: " Challenger ". [Cyphocaris micronyx] Stebbing 1888 : 656 (transferred to Cyphocaris anonyx by Stebbing 1906). SYNTYPE: Reg. no. 1889 : 5 : 15 : 17 (4 slides). LOCALITY: Sta. 295. Off the West coast of South America, 38° 07' S 94° 04' W, 1500 fm, 5 November 1875. SYNTYPE: Reg. no. 1889 : 5 : 15 : 18 (i slide). LOCALITY: Sta. 335. Near Tristan da Ctmha, 32° 24' S 13° 05' W, 1425 fm, 6 March 1876. COLLECTION: " Challenger ". EUONYX Norman Euonyx chelatus Norman 1867 : 202. SYNTYPES: Reg. no. 1911 : n : 8 : 13684-13686 (3 specimens, alcohol) 1911 : ii : 8 : M843-M845 (3 slides). LOCALITY: Sleat Sound, Isle of Skye, Scotland, July 1866. COLLECTION: Norman. Euonyx normani Stebbing 1888 : 669. HOLOTYPE: Reg. no. 1889 : 5 : 15 : 21 (specimen in alcohol + 8 slides). LOCALITY: Sta. i7oA. Near Kermadec Island, South Pacific, 29° 45' S 178° n' W, 630 fm, 14 July 1874. COLLECTION: "Challenger". HIPPOMEDON Boeck Hippomedon denticulatus (Bate) 1857 : 139 {Anonyx denticulata}. SYNTYPES: Reg. no. 1952 : 5 : 7 : 8-9 (6 specimens, alcohol). LOCALITY: Moray Firth. COLLECTION: Bate. AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 359 Hippomedon geelongi Stebbing 1888 : 635. HOLOTYPE: Reg. no. 1889 : 5 : 15 : 12 (specimen in alcohol + 7 slides). LOCALITY: Sta. 161. Off Melbourne, Australia, 33 fm, i April 1874. COLLECTION: "Challenger". Hippomedon holbolli var. abyssicola Sars 1885 : 142. SYNTYPE: Reg. no. 1911 : u : 8 : 12925 (i specimen, alcohol). LOCALITY: Norwegian North Atlantic Expedition Sta. 40. 63° 22' N 05° 29' W, 2222 m, 18 July 1876. COLLECTION : Norman. Hippomedon incisus Barnard 1930 : 325. SYNTYPES: Reg. no. 1930 : 8 : i : 16-17 (2 specimens in alcohol + i slide). LOCALITY: Sta. 242. Off Akaroa Heads, New Zealand, 10 m, 2 April 1912. COLLECTION: " Terra Nova ". Hippomedon longimanus (Stebbing) 1888 : 643 [Platamon longimanus']. HOLOTYPE: Reg. no. 1889 : 5 : 15 : 14 (part) (specimen in alcohol + 7 slides). PARATYPE: Reg. no. 1889 : 5 : 15 : 14 (part) (i specimen, alcohol). LOCALITY: Sta. i. Off Cape Finisterre, Spain, 41° 58' N 09° 42' W, 1125 fm, 30 December 1872. COLLECTION: " Challenger ". ICHNOPUS Costa [Ichnopus affinis] Heller 1867 : 19 (transferred to Ichnopus taurus Costa by Delia Valle 1893). SYNTYPE: Reg. no. 1911 : u : 8 : M655 (i slide). LOCALITY: Adriatic Sea, 1866. COLLECTION: Norman. [Ichnopus serricrus] Walker 1909 : 328 (transferred to Ichnopus taurus Costa by Spandl 1924)- SYNTYPES: Reg. no. 1909 : i : 29 : 14-16 (2 specimens, alcohol). LOCALITIES: Coco Is., Cargados Islands, Indian Ocean, surface, 30 August 1905. Amirante Is., Indian Ocean, 22-85 fm, n October 1905. SYNTYPE: Reg. no. 1909 : 2 : 13 : 3 (i slide). LOCALITY: Amirante Islands, Indian Ocean, 25-85 fm, n October 1905. COLLECTION: " Sealark " Expedition. KERGUELENIA Stebbing Kerguelenia antarctica Barnard 1930 : 318. SYNTYPES: Reg. no. 1930 : 8 : i : 1-2 (2 specimens in alcohol + i slide). LOCALITY: Sta. 356. McMurdo Sound, Ross Sea, 92 m, 22 January 1913. COLLECTION: " Terra Nova ". 360 M. H. THURSTON & ELIZABETH ALLEN Kerguelenia compacta Stabbing 1888 : 1220. SYNTYPES: Reg. no. 1889 : 5 115 : 159 (i specimen in alcohol + 2 slides). LOCALITIES: Off Christmas Harbour, Kerguelen, South Indian Ocean, 120 fm. Sta. I49H. Off Cumberland Bay, Kerguelen, South Indian Ocean, 127 fm, 29 January 1874. COLLECTION: " Challenger ". Kerguelenia palpalis Barnard 1932 : 28. HOLOTYPE: Reg. no. 1936 : n : 2 : i (alcohol). LOCALITY: Sta. 175. Bransfield Strait, South Shetland Islands, 63° 17' S 59° 48' W, 200 m, 2 March 1927. COLLECTION: " Discovery " Committee. LEPIDEPECREELLA Schellenberg Lepidepecreella bidens (Barnard) 1930 : 321 [Paracyclocaris bidens]. SYNTYPES: Reg. no. 1930 : 8 : i : 5-8 (7 specimens in alcohol + i slide). LOCALITY: Sta. 90. Off Three Kings Islands, New Zealand, 183 m, 25 July 1911. COLLECTION: " Terra Nova ". Lepidepecreella ovalis Barnard 1932 : 61. SYNTYPES: Reg. no. 1936 : ir : 2 : 329-330* (2 specimens, alcohol). LOCALITY: Sta. 45. 2-7 miles S 85° E of Jason Light, South Georgia, 238-270 m, 6 April 1926. SYNTYPES: Reg. no. 1936 : n : 2 : 331-333 (1+2+2 specimens, alcohol). LOCALITIES: Sta. 42. Off Cumberland Bay, South Georgia, 120-204 m, i April 1926. Sta. 140. Stromness Harbour to Larsen Point, South Georgia, 122-136 m, 23 December 1926. Sta. WS 33. 54° 59' S 35° 24' W, 130 m, 21 December 1926. COLLECTION: " Discovery " Committee. LEPIDEPECREOIDES Barnard Lepidepecreoides xenopus Barnard 1931 : 426. SYNTYPE: Reg. no. 1936 : n : 2 : 334* (i specimen, alcohol). LOCALITY: Sta. 181. Schollaert Channel, Palmer Archipelago, Graham Land, 160-335 m» I2 March 1927. SYNTYPES: Reg. no. 1936 : n : 2 : 335-336 (i + i specimens, alcohol). LOCALITIES: Sta. 177. 27 miles SW of Deception Island, South Shetland Islands, 1080 m, 5 March 1927. Sta. WS 33. 54° 59' S 35° 24' W, 130 m, 21 December 1926. COLLECTION: " Discovery " Committee. AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 361 LEPIDEPECREUM Bate & Westwood Lepidepecreum cingulatum Barnard 1932 : 60. SYNTYPES: Reg. no. 1936 : n : 2 : 326-328 (4 specimens, alcohol). LOCALITY: Sta. 165. Dove Strait, Signy Island, South Orkney Islands, 24-36 m, 18-20 February 1927. COLLECTION: " Discovery " Committee. Lepidepecreum foraminiferum Stebbing 1888:686. SYNTYPES: Reg. no. 1889 : 5 : 15 : 26 (4 specimens in alcohol + 6 slides). LOCALITY: Sta. I49H. Off Cumberland Bay, Kerguelen, 127 fm, 29 January 1874. COLLECTION: " Challenger ". LYSIANASSA Milne Edwards Lysianassa coelochir (Walker) 1904 : 243 [Lysianax coelochir]. SYNTYPES: Reg. no. 1905 : 2 : 18 : 17-18 (2 specimens, alcohol). 1905 : 2 : 18 : 312 (i slide). LOCALITY: Ceylon. COLLECTION: Herdman. Lysianassa urodus (Walker & Scott) 1903 : 220 [Lysianax urodus]. SYNTYPES: Reg. no. 1914 : 3 : 31 : 7-8 (2 specimens, alcohol). LOCALITY: Abd-el-Kuri, Socotra. DONOR: A. O. Walker. Lysianassa variegata (Stimpson) 1856 : 394 [Anonyx variegata]. SYNTYPE : Reg. no. 68 : 107 (16) (i specimen). LOCALITY: Simon's Bay, Cape of Good Hope. COLLECTION: Stimpson. [Lysianassa woodmasoni] Giles 1890 : 68 (transferred to Glycerina tenuicornis (Haswell) by Pirlot 1936). SYNTYPES: Reg. no. 1968 : 53 (i slide). LOCALITY : Macpherson's Strait, Andaman Islands, 17 fm. COLLECTION: Giles. METACYPHOCARIS Tattersall Metacyphocaris helgae Tattersall 1906 : 29. SYNTYPE: Reg. no. 1911 : n : 8 : 13692 (i specimen, alcohol). LOCALITY : 40 miles N by W of Eagle Island, Co. Mayo, Eire, 630 fm, February 1905. COLLECTION: Norman. ZOOL. 17, 9. 23§§§ 362 M. H. THURSTON & ELIZABETH ALLEN SYNTYPES: Reg. no. 1947 : n : 4 : 11-13 (4 specimens, alcohol). LOCALITY: 40 miles N by W of Eagle Island, Co. Mayo, Eire, 750 fm, n August 1904. SYNTYPES: Reg. no. 1947 : n : 4 : 14-16 (3 specimens, alcohol). LOCALITY: 53° 07' N 15° 06' W, 750 fm, 12 May 1905. SYNTYPE: Reg. no. 1947 : n : 4 : 17 (i specimen, alcohol). LOCALITY: 50 miles N by W of Eagle Island, Co. Mayo, Eire, 600 fm, 20 May 1905. COLLECTION : Tattersall. MICROLYSIAS Stebbing Microlysias indica Barnard 1937 : J44- SYNTYPES: Reg. no. 1938 : i : 3 : 19-22 (5 specimens, alcohol). LOCALITY: Sta. 53. South Arabian Coast, 19° 23' N 57° 53' E, 13-5 m, 2 November 1933- COLLECTION: " John Murray ". Microlysias xenokeras Stebbing 1918 : 64. SYNTYPES: Reg. no. 1928 : 12 : i : 2055-2061 (43 specimens, alcohol). LOCALITY: Vetch's Pier, Durban, South Africa, 2 fm, 9 July 1917. From sea squirt. DONOR: T. R. R. Stebbing. NANNONYX Sars Nannonyx integricauda (Stebbing) 1888 : 695 [Ambasia integricauda]. HOLOTYPE: Reg. no. 1889 : 5 : 15 : 29 (4 slides). LOCALITY: Sta. 1490. Royal Sound, Kerguelen, 28 fm, 20 January 1874. COLLECTION: " Challenger ". Nannonyx spinimanus Walker 1895 : 292. SYNTYPE: Reg. no. 1914 : 3 : 31 : 9 (i specimen, alcohol). LOCALITY: Pwll Fanog, Menai Strait, Wales, 10-12 fm, 2 April 1894. SYNTYPES: Reg. no. 1907 : 4 : 8 : 5 (i slide). 1908 : 3 : 10 : 7 (i slide). LOCALITY: Menai Bridge, Wales, 7-10 fm, 31 May 1894. SYNTYPE: Reg. no. 1908 : 3 : 10 : 8 (i slide). LOCALITY: Britannia Bridge, Menai Strait, Wales, 7-10 fm, 31 May 1894. DONOR: A. O. Walker. The depths given here are those on the bottle or slide label. In his report Walker gives the depth as 5-8 fms. AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 363 NORMANION Bonnier [Normanion amblyops] Sars 1895 : 674 (transferred to Normanion quadrimanus by Stebbing 1906). SYNTYPES: Reg. no. 1911 : n : 8 : 12768-12772 (5 specimens, alcohol). LOCALITY: R0dbjerget, Trondhjemsfjord, Norway, 200-300 fm. On skins of live Gadus aeglefinus [now Melanogrammus aeglefinus (L)] and Spinax niger [now Etmopterus spinax (L)]. COLLECTION: Norman. Normanion quadrimanus (Bate and Westwood) 1868 : 503 [Opis quadrimanus]. SYNTYPE: Reg. no. 1911 : n : 8 : M654 (x slide). LOCALITY : Coast of Banffshire, 1864, collected Mr. Edward. COLLECTION: Norman. ONESIMOIDES Stebbing Onesimoides carinatus Stebbing 1888 : 648. HOLOTYPE : Reg. no. 1889 : 5 : 15 : 15 (specimen in alcohol + 6 slides). LOCALITY: Sta. 184. Off the NE coast of Australia, 12° 08' S 145° 10' E, 1400 fm, 29 August 1874. COLLECTION : " Challenger ". ONISIMUS Boeck Onisimus brevicaudatus Hansen 1887 : 216. SYNTYPES: Reg. no. 1891 : i : 6 : 21-25 (5 specimens, alcohol). LOCALITY: Kara Sea, 71° 13' N, no m, " Dijmphna " Expedition. DONOR: Copenhagen Museum. SYNTYPES: Reg. no. 1911 : n : 8 : 13644-13648 (5 specimens, alcohol). LOCALITY: Kara Sea, 71° 13' N, no m, " Dijmphna " Expedition. COLLECTION: Norman. SYNTYPES: Reg. no. 1928 : 12 : i : 1623-1628 (7 specimens, alcohol). LOCALITY: Kara Sea, 71° 13' N, no m, " Dijmphna " Expedition. COLLECTION: Stebbing. Onisimus caricus Hansen 1887 : 214. SYNTYPES: Reg. no. 1891 : i : 6 : 18-20 (3 specimens, alcohol). LOCALITY: Kara Sea, " Dijmphna " Expedition. DONOR: Copenhagen Museum. SYNTYPES: Reg. no. 1911 : n : 8 : 13649-13652 (4 specimens, alcohol). LOCALITY: Kara Sea, " Dijmphna " Expedition. COLLECTION : Norman. 364 M. H. THURSTON & ELIZABETH ALLEN SYNTYPES: Reg. no. 1928 : 12 : i : 1615-1622 (part) (4 specimens, alcohol). LOCALITY: Kara Sea, " Dijmphna " Expedition. COLLECTION: Stebbing. ORCHOMENE Boeck Orchomene amblyops Sars 1890 : 65. SYNTYPES: Reg. no. 1911 : n : 8 : 13087-13096 (n specimens, alcohol). LOCALITY: Trondhjemsfjord, Norway. COLLECTION: Norman. The absence of a date on the label of these specimens prevents absolute certainty as to their type status. Orchomene batei Sars 1882 : 81. SYNTYPES : Reg. no. 1911 : n : 8 : M73I-732 (2 slides). LOCALITY: West Norway. COLLECTION: Norman. The absence of a date, both on the slides and in the accession register, makes it impossible to be certain that these specimens are part of Sars' type series. Orchomene goniops Walker 1906 : 455. SYNTYPES: Reg. no. 1907 : 6 : 6 : 94-95 (2 specimens, alcohol). 1907 : 6 : 13 : 8 (i slide). LOCALITY: McMurdo Sound, Ross Sea, 21 August 1903. COLLECTION : National Antarctic Expedition. [Orchomene musculosus] Stebbing 1888 : 673 (transferred to Orchomenella chilensis (Heller) by Chilton 1912). HOLOTYPE: Reg. no. 1889 : 5 : 15 : 22 (specimen in alcohol + 6 slides). LOCALITY: Sta. 230. South of Japan, 26° 29' N 137° 57' E, surface, 5 April 1875. COLLECTION: " Challenger ". ORCHOMENELLA Sars Orchomenella abyssorum (Stebbing) 1888 : 676 [Orchomene abyssorum]. HOLOTYPE: Reg. no. 1889 : 5 : 15 : 23 (alcohol, dissected). LOCALITY : Sta. 323. South Atlantic Ocean, E of Buenos Aires, 35° 39' S 50° 47' W, 1900 fm. COLLECTION: " Challenger ". Orchomenella cavimanus (Stebbing) 1888 : 679 [Orchomene cavimanus]. HOLOTYPE: Reg. no. 1889 : 5 : 15 : 24 (part) (specimen in alcohol -f 7 slides). LOCALITY: Kerguelen, South Indian Ocean, no further data. PARATYPES: Reg. no. 1889 : 5 : 15 : 24 (part) (i + i specimens, alcohol). LOCALITIES: Betsy Cove, Kerguelen, South Indian Ocean, surface. Sta. I49H. Off Cumberland Bay, Kerguelen, South Indian Ocean, 127 fm. COLLECTION: " Challenger ". AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 365 Orchomenella chelipes Walker 1906 : 456. SYNTYPES: Reg. no. 1907 : 6 : 6 : 136-143 (4 specimens, alcohol). 1907 : 6 : 13 : 9 (i slide). LOCALITY: McMurdo Sound, Ross Sea, 28 February 1902. COLLECTION : National Antarctic Expedition. Orchomenella chilensis (Heller) 1868 : 129 [Anonyx chilensis]. PARALECTOTYPES : Reg. no. 1966 : 8 : 14 : 1-24 (24 specimens, alcohol). LOCALITY: Chile, " Novara " Expedition. DONOR : Vienna Museum. Orchomenella franklini Walker 1903 : 47. HOLOTYPE: Reg. no. 1903 : 10 : 5 : 10 (alcohol). LOCALITY: Franklin Island, Ross Sea. COLLECTION: " Southern Cross ". Orchomenella nodimanus (Walker) 1903 : 44 \Orchomenopsis nodimanus']. SYNTYPES: Reg. no. 1902 : n : 5 : 46-56 (ca 100 specimens in alcohol + 2 slides). LOCALITY: Cape Adare, 26 fm, 4-5 November 1899. COLLECTION: " Southern Cross". Orchomenella pinguides Walker 1903 : 46. SYNTYPES: Reg. no. 1902 : n : 5 : 39-45 (ca 130 specimens in alcohol + 3 slides). LOCALITY : Cape Adare, 26 fm, 4-5 November 1899. COLLECTION : " Southern Cross ". Orchomenella plebs Hurley 1965 : 107. PARATYPES: Reg. no. 1968 : 128 (12 specimens, alcohol). LOCALITY: Sta. A. 32. White Island, Ross Ice Barrier, 15 November 1961. DONOR: D. E. Hurley. Orchomenella rotundifrons Barnard 1932 : 72. SYNTYPES: Reg. no. 1936 : n : 2 : 523-529 (n specimens, alcohol). LOCALITY: Sta. 174. Deception Island, South Shetland Islands, 5-10 m, 28 February — 2 March 1927. COLLECTION: " Discovery " Committee. [ORCHOMENOPSIS Sars] [Orchomenopsis proximo] Chevreux 1903 : 93 (transferred to Orchomenella chilensis (Heller) by Chilton 1912). SYNTYPES: Reg. no. 1965 : 12 : 2 : 1-8 (16 specimens, alcohol). LOCALITY: " Princesse Alice" Sta. 1138. Off the Cape Verde Islands, i6°44'N 27° 08' W, 692 m, 21-22 July 1901. DONOR : Monaco Museum. 366 M. H. THURSTON & ELIZABETH ALLEN [Orchomenopsis rossi] Walker 1903 : 45 (transferred to Orchomenella chilensis (Heller) by Chilton 1912). SYNTYPES: Reg. no. 1902 : n : 5 : 85-90 (31 specimens in alcohol + i slide). LOCALITY: Ross Sea, 78° 35' S, near surface, 18 February 1900. COLLECTION: " Southern Cross ". PERRIERELLA Chevreux & Bouvier Perrierella audouiniana (Bate) 1857 : J38 \Lysianassa audouiniana}. HOLOTYPE : Reg. no. 1952 : 5 : 7 : i (alcohol). LOCALITY: Plymouth Sound, England. COLLECTION: Bate. PODOPRION Chevreux Podoprion bolivari Chevreux 1891 : 6. SYNTYPES: Reg. no. 1907 : 12 : 2 : 100-103 (4 specimens, alcohol). LOCALITY: Vigo Bay, Spain. DONOR : University College, Dundee. SYNTYPES: Reg. no. 1911 : n : 8 : 13660 (i specimen, alcohol). 1911 : ii : 8 : 13661-13665 (5 specimens, alcohol). LOCALITY : Vigo Bay, Spain, 20 m. COLLECTION: Norman. SYNTYPES: Reg. no. 1928 : 12 : i : 1602-1604 (4 specimens, alcohol). LOCALITY: Vigo Bay, Spain. COLLECTION: Stebbing. PODOPRIONIDES Walker Podoprionides incerta Walker 1906 : 457. HOLOTYPE: Reg. no. 1907 : 6 : 6 : 200 (i specimen, alcohol). 1907 : 6 : 13 : 10 (i slide). LOCALITY: Hole 12, Winter Quarters, McMurdo Sound, Ross Sea, 29 August 1902. COLLECTION : National Antarctic Expedition. PSEUDALIBROTUS Delia Valle Pseudalibrotus caspius Sars 1896 : 422. SYNTYPES: Reg. no. 1911 : 11 : 8 : 13599-13600 (2 specimens, alcohol). LOCALITY : Central Caspian Sea, 250 fm. COLLECTION: Norman. AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 367 PSEUDORCHOMENE Schellenberg Pseudorchomene coatsi (Chilton) 1912 : 477 [Orchomenopsis coatsi]. SYNTYPES: Reg. no. 1912 : 10 : 5 : 4-18 (15 specimens, alcohol). LOCALITY: Sta. 411. Off Coats Land, 74° 01' S 22° oo' W, 161 fm, 12 March 1904. COLLECTION: " Scotia ". DONOR: Dr. W. S. Bruce. SYNTYPES: Reg. no. 1912 : 10 : 14 : 49-53 (7 specimens, alcohol). LOCALITY: Sta. 411. Off Coats Land, 74° 01' S 22° oo' W, 161 fm, 12 March 1904. COLLECTION: " Scotia ". DONOR: Prof. C. Chilton. SCOPELOCHEIRUS Bate Scopelocheirus crenatus Bate 1857 : 138. SYNTYPES: Reg. no. 1952 : 5 : 7 : 13 (26 specimens, alcohol). LOCALITY: Plymouth Sound. COLLECTION: Bate. SHACKLETONIA Barnard Shackletonia robusta Barnard 1932 : 29. HOLOTYPE: Reg. no. 1936 : n : 2 : 2 (alcohol). LOCALITY: Sta. 170. Off Cape Bowles, Clarence Island, South Shetland Islands, 61° 25' S 53° 46' W, 342 m, 23 February 1927. PARATYPE: Reg. no. 1936 : u : 2 : 3 (i specimen, alcohol). LOCALITY: Sta. 123. Off mouth of Cumberland Bay, South Georgia, 230-250 m, 15 December 1926. COLLECTION: " Discovery " Committee. SOCARNELLA Walker Socarnella bonnieri Walker 1904 : 239. SYNTYPES: Reg. no. 1905 : 2 : 18 : 306-307 (2 slides). LOCALITY: Galle, Ceylon. On reef with ascidians. COLLECTION: Herdman. SOCARNES Boeck Socarnes erythrophthalmus Robertson 1892 : 200. SYNTYPES: Reg. no. 1911 : n : 8 : 12714-12728 (15 specimens, alcohol). 1911 : u : 8 : M682 (i slide). LOCALITY: Cumbrae, Firth of Clyde, Scotland, 1888. COLLECTION: Norman. 368 M. H. THURSTON & ELIZABETH ALLEN SOCARNOIDES Stebbing Socarnoides kergueleni Stebbing 1888 : 691. SYNTYPES: Reg. no. 1889 : 5 : 15 : 27 (2 specimens in alcohol + 5 slides). LOCALITY: Sta. i4gE. Greenland Harbour, Kerguelen, South Indian Ocean, 30 fm, 21 January 1874. SYNTYPE: Reg. no. 1889 : 5 : 15 : 28 (i specimen in alcohol + 2 slides). LOCALITY: Sta. 149!!. Off Cumberland Bay, Kerguelen, South Indian Ocean, 127 fm, 29 January 1874. COLLECTION: " Challenger ". SOCARNOPSIS Chevreux Socarnopsis crenulata Chevreux 1910 : 165. SYNTYPES: Reg. no. 1912 : 4 : 4 : 1-4 (4 specimens, alcohol). LOCALITY: Bougie, Algeria. DONOR: E. Chevreux. SOPHROSYNE Stebbing Sophrosyne murrayi Stebbing 1888 : 652. HOLOTYPE: Reg. no. 1889 : 5 : 15 : 16 (specimen in alcohol + 2 slides). LOCALITY : Off Christmas Harbour, Kerguelen, South Indian Ocean. COLLECTION: " Challenger ". STOMACONTION Stebbing Stomacontion insigne Barnard 1932 : 33. HOLOTYPE: Reg. no. 1936 : n : 2 : 9 (alcohol). 1936 : ii : 2 : 3608 (alcohol, mouth parts relaxed from slide.) LOCALITY : Sta. 160. Near Shag Rocks, South Atlantic Ocean, 53° 43' S 40° 57' W, 177 m, 7 February 1927. COLLECTION: " Discovery " Committee. Stomacontion kergueleni (Stebbing) 1888 : 720 [Acontiostoma HOLOTYPE: Reg. no. 1889 : 5 : 15 : 36 (4 slides). LOCALITY: Sta. 1490. Royal Sound, Kerguelen, South Indian Ocean, 28 fm, 20 January 1874. COLLECTION : " Challenger ". Stomacontion pepinii (Stebbing) 1888 : 716 [Acontiostoma pepinii]. SYNTYPES: Reg. no. 1889 : 5 : 15 : 35 (2 specimens in alcohol + 5 slides). LOCALITY: Sta. 1496. Royal Sound, Kerguelen, South Indian Ocean, 28 fm, 20 January 1874. COLLECTION : " Challenger ". AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 369 TMETONYX Stebbing Tmetonyx bruuni Reid 1951 : 196. PARATYPES: Reg. no. 1961 : 8 : i : 487-495 (9 specimens, alcohol). LOCALITY: Sta. 160. Off Bathurst, Gambia, West Africa, 14 m, 24 April 1946. COLLECTION: " Atlantide ". Tmetonyx cicadoides (Stebbing) 1888 : 612 [Anonyx cicadoides]. SYNTYPES: Reg. no. 1889 : 5 : 15 : 3-5 (16 specimens in alcohol + 17 slides). LOCALITIES : Sta. 149. Accessible Bay, Kerguelen, South Indian Ocean, 20 fm, 9 January 1874. Sta. 1490. Royal Sound, Kerguelen, South Indian Ocean, 28 fm, 20 January 1874. Sta. 149!!. Off Cumberland Bay, Kerguelen, South Indian Ocean, 127 fm, 29 January 1874. COLLECTION: " Challenger ". Tmetonyx longitelson Barnard 1932 : 54. SYNTYPES: Reg. no. 1936 : n : 2 : 317-319 (3 specimens, alcohol). LOCALITY: Sta. 159. 53° 52' S 36° 08' W, 160 m, 21 January 1927. COLLECTION: " Discovery " Committee. Tmetonyx miersi (Stebbing) 1888 : 631 [Hippomedon miersi]. HOLOTYPE: Reg. no. 1889 : 5 : 15 : n (i specimen in alcohol + 7 slides). LOCALITY: Sta. 162. Off East Moncoeur Island, Victoria, Australia, 39° 10' S 146° 37' E, 38 fm, 2 April 1874. COLLECTION : " Challenger ". Tmetonyx nobilis (Stimpson) 1853 : 50 [Anonyx nobilis]. SYNTYPES: Reg. no. 68 : 107 (9, 23) (3 + 2 specimens, alcohol). LOCALITY: Nantucket Island, Mass., U.S.A. COLLECTION: Stimpson. Tmetonyx stebbingi (Walker) 1903 : 52 \Hoplonyx stebbingi]. SYNTYPE: Reg. no. 1902 : n : 5 : 126 (i specimen, alcohol). LOCALITY : Cape Adare, South Victoria Land. SYNTYPES: Reg. no. 1903 : 10 : 5 : 11-12 (2 specimens in alcohol + i slide). LOCALITY: Cape Adare, South Victoria Land, 26 fm, 4-5 November 1899. COLLECTION: " Southern Cross ". Barnard (1958) on the authority of Chilton (1912) listed Hoplonyx stebbingi Walker under Tryphosites. It has since been shown (Hurley 19650) that two species were present, Tmetonyx stebbingi (Walker) and Tryphosites capadarei Hurley. TRISCHIZOSTOMA Boeck Trischizostoma serratum Barnard 1925 : 320. SYNTYPE: Reg. no. 1928 : 4 : 13 : 13 (i specimen, alcohol). LOCALITY : Off Natal, South Africa. DONOR: South African Museum. 370 M. H. THURSTON & ELIZABETH ALLEN TRYPHOSA Boeck Tryphosa adarei Walker 1903 : 49. SYNTYPES: Reg. no. 1902 : n : 5 : 127-135 (12 specimens in alcohol + 2 slides). 1903 : 10 : 5 : 16 (2 specimens, alcohol). LOCALITY: Cape Adare, South Victoria Land, 26 fm, 4-5 November 1899. COLLECTION: " Southern Cross ". Tryphosa analogica Barnard 1932 : 52. SYNTYPES: Reg. no. 1936 : n : 2 : 298-300* (3 specimens, alcohol). LOCALITY: Sta. WS 25. Undine Harbour, South Georgia, 18-27 m» r7 December 1926. SYNTYPES: Reg. no. 1936 : n : 2 : 301-305 (3+3 + 1 specimens, alcohol). LOCALITIES: Sta. 144. 53°48'S 35° 37' W, 728m, 8 January 1927. Sta. 156. 53° 5i' S 36° 21' W, 200-236 m, 20 January 1927. Sta. 159. 53° 52' S 36° 08' W, 160 m, 21 January 1927. COLLECTION: " Discovery " Committee. Tryphosa angulata Sars 1891 : 78. SYNTYPES: Reg. no. 1911 : n : 8 : 13257-13261 (5 specimens, alcohol). LOCALITY: Trondhjemsfjord, Norway. COLLECTION: Norman. No date is recorded on the bottle label or in the accession register, and without this, the type status of these specimens cannot be confirmed. [Tryphosa antennipotens] Stebbing 1888 : 617 (transferred to Uristes gigas Dana by Stebbing 1906). HOLOTYPE: Reg. no. 1889 : 5 : 15 : 6 (i specimen in alcohol + 6 slides). LOCALITY: Sta. 150. Off Heard Island, South Indian Ocean, 52° 04'$ 71° 22'E, 150 fm, 2 February 1874. COLLECTION: " Challenger ". Tryphosa castellata Barnard 1932 : 53. SYNTYPES: Reg. no. 1936 : ri : 2 : 306-307 (2 specimens, alcohol). LOCALITY: Sta. 222. St. Martin's Cove, Hermite Island, Cape Horn, 30-35 m, 23-24 April 1927. COLLECTION: " Discovery " Committee. [Tryphosa ciliata Sars] 1882 : 81 (transferred to Orchomenella nana (Krayer) by Bonnier 1893). SYNTYPE: Reg. no. 1911 : n : 8 : M752 (i slide). LOCALITY: South Norway. COLLECTION: Norman. This slide has no date on its label, so it is not possible to confirm the type status of the specimen. AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 371 Tryphosa cucullata Walker 1904 : 244. HOLOTYPE: Reg. no. 1905 : 2 : 18 : 25 (alcohol). LOCALITY: Kondatchi Paar, Ceylon, 17 November 1902. COLLECTION: Herdman. Tryphosa kergueleni (Miers) 1875 : 74 [Lysianassa kergueleni]. SYNTYPES: Reg. no. 76 : 27 (6 specimens, alcohol). LOCALITY: Royal Sound, Kerguelen, South Indian Ocean. DONOR: Rev. A. E. Eaton. Tryphosa major Barnard 1932 : 50. SYNTYPES: Reg. no. 1936 : n : 2 : 261-263* (4 specimens, alcohol). LOCALITY: Sta. 170. Off Cape Bowles, Clarence Island, South Shetland Islands, 61° 25' S 53° 46' W, 342 m, 23 February 1927. SYNTYPES: Reg. no. 1936 : u : 2 : 264-270 (7+3 specimens, alcohol). LOCALITIES: Sta. 159. 53° 52' S 36° 08' W, 160 m, 21 January 1927. Sta. 175. Bransfield Strait, South Shetland Islands, 63° 17' S 59° 48' W, 200 m, 2 March 1927. COLLECTION: " Discovery " Committee. Tryphosa murrayi Walker 1903 : 50. SYNTYPES: Reg. no. 1902 : n : 5 : 136-137 (2 specimens, alcohol). LOCALITY : Cape Adare, South Victoria Land. COLLECTION: " Southern Cross ". Tryphosa triangularis Barnard 1932 : 51. SYNTYPES: Reg. no. 1936 : n : 2 : 275-283* (10 specimens in alcohol + 5 slides). LOCALITY: Sta. 159. 53°52'S 36° 08' W, 160 m, 21 January 1927. SYNTYPES: Reg. no. 1936 : n : 2 : 284-297 (2+6+3 + 8+4 specimens, alcohol). LOCALITIES: Sta. 140. Stromness Harbour to Larsen Point, South Georgia, 122-136 m, 23 December 1926. Sta. 141. East Cumberland Bay, South Georgia, 17-27 m, 29 December 1926. Sta. 144. Off entrance of Stromness Harbour, South Georgia, 155-178 m, 5 January 1927. Sta. 145. Stromness Harbour, South Georgia, 26-35 m» 7 January 1927. Sta. WS 25. Undine Harbour, South Georgia, 18-27 m> 17 December 1926. COLLECTION: " Discovery " Committee. Tryphosa trigonica (Stebbing) 1888 : 630 \Hippomedon trigonicus']. SYNTYPE: Reg. no. 1889 : 5 : 15 : 10 (part) (i specimen in alcohol + 5 slides). LOCALITY : Kerguelen, South Indian Ocean, no further data. SYNTYPES: Reg. no. 1889 : 5 : 15 : 10 (part) (4 specimens, alcohol). LOCALITY: Kerguelen, South Indian Ocean, 120 fm. COLLECTION: " Challenger ". 372 M. H. THURSTON & ELIZABETH ALLEN SYNTYPES: Reg. no. 1928 : 12 : i : 1719-1721 (3 specimens, alcohol). LOCALITY : Kerguelen, South Indian Ocean, 30 fm. COLLECTION: Stebbing. TRYPHOSELLA Bonnier Tryphosella albino, Barnard 1932 : 48. HOLOTYPE: Reg. no. 1936 : n : 2 : 249 (alcohol). LOCALITY: Sta. 170. Off Cape Bowles, Clarence Island, 61° 25' S 53°46'W, 342 m, 23 February 1927. COLLECTION: " Discovery " Committee. Tryphosella barbatipes (Stebbing) 1888 : 621 \Tryphosa barbatipes]. SYNTYPES: Reg. no. 1889 : 5 : 15 : 7 (2 specimens in alcohol + 8 slides). LOCALITY: Sta. 149 H. Off Cumberland Bay, Kerguelen, South Indian Ocean, 127 fm, 29 January 1874. COLLECTION: " Challenger ". TRYPHOSITES Sars Tryphosites capadarei Hurley i9&5a : 177. HOLOTYPE: Reg. no. 1965 : 12 : 2 : 9 (alcohol). LOCALITY : Cape Adare, South Victoria Land, 26 fm, 4-5 November 1899. COLLECTION: " Southern Cross ". Tryphosites chevreuxi Stebbing 1914 : 355. SYNTYPES: Reg. no. 1928 : 12 : i : 1729-1737 (9 specimens, alcohol). LOCALITY: Roy Cove, Falkland Islands, 8 fm. COLLECTION: Stebbing. VALETTIA Stebbing Valettia coheres Stebbing 1888 : 724. HOLOTYPE : Reg. no. 1889 : 5 : 15 : 37 (specimen in alcohol + 4 slides). LOCALITY: Sta. 156. Antarctic Ocean, 62° 26' S 95° 44' E, 1975 fm, 26 February 1874. COLLECTION: " Challenger ". WALDECKIA Chevreux Waldeckia kroyeri (White) 1847 : 124 [Ephippiphora kroyeri]. SYNTYPE: Reg. no. 45 : 47 (i specimen, dry). LOCALITY: Tasmania. DONOR: Admiralty. AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 373 Family STEGOGEPHALIDAE ANDANIA Boeck [Andania abyssorum] Stebbing 1888 : 739 (transferred to Andaniotes corpulentus (Thomson) by Stebbing 1897). HOLOTYPE: Reg. no. 1889 : 5 : 15 : 42 (7 slides). LOCALITY: Sta. 168. Off New Zealand, 40° 28' S 177° 43' E, noo fm, 8 July 1874. COLLECTION: " Challenger ". ANDANIEXIS Stebbing Andaniexis australis Barnard 1932 : 76. SYNTYPE: Reg. no. 1936 : n : 2 : 588-591* (4 specimens, alcohol). LOCALITY: Sta. 87. 33° 53' S 09° 26' E, iooo(-o) m, 25 June 1926. SYNTYPE: Reg. no. 1936 : n : 2 : 592 (i specimen, alcohol). LOCALITY: Sta. 89. 34° 05' S 16° oo' E iooo(-o) m, 28 June 1926. COLLECTION: " Discovery " Committee. ANDANIOTES Stebbing Andaniotes linearis Barnard 1932 : 80. SYNTYPES: Reg. no. 1936 : u : 2 : 644-648* (6 specimens, alcohol). LOCALITY: Sta. 156. 53° 51' S 36° 21' W, 200-236 m, 20 January 1927. SYNTYPES: Reg. no. 1936 : n : 2 : 649-656 (1+3 + 2 + 1 + 1+1+2 speci- mens, alcohol). LOCALITIES : Sta. 39. East Cumberland Bay, South Georgia, 179-235 m, 25 March 1926. Sta. 42. Off entrance of Cumberland Bay, South Georgia, 120-204 m, I April 1926. Sta. 144. Off entrance of Stromness Harbour, South Georgia, 155- 178 m, 5 January 1927. Sta. 148. Off Cape Saunders, South Georgia, 132-148111, 9 January 1927. Sta. 149. Entrance of East Cumberland Bay, South Georgia, 200-234 m, 10 January 1927. Sta. 175. Bransfield Strait, South Shetland Islands, 63° 17' S 59° 48' W, 200 m, 2 March 1927. Sta. 190. Bismarck Strait, Palmer Archipelago, 90-130 m, 24 March 1927. COLLECTION: " Discovery " Committee. Andaniotes simplex Barnard 1930 : 328. SYNTYPES: Reg. no. 1930 : 8 : i : 92-94 (3 specimens in alcohol + i slide). LOCALITY: Sta. 90. Off Three Kings Islands, New Zealand, 18301, 25 August 1911. COLLECTION: " Terra Nova ". 374 M- H- THURSTON & ELIZABETH ALLEN BATHYSTEGOCEPHALUS Schellenberg Bathystegocephalus globosus (Walker) 1909 : 329 [Stegocephalus globosus]. SYNTYPES: Reg. no. 1909 : i : 29 : 17-31 (part) (2 + 17+2 specimens, alcohol). LOCALITIES: Indian Ocean, 200 fm, 200 fm, and 750 fm, 16 October 1905. SYNTYPES: Reg. no. 1909 : i : 29 : 17-31 (part) (2 specimens, alcohol). LOCALITY: Indian Ocean, 250-500 fm, 17 September 1905. SYNTYPE: Reg. no. 1909 : 2 : 13 : 4 (r slide). LOCALITY: Indian Ocean, 200 fm, 16 October 1905. COLLECTION: " Sealark ". SYNTYPES: Reg. no. 1925 : 9 : 8 : 122-123 (2 specimens, alcohol). LOCALITY: Indian Ocean, 200 fm, 16 October 1905. COLLECTION : Walker. EUANDANIA Stebbing Euandania gigantea (Stebbing) 1888 : 730 [Andania gigantea]. SYNTYPE : Reg. no. 1889 : 5 : 15 : 39 (i specimen, alcohol). LOCALITY: Sta. 146. Near Marion Island, South Indian Ocean, 46° 46' S 45° 3i'E, 1375 fm, 29 December 1873. SYNTYPE : Reg. no. 1889 : 5 : 15 : 40 (i specimen in alcohol + 13 slides). LOCALITY: Sta. 147. East of Marion Island, South Indian Ocean, 46° i6'S 48° 27' E, 1600 fm, 30 December 1873. COLLECTION: " Challenger ". PARANDANIA Stebbing Parandania boecki (Stebbing) 1888 : 735 [Andania boecki}. HOLOTYPE: Reg. no. 1889 : 5 : 15 : 41 (specimen in alcohol + 8 slides). LOCALITY: Sta. 120. Off Pernambuco, 08° 37' S 34° 28' W, 675 fm, 9 September i873. COLLECTION: " Challenger ". PHIPPSIA Stebbing Phippsia gibbosa (Sars) 1882 : 85 [Stegocephalus gibbosus], SYNTYPE: Reg. no. 1911 : n : 8 : Mg8i (i slide). LOCALITY: West Norway. COLLECTION: Norman. The absence of date and precise locality leaves some doubt as to the validity of the type status of this specimen. AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 375 PHIPPSIELLA Schellenberg Phippsiella rostrata Barnard 1932 : 76. SYNTYPES: Reg. no. 1936 : n : 2 : 585-587 (3 specimens, alcohol). LOCALITY: Sta. 158. 53° 51' S 36° 2i'W, 200-236 m, 21 January 1927. COLLECTION: " Discovery " Committee. STEGOCEPHALOIDES Sars Stegocephaloides attingens Barnard 1916 : 131. SYNTYPES: Reg. no. 1928 : 4 : 13 : 41-44 (4 specimens, alcohol). LOCALITY: South Africa. DONOR: South African Museum. Stegocephaloides auratus (Sars) 1882 : 86 [Stegocephalus auratus]. SYNTYPE: Reg. no. 1911 : n : 8 : MgSo (i slide). LOCALITY: Finmark. COLLECTION: Norman. The validity of the type status of this specimen is in doubt. The locality given in the original description (Sars 1882) is " Norvegiae occidental's ". Later, (Sars 1891) this is repeated, but three separate localities are given, of which Mager0 is in Finmark. Stegocephaloides australis Barnard 1916 : 129. SYNTYPES: Reg. no. 1928 : 4 : 13 : 45-48 (4 specimens, alcohol). LOCALITY : South Africa. DONOR: South African Museum. Family AMPELISCIDAE AMPELISCA Kr0yer Ampelisca abyssicola Stebbing 1888 : 1047. HOLOTYPE: Reg. no. 1889 : 5 : 15 : 121 (specimen in alcohol + 6 slides). LOCALITY: Sta. 24. Off Culebra Island, St. Thomas, Virgin Islands, i8°39'N 65° 05' W, 390 fm, 25 March 1873. COLLECTION: " Challenger ". Ampelisca acinaces Stebbing 1888 : 1036. HOLOTYPE: Reg. no. 1889 : 5 : 15 : 119 (specimen in alcohol + 7 slides). LOCALITY: Sta. 1636. Port Jackson, Sydney, Australia, 35 fm, 3 June 1874. (Slide label gives locality as Sta. i63A, 30 fm.) COLLECTION: " Challenger ". 376 M. H. THURSTON & ELIZABETH ALLEN Ampelisca amblyops Sars 1891 : 180. SYNTYPES: Reg. no. 1911 : n : 8 : 15027 (2 specimens, alcohol). LOCALITY: Christianiafjord, Norway. COLLECTION : Norman. Ampelisca anomala Sars 1882 : 108. SYNTYPES: Reg. no. 1911 : n : 8 : 15025-15026 (2 specimens, alcohol). LOCALITY : West Norway. COLLECTION : Norman. Considerable doubt must be attached to the type status of these specimens as Sars had only two examples when he descibed the species. Ampelisca birulai Briiggen 1909 : 17. SYNTYPES: Reg. no. 1936 : 3 : 18 : 120-122 (3 specimens, alcohol). LOCALITY : Siberian Sea. DONOR: Academy of Sciences, Leningrad. Ampelisca brachyceras Walker 1904 : 251. SYNTYPES: Reg. no. 1905 : 2 : 18 : 42 (i specimen, alcohol). 1905 : 2 : 18 : 322 (i slide). LOCALITY: Ceylon, November, 1902. COLLECTION: Herdman. Ampelisca bransfieldi Barnard 1932 : 83. SYNTYPES : Reg. no. 1936 : n : 2 : 708-709 (2 specimens, alcohol). LOCALITY : Sta. 177. 27 miles SW of Deception Island, South Shetland Islands, 1080 m, 5 March 1927. COLLECTION: " Discovery " Committee. Ampelisca brevicornis var. cavicoxa Reid 1951 : 208. SYNTYPES: Reg. no. 1961 : 8 : i : 503-504 (2 specimens, alcohol). LOCALITY: Sta. 161. Off Bathurst, Gambia, 18 m, 27 July 1946. COLLECTION: " Atlantide ". Ampelisca byblisoides Barnard 1925 : 335. SYNTYPES: Reg. no. 1928 : 4 : 13 : 54-57 (4 specimens, alcohol). LOCALITY: South Africa. DONOR: South African Museum. {Ampelisca chevreuxi] Walker 1904 : 254 (transferred to Ampelisca zamboangae Stebbing by Pirlot 1936). HOLOTYPE: Reg. no. 1905 : 2 : 18 : 50 (alcohol). 1905 : 2 : 18 : 329 (i slide). LOCALITY: 12 miles W of Vankala, Ceylon, 7^-9 fm, 6 March 1902. COLLECTION: Herdman. AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 377 AmpeliscachiltoniStebbing 1888 : 1042. SYNTYPES: Reg. no. 1889 : 5 : 15 : 120 (4 specimens in alcohol + 8 slides). LOCALITY: Sta. 167. Off New Zealand, 39° 32' S 171° 48' E, 150 fm, 24 June 1874. (slide label gives Sta. i67A, ro fm.) COLLECTION: " Challenger ". Sta. i67A is not mentioned in Stebbing's report. It was collected in Queen Charlotte Sound, South Island, New Zealand, on 27 June 1874. This locality is ca. 280 km SE of the position of Sta. 167. Ampelisca compacta Norman 1882 : 688. HOLOTYPE: Reg. no. 1911 : n : 8 : Mg63 (i slide). LOCALITY: " Knight Errant " Sta. 7. Off Lewis, Scotland, 59° 37' N 07° 19' W, 530 fm, ii August 1880. COLLECTION: Norman. Ampelisca cy clops Walker 1904 : 253. SYNTYPES: Reg. no. 1905 : 2 : 18 : 49 (2 specimens, alcohol). 1905 : 2 : 1 8 : 327-328 (2 slides). LOCALITY: Ceylon. COLLECTION: Herdman. Ampelisca fusca Stabbing 1888 : 1052. SYNTYPES: Reg. no. 1889 : 5 : 15 : 122 (3 specimens in alcohol + 7 slides). LOCALITY: Sta. 142. Off Cape Agulhas, South Africa, 35° 04' S 18° 37' E, 150 fm, 18 December 1873. COLLECTION: " Challenger ". Ampelisca hemicryptops Barnard 1930 : 329. SYNTYPES: Reg. no. 1930 : 8 : i : 105-108 (4 specimens, alcohol). LOCALITY: Sta. 339. McMurdo Sound, Ross Sea, 256 m, 24 January 1912. SYNTYPES: Reg. no. 1930 : 8 : i : 109-113 (3+2 specimens, alcohol). LOCALITIES: Sta. 194. Gates Land, Antarctica, 329-366 m, 22 February 1911. Sta. 314. McMurdo Sound, Ross Sea, 406-441 m, 23 January 1912. COLLECTION: " Terra Nova ". [Ampelisca ingens] Bate 1862 : 92 (transferred to Ampelisca eschrichti Kr0yer by Sars 1891). SYNTYPES: Reg. no. 68 : 107 (6) (3 specimens, alcohol). LOCALITY: Grand Manan, New Brunswick, Canada. COLLECTION: Stimpson. Ampelisca japonica Bate 1862 : 94. HOLOTYPE: Reg. no. 68 : 107 (18) (alcohol). LOCALITY: Japan. COLLECTION: Stimpson. 37« M. H. THURSTON & ELIZABETH ALLEN Ampelisca limicola (Stimpson) 1853 : 57 [Pseudophthalmus limicolus]. SYNTYPE: Reg. no. 68 : 107 (19) (i specimen, alcohol). LOCALITY: Charleston Harbour, S. Carolina, U.S.A. COLLECTION: Stimpson. A mpelisca palmata Barnard 1916 : 136. SYNTYPES: Reg. no. 1928 : 4 : 13 : 49-53 (10 specimens, alcohol). LOCALITY : South Africa. DONOR : South African Museum. A mpelisca pusilla Sars 1891 : 181. SYNTYPES: Reg. no. 1911 : n : 8 : 15033-15042 (14 specimens, alcohol). LOCALITY: Trondhjemsfjord, Norway. COLLECTION: Norman. Probably type material. Ampelisca sarsi Chevreux 1888 : 666. SYNTYPES: Reg. no. 1911 : n : 8 : 15068-15069 (2 specimens, alcohol). LOCALITY: Le Croisic, France. COLLECTION: Norman. Ampelisca scabripes Walker 1904 : 250. SYNTYPES: Reg. no. 1905 : 2 : 18 : 39-41 (3 specimens, alcohol). LOCALITY: Periya Paar, Ceylon. SYNTYPES: Reg. no. 1905 : 2 : 18 : 321 (i slide). LOCALITY: Ceylon. COLLECTION: Herdman. Ampelisca statenensis Barnard 1932 : 84. HOLOTYPE: Reg. no. 1936 : n : 2 : 710 (alcohol). LOCALITY: Sta. WS 88, 54° oo' S 64° 57' W, 118 fm, 6 April 1927. COLLECTION: " Discovery " Committee. Ampelisca tridens Walker 1904 : 249. SYNTYPES: Reg. no. 1905 : 2 : 18 : 33-38 (7 specimens, alcohol). LOCALITY: Sta. LVIII, Gulf of Manaar, Ceylon, 10 March 1902. SYNTYPES: Reg. no. 1905 : 2 : 18 : 318-320 (3 slides). LOCALITY: Ceylon. COLLECTION: Herdman. SYNTYPES: Reg. no. 1925 : 9 : 8 : 163-172 (13 specimens, alcohol). 1925 : 9 : 8 : 176-181 (6 specimens, alcohol). LOCALITY: Sta. XXXII, Ceylon. AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 379 SYNTYPES: Reg. no. 1925 : 9 : 8 : 173-175 (3 specimens, alcohol). 1925 19:8: 182-191 (ca. 50 specimens, alcohol). 1925 19:8: 192-195 (4 specimens, alcohol). LOCALITY: Ceylon. SYNTYPES: Reg. no. 1925 : 9 : 8 : 196-205 (15 specimens, alcohol). LOCALITY: E. Cheval Paar, Ceylon, November 1902. COLLECTION: Walker. Ampelisca zamboangae Stebbing 1888 : 1057. HOLOTYPE: Reg. no. 1889 : 5 : 15 : 123 (4 slides). LOCALITY: Off Samboangan, Philippine Islands, surface, 18 February 1875. COLLECTION: " Challenger ". BYBLIS Boeck Byblis abyssi Sars 1879 : 456. SYNTYPE: Reg. no. 1911 : n : 8 : 15133 (i specimen, alcohol). LOCALITY: Norwegian North Atlantic Expedition Sta. 31. Norwegian Sea. 63° 10' N 05° oo' E, 763 m, 29 June 1876. COLLECTION: Norman. Byblis affinis Sars 1891 : 186. SYNTYPES: Reg. no. 1911 : u : 8 : 15126-15131 (7 specimens, alcohol). LOCALITY: Trondhjemsfjord, Norway, 40-60 fm. COLLECTION: Norman. Byblis erythrops Sars 1882 : 109. SYNTYPE: Reg. no. 1911 : n : 8 : ^1969 (i slide). LOCALITY: Bejan, West Norway. COLLECTION: Norman. Byblis lepta (Giles) 1888 : 223 [Ampelisca lepta]. SYNTYPES: Reg. no. 87 : 14 (ca. 40 specimens, alcohol). 1909 : 4 : 3 : i (3 slides). LOCALITY : Swatch-of-no-Ground, Bay of Bengal, 107 fm. COLLECTION: Giles. Byblis longicornis Sars 1891 : 185. SYNTYPE: Reg. no. 1911 : n : 8 : 15132 (i specimen, alcohol). LOCALITY: Norwegian North Atlantic Expedition Sta. 260. Porsangerfjord, Norway, 70° 55' N 26° n' E, 232 m, 25 June 1878. COLLECTION: Norman. 380 M. H. THURSTON & ELIZABETH ALLEN BYBLISOIDES Barnard Byblisoides juxtacornis Barnard 1931 : 426. SYNTYPES: Reg. no. 1936 : n : 2 : 735-740* (8 specimens, alcohol). LOCALITY: Sta. 181. Schollaert Channel, Palmer Archipelago, Graham Land, 278-500 m, 12 March 1927. SYNTYPE: Reg. no. 1936 : n : 2 : 741 (i specimen, alcohol). LOCALITY : Sta. 186. Fournier Bay, Anvers Island, Palmer Archipelago, Graham Land, 295 m, 16 March 1927. COLLECTION: " Discovery " Committee. HAPLOOPS Liljeborg Haploops securiger Barnard 1931 : 426. SYNTYPES: Reg. no. 1936 : ir : 2 : 742-746* (7 specimens, alcohol). LOCALITY: Sta. 123. Off entrance of Cumberland Bay, South Georgia, 230-250 m, 15 December 1926. SYNTYPES: Reg. no. 1936 : n : 2 : 747-796 (17+6+4 + 2 + 21 +21+6 + i specimens, alcohol). LOCALITIES: Sta. 27. West Cumberland Bay, South Georgia, no m, 15 March 1926. Sta. 42. Off entrance of Cumberland Bay, South Georgia, 120-204 m, I April 1926. Sta. 140. Stromness Harbour to Larsen Point, South Georgia, 122- 136 m, 23 December 1926. Sta. 144. Off entrance of Stromness Harbour, South Georgia, 155-178 m, 5 January 1927. Sta. 146. Off South Georgia, 53° 48' S 35° 37' W, 728 m, 8 January 1927. Sta. 153. Off South Georgia, 54° 08' S 36° 27' W, 106 m, 17 January 1927. Sta. 170. Off Cape Bowles, Clarence Island, South Shetland Islands, 61° 25' S 53° 46' W, 342 m, 23 February 1927. Sta. WS 33. Off South Geor- gia, 54° 59' S 35° 24' W, 130 m, 23 December 1926. [TETROMATUS Bate] [Tetromatus bellianus] Bate 1856 : 58 (nom. nud.), 1857 : 139 (transferred to Ampelisca brevicornis (Costa) by Sars 1891). SYNTYPES: Reg. no. 1952 : 5 : 7 : 27-29 (3 specimens, alcohol). LOCALITY: Moray Firth, Scotland. COLLECTION : Bate. Family HAUSTORIIDAE BATHYPOREIA Lindstrom Bathyporeia elegans Watkin 1938 : 222 [Bathyporeia pelagica Sars 1891 : 129]. SYNTYPES: Reg. no. 1968 : 50 (2 specimens, alcohol). LOCALITY: Jaederen, Norway. COLLECTION : Zoological Museum, Oslo. AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 381 Bathyporeia pelagica (Bate) 1856 : 59 (nom. nud.), 1857 : X46 [Theristes pelagica]. HOLOTYPE: Reg. no. 1952 15:7: 182 (alcohol). LOCALITY: Moray Firth, Scotland. COLLECTION: Bate. [Bathyporeia robertsoni] Bate 1862 : 173 (transferred to Bathyporeia pilosa Lindstrom by Watkin 1938). SYNTYPES: Reg. no. 1952 : 5 : 7 : 183-184 (2 specimens, alcohol). LOCALITY : Cumbrae, Firth of Clyde, Scotland. COLLECTION: Bate. CARDENIO Stebbing Cardenio paurodactylus Stebbing 1888 : 806. SYNTYPES: Reg. no. 1889 : 5 : 15 : 58 (3 specimens in alcohol + 4 slides). LOCALITY: Betsy Cove, Kerguelen, South Indian Ocean, surface, 10 January 1874. COLLECTION: " Challenger ". Also in the Museum Collection is the specimen on which Barnard (1932 : 91, fig. 43) based the description of the male of the species. He designated this specimen as " Type of <$ " but in fact it has no type status. HAUSTORIELLA Barnard [Haustoriella psammophila] Barnard 1931 : 426 (transferred to Phoxocephalopsis zimmeri Schellenberg by Barnard 1932). SYNTYPES: Reg. no. 1936 : n : 2 : 804-805 (2 specimens, alcohol). LOCALITY: Sta. WS 96. NW of Falkland Islands, 48°oo'S 64°45'W, 96m, 17 April 1927. COLLECTION: " Discovery " Committee. PLATYISCHNOPUS Stebbing Platyischnopus herdmani Walker 1904 : 247. SYNTYPES: Reg. no. 1905 : 2 : 18 : 28-31 (4 specimens, alcohol). 1905 : 2 : 18 : 315-317 (3 slides). 1925 : 9 : 8 : 250-252 (2 specimens, alcohol). LOCALITY: East Cheval Paar, Ceylon, November 1902. COLLECTION: Herdman. Platyischnopus mirabilis Stebbing 1888 : 830. SYNTYPES: Reg. no. 1889 : 5 : 15 : 63 (4 slides). LOCALITY: Port Jackson, Australia, 2-10 fm. COLLECTION: " Challenger ". 382 M. H. THURSTON & ELIZABETH ALLEN PONTOPOREIA Kr0yer [Pontoporeia hoyi] Smith 1874 : 647 (transferred to Pontoporeia affinis Lindstrom by Gurjanova SYNTYPES: Reg. no. 1911 : n : 8 : 13737-13746 (16 specimens, alcohol). LOCALITY: Lake Superior, U.S.A., 50 fm, 1871. COLLECTION: Norman. UROTHOE Dana Urothoe bairdi Bate 1862 : 114. SYNTYPES: Reg. no. 1952 : 5 : 7 : 32-34 (6 specimens, alcohol). LOCALITY: Moray Firth, Scotland. COLLECTION: Bate. Urothoe brevicornis Bate 1862 : 116. SYNTYPES: Reg. no. 1952 : 5 : 7 : 35-37 (6 specimens, alcohol). LOCALITY: Tenby, Wales. COLLECTION: Bate. Urothoe elegans (Bate) 1856 : 58 (nom. nud.), 1857 : 145 [Gammarus elegans]. SYNTYPE : Reg. no. 1952 : 5 : 7 : 139 (i specimen, alcohol, badly mutilated). LOCALITY: Plymouth, England. COLLECTION: Bate. Urothoe grimaldii Chevreux 1895 : 428. SYNTYPES: Reg. no. 1911 : n : 8 : 14074-14077 (3 specimens, alcohol). LOCALITY : Coast of Morocco. COLLECTION: Norman. Urothoe marina (Bate) 1857 : 140 [Sulcator marinus]. HOLOTYPE: Reg. no. 1952 : 5 : 7 : 141 (alcohol). LOCALITY: Banff, Scotland. COLLECTION: Bate. Urothoe ruber Giles 1888 : 246. SYNTYPES: Reg. no. 1968 : 54 (2 slides). LOCALITY: Banks of the Chittagong River. The slide label gives the locality as Mutlah Flats. COLLECTION: Giles. Urothoe spinidigitus Walker 1904 : 245. HOLOTYPE: Reg. no. 1905 : 2 : 18 : 26 (alcohol). 1905 : 2 : 18 : 314 (i slide). LOCALITY: Cheval Paar, Ceylon, November 1902. COLLECTION: Herdman. AMPHIPOD TYPE MATERIAL IN THE BRITISH MUSEUM 383 UROTHOIDES Stebbing Urothoides lachneessa (Stebbing) 1888 : 825 [Urothoe lachneessa]. SYNTYPES: Reg. no. 1889 : 5 : 15 : 62 (part) (2 specimens, alcohol). LOCALITY: Off Cumberland Bay, Kerguelen, South Indian Ocean, 120 im. SYNTYPES: Reg. no. 1889 : 5 : 15 : 62 (part) (6 slides). LOCALITY : Off Christmas Harbour, Kerguelen, South Indian Ocean, 120 fm. COLLECTION: " Challenger ". The locality " off Christmas Harbour " is not mentioned by Stebbing (1888) under this species. It is probable, however, that this locality and " off Cumberland Bay " are the same, as the entrances of the two bays are only 9 km apart. Urothoides oniscoides Barnard 1932 : 93. HOLOTYPE: Reg. no. 1936 : n : 2 : 803 (alcohol). LOCALITY: Sta. 196. Bransfield Strait, South Shetland Islands, 62° 17' S 58° 21' W 720 m, 3 April 1927. COLLECTION: " Discovery " Committee. REFERENCES BARNARD, J. L. 1958. 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Pelagic Amphipoda of the Irish Atlantic Slope. Scient. Invest. Fish Brch Ire., 1905, 4 (8) 1-39. WALKER, A. O. 1895. Revision of the Amphipoda of the L.M.B.C. District. Proc. Trans. Lpool biol. Soc., 9 : 287-320. 1903. Amphipoda of the " Southern Cross " Antarctic Expedition. /. Linn. Soc., 29 : 38-64. 1904. Report on the Amphipoda collected by Professor Herdman at Ceylon in 1902. in Report to the Government of Ceylon on the Pearl Oyster Fisheries in the Gulf of Manaar. Part 2. Supplementary Rept., (17) : 229-300. 1906. Preliminary descriptions of new species of Amphipoda from the " Discovery " Antarctic Expedition, 1902-4. Ann. Mag. nat. Hist., (7) 17 : 452-458. — 1907. Amphipoda. National Antarctic Expedition 1901-1904. Zoo/., 3 : 1-38. 1909. Amphipoda Gammaridea from the Indian Ocean, British East Africa, and the Red Sea. Trans. Linn. Soc. Lond., (2) 12 Zoo/., (18) : 323-344. & SCOTT, A. 1903. Decapod and sessile-eyed crustaceans from Abd-el-Kuri. pp. 216- 232 in Forbes, H.O. The Natural History of Socotra and Abd-el-Kuri. xlvii + 598 pp. WATKIN, E. E. 1938. A revision of the amphipod genus Bathyporeia Lindstrom. /. mar. biol. Ass. U.K. 23 211-236. [WHITE, A.] 1847. List of the specimens of Crustacea in the collection of the British Museum. viii +143 pp. (British Museum, London). 18470. Descriptions of new or little-known Crustacea in the collection at the British Museum. Proc. zool. Soc. Lond. 15 : 118-126. 386 M.H. THURSTON & ELIZABETH ALLEN INDEX Square brackets enclosing only the generic name indicate species whose generic position has been changed since the original description of the species. Square brackets enclosing both the generic and specific names mark those species now considered to be junior synonyms. Round brackets indicate names of valid species which are not represented in the collections of the British Museum (Natural History) by type material, but which include type material of species now placed as junior synonyms. abyssi, Byblis, 379 abyssicola, Ampelisca, 375 abyssicola, Hippomedon holbolli var., 359 [abyssorum, Andania], 373 abyssorum, [Orchomene], 364 abyssorum, Orchomenella, 364 acinaces, Ampelisca, 375 adarei, Tryphosa, 370 affinis, Byblis, 379 [affinis, Ichnopus], 359 (affinis, Pontoporeia) , 382 albina, Tryphosella, 372 amblyops, Ampelisca, 376 [amblyops, Normanion], 363 amblyops, Orchomene, 364 [ampulloides, Anonyx], 354 analogica, Tryphosa, 370 angulata, Tryphosa, 370 annulatus, Anonyx, 355 anomala, Ampelisca, 376 (anonyx, Cyphocaris), 358 antarctica, Kerguelenia, 359 antarcticus, Aristias, 355 [antennipotens, Tryphosa], 370 attingens, Stegocephaloides, 375 audouiniana, [Lysianassa] , 366 audouiniana, Perrierella, 366 auratus, Stegocephaloides, 375 auratus, [Stegocephalus], 375 australis, Andaniexis, 373 australis, Stegocephaloides, 375 bairdi, Urothoe, 382 barbatipes, [Tryphosa], 372 barbatipes, Tryphosella, 372 batei, Orchomene, 364 bathycephala, Amaryllis, 354 [bellianus, Tetromatus], 380 bidens, Lepidepecreella, 360 bidens, [Paracyclocaris], 360 birulai, Ampelisca, 376 bispinosus, Anonyx, 355 boecki, [Andania], 374 boecki, Parandania, 374 bolivari, Podoprion, 366 bonnieri, Socarnella, 367 brachyceras, Ampelisca, 376 bransfieldi, Ampelisca, 376 brevicaudatus, Onisimus, 363 (brevicornis, Ampelisca), 380 brevicornis var. cavicoxa, Ampelisca, 376 brevicornis, Urothoe, 382 bruuni, Tmetonyx, 369 byblisoides, Ampelisca, 376 capadarei, Tryphosites, 372 caricus, Onisimus, 363 carinatus, Onesimoides, 363 caspius, Pseudalibrotus, 366 castellata, Tryphosa, 370 cavicoxa, Ampelisca brevicornis var., 376 cavimanus, [Orchomene], 364 cavimanus, Orchomenella, 364 challengeri, Cyphocaris, 358 chelatus, Euonyx, 358 chelipes, Orchomenella, 365 [chevreuxi, Ampelisca], 376 chevreuxi, Tryphosites, 372 chilensis, [Anonyx], 365 chilensis, Orchomenella, 364, 365, 366 chiltoni, Ampelisca, 377 cicadoides, [Anonyx], 369 cicadoides, Tmetonyx, 369 [ciliata, Tryphosa], 370 cingulatum, Lepidepecreum, 361 coatsi, [Orchomenopsis], 367 coatsi, Pseudorchomene, 367 coelochir, Lysianassa, 361 coelochir, [Lysianax], 361 coheres, Valettia, 372 INDEX 387 collinus, Aristias, 355 compacta, Ampelisca, 377 compacta, Kerguelenia, 360 (corpulentus, Andaniotes), 373 crenatipalmatus, Cheirimedon, 357 crenatus, Scopelocheirus, 367 crenulata, Socarnopsis, 368 cucullata, Tryphosa, 371 cyclops, Ampelisca, 377 denticulata, [Anonyx], 358 denticulatus, Hippomedon, 358 elegans, Bathyporeia, 380 elegans, [Gammarus] , 382 elegans, Urothoe, 382 erythrophthalmus, Socarnes, 367 erythrops, Byblis, 379 (eschrichti, Ampelisca), 377 exiguus, Anonyx, 355 falklandica, Arugella, 356 falklandica, [Lysianassa] , 356 [faroensis, Cyclocaris], 357 foraminiferum, Lepidepecreum, 361 fougneri, Cheirimedon, 357 franklini, Orchomenella, 365 fusca, Ampelisca, 377 geelongi, Hippomedon, 359 georgiensis, Ambasiopsis, 354 gibbosa, Phippsia, 374 gibbosus, [Stegocephalus], 374 gigantea, [Andania], 374 gigantea, Euandania, 374 (gigas, Uristes), 370 globosus, Bathystegocephalus, 374 globosus, [Stegocephalus], 374 goniops, Orchomene, 364 grimaldii, Urothoe, 382 (guilelmi, Cyclocaris), 357 [hanseni, Cheirimedon], 357 (hanseni, Orchomene), 355 haswelli, [Amaryllis], 357 haswelli, Bathyamaryllis, 357 helgae, Metacyphocaris, 361 hemicryptops, Ampelisca, 377 herdmani, Platyischnopus, 381 holbolli var. abyssicola, Hippomedon, 359 [hoyi, Pontoporeia] , 382 incerta, Podoprionides, 366 incisus, Hippomedon, 359 indica, Microlysias, 362 indicus, Anonyx, 355 [ingens, Ampelisca], 377 insigne, Stomacontion, 368 integricauda, [Ambasia], 362 integricauda, Nannonyx, 362 japonica, Ampelisca, 377 juxtacornis, Byblisoides, 380 kergueleni, [Acontiostoma], 368 kergueleni, [Lysianassa], 371 kergueleni, Socarnoides, 368 kergueleni, Stomacontion, 368 kergueleni, Tryphosa, 371 kroyeri, [Ephippiphora], 372 kroyeri, Waldeckia, 372 lachneessa, [Urothoe], 383 lachneessa, Urothoides, 383 lepta, [Ampelisca], 379 lepta, Byblis, 379 limicola, Ampelisca, 378 limicolus, [Pseudophthalmus], 378 linearis, Andaniotes, 373 lobata, Allogaussia, 353 longicornis, Byblis, 379 longimanus, Hippomedon, 359 longimanus, [Platamon], 359 longitelson, Tmetonyx, 369 [magellanicum, Acontiostoma], 353 major, Tryphosa, 371 marina, Urothoe, 382 marinus, [Sulcator], 382 marionis, Acontiostoma, 353 megalurus, Crybelocephalus, 357 [melanophthalmus, Anonyx], 355 [micronyx, Cyphocaris], 358 miersi, [Hippomedon], 369 miersi, Tmetonyx, 369 mirabilis, Platyischnopus, 381 murrayi, Sophrosyne, 368 murrayi, Tryphosa, 371 musculosus, [Orchomene], 364 (nana, Orchomenella), 370 navicula, Allogaussia, 354 nobilis, [Anonyx], 369 nobilis, Tmetonyx, 369 nodimanus, Orchomenella, 365 nodimanus, [Orchomenopsis], 365 normani, Euonyx, 358 (nugax, Anonyx), 354 388 M. H. THURSTON & ELIZABETH ALLEN obesum, Acidostoma, 353 obesus, [Anonyx], 353 (odhneri, Paralysianopsis), 356 oniscoides, Urothoides, 383 ovalis, Lepidepecreella, 360 pacificus, Aristias, 356 palmata, Ampelisca, 378 palpalis, Kerguelenia, 360 paurodactylus, Cardenio, 381 pelagica, Bathyporeia, 381 pelagica, [Theristes], 381 pepinii, [Acontiostoma], 368 pepinii, Stomacontion, 368 (pilosa, Bathyporeia), 381 pinguides, Orchomenella, 365 plebs, Orchomenella, 365 proxima, [Orchomenopsis] , 365 [psammophila, Haustoriella], 381 punctatus, Anonyx, 355 pusilla, Ampelisca, 378 quadrimanus, Normanion, 363 quadrimanus, [Opis], 363 [rhinoceros, Austronisimus] , 356 [robertsoni, Bathyporeia], 381 robusta, Shackletonia, 367 rossi, [Orchomenopsis], 366 rostrata, Phippsiella, 375 rotundatus, Chironesimus, 357 rotundatus, [Lakota], 357 rotundifrons, Orchomenella, 365 ruber, Urothoe, 382 sarsi, Ampelisca, 378 scabripes, Ampelisca, 378 securiger, Haploops, 380 serratum, Trischizostoma, 369 [serricrus, Ichnopus], 359 setosus, Aroui, 356 simplex, Andaniotes, 373 spinidigitus, Urothoe, 382 spinimanus, Nannonyx, 362 statenensis, Ampelisca, 378 stebbingi, [Hoplonyx], 369 stebbingi, Tmetonyx, 369 symbiotica, Aristias, 356 tahitensis, Cyclocaris, 358 (taurus, Ichnopus), 359 (tenuicornis, Glycerina), 361 tenuipes, Amaryllis, 354 tenuipes, [Vijaya], 354 triangularis, Tryphosa, 371 tridens, Ampelisca, 378 trigonica, Tryphosa, 371 trigonicus, [Hippomedon], 371 uncinata, Ambasiopsis, 354 urodus, Lysianassa, 361 urodus, [Lysianax], 361 variegata, [Anonyx], 361 variegata, Lysianassa, 361 [woodmasoni, Lysianassa], 361 xenokeras, Microlysias, 362 xenopus, Lepidepecreoides, 360 zamboangae, Ampelisca, 376, 379 (zimmeri, Phoxocephalopsis) , 381 MICHAEL H. THURSTON, B.Sc. NATIONAL INSTITUTE OF OCEANOGRAPHY WORMLEY GODALMING SURREY ELIZABETH ALLEN Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON, S.W.7 INDEX TO VOLUME 17 The page numbers of the principal references and the new taxonomic names are printed in bold type. For Number 6 page references are given only for the three phyla considered and orders of the Polyzoa as the paper itself is an index. For index to Number 9 see page 386. abessinicus, Mesopicos abingoni, Campethera abyssinicus, Dendropicos Acoleidae Actinonema . Actitis .... Acuminodeutopus . adriatica, Euckromadora adriatica, Spilophora adriatica, Steineridora . Aegithina afer, Lemboides affinis, Apus . affinis, Micropus 22 20, 21 19 336-341 185, 188, 189 284, 317, 330 • 95 155, 168, 170 202 152, 170, 199 285, 297, 310 95 •284,319,333,335 335 africana, Clupea . 263, 264, 268-270, PI. 2 africana, Ilisha . . 264, 268-270, PI. 2 africana, Agama agama ... 73 Agama ..... 65-90, 6 pis agama, Agama 67, 68, 69, 70-75, 77» ?8, 81, 82, 87, PI. i, 4, 5 agama, Agama agama .... 68 agama africana, Agama .... 73 agama agama, Agama .... 68 agama benueensis, Agama 67, 68, 75, 78, 79, 88 agama boensis, Agama . . . 78, 79, 85 agama mucosoensis, Agama ... 74 agama savatieri, Agama . . 72, 73 Agamidae ..... 65-90 agmen, Mesopicos . .... 22 albaredai, Gyrocoelia .... 338 albolarvatus, Dendrocopos . . 13, 30 Albula ....... 264 Albulidae ...... 264 algicola, Microdeutopus . 97, 109, no, 114, 115, 116, 117-iig, 134 alii Austranema 152, 169, 187, 195-197, 201 alii, Nygmatonchus 152, 153, 155, 158, 175, 185, 186, 187, 201 allomyodes, Raillietina . . 283, 299-304 Alosa, ..... 264, 267, 274 alosa, Alosa ..... 264, 267 alosa, Clupea .... 264, 267 Amazona ...... 288 amazonica, Amazona .... 288 ambigua, Paronia ..... 288 ambigua, Parvitaenia .... 325 amokurae, Euchromadora . . .182 amokurae, Graphonema 152, 168, 183, 200 amokurae, Spilophora . . . 170, 202 amokuroides, Graphonema . . 183, 201 amokuroides, Spilophora . 182, 183, 202 Amphideutopus ..... 95 Amphithoe ...... 98 analis, Dendrocopos . . . 31-32 Anas ...... 284, 315 Andrepigynotaenia . . . 339, 341 Angularella ...... 327 anomala, Autonoe . . . .no anomalus, Gammarus . . . .no 'anomalus' group of Microdeutopus 97, 109-1 10 anomalus, Microdeutopus 93, 95, 96, 97, 101, 109, 110-H4, 115, 116, 117, 118, 119, 134, 139 Anomotaenia 283, 317, 318, 319-324, 325, 327, 328 Anoplocephalidae .... 286-287 Anseriformes . . . . .284 antarctica, Euchromadora . . .188 Anthreptes ..... 285, 314 Aora 95, 139 Aorcho ..... 94, 95 Aoridae ..... 91-148 Aoroides ...... 95 Apodiformes . . . . .284 Apus . . . 284, 319, 320, 333, 335 apus, Apus ...... 320 apus, Cypselus ..... 335 archaica, Euchromadora . 168, 170, 201 archaica, Steineridora . . 152, 170, 199 arctica, Euchromadora . . . 189, 201 arcticus, Lembos . . . . .139 arcticus, Microdeutopus . . . 139 arcticus, Picoides . . . . -33 ardeae, Parvitaenia .... 325 Ardeola 317 ardeolae, Dilepis . . . 283, 316-317 ardeolae, Parvitaenia .... 325 arizonae, Dendrocopos .... 30 armata, Stimpsonella . . . .129 armatus, Microdeutopus 95, 97, 109, 129-131, 132, 134, 135 armatus, Stimpsonella .... 96 arquatus, Numenius . . . .316 aruensis, Raillietina . . . 302, 306 Ascometra .... 283, 331-333 assimilis, Dendrocopos . . . 40, 42 assumptionis, Centropus toulou . . 285 Asyndesmus . . . . . 3, 6, 14 atherinoides, Clupea . 264, 273-274, PI. 3 atherinoides, Pterengraulis 264, 273-274, PI. 3 atratus, Dendrocopos . . . 31-32, 41 390 INDEX atrigularis, Stenocercus . auratus, Colaptes . aurifrons, Melanerpes aurita, Parvitaenia australiensis, Gyrocoelia australis, Lemboides Austranema 152, 169, 172, 195- Autonoe awokera, Picus azurea, Lacerta azurea, Urocentron azureum, Uracentron azureum, Urocentron azureus, Doryphorus azureus, Ophessa . azureus, Stellio azureus, Uranocentron azureus, Uromastyx 47, 48 . 58 3, 6, 12 i6 • 325 • 338 . 139 178, 185-188, 189, •197, 200-2OI, 202 . 96, 98, 101, no . 24, 25, 26 47, 49, 50, 5i 50 . 50-53, 57, 59, 62 50 50, 55 . • . -50 50 . 5° . 50 baelama, Clupea . . . .264, 274 baelama, Thrissina . . . 264, 274 balia, Campethera punctuligera . . 20 beauforti, Paronia .... 288 bengalensis, Centropus . . . .285 benghalense, Dinopium . . . . 9, 41 bennettii, Campethera . . .9,20,21,23 bennettii scriptoricauda, Campethera 20, 21 bennettii vincenti, Campethera . . 20 benueensis, Agama 68, 73, 75, 78-83, 85, 87, 88, PI. 2, 4, 5 benueensis, Agama agama 67, 68, 75, 78, 79, 88 berlepschi, Dendrocopos mixtus . . 30 bernicla, Branta 33 beryllinus, Coryllinus .... 288 beryllinus, Coryllis .... 288 bibronii, Agama ..... 74 bicornata, Neochromadora . . 185, 201 bicornata, Nygmatonchus . 152, 184, 200 bidens, Microdeutopus .... 98 bidentatus, Microdeuteropus . . .139 biuterina, Paronia .... 288 Block, M. E. . . . . 261-279 blochii, Clupeonia . . . 267, 268 Blythipicus . . . . . 3, 9, 12 bocki, Paronia . . 283, 286-287, 288 bodkini, Kowalewskiella . 329, 330, 331 bodkini, Raillietina .... 329 bodkini, Skrjabinia .... 329 boensis, Agama . . . 67, 83, 85, 88 boensis, Agama agama . . .78, 79, 85 borealis, Dendrocopos . . . 15, 29, 30 boschas, Anas . . . . -315 boschas domestica, Anas . . .284 bothrioplites, Taenia . . . 287, 290 botrioplitis, [Taenia] .... 292 boueti, Agama ..... 74 bovieni, Notopentorchis . . 334, 335 bovieni, Paruterina .... 334 Brachypternus ..... 35 brachysoma, Ilisha . . . 270, 271 brachyurus, Micropternus . . . 4, 6 Branta ...... 33 brasiliensis, Clupea .... 264 brevicaudata, Stellio .... 50 brevicaudatum, Urocentron ... 50 brevicaudatus, Doryphorus ... 50 brevicaudatus, Ophessa ... 50 brevicaudatus, Stellio .... 50 brevicaudatus, Uranocentron . . 50 brunnichii, Clupea . . . 265, 274 burhini, Infula .... 338, 339 Burhinus ...... 318 Burt, M. D. B 281-346 burti, Paricterotaenia . . 283, 3 15-31 6 Butastur . . . . . 330 Butorides . . . 284, 316, 317, 324 buzzardia, Kowalewskiella . . . 330 bycanistis, Raillietina .... 296 cabindae, Neomicrodeutopus cacatuina, Raillietina Cacomantis . caeruleus, Uromastyx caflfer, Apus . cafifrapi, Sphaeruterina . cailliautii, Campethera . calcaruterina, Paronia calcostetha, Nectarinia . calidris, Totanus . callonotus, Veniliornis Callolophus . calyptomenae, Raillietina cambriensis, Lembos Campephilus Campethera 4, 9, 12, 13, 17, 18, candidus, Leuconerpes . candidus, Melanerpes canente, Hemicircus canus dedemi, Picus canus, Picus . Capitoniformes carinatus, Hypodontolaimus caroli, Campethera carolinus, Melanerpes carrinii, Paronia carrinoi, Paronia . carrioni, Stenocercus castaneus, Celeus . castor, Doryphorus castor, Uracentron castor, Urocentron cathpharius, Dendrocopos Celeus .... centralis, Mesopicos Centropus Centurus Charadriiformes 95 300 . 284, 327 50 • 335 • 335 21 . 288 . 285, 314 • 330 13 24 . 300 . . 96, 109 • 5, 12, 39-40 19, 20-22, 23,24, 25 . . .16 . 9, 16 . . 6, 7, 14 25, 33 . 3, 24, 25, 26, 33 .308 . 198 . . 9, 12, 13 3 . 287 286, 287 . 58 4 • 55, 57, 58 47, 48 55 31 . 4> 9 . i - 22 284, 285, 331, 333 . 5, 16 . . . .284 INDEX 391 Charadrius . 284, 305, 310, 315, 336, 339 Chatoessus ...... 265 Cheilostomata . . 209-2 30, 288-243 chelifer, Microdeutopus 96, 97, 107, 109, 130, 131, 132-135, 139 chelifera, Stimpsonella . . . 96, 132 chelifera, Stimpsonia . . . 96, 132 chersonesus, Chrysocolaptes lucidus . 36 chilensis, Innocuonema 152, 183, 200, 201 chinensis, Clupea . chinensis, Streptopelia . Chirocentridae Chirocentrus chitwoodi, Chromadorita chlorigaster, Gecinus chlorigaster, Picus chlorolophus, Gecinus . chlorolophus, Picus chlorolophus (rodgeri,) Picus Choanotaenia Chromadora . Chromadoridae . 267 . 284, 297 264, 265, 266-273 264, 265 . 182 24 24, 25, 27-28 24 24, 25, 27-28 28 • 329 168, 182, 183, 190, 201 152, 153, 156, 189-191 Chromadorinae Chromadorita chrysauchen, Melanerpes Chryserpes 153, 191 . 182 17, 18 16 Chrysocolaptes Chrysophlegma chrysopogon, Megalaema chrysopogon, Megalaima Chrysoptilus Ciconiiformes cingulifera, Kowalewskiella circumcincta, Raillietina clavipera, Parvitaenia clivosa, Graphonema clovosa, Graphonema clovosa, Innocuonema Clupanodon . Clupea Clupea (of Brunnich) Clupeidae Clupeonia cochlearii, Parvitaenia . coeruleus, Uracentron Coilia . 3, 9, 12, 13, 35-36, 37, 42 . 24 288, 305 • 285 • 9, i? 284, 296, 306 • 329, 330 295, 296 • 325 . 182 . 201 152, 183, 2OO . 268 261-279, Pis 1-3 • 274 264, 265, 266-273 267, 268 • 325 50 264, 276 Colaptes . 3, 4, 5, 6, 9, 12, 13, 17, 23, 42 colesi, Austranema . . 152, 172, 201 colesi, Euchromadora . 155, 175, 186, 187, 201 coliiformes . . . . .288, 307 Colius 288 Collocalia ...... 335 collocaliae, Notopentorchis . 283, 333-335 colonorum, Agama . . . 69, 72 Columba . . . 284, 288, 294, 297 columbae, Paronia .... 288 columbiae, Aoroides 95 columbiella, Raillietina . . . 301, 302 Columbiformes 284, 288, 296, 300, 302, 303, 306, 307 come, Nematalosa concretus, Hemicircus concretus sordidus, Hemicircus Coraciiformes Coremapus . coryllidis, Paronia Coryllinus . Coryllis . crassa, Chromadorita crassa, Gyrocoelia . crassicaudatus, Stenocercus cruentatus, Melanerpes . cryptacantha, Raillietina Crypturiformes . . , Ctenostomata Cuculiformes . culminatus, Rhamphastos curvirostra, Treron Cyanops . cyathiformis, Anomotaenia Cyclophyllidea Cyclopsittacus Cyclostomata cyprinoides, Clupea cyprinoides, Megalops Cypselus . • 273 . 10, 13, 14 10 296, 300 95, 96, 101 . ' . 288 . 288 . 288 . 182 • 338, 339 . 58 9 300 302 233-235, 243-244 . 284 . 288 284, 304 . 288 . 321 . 281-346 . 288 230-232, 244-245 . 265 . 265 • 335 daggetti, Sphyrapicus varius . . . 6, 1 1 damnoniensis, Lembos . 96, 109, no, 114 damnoniensis, Microdeutopus 97, 109, no, 112, 113, 114-117, 118, 119, 134 darjellensis, Dendrocopos . . 30-3 1 darjellensis desmursi, Dendrocopos . 30 Davainea . . . 287, 290, 308, 309 Davaineidae ..... 287-3 10 debilis, Raillietina .... 306 dedemi, Picus ..... 25 dedemi, Picus canus . . . 25, 33 delgadus, Aorcho ..... 95 Dendrocopos 3, 4, 6, 7, 9, 12, 13, 14, 15, 17, 19, 28-32, 33, 34, 35, 40, 41, 42 Dendropicos . . 4, 12, 17, 18, 19, 31, 42 denticulata, Euchromadora 168, 170, 171, 188, 2OI dentex, Clupea .... 264, 266 depressa, Anomotaenia . . 283, 319-321 depressa, Taenia . . . . .319 dicolorus, Rhamphastos . . . 288 Dicriconema . . 153, 154, 187, 201 dikeniensis, Sphaeruterina . . . 335 Dilepididae .... 316-335 Dilepis .... 283, 316-317 Dinopium . . 3, 9, 12, 13, 33, 35-36, 37, 42 diophthalmus, Cyclopsittacus . . .288 ditchela, Pellona . . . . .271 dolichocephalus, Amphideutopus . . 95 dolloi, Pristigaster . . . .269 domestica, Anas boschas . . 284, 315 domestica, Columba livia . 284,294,297 392 INDEX dorab, Chirocentrus dorab, Clupea dorae, Dendrocopos Doryphorus . Dryocopus . Dryopoides . dubia, Euchromadora dubia, Steineridora (?) dussumieri, Thryssa . 264, 266 . 266 31 47.50,55.57.58.61,62 3, 9, 12, 37-38, 42 95 168, 170, 201 152, 170, 199 . 276 echinobothrida, Raillietina 283, 287-2go, 292, 293 efulensis, Campethera . . . .21 eileenae, Euchromadora 152, 155, 162-163, 168, 169, 192, 193-194, 199, 202 elachus, Dendropicos . . . .19 elliotii, Mesopicos ..... 23 elliotii, Polipicos . . . . 18, 23 elliotii johnstoni, Mesopicos . . .23 elliotii sordidatus, Mesopicos ... 23 encrasicholus, Clupea .... 273 encrasicolus, Clupea . . . 264, 273 encrasicolus, Engraulis . . 264, 273, 274 Engraulidae . . . 264, 265, 273-279 Engraulis . . . 264, 273, 274, 276 Entoprocta .... 235, 245-248 erythrocephalus, Chrysocolaptes lucidus . 35 erythrocephalus, Melanerpes . . . 6, 14 erythrops, Dryocopus .... 37 erythrops, Phloeoceastes . . 38, 39 erythropygasus, Picus .... 24 erythropygius, Picus . . . 25, 27 erythrorhynchus, Rhamphastos . . 288 erythrothorax, Lorius .... 288 Etheridge, R. 45-64 Ethmolaiminae . . . . 153, 191 Euchromadora .... 149-2O4 everetti, Dinopium javanense 35 facile, Anomotaenia facile, Liga .... farciminosa, Hymenolepis fasciatus, Picoiides tridactylus fasciolaris, Fimbriaria fausti, Gyrocoelia . fernandinus, Nesoceleus . festivus, Chrysocolaptes . festucosus, Halex . filiformis, Raillietina Fimbriaria .... flabralis, Raillietina flaccida, Graphonema flaccida, Innocuonema flaccida, Raillietina flaminiata, Raillietina flavescens, Celeus . flaviceps, Doryphorus flaviceps, Uracentron 47, 48, 53, flaviceps, Uranocentron . 317, 318, 319 283, 317-319 • 3i3 • 33 . 283, 315 • 338 6 • 9, 35 . 266, PI. i • 3°2 . 283, 315 300 l82, 183, 201 152, 200 • 302 • 3°° 9 55 54, 55-58, 59, 62 55 flaviceps, Urocentron flavifrons, Melanerpes flavigula, Melanerpes flavinucha, Chrysophlegma flavinucha, Picus . formicivorus, Melanerpes fossiferus, Nygmatonchus fragilis, Pareuchromadora fragilis, Raillietina frayi, Raillietina . . . frigida, Liga . . . . fringillarum, Hymenolepis fuliginosus, Mulleripicus . fulvia, Raillietina . fulvus, Mulleripicus . . fumidus, Dendrocopos . . funebris, Picoides tridactylus . fuscescens, Dendropicos . 152, 55 . 17 17 . 24 . 24, 25, 26 • 9, 17, 18 184, 185, 200 189, 202 300 302 • 319 311, 312, 313 12 . 300 13, 14 • 30 • 33 12, 19 gabonensis, Dendropicos gabonica, Pellona . galeatus, Dryocopus galeatus Phloeoceastes . Galliformes . Gallus gallus, Gallus Gammaridea gaulica, Euchromadora 19 . 269 • 37 . 9, 38 285, 296, 306, 307 285, 287, 295 285, 287, 295 91-148 152, 155, 161, 162, 163, 165, 168, 199 9, 12, 36, 37 . 24 • 3. 4, 5. 18, 23, 42 J52, 155, 166, 178, 183, 192, 195, 196, 200, 202 glandularis, Parvitaenia . . . 325 glareola, Tringa . 284, 295, 322, 328, 329, 330 Gecinulus Gecinus Geocolaptes . georgei, Graphonema glareolae, Choanotaenia . Gnathobolus goertae, Mesopicos Goodwin, D. goura, Raillietina . gracilimembris, Agama . Grandidierella grandimana, Autonoe grandimanus, Microdeutopus Grandison, A. G. C. grantia, Gecinulus Graphonema 152, 155, 166, 329, 331 263, 264, 271 22, 24 . 1-44 . 306 67, 68, 85-88, PI. 3, 6 95. 96, 139 . 98 . 98 65-90 37 178, 180-183, 192, 195, 196, 200, 201, 202 grayi, Ardeola . . . . 317 grillotalpa, Microdeutopus . 98, 117, 119 griseocephalus, Mesopicos . . 22, 24 griseocephalus persimilis, Mesopicos 22, 23 grobbeni, Raillietina . 287, 290, 293, 306 gryllotalpa, Microdeuteropus ... 98 gryllotalpa, Microdeutopus 93, 94, 95, 96, 97, 98-iQi, 102, 103, 104, 106, 108, 109, 114, 119, 134, 139 INDEX 393 guatemalensis, Phloeoceastes . . 11, 38, 39 guentheri, Uracentron 47, 48, 52, 53-54, 5^, 59, 62 guentheri, Urocentron 53 guttacristatus, Chrysocolaptes lucidus . 9, 36 gutteralis, Hirundo rustica . . . 327 Gyrocoelia .... 283, 336-339 haematogaster, Phloeoceastes . . . 37 haemorrhous, Molpastes . . . 288 Halex 266, PI. i hamiltonii, Clupea .... 277 hancocki, Microdeutopus 94, 97, 119, 121, 122, 123, 124, 125, 127, 128, 134, PI. i Hansenella . . . . . 119, 138 harengus, Clupea .... 264, 266 harterti, Dendrocopos major ... 32 haswelli, Microdeutopus 93, 95, 97, 100, 103, 105, 107-109, 138 6, 7, 10, 13, 14 • 313 22 4 95 264, 267-268, PI. i . 40 205-260 285, 327, 335 49 188, 201 154, 175. 201 . 310-315 283, 310-315 153, 191 174, 188, 190, 191, 196, 197-198, 202 • 329, 331 329, 330 . 284, 317 • 330 Hemicircus . hemignathi, Hymenolepis herberti, Campethera nivosa . herminieri, Melanerpes . heteruropus, Acuminodeutopus Hilsa .... himalayensis, Dendrocopos Hincks, T. . Hirundo Hoplocercus . hupferi, Euchromadora . hupferi, Odontocricus Hymenolepididae . Hymenolepis Hypodontolaiminae Hypodontolaimus 152, 15^ hypoleuca, Choanotaenia hypoleucia, Choanotaenia hypoleucos, Actitis hypoleucos, Tringa Ilisha . . . 264, 268-271, PI. 2 ilisha, Hilsa ...... 267 imperialis, Campephilus . . . . 5, 12 indicus, Butastur ..... 330 indicus, Stolephorus . . . .274 inflatispiculum, Euchromadora . 189, 201 Infula 338, 339 Inglis, W. G. . . . . 149-204 Innocuonema . . 152, 183, 200, 201 insularis, Centropus toulou . . .285 intermedium, Dinopium javanense . . 42 Ipocrantor . . . . n, 39-40, 42 Ipophilus . . . . . 1 8, 19 Iserti, Pellona ..... 268 iserti, Pellona .... 269, 270 javanense, Dinopium . . 9, 35, 36, 42 javanense everetti, Dinopium ... 35 javanense intermedium, Dinopium . . 42 javensis, Dryocopus . . . 37, 38 javanica, Notopentorchis . . 334, 335 javanica, Paruterina . . . 334, 335 johnstoni, Mesopicos elliotii ... 23 johnstoni, Polipicos . . . .18 johri, Raillietina . . 283, 291-295, 296, 310 kantipura, Raillietina kelee, Clupea kelee, Hilsa . kempii, Thryssa kiewietti, Gyrocoelia kirkii, Veniliornis . kizuki, Dendrocopos konigi, Mesopicos . korkei, Raillietina . Kowalewskiella kraemmeri, Lembopsis . kraemmeri, Microdeutopus krameri manillensis, Psittacula Lacerta leiopoae, Raillietina Lemboides . Lembopsis Lembos leptosoma, Taenia . leschenaultii, Charadrius 306 267 264, 267-268, PI. i . 276 • 338 17 29, 30 22 . 306 328-331 • 139 95, *39 . 294 47, 283, 295, 310, 49, 50, 5i . 306 95, 139 95, 126, 139 93. 95. 96, ioi, 109, 139 . 287 284, 305, 310, 315, 336, 339 Leuconerpes . leucopogon, Phloeoceastes leucopterus, Dendrocopos leucotos, Dendrocopos leucotos owstoni, Dendrocopos lewis, Asyndesmus Liga .... lignarius, Dendrocopos . lineatus, Dryocopus lineatus, Phloeoceastes . livia domestica, Columba longiannulata, Kowalewskiella longicaudata, Actinonema longicaudata, Euchromadora . longicornis, Hansenella . longipennis, Macropteryx longipes, Autonoe . longipes, Lembos . longipes, Microdeutopus loricata, Euchromadora . loricata, Spilophora loricata, Steineridora Lorius .... lucidus, Chrysocolaptes 16, 42 ii 31, 32 • 6. 33, 34 33, 34 . 6, 14 283, 317-319 30 37 9, 37, 38, 39, 41-42 284, 294, 297 329, 33° . 189 188, 189 119, 138 • 335 101 95, 139 • 139 155, 165, 168 170, 202 152, 166, 199 . 288 . 35, 36, 42 lucidus chersonesus, Chrysocolaptes . . 36 lucidus erythrocephalus, Chrysocolaptes . 35 lucidus guttacristatus, Chrysocolaptes . 9, 36 lucidus strictus, Chrysocolaptes . . 35 394 INDEX luederitzi, Euchromadora lugubris, Dendropicos lutzi, Raillietina Lymnocryptes • 154 19 302-303 . 316 macei, Dendrocopos . . . 31-32 macrodentatus, Hypodontolaimus . .198 macromanus, Lembos .... 95 macropeos, Parvitaenia .... 325 macrophallica, Parvitaenia . . . 325 macrophallus, Infula .... 339 Macropteryx ..... 335 macroscolecina, Raillietina . . . 307 macularia, Actitis ..... 330 maculatus, Microdeutopus . . .139 maculosa, Campethera . . . . 9, 21 magellanicus, Ipocrantor . . n, 39, 42 magna, Parvitaenia .... 325 magniovata, Hymenolepis . . -313 magniuncinata, Vitta . . . 325, 327 mahonae, Hymenolepis . 283, 310-313 major, Dendrocopos . 3, 7, 15, 28, 31, 32 maj or harterti, Dendrocopos ... 32 major parroti, Dendrocopos . . .32 major tianshanicus, Dendrocopos . . 32 malabarica, Thryssa . 264, 274-276, PL 3 malabaricus, Clupea 263, 264, 274-276, PL 3 malabaricus, Thryssa .... PL 3 malacensis, Anthreptes . . . 285, 314 manillensis, Psittacula krameri . . 294 marmoratus, Stenocercus . . -58 martius, Dryocopus . . '3, 37, 38 meadi, Euchromadora . 152, 168, 188, 199 mediterranea, Euchromadora . 155, 179, 201 mediterranea, Protochromadora 152, 179, 200 medius, Dendrocopos . . . 3, 6, 31 Megalaema ..... 288, 305 Megalaima . . . . . .285 Megalopidae ...... 265 Megalops ..... 265, 266 megaloptera, Ilisha . . . .271 megnae, Grandidierella . . . .139 megnae, Microdeutopus . . . 96, 139 Meiglyptes . . . 9, 13, 285, 305 Melanerpes 3, 4, 5, 6, 9, 14, 16, 17, 18, 28, 42 melanoleucos, Phloeoceastes . n, 37, 38, 39 melanota, Ilisha ..... 269 melastoma, Clupea . 264, 265, 270-271 melastoma, Ilisha . . . 264, 270-271 melastoma, Pellona . . . .270 mentalis, Chrysophlegma . . .24 mentalis, Picus . . . . 24, 25 meridiana, Euchromadora . . 188, 201 merulinus, Cacomantis . . . 284, 327 Mesopicos 3, n, 12, 13, 17, 18, 19, 22-23, 24> 25 meyeri, Uracentron . . . 47, 48 meyeri, Urocentrum .. . .58 michaelseni, Raillietina .... 307 micracantha, Raillietina . . . 296 Microdentropus Microdeuteropus Microdeutopus microphallica, Parvitaenia Micropternus Micropus micropus, Ilisha micropus, Notopentorchis micropus, Pellona milligani, Gyrocoelia minax, Microdeuteropus minax, Microdeutopus mineaceus, Callolophus . mineaceus, Chrysophlegma mineaceus, Picus . minimus, Lymnocryptes minor, Clupea minor, Dendrocopos minutiuncinata, Vitta minutus, Nygmatonchus mixtus, Dendrocopos . 96 . 96, 98, 107, 139 91-148, PL i • 325 . 3, 4, 6, 9, 25 • 335 . 271 • 335 270 • 336, 338 . 98 96, 98 24 24 24, 25 • 3i6 266, 267 • 3, 29 • 327 152, 184, 200 30 mixtus berlepschi, Dendrocopos . . 30 Molpastes ...... 288 mortoni, Microdeuteropus . . .139 mucronatus, Gnathobolus . 264, 271, PL 2 mucronatus, Odontognathus . 264, 271, PL 2 mucosoensis, Agama agama ... 74 Mulleripicus . . .12, 13, 14, 38 mutabilis, Anomotaenia .... 328 Myers, A. A. . . . . 91-148 mystacina, Clupea . . . .277 mystax, Clupea . 264, 265, 276-277, PL 3 mystax, Thryssa . . 264, 276-277, PL 3 mystus, Clupea . . . 264, 276, 278 mystus, Coilia .... 264, 278 mystus, Engraulis . . . . .276 nagpurensis, Raillietina • 303- 307 namaquus, Thripias 23 nannalai, Clupea . . 268 nanus, Dendrocopos 19, 29 nasus, Chatoessus 272 nasus, clupea . 263, 264, 272-273, PL 2 nasus, Nematalosa 264, 272-273, H. 2 Nectarinia . 285, 314 Nematalosa . 264, 272.273, PL 2 Neoangularia . 321 Neochromadora 184, 185, 190, 201 Neoliga • 321 Neomegamphopus 95 Neomicrodeutopus • 95 Nesoceleus . 3. 6, 7. H nigrigularis, Trichoglossus . . .288 nivosa, Campethera . 9, 12, 13, 19, 21-22 nivosa herberti, Campethera . . .22 nivosa poensis, Campethera . . .22 noguchii, Sapheopipo . . -4, 33-35 norvegica, Spilophora . . 182, 183, 202 notata, Campethera . . . .21 INDEX 395 Notopentorchis . . . 283, 333-335 Notopteridae ..... 265 Notopterus ...... 265 notopterus, Notopterus .... 265 nova, Raillietina penetrans . . . 303 novaehollandiae, Trichoglossus . . 288 nripendra, Raillietina .... 307 nubica, Campethera . . . 9, 20, 21 nuchalis, Sphyrapicus varius ... 6 Numenius . . . .33. 284, 316, 321 nuttallii, Dendrocopos .... 29 Nygmatonchus 152, 153, 155, 158, 175, 184- 185, 186, 191, 200, 201 nymphaea, Anomotaenia . 283, 321-322 nymphaea, Taenia . . . .321 obsoletus, Dendrocopos . . . 19, 42 obsoletus, Dendropicos . . . 19, 42 obsoletus, Ipophilus . . . 18, 19 obsoletus, Yungipicus . . . .19 oculatus, Amphideutopus ... 95 Odontocricus . . 153, 154, 175, 201 Odontognathus . . 263, 264, 271, PL 2 oedicnemus, Burhinus . . . .318 oglina, Megalops ..... 266 oglinum, Opisthonema . 264, 266-267, PL i oligocantha, Raillietina .... 302 olivaceus, Geocolaptes . . . 4, 5, 28-24 Ophessa ...... 50 Opisthonema . . 264, 266-267, PI- I oreites, Mesopicos ..... 22 orguion, Paradryope .... 95 ornata, Rhips . . . . .184 Orthoskrjabinia ..... 333 owstoni, Dendrocopos leucotos . 33, 34 pachyderma, Graphonema . . .180 pachydermatum, Actinonema . . 185, 189 pagollae, Gryrocoelia . . . 336, 338 palluma, Lacerta ..... 59 palluma, Phymaturus . . . 58, 59 palluma, Uracentron . . . 47, 48 palluma, Urocentron . . . 58, 59 pandispiculatus, Hypodontolaimus . .198 paradoxa, Gyrocoelia . . . 336, 338 paradoxa, Spilophorella . . . .190 Paradryope ...... 95 paraechinobothrida, Davainea 287, 290, 292 parafricana, Euchromadora . 179, 180, 201 parafricana, Protochromadora 152, 179, 200 paragama, Agama 75-78, 79, 87, 88, PL i, 4, 5 paraheterophya, Chromadora . 182, 183, 201 paramicrorhyncha, Anomotaenia . . 322 Parapinnanema 152, 155, 171-177, I78, 191, 192, 194, 199 Pareuchromadora .... 189, 202 Paricterotaenia , 283, 315-316, 322, 323, 324 Paronia . . . 283, 285-287, 288 Paroniella parroti, Dendrocopos major Paruterina . parvirostris, Vitta . Parvitaenia . parviuncinata, Raillietina Passeriformes Passerilepis . passeris, Passerilepis paucitesticulata, Raillietina pectinata, Austranema . pectinata, Euchromadora Pellona penetrans, Davainea penetrans nova, Raillietina permista, Campethera permista, Raillietina permutabilis, Euchromadora . 305-310 32 • 334. 335 • 327 283, 324-325 283, 295-297 285, 288, 300, 303, 308, 313 . 312 • 312, 313 . 300 152, 187, 201 187, 201 268, 269, 270, 271 287, 290, 292 • 303 . 9, 21 • 303. 307 152, 168, 199 persimilis, Mesopicos griseocephalus . 22, 23 perversa, Gyrocoelia perverse, Gyrocoelia phaeopus, Numenius Phloeoceastes Phymaturus . Picidae Piciformes Picoides Piculus 283, 284, 336-339 . 338 33. 284, 321 • 5. 9. ii, 37-39.41-42 58,59 • i-44 285, 288, 302, 303, 307 3, 8, 12, 13, 28, 29, 32-33 9 Picus 3, 4, 5, 9, 12, 13, 15, 17, 24-28, 33, 37, 38, 40, 41 pilchardus, Clupea . . . 264, 266 pilchardus, Sardina . . . 264, 266 pileatus, Dryocopus . . . 37, 38 pileatus, Phloeoceastes . . 5, 9, 38 pintneri, Raillietina .... 307 pitius, Colaptes ..... 6 Platynotus ..... 49, 60 Plica ...... 49, 60 poecilolaemus, Dendropicos . . .19 poecilosomoides, Chromadora . . .190 poensis, Campethera nivosa . . .22 poiken, Clupea ..... 268 Polipicos . . . . . 1 8, 23 pollens, Phloeoceastes . . . .11 polychalix, Raillietina 291, 294, 296, 303 . 198 6 12 269 . 188 • 339. 34i • 93, 96, 109, no • 34i 172, 177-i8o, 199- 200, 201 provincialis, Raillietina .... 296 Pseudangularia . . . . .321 pseudocyclorchida, Parvitaenia . . 325 pseudoechinobothrida, Raillietina 287, 290, 293 Psittaciformes 288, 296, 300, 302, 303, 306, 307 poncticus, Hypodontolaimus portoricensis, Melanerpes principalis, Campephilus Pristigaster . Prochromadorella . Progynotaenia propinquus, Microdeutopus Proterogynotaenia Protochromadora 152 396 INDEX Psittacula ...... 294 Pterengraulis . . 264, 273-274, PI. 3 Pterobranchia ..... 235 Ptilonopus ...... 288 Ptycholaimellus . . . . .198 pubescens, Dendrocopos . . . 3, 29 pulverulentus, Mulleripicus . . 12, 13, 14 punctuligera, Campethera . . 9, 20, 21 punctuligera balia, Campethera . . 20 puniceus, Gecinus ..... 24 puniceus, Picus . . . 24, 25, 27, 28 purava, Thryssa . . . . .277 purpurea, Parvitaenia .... 325 pusilla, Spilophora . . 182, 183, 204 pycnonoti, Paronia . . . 286, 288 Pycnonotus ...... 288 pyrrhogaster, Mesopicos . . .23 pyrrhogaster, Thripias . . . .23 pyrrhotis, Blythipicus .... 9 283, rabieri, Picus radiolatus, Melanerpes rafflesii, Dinopium rafflesii, Gauropicoides Raillietina ramsayi, Cyanops . rectunguis, Centropus Rhamphastiformes Rhamphastos Rhips .... Rhyps .... robustus, Phloeoceastes rodgeri, Picus chlorolophus roosevelti, Neomegamphopus roseiventris, Stenocercus ruber, Sphyrapicus varius rubiginosus, Blythipicus rubrifrons, Melanerpes . rueppelli, Agama . rupicola, Colaptes . rustica, Anomotaenia rustica, Hirundo rustica, Vitta rustica gutteralis, Hirundo ruwenzori, Mesopicos Ryland, J. S. . 24, 25, 26 17 . 9, 36, 37 • 36 284, 285, 287-310 . 288 . 285 . 288 . 288 . 184, 185 . 185 . - . ii, 38 28 95 . 58 ii 12 17 . 84 4. 5, 6 • 325, 327 • 285, 335 283, 325-327 • 327 23 205-260 sabangensis, Chromadora sabulicola, Chromadora . salenskii, Amphithoe sanguinolenta, Agama sankaranica, Agama 67, 68, 73: Sapheopipo . Sardina Sardinella . v savatieri, Agama agama scalaris, Dendrocopos 83-85, 87, l82, 2OI I9O . 98 . 69 78, 79, 82, 88, PL 3, 4, 5 4. 33-35, 42 264, 266, 274 • 274 72, 73 29 scampae, Euchromadora 155, 177, 179, 201 scampae, Protochromadora . 152, 172, 178, i 80, 200 'schmitti' group of Microdeutopus 97, 109, 119, 127 schmitti, Microdeutopus 94, 97, 119, 120-123, 124, 125, 128, 134, 138, PI. i Schneider, J. G. . . . 261-279 schulzi, Phloeoceastes . . -9, 38, 39 scriptoricauda, Campethera . . .? 20 scriptoricauda, Campethera bennettii 20, 21 scriptus, Nygmatonchus . 152, 184, 185, 200 septentrionalis, Euchromadora . .188 setifer, Pareuchromadora . . 189, 202 setirostris, Clupea . . . 264, 277-278 setirostris, Thryssa . . 264, 277-278 sequens, Raillietina 283, 297-299, 300, 301, 304 serpentulus, Hymenolepis . . -313 Sharpei, Picus viridis .... 25 Shipleya . . . . . -339 shirleyae, Austranema 152, 172, 179, 187, 201 shirleyae, Euchromadora . 155, 186, 187, shorii, Dinopium . siamensis, Paroniella siamensis, Raillietina simonsi, Stenocercus sinensis, Centropus sinensis, Clupanodon sinensis, Clupea sinensis, Pycnonotus . g, 36 305-3io 283, 305-310 - 58 . 285 . 268 264, 267-268, PI. i 288 slacksmithi, Hypodontolaimus . 152, 155, 174, 190, 196, 197-igS, 202 sordidatus, Mesopicos elliotii ... 23 sordidus, Hemicircus concretus . . 10 spectabilis, Chromadora . 182, 183, 201 Sphaeruterina ..... 335 sphecotheridis, Davainea . 308, 309, 310 Sphyrapicus . . . .3,6,11,12,14,16 spilogaster, Veniliornis . . . . 4, 30 Spilophora . . . 170, 182, 183, 202 Spilophorella . . . . 153, 190 spinicarpus, Lembopsis .... 95 spinosus, Doryphorus . . . 61, 62 spiralis, Raillietina .... 296 sporadhi, Microdeutopus 97, 108, 136-138 Sprattus .... 264, 266, 274 sprattus, Clupea .... 264, 266 sprattus, Sprattus .... 264, 266 squamatus, Picus . . . -13, 24, 27 stagnalis, Totanus . . . . 330 stagnatilidis, Choanotaenia . 329, 330, 331 stagnatilis, Tringa . . . -33° stateni, Euchromadora . . 168, 171, 201 stationis, Microdeutopus 95, 97, 100, 103, 104-IO7, 108, 109, no, 118, 122, 134, 139 Steineridora 152, 166, 168-171, 179, 199, 2OI, 202 Steironotus . . . . . .61 stellifera, Paricterotaenia . . . . 316 INDEX 397 Stellio Stenocercus . stierlingi, Dendropicos stierlingi, Ipophilus Stimpsonella Stimpsonia . Stolephorus . Streptopelia . striata, Euchromadora striatus, Butorides striatus, Colius striatus, Melanerpes stricklandi, Dendrocopos strictus, Chrysocolaptes lucidus Strobilurus . stylosa, Hymenolepis 47, 50 58, 60 19 18, 19 96, 131 96, 135 • 274 284, 297 152, 155, 162, 164, 168, 191-193, 199 284, 316, 317, 324 . 288 . 4, 16-17 30, 33 35 45-64 • 313 suavissimus, Cyclopsittacus . . . 288 suilla, Chromadora . . 182, 183, 201 superciliaris, Melanerpes . . -17 susanae, Kowalewskiella 283, 295, 310, 328- swainsoni, Trichoglossus synura, Clupea syriacus, Dendrocopos 287 152 5. 12, 13 264, 264, Taenia taeniolaema, Campethera taiwanensis, Raillietina . temminckii, Dendrocopos tentabunda, Chromadora tentabunda, Innocuonema tenuipes, Microdeuteropus tenuis, Dicriconema tenuis, Lembos tenuis, Microdeutopus Thripias thrissa, Clupanodon thrissa, Clupea Thrissina thrissoides, Clupea Thryssa thyroideus, Sphyrapicus tianshanicus, Dendrocopos major tiphia, Aegithina . titii, Microdeutopus tittii, Microdeutopus toco, Rhamphastos tokiokai, Euchromadora toli, Hilsa .... torquatus, Steironotus . torquatus, Strobilurus . Totanus .... toulou, Centropus . toulou, Centropus toulou toulou assumptionis, Centropus toulou insularis, Centropus toulou toulou, Centropus Treron . . . 284 284 . 288 265 3. 4. 40 , 288, 319, 321 . 21 301, 303, 304 3 l82, 201 , 183, 200, 201 • 139 187, 201 • 139 96, 139 17, 18, 19, 23 266, 267 266-267, PL i . 264, 274 • 265 274-278, PL 3 12, 14 • 32 285, 297, 310 • 139 . 288 152, 168, 199 267, 268 61 48, 60, 61-62 • 330 . 285, 331, 333 . 285 . 285 . 285 . 285 , 291, 299, 304 trichoglossi, Paronia .... 287 trichoglossi, Taenia .... 288 Trichoglossus ..... 288 Trichopicus ...... 9 trichopus, Microdeutopus 97, 119, 121, 122, 123, 124, 125, 126-127, 128, 129, 134 tridactylus, Pico'ides . . 8, 28, 29, 33 tridactylus fasciatus Picoides 33 tridactylus funebris, Picoides . . . 33 tridactylus trydactylus, Picoides . . 33 tridens, Lembopsis . . . .139 tridens, Microdeutopus . . . 95, 139 Tringa . . . 284, 295, 322, 328, 329, 330 tringae, Anomotaenia . . 283, 322-324 tringae, Paricterotaenia . . . 322, 323 Tripsurus . . . . . .16 trissa, Clupea . . . 264, 266-267 trivialis, Anomotaenia . . 317, 318, 319 triza, clupea . . . . . .266 Tropidurines . . . . 47, 49 Tropidurus . . . . . 49, 60 trydactylus, Picoides ... 28, 29 trydactylus, Picoides tridactylus . . 33 tukki, Meiglyptes . . . .285, 305 tullbergi, Campethera . . . .21 typica, Aora . . . . 95, 139 Unciola undulatoides, Vitta unicolor, Collocalia Uracentron . Uranocentron Urocentron . Urocentrum . Uromastyx . • 95 • 327 • 335 45-64 • 47. 50, 55 47. 50, 55, 58, 59 47, 58 47, 50 vaillantii, Picus . . . .24, 25-26 validus, Chrysocolaptes . . . • 9, 35 valliclusa, Raillietina weissi . . 303, 304 variabilis, Paronia ..... 288 varius, Sphyrapicus . . 3, 6, n, 12, 14 varius daggetti, Sphyrapicus . . . 6, n varius nuchalis, Sphyrapicus ... 6 varius ruber, Sphyrapicus . . .11 Veniliornis . . . . 4, 12, 13, 17, 30 vernans, Treron . . . 284, 291, 299 versiculata, Autonoe . . . . 101 versiculatus, Coremapus . . 96, 101 versiculatus, Lembos . . . 96, 101 versiculatus, Microdeutopus 95, 96, 97, 99, 100, 101-104, 106, 108, 109 vesiculigera, Notopentorchis . . . 335 viguieri, Lembos ..... 95 villosus, Dendrocopos . . 3, 28, 29, 33 vincenti, Campethera bennettii . . 20 vireus, Megalaema .... 288 viridanus, Picus . . . . 24, 27 viridis, Gecinulus 37 INDEX viridis, Picus 3, 4, 15, 24, 25, 26, 37, 38, 41 viridis sharpei, Picus .... 25 vitrirostris, Thryssa . . . .276 Vitta 283, 325-327 vittatus, Picus . . . .24, 27, 40 volzi, Davainea . . . 287, 290, 292 vulgaris, Chromadora . . . .168 vulgaris, Euchromadora 152, 155, 157, 160- 161, 165, 176, 199 vulgaris, Graphonema 152, 180, 181, 183, 195, 200 vulpes, Albula ..... 264 websteri, Lembos . websterii, Lembos . websterii, Microdeutopus weidholzi, Agama . weissi, Raillietina . weissi valliclusa, Raillietina wellsi, Campethera • 95 96, 139 • 139 67, 85, 86, 88 301, 303, 304 • 303, 304 21 werneri, Raillietina .... 307 werneri, Uracentron 47, 48, 52, 54, 55, 59, 62 westwoodi, Dryopoides .... 95 Whitehead, P. J. P. . . . 261-279 wilsoni, Austranema .... 202 wilsoni, Parapinnanema 152, 155, 171, 174, 176, 177, 178, 191, 192, 194, 199 Woodpeckers ..... 1-44 xanthocephalus, Mesopicos xanthocephalus, Thripias xanthopygaeus, Picus Yungipicus . zavattarii, Paronia , zosteropis, Hymenolepis • 23 23 24, 27 288 PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING