ss BUOLEERIN OF THE BRITISH MUSEUM (NATURAL MISTORY) ZOOLOGY Wolk 2 1971-1973 BRITISH MUSEUM (NATURAL HISTORY) LONDON: 1977 DATES OF PUBLICATION OF THE PARTS INO: i 5 October 1971 Nov 2%. 31 December 1971 INGE 3} 0 31 December 1971 No. 4 . 3 March 1972 INOS in 3 March 1972 No.6. 29 March 1972 No7): 30 March 1972 INO! 8) = 14 June 1972 No.9 . 16 March 1973 ISSN 0007-1498 Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4,5NU CONTENTS ZOOLOGY VOLUME 22 Hyoid and ventral gill arch musculature in eae ag fishes. By P. H. GREENWooD (Pls 1-2) The clupeoid fishes described by Francis aa By P. K. ee & P. J. P. WHITEHEAD Fine structure of Bodo saltans and Bodo eds ere Protozoa) and their affinities with the lees ae oy B. E. BRooKER (Pls 1-6) The type specimens and identity of the species Bean in ae genus Lithobius by C. L. Koch and L. Koch from ee to a By E. H. Eason : 5 : : : Contributions to the life-histories and development of Sones minutus Rudolphi, 1819 and C. heterochrous Bees 1802. 2) D. I. GiBson 2 Bats from Thailand a Geese. By Ne 135 Lebate & K. THONGLONGYA. c 6 A redescription of Suen servatus (Fage) Sue nov. (Mysidacea Lepidomysidae) from the material collected on Aldabra Atoll, with a key to the species of Lepidomysidae. By R. W. INGLE Recent records of mammals (other than bats) from Ethiopia. By G. B. CorBet & D. W. YALDEN . : . : 5 The shell structure and mineralogy of the Bivalvia. II. Lucinacea— Clavagellacea conclusions. By J. D. TayLor, W. J. KENNEDY & A. Hatt (Pls 1-15) ‘ Index to Vol. 22 103 I5I 171 197 211 253 295 eS . MUSCULATURE IN : . E _ OSTEOGLOSSOMORPH FISHES P. H. GREENWOOD ee 2) "BULLETIN. OF ° | E ne (NATURAL HISTORY) Vol. 22 No. 1 _ LONDON : 1971 HYOID AND VENTRAL GILL ARCH MUSCULATURE IN OSTEOGLOSSOMORPH FISHES BY PETER HUMPHRY GREENWOOD Pp. 1-55; 21 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 22 No. 1 LONDON : 1971 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, 1s issued in five series, corresponding to the Departments of the Musewm, and an Historical sertes. 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. 22 No. i of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). © Trustees of the British Museum (Natural History), 1971 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 5 October, 197% Price {1-80 HYOID AND VENTRAE GILL ARCH MUSCULATURE IN OSTEOGLOSSOMORPH FISHES By P. H. GREENWOOD CONTENTS Page INTRODUCTION 0 : 4 MATERIALS AND METHODS . . : : : é : : : 7 HIoODONTIDAE: Hiodon alosoides ; 9 OSTEOGLOSSIDAE: Osteoglossum bicirr, aR 0 0 9 g 3 12 Scleropages leichardti . F ° ‘5 3 a 13 Heterotis niloticus 0 5 : : 6 : 14 Avapaima gigas. ; ¢ ¢ : : . 14 PANTODONTIDAE: Pantodon buchholzi ¢ : - : : 0 16 NOTOPTERIDAE: Papyrocranus afer . : a é F é 6 18 Xenomystus nigri . 21 Papyrocranus and Xenomystus eocpered an Notopterus 22 MorRMYRIDAE: Mormyrus kannume 23 Mormyrus caschive 27 Mormyrus lacerda 28 Mormyrus hasselquisti 28 Cyphomyrus discorhynchus . 30 Marcusenius cyprinoides . : . 5 : a 31 Marcusenius victoriae ci : F F Z : 32 Gnathonemus longibarbis . 0 , 4 0 : 32 Campylomormyrus elephas . @ 0 5 : 6 33 Petrocephalus bane. 0 A 2 OD 35 Petrocephalus catostoma * - ; A é a 37 Isichthys henryi : é : é é a : 37 Mormyrops anguilloides : 5 : é : 5 38 Hyperopisus bebe : : : : 0 : : 40 GYMNARCHIDAE: Gymmnarchus niloticus : ‘ F F i 9 44 SUMMARY AND DISCUSSION. ° 0 : ° 6 : c 48 Phyletic relations within the Sopreaity F : Z 5 4 . 53 ACKNOWLEDGEMENTS ¢ cs : : : o 5 . 54 REFERENCES . 5 F o c 3 6 9 el ‘i 0 54 ABSTRACT THE ventral hyoid and gill arch musculature in several representatives of all osteo- glossomorph families is described and compared. The results of this survey show clearly the unusual specializations of the Mormyridae, and add ‘evidence to the 4 P. H. GREENWOOD suggested relationships of the Mormyridae to the Notopteridae. The Osteo- glossidae and Pantodontidae show a different (and more usual) type of specialization. The Hiodontidae have a basic teleostean pattern in the hyoid muscles, not far removed from the Amza pattern. Based on these and other characters, a new intragroup classification is suggested for the Osteoglossomorpha (including the reduction of the Gymnarchidae to sub- familial status within the Mormyridae). INTRODUCTION Although the ventral gill arch and hyoid musculature has been described (with varying degrees of thoroughness) for some osteoglossomorph fishes (Holmquist, 1911; Munshi, 1960; Bishai, 1967; Nelson, 1969), to date there has been no fully compara- tive account of these muscles; furthermore, only a few species have been investigated. The present study is an attempt to fill both these gaps, and is part of a continuing investigation of intragroup relationships among the Osteoglossomorpha. That the musculature might provide some information on this subject is suggested by Nelson’s (1969) examination of dorsal and ventral gill arch muscles in certain notopterid, mormyrid, osteoglossid and hiodontid species. Nelson’s results are somewhat equivocal (and some are modified by my study), but if the hyoid musculature is also taken into account, a rather different picture emerges. Nomenclature. For the gill arch muscles I have followed the nomenclature used by Nelson (1967 and 1969) which, in turn, was based on the terminology of Vetter (1878) and Edgeworth (1935). The hyoid muscles provide something of a nomenclatural problem. The major muscle connecting the hyoid bar with the lower jaw is generally called either pro- tractor hyoideus or geniohyoideus. (See Holmquist, 1g11; Dietz, 1912, Edgeworth, 1928 and 1935; Munshi, 1960, and Osse, 1969 for discussions of this problem). It is clear that on grounds of homology and ontogeny (Edgeworth, op. cit.) the muscle should not be called a geniohyoideus in teleost fishes. Associated with this muscle there is usually a much smaller, transverse muscle, the so-called intermandibularis, which lies anteriorly between the rami of the jaws. As Holmquist (of. cit.) suggested, and Edgeworth (1928) later demonstrated embryologically, the protractor hyoideus is a compound muscle derived from an intermandibularis component anteriorly (the so-called posterior intermandibularis as distinct from the transverse anterior intermandibularis), and an interhyoideus component posteriorly. Among the osteoglossomorphs studied, the protractor hyoideus shows, at least superficially, varying degrees of complexity or unity. In many osteoglossoids it appears to be a single muscle, in hiodontids a single muscle obviously of compound origin, and in mormyrids and notopterids a group of distinct muscles. Thus, in osteoglossoids I shall refer to this muscle as the protractor hyoideus, and in the notopterids and mormyrids the components will be named (vz. interhyoideus and posterior intermandibularis). The use of the term “‘protractor’ here is purely OSTEOGLOSSOMORPH HYOID MUSCULATURE 5 nominal and does not imply any functional attributes (see Osse, op. cit.). In most species the anterior intermandibularis is clearly identifiable despite its great variation in size and area of attachment. There is general agreement on the nomenclature for that part of the constrictor hyoideus ventralis associated with the branchiostegal rays, namely the hyohyoideus, and that term is used in this paper. lic. 1. Outline drawings, not to scale, of: A. Huiodon alosoides. B. Osteoglossum bicivrhosum. C. Scleropages leichardti. D. Heterotis niloticus. E. Avapaima gigas. F. Pantodon buchholzi. G. Papyrocranus afer. HH. Xenomystus nigri. P. H. GREENWOOD Fic. 2. Outline drawings, not to scale, of various mormyrid species: A. Mormyrus kannume. B. Mormyrus caschive. C. Mormyrus lacerda. D. Mormyrus hasselquisti. E. Cyphomyrus discorhynchus. F. Marcusenius cyprinoides. G. Marcusenius victoriae. H. Gnathonemus longibarbis. 1. Campylomormyrus elephas. J. Petvocephalus bane. K. Isichthys henryi. L. Mormyrops anguilloides. M. Hyperopisus bebe. N. Gym- narchus niloticus. OSTEOGLOSSOMORPH HYOID MUSCULATURE 7 MATERIALS AND METHODS Species Hiodon alosoides Hiodon alosoides Osteoglossum bicirrhosum Scleropages leichardti Scleropages formosus Heterotis niloticus Arapaima gigas Pantodon buchhola Papyrocranus afer Papyrocranus afer Xenomystus nigri Xenomystus nigri Xenomystus nigrt Notopterus kapirat Notopterus kapirat Mormyrus kannume Mormyrus lacerda Mormyrus hasselquisti Cyphomyrus discorhynchus Marcusenius cyprinotdes Marcusenius victoriae Marcusenius victoriae Gnathonemus longibarbus Campylomormyrus elephas Campylomormyrus elephas Petrocephalus bane Petrocephalus bane Petrocephalus bane Petrocephalus catostoma Isichthys henryi Tsichthys henryt Mormyrops deliciosus Mormyrops anguilloides Hyperopisus bebe Hyperopisus bebe Gymnarchus niloticus Gymnarchus niloticus Gymnarchus niloticus B.M.(N.H.) register number 1965.7.2 : I-3 1965.11.23 : 1-8 1926.10.27 : I-2 1966.9.23 : 5 1966.9.5 : I unregistered unregistered 1913.3.12 : I 1969.3.26 : 27 unregistered unregistered unregistered unregistered 1931.7.20 : Q-I7 1938.2.22 : II unregistered 1965.3.15 : 316-320 unregistered 1966.7.29 : 2-4 1g61.12.1 : 48-74 1961.6.13 : 16 1962.2.6 : 26 1928.5.24 : 2 1928.7.30 : 3-4 I919.9.10 : 61 1907.12.2 : 231-232 1905.3.15 : I-2 unregistered 1961.6.21 : 7-18 1958.9.18 : 5-6 unregistered 1969.3.26 : 28 unregistered 1948.6.30 : I-4 1969.3.25 : 34 1969.3.26 : 49 1948.6.30 : 21 1902.11.10 : 56 Dissections were made on the following specimens; for most species dried skeletons or alizarin transparencies were also available. Standard length, mm. 185, 143, TI2, 150, 136 124, I10, 108, 105 286 225 145, 140, 140, 130 167, 155 210, 200 137, 125 132 134 150 104 112 76, 73, 72, 68, 60 200 270, 235 205 190, 195 379 330 290 240 145, 140, 140 A: AIM: Apn: ASCh: Al Bb 1-3: BhTp: BrR: BrM: Bt: CbI-V: CbM: Ch: Che: ChUh: (ibe Cr CTM: bD: Gr: HaBM: HbI: HbIL: Hhy: Hhya: Hp: Hpic: IH: IHhy: Ihyi: Ihyim: Pp. H. GREENWOOD Species B.M.(N.H.) Standard length register number mm. Gymmnarchus niloticus 1953-7-10 : 5 400 Amia calva unregistered 48 (head only) Albula vulpes 1949.11.29 : 1-4 177, 157, 152 Elops saurus 1961.8.31 : 45 134 ABBREVIATIONS USED IN TEXT FIGURES Articular Thyl: interhyoideus muscle, lateral anterior intermandibularis muscle division aponeurosis Ihyvl: interhyoideus, ventrolateral muscle articular surface for ceratohyal on IOp: interoperculum 1st basibranchial (Nelson’s [1968] Le: ligament from basihyal tooth plate terminology) anterior transversus muscle basibranchial, 1st—3rd arch basihyal tooth plate branchiostegal ray branchiostegal membrane basihyal, and its tooth plate, with connective tissue cover ceratobranchial (arches I-V) muscle connecting 4th and 5th ceratobranchials ceratohyal cartilaginous part of ceratohyal muscle between ceratohyal and urohyal cleithrum connective tissue surrounding lateral and ventral edges of basi- hyal and its anterior tooth plate connective tissue mass covering posteroventral end of basihyal tooth plate and articulation of the urohyal dentary gill rakers hypaxial body musculature hypobranchial (arch 1) of second arch hyohyoideus muscle anterior portion of hyohyoideus muscle hypohyal hypohyal (cut through) interhyoideus muscle inferior hyohyoideus muscle interhyoideus muscle, inner (medial) division interhyoideus muscle, innermost division (to urohyal) LUh: ligament from urohyal to hypohyals Max: cut end of maxilla Mb: mental barbel MC: mental cartilage Ob 1-3: obliquus muscle (1st—3rd gillarches) PhCA and P: external and internal pharyngocleithralis muscles PIM: posterior intermandibularis muscle PIMm: lateral muscle bands of posterior intermandibularis muscle PIMms: posterior muscle slips of posterior intermandibularis muscle PIMt: tendinous portion of posterior intermandibularis muscle POp: preoperculum Proll; bony process from second hypo- branchial QO: quadrate R: rectus muscle RC; rectus communis muscle S: sulcus between posterior inter- mandibularis and interhyoideus muscles Sc: lateral line sensory canal in dentary SH: sternohyoideus muscle SHI: sternohyoideus, lower division SHu: sternohyoideus, upper division SHhy: superior hyohyoideus muscle SOpBr: subopercular branchiostegal rays it teeth TIhy: tendon from interhyoideus muscle TIhy 1: tendon from lateral division of interhyoideus muscle TSH: tendon from sternohyoid TSHBb: tendon from sternohyoid muscle to second basibranchial Uh: urohyal T-Vi; gill arches OSTEOGLOSSOMORPH HYOID MUSCULATURE 9 Family HIODONTIDAE Hiodon alosoides (Rafinesque) (Text-fig. 1A) Protractor hyoideus complex (text-fig. 3). Although at first sight there appears to be only a single muscle forming the floor of the mouth, closer examination of fibre direction shows that it is a compound of: (1) a large posterior sheet (fibres running obliquely anteromedially towards the median aponeurosis) (2) a much smaller, oval sheet anteriorly (fibres transverse to postero-medial) and (3) on each side, joining the former two sheets, a narrow elongate muscle with almost longitudinally directec fibres. The various components are tightly joined through narrow aponcuroses, and there is some exchange of fibres between the different muscles. The posterior sheet originates on the ceratohyal, with a few fibres stemming from the bases of the fourth and fifth branchiostegal rays; a fine median aponeurosis is visible along the entire length of the muscle. A sheet of dense connective tissue joins the lateral margins of this muscle to the ventral margin of the dentary on each side. Anteriorly the muscle ends aponeurotically on a broad sheet of connective tissue extending between the rami of the jaw, and lying dorsal to the other muscles of the protractor hyoideus complex. The posterior sheet joins, aponeurotically, the oval median muscle and, on each side, the slender elongate muscles. Thus, it has no direct insertion onto the lower jaw. In contrast, both the median oval and the lateral rectangular muscles attach directly to the median ventral face of each dentary. Nt BrR ag XC \ ( ieee Fic. 3. Hiodon alosoides: ventral hyoid muscles after removal of the skin. 10 Po HH. GREENWOOD From its innervation (a branch of the hyohyoideus VII nerve) and its origin on the ceratohyal, I would identify the posterior muscle sheet as the conjoined left and right interhyoidei. The anterior complex of three muscles (all innervated by a branch of the mandibular V nerve) would seem to be, from their topographical positions, an unpaired median anterior intermandibularis (the oval muscle) and a posterior inter- mandibularis divided by the backward extension of the anterior intermandibularis. There is a striking similarity between the protractor hyoideus complex in Hiodon and the condition found in Amia, Elops and Megalops (personal observations on A. calva, E. saurus and M. cyprinoides; also Allis, 1897; and Liem, 1967). In all, there is a single anterior intermandibularis, paired (but medially joined in Ama and in the elopoid genera) posterior intermandibulares (the geniohyoideus of Liem of. cit.), and an extensive, paired interhyoideus. Hiodon differs from the others chiefly in having the interhyoid components more compact and closely associated with each other. Liem (op. cit.) describes musculose insertions of all components onto the gular plate, but I was unable to confirm this in my own dissections. Indeed, the muscles seem to be completely free from the gular plate which is attached only to the connective tissue covering the muscles. The protractor hyoideus in all four genera differs markedly from that in Albula (personal observations; see also Holmquist, 1g11; Liem, op. cit.), which represents a relatively specialized condition (see also p. 44 for further comments on Albula and Gymnarchus). Hyohyoideus. The superior (interbranchiostegal) part of this muscle is weakly developed, and is largely tendinous (text-fig. 3). The inferior part is much better developed. It originates on the first (7.e. lowermost) branchiostegal ray of each side and inserts, mainly, on the hypobranchial of the opposite side (left muscle lying below the right). From the medial side of each inferior hyohyoideus a short slip of muscle inserts, through a long shared tendon, onto the basihyal plate about half way along its length. The tendon is closely applied to the ventral face of the basihyal plate even before its actual point of insertion. Sternohyoideus (text-fig. 4). The main, ventrally situated part of this muscle inserts directly onto the urohyal, and has its origin aponeurotically, from the ventral body musculature. No ventral part of the sternohyoid originates on the cleithrum, but there is a smaller dorsal component originating on the horizontal limb of that bone. From about the middle of this upper segment a broad-based, almost com- pletely tendinous slip runs forward and upwards to insert on the basibranchial of the second gill arch. Apart from this link, there is no connection between the sterno- hyoideus and the branchial skeleton. The possible significance of this tendon in the evolution of the tendon-bones and ventral bony processes associated with the second gill arch in all other osteoglossomorphs (see Greenwood, Rosen, Weitzman, and Myers, 1966; Nelson, 1968) will be discussed later (page 51). Ventral gill arch muscles (text-fig. 4). Well-developed obliqui muscles are present on the first three gill arches. The rectus communis is a large muscle at its origin from the ceratobranchial of the fourth arch, but it becomes tendinous as it passes below the medial end of the second obliquus, and remains tendinous until its insertion on the second basibranchial. This tendon also attaches to a small process from the OSTEOGLOSSOMORPH HYOID MUSCULATURE 11 second hypobranchial, and is very closely associated with, but distinct from the ligament joining the first basibranchial to the second hypobranchial. A poorly- defined rectus is present between the fourth ceratobranchial and the third hypo- branchial; it is barely distinguishable from the larger and laterally situated rectus communis. Well developed anterior and posterior transversi link the proximal ends of the fourth and fifth ceratobranchials respectively. External and internal pharyngocleithrales are present. The origin of the external division is medial to the sternohyoideus, but the internal muscle originates from the cleithrum at the same level as the sternohyoideus, and superficially resembles a division of that muscle. AT PhCP PhCA Uh TSHBb SH HaBM CL. sss) Fic. 4. Hiodon alosoides: ventral gill arch musculature and sternohyoideus in left lateral view. 4 12 P. H. GREENWOOD Family OSTEOGLOSSIDAE Subfamily OSTEOGLOSSINAE (Nelson, 1968) Osteoglossum bicirrhosum Vandelli (Text-fig. 1B) Protractor hyoideus: is a stout muscle, unpaired except posteriorly over its origins (on each ceratohyal and, in part, aponeurotically from the inferior hyohyoideus of each side). It inserts, through a very short tendinous portion, onto the anterior part of the dentary around the symphysis. This anterior section of the muscle is split, horizontally, by the passage of a stout, transversely aligned anterior inter- mandibularis; that part of the protractor lying above the intermandibularis is thicker. There is no median longitudinal aponeurosis, but there is a distinct tendinous inscription running transversely at about the middle of the muscle. Holmquist (1911) and, later, Edgeworth (1935, fig. 277) described the muscle as divided horizontally into a broader, dorsal, interhyordeus portion, and a narrower, ventral (and medial) posterior intermandibularis portion; both authors also figure the transverse inscription. Despite careful dissection and probing, I could not find any such horizontal division in the muscle. Indeed, in the two specimens I dissected (28-6 and 34-0 cm standard length) the muscle could better be interpreted as being transversely and vertically divided. For example, that section lying anterior to the inscription is innervated by a branch of the mandibular V nerve, but most of the muscle behind the inscription is supplied by a branch of the hyohyoideus VII. This would seem to imply that the anterior part is derived from the posterior inter- mandibularis, and the posterior part from the interhyoideus However, since the inscription does not penetrate deeply into the muscle (7.c. it is not a complete, plate- like aponeurosis) and because the branch of the trigeminal nerve extends behind it, I would not be prepared to delimit the component parts on adult morphology alone. My uncertainty is reinforced by the condition of the muscle in Pantodon buchholz (see page 16), where it seems to approach closely the condition described by Holm- quist for Osteoglossum bicirrhosum. Hyohyoideus. The superior portion (between the branchiostegal rays) is moder- ately developed. The inferior portion, although narrow, is fully muscular. It originates entirely from the first branchiostegal ray and has a tendinous insertion mainly onto the hypohyal of the opposite side; a few fibres, however, have a tendi- nous insertion onto the hypohyal of their own side. (It may be noted that Holmquist [1911, fig. 11] shows the right inferior hyohyoideus overlapping the left, but in all specimens I have examined [and in all other osteoglossids] left overlaps right). Sternohyoideus (text-fig. 5). The greater part of this muscle originates on the dorsal surface of the horizontal limb of the cleithrum; a small part stems from the anterior tip of the conjoined cleithra. The sternohyoideus inserts onto the urohyal, but that part passing below the first gill arch is closely attached to the hypobranchial OSTEOGLOSSOMORPH HYOID MUSCULATURE 13 by a thick connective tissue fascia. The muscle is also closely attached to the inner aspect of the ventrally directed bony processes on the second hypobranchials; each process is, however, entirely superficial to the muscle. Ventral gill arch muscles (text-fig. 5). Obliqui muscles (linking cerato- and hypo- branchial elements) are present on the first three gill arches. A small rectws muscle connects the fourth ceratohyal with the third hypobranchial. No rectus communis is developed. The proximal ends of the fourth and fifth ceratobranchials are joined, respectively, by the anterior and posterior transversi. The fifth ceratobranchial is also connected to the cleithrum by the strong external and internal pharyngocleithrales. The internal pharyngocleithralis is subdivided into a narrow posterior and a much broader anterior part. Both pharyngocleithrales have their origins medial to the sternohyoideus. Scleropages leichardti Giinther (Text-fig. 1C) In most details, the musculature of S. leichardti is identical with that described for Osteoglossum bicirrhosum. Comments made above on the morphology of the protractor hyoideus muscle and its components apply equally to the muscle in Scleropages. HbI HbI Gr PhCP HaBM Hpic Bb1 Pri SH ees Fic. 5. Scleropages leichardti: ventral gill arch muscles and sternohyoideus in left lateral view. The hypohyal has been partly cut away. 14 P. H. GREENWOOD Among the ventral gill arch muscles, the only intergeneric difference noted was that in Scleropages the proximal end of the rectus muscle is fused with the hypo- branchial head of the obliquus of the third arch (text-fig. 5). Subfamily HETEROTINAE (Nelson, 1968) Heterotis niloticus (Cuvier) (Text-fig. 1D) Protractor hyoideus. This stout muscle originates on the ceratohyal and lower three branchiostegal rays of each side, but is a single element over its anterior half. It inserts directly, through short left and right heads, on either side of the dentary symphysis. Slightly anterior to the point of tusion between left and right halves, there is a very distinct transverse inscription; no other division of the muscle can be detected by dissection. The anterior part of the protractor is supplied by a branch of the mandibular V nerve, the posterior part by a branch of the hyohyoideus VII. The latter branch emerges from the medial side of the branchiostegal membrane at the base of the first ray; the main nerve continues forward, and supplies the inferior hyohyoideus muscle. Heterotis is unusual in having a strong connective tissue link between the pro- tractor hyoideus and the hypohyals (at a point near the union of the protractor’s two halves). A small and narrow anterior intermandibularis lies dorsal to the protractor hyoideus and does not pass through any part of it (cf. Osteoglossum and Scleropages). Sternohyoideus. This stout muscle originates almost entirely from the dorsal surface of the horizontal limb of the cleithrum; a few ventral fibres stem from the anterior tip of that bone. It inserts onto the urohyal but is also firmly attached to the hypobranchial of the second arch and, less intimately, to the first hypobranchial as well. The ventral processes from the second hypobranchials are partly buried in the sternohyoid. Ventral gill arch muscles. The most outstanding feature of these muscles is the development of a rectus communis from the fourth ceratobranchial to the base of the process on the second hypobranchial; there is also a tendinous connection between this muscle and the third hypobranchial. In all other respects the gill arch muscles (including the vectws) are like those of Osteoglossum and Scleropages. The pharyngocleithrales of Heterotis, however, are simpler since the internal muscle is undivided. Arapaima gigas (Schinz) (Text-fig. IE) Protractor hyoideus: is a short, largely paired muscle originating from the cerato- hyal and the basal parts of the second and third branchiostegal rays. Only about OSTEOGLOSSOMORPH HYOID MUSCULATURE 15 the anterior third of the muscle is unpaired, the two halves meeting along a weak aponeurosis. Slightly anterior to this junction there is a transverse tendinous inscription. The protractor inserts, through left and right musculose heads, on either side of the dentary symphysis. A small and narrow anterior intermandibularis passes through the muscle a little posterior to its insertion. It is impossible, by inspection or dissection, to determine the extent of the protractor’s component muscles, nor is it possible to determine the manner of their fusion. Hyohyoideus. The superficial part is moderately well-developed and largely muscular. The inferior division is short and relatively stout; it inserts through a long tendon onto the hypohyal of the opposite side. Sternohyoideus. Undoubtedly the sternohyoideus is the most characteristic muscle of Avapaima gigas (see text-fig. 6). It is divided, horizontally, into a small anterior and dorsal division, and a much larger unpaired ventral part. The latter originates (as is usual) on the horizontal limb of the cleithrum, and inserts on either side of the peculiarly shaped urohyal. In cross-section, this bone is shaped like an inverted T, with the arms extended to such a degree that the anterior half of the lower sternohyoid is completely covered by bone. A broad ligament from either side of the urohyal attaches it, ventrally, to the ceratohyals and hypohyals. The small ventral process of the second hypohyal barely contacts the dorsal part of this ventral sternohyoid division. Indeed, the process in Avapaima is the shortest found in any osteoglossid, and has the least intimate contact with the sternohyoid. Uh TSH Bt Smm CbI BrM BrR Gr SHu Cb eee Fic. 6. Avapaima gigas: ventral view of the sternohyoid, to show its upper and lower divisions and their relationship with the urohyal, cleithrum and first hypobranchial. Semi-schematic; the lower jaw is not depicted. Cb=ceratohyal. 16 Pp. H. GREENWOOD The dorsal division of the sternohyoid is paired. Each half originates on the ventral face of the first hypobranchial, and extends over almost the entire length of that bone (thus underlying the first obliquus muscle). The left and right halves meet, aponeurotically, in the midline; the broad tendon originating at this union inserts, somewhat asymmetrically, on the inner aspect of the ceratohyals near their tips. Although some part of the tendon inserts on the right ceratohyal, by far the greater part is attached to the left bone. No other osteoglossid shows such clear-cut subdivision of the sternohyoid, or such asymmetry in its insertion. In fact, insertion of even part of the sternohyoid onto the ceratohyals is most unusual in teleosts. Nevertheless, it is difficult to identify the upper division of this muscle in Avapaima as other than part of the sterno- hyoideus. That other osteoglossids have a close association of the sternohyoid with the first and second hypobranchials, and that there is an incipient division of the muscle in Heterotis (see p. 14), all seem to support the recognition of a divided sternohyoid in Arapaima. Ventral gill arch muscles: in Avapaima have the typical osteoglossid pattern ; there is no rectus communis. Obliqui are present on the first three arches; the obliquus of the third arch is closely associated with the well-developed vectws running almost in the midline from the fourth ceratobranchial to the third hypobranchial. Anterior and posterior tyansversi are present, as are stout and undivided internal and external pharyngo- cleithrales. Family PANTODONTIDAE Pantodon buchholzi Peters (Text-fig. 1F) Protractor hyoideus (text-fig. 7): is a single muscle over its anterior half, but is paired posteriorly, with the left and right halves originating on the ceratohyal and lower three branchiostegal rays of their side. The muscle is marked by a transverse inscription at the point where the two halves unite. At this point, the unpaired portion is visibly separable into a broad dorsal section (inserting on either side of the dentary symphysis) and a much narrower, more compact, median and ventral part which inserts, tendinously, onto the sym- physis itself. Slight pressure with a probe along the horizontal sulcus demarcating the two parts separates them back to the level of the transverse inscription. Beyond this point there is considerable interchange between the parts, and the sulcus itself is no longer distinct. A well-developed, stout, transverse anterior intermandibularis lies between, and separates, the dorsal and ventral parts of the protractor anteriorly. In many respects, the condition of the protractor hyoideus in Pantodon resembles that described for Osteoglosswm by Holmquist (op. czt.) and Edgeworth (1935); it will be recalled that I found a rather different arrangement in that genus (see p. 12). OSTEOGLOSSOMORPH HYOID MUSCULATURE 17 Pantodon differs from Holmquist’s description of Osteoglossum in that the muscles cannot be separated posteriorly beyond the transverse inscription. Holmquist and Edgeworth identify the entire length of the median, ventral section of the muscle in Osteoglossum as a protractor hyoideus (i.e. a posterior intermandibu aris in the terminology used here), and the overlying, broader part as the interhyoideus component. Identifying the components of the protractor hyoideus in Pantodon is not easy, particularly since the innervation cannot readily be traced within the muscle. Judging from the position of the upper and lower insertions, and from the fact that the ventromedial segment is so clearly circumscribed, I would identify it as the posterior intermandibularis; the much larger muscle above, lateral to and behind it would then be the interhyoideus portion. In other words, a situation like that in the protractor hyoideus of Salmo salar (see Holmquist, 1g11, and Dietz, 1912). Hyohyoideus. Both the superior and inferior divisions are well-developed, the latter originating on the first branchiostegal ray, and inserting tendinously on the hypohyal of the opposite side. Sternohyoideus. The lower third of this muscle originates on a broad aponeurosis with the ventral body musculature; the remainder stems from the dorsal surface of the horizontal limb of the cleithrum. Anteriorly, the sternohyoid inserts around the small urohyal which is completely embedded in the muscle. The medial face of Fic. 7. Pantodon buchholzi: ventral hyoid musculature. A. In ventral view. B. In oblique ventro-lateral view. The transverse inscription (aponeurosis) is shown, but not labelled, in both views. 18 P. H. GREENWOOD each process from the second hypobranchial is firmly but superficially attached to the sternohyoid, which is also closely attached by connective tissue to the ventral side of the first hypobranchial. Ventral gill arch muscles. As in Heterotis (but not other osteoglossids) a distinct rectus communis is present; the small vectus from the fourth ceratobranchial to the third hypobranchial is distinct from the rectus communis but is closely applied to it. Obliqui are present on the first three arches; anterior and posterior tvansversi are well developed, and the pharyngocleithrales are simple but relatively stout. Family NOTOPTERIDAE Munshi has given a detailed description of the cranial muscles in the Asiatic species Notopterus chitala, and Nelson (1969) has listed the branchial muscles of the same species. In view of this previous work, I have concentrated on the two African species, Papyrocranus afer and Xenomvystus nigri (see also Greenwood, 1963 and Nelson op. cit.). Some comments on Munshi’s description of the hyoid muscles in N. chitala, and a general comparison of the hyoid and gill musculature in the three genera follow the separate accounts for Papyrocranus and Xenomystus. Papyrocranus afer (Ginther) (Text-fig. 1G) Ventral hyoid musculature (text-fig. 8). The most superficial (7.e. ventral) muscle has its origin, on each side, equally from the ceratohyal and from the basal part of branchiostegal rays 2 to 6. The two halves of this muscle join to form a single element over about the anterior half of their length. Anteriorly, the muscle inserts onto the dentary through dorsal and ventral heads; the single, narrow and tendinous ventral insertion is onto the symphysis, while the broader, more musculose dorsal insertion is double and lies on either side of the symphysis. This ventral muscle is innervated solely by a branch of the mandibular V nerve. Careful dissection shows that no branch of the hyohyoideus VII runs to it (see below under interhyoideus). With respect to its innervation, the muscle differs from the topographically similar muscle (the protractor hyoideus) in osteoglossid and pantodontid fishes. Munshi (op. cit.) identifies the muscle in Notopterus as a posterior intermandibularis and I would agree with his identification, both on the grounds of the muscle’s innervation solely from the trigeminal nerve, and because distinct interhyoideus muscles (innervated from the facialis nerve) are also present. It will be recalled . | OSTEOGLOSSOMORPH HYOID MUSCULATURE 19 SHhy AIM IHhy BrM LA PIM B Che Ch IH BhTp Lom oe CTtcut) D SHhy Hp THhy(left) Ch L Fic. 8. Papyrocranus afer: A. ventral hyoid muscles, after removal of the skin. B. ventro-lateral view of head to show the interhyoideus muscles. The right half of the lower jaw removed, as is the cheek and jaw musculature of that side. 20 P. H. GREENWOOD that in osteoglossoids the protractor hyoideus has a double (trigeminal and facialis) innervation, and that in these fishes no separate interhyoideus is present. A small and weak anterior intermandibularis muscle lies between the dorsal and ventral insertions of the posterior intermandibularis. Its fibres are transversely orientated, and attach to the median face of the dentary on either side of the symphysis. Interhyoideus (text-fig. 8). This paired muscle appears to be further divided on each side into a shallow lower portion, and a deeper upper part (see fig. 8B). Anteriorly, however, the two parts have a common tendinous insertion onto the dorsal surface of the hypohyal of the same side. The upper division originates entirely from the ceratohyal, but the lower part originates from the ventral margin of the ceratohyal and, in minor part, from the heads of the first two branchiostega. rays. The interhyoideus is innervated by a branch of the hyohyoideus VII nervel This nerve enters the lower muscle division after emerging from behind the branchio- stegal rays and passing along the upper margin of the first ray. Hyohyoideus. The superior division is rather poorly developed, and is almost completely tendinous. The inferior part is also tendinous but has its medial third strongly muscular (fig. 8B). This muscular section originates from the distal third of the first branchiostegal ray. The whole inferior hyohyoid inserts, tendinously, on the hypohyal of the opposite side (the left muscle passing ventral to the right). Sternohyoideus (text-fig. 9). The large sternohyoideus originates entirely from the dorsal surface of the horizontal limb of the cleithrum. Ventrally, part of the muscle inserts, through a pair of tendons, onto the hypohyals and the anteroventral tip of each ceratohyal (a most unusual arrangement; see also Avapaima, page 15). However, the bulk of the sternohyoid inserts onto and around the small urohyal which is completely embedded in the muscle. (It should be noted that a pair of strong ligaments from the ventral face of the basihyal tooth-plate also attach to the urohyal and are, in consequence, partly covered by the sternohyoid.) The median face and posterior margin of each ventral process (tendon bones) from the second basibranchial are firmly but superficially attached to the lateral face of the sternohyoid on each side. Anterior to this point the muscle closely approaches the first basibranchial but is not attached to it. Ventral gill arch muscles (text-fig. 9) | Well-developed ob/iqui are present on the first three gill arches; the muscles of all three arches insert on the respective cerato- branchial, but those on the first and second arch have a double origin, from the hypo- and basibranchial. A distinct vectus runs from the third hypobranchial to the ceratobranchial of the fourth arch where it comes into close contact with the head of a stout rectus communis connecting that arch with the base of the ventral process (tendon bone) articulating with the second basibranchial. The prox mal ends of the fourth and fifth ceratobranchials are joined, respectively, by the broad anterior and posterior transverst. The external and internal pharyngocleithrales are well-developed, simple muscles whose origins lie medial to the sternohyoideus. OSTEOGLOSSOMORPH HYOID MUSCULARE 21 Xenomystus nigri (Ginther) (Text-fig. 1H) In all major details the musculature of this species is like that of Papyrocranus afer. The most noticeable differences may be listed briefly. Posterior intermandibularis. The origin is mainly from the ceratohyal, with only about one quarter stemming from the first branchiostegal ray. It inserts through a single broad head, an arrangement possibly correlated with the much weaker and rather ill-defined anterior intermanibularis in this species. As in Papyrocranus the posterior and anterior intermandibulares are innervated by a branch of the mandi- bular V nerve. Hyohyoideus. The inferior divisions fuse in the midline before inserting onto the hypohyals. Interhyoideus: in this species is a pair of undivided muscles fused anteroventrally, and inserting onto the hypohyals through a short tendon which wraps around the face of these bones. Cbr HbI Hpic BhTp LUh SH Uh TSH Chileft) Smm Fic. 9. Papyrocranus afer: lateral view of gill arch muscles and sternohyoideus. The anterior part of the sternohyoid has been dissected away on the right side to show the embedded urohyal. The tendon (TSH) inserts onto the right ceratohyal (removed) and right hypohyal (partly removed). 22 P. H. GREENWOOD Gill arch muscles. 1 was unable to locate a separate rectus in Xenomystus. The rectus communis is present and large; like the obliqui it appears to be relatively larger in this genus. As in Papyrocranus, the obliqui of the first two arches originate from both the hypo- and the basibranchials. PAPYROCRANUS AND XENOMYSTUS COMPARED WITH NOTOPTERUS Basically, the musculature in the three genera is very similar (see Munshi, 1960 for Notopterus). In the two Notopterus species I examined (N. chitala and N. kapirat), the interhyoideus of each side is undivided (i.e. like that of Xenomystus); in N. kapirat it is indistinguishable ventrally from the upper, anterior part of the inferior hyohyoideus. According to Munshi (oP. czt.) the left and right interhyoidei of N. chitala fuse medially. But, in his figure the muscles are shown as fan-like, horizon- tally disposed structures continuous with the inferior hyohyoidei, and running obliquely towards the midline; here the posterior portion ot the left interhyoideus overlaps that of the right muscle. In the single specimen I dissected, the inter- hyoidei are vertically aligned (as in the other species considered), insert separately and are quite distinct from the inferior hyohyoidei. In other words, a situation identical with that found in Papyrocranus. I would contest Munshi’s identification of the muscle running from each basi- branchial process (tendon bone) to the urohyal. This he calls the pharyngohyoideus (= rectus communis in the nomenclature used above). However, the figure shows the muscle as only partly distinct from the sternohyoideus (Munshi’s rectus cervicus) below it, and no fibres are shown connecting with the upper posterior face of the tendon bones (as do those of the rectus communis). In the specimen I have exam- ined (as in N. kapirat, Papyrocranus and Xenomystus also) it is not possible to separate the fibres attached to the tendon bone from those of the underlying sterno- hyoid. Incontrast, the muscle I have identified as a rectus communis is distinguish- able from the sternohyoideus even at its insertion onto the tendon bone where the two muscles are closely apposed. Thus it seems likely that the muscle Munshi identifies as a rectus communis is, in fact, part of the sternohyoideus. Family MORMYRIDAE The bauplan of the hyoid and gill arch musculature is remarkably constant in the Mormyridae, despite the great range of variation in head shape and jaw form of these fishes. The more outstanding features of the mormyrid musculature may be summarized as follows :— (i) The muscles of the protractor hyoideus complex show the greatest degree of individuality and subdivision of any osteoglossomorph fishes; the interhyoideus muscles are enlarged and insert onto the lower jaw. (ii) There is considerable hypertrophy of the hyohyordeus with the consequent loss of distinct superior and inferior divisions; the left and right halves of the muscle I ee OSTEOGLOSSOMORPH HYOID MUSCULATURE 23 join medially and the muscle is attached to the interopercula and ceratohyals as well as to the urohyal. This specialized musculature is associated with the peculiar branchial specializations of all mormyrids. For example, there is no discrete and expandible branchiostegal membrane, the rays being buried in the hyohyoideus, and the whole mass covered by skin continuous with that of the body; in consequence, the opercluar aperture is greatly restricted and lies entirely above the branchiostegal rays. (iii) In the ventral gill arch musculature the obliqui have extensive areas of attachment, and the weakly developed pharyngocleithrales lie external to the sterno- hyoideus. The anterior part of the sternohyoideus is closely associated with the corresponding ventral elements of the gill arches, although there is never a direct musculose or tendinous insertion onto these bones. Surprisingly, little attention has been paid to the cephalic and branchial muscula- ture of mormyroid fishes. Holmquist (1911), drawing on his investigation of the hyoid musculature in Gymnarchus niloticus, noted certain peculiarities, but he was unable to extend his observations because of lack of comparative material. Nelson (1969) briefly commented on the gill arch muscles in Mormyridae and compared these with those of other osteoglossomorphs. The most comprehensive treatment is that of Bishai (1967) on Mormyrus caschive. Unfortunately, the nomenclature used by this author makes direct comparison very difficult, and I suspect that his description of the gill arch musculature is erroneous in many respects (see below, page 28). Since the bauplan of the hyoid and ventral gill arch muscles is so similar in all mormyrids, I shall give a detailed description for one species only; deviations from this pattern will be noted for the other species examined. Mormyrus kannume Forsk. (Text-fig. 2A) The snout in this species is slender, moderately decurved, produced and tubular, with the mouth small and terminal. Posterior intermandibularis (text-fig. 10A): is a fairly thick muscle with originates, aponeurotically, over the anterior part of the hyohyoideus, and directly from the interoperculum of each side. It inserts through two muscular heads on either side of the mental cartilage (that is, it does not attach directly to the dentary). At about its midpoint there is a faint, longitudinal aponeurosis which extends anteriorly; posterior to this point both halves of the muscle are contiguous medially throughout their lengths. The posterior intermandibularis is broad and extends laterally almost to the ventral margin of each dentary, to which it is attached by a connective tissue sheet. Anterior intermandibularis (text-fig. 10B): lies immediately above the posterior division. It is a relatively thin but expansive muscle extending longitudinally from the level of the posterior interopercular margin to a little behind the mental cartilage. P. H. GREENWOOD Fic. 10. Moyrmyrus kannume: ventral hyoid muscles. A. After removal of the skin. B. Posterior intermandibularis removed. C. Anterior intermandibularis removed; right lateral division of the interhyoideus muscle removed. OSTEOGLOSSOMORPH HYOID MUSCULATURE 25 Laterally it inserts, directly, onto the interoperculum, articular and the dentary. All its fibres are transversely arranged; there is a faint but distinct median aponeurosis extending along the entire length of the muscle. Both the anterior and posterior intermandibularis are innervated by a branch of the mandibular V nerve. Interhyoideus (text-fig. 10C). A bilaterally paired muscle originating on the hyoid arch but inserting on the medial face of each dentary and articular, is identified as the interhyoideus because of its origin (the epi- and ceratohyal) and because of its innervation (a branch of the hyohyoideus VII). The lateral (7.e. outer) muscle of each pair is the larger; it has a narrow origin from the upper part of the ceratohyal and part of the epihyal, and inserts tendinously onto the ventral margin of the dentary immediately lateral to the symphysis. The inner and smaller muscle has a much wider origin (entirely from the cerato- hyal) ventral to that of the lateral division. It inserts on the medial face of the dentary, and also the articular, considerably behind and above that of the outer division. Hyohyoideus (text-fig. 10): is a stout, thick muscle with the left and nght halves meeting medially along a fine aponeurosis. The branchiostegal rays are almost completely embedded in muscle; it is impossible to recognize separate superior and inferior divisions. The posterior part of the hyohyoideus is attached to the sub- opercular branchiostegal rays,! the middle section to the interoperculum, and the forward part to the anterior face and ventral margin of the ceratohyal laterally; medially the muscle is attached to the urohyal. All these connections are musculose. Externally, the hyohyoideus is covered by skin which is continuous with that of the body and lower surface of the jaw. In other words, there is no separate branchio- stegal membrane. Internally, the medial face of the hyohyoideus is bound to the overlying ventral body musculature by an extensive connective tissue union. Sternohyoideus: is a large muscle originating mainly from the dorsal face of the horizontal cleithrum, but with a small ventral portion continuous with the hypaxial body muscles. The sternohyoid inserts on the urohyal but its antero-dorsal section is closely associated, through a connective tissue fascia, with the second basi- branchial. The ventral processes from the second hypobranchials are, proximally, embedded in the sternohyoid; distally they lie outside the muscle but closely attached to its lateral face. Ventral gill arch musculature (text-fig. 11). Well-developed obliqui are present on the first three gill arches; all are broad muscles extending onto the ceratobranchial well beyond its articulation with the small hypobranchial. From the first obliquus there is a distinct postero-medially directed slip of muscle which inserts onto the second hypobranchial. From the second obliquus a similar slip runs to, and inserts on, the second basibranchial. The third obliquus links only the cerato- and hypo- branchial of its arch. 1In mormyrids the upper two branchiostegal rays are blade-like bones which have lost their articu- lation with the hyoid arch. Instead, the two bones are firmly attached to one another, and the upper bone is immovably attached to the lower margin of the operculum. Together, these two rays form an apparently immovable pseudosuboperculum. 26 P. H. GREENWOOD A moderately developed rectus communis extends between the fourth cerato- branchial and the proximal end of the ventral process from the second hypobranchial. The muscle is closely applied to, but distinct from, the dorsal margin of the sterno- hyoideus. A short, broad and nearly triangular muscle joins the fourth and fifth ceratobranchials near their proximal ends. I cannot be certain about the identity of this muscle. It could be a displaced obliquus of the fourth arch or, more likely, part of this obliquus since there is a large transversus associated with the arch (all this assuming that obliqui are, primitively, associated with each arch; see Nelson, 1967). The anterior transversus is moderately broad; it links the ventral tips of the fourth ceratobranchials. The posterior transversus runs obliquely forward so that it has the shape of a V, the apex inserting on the cartilaginous block lying between the ventral tips of the third and fourth ceratobranchials (the fourth basibranchial of Nelson, 1968). The arms of the V are closely applied to the ventral surface of each cerato- branchial. HbI CbI Ob2 RC Ob3 BhTp AsCH HbIL SH LoS} Fic. 11. Moymyrus kannume: ventral gill arch muscles and sternohyoideus in left lateral view. Not all the connective tissue surrounding the basihyal and urohyal has been dissected away. OSTEOGLOSSOMORPH HYOID MUSCULATURE 27 External and internal pharyngocleithrales are present, but are not strongly developed. An unusual feature of these muscles in all mormyrids is their origin external to the sternohyoideus Both divisions, but especially the external one, have a deep and broad insertion on the fifth ceratobranchial. Mormyrus caschive Linn. (Text-fig. 2B) Bishai i(1967) has given a detailed account of the cranial and branchial muscles in this species Unfortunately, I have been unable to dissect a specimen, but since in osteologcal and other features M. caschive is very similar to M. kannume, it seems reasonable to assume that the musculature would also be similar. Thus, it is surprising to find a number of departures from the M. kannume condi- tion in the ventral musculature of M. cashive as described by Bishai. Because muscle nomenclature used by this author differs from that usually employed, and because of the anatomical differences noted, I shall give a list of synonyms for the muscles involved and also comment on the apparent discrepancies between my observations on M. kannume and Bishai’s on M. caschive. In general, I would agree with Bishai’s description of the superficial hyoid muscles. His depressor labii inferioris muscle is my posterior intermandibularis, and his inter- mandibularis is my anterior intermandibularis. The posterior intermandibularis of M. caschive (cf. text-fig. 10A with Bishai’s fig. 4) is narrower anteriorly and seems to extend further posteriorly, covering the entire hyohyoideus (i.e. the interbranchio- stegalis muscle of Bishai). There is also substantial agreement in the arrangement of the deep ventral muscles (Bishai’s geniohyoideus internus and externus, which are my inner and outer divisions of the interhyoideus). According to Bishai’s figure 5, there are three divisions of this muscle, the innermost of which is not labelled. However, I suspect that the object which Bishai has labelled “‘geniohyoideus externus”’ is, in fact, a cut section of skin (or even cheek muscle) ; further, it seems that his “internus’”’ muscle should be labelled ‘“‘externus”, and that the short unidentified innermost muscle is the internal division of his geniohyoid series (i.e. my interhyoid series). The same figure also shows a condition of the sternohyoideus which I find difficult to accept. However, I think the figure is explicable when one considers Bishai’s description of the ventral branchial muscles (op. cit., page 21, and fig. 8). Here the author describes three large muscles, originating on each side from the cleithrum, and inserting on the urohyal and the ventral processes of the second hypobranchials. These muscles are identified by Bishai as ‘‘anterior portions of the pharyngoclavi- cularis muscle’. Apart from an abuse of the term pharyngoclavicularis for muscles with these topographical relationships, it seems that Bishai failed to recognize their true identity as parts of the sternohyoideus (see page 25). Bishai’s pharyngocleithralis posterior apparently consists of both the external and internal divisions of this muscle. 28 P. H. GREENWOOD I cannot find separate muscles in M. kannume corresponding to Bishai’s pharyngo- arcualis anterior, and obliquus ventralis anterior and posterior (of the first gill arch, see his fig. 8). Indeed, it seems that he has misinterpreted the double-headed condition of the first obliquus muscle which inserts, mainly, on the first hypo- branchial but also has a slip passing to the second hypobranchial (see page 25). Bishai does not describe a rectus communis muscle but his fourth obliquus ventralis superioris could well be part of a rectus communis (v7z. that portion near its origin on the fourth arch and below the third arch). Likewise, Bishai’s obliquus ventralis superioris 5 seems to correspond to the small muscle, present in most mormyrids, which links the ceratobranchials of the fourth and fifth arches (see page 26). The anterior transversus (i.e., the fourth transversus ventralis of Bishai) requires no comment, but I believe that Bishai has misidentified the posterior transversus, and called it the pharyngoarcualis posterior (see page 51 for a discussion of the posterior transversus in mormyrids). Mormyrus lacerda Casteln. (Text-fig. 2C) In this species the snout is relatively short, broad and but slightly decurved; the mouth is broad and terminal in position. With few exceptions, the musculature is like that of Mormyrus kannume. The anterior intermandibularis inserts onto the dentary and angular only; over its posterior third it fails to reach the lateral margins of the head. However, its postero- lateral tips are attached to the ceratohyal near the origin of the interhyoideus. The posterior intermandibularis, relative to that of M. kannwme, is somewhat less substantial over its posterior half. The pharyngocleithrales are complex. There are three distinct but contiguous heads on the cleithrum, all originating lateral to the sternohyoideus. At about the midpoint between girdle and ceratobranchial, the three separate muscles fuse, become tendinous and then, as a single element, become muscular again. Presum- ably this muscle should be considered as fused external and internal pharyngo- cleithrales. Mormyrus hasselquisti Val. (Text-fig. 2D) In this species the snout is short, broad and but very slightly decurved. The mouth is broad and terminal. The musculature of M. hasselquisti is virtually identical with that of M. lacerda (text-fig. 12). The anterior half of the posterior intermandibularis is, however, not quite so broad. It is separated from the ventral margin of the articular and dentary by a distinct connective tissue band through which the underlying anterior inter- mandibularis can be seen. OSTEOGLOSSOMORPH HYOID MUSCULATURE Fic. 12. Mormyrus hasselquisti: ventral hyoid muscles. A. Right half of posterior intermandibularis removed. B. Right half of the anterior intermandibularis removed. 30 P. H. GREENWOOD Cyphomyrus discorhynchus (Peters) (Text-fig. 2E) The snout in this species is short, broad and strongly decurved, with the small mouth situated subterminally and its opening directed somewhat ventrally. The genus Cyphomyrus, once part of the large and probably artificial assemblage of species in the genus Marcusenius, was separated out by Myers (1960). There are several small but none the less characteristic features in the musculature of C. discorhynchus. The posterior intermandibularis (text-fig. 13) is well developed, but anteriorly it does not extend to the lateral margins of the lower jaw. That section of the muscle originating on the interoperculum has its fibres more closely packed than are those in the larger section originating from an aponeurosis over the hyohyoideus muscle. The two parts of the intermandibularis can be separated easily by gentle traction. TIhy AIM eS MC PIM Hhy Fic. 13. Cyphomyrus discorhynchus : oblique ventro-lateral view of the hyoid musculature, after removal of the skin. OSTEOGLOSSOMORPH HYOID MUSCULATURE 31 The anterior intermandibularis (text-fig. 13) is a short muscle showing some individual variability in length. In only one of the three fishes examined does this muscle extend anteriorly to the level of the posterior intermandibularis insertion ; in one fish it extends posteriorly to the interoperculum (to which it is attached), in another it does not quite reach that point, whilst in the third fish, the muscle is attached to the interoperculum on one side but does not contact that bone on the other side. In all three specimens a slip from the posterior margin ot the muscle passes upwards on each side to insert on the anterior face of the ceratohyal. The major part of the sternohyoideus originates, aponeurotically, from the ventral body musculature; only a small part of the muscle originates from the cleithrum. The ventral gill arch musculature is unusual in having, in addition to a rectus communis, a short but discrete muscle extending between the ventro-lateral ends of the second and third ceratobranchials. Marcusenius cyprinoides (Linn.). (Text-fig. 2F) This species is usually classified in the genus Gnathonemus, but Taverne’s (1968) recent investigations have shown that it and other short-snouted species are more properly included in the genus Marcusenius Gill as revised by Taverne (of which M. cyprinoides is the type species). The snout of M. cyprinoides is relatively short and broad, and is not strongly decurved. The small mouth is terminal in position but has its opening directed somewhat dorsally. There is a noticeable submental protruberence into which the lower lip passes insensibly. The posterior intermandibularis has the typical mormyrid origin from the inter- operculum and from above the hyohyoideus. It inserts, however, entirely onto the enlarged submental cartilage; that is, it has no direct connection with the dentary. The anterior intermandibularis, in contrast with that of the other species described above, is greatly reduced in size. Its length is only slightly greater than that of the eye, and it is confined to the articular region of the lower jaw. It inserts onto the articular of each side just before the articular-quadrate joint. The medial section of this muscle is tendinous. The paired interhyoideus muscles, and the hypertrophied hyohyoideus are typical. The sternohyoideus originates mainly from the dorsal surface of the horizontal limb of the cleithrum, but a small ventral part arises, aponeurotically, from the hypaxial body musculature. As in other mormyrid genera, the anterior part of the sterno- hyoid is closely associated with the ventral region ot the first two gill arches. The ventral processes from the second hypobranchials are completely embedded in the sternohyoideus. The ventral gill arch muscles are typical (see under M. kannume) except that there is, apparently, no short, triangular muscle connecting the fourth and fifth cerato- branchials. 32 P. H. GREENWOOD Marcusenius victoriae (Worthington) (Text-fig. 2G) This species closely resembles M. cyprinoides (and like that species was formerly included in the genus Gnathonemus). The musculature too is virtually identical with that of M. cyprinoides, although the anterior intermandibularis (text-fig. 14) is a little further reduced in size, and the muscle connecting the fourth and fifth ceratobranchials is present. Gnathonemus longibarbis (Hilgendorf) (Text-fig. 2H) The snout is moderately short (especially as compared with other Gnathonemus species; see Taverne [1968]), the mouth small and terminal, and there is a long tubular submental barbel. The posterior intermandibularis is an extensive muscle, originating, as is usual, over the hyohyoideus and from the interopercula. It has strong connective tissue attachments (in which some muscle fibres occur) to the ventral margin of the I Smm Fic. 14. Marcusenius victoriae: ventral hyoid musculature. The right half of the posterior intermandibularis muscle removed to show overlying muscles. OSTEOGLOSSOMORPH HYOID MUSCULATURE 33 articular, dentary and preoperculum. BULLETIN OF 1 MUSEUM (NATURAL HISTORY) ee eee Vol. 22 No. 2 Vise en - LONDON : 1971 ergy THE CLUPEOID FISHES DESCRIBED BY FRANCIS DAY BY PURNESH KUMAR TALWAR AND PETER JAMES PALMER WHITEHEAD Pp. 57-85, 2 Plates, 3 Text-figures BULLETIN OF : THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 22 No. 2 LONDON : 1971 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, 1s 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. 22, No. 2 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). @©Trustees of the British Museum (Natural History), 1971 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 31 December 197% Price £1.20 THE CLUPEOID FISHES DESCRIBED BY FRANCIS DAY By P. K. TALWAR & P. J. P. WHITEHEAD ABSTRACT Francis Day described six new clupeoid fishes: Spratelloides malabaricus, Clupea sindensis, Clupea variegata, Chatoessus modestus, Pellona sladeni and Engraulis auratus. For the first of these the new genus Dayella is proposed. Except for the last, all are considered valid. Lectotypes have been chosen from among Day’s figured specimens now in the Zoological Survey in Calcutta, except in the case of E. auratus for which a British Museum specimen has been chosen. The history of Day’s fish collections is briefly outlined; the Calcutta specimens are considered to be the most important, followed by those in Sydney, Vienna, Leiden, Berlin, Florence and Chicago. Apart from some small collections prior to 1870, the British Museum received only the remainders (in 1889). INTRODUCTION Francis Day (1829-1889) listed 55 clupeoid species in his Fishes of India (1875-8), of which 47 were illustrated, and 46 species are here recognized as valid (Table 1), He described six new species of clupeoid fishes, of which all but one are valid. Day’s descriptions and figures are generally good but many diagnostic features essential to modern clupeoid systematics are omitted, Unfortunately, Day did not specify which specimens were used in his original descriptions and a major problem has been to decide in which institution his types are deposited. The full history of Day’s collections is complex and will be described elsewhere (Whitehead & Talwar, in preparation) but a brief resumé can be given here. Day’s first ichthyological work, the Fishes of Malabar (1865) resulted from his stay in Cochin (1859-64) and from this time stemmed small collections sent to Albert Giinther at the British Museum. Day subsequently investigated the fisheries in almost every large river and along most of the coast of India and Burma, making large collections and finally teturning to England in 1874 to work on his specimens and write his monumental Fishes of India (1875-1888). Unfortunately, a series of bitter quarrels developed between Day and Giinther, with the result that Day donated or sold much of his collection to other museums, the British Museum once again receiving material only in the year of Day’s death. The following is a sum- mary of the distribution of Day’s specimens: 1864-1870 British Museum (r15 lots, c. 400 specimens. Day types specified in letters but not in Register) 1865 East India Museum, London (7 species, including Engraulis auratus) 60 PK. FALWAR & PP. i. Po WHITEHEAD 1875-1879 Rijksmuseum van Natuurlijke Historie, Leiden (11 lots, c. 500 specimens. Day types claimed in Register) 1876-7 Indian Museum, Calcutta (figured specimens, now in Zoological Survey of India) 1874-1882 Zoologisches Museum, Berlin (many lots, 296 specimens. Day types claimed in Register) 1881-1884 Florence (three lots, 333 specimens, 3 types claimed in Register) 1883 Australian Museum, Sydney (Day collection from International Fisheries Exhibition, London. Day, Bleeker and Blyth types claimed by Whitley, 1958) 1886 Naturhistorisches Museum, Vienna (1000 specimens, 815 species— Day and Bleeker types claimed in Annual Report) 1889 British Museum (c. 5,000 specimens. Day types subsequently recognized) ; also Leningrad (see p. 85) 1899 Field Museum of Natural History, Chicago (452 Day specimens from British Museum sent in exchange by Boulenger) The Calcutta specimens include those used by Day in illustrating the Fishes of India (specified as such in Registration Book) and these are being listed by Talwar & Chakrapany (in press). We have concluded that, unless a valid lectotype has been chosen already, the figured Calcutta specimens are the most suitable. In his final letter of reconciliation to Giinther, Day specified that his type collection of Indian fishes went to Calcutta, his No. 2 to Sydney, No. 3 to Vienna, while Florence, Berlin and Leiden had large numbers of specimens. Thus the British Museum received his remainders, except for types in the pre-1870 lots. This order should be followed in making a lectotype selection. In the descriptions given here, measurements follow those of previous clupeoid studies (e.g. Whitehead, Boeseman & Wheeler, 1966). Synonomies include references based on Day material or those relevant to the discussion of Day’s species. All further synonyms are given by Whitehead (in press). The following abbreviations are used: AMS Australian Museum, Sydney BMNH British Museum (Natural History), London FMNH Field Museum of Natural History, Chicago MNHN Muséum National d’Histoire Naturelle, Paris NHMV _ Naturhistorisches Museum, Vienna RMNH Rijksmuseum van Natuurlijke Historie, Leiden ZMB Zoologisches Museum, Berlin ZSI Zoological Survey of India, Calcutta ACKNOWLEDGEMENTS For information on the Day collections sent to Leiden, Berlin, Vienna, Florence, Sydney and Chicago, and for the loan of clupeoid specimens from Leiden, Paris and Berlin, we are indebted to Dr M. Boeseman of the Rijksmuseum van Natuurlijke Historie, Dr C. Karrer of the Zoologisches Museum, Dr P. Kahsbauer of the Natur- historisches Museum, Prof. Leo Pardi of the Museo Zoologico dell’Universita, THE CLUPEOID FISHES DESCRIBED BY FRANCIS DAY 61 Dr J. R. Paxton of the Australian Museum, Dr L. P. Woods of the Field Museum of Natural History in Chicago, and Dr M.-L. Bauchot of the Muséum National d'Histoire Naturelle in Paris. The senior author also wishes to thank the Director, Dr A. P. Kapur, and the Superintending Zoologist, Dr A. G. K. Menon, of the Zoological Survey of India, for providing necessary facilities. Family CLUPEIDAE (Subfamily CLUPEINAE) I. Clupea sindensis Day, 1878 = Sardinella sindensis (Day, 1878) (Plate ra) Clupea sindensis Day, 1878, The Fishes of India: 638, pl. 163 (2) (Seychelles, Sind, Bombay; figure (life-size) on Karachi specimen, shown 95:5 mm S.L.); Idem, 1889, Fauna British India, Fishes, 1 : 374 (repeat). Sardinella sindensis: Chan, 1965, Jap. J. Ichthyol., 13 (1-3) : 11, fig. 21 (key, 44 specimens ex Philippines); Whitehead (in press), Symp. Indian Ocean Adj. Seas. Mar. biol. Ass. India (key, synopsis, fig.). MATERIAL. a. ZSI.2630, a fish 95-5 mm S.L., ex Karachi (stated to be basis for Day’s figure— LECTOTYPE) b. ZSI.2614, a fish 90-0 mm S.L., ex Karachi (PARALECTOTYPE) c. AMS.B7642, a fish 118-5 mm S.L. (140-5 mm tot. 1.), ex Bombay (claimed as type by Whitley, 1958—? PARALECTOTYPE) NHMYV (no specimens) RMNH (no specimens) ZMB (no specimens) BMNH.1889.2.1.1919-24, five fishes 77-5-99°5 mm S.L., ex Sind (registered as Clupea venenosa; one specimen, a skeleton) ga rho a Whitley (1958) gave wholesale endorsement to the type designations made in the published list of Day material bought by the Australian Museum in 1883 (Anon., 1885, 1886). This material, which had been shown at the International Fisheries Exhibition in London in 1883, had already been catalogued (Day, 1883) and, al- though some specimens were marked as the types of Bleeker and Blyth species, none was indicated as a Day type. Day appears to have described the present species on more than one specimen and while the Sydney specimen may be part of the syntypical series, we feel it preferable to designate as lectotype the one speci- men that definitely contributed to the original description, vzz. the Calcutta figured specimen, particularly since Day himself drew the figure. Day (1878, 1889) tentatively included Meletta venenosa Valenciennes in his synonymy, hence the inclusion of the Seychelles in his distribution of the species. The Valenciennes species is Herklotsichthys punctatus (Rippell) (Whitehead, 1967 : 62 15 Jee ANINIBMIINGRS Go 12) odes AVIS CID ODL NID) 35). The British Museum specimens were not relabelled ‘sindensis’ until examined by Regan (1917a); it is unlikely, therefore, that they were used by Day in his description. Description. Based on the LECTOTYPE, a fish 95:5 mm S.L., ex Karachi, ZSI1.2630 (basis for pl. 163 (2) of Fishes of India) (in parenthesis are given measure- ments for the Calcutta PARALECTOTYPE, ZSI.2614). Br. St. 6, D iii 13 (14), Piz4, Vi7, Ali 16, g.r. 36 + 65 (37 + 63), scutes 18 + 14, scales in lateral series 42 (43), transverse II, pre-dorsal 15 (?). In percentages of standard length: body depth 25-7 (23:9), head length 26-7 (22:5); snout length 6-5 (6-9), eye diameter 6-3 (6-9), upper jaw length 9-9 (10-6), lower jaw length 12-0 (10-6); pectoral fin length 16-2 (15-0), pelvic fin length 9-9 (8-1), length of anal fin base 15-7 (14:5); pre-dorsal distance 46-I (43-3), pre-pelvic distance 48-2 (48-9), pre-anal distance 76-4 (77°8). Body fairly compressed, its width about 2} times in its depth, the latter more or less equal to head length; belly keeled, scutes partly concealed by scales on either side. Snout equal to or a little greater than eye diameter. Upper jaw reaching to vertical from anterior third of eye; two supra-maxillae, the Ist (anterior) about 5 times as long as deep, the 2nd (posterior) with upper and lower expanded parts similar in shape and size, the whole almost circular; no hypomaxilla; expanded portion of maxilla with faint longitudinal ridges, lower edge of maxilla with fine denticulations posteriorly. Lower jaw profile rising steeply, its depth half its length. Pre-maxillae and vomer edentulous, but fine teeth on either side of dentary sym- physis, a median line of conical teeth on tongue and fine teeth on palatines and ecto- and endo-pterygoids. Gillrakers fine and slender, close-set, the longest about 4 of eye diameter and equal to length of corresponding gill filaments. Pseudobranch present, exposed, with a dozen filaments, its length about equal to eye diameter. Cleithral lobe and bilobed dermal outgrowths from cleithrum well developed. Operculum about twice as high as wide, its lower margin almost horizontal; sub-operculum rectangular. Opercular series and cheek covered by adipose tissue overlying ramifications of sensory canal system. Fronto-parietal region with cuneiform area bearing 8 (9) longitudinal striae; supra-orbitals with about four longitudinal striae. Dorsal fin origin much nearer to snout than to caudal base; lower part of fin invested in scaly sheath. Pectoral fin tips failing to reach pelvic base by more than one eye diameter, failing to reach vertical from dorsal origin by 14 eye diameters; no axillary scale but scales above first ray truncated to leave shallow depression for reception of fin. Pelvic fin base below middle of dorsal base, nearer to pectoral base than to anal origin; axillary scale present, almost length of fin. Anal fin slightly nearer to caudal base than to pelvic base; last two rays enlarged, about twice length of antepenultimate ray. Scales: unexposed portion of scale with one major and four (anterior scales) to six (posterior) minor vertical striae, the former continuous, the latter interrupted at centre of scale; exposed portion of scales with eroded and slightly crenellated posterior border, faint horizontal ridging and small perforations. Pre-dorsal THE CLUPEOID FISHES DESCRIBED BY FRANCIS DAY 63 medial ridge covered by overlapping scale rows on either side. Alar scales absent (probably lost; present in some specimens of BMNH.1889.2.1.1919-24). Colour: in alcohol, upper } of body slate-coloured, remainder of flanks silvery- gold. Fins hyaline, but dark spot at base of anterior dorsal rays. Inside face of operculum somewhat dusky. Note. Sardinella sindensis, together with S. gibbosa (Bleeker), can be separated from other species of Sardinella by its slightly higher post-pelvic scute count (15-16, rarely 14 or 17-18; cf. 12-14, rarely 11 or 15—see key in Whitehead, in press). This slight distinction held true in 44 and 159 specimens (respectively) examined by Chan (1965), and also in British Museum material, and it is unfortunate that both lectotype and paralectotype of S. sindensis have the lower count of 14. One out of five other Day specimens (BMNH.1889.2.1.1919-24) has 14 post-pelvic scutes. If scute number is diagnostic, then S. sindensis can be separated from S. gzbbosa by its slightly higher range for gillraker numbers (58-72 at 69-122 mm S.L.; cf. 43-63 at go-150 mm S.L.—figures from Whitehead, in press). Specimens with only 14 post-pelvic scutes can be distinguished from S. albella (Valenciennes) and S. fimbriata (Valenciennes) by their more slender body (24:5-27:8% of S.L. (Chan, 1965); cf. 32-35 and 28-34°%, respectively—Whitehead, in press). Sardinella brachysoma Bleeker and S. zunasi (Bleeker) are also slightly deeper species which are further distinguished by the numerous overlapping or continuous vertical striae on the posterior scales. (Subfamily PELLONULINAE) DAYELLA gen. nov. TYPE SPECIES: Spratelloides malabaricus Day. Diacnosis: clupeid fishes with 5-6 branchiostegal rays, a short anal fin (less than 20 rays), small unkeeled pre-pelvic scutes but no post-pelvic scutes, eight pelvic rays, a single (posterior) supra-maxilla, gillrakers present on posterior face of 3rd epibranchial, and posterior frontal fontanelles occluded in adults. A single species known. 2. Spratelloides malabaricus Day, 1873 = Dayella malabarica (Day, 1873) Spratelloides malabaricus Day, 1873, Proc. zool. Soc. Lond.: 240 (‘Sea, ascending rivers in Malabar, and attaining 3 inches in length’); Idem, 1878, Fishes of India: 648, pl. 161 (5) (‘Western Coasts of India, in rivers and estuaries’; up to 3 inches, figure (? life size) 55:3 mm S.L.); Idem, 1889, Fauna British India, Fishes, 1 : 400, fig. 124 (repeat). MATERIAL. a. ZSI.2246, a fish 51-0 mm S.L., ex Malabar (stated to be basis for Day’s figure although 4:3 mm shorter—LECTOTY PE) b. RMNH.2726, a fish 58 mm S.L., ex Malabar—PARALECTOTYPE 64 P. K. TAEWARN& PP. J.P’ WHITEBEAD c. BMNH.1889.2.1.2048, a fish 47-3 mm S.L., ex Malabar, stained with alizarin —PARALECTOTYPE d. Zool. Inst. Leningrad, 8220, a fish 48°3 mm S.L., ex Canara —PARALEC- TOTYPE The following specimens are Ehirava fluviatilis. e. AMS.B8288, a fish 44:0 mm S.L. (52:0 mm tot. 1.), ev Malabar (claimed as type by Whitley, 1958) f. NMV (no specimens) g. ZMB.10413, three fishes 28-9-35-4 mm S.L., ex Malabar. h. BMNH.1889.2.1.2051, two fishes 40-0-49:I mm S.L., ex Malabar (removed from jar containing BMNH. paralectotype) i. BMNH.1889.2.1.2050, one fish 53-0 mm S.L., ex Malabar (also removed from BMNH paralectotype jar) j. BMNH.1889.2.1.2052-5, four fishes 46:5-56:2 mm S.L., ex Canara (one fish 50-0 mm stained with alizarin; three fishes donated to the Musée Royale de l'Afrique Centrale, Turvuren) . FMNH.2379, a fish 49:0 mm S.L., ex Canara (numbered 240 and donated by G.A. Boulenger from BMNH collection) an The two species included in the Day material are superficially very similar and hitherto Deraniyagala’s fluviatilis has been considered a synonym of Day’s mala- baricus (e.g. in Whitehead, 1963). It was not until the single true specimen of Dayella malabarica in the British Museum was stained with alizarin and re-examined that the Day material was found to be mixed. Although specimens of E. fluviatilis predominate, Day’s original description seems to have been based on D. malabarica since Day states that the ‘dorsal commences slightly before the origin of the ventral’. In D. malabarica the dorsal origin is well before the pelvic base, the latter lying below the first branched dorsal ray. In E. fluviatilis the pelvic base is before, below or only just behind the first wnbranched dorsalray. The statement is repeated in Day’s second description and in his figure (Day, 1878 : pl. 161) the dorsal origin is clearly well before the pelvic base. The specimen in Calcutta is slightly smaller than Day’s figure but is presumed to have been the model and is here chosen as lectotype of Spratelloides malabaricus. The holotype of Ehivava fluviatilis Deraniyagala is in London (BMNH.1929.7.1.1) ; the specimen in the Zoological Survey of India (ZSI.F11043/1), claimed as a paratype by Menon & Yazdani (1963), is from Moratua (Western Province of Ceylon), a locality not mentioned in the original description. DESCRIPTION. Based on the LECTOTYPE, a fish 51-0 mm S.L., ex Malabar, ZSI1.2246 (basis for pl. 161 (5) ot Fishes of India). Figures for the Leiden and British Museum specimens (49:5 and 47:3 mm S.L.—lots b and c above) are given in parenthesis. Bye SUG (6) (Gy 5) ID) tht aese (Gere, sea), 12) at Fe (Ger), aes), Wah, (Ge, a) eX vob) aeY (02105), (C nr. (n.r., 10 + 9g), g.r. II + 27 (10 + 24, Io + 27), scutes ? o (4, 1), scales in lateral series 38 (n.r., 36), transverse g (n.r., n.r.), vertebrae 40 (BMNH alizarin specimen). THE CLUPEOID FISHES DESCRIBED BY FRANCIS DAY Fic. 1. Pre-pelvic scutes in three Indo-Pacific pellonulines. a. Dayella malabarica, 473mm S.L., BMNH.1889.2.1.2048. b. Dayella malabarica, 49-5 mmS.L., RMNH.8585. c. Gilchristella aestuarius, 52:3 mm S.L., BMNH.1915.7.6.3. d. Ehivava fluviatilis, 491mm S.L., BMNH.1889.2.1.2051. 65 66 PO oK. TALWAR & PB. J: PS WHITEREAD In percentages of standard length: body depth 22-5 (20-0, 20-0), head length 27°7 (24-0, 26-0); snout length 8-3 (7-5, 7-4), eye diameter 8-3 (8-0, 7-8), post-orbital distance 9-9 (8-5, 8-7), length of upper jaw 9-9 (8-7, 9-7), length of lower jaw 14:7 (12-3, 13-1); pectoral fin length 16-7 (17-1, 15-8), pelvic fin length 13-7 (13-3, 14:5), length of anal base 18-6 (15-3, 14°5); pre-dorsal distance 49-0 (47:9, 48:5), pre-pelvic distance 52-9 (50-4, 49°5), pre-anal distance 78-4 (77:3, 75:0). Body fairly compressed, its width almost 3 times in its depth, the latter a little less than head length; belly rounded, fully scaled but the scales underlain by 4 (Leiden) or 1 (BMNH) plate-like scutes bearing rudimentary lateral arms (fig. 1a, b), the scutes not reaching back to the main pelvic scute (which has normal lateral arms). Snout equal to or a little shorter than eye diameter. Jaws unequal, the lower projecting slightly. Upper jaw reaching to vertical from anterior eye border or anterior pupil margin, ventral expanded portion of maxilla beginning abruptly and not tapered smoothly into slender anterior limb of bone, the entire edge of the expanded portion finely denticulated (fig. 2); a single (posterior) supra-maxilla, the lower part of the expanded portion deeper and longer than the upper part (Harengula shape), its depth about + eye diameter. Lower jaw rising fairly steeply in the first third of its length; 6-7 small conical teeth on either side of symphysis. A single row of small conical teeth on pre-maxillae, separated by a median diastema. Fine teeth on tongue, scattered on antero-median process of palatine and along outer edge of that bone. Fic, 2. Dayella malabarica, upper and lower jaws, alizarin stained specimen, 47-3 mm S.L., BMNH.1889.2.1.2048. THE CLUPEOID FISHES DESCRIBED BY FRANCIS DAY 67 Gillrakers fine, slender, 24 times in eye diameter and a little longer than corres- ponding gill filaments; about 7 short, stumpy gillrakers on posterior face of 3rd eipibranchial. Pseudobranch present, exposed, about } eye diameter, with about a dozen filaments. Cleithral lobe present at lower angle of gill opening, fairly well developed. Operculum about 14 times as deep as broad, its posterior margin with a deep indentation, its lower margin rising at an angle of about 15° to the horizontal; suboperculum rectangular except for rounding of postero-ventral angle. Cutaneous sensory canals branching over entire opercular series and cheek. Fronto- parietal rea smooth, posterior frontal fontanelles retained and minute, I-o mm in length (oa-5 mm in BMNH specimen—fig. 3c; virtually occluded in Leiden specimen). Dorsal fin origin nearer to snout tip than to caudal base by ? eye diameter and a little in advance of vertical from pelvic base. Pectoral fin tips failing to reach pelvic base by 1 (14) eye diameters; no axillary scale. Pelvic fin base below first branched dorsal ray and about equidistant between pectoral base and anal origin; axillary scale present, about } of fin length (Leiden specimens). Anal fin origin a little nearer to caudal base than to pelvic base; last two rays normal, not separated from rest of fin. Scales: deeper than broad, with distinct anterior ‘shoulders’; unexposed portion with a single continuous striation, preceded by 3-4 short radiating striae (absent on anterior scales, joined to form a loop on some posterior scales) ; exposed portion of scale without striae, its posterior margin slightly eroded and produced medially. Colour: in alcohol, uniform light brown with a faint silvery midlateral stripe not quite as broad as eye; fins hyaline. Note. The new genus Dayella is a member of the pellonuline complex that comprises the monotypic Indo-Pacific genera Ehivava, Gilchristella, Sauvagella and Spratellomorpha. These five genera are distinguished from all other Indo- Pacific pellonulines by their lack of post-pelvic scutes; in addition, pre-pelvic scutes are either absent or extremely poorly developed. The separation of the five species at generic level is arguable but the features that distinguish them are non-meristic and appear to be of some significance in other groups of clupeids. Dayella can be identified from the following key. INDO-PACIFIC PELLONULINAE THAT LACK POST-PELVIC SCUTES I. Anal fin entire, last two rays not separate A. Gillrakers present on posterior face of 3rd epibranchial I. Posterior frontal fontanelles minute (4 eye diameter) or completely occluded in adults (fig. 3c); pre-pelvic scutes 1-4, rudimentary, lateral arms barely developed (fig. ra, b); pelvic base well behind dorsal origin; gillrakers 24-27 Dayella malabarica (Day) 2. Posterior frontal fontanelles larger (4-4 eye diameter), retained in adults (fig. 3a, b); pelvic base before dorsal origin a. No pre-pelvic scutes; gillrakers Ig (at 40 mm S.L.) Sauvagella madagascariensis (Sauvage) 68 PK. TALWAR & P. J. P) WHITEHEAD b. Up to 9 pre-pelvic scutes, with thin lateral arms i. Anterior arm of supra-occipital broadening anteriorly (fig. 3b); gillrakers 39-45 Gilchristella aestuarius (Gilchrist) ii. Anterior arm of supra-occipital very slender anteriorly (fig. 3a); gillrakers 40-60 Gilchristella sp. (see below) B. Gillrakers absent on posterior face of 3rd epibranchial; posterior frontal fontanelles large, anterior arm of supra-occipital broadening anteriorly (fig. 3d); pre-pelvic scutes present, poorly developed but with distinct lateral arms (fig. 1d); pees base below or before dorsal origin; gillrakers 26-30 . : : F Ehirava fluviatilis Deraniyagala II. Anal fin split, the Hast two rays separate from rest of fin; gillrakers present on posterior face of 3rd epibranchial; posterior frontal fontanelles large, probably retained in adults, similar to those of Ehirava; gillrakers 26-31 Spratellomorpha bianalis (Bertin) The four previously described Indo-Pacific genera were formerly placed in the round herrings or Dussumieriidae (Whitehead, 1963). The subsequent discovery of partially scuted (Laeviscutella, Sierrathrissa) or non-scuted (Congothrissa) forms amongst the otherwise fully scuted West African Pellonulinae suggested that both Indo-Pacific and African genera were members of a clupeid group that showed progressive stages in scute loss. This appeared to be correlated with trends towards reduction in supra-maxillae and numbers of branchiostegal rays, together with a retention of the posterior frontal fontanelles by adults (Poll, Whitehead & Hopson, 1965). The non-scuted genera Spratelloides (Indo-Pacific) and Jenkinsia (Western Atlantic) may eventually join this group, although their very characteristic W-shaped pelvic scute seems to link them with the ‘true’ round herrings Dussuwmieria and Etrumeus—whose high and presumably primitive branchiostegal count implies yet another route to scute loss (or perhaps the primitive absence of scutes, at least in this branch of the clupeids). For the present, the five poorly or non-scuted genera shown in the key above are placed in the tribe Ehiravini of the subfamily Pellonulinae. In Dayella, the scutes are more rudimentary than in any other clupeid genus. They are thin, difficult to find in unstained material and those with small lateral arms could easily be mistaken for scales. Their resemblance to scales is increased by the relatively large size of the expanded portion of the scute when compared with those of other genera (fig. ta-d). The variation in shape of these scutes implies that these are structures on the way to being lost and not an early stage in the evolution of scutes. Dayella appears to be most closely allied to Gilchristella, Sauvagella and Ehirava, differing from them chiefly in its occluded posterior frontal fontanelles and its less advanced pelvic base. The absence of gillrakers on the posterior face of the 3rd eipibranchial seems to hold some significance elsewhere in the Clupeidae, but this THE CLUPEOID FISHES DESCRIBED BY FRANCIS DAY 69 may not mean that Ehivava is necessarily remote from the other genera of the group. The status of Sauvagella madagascariensis is uncertain. Re-examination of a syntype of this species (40 mm S.L., MNHN.3794) has confirmed that even the rudimentary scutes of Dayella are not present. This fish has only 109 gillrakers and the posterior frontal fontanelles are large, suggesting that it is a juvenile; gillrakers may increase with size. In a redescription of the species (Whitehead, 1963), eleven South African specimens were included, from the Buffalo river, Cape Province (BMNH.1878.1.22.25 and 33-43) and the stated gillraker count of 40-56 referred to these specimens only. Careful removal of the belly scales now shows that these fishes have up to 7 thin and barely apparent scutes with fairly long lateral arms. Thus, they are clearly distinct from Sauvagella (as far as can be judged from the very small types) and for the moment they appear to be an undescribed species ot Gilchristella. From G. aestuarius they differ, however, in having a very slender P Fic. 3. Posterior frontal fontanelles in four Indo-Pacific pellonulines, dorsal view showing frontals (fr.), parietals (p.) and anterior arm of supra-occipital (s. occ.), the fontanelles black. a. Gilchristella sp., 52-6 mm S.L., anomalous Buffalo River specimen, BMNH.1878.1.22.25. b. Gilchristella aestuarius, 54-5 mm S.L., BMNH.19109.9.12.3. c. Dayella malabarica, 47-3 mm S.L., BMNH.1889.2.1.2048. d. Ehivava fluviatilis, 50:0 mm S.L., BMNH.1889.2.1.2052. 70 TPE Is ABYMEM VINE (4 125 fg 1, \WiISWlsh olen d/N1D) anterior arm of the supra-occipital (fig. 3a) and a higher gillraker count (40-60 cf. 39-45). In general, the members of the Ehirava complex are rather poorly known and would repay further study when more specimens are available. (Subfamily ALOSINAE) 3. Clupea variegata Day, 1869 = Gudusia variegata (Day, 1869) (Plate 1c) Clupea variegata Day, 1869, Proc. zool. Soc. Lond.: 623 (Irrawaddy and its branches; many specimens, to 7 inches); Idem, 1878, Fishes of India: 639, pl. 161 (4) (repeat; figure of fish 152 mm S.L., presumably life-size); Idem, 1889, Fauna British India, Fishes, 1 : 375 (repeat). Gudusia variegata: Regan, 1917, Ann. Mag. nat. Hist., (8) 19 : 308 (on single Day specimen in British Museum); Motwani, Jayaram & Sehgal, 1962, Tvop. Ecol., 3 (1-2) : 17-43 (Brahmaputra at Jogighopa, Goalpara District); Whitehead, 1965, Bull. By. Mus. nat. Hist. (Zool.), 12 (4) : 150, fig. 11 (Day specimen redescribed; Day’s figure reproduced) ; Idem, (in press), Symp. Indian Ocean Adj. Seas. Mar. biol. Ass. India (key, note on synonymy). ? Clupea suhia Chaudhuri, 1912, Rec. Indian Mus., 7 : 436, pl. 38 (1) (river Gandak in Saran, Bihar). ? Gudusia godanahiai Srivastava, 1968, Fishes Easteyn Uttar Pradesh: 6, fig. 4a, b (Gorakhpur, Uttar Pradesh). MATERIAL. a. ZSI.2245, a fish 150:0 mm S.L., ex Irrawaddy river (stated to be basis for Day’s figure—LECTOTY PE) b. ZSI (Duplicate Cat.) 43, a fish 158 mm S.L., ex Bassein river (labelled Clupea burmanica) c. ZSI. (Duplicate Cat.) 168, a fish 78 mm S.L., ev Mandalay, coll. Major E. B. Sladen (labelled Clupea burmanica) d. AMS.B7676, a fish 158-5 mm S.L. (191 mm tot. 1.), ex Bassein (claimed as type by Whitley, 1958. NHMV (no specimens) RMNH.2586, a fish 106-4 mm S.L., ex Bassein. ZMB (no specimens) BMNH.1870.6.14.38, a fish 155 mm S.L., ex Bassein. Bog rh © For reasons given under the previous two species, the figured specimen in Calcutta is chosen as lectotype. The two specimens labelled ‘buymanica’ ate G. variegata but are probably not syntypes since their meristic counts exceed the values given in the original description (but are consistent with the ranges given in the Fishes of India). Day did not publish the name burmanica but may have initially intended to use it for this species. The British Museum specimen, although presented in 1870 and thus the first of this species to be given away by Day, has too low an anal and pectoral count (111 22 and i113; cf. 111 26 and i15) to have figured in the original THE CLUPEOID FISHES DESCRIBED BY FRANCIS DAY 71 description, which in any case seems to have been based on a single specimen (although many were taken). For this reason, the Sydney and Leiden specimens cannot be regarded as paralectotypes. DEscRIPTION. Based on the LECTOTYPE, a fish 150-0 mm S.L., ex Irrawaddy river, ZSI.2245 (basis for pl. 161 (4) of Fishes of India). Br. St. 6, Diii 12, Pi 15, Vi7, A iii 26, g.r. 210 (approx.), scutes Ig + II, scales in lateral series go, transverse 35. In percentages of standard length: body depth 44-3, head length 29:3; snout length 5-5, eye diameter 6-7, length of upper jaw 11-7, length of lower jaw 13-0, operculum height 14-3, its breadth 8-3; pectoral fin length 20-0, pelvic fin length 10-0, length of anal fin base 23-0; pre-dorsal distance 52-0, pre-pelvic distance 54-7, pre-anal distance 74:0. Body strongly compressed, its width 33 times in its depth, the latter 1} times head length; belly keeled, tips of scutes projecting slightly from sheath of scales on either side. Snout shorter than eye diameter; pre-orbital distance (including eye) } of post-orbital distance. Lower jaw included when mouth closed: pre- maxillae rising steeply to form very distinct notch in upper jaw. Maxilla reaching to just beyond vertical from eye centre, expanded portion without longitudinal ridges or striae and smooth along lower edge; two supra-maxillae, the rst (anterior) about 6 times as long as deep and almost equal in length to eye diameter, the 2nd (posterior) with slender anterior shaft and lower part of expanded portion larger than upper. Lower jaw rising rather gently in the first third of its length, its depth about 3 times in its length; no teeth present. No teeth in upper jaw nor within mouth. Gillrakers fine, straight or slightly curved, slightly longer than corresponding filaments; filaments of anterior hemibranch of 1st arch 3-? length of those of posterior hemibranch; many fine gillrakers present on posterior face of 3rd epibranchial. Medio-pharyngobranchial present, about 2 eye diameter, bearing many short gillrakers. Pseudobranch present, attenuated, about 1# times eye diameter, with distinct ventral ridge and a groove below it. Cleithral lobe barely developed, hardly breaking outline of gill opening. Operculum not quite twice as deep as broad, its lower edge rising steeply; sub-operculum crescentic; lower third of an- terior operculum margin not overlapped by pre-operculum, leaving a small triangular area covered only by skin. Cutaneous sensory canals branching through the adipose tissue covering the suborbitals, operculum, sub-operculum and scales behind head. Adipose eye-lid with vertical slit exposing 2 of pupil. Dorsal surface of head covered by fairly thick skin but a pair of cuneiform fronto-parietal areas with about six longitudinal striae left exposed. Dorsal fin origin slightly nearer to snout tip than to base of caudal; a very low scaly sheath along base. Pectoral fin tips failing to reach pelvic base by about 2 eye diameter; axillary scale present, half length of fin. Pelvic fin base below vertical from dorsal origin and a little nearer to pectoral base than to anal origin; axillary scale present, half length of fin. Anal origin equidistant between pelvic and caudal bases; anal base longer than pectoral fins and greater than the distance 72 P. K. TALWAR & P. J. P. WHITEHEAD snout tip to posterior border of pre-operculum. Caudal fin (broken) slightly longer than head length, lower lobe longer than upper. Scales: almost circular, but becoming more elongate on posterior part of body; a single vertical striation continuous across scale, preceded by o (anterior scales) to 3 (posterior scales) short and irregular striae interrupted at centre of scale; exposed border irregular, becoming pectinate in posterior scales. Minute scales covering caudal except for hind border. Colour: in alcohol, back brown, flanks golden; a series of brown spots along upper flank, some expanded vertically, those behind dorsal extending right across back; a dark humeral spot. Note. The genus Gudusia (often misspelt Gadusia in the literature) at present includes two species, G. chapra (Ham. Buch.) and G. variegata. The latter has a deeper body (depth greater than 40% of S.L.), a shorter head (head length less than 28% of S.L.), and more anal finrays (111 22-26; cf. iii 19-22). In addition, G. variegata has a very prominent series of black spots along the flank, whereas G. chapra is usually described as having a dark shoulder spot, sometimes absent, and faint or no spots along the flank (Whitehead, 1965; Srivastava, 1968). Rather few specimens of G. variegata have been described, however, and some of these were misidentifications. Thus, specimens identified as G. variegata from Akyab by Lloyd (1907) include at least one fish (ZSI.1491/1) that is Hilsa kelee, while juveniles reported as G. variegata from the Mandalay fish market by Jenkins (1910)"include specimens (ZSI.1770/1) of a species of Hilsa (Tenualosa). Gudusia variegata is usually considered a Burmese species (Regan, 1917b; Fowler, Ig4I : 635), but both Chandhuri’s Clwpea suhia and Srivastava’s Gudusia godanahiar (Ganges drainage) had very prominent black spots along the flanks which were, indeed, the main reason for distinguishing these nominal species from the sympatric and unspotted G. chapra (not a sexual feature according to Srivastava, 1968). Gudusia variegata has also been reported from the Brahmaputra, by Motwani et alii (1962), presumably because of the strong pattern of spots since G. chapra was also recorded from the same area (no descriptions given, however). If these Indian records truly relate to G. variegata, then some modification must be made to the key since Srivastava’s G. godanahiai were rather slender (depth 33-3-38:7% of S.L.) and thus within the range of G. chapra (31-0-40:0% in 30 specimens—Whitehead, 1965: fig. 13); the head length (27-4-31-7% of S.L.) of Srivastava’s specimens, however, agreed with current definitions of G. variegata, but the low anal count of iii 20 was that of G. chapra. Srivastava distinguished his new species by the presence of 14 pectoral rays (13 in his G. chapra), but Regan (1917b) recorded 13-14 pectoral rays in the British Museum material of G. chapra and there is probably overlap between the two species. For the present, the status of G. variegata and its possible synonyms must remain uncertain until more material has been examined. THE CLUPEOID FISHES DESCRIBED BY FRANCIS DAY 73 (Subfamily DOROSOMATINAE) 4. Chatoessus modestus Day, 1869 = Gonialosa modesta (Day, 1869) Chatoessus modestus Day, 1869, Proc. zool. Soc. Lond.: 622 (Bassein river as high as Een-gay-gyee Lake; many specimens, up to 5} inches); Idem, 1878, Fishes of India: 633, pl. 160 (1) (also Selwein at Moulmein; figure of fish 100-r mm S.L., ? life size); Idem, 1889, Fauna British India, Fishes, 1 : 386 (repeat). Gonialosa modesta: Regan, 1917, Ann. Mag. nat. Hist., (8) 19 : 315 (on Day material in British Museum); Menon & Yazdani, 1963, Rec. 2001. Surv. India, 61 : 98. MATERIAL. a. ZSI.2695, a fish 98-0 mm S.L., ex Bassein river (stated to be basis for Day’s figure—LECTOTY PE) b. ZSI.F8022/1 and 8023/1, two fishes 58-o-r01-0 mm S.L., ex Bassein river (= G. manminna) c. AMS.B7637, a fish 105-0 mm S.L. (127:5 mm tot. 1.), ex Burma (claimed as type by Whitley) NHMYV (no specimens) RMNH.2585, a fish 116-0 mm S.L., ex Moulmein (claimed as type in register) ZMB (no specimens) g. BMNH.1880.2.1.1879, a fish 82-6 mm S.L., ex Burma Menon & Yazdani (1963) erroneously listed the first of the three Zoological Survey fishes as holotype, and the other two as paratypes, but Day did not indicate a holo- type nor did he give an exact length measurement. The first Calcutta specimen, used for Day’s figure, is here designated lectotype; the other two are G. manminna and thus do not agree with the original description, which in any case shows no ranges for meristic and morphometric values, suggesting that only a single fish was measured (although many were caught). For this reason, the Sydney, Leiden and London specimens are not regarded as paralectotypes. DEscRIPTION. Based on the LECTOTYPE, a fish 98-0 mm S.L., ex Bassein river, | ZSI.2695 (basis for pl. 160 (1) of Fishes of India). Br. St. 6, D iii 13, Pir5, Vi7, A iii 25, g.r. 150 (approx.), scutes 17 + 12, scales in lateral series 47, transverse 17. In percentages of standard length: body depth 48-5, head length 27-6; snout length 6-4, eye diameter 7:9, post-orbital distance 13-3, length of upper jaw 7:1, length of lower jaw 9-7; pectoral fin length 23-0, pelvic fin length 10-2, length of anal base 26-8; pre-dorsal distance 52-0, pre-pelvic distance 50-0, pre-anal distance 72°4. Body compressed, its width 4,3, times in its depth, the latter almost twice head length; belly keeled, tips of scutes projecting below scaly sheath; profile of back concave beyond nape, rising abruptly to dorsal origin, belly profile evenly convex. Snout shorter than eye diameter. Mouth sub-terminal, transverse, with snout projecting strongly; pre-maxillae meeting at an angle to form a distinct notch in tho A 74 Pp. K. TALWAR & P. J. P. WHITEHEAD upper jaw. Maxilla slender, slightly expanded and curved downwards distally, reaching to vertical from anterior border of eye; a single narrow supra-maxilla, its length 2-5 mm and depth 0-5 mm. Lower jaw with dentaries meeting at an obtuse angle, the edge of each dentary flared or reflected outwards in front of tips of maxillae. No teeth in jaws. Gillrakers very fine, close-set, shorter than corresponding gill filaments and slightly more than } eye diameter; gill filaments of anterior hemibranch equal to those of posterior hemibranch. Numerous fine and close-set gillrakers on posterior face of 3rd epibranchial. Pseudobranch present, exposed, its length almost one eye diameter; about 20 filaments present. Cleithral lobe present, breaking outline of gill opening but not strongly developed. Operculum 1} times as deep as broad, its lower border rising steeply (about 40°); suboperculum long and narrow, its posterior border rounded. Cutaneous sensory canals branching over cheek, oper- cular series and nape. Adipose eye-lid with vertical slit exposing }4 of eye. Dorsal surface of head with a pair of cuneiform fronto-parietal areas bearing 6 longi- tudinal striae, the two areas linked posteriorly by a transverse bony ridge. Dorsal fin origin slightly nearer to snout than to base of caudal fin; final finray not elongated. Pectoral fin tips reaching to beyond pelvic base; axillary scale present, } length of fin. Pelvic fin base in front of dorsal origin and nearer to pectoral base than to anal origin; axillary scale present, § length of fin. Anal origin nearer to pelvic than to caudal base. Lower lobe of caudal larger than upper. Scales: unexposed portion with one major vertical striation and 3 (anterior scales) to 4-5 (posterior) minor vertical striae interrupted at scale centre. Exposed portion without striae, posterior margin of scale not eroded, perforated or fimbriated. Colour: in alcohol, upper parts of body light brown, lower parts silvery; a dark humeral spot present. Fins hyaline. Note. The two species of Gonialosa, G. modesta and G. manminna (Ham. Buch.), have been separated on body depth (40-50% of S.L. in modesta ; 30-39 % in manminna) and number of scales along the flank (45-47 and 55-65 respectively—Whitehead, 1962; in press). Gudusia manminna (from the Ganges and Brahmaputra and their tributaries) is fairly well represented in collections and the literature, but G. modesta (recorded only from Burma) is not. Larger collections may show modesta to be merely a subspecies of G. manminna. (Subfamily PRISTIGASTERINAE) 5. Pellona sladeni Day, 1869 = Ilisha sladeni (Day, 1869) Pellona sladeni Day, 1869, Proc. zool. Soc. Lond.: 623 (Irrawaddy at Mandalay; specimens up to 7 inches); Idem, 1878, Fishes of India: 645, pl. 164 (1) (repeat; ‘A single example obtained, 7 inches in length’; figure shows fish of 146 mm S.L.); Idem, 1880, Fauna British India, Fishes, 1 : 383 (repeat). Ilisha sladeni: Norman, 1923, Ann. Mag. nat. Hist., (9), 11 : 6 (Day specimen and two others described). THE CLUPEOID FISHES DESCRIBED BY FRANCIS DAY 75 MATERIAL. a. ZSI.2672, a fish 210 mm S.L. (ca. Io inches tot. 1.), ex Irrawaddy (stated to be basis for Day’s figure—LECTOTY PE) b. ZSI (Duplicate Cat.) 298, a fish 189 mm S.L. (ca. 8} inches tot. 1.), ex Mandalay (? PARALECTOTY PE) AMS (no specimens) NHMV (no specimens) RMNH (no specimens) ZMB (no specimens) BMNH.1870.6.14.36, a fish 209 mm S.L. (almost ro inches tot. 1.), ex Mandalay. mM ro Ae As in previous cases, Day clearly collected more than one specimen, but he gave no ranges for meristic or morphometric values in his original description. In the Fishes of India, however, he stated ‘single example obtained’; he may perhaps have been referring to the fish that he himself had drawn for the Fishes of India, since it was now eight years since he had given his other specimen to the British Museum. All three extant specimens are larger than either the figure or the maximum length stated (7 inches). They also differ in having more pre-pelvic scutes (23 or 24; cf. 20) and more pectoral rays (14 or 15; cf. 11). Day altered his scute count to 23 in the Fishes of India (but not the pectoral count) and it must be presumed that the earlier counts were errors. The larger Calcutta fish, the figured specimen, is chosen as lectotype, on the assumption either that Day lost the original (smaller) specimen or that the maximum length of 7 inches was also an error. DEscRIPTION. Based on the LECTOTYPE, a fish 210 mm S.L., ex Irrawaddy river, ZSI.2672 (basis for pl. 164 (x) of Fishes of India) (measurements of ZSI para- lectotype given in parenthesis). Br. St. 6, D iii 10, Pi 13, Vi6, A iii 4I, g.1. Io + I + 21, scutes 23 + 10, scales in lateral series 48, transverse 10. In percentages of standard length: body depth 21-9 (25-4), head length, 24:5 (27:5); snout length 5-5 (5-8), eye diameter 6-0 (6-6), length of upper jaw 11-0 (11-9), length of lower jaw 11-4 (13-0); pectoral fin length 19-8 (21-7), pelvic fin length 4°8 (6-6), length of anal fin base 30-0 (29-4) ; pre-dorsal distance 59°I (59:3), pre-pelvic distance 39-I (44-0), pre-anal distance 65:2 (67-2). Body strongly compressed, its width 4 times in its depth, belly strongly keeled, the tips of the scutes projecting below scaly sheath, especially behind pelvic fin base; anterior four scutes on isthmus. Dorsal profile slightly concave before nape, ventral profile evenly convex, the two almost parallel between pectoral base and dorsal origin. Snout a little shorter than eye diameter. Lower jaw strongly projecting, about 4 eye diameter beyond snout when mouth closed. Maxilla teaching to vertical from anterior pupil border, fine denticulations along its lower edge; no hypo-maxilla; two supra-maxillae, the rst (anterior) 5 times longer than deep and about 1} eye diameter, the 2nd (posterior) with lower lobe of expanded portion much larger than upper. A single series of fine teeth on pre-maxillae, with median diastema, small conical teeth present on either side of dentary symphysis. No teeth on vomer but fine teeth on tongue, palatines and ecto- and endo-pterygoids. 76 Pr Ke DAL WAR Te Ps Po WHITE EE AD Gillrakers fairly slender, the longest + eye diameter and 14 times length of cor- responding gill filaments; no gillrakers on posterior face of 3rd epibranchial. Pseudo- branch present, exposed, its length } eye diameter; ventral margin not ridged. No cleithral lobe. Operculum elongated posteriorly, its ventral margin equal to its height and rising at an angle of about 20°; sub-operculum eliptical, long and narrow, its height 3,4, times in its width; lower border of sub-operculum and hind border of inter-operculum almost parallel to upper profile of head, to leave a broad triangular area below (bounded posteriorly by base of pectoral fin). Dorsal surface of head with two prominent longitudinal striae, diverging slightly posteriorly, flanked by two small lateral striae over eyes. Dorsal fin origin set far back on body, equidistant between caudal base and posterior margin of operculum. Pectoral fin tips reaching almost to tips of pelvics; axillary scale present, } length of fin. Pelvic base nearer to pectoral base than to anal origin by 2 eye diameters; axillary scale present, about 4 length of fin. Anal fin origin below vertical from anterior third of dorsal base; base of fin covered by low scaly sheath. Scales: unexposed portion with a single complete W-shaped vertical striation, preceded by 3 (anterior scales) to 6 (posterior) shorter striae interrupted at centre of scale; exposed portion with about 16 very short radiating striae at edge of scale, not discernible in posterior scales. Colour: in alcohol, upper 4 of body brown, rest of flanks silvery; fins hyaline, hind margin ot caudal dusky. Inner face of operculum slightly dusky. Note. Ilisha sladeni closely resembles J. pristigastroides (Bleeker), with which it has been synonymized (Whitehead, 1970), but comparison of Day’s material with Bleeker’s type in the British Museum (1867.11.28.12—redescribed in Whitehead et alii, 1966) suggests that I. sladeni is distinct. In both species the anal origin is well before the vertical from the midpoint of the dorsal base, a feature used in keys to separate these species from all other Ilisha (Whitehead et alit, loc. cit.; Whitehead, 1970). The type of I. pristigastroides is a smaller fish (151 mm S.L.; cf. 189-210 mm in the Day material), but this does not account for its deeper body (30:8% of S.L.; cf. 21-9, 25-4 and 22-4% in the Day material) since a larger Bleeker specimen (302 mm S.L., BMNH.1867.11.28.9) is still deeper-bodied (30:9% of S.L.) than a similar large specimen of J. sladeni (25-9% ina fish of 308 mm S.L., ex Sittang river, Burma, BMNH.1891.11.30.402). The Bleeker type also has a relatively longer anal base (41:1% of S.L.; cf. 30-0, 29:4 and 30-8%), the anal origin being set further back on the body (equidistant between caudal base and eye centre; cf. nearer to caudal base than to pectoral base), and the dorsal origin is also corres- pondingly less far back on the body. In spite of the more compact body in I. pristigastroides, there are more pre-pelvic scutes (26) than in the elongate J. sladeni (23-24). All these features also serve to separate the two larger specimens mentioned above. A striking feature of I. sladeni, to some extent shared by the type of J. pristi- gastrotdes, is the very elongate appearance of the head, shown well in Day’s drawing (see Pl. 2). This is partly due to the length of the head but more particularly to THE CLUPEOID FISHES DESCRIBED BY FRANCIS DAY 77 the more squat operculum (its height just over 24 times in head length; cf. 2-2} times in other species). Also, the depth of the head, taken at right angles to the profile at the occiput, is much less (1 ? times in head length) than in other species (1j- 14 times). A further difference between I. sladeni and I. pristigastroides is in the form of the swimbladder. In J. sladeni the swimbladder terminates at the posterior end of the body cavity (BMNH specimens of 209 and 308 mm S.L.), whereas in I. pristigastroides there is a postcoelomic, tapering prolongation down the right side of the body lateral to both the haemal spines and the anal pterygiophores and reach- ing as far as the level of the 15th branched anal ray. The condition in J. sladeni appears to be unique amongst Indo-Pacific members of J/isha but it is found in the South American J. furthii (and probably also in the related I. amazonica—no BMNH specimens). The asymmetrical postcoelomic prolongation in J. pristi- gastroides is similar to that found in the Indo-Pacific J. elongata and I. megaloptera; in I. africana (West Africa) and I. indica (Indian Ocean) the prolongation of the swimbladder is bifid. Family ENGRAULIDAE 6. Engraulis auratus Day, 1865 = Thryssa dussumieri (Valenciennes, 1848) (Plate 2) Engraulis dussumieri Valenciennes, 1848, Hist. Nat. Poiss., 21: 69 (no locality; putative neotype described by Whitehead, 1967 : 142); Day, 1878, Fishes of India: 627, pl. 158 (4) (E. auratus in synonymy); Idem, 1889, Fauna British India, Fishes, 1 : 391. Thryssa dussumieri: Whitehead (in press), Symp. Indian Ocean Adj. Seas. Mar. biol. Ass. India (key, synopsis, fig.). Engraulis auratus Day, 1865, Proc. zool. Soc. Lond.: 312 (Cochin on Malabar coast; on specimen 438; inches = 117 mm); Idem, 1865, Fishes of Malabar: 238, pl. 19 (2) (repeat; fig. (? life-size) 44 inches = 114-7 mm tot. 1.). MATERIAL. a. ZSI (no specimens) b. AMS (no specimens and none in Great Fisheries Exhibition Catalogue by Day, 1883) NHMV (no specimens) RMNH (no specimens) ZMB.10412, a fish 89-7 mm S.L. (109-2 mm tot. 1.), ex Bombay BMNH.1867.5.30.13, a fish 83:1 mm S.L. (99-4 mm tot. 1., caudal lobes damaged, estimated 103-3 mm), ex Madras, coll. Day BMNH.1889.2.1.1779, a fish 90-6 mm S.L. (112-7 mm tot. 1., caudal complete), ex Malabar, coll. Day (outside label altered from E. auratus to mystax Gthr.) (label inside jar, Engraulis auratus Malabar), LECTOTYPE ho ao ga 78 P. K. TALWAR & P. J. P. WHITEHEAD h. BMNH.1889.2.1.1780, a fish 55:2 mm S.L. (68-9 mm tot. 1., caudal complete), ex Canara, coll. Day (label in jar, Engraulis auratus Canara) In the Fishes of Malabar (p. vi) it is stated that a specimen of E. auratus had been deposited in the East India Museum (but apparently not in the British Museum). Manuscript catalogues and lists of zoological material presented to the museum of the East India Company are now in the British Museum (Natural History). One list of fishes 1s headed “The following families are from Day’s Malabar Fishes’ and it includes Engraulis auratus from Malabar (preceded by the number 5, which seems to be an indication of the number of specimens). When the India Museum was dispersed in 1879, at least two of Day’s presentations (birds and fish skins) were sent back to Day. Since no fishes were given to the British Museum at that time, Day probably also received back his spirit specimens, including Engraulis auratus. It is possible, therefore, that the specimen (or one of them) is that now in London and presented in 1889 (specimen g above). There is a discrepancy of 2-3 mm between the length of the single specimen described in the original description and the length of the figure, which suggests that the figure was not exactly life-size. Since all four extant specimens are too small and there can be no certainty which, if any, contributed to the original des- cription, we have chosen the British Museum Malabar specimen (g above) as lecto- type. DEscRIPTION. Based on the LECTOTYPE, a fish 90-6 mm S.L. (112-7 mm tot. 1.) in good condition, ex Malabar, BMNH.1889.2.1.1779. Br. St. 11, DI ili 10, Pi rz, Vi 6, A iii 32, g.r. 15 + 18, scutes 15 + 7. In percentages of standard length: body depth 27-8, head length 26-9; snout length 3°6, eye diameter 6-4, length of upper jaw 41-1, length of lower jaw 19:3; pectoral fin length 18-1, pelvic fin length 11-1, length of anal base 33-0; pre-dorsal distance 49:0, pre-pelvic distance 42-2, pre-anal distance 59-3. Body compressed, its width 3 times in its depth, the latter only slightly greater than head length; belly not strongly keeled, all but tips of scutes concealed by scaly sheath; head profile rising steeply from snout to nape and then more gradually to dorsal origin, belly profile evenly convex. Snout short, about = eye diameter. Upper jaw very long, the right maxilla pointed posteriorly, reaching to pelvic base and about # along pectoral fin (tip of left maxilla broken off); expanded portion of maxilla tapering rapidly behind 2nd supra-maxilla but with a membrane along upper edge; 2nd (posterior) supra-maxilla with upper part of expanded portion larger than lower; no Ist (anterior) supra-maxilla. Dentary symphysis below mid- point between eye and tip of snout; articulation of lower jaw 4 eye diameter behind 2nd supra-maxilla. A single series of fine conical teeth on dentaries, pre-maxillae and along lower edges of maxillae except near tip; two (right) and three (left) conical teeth on vomer; fine granular teeth on tongue, palatines and on endo- and ecto-pterygoids. Gillrakers fine, slender, the longest 14 times length of corresponding gill filaments and almost equal to eye diameter; 8 short, triangular rakers on posterior face of 3rd epibranchial: gillraker serrae on both Ist and 2nd arches in distinct clumps, THE CLUPEOID FISHES DESCRIBED BY FRANCIS DAY 79 with the longest serae in the middle of each clump. Pseudobranch present, con- cealed by skin but with small posterior opening. Isthmus silvery, not bearing anterior members of scute series, tapering evenly to just behind posterior margin of branchiostegal membrane. Operculum 3} times as deep as broad, its posterior margin evenly rounded and not completely covering gill opening. Dorsal surface of head covered by thick layer of skin with numerous pores; posterior frontal fontanelles present, long and narrow, 2:1 mm by 0-5 mm (right). Dorsal fin origin nearer to snout than to caudal base by 1 eye diameter; fin preceded by a small scute-like plate bearing a retrorse spine. Pectoral fin reaching just over 3 along pelvic fin; axillary scale present, just over half length of fin. Pelvic fin base I eye diameter before vertical from dorsal origin and much nearer to pectoral base than to anal origin; axillary scale present, # length of fin; a second triangular scale present, below fin, } length of fin. Anal origin 3 eye dia- meter behind vertical from last dorsal ray. Caudal peduncle a little deeper than long. Scales : distinct anterior and posterior ‘shoulders’ to scale; unexposed portion with 10-12 irregular vertical striae, not interrupted at centre of scale, exposed portion with one semicircular striation but more frequently reticulated, especially in posterior scales, the reticulations finally covering the whole scale. Many elongate scales at base of caudal but no true alar scales. Colour: general body colour silvery/gold, but brown where scales lost ; dark brown venulose humeral area with peppering of dark pigment across back (as in Day’s figure—see Pl. 2c). Fins hyaline except for narrow dark posterior border to caudal. Note. The long maxilla, absence of the rst supra-maxilla and distinct clumping of the gillraker serrae are characteristic of T. hryssa dussumieri (descriptions in Whitehead, 1967 : 142, fig. 14c and 1968 : 23, fig. 2a), and Day (1878) later placed his auratus in the synonymy of that species. Pp. K. TALWAR & P. j- P. WHITEHEAD 80 (vruanjord san (Ztgr “Te A) * se osye) * * vivIvAoy] vsojVNIS| (I) zQI gtO ayy vadnia Legt “eA Aajswso1a, vyjaUipéavs — 9€9 4ajsvsoia) vagnig (vsouauan se) bz-6161'1'z (gZgr ‘Aeq) 6rSz7HNWY +SSorgWz 6ggsr'HWNaA zbola'sNV of9z [SZ sisuapurs vyjauipavs (z) €g1 gto stsuapurs vagn{D (4¥gr “ye A) * * * * * * vywiquiy vyjauripaws §(€) 191 LEO vywuquy vadna zSgi “IIE * * vuoskyovag vyjauripavs (€) €gt S€g wmosdyavaq vadnjo (6zgr “ang) * * * vanunjau vyauipavs (S$) tgr 9€9 vpnvriv vadnia (6zgr “and) * DANUDIAUL DIJIUIPADS — tg vanunjam vagdnia Legr “eA * * * * * sdaa1suo) vyjauripawg (z) 191 LEQ sdansuo, vadnja (4€g1 “ddnay) * (stsuasanjou se) * * snyojaund sXyjyoisjopysazy (1) E91 9£9 vazunyy vadnjg (eeutedny9) Legit “Tea 1yjassvy (sapiosdoja se) * * vinav viaauunssnq (S$) 991 Ltg Diuatuinssnq Lrgr “TRA * . * * * pinoy visatunssnq (bt) 991 Lhg DvnIv viAaLUNssNng (aeurierurmnssnq) avdiadn To (SZZ1 “yssi0,q) * x * * * qvaop snaquazoary (€) 991 zSQ quaop snAquaz0Ai14D AVGININADONIHOD urpiog. PUUeTA uopuoTy Saupss e}WNIeg vipuy fo saysty uapiay “sn sn “sn “sn *f@AINS uor}eOyIyUSP] ‘sy «6 oded ut seeds “snusy{Qy [007 -3styanyzeN Jug. “qsny [007 (sadA} ajqissod 10f uaAt3 Joquinu Uor}eI4sI8e1) WOTDa][OO ut Juesaid, uepley pue ulpieg ‘euusr,~ ‘uopuo7y ‘AaupAs ‘ezQn9]e9 Ul [eLIazeu Aeq pue wpuy fo saysry sAeq ut saysy proadnyo ayy, I alavy 81 THE CLUPEOID FISHES DESCRIBED BY FRANCIS DAY 98St HN 9z4z HNWY €rbtorgwz * (£g -d ose aes) (voyasip se) * 6Lgr1z “6881' HNWA (sn1j409 se) * gt'bi-9 ‘oLgr HNWA S—ofoz'1°z “638 HNWA gboz'1'z “638l HNWA (voysjtp se) * * * LEolqSWV $69z'ISZ * * 9Lola'SWV SbzzISZ 88784 SNV 9hzz SZ (6€gr “ureas) woipur DYysuT (6€gr “urems) vaipur nysyy (6€gr “ureas) vILpuL vYSUT (o€gr “uueg) Dywsuoja vysyy (o€gr “uuag) vywsuoja vysyy Zhgr “ea vjayIpp vuonjag Leer “Tea wjayrpip vuonag (zzgi “yong ‘urezz) DPUNIv_YI DULOJSOJUOPOU Fr (69g1 ‘Aeq) DISapoul vsojv1U0+ (zzgr “yong ‘wezz) Durnuunu vsoj]MI1U0y (S621 ‘yoorq) SNSDU VSO]DIVIMa (69gr ‘feq) Djvsa1Ava wisnpNy (zzgr “yong ‘wezq) vadvys visnpny (4g “yea) 40} DSILH (zzgi “yong ‘urezz) YSU OSIVET (6zgr “ang) 99124 DSIUT 6z61 ‘ereseAtueiaq syyoiany vavary a (€Zgr ‘Aeq) DIVADQvIVUL VIaKD ZZBI “Yong “wey DUAOQOS DILAOD (§) (S) I9QI zg sho zbg vutoskyonaq vuojag snyoum vuoyag vIupur wuojjag Yynvuayssay Duojag vyosuoja vuoYag tuanaoy vuonag vjaynip vuoyag (seuriezses1yst1q) vpunovy9 snssaojvyD Snysapout snssaojvy 9 Dunuunue snssaojvyD Snspu snssaojvyD (aeuryeurI0so10q) vywsarava vadnt9 vadoys vadnia 104 vadnya vysyr vadnyo vyansouvy vadrig (eeutsory) SNILADQDIDUL sapropjaywads (eeurpnuojeg) DUAOGOS VILAOD PAK. TALWAR & PP. j. P. WHITEBEAD 82 * (snqvann) ziFol'gqwz * (snqoanv) E1rof'S ‘Lost HNWA * (uqjassna se) * gf'b19 ‘ofgl HNWA * zL9zISZ (oSgr ‘10}ue5) sdazinasq vuurdryas (zzgr “yong ‘urezyz) vsvyd vuurdyas (grgr “TeA) Ayo vuurdyas (6rgr “111q) sisuajpmuiDy vsstAy T (S6Zr “yoorq) DILADQujvUL DSSAAY (S€gr ‘Aviy) 1UuopuUDy vSshAy T (grgr “Te A) taatunssnp vsshAy T (zzgr “yong “ureyz) vavang vsshay J. (rogr “uy9s) xvjstua vsstay T (zg “ssnorg) SUAJSOAIJAS DSSKAY T. (SZZ1 “yss10,7) DULYIAVG DUISSIAY T, (€ogr ory) UUwosAaMmor snsoYygajojS (€zgr ‘sseyy ura) snovpur snaoydajoys (zSgr “r4Iq) 14q snsoydajojs 1€gr ‘Avi DUDIaSSNA DPUOIDY (6zgr “ang) asoopan} snaazgoyssidc, (6981 ‘Aeq) 1mapyjs DYSYT (6€gr ‘urems) vaaqojwsaut DYsty (6€gr ‘“urems) vaajqoppsaum vysyy 829 fto Sto sdarinadq synvasug vavay SYNDATUA Ayo} synvssug sisuajpuuDy synvasud snaLnqnjpur synvAsUq UUWONUMDY SYNDATUT lalunssnp SYNnvASUT vavand sinvasuq xvjshut synvssug SUISOAYAS SINDASUT punjaog synvasugq SnuDuossauuLor sunvasuq snoipUr synvAsuy 14q SYNDATUT HVATITNVYONA DUDIIJASSNA DPUOIDAT Aoozany snsazqoyjsidQ luapyys DUuOTIaIT vaasyy Duojag vaajgopsaum vuonag 83 THE CLUPEOID FISHES DESCRIBED BY FRANCIS DAY setoads z€ ‘uaploT saroeds gr “LLgr ‘1'z'6ggr ‘TN jo suaurtoads s,Aeq Jo auQ “snswu- py 04 asor soroeds €z ‘PUUST A soroeds 1b ‘uopuoT satoads oF saroads 6€ Aeq Aq peztuSosex asoy ysduoure saroads prea ob shgi “Tea taaluinssnp v1y109 grgr “Tea ¢wpypucar vytog grgr “Tea wouter vytog (zzgr “yong ‘wrezz) YVADIUIDA D1Y10D (zzgi “yong -urezz) YVADIUMIDA DYL0D (tzgr “yong ‘wezz) YVAVIUMIDA 117109 (z) 661 Wd ‘ereur) xa ‘sofoods mou sry} st wasn Ysytg 94} UT susvu: N 9 ‘psojvjvuan Jo seeds Mou v Suiqiiosep st UOS[aN YJoIeD “Id : aj0N eq Aq pays seroeds proednyo S¢ taaunssnp v1Y4109 stsuaausog vYyt0D wouter vyiog sypuusasvaponb mi109 S1dojuna vY10D YVADIULDA DI10D) 84 P. K. TALWAR & P. J. P. WHITEHEAD REFERENCES ANON. 1885. Australian Museum (Report of the Trustees for 1884). Govt. Printer, Sydney, 46 pp. App. XI (p. 29)—report by E. P. Ramsey; (pp. 42—46)—list of Day specimens. 1886. Australian Museum (Report of the Trustees for 1885). Govt. Printer, Sydney, 17 pp. (p. 5)—list of Day’s specimens, cases II and III. Cuan, W. L. 1965. z 89 INTRODUCTION c : A “ 2 5 : : : 89 MATERIALS AND METHODS . 5 : : : * 3 : : 90 RESULTS ; 4 0 C 0 6 5 6 5 : " 90 Light microscopy O 2 : 7 3 : : é go Bodo saltan. 5 5 : : 4 F a 90 Bodo caudatus . : 6 c g é é 92 Electron microscopy . é é A c ° ¢ ¢ a 92 Flagellar pocket . 5 : 0 : 92 Basal bodies and flagella. . : 7 3 92 Alimentary system . é 3 é c ¢ 93 Microtubular systems : c c : 5 94 Kinetoplast-mitochondrion c 5 < : 95 Nucleus . 6 ; 2 6 4 4 96 Endocytoplasmic bacteria . . 5 c ° 96 Cytoplasmic membrane systems . ¢ : : 96 DIscussIoN. 5 0 ¢ : ¢ : : : 5 5 97 ACKNOWLEDGEMENT A : : 5 6 . : A : IOL REFERENCES . 0 : 6 ; 5 5 : : 6 IoI SYNOPSIS Characters specific to each of two species of Bodo are described for the first time. Bodo saltans possesses cytoplasmic bacteria, hair-like appendages (mastigonemes) on the anterior flagellum and circumbuccal lappets surrounding the opening of the alimentary system. In B. caudatus, an electron dense band separates the kinetoplast from the basal bodies of the flagella. In addition, clear differences exist between the microtubular systems associated with the buccal cavity and cytopharynx. It is suggested that the mastigonemes and circumbuccal lappets of B. saltans are responsible for the capture of food organisms. In both species the alimentary system is a membrane-lined tube surrounded by a number of microtubules. The single mitochondrion is dilated in the vicinity of the basal bodies and contains a prominent kinetoplast. Since both these organelle systems closely resemble those found in members of the related Trypanosomatidae, the possible origin of this family from Bodo or a Bodo-like flagellate is discussed. INTRODUCTION Bodo is a cosmopolitan flagellate found in fresh and brackish waters and in some soils. This and other members ot the Bodonidae are of interest chiefly because of their close relationship to the economically and medically important trypanosomes. go B. E. BROOKER This relationship is based on the presence of a mass of DNA—the kinetoplast— situated in a dilatation of the single mitochondrion. In trypanosomes, this organelle has been regarded as a genetic system containing the information required for the synthesis of mitochondrial enzymes (Steinert, 1g60) but in Bodo its function has not been examined. Of the many species of Bodo which have been described, Bodo saltans Ehrenberg, 1831 and Bodo caudatus Stein, 1878 are probably the two most commonly found. They occur in quite different habitats for whereas B. caudatus tends to be coprozoic, B. salians is usually found in freshwater. Separation of the two species depends on such characters as body shape and size, length of the flagella and the position of the nucleus relative to that of the kinetoplast. However, the present fine structural study describes a number of clearly defined qualitative differences between B. saltans and B. caudatus which would not have been visible to the earlier light microscopists. MATERIALS AND METHODS Bodo saltans was isolated from a sample of fresh water taken from a pond near Slapton Ley, Devon. From this isolate, clone cultures were established on 0-1% w/v ‘Oxoid’ dehydrated liver intusion (pH 4-6) and maintained at 25°C. Bodo caudatus was isolated from an infusion of pig faeces which was obtained from Winches Farm, near St. Albans, Hertfordshire. Clone cultures from this isolate were maintained at 25°C on 0:2% w/v ‘Oxoid’ beef extract (pH 5-8). In both cases, cultures were agnotobiotic. For light and electron microscopy, cultures were harvested after 3 days growth. Light microscopy.—Phase contrast observations were made using a Leitz Ortholux microscope fitted with a Heine condenser. Flagellates were examined either alive or after fixation with 1°/, osmium tetroxide. Smears fixed in Schaudinn’s fluid were stained with iron haematoxylin and examined by bright field microscopy. Electron microscopy.—Flagellates were collected by centrifugation at 1,000 r.p.m. for 10 minutes and the resulting pellet fixed for 5 minutes at room temperature in 1% osmium tetroxide buffered to pH 7-4 either with o-1 M Sorensen’s phosphate buffer or 0-1 M veronal acetate (Michaelis). Before dehydration, the pellet was treated with 1°% uranyl acetate in 25% ethanol for 30 minutes. After dehydration in ethanol-water mixtures and absolute ethanol, the pellet was treated with propylene oxide or toluene and embedded in Araldite. Sections were cut using a Porter-Blum MT2 ultramicrotome and stained in lead citrate prior to examination in an EM 6B electron microscope. Some flagellates were fixed as before, washed in distilled water and dried onto grids. They were then placed in a coating unit and shadowed with gold/palladium at an angle of 30°. Negatively stained preparations were made using sodium phosphotungstate at pH 7-0. RESULTS LicHt mMiIcroscopy.—Bodo saltans.—The body ot this flagellate is oval in shape (5-8 um long and 2-5 um wide) and has two flagella of unequal length which arise from the bottom of a depression (the reservoir or flagellar pocket) near the anterior STRUCTURE OF BODO SALTANS AND B. CAUDATUS gl end of the cell (Fig. 1). During locomotion in which the body rotates on its own axis, the shorter anterior flagellum is extremely active while the posterior flagellum remains stationary or undergoes slight movement. When stationary, the flagellate often attaches itself to some object by the tip of its trailing flagellum and may then exhibit rapid oscillations. This behaviour is characteristic of B. saltans. The kinetoplast lies near to the basal bodies of the flagella and the nucleus, which con- \ WH tah | ) LAPPETS FLAGELLAR POCKET KINETOPLAST CYTOPHARYNX BUCCAL CAVITY ] y \ 2, CONTRACTILE Yr \ VACUOLE \ CYTOPLASMIC BACTERIA MITOCHONDRION FLAGELLA NUCLEUS FOOD VACUOLE Fic. 1. Diagram of Bodo saltans showing the arrangement of the main organelles. tains a conspicuous central karyosome, is central or mid-ventral in position. The single mitochondrion is markedly siderophilic and appears as a thin filament which originates from one side of the kinetoplast, describes a figure of eight or a loop within the cell and terminates at the opposite side of the kinetoplast. A single, round contractile vacuole appears at intervals at the anterior end of the living cell. It is situated just below and to one side of the flagellar pocket into which it periodic- ally discharges its contents. The posterior half of the flagellate is occupied by a variable number of food vacuoles. 92 B. E. BROOKER The opening of the alimentary system, the buccal cavity, is marked by a small vacuole on the ventral surface near the anterior end of the flagellate. Ingestion of bacteria is a very rapid process and can be observed satisfactorily only during the short periods of quiescence when it is attached to the substratum by its posterior flagellum. During ingestion, a bacterium is drawn into the buccal cavity and rapidly passes ventro-dorsally along a path which corresponds exactly with that of the cytopharynx (see later). A large vacuole then appears at a point which prob- ably corresponds to the end of the cytopharynx and the bacterium passes into it. This food vacuole then slowly moves to the posterior end of the cell. Bodo caudatus. When harvested in the logarithmic phase of growth, B. caudatus is long and narrow with a convex dorsal and concave ventral surface (8-14 ym long and 4-6 um wide). The two flagella are of unequal length and arise trom the bottom of the flagellar pocket near the anterior end of the cell. The anterior flagellum is the shorter but most active during locomotion. Situated in the anterior half of the cell, the nucleus frequently lies very close to the kinetoplast. As in B. saltans, the single mitochondrion is continuous at both its ends with the kineto- plast and describes a loop-like circuit of the cell. Almost the entire posterior half of the flagellate is occupied by food vacuoles and an active contractile vacuole lies close to the flagellar pocket. Although anteriorly a small buccal cavity is clearly visible, the ingestion ot bacteria is very difficult to observe because of the relentless swimming habit of this flagellate. ELECTRON MICROscopy.—Flagellar Pocket. In both species of Bodo, the flagellar pocket is a lateral depression at the anterior end of the body. Because the two flagella emerge from the cell at the bottom of this structure (Fig. 1, Pl. 1, Fig. A), Pitelka (1961) referred to it as the circumflagellar depression. The flagellar pocket is lined with a unit membrane continuous with that covering the rest of the cell and below this, there lie a number of pellicular microtubules. Further details of these are given below. Basal bodies and flagella. The two basal bodies are embedded in the cytoplasm at the base of the flagellar pocket and are therefore antero-lateral in position. They are structurally differentiated into 2 regions, an intracytoplasmic proximal portion and an extracytoplasmic transition zone which connects the intracytoplasmic portion to the flagellar shaft and is bounded by the flagellar membrane (PI. 1, Fig. A). The proximal end of the basal bodies lies very close to the surface of the kinetoplast capsule (Pl. 1, Fig. A). The gap between these two structures is some- what variable in width but is generally in the order of 100 nm and direct contact has never been observed. Under some conditions of fixation, a reticulum of fine filaments (8 nm wide) fills each basal body. The junction of the basal body transition zone with the flagellum is marked by two transverse basal plates which are thickened peripherally (Pl. 1, Fig. B). The two central tubules of the axoneme originate just above the distal basal plate and the paraxial rod—a structure which runs parallel to the axoneme for about three quarters of its length—arises from a lateral extension of the proximal basal plate. The basal plate extensions of each flagellum face one another and the overlying flagellar STRUCTURE OF BODO SALTANS AND B. CAUDATUS 93 membranes are joined by a wide but very thin extracellular striated band (PI. 1, Figs B and C). Of the three closely spaced striations, only the central one (5 nm wide) is prominent. Those lying on either side of it are more diffuse and can be satisfactorily resolved only in longitudinal sections of the band. The transition zone of the basal bodies consists of 9 peripheral doublets of tubules each of which is joined to the adjacent flagellar membrane by a narrow connective. The intracytoplasmic portion is composed of g tubule triplets and is open proximally. The proximal ends ot the basal bodies are connected by three striated rootlets (Pl. 1, Fig. D). Two of these arise from a single triplet on the basal body of the posterior flagellum and diverge slightly as they approach and insert onto the other basal body. A third rootlet running approximately parallel to the other two also connects the basal bodies. All three rootlets have a major period of 50 nm. The prominent bands delimiting the major repeating unit are 25 nm wide and between these lies a narrower 5 nm wide band. There is in addition, a ‘Y’ shaped rootlet which arises from the basal body of the anterior flagellum and passes antero-laterally to insert onto a group of 2-3 short microtubules running parallel to the cytopharynx (PI. r, Fig. D). Both flagella contain the familiar ‘9+2’ arrangement of tubules and a paraxial rod (Pl. 1, Fig. E). After running parallel to the axoneme for about three-quarters of its length, the paraxial rod tapers before terminating. Details of its structure are difficult to resolve but it appears to have a lattice-like architecture. The anterior flagellum of Bodo saltans, unlike that of B. caudatus, bears mastigonemes. These hair-like appendages are arranged in bundles along one side of the flagellum and have been reported elsewhere (Brooker, 1965). Metal-shadowed preparations of B. saltans show that the posterior flagellum bears a number of parallel transverse striations whose separation is 14 nm (PI. 1, Fig. F). They are found only after the flagellum has emerged from the flagellar pocket and extend distally for a distance of 3um. In B. caudatus similar striations first appear at a level corresponding to the junction of the basal body with the flagellum but because they extend distally only for a distance of about 1 ym, they are not visible in shadowed material. In sectioned material, the striations are seen as periodic thickenings (14 nm wide) of both leaflets of the flagellar membrane (PI. 1, Fig. E, Pl. 2, Fig. A). Such thickenings occupy about 25% of the circumference of the flagellar membrane (PI. 1, Fig. E). On passing distally, this percentage gradually decreases so that in shadowed prepara- tions of B. saltans, the array of striations appears to taper to an end (PI. 1, Fig. F). Alimentary system. The ingestion of food particles by Bodo takes place by way of permanent oral structures—the buccal cavity and cytopharynx—which lie to the right of the flagellar pocket (Pl. 2, Fig. B). The position at which the buccal cavity opens on to the surface of the cell differs in the two species. In B. saltans, it opens . antero-ventrally (Pl. 2, Fig. C) but in B. caudatus it is found at the extreme anterior end of the flagellate (Pl. 2, Fig. D). Very rare sections in which a bacterium is found in the buccal cavity (PI. 1, Fig. A) suggest that this organelle is capable of considerable distension. Frequently, the membrane lining the buccal cavity has a pronounced cell coat (Pl. 2, Fig. C) which takes the form of bundles of fine filaments projecting perpendicularly from the membrane. A number of structures, referred 94 B. E. BROOKER to here as circumbuccal lappets, surround the opening of the buccal cavity of B. saltans (Pl. 2, Fig. B). They are flat, triangular projections (Pl. 3, Fig. A) which are joined at their bases to form a ring. Each lappet is composed of many fine filaments and arises from an intracellular band at the margin of the buccal cavity. These structures appear to be totally absent from B. caudatus. From the left side of the buccal cavity arises an elongate tube lined by a unit membrane and surrounded by a number of microtubules (Pl. 2, Fig. B). In accord- ance with the terminology used for the oral apparatus of ciliates, this organelle will be referred to as the cytopharynx. In Bodo saltans the cytopharynx approaches the kinetoplast, passes underneath and to one side of it and on reaching the dorsal surface of the body, curves to one side (Pl. 3, Fig. C). In B. caudatus, it passes posteriorly just below the cell membrane (Pl. 2, Fig. D). Although the cytopharynx terminates just posterior to the kinetoplast, the microtubules associated with it frequently reach the posterior margin of the nucleus before terminating. In both species, the diameter of the lumen decreases distal to the buccal cavity. Numerous vesicles of variable diameter are always found underneath and to the right side of the cytopharynx in B. saltans (Pl. 2, Fig. B, Pl. 3, Fig. C). Although similar vesicles are occasionally found in B. caudatus, they are generally smaller and less numerous. Their origin is not clear, but many longitudinal sections of the cyto- pharynx suggest that they arise as invaginations of the cytopharyngeal membrane. Both light and electron microscope observations suggest that ingestion of food particles takes place at the blind end of the cytopharynx. Since the diameter of the cytopharynx is at all points along its length smaller than that of food organisms found in the food vacuoles, ingestion must be accompanied by considerable dis- tension of the cytopharynx. Ingestion results in the formation of a number of food vacuoles which migrate to the posterior half of the cell. The food vacuoles are bound by a single unit membrane and may contain one or more food organisms (Pl. 5, Fig. D). When digestion is complete, only a diffuse mass of undigestible material remains in the vacuole. Since such vacuoles do not accumulate in the cytoplasm, it seems likely that undigestible material is voided by the coalescence of the cell membrane with that of the food vacuole. However, this has never been observed. Microtubular systems. Microtubules of external diameter 20-25 nm are associated with several organelles in the anterior half of the flagellate. Many, but not all, of the tubules which are associated with the flagellar pocket, cytopharynx and body, appear to arise from the proximal ends of the two basal bodies. Bodo saltans. The microtubules which are to surround the cytopharynx pass along the flagellar pocket in two groups. At the opening of the flagellar pocket, three members of each group approach each other (Pl. 3, Fig. B) and continue as microtubule doublets. They curve over the narrow bridge ot cytoplasm separating the flagellar pocket from the alimentary system and enter the walls of the buccal cavity where they adopt a ‘U’ shaped configuration. From the left side of this cavity, the microtubules follow the course of the cytopharynx and become arranged around it in two groups. Associated with the floor and left side of the cytopharynx is a group of 5 microtubules and with the roof a group of 3 (Pl. 3, Fig. D). The right STRUCTURE OF BODO SALTANS AND B. CAUDATUS 95 side of the organelle is free of tubules. The middle 3 tubules of the group of 5 are double and joined on their left side to the cytopharyngeal membrane by a short connective. Although 8 microtubules normally run parallel to the cytopharynx, 9 or Io are sometimes found (Pl. 3, Fig. E). The microtubules of each group are connected by a number of fine filaments which occur at intervals (13 nm) along their length (Pl. 4, Fig. A). Beyond the blind end of the cytopharynx, the connectives of the microtubule doublets disappear but the arrangement of the two groups of tubules remains constant until they terminate near the dorsal surface of the flagellate. Beneath the cell membrane at the anterior end of the cell lie a number of pellicular microtubules. Although some of these arise from the basal bodies of the flagella, many appear to have their origin near the opening of the flagellar pocket. From here, the microtubules spiral round the anterior end ot the cell and on approaching the right side of the buccal cavity dip below the cell surface (Pl. 4, Fig. B). They then describe a semi-circular path as a curved band of 15-20 tubules (Pl. 4, Fig. C). Passing below the buccal cavity (Pl. 3, Fig. C) they move anteriorly and terminate in the cytoplasm between the cytopharynx and the flagellar pocket. Bodo caudatus. The arrangement of microtubules associated with the buccal cavity resembles that described for B. saltans. The tubules which are to surround the alimentary system pass along the wall of the flagellar pocket and enter the buccal cavity. Here, transverse sections show that the tubules are arranged in a line along the left wall (Pl. 4, Fig. D, Pl. 5, Fig. A). As the tubules follow the cytopharynx from the left side of the buccal cavity, they distribute themselves around the cytopharyngeal membrane (PI. 5, Fig. B) in 2 overlapping groups. One group associated with the roof of the cytopharynx usually contains 4 microtubules; the other group, which lies next to the floor and left side of the organelle, may con- tain 4, 5 or 6 tubules three of which are double and joined as in B. saltans to the cytopharyngeal membrane by a short connective. Such connectives disappear a short distance from the buccal cavity. Another set of microtubules emerges from the flagellar pocket, spirals round the anterior end of the cell and comes to occupy the right hand wall of the buccal cavity just below the lining membrane (PI. 4, Fig. D). The 15-20 microtubules in this set are joined by intertubular connectives and for most of their length travel posteriorly parallel to the buccal cavity and cytopharynx. A limb of the single mitochondrion which passes beneath the buccal cavity modifies the path taken by some of the tubules. As they travel posteriorly, the tubules gradually move away from the cytopharynx (Pl. 5, Figs A, B) and towards the cell membrane until, at a level beyond the blind end of the cytopharynx, they are seen as a row of pellicular micro- tubules. These tubules travel some distance posteriorly before terminating. In addition to this major set, an equally conspicuous row of very short parallel tubules is found on either side of the buccal cavity (Pl. 4, Fig. D). Kinetoplast-mitochondrion.—Lying just below the cell membrane, the single mitochondrion describes a loop-like circuit of the cell (Fig. 1). In Bodo saltans, this organelle travels above and parallel to the cytopharynx, describes a semi-circular path around the right side of the buccal cavity (Pl. 5, Fig. D) and passes to the extreme posterior end of the cell before returning anteriorly to the basal body region. 96 B. E. BROOKER The whole circuit of the mitochondrion frequently takes the form of a figure of eight. The form of the mitochondrion in B. caudatus is similar to that of B. saltans except that in the posterior region of the former it is frequently seen as a branching structure. The cristae are predominantly plate-like and arise from the inner mitochondrial membrane. The granular matrix of the mitochondrion is dense but sometimes contains irregularly shaped bodies of much greater electron density (PI. 5, Fig. C). In the region of the basal bodies, a prominent spherical dilatation of the mito- chondrial tube houses the kinetoplast. This will be referred to as the kinetoplast capsule. The kinetoplast is composed of a complex reticulum of fine filaments (2°5-3-0 nm thick) and contains a large number of irregularly distributed electron dense nodes (PI. 6, Fig. A). It is separated from the wall of the mitochondrion by a number of cristae embedded in a thick layer of mitochondrial matrix (Pl. 6, Fig. A). The basal bodies of the flagella lie very close to the surface of the mitochondrial kinetoplast capsule but are never seen in contact with it. In Bodo caudatus, the basal bodies are separated from the capsule by a flat electron dense pad (PI. 1, Fig. A). Nucleus.—tThe relative position of the nucleus within the:cell is one of the features by which the two species of Bodo may be separated. In B. saltans it is found mid- ventrally some distance from the kinetoplast capsule whilst in B. caudatus, it is always found very close to this part of the mitochondrion. The nucleus is bound by a nuclear envelope composed of two membranes of which the outer is continuous with the granular endoplasmic reticulum. A prominent and finely granular nucleolus of variable shape occupies the centre of the nucleus (Pl. 5, Fig. D). In B. saltans a layer of condensed chromatin is often seen attached to the inner membrane of the nuclear envelope but in B. caudatus, this component of the nucleus appears to be absent or at least is not visualized by the techniques used in this study. Endocytoplasmic bacteria.—Structures have been observed in the cytoplasm of Bodo saltans which, on morphological grounds, have been tentatively identified as bacilliform bacteria (approximately I ym long and 0-3 wm wide). They are found in all individuals of the flagellate population and are always situated in the anterior half of the cell (Pl. 2, Fig. C, Pl. 4, Fig. C). Although the largest number of bacteria seen in one cell profile is four, the total population is probably much larger. They can be easily distinguished from food organisms since the latter are separated from the cytoplasm of the flagellate by the membrane of the food vacuole and are usually found at the posterior end of the cell (Pl. 5, Fig. D). Each bacterial cell is bounded by a cell membrane 8 nm thick which bears on its outer surface a thin layer of filamentous material in direct contact with the host cytoplasm. Extensions of a layer of dense material lying beneath the cell membrane project into the electron lucent central portion of the bacterium which is traversed by fine fibrils. Deep, mid-length constrictions of the bacteria suggestive of division are commonly encountered and appear independently of the host cell division cycle (Pl. 6, Fig. C). Cytoplasmic membrane systems.—The contractile vacuole is situated on the left side of the flagellate just below and to one side of the flagellar pocket (Pl. 4, Fig. D). Surrounding the vacuole is a number of vesicles and tubules which serial sections STRUCTURE OF BODO SALTANS AND B. CAUDATUS 97 show to be continuous with the lumen of the vacuole (PI. 6, Fig. B). After systole, the membrane lining the vacuole appears rounded in section. At discharge, the membranes between the flagellar pocket and contractile vacuole coalesce and the contents of the vacuole are discharged into the flagellar pocket. The thin layer of cytoplasm between the membranes of the flagellar pocket and contractile vacuole is traversed by a number of concentrically arranged circular septa. Coated vesicles are commonly encountered in the vicinity of the vacuole and often they are seen with their membrane confluent with that of the vacuole. The Golgi apparatus is situated directly below the cytopharynx (Pl. 4, Fig. C, Pl. 5, Fig. D) and to the right side of the contractile vacuole and is composed of a stack of 6-10 compressed saccules. Although vesicles of both the smooth and coated type are actively proliferated from the margins of all the Golgi saccules, there is frequently a notable concentration in the region of the distal saccule. Cisternae of granular endoplasmic reticulum arise from the outer membrane of the nuclear envelope and ramify throughout the cell. One limb of this reticulum is permanently associated with the posterior margin of the cytopharynx and runs parallel to it for most of its length (Pl. 3, Fig. D). DISCUSSION Since, at the level of the light microscope, Bodo saltans and B. caudatus appear to possess the same major organelle systems, separation of the two species is based on differences in the spatial arrangement of these organelles and on differences in the size and shape of the body. Whilst confirming the validity of such criteria, the present study has shown that separation is also possible using characters which are beyond the resolution of the light microscope. Thus in B. saltans, circumbuccal lappets, endocytoplasmic bacteria and mastigonemes on the anterior flagellum are consistently present but are never found in B. caudatus. Similarly, the dense layer of material which separates the basal bodies of the flagella from the kinetoplast capsule is present only in B. caudatus. Although Pitelka (1961) was only able to make a tentative identification of the flagellate she studied, it is clear that it was B. saltans for her pictures show the cytoplasmic bacteria and circumbuccal lappets of this species. The alimentary system and the microtubules associated with it have been briefly reported by Pitelka (1961). She pointed out that the non-contractile rostral vacuole described by Hollande (1942) corresponded to the cup-like depression at the opening of the alimentary system, a structure which has been referred to as the buccal cavity in the present study. Sections of the alimentary system containing partially ingested bacteria suggest that the buccal cavity and cytopharynx are capable of considerable distension. This conclusion is supported by the observations made by Sinton (1912) on a flagellate which he referred to as Prowazekia urinaria but which, from his description, was probably Bodo caudatus. He observed that the flagellate was able to ingest not only large bacteria but also red blood cells and that on these occasions the buccal cavity was capable of being greatly distended. As Sinton describes it, the path taken by the bacteria through the cell during ingestion 98 B. E. BROOKER corresponds exactly to the course of the cytopharynx described in the present study. A similar conclusion has been drawn from the light microscope observations of feeding in B. saltans. These results are contrary to Pitelka’s assertion that bacteria do not pass along the cytopharynx. Although the function of the microtubules associated with the alimentary system is unknown, it is possible that they confer a degree of elasticity on the organelle which enables it to return to its normal shape and size once ingestion is complete. Schuster (1968) suggested a similar function for the cytopharyngeal tubules of the cryptomonad flagellate Cyathomonas truncata and extended this proposal to the case of Bodo. The capture of prey by Bodo caudatus has been described by Sinton (1912). According to this author, the distal portion of the anterior flagellum is capable of grasping bacteria and propelling them to the opening of the buccal cavity by coiling movements. Infolding movements of the edges of the buccal cavity then initiate ingestion. This mechanism is possible in B. caudatus only because the buccal cavity opens anteriorly and is therefore ideally situated to receive bacteria carried to it by the anterior flagellum. Because the buccal cavity of B. saltans opens antero- ventrally, the mechanism described above for B. caudatus does not appear adequate to explain the capture of food organisms. Instead, a mechanism involving the mastigonemes of the anterior flagellum is proposed. Although the mastigonemes do not appear to play a major role in locomotion (Holwill, 1966), it is possible that during the oar-like movements of the anterior flagellum (Holwill, 1966) they exert a component force in the direction of the buccal cavity which sweeps food organisms towards it. Such a mechanism may bring bacteria to the vicity of the buccal cavity but the initiation of ingestion probably depends on movements of the margins of the buccal cavity as described by Sinton (1912) for B. caudatus. In this process, participation by the circumbuccal lappets may be important. It is visualized therefore that the mastigonemes and the circumbuccal lappets are functionally integrated to form a system responsible for the capture and ingestion of food organisms. Schuster (1968) has described a filamentous fringe surrounding the opening of the cytopharynx of Cyathomonas which, like the circumbuccal lappets of B. saltans, has its origin beneath the cell membrane and is believed to assist in feeding. In her study of Bodo saltans, Pitelka (1961) suggested that the cytopharynx was a modified intracytoplasmic flagellum. This suggestion was based on the observation that the cytopharynx is surrounded by 9 microtubules and arises close to the surface of the kinetoplast near the basal bodies of the flagella. However, it has been shown here that the cytopharynx passes beyond the kinetoplast capsule and although occasionally surrounded by 9g or ro tubules, 8 is the usual number. In view of these findings, the homology attempted by Pitelka must be considered doubtful. Vesicles of various sizes are associated with the cytopharynx of Bodo (Pitelka, 1961), Ichthyobodo(Costia)necator (Joyon and Lom 1966, 1969) and Cyathomonas truncata (Mignot, 1965; Schuster, 1968). In the case of Cyathomonas, Mignot (1965) believed that these vesicles arise from the Golgi apparatus and Schuster (1968) has suggested that they contain digestive enzymes which are ultimately emptied into the food vacuoles. In Bodo however, profiles showing an undulatory cytopharyngeal STRUCTURE OF BODO SALTANS AND B. CAUDATUS 99 membrane strongly suggest that the vesicles arise by pinocytosis although in the absence of tracer experiments this can only be conjecture. Although in most respects the flagella of Bodo closely resemble those described from other kinetoplastid flagellates (Pyne, 1960; Anderson and Ellis, 1965; Vicker- man, 1969), they do possess two features, namely the striations of the posterior flagellum and the extracellular interflagellar connective, which are not shared by other members of this group. The significance of these structures is not known, but an extracellular connection between the 2 flagella may go some way to explaining synchrony of these organelles during movement. The paraxial rod of both flagella arises from the proximal basal plate but in the closely related trypanosomatids it only becomes recognizable a short way along the flagellum. The mitochondrion was probably first visualized by Whitmore (1911) who described a fibril from Bodo asiaticus (= B. caudatus) which ran from the kinetoplast to the posterior end of the cell. Alexeieff (1912) observed this ‘fibrille sidérophile’ in B. caudatus and B. edax but reported that it could only be seen in flagellates from young cultures. A detailed study of this structure was made by Hollande (1936, 1942). He noted that it was a constant feature of all species of Bodo but that there was a species difference in the extent to which it was developed. Because the ‘cote’ or ‘cordon siderophile’, as Hollande called it, was best developed in B. saltans, he paid more attention to this species and described in some detail the path taken by it through the cell. As noted by Pitelka (1961), it seems probable that Hollande was describing the mitochondrion since electron microscopy shows that this is the only organelle which follows an identical path through the cell. The enclosure of the DNA-containing kinetoplast in a dilatation of the mito- chondrial tube is a feature uniting Bodo with members of the Trypanosomatidae. However, they differ in the organization of the kinetoplast for whereas in the Trypanosomatidae the kinetoplast is disk shaped with its component fibrils arranged antero-posteriorly, in Bodo it is spherical with its fibrils forming a reticulum. Although in all kinetoplastid flagellates the basal bodies of the flagella lie very close to the surface of the kinetoplast capsule, no physical connection between the two has been found. Such a connection has been sought because in many trypanosomatids the two structures are linked in morphogenesis and appear connected after cell rupture (Simpson, 1968). The present study suggests that in the case of Bodo caudatus no direct connection is possible because the two structures are separated by a thick electron dense pad. The observations made by Simpson (1968) on Leish- mania tarentolae led him to suggest that the kinetoplast capsule is attached to the basal body by an EDTA-sensitive cytoplasmic cement. There seems no reason why this explanation cannot be extended to other kinetoplastid flagellates. Electron dense bodies similar to those found in the matrix of the mitochondrion have also been described from Crithidia fasciculata (Brooker, 1971) and in Tetra- hymena pyriformis Levy and Elliott (1968) found that they become more numerous when the ciliates are starved. Although their significance is unknown, it is inter- esting that these bodies resemble the altered kinetoplast DNA of trypanosomatid flagellates which have been exposed to acriflavine (Kusel, Moore and Weber, 1967; Hill and Anderson, 1969). Although dyskinetoplastic Bodo caudatus was obtained 100 B. E. BROOKER by Robertson (1929) using acriflavine, the electron miscroscopy of the kinetoplast after this treatment has not been studied. The endocytoplasmic bacteria of Bodo saltans appear to be a constant feature of this species. Profiles showing constrictions across the equator of some bacteria support the assumption that they multiply to keep pace with division of the flagellate. Gram stained preparations of B. saltans provide little useful information since it is difficult to distinguish endocytoplasmic bacteria from food organisms. However, since the walls of Gram negative bacteria are very thin (Kellenberger and Ryter, 1958) compared with those of Gram positive bacteria (Glauert, 1962) the cytoplasmic bacteria of Bodo are judged to be Gram negative. Bodies thought to be bacteria constantly occur in or on many species of protozoa as shown by the review of Kirby (1941). In most cases their identity and function is unknown. Since, in the case of Bodo saltans, there is no evidence to suggest that the flagellate benefits or is harmed by the association it is impossible at this stage to decide whether the bacteria are parasitic or symbiotic. However, in another kinetoplastid flagellate, namely Crithidia oncopelti, cytoplasmic bacteria or ‘polar bodies’ have been described by Newton and Horne (1957) which appear to provide the flagellate with lysine (Gill and Vogel, 1962). Although there now appears to be general acceptance of the theory which holds that trypanosomes originated from the intestinal flagellates of insects (Hoare, 1948; Baker, 1963), the origins of the family still seem unclear. The presence in all trypanosomatid flagellates of a barren basal body in addition to that which produces the single motile flagellum (Rudzinska and Vickerman, 1969) may indicate origin from a biflagellate ancestor. Since at some stage this is likely to have been a free living flagellate, possible descent from Bodo or a Bodo-like organism is worth a brief consideration. Although the kinetoplast-mitochondrion is an obvious character uniting both groups of flagellates, it appears that an organelle comparable to the cytopharynx of Bodo has also been retained in a more or less modified form by many trypanosomatids (Brooker, 1971). In trypanosomatids, it is a deep (cytopharynx) or shallow (cytostome) invagination of the cell membrane associated, as in Bodo, with a number of microtubules. In both cases this organelle is endocytotic but whereas in Bodo it is primarily concerned with the ingestion of bacteria, in those trypanosomatids which have been examined it appears to be pinocytotic (Steinert and Novikoff, 1960; Preston, 1969; Brooker, 1971). Qualitative and quantitative differences in the nature of food ingested by Bodo and trypanosomatids may go some way to explain observed differences both in cell shape and the spatial arrangement of some organelles. Thus, the cytopharynx of Bodo, unlike that of trypanosomatids, is situated some distance from the flagellar pocket in order to facilitate prey capture and virtually the entire posterior half of the cell is devoted to the accommodation of food vacuoles. It is proposed therefore that the adoption of a parasitic mode of life by a Bodo-like flagellate and the subsequent abandonment of bacterophagic nutrition could have produced changes in body form which, together with other physiological adaptations led to the emergence of an ancestral trypanosomatid. Although such changes would have resulted in the retention of the cytopharynx, it STRUCTURE OF BODO SALTANS AND B, CAUDATUS 101 is assumed that the loss of one flagellum and development of the relatively sparse microtubular system occurred at some later stage. ACKNOWLEDGEMENTS I gratefully acknowledge the technical assistance given by Mr. C. G. Ogden who was responsible for many of the electron micrographs used in this paper. KEY TO ABBREVIATIONS USED IN THE PLATES ax axoneme G Golgi apparatus b bacterium ger granular endoplasmic reticulum bb basal body ifc interflagellar connective be buccal cavity k kinetoplast cb cytoplasmic bacterium m mitochondrion cbl circumbuccal lappets mast mastigonemes cv contractile vacuole mt microtubule cyt cytopharynx n nucleus f flagellum pr paraxial rod fp flagellar pocket rt rootlet fv food vacuole ves vesicle REFERENCES ALEXEIEFF, A. 1912. Sur la revision du genre Bodo Ehrbg. Arch. Protistenk. 26 : 413-419. Anverson, W. A. & Erris, R. A. 1965. Ultrastructure of Trypanosoma lewisi: flagellum, microtubules and the kinetoplast. J. Protozool. 12 : 483-499. Baker, J. R. 1963. Speculations on the evolution of the family Trypanosomatidae Doflein, 1got. Expl. Parasit. 13 : 219-233. Brooker, B. E. 1965. Mastigonemes in a bodonid flagellate. Expl. Cell Res. 37 : 300-305 1971. The fine structure of Crithidia fasciculata with special reference to the organelles involved in the ingestion and digestion of protein. Z. Zellforsch. 116 : 532-563. Git, J. W. & VocEL, H. J. 1962. Lysine synthesis and phylogeny: biochemical evidence for a bacterial-type endosymbiote in the protozoon Herpetomonas (Strigomonas) oncopelti. Biochim. biophys. Acta 56 : 200-201. GravErtT, A.M. 1962. Fine structure of bacteria. By. med. Bull. 18 : 245-250. Hi1t, G. C. & ANDERSon, W. A. 1969. Effects of acriflavine on the mitochondria and kineto- plast of Crithidia fasciculata. Correlation of fine structural changes with decreased mito- chondrial enzyme activity. J. Cell Biol. 41 : 547-561. Hoare, C. A. 1948. The relationship of the haemoflagellates. In Proceedings of Fourth International Congress of Tropical Medicine. 2 : 1110-1116. HotianpeE, A. 1936. Sur la cytologie d’un flagellé du genre Bodo. C.r. Séanc. Biol. 123: 651-653. 1942. Etude cytologique et biologique de quelques flagellés libres. Archs. Zool. exp. gén. 83 : 1-268. Hotwitt,M.E.J. 1966. Physical aspects of flagellar movement. Physiol. Rev. 46 : 696-785. Joyon, L. & Lom, J. 1966. Sur l'ultrastructure de Costia necatrix Leclercq (Zoofiagellé); place systématique de ce Protiste. C.v. Acad. Sci. (Paris) 262 : 660-663. 1969. Etude cytologique, systématique et pathologique d'Ichtyobodo necator (Henneguy, 1883) Pinto, 1928 (Zooflagelle). J. Protozool. 16 : 703-710. KELLENBERGER, E. & RyTeR, A. 1958. Cell wall and cytoplasmic membrane of Escherichia coh. J. biophys. biochem. Cytol. 4 : 323-326. 102 B. E. BROOKER Kirspy, H.W. 1941. In Protozoa in Biological Research. Eds. Calkins, G. N. and Summers, F. N. New York. KusEL, J. P., Moore, K. E. & WEBER, M.M. 1967. The ultrastructure of Crithidia fasciculata and morphological changes induced by growth in acriflavin. J. Protozool. 14 : 283-206. Levy, M. R. & Extiotr, A. M. 1968. Biochemical and ultrastructural changes in Tetva- hymena pyriformis during starvation. J. Protozool. 15 : 208-222. Micnot, J. P. 1965. Etude ultrastructurale de Cyathomonas tvuncata From. (Flagellé Cryptomonadine). J. Microscopie 4 : 239-252. Newton, B. A. & Horne, R. W. 1957. Intracellular structures in Stvigomonas oncopelti 1. Cytoplasmic structures containing ribonucleoprotein. Expl. Cell. Res. 13 : 563-574. Piretka, D. R. 1961. Observations on the kinetoplast-mitochondrion and the cytostome of Bodo. Expl. Cell Res. 25 : 87-93. Preston, T. M. 1969. The form and function of the cytostome-cytopharynx of the culture forms of the elasmobranch haemoflagellate Tyypanosoma vaiae Laveran and Mesnil. J. Protozool. 16 : 320-333. Pyne, C. K. 1960. L’ultrastructure de l’appareil basal des flagelles chez Cryptobia helicis (Flagellé, Bobonidae). C.v. Acad. Sci. (Paris) 250 : 1912-1914. Ropertson, M. 1929. The action of acriflavine upon Bodo caudatus. Parasitology 21 : 375- 410. Ruopzinska, M. A. & VICKERMAN, K. 1969. The fine structure. In Infectious blood diseases of man and animals. New York 1 : 217-306. ScuustErR, F. L. 1968. The gullet and trichocysts of Cyathomonas truncata. Expl. Cell Res 49 : 277-284. Stmpson, L. 1968. Behaviour of the kinetoplast in Leishmania tarentolae on cell rupture. J. Protozool. 15 : 132-136. Sinton, J. A. 1912. Some observations on the morphology and biology of Prowazekia uvinarvia (Bodo urinarius, Hassall). Ann. trop. Med. Parasit. @ : 245-269. STEINERT, M. 1960. Mitochondria associated with the kinetonucleus of Trypanosoma mega. J. biophys. biochem. Cytol. 8 : 542-546. STEINERT, M. & Novikorr, A. B. 1960. The existence of a cytostome and the occurrence of pinocytosis in the trypanosome Trypanosoma mega. J. biophys. biochem. Cytol. 8 : 563-569. ViIcKERMAN, K. 1969. ‘The fine structure of Trypanosoma congolense in its bloodstream phase. J. Protozool. 16 : 54-69. WuitmoreE, E. 1911. Prowazehkia asiatica (syn. Bodo asiaticus Castellani and Chalmers). Arch. Protistenk. 22 : 370-378. B. E. BRooKER NUFFIELD INSTITUTE OF COMPARATIVE MEDICINE THE ZOOLOGICAL SOCIETY OF LONDON REGENT’S PARK Lonpon, N.W.1 >" PLATE 1 Fic. A. Longitudinal section through the anterior end of B. caudatus. The buccal cavity is seen in transverse section. A bacterium in an early stage of ingestion occupies the greatly distended buccal cavity. Note the dense pad (arrowheads) lying between the basal body of the flagellum and the kinetoplast capsule. 29,500. Fic. B. Longitudinal section through the basal plate region of two flagella showing the striated interflagellar connective. 96,000. Fic. C. Transverse section of two flagella at the level of the basal plates which are joined by the interflagellar connective. x 54,000. Fic. D. Transverse section of the basal bodies of B. saltans. They are connected by striated rootlets. The rootlet passing from the basal body of the anterior flagellum to the vicinity of the cytopharynx can just be seen (arrow). This cell is in the early stages of division and the daughter basal bodies are already forming. A rootlet connects the basal body of the anterior flagellum with its daughter. x 56,500. Fic. E. Transverse section of the posterior flagellum of B. caudatus showing the axoneme and the paraxial rod. On one side, a flagellar striation is seen as a thickening of the flagellar membrane associated with sub-membrane material. 56,500. Fic. F. Posterior flagellum of B. saltans showing the parallel striations. Metal shadowed. 44,500. Bull. Br. Mus. nat. Hist. (Zool.) 22, 3 PE Age 1 PALINANS, 72 Fic. A. Longitudinal section through the posterior flagellum of B. caudatus showing the striations of the flagellar membrane. ™ 56,500. Fic. B. Transverse section through B. saltans showing the relationship of cytopharynx to the flagellar pocket. Small portions of the circumbuccal lappets are visible. One of the basal body rootlets appears clearly. Note the abundant vesicles lying on one side of the cytopharynx. 29,500. Fic. C. Longitudinal section of the buccal cavity and part of the cytopharynx of B. saltans. Note the pronounced cell coat (arrowheads) of the buccal cavity membrane, the cytoplasmic bacteria and position of the lappets. The anterior end of the flagellate is at the top of the micrograph. 30,000. Fic. D. Longitudinal section of B. caudatus showing the anteriorly directed buccal cavity. The kinetoplast and nucleus lie very close to each other in this species. The anterior end of the flagellate is to the right of the micrograph. 22,000. PAT Bes p 8} Mus. nat. Hist. (Zool.) 22, Bull. Br. PLATE 3 Fic. A. Negatively stained B. saltans showing the position of the circumbuccal lappets relative to the mastigonemes of the anterior flagellum. The position of the flagellum here corresponds to the bottom of the effective stroke. 28,500. Fic. B. Section through the flagellar pocket of B. saltans demonstrating the pairing of microtubules shortly before they enter the buccal cavity. ™ 37,500. Fic. C. Longitudinal section of B. saltans showing the undulatory path of the cytopharynx. The Golgi apparatus lies directly beneath this organelle. Note the numerous vesicles associated with the cytopharynx and the undulation of the cytopharyngeal membrane. A band of micro- tubules circumscribes the floor of the buccal cavity. ™ 45,000. Fic. D. Transverse section of the cytopharynx of B. saltans showing eight microtubules of which three are paired. Note the vesicles lying beside it. 66,000. Fic. E. Transverse section of the cytopharynx of B. saltans showing that it is occasionally surrounded by nine microtubules. 40,000. Bull. Br. Mus. nat. Hist. (Zool.) 22, 3 BE ARES PLATE 4 Fic. A. Tangential section of the cytopharynx in B. saltans. Note the connectives joining adjacent tubule pairs. » 56,700. Fic. B. Longitudinal section of B. saltans which passes through the flagellar pocket and cytopharynx transversely. The microtubule dipping below the surface of the cell is one of many which eventually pass under the buccal cavity as shown in Fig. C. Note the position of the cytopharynx and the vesicles lying one one side of it. 30,000. Fic. C. Transverse section of the buccal cavity of B. saltans. Note the tract of microtubules passing under the floor of this organelle. 30,000. Fic. D. Longitudinal section of B. caudatus which passes through the buccal cavity and flagellar pocket transversely. The contractile vacuole lies next to the flagellar pocket. Micro- tubules destined to surround the cytopharynx are arranged in a row in this section. Note the row of tubules associated with the opposite wall of the buccal cavity. » 40,000. Bull. By. Mus. nat, Hist. (Zool.) 22, 3 PLATE 4 PLATE 5 Fic. A. Transverse section of the buccal cavity posterior to that shown in Pl. 4, Fig. D. The microtubules associated with the buccal cavity (between arrows) progressively move away from it. ™ 50,000. Fic. B. Transverse section through the beginning of the cytopharynx of B. caudatus. The microtubules are arranging themselves around its walls. Microtubules associated with the cytopharynx are the same set as shown in the previous figure (between arrows). 50,000. Fic. C. Section of the mitochondrion showing the electron dense bodies which are sometimes found in the matrix. 24,000. Fic. D. Longitudinal section through B. saltans showing the mitochondrion passing to one side of the buccal cavity before travelling to the posterior end of the cell. Other features include the nucleus, food vacuoles and Golgi apparatus. ™ 15,500. Bull. Br. Mus. nat. Hist. (Zool.) 22, 3 PLATE 5 IPALYNADID, (6) Fic. A. Section through the flat plane of the kinetoplast of B. caudatus. Note that the kinetoplast is composed of a reticulum of fine filaments with electron dense nodes interspersed between them. The matrix of the mitochondrion is visible at the periphery of the kinetoplast. 40,000. Fic. B. Contractile vacuole and spongiome of B. saltans. ™ 24,000. Fic. C. Cytoplasmic bacterium of B. saltans in division. 46,000. Bull. Br. Mus. nat. Hist. (Zool.) 22, 3 f ie PLATE 6 we he ie, A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) Kay, E. Atison. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 1965. (Out of Print.) £3.75. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180; 11 Plates, 15 Text-figures. 1967. {4. TayLor, J. D., KENNEDY, W. J. & Hatt, A. The Shell Structure of Mineralogy at the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates, 77 Text-figures. 1969. £4.50. Haynes, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur) 1962-1964. (In press.) J Bt Printed in England by Staples Printers Limited at their Kettering, Northants, establishment THE TYPE SPECIMENS AND IDENTITY OF ‘THE SPECIES DESCRIBED IN THE GENUS PITHOBIOS BY C.L. KOCH AND L. KOCH FROM 1841 TO 1878 (CHILOPODA : LITHOBIOMORPHA) E. H. EASON Cc BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 22 No. 4 LONDON : 1972 THE TYPE SPECIMENS AND IDENTITY OF THE SPECIES DESCRIBED IN THE GENUS LITHOBIUS BY C. L. KOCH AND L. KOCH FROM 1841 TO 1878 (CHILOPODA : LITHOBIOMORPHA) BY EDWARD HOLT EASON Bourton Far Hill, Moreton-in-Marsh, Gloucestershire Pp. 103-150; 7 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 22 No. 4 LONDON : 1972 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), ‘instituted in 1949, 1s 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. 22, 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), 1972 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 3 March, 1972 Price £1.40 THE TYPE SPECIMENS AND IDENTITY OF THE SPECIES DESCRIBED IN THE GENUS LITHOBIUS BY C.L. KOCH AND L. KOCH FROM 1841 TO 1887 (CHILOPODA : LITHOBIOMORPHA) By E. H. EASON INTRODUCTION Cart Lupwic Kocu (C. L. Koch) described fifteen nominal species of Lithobius. His son, Dr. Ludwig Carl Christian Koch (L. Koch) redescribed most of his father’s species and described thirty-two more and one variety. Subsequent authors have placed various interpretations on L. Koch’s work, Latzel (1880) and Haase (1880) giving the earliest detailed and accurate accounts of what they considered to be the Kochs’ western European species. These two authors, who were in correspondence with L. Koch and had access to some of his material, are in general agreement with one another and Latzel is universally recognized as the first authoritative reviser of both C. L. Koch’s and L. Koch’s species, although he dealt only with those found in the Austro-Hungarian Monarchy. But in spite of Latzel’s and Haase’s work there is still uncertainty as to the identity of some of these species whereas most of those originally described by L. Koch from Greece and from Tinos in the Aegean Archipelago have never been revised and their identity has not hitherto been established. Carl Koch gave a very brief account of the Koch Collection of Arachnida and Myriapoda (C. Koch, rgro) but he did not mention any species by name and merely enumerated those of each class attributable to C. L. Koch and to L. Koch. I have recently been able to examine the specimens of Lithobius in this Collection, the bulk of which is preserved in the British Museum (Natural History) and the remainder in the Zoological Museum, Berlin. It includes the original material of two of C. L. Koch’s and twenty-three of L. Koch’s species together with the specimens of ten of C. L. Koch’s on which L. Koch seems to have based his redescriptions. The type specimens and identity of L. grossipes C. L. Koch and L. litoralis L. Koch have been discussed in a previous publication (Eason, 1970a) and in the present paper an attempt is made to determine the identity of the other species. SPECIES DESCRIBED BY C. L. KOCH The original descriptions of these species (1841, 1844, 1847) are very inadequate by modern standards, relying to a large extent on details of colour and other super- ficial features and omitting many of the characters now recognized as of taxonomic importance. Although habitat is mentioned, type localities are indefinite. In a later publication (C. L. Koch, 1863) amplified descriptions, coloured plates and line drawings are provided but in only a few cases are they of any real value. L. Koch 106 E. Ho EASON (1862) was the first to redescribe these species more adequately and it is reasonable to assume that he interpreted most of his father’s original descriptions correctly. According to C. Koch (1910) only a small minority of C. L. Koch’s species of Myriapoda are represented in the Koch Collection by type specimens and L. Koch did not use the original material of his father’s species of Lithobius for many of his redescriptions. Only L. grossipes and L. melanocephalus seem to have been redes- cribed from type specimens, both of which are present in the Collection. Of the other species, eleven were almost certainly redescribed from specimens L. Koch either collected himself, mostly from Nuremberg and the surrounding Franconian Jura, or had sent him by other collectors: all these specimens except those of L. impressus are present in the Collection. The remaining species, L. glabratus and L. varius, were unknown to L. Koch. The original material belonging to all but two of C. L. Koch’s species had probably already been either lost or badly damaged before 1862. The fact that C. L. Koch’s later descriptions and illustrations did not appear until 1863 is no evidence of the continued existence of these specimens; the author died in 1857 so his book must have been compiled from earlier work and suffered delay in publication. It is therefore necessary to select neotypes from L. Koch’s material for six of C. L. Koch’s species whose identity needs to be established in order to ensure stability of nomenclature. SPECIES DESCRIBED BY L. KOCH The original descriptions of most of these species (1862, 1867, 1878) are fairly adequate. The number of antennal articles, prosternal teeth, coxal pores, tergal projections, and ventral spines on the fifteenth legs are all recorded, the ocelli are figured in many cases and the female gonopods are described where females were available. A notable omission, however, is any mention of coxolateral spines (VaC) or accessory apical claws on the fifteenth legs. Fairly definite type localities are given for most of the species but type specimens are not designated. Of the twenty-three species and one variety represented by original material in the Koch Collection, eight were based on single specimens (holotypes) and sixteen on more than one specimen (syntypes): it is necessary to select lectotypes from only seven of the latter in order to ensure stability of nomenclature. Three further species are represented in the Collection by specimens apparently identified by L. Koch but which do not seem to belong to the original material. Two of these together with a further six which are not represented were all originally described from borrowed specimens which may never have formed part of the Collection. The chiet source of these specimens was the collection of Graf von Keyserling of Munich. Part of the latter, consisting largely of insects, has been acquired by the British Museum (Natural History) but all attempts to trace the specimens of Lithobius both here and in the principal museums of Germany and Austria have failed. METHODS All the species described in the genus Lithobius by C. L. Koch and L. Koch are recorded in order of their dates of publication whether or not there is any material available for examination. Where more than one description applying to the THE LITHOBIUS SPECIES OF C. L. AND L. KOCH 107 same zoological species occurs in the same work, whichever name Latzel (1880), their first reviser, believed to attach to the best description is regarded as having priority even though it may appear on a later page (Code, article 24a). All the specimens in the Koch Collection labelled with the name of a species of Lithobius originally described either by C L. Koch or L. Koch have been examined. They are all preserved in spirit, each tube of specimens containing labels bearing the identity, usually the locality and sometimes the habitat and name of the collector. The labels belonging to the specimens in the Zoological Museum, Berlin have all been rewritten but at least one label in each of the tubes in the British Museum (Natural History) appears to be in L. Koch’s hand. In some instances L. Koch placed specimens of different species bearing a super- ficial resemblance to one another under the same name. Although he made rel- atively few mistakes over males which usually have the most characteristic features, he had difficulty in identifying females of similar species. For example, he failed altogether to recognize females of L. muticus C. L. Koch, placing nearly all of them with L. mutabilis L. Koch and identifying as L. muticus females of L. pelidnus Haase. On the other hand he never placed widely different stadia of the same species together and regarded many of them as taxonomically distinct. In the present study each specimen is recorded under the name given it by L. Koch but those he misdetermined are also given their correct identity and placed in separate tubes, retaining their original registered numbers. All labels are recorded exactly as written by Koch. Where they have been rewritten by some recent museum worker this is indicated: where they are difficult to interpret extra words of explan- ation are inserted and enclosed in square brackets. Descriptions of specimens are only given when some special feature or aberration requires emphasis, or when an adequate account of the species in question is not to be found in the literature; otherwise reference is made to a published description. When it is necessary to tefer to an immature post-larval stadium, Verhoeff’s (1905) terms are used for the larger species such as L. forficatus in which the life-history is well-known, but for the smaller species to which Verhoeff’s terms cannot be strictly applied, the specimen are allotted to stadia corresponding to those described for L. variegatus Leach (Eason, 1964). Selected neotypes and lectotypes are labelled as such and placed in separate tubes. Selection of the former presents no difficulty but some of the syntypical series of L. Koch’s species for which lectotypes are selected consist of specimens belonging to more than one zoological species; here a lectotype is selected from the specimens answering most closely to the original description of the species in question, or where there is nothing to choose between them in this respect, from those belonging to the zoological species to which it is desirable to attach Koch’s name in order to cause the least confusion in current nomenclature. The originally published type localities of most of C. L. Koch’s and many of L. Koch’s species are vague or equivocal: in these cases the designate type locality is that of the type specimen. Conclusions as to the status and present generic classification of all the species described by C. L. Koch and L. Koch in the genus Lithobius together with their published and designate type localities, are summarised in Table 1. 108 E. H. EASON 1. Lithobius impressus C. L. Koch Lithobius impressus C. L. Koch, 1841 : 224. 1863, 1: 115, fig. 105a, b&c. L. Koch, 1862 : 36, fig. 7a & b. Type Locatity. Algerian coast. yy 66 MATERIAL EXAMINED. “‘L. impressus [rewritten] “Corsica, [/eg.] E. Simon” B.M.(N.H.) Reg. no. 13.6.18.326-340. Fifteen specimens which answer to the original description of Lithobius impressus corsicus Léger and Duboscq, 1903. REMARKS. Since the type locality is Algeria and L. Koch’s description was based on specimens from Algiers and Oran borrowed from the Kyeserling Collection, none of the above specimens from Corsica can be selected as neotype. L. Koch must have identified them after finishing his book without attaching any significance to the hooked spine (DpP) on the 14th male prefemur which is characteristic of subsp. corsicus (Léger & Duboscq, 1903: 316, fig. 1) but never found in the North African form. Silvestri (1897) regarded L. nudicornis Gervais as the correct name for this species but most authors reject mudicornis which was described very scantly from a Sicilian specimen (Gervais, 1837). On the other hand impressus is also rejected by many authors in favour of L. elongatus Newport which was published some years later (Newport in Lucas, 1849). However, quite apart from the fact that impressus takes precedence over elongatus there are good reasons for supposing that these names apply to two distinct subspecies (Eason, 1971) and the valid name for the common Algerian form, which belongs to the genus Ewpolybothrus Verhoeff and the subgenus Allopolybothrus Verhoeff as emended by Jeekel (1967), is Eupolybothrus (Allopolybothrus) impressus impressus (C. L. Koch). 2. Lithobius dentatus C. L. Koch Lithobius dentatus C. L. Koch, 1844 : 22, fig. 22. 1847: 148. 1863, 1: 117, fig. 106a, b&c. L. Koch, 1862: 54, fig. 18. TYPE LOCALITY. Germany. MATERIAL EXAMINED. “‘Lithobius dentatus C. L. Koch, Nirmberg, leg. L. Koch [rewritten] Zool.Mus.Berlin: Kat.Nr.333. A male and a female, both mutilated. “Lithobius dentatus C.K.” “Eichstaedt, Happurg, Ntirmberg” B.M.(N.H.) Reg. no. 13.6.18.130-136. Six more or less mutilated specimens. “Lithobius dentatus C.K.” “Kamthen [Carinthia, Austria]’’ B.M.[N.H.] Reg. no. 13.6.18.137. Two males and a temale in fair condition. “Lithobius dentatus C.K.” “Meran [Merano, Italy], [/eg.] Milde” B.M.(N.H.) Reg. no. 13.6.18.138. A mutilated male. TYPE SPECIMEN. The best preserved specimen from Germany, a male 12 mm long answering to Latzel’s (1880 : 81) description of L. dentatus var. alpestris, is here formally designated as the neotype (B.M.(N.H.) 13.6.18.135). Of the localities given for this specimen, Happurg is 25 km. from Nuremburg and Eichstatt (= Eich- THE LITHOBIUS SPECIES OF C. L. AND L. KOCH 109 staedt) is only 60 km. distant, so the designate type locality may be given as ““Nurem- berg district”. REMARKS. Since neither C. L. Koch nor L. Koch mentioned the dorsal sulci on the 14th and 15th male tibiae, Latzel believed he had discovered a new variety characterized by these sulci which he called alpestris. However, the tibial sulci are quite distinct in the neotype and in the only other males in the Collection with the 14th and 15th legs intact ; they were also noted by Haase (1880 : 24) as characteristic of the species so it is clear that they were merely overlooked by the Kochs and var. alpestris should be disregarded. 3. Lithobius calcaratus C. L. Koch Lithobius calcavatus C. L. Koch, 1844 : 23, fig. 23. 1863, 2: 45, fig. 168 a&b. L. Koch, 1862 : 70, fig. 30. TYPE LOCALITY. Germany. MATERIAL EXAMINED. “‘Lithobius calcaratus C. L. Koch, Franconia (Jura), leg. L. Koch [rewritten] Zool. Mus. Berlin: Kat. Nr. 337. Four males and a female. “Lithobius calcaratus C.K.” ‘‘Niirnberg’’ B.M.(N.H.) Reg. no. 13.6.18.18-33. Eight males and eleven females. “Lithobius calcaratus C.K.” “frank. Jura’’ B.M.(N.H.) Reg. no. 13.6.18.34-44. Seven males and a female together with three females ot L. pelidnus Haase. “Lithobius calcaratus C.K.” ‘“Miinchen’” B.M.(N.H.) Reg. no. 13.6.18.45. A female of L. muticus C. L. Koch. “Lithobius calcaratus C.K.” ‘Rom, [{/eg.] Seidlitz’’ B.M.(N.H.) Reg. no. 13.6.18. 46-52. Three males and two females together with four females of L. erythrocephalus C. L. Koch. “calcaratus? Niirnberg” B.M.(N.H.) Reg. no. 13.6.18.53-55 (part). A female each of L. muticus, L. mutabilis L. Koch and L. pusillus pusillifrater Verhoeft. “Lithobius calcaratus C.K.” ““Niirmberg, missbildung der Analbeine’”’ B.M.(N.H.) Reg. no. 13.6.18.53-55 (part). A male with the left 15th leg missing and the right 15th leg imperfectly regenerated without a femoral process. TYPE SPECIMEN. A well-preserved male 10-5 mm long from Nuremberg answering to Latzel’s (1880 : 105) description of L. calcaratus is here formally designated as the neotype (B M.(N.H.) 13.6.18.18). Remarks. All the above males, which are either fully mature or 4th post-larval stadia with the femoral process on the 15th leg at least partly developed, are cor- rectly labelled as we would expect, since they are easily identified from C. L. Koch’s original description and illustration; but of the twenty-six females L. Koch labelled “L. calcaratus’’, no fewer than eleven were misdetermined. In fact, females of this species are quite easy to identify owing to the characteristic arrangement of the ocelli and the spinulation of the legs, and Koch may have been misled by colour, a rather variable feature to which he paid undue attention in his descriptions. In one of the males from the Franconian Jura (B.M.(N.H.) 13.6.18.41) both 15th r10 E. H. EASON legs are missing but a femoral process is present on the right 14th leg. This last character was used by Matic (1961) to define Lithobiws lanzae Matic which therefore seems to be based on an aberrant specimen of L. calcaratus. 4. Lithobius communis C. L. Koch Lithobius communis C. L. Koch, 1844 : 24, fig. 24. 1863, 2: 47, fig. 169a&b. L. Koch, 1862 : 80, fig. 37. TYPE LOCALITY. Germany. MATERIAL EXAMINED. “‘Lithobius communis C. L. Koch, Nurnberg, leg. L. Koch [rewritten] Zool. Mus. Berlin: Kat. Nr. 341. A male and a female, both mutilated 3rd post-larval stadia of L. mutabilis L. Koch. “Lithobius communis C.K.” “frank. Jura’ B.M.(N.H.) Reg. no. 13.6.18.64-86. Twelve immature males of L. mutabilis, four being 2nd post-larval stadia and eight being 3rd post-larval stadia. In addition there are four males and three females of L. pusillus pusillifrater, three immature males of L. pelidnus and two immature males of L. muticus. “Lithobius communis C.K.” “‘[hab.] Haspelmooz’’ B.M.(N.H.) Reg. no. 13.6.18.87. A mutilated female 3rd post-larval stadium of L. mutabilis and a female 4th post- larval stadium probably belonging to L. lapidicola Meinert (sensu Jeekel, 1964 non Latzel, 1880). “Lithobius communis C.K.” “Cusel [Rhineland Palatinate]” B.M.(N.H.) Reg. no. 13.6.18.88. A mutilated male 3rd post-larval stadium of L. mutabilis. REMARKS. Of the above specimens, only the 3rd post-larval stadia of L. mutabilis agree exactly with L. Koch’s description of L. communis. All that needs to be added to this description is that the ventral spine on the 15th tibia is VaT, an accessory apical claw is present on the 15th leg, and T.13 bears feeble posterior projections. Latzel (1880 : 102) suggested that L. Koch’s description was based on immature specimens of L. mutabilis with the possible inclusion of males of L. muticus and L. pelidnus ; he added that he had actually examined two of L. Koch’s specimens of “communis” and found them to be immature males of L. mutabilis and L. pelidnus. However, neither the specimens of muticus nor those of pelidnus labelled “communis” by L. Koch have either the full complement of three ventral femoral spines or the single ventral tibial spine on the 15th leg as recorded by him for this form; nor are these spines present in L. pusillus pusillifratey in which the antennae are much shorter than in the example of L. communis illustrated by C. L. Koch (1863: fig. 16ga). This last illustration does, in fact, resemble L. mutabilis more than any other species. Latzel’s tentative suggestion that L. communis C. L. Koch, 1844 is a synonym of L. mutabilis L. Koch, 1862 is therefore almost certainly justified and has been accepted by Haase (1880) and subsequent authors. But the name L. communis has not actually been used except as a junior synonym since Rosicky (1876) redescribed the species, and its revival would only cause confusion. It is intended therefore to ask the International Commission on Zoological Nomenclature to use its plenary powers THE LITHOBIUS SPECIES OF C. L. AND L. KOCH III to supress the name communis C. L. Koch 1844 as published in the binomen Lithobius communis C. L. Koch, and to supress the name minutus C. L. Koch 1847 as published in the binomen Lithobius minutus C. L. Koch (see p. 118), as so to validate Lithobius mutablis L. Koch. 5. Lithobius grossipes C. L. Koch Lithobius grossipes C. L. Koch, 1847 : 146. 1863, 1 : 67, fig. 57a, b& c. L. Koch, 1862 : 32, fig. 4. TYPE LOCALITy. Triest. TYPE SPECIMEN. The holotype. B.M.(N.H.) Reg. no. 13.6.18.262. Remarks. All the specimens identified as L. grossipbes by L. Koch are in the British Museum (Natural History) and were discussed in a previous publication (Eason, 1970a). Reasons for believing that a female from Idrija is the holotype were given and it was shown that Eupolybothrus grossipes (C. L. Koch) is a valid species and not a synonym of E. fasciatus (Newport) as was previously supposed. 6. Lithobius punctulatus C. L. Koch Lithobius punctulatus C. L. Koch, 1847: 147. 1863, 1: 68, fig. 58a & b.? L. Koch, 1862 : 30, fig. 3. TYPE LOCALITY. Triest. MATERIAL EXAMINED. “‘L. punctulatus [rewritten]’’ ““Dalmatien’’ B.M.(N.H.) Reg. no. 13.6.18.586. A mutilated female of Eupolybothrus sp. 25 mm long with no legs and broken antennae. “L, punctulatus [rewritten]” “Griechenland’”’ B.M.(N.H.) Reg. no. 13.6.18.587— 588. Two-cleared fragments of Eupolybothrus sp. with neither legs nor antennae, one from the head to T.9, the other from T.4 to the end of the body. Although obviously from different individuals, these fragments both seem to belong to the same species. Remarks. L. Koch’s account of L. punctulatus was based on examples from Dalmatia and Greece with all their legs missing so there is little doubt that the above specimens are those in question. This account describes the tergal projections as short and broad which led Latzel (1880 : 56) to suspect that L. Koch was describing examples of Eupolybothrus leptopus (Latzel): but the shape of these projections in all three specimens is quite consistent with a diagnosis of one of the fasciatus- grossipes group of species. The Dalmatian specimen, however, has II, 19, 21 and 21 coxal pores on the 12th to 15th legs respectively and this reduced number of pores, particularly those of the 12th, relative to those found in most examples of E. grossipes and related species of comparable size, is indeed rather suggestive of E. leptopus. On the other hand the fragment from Greece (with 24, 30, 32 and 27 coxal pores) is, in view of its locality, more likely to belong to E. litoralis (L. Koch), a species very close to E. grossipes. 112 E. H. EASON C. L. Koch’s descriptions and illustrations of L. punctulatus are difficult to inter- pret; there is no special reason to suppose that they apply either to E. leptopus or E. litoralis. Meinert (1872) used the name L. punctulatus C. L. Koch (probably correctly) to apply to E. grossipes but Latzel (1880 : 52) argued that C. L. Koch’s original (1847) and subsequent (1863) descriptions of punctulatus do not apply to a species of Eupolybothrus at all but to Lithobius validus Meinert, 1872. Latzel’s opinion has been accepted by some authors although Meinert’s name for the latter species continues to be used by others. Owing to the uncertainty surrounding the identity of C. L. Koch’s original specimen and of those L. Koch used for his re- description, L. punctulatus should be rejected as a nomen dubium and the species with which Latzel equated it should be known as Lithobius validus Meinert. 7. Lithobius montanus C. L. Koch Lithobius montanus C. L. Koch, 1847 : 148. 1863, 2: 8, fig. 132a&b. L. Koch, 1862 : 27, fig. 1. TYPE LocaLity. South Tyrol. MATERIAL EXAMINED. ‘“‘Lithobius grossipes C.K.” “‘Seiseralpe [an alpine hut in Italy], [/eg.] Gredler” B.M.(N.H.) Reg. no. 13.6.18.266. A male of Eupolybothrus grossipes 35 mm long, agreeing in detail with L. Koch’s description of L. montanus. Remarks. L. Koch’s rediscription of this form was based on a single male sent him by Prof. P. Gredler from “‘Seiseralpe’’ in South Tyrol so the above specimen is undoubtedly the one in question. C. L. Koch in his original (1847) and subsequent (1863) accounts of L. montanus described the colour as uniform reddish-brown, and his coloured plate (1863: fig. 132a) is of a pale brown specimen without the dark dorsal pattern he illustrated in a comparable coloured plate of L. grossipes (1863: fig. 57a). L. Koch described the colour of L. montanus as paler anteriorly than posteriorly, making no mention of a dark pattern, and it seems that this relatively pale colour together with a trivial structural aberration in the holotype of L. gros- sitpes mentioned in his key led him to copy his father in supposing that montanus and grossipes were distinct species. Sometime between finishing his book and completing his collection he must have decided to discard these two characters as a means of differentiating species, changing the name of his specimen from montanus to grossipes. Latzel (1880 : 48), while recognizing L. montanus as a synonym of L. grossipes, pointed out that it differs in colour from his own specimens, and Dalla Torre (1882) and Attems (1929) both retained the name montanus for a pale variety of L. grossipes. Although the holotype of L. grosstpes has been dried and the original colour pattern had probably been lost before it was examined by L. Koch, many preserved specimens of L. grossipes show more evidence of a dark pattern than does L. Koch’s specimen of montanus. However, colour is a poor taxonomic character in the Lithobiidae and there is no justification for regarding L. montanus as other than a synonym of L. grossipes. THE LITHOBIUS SPECIES OF C. L. AND L. KOCH 113 8. Lithobius glabratus C. L. Koch Lithobius glabratus C. L. Koch, 1847 : 149. 1863, 1 : 131, fig. 121a & b. TYPE LOCALITY. Bavaria. Remarks. L. Koch (1862) was not familiar with this species and there is no material referable to L. glabratus in the Koch Collection. Latzel, however, was satisfied as to its identity and gave a full description (Latzel, 1880 : 74). Pocock (1890), after examining the type specimen of L. melanops Newport, 1845, proposed L. glabratus as a synonym and this species is now universally known as L. melanops Newport (see also Eason, 1971). 9. Lithobius agilis C. L. Koch (Fig. 1) Lithobius agilis C. L. Koch, 1847: 149. 1863, 1 : 132, fig. 122a&b. L. Koch, 1862: 52, fig. 17. TYPE Locality. Bavaria. MATERIAL EXAMINED. “‘Lithobius agilis C. L. Koch, Nurnberg, leg. L. Koch [rewritten]’’ Zool. Mus. Berlin: Kat. Nr. 334. A male and a female, both mutilated. “Lithobius agilis C. Koch” “Mégeldorf, [hab.] Erlenwalddren” B.M.(N.H.) Reg. no. 13.6.18.4-8. Three males and two females in fair condition. TyPE SPECIMEN. A fairly well-preserved female 9-5 mm long from Mogeldorf near Nuremburg answering to Latzel’s (1880 : 78) description of L. agzlis is here formally designated as the neotype (B.M.(N.H.) 13.6.18.4). Remarks. The spurs on the gonopods of the neotype (Fig. 1) and the only other female in the Collection with intact gonopods (Kat. Nr. 334) are just as slender as those figured by Loksa (1948: fig. 3) as characteristic of L. agilis pannonicus Loksa from Hungary, although they are rather less expanded in the distal one third with the extremity less obviously serrate. Neither L. Koch nor Latzel figured these spurs but they both described them as long and slender. Loska may have been misled into assuming that the typical form of the species bears relatively short, stout spurs by Brolemann’s (1930: fig. 429) figure of a specimen of L. agilis from the Pyrenees in which the external spur is barely three times longer than broad. The spurs of specimens of this species from Austria (Eason, 1964: fig. 414) are intermediate in shape between those figured by Brolemann and those of the neotype, so there is little justification for naming a subspecies on the basis of this character. to. Lithiobus curtipes C. L. Koch Lithobius curtipes C. L. Koch, 1847: 150. 1863, 2:7, fig. 131a&b. L. Koch, 1862 : 68, fig. 29. TYPE LOCALITY. Bavaria. MATERIAL EXAMINED. “‘Lithobius curtipes C. L. Koch, Ntrnberg, leg. L. Koch 114 E. H. EASON {rewritten} Zool. Mus. Berlin: Kat. Nr. 339. Four males and two females together with a male of L. aeruginosus L. Koch. “Lithobius curtipes C. Koch” “Niirnberg, [hab.] Haspelmooz”’ B.M.(N.H.) Reg. no. 13.6.18.111-121. Four males and four females together with a male and a female of L. aeruginosus and an immature male of L. crassipes L. Koch. “curtipes, Franzensbad [Frantiskovy Lazne, Czechoslovakia] B.M.(N.H.) Reg. no. 13.6.18.122. A single female. “Lithobius curtipes C. Koch” “Lithauen [Lithuania] B.M.(N.H.) Reg. no. 13.6.18. 123-126. Three males and a female. “curtipes?, Nirnberg’”’ B.M.(N.H.) Reg. no. 13.6.18.127-128. Two immature males. “L. curtipes?, Bohmen [Bohemia] [rewritten]’’ B.M.(N.H.) Reg. no. 13.6.18.633- 636. Two males and two females. TYPE SPECIMEN. A well-preserved male 9 mm long from Nuremberg answering to Latzel’s (1880 : 130) description of L. curtipes is here formally designated as the neotype (B.M.(N.H.) 13.6.18.111). REMARKS. There is no doubt at all that the above specimens, other than the three examples of L. aeruginosus and the single one of L. crassipes, belong to the species known to western European authors as L. curtipes: the 15th tibial projection in the neotype and the other adult males is the same as that found in British speci- mens (Eason, 1951: fig. 1). The inclusion by L. Koch of examples of L. aeruginosus among those of L. curtipes may be accounted for by the fact that he based his original description of L. aevuginosus on immature males (one of them actually belonging to L. curtipes) and was unaware of the true nature of this species. In order to understand the controversy surrounding the identity of L. curtipes it is necessary to consider the different interpretations placed by various authors on L. Koch’s description. Since C. L. Koch only examined females he did not mention the 15th tibial projection, which is only found in males, in either of his descriptions, but L. Koch described it as “einen kurzen kegelformigen Forsatz.’’ This is both imprecise and misleading, since the projection barely assumes a conical shape even when fully developed; but L. Koch also described and figured the arrangement of the ocelli (L. Koch, 1862: fig. 29) which is fairly characteristic. Stuxberg (1871 :501) described the tibial projection rather more accurately as “en tydligt utskjutande vundad process” (a distinct projecting rounded process) as well as describing the arrangement of the ocelli. Confusion began when Meinert, after examining the specimens on which Porat (1869) quite correctly based his records of L. curtipes, noted that the ocelli were not arranged exactly as figured by L. Koch and that the projection was borne on the 5th article of the 15th leg and not on the 4th as L. Koch stated, and assumed that these specimens did not belong to L. curtipes but represented a form of L. crassipes (Meinert, 1872: 341). Meinert also assumed that the projection he found on Porat’s specimens was of a different shape from the conical projection described by L. Koch which he understood to refer to a more clearly differentiated process such as that found on the 15th femur of males of L. calcaratus: but his description of THE LITHOBIUS SPECIES OF C. L. AND L. KOCH 115 this projection — “‘dannedes kun af et fremspringende Hjorne af Leddets Bagrand— en Rende forstattes ind paa Fremragningen’’ (formed only by a projecting corner of the posterior edge of the article, with a groove running -onto the projection) — is the earliest really accurate account in the literature and there is no doubt that Porat’s specimens did belong to L. curtipes. The arrangement of the ocelli in this species can be misleading as they may occur in irregular rows rather than in a rosette (Eason, 1964: 238) and the confusion over numbering the articles of the 15th legs must have been due either to an error on the part of Koch or to a misprint. Stux- berg (1876 : 25) perpetuated Meinert’s mistake and attributed his own earlier (1871) description of L. curtipes to L. crassipes. Both Latzel (1880 : 131) and Haase (1880 : 39) rectified Meinert’s mistake and gave adequate accounts of L. curtipes under its correct name, but they both repeated Koch’s rather misleading expression, “kurzen kegelformigen Forsatz’’, in describing the tibial projection, although Latzel supplemented this by mentioning the groove on the dorsal surface. Porat (1889) described the species correctly but gave no details of the shape of the tibial projection. It was, no doubt, the failure of most of these early western European authors to give really full and accurate accounts of the projection, and the fact that the best description, Meinert’s, was attributed to L. crassipes, that led Muralewitsch (1926) and Loksa (1947) to apply the name L. curtipes C. L. Koch to an eastern European species which does not occur in Bavaria and which was originally described by Sseliwanoff (1880) from the Crimea. This species, L. pusillus Sseliwanoff, is very similar to L. curtipes but the rounded tibial projection is replaced by a small cylindrical spur very much the same in structure as the femoral process found in L. calcaratus, a structure wrongly envisaged by Meinert as occurring in L. curtipes. Sseliwanoff (1880), who wrote in Russian, as well as describing L. pusillus, gave a clearly recognizable account of L. curtipes which he named as a new species, L. vicinus Sseliwanoff. Loksa (1947), who cannot have been familiar with Sseliwanoff’s work, gave another very adequate account of L. curtipes which he named as another new species, L. baloghi Loksa. Some modern eastern European authors have followed Loksa’s nomenclature, naming L. curtipes, which occurs at least as far east as the Caucasus, as L. baloghi, and L. pusillus Sseliwanoff as L. curtipes. Although Sseliwanoff’s paper describing L. pusillus appeared in volume 11 of Trudy Russkago Entomologicheskago Obshchestva, the volume for 1878, it was not actually published until 1880 and Garbowski (1897), assuming the name to be pre- occupied by L. pusillus Latzel, 1880, proposed the new name sseliwanoffi for Sseli- wanoff’s species ; but this proposal has never been followed. L. pusillus Sseliwanoff seems to have been described repeatedly by various authors either as a new species or as a subspecies of L. curtipes. L. ferganensis Trotzina may prove to be its valid name (Lignau, 1914) but Trotzina’s (1893) description is not altogether clear. L. curtipes turkestanicus Attems, the original description of which is accompanied by an illustration of the tibial projection (Attems, 1904: fig. 2), is undoubtedly identical with L. pusillus Sseliwanoff and the species has recently become known as L. turkestanicus Attems, while the true L. curtipes continues to be known as L. baloghi in eastern Europe. 116 E. H. EASON A final point of nomenclatural interest is that L. curtipes C. L. Koch was desig- nated by Verhoeff (1905) as the type species of the subgenus Monotarsobius to which L. turkestanicus also belongs. 11. Lithobius erythrocephalus C. L. Koch Fig. 2 Lithobius erythrocephalus C. L. Koch, 1847: 150. 1863, 2: 22, fig. 145a, b & c. L. Koch, 1862 : 68, fig. 39. TYPE LOCALITY. Bavaria. MATERIAL EXAMINED. “‘Lithobius erythrocephalus C. K.” “Happurg” B.M. (N.H.) Reg. no. 13.6.18.139-141. Two males and a female. “Lithobius erythrocephalus C.K.” “[hab.] Glaishammer Walddren” B.M.(N.H.) Reg. no. 13.6.18.142-151. Nine females together with a female of L. mutabilis. “Lithobius erythrocephalus C.K.” ‘““Bozen [Bolzano, Italy]’’ B.M.(N.H.) Reg. no. 13.6.18.152. A single female. TYPE SPECIMEN. A well-preserved female 13 mm long from Happurg in the Franconian Jura answering to Latzel’s (1880 : 110) description of L. erythrocephalus is here formally designated as the neotype (B.M.(N.H.) 13.6.18.139). REMARKS. A number of subspecies of L. evythrocephalus have been described depending for their definition on the shape of the spurs on the female gonopods, the sculpturing of the male 15th tibiae, the shape of the short tergites, and the number of antennal articles. The genital spurs of the neotype are figured (Fig. 2) and those of the other females recorded above are of much the same shape. The 15th tibiae of the males from Happurg are oval in cross-section, without secondary sexual char- acters. All specimens of both sexes have feeble posterior projections on T.13 and their antennal articles vary from 29 to 31. The neotype and other females all agree with the modern conception of the nom- inate subspecies, but Dobroruka (1962) stated that the males of this subspecies have flattened 15th tibiae. This feature is not shown by the males from Happurg (the de- signate type locality) although other males of L. erythrocephalus in the Collection from the Franconian Jura labelled “‘mutabilis’” by L. Koch (B.M.(N.H.) Reg. no. 13.6.18. 459-460) do have markedly flattened 15th tibiae: it thus seems that this is an unstable character, not associated with any particular subspecies (see also Eason, 1970b). 12. Lithobius muticus C. L. Koch Lithobius muticus C. L. Koch, 1847: 151. 1863, 1: 118, fig. 107a & b. L. Koch, 1862 : 79, fig. 36. TYPE LOCALITY. Bavaria. MATERIAL EXAMINED. “muticus, Niirnberg’” B.M.(N.H.) Reg. no. 13.6.18 532-533. Iwo females of L. calcaratus. “L. muticus [rewritten]? “Eichstaedt, Happurg” B.M.(N.H.) Reg. no. 13.6.18. THE LITHOBIUS SPECIES OF C. L. AND L. KOCH 1h dy) 534-550. Six males of L. muticus, six females of L. pelidnus, and four females and one male of L. mutabilis, “L. muticus [rewritten]” “[hab.] Valzner Weiher” B.M.(N.H.) Reg. no. 13.6.18 551-562. Four males of L. muticus and eight females of L. pelidnus. “L. muticus [rewritten] “Miinchen’”’ B.M.(N.H.) Reg. no. 13.6.18.536-566. Two males and a female of L. muticus and a female of L. mutabilis. TYPE SPECIMEN. A male 13 mm long from Eichstatt or Happurg, both in the Franconian Jura, is here formally designated as the neotype (B.M.(N.H.) 13.6.18. 534). This is ‘the largest and best preserved available specimen of L. muticus, showing the characteristic broad head and the small setose swelling on the 14th tibia, and agreeing in all respects with Latzel’s (1880 : 116) description of males of this species. Remarks. The characteristic swelling on the 14th tibia of the male was not mentioned either by C. L. Koch or L. Koch in their published descriptions of L. muticus, but L. Koch did mention it in private correspondence with Latzel (Latzel, 1880 : 119) so there is no doubt about the identity of the males on which he based his description. The same cannot, however, be said of the females. In his key L. Koch contrasted the incurved internal pair of spurs on the gonopods of L. mutabilis with the straight spurs of those of L. muticus in order to differentiate between females of these two species; he also described the claw of the gonopod of L. muticus as tripartite. In fact, incurving of these spurs, although not invariable, is usually more marked in muticus than in mutabilis and the claw of muticus has the external denticle so reduced as to appear bipartite. It seems, therefore, that L. Koch did not have females of L. muticus before him when he described this species but those of L. pelidnus and L. mutabilis in which the internal spurs, partic- ularly in pelidnus, are often straight and the claw always tripartite. Most of the females of L. muticus in the Collection were identified by L. Koch as L. mutabilis. C. L. Koch’s original description of this species is unsatisfactory but his illustra- tions of a female (C. L. Koch, 1863: fig. 107) resemble the species regarded as L. muticus more than any other and there is no reason to dispute its identity. 13. Lithobius varius C. L. Koch Lithobius varius C. L. Koch, 1847 : 151. 1863, 1 : 128, fig. 118a & b. TYPE LOCALITY. Bavaria. Remarks. L. Koch (1862) was not familiar with this species and there is no mat- erial referable to L. varius in the Koch Collection. C. L. Koch’s illustrations of a female (C. L. Koch, 1863: fig. 118) are quite consistent with Latzel’s (1880 : 126) suggestion that this species may be identical with L. aeruginosus L. Koch, but there is no certainty of this and L. varius should be rejected as a nomen dubium. 118 E. H. EASON 14. Lithobius minutus C. L. Koch Lithobius minutus C. L. Koch, 1847 : 152. 1863, 1: 129, fig. r119a & b. L. Koch, 1862 : 84, fig. 40. TYPE LOCALITY. Bavaria. MATERIAL EXAMINED. ‘“‘Lithobius minutus C. L. Koch, Niirnberg, leg. L. Koch [rewritten] Zool. Mus. Berlin: Kat. Nr. 342. Twelve badly mutilated immature specimens of Lithobius sp. with neither legs nor antennae, probably 1st and 2nd post-larval stadia of L. mutabilis. “L. minutus [rewritten] “frank. Jura” B.M.(N.H.) Reg. no. 13.6.18. 387-400. Fourteen 1st post-larval stadia of L. mutabilis together with a 2nd post-larval stadium of another species, probably L. pusillus pusillifrater. “L. minutus [rewritten] “[hab.] Haspelmooz’”’ B.M.(N.H.) Reg. no. 13.6.18.401. A 4th larval stadium and three badly mutilated 1st post-larval stadia, all probably belonging to L. pusillus pusillifrater. “TL. minutus [rewritten] “Nirnberg’” B.M.(N.H.) Reg. no. 13.6.18.402—-433. Thirty-two immature specimens ot L. mutabilis ranging from a 4th larval stadium to 2nd post-larval stadia. REMARKS. Of the above specimens, only the Ist post-larval stadia of L. mutabilis agree exactly with L. Koch’s description of L. minutus. Latzel (1880 : 228) gave L. minutus as a synonym of L. mutabilis and Haase (1880 : 32) redescribed the form in some detail, coming to the same conclusion. Although it is undoubtedly a senior synonym of L. mutablis (and a junior synonym of L. communis), the name has not been used by subsequent authors, and, in order to validate L. mutabilis, it is intended to ask the International Commission on Zoological Nomenclature to use its plenary powers to suppress the name minutus C. L. Koch 1847 as published in the binomen Lithobius minutus C. L. Koch, and to supress the name communis C. L. Koch 1844 as published in the binomen Lithobius communis C. L. Koch (see also p. III). 15. Lithobius inermis L. Koch Lithobius ineymis L. Koch in Rosenhauer, 1856 : 415. L. Koch, 1862 : 65, fig. 26. TypE Locality. Malaga, Spain. Remarks. L. ineymis was originally described from a specimen borrowed from the Rosenhauer Collection. Dr. Egon Popp, curator of the Zoologische Staats- sammlung, Munich, informs me that many of Rosenhauer’s specimens were sold to the Alte Akademie, Munich, the precursor of the Staatssammlung, but that most of them were destroyed during the Second World War. Whether the type specimen of L. inermis was among this material is not known, but it has not been found in Munich and there is no record of Rosenhauer having deposited any of her specimens elsewhere. Koch’s earlier (1856) description of this species is very scanty but his subsequent (1862) one is recognizable and Brolemann (1926 : 264) described it very fully. THE LITHOBIUS SPECIES OF C. L. AND L. KOCH 119 16. Lithobius festivus L. Koch Lithobius festivus L. Koch 1862 : 29, fig. 2. TYPE Locality. Garmisch, Bavaria. MATERIAL EXAMINED. Among the specimens Koch correctly identified as L. grossipes the holotype and most of the specimens from South Tyrol bear an additional name, ‘‘festivus’’, in Koch’s hand. But the only specimen of this species in the Collection from Garmisch is merely labelled “Lithobius grossipes”’ ‘““Garmisch”’ (see also Eason, 1970a). TYPE SPECIMEN. L. festivus was originally described from a male and a female from Garmisch borrowed from the Keyserling Collection. The specimen referred to above, a male pseudomaturus 24 mm long, in spite of being labelled “‘L. grossipes”’ and having 49 antennal articles (Koch gave 46-47), is in agreement with Koch’s description in other respects and is undoubtedly the male in question, the sole surviving syntype (B.M.(N.H.) 13.6.18.293). The female syntype must have been returned to the Keyserling Collection and has not been found. Remarks. C. L. Koch based Lithobius grossipes on an aberrant specimen and L. montanus (= grossipes) on a specimen without the dark dorsal pattern on the tergites frequently found in grossipes (see p. 112). L. festivus was based on smaller specimens with the dorsal pattern distinct but with the ocelli of the superior row round, and not oval as in large adults. It is not surprising, therefore, that L. Koch at first thought that he was dealing with three distinct species. But it is clear from his labelling of his specimens that he later realised their true identity although he may, at one stage, have regarded festivus as a variety of grossipes and only finally as a true synonym. 17. Lithobius transmarinus L. Koch Lithobius transmarinus L. Koch, 1862 : 33, fig. 5. TYPE LOCALITY. New Orleans, U.S.A. Remarks. The identity of L. transmarinus is discussed along with that of the next species. 18. Lithobius mordax L. Koch Lithobius mordax L. Koch, 1862 : 34, fig. 6. TYPE LOCALITY. New Orleans, U.S.A. Remarks. L. transmarinus and L. mordax seem each originally to have been described from a single specimen, a female and a male respectively, borrowed from the Keyserling Collection: neither of these has been found. There is little in the original descriptions of these two species to suggest that they are distinct from one another and some authors have regarded them as identical. Bollman (1893) believed them both to be synonyms of L. spinipes Say, 1821, but Brolemann (1896) disputed this synonymy and regarded them both as distinct species. Chamberlin 120 E. H. EASON (1911) at one time believed transmarinus to be the female of mordax but in a later publication (Chamberlin, 1925b) he described them as separate species of Neolitho- bius Stuxberg. These descriptions of Chamberlin’s seem to apply to two distinct species and their validity has not recently been disputed. 19. Lithobius trilineatus L. Koch Lithobius trilineatus L. Koch, 1862 : 37, fig. 8. Type LocaLitry. Bahia, Brazil. MATERIAL EXAMINED. “L. trilineatus, Bahia [rewritten]” B.M.(N.H.) Reg. no. 13.6.18.651. A female of L. fortificatus 20 mm long with the r4th and 15th legs missing. Type spECIMEN. L. trilineatus was originally described from a male and a female from Bahia borrowed from the Keyserling Collection. The above specimen of L. forficatus agrees with this description and is undoubtedly the female in question, the sole surviving syntype. The male syntype must have been returned to the Keyserling Collection and has not been found. Remarks. Koch distinguished this form from L. forficatus by means of a number of sulci he observed on the 15th legs, but these are not reliable characters in L. forficatus. Synonymy of L. trilineatus with L. forficatus was first proposed by Fed- rizzi (1877) and has never been disputed. As Brolemann (1909) pointed out, the species must have been introduced to Brazil. 20. Lithobius forficatus var. villosus L. Koch Lithobius forficatus var. villosus L. Koch, 1862 : 41. TypE Locality. Bavarian Alps. MATERIAL EXAMINED. “‘Lithobius forficatus L.” “‘var. villosus, Bayer. Alpen’”’ B.M.(N.H.) Reg. No. 13.6.18.242. A male of L. forficatus 22 mm long with the 15th legs missing. “forficatus var. villosus, Alpen” B.M.(N.H.) Reg. no. 13.6.18.245-246. A male and a female of L. forficatus. TYPE SPECIMEN. L. forficatus var. villosus was originally described from a single male and although the male labelled “Bayer. Alpen” has 10, 9, 9 and 7 coxal pores on the rath and 15th legs respectively (Koch gave 9, 9, 9, 6) it agrees with this description in other respects and is undoubtedly the holotype. Remarks. Koch distinguished var. villosus from the typical form of the species by the larger number of ocelli and coxal pores, the longer 15th legs, and a number of other quite trivial characters. Although the 15th legs of the holotype are missing, those of the other two specimens of villosus in the Collection are barely longer than is usual in L. forficatus. The other characters of the holotype also fall well within normal limits for the species and villosws has never been regarded as a valid variety or subspecies. RHE LITHOBLUS SPECIES OFC. L. AND L. KOCH 121 21. Lithobius parisiensis L. Koch Lithobius parisiensis L. Koch, 1862 : 42, fig. ro. TYPE LOCALITY. Paris. Remarks. The original description of L. parisiensis was based on a single speci- men borrowed from the Keyserling Collection which has not been found. It was described, like L. trilineatus, as having sulci on the 15th legs, but as differing from both L. trilineatus and L. forficatus in having more ocelli (39), prosternal teeth (8 + 8) and coxal pores (9, 10, 9,7). Stuxberg (1871) suggested L. parisiensis as a possible synonym of L. forficatus but Haase (1880), who had examined Keyserling’s specimen, believed it to be a distinct species. However, all the distinctive features of this form are sometimes found in large specimens of L. forficatus and, although there is every possibility ot an introduced centipede being found in Paris, there is no known species of Lithobius to which the description of parisiensis might apply other than forficatus and there is little doubt that Stuxberg was correct. 22. Lithobius muscorum L. Koch Lithobius muscorum L. Koch, 1862 : 43, fig. 11. TYPE LOCALITY. Germany. MATERIAL EXAMINED. “L. muscorum [rewritten]” ‘“‘[hab.] Valzner Weiher” B.M.(N.H.) Reg. no. 13.6.18.434. A female pseudomaturus of L. forficatus 14 mm long. TYPE SPECIMEN. The original description of L. muscorum was based on a single female and agrees well with the above specimen of L. forficatus which is undoubtedly the holotype. No locality was given in the original description, nor is there a locality label accompanying the holotype, so the presumption is that the specimen was found somewhere in Germany. Remarks. Of the features Koch regarded as characteristic of this form, the circular coxal pores are commonly found in immature stadia of L. forficatus, the rather short antennae with only 33 articles are just within normal limits for the pseudomaturus of this species and the shape of the tergal projections which Koch mentions in his key is quite unremarkable in the holotype. Synonymy of L. muscorum with L. forficatus was first proposed by Stuxberg (1871) and has never been disputed. 23. Lithobius hortensis L. Koch Lithobius hortensis L. Koch, 1862 : 45, fig. 12. TYPE LocALITIES. Nuremberg; Landstuhl, Rhineland Palatinate. MATERIAL EXAMINED. “Lithobius hortensis L. Koch, Syntypen, Niirnberg, leg. L. Koch [rewritten]” Zool. Mus. Berlin: Kat. Nr. 335. Two male pseudo- maturus and a small adult female of L. forficatus. 122 E. H. EASON “L. hortensis [rewritten]’’ ““Niirnberg, [hab.] in Garten’? B.M.(N.H.) Reg. no. 13.6.18.303-318 (part). Five males and eleven females of L. forficatus, all either praematurus, pseudomaturus or small adults. “L. hortensis [rewritten]”’ “Ntirnberg” B.M.(N.H.) Reg. no. 13.6.18.303-318 (part). Four exuviae of L. forficatus. Two appear to be from adult males, one from a male pseudomaturus and the other from a female pseudomaturus. “L, hortensis [rewritten]” “Landstuhl” B.M. (N.H.) Reg. no. 13.6.18.319-321. Two male pseudomaturus and a small adult female of L. forficatus. TYPE SPECIMENS. The original description of L. hortensis was based on a number of specimens of both sexes and all the above examples of L. forficatus, except the exuviae, seem to belong to the syntypical series. REMARKS. Koch distinguished this form from L. muscorum, with which it agrees in having circular coxal pores, by the longer antennae and the shape of the tergal projections neither of which have any taxonomic significance. In fact, the coxal pores in some of the larger and more mature syntypes are oval but in no case are they oblong or split-shaped as in most large adults of L. forficatus. Synonymy of L. hortensis with L. forficatus was first proposed by Meinert (1868) and has been accepted by most authors. Latzel (1880 : 61), however, suggested that adults of L. forficatus with circular coxal pores might be regarded as a variety (subspecies) and Verhoeff (1937) considered this form, which he called L. forficatus var. hortensis, to be predominant in the Mediterranean region. But there is no justification for retaining hortensts as the name of a subspecies or even a variety since the shape of the coxal pores in adults of L. forficatus varies continuously, showing every gradation in shape from circular, oval, oblong to slit-shaped even in specimens from the same locality. 24. Lithobius sordidus L. Koch Lithobius sordidus L. Koch, 1862 : 47, fig. 13. Type Locatity. Munich district. RemarKS. The original description of L. sordidus was based on a single female borrowed trom the Keyserling Collection which has not been found. It seems to apply to a pseudomaturus of L. picews L. Koch with only 43 antennal articles, no ventral spines on the 15th tibia and only 2 + 2 spurs on the gonopods. Latzel’s (1880 : 64) proposal of L. sordidus as a synonym of L. piceus is probably justified. 25. Lithobius fossor L. Koch Lithobius fossor L. Koch, 1862 : 48, fig. 14. TyPE LocALity. Gritz, near Nuremberg; Ehrenbiirg, Franconian Jura. MATERIAL EXAMINED. “‘Lithobius fossor L.K.” “Gritz [Gritz]” B.M. (N.H.) Reg. no. 13.6.18.248. A male pseudomaturus of L. piceus 11.5 mm long. “Lithobius fossor L. Koch” “Ehrenbiirg” B.M.(N.H.) Reg. no. 13.6.18.249. A male pseudomaturus of L. piceus 12-5 mm long. THE LITHOBIUS SPECIES OF C. L. AND L. KOCH 123 TYPE SPECIMENS. Although neither of the above specimens of L. piceus agrees with the original description of L. fossor in every detail, there is little doubt that they are the two males on which this description was based and are therefore the syntypes. REMARKS. Koch distinguished this form from L. sordidus by the more numerous antennal articles and the presence of two ventral spines on the 15th tibia. The syntypes have 52 and 47 antennal articles respectively (Koch gave 49) and VaT is the only ventral spine on the 15th tibia of either specimen. A second ventral spine on this article (VmT) is most unusual in L. picews and it is unlikely that Koch had any specimens before him other than these two syntypes when he wrote his rather inaccurate description. Synonymy of L. fossor with L. piceus was first proposed by Latzel (1880 : 64) and has never been disputed. 26. Lithobius piceus L. Koch Bigs 7, Lithobius piceus L. Koch, 1862 : 49, fig. 15. TYPE LOCALITY. Garmisch, Bavaria. MATERIAL EXAMINED. “‘piceus, frank. Jura’ B.M.(N.H.) Reg. no. 13.6.18.577. A female pseudomaturus. “piceus?, Niirnberg’”’ B.M.(N.H.) Reg. no. 13.6.18.578-579. Two female pseudo- maturus. Remarks. L. picews was originally described from a mature female borrowed from the Keyserling Collection which has not been found. None of the above speci- mens belong to the original-material and must have been named by Koch after he had finished writing his book. Garmisch, the type locality of L. piceus, is only some 200 Km. from Nuremberg and the Franconian Jura and there is no reason to doubt that the specimens from these localities are identical with the form originally described, but owing to their immaturity none of them is suitable for selection as a neotype. Koch distinguished L. piceus from L. sordidus and L. fossor by the more numerous antennal articles, ocelli and coxal pores, all features of maturity. Like fossor, piceus was described as having two ventral spines on the 15th tibia, presumably VmT in addition to the usual VaT: although Haase (1880) mentioned an occasional second ventral spine on this article, it was not mentioned either by Latzel (1880) or Brolemann (1930) in their descriptions ot L. picews. Tobias (1969) examined 80 examples of this species from the Pyrenees and found 15VmT on one side of one individual only, so that either the type specimen was unusual or, as in the case of L. fossor, Koch was mistaken. Another character Koch used to distinguish between piceus and sordidus was the number of spurs on the female gonopods, 3 + 3 in the former and 2 + 2 in the latter. The female pseudomaturus of piceus frequently has the full complement of 3 + 3 spurs but the internal pair are very small or may be absent. The female in the 124 E. H. EASON Collection from the Franconian Jura has 3 + 3 spurs but those from Nuremberg have 3 + 2 and 2 + 2, and it was probably this that led Koch to be uncertain of their identity. Perhaps the most significant feature shown by the above examples of L. piceus, as well as by both of those of L. fossor, is the relative slenderness of the tarsi and metatarsi of the 15th legs, the distal extremity of the tibia being broader than the base of the tarsus in the ratio 3 : 2 (Fig. 7). Brolemann’s figure of L. piceus gract- litarvsis Brolemann (Brolemann, 1930 : fig. 400) shows about the same relative change in breadth between these two adjacent articles and it seems that he (Brole- mann, 1898) described this subspecies from the Pyrenees on the assumption that it differed from the typical form in this respect. However, in addition to Koch’s specimens, English examples and those from Italy (Eason, 1964: figs 342 & 343), the latter answering to the description of L. piceus verhoefi Demange, 1958, show that same abrupt transition in the breadth of the leg at the 15th tibiotarsal articu- lation so it seems that this character is widespread throughout the species. More- over, L. piceus gracilitarsis was recorded by Negrea (1965) from Transylvania and by Folkmanova (1951, 1954) and Folkmanova and Lang (1955, 1960) from a number of localities in Czechoslovakia and southern Poland, and although Matic and Dara- bantu (1968) suggested that the slender tarsi of the specimens on which some of these records were based may be features of immaturity, this is not the case in English specimens. On the other hand Brolemann (1930 : 262) described the Alpine form of L. piceus (which he regarded as the nominate subspecies) as having no abrupt transition in breadth between the 15th tibia and tarsus. It we assume that most records of L. pices refer to this latter form, the comparative distribution of L. piceus and L. p. gracilitarsis does not suggest that we are dealing with two sub- species but that the shape of the 15th legs in L. prcews is variable and that subsp. gracilitarsis has no real status. There is also the possibility that some records of L. piceus piceus in the literature may refer to a closely related species, L. peregrinus Latzel, 1880, in which the 15th tarsi and metatarsi are relatively stout. 27. Lithobius coriaceus L. Koch Lithobius coviaceus L. Koch, 1862 : 51, fig. 16. TYPE LOCALITY. Germany. MATERIAL EXAMINED. “‘Lithobius coriaceus L.K.” “frank. Jura’ B.M.(N.H.) Reg. no. 13.6.18.88-91. Two agenitalis II and a male immaturus of L. forficatus. “Lithobius coriaceus L.K.” “Gritz [Griitz, near Nuremberg], [hab.] Garten” B.M.(N.H.) 13.6.18.92-96. Two agenitalis I, an agenitalis II and two male imma- turus of L. forficatus. “Lithobius coriaceus L.K.’”’ “Dietenhofen [Franconian Jura]’’ B.M.(N.H.) Reg. no. 13.6.18.97. An agenitalis II of L. forficatus. TYPE SPECIMENS. The original description of L. coriaceus was based on a number of specimens and all the above examples of L. forjicatus seem to belong to the syn- typical series. THE LITHOBIUS SPECIES OF C. L. AND L. KOCH 125 Remarks. Koch had both sexes of the stadium immaturus as well as agenitalis (which he took to be males) before him when he described this form, so the above series which does not include females is incomplete. Of the key characters given by Koch, 4 + 4 prosternal teeth is exceeded in one immaturus (4 + 5) while two ventral spines are never found on the 15th tibia of the agenitalis of L. forficatus, so his description is not altogether accurate. Synonymy of L. coriaceus with L. forficatus was first proposed by Stuxberg (1871) and has never been disputed. 28. Lithobius velox L. Koch Lithobius velox L. Koch, 1862 : 56, fig. 19. Type Locatities. Landstuhl, Rhineland Palatinate; Franconia; Vienna district. MATERIAL EXAMINED. “‘L. velox [rewritten]’’ “Landstuhl’’ B.M.(N.H.) Reg. no. 13.6.18.616-617._ A male and a female of L. melanops Newport. “L. velox [rewritten] ‘“Dietenhofen [Franconian Jura]’” B.M.(N.H.) Reg. no. 13.6.18.618. A male 4th post-larval stadium of L. melanops. “T. velox [rewritten]” “Wien” B.M.(N.H.) Reg. no. 13.6.18.619. A mutilated female of L. melanops. TYPE SPECIMENS. The original description of L. velox was based-on a number of specimens of both sexes and all the above examples of L. melanops certainly belong to the syntypical series. REMARKS. The size, the number of ocelli, and the number of coxal pores both in Koch’s description and in the syntypes are all close to the lower normal range for adults and the upper range for 4th post-larval stadia of L. melanops. Meinert (1868) suggested L. velox as a possible synonym of L. bucculentus L. Koch, under which he was probably describing examples of L. melanops. Synonymy of L. velox with L. glabratus (= melanops) was first proposed by Latzel (1880 : 74) and has never been disputed. 29. Lithobius bucculentus L. Koch Lithobius bucculentus L. Koch, 1826 : 57, fig. 20. Type LocaLity. Munich district. MATERIAL EXAMINED. ‘“‘bucculentus Mein.(?), Ratzes [Rasa, Italy] leg. Milde”’ B.M.(N.H.) Reg. no. 13.6.18.17. A single male of L. tricuspis Meinert. Remarks. Koch’s original description of L. bucculentus as having sharp posterior projections on T.g, 11 and 13, and the antennae and 15th legs both over half the body-length is much more suggestive of L. tricuspis than of L. melanops (of which it has hitherto been regarded as a synonym), but this description was based on a male from Munich borrowed from the Keyserling Collection which has not been found, and the specimen from Rasa in the South Tyrol is not the holotype; nor can the latter be selected as neotype as it was taken too far from the type locality and Koch was uncertain of its identity. This uncertainty may have been due to the presence, on the specimen from Rasa, of a ventral spine (VaT) on the 15th tibia, a spine which 126 E. H. EASON may be absent in L. tricuspis (Eason, 1965) and which Koch did not mention in the original description. Although 15 VaT rarely if ever occurs in L. melanops, Meinert (1868) did mention a single variable ventral spine of the 15th tibia in his redescription of L. bucculentus, possibly because he based it on examples of tricuspis as well as of melanops, and this may have led Koch to attach Meinert’s name to the specimen from Rasa which he probably examined after he had read Meinert’s paper. Stuxberg’s (1871) description of L. bucculentus is general enough to include all species of Lithobius with more than seven ocelli on each side, 2 + 2 prosternal teeth, and posterior projections on T.g, rr and 13: he included all the nominal species known to him which are embraced by this definition in his synonymy. Meinert’s (1868) and Haase’s (1880) descriptions of this species are identical with each other, much more restricted than Stuxberg’s, and more likely to refer almost exclusively to L. melanops. Latzel (1880: 74) gave a full description of L. glabratus (= melanops) and proposed L. bucculentus as a synonym. But it is fairly certain that all these authors were mistaken and that L. bucculentus is the senior synonym of L. tricuspis Meinert, 1872. However, the name has not been used for well over fifty years and to revive it would cause confusion. It is intended therefore to ask the International Commission on Zoological Nomenclature to use its plenary powers to suppress the name bucculentus L. Koch 1862 as published in the binomen Lithobius bucculentus L. Koch, so as to validate Lithobius tricuspis Meinert. 30. Lithobius melanocephalus C. L. Koch Lithobius melanocephalus C. L. Koch in L. Koch, 1862 : 58, fig. 21. C. L. Koch, 1863, 1 : 130, fig. 120a & b. Type LocaLity. Ehrenbirg, Franconian Jura. MATERIAL EXAMINED. “‘L. melanocephalus [rewritten]’’ “Ehrenbiirg’” B.M. (N.H.) Reg. no. 13.6.18.384-385. A male and female of L. melanops. TYPE SPECIMENS. The original description of L. melanocephalus was based on a male and a female and the above two examples of L. melanops are undoubtedly the syntypes. Remarks. L. Koch attributed this species to his father and the later description (C. L. Koch, 1863) was no doubt made from one of C. L. Koch’s specimens: but the first description to be published, based on L. Koch’s two specimens, must stand as the original. There is no reason to suppose that C. L. Koch’s illustration of L. melanocephalus refers to L. dentatus as Latzel (1880 : 76) suggested; in this figure (C. L. Koch, 1863: fig. 20a) the tergal projections do indeed resemble those of L. dentatus but the antennae are only two-fifths of the body-length with 34 articles which is typical of L. melanops and quite unlike L. dentatus in which the antennae are about three-fifths of body-length with 50 to 60 articles: C. L. Koch is likely enough to have made a slight error in outlining the shape of the tergites but is much less likely to have been mistaken over the antennae. The size, the number of ocelli, and the number of coxal pores both in L. Koch’s description and in the syntypes are close to the upper normal range for adults of THE LITHOBIUS SPECIES OF C. L. AND L. KOCH 127 L. melanops. L. Koch described L. melanocephalus as resembling L. bucculentus (= tricuspis) in having three ventral spines on the 15th femur but differing in having blunt tergal projections; although a third ventral femoral spine (15VpF), which is almost invariable in L. tricuspis, is quite common in large specimens of L. melanops, it is only present on the left 15th leg of the female syntype (the 15th legs of the male syntype are missing); the tergal projections in the syntypes as in all specimens of L. melanops are, of course, noticeably blunter than in L. tricuspis. Meinert (1872) and Haase (1880) suggested L. melanocephalus as a possible synonym of L. bucculentus under which they were probably describing examples of L. melanops. Latzel (1880 : 74), in spite of his doubt about the identity of C. L. Koch’s figure had no doubt about L. Koch’s description and was the first to propose L. melano- cephalus as a synonym of L. glabratus (= melanops). 31. Lithobius venator L. Koch Lithobius venatoy L. Koch, 1862 : 59, fig. 22. Type Locatity. Ehrenbitirg, Franconian Jura. MATERIAL EXAMINED. “L. venator [rewritten]’’ “Ehrenbirg’ B.M.(N.H.) Reg. no. 13.6.18.620. A female of L. melanops 13 mm long. “L. venator [rewritten]’’ ‘“Nirnberg’ B.M.(N.H.) Reg. no. 13.6.18.621-629. Ten more or less mutilated specimens of L. mgrifrons Latzel and Haase. “L. venator [rewritten]” “‘[?]’” B.M.(N.H.) Reg. no. 13.6.18.630-631. A male and a female of L. melanops. “nigrifrons? venator?, Nurnberg” B.M.(N.H.) Reg. no. 13.6.18.570-573. Four mutilated Ist post-larval stadia of L. melanops. TYPE SPECIMEN. The original description of L. venator was based on a single female and although the above female of L. melanops from Ehrenbtirg has 36 antennal articles (Koch gave 38) and 1 + 4, 4, 2 ocelli (Koch gave 1 +4, 4, 3 and figured I + 4, 3, 2) it agrees with this description in other respects and is undoubtedly the holotype. REMARKS. Koch distinguished this form from L. melanocephalus by the fewer ocelli and the absence of the third ventral spine on the 15th femur, and from L. velox by the shape of the internal margins of the tergal projections; none of these characters has any taxonomic significance. Koch’s labelling of examples of L. nigrifrons as L. venator is not surprising as this species answers equally well to the original description of L. venator and Latzel (1880 : 73) suspected that migrifrons and venator might be identical. But Koch, in private correspondence with Latzel (Latzel, 1880 : 73), confirmed that his three specimens of L. venator (no doubt the holotype and the two specimens from an indecipherable locality) had accessory apical claws on the 15th legs, claws which are present in L. melanops but not in L. mgrifrons. The identity of L. venator has not hitherto been definitely established. Meinert (1868) suggested it as a possible synonym of L. bucculentus (i.e. L. melanops). Stux- berg (1871) gave the same synonymy but also included such diverse species as 128 E. H. EASON L. dentatus, L. agilis and L. intrepidus Meinert as synonyms so one cannot be sure which species he had in mind. Latzel (1880 : 74) suggested L. venator as a possible synonym of L. glabratus (= melanops) but he was not altogether satisfied as to its identity. The only descriptions of species under L. venator L. Koch, other than the original, are those of Porat (1869) and Sseliwanoff (1880); these are both difficult to interpret but they probably refer to L. melanops. Attems (1927) regarded venator as a subspecies of L. melanops without the accessory apical claw on the 15th leg, but this use of the name is quite wrong. L. venator is definitely a synonym of L. melanops. 32. Lithobius minimus L. Koch Lithobius minimus L. Koch, 1862 : 61, fig. 23. TYPE LOCALITY. Germany. MATERIAL EXAMINED. “‘L. minimus [rewritten] ““Mogeldorf [near Nuremberg], [hab.] Erlenwalddren” B.M.(N.H.) Reg. no. 13.6.18.386. A female 3rd _post- larval stadium 6-4 mm long with both 15th legs missing. This specimen belongs to one of the lapidicola-borealis group of species characterized by rather feeble pos- terior projections on T.11 and 13 only; it may very tentatively be referred to L. salicis Verhoeff, but the absence of the 15th legs and the immaturity of the specimen make definite identification impossible. Remarks. The original description of L. minimus disagrees altogether with the above specimen which has 35 antennal articles (Koch gave 22), I + 4, 3, I ocelli (Koch gave I + 2, 1), 2, 2, 2, 2 coxal pores (Koch gave I, I, I, I), and 0, 1, 3, 2, I ventral spines on the 14th leg (Koch gave 0, 0, I, I, 0 on the 15th). The extent of this disagreement can hardly be accounted for by a careless description and the specimen must have been labelled in error by Koch himself or during some rearrangement of his Collection. Latzel did not deal with this species but Meinert (1872) suggested L. minimus as a possible synonym of L. bucculentus (i.e. L. melanops). Koch’s description, how- ever, agrees better with the 1st post-larval stadium of L. agilis. But there can be no certainty about the identity of L. minimus and it should be rejected as a nomen dubium. 33. Lithobius immutabilis L. Koch Lithobius immutabilis L. Koch, 1862 : 62, fig. 24. TYPE LOCALITY. Germany. MATERIAL EXAMINED. “‘Lithobius immutabilis L. Koch, Syntypen, Nurnberg, leg. L. Koch [rewritten]’”’ Zool. Mus. Berlin: Kat. Nr. 331. Two mutilated 1st post-larval stadia of L. dentatus. “L. immutabilis [rewritten] ‘“Mégeldorf [near Nuremberg], [hab.] Erlengebtisch”’ B.M.(N.H.) Reg. no. 13.6.18.322-325. A 4th larval stadium and four Ist post- larval stadia of L. dentatus, all more or less mutilated. THE LITHOBIUS SPECIES OF C. L..AND L. KOCH 129 TYPE SPECIMENS. The original description of L. immutabilis was based on a number of specimens and all the above examples of L. dentatus seem to belong to the syntypical series. REMARKS. Koch’s description is clearly of a series of immature specimens and applies equally well to a number of species. Meinert (1872) and Latzel (1880 : 74) suggested L. immutabilis as a possible synonym of L. bucculentus (i.e. L. melanops) and L. glabratus (= melanops) respectively. Only Haase (1880), who may well have seen the original material, suggested its true synonymy with L. dentatus. 34. Lithobius macilentus L. Koch Lithobius macilentus L. Koch, 1862 : 63, fig. 25. TYPE LOCALITIES. Nuremberg; Franconian Jura; Bolzano, Italy. MATERIAL EXAMINED. “‘Lithobius macilentus L. Koch, Syntypen?, Niirnberg, leg. L. Koch [rewritten]’’ Zool. Mus. Berlin: Kat. Nr. 332. A female of L. erythroce- phalus and two very mutilated specimens, a male and a female, probably 4th post- larval stadia of L. mutabilis. “L. macilentus [rewritten]” ‘“‘[hab.] Valzner Weiher, Gritz [Griitz, near Niirem- berg] B.M.(N.H.) Reg. no. 13.6.18.380-382. Two female 3rd post-larval stadia of L. aulacopus Latzel with 37 and 35 antennal articles respectively, together with one 1st post-larval stadium of L. tricuspis with 26 antennal articles, all three more or less mutilated. “L. macilentus [rewritten]? ‘‘macilentus, Botzen [Bolzano]” B.M.(N.H.) Reg. no. 13.6.18.383 (part). A mutilated 1st post-larval stadium ot L. tricuspis with antennae missing. “L. macilentus [rewritten]’‘‘Happurg [Franconian Jura]” B.M.(N.H.) Reg. no. 13-6.18.383 (part). A badly mutilated fragment barely recognizable as belonging to a species of Lithobius. y) 66 yee TYPE SPECIMENS. The original description of L. macilentus was obviously based on two different species and there is no doubt that the above specimens from Griitz and Bolzano, and possibly also the fragment from Happurg, belong to the syntypical series. The same cannot be said of those in the Berlin Museum which bear no tesemblance to Koch’s description and must have been labelled by mistake. That part of Koch’s description applying to the larger specimens with more than 32 antennal articles (females) together with his figure of the ocelli clearly refers to an immature example of L. aulacopus; the least defective ot these, a 3rd _post-larval stadium 8mm long with 37 antennal articles, answering exactly to Koch’s description of the larger form and also to Latzel’s (1880 : 85) description of the ‘‘juvenis’’ of L. aulacopus, is here formally designated as the lectotype (B.M.(N.H.) 13.6.18.380). Remarks. Meinert (1872) suggested L. macilentus as a possible synonym of L. agilis. Fedrizzi (1877) gave a rather diffuse account of the species which is more than mere repetition of Koch’s description and, although probably composite, seems to include L. aulacopus. Latzel (1880 : 80) recognised the inclusion of immature examples of two separate species by Koch in his original description of L. macilentus 130 E. H. EASON and, probably following Meinert, proposed L. agilis as the senior synonym to apply only to the smaller specimens with fewer than 32 antennal articles which Koch regarded as males; he made no suggestion as to the identity of the larger females with more than 32 antennal articles and dismissed Koch’s description and figure of the ocelli as unreliable, going on to describe L. aulacopus as a new species (Latzel, 1880 : 84). Meinert’s and Latzel’s failure to guess the identity of the smaller specimens is quite understandable but both of them overlooking Koch’s very adequate description of the larger females is surprising. Haase (1880) followed Latzel’s synonymy and all subsequent authors have accepted L. macilentus as a synonym of L. agilis. Lohmander (1957 and in /itt.) pointed out that L. aulacopus is a junior synonym of L. intrepidus Meinert, 1868. Most authors, however, continue to use the name aulacopus aad the nomenclature of this species is unsatisfactory: it should now be known as L. macilentus L. Koch. 35. Lithobius alpinus L. Koch Fig. 3 Lithobius alpinus L. Koch, 1862 : 66, fig. 27. TYPE LOCALITY. Seiseralpe, an alpine hut in the Italian Tyrol. MATERIAL EXAMINED. “‘Lithobius alpinus in Seiseralpe leg. Gredler’’ B.M.(N.H.) Reg. no. 13.6.18.12. A female of L. lucifugus L. Koch 15 mm long with oval coxal pores and most of the legs missing. TYPE SPECIMEN. The original description of L. alpinus was based on a single defective female and agrees exactly with the above specimen of L. lucifugus which is undoubtedly the holotype. REMARKS. Koch used the shape of the coxal pores as a key character, separating those forms such as L. alpinus with oval coxal pores from species in which the pores are circular. Latzel (1880 : 122) realised that these pores may be either circular or oval in L. lucifugus and suggested L. alpinus as a possible synonym of this species. Borek (1967) argued that L. alpinus must be a species distinct from L. lucifugus owing to its small number of antennal articles (30), but this argument is not justified ; the number of antennal articles is very variable in L. lucifugus as Borek himself pointed out. The holotype shows a pair of small paramedian prosternal teeth not noted by Koch, in addition to the usual 2 + 2 (Fig. 3). These extra teeth are characteristic of L. lucifugus var. latzeli Verhoeff, 1935 which Verhoeff (1937) later raised to a sub- species; but the form and number of prosternal teeth in /ucifugus are very variable, even in examples from the same locality, and there is no justification for regarding latzeli as a subspecies. Should it be thought necessary to give varietal status to specimens with these extra prosternal teeth, the name Jatzeli Verhoeff should be used, since the name alpinus must be rejected as a junior homonym of Lithobius alpinus Heer, 1845, which refers to an immature specimen of uncertain identity from the Swiss Alps. THE LITHOBIUS SPECIES OF C. L. AND L. KOCH 131 36. Lithobius granulatus L. Koch Lithobius granulatus L. Koch, 1862 : 67, fig. 28. TyPE LOCALITY. Unknown. MATERIAL EXAMINED. ‘“‘Lithobius granulatus L.K.” “Patria?” B.M.(N.H.) Reg. no. 13.6.18.258. A male of L. lucifugus 19 mm long with oval coxal pores and with both antennae and all the legs missing. TYPE SPECIMEN. The original description of L. granulatus was based on a single male with antennae and legs missing and agrees exactly with the above specimen of L. lucifugus which is undoubtedly the holotype. Remarks. As in the case o1 L. alpinus the oval coxal pores of L. granulatus were used by Koch as a key character. The only trace of an appendage borne by the holotype consists of the first and second articles of the right antenna, the second of which appears unusually elongate, and it was this slight aberration which led Koch to regard L. granulatus as a species distinct from L. alpinus. There is no previously proposed synonymy for this form and very little mention of it in the literature. Sseliwanoff’s (1880) description of L. granulatus L. Koch refers to some species other than L. lucifugus. L. granulatus Meinert, 1872 is a homonym referring to a South American species. 37. Lithobius crassipes L. Koch Lithobius crassipes. L. Koch, 1862 : 71, fig. 31. Type LocaLity. Nuremberg district. MATERIAL EXAMINED. “‘Lithobius crassipes L. Koch, Syntypen?, Franconia (Jura), leg. L. Koch [rewritten]? Zool. Mus. Berlin: Kat. Nr. 340. A male and two females. “No. 272 Lithobius crassipes L.K. Types [rewritten]”’ “frank. Jura” B.M.(N.H.) Reg. no. 13.6.18.98-110. Four males and nine females ranging in the degree of their maturity from 3rd post-larval stadia to adults. “crassipes, Niirnberg’’ B.M.(N.H.) Reg. no. 13.6.18.259-261. A male and a fe- male together with a female of L. curtipes. TYPE SPECIMENS. The original description of L. crassipes was based on a number of specimens of both sexes. All the above specimens from the Franconian Jura, much of which lies in the Nuremberg district, seem to belong to the syntypical series, but those labelled ‘‘Niirnberg’’ were probably added to the Collection after the description had been written. A well-preserved adult female 8-5 mm long answering to Latzel’s (1880 : 128) description of L. crassipes is here formally desig- nated as the lectotype (B.M.(N.H.) 13.6.18.98). Remarks. The single female of L. curtipes from Nuremberg does not show the characteristic arrangement of the ocelli very clearly, so it is not surprising that Koch should have included it among his specimens of L. crassipes. 132 1D dnl IDS ORY 38. Lithobius sulcatus L. Koch Fig. 4 Lithobius sulcatus L. Koch, 1862 : 73, fig. 32. Type LocaLity. Nuremberg. MATERIAL EXAMINED. “‘L. sulcatus [rewritten]’’ ‘“‘Happurg [Franconian Jura]” B.M.(N.H.) Reg. no. 13.6.18.599-600. A male 2nd post-larval stadium of L. agilis and another immature male, probably a 2nd post larval stadium of L. crassipes. “L. sulcatus [rewritten] “Gritz [Gritz], [hab.] Walddren bin. Glaishammer”’ B.M.(N.H.) Reg. no. 13.6.18.601-607. Three specimens answering to Koch’s description of L. sulcatus together with a female 3rd post-larval stadium of L. aevuginosus and three other immature specimens, probably a Ist post-larval stadium and female 2nd post-larval stadium of L. curtipes and a Ist post-larval stadium of L. crassipes. TYPE SPECIMENS. The original description of L. sulcatus was based on a number of immature specimens and all those from Griitz, near Nuremberg, some of which bear only a superficial resemblance to one another, seem to belong to the syntypical series. One of those answering to Koch’s description is here formally designated as the lectotype (B.M.(N.H.) 13.6.18.601). REMARKS. Koch can have examined only three specimens at all carefully when writing his description of L. sulcatus ; most of the others labelled “L. sulcatus’ agree with this description in a few characters only and the example of L. agilis not at all. Latzel did not deal with the species, there is little reference to it in the literature, and it has never been redescribed although Attems (1909) referred it to the subgenus Monotarsobius. The following description is based on the lectotype and the other two specimens from Grtitz with which it is conspecific. Description. Length: 4-6 to 4.8mm. Antennae: 1-4 to 1-5 mm long; of 21 or 22 articles. Ocelli: 3 or 4; a relatively small posterior ocellus, a rather larger inter- mediate one and one or two much smaller anterior ocelli (Fig. 4). Prosternum: with 2 -+ 2 teeth and a pair of well-developed lateral spines. Tergites: slightly wrinkled; general shape as in species of Monotarsobius with posterior angles of T.9, 11 and 13 obtusely rounded. Coxal pores: 1, 1,1, 1. Legs: tarsus and meta- tarsus fused on rst to 11th; the 14th and 15th moderately thickened; 15th accessory apical claw present. Genitalia: undeveloped. Spinulation: Ventral Dorsal G i 12 F ar (c t 12) F ar i — — — — m (a) 2 — — — — m (a) 3 - -—- — — —- —- = Pp ® 4 m =, op ey eae 5 — — — m m — -- —- a—p a—p @ = = ae = ap ap 7) — — — m m — — — a—p a—p Li LIHOBRLUS SPECIES OG, L. AND LE. KOCH 133 Spinulation: Ventral Dorsal Cc t 12) F 2 Cc t 12) F z 8 — — — m m — — — a—p a—p 9 — — — m m — — — a—p a—p 10 eat = — m m = — (Pp) Pp = II = — — m m — — 2) (p) — 12 = — m m m — — (mp) (p) — 13 — — m m — I4 = = m m = 15 + —_ m m — The letters enclosed in brackets indicate variable spines. IpENTITY. These specimens are Ist post-larval stadia of a species of Monotar- sobius which, in its adult form, would certainly have antennal articles considerably in excess of twenty. They would therefore answer to L. microps Meinert, 1868 (= L. duboscqui Brolemann), one of the few species of Monotarsobius with more than twenty antennal articles and a 15th accessory apical claw, were it not for the pres- ence of such spines as DaF and DpF which are rarely found even in adults of L. mucrops (see also Jeekel, 1964). Verhoeff (1931, 1934, 1937) attached the name microps to one or more European species of Monotarsobius which differ from the true microps in having a more profuse spinulation. In using the name in this way Verhoeff was following Meinert who, in his redescription ot L. microps (Meinert, 1872), included at least one other species in addition to the true microps. But in spite ot “L. microps’” figuring in numerous keys and brief descriptions it seems never to have been properly described except by Brolemann (1930), and there is some doubt as to whether Brolemann was describing the same species as Verhoeff. On the other hand a bewildering number of subspecies of “‘microps’’ have been described by Verhoeff and other authors from various parts of Europe, some in considerable detail. Although the specimens of L. sulcatus may possibly be examples of L. mucrops with unusual spinulation, they are more likely to be identical with the misnamed “‘L. microps’’ of Verhoeff or one of its subspecies. But a full description of the species must await discovery of adults in the neighbourhood of Nuremberg. 39. Lithobius aeruginosus L. Koch Lithobius aeruginosus L. Koch, 1862 : 74, fig. 33. Type LocaLity. Nuremberg district. MATERIAL EXAMINED. “‘Lithobius aeroginosus L.K.” “frank. Jura’ B.M.(N.H.) Reg. no. 13.6.18.1. A male 4th post-larval stadium 5:8 mm long. “Lithobius aeruginosus L.K.” B.M.(N.H.) Reg. no. 13.6.18.2. A male 4th post- larval stadium of L. curtipes 7 mm long. TYPE SPECIMENS. The original description of L. aeruginosus was based on at least two males and the above two specimens seem to be the syntypes. The one labelled “frank. Jura’’ answers more closely to this description and also agrees with Latzel’s (1880 : 126) description ot L. aeruginosus; it is here formally designated as the lectotype (B.M.(N.H.) 13.6.18.1). 134 E. H. EASON Remarks. Although both the above specimens have three ventral spines on the 15th prefemur (Koch gave one) they answer fairly well to Koch’s description in other respects. The male of L. curtipes, however, has the characteristic tibial projection feebly but quite distinctly developed on the 15th leg, a second ventral spine (VaF) on the 15th femur, and the ocelli, though not arranged as in adults of L. curtipes, are in a somewhat irregular line and not in a precisely straight line as in the lectotype: Koch evidently overlooked all these details and seems to have depended more on the lectotype for his description. The 15th legs of the lectotype are unusually long for this species (two-fifths of the body-length), a feature noted by Koch and attributed by Latzel (1880 : 128) to immaturity. But Koch regarded it as characteristic of the species and this may have led him to identify the other examples of L. aeruginosus in the Collection, all of which have relatively short legs, as L. curtipes or L. sulcatus. 40. Lithobius mutabilis L. Koch Lithobius variegatus: C. L. Koch, 1844: 21, fig. 21. 1863, 2: 21. fig. 144a & b (non Leach 1814) Lithobius mutabilis L. Koch, 1862 : 75, fig. 34. TYPE LOCALITY. Germany. MATERIAL EXAMINED. “‘Lithobius mutabilis L. Koch, Syntypen?, Nurnberg, leg. L. Koch [rewritten]’’ Zool. Mus. Berlin: Kat. Nr. 329. Four males and four females together with a female of L. muticus, all more or less mutilated. “L. mutabilis [rewritten] “frank. Jura’ B.M.(N.H.) Reg. no. 13.6.18.435-458. Five males and nine females together with five males and three females of L. pelidnus and a male and female of L. muticus. “mutabilis, frank. Jura’? B.M.(N.H.) Reg. no. 13.6.18.459-460. Two males of L erythrocephalus. “mutabilis, Nirnberg’’ B.M.(N.H.) Reg. no. 13.6.18.461-520 (part). Twenty-four males (including 3rd and 4th post-larval stadia) and twenty-three females together with seven males and five females of L. pelidnus and an adult male, a male 3rd post- larval stadium and eight females of L. muticus. “L. mutabilis [rewritten]”’ ““Niirnberg—Einlegend 2 mit Ei” B.M.(M.H.) Reg. no. 13.6.18.461-520 (part). A single female. “L. mutabilis [rewritten] “Miinchen“ B.M.(N.H.) Reg. no. 13.6.18.521-526. A female 4th post-larval stadium of L. mutabilis, a male and a female of L. muticus, two females of L. lapidicola Meinert (sensu Jeekel, 1964 non Latzel, 1880), and two rather defective females, probably belonging to L. subtilis Latzel. “mutabilis, Tirol’ B.M.(N.H.) Reg. no. 13.6.18.527. A single male. “L. mutabilis [rewritten]”’ ‘““Bohmen [Bohemia] B.M.(N.H.) Reg. no. 13.6.18. 528-531. Two males and two females. “mutabilis, Franzensbad [Frantiskovy Lazne, Czechoslovakia]” B.M.(N.H.) Reg. no. 13.6.18.632. A single male. TYPE SPECIMENS. The original description of L. mutabilis was based on two distinct species, L. mutabilis as described by Latzel (1880 : 97) and L. pelidnus THE LITHOBIUS SPECIES OF C. L. AND L. KOCH 135 Haase, 1880. The specimens in the Berlin Museum and those in the British Museum (N.H.) from the Franconian Jura (13.18.6.435—458) all seem to belong to the syn- typical series and a well-preserved male 10 mm long agreeing with Latzel’s description of L. mutabilis is here formally designated as the lectotype (B.M.(N.H.) 13.6.18.435). Remarks. L. Koch intended his description of L. mutabilis to apply to L. variegatus Leach as described by C. L. Koch in 1844 and he renamed it because he realised that Leach’s (1814) description referred to an altogether different species. Although one cannot say whether C. L. Koch had before him examples of the species now recognized as L. mutabilis, those of L. pelidnus, or a mixture of the two when he wrote his description of L. variegatus, and his illustration of the latter (C. L. Koch, 1863: fig. 144) could apply to either species, the obvious course is to select a specimen of the former as lectotype in order to preserve current nomenclature. Females of L. mutabilis and L. pelidnus are very difficult to distinguish from one another and it has already been shown how L. Koch came to confuse them with those of L. muticus (see p.117), so most of his misdeterminations are easily explained: but the two males of L. erythrocephalus from the Franconian Jura and the two unexpected species from Munich bear only a superficial resemblance to L. mutabilis and cannot have been examined very carefully. Folkmanova (1949) pointed out that many of the infraspecific forms of L. muta- bilis enumerated and keyed by Verhoeff (1935) are based on unstable characters and are therefore without validity. Of Koch’s specimens, the single male from the Tyrol is quite without posterior projections on T. 11 and 13 and agrees in other respects with Verhoeff’s definition of L. mutabilis mutabilis; but all his other specimens including the lectotype have at least traces of posterior projections on T.11 and small but quite distinct projections on T.13, and would therefore run to L. mutabilis var. carpathicus in Verhoeff’s key. Although L. mutabilis and L. pelidnus are both fairly adequately described by Latzel (1880), the best descriptions of these two species, which are accompanied by illustrations of the male 15th legs upon which their differentiation largely depends, are those of Matic (1966). 41. Lithobius cinnamomeus L. Koch Lithobius cinnamomeus L. Koch, 1862 : 77, fig. 35. TYPE LOCALITY. Germany. MATERIAL EXAMINED. “Lithobius cinnamomeus L.K.” “Happurg [Franconian Jura]’”’ B.M.(N.H.) Reg. no. 13.6.18.60-63. Two males and two females, all either 4th post-larval stadia or small adults of L. muticus. TYPE SPECIMENS. The original description of L. cinnamomeus was based on a number of specimens of both sexes and all theabove examples of L.muticus definitely belong to the syntypical series. Remarks. As well as describing this form as being smaller, paler, and with fewer antennal articles and fewer ocelli than L. muticus, Koch noted the incurved internal pair of spurs on the female gonopods which he contrasted with the straight spurs 136 E. H. EASON which he believed, mistakenly as we have seen, to occur in L. muticus (see p. 117). Although he made no mention of the swelling on the male 14th tibia in his description, he did confirm its presence in L. cimnamomeus in private correspondence with Latzel (Latzel, 1880: 119). Latzel proposed L. cimnamomeus as a possible synonym of L muticus but he remarked on the large head of the latter compared with the rel- atively small head of the former, and suggested that the two might possibly prove to be distinct species. But it is only in the largest males of L. muticus that the shape of the cephalic shield is really distinctive and there is no doubt that L. cin- namomeus is a synonym of L. muticus. 42. Lithobius lucifugus L. Koch Fig. 5 Lithobius lucifugus L. Koch, 1862 : 82, fig. 38. TyPE LOCALITY. Bolzano, Italy. MATERIAL EXAMINED. “L. lucifugus [rewritten]” ““Botzen [Bolzano]’’ B.M.(N.H.) Reg. no. 13.6.18.369. A single male 15 mm long with circular coxal pores. “L. lucifugus [rewritten] ‘““Ratzes [Rasa, Italy], [leg.] Milde” B.M.(N.H.) Reg. no. 13.6.18.370. A female of L. pelidnus. “lucifugus?, Nurnberg” B.M.(N.H.) Reg. no. 13.6.18.371. A male 3rd post- larval stadium of L. mutabilis. “TL. lucifugus? cinnamomeus? Tirol [rewritten]’’ B.M.(N.H.) Reg. no. 13.6.18.372. Two males and three females ot L. lucifugus. TYPE SPECIMEN. The original description of L. lucifugus was based on a single male and agrees exactly with the above male from Bolzano which is undoubtedly the holotype. REMARKS. The circular coxal pores of the holotype explain why L. lucifugus is so far removed, in Koch’s system, from L. alpinus and L. granulatus with which it is conspecific (see p. 130). The prosternum of the holotype (Fig. 5) with 2 +4 2 teeth establishes this number of teeth as typical, but the appearance of 2 + 3 prosternal teeth on one of the specimens from the Tyrol is evidence of their varia- bility in L. lucifugus. Koch's inclusion of a female of L. pelidnus under L. lucifugus is a further example of the mistakes he-made in placing females of similar species. L. lucifugus was fully described by Latzel (1880 : 120) and Brolemann (1930). 43. Lithobius lubricus L. Koch Lithobius lubricus L. Koch, 1862 : 86, fig. 41. Type LocaLity. Nuremberg district. MATERIAL EXAMINED. “‘L. lubricus [rewritten]? “frank. Jura” B.M.(N.H.) Reg. no. 13.6.18.373-374. Three post-larval stadia of L. calcaratus, one belonging to the 1st, a male to the 2nd and a female to the 3rd. “L. lubricus [rewritten]” “Dietenhofen [Franconian Jura]’’ B.M.(N.H.) Reg. no. 13.6.18.375. A male 3rd post-larval stadium of L. calcaratus. THE LITHOBIUS SPECIES OF C. L. AND L. KOCH 537; “L. lubricus [rewritten]’’ ““Gritz [Griitz, near Nuremberg], [hab.] Valzn. Weiher, Glaishammer” B.M.(N.H.) Reg. no. 13.6.18.376-379. Four post-larval stadia of L. calcaratus, one belonging to the 1st, and two males and a female to the 3rd. TypE SPECIMENS. The original description of L. Jubricus was based on a number of specimens of both sexes. All the above examples of L. calcaratus come from the neighbourhood of Nuremberg and seem to belong to the syntypical series. Remarks. Koch’s failure to identify these specimens as L. calcaratus is easily understood, as the characteristic femoral process on the male 15th leg does not become obvious during the development of this species until the 4th post-larval stadium (see p. 109). Synonymy of L. lubricus with L. calcaratus was first proposed by Stuxberg (1871) and has never been disputed. 44. Lithobius carinatus L. Koch Fig. 6 Lithobius cavinatus L. Koch, 1862 : 87, fig. 42. TYPE LOCALITY. Greece. MATERIAL EXAMINED. ‘‘Lithobius carinatus L.K.” “Griechenland” B.M.(N.H.) Reg. no. 13.6.18.56-58. Three imperfectly cleared males with the antennae and all the legs missing. “Lithobius carinatus L.K.” ‘‘Patria?’”’ B.M.(N.H.) Reg. no. 13.6.18.59. A single well-preserved male. TYPE SPECIMENS. The original description of L. carinatus was based on a number of males. The above specimens from Greece seem to constitute the syntypical series and must have been examined by Koch before they lost their appendages. One of them, a male 24 mm long, is here formally designated as the lectotype (B.M.(N.H.) 13.6.18.56). Remarks. All the above specimens as well as Koch’s description are clearly referable to the common Greek species known as L. macrops Karsch, 1888. Although this description is quite adequate it has been overlooked by most authors: only Attems (1926) has recognized L. carinatus as the correct name for L. macrops. The striking difference between the relatively dense setae on the 15th prefemur and femur, and the very much sparser setae on the corresponding tibia, tarsus and metatarsus which Koch described, cannot be confirmed in the type specimens owing to their mutilation, but is present in the male from an unknown locality. However, three males and a female of this species (B.M.(N.H.) Reg. no. 89.3.29.36-38) from Athens, the type locality of L. macrops, are variable in respect of this character: one male and the female are similar to Koch’s specimens but with more setae at the base of the 15th tibia, whereas the other two males have setae of much the same density on all the articles of the 15th legs: none of these specimens from Athens shows the 138 E. H. EASON sharp difference between the setae of the femur and tibia which is so striking in Koch’s specimen. It would, in fact, be quite reasonable to regard L. carinatus and L. macrops as subspecifically distinct if we knew the exact localities in Greece attaching to L. carinatus and if it were not for other specimens of the species (B.M.(N.H.) Reg. nos. 1905.8.24.77 and 03.8.25.23-25) from unknown localities in Greece showing various degrees of differentiation between the more setose proximal and the almost glabrous distal articles of the 15th legs. Further, Matic figured a specimen of L. macrops from Athens with a glabrous 15th metatarsus (Matic e¢ al., 1968: fig. 1B). L. carinatus should, therefore be regarded as the senior synonym of L. macrops. Although this is an abundant and distinctive species the only really full account in the literature is that of Matic (Matic et al., 1968), and because Koch’s specimens differ in detail from that account they are described below. The characters of the appendages are taken from the specimens from an unknown locality. DESCRIPTION. Size: 20 to 25 mm long and about 2:5 mm broad at T.10. Colour: dull yellow. Head: broader than long. Antennae: one-third of body- length; of 32 irregular articles, some broader than long, others slightly elongate, appearing only very sparsely setose although many of the setae may have been lost. Ocelli: a large posterior ocellus, an intermediate ocellus of much the same size and two much smaller anterior ocelli, exactly as figured by Matic (Matic et al., 1968: fig. rE) ; organ of Témésvary rather smaller than smallest ocellus. Prosternum: with 2 + 2 teeth and a pair of lateral spines; lateral to the lateral spine the anterior border forms a narrow but distinct shoulder, sometimes amounting to a rounded node (Fig. 6). Tergites: the posterior angles of the large tergites all rounded, those of T.9 obtuse, those of T.11 right-angled and those of T.13 very slightly pro- duced; T.14 relatively broad; intermediate tergite (T.16) truncate. Sternites: S.5 to S.15 beset with minute setae; many of these setae have been lost but their in- sertions are visible. Coxal pores: 4, 3, 3, 3; circular; the medial pore on the 12th coxa is much smaller than the others and may be hidden by the adjacent sternite. Legs: the 14th and 15th short and stout, less than one-third of body-length; 15th prefemur and femur densely setose; the three distal articles of the 15th leg almost glabrous; setae of the r4th leg arranged in much the same wayas on the I 5th, but their differentiation is less marked; two ventral rows of stout seriate setae on the Ist to 13th metatarsi, extending onto part of the adjacent tarsus in some legs; 15th acces- sory apical claw about two-fifths of length of the principal claw which is short and stout. Gonopods: of a single article. Spinulation : Ventral Dorsal € t 12) F ag (E t P F ay 14 — m amp amp am — — mp Pp p 15 — m amp am — = = Pp Pp — No coxolateral spines. All spines rather short and stout. Koch recorded a third ventral spine on the 15th femur and a single ventral spine on the 15th tibia. THE LITHOBIUS SPECIES OF C. L. AND L. KOCH 139 45. Lithobius pubescens L. Koch Lithobius pubescens L.. Koch, 1867 : 898. Type Locality. Tinos, Aegean Archipelago. MaTERIAL EXAMINED. “‘L. pubescens [rewritten]’”’ “Tinos Erber” B.M.(N.H.) Reg. no. 13.6.18.580. A male of L. carinatus 18 mm long. “L. pubescens [rewritten] ‘“Syra Erber” B.M.(N.H.) Reg. no. 13.6.18.581-582. A male and a female of L. cavinatus 16 mm and 12 mm long respectively. “L. pubescens [rewritten]’’ “Smyrna Erber’ “181 [printed]’”’ B.M.(N.H.) Reg. no. 13.6.18.583-585. A male and two females of L. carinatus 16 to 19 mm long. TYPE SPECIMEN. The original description of L. pubescens was based on a male andafemale. The latter has not been found but the above male of L. carinatus from Tinos agrees exactly with the description and is here formally designated as the lectotype. Remarks. Although Koch made no comparsion between L. pubescens and L. carinatus, his brief description of the former is quite clear and Karsch (1888) re- marked on the similarity between L. pubescens and L. macrops (= carinatus). The principal features which seem to have led Koch to describe L. pubescens as a distinct species are, as its name implies, the strongly setose antennae and 15th legs and the numerous minute setae on the posterior sternites of the male. The distal articles of the legs, particularly those of the 14th and 15th, of the lectotype and the specimens from Syria and Smyrna are certainly more densely setose than those of the Greek specimens of L. carinatus, but there is little true difference between the two groups of specimens in the setation of the antennae and the extent and density of the sternal setae. Most of these setae have been lost in Koch’s specimens of L. carinatus and may not have been present even when he examined them originally. There would, in fact, be little reason for regarding pubescens as distinct from carinatus were it not for a marked difference in size. Of the published figures for the lengths of Greek specimens, Karsch (1888) gave 23 mm, Verhoeff (1899) gave 20:5 to 21-5 mm and Matic (Matic et al., 1968) gave 22 to 30 mm; Koch’s specimens of L. carinatus are 20 to 25 mm and the other adult Greek specimens in the British Museum (N.H.) already referred to are 21 to 25 mm long ; two smaller females (B.M.(N.H.) 03.8.25.24.and 25) from Greece are 13 mm and 14 mm long but they are obviously immature with 3, 3/2, 2, 2 coxal pores and only 2 + 2 very unequal spurs on the gonopods. Comparable figures for the lengths ot specimens from Asia Minor and the Levant are that of Porat (1894) for a Syrian specimen (15 mm) and those of Verhoeff (1925, 1941, 1943) for specimens from Jaffa (13 to 14 mm), the Taurus Mountains (15 mm) and Alexandretta (19 mm); and of Koch’s specimens of L. pubescens, the lengths of which have already been given, the female from Syria only 12 mm long seems to be mature with 4, 3, 3, 3 coxal pores and fully developed gonopods with 3 + 3 spurs. There seems, therefore, to be some justification for retaining the name pubescens for a subspecies of L. cavinatus occurring in the Aegean Archipelago, Asia Minor and the Levant and differing from the nominate subspecies in being less than 20 mm long 140 E. H. EASON with the tarsi and metatarsi of the legs, particularly those of the 14th and 15th, more strongly setose. 46. Lithobius litoralis L. Koch Lithobius litoralis L. Koch, 1867 : 899. Type Locatity. Tinos, Aegean Archipelago. TYPE SPECIMEN. The holotype. B.M.(N.H.) Reg. no. 13.6.18.368. Remarks. This species has been discussed in a previous publication (Eason, 1970a) in which it was shown to be a valid species of Eupolybothrus Verhoeff and not a synonym of E. fasciatus (Newport) as was previously supposed. 47. Lithobius nigripalpis L. Koch Lithobius nigripalpis L. Koch, 1867 : 899. Type LocaLity. Tinos, Aegean Archipelago. MATERIAL EXAMINED. “‘L. nigripalpis [rewritten] “Tinos Erber’’ B.M.(N.H.) Reg. no. 13.6.18.575. A single male. TYPE SPECIMEN. The original description of L. nigripalpis was based on a single male and agrees fairly well with the above specimen which is undoubtedly the holo- type. REMARKS. There has been uncertainty about the identity of this species ever since Verhoeff (1899) redescribed it as a subspecies of L. forficatus with either a simple apical claw on the 15th leg or with only a minute accessory claw. In subse- quent keys and brief accounts (Verhoeff, 1925, 1937 etc.) the 15th legs of L. nigripalpis have always been described as having a simple claw. In fact, not only the holotype but also three specimens (a male and two females) from the Verhoeff Collection in the British Museum (N.H.) labelled “Lithobius forficatus nigripalpis Koch, Greece” (Reg. no. 03.8.25.64-66) all have small but distinct 15th accessory apical claws, and there is no doubt that all four specimens belong -to L. bulgaricus Verhoeff, 1925, which thus becomes a junior synonym of L. nigripalpis. It may be that when he wrote his account of bulgaricus, Verhoeff had already sold his material belonging to nigripalpis to various museums and had no specimens available for re-examination ; otherwise he would hardly have described L. bulgaricus as distinct. Having decided that L. nigripalpis is identical with L. bulgaricus it remains to arrive at its taxonomic status. It clearly belongs to the piceus-peregrinus group of species and, contrary to Verhoeff’s mistaken conception of the form as having close affinity with L. forficatus, most authors have regarded it as no more than a variety or even a synonym of one of the species of this group. Latzel (1880 : 65) included L. nigripalpis among the doubtful synonyms of L. picews; Attems (1905) regarded L. nigripalpis and L. peregrinus as varieties of the same species; and Mural- witsch (1926) believed L. forficatus nigripalpis of Verhoeff to be identical with L. viriatus Sseliwanoff, 1880, another member of the piceus-peregrinus group. THE LITHOBIUS SPECIES OF C. L:. AND L. KOCH 141 Although Matic (1964) proposed nigripalpis as a variety of L. bulgaricus, he has recently (Matic, 1966; Matic & Darabantu, 1968) listed L. nigripalpis among the synonyms of L. piceus and described L. bulgaricus separately. L. bulgaricus was originally described as a subspecies of L. piceus (Verhoeff, 1925) but Matic (1966) found these two forms to be sympatric in parts of Rumania so they can hardly belong to the same species. L. nigripalpts is, in fact, closer to L. pere- grinus than to L. piceus but the absence of a prosternal diastema and the absence of denticles on the claw of the female gonopod (both of which are present in L. pere- grinus) are sufficient grounds for regarding it as a true species. The original record from Tinos, Verhoeff’s (1899) records from the island of Aegina in the Saronic Gulf and the adjacent mainland of Attica (probably based partly on the specimens B.M.(N.H.) Reg. no. 03.8.25.64-66), Verhoeff’s (1925) original record of L. bulgaricus from Ruschuk on the Danube, and the distribution Matic (1966) gave for L. bulgaricus show this species to be widespread in the eastern and southern Balkans as well as in the Aegean Archipelago. Records of L. piceus olympicus Vethoeff from the south Marmoran coast (Verhoeff, 1944), and of L. politicus Chamberlin from southwest Anatolia (Chamberlin, 1952), both probably refer to L. nigripalpis and suggest that the species is also widespread in Asia Minor. Attem’s (1905) description of L. peregrinus from Erdschias-Dagh (Asia Minor) and Verhoeff’s (1944) description of what he regarded as the true L. nigripalpis from Ankara must both have been based on specimens of nigripalpis in which the 15th accessory apical claw was either absent or so small as to be overlooked. The only really full account of this species in the literature is that of Matic (1966) under L. bulgaricus. Because the holotype appears to be a pseudomaturus and therefore different in detail from Matic’s account it is described below. Description. Length: 19 mm. Colour: dark brown. Antennae: 8 mm long; of 48 articles. Ocelli: 1 + 4, 3, 2. Prosternum: with 4 + 4 teeth and the lateral spines lateral to the external teeth. Tergites: the posterior angles of T.8 and 10 rounded, those of T.12 blunt, those of T.14 angulated; posterior angles of T.g, 11 and 13 with prominent projections, those of T.13 being very long and sharp; pos- terior border of intermediate tergite strongly emarginate; the shape of the-tergites is in marked contrast to that in L. picews in which the posterior angles of T.10, 12 and 14 are sharp and slightly projecting. Cowal pores: 5, 5, 5, 4; circular. 15th legs: 7 mm long; stout; a feeble external sulcus on prefemur, femur and tibia; accessory apical claw about a quarter the length of principal claw. Spinulation : Ventral Dorsal Cc t 12 F ap (e t 12) F at I — — mp amp am — — mp a—p a 2 — = mp amp am = = amp a-—p a—p 3 = — mp amp am = _ amp a-—p a-p 4 = = mp amp am _— — amp a-—p a-—p 5 = —_— mp amp am a — amp a-p a—p 6 —_— —_— mp amp am — — amp a-—p a—p 7 —_— — amp amp am — — amp a-—p a-—p 142 E. H. EASON Spinulation : Ventral Dorsal Cc t 12 F an € t P F T 8 —_ — amp amp am — —_ amp a—p a-—p 9 -- —- amp amp am = -= amp a—p a-—p 10 _— = amp amp am a _- amp a-p a—p II _- _ amp amp am a = amp a—p a—p 12 — — amp amp am a — amp a—p a—p 13 a m amp amp am a — amp P a-p 14 a m amp amp am a — amp P p 15 a m amp amp a a — amp Pp — 15 VaC is duplicated on the right leg. 48. Lithobius asperatus L. Koch Lithobius asperatus L. Koch, 1878 : 788. TYPE LOCALITY. Japan. Remarks. L. asperatus was originally described from specimens belonging to a collection made by Dr. Albrecht von Roretz which has not been traced. There is, however, no doubt as to the identity of this species which is very common in Japan and much of eastern Asia and whose life-history has been studied in as much detail as that of any species of Lithobiidae (Murakami, 1958). L. asperatus has been redescribed by Haase (1887) from the Phillipines and by Attems (1909) from Japan. Chamberlin (1920) considered that Attem’s description, which gave 13 ocelli (Koch gave 23, Haase gave Ig to 23) and a ventral spine on the 15th tibia (neither Koch nor Haase gave this spine), applied to another species with fewer ocelli and more spines which he named Bothropolys spinostor on the basis of Attem’s description. But specimens in the British Museum (N.H.) from Japan (Reg no. 1937.9.9.55) and southeast Korea (Reg. no. 93.3.27.6) have 24 and 22 ocelli respectively and in both, the spinulation of the r4th and 15th legs is exactly as described by Attems: there is therefore no correlation between the number of ocelli and the spinulation, and B. spinosior is not a valid species. Another of Attem’s descriptions of L. asperatus, based on a single male from the Hawaiian Islands (Attems, 1903), was questioned by Chamberlin (1920) with more justification: this Hawaiian specimen had very deficient spinulation compared with the typical L. asperatus and Attems himself had already referred it to a separate species, Bothropolys maluhianus (Attems 1914): but Chamberlin, who cannot have read Attem’s later paper, renamed it B. oahuanus. L. asperatus belongs to the genus Bothropolys Wood as emended by Chamberlin (1925a) who divided the genus into Bothropolys s.str. and Poropolys, the former with and the latter without posterior projections on T.6 and 7. In B. asperatus the projections on T.6 are very feeble and were not even mentioned by Koch in his original description, while those on T.7, though distinct in some specimens, are so feeble in the example from Japan in the British Museum (N.H.) that one cannot say to which of Chamberlin’s subgenera it belongs: Poropolys should therefore be disregarded. THE LITHOBIUS SPECIES OF C. L. AND L. KOCH iM a 0-1 mm 0-1 mm , 05 05mm romm 4 \ t 5 ra 05mm 10mm Fics 1-7. Fig.1. L. agilis, spurs of left gonopod of neotype, ventral. Fig.2. L. erythro- cephalus, spurs of gonopods of neotype, ventral. Fig. 3. L. alpinus, dental margin of prosternum of holotype, ventral. Fig. 4. L. sulcatus, ocelli of lectotype. Fig. 5. L. lucifugus, dental margin of prosternum of holotype, ventral. Fig. 6. JL. carinatus, dental margin of prosternum of lectotype, ventral. Fig. 7. L. piceus, left 15th leg of female (Franconian Jura), dorsal. (4hgr ‘yooy “T 'D) sadissoad snayjoqhjodny = adAquss [erreaeg] yostmiey zogr “MI snaysaf ((eanf weruoouer7] Sinddezz) “IYS "Ss sn1qoyjvT prea adAjoon euearqg Lbgi “yyy smjvydas0ayjhaa “YS "Ss sn1qoyjrT Pea (70114sStp Srequrein yy) edAzoany Aueuey bgt “MTD snynjuap ‘aou ‘uAs “Lb61 ‘esyoT (snigosanjouopy) 1ys0zvq (snisqoanjouopy) “TJ = SNIQOYIVT pea (B1aquioinyy) adAzoan eueaeg Ltgi “MTD sadyano (sn2qosanjouo py) sniqoyjvy prea (einf{ ueruoouesy) adAyz0}0077 POII4SIp SioquieinyNy zggr "MT sadissvaa (g$21 ‘uury) smpvoysof-7J = (eanf{ uermoours7) sodAquds Aureus zggI “MT snaov1409 (orr ‘d aas) ZOQI ‘YOY “T syzqvinu “J = — Aueuay +ttgt MTD stunuumoo A ((einf ueruosuer7] Sinddeyy) 9 Lbgr ‘yooy "J “OD snoynu “JT = sadAqyuds Aueuag zggt “MI Snamompuura ee SSgi ‘yosrey sdossvm “TJ = : “I}S "S sn1qoyjrT pea adAy0},09T a09015) zggI “MT snpourava 1G 5 “IS “S sniqoynT prea (S1equioinyy) adAzooyy Aueueg +bgr MT snywsv2jv9 <3) (gz1 ‘d aas) -aou ‘udS “zLgi ‘rouley sidsna14 “T= — ZOL4SIp yOtunyY z9gt “MI snjuajynzong stjodoayjog pea — uedef gdgr “ST snjqvsadsv “aou “uA ‘SE61 ‘yaoyI0A Yyazv) “IVA snanfion *T = Steir ‘1aaF{ snurdjy “7 uou adAjojoH {jorAy, weezy] edjerastag zggr “MT snuidjo ((Brequieiny] JrOpyesoqn) “ays “S sn1qoynT prea adAjoony eueaeg bgt M'TO syn (snzqosanjouo jy) sniqoyjvy pyea (einf{ wermooursg) adAéq0},097 USIP Siaquieinyy zggr “MI Snsoutsnaav UOTPBOYISSE]D OIAUay) snjejs pue AyIpyeA Teuezeur ad4q 93 eusIsaq peystqnd se Ayipeoo] adé yp ayeq seroeds [euTWION, TT 144 I alavy 145 L. AND L. KOCH THE LITHOBIUS SPECIES OF C. sn1qoys2}0aN prea =z [y's'A] sueeiO MON 72981 “IT #epsou sadissoas snayjogdjodny = — yorsy yynog $= Ltgr MT O snuvjuom (giz ‘d 9as) sazquinu “7 = — eueaeg bgt MT O sninum uniqnp uamou —_— Aueuray zggr YT sna sdouvjau “J = sedAquds (einf{ ueruoourrg) Squeiyy «zOgI “TO smpvydazouvjam ‘aou ‘uAS Oger ‘jaz}e'] sndosvjnv “JT = ([81equiainyy] zynI5) [Ajeqy]] ouezjog “YS 'S sn1qoypvT prea adA30399°T ‘einf{ wetuoourry {S1oquioinNy z9gI YT snquajovu “IYS "S sn1goyjrT prea adA}oj0oH] (Ayez]] Ouezjog zggr “MT snanfion} tter Yyooy "I ‘OD snjvsv9j09 «JT = sadAquds POUYSIP S1oquieinyY = zOgI “MI snauaqny “ys ‘Ss snayjoqdhjodny prea adAjo[0F{ (osejedrys1y uvosay] soury, ogt "MT S1p403y *IYS 'S sn1goyjrT prea —_— {ureds] eseyepy 9Sgt "MT stmaaur (snayjoqajodon y) snayjoqajogny prea — ysvoo ueesty bgt MTD snssaadur ther ‘YoY “I “O snyvjuap “JT = (S1oqureinyy) sedAyudg Aueuriay zggr MT syzqninmun (eyeuTeq purpeuryy) [yNyspueT snyooyfaof *J = sodAyudg !SIaqUIGINNY ZOgT “SUT stsuazsoy ([erarjsosno x] eftzpy) “Is “Ss snayjoghjodny prea adAj0[07] ysorry, «= LbgI “M'TO sadissoad “aou “uAS *ZQgI ‘YOoy “I snsnfiony “JT = ad Ao0FY uMOUyUL, zOgI yUvT snqojnuvad Stgr y1odmony sdounjam “JT = — euevaeg Lbgr SUT O snyvaqnia {einf{ ueruoours.y] Simqueryy ZOQI ‘YOoy “TJ snag -T = sedAquds {(B1aquieinyy reou) zyNIg TORI “MT 4ossof “ST Snsopita snywoysof “JT = adAjo[0F{ sdiy ueeareg zgogi “rea snqootfsof UOT}PLOYISSe]D Ol1eues) snze}s pue AzIpyeA yeuszyeu od4y oyeustsaq peysrqnd se Ayryeo0] adAy, ayeq soroeds [eUuIWION, a i sdounjaum “JT = ad Ajo] 07] (einf{ ueruoouesg) Sinqueryq = z9gI “MT 40jnuan qOU4sIp PUUOTA ‘eIUOoOUeIT sdouvnjam "JT = sedAquds ‘(ayeureg puejeutyy) [ynyspuey zogr "MT *ojaa uniqnp uamou — eurvaeg Lbgr “MTD sntava snyooyfaof -J = adAquds {nzeig] eryeg zogi “MT snyvaunts sn1qoyj1oaN prea — Cvs'n] sues19 men z9gI “MT snuivmsunag A (g9g1 ‘y19UTey, UOT) 2 (snigosavjouopy) ¢1€61 ‘aoyIa, : sdosim “TJ = ([S1aquieinyy] zyn15) a Sn1qgoyjvy prea adA40300'T Biaquiainyy zggt “MT snjzogjns ea] : snang “JT = = PUASIpP YOUN, + zOgI ST snpipaos q . uniqnp uamou — ysoury, §=Ltgi “M'TO snivjnyound Q “AOU “QUIOD “zggI ‘YOY “T SnyDUIAdI “TJ JO setoadsqns ad440}09'T {osejadryory uvasey] soury Logr “MT suassaqnd ‘IYS "S 9n1QOY}1T prea — [erreaeg] yostmrey 9 zggt “MT snaod snyvoyasof *T = — sueq zogi “MT stsuatstavd ‘aou ‘uAS “Sz61 ‘yooyIaA snaiavsng snaaidg “7 = “IYS 'S sniqoyntT prea adAjooxy {osejadryory uevasey] soury Logi MT stdjudussin “IYS "S sn1qoyj1T pyea (einf{ uermoouerq) odAjzoan eueaeg bgt MTS snaynu “IS "S sn1qoyjrT prea (eainf{ ueruooues.yz) adAj40}00T Aueway z9gI MT syiquinu snqvotfaof *JT = adAjo[0H{ Aureus zggi MT nso2snm S wol}eOYISse[O oLIeUaD snjzejs pue AzIpye, Telisyeur odAj 97eusIseq peysqnd se Ayre00] adAT, ayeq soroeds [eurmon 14 —_—C ll? nn "ee THE LITHOBIUS SPECIES OF C. L. AND L. KOCH 147 ACKNOWLEDGEMENTS My sincere thanks are due to the Trustees of the British Museum (N.H.) for lending me specimens; to Dr. G. O. Evans, Dr. J. G. Sheals and Mr. K. H. Hyatt of the Arachnida Section of the Department of Zoology of the Museum for their per- sonal help; to the Zoological Museum, Berlin and Dr. M. Moritz tor lending me specimens; and to Professor O. Kraus of the Zoological Museum, Hamburg and Dr. E. Popp of the Zoologische Staatssammlung, Munich for their help in tracing the history of the Koch and Keyserling Collections. REFERENCES AttEems, C.G. (1903). Beitrage zur Myriopodenkunde. Zool. Jb. (Syst.), 18 : 63-154. (1904). Central und hoch-asiatische Myriopoden. Zool. Jb. (Syst.), 20 : 113-130. (1905). Myriopoden. In Ergebnisse einer naturwissenschaftlichen Reise zum Evdschias- Dagh (Kleinasien). Penther, A. & Zederbauer, E. (eds.). Annln naturh. Mus. Wien, 20: 163-167. (1909). Die Myriopoden der Vega-Expedition. Ark. Zool. 5(3) : 1-84. (1914). Die indo-australischen Myriopoden. Arch. Naturgesch. 80, Abt.A, 4 Heft : 1-398. (1926). Etude sur les Myriopodes recueillis par M. Henri Gadeau de Kerville pendant son voyage zoologique en Syrie (Avril—Juin 1908). Rouen : imprimerie Lecerf. (1927). Myriopoden aus dem nordlichen und éstlichen Spanien, gesammelt von Dr. F. Haas in den Jahren 1914-1919. Abh. senckenb. naturforsch. Ges. 39 : 233-290. (1929). Die Myriopodenfauna von Albanien und Jugoslavien. Zool. Jb. (Syst.), 56: 269-356. Botiman,C.H. (1893). The Myriapoda of North America. Bull. U.S. natn. Mus. 46 : 1-210. Borex, V. (1967). Beitrag zur Kenntnis der Variabilitat der Art Lithobius lucifugus L. Koch, 1862 (Chilopoda). Vést. ¢sl. Spol. zool. 31 : 109-115. BroLemann, H. W. (1896). Liste de Myriapodes des Etats-Unis, et principalement de la Caroline du Nord. Annls Soc. ent. Fr. 65 : 43-70. (1898). Myriapodes d’Ahusquay (Basses Pyrénées). Feuille jeun. Nat. (3) 28 : 200-203. (1909) Os Myriapodos do Brazil. In Catalogos da Fauna Brazileiva 2. Sao Paulo. (1926). Myriapodes des Pyrénées-Orientales. Bull. Soc. Hist. nat. Toulouse, 54 : 233-267. (1930). Myriapodes. Chilopodes. Faune Fr. 25 : 1-405. CHAMBERLIN, R.V. (1911). The Lithobiomorpha of the Southern States. Ann. ent. Soc. Am. 4 : 32-50. —— (1920). The Myriapoda of the Australian region. Bull. Mus. comp. Zool. Harv. 64 :1-269. (1925a). The Ethopolidae of America north of Mexico. Bull. Mus. comp. Zool. Harv. 57 : 383-437. (1925b). The genera Lithobius, Neolithiobus, Gonibius and Zinapolys in America north of Mexico. Bull. Mus. comp. Zool. Harv. 57 : 439-504. (1952). On the Chilopoda of Turkey. Istanb. Univ. FenFak. Mecm. (B)17 : 183-258. Dara Torre, K.-W. von (1882). Beitrage zur Arthropoden-Fauna Tirols. Ber. naturw.- med. Ver. Innsbruck, 12 : 32-73. DEMANGE, J.-M. (1958). Sur quelques Myriapodes cavernicoles de France et de Suisse. Revue suisse Zool. 65 : 843-855. DosroruKa, L. J. (1962). Uber Lithobius erythvocephalus C. L. Koch, 1847 (Chilopoda). Zool. Anz. 168 : 43-45. Eason, E. H. (1951). Notes on the Chilopoda (centipedes) of Warwickshire and Worcester- shire. Ann. Mag. nat. Hist. (12)4 : 257-268. (1964). Centipedes of the British Isles. London: Warne. —— (1965). On Lithobius tricuspis Meinert (Chilopoda, Lithobiidae) in Britain. Ann. Mag. nat. Hist. (13)8 : 285-295. 148 E. H. EASON Eason, E. H. (1970a). A redescription of the species of Eupolybothrus Verhoeff s.str. preserved in the British Museum (Natural History) and the Hope Department of Zoology, Oxford (Chilopoda, Lithobiomorpha). Bull. By. Mus. nat. Hist. (Zool.) 19 : 289-310. —— (1970b). The Chilopoda and Diplopoda of Iceland. Entomologica scand. 1 : 47-54. —— (1971). The type specimens and identity of the species described in the genus Lithobius by George Newport in 1844, 1845 and 1849. Bull. Br. Mus. nat. Hist. (Zool.) 21 : 297-311 Feprizz1,G. (1877). ILitobiltaliani. Atti Accad. scient. veneto-trent.-istriana, 5(2) : 184-233. FOLKMANOVA, B. (1949). Subspecie druhu Lithobius mutabilis Koch v naSich zemich. Vést. ésl. zool. Spol., 13 : 56-68. (1951). Onékterych Chilopodech novych pro Moravii. Sb. Klubu priv. Brné, 29 : 98-104. —— (1954). Prispévek k poznani slezskych stonozek z Beskyd. Privodov. Sb. ostvav. Kraje, 15 : 194-219. FoLtkmanova, B. & Lana, J. (1955). Stonozky vrchu Kotouce u Stramberka. Pyivodov. S. ostrav. Kyvaje, 16 : 506-512. —— (1960). Pyispévek k poznani stonozek Rychlebskych hor. Pyivodov. Cas. slezsky, 21 : 355-372. GaRBOWSKI, T. (1879) Phyletische Deutung der Lithobiusformen. Zool. Jb. (Syst.), 9: 244-270. Gervais, M. P. (1837). Etudes pour servir a l'histoire naturelle des Myriapodes. Annis Sci. nat. Zool., (2)7 : 35-60. Haase, E. (1880). Schlesiens Chilopoden; 1. Chilopodaanamorpha. Inaugural-Dissertation. Breslau: A. Neumann. —— (1887). Die Indisch-Australischen Myriopoden; 1. Chilopoden. Abh. Ber. K. zool. anthrop.-ethn. Mus. Dresden, 1(5) : 1-118. HeEErR, O. (1845). Ueber die obersten Granzen des Thierischen und Pflanzlichen lebens in unseren Alpen. Ziivch. Jug. naturf. Ges. Ziivich, 47 : 1-19. JeexeL, C. A. W. (1964). Beitrag zur Kenntnis der Systematik und Okologie der Hundert- fiisser (Chilopoda) Nordwestdeutschlands. Abh. Verh. naturw. Ver. Hamburg, N.F.8: ITI-153. (1967). On two Italian Lithobius species described by Silvestri, with taxonomic notes on the genus Eupolybothvus Verhoeff (Chilopoda, Lithobiidae). Beaufortia, 14 : 165-175. Karscu, F. (1888). Verzeichniss der von Herrn. E. v. Oertzen in den Jahren 1884-1885 in Griechenland und auf Kreta gesammelten Myriopoden. Berl. ent. Z. 32 : 220-224. Kocn, C. (1910). Die von Dr. Ludwig Koch in Niivnberg hinterlassene Avachniden — und Myriapoden —Sammlung. Niirnberg: J. L. Stich. Kocn, C. L. (1841). Arachnida und Myriapoda. In Wagner's Reisen in dey Regentschaft Algier 3. Leipzig. —— (1844). Deutchlands Crustaceen, Myviapoden und Avachniden 40. Regensburg. (1847). System der Myriapoden. In Kyritische Revision dey Insectenfauna Deutschlands 3. Panzer, G. & Herrich-Schaffer, A. (eds.). Regensburg. (1863). Die Myriapoden i & 2. Halle: H. W. Schmidt. Kocn, L. (1862). Die Myviapodengattung Lithobius. Niirnberg: J. L. Lotzbeck. —— (1867). Zur Arachniden—und Myriapoden-—Fauna siid-Europas. Verh. zool. — bot. Ges. Wien, 17 : 857-900. —— (1878). Japanesische Arachniden und Myriapoden. Verh. zool.—bot. Ges. Wien, 27: 735-797- LatzeL, R. (1880). Die Myriopoden der Osterreichisch — Ungarischen Monarchie, 1. Die Chilopoden. Wien : Holder. Leacu, W.E. (1814). Crustaceology. In Brewster's Edinburgh Encylcopaedia 7. Edinburgh. Liécer, L. & Duposcg, O. (1903). Recherches sur les Myriapodes Corse et leurs parasites. Archs Zool. exp. gén. (4)1 : 307-358. Licnau, N.G. (1914). Vielfiissler aus Abchasien. Ezheg. zool. Muz. 19 : 349-368. LoHMANDER, H. (1957). Faunistiskt faltarbete I Nord-och Vastjylland 1954 och 1956. Arsty. Géteborgs naturhisto. Mus., 1957 : 29-86. THE LITHOBIUS SPECIES OF C. L. AND L. KOCH 149 Loxsa, I. (1947). Beitrage zur Kenntnis der Steinlaufer—, Lithobiiden — Fauna des Kar- patenbeckens, 1. Fragm. faun. hung. 10 : 73-85. —— (1948). Beitrage zur Kenntnis der Steinlaufer-, Lithobiiden — Fauna des Karpaten- beckens, 2. Fragm. faun. hung. 11 : 1-11. Lucas, H. (1849). Histoire naturelle des animaux articulés. In Exploration scientifique de V Algerie pendant les années 1840, 1841, 1842. Zoologie 1. Paris. Matic, Z. (1961). Chilopodi, specialmente cavernicoli, raccolti in Toscana da Paola e Bene- detto Lanza e da Giorgio Marcucci. Nota II. Monitore zool. ital. 69 : 60-65. (1964). Nota critica asupra unor specii de Lithobiidae (Chilopoda) din fauna Republicii Populare Romine. Studi. Cerc. Biol. Cluj, Zool. 16 : 187-191. (1966). Chilopoda : Anamorpha. In Fauna Republicii Socialiste Romédnia. 6 Bucuresti. Matic, Z., CLicuicr, M. & DaRABANTU, C. (1968). Contributio alla conoscenza dei Chilopodi di Grecia. Boll. Sed. Accad. gioenia Sci. nat. (4)9 : 307-317. Matic, Z. & DaRABANTU, C. (1968). Note critique sur quelques espéeces du genre Lithiobius (Chilopoda, Lithobiidae). Izv. zool. Inst. Sof. 26 : 103-117. Meinert, F. (1868). Danmarks Scolopender og Lithobier. Naturh. Tidsskr. (3)5 : 241-268. (1872). Myriapoda Musaei Havniensis: bidrag til myriapodernes morphologi og syste- matik; II Lithobiini. Natuvh. Tidsskr. (3)8 : 281-344. Murakaml, Y. (1958). The life-history of Bothropolys asperatus (L. Koch). Zool. Mag. Tokyo, 67 : 217-223. (in Japanese). Muratewitscu, W. S. (1926). Ubersicht iiber die Chilopodenfauna des Kaukasus. Zool. Anz. 69 : 27-44. NecrEA,S. (1965). Contribution a l’étude de certains Lithobiidae (Chilopoda) des grottes de Roumanie. Int. J. Speleol. 1 : 287-305. Newport, G. (1845). Monograph of the class Myriapoda, order Chilopoda. Tvans. Linn. Soc. Lond. 19 : 349-439. Pocock, R. I. (1890). Contributions to our knowledge of the Chilopoda of Liguria. Annali Mus. civ. Stor. nat. Giacomo Doria, 29 : 59-68. Porat, C. O. von (1869). Redogorelse for en under sommaren 1868 utford zoologisk resa till Skane och Blekinge. Ofvers. VetenskAkad. Férh. Stockh. 26 : 631-653. (1894). Myriapodes récoltés en Syrie par le Docteur Théodore Barrois. Revue biol. N Fr. 6 : 62-79. RosENHAUER, W. G. (1856). Die Thieve Andalusiens. Erlangen: T. Blaesing. Rosicky, F. V. (1876). Die Myriopoden Béhmens. Arch. naturw. LandDurchforsch. Bohm. 3, Abt. 4, No. 7 : 1-44. Say, T. (1821). Descriptions of the Myriapoda of the United States. J. Acad. nat. Sci. Philad. 2 : 102-114. SILVEsTRI, F. (1897). Contributio alla conoscenza dei Chilopodi e Diplopodi della Sicilia. Boll. Soc. ent. ital. 29 : 233-26r. SSELIWANoFF, A. (1880). Materiali k izucheniyu russkikh tisyachenogikh. Tvudj russk. ént. Obshch. 11 : 3-26. STUXBERG, A. (1871). Bidrag till Skandinaviens Myriopodologi II. Sveriges Chilopoder. Ofvers. VetenskAkad. Férh. Stockh. 28 : 493-512. (1876). Myriopoder fran Sibirien och Waigatsch 6n samlade under Nordenskidldska expeditionen 1875. Ofvers. VetenskAkad. Férh. Stockh. 33(2) : 11-38. Topras, D. (1969). Grundsatzliche Studien zur Art-Systematik der Lithobiidae (Chilopoda: Lithobiomorpha). Abh. senckenb. naturforsch. Ges. 523 : I-51. Trorzina, A. (1893). Vier neue Lithobius—Arten aus Central-Asien. Horae Soc. ent. Ross. 28 : 247-253. VERHOEFF, K. W. (1899). Beitrage zur Kenntniss paldarktischer Myriopoden. XI. Auf- satz: neue und wenig bekannte Lithobiiden. Verh. zool. — bot. Ges. Wien, 49 : 451-459. (1905). Uber die Entwicklungsstufen der Steinlaiifer, Lithobiiden, und Beitrage zur Kenntnis der Chilopoden. Zool. Jb. (Supplement), 8 : 195-298. 150 E. H. EASON VERHOEFF, K. W. (1925). Beitrage zur Kenntnis der Steinlaufer, Lithobiiden. Arch. Naturgesch. 91: 124-158. (1931). Chilopoden der Insel Elba. Zool. Anz. 95 : 302-312. — (1934). Beitrage zur Systematik und Geographie der Chilopoden. Zool. Jb. (Syst.), 66 : 1-112. —— (1935). Quer durch Schwarzwald und schweizerischen Jura (Chiemgau), Chilopoden. Verh. naturw. Ver. Karlsruhe, 31 : 181-208. —— (1937). Chilopoden-—Studien. Zur Kenntnis der Lithobiiden. Avch. Naturgesch. N.F.6 : 171-257. ——, (1941). Asyanin zoogeografiyasi ve hayvan sistematige hakkinda. Asiatische Beitrage. Istanb. Univ. FenFak. Mecm. (B)6 : 85-117. (1943). Uber Chilopoden der Tiirkei. III. Aufsatz. Zool. Anz., 143 : 116-140. ——, (1944). Asya hayvanlari ve zoogeografiyasi hakkinda. Asiatische Beitrage VIII. Istanb. Univ. FenFak. Mecm. (B)9 : 307-347. Dr. E. H. EAson Bourton Far HILit MorETON-IN-MARSH GLOUCESTERSHIRE A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) Kay, E. Arison. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 1965. (Out of Print.) £3.75. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180; 11 Plates, 15 Text-figures. 1967. £4. Taytor, J. D., KENNEDY, W. J. & Hatt, A. The Shell Structure of Mineralogy at the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates, 77 Text-figures. 1969. £4.50. Haynes, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. aes) 1962-1964. (In press.) aut Printed in England by Staples Printers Limited at their Kettering, Northants, establishment - CONTRIBUTIONS TO THE LIFE- os HISTORIES AND DEVELOPMENT OF CUCULLANUS MINUTUS o> RWDOLPHI, 1819 AND C. HETEROCHROUS RUDOLPHI, 1802 _ (NEMATODA : ASCARIDIDA) D. I. GIBSON CONTRIBUTIONS TO THE LIFE-HISTORIES AND DEVELOPMENT OF CUCULLANUS MINUTUS RUDOLPHI, 1819 AND C. HETEROCHROUS RUDOLPHI, 1802 (NEMATODA : ASCARIDIDA) BY DAVID IAN GIBSON Pp 151-170; 3 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 22 No. 5 LONDON : 1972 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) instituted im 19409, 1s issued in five series corresponding to the Departments of the Musewm, 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. 22 No. 5 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.) © Trustees of the British Museum (Natural History), 1972 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 3 March, 1972 Price 70p CONTRIBUTIONS TO THE LIFE-HISTORIES AND DEVELOPMENT OF CUCULLANUS MINUTUS RUDOLPHI, 1819 AND C. HETEROCHROUS RUDOLPHI, 1802 (NEMATODA : ASCARIDIDA) By D. I. GIBSON CONTENTS Page 1. SYNOPSIS . 3 ° 5 é : a : b é a 153 2. INTRODUCTION . : 6 a ‘ o 0 é 4 0 153 3. Meruops . : 2 : : : : . c 5 154 4. RESULTS AND DISCUSSION . ‘ : é F ¢ ° 155 A. Cucullanus minutus é 0 3 6 0 é 155 (i) Larval and adult development 3 0 : Q 155 (ii) Life-history . . 0 - 5 : . 6 158 (ili) Discussion . 0 , é é : 5 5 158 B. Cucullanus hetevochrous . é : : a 0 F 161 (i) Larval and adult development ‘ c 5 161 (ii) Life-history . 5 é < : : 6 : 164 (iii) Discussion . : : 4 9 ¢ : : 164 5. FINAL DISCUSSION é 6 : ; 9 a ¢ : ¢ 167 >. ACKNOWLEDGEMENTS . p : : : : Q 2 : 168 7. REFERENCES 7 0 : 3 6 é ¢ 0 F : 168 1. SYNOPSIS The larval stages of the nematodes Cucullanus minutus and C. heterochrous from the flounder Platichthys flesus (L.) are described, those of C. heterochrous have not been previously recorded, Information on the incidence of infestation of flounders with these larvae throughout the year is used to make a significant contribution to our knowledge of the life-histories of these two species. The work also indicates that C. minutus and C. heterochvous are normally geographically isolated from each other, and that this is associated with temperature and their life-histories. 2. INTRODUCTION Cucullanus minutus and C. heterochrous were two of the commonest nematodes found in the flounder Platichthys flesus (L). from the estuary of the River Ythan, Aberdeenshire and from the sea off Aberdeen during the course of a survey of its helminth parasites. From previous work on the species of this genus, it appeared that the most obvious topics worthy of investigation were their life-histories and, 154 D. I. GIBSON especially in the case of C. heterochrous, the morphology of the larval stages. Janis- zewska (1939) has described the third- and fourth-stage larvae of C. minutus, but did not determine any of this parasite’s earlier life-history. With regard to C. heterochrous, no information on its life-history is available in the literature, and its larval stages do not appear to have been recorded previously. This is surprising considering how common and widespread these species are in flatfish. A study of the literature revealed that there is little information on the life- histories of many members of the genus Cucullanus, and that no life-history has been satisfactorily explained throughout the complete cycle. The only explanation of a complete life-cycle is that of Vessichelli (1910) for C. stelmuioides (Vessichelli, Ig10). Vessichelli states that the eggs are laid and hatch in the gut-lumen of the lamprey Petromyzon planeri (Bloch). The larvae penetrate the gut-wall, encyst and remain there during the period of gut reduction before spawning. After spawn- ing the lampreys perish, and lamprey larvae become infested by feeding upon the dead bodies of the adult lampreys. According to Shulman (1957) Vessichelli’s work has not been confirmed and requires further investigation. Jamiszewska (1939) showed that some of the larval stages of C. minutus were present in the gut-wall of flounders and that the final stage larvae migrated into the gut-lumen, but she could not ascertain how these larval stages came to be present in the gut-wall. Le- Van-Hoa & Pham-Ngoc-Khue (1967), when describing C. chabaudi, showed in the laboratory that the eggs of this species embryonate in two days, the first moult occurs inside the egg after four days and the second-stage larvae hatched from the eggs after five to six days. These authors then stated that the second-stage larvae continue to grow inside the swim-bladder of a fish (Pangasius pangasius Hamilton Buchanan), the second moult occurs in the liver, and the third and fourth larval stages are present in the gall-bladder and bile-duct. The fourth-stage larvae then migrate along the bile-duct to the intestine where the adult nematodes are found. These authors did not state how the second-stage infested the fish, and did not describe the larval stages. Finally, preliminary comments upon the life-histories of C. minutus and C. heterochrous, which are enlarged upon below, were made by MacKenzie & Gibson (1970). 3. METHODS The mean incidence of infestation of the adult and larval stages of these two nema- todes were obtained during a survey of the helminth parasites of the flounder. Seven hundred and forty flounders from the estuary of the River Ythan, Aber- deenshire (termed ‘estuarine flounders’) and one hundred and seventy flounders from the sea off Aberdeen (termed ‘marine flounders’) were examined. These results are represented in fig. 3. Most of the larvae were removed from the mucosa and sub-mucosa of the gut-wall by scraping, and the remainder deep in the sub- mucosa were found by examining sections of squashed gut-wall with transmitted light. The difference between mature and immature adult nematodes was taken to be, in the case of the females, the presence or absence of eggs, and, in the case of LIFE-HISTORIES AND DEVELOPMENT OF CUCULLANUS 155 the males, the presence of clearly defined spicules when viewed under a low powered (x Io) microscope; but in most cases the mature and immature could easily be distinguished by size. In the hatching experiments the adult worms were removed from the host, left over-night in 40% seawater, and the eggs produced were removed and kept in 80% seawater in a solid watch-glass. 4. RESULTS AND DISCUSSION A. Cucullanus minutus The only previous work on the life-history of this species was done by Janis- zewska (1939) who described in detail the larval stages recovered from the gut-wall of flounders from the Baltic Sea. The free-living larvae have, however, never been described. (i) Larval and adult development. a. The egg: Length : 61-68 ym Breadth: 34-38 um The eggs were prolate spheroids and contained an unsegmented ovum when they left the female. The ova were seen to cleave into two- and four-cell stages, and then after two days the morula and blastula stages were visible. After four days a nematode larva was visible, coiled and moving within the egg-shell, but hatching did not take place until about seven days at 19°C. (133 degree days). Embryon- ation was very slow at low temperatures and was not observed below 7°C. b. The first- and second-stage larvae: Length : 304-374 um Breadth : 13 pm The free-living larvae which hatched from the eggs (fig. 1) may have been the first- or second-stage larvae. No sign of a moulted cuticle was visible around the hatched larvae, though if such a cuticle had been tight fitting it may have been present. If this species resembles C. chabaudi (Le-Van-Hoa & Pham-Ngoc-Khue, 1967) then the first moult occurs within the egg, or if it resembles C. heterochrous (see below) the first larval cuticle is probably not lost until several days have passed in the free-living stage. Either way it is probable that the first- and second-stage larvae are very similar in appearance. Very little of the internal anatomy of these larvae was visible, though the anterior part of the oesophagus could be seen, showing that it was poorly developed compared with the adult. The remainder of the larva was obscured by the nuclei which, at this stage, are tightly packed together. These larvae were very active and capable of swimming. Unlike C. chabaudi, no second-stage larvae were ever recovered from the fish host. 156 D. I. GIBSON c. The third-stage larva (fig. 1): Length : 600-1,237 wm Breadth : 40-80 um Oesophagus : 245-306 ym Tail : 71-106 ym This stage was described in detail by Janiszewska (1939) as the first larval stage from the intestinal wall. These larvae were situated in the mucosa and sub-mucosa of mainly the anterior intestine. The anatomy of this stage was as described by Janiszewska (1939), except that I believe that the thickening of the cuticle on the lips close to the mouth might in fact act as a boring tooth similar to that in the in- festive stages of some anisakine nematodes. d. The fourth-stage larva (figs 1 & 2): Length : 720-1,210 ym Breadth : 58-100 um Oesophagus : 278-346 ym Tail : 80-106 pm The fourth-stage larvae, or pre-adults, were described in detail by Janiszewska (1939) as the second larval stage from the intestinal wall. These larvae were found wandering through the gut-wall of the flounder feeding upon the tissues as they moved. e. The immature adult (fig. 2): Length : 600 (contracted) —2,100zm Breadth : I13-332 pm Oesophagus : 308-572 um Tail : 75-120 um These specimens were normally less than 2 mm. in length. It was not determined whether the final moult took place actually inside the body-wall or in the gut- lumen. The immature adults are similar in appearance to the mature adults, except that the sexual organs are not fully developed. The oral spines make their first appearance in the adult stage. These nematodes are capable of a great deal of contraction, and folds in the cuticle may form collars around the body. The behaviour of these immature adults is the same as that of the mature specimens. Fic. 1. Cucullanid larvae. A. The second-stage larva of Cucullanus heterochrous within the first-stage larval cuticle. B. The third-stage larva of C. heterochrous from the flounder. C. The pre-adult of C. heterochrous. D. The first- or second-stage larva of C. minutus. E. The third-stage larva of C. minutus from the flounder. F. The pre- adult of C. minutus. (ct, cuticular thickening; ep, excretory pore; gp, genital primordium; mc, moulted cuticle; nr, nerve ring; p, oesophagus; pc, large oesophageal cell) I Loy LIFE-HISTORIES AND DEVELOPMENT OF CUCULLANUS a i=] i=] = 158 D. I. GIBSON f. The mature adult: Length : 2,000-3,800 um Breadth : 210-515 pm Oesophagus : 430-580 pm Tail : 70-160 um The mature adults have been described in detail by Gendre (1926), Tornquist (1931) and Berland (1970), and figured by MacKenzie & Gibson (1970). The females were generally longer and in most cases broader than the males. The adults live by moving freely amongst and feeding upon gut-contents, or, when food is not available, by attachment to and feeding upon the gut-wall of the flounder. Attach- ment to the gut-wall is brought about by the sucking action of the muscular oesoph- agus, by the collarette and associated oral spines situated on the lips and possibly by the spiny nature of the pseudobuccal cavity. Aspects of the behaviour of this species were discussed by MacKenzie & Gibson (1970). (ii) Life-history. In flounders from the River Ythan third-stage larvae were present in small numbers throughout the year, but increased infestation occurred during the early spring (fig. 3), especially during March and April. Fourth-stage larvae were first found in March, but their numbers increased to a maximum during May and June and then fell to a very low level of infestation from August to November. Immature adults were first recovered in April, their numbers increased to a peak in June and July, and then their numbers fell in the autumn. Mature adults were first found in May, their numbers increased to a peak in July and August, and then fell off during the autumn with one or two individuals remaining until December. In the marine flounders examined, the third-stage larvae were first recovered in January, the infestation had increased by March, but fell again by June (fig. 3). Fourth-stage larvae were first found in the March sample, the infestation was heavier in June, but had disappeared by September. Juvenile adults were recovered only from the June sample, and mature adults were found in the September and, to a greater extent, in the November samples. (iii) Discussion. The major period of egg-production of C. minutus in the River Ythan is during the summer months. This means that the eggs probably have time to embryonate and hatch during the late summer and autumn. This suggests that there is a gap of at least four to five months between the hatching of the eggs and the appearance of Fic. 2. Cucullanid pre-adults and immature adults (all drawings are of female worms). A. Pre-adult Cucullanus heterochrous. B. Immature adult C. heteyochrous. C. Pre- adult C. minutus. D. Immature adult C. minutus. (ep, excretory pore; du, developing uterus; gp, genital primordium; os, oral spines; pa, papilla; pc, large oesophageal cell; v, vulva) 159 LIFE-HISTORIES AND DEVELOPMENT OF CUCULLANUS Oo TAR 160 D. I. GIBSON the third-stage larvae in the flounders. As in the case of C. heterochrous discussed below, there are a number of possible explanations for this apparent gap : (1) an intermediate host is involved, (2) small second-stage larvae were too small to be seen during the flounder examinations, (3) there is an earlier site of infestation where the second-stage larvae develop and moult into the third-stage larvae, and (4) the second-stage larvae develop and grow in the free-living state. With regard to the first possibility, Wilker (1930) suggested that decapods and cumaceans were involved as intermediate hosts, but Janiszewska’s (1939) attempts to infest such crustaceans all failed. Markowski (1966) suggested that Nerezvs diversicolor Miller might serve as an intermediate host for C. minutus, and I too suspected this animal because it occurs in large numbers in muddy estuaries. However, my attempts to infest this species have all failed. Similarly, I have failed to infest or find cucullanid larvae in Neomysis integer (Leach), Corophium volutator (Pallas), gammarids and Crangon vulgaris L., though some development in the free-living state may be necessary before the larvae become infestive. One of the main arguments for the presence of an intermediate host is the accumulation of the majority of the larvae in the anterior intestinal wall of the flounder. The possibility that the second-stage larvae are present in the gut-wall but have not been seen is unlikely because both Janiszewska (1939) and myself have failed to detect them. The possibility that there is an earlier site of infestation in the flounder cannot be overlooked, though none were found by Janiszewska (1939) or by myself, since Le-Van-Hoa & Pham- Ngoc-Khue (1967) did find such a site when studying C. chabaudi. However, as suggested below, the appearance of the third-stage larvae occurs during the period when most of the large flounders are at sea on their spawning migration. Therefore, if the flounders are being infested by the parasite at that time, this accounts for the fact that C. minutus tends to infest the small flounders more heavily than the larger ones (unpublished information) and also accounts for the absence of these larvae in O-group flounders (those less than one-year old). If an earlier infestation of the flounders does occur then alternative explanations must be found for these two results. Evidence for the final possibility, i.e. that the larvae live for some months in the free-living state, is not very forthcoming either, except that free-living larvae are produced when the eggs hatch and that these can be kept alive for more than a week in 80% seawater. As in the case of C. heterochrous discussed below, it is not yet possible to favour any of these alternatives too strongly, though the first and the last possibilities do seem more likely. It is possible, there- fore, that the first stage-larvae moult into the second-stage larvae, which is probably the infestive stage, and these grow, with or without the use of an intermediate host, and infest the flounder mainly in the winter and early spring. Janiszewska (1939) stated that the infestation of the flounders with the third- stage larvae commenced in August, which is earlier than the major period of infest- ation recorded in flounders from the River Ythan. This difference might be ex- plained by the fact that on the coast of Poland, where Janiszewska (1939) was working, the sea will be warmer in summer than the waters of the River Ythan, and therefore development will be faster. The fact that temperature does affect the development of C. minutus is discussed below. In warmer waters, therefore, it is LIFE-HISTORIES AND DEVELOPMENT OF CUCULLANUS 161 possible that infestation occurs in the autumn and mature specimens are found in the spring and summer, while in cooler waters the main period of infestation occurs in the early spring and mature adults are found in the late summer and autumn. In marine flounders from the Aberdeen area most of the mature adults were not found until the autumn, and this may account for the fact that third-stage larvae were not found in these fish in the autumn (fig. 3). The fact that the development of this parasite appears to take so long in marine flounders from the Aberdeen region may mean that in most cases the winter arrives and the parasite dies before egg pro- duction has really got underway. This could account for the small numbers of this parasite in these marine flounders. B. Cucullanus heterochrous The only previous comments on the life-history of this species were given by MacKenzie & Gibson (1970), and there is no previous record of the larval stages. (i) Larval and adult development. a. The egg: Length: 75-102 um Breadth: 44-52 um The ova in the eggs cleaved in a similar manner to those of C. minutus, and larvae developed within the eggs after the same period of time. The eggs hatched after seven days at 19°C. and, like C. minutus, embryonation was not observed below 7°C. b. The first-stage larva: Length : 400-460 um Breadth: 12-14 ym The first-stage larva developed within the egg, and after hatching was extremely active and capable of swimming. Any details of the anatomy of these larvae were difficult to see because of its small size and heavy concentration of nuclei. However, the anterior oesophagus could be distinguished in some specimens showing that it was very different from that of the adults because the musculature was much less developed. c. The second-stage larva (fig. I): These larvae were so similar to the first stage-larvae that they could only be distinguished when parts of the moulted cuticle were still present. Such forms were found at 19°C. two days after hatching, though when the actual moult occurred is 162 D. I. GIBSON not known: this may even have been inside the egg. The oesophagus could be seen more clearly in the second-stage larvae, and was about 165 wm in length. The remainder of the alimentary canal could not be seen, and there was no anus visible. d. The third-stage larva (fig. 1): Length : 660-955 wm Breadth : 25-27 um Oesophagus : 240-293 um Tail : 78-93 um These larvae were found encysted in the gut-wall of the flounder within small round cysts. I have regarded this stage as the third-larval stage because (I) it is parasitic, (2) it is much larger than the second-stage larva, (3) there are morpho- logical differences between the second-stage larva and this larva, and (4) nothing resembling a second-stage larva was ever recovered from the flounder. The oesophagus of these larvae was better developed than in the previous stages, and the large oesophageal cells were visible, as they were in the corresponding stage of C. minutus. In the lip-region cuticular thickenings, which may act like boring teeth, were visible. In female specimens (fig. 1) a genital primordium could be seen, though, unlike the equivalent stage of C. minutus, I could not ascertain the position of the excretory gland or the excretory pore. The intestine appeared to be in the form of a tube composed of a large number of cells, and for the first time a definite anus could be seen. The third-stage larva of the closely-related C. civratus Miller, 1777, was recently described by Berland (1970); but this description resembles the fourth-stage larva of C. heterochvous. Berland’s argument for calling this larva the third-stage larva was because the development of the gonads resembles the third-stage larva of Contracaecum aduncum (Rudolphi, 1802), but there is no vulva present in C. minutus or C. heterochrous until the fourth larval stage. Therefore, because of this and the structure of the head, I consider Berland’s specimens to be fourth-stage larvae. e. The fourth-stage larva (figs I & 2): Length : 620-1,600 um Breadth : 27-40 pm Oesophagus : 240-359 ym danle 89-133 pm In these larvae, which were observed moulting from the third-stage larvae within the cysts in the gut-wall, the cuticular thickenings on the lips were lost and the oesophagus became similar in shape to that found in the adults. The fourth-stage larvae, or pre-adults, left the cysts and were recovered wandering through the tissues LIFE-HISTORIES AND DEVELOPMENT OF CUCULLANUS 163 of the gut-wall, feeding as they went. The lips possessed sharp edges which may have been associated with the movement through the tissues, because no oral spines or collarette were present at this stage. The large oesophageal cells were still visible, though they were not as clear as in the previous stage. In the female worms a developing uterus and vulva could be seen, though the vulva did not com- municate with the exterior. The excretory pore was situated about half way between the nerve ring and the posterior edge of the oesophagus. f. The immature adult (fig. 2): Length: 1I-6mm., e.g. Length: 5,380 um Breadth : 150 um Oesophagus : 800 um Tail : 173 um Oral spines and the collarette were present at this stage, and the pre-anal sucker and anal papillae were visible in males under a high powered microscope. In the females the uterus and ovaries were developing and the vulva could be seen to be connected to the exterior. These immature adults were found free inthe lumenof the gut or attached to the gut-wall. It was not ascertained where in fact the final moult occurred or whether it was the pre-adults or the immature adults which migrated into the gut-lumen. g. The mature adult: Mature adults were described by Térnquist (1931) and Berland (1970). My specimens reached up to 12 mm., in the case of the females, and 10 mm. in length in the case of the males. These nematodes wandered through the intestinal contents of the flounder feeding as they went, though, like C. minutus, when food was not available they fed upon the gut-wall. Details of the migrations of these nematodes in the gut-environment have been given by MacKenzie & Gibson (1970). In the past there has been some argument as to the nature of the structures which I have referred to as the oral spines. Gendre (1926), when discussing C. minutus, and Berland (1970) have commented that they may act as supporting ‘ribs’ for the cuticular collarette, and Berland (1970) also suggested that they may serve as an inter-locking device when the mouth is closed. My electron micrographs of C. heterochrous have shown that the spines, or denticulations, are, for at least the anterior-most part of their length, free of the collarette. They, therefore, probably aid the attachment of the parasite to the gut-wall, and may serve to split this wall and allow the muscular oesophagus to devour parts of the mucosa and sub-mucosa of the host. 164 D. I. GIBSON (ii) Life-history: The smallest larvae found in the flounders were the third-stage larvae. These were first recovered from the River Ythan in January, reaching a maximum in April, and then decreasing in numbers to zero in August, (Fig. 3). In marine flounders, however, these were found earlier, in November, when 8-8% of the fish examined contained small numbers of larvae. Due to the small size of these larvae small infestations may have been overlooked in the estuarine flounders, and this might account for this difference, but as mentioned below it is probable that there are no such larvae present in the Ythan estuary during these months. Fourth- stage larvae were first found in estuarine flounders during March. Their number increased to a maximum in May, and then decreased during the summer. In marine flounders their numbers increased during the late autumn and they were present in large numbers during the winter and spring. The dip in the level of infestation in marine flounders during the late winter may have been caused by the estuarine flounders entering the marine population at that time. Thus, as in the case of the third-stage larvae, the pre-adults are present in marine flounders long before they are in estuarine flounders. Immature adults made their first appearance in the estuarine flounders during March, their numbers increased to a maximum intensity in June and a maximum incidence in July, and then their numbers decreased to zero by the following Jan- uary. In marine flounders the numbers increased during the spring, as in Ythan flounders, but did not reach a peak until the autumn. This was possibly because the sea is cooler than the waters of the River Ythan during the summer months, and therefore development takes longer. These worms began to mature in estuarine flounders in September, mature adults increased in numbers to a maximum during the winter months and then decreased throughout the following spring. A similar effect was found in the marine flounders. (iii) Discussion The discrepancy in the periods of infestation with these larvae between the estuar- ine and the marine flounders could be the effect of temperature, since the waters of the River Ythan were warmer than the sea during the summer and cooler during the winter (H. D. Dooley, personal communication). However, there is a possibility that infestation with this parasite occurs much more frequently in the sea. This is suggested by the following facts: (1) infestation with this parasite is much heavier in marine flounders; (2) infestation with third-stage larvae proceeds earlier in marine flounders; and (3) this species tends to infest only the larger of the estuarine flounders (unpublished results) possibly because only the larger flounders participate in the spawning migration. It therefore appears that infestation of the marine flounders might occur before the arrival of the estuarine flounders in the sea, and the warmer temperature of the sea during the winter months probably allows faster development of these larvae in marine flounders. The sudden increase in the third- stage larval population in estuarine flounders in April is, therefore, probably assoc- iated with the return of some of the larger fish from the sea. LIFE-HISTORIES AND DEVELOPMENT OF CUCULLANUS 165 INCIDENCE PERCENTAGE 4 ra aA JFMAMJJASOND J Ss icaniah ie ako uuibes MONTHS MONTHS --@-- Ythan Flounders -—O Marine Flounders Fic. 3. The effect of season on the incidence of the larval and adult stages of Cucullanus heterochrous and C. minutus in flounders from the Ythan estuary and from the sea off Aberdeen. 1. The third-stage larvae of C. heterochrous. 2. Pre-adults of C. hetero- chrous. 3. Immature adults of C. heterochrous. 4. Mature adults of C. heterochrous. 5. Third-stage larvae of C. minutus. 6. Pre-adults of C. minutus. 7. Immature adults of C. minutus. 8. Mature adults of C. minutus. 166 D. I. GIBSON The difference in size between the free-living second-stage larvae and the third- stage larvae recovered from the flounders suggests that: (1) smaller second- or third-stage larvae were missed during the examinations of the gut-wall; (2) there is an earlier site of infestation in the flounder; (3) there is an intermediate host; or (4) the second-stage larvae undergo a period of growth in the free-living state. Dealing with each point in turn, the first suggestion seems unlikely as no equivalent larvae were found by Janiszewska (1939) or myself in C. minutus. However, the second suggestion that there is an earlier site of infestation is a possibility, because Le-Van-Hoa & Pham-Ngoc-Khue (1967) found second-stage larvae of C. chabaudi, which migrated to the liver for the second moult, growing in the swim-bladder. It is therefore possible that such larvae might occupy a similar site in the body of the flounder. However, as stated below, it is most probable that most of the eggs hatch in the late spring and early summer, and therefore these larvae would be expected to infest the flounders at least by the autumn, long before the estuarine flounders enter the sea. Thus the high infestation in estuarine flounders returning to the estuaries in spring and the absence of larvae in O-group flounders cannot be explained by this suggestion, unless an earlier site of infestation is associated with the third and fourth suggestions. The third suggestion, that an intermediate host is involved, is doubtful because there is no evidence for the presence of such a host either in this species or in the other species of the genus discussed earlier. The life-cycle of the Camallanidae, nematodes with a similar mode of life, does in fact involve an inter- mediate host, but these are no longer thought to be closely related to the Cucullanidae (Inglis, 1967). The final suggestion, that the second-stage larvae grow in the free- living state, is suggested by the fact that free-living second-stage larvae were kept alive for more than a week in the laboratory. Both the third and the final sug- gestions would account for the apparent gap of several months between hatching and the appearance of the third-stage larvae in the flounders, the infestation acquired by the estuarine flounders when they entered the sea, and the absence of these larvae in O-group flounders. However, there is no further evidence for the latter sug- gestion, and, as stated by MacKenzie & Gibson (1970), the free-living larvae would penetrate neither the skin nor the gut-wall of the flounder. Though the larvae used for these latter experiments were recently hatched specimens, and it is possible that some development in the free-living stage is required before the larvae become infestive. It 1s clear that, at this stage, it is not yet possible to favour strongly any one of these suggestions until more information is available. Circumstances beyond my control hampered further work on this problem, but in my opinion the latter suggestion seems at present to be the most likely possibility, though the presence of an earlier site of infestation or possibility of an intermediate host cannot yet be overlooked. In spite of the gap with regard to the actual method of infestation of the flounder, these results do give a good indication of what takes place during the life-history of this parasite. In my opinion, using the information given above, the life-history of C. heterochrous may occur as follows: Most of the eggs are produced during the winter and early spring months, they hatch in the late spring and early summer when the water becomes warm enough LIFE-HISTORIES AND DEVELOPMENT OF CUCULLANUS 167 (7°C.) and release free-living first-stage larvae. These require a number of days to moult into the second-stage larvae, which live on the sea-floor, grow and become infestive in the late autumn, winter and following spring. This coincides with the appearance of the third-stage larvae in the flounders. Most of these larvae moult into fourth-stage larvae (pre-adults) around April to May (though perhaps much earlier in marine flounders), and from then until June to July they grow inside the gut-wall feeding upon the tissues. During these latter two months many of the pre- adults moult into small immature adults, which, in the lumen of the gut, proceed to grow and slowly mature until the late autumn, winter and early spring when they begin to produce eggs. The mature adults continue to grow and produce eggs until their numbers decline in the late spring and summer. Nearly all the old adults are lost by the autumn. 5. FINAL DISCUSSION In both C. minutus and C. heterochrous in flounders from the River Ythan the main period of apparent infestation coincides with the spawning migration. The fact that larvae of C. minutus tended to infest the smaller flounders and those of C. heterochrous the larger flounders (unpublished information) might be explained thus: C. minutus is a very common parasite of flounders from the Ythan estuary (Gibson, 1972), and as the smaller flounders do not take part in the spawning migration they are much more available to the parasite at this time, and, similarly, C. heterochrous is a very common parasite of flounders from the sea off Aberdeen (Gibson, 1972) and, as it is only the larger flounders which migrate into the sea, they will stand a better chance of becoming infested. It therefore appears that large flounders may migrate into the sea and become infested with C. heterochrous while the small flounders remain in the estuary and become more heavily infested with C. minutus. In marine flounders C. minutus does not appear to develop to maturity until ‘several months later than it does in estuarine flounders. This may be caused by the fact that the water-temperature of the Ythan estuary is warmer than that of the sea during the summer. The possibility that the development of this parasite is affected by temperature is also suggested by the fact that mature specimens were obtained from flounders at Plymouth in April, two months earlier than in flounders from the Ythan estuary. Further evidence can be deduced from the work of Markowski (1966), who found C. minutus in flounders from a brackish water reservoir containing water partially heated by the cooling water of a power-station, and in the sea a few yards away only a small number of flounders were infested with this parasite. When the zoogeography of this species is studied it is noticeable that C. minutus has not been found in northern waters except by Rudolphi (1819) and Janiszewska (1939) in the Baltic Sea. von Linstow (1904) reported Dacnitis fusiformis Molin, a synonym of C. minutus, from the Murman coast ; but his specimens were from the rectum of the flounder and, as shown by MacKenzie & Gibson (1970), C. minutus is very rarely found in the rectum of the flounder, whereas C. heterochrous is most commonly found in that region of the gut. C. minutus has normally been found in a more southerly geographical area, e.g. Gendre (1926) in French estuaries, 168 D. I. GIBSON Stossich (1890, 1898) and Mola (1928) in the Mediterranean Sea, and Butzkaya (1952), Radulescu & Vasiliu-Suceveanu (1956), Bykhovskaya-Pavlovskaya et al. (1964), Komarova (1966), Markevitch (1967) and Naidenova (1970) in the Black Sea and its associated estuaries. On the other hand C. heterochrous, ‘the winter species’, has been recorded mainly in the North Sea, the Russian Arctic, the Siberian coast and the Gulf of the St. Lawrence, e.g. Nicoll (1907), Baylis (1928), Wilker (1930), Schuurmans Stekhoven (1935), Kreis (1952), Polyanski (1955), Zhukov (1960), Strelkov (1960), Berland (1961, 1970) and Ronald (1963). This species has been recorded further south only on a small number of occasions, e.g. Stossich (1892, 1898),* as Dacnitis foveolatus (Rud.), in the Mediterranean Sea and Gendre (1927) off the north-west coast of Africa. It therefore appears that C. minutus is a parasite of flatfish living in warmer water than C. heterochrous, and that Aberdeen is close to the northern limit of C. minutus in a marine environment and is in a position where the ranges of these two parasites overlap. Further north it is probable that, except in estuarine conditions which are warmer than the sea during the summer, C. minutus would not be capable of completing its full life-cycle in one year. The Baltic Sea may be an exception to this, as its isolation from the Atlantic Ocean and shallowness means that it is warmer in summer than equivalent latitudes of the North Sea, and, with its lower salinity, it is therefore suitable for the development of this parasite. The slow development and relatively long life of C. heterochrous contrasts with the fast development and short life of C. minutus, for if the above interpretations of the results are correct, the life-history of C. heterochrous takes two years for completion and that of C. minutus only one year. This may explain the fact that C. hetero- chrous is able to occupy a more northerly geographical range than C. minutus. 6. ACKNOWLEDGEMENTS I should like to thank Dr. H. H. Williams for his advice and encouragement during the course of this work. I also wish to thank Professor V. C. Wynne- Edwards for the use of the facilities of his department at the University of Aberdeen where most of this work was carried out and the Scientific Research Council for a grant. Iam grateful to Dr. J. P. Harding and Mr. S. Prudhoe for reading the manu- script. 7. REFERENCES Bay.is, H. A. 1928. Records of some parasitic worms from British vertebrates. Ann. Mag nat. Hist. (to) 1 : 329-343. BERLAND, B. 1961. Nematodes from some Norwegian fishes. Sarsia 2 : 1-50. 1970. On the morphology of the head in four species of the Cucullanidae. Sarsia 43 : 15-64. ButzKaya, N. A. 1952. Parasitic fauna of the Black Sea economic fishes in the pre-estuarine area of the Danube. Tvrud. Leningr. Soc. Nat. 71 (4) : 30-52 (In Russian). BykHOvsKAYA-Paviovskaya, I. E. et al. 1964. Key to the parasites of freshwater fish of the U.S.S.R. Israel Program for Scientific Translation, Jerusalem. * Stossich’s specimens were not from flatfish and may therefore not have been C. heteyochrous. LIFE-HISTORIES AND DEVELOPMENT OF CUCULLANUS 169 GENDRE, E. 1926. Sur un nématode peu connu, Cucullanus fusiformis Molin. P.-v. Soc. linn. Bordeaux 78 : 39-49. 1927. Parasitologia Mauritanica. Helmintha (2): Nématodes parasites des poissons de la c6té de Mauritanie (Suite). Bull. Com. Etud. hist. scient. Afr. occid. fr., N.S. 21: I-I4. Gipson, D. I. 1972. Flounder parasites as biological tags. J. Fish Biol. 4 (r) : (In Press). Inciis, W. G. 1967. The relationships of the nematode superfamily Seuratoidea. /. Helminth. 41 : 115-136. JANISZEWska, J. 1939. Studien iiber die Entwicklung und die Lebenweise der parasitischen Wiirmer in der Flunder (Pleuvonectes flesus L.). Mém. Acad. pol. Sci., ser. B, 14, 1-68. Komarova, T. I. 1966. Helminth fauna of edible fish of the Dnieper estuary. In: Mazur- movich (edit.). Parasites, intermediate hosts and vectors. Akad. Nauk Uky. SSR. Kiev, sey. Problemy Pavasit. 6 : 57-66 (In Russian). Kreis, H. A. 1952. Beitrage zur Kenntnis parasitischer Nematoden. 10. Parasitische Nematoder aus der Umgebung der Faréer. Vidensk. Meddy dansk naturh. Foren. 114 : 251-307. Le-Van-Hoa & PHAM-Ncoc-Kuur. 1967. Morphologie et cycle évolutif de Cucullanus chabaudi n. sp., parasites des poissons, Pangasius pangasius H.B. (P. buchanani) du Sud Viet-Nam. Bull. Soc. Path. exot. 60 : 315-318. Linstow, O F. B. von. 1904. Entozoa des zoologischen Museums der Kaiserl-Akademie der Wissenschaften zu St. Petersburg. II. Ezheg. zool. Muz. 8 : 265-2094. MacKenzie, K. & Gipson, D. I. 1970. Ecological studies of some parasites of plaice Pleuro- nectes platessa L and flounder Platichthys flesus (L.). Symp. By. Soc. Parasit. 8 : I-42. Markevitcn, A. P. 1967. Analysis of parasitic fauna of Ukrainian fish from ecological and geographical respects. Proc. Zool. Soc. U.A.R. 2 : 169-179. MARKowSKI, S. 1966. The diet and infection of fishes in Cavendish Dock, Barrow-in-Furness. J. zool. Lond. 150 : 183-197. Mora, P. 1928. Vermi parassiti dell’ittiofauna italiana. Contributo alla patologia ittica. Boll. Pesca. Piscic. Idriobiol. 4 : 395-443. NartpENovA. N. N. 1970. The parasite fauna of fish in the family Gobiidae from the Azov Sea. In: Delyamure (edit.). Parasites of marine molluscs, fish and mammals. Akad. Nauk. Ukr. SSR. Biologiya Morya. Kiev. 20 : 74-84 (In Russian). Nicotr, W. 1907. A contribution towards a knowledge of the entozoa of British marine fishes. Part I. Ann. Mag. nat. Hist. (7) 19 : 66-94. PoLyanski, Y. I. 1955. Parasites of the fish of the Barents Sea. Tyudy zool. Inst., Leningy. 19: 5-170 (In Russian). (Translation: Israel Program for Scientific Translation, Jerusalem, 1966). Raputescu, I. & Vasitiu-Suceveanu, N. 1956. Contributiuni la cunoasterea parazitilor pestilor din complexul lagunar Razelm-Sinoe. Anal. Inst. cerc. pisc. 1 : 309-333 (In Roumanian, French & Russian). Ronatp, K. 1963. The metazoan parasites of the Heterosomata of the Gulf of the St. Lawrence. VII. Nematoda and Acanthocephala. Can. J. Zool. 41 : 15-21. Ruporput,C.A. 1819. Entozoorum synopsis cui accedunt mantissa duplex et indices locupletiss- imi. Berolini. SCHUURMANS STEKHOVEN, J. H. 1935. Nematoda parasitica. Tierwelt N.-u. Ostee. Tiel 5. c. 28 : 1-50. SHULMAN, S.S. 1957. Material on the parasitofauna of lampreys from the basins of the Baltic and White Seas. Bull. All-Union Sci. Res. Inst. Lake and River Fisheries 42 (In Russian). (Translation in: Parasites and diseases of fish. Edit., Petrushevskii,G. K. Israel Program for Scientific Translation, Jerusalem, 1966.) Stossicu, M. 1890. Brani di elmintologia tergestina. Boll. Soc. adviat. Sci. nat. 12 : 39-47. 1892. Osservazioni elmintologiche. Glasn. hrv. narodsl. Drust. 7 : 64-73. —— 1898. Saggio di una fauna elmintologica di Trieste e provincie contermini. Prog. Civ. Scuola R. Sup., Trieste, 1-162. 170 Di lee Gals SON StrRELKoV, J. A. 1960. Endoparasitic worms of marine fishes of east Kamchatka. Tyudy zool. Inst., Leningy. 28 : 146-196 (In Russian). TorNoutist, N. 1931. Die Nematodenfamilien Cucullanidae und Camallanidae, nebst weiteren Beitragen zur Kenntnis der Anatomie und Histologie der Nematoden. Géteborgs K. Vetensk.-o. VitterhSamh. Handl. 5f, s.B, 2 : 1-441. VESSICHELLI, N. 1910. Di un nuovo Dacnitis parasitta del Petromyzon planeri Monitore zool. ital. 21 : 304-307. WULKER, G. 1930. Uber Nematoden aus Nordseetieren I. Zool. Anz. 87 : 293-302. ZuuKov, E. V. 1960. Endoparasitic worms of fishes from the Sea of Japan and South Kuril shallow-waters. Trudy zool. Inst., Leningr. 28 : 3-146 (In Russian). Dr D. I. Gipson Department of Zoology British Museum (Natural History) CROMWELL Roap Lonpon SW7 5BD A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) Kay, E. Arison. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 1965. (Out of Print.) £3.75. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180; 11 Plates, 15 Text-figures. 1967. {£4._ TayLor, J. D., KENNEDY, W. J. & Hatt, A. The Shell Structure of Mineralogy at the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates, 77 Text-figures. 1969. £4.50. Haynes, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur) 1962-1964. (In press.) Bri Printed in England by Staples Printers Limited at their Kettering, Northants establishment TS BATS FROM THAILAND AND CAMBODIA 4) eect 11 Be ee AND KITT! THONGLONGYA BULLETIN OF MUSEUM (NATURAL HISTORY) Rye ar + Vol. 22 No. 6 LONDON: 1072 BATS FROM THAILAND AND CAMBODIA BY JOHN EDWARDS HILL AND KITTI THONGLONGYA Pp. 171-196; 4 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 22 No. 6 LONDON : 1972 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted im 1949, 1s 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. 22, No. 6 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). © Trustees of the British Museum (Natural History), 1972 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 29 March, 1972 Price 95p BATS FROM THAILAND AND CAMBODIA &, By J. E. HILL & KITTI THONGLONGYA { \ Ne, | INTRODUCTION Tue chiropteran fauna of southeast Asia as a whole is as yet not fully understood and knowledge of the bats of Thailand and Cambodia in particular rests largely on the efforts of a limited number of collectors. In recent years a more active interest in the bats of the region has become apparent and as a result a number of specimens have been received for determination at the British Museum (Natural History). Some of these have led towards the solution of taxonomic problems of long standing: others represent new records or range extensions for Thailand and Cambodia or are examples of taxa rare in collections. The majority of the specimens originate from Thailand and have been obtained over several years by Dr. J. T. Marshall of the United States Army Medical Component, South East Asia Treaty Organization. These were examined initially in Thailand by the junior author and are now in the collection of the British Museum (Natural History), except for a number of duplicate specimens sent to the Smithsonian Institution, Washington. A few specimens collected by the junior author are from the Thai National Reference Collection of which he is Curator. Cambodian specimens obtained by Mr. J. M. Klein are from the Museum National d’Histoire Naturelle, Paris, a small series of duplicates being retained in the British Museum (Natural History). Measurements are in millimetres: the minimum, maximum and mean (in parentheses) are given for series. ACCOUNT OF SPECIES Rousettus amplexicaudatus amplexicaudatus (E. Geoffroy, 1810) TuHaitanD: Doi Pha Hom Pok, Fang, Chiangmai, 6,800 feet, c.18° 47’ N., 98° 59’ E. 1 (young). Phu Nam Tok Tap Kwang, Kaeng Khoi, Sara Buri, c. 14° 42’ N., 100° 52’ E. 2 $4 (x young). The specimen from Doi Pha Hom Pok confirms the record by Chasen (1940 : 29) of R.a. amplexicaudatus from the mountains of northern Thailand, a region apparently on the northwestern border of its distribution. Rousettus leschenaulti leschenaulti (Desmarest, 1820) THAILAND: Koh Klet, Pak Klet, Nonburi, 13° 50’ N., 100° 29’ E. 1. Bang Phra, Siracha, Chonburi, 13° 12’ N., 100° 57’E. 84 (6 young), 6 99 (young). Recorded hitherto from the mountainous northern part of Thailand by Chasen (1940: 29). Specimens (B.M. 14.12.8.31-34) reported from Tagoot, Tenasserim, Burma as R. leschenaulti by Wroughton (1915: 702) are in fact R. amplexicaudatus. 174 Wo 185 FEMI, re ANG TRCN Pteropus hypomelanus Temminck, 1853 THAILAND: Tao Poon, Koh Som, Koh Samui, Surat Thani. 1 (young), 2 99 (I young). The small islet of Koh Som is close by the larger island of Samui, in the Bight of Bandon off the east coast of southern Thailand. No collector of mammals appears to have visited it hitherto although Koh Samui was visited by H. C. Robinson and C. B. Kloss in 1913 on behalf of the Federated Malay States Museum. No Ptevopus were encountered (Robinson and Kloss, 1g15a) and the genus was first reported from the islands by Marshall and Vandee (1970 : 504) who recorded these and other specimens obtained in 1968. Pteropus hypomelanus is known to occur on a number of the other small islands off the east coast of the Malay Peninsula and on the Natuna Islands. The back of the adult female from Koh Som is black, streaked with grey, the mantle bright chestnut and the head greyish, tinged with buff. The underparts are predominantly grey brown or hair brown, excepting for a band of black across the base of the throat. The young male and female have less grey on the back, the mantle bright chestnut in the male and deep rufous in the female, both with blackish brown crown and nape and with black underparts. Measurements of the adult female: length of forearm 141; greatest length of skull 64-3; condylobasal length 62:3; palatal length 36-9; least interorbital width 8-4; postorbital width 7-6; zygomatic width 31-6; braincase width 21-6; mastoid width 19-9; c1-c! 11-0; m1-m1 17:6; c-m2 24-3; length of mandible 48-9; c—-mg3 27-4. It is difficult to refer these specimens to any ot the subspecies of P. hypomelanus so far described from the islands of the South China Sea. The darker back and greyish or black brown crown and nape set them apart from the pale subspecies P.h. lepidus from the majority of the east coast islands of the Malay Peninsula and P.h. canus and P.h. annectens from the North and South Natuna Islands: although the colour of the back resembles that of P.h. condorensis from Con Son Island (Pulo Condore), in this subspecies the crown and nape are chestnut brown like the mantle. Andersen (in Kloss, 1916 : 38) reports specimens of P.h. condorensis from certain of the small islands off the southeastern coast of Thailand. These are not in the collection of the British Museum (Natural History) and although the same size as the adult from Koh Som, from the description some examples differ from the Koh Som specimens in the colour of the underparts. Unfortunately, Andersen did not mention the colour of the crown and nape. Insufficient material is available to demonstrate the extent of colour variation in condorensis, and for the present specimens from Koh Som are left unallocated to subspecies. They are very similar in colour and size to P.h. geminorum, so far known from the islands of the west coast of the Peninsula from Paya, off Kedah, States of Malaya, north to the Mergui Archipelago, whence there are specimens in the British Museum (Natural History) from the islands of Barwell, Sir John Hayes, Malcolm and the Gregory group. The adult female from Koh Som corresponds to the hair brown phase of geminorwm noted by Andersen (1912 : 107): the bright chestnut mantle contrasts with the dark seal brown mantle usual in this phase of geminorum, but a BATS FROM THAILAND AND CAMBODIA 175 similar condition is exhibited by a specimen from Sir John Hayes Island. The two younger specimens correspond to the black-bellied phase noted by Andersen (p. 106). Pteropus hypomelanus is found predominantly on islands and has yet to be recorded from the mainland of the Malay Peninsula. It is of interest therefore to report the occurrence of similar populations on small islands lying off the eastern and western coasts at approximately the same latitude. Cynopterus sphinx angulatus Miller, 1908 THAILAND: Doi Pha Hom Pok, Fang, Chiangmai, 6,800 feet, c. 18° 47’ N., 98° 59’ E. 1, 2 99. Doi Ithanon, Chom Thong, Chiangmai, 1,700 metres. 18° 35 'N., 98° 29'E. 1. Phu Nam Tok Tap Kwang, Kaeng Khoi, Sara Buri, c.14°21’N.,100°52'E. 19. Bang Phra, Siracha, Chonburi, 13° 12’ N., 100° 57’ E. 22 $g (3 young), 25 99. Koh Klet, Pak Klet, Nonburi, 11 miles north of Bangkok. 1, 1 9. Lumpinee, Bangkok. 1,1 9. CAMBODIA: Prek Tasek, 8 kilometres northwest of Phnom Penh. 1 9. Siem Reap. I 9. In his monographic study of the Megachiroptera, Andersen (1912) recognized a single species of Cynopterus, C. brachyotis, in the Malay Peninsula, with two sympatric subspecies, one, C.b. brachyotis, with shorter ears and rostrum, the other, C.b. angulatus, having longer ears and rostrum. According to Andersen, both occur in Sumatra and throughout much of the Peninsula but C.b. angalatus alone extends north of the Peninsula as far as northern Thailand. Neither was thought by Andersen to have any close affinity to the Indian and Javan species C. sphinx which he thought occurred on the mainland as far east as northern Thailand (where according to Andersen it is to be found sympatrically with C.b. angulatus), although a subspecies, C.b. titthaecheilus, was recognized by this author in Java. However, Kloss (1911 : 185) listed angulatus as a distinct species, noting an affinity to C. sphinx and suggesting that angulatus should be regarded as the southern race of the Indian species. Later, Robinson and Kloss (1915a : 133, Ig15b : 114) reiterated this suggestion as a logical conclusion, although in both instances these authors follow Andersen in listing angulatus as a subspecies of brachyotis. This led to an exchange of opinion-by Andersen and Kloss (1915 : 220) in which the views of Andersen seem rather doubtfully to prevail. His opinion is published again in Kloss (1916 : 40), who provides further (p. 41) comments on specimens from the islands off southeastern Thailand among which Andersen had identified both C.b. brachyotis and C.b. angulatus. Kloss clearly remained un- convinced and later (1917 : 300; Robinson and Kloss, 1918 : 26; Kloss, 1919 : 361) reverted to the view that angulatus represented a distinct species. Subsequently, Chasen (1940 : 25) considered angulatus to be a large subspecies of brachyotis occurring only in the north of the Malay Peninsula and in some nearby islands, a view adopted by Hill (1961 : 630) and Medway (1969 : 12). It is evident from the literature that specimens from southern Thailand are critical to any study of the status of angulatus. Sanborn (1952 : 2) for example 176 Vo Vin eMC IL, ee ihs ISA Atal referred specimens from Kwan Don Setul (=Satun) Province to this form but pointed out that some of the specimens examined had ears and rostra which agreed with C. sphinx. In fact, this series reported by Sanborn from a locality almost on the border with Malaya has measurements (length of forearm 61-0~-72-4, of ear 16:5—22:0) indicating that it includes both angulatus and brachyotis. These appear to occur together in Perak, in the northern States of Malaya (Andersen, 1912 : 614; Hill, rg6r : 630); on Koh Lak, off the northeastern part of the Malay Peninsula in southern Thailand (Gyldenstolpe, 1916 : 9); on the islands off southeastern Thailand (Andersen, in Kloss, 1916 : 40); and now, in the present collection, at Bang Phra, Siracha, yet further north. Furthermore, specimens ot brachyotis have been obtained at Khao Luang, Nakhon Si Thamrat, on the eastern side of the Peninsula, which is north of Trang, the type locality of angulatus, on the opposite side of the Peninsula. More material than was available to Andersen now exists in the collection of the British Museum (Natural History) : in particular, there are good series of specimens from central Burma, from Tenasserim (Wroughton, 1915 : 703) and the Mergui Archipelago (Lindsay, 1927 : 44), from central and northern Thailand, and from the Malay Peninsula (Hill, IQ6I : 630). The specimens now available from this critical area divide into two groups, one of larger examples with longer forearms and ears, the other group with generally shorter forearms and shorter ears (the length of the ear as given here is from the meatus, measured personally or for dry specimens by the collector). These correspond to angulatus and brachyotis as these are defined by Andersen (1912). It is difficult to postulate so extensive an area of intergradation in order to maintain the opinion that angulatus and brachyotis are conspecific. There are two alternatives to the view that angulatus is a subspecies of brachyotis, namely that it is a species distinct from either brachyotis or from sphinx; or that it is a subspecies of sphinx. The latter alternative is adopted here and is supported by the long series of specimens from Burma and Thailand which show that the criterion ot rostral length used by Andersen to separate angulatus and C. sphinx sphinx is by no means as exclusive as was thought by that author. Andersen (pp. 600, 612) considered that angulatus might be distinguished from C.s. sphinx by a shorter rostrum, its length in angulatus being less than one quarter of the total length of the skull, in sphinx one quarter or more. The rostral lengths of specimens from India (C.s. sphinx, C.s. gangeticus), Burma (C.s. sphinx), Thailand, Malaya and Sumatra (angulatus) and Java (C.s. titthaecheilus) are compared graphically against total skull length in Figure 1, the dotted line demonstrating those points at which the rostral length is one quarter of the total skull length. There is no clear separation between Indian, Burmese and Thai specimens and, indeed, in sphinx from Ceylon and titthaecheilus from Java the length of the rostrum is not infrequently less than one quarter of the total length of the skull. The specimens obtained by Dr. Marshall in Thailand are very slightly smaller than C.s. sphinx and have relatively slightly shorter rostra, in these features resembling specimens from Tenasserim and the Malay Archipelago. There appears to be no exclusive external or cranial feature by which C. sphinx BATS FROM THAILAND AND CAMBODIA 177 can be separated from C. brachyotis, a difficulty recognized by Andersen (1912 : 609) who noted that the numerous subspecies of C. brachyotis taken together could be separated from the modifications of C. sphinx only by their relatively shorter ears. Cynopterus sphinx is distributed from India and Ceylon through Burma, Thailand and Indochina to Hong Kong: it extends in the Malay Peninsula as far south as Perak and thence to Sumatra, Java and Timor. Occurring more frequently on islands, C. brachyotis extends from Borneo and the Philippine Islands to Java, Sumatra, the Malay Peninsula and its adjacent islands northwards at least as far as southern Thailand and North Vietnam (C.b. hoffeti Bourret, 1944), occurring also on the Nicobar Islands and Ceylon. Although on occasion some subspecies (C.b. scherzert from the Nicobar Islands, C.b. insularwm from Kangean and Mata Siri Islands, off Java) approach or equal the smaller of C. sphinx in size, such subspecies have the characteristically smaller ears of brachyotis. There is some overlap between the two species in the area of sympatry from Thailand to Sumatra in both length of forearm and ear, and although a definitive line cannot be drawn for either character, the majority of specimens can be separated readily by a combination of these parameters, which are compared for Burma, Tenasserim, Thailand and Indochina in Figure 2 and for Malaya and Sumatra in Figure 3. A comparable comparison of specimens of sphinx and brachyotis from Ceylon and Java appears in Figure 4. On the mainland and in Sumatra, the 9.5 2 AAS i Z 2 v 5 Fa e 9.0 v A UA Re v 5 Zee rv v a os v v se 4 8.5 voy ay Be on Z v vv vun- v Pa a Vv ees § Oo =O” $8.0 a 5% R m”°@oo U vou” oon a ~~ vem 0 oO Vv INDIA = s Nard es @ CEYLON 7.5 a @go oe = of s as © BURMA 2 Lie a8 = v m THAILAND, INDOCHINA, ih 2 TENASSERIM, MALAYA & SUMATRA s a an s 7.0 en s A JAVA me ne --- ROSTRAL LENGTH ONE QUARTER TOTAL LENGTH OF SKULL 65 29 20 n 32 33 a4 35 36 37 38 TOTAL LENGTH OF SKULL Fic. 1. Length of skull and rostrum (front of orbit to nares) in Cynopterus sphinx. 178 j; BRIDE a Kaonos forearm length of specimens referable to sphinx generally exceeds 65 mm. and the length of the ear is greater than 19:0 mm., while specimens referable to brachyotis have forearm and ear lengths only rarely exceeding these values. The large number of specimens from Thailand show that C.s. angulatus has ears that in length considerably exceed the maximum value of 18-0 mm. which Andersen (1912 : 612) noted for angulatus when he allocated it to brachyotis as a subspecies. In this connection it should be noted that Andersen stated (p. 612) of angulatus that the measurement of the ear is from the orifice, yet in the typical series measured by Miller (1898 : 318) the length of the ear from the meatus is given as 18-21 mm. and from the crown 15-18 mm., only the latter being under the maximum given by Andersen for angulatus. Still further, the table of measurements in a British Museum (Natural History) copy of Miller (1898) has been annotated (p. 318) by Andersen to the effect that he has examined the first six specimens tabulated. The BURMA. TENASSERIM. THAILAND & INDOCHINA © Cynopterus sphinx a Cynopterus brachyotis 72 ©) \0' 0: 6, @eeos ef . eee . . u . ° eo © © eeece owe . . . . . ° Pao ee meee © am wetees am © ° A R 19 i ee oc ec @e . ee rs 18 ae eo ° . a a v7 s s . a s 16 ee 1s a 57 59 61 63 65 67 69 7\ 73 75 FOREARM Fic. 2. Length of forearm and ear in Cynoplerus sphinx and C. brachyotis. i BATS FROM THAILAND AND CAMBODIA 179 first of these (U.S. National Museum 83,524) with forearm of 61 mm. and ear 18-4 mm. he notes as C.s. (sic) brachyotis, the remainder as true angulatus, thus demonstrating that the two forms occur together at the type locality of angulatus, Trang, in southern Thailand. The montane subspecies C. brachyotis altitudinis Hill, 1961, from Mount Brinchang, Perak has generally longer ears than C.b. brachyotis, reaching a maximum of 20 mm., but has a short forearm only exceptionally reaching a maximum length of67mm. Furthermore, it is linked to C.b. brachyotis from the surrounding lowlands by intermediates (Hill, 1961 : 632). Chasen (1940 : 25) listed the large C. major Miller, 1902, from Nias Island, off west Sumatra as a subspecies of sphinx. Andersen (1912 : 630) considered it a representative of ‘C. brachyotis angulatus’ on account of its ears which he noted (p. 629) as 16-5-18 mm. in length, but Chasen (p. 29) measured the length of the ear at 18-5-20 mm. Cynopterus brachyotis brachyotis (Miller, 1838) THAILAND: Khao Luang, Nakhon Si Thamrat, 1,300 feet, 8° 26’ N., 99° 58’ E. 3 dd (I young), 1 9. Bang Phra, Siracha, Chonburi, 13° 12’ N., 160° 57’ E. Ig, 5 99. MALAYA & SUMATRA 22 e e © Cynopterus sphinx e a 4 Cynopterus brachyotis brachyotis © e e 20 e e e e s ” 3 e a s 2 e 5 a a a ne R F , yo Ty a yap SOT Ale ack a ae a a a a va Ay a ay a s aaa a a “a a 4 OU ANP ta mt a a “a s ie a a a Ah. a aa “ cae “ 16 aa ey ae a a rs a a a or a a - 56 58 60 62 64 66 68 70 72 74 76 FOREARM Fic. 3. Length of forearm and ear in Cynopterus sphinx and C. brachyotis. “syodyonag *D pue xuryds snaajgoudkD ur 1e9 pue wiuieeloj Jo yySueT “bh “OI = _ wavadod ’ £8 18 62 “ZL GZ eZ nz 69 £9 s9 £9 19 6S Zs v v ot y: v Vi ava.Y u I H 3 : is - V4 SR RAID ALY ab i , e & 3 e vy vv 6 ) is ° q = il ¥ © v oz . oo 2) ° ° e ° ° ° ° ° ° ° e itd ° ° ° ° ° oo ° ° ° wz Y siyoAyr01q sniajdoudy 9 oo ° {a} 4 e xuiyds sniaydoudy NOTA3D VAVE ir 180 wit BATS FROM THAILAND AND CAMBODIA 181 These specimens agree exactly with C.b. brachyotis from the southern part of the Malay Peninsula: those from Bang Phra constitute the northernmost record in Thailand: C.b. brachyotis occurs also on the islands of Chang, Mehsi East, Kra, Klum, Kut and Lak in the Gulf of Siam. Megaerops ecaudatus (Temminck, 1837) THAILAND: Khao Luang, Nakhon Si Thamrat, 1,300 feet, 8° 26’ N., 99° 58’ E. Ig,1¢d. Doi Pha Hom Pok, Fang, Chiangmai, 6,800 feet, 18° 47’ N., 98° 59’ E. 16. This species apparently has not been recorded hitherto from Thailand although known from Malaya (Bonhote, 1903 : 15; Hill, 1961 : 636) and both North and South Vietnam (Van Peenen, Ryan and Light, 1969 : 41). Chironax melanocephalus (Temminck, 1825) THAILAND: Khao Luang, Nakhon Si Thamrat, 1,300 feet, 8° 26’ N., 99° 58’ E. 1Q. No skull is available to confirm the identification of this specimen as C. melanocephalus, but it has the characteristically darker head of that species with bright orange patches at the sides of the throat just anterior to the shoulders. Chironax melanocephalus has been reported from Selangor (Chasen, 1940 : 30, Hill, 1961 : 640) and, more recently, specimens (B.M.67. 1484-1488) have been received from Gunong Benom, Pahang, but the species has not before been reported from Thailand. The specimen from Khao Luang agrees well with one from Pahang but the head is a little less intensely black, the underside rather more brownish and the orange throat patches less prominent. Length of forearm 47:3. Sphaerias blanfordi (Thomas, 1891) THAILAND: Doi Pha Hom Pok, Fang, Chiangmai, 6,800 feet, 18° 47’ N., 98°59’ E. I 9 (young adult). This species has evidently an extensive range through the montane areas of southeastern Asia. The original material consisted of three examples (one, B.M.90.4.7.6, in the collections of the British Museum (Natural History)) collected by Fea at Leito, Cheba, Karin Hills, Burma at 1,000 metres, while subsequently Allen and Coolidge (1941 : 136) recorded two specimens from Mount Angka (Doi Inthanon), in northern Thailand, at 4,000 feet. More recently, Bhat (1968 : 471) has recorded S. blanfordi from various Indian localities in Uttar Pradesh, at elevations ranging from 800 to 2,710 metres. Measurements of the specimen from Doi Pha Hom Pok: length of forearm 52-0; total length of skull 27-8; condylobasal length — ; condylocanine length — ; length orbit to nares 7-0; lachrymal width 70; least interorbital width 5-3; postorbital width 6-7; zygomatic width 15-5; diameter of orbit 6-5; braincase width 11-8; mastoid width 11-3; cl-cl 5-8; least width between bases of canines 3-1, p*-p4 5-3, least width between p4-p4 4-7; m1—m! 7-5 ; least width between m1-m! 5-6; width mesopterygoid fossa 3-8; c-m! 8-6; length of mandible 19-9; coronoid height 7-7; c—mg 9°8. 182 jpg TMi, Ce an IACtar abit Macroglossus minimus sobrinus Andersen, 1911 THAILAND: Doi Inthanon, Chom Thong, Chiangmai, 1,700 metres, 18° 35’ N., Of Zo) 19, 10) Ban Bangmakham, Koh Samui, Surat Thani, 9° 30’ N., 100° 00’ E. 14., Khao Luang, Nakhon Si Thamrat, 1,300 feet, c. 8° 26’ N., 99° 58’ E. 14,12 (young). Ellerman and Morrison-Scott (1951 : 101) do not include Thailand in the distri- bution of M. minimus although the species is listed by these authors from Tenasserim. However, Horsfield (1851 : 29) lists a specimen in the Museum of the East India Company collected in Thailand by Finlayson, apparently the same specimen listed subsequently from the Indian Museum by Anderson (1881 : 107), and Bonhote (1903 : 15) records a specimen from Patani, in southern Thailand. Taphozous theobaldi theobaldi Dobson, 1872 THAILAND: Sara Buri. 14. Khao Lom Phat, 21 kilometres eastnortheast of Sara Buri. 1 9. Described from Tenasserim, T. theobaldi was listed from Thailand by Pousargues (1904 : 544) and has been recorded subsequently from North Vietnam (Bourret, 1944 : 9) and South Vietnam (Van Peenen, Ryan and Light, 1969 : 50). Measure- ments (gf, 9): length of forearm 74:0, 74:1; greatest length of skull 24-2, 23-6; condylocanine length 23-9, 23:5; least interorbital width 7-5, 7-0; postorbital width 5:2, 5°35 zygomatic width 14-3, 14:1; braincase width 10-8, 11-1; mastoid width 12°8, 12:6; cl-c! 4-9, 4:8; m3-m$ 10-2, 10-1; c—m3 10-7, 10:5; length of mandible 18-7, 18-5; c-mg 11°8, 11-7. Taphozous longimanus longimanus Hardwicke, 1825 CamBopiA: Prek Phnau, 6 kilometres northeast of Phnom Penh. 2 99. Cambodia, no certain locality. 5 99. Taphozous longimanus has not hitherto been reported from Cambodia although known to occur in Tenasserim (Wroughton, 1915 : 706) and further north in Burma (Wroughton and Davidson, 1918 : 477, Wroughton, 1918b : 25). The wings of these specimens are dusky, and they are referred to the nominate subspecies rather than to the pale-winged T./. albipinnis from the Malay Peninsula, Sumatra and Borneo. Length of forearm 62-0, 62-3: of five females of uncertain provenance 60-5-63:1 (61-5). Rhinolophus affinis macrurus Andersen, 1905 THAILAND: Doi Inthanon, Chom Thong, Chiangmai, 18° 35’ N., 98° 29’ E. 14. Ban Papae, Mae Sariang, Mae Hong Son, 2,365 feet, c. 18° 45’ N., 97° 55’ E. 14. Now Senckenberg Museum No. 33816. Longer tails (c. 26-29 mm.) refer these specimens to R. a macrurus rather than to R. a superans from the Malay Peninsula. BATS FROM THAILAND AND CAMBODIA 183 Rhinolophus malayanus Bonhote, 1903 THAILAND: Phu Nam Tok Tap, Kwang, Kaeng Koi, Sara Buri c. 14° 12’ N., TOOg52, ETO: First described from Biserat, in Jalor, southern Thailand, the range ot this species has since been extended to include Perlis, in the northern States of Malaya (Hill, in press), Pulau Langkawi, off the west coast of the Malay Peninsula (Hill, in press), Koh Lak, off the east coast of southern Thailand (Gyldenstolpe, 1916 : 13); Laos (Andersen, 1905: 89, Tate and Archbold, 1939: 6; Phillips, 1967: 634) and tentatively North Vietnam (Osgood, 1932 : 218). Measurements of the specimen from Sara Buri: length of forearm 40-1: greatest length of skull 18-3; greatest length to canine 17:3; condylocanine length 15-2; supraorbital length (distance from junction of supraorbital ridges to median anterior point of nasal swellings) 5-0; least inter- orbital width 1-9; zygomatic width 8-7; braincase width 7-4; mastoid width 8-3; cl_cl 4:2; m3—-m3 6-4; c-m 6-7; length of mandible 11-3; c—mg 7:0. Phillips (1967 : 634) considers that on the basis of the published description it is probable that Rhinolophus chaseni Sanborn, 1939 from Con Son Island (Pulo Condore), off the southeastern coast of South Vietnam will prove conspecific with R. malayanus. Elsewhere, one of us (Hill, in press) has indicated that chaseni (probably the bat reported from Con Son as R. minor by Pousargues, 1904 : 544) is not closely related to malayanus as was suggested by its describer but instead should be considered a subspecies of R. borneensis. Rhinolophus malayanus differs from R. borneensis chiefly in the form of the anterior nasal swellings: in malayanus the median swellings are large and much inflated, extending laterally down the sides of the rostrum to the extent that the lateral swellings are relatively small, while in borneensis the median swellings are smaller and less inflated, not extending laterally down the sides of the rostrum, with the lateral swellings conspicuously larger than in malayanus. Examination of the holotype of chaseni shows it to belong with borneensis rather than with malayanus. Rhinolophus shameli Tate, 1943 THAILAND: Ban Bon Dan, kilometre 58, Route 23, Korat, 400 metres. I 3. CamBopiA: Preah Khan, Siem Reap, 3 gd. 1 9. Shamel (1942 : 319) referred two specimens from Thailand in the Smithsonian Institution to R. coelophyllus Peters, 1867, a species rare in collections and character- ized by the unique hood-like appearance of the posterior nose-leaf, the connecting process entering a vertical fissure in its anterior face. The two specimens reported by Shamel, however, differed sufficiently from each other in a number of respects that this author described them individually. Subsequently, Tate (1943 : 2) considered that one, from the island of Chang in the Gulf of Siam, merited sub- specific separation as R.c. shameli when compared with the other specimen, obtained at Chiangmai in northern Thailand. Apart from these specimens, R. coelophyllus has been reported from the Salween River (the type locality) ; from Tsagine (Sagain), Upper Burma (Ellerman and Morrison-Scott, 1951 : 123); from Koh Lak off the 184 Jo BeHiD & ely Kon east coast of southern Thailand (Gyldenstolpe, Ig11 : 16) and from the States of Malaya (Chasen, 1940: 42). The specimens now reported from Thailand and Cambodia, however, together with others obtained recently on Pulau Langkawi, off the west coast of the Malay Peninsula, suggest that shameli represents a distinct species. According to Shamel (1942 : 319) the specimen from Koh Chang (shamelz) is much more brightly coloured than that from Chiangmai and differs also in the configuration of the upper surface of the rostrum, which posteriorly to the nasal swellings is flat and not excavated, the margins formed by the division and forward extension of the sagittal crest not ridged or beaded. In the specimen from Chiangmai this region is scooped out, the margins of this depression formed by strong, beaded supraorbital ridges developed from the forward extension and division of the sagittal crest. Tate (1943 : 3) in describing the specimen from Koh Chang as shameli points out that it is considerably larger than the example from Chiangmai, as can be seen from the measurements published by Shamel, and also that it has hyopsodont teeth, these features together with its more brilliant coloration forming the basis of his diagnosis. The specimen reported here from Ban Bon Dan agrees closely in colour with the description by Shamel (1942 : 319) of the example from Koh Chang subsequently designated as the holotype of shameli by Tate. The head is dark brown to the anterior bases of the ears: the remainder of the dorsal surface is bright golden brown, anteriorly slightly more brilliant, posteriorly a little browner. The individual hairs are pale cream for most of their length, tipped with the brighter colour. The ventral surface is paler, overall orange buff, lacking brown, the hairs orange buff at the base and for most of their length, with brighter tips. Cranially, the specimen agrees exactly with the description by Shamel of the Koh Chang example, the upper surface of the rostrum flat and not margined by prominent supraorbital ridges. Specimens from Cambodia (whence neither coelophyllus nor shameli have been reported hitherto) and Upper Burma agree in size and rostral features with shameli but differ in colour. These specimens are brown dorsally, the individual hairs pale greyish white at the base: the ventral surface by contrast is very much paler, the throat, chest and belly pale buff, the hairs creamy white at the base and tipped with buff or buff brown. The flanks and inguinal region are slightly darker, the hair tips brown and lacking any buff. Since in all other respects these specimens agree exactly with shameli they support the view that this species exists in a brownish phase and a contrasting brighter, more reddish phase, a phenomenon not uncommon in Rhinolophus. The collections of the British Museum (Natural History) include one example from Kedah and two from Pulau Langkawi which agree closely in colour with the specimens from Cambodia and Upper Burma referred here to shameli and with the description of a single specimen from Chiangmai reported by Shamel (1942 : 319): those from Langkawi (in alcohol) are very slightly browner ventrally than those from Cambodia. However, the three specimens are very much smaller than shameli and the upper surface of the rostrum is in each deeply excavated behind the nasal swellings, the depression enclosed laterally by prominent supraorbital ridges formed BATS FROM THAILAND AND CAMBODIA 185 from the anterior division of the sagittal crest, while all have less massive teeth. In these respects, they thus resemble precisely the specimen described from Chiangmai by Shamel which Tate (1943 : 3) noted was ‘probably referable to the type from Salween River, Burma’. The type specimen of coelophyllus is small (forearm length 42, according to Peters, 1867a : 427) and for the present this name is used for the smaller species to which the Chiangmai, Kedah and Langkawi specimens belong. Those from Koh Lak reported by Gyldenstolpe (1911 : 16) are cranially a little smaller and may represent an undescribed subspecies of R. coelophyllus as Tate (p. 3) suggests. Measurements of R. shameli and R. coelophyllus are compared in Table 1. Rhinolophus acuminatus Peters, 1871 THAILAND: Chonburi, 13° 22’ N., 100° 59’ E. 1, 2 99 (x young). Bang Phra, Siracha, Chonburi, 13° 12’ N., 100° 57'E. 293 (1 now Senckenberg Museum No. 33817). I 2CamBopIA: Preah Khan, Siem Reap. 34,1 9. Rhinolophus acuminatus is recorded from Thailand and Laos by Dobson (1878 : 878) and Pousargues (1904 : 544) and from Thailand by Shamel (1942 : 321) and Sanborn (1952 : 3): these specimens from Preah Khan constitute the first record of the species from Cambodia. Specimens from Thailand and Cambodia agree very closely in colour and size with an extensive series of R.a. acuminatus (to which Thai specimens are referred by Shamel and by Sanborn) from Java. Six examples (including three from Cambodia) are in the dark or grey phase, the dorsal pelage greyish brown, the hairs tipped light grey or silver, especially anteriorly: the ventral surface is pale grey. Three (including one from Cambodia) display the red or russet phase and are dorsally reddish brown, ventrally a little paler. A single example exhibits a condition midway between these extremes: the anterior part of the back is greyish brown, tinged with russet, the posterior back russet, while the ventral surface is suffused with russet yet retains much of its greyish tinge. Although there is a good series of R.a. acuminatus in the British Museum (Natura] History), the remaining subspecies, R.a. audax from Lombok, the Sumatran R.a. sumatranus, R.a. calypso from Enggano Island, off Sumatra and R.a. circe from nearby Nias Island are as yet only very poorly represented. Furthermore, Chasen and Kloss (1932 : 48) identify as R. acuminatus a specimen from Sabah, Borneo which Chasen (1940 : 39, footnote) suggested might represent an undescribed subspecies, provisionally associating it with R.a. sumatranus. Allen and Coolidge (1940 : 136) did not allocate specimens from Mount Kinabalu to subspecies, but Medway (1965: 54) lists Bornean specimens as R.a. sumatranus. In these circumstances it is difficult to assess subspecific variation in R. acuminatus and for the present mainland specimens are left unallocated. Rhinolophus borneensis Peters, 1861 CamBopiA: Preah Khan, Siem Reap. 26. There appears to be but one previous report of R. borneensis from the mainland of southeastern Asia, by Robinson and Kloss (1915b : 116), who tentatively identified 186 eT Yo IMeysue’y yepes purpeyy ‘eusurryD PIpoquie) purpleyy ‘ye10y, vung iaddy, Sury9 yoy ‘edAjojoHy uolze00'T i Ge MOG aS Se Ora e Biarsi sanjea asay} 10} $-g pue o.gb saats (9 : ch61) ayey + jaweys Wor » od 8-9 1-g 1-8 Lb gt o£ or Sg — — 1g — gL 1.9 z+ G.G1 g-Lt GL 611 zl g9 Lb c6 LG Lg 96 61 6F OF go1 g-gt ELI ¥-61 of — €2 Lo Lr 26 3S 98 4-6 12 6F OS Lor L.gr eli ¥-6r (yf (omiat AA eyfey Ci? tate) CHG ASP SH) aA te (oy? Atlee (epispe gL — $s. zG ug 06 o2 — 72S — — G.Lr 061 rg — SL o4 of 66 9S og 46 gr €§ +6 Lr 0.02 S.gr L.0z €¢ — 62 12 gb 96 25 Lg 66 rez $5 SG Lr 1-02 9-g1 Loz rg — gf zl gr +6 of $g 66 gx €S — F-4r S.6r 1-g1 6.61 £9 gzi gl €2 1-6 96 69 99 66 6r GS G.G G./1 0.0% 1-81 ¥-02 pes nance (5/2 Phy) wel Gey HS Aas WS) hea CS NES /Atope ashe 62 921 92 69 1S 4-6 09 68 S6 — GE G6. 9.41 g.61 ¥-g1 L.0z cg — log — — — 99 Sg 06 of — gS — — 81 Oz ee te hielety 3 2 £€2 90 em eNS9R Bw Fe GO0 SO GO Ba > 822 7 4 BESE SEES 28 2 EE as os ae ge 1 e¢ sete Eh oe ape eter Gh tee feaey el is c ¥ i) is) fp Siete (stip ong le set a: 6 & 25 2. fe as come) OMT o ey 2a aoe eae eeeses 1h eee > S ne 3 gg = ag i = snppdygojaoa *yy ‘yaunys snydojouryy yO syusWeINSeay] I alaVy wiied10y Fo *0 OF OF €o OF *0 OF £0 Fo *0 fo 0 OF XOS adjoysuepfAy ZZ8'89 1Z8"89 ‘I'1'01'g6 +09¢L97 WN'S' snqjdygojaoa ay “snyq Sued Teysrey “Lf zgzS:gl brb 16 ¥SSzLov WN'S' yaunys “yy BATS FROM THAILAND AND CAMBODIA 187 a specimen from Khao Nawng, Bandon, southern Thailand with this species. This specimen, however, later became the holotype of R. robinsoni Andersen, 1918. Rhinolophus borneensis spadix from the Natuna and Karimata Islands seems only barely separable (Andersen, 1905 : 87) from the Bornean subspecies R.b. borneensis by virtue of its very slightly larger ears. Both subspecies, like the Cambodian specimens, are a little larger in some respects than the holotype of R.b. chaseni Sanborn, 1939 (see above, under R. malayanus) from Con Son Island (Pulo Condore), but measurements of a series of chasent given by Van Peenen, Ryan and Light (1969 : 59) and by Van Peenen, Cunningham and Duncan (1970 : 421) (in which by a lapsus the length of the forearm is given as the greatest skull length, the latter appearing as the zygomatic breadth) approach or overlap those of mainland speci- mens. It may well be, therefore, that chaseni will prove to be the correct subspecific name for the mainland population. Specimens reported as Rhinolophus sp. from North Vietnam by Osgood (1932 : 219) seem perhaps referable to R. borneensis. In size they agree with those reported from Cambodia, and Osgood notes that the development of the nasal swellings is perhaps less than in malayanus, the skulls agreeing in general robustness with stheno. In borneensis the median nasal swellings are smaller than those of malayanus (see above, under R. malayanus), and, so far as mainland specimens are concerned, the skull is generally more robust, similar in this respect to stheno. Measurements of the Cambodian specimens, with those of the holotype of R.b. chaseni (in parentheses) : length of forearm 44-1, 43-4, (41-3); greatest length of skull 19-7, 193 (18-7); condylocanine length 17-3; 17-6 (16-5): rostral width 5-3, 5-3, (5:3); width across nasal swellings 5-5, 5:5, (5:5); least interorbital width 2:5, 2-3 (2:5): zygomatic width 9-9, 9:9 (9:3); braincase width 8-3, 8-4 (8-2); mastoid width 9:5, 9:5, (8:8); clcl 4:8, 4-9 (4-9); m3—m8 7-2, 6-8, (6-8); c-m3 7-7, 7-4, (7-3); length of mandible — , 12-7, —; c-mg 8:3, 8:0, (7°8). Coelops frithii Blyth, 1848 THAILAND: Khao Luang, Nakhon Si Thamrat, 1,300 feet, 8° 26’ N., 99° 58’ E. Ig. The elongate, narrow outline of the lappets projecting from the supplementary leaflets flanking the anterior noseleaf suggests that this specimen should be referred to C. frithit rather than to C. robinsoni in which these lappets are rounded and wider. Unfortunately, the skull, which would provide a more definite indication, is missing. Length of forearm 38-2. Robinson and Kloss (1915b : 116) record two specimens from Khao Nawng, Bandon, southern Thailand as C. robinsoni. These, formerly in the collection of the Federated Malay States Museum (531/12, 532/12), are now in the collections of the British Museum (Natural History) (B.M. 68. 605-606). Elsewhere, one of us (Hill, in press), in a review of the species of Coelops, has shown these to be examples of C. frithit, which is recorded from Chiangmai by Gyldenstolpe (1916 : 15) and listed from Laos by Pousargues (1904 : 544). 188 je Ea RIEL ie Ker Myotis hasseltii continentis Shamel, 1942 CampopiA: Prek‘Phnau, 6 kilometres northeast of Phnom Penh. 3 g¢, 3 29. Large-footed Myotis of southeastern Asia stand in need of revision, but at the present time there is insufficient material in the collections of the British Museum (Natural History) from a suitably wide range of localities for this to be undertaken in detail. Specimens attributed to the named forms horsfieldii, hasseltii, and adversus have been reviewed recently (Hill, in press) and there seems little doubt that M. horsfieldii (Temminck, 1840) must be considered a distinct species (Medway, 1965 : 60) on account of its small size, blackish coloration and narrow braincase when compared with hasseltii and adversus. Myotis deignant Shamel, 1942, from Chiangmai, northern Thailand seems likely to be a subspecies of horsfieldit. Myotis hasseltii (Temminck, 1840) applies apparently to larger specimens in which the pelage is short, the post-palatal extension short and lacking thin bony laminae to support the post-palatal spine, in which i? and 18 are relatively massive, the second upper premolar (pm?) is minute, usually intruded from the toothrow and the second lower premolar (pmg) very small, intruded from the toothrow sometimes to the extent that pmg and pm, are in contact or nearly so. Myotis adversus (Horsfield, 1824) is characterized by dense, woolly pelage, a long post-palatal extension with thin bony laminae supporting the post-palatal spine, pm? less reduced, usually not much intruded from the toothrow and with pmg although reduced, in the toothrow or only very slightly intruded. Myotis hasseltii is distributed from Ceylon, Thailand and Indochina to Malaya, Java and Borneo. Specimens from Thailand and Cambodia agree closely with the description of Myotis adversus continentis Shamel, 1942 : 323 which apparently is referable to hasseltiit rather than to adversus. Shamel says ‘fur velvety and short’: pm? is crowded inwards, with pm? and pm? in contact, while pmg is small, but in the holotype standing in the toothrow. According to Medway (1965 : 62) M.h. macellus (Temminck, 1835) is the Bornean subspecies. Myotis adversus occurs in Java (M.a. adversus) and Borneo (M.a. cavimatae Miller, 1906) east to Australia (M7.a. moluccarum (Thomas, 1915) ). Minimum, maximum and mean length of forearm in five specimens of M. hasseltit continentis from Cambodia 36-8—39-2 (38-2); measurements of three skulls (g, 29): greatest length 15:3, 15-7, 15:8; condylobasal length 14-4, 14:6, 14-6; least interorbital width 4-2, 4-1, 4:0; zygomatic width 10-2, 9-9, 10-2; width of braincase 7-8, 7-6, 7-°8; mastoid width 8-6, 8-4, 8-6; cl-c! 4-3, 4:3, 4:3; m’—m3 6-1, 6-2, 6:0; c-m3 5:5; 5-6, 5-7; length of mandible —, 11-1, 10-9; c—mg 6-1, 6-1, 6:2. Myotis annectans (Dobson, 1871) THAILAND: Doi Pui, Chiangmai, 1,250 metres. 1 9. Collected by Kitti Thonglongya and now in the Thai National Reference Collection. Topal (1g70a) has recently examined the holotype of Pzpzstrellus annectans Dobson, 1871: 213 from the Naga Hills, Assam, in the Indian Museum, Calcutta and concluded that despite the absence of the small second premolars (pm$) it should be referred to Myotis. Furthermore, Topal has concluded that it is synonymous BATS FROM THAILAND AND CAMBODIA 189 with Myotis primula Thomas, 1920 : 248 from Pashok, near Darjeeling, northeastern India, which it antedates by many years. A result of this conclusion is that the subgeneric name Megapipistrellus Bianchi, 1917 with type species Pipistrellus annectans Dobson must be transferred to Myotis. Apart from the original, the only other record of P. annectans seems to be the report by Schneider (1905 : 80) of three specimens identified by Leche from the Upper Langkat, Sumatra and at that time in the Zoological Institute of the University of Stockholm. Schneider gave no diagnostic features but subsequently the specimens were described in detail by Arnbiack-Christie-Linde (1909 : 574). They are, however, much too small to be referred to annectans. The specimen from Chiangmai agrees exactly with the description of Pipistrellus annectans by Dobson and provides an opportunity to describe the coloration of the species, the holotype being in alcohol, and also to make a direct comparison with the holotype of Myotis primula. Dorsally, the Chiangmai specimen is rich dark chestnut brown, the individual hairs dark brown at the base, tipped with paler brown. The ventral surface, including the underside of the head and throat, is greyish for the most part, with brown underlay, the hairs dark brown at the base, heavily tipped with greyish white. The hair tips on the belly are ochraceous or orange to produce an orange yellow median patch. The pelage is rather long and woolly, extending slightly on to the endopatagium but hardly at all on to the uropatagium. The face is densely hairy, excepting the nostrils and the areas around the lips and eyes, the pelage forming a dense fringe on the forehead. Ears translucent distally, less so proximally, of moderate size with rounded tips. The anterior margin of the ear is strongly convex proximally, less so distally, the posterior margin sharply concave just below the tip, becoming convex, slightly concave again near the base of the tragus, with a small antitragal lobe. The tragus is long and tapering, with an acute, rounded tip, its anterior margin slightly concave, the posterior margin convex and with a small rounded basal lobe. There is no post-calcarial lobe and the extreme tip of the tail is free of the membrane. The skull has an elongate, uninflated braincase, relatively wide unconstricted interorbital region and slight sagittal crest. The rostrum is low and narrow anteriorly, with shallow median frontal sulcus and short, U-shaped anterior narial emargination. The large anteorbital forearm is separated from the orbit by a moderate bar of bone with a small upper subsidiary foramen within the orbit. The palate is sharply domed, narrowed anteriorly, with U-shaped anterior emargination and wide ligulate post-palatal spine. The inner upper incisor (i?) is large, slightly longer than wide, with large anterior cusp and lower posterior cusp: i? is wider than long, transverse to the toothrow, with a heavy cusp rising from a narrow cingulum and separated from the canine by a narrow diastema. Lower incisors with no especial peculiarities, canines low, c! about the same height as pm‘, c; similarly about the same height from the cingulum as pmy. Anterior upper premolar (pm?) large, in toothrow, touching cl and pm4: pmg not reduced or displaced, in contact with c; and pmy. No trace of pm in either jaw. The specimen agrees precisely with the diagnosis by Dobson of P. annectans, which it obviously represents. A direct comparison with the holotype and other material of Myotis primula 190 Jo EPH een. WS any Thomas, 1920 shows exact agreement in almost every respect. The ears in primula are not translucent distally as they are in the Chiangmai specimen which is very slightly more orange ventrally than primula but in every other external feature there is complete correspondence. The skulls and teeth agree precisely excepting only for the presence of a minute pm? in primula: its degree of intrusion, however, varies in the specimens examined and although in the holotype (B.M.16.3.25.30) the anterior (pm3) and posterior (pm{) premolars do not touch, in two others (B.M.20.7.27.2-3) these teeth are in contact or nearly so. There can be no doubt that annectans is in fact primula in which these teeth have never appeared: the ears and tragus of annectans and primula refer the taxon which they represent indubitably to Myotis, as Topal has concluded. Measurements of M. annectans trom Chiangmai are compared with those of M. primula in Table 2. TABLE 2 Measurements of Myotis annectans ‘M. primula’ 3 M. annectans 3 Holotype ‘M. primula’ 3 ‘M. primula’ 2 Chiangmai, B.M.16.3.25.30 B.M.20.7.27.2 B.M.20.7.27.3 Thailand Assam Assam Assam Length of forearm 43°3 47:0 40°5 45°2 Greatest length of skull 17:0 173 16-7 oS Condylobasal length 16°4 16-7 15°9 — Condylocanine length 15'3 15:60 149 — Least interorbital width 4:2 43 43 4:2 Zygomatic width II-2 II-5 — — Braincase width 8-0 8-4 8-6 7:8 Mastoid width 8:8 8-6 8-5 8-4 ci-cl 45 48 48 46 m3—m3 76 73 74 72 c-m3 6-7 6-9 6-6 6-7 Length of mandible 12°3 12:7 — — c-mg3 73 73 71 72 Myotis siligorensis thaianus Shamel, 1942 THAILAND: Tham Tab Tao, Fang, Chiangmai. 299. Thai National Reference Collection. These are apparently the first of M.s. thaianus to be reported since its description from Chiangmai by Shamel (1942 : 323). Measurements: length of forearm 31:7, 31°9; greatest length of skull 12-8, 12-7; condylobasal length 11-8, —; condylocanine length 11-1, —; least interorbital width 2:8, 2-9; zygomatic width 7-0; —; braincase width 5-8, 6-0; mastoid width 6-4, 6-6; depth of braincase 4-7, —; cl-c! 3-0, 3:0; m3—m8 4:6, 4-7; length of entire toothrow 5:4, 5:3; c-m® 4:5, 4:4; length of mandible 8-4, 8-6; c-m3 4:9, 4:8. BATS FROM THAILAND AND CAMBODIA 191 Pipistrellus cadornae Thomas, 1916 THAILAND: Petchabun, Thung Salang, Luang, 450 metres. 1g. Thai National Reference Collection. The first of this species to be reported from Thailand, this specimen is in excellent agreement with the holotype and with those reported from Upper Burma by Hill (1962 : 133). Measurements: length of forearm 33-2, greatest length of skull 13-6; condylobasal length 12-9, condylocanine length 12-6; least inteorbital width 3-7, width of braincase 6-7; mastoid width 7:5; cl—c! 4-5, m8—m8 5-7 c—m3 4:5; c—mg 4:7. Hesperoptenus tickelli (Blyth, 1851) Tuairanp: Nakhon Rat Sima (= Khorat). 1 9. Hesperoptenus tickelli has been recorded hitherto from no further east than Bengal on the Asian mainland and from the Andaman Islands so that its presence in eastern Thailand represents a considerable extension of range. In colour this specimen agrees more closely with Indian examples than with specimens from Ceylon and the Andaman Islands, which are very slightly darker dorsally, especially on the hindback. Measurements: length of forearm 52:3; greatest length of skull 18-5; condylobasal length 18-3; least interorbital width 5-0; zygomatic width 14-1; width of braincase 9-9; mastoid width 11-1; cl-c! 6-3; m3—m8 9-2; c-m? 7-4; length of mandible 14-1; c—mg 8-0. Hesperoptenus blanfordi (Dobson, 1877) THAILAND: Khao Luang, Nakhon Si Thamrat, 450feet. 8°26'N.,99°58’E. 16. This rare species has been recorded once previously from Thailand, by Robinson and Kloss (1915b: 116), who listed a specimen from Khao Nawng, Bandon. Measure- ments of the specimen from Khao Luang: length of forearm 25-8, greatest length of skull 12-2; condylobasal length 11-6; least interorbital width 4-3; zygomatic width —; width of braincase 7-0; mastoid width 7-2; cl-c! —; m3—m3 6-2; c-m3 3-9; length of mandible 8-4; c-m3 -—. Scotophilus kuhlii gairdneri Kloss, 1917 THAILAND: Koh Klet, Pak Klet, Nonburi, 11 miles north of Bangkok. 1 9. There is some uncertainty in the literature as to the correct classification of Asian Scotophilus, but there are, however, several partial studies of the Asian species. Briefly discussed by Sody (1928 : 86), the Asian forms were reviewed in some detail by Tate (1942 : 283) while Shamel (1942 : 325) provided a succinct examination of specimens in the United States National Museum which has remained largely over- looked by subsequent authors More recently, Siddiqi (1961) reviewed the Indian and some southeast Asian forms, while Peterson (1968) has reviewed the outstanding problems in the genus so far as southeastern Asia is concerned The consensus of opinion is that two species should be recognized, a larger and a smaller, sympatric in some places (excluding the very large Scotomanes (Parascoto- manes) beauliewt Bourret, 1942 from North Vietnam, listed in Scotophilus by 192 fi Da Te WALR Be ce Ihe CIPD Ellerman and Morrison-Scott, 1951 : 180 but considered synonymous with Ja 10 by Topal, 1970b : 342). Tate, Shamel and Siddiqi assign smaller specimens to Scotophilus temminckii Horsfield, 1824: larger specimens are referred to S. heathi Horsfield, 1831 by Tate and Siddiqi and to S. kuhlii Leach, 1822 by Shamel. Examination of specimens in the British Museum (Natural History) confirms this division into two size groups. A major difficulty arises in the allocation of names to these groups through the uncertain status of Scotophilus kuhlit Leach, 1822, the type species of Scotophilus, neither its specific identity nor its locality having been satisfactorily established. The name was considered indeterminable by Sody, Tate (who suggested a further study of the type specimen) and Siddiqi: over the years it has been used both for the smaller and the larger species. However, Peters (1867b : 679) thought the juvenile type specimen to be an example of S. temminckii (Horsfield, 1824), a conclusion reiterated by Dobson (1875 : 368) and by Blanford (1888 : 267), who also noted that an examination of the specimen by Thomas showed its upper incisors to be like those of temminckii. The type specimen was re-examined recently at the request of Dr R. L. Peterson, of the Royal Ontario Museum, who has published in part (1968 : 1081) the results of this survey. In brief, the holotype consists of a skull and. specimen in alcohol, labelled India. It is a very young individual, the third upper molar just piercing the gum, with the two tricuspid milk incisors still present on each side of the jaw. On the right the unicuspid permanent incisor is emergent alongside the inner of these: on the left the permanent tooth has been lost from the damaged socket, only the two milk incisors remaining. Specimens of similar age in the collections of the British Museum (Natural History) are very like this, and the holotype seems at one of the normal stages of dental development in the genus. The dimensions of the teeth indicate that it represents the smaller of the two Asiatic species undef consideration. The available evidence suggests that it originated from India: the name kuhlii should therefore replace temmincki for the smaller species, while wroughtoni Thomas, 1897, hitherto considered the Indian subspecies, will become a synonym of S.k. kuhlii. Siddiqi (1961 : 452 listed castaneus Gray, 1838 from West Bengal, East Pakistan to southwestern China and the Malay Peninsula. This author, however, did not examine gairdneri Kloss, 1917 from Thailand. Sub- species allocated to kuhlii are listed in full by Tate (1942 : 285) and Shamel (1942 : 327): for the present gairdneri is retained for Siamese and Cambodian specimens on account of their slightly darker dorsal coloration when compared with Indian specimens (kuhliz) : castaneus from the Malay Peninsula is dorsally similar to gairdnert but is darker ventrally. There is a wide degree of individual colour variation, however, and these differences are small: at best it seems that the subspecies are only weakly separable. Length of forearm (eight specimens) 49-I-51-2 (50-3). Scotophilus heathi (?) watkinsi Sanborn, 1952 THAILAND: Bang Phra, Siracha, Chonburi, 13° 12’ N., 100° 57’ E. 1. Koh Klet Pak Klet, Nonburi, 11 miles north of Bangkok. 1 9. Bangkok. 1 sex uncertain (skull only). BATS FROM THAILAND AND CAMBODIA 193 The collections of the British Museum (Natural History) include extensive series of S. heathi from Indian and Burmese localities (listed by Wroughton (1918a : 594) as S. kuhlit) but the representation from Thailand and Indochina is very much poorer. As in S. kuhliz, there is considerable individual colour variation and for the present no attempt has been made to determine subspecific boundaries. These specimens are similar in size to those from Pak Nam Pho, Nakon Sawan Province, central Thailand described as S. solutatus watkinsi by Sanborn (1952 : 4) to which they are provisionally referred. The description and measurements lead this to be allocated to S. heathi, and it may represent a valid subspecies in Thailand. Measurements (g, 2, sex uncertain): length of forearm 61:2, 61-4, —; greatest length of skull 22-6, 22-5, 22:0; condylobasal length 20-8, 20-9, 20-3; palatal length 7-7, 7°8, 7-6; rostral width between anteorbital foramina 8-3, 8-3, 7-9; least inter- orbital width 5-4, 5:3, 5:3; zygomatic width 16-4, 16-5, 15-9; mastoid width 14-2, 14:4, 13:8; braincase width 10-6, 10-7, 10-2; cl-cl 7-6, 7-7, 7-4; m3—m3 10:3, I0:2, 9:8; com 7-6, 7-7, 7:5; length of mandible 15-5, 16-0, 15:3; c-mg3 8°8, 8:9, 8-7. Murina cyclotis cyclotis Dobson, 1872 THAILAND: Tham Tab Tao, Fang, Chiangmai. 1 ¢ (young adult). The genus Murina was recorded from Thailand by Shamel (1942 : 327) who identified two specimens from the northern part of the country as M. toxopei Thomas, 1923 (= M. florium toxopet) known otherwise from the island of Buru, in the Moluccas. This specimen from Chiangmai is quite clearly referable to M. cyclotis cyclotis as it is described by Hill (1963 : 53), agreeing closely with specimens in the British Museum (Natural History) from northem Burma. It is probable that the specimens described and measured by Shamel also represent M.c. cyclotis with which from the account they agree in most respects. Tadarida plicata plicata (Buchanan, 1800) THAILAND: Phu Nam Tok Tap Kwang, Kaeng Khoi, Sara Buri, c. 14° 42’ N., 100° 52’ E. 16. CamBopiA: Angkor Wath, Siem Reap, Cambodia. 1 9. Tonle, 25 kilometres southeast of Phnom Penh, Cambodia. 1,5 99. Yoshiyuki (1966 : 40) reported T.p. plicata for the first time from Cambodia. Length of forearm in the Thai example 47-7; in specimens from Cambodia 46-9- 49°0 (48-0). SUMMARY Collections of bats from Thailand and Cambodia examined recently at the British Museum (Natural History) have included a number of species new to these countries or of taxonomic significance. Megaerops ecaudatus, Chironax melanocephalus, Myotis annectans, Pipistrellus cadornae and Hesperoptenus tickelli are reported for the first time from Thailand, together with Pteropus hypomelanus from the islet of Koh Som in the South China Sea. The first specimens of Taphozous longimanus 194 ji. Ee MIL se) ie er any longimanus, Rhinolophus borneensis and R. acuminatus trom Cambodia are recorded, with further examples of Tadarida plicata plicata, known hitherto in that country from one reported occurrence. Further specimens reported of species poorly known from Thailand include Rousettus amplexicaudatus, R. leschenaulti leschenaulti, Sphaerias blanfordi, Macroglossus minimus sobrinus, Taphozous theobaldi theobaldi, Rhinolophus malayanus, R. acuminatus, Coelops frithiit, Myotis hasselti continentis, M. siligorensis thaianus, Hesperoptenus blanfordi and Murina cyclotis cyclotis. An examination ot Cynopterus angulatus Miller, 1908 shows it to be a subspecies of C. sphinx rather than of C. brachyotis. Further specimens of Rhinolophus coelophyllus shameli Tate, 1942 indicate that shameli is specifically distinct from R. coelophyllus. R. chaseni Sanborn, 1939 is thought to be a subspecies of R. borneensis. Large- footed Myotis of southeastern Asia are discussed and continentis Shamel, 1942 thought to be a subspecies of M. hasseltii rather than of M. adversus as it was described. A specimen referable to Pipistrellus annectans Dobson, 1871 confirms the view that this taxon should be referred to Myotis and that M. primula Thomas, 1920 is its synonym. The holotype of Scotophilus kuhlit Leach, 1822 has been examined and this name applied to the smallest of the Asiatic species of Scotophilus, formerly referred to S. temminckiw (Horsfield, 1824). REFERENCES ALLEN, G. M. & CootipGe, H. J. 1940. Mammal and bird collections of the Asiatic Primate Expedition, 3. Mammals. Bull. Mus. Comp. zool. Harv. 87 : 131-166. ANDERSEN, K. 1905. On some bats of the genus Rhinolophus, with remarks on their mutual affinities, and descriptions of twenty-six new forms. Proc. zool. Soc. Lond. 2 : 75-145, 22 figs., 2 pls. 1912. Catalogue of the Chiroptera in the collection of the British Museum. I. Mega- chivoptera. London. 1916. In Ktoss, C. B. Ona collection of mammals from the coast and islands of south- east Siam. With an account of the fruit bats, by Dr Knud Andersen. Proc. zool. Soc. Lond. 27-75, 1 pl. —& Ktoss,C.B. 1915. Remarks on some races of Cynopterus. J. fed. Malay St. Mus. 5 : 220-222. ANDERSON, J. 1881. Catalogue of the Mammalia in the Indian Museum, Calcutta. Part I. Primates, Prosimiae, Chiroptera, and Insectivova. Calcutta. ARNBACK-CHRISTIE-LINDE, A. 1909. On intermediate forms among Chiroptera. Zool. Anz. 34 : 572-582, 4 figs. Buat, H. R. 1968. Sphaerias blanfordi (Thomas, 1891) from Himalayan region of Uttar Pradesh: an addition to the Chiropteran fauna of India. J. Bombay nat. Hist. Soc. 65 : 471-473, I tab. BLANForRD, W.T. 1888. Notes on Indian Chiroptera. J. Asiat. Soc. Beng. 57, (2) : 260-272. Bonuote, J. L. 1903. Anthropological and Zoological results of an expedition to Perak and the Siamese Malay States, 1901-1902. Undertaken by Nelson Annandale and Herbert C. Robinson. Report on the mammals. Fasciculi Malayenses 1 : 1-44, 4 pls. Bourret, R. 1944. Mammiféres récemment entrés dans les collections du laboratoire de zoologie de l’Ecole Superieure des Sciences — description d’une varieté nouvelle. Notes Trav. Ec. sup. Sci. Hanoi 3 : 1-17. CuHaAsEN, F. N. 1940. A handlist of Malaysian mammals. Bull. Raffles Mus. No. 15: 1-xx, 1-209, map. —— & Ktoss, C. B. 1932. Mammals from lowlands and islands of North Borneo. Bull. Raffles Mus. No. 6 (1931) : 1-82. BATS FROM THAILAND AND CAMBODIA 195 Dogson, G. E. 1871. Notes on nine species of Indian and Indo-Chinese Vespertilionidae, with remarks on the synonymy and classification of some other species of the same family. Proc. Asiat. Soc. Beng. 210-215. 1875. On the genus Scofophilus, with description of a new genus and species allied thereto. Proc. zool. Soc. Lond. 368-373. 1878. Notes on recent additions to the collection of Chiroptera in the Muséum d'Histoire Naturelle at Paris, with descriptions of new and rare species. Proc. zool. Soc. Lond. 873-880. ELLERMAN, J. R. & Morrison-Scort, T. C. S. 1951. Checklist of Palaearctic and Indian mammals, 1758-1946. 1st. ed. London. GYLDENSTOLPE, N. 1916. Zoological results of the Swedish Zoological Expeditions to Siam Tg1I—-1912 & 1914-1915. K. svenska Vetensk Akad. Handl. 57, (2) : 1-59, 6 pls. HorsFIELD, T. 1851. A catalogue of the Mammalia in the Museum of the Hon. East-India Company. London. Hit, J.E. 1961. Fruit-bats from the Federation of Malaya. Proc. zool. Soc. Lond. 136 : 629- 642, 1 fig., 4 tabs. —— 1962. Notes on some insectivores and bats from Upper Burma. Proc. zool. Soc. Lond. 139 : 119-137, 5 tabs. 1963. Notes on some tube-nosed bats, genus Muvina, from southeastern Asia, with descriptions of a new species and a new subspecies. Fedn. Mus. J. N.S. 8 : 48-59, 4 pls. (in press). New records of Malayan bats, with taxonomic notes and the description of a new Pipistrellus. Bull. Br. Mus. nat. Hist. (Zool.) 23 : 21-42, 3 tabs. Koss, C. B. 1911. Ona collection of mammals and other vertebrates from the Trengganu Archipelago. J. Fed. Malay St. Mus. 4 : 175-212. 1916. Ona collection of mammals from the coast and islands of south-east Siam. With an account of the fruit-bats, by Dr. Knud Andersen. Proc. zool. Soc. Lond. 27-75, 1 pl. 1917. Ona third collection of Siamese mammals. J. nat. Hist. Soc. Siam 2 : 288-318. 1919. On mammals collected in Siam. J. nat. Hist. Soc. Siam 3 : 333-407, 1 pl. Linpsay, H.M. 1927. Bombay Natural History Society’s Mammal Survey of India, Burma and Ceylon. Report No. 39, Mergui Archipelago. J. Bombay nat. Hist. Soc. 31 : 42-48. MarsHaLt, J. & VANDEE, N. 1970. Mammals of Samui Island, Thailand. Nat. Hist. Bull. Siam Soc. 23 : 501-507, 1 pl., 1 tab. Mepway, Lorp. 1965. Mammals of Borneo. Field keys and an annotated checklist. Io Malay. Brch R. Asiat. Soc. 36, (1963) : i-xiv, 1-193, 9 figs., 34 pls., 5 tabs., map. —— 1969. The wild mammals of Malaya and offshore islands including Singapore. Kuala Lumpur; Singapore; London. MILLER, G. S. 1898. List of bats collected by Dr. W. L. Abbott in Siam, Proc. Acad. nat. Sci. Philad. 316-325. —— 1906. Seven new Malayan bats. Proc. biol. Soc. Wash. 19 : 61-65. Oscoop, W. 1932. Mammals of the Kelley—Roosevelts and Delacour Asiatic Expeditions. Publs. Field Mus. nat. Hist. (Zool. Ser.) 18 : 193-339, 30 figs., 2 pls. PETERS, W. 1867a. On some mammals collected by Capt. A. C. Beavan, C.M.Z.S., at Moulmein, Burmah. Proc. zool. Soc. Lond. (1866) : 426-430, 1 pl. —— 1867b. Fernere Mittheilungen zur Kenntnifs der Flederthiere namentlich iiber Arten des Leidener und Britischen Museums. Mber. k. preuss. Akad. Wiss. (1866) : 672-681. Peterson, R.L. 1968. Notes on an unusual specimen of Scotophilus from Vietnam. Can. fe Zool. 46 : 1079-1081, 1 fig. Puituirs, C. J. 1967. A collection of bats from Laos. J. Mammal. 48 : 633-636, 1 tab. PousarcuEs, E. DE. 1904. Mammiferes de I'Indo-Chine. In Pavie, A.J.M. Mission Pavie Indo-Chine 1879-1895. Etudes Diverses. III Recherches sur l’Histoive Naturelle de ?’Indo- Chine Orientale. Paris. Roxrnson, H.C. & Kioss,C.B. 1915a. The zoology of Koh Samui and Koh Pennan. J. fed. Malay St. Mus. 5 : 128-152. 196 [je to ISBWUAl, ee AN, eae Roprnson, H. C. & Kross, C. B. t915b. On a collection of mammals from the Siamese Province of Bandon, N.E. Malay Peninsula. J. fed. Malay St. Mus. 5 : 111-127. 1918. Results of an expedition to Korinchi Peak, SumatraI. Mammals. J. fed. Malay St. Mus. 8 (2) : 1-80, 1 pl. SANBORN, C. C. 1952. The mammals of the Rush Watkins Zoological Expedition to Siam. Nat. Hist. Bull. Siam. Soc. 15 : 1-20. SCHNEIDER, G. 1905. Ergebnisse zoologischer Forschungsreisen in Sumatra. Saugetiere (Mammalia). Zool. Jb. (Syst.) 23 : 1-172, 1 fig., 3 pls., 2 maps. SHAMEL, H.H. 1942. A collection of bats from Thailand (Siam). J. Mammal. 23 : 317-328. Srppig1, M.S. U. 1961. Notes on the status of bats of the genus Scotophilus from southern Asia with key to the recognized forms. Ann. Mag. nat. Hist. (13), 3, (1960) : 449-454. Sopy, H. J. V. 1928. Twee nieuwe subspecies van Pachyotis temmincki Horsf. Natuurk. Tijdschr. Ned.-Indié, 88 : 86-91. Tate, G. H. H. 1942. Results of the Archbold Expeditions. No. 47. Review of the Vespertilionine bats, with special reference to genera and species of the Archbold Collections. Bull. Am. Mus. nat. Hist. 80 : 221-297, 5 figs. 1943. Results of the Archbold Expeditions. No. 49. Further notes on the Rhinolophus philippinensis group (Chiroptera). Am. Mus. Novit. No. 1219 : 1-7. — & ARcHBOLD, R. 1939. Results of the Archbold Expeditions. No. 24. Oriental Rhinolophus, with special reference to material from the Archbold Collections. Am. Mus. Novit. No. 1036 : 1-12, 1 tab. THomas, O. 1920. Scientific results from the Mammal Survey. No. XXIII. A new bat of the genus Myotis from Sikkim. J. Bombay nat. Hist. Soc. 27 : 248-249. Topat, G. 1970a. On the systematic status of Pipistvellus annectans Dobson 1871 and Myotis primula Thomas 1920 (Mammalia). Anmls hist.-nat. Mus. natn. hung. Zool. 62 : 373-379, 2 tabs. 1970b. The first record of Ja io Thomas, 1902 in Vietnam and India, and some remarks on the taxonomic position of Pavascotomanes beaulieui Bourret, 1942, Ia longimana Pen, 1962 and the genus Ja Thomas, 1902 (Chiroptera, Vespertilionidae). Opusc. zool. Bpest 10 : 341-347, 2 figs., 2 tabs. VAN PEENEN, P. F. D., Ryan, P. F. & Licnt, R.H. 1969. Preliminary Identification Manual for mammals of South Vietnam. Washington. ——, Cunnincuam, M. L. & Duncan, J. F. 1970. A collection of mammals from Con Son Island, Vietnam. J. Mammal. 51 : 419-424, 1 fig. WroucuHTon, R. C. 1915. Bombay Natural History Society’s Mammal Survey of India, Burma and Ceylon. Report No. 17. South Tenasserim. jJ. Bombay nat. Hist. Soc. 23 : 695-720. 1918a. Summary of the results from the Indian Mammal Survey of the Bombay Natural History Society. J. Bombay nat. Hist. Soc. 25 : 547-598. 1918b. Summary of the results from the Indian Mammal Survey of the Bombay Natural History. J. Bombay nat. Hist. Soc. 26 : 19-58. — & Davinson, W. M. 1918. Bombay Natural History Society’s Mammal Survey of India, Burma and Ceylon. Report No. 29. Pegu. J. Bombay nat. Hist. Soc. 25 : 472-481. Yosuivuki, M. 1966. On Tadarida plicata plicata newly collected from Cambodia. J. mammal Soc. Japan 3 : 40-42. LF , 9 Kittt THONGLONGYA {5 1d5leheae, THAI NATIONAL REFERENCE COLLECTION Department of Zoology APPLIED SCIENTIFIC RESEARCH CORPORATION British Museum (NATURAL History) oF THAILAND CROMWELL ROAD 196, PHAHON YOTHIN Roap London, SW7 5BD Banc KHEN, BANGKOK — 9 A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) Kay, E. Attson. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 1965. (Out of Print.) £3.75. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180; 11 Plates, 15 Text-figures. 1967. £4. Taytor, J. D., Kennepy, W. J. & Hatt, A. The Shell Structure and Mineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates, 77 Text-figures. 1969. £4.50. Haynes, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur) 1962-1964. (In press.) 5 laa Printed in England by Staples Printers Limited at their Kettering, Northants, establishment 7Taw A REDESCRIPTION OF q PELAEOMYSIS SERVATUS (FAGE) ey COMB, NOV, (MYSIDACEA : LEPIDOMYSIDAE) .OM THE MATERIAL COLLECTED ON ALDABRA ATOLL, a WT A KEY HhO THE SPECIES OF LEPIDOMYSIDAE Sh hy * es C 4S ys ff MRE say, 7 V/ AGR 29 A PUTED BULLETIN OF MUSEUM (NATURAL HISTORY) ‘a 3 my Vol. 22 No. 7 eee Eh LONDON : 1972 AoREDESCRIPTION OF SPELAEOMYSIS SERVATUS (FAGE) COMB. NOV. (MYSIDACEA : LEPIDOMYSIDAE) FROM THE MATERIAL COLLECTED ON ALDABRA ATOLL, WITH A KEY TO THE SPECIES OF LEPIDOMYSIDAE BY RAYMOND WILLIAM INGLE Pp 197-210; 5 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 22 No. 7 LONDON : 1972 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) instituted in 1949, its issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at regular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 10965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 22 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), 1972 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 30 March, 1972 Price 65p A REDESCRIPTION OF SPELAEOMYSIS SERVATUS (FAGE) COMB. NOV. (MYSIDACEA : LEPIDOMYSIDAE) FROM THE MATERIAL COLLECTED ON ALDABRA ATOLL, WITH A KEY TO THE SPECIES OF LEPIDOMYSIDAE By R. W. INGLE SYNOPSIS The crustacean Spelaeomysis servatus (Fage) is redescribed from specimens collected on Aldabra Atoll. The species has been transferred from the genus Lepidomysis Clarke. A key is provided for the identification of the known species in the genus Spelaeomysis. INTRODUCTION Amonc the crustaceans collected by Dr. K. G. McKenzie during phase III of the Royal Society of London Expedition to Aldabra (see McKenzie, 1971), were a number of mysidaceans that agree substantially with the descriptions and figures by Fage (1924, 1925) of a species designated by him as Lepidophthalmus servatus. For nomenclatorial reasons the generic name used by Fage was replaced by Lepidops (Zimmer, 1927) and later by Lepidomysis (Clarke, 1961). The type material of this species is from S. Zanzibar, E. Africa. As far as I am aware this material from Aldabra constitutes the second occurrence of this species and extends its range some 400 miles south eastward of the type locality. Unlike the type material, these Aldabran specimens are well preserved. Adults and juveniles of both sexes are represented, although there are no ovigerous females in the sample. The opportunity has been taken, therefore, to redescribe the species from this fresh material and to compare it with the other known species of Lepidomy- sidae, Spelacomysis bottazzii Caroli, 1924, S. quinterensis (Villalobos, 1951), and S. longipes Pillai and Mariamma, 1964. These comparisons have shown that Lepi- domysis cannot be retained as a separate genus and that L. servatus, the only species in the genus, must be transferred to Spelaeomysis. The reasons for this decision are discussed on p. 207. The definition of the genus Spelaeomysis is amended slightly and a key is provided to separate the species belonging to this genus. 200 R. W. INGLE Fic. 1. Spelaeomysis servatus (Fage). Male, 5-5 mm, t.l. Aldabra Atoll. A. Dorsal view showing chromatophore pattern, thoracic limbs omitted. B. Lateral view showing projecting scale of penultimate thoracic segment. C. Left mandible, ventral aspect, lower inset — distal portion of right mandible. D. Right antennule, peduncle and proximal flagella segments, ventral aspect. E. Left antenna, peduncle and proximal flagellum segments, ventral aspect. F. Telson, dorsal aspect. Scales in mm. A REDESCRIPTION OF SPELAEOMYSIS SERVATUS (FAGE) 201 SYSTEMATICS Family LEPIDOMYSIDAE Clarke, 1961 : 251 Spelaeomysis servatus (Fage) comb. nov. Lepidophthalmus servatus Fage, 1924 : 2127; 1925 : 525, figs 1-7. Lepidops servatus; Zimmer, 1927 : 644; Gordon, 1960 : 299, 308, figs 20, 21, 27, 28. Lepidomysis servatus; Clarke, 1961 : 251. MATERIAL EXAMINED. 20 3g, 3:0-5°5 mm, 17 99, 4:0-6-0 mm, (total lengths measured from distal margins of eyes to distal margins of telsons). Station 30, Aldabra Atoll, Indian Ocean, lat. 9° 24’S., long. 46° 20’ E, local coordinates used by Royal Soc. Exped. 3755, 0525. Tide-dominated saline seepage pool in Pemphis scrub. Salinity 23-5°/oo. Associated fauna: Civolana sp. marine molluscs, marine alga Caulerpa verticillata. Collected by Dr. K. G. McKenzie, 1.2.1968. Description. Male (fig. 1A), body sub-cylindrical, carapace lateral margins strongly deflected ventrally, posterior expansions reaching to middle of first abdom- inal segment, antero-lateral expansions extending beyond rostral apex, rostrum broadly rounded. Carapace with a pair of faint longitudinal grooves, extending forward from posterior margin of carapace to its middle and then strongly deflected ventrally. Last two thoracic segments dorsally exposed, penultimate segment with a well developed scale overlapping the postero-median carapace margin and shown in lateral aspect in fig. 1B. Outline of thoracic and abdominal segments continuous. First abdominal segment shortest, sixth segment longest. Postero-ventral margins of segments 3-5 each with a single broad lamella arising between the pleopod bases and increasing in size respectively on each segment. Telson (fig. 1F) linguiform, posterior margin with a long median spine flanked on either side by one short and two long spines; lateral margins unarmed proximally, but with a row of 9 or Io spines in the distal two thirds, increasing in length posteriorly. Each eyestalk (eye-plate) flattened dorso-ventrally and sub-triangular, narrowing anteriorly with inner margin curved, the cornea well developed. Antennule (fig. 1D), first peduncular segment longer than second segment, third sub-equal to second, inner flagellum reaching to middle of third abdominal segment, outer flagellum slightly shorter than inner one and with aesthetascs on inner margins of segments, decreasing in length distally. Antenna (fig. 1Z), sympod three-segmented, first to third seg- ments incompletely separated, third segment with a sub-acute process on outer distal margin. Exopod (scale) sub-oval, about half as long again as first segment of endopod peduncle, whole margin of scale setosed except near base. Peduncle of endopod formed of two sub-equal segments, flagellum reaching to second abdominal segment. Labrum outline shown in fig. 2D. Left mandible (fig. 1C), with a strongly bifid incisor process, each bifurcation faintly dentate, lacinia mobilis prominent and dentate, four penicils (spines) present, molar process truncate. Right mandible with lacinia mobilis bifid (inset to fig. 1C), and with molar process differing in shape from that of left mandible. Palp stout and three-segmented, first segment short, second longest and with pectinate and simple setae, distal segment narrowest and with long pectinate and simple setae. Maxillule (fig. 2B), proximal endite with five setae (three distal and two proximal) and four weakly 202 R. W. INGLE Fic. 2. Spelaeomysis servatus (Fage). Male, 5-5 mm. t.l. Aldabra Atoll. A. Labium. B. Right maxillule, right inset—distal portion of major lobe. C. Right maxilla. D. Labrum. E. First right thoracic limb. All from ventral aspect. Scales in mm. A REDESCRIPTION OF SPELAEOMYSIS SERVATUS (FAGE) 203 pectinate long spines distally. Distal endite with three subterminal setae, fourteen or fifteen strong spines (some pectinate) and one or two simple setae. Maxilla (fig. 2C), exopod oval and with twenty nine or thirty long plumose setae, distal lobe well developed, proximal lobe undifferentiated, distal endopod segment (en) smaller than proximal segment, latter with a slightly expanded inner margin, endite of second segment (e2) of sympod and bifid endite of third segment (e3) well developed. Maxilla with simple and pectinate setae of types shown in insets to fig. 2C. Labium (fig. 2A), symmetrical and deeply cleft, inner margins of each lobe setose. First thoracic limb (fig. 2E), with coxa not clearly demarcated from basis (0), inner margin of basis expanded. First, second (2) and fourth endopod segments with inner margins expanded, fifth (5) segment truncate. Inner margin of fourth segment invested with a row of stout pectinate setae. Outer distal margin of third segment with three long spines, remaining segments of endopod with plumose or with simple setae as shown in figure. Exopod (ex) reduced to a lanceolate scale bearing two terminal setae and a row of fine short hair-like setae on both margins, epipod long. Second thoracic limb (fig. 3A), coxa demarcated from basis, first endopod segment almost as long as second segment, third segment longest, fourth longer than fifth. Inner margins of third to fifth segments with stout pectinate and plumose setae. Exopod well developed and nine-segmented, basal segment longest, seven distal segments each with a pair of plumose setae. Third, fourth, fifth and sixth thoracic limbs all similar in shape. Fifth limb (fig. 3B), with coxa not differentiated from basis, length of first endopod segment about one third the length of second segment, third segment about two and a half times length of second, fourth slightly shorter than third, claw about one fifth length of fifth segment. Exopod long, basal segment longest, ten distal segments (not shown in figure) each with a pair of long plumose setae. Seventh thoracic limb (fig. 3C) with coxa not differentiated from basis. First endopod segment slightly shorter than second (2), third almost three and a half times length of second, fourth slightly longer than third and with an outer proximal row of six pectinate setae, fifth segment about two thirds length of fourth, claw about half the length of fifth segment. Exopod slightly shorter than those on preceding limbs. Eighth thoracic limb (fig. 3D), with coxa slightly differ- entiated from basis. Coxal segment with inner broad setose penial lobe. First and second endopod segments equal in length, third segment about three and a half times the length of second, fourth much longer than third and with outer proximal row of seven and inner distal row of three pectinate setae, fifth segment shorter than fourth, inner margin with a row of fifteen long pectinate setae, claw almost half length of fifth segment. Exopod with distal segments not clearly differentiated. First to fifth pleopods biramous and invested with plumose setae. Endopod one-segmented and increasing in length progressively from first to fifth pleopods. Exopods of pleopods three-segmented with first segment shortest. First pleopod (Fig. 4A) with second and third segments of exopods equal in length, second segment with a small longitudinal setose ridge on inner ventral surface. Second pleopod (fig. 4B), with second exopod segment longest and broadly expanded, inner margin with a proximal row of graded setae and a distal row of retinaculae; a ventral 204 R. W. INGLE Fic. 3. Spelaeomysis servatus (Fage). Male, 5-5 mm, t.l. Aldabra Atoll. A. B. C. D. Second, fifth, seventh, eighth right thoracic limbs respectively, ventral aspects. Scales in mm. A REDESCRIPTION OT SPELAEOMYSIS SERVATUS (FAGE) 205 a or Sas- Fic. 4. Spelaeomysis servatus (Fage). Aldabra Atoll. Male 5-5 mm,t.l. A. B. C. First, second, third right pleopods respectively. D. Right uropod. Female 6 mm, t.l. E. F. Seventh and eighth thoracic limbs respectively. G. Second right pleopod. All ventral aspects. Scales in mm. 206 R. W. INGLE oblique sinuous ridge arising from ventral surface of second segment invested with long setae. Pleopods three (Fig. 4C) to five similar in shape. Uropods longer than telson, ventral distal margin of sympod prolonged backwards and shaped as depicted in fig. 4D and with a long curved spine. Inner margin with short setae. Exopod with distal transverse suture, endopod one-segmented. Outer exopod margin, proximal to suture, with a graded row of non-plumose setae, rest of exopod margin with plumose setae. Endopod outer and distal margins invested with plumose setae. Female. The female differs from the male in the following features. (1) Second to eighth thoracic limbs each bear an oostegite arising from inner surface of coxa (figs. 4, E,F), and increasing in size posteriorly. (2) Seventh thoracic limb (fig. 42), with first endopod segment much shorter than second, third segment slightly less than three times length of second, fourth slightly shorter than third and with outer proximal row of four prominent pectinate setae, fifth segment about three- quarters length of fourth, claw about one fifth length of fifth segment. (3) Eighth thoracic limb (fig. 4F), with first endopod segment shorter than second, third about two and a half times the length of second, fourth with eight pectinate setae on outer proximal margin and five pectinate setae on inner distal margin, (shorter than those of male), claw slightly less than a quarter length of fifth segment. (4) Second pleopod (fig. 4G), with a four-segmented exopod second segment shorter than either third or fourth, fourth invested with a long stout setae. Juveniles. The smallest specimens (g 3 mm, 9 4 mm), have the proportions of the segments of thoracic limbs similar to those described for the adult male. In par- ticular, the claws of thoracic limbs seven and eight are nearly half the length of segment five of these respective limbs. Compared with the adults, juveniles have about half the number of setae on the fourth segment of thoracic limbs seven and eight, and in juvenile females only half the adult number of setae are present on the fifth segment of limb eight. The telson spines of juveniles are longer. The median spine is more than one third the telson length, whilst in adults it is less than one third of the length. REMARKS. The specimens of Spelaeomysis servatus from Aldabra, the original descriptions of the species by Fage (1924, 1925), and four specimens (3 9? and 1 g) of the type series examined, all agree, except in the following features. (1) The eyestalks of the Aldabran specimens and of the four types are subquadrate, but do not have the inner distal margins acutely sloping as depicted by Fage (1925, fig. II). (2) The dorsal surfaces of segments two and three of the antennal sympod of the Aldabran and type specimens are not invested with setae as shown by Fage in his fig. III. (3) The structure of the male’s second pleopod of the Aldabran material and of the type specimen examined does not agree with the figure of this appendage given by Fage in his fig. 1V. This difference may be due to an abnormal flattening of his material after mounting as a micro-preparation. (4) Fage figures very short telson spines (his fig. VII), the median spines measuring only about one sixth of the telson length. These spines on the Aldabran and the type specimens examined measure about one third of the telson length. A REDESCRIPTION OF SPELAEOMYSIS SERVATUS (FAGE) 207 Discussion. Spelaeomysis servatus was first collected in the surface lake water of a coralline grotto on Zanzibar Island, E. Africa (see Allaud & Jeannel, 1914: 381). The specimens from Aldabra Atoll are from a limestone seepage pool and were also collected near the surface of the water during a flowing tide (personal communi- cation from Dr. K. G. McKenzie). Both habitats are similar and of the type in which members of the hypogean fauna are often found. Although the Aldabran specimens were taken from an exposed pool, it seems unlikely that this is their normal habitat. It is possible that they were carried into this surface pool by the upwelling of water caused by the incoming tide passing through the subterranean limestone interstices. Whether, on Aldabra, this mysid inhabits hypogean water that is cut off from the surrounding sea (and is of a lower salinity), or whether it is a true marine subterranean dweller, remains to be investigated. At present there is little information available about the biotopes of the other species of Spelaeomysis. Gordon (1960 : 308) defined the family Lepidopidae (now Lepidomysidae). This family contains two genera at present, Lepidomysis Clarke, (1961) and Spelaeomysis Caroli, (1924). The genus Spelaeomysis was redefined by Pillai & Mariamma, (1964 : 223) and all the features listed in their definition, except for ‘‘eye-plates without visual elements’, are present in material examined of L. servatus. The presence or absence of visual elements does not constitute a valid generic character, as in the Mysidacea these elements can be developed, reduced or can be absent in different species within the same genus (Pillai & Mariamma, 1964: 119). There is no justification, therefore, for maintaininig two genera in the family Lepidomy- sidae, and the species Lepidomysis servatus is now transferred to the genus Spelaeom- ysis Caroli. This last generic name has precedence over Lepidomysis Clarke, 1961. The family Lepidomysidae and the genus Spelacomysis can now be defined as follows: Body depressed or sub-cylindrical. Carapace produced antero-laterally into two rounded lobes and postero-laterally into large wings. Last two thoracic segments dorsally exposed, penultimate segment with anteriorly directed dorsal scale over- lapping postero-median margin of carapace. Sixth abdominal segment slightly longer than fifth. Telson triangular or linguiform with margins partly or completely spinose. One or two eye-plates with or without visual elements. Labium with lobes well separated. Mandible with well developed incisor and molar processes; palp large and with robust pectinate spines on outer margin of second and third segments. Maxillule with a two-segmented reflexed palp bearing a long distal seta. Maxilla with inner margin of endopod segment expanded. Endopod of first thoracic limb with well developed endites, exopod reduced to a small unsegmented scale, epipod large. Endopod of second thoracic limb stout, exopod long and seg- mented. Thoracic limbs three to eight sub-equal or eighth very long. Coxa of eighth limb of male with penial lobe (?absent in S. bottazzii). Coxae of second to eighth thoracic limbs of female with oostegites. Pleopods alike in both sexes, except for sexual dimorphism of second pair, increasing in size from first to fifth pair, biramous, with one segmented-endopod and three-segmented exopod (four-segmented on second pair in female). Chitinous lamellae on postero-ventral margins of abdomi- nal segments three to five. Uropod with a backward prolongation of sympod, exopod with a lateral suture (very faint in S. bottazzii), endopod without statocyst. 208 R. W. INGLE Fic. 5. A. Spelaeomysis longipes Pillai & Mariamma. Male, 5-6 mm, t.l. paratype, Kottayam, Kerala, India. Anterior portion of carapace, eye-plate and peduncular segments of antenna, dorsal aspect. B. Spelaeomysis quinterensis (Villalobos) male 7-5 mm, t.l. paratype, Gruta de Quintero, Mexico. First left thoracic limb, dorsal aspect. Spelaeomysis bottazzii Caroli, female 10-0 mm, t.l. Terra d’Otranto, Italy. C. Telson, dorsal aspect. D. First left thoracic limb, dorsal aspect. Scales in mm. A REDESCRIPTION OF SPELAEOMYSIS SERVATUS (FAGE) 209 KEY TO DETERMINATION OF THE SPECIES OF THE GENUS SPELAEOM YSIS Carott 1 (2) A pair of eye-plates present (fig. 14). Antennule, basal segment, outer distal margin not produced F . c 4 : 5 c : 3 2 (1) A single eye-plate present (fig. 5A). " Antennule, basal segment, outer distal margin produced as a conical acute process (fig. 54). Spelaeomysis longipes Pillai & Mariamma (Kerala) 3 (4) First thoracic limb (fig. 2E), fifth segment (5) stout, length equal to or slightly exceeding breadth, inner margins of basis (b) and of endopod first segment expanded as broad endites : : , 2 ° : . 0 c 5 4 (3) First thoracic limb (fig. 5B), fifth segment slender, length greater than breadth, inner margins of basis and that of endopod first segment expanded as narrow endites . : Spelaeomysis quinterensis (Villalobos) (Yucatan) 5 (6) Distal two thirds of telson. margin with spines, apex rounded (fig. 1F). Eye- plates with visual elements (fig. 14). First thoracic limb (fig. 2£), fifth segment (5), length equal to breadth, fourth segment inner margin with strongly produced endite. Spelaeomysis servatus (Fage) (Zanzibar, Aldabra) 6 (5) Whole of telson margin with spines, apex sub-acute (fig. 5C). Eye-plates without visual elements. First thoracic limb, fifth segment, length exceeding breadth (fig. 5D), fourth segment with inner margin not strongly produced. Spelaeomysis bottazzii Caroli (Otranto) ACKNOWLEDGEMENTS I must sincerely thank Dr. O. S. Tattersall for reading through the manuscript of this paper and for making available to me a part of Villalobos’ paper translated into English. I am also grateful to Dr. K. G. McKenzie tor providing information on the Aldabran habitat of S. servatus. REFERENCES Attaup, CH. & JEANNEL, R. 1914. In: R. Jeannel & E. G. Racovitza. Enumération des grottes visitées 1911-1913. Avrchs Zool. exp. gén. (Biospeologica XX XIII) 53 : 325-558. Carout, E. 1924. Su di un misidaceo cavernicolo (Spelaeomysis bottazzii n.g., n.sp.,) di Terra d’Otranto. Atti Accad. naz. Lincei Re. 33 : 512-513. CLaRKE, WM. D. 1961. Proposal of anew name, Lepidomysis, for the preoccupied mysidacean generic name Lepidops Zimmer, 1927. Crustaceana 2 : 251-252. Face, L. 1924. Sur un type noveau de Mysidacé des souterraines de l’ile de Zanzibar. C.r. hebd. Seanc. Acad. Sci., Pavis 178 : 2127-2129. 1925. Lepidophthalmus servatus Fage. Type noveau de Mysidacé des eaux souterraines de Zanzibar. Aychs Zool. exp. gén. (Biospeologica LI) 63 : 525-532. Gorpon, I. 1960. On a Stygiomysis from the West Indies, with a note on Spelaeogriphus (Crustacea, Peracarida). Bull. By. Mus. nat. Hist. (Zool.) 6 : 285-324. 210 R. W. INGLE McKenziz, K. G. 1971. Entomostraca of Aldabra, with special reference to the genus Heterocypris (Crustacea, Ostracoda). Phil. Tvans. R. Soc. B 260 : 257-297 Pitrar, N. K. & Mariamma, T. 1964. On a new Lepidomysid from India. Crustaceana 7 : 113-124. VittaLopos, A. 1951. Un nuevo misidaceo des las Grutas de Quintero en el Estado de Tamaulipas. An. Inst. Biol. Univ. Méx. 22 : 191-218. ZimMER, C. 1927. Mysidacea. In: W. Kukenthal & T. Krumbach, Handbuch der Zoologie, 3 (1) : 607-750, (W. de Gruyter, Berlin, Leipzig). Dr. R. W. INGLE Department of Zoology British MusEuM (NATURAL History) CROMWELL Roap Lonpon, SW7 5BD A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) Kay, E. Arison. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 1965. (Out of Print.) £3.75. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180; 11 Plates, 15 Text-figures. 1967. {4. TayLor, J. D., KENNEDy, W. J. & Hatt, A. The Shell Structure and Mineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates, 77 Text-figures. 1969. £4.50. Haynes, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur) 1962-1964. (In press.) id sae Printed in England by Staples Printers Limited at their Kettering, Northants. establishment _ RECENT RECORDS OF MAMMALS (OTHER THAN BATS) FROM ETHIOPIA G. B. CORBET AND D. W. YALDEN BULLETIN OF ISH MUSEUM (NATURAL HISTORY) ame ices 2a No. 8 3 LONDON : 10972 RECENT RECORDS OF MAMMALS (OTHER THAN BATS) FROM ETHIOPIA BY GORDON BARCLAY CORBET AND DEREK WILLIAM YALDEN University of Manchester xo" Muse S Coen HIST- oe 49 JUN 1972 Pp. 211-252; 3 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 22 No. 8 LONDON : 1972 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), «instituted in 1949, ts issued in five series, corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become veady. 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. 22, 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), 1972 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 14 June 1972 Price £1.55 \S PRESEnTey c XS eZ RECENT RECORDS OF MAMMALS (OTHER THAN BATS) FROM ETHIOPIA By G. B. CORBET & D. W. YALDEN CONTENTS Page SYNOPSIS a : < ; C 6 0 0 0 ¢ 213 INTRODUCTION c F 3 5 4 5 : : C 5 213 COLLECTING LOCALITIES The 1964 (Royal Military Academy) Expedition . a . 214 The 1966 (Royal Military Academy) laces 0 0 3 0 216 The 1968 (Great Abbai) Expedition. - a a 216 The Wellcome Parasitology Unit : 6 A 5 , é 222 SYSTEMATIC LIST Insectivora 9 . . 4 . a 5 5 3 0 223 Primates . . 2 : : 0 ¢ é 4 : 4 22 Lagomorpha. 7 6 ¢ 5 d ¢ : a 5 226 Rodentia . c 9 2 A 2 0 : : : 227 Carnivora . ¢ : 4 0 6 6 6 : ; 239 Hyracoidea ¢ 0 A - : : ° 0 : ¢ 241 Tubulidentata . 3 c Q 5 : : 0 0 0 242 Perissodactyla . . F 2 : 6 3 : 242 Artiodactyla. 9 . ‘ a a : A 0 242 DISCUSSION. 0 0 é a a 0 c 0 : 245 ACKNOWLEDGEMENTS 3 c ¢ a ° 0 : é é 249 GAZETTEER 3 D 4 : o 3 : . 3 ; 5 249 REFERENCES . 5 5 : . 9 a : : : 3 250 SYNOPSIS Recent expeditions to Ethiopia collected about 400 mammal specimens for the British Museum (Natural History), and these are identified and discussed. Four species are apparently new to Ethiopia, the shrews Crocidura niobe and Suncus etruscus, the murid Arvicanthis somalicus, and the cane rat Thryonomys gregorianus. In addition, there are several species that have been reported from Ethiopia only rarely, including Tachyoryctes macrocephalus, Tatervillus emini, Lemniscomys striatus, Praomys fumatus and Proteles cristatus. The peculiar zoogeographic relationships of the Ethiopian (i.e. Abyssinian) fauna are discussed in the light of these and previously published records. While an exhaustive examination of the mammalian fauna is not yet possible, it is evident that parallels exist with the better known avian and lepidopteran faunas. INTRODUCTION THE mammals of Ethiopia are relatively poorly known, and there has never been any comprehensive published account. Many East African species were first described from Ethiopian material, e.g. by Cretzschmar (1826) and Heuglin (1877), but these collections came mainly from the coast of the Red Sea. The main collections from Ethiopia in the British Museum are those made by P. Zaphiro in 1905, especially in 214 G. B. CORBET & D. W. YALDEN the southern half of the country, and by Major R. E. Cheesman about 1926 to 1932, mainly in the central highlands, but neither of these were reported upon except to describe individual novelties, e.g. by Thomas (1928). De Beaux (1925, 1943) added further records, and recently Ingersol (1968) has collected the mammals of the Harar area. Blower (1968) has given a more general account of the wildlife of Ethiopia. In 1964 and 1966, adventure training expeditions from the Royal Military Acad- emy, Sandhurst, collected mammals for the British Museum, principally in the region around Lakes Shamo and Abbaya in the Great Rift Valley, south of Addis Ababa (Anon., 1965, 1967). In 1968, the Great Abbai Expedition collected further specimens, particularly along the valley of the Blue Nile (=Great Abbai), and also more widely in central Ethiopia (Anon., 1970; Blashford-Snell, 1970). It is this last collection that forms the main basis for this paper, but relevant information from the other collections is included. D.W.Y. was a member of the Great Abbai Expedition and is responsible for the reports of field observations. G.B.C. identified the collection and supplied the taxonomic notes. The Chiroptera are not included here, as they have been considered separately by Hill and Morris (1971). COLLECTING LOCALITIES Localities in Ethiopia are frequently difficult to trace, sometimes because of changes of name, but also due to the phonetic rendering of native names in different European languages (English, French, German and Italian, at least). The main localities (and alternative names) are therefore given here with geographical co- ordinates, altitude, and the dates of collecting at each. The co-ordinates were taken from the 1 : 100000 scale world maps, series GSGS 4646, sheets NC36, 37 and 38, although in some instances there were apparent discrepancies between these maps and astrofixes taken in 1968. In the absence of a botanist on the expeditions, ecological notes are necessarily vague. Since some of the localities, at least, have been little investigated, it has seemed worthwhile to mention also the sight records of larger mammals. The 1964 (Royal Military Academy) Expedition The 1964 Sandhurst expedition collected about 40 mammals from two areas, one in the region of the Rift Valley Lakes, the other further north near Lake Tana. Soddu, 6° 45'N, 37° 40’E, altitude 1500m. Collecting here extended from 8-23 August 1964, and was carried out in the area south of Soddu toward the north shore of Lake Abbaya—specimens are referred in the taxonomic list to Soddu, in order to distinguish them from material collected in 1966. This area is in the rift valley and is rather dry, with some cultivated patches but much thick thorn scrub, isolated trees, and some rocky outcrops. Little Abbai, 11° 20'N, 37° 00’E, altitude 2000 m. Collecting in the valley of the Little Abbai extended from 27th August to 5th September 1964. This area is on the main Ethiopian plateau, between the towns of Danghila and Bahar Dar, and 215 VSAM 1:6 is paeree y SVIVINVSL we" Vv MAMMALS FROM ETHIOPIA (2350) NVv8n vuss0 7 Iveaav Le 41v34u9 vuesd-, < VvuvavioNna; ‘ Nees s! TN VLOW | \svtiHonya Ground over 1500 m Map of Ethiopia showing principal collecting localities. Fic. 1. is shaded. 216 G. B. CORBED & DD. W. YALDEN south of Lake Tana. With the higher rainfall, rather more of the land is cultivated but there are also patches of scrub and some rocky outcrops. The 1966 (Royal Military Academy) Expedition The 1966 Sandhurst expedition collected about 90 mammals under the direction of Dr P. A. Morris from the Rift Valley Lakes region between 6th August and 3rd September 1966. Lake Abbaya Base camp was established at the S.W. corner of Lake Abbaya (=L. Margherita) near Collufu River at 6° 04’N, 37° 40’E, altitude 1500 m. Arba Minch, the provincial capital of Gemu Gofa, is 10 km west of the base camp site. There are small areas of cultivated land, but the dominant vegetation is thorn scrub. At the time of collecting, the area was extremely dry and dusty until the very end of the period when the rainy season began. Darsi River flows into Lake Abbaya about 12 km north of the base camp. The lake shore region is very swampy, particularly at the mouth of the Darsi, and some collecting was carried out in this region. Some collecting was also carried out in the Bonchie (Bonghe) Valley, 6° 05’N, 37° 20'E, some 35 km west of the base camp, on the lower slopes of Mount Gughe, at about 2700 m. This is an area of riverine forest, with closed canopy, plentiful creepers, and sparse, though lush, ground cover, but with some cultivation. Lake Shamo Lake Shamo (=Chamo, Ciamo; also =L. Ruspoli) lies immediately south of Lake Abbaya in the Rift Valley. Some collecting was carried out in an area known colloquially as the ‘White Grasses’, east of the isthmus between the two lakes, at 5° 58'N, 37° 55’E, at an altitude of 1500 m. This is the area shown on the map by Blower (1968) as the proposed Lake Chand (a misprint for Chamo?) Game Reserve. It is an extensive rolling plain with stony soil and dry, white, grass sharply delimited from the surrounding thorn scrub and hillsides. Sight records here included herds or spot counts of 96 common zebra, Equus burchelli boehmi, 16+ waterbuck, Kobus defassa, 40+ Grant's gazelle, Gazella granti, 104 hartebcest, Alcelaphus buselaphus swaynet and 9+ warthog, Phacochoerus aethiopicus. The herd of A.b. swaynei is believed to be the largest herd known of this race which is on the verge of extinction (Simon, 1968). The 1968 (Great Abbai) Expedition This expedition was in the field during August and September 1968. The main object was a navigation of the Blue Nile (Great Abbai) from near its source at Lake Tana to a point near the border of Ethiopia and the Sudan. Most of the zoological collecting was therefore carried out in the Blue Nile Gorge, but specimens were also obtained near the various base camps established on the plateau, and elsewhere in Ethiopia. MAMMALS FROM ETHIOPIA 217 The zoological team on the Great Abbai Expedition (Dr P. A. Morris, Dr M. J. Largen, Mr H. King and D.W.Y.) were responsible for most of the collecting in- cluding that in the Blue Nile Gorge, at Bahar Dar, Harar, and Awash. Collecting at Ghimbi, and much of that at Debra Markos, was undertaken by other members of the expedition. Most small mammals were collected with various types of break- back trap, of ‘rat’ and ‘mouse’ sizes, while a few were caught by hand or killed with dust shot. Larger mammals were shot at night. Hunting was unsuccessful in the Blue Nile Gorge, primarily because of the difficulty of moving about quietly on difficult terrain in the dark, but perhaps also due to a genuine scarcity of larger mammals. Elsewhere, such hunting was most productive. A general account of the progress and organization of the expedition is given by Blashford-Snell (1970), Snailham (1970) and Anon. (1970). Debra Markos (Debre Markos) The main base for the expedition was established beside the airfield at Debra Markos, the provincial capital of Gojjam, I0° 20’N, 37° 50’E, altitude 2500 m. Limited collecting was carried out here from 4-27 August 1968. This area is part of the main Ethiopian plateau, and is intensively farmed. Mostly the farmland is pasture for cattle and sheep, but there is also some arable farming. Very little native vegetation remains; the native trees have been replaced by introduced Eucalyptus sp., and there are no hedgerows, or even walls, between the fields, which are demarcated instead (if at all) by ditches. The airfield itself carried some low, overgrazed, Acacia scrub. No wild ungulates were seen, but spotted hyaenas, Crocuta crocuta, were common in and around the town. Two jackals, Canis aureus, were seen on the airstrip, and a burrow of aardvark, Orycteropus afer, was found. Among the rodents trapped in the area were Avrvicanthis niloticus, Lophuromys flavopunctatus, Mus mahomet and Otomys typus. A limited amount of collecting was also carried out along the main road which runs north-west from Debra Markos to Bahar Dar through Engiabara and Danghila (11° 15'N, 36° 55’E) and southeast to Shafartak Bridge through Degen (—Dejem, Gunghi) (10° 10’N, 38° 05’E). The whole of this stretch of road runs along the Ethiopian plateau at around 2500 m, through similar pastoral country. Blue Nile Gorge The main collections in the Blue Nile gorge were made during stops on the journey by river from Shafartak road bridge westwards for about 400 km to a point near Sirba. Within this stretch of the river, a series of ‘forward bases’ (F.B.1, F.B.2, ¥.B.3) were established and most collecting was done at these, but some specimens were obtained at other, temporary, stops. Shafartak Bridge (Bridge of Gojjam) 10° 06'N, 38° 17’E, altitude 1150 m. Guder River, junction with the Blue Nile, 9° 50’N, 37° 41’E, altitude 1000 m. This was the location of Forward Base One (F.B.1), 85 km west of Shafartak. Col- lections were made from 8-11 August 1968. The gorge in this area, known col- 218 G. B. CORBET & D. W. YALDEN 7 Zz ° ° i 2 ee x ‘ Fe | 4: < a Ivaay 1¥349 t < | |. fsa = 7) im A > ie] for = WwW am ° eae < Sud WZ co 3 ) ais | o s i = 5 IS Sy s ; a ) , < ; \ 9 j ws w an 3 ee iM oe i aes 8 ITLLITAW ae y i> San er area e ex oO « a < < = 3 yIN100 oO z =) a te = = “aS rN aS canth ° . Ko) rs 2 m z S = ~ Se 2 HS = ao = i Ls = > SS $ 2 oe < a cs a ul ° wo ra) Fic. 2. Map of the Blue Nile showing localities studied during the Great Abbai Expedition. MAMMALS FROM ETHIOPIA 219 loquially as the Black Gorge (Blashford-Snell, 1970), is extremely steep-sided, making progress either along or away from the river very difficult. The gorge is entirely uninhabited in this area, except for a Water Resources Board station about 2 km upstream on the Guder (whose flow it monitors). The vegetation consists of a small strip of riverine forest, only one to a few trees wide, and a dry semi-closed thorn scrub away from the rivers and water courses. At the time of collecting, the vegetation was all extremely green, but aerial reconnaissance during the previous dry season revealed a very difficult picture ; then, there was only a thin strip of green riverine forest along the rivers, and brown vegetation behind, with the ground cover completely desiccated, and the thorn bushes leafless. During the journey downstream from Shafartak to F.B.1, a single leopard, Panthera pardus, was seen beside the Blue Nile, and troops of olive baboons, Papio anubis, were seen, 6 km and 3 km east of the mouth of the Mugher River, 9° 50’N, 37° 55'E. Another troop was reported from the higher ground above F.B.1, and the villagers there also claimed that there was a group of six lions, Panthera leo, but this could not be confirmed. Within the gorge, in the immediate neighbourhood of the Forward Base, no trace of larger ungulates or carnivores were seen. Quills of porcupine, Hystrix cristata, were found in the area, and at the mouth of the Mugher River. The only small mammals caught were Acomys sp. and Dendromus mystacalis. Fincha (=Fingiar) River, junction with the Blue Nile 10° 03'N, 37° 20’E, altitude 1000 m. This was a temporary base, where there was some collecting from 11-13 August 1968. The general terrain and vegetation were the same as F.B.1. In moving from F.B.1 to the Fincha River on 11 August, two klipspringer, Oreotragus oreotragus, were seen on the cliffs, and tracks of Papio anubis were found on the river bank. A single Colobus guereza was seen in the riverside trees about 1 km west of F.B.1, and two warthog, Phacochoerus aethiopicus, were seen en route. At the mouth of the Fincha, quills of Hystrix cristata were again found, and at least two hyraxes, Procavia habessinica were seen. The only small mammal trapped here was the sole dormouse, Graphiurus murinus, obtained by the expedition. Mabil. Forward Base Two was established on the bank of the Blue Nile at 10° 19/N, 36° 45’E, about 8 km S.W. of Mabil village, and ro km W. of the Dolnik River at 900 m. Collecting here occupied 14-20 August 1968. Travelling west from the Fincha to F.B.2, on the r4th August, the first 10 km of the journey continued through steep-sided gorge, and a further two klipspringer were seen. These were the last ones seen along the river, however, for further west- wards the valley opens out. The vegetation remains essentially the same, with just a thin strip of tall trees and a deciduous thorn scrub hinterland. The valley remains uninhabited, but small patches of cultivated land (mostly with maize) suggest temporary, perhaps seasonal farming. With the river flowing more slowly, hippo- potamus, Hippopotamus amphibius, were seen for the first time, two east and two west of the Wamet (Uamet) ford at 10° 09’N, 37° 12’E, and at least nine small troops of Papio anubis. Another single Colobus was noted 1 km west of Wamet. Sight records at F.B.2 included single specimens of the mongoose, Ichneumia albicauda and a male bushbuck, Tragelaphus scriptus, while most of the small mam- mals caught were Mus sp., probably Mus pasha and Mus tenellus. 220 G. B. CORBET & D. W. YALDEN Azir River. A temporary base was established from 20-22 August 1968 near the Azir River at 10° 29'N, 36° 25’E, altitude 800 m. The Blue Nile from F.B.2 to the Azir River continues through the rather wider valley. A party of Colobus was seen at the mouth of the Dim River (10° 30’N, 36° 26’E) and there was another party in the vicinity of the temporary base. There were also small groups of vervet monkeys, Cercopithecus aethiops, at both these localities. A single warthog was seen at the base, and tracks were numerous, as were those of hippopotamus. - A number of mice, Praomys natalensis, were trapped here. Caroarsa River. A temporary base was established 10 km east of the Caroarsa River, at 10° 07’N, 36° 12’E, altitude 600 m. Collections date from 23-24 August. From the Azir River to some distance west of the Caroarsa, the sides of the Blue Nile Gorge close in to give a steep-sided rocky valley again, known colloquially as the Western Cataracts. However, the river is not so consistently turbulent as in the Black Gorge and a dozen hippopotamus were counted during the journey west to the Caroarsa on 22 August. There was a small party of Cercopithecus aethiops in the vicinity of the temporary base, and at least one, perhaps two, Colobus. Traces of porcupine and hyrax (?Procavia habessinica) were found, while there were tracks of leopard on the river bank. Ungulate tracks, possibly bushbuck, were also noted. Only three rodents were trapped, single specimens each of Lemniscomys striatus, Mus tenellus and Acomys ?dimidiatus. The vegetation at this locality was more lush than elsewhere, a deeper band of riverine forest stretching back for at least too m from both banks of the river. As indicative of the added moistness in the area, it was the only site along the Blue Nile from Shafartak westwards where mosquitoes were troublesome. Didessa River. Forward Base Three was established on an island at the mouth of the Didessa River, 10° 05/N, 35° 38’E, altitude 450 m. Collecting extended from 26-29 August. The island and a thin strip along the river banks constituted a riverine forest, but up the valley sides this gave way as elsewhere to thin scrub. There was a flattish damp area with tall (over 2 m) grass penetrated by the trackways of hippopotamus, and inhabited also by what appeared to be cane rats, Thryonomys sp. From the Caroarsa westwards the Blue Nile runs in a widening valley and flows more gently. The first permanent habitations were noted in the Western Cataracts, and the valley became more densely populated, and more intensively cultivated, from there westwards. The nearest village to F.B.3 was about 5 km west, and there were maize fields on the opposite river bank. Small numbers of Papio anubis, Colobus and hippopotamus were recorded during the journey to the Didessa, and one or two hippopotamus were heard during the nights in the rivers beside F.B.3.. A number of Praomys natalensis were caught in the native village nearby, while trapping in the river bank vegetation yielded several Lemniscomys striatus, Crocidura ?sericea, and Acomvys sp. Sirba, 10° 05'N, 35° 30’E, marked the western end of the journey downstream. No terrestrial mammals were collected there but four oribi, Ourebia ourebi, were seen on the mission airstrip at night on 30th August. The banks of the Blue Nile between the Didessa and Sirba are intensively cultivated, lacking anything more than the odd MAMMALS FROM ETHIOPIA 221 remnant tree from the riverine forest, and no large mammals were seen during the last journey down river. Ghimbi, Wollega Province Ghimbi, 9° 10'N, 35° 50’E, altitude 2150 m, was a collecting area from 22-31 August 1968. Ghimbi is on the densely populated and intensively cultivated plateau, on the main road west from Addis Ababa. Some collecting was carried out in the town, and in the surrounding maize fields; other specimens came from rough bushland along the main road up to 30 km out of the town. Two specimens came from an area of savannah 24 km north of Ghimbi, at 9° 30'N, 35° 50’E, altitude 1600 m. The main rodents trapped here were Rattus vattus and Lophuromys flavo- punctatus, while among carnivores shot here were Felis serval, I chneumia albicauda and Viverra civetta. In addition specimens of Orycteropus afer, Crocuta crocuta, Colobus guereza, Canis ?aureus and Phacochoerus aethiopicus were reported seen in the area. Bahar Dar, Gojjam Province Bahar Dar (Bahr Dar, Bahadar) is located on the main road north from Addis Ababa to Gondar, at the point where it bridges the Blue Nile near the source of the river, I1° 35’N, 37° 25’E, altitude 1830 m. Although on the plateau this area is not so intensively farmed as the environs of Ghimbi and Debra Markos; more- over, there is little arable farming, perhaps because the ground is waterlogged in the wet season, or because the soil is shallow, and instead pastoral farming pre- dominates. Numerous thickets and patches of thorn scrub remain, there is a thin strip of riverine forest along the Nile, and native tree species have not been replaced by the almost ubiquitous Eucalyptus. Collecting around Bahar Dar occupied 11-14 September 1968; it was concentrated in an area 15 km south-east of the town, along the road that follows the south bank of the Nile to the Tississiat Falls. Species seen in this area but not collected include Cercopithecus aethiops, Papio anubis, Felis serval, Genetta sp. and Tragelaphus scriptus. The principal rodent trapped was Lophuromys flavopunctatus, and specimens of the rodents Praomys albipes, P. natalensis, Dasymys incomptus and three species of crocidurine shrew were obtained. Harar, Harar Province Harar itself is on the high ridge that runs eastwards from the eastern wall of the rift valley, but collecting was carried out on the lower ground further east, along the road to Jiggigga. This area is rather sparsely populated, and is mostly very thin thorn scrub with little or no ground cover; some parts are very dry and rocky. Bisidima River, 9° 15’N, 42° 12’E, altitude 1500 m. This locality was collected 20-21 September. There is some cultivation of maize, and collecting areas included the Opuntia hedges around these fields and rough thorn scrub. Trapping yielded specimens of Rattus rattus, Praomys natalensis, Lophuromys flavopunctatus and Crocidura doriana. 222 G. B. CORBET & D. W. YALDEN Valley of the Rocks, 9° 15’N, 42° 20’E, altitude 1300 m. An area of sparse Acacia woodland, with many boulders, cliffs and scree slopes, collected 21 September. Two klipspringer, Oreotragus oreotragus, were seen here, and Procavia was collected. 60 km East of Harar, 9° 14'N, 42° 32’E, altitude 1200 m. This is a sparsely inhabited area of thin Acacia scrub and very little ground cover, which was studied 21-22 September. Among mammals seen in this area were Felis silvestris, Canis ?aureus, Ichneumia albicauda, and a large troop of sacred baboons, Papio hamadryas. The only small mammals caught here were a gerbil Taterillus 2?emini and a rat Praomys fumatus. Awash, Shoa Province Awash (=Auasc, Aouche) is well known as a National Park. It lies in the Great Rift Valley in an extension northwards of the more familiar East African Acacia savanna, mostly grass with scattered trees and bushes. There is a strip of riverine forest along the Awash River. The area is still moderately active volcanically, with recent lava flows and lava bubble caves, and is dominated by the quiet cone of Mount Fantalle which rises to 2000 m. The lava bubble caves are of particular importance in the present context. Some of the lava bubbles have opened at the sides, to give normal caves which were used by bats and porcupines; others have collapsed in the top to form pitfall traps, and two of these, at 8° 56’N, 39° 57’E, yielded more than a hundred skulls of various animals. Most collecting was carried out on the north bank of the Awash River at 8° 50’N, 40° o1’E, and a few specimens came from the area of the hot springs at Filhoa, 9° 00’N, 39° 58’E. Collecting lasted from 23-28 September 1968. Naturally, in such a rich area for game sight records of many species were obtained. The principal ungulates were oryx, Oryx gazella beisa, and Soemmerring’s gazelle, Gazella soemmerringt. Smaller numbers of waterbuck, Kobus defassa, lesser kudu, Tragelaphus imberbis, greater kudu, T. strepsiceros, and warthog, Phacochoerus aethiopicus, were seen, together with three gerenuk, Litocranius walleri. An aard- vark was killed on the road in the Park; two leopards and two lions were seen, and a single hippopotamus occasionally wandered through the camp at night. A small troop of Papio anubis was seen frequenting the riverine forest along the Awash River, and small numbers of P. hamadryas were seen in the drier hinterland of the Park. A single zorilla, Ictonyx striatus, seen at night was the first record of that species in the locality. The principal small mammals caught were the gerbils Taterillus Pemint, Gerbillus ?harwoodi and Tatera robusta, spiny mice, Acomys dimidiatus mullah, and the shrew Crocidura ?sericea. Wellcome Parasitology Unit In addition to these collections, a number of specimens recently collected by the Wellcome Parasitology Unit based in Addis Ababa have been included. These came from the following localities. Dessie (Dese), Welu Province, 10° 08’N, 39° 43’E, altitude 2700 m. MAMMALS FROM ETHIOPIA 223 Sabeta, Shoa Province, 8° 55’N, 38° 40’E, altitude 2500 m. Addis Alem, Shoa Province, 9° 03'N, 38° 25’E, altitude 2400 m. SYSTEMATIC LIST The taxonomy of most groups of the smaller mammals of Africa is in a very pro- visional state, especially at the level of delimiting the species. These problems can be solved only by comprehensive revision of each genus on a continental scale, which is out of place in a regional list such as this. The following list is based primarily upon the collection made by the Great Abbai Expedition in 1968, but it also includes mention of all specimens collected by the expeditions from the Royal Military Academy in 1964 and 1966 and some other Ethiopian specimens recently received at the British Museum. Identification has been attempted at the subspecific level only where the subspecies seem clearly defined or, more often, where the specimens can be confidently allocated to a form named from Ethiopia whose specific relationship is doubtful. Order INSECTIVORA Family MACROSCELIDIDAE—FElephant shrews Elephantulus rufescens (Peters) Lake Abbaya: one collected at Cullufu River, between Lake Abbaya and Lake Chamo. This represents the north-western extremity of the range of E. rufescens, which is otherwise known in Ethiopia only from the southern border (Mega, Farda Robo, Murr) and from the Harar district, and extends south in the drier, short-grass steppes to Tanzania (Corbet & Hanks, 1968). This specimen differs from all other forms of E. rufescens in being very grey, with the mid-dorsal line very dark and the post- ocular spots and upper surface of tail almost black. Family SORICIDAE—Shrews Suncus etruscus (Savi) Bahar Dar: one caught by hand under a stone in wet grassland. This appears to be the first record of this minute shrew (or any pygmy Suncus) from Ethiopia. It is an adult male with the following dimensions: head and body 49 mm; tail 29 mm; hind feet 7 mm; condylo-incisive length 12-6 mm (13-2-13-7 mm in sample of 13 other S. etruscus) ; upper tooth row 5-3 mm (5:3—5-8 mm in 20 others) ; width across M2—M2 3-4 mm (3-6-4:0 mm in 22 others). It resembles the Mediter- ranean S. etruscus, and differs from the East African S. infinitesimus (Heller), in having the braincase exceedingly flat, with a depth of only 2-4 mm at the basi- sphenoid. The only other records of S. etvwscus from south of the Sahara appear to be one from northern Nigeria in the British Museum collection, reported by Morrison-Scott (1946), and one from Guinea (Heim de Balsac & Lamotte, 1957). The Nigerian 224 G. B. CORBET & D. W. YALDEN skull also has a short tooth-row (5:3 mm), but the rostrum is wider (M2—M2 3:8 mm) and the length of the skull is not determinable. The Ethiopian and Nigerian speci- mens differ from most available specimens from the main part of the species’ range around the Mediterranean in the large size of the fourth unicuspid teeth and the lack of a concavity on the anterior margin of the large upper premolar, but both have the very flat skull of S. etruscus, and there seem to be no grounds tor the view of Petter & Chippaux (1962) that the Nigerian one should be referred to S. infinitesimus. Crocidura doriana Dobson Bahar Dar: two trapped in thicket in wet grassland. Debra Markos: two trapped in grazed Acacia scrub. Harar: one from Bisidima River. Awash: several skulls from the lava caves. The two from Debra Markos are very large, exceeding any other specimens from Ethiopia available in the collection of the British Museum: upper tooth-rows 14:7 and 148 mm, condylobasal lengths 32:2 and 32-4 mm. The maxima for other Ethiopian skulls are 14-2 mm (n = 17) and 31-8 mm (n = 11) respectively, and Osgood (1936) gave 13-8 mm as the maximum length of upper tooth-row in 28 skulls from Ethiopia. It is probable that this form should be treated as a race of C. occzdentalis (Pucheron) or, with occidentalis, as a race of C. flavescens (I. Geoffroy). Crocidura cf. sericea Sundevall Awash: eight collected, possibly of this species; a further 42 damaged skulls were collected from the lava caves. Blue Nile Gorge: two from the mouth of the Didessa River. These are medium-sized shrews with a pale grey ventral pelage. Measurements of upper tooth-row of all the Awash skulls, totalling 49, give a unimodal distribution with a mean length of 9:02 mm (S.D. + 0:37, range 8-o-10-1). These all show good agreement with specimens from Sudan that have been named C. sericea, and with the types of C. hindei Thomas (Kenya) and C. h. marrensis Thomas & Hinton (Jebel Marra, Sudan). Setzer (1956) allocated larger specimens from central Sudan to C. s. sericea and treated the smaller marrensis and C. lutrella Heller (from Lado Enclave) as races. The Ethiopian specimens agree more closely in size with lutrella and marrensis than with the specimens allocated by Setzer to C. s. sericea which had upper tooth-rows of 10-3 and 10-6 mm. Crocidura niobe Thomas Ilubabor Province: one collected by Drs Largen, Morris and Yalden east of Abiu (west of Gore), at 1500 m in forest in January 1971. This species of shrew was apparently known only from the east side of Ruwenzori, Uganda between 2000 and 2500 m (Thomas, 1906), although it is possible that other MAMMALS FROM ETHIOPIA 225 forms from the Congo and from West Africa may be conspecific. This locality is on the south-western edge of the Ethiopian plateau. It is very dark greyish brown above and below, and the feet and tail are also dark. The tail is extremely slender, about 80 °%, of the length of head and body, and has very sparse vibrissae on the proximal half only. Measurements are: head and body 70; tail 55; hind feet (without claws) 12; ear 9; condylobasal length 19-5; upper tooth- row 8-9. Crocidura bicolor Bocage Little Abbai: one collected. Bahar Dar: one caught under a stone in wet grassland with thickets. This is a small shrew with very dark greyish brown dorsal pelage and moderately dark grey ventral pelage. There is a further specimen in the British Museum from ‘I40 miles S. of Addis Ababa’ and Osgood (1936) recorded specimens (as C. b. nana Dobson) from Hadama on the Awash River and from Addis Ababa. Order PRIMATES Family LORISIDAE Galago senegalensis E. Geoffroy—-Lesser bush-baby Ghimbi: one collected 5 km east of the town on the edge of cultivation. Lake Chamo: two collected on north side of lake. Family CERCOPITHECIDAE Cercopithecus aethiops (L.)—Vervet monkey Soddu: three collected. Little Abbai: one collected. Lake Abbaya: eight collected. Blue Nile Gorge: small parties of six to eight seen at the Azir, Dim and Caroarsa river mouths, in or near riverine forest. Bahar Dar: a party of about six seen in riverine forest. Colobus guereza Riippell—Black and white colobus Soddu: one collected. Little Abbai: one collected. Blue Nile Gorge: fairly widespread, with sightings from practically the whole length of the river from the Guder westwards to the Didessa, involving about 20 animals altogether. Papio anubis (Fischer)—Olive baboon Lake Abbaya: one collected near Soddu, and two from Arba Minch. 226 G. B. CORBET & D. W. YALDEN Blue Nile Gorge: seen along the whole length of the river, except perhaps in the Black Gorge, in groups of up to 20. Bahar Dar: a troop of nine seen on two occasions beside the road to Tississiat. Awash: a small troop seen in the gallery forest along the Awash River, and reported to raid the camp site occasionally. Papio hamadryas (L.)—Sacred baboon Harar: a troop of about 120 seen crossing the Jiggigga road about 60 km east of Harar. Awash: a small troop on the lower northern slopes of Mount Fantalle. The ecological and ethological relationship between P. hamadryas and P. anubis in the Awash area has been studied by Kummer et al. (1970) who reported a zone of hybridization about 15-20 km wide. This, however, is narrow in relation to the mobility of these animals and suggests a considerable degree of reproductive isola- tion that justifies the retention of species rank for these two forms. There is an ecological separation of the two species, with P. hamadryas in the drier regions away from the river. Kummer (1968) has suggested that the juxtaposition of the two species is fairly recent, due to an eastward extension of the range of P. anubis. Theropithecus gelada (Riippell)—Gelada baboon Debra Libanos, Shoa: the small colony here is a well known tourist attraction, and lives on cliff-tops not far from the main road north from Addis Ababa. These animals were seen on several occasions, but were the only ones observed. Order LAGOMORPHA Family LEPORIDAE Lepus habessinicus Hemprich & Ehrenberg—Abyssinian hare Harar: @ collected at Bisidima River; gand collected 60 km east of Harar on Jiggigga Road. Awash: gand 9 collected. Lake Abbaya: 9 collected near Soddu northwest of lake. These specimens agree with L. habessinicus as recognized by Petter (1963), rather than his L. capensis starcki, in having only narrow margins of black outside the tips of the ears and in having deep, cement-filled grooves in the upper incisors. This species is closely related to L. capensis, if indeed it is distinct at all, and it differs from L. whytei (below) in the very soft pelage, longer ears, black and white tail and in having the anterior faces of the upper incisors in the same (transverse) plane. Lepus ?whytei fagani Thomas—Bush hare Bahar Dar: four collected in wet grassland with scattered thickets (two of them pregnant with one and two foetuses). Ghimbi: three collected in bush country close to cultivation. MAMMALS FROM ETHIOPIA 227 These specimens agree closely with the type of fagani (from Lake Tana). This form was at first allocated to ‘L. crawshayi’ by Petter (1959) but later to L. habes- sinicus (Petter, 1963, 1967). The type of fagani seems clearly referable to the East African group commonly called L. crawshayi rather than to the capensis group to which habessinicus belongs; it and the specimens in the present collection have harsher pelage, brown on the sides of the tail, short ears, and upper incisors with oblique anterior faces. However, the type of crawshayi de Winton (from central Kenya) seems to belong to the capensis group, and the earliest name for what Petter called L. crawshayi is probably L. whytei Thomas, 1894 from Malawi. Throughout East Africa L. whyter seems to be the short-eared, harsh-furred bush hare, whilst L. capensis is the long- eared, soft-furred species of the more open plains. All the present specimens of L. habessinicus are from the lower, drier areas in the south and east while the L. ?whytei all came from the plateau. Order RODENTIA Family SCIURIDAE—Squirrels Heliosciurus gambianus multicolor (Riippell)—-Sun squirrel Blue Nile Gorge: one collected at Mabil. This squirrel seems identical to a large series in the British Museum from the upper part of the Blue Nile just below Lake Tana, whence it has also been reported by de Beaux (1925). These are lighter than two from Kaffa, presumably referable to H. g. kaffensis Neumann. The species does not appear to have been recorded on the Sudanese part of the Blue Nile. Xerus erythropus leucoumbrinus (Riippell)—Striped ground squirrel Lake Abbaya: two from Soddu; one from Arba Minch. Lake Chamo: one from northwest of the lake. These localities are probably close to the northeastern limit of this species, the unstriped X. rutilus alone being found in the more eastern parts of the country. Xerus rutilus (Cretzschmar)—Unstriped ground squirrel Harar: Valley of the Rocks, 40 km east of Harar. At least three unstriped ground squirrels were seen in a rocky area of open Acacia woodland. They are presumably referable to this species, which was recorded from this area by Ingersol (1968). Family CRICETIDAE Subfamily GERBILLINAE—Gerbils Gerbillus (Dipodillus) ?harwoodi Thomas Awash: seven trapped in various parts of the region ; a number of cranial fragments, probably of this same species, were found in each of the lava caves examined. 228 G. B. CORBET & D. W. YALDEN These small gerbils show good agreement with the type of harwoodi (from Naivasha, Kenya), but this must be treated as provisional pending a much-needed revision of this group. Tatera robusta (Cretzschmar) Awash: one trapped on north bank of river; a large number of skulls, probably of this species, from the two lava caves. The greyish dorsal pelage of the only available skin gives this gerbil a closer resem- blance to Sudanese 7. 7. vobusta than to the more yellowish Somalian forms such as shoana and phillipsi. Setzer (1956) used the subspecific name taylori Hatt (Red Sea coast) for this species in eastern Sudan, but the skin from Awash differs from taylor and resembles vobusta in having the dorsal hair of the tail black rather than brown. Tatera valida soror Allen Blue Nile Gorge: one from 10 km west of Mabil; one from the mouth of the Didessa River; mandibles, probably of the same species, from a cave at the mouth of the Guder River. These agree well with the description of sovor, from Fazogli on the Sudanese Blue Nile, which was allocated to T. valida by Davies (1968), although this must be considered tentative. Taterillus ?emini (Thomas) Harar: one from 60 km east of Harar on the Jiggigga road. Awash: two from the north bank of the Awash River. Both of these gerbils were in very dry, open Acacia scrub, the one at Harar close to maize cultivation, those at Awash close to riverside woodland. There are no previous examples of this genus from Ethiopia in the British Museum except for the much smaller T. harringtoni from the Kenya border near Lake Rudolph. The specimens from Harar and Awash are richly coloured, with almost no grey showing in the pelage. They resemble T. emini from Sudan and Uganda in size, but differ in the posterior palatal vacuities which are unusually short for any Taterillus. Measurements are as follows (Harar followed by Awash): head and body 117, 119; tail (from pelvis) 127, 167; hind foot (without claw) 31, 32; ear 19, 16; condylobasal length 31-1, 33:7; upper molar row 4-9, 4-8; posterior palatal vacuities 3-0. 3:1. Family MURIDAE Genus ACOMYS—-Spiny mice Acomys ?cahirinus (Desmarest) Blue Nile Gorge: two from mouth of Guder River, trapped on a rocky hillside in thick bush. Lake Abbaya: one from N.E. of Lake Chamo. See remarks under A. ?dimidiatus below. MAMMALS FROM ETHIOPIA 229 Acomys ?dimidiatus (Cretzschmar) Blue Nile Gorge: one 10 km east of Caroarsa River, in riverine forest; three at mouth of Didessa River, in riverine forest and long grass. Awash: four trapped in dry open Acacia scrub close to riverine woodland; remains of two animals from one of the lava caves. The taxonomy of the spiny mice of the genus Acomys in this region is very pro- visional. The Awash animals agree very closely with the type of mullah Thomas from Harar, which Setzer (1968) referred to A. dimidiatus. These have large skulls, the only one with moderately worn teeth having a condylobasal length of 28-2 mm and with upper tooth-rows of 4:5, 4:5 and 4-6 mm. The Acomys from the Blue Nile appear to fall into two groups. Two from the mouth of the Guder River are smaller, with upper molar rows of 4-1 and 4-3 mm and condylobasal lengths of 25-8 and 25:5 mm (both are adults with well worn teeth). The remaining four (Caroarsa and Didessa Rivers) have upper molar rows of 4:6, 48, 4:8 and 4-9 mm and the two with worn teeth have condylobasal lengths of 27-7 and 28-¢4mm. Skins of these two groups are not distinguishable. The difference in size strongly suggests the presence of two species, but in the absence of both at the same locality and in more adequate numbers this cannot be considered certain. The smaller one is tentatively referred to A. cahivinus and agrees well, in size and in colour, with A.c. cineraceus Fitzinger & Heuglin from eastern Sudan to which Setzer (1956) referred animals from the Sudanese Blue Nile. If Setzer (1968) was correct in allocating mullah from Harar to a larger species, A. dimidiatus, widely sympatric with A. cahirinus, then the four larger animals from the Blue Nile may represent A. dimidiatus, but this must be considered very provisional until these two species can be more adequately delimited. The single animal from Lake Chamo has a very greyish pelage and a small skull (upper molar row 4-3 mm, condylobasal length 25-0 mm) and is likely to represent A. cahirinus. Genus ARVICANTHIS—Grass rats Arvicanthis niloticus abyssinicus (Rippell) Debra Markos: four trapped in low, grazed Acacia scrub. The single female was pregnant (four embryos). Dessie: ten from 18 km N.W. of Dessie, trapped on a rocky bank beside a marsh at 2700 m (from Wellcome Parasitology Unit no. 2). Addis Alem: one collected by Dr Aklelu Lemma. The animals from Debra Markos could be referred to fluvicinctus Osgood, but this is unlikely to be separable from abyssinicus. It was based entirely on the longer upper molar row of animals from west of the Blue Nile and the present series from Debra Markos and Dessie, on either side of the Blue Nile, show considerable overlap in this character. The specimens from Debra Markos are, however, distinguished from the Dessie series by an overall suffusion of yellowish brown whereas the Dessie animals are almost pure grey. 230 G. B. CORBET & D. W. YALDEN These are all dark-bellied animals and most have a distinct narrow mid-dorsal stripe. Other Ethiopian forms that are probably conspecific are blicki Frick (S. Chilalo Mts), and satuvatus Dollman (Guma, Didessa River). Arvicanthis lacernatus (Rippell) Harar: one from the Bisidima River, trapped in grazed thorn scrub. Dessie: seven from 31 km N.W. of Dessie, trapped in a dry river valley at 2000 m (from Wellcome Parasitology Unit no. 2). Lake Abbaya: seven from Arba Minch, S.W. of lake. Awash: parts of ten skulls of a large Avvicanthis, probably this species, from one of the lava caves. One skin and skull, of an animal trapped in the National Park, received on loan from the park museum. There seems to be good reason to recognize two species of large Avvicanthis in Ethiopia and in adjacent regions of Sudan, Uganda and Kenya, although much remains to be done to assess their variability. The present species has the under- parts grey, rather sharply demarcated from the brown of the flanks. The grey of the belly may be quite dark, as in the series from the Dessie region, or may be suf- fused with brown in the midline as in the animal from Harar, but it is never speckled like the dorsal pelage as it isin most A. niloticus. A. lacernatus lacks a clearly defined dorsal stripe, and there is a tendency for the rump and hind legs to be unspeckled and tinged with orange brown. These characteristics are seen in fairly extreme form in the specimen from Harar; in the series from Lake Abbaya the brown on the rump is less distinct whilst in the series from Dessie it is absent altogether and the ventral pelage is a uniform dark grey. The brown colour on the rump in this species is very evenly distributed and seems quite distinct from the sporadic erythrism that seems to be a characteristic of A. niloticus. The specimen of A. niloticus from Addis Alem, for example, has on the belly a large irregular patch of hair with the basal parts chocolate brown instead of the usual slaty grey, and the tips yellowish brown instead of pale cream. The specimen from Harar agrees very closely with the type of pellicews Thomas (from Lake Tana) which Osgood (1936) maintained was a synonym of lacernatus. Other Ethiopian forms that are probably conspecific are zaphivi Dollman (Guma, Didessa River), mearnsi Frick (Awash) and perhaps raffertyi Frick (Gardula). It is possible that another synonym this species is Isomys testicularis Sundevall, 1843 from Bahr-el-Abiad, Sudan. There is no evidence amongst the present collection of the two species being strictly sympatric in Ethiopia, but de Beaux (1930) recorded both from a single locality in Eritrea, and Heller (1g11) described both species from Rhino Camp, N.W. Uganda (as A. jebelae and A. rubescens). Arvicanthis somalicus Thomas Awash: parts of 69 skulls from the lava caves; one animal shot by day in dry Acacia scrub, but only the skull preserved; six animals subsequently trapped by MAMMALS FROM ETHIOPIA 231 D.W.Y. in December 1970 at Metahara, in heavily grazed grassland with patches of Acacia scrub. This species, described from northern Somalia, does not appear to have been recorded hitherto in Ethiopia. Allen (1939) listed it as a distinct species, with chanleri and reptans from northern Kenya as races, but Misonne (1968) recognized only one species of Arvicanthis, presumably considering somalicus to be no more than subspecifically distinct from A. nzloticus. The skulls from the Awash caves were the first indication of two sympatric species differing in size (fig. 3), and agreeing respectively with specimens of A. somalicus A.n. abyssinicus: Ethiopia a lL A. lacernatus: Ethiopia EEE er Awash caves, Ethiopia white-Awash A. somalicus black-Somalia eS 5:0 5-4 5:8 6:2 6-6 7:0 7-4 Fic. 3. Length of upper molar row (at maximum of crown) of Arvicanthis spp. 232 G. B. CORBET & D. W. YALDEN from Somalia and of A. lacernatus from elsewhere in Ethiopia. This was confirmed by the trapping of six specimens of A. somalicus nearby at Metahara in 1970. These specimens agree very closely with A. somalicus from Somalia, including the type, except that the pelage is less yellow. The difference in size, especially of the tooth-row, is sufficiently clear to leave no doubt that there is sympatry of two species at Awash. (Besides the large skulls in the cave sample, we have subsequently seen a specimen of A. lacernatus trapped at Awash, which had a tooth-row of 6:3 mm.) A. somalicus can therefore be considered specifically distinct from A. niloticus and A. lacernatus, and this record extends its known range by about 400 km westwards from the nearest localities in northwestern Somalia represented by specimens in the British Museum (around Hargeisa). The types of chanleri Dollman and reptans Dollman, both from the Northern Uasso Nyiro district of Kenya, also fall well within the range of these Ethiopian specimens, with molar rows of 5-6 and 5:5 mm respectively, but considerably more work needs to be done to establish to what extent these small forms are sympatric with larger ones in Kenya. Externally A. somalicus and A. lacernatus are very similar at Awash. The best distinguishing characters are probably the greater amount of yellow in the dorsal pelage of A. lacernatus, especially marked on the rump, and the longer hind feet (22-23 mm in A. somalicus; about 30 mm in A. lacernatus). The five available skins of A. somalicus from Awash are all of rather young animals, with the teeth scarcely worn, and therefore the other external measurements (head and body 93-106 mm; tail 88-105 mm) are probably below adult size. The ventral pelage of A. somalicus resembles that of A. lacernatus in being pale grey, rather sharply demarcated from the brown of the flanks. Dasymys incomtus (Sundevall)—Shaggy rat Bahar Dar: one trapped in wet grassland with scattered thickets. Bahar Dar is close to the type locality of D. 7. griseifrons Osgood, but this specimen does not show the grey forehead and face described for griseifrons. The only other examples of this species from Ethiopia in the British Museum are single specimens from Nono, upper Omo and Wodessa River, Guma. Delany & Neal (1963) also reported this species from damp habitats in Uganda (‘swamps, reed beds and river valleys’) and mentioned that it occurs above 4000 m on Ruwenzori. Lemniscomys striatus (L.)—Striped mouse Blue Nile Gorge: one 10 km east of Caroarsa River; four at mouth of Didessa River. These were all trapped in long grass on the bank of the river. MAMMALS FROM ETHIOPIA 233 The only previous records of this species in Ethiopia appear to be from Nono, 150 km south of the Blue Nile (the type of wroughtoni Thomas), from the Urgessa River (an upper tributary of the Didessa River) and from Delbena River, Konso (?near Mega in the extreme south). The two skins from the Blue Nile have the nor- mal yellowish brown ground colour with only the mid-dorsal stripe dark, closely resembling L.s. massaicus (Pagenstecher) of Kenya and the skin from Konso, and quite different from the adjacent L.s. wroughtoni which has all the dark stripes virtually black. The type and only available specimen of wroughtont came from 3600 m; the single specimen from the Urgessa River, at 2300 m, is considerably less dark, in fact intermediate between wroughtoni and the normal specimens from the Blue Nile at the mouth of Didessa River at about 800 m. The Blue Nile animals resemble the East African L.s. massaicus and differ from West African forms, e.g. L.s. striatus, in having the lateral white stripes very bold and continuous. They differ from most L.s. massaicus in having the second white stripes (away from the mid-dorsal line) very well marked whereas these stripes are very fragmentary in massaicus. Lophuromys flavopunctatus Thomas—Harsh-furred rat Bahar Dar: seven trapped in thickets in wet grassland. Debra Markos: one trapped in low, grazed Acacia scrub. Ghimbi: two adults and three nestlings. Harar: two trapped amongst thorn scrub at Bisidima River. Dessie: two from 18 km N.W. of Dessie at 2700 m, trapped on the edge of a marsh (from the Wellcome Parasitology Unit no. 2). Addis Ababa: one from the British Embassy compound (from Wellcome Parasit- ology Unit no. 2); one from Meta Abo, Sabata, c. 20 km S.W. of Addis Ababa. The specimens from Harar appear to differ from the series from the highlands only in the lighter and brighter orange-brown colour of the proximal zone of the dorsal hairs. These can probably all be attributed to L.f. favopunctatus. The single skin from Addis Ababa differs from all the others in being more coarsely speckled above due to longer subterminal light bands on the hairs. The locality at Harar, although close to the River Bisidima, is an unusually dry habitat for this species which is more characteristic of montane forest or moist scrub. One female from Harar and that from Debra Markos were pregnant, each with five embryos. Genus MUS—Pygmy mice Mus mahomet Rhoads Debra Markos: two trapped in small thicket surrounded by grazed grassland. Dessie: one from 18 km N.W. of Dessie at 2700 m on a steep rocky hillside grazed by goats; two from 31 km N.W. of Dessie at 2000 m (all from the Wellcome Para- sitology Unit). 234 G. B. CORBET & D. W. YALDEN This is a rather distinctive species, apparently confined to Ethiopia and the adjacent part of Somalia. It is a fairly large grey-bellied form with yellow lines bordering the grey and a faint mid-ventral yellowish stripe. Mus tenellus (Thomas) Blue Nile Gorge: four from 10 km west of Mabil, trapped in dry scrub; one 10 km east of Caroarsa River, trapped amongst tall, wet grass. Awash: two, probably of this species, in dry Acacia scrub close to riverine wood- land. The group of small, white-bellied Mus (or Leggada) badly needs revision. These have been compared with the type of fenellus (from Roseires on the Sudanese Blue Nile) and there seems little doubt that at least the Blue Nile animals can be allocated to that form. Mus ?pasha (Thomas) Blue Nile Gorge: eight from 10 km west of Mabil, trapped in grassland adjacent to permanent riverine vegetation. These are white-bellied mice considerably larger than M. tenellus, with head and body length of 69-85 mm (49-65 mm in M. tenellus), hind feet of 14-15 mm (12-13 mm in M. tenellus), and upper molar row of 3-7 mm (two specimens; 3-I and 3:0 mm in two M. tenellus). Leggada pasha was described from the Uele District, N.E. Congo but was treated by Osgood (1936) as a race of Mus proconodon Rhoads from Somalia. Setzer (1956) did not record either of these forms from the Sudan. Genus PRAOMYS—Soft-furred rats Praomys albipes (Rippell) Debra Markos: one caught by hand in camp amongst low Acacia scrub. Bahar Dar: two trapped in thickets in wet grassland. Blue Nile Gorge: one caught on a boat, 30 km below Portuguese Bridge near Mota at 11° 12'N, 38° 05’E, altitude 1400 m. Addis Ababa: nine from about 48 km west of Addis Ababa (from Wellcome Parasitology Unit). Dessie: one from 31 km N.W. of Dessie (from Wellcome Parasitology Unit). Lake Abbaya: eight from the Bonchi Valley, west of the lake at 2900 m. This is a clearly defined species, formerly placed in the genus Myomys, confined to the Ethiopian Highlands except for an isolated and only doubtfully conspecific form, P.a. fuscirostris, in Sudan. These specimens, like all those recorded from Ethiopia, belong to P.a. albipes. Allare from the plateau, between 1400 and 3000 m. Praomys fumatus brockmani (Thomas) Harar: one trapped in thorn scrub beside a dry river bed 60 km east of Harar on Jiggigga road. MAMMALS FROM ETHIOPIA 235 This specimen agrees well with the type of brockmani from northern Somalia and is distinctly paler than P.f. allisoni Hayman from central Ethiopia. This species was recorded from the Harar area by Ingersol (1968). Praomys (Mastomys) natalensis (Smith) Blue Nile Gorge: one skull found at mouth of Guder River; one trapped in dry scrub 10 km west of Mabil; four trapped in long grass at the edge of riverine forest at the mouth of the Azir River; eleven from the mouth of the Didessa River, in long grass on the river bank and in maize cultivation near the village. Bahar Dar: one trapped in thicket in wet grassland. Ghimbi: one 24 km north of Ghimbi, in long grass. Harar: two from Bisidima River, in dry, grazed thorn-scrub. Awash: four from dry Acacia scrub; a few skulls from the lava caves. Lake Abbaya: eight from the following localities. Bonchi Valley, west of lake, 2900 m; Collufo River, between Lake Abbaya and Lake Chamo; mouth of Darsi River. Little Abbai: six collected. Osgood (1936) recognized two species of ‘Mastomys’ in Ethiopia, which he called M. coucha lateralis (Heuglin) and M. macrolepis (Sundevall). The latter he described as having a longer tail, shorter hind feet and paler ventral pelage, and recorded from the Blue Nile Gorge and Lake Tana (the type locality is Roseires on the Blue Nile in Sudan). Setzer (1956) listed macrolepis as a race of M. natalensis (M. coucha), although he recognized two sympatric species of Mastomys elsewhere in Sudan. In the present collection the animals from the Nile Gorge have the tail longer than the head and body (measured from the anus) with four exceptions out of 16 (mean 102°4 o of head and body); those from Awash and Harar have the tail shorter than the head and body in five out of six animals (mean 94°7%). The Nile animals have the ventral pelage paler than the eastern ones, but there is no difference in length of hind foot: mean of 24-6 mm in the gorge, 24-5 mm in the eastern samples. The single skin from Bahar Dar on Lake Tana is dark-bellied with a short tail as in the eastern samples. These are all tentatively referred to P. natalensis, but this group badly needs revision, especially in view of the demonstration of sibling species with very different chromosome complements elsewhere in Africa (Matthey, 1966). Rattus rattus alexandrinus (Geoffroy)—House rat Ghimbi: three collected in or around the town. Harar: three from the Bisidima River. Lake Abbaya: six from near Soddu. These are called R.r. alexandrinus following the cevision by Schwarz and Schwarz (1967) who treated Mus samharensis Heuglin (Ethiopia) and Mus kijabius Allen (Kenya) as synonyms. 236 G. B. CORBET & D. W. YALDEN Dendromus mystacalis Heuglin Blue Nile Gorge: one from mouth of Guder River, caught by hand in the camp near steep rocky hillside with thick bush; one at mouth of Didessa River, caught by hand under bushes ona smallisland. Both were in the riverine forest strip. This is a poorly known species in Ethiopia although it appears to have a wide distribution through much of Africa. These specimens have a very slight darkening in the mid-dorsal area but no black stripe, and a similar specimen is in the British Museum from 230 km south of Addis Ababa. Specimens from Dangila and Lake Tana on the other hand mostly have a clear dorsal stripe. Otomys typus (Heuglin) Debra Markos: one trapped by day in low, grazed Acacia scrub. This specimen represents the nominate subspecies, described from the Simien Mountains. Family GLIRIDAE—Dormice Graphiurus murinus ?internus Dollman Blue Nile Gorge: one at the mouth of the Fincha River, trapped in riverine forest, 45 cm off the ground in dry bushes. There are very few previous records of Graphiurus from Ethiopia. These appear to be from Mt. Amar Cocche, Omo Valley, recorded as Claviglis brockmant internus by de Beaux (1943); Saganeita, Eritrea, recorded as probably C. orobinus by de Beaux; Guri Dagono and Sidamo, Arussi, recorded as G.m. satwratus by Hayman (1960) ; and Gara Mullata, Harar, recorded by Ingersol (1968) without allocation to subspecies. Only the skin from Guri Dagono has been available for comparison. The present specimen appears identical, but with respect to colour these specimens are much closer to the type of G.m. internus Dollman from northern Kenya than to the darker G.m. saturatus from Mount Elgon. The skull is, however, rather large (condylobasal length 23:3 mm as compared with 20-7 in the type of internus, a slightly older animal judging by the teeth). This genus has never been adequately revised and the allocation of all named forms from northeastern Africa to G. murinus (type locality Cape of Good Hope) must be considered very tentative. A form described from Senaar on the Blue Nile in Sudan, Myoxus orobinus Wagner, 1845, may be relevant, but no further specimens have been reported from that region of Sudan and the original description is in- adequate. Family RHIZOMYIDAE—Mole-rats Tachyoryctes splendens (Rippell) Dessie: one from 18 km N.W. of Dessie (from the Wellcome Parasitology Unit no. 2). Addis Ababa: one from Meta Abo, Sabata, c. 20 km S.W. of Addis Ababa. MAMMALS FROM ETHIOPIA 237 Lake Abbaya: two from the Bonchi Valley at 2900 m; one from Mount Gugi (2400 m); and three from Arba Minch. These can probably all be referred to T.s. splendens, being rather smaller than T-s. cheesmani Thomas from Lake Tana. Tachyoryctes macrocephalus hecki Neumann and Rimmler Bale Mountains: one (skin and skull) from Dinshu (=Gurie), Ueb Valley, at 3500 m (from Mr J. H. Blower); a fragmentary skull from an eagle’s nest 30 km south of Dinshu at 3800 m (sent by Mr L. H. Brown); a complete skull from Batu (6° 55’N, 39° 47’E, 4150 m) collected by Mr H. F. Mooney. Mr Blower (in litt.) also saw colonies of this species at 3800 and 4000 m in the same area, apparently confined to ‘open moorland where the grass is very short, often in relatively damp places, some- times near streams’. This is a very large mole-rat, about twice the size of the common T. splendens. Both the nominate race (from Shoa) and T.m. hecki appear to be known only from the original specimens, namely three syntypes of macrocephalus (in Berlin, Frankfurt, and London) and the type of heckz in Berlin. The type locality of hecki was given as ‘Abakkara, etwa 150 km westlich des Abassi Sees in Grenzgebiet zwischen Djamdjam und Arussi Galla gelegen’. As- suming that ‘Abassi See’ is Lake Awusa, the ‘westlich’ would seem to be an error for ‘ostlich’. This would place it not only between Djamdjam and Arussi but close to the locality of the records reported here. The present specimens agree with the description of hecki, and differ from the London syntype of macrocephalus, in having the rostrum and upper incisors rela- tively small (Table 1). This difference is very marked and suggests that these might prove to be genuinely discrete subspecies or even species, perhaps isolated by altitude. Even in T.m. macrocephalus the incisors are relatively smaller than in the small T. splendens, suggesting a less fossorial habit, and this is supported by the sight records, the animals being described as marmot-like and frequently visible on the surface (J. H. Blower, in Jitt.). There is an old record of ‘Tachyoryctes macrocephalus’ from Kita in the upper part of the Senegal basin (now in Mali) (Rochebrune, 1883). It was described as ‘plus commune ... dans les environs de Kita, ot elle habite les crevasses des rochers et les pentes sablonneuses des collines boisées; les Européens la désignent sous le nom de Marmotte’. It seems probable that this account really refers to the gundi, Felovia vae (Lataste), which was not otherwise mentioned by Rochebrune and was first dis- covered by Lataste in the Felou Hills, about 250 km northwest of Kita, and des- cribed by him in 1886. Ignoring this probably erroneous record, T. macrocephalus is endemic to the Ethiopian plateau. 238 G. B. CORBET & D. W. YALDEN TABLE I Measurements of Tachyoryctes macrocephalus T. m. hecki T. m. macrocephalus —— SEE) SS SSS Type of Senck. BM 68.856 BM 70.746 hecki* BM 71.876 BM 42.8.15.1 Mus. 728* Berlin* Length of head and body (from dry skin) c. 400 Length of hind feet 33 34 Condylobasal length 64:0 Width across pre- maxillae TI-2 12:2 c. 10°9 11-8 I4°1 Combined width of upper incisors (near tip) TP 7:0 7:2 7:0 o5 8-6 05 Interorbital width 57 6-0 6-2 6-2 57 63 Length of nasals 29°6 30°2 28-3 27°9 Upper molar row (alveolar) 12-0 11-8 1370 12°1 12-0 12°2 Lower molar row (alveolar) 13°5 13°8 13°8 13°I 13°3 13'7 Length of mandible 43 C. 42 43 c. 45 43 * From Neumann & Riimmler, 1928. Family THRYONOMYIDAE—Cane rats Thryonomys gregorianus (Thomas) Blue Nile Gorge: one damaged skull from a cave 10 km W of Mabil; possible sightings at the Didessa River. Wollega Province: one specimen from Bako in Addis Ababa Museum (skull and colour photo of mounted skin examined by G.B.C.). Ilubabor Province: one skin from Gambela collected by Dr J. Ash in March 1969. The Blue Nile specimen was on the surface of the floor of a cave covered with bat and hyrax guano and therefore must be very recent. These appear to be the first records of Thryonomys from Ethiopia, and in Sudan the only record of T. gregorianus is from the Didinga Mountains on the southern border. These skulls agree closely with the type of gregorianus (from central Kenya) and differ from the type of harrisont Thomas & Wroughton from southern Sudan in having the nasals short and wide; the posterior processes of the premaxillae wide and rounded (in dorsal view); and the zygomatic parts of the lachrymals elongate. The skin from Gambela could con- ceivably be 7. harrisoni if that is distinct. Both skins have the short tail (not ex- tending beyond the hind feet when these are extended backwards) characteristic of T. gregorianus and T. harrisoni in contrast to the very much longer tail of T. swinderianus. The skin from Gambela is of a young animal with the following measurements: head and body 329; tail 73; hind feet 71; and ear 25 mm. Setzer (1956) treated T. gregorianus and T. harrisoni as specifically distinct, re- cording both from southern Sudan. Misonne (1968) recognized only two species in MAMMALS FROM ETHIOPIA 239 the genus, T. swinderianus and T. gregorianus, presumably treating harrisoni as conspecific with gregorianus, but in view of the cranial differences and the small numbers of specimens available it seems wise to treat them as provisionally distinct. A subfossil Thryonomys from a mesolithic site at Khartoum, Sudan has been described as T. arkelli by Bate (1947, 1949). Bate considered it more closely related to the larger T. swinderianus than to T. gregorianus or T. harrisoni and a re-examina- tion of the type supports this view. It differs from the types of T. gregorianus and T. harrisoni and from the Ethiopian specimens in its slightly larger size, prominent masseteric knob and robust anteroventral root of the zygomatic arch. Family HYSTRICIDAE—Porcupines Hystrix cristata L.—Crested porcupine Lake Chamo: one from northeast of lake, in the ‘White Grass’ area. Blue Nile Gorge: quills found at the Mugher, Guder, Fincha and Caroarsa Rivers. Awash: quills found in the lava caves. H. cristata is here considered to include the East African galeata (Corbet & Jones, 1965). Order CARNIVORA Family CANIDAE Lycaon pictus (Temminck)—Hunting do § dog Lake Abbaya: one shot near Soddu in 1964. Canis aureus L.—Golden jackal Luma, Didessa Basin: a skull received from Mr John Blower is probably of this species. The animal was shot in a cultivated clearing in bamboo forest at 9° 40’N, 35° 55'E, 2000 m. Dangila: one shot between Dangila and Engiabara. Little Abbai: two collected. Otocyon megalotis (Desmarest)—Bat-eared fox Harar: one from the Bisidima River; one from 60 km east of Harar on the Jiggigga road; eight others seen in this area. Awash: a skull was found on the floor in one of the lava caves. Family MUSTELIDAE Ictonysx striatus (Perry)—Zorilla Awash: one seen at night. Addis Ababa: one dead on the main road south-east from Addis Ababa to Shas- hamane. 240 GAB CORBET D2 Wis YoAIeD BN, Family VIVERRIDAE Genus GENET T A—Genets Genetta genetta (L.) Harar: one from the Bisidima River; one from 60 km east of Harar on the Jiggigga road. Genetta ?tigrina (Schreber) Ghimbi: one from 24 km east of Ghimbi. Lake Chamo: one from northwest of lake. Lake Abbaya: one from Arba Minch, southwest of lake. The thiee skins collected are very similar in colour and pattern. According to Coetzee (1967) the Ethiopian race is G.t. schradert Matschie. The taxonomy of this group of genets is in a very provisional state. It is possible that the northern forms of the tigvina group, including schraderi, represent a separate species, G. rubiginosa. A third species of genet recorded from Ethiopia, but not represented in the present collections, is G. abyssinica (Rippell) which Coetzee (1967) separated subgenerically from G. genetta and G. tigrina. Ichneumia albicauda (G. Cuvier)—White-tailed mongoose Bahar Dar: one collected. Ghimbi: three collected. Lake Abbaya: two from Arba Minch. Lake Chamo: three from northwest of lake; two from 12 km south of Gardulla, south of the lake. Blue Nile Gorge: one seen at Mabil. Herpestes sanguineus Riippell—Slender mongoose Lake Chamo: one from west side of lake; one from northeast of lake. Viverra civetta Schreber—Civet Ghimbi: one from to km east of town. Lake Abbaya: one from near Soddu; one from Arba Minch. Family HYAENIDAE Proteles cristatus (Sparrman)—Aardwolf Awash: the remains of five animals were recovered from one of the lava caves, one as a complete skeleton and four skulls. The aardwolf has a very wide distribution in Africa, but it is a very elusive animal and is seldom recorded. Ingersol (1968) reported it from the Afden Plain just east of the Awash and at Amarresa in the Harar district. MAMMALS FROM ETHIOPIA 241 Crocuta crocuta Erxleben—Spotted hyaena Harar: about 15 hand-fed by the ‘hyaena man’ at dusk outside the city. Ghimbi: one dead on the main road east of the town. Debra Markos: up to six seen around the town. Family FELIDAE Felis silvestris libyca Forster—Wild cat Harar: one from the Bisidima River. Bahar Dar: one collected. Blue Nile Gorge: one from Shafartak. Felis serval Schreber—Serval Degen, Gojjam: one collected. Ghimbi: one collected. Little Abbai: two collected. These all have the ground colour of the pelage yellowish as in F’. s. hinder Wrough- ton from Kenya which they closely resemble, and are not nearly so pale as in the type of Fs. tanae (Pocock, 1944) from Lake Tana. Panthera pardus (L.)—Leopard Blue Nile Gorge: one seen beside the river, between Shafartak and the Mugher; traces near the Caroarsa River. Awash: two seen. Panthera leo (L.)—Lion Awash: two seen hunting together. Blue Nile Gorge: unconfirmed report of a party near the mouth of the Guder River. Order HY RACOIDEA Procavia habessinica (Hemprich & Ehrenberg)—Large-toothed rock hyrax Harar: two from Valley of the Rocks, 40 km east of Harar; two from 60 km east of Harar (all P.h. erlangeri Neumann). Lake Langano: two from west side of lake (P.h. alpini (Gray)). These two groups are so different in appearance that their conspecificity must be considered doubtful. The P.h. erlangeri agree closely with a series in the British Museum from Dire Dawa, about 50 km N.W. of Harar, and specimens were recorded from Harar by Neumann. These all have the head black with a rufous tinge, the dorsal pelage mottled yellow and black and the dorsal glandular spot scarcely visible. A very young animal (head and body 260 mm) has the head equally black but there is less yellow in the rest of the dorsal pelage. 242 G. B. CORBET & D. W. YALDEN The skins of P.h. alpint have the head much lighter, the dorsal pelage greyish brown and finely speckled, and the glandular streak prominently yellow. The yellow glandular streak distinguishes this form from P.h. habessinica from N.E. Ethiopia and from P.h. scioana (Giglioli), a very dark form from Ankober, Shoa, as well as from P.h. erlangert. In view of the many problems remaining to be solved in this genus it seems pre- mature to assume these to be conspecific with the South African P. capensis as suggested by Ellerman & Morrison-Scott (1951). Heterohyrax brucei brucei (Gray)—Small-toothed rock hyrax Harar: two from 60 km east of Harar, at precisely the same locality as the Pro- cavia habessinica. Blue Nile Gorge: mandibles found at mouth of Guder River. Lake Abbaya: two from Arba Minch; two from Arba Mondi, south of lake. The specimens from Harar agree closely with the type of brucei. The form hararensis Brauer from near Harar was listed by Allen (1939) as a synonym of somalica Thomas from N. Somalia, described as smaller and paler than the type of brucet. It is unlikely that somalica will prove a valid subspecies, but in any case the specimens from Harar seem closer to brucei than to somalica. H.b. hararensis was diagnosed as smaller than brucei and distinguished from both brucei and somalica by greyish white ventral pelage. In fact the present specimens have quite dark yellow- ish buff undersides, one of them with an irregular pure white area of the chest and in front of the genitalia. The dorsal glandular streak is bright yellowish brown. The specimens from Lake Abbaya in the Rift Valley are scarcely distinguishable from those from Harar. Order TUBULIDENTATA Orycteropus afer (Pallas)—Aardvark Debra Markos: burrows seen. Awash: one killed by a car on the track in the park. Order PERISSODACTYLA Family EQUIDAE Equus burchelli boehmi Matschie—Common zebra Lake Chamo: up to 96 counted in the ‘White Grass’ area. Order ARTIODACTYLA Family HIPPOPOTAMIDAE Hippopotamus amphibius L.—Hippopotamus Blue Nile Gorge: seen from the Fincha River westwards, including Wamet Ford (4), Azir to Caroarsa (12), Didessa (tracks, and several heard at hight). Heard also MAMMALS FROM ETHIOPIA 243 in the vicinity of Tississiat Falls. A skull found at the mouth of the Azir River was preserved. Awash: one seen on several occasions. Lake Abbaya: two skulls were found and preserved. Family SUIDAE Phacochoerus aethiopicus (Pallas)—Wart hog Lake Abbaya: six collected, two near Soddu and four from near Arba Minch. Lake Chamo: group of nine seen in the ‘White Grass’ area. Awash: two family parties of 4+ seen; a skull was found in one of the lava caves. Potamochoerus porcus (L.)—Bush pig Lake Abbaya: one from south of the lake. Family BOVIDAE Madoqua phillipsi hararensis Neamann—Phillip’s dik-dik Harar: three from 60 km east of Harar on the Jiggigga road. Awash: numerous sight records, each of one or two animals only. Madoqua guentheri Thomas—Giinther’s dik-dik Lake Chamo: four from the ‘White Grass’ area. Sylvicapra grimmia (L.)—Common duiker Bahar Dar: one collected, and several others seen. Little Abbai: two collected. Lake Abbaya: one from near Soddu. Sire, Wollega: skull preserved of a female killed on the main road from Ghimbi to Addis Ababa, between the towns of Sire and Bako at 9° 08'N, 37° 05’E. Ourebia ourebi (Zimmerman)—Oribi Blue Nile Gorge: four seen at night on the air-strip at Sirba. Oreotragus oreotragus (Zimmerman)—Klipspringer Blue Nile Gorge: two seen between the Guder and Fincha Rivers and two between the Fincha River and Wamet Ford. Harar: two seen in the Valley of the Rocks. Kobus defassa (Rippell)—Waterbuck Awash: several, including a party of four. Lake Chamo: a herd of 16 in the ‘White Grass’ area. 244 G. B. CORBET & D. W. YALDEN Alcelaphus buselaphus swaynei (Sclater)—Common hartebeest Lake Chamo: up to 104 counted in the ‘White Grass’ area. This subspecies is considered to be extinct in Somalia, and perhaps only 200 exist in Ethiopia (Simon, 1968). Tragelaphus scriptus (Pallas)—Bushbuck Ghimbi: two from 16 and 20 km east of Ghimbi. Lake Abbaya: two from Arba Minch. Blue Nile Gorge: one seen at Mabil. Bahar Dar: one seen. Awash: one seen. Tragelaphus imberbis Blyth—Lesser kudu Awash: seen in small numbers. Tragelaphus strepsiceros (Pallas)—Greater kudu Lake Chamo: two from between Lake Chamo and Lake Abbaya. Lake Abbaya: one from Borodda, N.W. of lake. Awash: up to 14 seen. Litocranius walleri (Brooke)—Gerenuk Awash: three seen. Gazella soemmerringi (Cretzschmar)—Soemmerring’s gazelle Awash: large numbers seen, in groups of 50-100. One was collected and a skull was retrieved from one of the lava caves. The specimen collected resembles the Somalian race, G.s. berberana Matschie, 1893, in its horn shape and extensive dark nose spot, and the Awash animals can probably be attributed to that race. This specimen does not agree well with Matschie’s (1912) description of G.s. erlangeri from Awash. Gazella granti Brooke—Grant’s gazelle Lake Chamo: two collected in the ‘White Grass’ area and 46 counted on one occasion. Oryx gazella beisa (Rippell)—Oryx Awash: large numbers seen, including one herd of 150. Parts of two skulls were retrieved from the lava caves. MAMMALS FROM ETHIOPIA 245 DISCUSSION Zoogeographically, the fauna of Ethiopia (usually termed, to avoid confusion, the Abyssinian fauna) is noted for its peculiar caste. While much of the fauna is of a general East African type, including species with a wide distribution even down to the Cape Province, there is also a small proportion of species of Palaearctic affinities and a significant roster of endemic species as well. Moreau (1966) for the birds and Carcasson (1964) for the butterflies have commented on these peculiarities of the Abyssinian fauna when seen from the viewpoint of continental zoogeography, while Carpenter (1935), approaching the subject from a study of the Abyssinian butterfly fauna particularly, and including also relevant information from other groups, has discussed the different taxa. The conspicuous endemicity undoubtedly results from the large area of high plateau included in Ethiopia and its isolation from neighbouring mountain areas— both Moreau and Carcasson note that two habitat types, montane woodland and montane grassland, contain most of the endemics in the groups they studied. The interplay of a variable Pleistocene climate and this montane area is presumably responsible for the Palaearctic element reaching Ethiopia—during glacial (pluvial) periods, montane habitats would have occurred at lower altitudes, and would have spread up the mountains along the Red Sea coast of the Sudan and Nubia. This spread of montane habitat in the glacial periods would also account for another peculiar zoogeographical relationship of some Abyssinian fauna, that element which is of general montane type and may as a result have affinities with the fauna of the Cameroon Highlands of West Africa. Any zoogeographical analysis of the mammalian fauna has hitherto been mainly confined to the larger mammals—see for example Blower (1968), Scott (1958) and Glass (1965)—and can be usefully illustrated by the maps showing the range of individual species throughout Africa in Dorst & Dandelot (1970). In the following analysis the smaller mammals are also considered, although many taxonomic problems remain to be solved before this can be quantified precisely at the levels of the species and subspecies. The East African savanna and steppe element This is the dominant element in southern and western Ethiopia, in the Rift Valley and in the valley of the Blue Nile. Many of the species concerned have wide distri- butions throughout the savanna zone from West Africa to Tanzania and further south, e.g. the giraffe Giraffa camelopardalis, the hartebeest Alcelaphus buselaphus and the striped mouse Lemniscomys striatus. Species of this group vary in the extent to which they penetrate Ethiopia. L. striatus occurs in the valley of the Blue Nile and its tributaries but does not appear to occur in the Rift Valley. A. buselaphus on the other hand extends through the Rift Valley to the Awash Valley. Grant’s gazelle Gazella granti shows another pattern in that it reaches from Kenya into the southern part of the Rift Valley but is replaced by a related species, G. soemmerringz, in the northern part of the rift. Small mammals falling into this general group, all reaching their north-eastern 246 G. B. CORBET & D. W. YALDEN limit in Ethiopia, are Graphiurus murinus, Praomys fumatus, Elephantulus rufescens and Galago senegalensis. Species with a similar range, but also reaching north of Ethiopia, are Acomys spp., Arvicanthis niloticus, Hystrix cristatus, Lepus capensis, Crocidura flavescens s.l, and Felis silvestris. The Somalian arid zone element Somalia is an area with a considerable degree of endemicity amongst the dry steppe fauna, and some of these species reach into eastern Ethiopia. The most sharply differentiated (endemic genera) are the gundi, Pectinator spekei, the naked mole-rat, Heterocephalus glaber, which reaches west as far as Shoa, the dibatag, Ammodorcas clarkei, which just reaches eastern Ethiopia, the beira, Dorcatragus megalotis, and the gerbil Microdillus peeli, the last two unrecorded from Ethiopia. Another group comprises species that are more closely related to widely distributed savanna or steppe species, e.g. Gazella soemmerringt which has a wide range, reaching round the northern edge of the Ethiopian highlands to the Sudan, the dik-dik Madoqua phillipsi and the grass-mouse Arvicanthis somalicus here recorded from Awash. Comparable Somalian species that do not appear to have been recorded in Ethiopia are the elephant shrew Elephantulus revoili and the hedgehog Evinaceus sclateri. The Palaearctic element This is a very small element in the mammalian fauna comprising the ibex, with Capra ibex nubiana in the Red Sea Hills and a southern isolate, C.2. walie, in the Simien Mountains; the wild ass Equus africanus in the Danikil Desert; and the shrew Suncus etruscus, here reported from the plateau but in its Mediterranean range a species of the drier lowlands. It is probable that the hedgehog Paraechinus aethiopicus, a species with a Saharan and Arabian distribution, is also present in northern Ethiopia. All these are tolerant of dry conditions. The ass probably had a continuous range from North Africa to Somalia in recent times, but the ranges of the ibex and the shrew are of a relict character, the latter showing a considerable degree of disjunction although it is an easily overlooked species and may still be found in intervening regions. Among the birds of Ethiopia, the most notable Palaearctic immigré is the chough Pyrrhocorax pyrrhocorax, while the endemic owl Asio abyssinicus is considered a close relative of the northern A. otws; other Palaearctic birds listed by Moreau (1966) include the grebes Podiceps caspicus and P. ruficollis, the lammergeier, Gypaetus barbatus, Alpine swift, Apus melba, and quail Coturnix coturnix, which occur also into Kenya. Among the butterflies Carcasson (1964) and Carpenter (1935) list seven or eight species of Palaearctic affinity, including the endemic ‘speckled wood’, Pararge madarakal, and Pieris brassicoides, found also in Tanganyika. The East African montane element In spite of the very great distances separating the montane forest and grassland habitats of Ethiopia from comparable habitats further south, e.g. in the Kenya MAMMALS FROM ETHIOPIA 247 highlands and Mount Elgon, there are several forms showing very little differentia- tion between these segments of their ranges. Examples amongst the rodents are Lophuromys flavopunctatus, Lophiomys imhausi, Otomys jacksoni and Tachyoryctes splendens. Some of these species, however, have moderately wide ecological requirements, e.g. L. flavopunctatus which is here recorded from seasonally dry habitats near Harar although it is more strictly confined to forest farther south. A lesser known rodent with a highly disjoint distribution is Colomys goslingi, known from swampy habitats in Cameroon, N.E. Congo, Kenya and the Ethiopian plateau where it is known from a single specimen previously believed to represent an endemic genus Nilopegamys (see Hayman, 1966). The mountain reedbuck, Redunca fulvoru- fula, also has a highly disjunct range with isolates in S. Africa and perhaps in Cam- eroon, but it is tolerant of fairly dry habitats. Amongst the birds and butterflies there are likewise a number of species with strikingly disjunct distributions in various montane areas in East Africa, and across in some cases to the Cameroon Mountains and in others as far as Cape Province. The weaver Cryptospiza salvadori, for example, occurs in isolated patches of Ethiopia, Uganda, Kenya and Tanzania, while the babbler Alcippe abyssinicus occurs in these areas and in the Cameroons as well (Moreau, 1966). Carpenter (1935) listed about ten butterflies with similar distributions. The endemic element Endemic Ethiopian species are confined to the highlands and especially to the high montane grasslands which have presumably had the longest period of isolation and until recent deforestation were presumably much more sharply isolated from any comparable lowland habitats than they are today. The species concerned are the gelada baboon, Theropithecus gelada, the Simien fox, Canis simensis, the mountain nyala, Tragelaphus buxtoni, the genet Genetta abyssinica, the giant mole-rat Tachyoryctes macrocephalus, and the murid rodents Stenocephalemys albocaudata, Muriculus imberbis, Dendromus lovati, Praomys albipes, and Pelomys dembeensis. All these are well defined species, and three are currently considered endemic genera, namely Theropithecus, Stenocephalemys and Muriculus. It is possible that further endemic species occur in the genera Crocidura, Procavia, Lophuromys, Arvicanthis, Praomys, Mus and Pelomys, but further revision is needed to clarify their relation- ships with neighbouring forms. All of these endemics belong to African rather than to Palaearctic groups, and all have their closest relatives in Ethiopia or in adjacent parts of East Africa. There are probably about 135 species of mammals in Ethiopia (excluding bats and marine species) and the eleven clear-cut endemics listed above therefore represent about 8-5 % of the fauna, a figure that is likely to increase somewhat with further revision. Of the estimated 124 non-endemic species, about 16 are montane species whose Ethio- pian populations are widely isolated from the rest of the range whilst the remainder, the great majority, are the more xerophilous species with ranges extending relatively unbroken into surrounding territories. Carcasson (1964) listed at least 8 endemic butterflies (Pararge maderakal, Papilio 248 G. B. CORBET & D. W. YALDEN aethiops, Mylothris mortone, Bicyclus aethiops, Charaxes phoebus, Acraea oscart, A. ungemachi, A. safie) while Carpenter (1935) listed in all about 526 species from above 1500 m in Ethiopia; neither of these figures are complete, but they suggest an endemicity in the fauna of perhaps 2°4. Among the endemic birds, Moreau (1966) mentioned the owl Asio abyssinicus, blue-winged goose Cyanochen cyanopterus, a swallow Hirundo megaensis, and the crow Zavattariornis stresemanni. In all, he suggested that there are 3 montane-forest and 21 montane non-forest endemic birds, while Urban & Brown (1971) listed 23 endemics in a total avian (breeding) fauna of 655 species. This would suggest an endemicity of about 3:5 %. Summary of Zoogeographical Affinities It is clear from what little information has been presented above that the mammal fauna of Ethiopia shares with the better known bird and butterfly faunas the zoogeographical peculiarities of a conspicuous endemic element, a Palaearctic element, and also an element of discontinuously distributed montane animals. There is a suggestion that among the mammals the endemic element is greater and the other two elements fewer, and it is tempting to speculate on the role of relative mobility of the mammals as against the birds and butterflies. Unfortunately the mammalian faunas of both Ethiopia and other areas of importance are not suf- ficiently well known to be categorical about this—it may be relevant to note here that this paper records four species new to Ethiopia (Crocidura niobe, Suncus etruscus, Thryonomys gregorianus and Arvicanthis somalicus) and records several others that have been previously noted fewer than six times (Zachyoryctes macrocephalus, Taterillus emini, Lemniscomys striatus, Praomys fumatus, Proteles cristatus). Fauna of the Blue Nile Gorge Since the main object of the 1968 expedition was to carry out a scientific survey in the gorge of the Blue Nile, some concluding comments on the small mammals ob- tained there are justified. One point that was very evident in the field was the scarcity of small mammals—over nine hundred trap-nights were required to obtain 52 small mammals, or 17 trap-nights per specimen (for comparison, at Awash, about 5 trap-nights per animal, and at Bahar Dar 15 per animal, were required). From the species caught, it is evident that the small mammal fauna of the gorge is comparable with that found at Awash and east of Harar; among the species or genera common to both are Crocidura ?sericea, Acomys dinudiatus, Heterohyrax brucei, Procavia habes- sinica (on sight records only from the Blue Nile Gorge), Tatera sp. (T. valida in the Gorge, T. robusta in the East), Mus tenellus, and Oreotragus oreotragus. By contrast, the collecting localities on the plateau (Ghimbi, Bahar Dar, Debra Markos) have in common Lophuromys flavopunctatus, and Praomys albipes was found at the two more northern of these. The presence in the Blue Nile Gorge of Tatera valida, Lemniscomys striatus, Mus ?pasha and Acomys spp. implies perhaps the in- cursion of an East African savanna element. While for human travel the Blue Nile Gorge represents a considerable barrier, so far as small mammals are concerned MAMMALS FROM ETHIOPIA 249 it would seem rather to be a highway allowing dry-country, lowland species to pene- trate deeply into the centre of Ethiopia. ACKNOWLEDGEMENTS Thanks are due to all those members of the Great Abbai Expedition and the previous expeditions who made these collections, and this report, possible. If this acknowledgement seems rather curt, full justice is done in the expedition reports. So far as the actual collection of mammals is concerned, particular thanks are due to Dr P. A. Morris, Dr M. J. Largen, and H. King. D.W.Y. would like to acknowledge the receipt of a staff travel grant from the University of Manchester. We are grate- ful also to Mr John Blower, Mr Leslie Brown, Dr R. Ashford and Dr Aklelu Lemma for sending specimens. GAZETTEER Altitude Dates of (metres) collecting Abiu, Ilubabor 8°11'N _35°22’E 1500 January 1971 Addis Ababa, Shoa g°00’N = 38°45/E 2300 Addis Alem, Shoa 9°03,N = 38°25'E 2400 Arba Minch, Gemu Gofa 6°04’N —- 337°40’E 1300 6 August-3 September 1966 Awash, Shoa 8°50’N _ 40°o1’E 950 23-28 September 1968 Awash (lava caves), Shoa 8°56’N = 339°57’E 950 28 September 1968 Azir River (mouth of) 10°290'N —- 336°25/E 800 20-22 August 1968 Bahar Dar, Gojjam 11°35'N 37°25/E 1830 11-14 September 1968 Bako, Wollega 9°08’N —- 337°05E 1800 Bisidima River, Harar g°15'N 42°12’E 1500 20-21 September 1968 Bonchie Valley, Gemu Gofa 6°05'N —- 37°20’E 2700 1966 Caroasa River (near mouth) 10°07’N = 336°12”E 600 23-24 August 1968 Collufu River (mouth),Gemu Gofa 6°04’N 37°40’E 1300 6 August-3 September 1966 Danghila, Gojjam 11°r5’N—- 36°55/E 2000 Darsi River, Gemu Gofa 6°15'N 37°50°E 1300 Debra Libanos, Shoa 9°45,N 38°50°E 2300 Debra Markos, Gojjam 10°20N 37°50’E 2500 4-27 August 1968 Degen, Gojjam to°10o’N = 38°05’E 2500 Dessie, Welo 10°08’N —- 39°43’E 2700 Didessa River (mouth of) 10°05'N- 35°38’E 450 26-29 August 1968 Dim River (mouth of) 10°30.N 36°26’E 800 22 August 1968 Engiabara, Gojjam 10°58’N - 36°58’E 2500 Fantalle (Mt), Shoa 8°58’N —- 339°54’E 2000 Filhoa, Awash, Shoa g°00’N = 339° 58’E 1000 23-28 September 1968 Fincha River (mouth of) 10°03'N- 37°20’E 1000 11-13 September 1968 Gambela, Ilubabor 8°15'N —- 334°35'E 600 Ghimbi, Wollega g°I0’'N = 35°50’E 2150 22-31 August 1968 Guder River (mouth of) 9°50°N 37°41’E 1000 8-11 August 1968 Gughe (Mt), Gemu Gofa 6°05’'N = 37°20’E 2700 Harar, Harar 9°20'N 42°08’E 2000 Jiggigga Road (60kmE.ofHarar) 9°14’N 42°32’E 1200 21-22 September 1968 250 G. B. CORBET & D. W. YALDEN Lake Abbaya, Gemu Gofa (S.W., near Arba Minch) 6°04’N- 337°45'E 1300 6 August—3 September 1966 Lake Abbaya, Gemu Gofa (N.W., near Soddu) 6°40'N —- 38°00’E 1500 8-23 August 1964 Lake Chamo, Gemu Gofa (White Grasses) 5°58 N 37°55 E 1500 1966 Little Abbai, Gojjam 11°20’N = 37°00’E 2000 27 August— 5 September 1964 Mabil, Gojjam 10°19’N —-36°45/E goo 14-20 August 1968 Mota, Gojjam 11°05’N-337°54’E 2500 Metahara, Shoa 8°53,N = 39°55/E 1000 Mugher River (mouth of) 9°50 N 37°55'E 1000 8 August 1968 Nono, Upper Omo 8°30'N 37°21"E 1600 Sabeta, Shoa 8°55'N 38°40’E 2500 Shafartak 10°06’N = s-38°17’E 1150 Sirba, Wollega 10°50°N—- 35°30’E 400 29-30 August 1968 Sire, Wollega 9°03'N—- 36°54’E 1800 Soddu, Sidamo 6°45N 37°50'E 1800 Tississiat Falls Tr-30°N) 37-3708: 1800 Valley of the Rocks, Harar 9°15 N 42°20'E 1300 21 September 1968 Wamet Ford 10°09’N 337° 12”E 1000 20 August 1968 REFERENCES ALLEN, G.M. 1939. G. B. CorBEet Department of Zoology British Museum (NatTuRAL History) CROMWELL Roap Lonpon, SW7 5BD D. W. YALDEN Department of Zoology UNIVERSITY OF MANCHESTER MANCHESTER, M13 9PL A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) Kay, E. Attson. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 1965. (Out of Print.) £3.75. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180; 11 Plates, 15 Text-figures. 1967. £4. Taytor, J. D., KENNEDY, W. J. & Hatt, A. The Shell Structure and Mineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates 77 Text-figures. 1969. £4.50. Haynes, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur) 1962-1964. (In Press). al - (22). Printedin England by Staples Printers Limited at their Kettering, Northants, establishment a THE SHELL STRUCTURE AND MINERALOGY OF THE BIVALVIA Il. LUCINACEA —- CLAVAGELLACEA CONCLUSIONS J. D. TAYLOR W. J. KENNEDY A. HALL Aah Mos Cc Cy ay NAT. HIST. BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 22 No. 9 LONDON: 1973 iit SHeELeE STRUCTURE AND MINERALOGY OF THE BIVALVIA II. LUCINACEA — CLAVAGELLACEA CONCLUSIONS BY JOHN DAVID TAYLOR WILLIAM JAMES KENNEDY Department of Geology & Mineralogy, University of Oxford ANTHONY HALL Department of Geology, Kings College, University of London Pp. 253-294; 15 Plates, 33 Text-figures, 22 Tables Eo RT ES, ‘ oN WAT./Hist, ~ - 2 MAY ‘973 | itieaeatat OGy ist BULLETIN--OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 22 No. 9 LONDON : 1973 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted im 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become veady. 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. 22, No. 9 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), 1972 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 16 March, 1973 Price £4-50 PEE SHELL STRUCTURE AND MINERALOGY OF THE BIVALVIA II. LUCINACEA — CLAVAGELLACEA CONCLUSIONS By J. D. TAYLOR, W. J. KENNEDY & A. HALL ABSTRACT THE shell microstructure of twenty six remaining bivalve superfamilies is described with the aid of acetate peels and electronmicroscopy. In the order Heterodonta most shells consist of outer crossed-lamellar and an inner complex crossed-lamellar layers. Three superfamilies, the Lucinacea, Tellinacea and Veneracea have all, or some members, with an additional outer layer of composite prismatic structure. Minor variations consist of the occurrence of homogeneous structure in many families, resulting from a reduction in grain size and loss of crystal form in each structure. The order Myoida is more varied with crossed-lamellar and complex crossed-lamellar shells in the Corbulidae, Gastrochaenacea and some Pholadacea. The Hiatellacea are mainly all homogeneous but Panopea has a three layered shell consisting of an outer simple aragonite prismatic layer and middle and inner homo- geneous layers. A very similar outer prismatic layer is found in some Pholadacea. In the Anomalodesmata two shell structure conditions are found, either a three layered shell, consisting of an outer layer of simple aragonite prisms and two nacreous layers or two homogeneous layers. Twelve shell structure characters can be used as an aid to superfamilial classifica- tion; but they must be used in conjunction with other characters and geological history. The shell structure characters have been superimposed upon a phylo- genetic tree derived from many characters; the points of variance and similarity of shell structure with this phylogeny are discussed in turn. It is suggested that the Arcoida may be more closely related to the Heterodonta than the Pteriomorphia. A ‘pholadomyacean’ stock has been in existence since the Ordovician and it is probable that both the Myoida and Anomalodesmata may be derived from this stock. CONTENTS Page INTRODUCTION . : : ; é A A 6 : : 256 SYSTEMATIC DESCRIPTIONS. F 5 a 5 a 0 6 5 256 LUCINACEA : 6 3 5 Fi z Fi , a A 256 CHAMACEA 3 : : : ; : : : , : : 258 LEPTONACEA . 4 , 3 f 5 4 0 6 0 5 259 CHLAMYDOCONCHACEA 0 ° é : : 3 ¢ , : 259 CYAMIACEA 5 : 0 5 9 : Q 5 : : - 260 CARDITACEA . ; : i F : : i : : : 260 CRASSATELLACEA : s A 3 . 5 0 0 é 261 CARDIACEA F 5 : : F . a é 3 é 263 TRIDACNACEA . a 0 a e : 2 . : : 3 264 256 TAYLOR, KENNEDY AND HALL CONTENTS—Continued Page MACTRACEA . A : 5 “ 2 5 c 5 . , 265 SOLENACEA 9 5 . . 3 5 5 . : F 265 TELLINACEA . é 6 - . A 6 ° ; “ 266 GAIMARDIACEA. 5 : 5 4 5 5 ; 6 : 6 268 ARCTICACEA . , : F 5 3 : a : 2 268 DREISSENACEA . z c ; = 5 5 : : : 268 GLOSSACEA 4 5 : D 5 5 ‘ . 5 ; : 269 CORBICULACEA . : 6 5 : 4 5 5 : F 5 269 VENERACEA . B 5 p 5 0 0 Z fi “ : 270 MYACEA . . a : 5 5 ; 3 : F 5 F 275 GASTROCHAENACEA . 5 F 5 c - 2 2 c 277 HIATELLACEA . , 5 é a : , 5 6 a é 277 PHOLADACEA . ; : 5 0 ; Hi : . 0 ; 279 PHOLADOMYACEA 6p 5 : 2 5 0 5 : . 282 PANDORACEA . 5 6 0 5 6 5 S : “ 282 POROMYACEA . S : : 3 5 : : . 7 5 284 CLAVAGELLACEA a 0 : F 6 7 5 : i 7 284 CONCLUSIONS . a : . : a , 2 a : . 286 ACKNOWLEDGEMENTS . : 6 : : ‘ ’ ; . 292 REFERENCES . - 5 : 5 : 6 5 4 é r 292 INTRODUCTION This is the continuation to Taylor, Kennedy & Hall (1969) in which we reviewed Bivalve shell structure, mineralogy and shell formation, and documented these features in the superfamilies Nuculacea to Trigonacea. Here we describe the distribution of structures in the remaining superfamilies and discuss the importance of shell structures in the classification of the Class. SYSTEMATIC DESCRIPTIONS Since the publication of the first part of this work was published the Treatise of Invertebrate Palaeontology, Part N Bivalvia has appeared, which uses a system of classification slightly modified from that of Newell (1965) which we used previously. As the ‘Treatise’ classification will stand for some time we have used it for the arrangement of superfamilies described below. Sub Class HETERODONTA Order VENEROIDA LUCINACEA (Plate 1, figs 1, 2 & 5; Text-figs r & 2) Fifteen species of this family have been examined mineralogically and ten struc- turally. The shell is aragonitic throughout. Three main shell layers are present in all the species examined. There is an outer composite prismatic layer, a middle crossed-lamellar layer, which forms much of the hinge plate and an inner complex crossed-lamellar layer which is bounded by the 1ojONppy eued 49h] IoAe] IaAVy] suol}eAIasqgQ MM ——_ Jeuuy = S|PPTIN Jojng = A8oyer90ury, Ayyeoo0'T soroeds eoerysoAyy es ap pe aemsud __arpamiyl _agyamg—aprmsud eee pring suusrid od4j-Teoer}soATY OeUISTIg HOY, +-pesso1a xejduiog -passorg aysoduiog = oytuoseiy —saqjayoAhos Durdasy DIYDPOD I9Ae] I9UUT 9Y} UT se~Nqny es1v] peie}ze0s pue suistid odA7 oryeurstid Ieyeurey, ~reyjewe, siyeursiid oytoeg (snoeuury) -[eoeIjsOAUI JO spueq UIY], Uses jON ‘uly, -pesso1o xefdurogj -pessoig oysoduiog )§=—s- ay UO StI -opu] nyojoundg v1ywpoD iojonppy ened IoAR] Ish] IoAP] suol}eAIOSqO mS rauuy = eTPP Jajng = ASoTeIOUTT, AyyeooT satoadg eIeISOAT VaHOVNIONT I alavy roqoNppy Tene take] 1akP] IaAvy suoljyRArlasqQ —————_, Jeuuy) =: PIPPIN qajng = ASoyerauryy AqyEI0T saroads PoRsoAW oneusud Ipyawry, aryemry, = = 9syeustrd —_ ‘ary, -passoio xajduioy -passorg aztsodur0d ayluosely ‘ds vatsvdy T | oreusud qeyeurey « reyewre,)=— a eusud (Aaqstid) — ‘uty -passoro xajdurog 9-passoig aysodmog a}mosery Jopenoy DUDY) DISMIINT oneusud Iepawey ieyewe, = soneuwsiud (qurT) ae ‘aIy. -passora xajduiog -passorg aztsoduiog ayuosery PYPIN sypurony sagrsoT oneurstid qeyjewry iepawe, = =oyeusud (aAaaq) _ ‘ary, -passoio xajduioy =-passorgQ «ayisoduiog «= ap uOSery Jopenoy vaudnga vpjao1awarq oneuisud qeyjeurey aeyaurey = a euIstad (aaaayy) } _— ‘ary -passoio xajduiog + -passoig aysoduiog = aytuoSe1y —sayjayoAag suasazaup vuaza ; oryewusud reyawey = reyjaure, = ayeursud. Jerang : _— ‘ary. -passoia xajdwiog -passo1g apsoduiog azmosery sorpuy 4seq pyoraquuf s1q4oD JaAv] Jauut o1yeusud oy} ur sreqjid peorzjsoAur ‘padojaaap aeyemey = reyemel, = o1yeursud purysuq Aapuezy 2 saqi0.¢ jo yuourdoyaaap aatsuayxy uaas JON -TIAM -passora xajduiog ~-passorg «aysoduiog = aq tuo Seay ‘Aeq 10], vjopun sisdouonT | roAP] sty} yo sjred yeursrew ay} jo [Ng ay} wL10F 1aAey] JauUT ayy ut suistd ad{4-peoes0Aur oneuisud aeyawe, reyeaurey = dteurstud (snaeuury) yo JaXv] aAtsua}Xy uaaS JON, ‘uly -passoio xajdui0y ~-passory © ayisodwog «= apruoSery ApMyey Sypadog DULINT JaAP] Iau ayy oreusud ur suusud adAq-;rorrjs0Aur padojaaap qeyeurey, «reyjaurey = 91yeustd uvdef adda Jo Juauidojaaap aatsua}xy uaas JON -II@A\ ~passoro xajduioy -passory aztsoduiog §~=— a TuoSeIy eUuIIYysO oid nummy oquin yyeaueq sustud oneusud qeyjawe, =reyeure, = oyeursud oaulog: WIS ad4q-feorysofu yo s1aheT uaas JON ‘um. -paesso1o xajduioy -passoig §ayisodmoa ajyuosery eMeIES sisuarhyf purnT JaAz] Jouut ay} ur suisud ad44-;eoe4s0Aur ojemsud qeyewrey «reyjawey, = s1yeursud (yur) jo staXv| JuaUTMOI = uaas JON QO. -passoro xajdwoy -passory apsodmog oayuosery apie, odeg vyjaquinjoo nuranT aoueLy ; 4 “IaIssOIy JaAv] JouUT 9y} UT suistid adAy o1yeustid deyewe, ieyawey, = o1yeusuid arresyea (yoreure Tq) -]eoeI3s0Aul Jo spueq AULT —uaas 30. ‘uly. -passoio xajduioy ~-passorg © ayisodmiog = ay uo Sey ‘aua00y «= MSOPJ9IUD) DLAQuELT yuasaid aq Avur sginqny, “reAP] Jouut ay} (Ausiqio,P) ur suisud ad.43-JeoeysoAur ornewusiud qeyamry =reyewe, = oneursud vynanstaponb JO SIaAP] DAISUa}XY] —UaaS FONT ‘uly, -passo1a xajduiog -passory aysodmog apuosery — sarpuy “Ay Dya21awary so]nqn} parajyeos pure suisud od43-;eoe1ys0Aur Jo ueq auy SUIe}UOD 71 aTayM ‘spuolser [euoquIn ay} Aojaq padojaaap ysaq st arnjona4s Iejjame] passor9-xajdui0o ‘z9Xe] JoUUT ay JO FING ayy oneusiud qeyjawry =reyjewe, = oeustid (snoeuury) pring susud ad43-jeorjsoAy oneuisi1g WML -passors xajdwmoy -passory aysodwoy aytuosery — sayjayofeg vurdaasy DLyoporD roAry JAUUT 8Yy} UT sarnqny aS1r] peso}}P0s pue stustid adAy oneusud reyeure, reyawey, = atyewsud oye (snoeuury) -eov1}s04ul Jo spurq uly], uaas joNT ‘ul. -passo1s xajdwog -passory aysodmoy oz1u0Sriy -opuy viojaund viywpod ioyONppy ened rode] tary] aaAey] : SUOIVBAIaSqQ —— ,—___Y- Jouuy = a ppy JaynQ = Adoyeszsuryy AqyeooT sorads roeI4soAyy VAOVNIONT ‘ I alavy " SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 257 trace of the pallial line. The outer composite prismatic layer is usually very thin but forms the ribs when present. The first order prisms lie with their long axes arranged radially to the umbo (Plate 1, fig. 1). In the middle crossed-lamellar layer the first order lamels are arranged with their long axes concentric to the shell margin. The lamels are often very fine when compared with some groups such as the Arcacea or Limopsacea but similar to those in the Carditacea and Astartacea. A prismatic pallial myostracum is present in all species examined, although variable in thickness. The inner shell layer is the most variable; complex crossed-lameilar structure is present in all species (Plate 1, figs 2 & 5) but varies in extent. The rest of the inner layer is made up of myostracal prisms which may occur as sheets, which alternate with the complex crossed-lamellar structure (Lucina fijiensis) or as large blocks (Codakia tigerina). These sheets of myostracal prisms may represent periods of temporary mantle attachment. Scattered large tubules are present in the inner layer of Codakia punctata; tubules are also present in Divaricella quadrisulcata and Codakia tigerina. CIR agg SS NOMA AIR Uy WA Fic. 1. Radial section of Lucina columbella. CP = composite prismatic, CL = crossed- lamellar, CCL = complex crossed-lamellar, PM = pallial myostracum. Fic. 2. Radial section of Corbis fimbriata. CP = composite prisms, CL = crossed- lamellar, CCL = complex crossed-lamellar, P = pallial myostracum. At 258 TAYLOR, KENNEDY AND HALL CHAMACEA (Text-fig. 3) The shell structure, anatomy, and evolution of the Chamacea have been discussed in some detail elsewhere (Kennedy, Morris & Taylor, 1970), as has the shell structure and mineralogy of the anomalous species Chama pellucida (Taylor & Kennedy, 1969). Amongst Recent species the shell is aragonitic throughout, with the exceptions of Chama pellucida and C. exogyra, which contain substantial amounts of calcite in a distinct outer layer. Two fossil species Chama gryphina (Miocene) and Chama hauert (Turonian) show traces of calcite but there is no evidence that this was an original feature of the shell. In all wholly aragonitic species examined, two main shell layers are present (text- fig. 3). There is an outer, crossed-lamellar layer and an inner complex crossed- lamellar layer bounded by the trace of the pallial line. In the outer layer the first order lamels are arranged concentrically to the shell margin; in many specimens part of the outer layer may be missing apparently as a result of abrasion. Myostraca are very well developed in the Chamacea, the pallial myostracum is thin, although always distinct and readily recognizable, extending throughout the hinge area. The adductor myostraca form thick pads and often interdigitate with the inner complex cross-lamellar layer as a result of slight shifting of attachment and body position during growth. In Chama iostoma other myostraca were seen presum- ably associated with the pedal muscles. The inner shell layer of the Chamacea is basically formed of complex crossed- lamellar structure, varying from fine to coarse textured and complicated by sheets of myostracal type prisms and myostracal pillars. These features show varying development both inter and intra specifically. Myostracal pillars often arise from Fic. 3. Anterior-posterior section through Pseudochama vadians. AM = adductor myo- stracum, PM = pallial myostracum, CL = crossed-lamellar, CCL = complex crossed- lamellar. SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 259 pallial and adductor myostraca. In Chama spondyliodes myostracal pillars are also present in parts of the outer crossed-lamellar layer. Seen on inner shell surface myostracal prisms are often arranged in rows radial from the umbo. The Chamacea are markedly tubulate but there is much variation in the abundance and distribution of these structures which are usually confined to the inner layer. Chama pellucida and C. exogyra differ from wholly aragonitic forms in possessing an outer prismatic calcite layer of unusual structure described by Taylor & Kennedy (1969). The remarkable occurrence of calcite was first noticed by Lowenstam (1954) but he considered (1964) that the outer crossed-lamellar layer of tropical species had changed to calcite prisms in the temperate C. pellucida. He used this example as evidence in the general temperature/mineralogy trends he demonstrated for invertebrate skeleta. However other temperate species of Chama seem to be wholly aragonitic. A table of the shell structure characters of the thirty species of Chama examined will be found in Kennedy et al (1970, p. 390). LEPTONACEA Most species of this superfamily are very small and thin shelled. Four species were examined mineralogically and by electron-microscopy. The shell is aragonitic. Two shell layers are present in all species examined; an outer crossed-lamellar layer and an inner complex crossed-lamellar layer. The two layers are separated by the prismatic pallial myostracum. TABLE 2 LEPTONACEA Species Locality Mineralogy Outer Inner Pallial layer layer myostracum Kellia suborbicularis Britain Aragonite Crossed- Complex crossed- Prismatic, Montagu lamellar lamellar thin Kellia pustula Indian Ocean Aragonite Crossed- Complex crossed- Deshayes lamellar lamellar Scintilli owent Karachi Aragonite Crossed- Complex crossed- Prismatic, Deshayes lamellar lamellar thin Scintilla rosea Indian Ocean Aragonite Crossed- Complex crossed- Prismatic, Deshayes lamellar lamellar thin CHLAMYDOCONCHACEA Lack of material prevented examination of specimens of this peculiar monogeneric superfamily which have internal shells contained within mantle sacs. 260 TAYLOR, KENNEDY AND HALL CYAMIACEA Three species of this small superfamily were examined structurally and mniner- alogically. In all three species the shell is aragonitic and consists of two layers both of them of fine granular homogeneous structure. The layers are separated by a thin prismatic pallial myostracum. TABLE 3 CYAMIACEA Species Locality Mineralogy Outer layer Inner Pallial layer myostracum Cyamium antarcticum Falkland Aragonite Homogeneous Homogeneous Prismatic (Philipp) Islands Cyamium laminiferum Antarctic Aragonite Homogeneous Homogeneous Prismatic (Lamy) Neodavisia cobbi Falkland Aragonite Homogeneous Homogeneous (Cooper & Preston) Islands CARDITACEA (Plate 2, figs 4, 5 & 6; text-figs 4 & 5) Fifteen species have been examined mineralogically and eleven structurally. The shell is aragonitic throughout. Two main shell layers are present, an outer crossed-lamellar layer which forms the bulk of the hinge and teeth, and an inner complex crossed-lamellar layer which is bounded by the trace of the pallial myostracum. The first order lamels of the outer layer are very fine and are mostly arranged concentrically, but in some species a reflected shell margin causes the lamels to appear to have a radial alignment (Plate Fic. 4. Radial section of Cardita vdviegata. CL = crossed-lamellar, CCL = complex crossed-lamellar, PM = pallial myostracum, T = tubules. S9[nqnj auy pue spueq oneur “slid ou ‘seqpid TeorijsoAur _2uy_yuepunge HIM ‘papueg _ ST IoAR] Te][9ule]-passo1o euL ‘ToAR] JoUUT 24} UI Juasard ere Surpueq yusurM01d pue sornqn} ourz ‘ouy AoA SI ade] TET[PUIL]-passoso ayy suol}eArasqgQ Poe IISOATY, VaOV.LIGUVD b atavy oryeurstid Telfourey Waas 307 ‘UWI -passox5 xe[duro5 AET]PwE]-passory auOSery UrewIg — vyyjs o1zeurstid TeyTourey wees Jon, ‘UIUL -passora xe{dui05 TelEwMe]-passory ouu0Sery POY "Syma, o1eustid Ieypourey ureqrig w99s JON ‘UIYL -passozp xo[duio5 7eT]ewE]-passor5y eU0Ser1y ‘ous00q oryeurstad Teyjourey woes Jon, ‘UIYL -passo1p xe[duo05 Te]Tewe]-passox5 971W0S1y sour ‘uaas 40) NO UIYL ~pessoro xajduro5 TeT[aure]-passo1y oyUOSe1y Teypourey puryeaz mon 498s JON uaas Jon, ~Passo19 xefduro> TeT[oue]-passor> a}UOse1y ‘puryyjony Toyonppy Teyreg Se IaAPJ douuy IaAey 19309 ASoperourpy Ayteoory (e1raurerg ep) onunyd sooraua A (211turerg ap) IUYIUOI vYyvIay T Aqiamog BIDIINS vyYpADD Premyory DIv2INID wypADD puesta 8 —aapaa ap yoreureT SYVAISNDY DYLpADD satoads onrusud reyaurey (aqrauretg, ap) uaas ON ‘ary -passor9 xajduroy aeyjeuref-passory —axtuo Sey Urey wypjsonund oneustud aeyeurry (eptaurelg 8p) iN uaas JON, ‘ung -passor9 xajdwiog = IeTJauIEI-passoig —_ ayTUOS ey ROLY 'S —_ByVAIULYILOI DIVIIYT, oneusiud reyawey Urey uaas JON, ‘aly -passo19 xajdwioy IeT[auIE[-passory «= ayTUOA IY ‘qua00q orjeuisud APyaurey uaas JON ‘uly, -passo19 xajduiog =re]Jaure]-passo1z ayuosiy sajnqn} auy pue spurq oneur -sud auy ‘sreqid peorsjsoAur aouRLy euy yURpuUNge YIM ‘papurq eee — Ayjourstp st aXe] Jouut oy. poo -auy Joyyer st roAR] Ja][oUeE] oneusud qeypouey TAISSOIS) -passo1o ay} JO aINjoNIys ay, waes JON ‘Uy, -passoro xajduiog = Ie]JaUIE[-passory = ayTUOR SIV OINRITED vyparaquer vip, sajnqny auy yuepunqe YIM ‘papueq Ajsuryiys st 1Ae] touur ayy, ouy Jayjzer st taAey Teyjawey oneusud qeyaurey 2M) -passord ay} JO aINjon4A}s ey, uaas JON, ‘uly, -passo19 xajdwog re[jaure,-passo1y = ay UO ey sopenoy suuffo vaaunpipana sroAP] YO Ul Sarnqny aunt -auty [ered ay} apts}no sake] 4ajno ay} yo sow ynoysnoryy quasaid osye are sreqpid asay.y, “raAv[ IAUUT ay} MOTAq SaTT Stq} a1ayM IaAP] 19}N0 ay} JO qied sou ay} ur pue ‘1aAPy] qouur ay} ur sreyjid peoesys oreustid TPTOWP] (exgin3nag) | -oAul JURpuNqe aie alay], eas JON ‘ul. -passoio xajdwioy sejjeurej-passory = a TuoS ery eAuay DIDFaADA DIYPADD JaAvy Iauut ayy ul juasaid are saynqny our “raAR] 1ayno ayy jo spred peurs -IPUI 9Yy} UI May B pur IoAP] aouut ay} ur sreqid jeorys -oXu yuRpunqe are aroy ‘auy Jayzer st JaAvy repjawey go1geusiud Teyjawey -passoro ay} JO a1NjONI}s ay], Uses JON ‘uly. ~passoio xajduiop © reyjaurey-passorg §=—_ aytuoSe1ry wqaamos vy daXe] auut ayy ur sajnqn} euy pue sraAey wistd auy Jo sa0R1} are a1oy ‘auy Jayzer st Jake] reyjaurey Te][ouLe] u Passor9 ay} JO aInjon4ys ay, aes JON UaES JON -passo19 xa[duioy Ae[[aurE[-passory ayuoSery —pueysuaand DIAOWUADUE DILPADD, JaXe] Jou! ayy Ur saynqny auy are alayy, = “auy 1oyyeI st Teyjourey y SIAL] Y}OG JO eINJONAYS AY], Uses JON as JON’ -passoso xajdwoy szejJauE]-passory ay UO purjsuaan?) DIDSSDAIUL DIL JoAv] JeUUT ay} Url saynqnz yuepunqe aie alay, “auy 1ay3RI st oneusud TeT[PUIE] s19AP] Y}Oq JO aInjonyjs ay, uaas ONT ‘uly, -xajduioo passo1y Ae[Jawe]-passory = aytuoSeay epHuop.y Dui sajnqn} auy pur s1aAry] sustuid adAy yeoexjsoAur ‘guy 1ayzeI ‘pasa} -yeoeIySOAUI JO -jeos ‘Burpueq yuourmoid spueq pure sreqid sey IaAP] Ie][auIe] passo19 yeoerjsoAur -xojdmoo ay} ‘auy Araa oryeuisuid YFEM Tepe] arginsn.ig St 19Av] Ae[[aUIe[-passoro ay, uaas ON ‘ur. -passoro xajdwoy reyjaure-passory = aytuoSery uea0gQ uerpuy Dppjnaiyo2 vip daAe] Jauut ay} Ur juasaid are saynqnz aulq “19AP] 1e]Jaure|-passo19 xajduos ay st se ‘auy A10A oewsud Ie]jawey St JoAV] Iv[JaUTE]-passoso ay, aes ON, ‘uy, -passo1o xajdwioj reyjauref-passory = ayuoSery puelSuq Many Sypadog DIY ‘zoAP] JoUUT ay} UL yUdSaId are Surpueq juautuoid pue sajnqn} aurq = ‘auy Aaa Te]JoUy puryesz Man St 1aXv] [JaurE[-passoss oy, Uwaas JON das JON, -passo19 xajdwiog 1e[Jaure]-passorg ay UOS ery ‘pueppny syvajsno << zoyonppy — eed suoljeArasqQ, ——— rahP] Iauuy aoe] 19jNQ = ASoyesouryy Aqyeoo"y PoRsysoAyy VaOVLIGUVD b alavy 10790PPV ened suorzeAtesqO ~~ roAvy IauUT roAv] JoynQ Asoreroury, AqryeooT satoeds BORIISOAT| *sono[y) ‘AoTqOOT (Aus31q10,P) opyuoseiy ‘sery “T uxanb ajanjs ACIATIIBOV ayuose1y purusei5 widuyya aanjsy pero) Jah] IOUUT ‘1e[[aUTe[-passo19 ‘uy ONT Ajeuy Artoa JoXe[ I9ING = Usaesj}ON ‘4oUTsTpuy snosusSowofF{ IeT[aUre[-passorg opuoSery eyeNsny My suardisap vjjajwssvAD Surpueq snonordsuos yytAA éoryeurstid JoAe] IOUUT ‘IeT[oUTe][-passo.1o “ary CUCEREL Ajouy AiaA JoAe]T 103NG uses JON ‘JOUTFSIPUT snoauesouloyy = IeTEWIL[-passo1g s}UOSeIy SOIPUT “AX wUiNADIIYUY YIJAZVSSVAD 10;ONPPY ened suol}yeAtasqO ———“———“ IoAR] IOUUT roXkeq JoynQ = ASoyes9Uty, Aqryeo0'T satoads eoei}soAT VaOVTITALVSSVYO ¢ aldvi suoIzeAIESqOQ, suisud adAq-jeoesjsoAur jo 4[1nq Ajaruea jsourye 19Ae] Jauuy stustud adAq-jeoe1}s0Aur yo 4[INq Ajarjue ysouye JaXey Java susud ad4j-[eoesofur Jo 3INq Ajarjue ysourye IaAe] Jouuy uolger euoquin UI aINjJONI}S Ie[Jaure, passo19 -xajduio9 YIM Sutrasuy-19yur ‘suistid adAq-[eoe1jsoAur jo #1INq Aysow ssXvy s9UUT dav] Iauut ay} FO ysour uLroj sursiid adA}-peoe.1s0Aqy sauoquin y}yRauaq Ajuo quasaid arnjon4s Teyjaurey-passoio xajdui05 Ajuo sauoquin y}eauaq yuasaid ainjonzjs snoauasowoy ‘suisiid adAq-peoerjsoAur o/3pinq Ajasre] Jaf] Jauuy SpreMut pers} UayA\ snoauasowo0y Surmosaq ‘re[[awe[-passo19 Ajauy Ara ra€ey 19309 daAe] Jauut ey} Ul Suipueq JueuTMOIg Surpueq snonoidsuos yyw JaXe] Jouut ‘re][aure]-passor9 Ajauy Ara 1AP{ 10309 surpurq snonoidsuos yy JoAP] JOUUT ‘reT[PtUE]-passor9 Ajauy Ara Jakry 193nO suoRAIaSqG 40j9nppy renred —______., ————' vorrysoAyy ornemsud — “UNL orneusud — “ary oryeustid = ury} oeusud “ary uaas JON “JouNjsIpuy oreusud uaas JON ‘ory goryeustid ‘uryy uaas JON ‘JOUT}SIPUT orpeursud = ‘ug orjeutsiud ‘UI oeusud ‘uy uaas JON uaas JON uaas JON uaas JON goryeumsud “uryy uaas JON ‘JOUT|SIpUyT goryeusiid “aryy uaas JON “JOUTSIpUy Jojonppy eyed © porlsoAyW raXry] 1auUy suisud jeoeiysofur Tele -passor9 xaydui05 susud yeov1ysoAur YPM reyaurey -passoio xajdui0>, suisud jeoeysofur UII snoouasouroyy suisud jeormsofur YIM reyaurey -passo19 xajdui09 susud yeoesysoAur YIU snoouasouloyyT suusud eors}soAur YIM Ieyjourey passo19 xajduio5 suisud TeorsysoAur YIN snoauasowopy snoauasoulo fy snoauasowoyy snoauasoulofy snoauasowopy snoauasowoy{ iaXrv] Jauuy sake] 1aynQ ASolerauTy, Ie[[aure|-passo1y APT[PUIE[-Passo1g Teyjaure[-passoig re[jeure]-passory Ie[[aure]-passory Iejjaure[-passo1y ie]Jaure]-passo1g re]Jaurey-passo1y Te][PUIe[-passo1g Ie][aure|-passory Tey[aure|-passo1y Ie][aULe[-passo1g rary] 1030O VAOVTTIALVSSVUO ¢ alavye ayuosery oyuosery ayuosery aymoseiy oymmosery aymosery ayuosery oyluosery ayuos1y ayuosery oyuosery ayuosery aywuosery apuosery ayluosery ayuosery opuoseiy ASopeso ur Aqre00T *sonoyy, ‘AappIO lL ‘sery "T purjusaisy, epHolt aouely ‘Avloued, ‘mene yT aouRLy ‘Aeroureq. ‘UeTZOyNT pod mt Ayer] ‘quad0[iq. purruee5 ureyagy ‘quacol[d ureyagy ‘quas0l Ayes] ‘qua00ld purjuasr5y sopenogy aoueLy ‘Avroured, ‘uerayny erpeaysny “AX sorpuy “AA Ayyeo0'T DINU] DIJIUISSVAD, DIDUOSIA} D]]9JDSSVAD, DIDIDILP B]JaIVSSVAD saroads (Au31q10,P) uxenb aanjsy Aeraysoryy dye aanjs ¥ peiu0) yorewey saAeysaqy (e3s09 ep) ; DIDIINS IAVIS (23809 ep) DyvIjns OUADIS : Aqiamosg DIDAIS PADIS (exarquof) 2Y DUO BpADIS AqiaMos wnbyqo 3janjs soAeysaqy DIDSSVAIUL 9JADIS (zayoerurnyos) SYD9409 BfADIS (Aqaamos) DS0GQ1G DIJI9{VSSDAINT (Aqaamos) DIVIPDA DI] 3VSSDAD yoreue’y DSO[JIUD] D1J2DSSVAD EXCL G suardisap vjjajwssvag 9A0a)T UNADIJYUD 1119ZDSSVAD, soroads we SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 261 2, fig. 4). The outer layer is frequently very thin or worn off in the umbonal area. A thin discontinuous prismatic pallial myostracum was seen in most species. In Cardita calyculata the prisms were seen in the umbonal area only. The inner layer is somewhat variable; the complex crossed-lamellar structure is rather fine and prominent banding is seen in most species (Plate 2, fig. 6). Myo- stracal pillars are developed in several of the species examined: in Cardita calyculata and Venericardia vmbricata they are restricted to the inner layer. In C. sowerbyi & C. marymorea there are abundant pillars in the inner layer and a few in the marginal parts of the outer layer (Plate 2, fig. 5). However in C. variegata there are abundant pillars in both inner and outer layers. Sheets of myostracal prisms are also present in the inner layer of C. calyculata, C. marmorea and V. imbricata. Tubules are present in all the Carditacea examined and occur in the inner layer of all species and in both layers of Cardita variegata. CRASSATELLACEA (Plate 1, figs 3, 4, 6 & 8; Plate 2, figs 3 & 4; text-figs 6 & 7) The superfamily Crassatellacea is represented by two living families, the Cras- satellidae and the Astartidae Sixteen species have been examined mineralogically and twelve optically; the shell is aragonitic throughout. In both families the shell consists of two layers; both have an outer crossed- lamellar layer but the inner layers differ. The inner layer of the Crassatellidae is homogeneous, whereas that of the Astartidae is largely built up of myostracal-type prisms, with only traces of complex crossed-lamellar or homogeneous structures. The outer crossed-lamellar layer of all the species examined is built up from very fine primary lamels, arranged concentrically with the shell margin (Plate 1, figs 3 & 4). The lamels are obvious in the outer parts of the layer but traced inwards they Fic. 5. Radial section of Venericardia imbricata. CL = crossed-lamellar, CCL = complex crossed-lamellar, PM = pallial myostracum, T = tubules. 262 TAYLOR, KENNEDY AND HALL become increasingly difficult to resolve and the layer appears homogeneous. Most of the hinge area is formed from crossed-lamellar structure. The inner layer of the Crassatellidae shows no obvious macro-features other than conspicuous banding; electron-microscopy shows the structure to consist of irregular granular structure. In the Astartidae the inner layer is largely built up of myo- stracal prisms (Plate 1, figs 7 & 8; Plate 2, figs 2 & 3). In Astarte borealis scanning microscopy shows the outcropping prisms (Plate 2, fig. 3) revealed as distinct bosses; these show surface features of parallel ridges and striae. In A. borealis and A. incrassata there is a narrow homogeneous sheet on the inside of the pallial myo- stracum separating the myostracum from the main prismatic part of the inner layer. In other species the prisms arise directly from the pallial trace. There is consider- able geometric selection of the prisms of the inner layer; those closest to the pallial myostracum are small and numerous but traced towards the inside of the shell there is a reduction in numbers and a resultant increase in size of the prisms. The two shell layers are separated by the trace of the pallial myostracum. In the Crassatellidae this is marked by a distinct unconformity of growth lines but actual myostracal prisms have been detected in only one species. Eucrassatella gibbosa which is also the only species examined with a distinct prismatic layer associated with the adductor muscles. In the Astartidae all the species possess a prismatic myo- ct So) Tain, ani cae f pp Il pl: a ~ > mw / | PM H- } C Fic. 6. Radial section of Eucrassatella gibbosa. CL = crossed-lamellar, H = homogeneous, PM = pallial myostracum. Fic. 7. Radial section of Astavte borealis. CL = crossed-lamellar, PM = pallial myo-stracum, MP = myostracal prisms. (Aqiamos) ayu0se1ry Jopenoy wnsoryuas wnipAévacyoDA TL (TessT) ayuosery — sayjayoAeS asuUarZzans WINIPADILOADT e gouely oeusiig }OUTSIPU. -pesso.to xadwiog = TeTawWL]-pesso1D apuose1y uleyIg DIDaNID VIPADIOYJUYI F 10,9NPPV ened suorzeAIasqO | ——_“ roAv] JoUUT rar] JaynQ Asoye1oUT, Ayyeo0T, satoads voriysoAT] 10}9NPPV Temled suol}eAIasqO —_ aoAR] IOUUT rahe] JoynQ ABO[eIOULN, Ayppeoo'T satoads vovizsoAy Vaoviduvo 9g a1aVL oyewsud ‘ug NWeUISU dVeUISTIg — qoursipuy oquin yyeauaq oneustiud spurq jeoes3zsoAur uty y = ‘ULL adury ay} Sumoy 91N}ON44s Ie|[aUle|-passor9 auy ‘Wo; paljays-uIy VY _ oryeulsigy oryeursiud = “UL oyeursud ‘uy. onewsud = “Uy - yourjsipuy TaAP] 193N0 ay} JO S[awe] Japs10-ysIy, ay} YQIM Ajinuuos ur aie 1aAP] Ie[[aure[-passo19 xajduroo ay} jo seg _ oryewisiig _ qourysipuy o1yeusud a “url onewsud = ‘UryL oeusid o1VRuIst “un IVeUSg JOUTYSIpUy royoNppy 1eured oH _ ———— BIPIISOAT suOIRAIASqO iojoNppy suoqeAlasqO BORIISOAT ajiuosely ayiuosery ayuoseiy aywosrry ayuosR1y -passoio 3 Ie[[aulL[-passo1g ayUOSeIy IP|Jawry passo19 xajdwog re]Jawel-passory = aytuoseIy reT[aUIey -passo19 xajdwi0p ©=TeTJaUIe][-passory — ayTUOSeIY IP[[PWey] -passo19 xajduioy Ie]JaUIL[-passory — ayuOS ey Ie] [eure] -passo19 xajduog ©re[Jaure[-passory ~—-aytuOSeIy IP][aUIE] -passo19 xejduiog = 1e]JaWIe|-passory a} uOSeIy IP][AUIE] -passoio xajdwiog =TeTJaUIe|[-passo1y — ayuOSeIy IeT[aurey -passoio xa[duiog = e][aWIE[-passory — aytuose1y Aepjaurey] -passo19 xajdwiog = e[Jaure[-passory = ayuosery Teyjourey xajduio) §rr]jawr|-passo1g oyu ery Ie][aur] -passoio xajdwog Ie[[aUIe|-passory — aytuoS ery ae]jawe] -passo19 xajduiog =e|JauIe]-passor1y — aquos ery Teyjawey -passor9 xajduroa aUIe]-pessoly a}uose1y Ie|[awey] -passoio xa[dwiog = 1e]JaUIe[-passory = ayTUOSeAy Ip]jaue] -passoio xajdwio9 «= re[JaurE[-passory = ayTUOSeIy daXey Jauuy aaXvy rayng = Asoyesouryy Jake] JouUT Jake] Jayng = Asoyessulyy VaOviddvo Q ATAVE iopenoq satfayoAag aouRly ‘Aesraureq ‘aerqjeyny purjusain ureyuag ‘gua00jzsla[q PITUR TY sopenoy satpaypAas Jopenoy purjuaaisy ‘ST “Oost sayquy voreure [ Avg urqnd satjayoAag Pury pury urepug ureyagy ureyigy Aypeo0T Aqyeoo'T (Aqiamos) wmnsoryuas UN wpavqAyIVA T (1essI) aSUatZaNS WNIPADIADT (yoreweT) uennbyqo wnipav20x07T snouqe.y wMNDYII WNIPADD, (Aqiamos) mosuryand pipav20yzUno y (aaaayq) UNJDUADIUL WUNIPADIYSD A (3u04S 3 ule]}19}) aSUajSOIOUDNS VIPADIOUOTA T, (poo) MNsOpnNIDUL WINIPADIAYIVA T, (Aqiamos) SAOSUOI WUNIPADIAYIVAT. (x91101N) snaipunjuaods sadiadas (Aqitamos) nnaisurd vapiaddog (snaeuuryq) uinqoadas WanIpAvILaDT (ure) WNSSDAD WNIPADILAID T (Aqaamos) aypajsny WinipAvrrnaDT (Aqaaaos) MnJoUdayY UUNIPADIDT (snaeuurq) vipavqiuay vIpAvIVUA FT (snoeuury) OpaUn WNnsDAT (snaeuury) ajnpa viUuUAIpojSVAID (snovuury) DIVULYII DULPAVIOYJUDI (snavuury) DIVIINID VDIPAVION JUDIE satoads satoads “ae SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 263 stracum beneath the pallial attachment, but no adductor myostracum was detected. In a recent review of the Crassatellacea (Boyd & Newell 1968) a diphyletic origin for the superfamily was suggested. Our observations on the differences between the Crassatellidae and the Astartidae would tend to support this suggestion. The features of the Permian Oriocrassatella elongata described as “crater-like blisters in umbonal cavity” (Boyd & Newell, 1968) appear to be cavities left by the solution of myostracal prisms in the inner layer. CARDIACEA (Plate 3, figs 1-4; text-fig. 8) Fifteen species of this group have been examined structurally and twenty mineralogically. The shell is aragonitic throughout. Two main shell layers are present, an outer crossed-lamellar layer which forms the hinge and an inner complex crossed-lamellar layer which is bounded by the trace of the pallial line. In the outer layer the first order lamels are quite large and are aligned concentrically everywhere except for the hinge (Plate 3, figs 1, 2 & 3). Transverse sections show that the strong ribbing of most species of this superfamily produces complex patterns of first order lamels in the outermost part of the outer layer. This layer becomes very thin in the umbonal area and is frequently eroded and lost. There is a prismatic pallial myostracum in most Cardiacea, (indistinct in some species) which separates the inner complex crossed-lamellar layer. In Laevicardium alternatum the inner and outer layers are in direct local contact and the blocks of laths in each layer show structural continuity. Sections cut through the adductor muscle scars of some species show lenses of myostracal prisms. Bands of myostracal prisms associated with pedal muscles were seen in the umbonal area of Trachycardium consors. Fic. 8. Radial section of Cevastoderma edule. CL = crossed-lamellar, PM = pallial myostracum, CCL = complex crossed-lamellar. 264 TAYLOR, KENNEDY AND HALL TRIDACNACEA (Plate 3, figs 5-8; text-fig. 9) This is a small superfamily closely related to the Cardiacea (Stasek, 1962); included genera are Tvidacna and Hippopus. Four species have been examined structurally and mineralogically. The shell is wholly aragonitic. The shell is very thick, with two shell layers, an outer crossed-lamellar layer and an inner complex crossed-lamellar layer which is bounded in extent by the trace of the pallial line. In the outer layer the first order lamels are large (Plate 3, fig. 6) and arranged concentrically in all but the hinge area. The strong ribbing however causes an apparent complex pattern of first order lamels (Plate 3, fig. 4). There isa thin prismatic pallial myostracum in all the species examined. The inner complex crossed-lamellar layer is somewhat variable in character; in the three species of Tridacna studied the structural elements are fairly coarse and interleaved with thin sheets of myostracal-prisms. In Hippopus however the structure is very fine with an almost homogeneous appearance and with many fine prismatic sheets (Plate 3, figs 7 & 8). Higher magnifications (Plate 3, fig. 7) show that the structure consists of sheets of fine needles. All species show very strong daily growth bands in both layers and show prismatic pedal and adductor myostraca. LZ L Fic.9. Radial section of Tridacna maxima. CL = crossed-lamellar, CCL complex crossed- lamellar, PM = pallial myostracum. 2 ee (poo) ayuosery uofAa9 DIDIUNA Ua]OS oryeurstid Teyjawey Ieypawey — ‘UIY., -passo1o xefdu0g ~—- passo9 Ajeurg = ayruoSery iopenoq ‘ds wnbyis snoauasowioy oryeustid ay eT[OUTeT 0} Ieyaurey (1a]8ueds) — UY L ‘snoauesoul0yy ~—--passozo Aouryy ayuosery eIpuy snsonjon] snyjayng oryveustid oy eTOUIL] Te][owey (suoeuury) — ‘Uy LL ‘snooueso0uIoyY —-passo1o Apoury — oqtuoSery Ule}yU onbyis sisuy oryeurstid 97 e] [Sure] Ie]pourey (sneeuurT) = ‘ur ‘snosuasouloyy ~—--passozo Ayaurg oyuosery UrIe}Ig sisua sisusy 1opONppy Teed suoI}eAIISqG ——_——qY Jade] Jouuy JaAR] 19jnG ASoyerouryy AyypeooT ouleN eoerzsoA Ty ~ = = $= DOP TEL eoevsjsoAw jepod oijyeustid o1jzeustid Tey OUIeT ‘raAv] IOUUT UI spueq UISIIg UL ‘uly, -passoio xejduiog = IeTJauIe][-pessoIg = optuOSeIW uwe900 ULIpUT DUAXDUL DUIDPIA T. 2 yoreureyT vorijsoAw jeped orewsud oneustid Ieyjowry] ‘raAe] IOUUT UT spueq UWISTIg ‘UI, ‘uly. -pesso1o xejdwiog =IeyJaue[-passo1g = oy 1UOSey soIpuy seq 99049 DUIDPIA [. snoeuury oreusid orewstid Ieypourey "I JaAe] IOUUT UT spueq WSsIg ‘UIYL ‘uly, -pesso19 xejdwog = reyJaure]-passo1g == oy UOSeIy eyerjsny sndoddiy sndodqiy 10JONppy Teed suolzeAIesqQ —_,—_- YY JoAR] IoUUT JaAey Joyng §= ASoyeIOUTTY Aqyeo0'[ satsads eoviqsoAy, VaAOVNOVCIUL 4 alavy (poo) ayuosery uofag, DIDIUNA, UIJOS oryeuisuid reTaue] AeyaUrey — ‘ur, -passox9 xajduiog = -passoi9 Ajourg = a} !MOSeTy aiopenoy “ds onbyis snoauasowoy oneuistid o7e][aUIET 0} rTeToUTey (aaj8uads) _ “uy ‘snoauasou1oy{ + = - passo19 Ajaut.y oyuosery eIpuy snsonjav] $njj91jND. oneusud Eraicas| aeyawey (suaeaurq) _ “ary. ‘snoauasowoy{ = -passor9 Ajouny aymosery urewug. onbyts sisuz orjyemusud OPET[IUIET reyourey (snoevuuryq) — “UL ‘snoauasowoy{ = -passor9 AjauLy ayuosery urequg sisua sIsuay 10j9NPppY rented suolyeArasqO Fs daXey rouuy Jakep Jang = ASoyesrauryy AypeooT aweN > eoviqsoAy _— WHOVNATOS 6 aTavL idduga aqluosery ropenoy DIDIIA DIUABPOAJIV AT (seXeysoq) ayuosery ueasQ UeIpuy MeyADUE DYPADD, oquin y}yeaueq ayqisIA voryysohu [epaq ‘suisud adAj-jvorjsokul Jo sjaays oeusud IP[PUIET (snoewuury) WA ‘aug Artaa JaXe] Jauuy = Waas JON, ‘uly, -passor9 xajdwioy 1e][auWe]-passoigQ — aytuoseIy urezigy mpyos vjnsids rakv] gauur ayy ut stusud od&y onemsud oeusud reyjauey (pinoy) -jeoerysoAul yo spueq Urq] ‘uly.L ‘un -passoro xajduiog reyJaure]-passorg «= ay Tuo ery iopenoy DIDINPUN Vay qaAry] aouut oy} ur sumstid odAy oneusud Te[[aUIe] Aqiamos x diuapoig¢ -[Poe1}SOAUI Jo spueq UIYT, Uaas JON ‘uty. -passor9 xajdwog rP[jaure-passorg = ayTuOSeIy iopenoy vpyjod vuynyy oneusud IET[aWIeL (Aer) uaas JON ‘uly, -passor9 xajdwoy =e] JaWIe]-passo1g oyuosery POLIOWY “AA Dj9]0XA VIJIUOAJIV IY oreusud qe]jaurey wea uaas JON, ‘uty, -passo1o xajduroy =e] [AWE|-passorD ayuosery atopenoy wasy? vjJINOAD LY oyneumsud osneurs.id qeTJSuUTe] Japurjos, uty. ‘UIyL -passo19 xejdui0g «re]JaWe]-passoIg © aytuosery = —Aeqanbuv1ry pa2vjoun DAVID IN onewsiud o1eusud AeTeUre] sesuy ‘younstipuy ‘jouNstpuy -passoio xajdwioy Ie[Jaue]-pessory © aytuoSery uosyoe[ JIOg vyonpoag vajovyyy aeyawey iddyryd uaas}ON uaes}ON -passoi9 xajdwiog re]jaure[-passo1g = ayluosery saurddyiyg I1UOZ0INA) DAZID I qeypewey (snoeuury) — uaas JON uaas JON -passo19 xejdwiog sejjeure[-passoig = e}tuodery ureyige DUYIYAOI DAJID IY onjeusiid Ap]Jouey (urjaur5) uaas JON ‘uly, -passor9 xajduiog reyfauIe|-passo1g = aptuOSeIW saqfayoAas DIDAQn]S vapojIv] A0JONPPY Tented SuOoIPeAIASGQ, ——___ -, —____' dade] Jouuy aah] Jayng = ASoyesrauryyy Ayyeo0T aweN PORISOATY VaOVELOVN g aTav, voensoAu yepad oneusud oewsud qeyjauey yorewey ‘zaAv] Jauur UT spueq Wsug ‘UIE ‘uly -passor9 xajdwiog —-IeT[aWIE]-passo1g apuosery uraoQ ueIpuy vsommnbs pusvpiayt vorsysoAw jepad oewsud oneustid qeypauULy SuIpoL ‘aaXe] JOUUI UT spueq WStI ‘un. ‘uly, -passo19 xejdwoy reT[auUrE]-passoIg © aytuosery uea0Q UPIpUL DUD DUIDpLAT vorjsoAum jeped onrusud sewsud TeTaMEy] yoreurey ‘z9AR] JOUUT UI spurq WSL UNL ‘UIY., -passoi9 xajdwioy ivyjauey-passory = a] Tuo ey sarpuy }Seq ‘DII0K9 DUIDD: oneusud oreursud rvypawey snoeuury — JaXv| JUN UI spueq WIsTIg ‘uy ‘UY, -passo19 xajdwiog —Ae]JawL]-passorg = — a} LUO IW eyeqsny sndoddiy sndoddiH : 10;9nppy Tented SuoIeAIasqO —___-,. ——___' aoXv] Iouuy qoke] taynQ = Adoyesauryy AyyeooT satoads vovi}soAqy VaIOVNOVORELL Latavy SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 265 MACTRACEA (Plate 4, figs 1 & 2; text-fig. ro) Thirteen species have been examined mineralogically and ten structurally. The shell is entirely aragonite. There are two shell layers in all species examined, an outer crossed-lamellar layer and an inner complex crossed-lamellar layer bounded by the pallial myostracum. In the outer layer, the first order lamels are arranged concentrically and in this super- family are characteristically very fine (Plate 4, fig. 2). The layer is usually very thin and worn in the umbonal region but forms most of the hinge. In Spisula solida the crossed-lamellae are finer than in most other species and appear homogeneous in the inner parts of the layer. The inner layer of this species also has a very fine structure and sheets of myostracal prisms are present. These sheets also occur in Raeta undulata. The separation of the two layers is sharp (Plate 4, fig. 1) but the pallial myostracum is thin and indistinct in most species. The adductor myostraca are also poorly defined. SOLENACEA (Plate 4, fig. 3) Seven species have been examined mineralogically and four structurally. The shell is aragonitic throughout. Two main shell layers are present, an outer crossed- lamellar layer which forms the hinge and an inner homogeneous layer bounded by the trace of the pallial line. In the outer crossed-lamellar layer the first order lamels are very fine and arranged concentrically to the shell margin over most of the shell. Locally this layer may appear homogeneous. A very thin prismatic pallial myo- stracum, best developed below the umbo is present in all species. The inner layer of all the examples appears homogeneous with a striking lamellate appearance. Electron-microscopy of the inner layer of Ensis siliqua shows that the apparent homogeneous layer is in fact built up from layers of very fine complex crossed- lamellar structure (Plate 4, fig. 3) which alternate with bands of fibrous appearance which may be organic matrix. Etching reveals the presence of a reticulum of organic matrix sandwiched between carbonate laths. Fic. to. Radial section of Mactvonella exoleta. CL = crossed-lamellar, CCL = complex crossed-lamellar, PM = pallial myostracum. 266 TAYLOR, KENNEDY AND HALL TELLINACEA (Plate 4, figs 4, 5 & 6; Plate 5, figs 1-7; Text-figs 11-15) Thirty-one species have been examined mineralogically and twenty-four optically. The shell is totally aragonite. Most of the species we have examined have three layered shells (Plate 4, fig. 4), with an outer composite prismatic layer, a middle crossed-lamellar layer and an inner layer which may be made of complex crossed-lamellar or homogeneous struc- tures. The inner layer is, as usual, bounded by the pallial trace. In Egeria radiata, Florimetis corrugata, Psammotea radiata, Macoma balthica, Tellidora burneti and two species of Solenotellina, there are however only two shell layers, an outer crossed- lamellar layer and an inner, complex crossed-lamellar layer which is bounded by the trace of the pallial line. Trueman (1942) describeda threelayeredshellin Tedlina tenuis. The outer composite prismatic layer of three layered species is usually very thin and is frequently worn away from the umbonal area. It consists of horizontal first order prisms, arranged radially from the umbo. Each prism is built up of fine needle-like second order prisms (Plate 4, figs 5 & 6) which are arranged in the characteristic divergent feathery pattern seen in longitudinal section. This layer is at its thickest development in the Donacidae and Semelidae, where the arrangement of first and second order prisms is very clear. The Donacidae develop strong internal marginal denticles with a resultant thickening of the outer shell layer (as in the Nuculidae). The middle layer of three layered shells and the outer layer of two layered shells is built of crossed-lamellar structure with rather fine concentrically arranged first order lamels (Plate 4, fig. 4; Plate 5, fig. 3). This layer forms the hinge in all species examined. The inner layer of most species examined is built of complex crossed-lamellar structure (Plate 5, figs 5 & 6) although as in the Solenacea the fabric is so fine as to appear homogeneous under light microscopy (Plate 5, fig. 7). Electron-microscopy reveals that this is very fine complex crossed-lamellar structure. In some species Quidnipagus palatam, Asaphis deflorata, and Psammotea radiata thin prismatic sheets are developed. In Semele tortuwosa these occupy most of the inner layer (Plate 5, figs 1 & 2). The inner layer is also often markedly lamellate caused by the presence of thick sheets of protein matrix which presumably account for the flexibility of these shells. Most species possess a thin prismatic pallial myostracum (Plate 5, fig. 2) although this may be indistinct in some and marked only by a sharp break in growth lines. Sections through adductor muscle scars reveal pads of myostracal prisms whilst thin prismatic bands of the trace pedal muscle attachment were seen beneath the hinge line in Psammotea occidens, Solenotellina diphos, and Tellina calcarina. A ‘so called’ new shell layer called the mosaicostracum has been recognized in the Tellinacea on the basis of surface morphology by Hamilton (1969). We are uncertain how this relates to the layers recognized herein. We have examined the fine structure of Semele tortuosa, Donax faba, hecuba scortum, Tellina radiata, Quidmipagus palatam, Asaphis deflorata and Scutarcopagia scobinata. eze][aure] 19AP] IOUUT azeT[aue] 1aAV] IoUUT ayeyjaurey IaAe] JIouuy pepuregq ‘euy 1aAe] IouUy oyeUISTIG suor}zeAIesqQ 10;onppy = = snoaussouloyy Ieypewey = Teyewry — oVeUISHI -passoio xajduiog = -passoxy TeT[ouey -passoio xajduioo = eyJowrey] — oryeuIsLIg /snosueso0ul0yy + -passo1g Ieyewe, = reTfourey _ O17VeUISHIg -passoio xejduiog -passoig Ieyewe, = Ie[jeurey o17eUISIIq -passo19 xefduiog = -passoig Temied IoAP] IoAel SS —— Jouuy = eT PPI eoelysoAy VaAOVNITTIAL OI aTavy lIopenoy Jopenoq saTjayoAes Ayeq] lopenoy SoIpuy “MA (snoeuury) DIADUIADI 111152145 uoss[O % Arqs[ig wss01Q SN4ANIaIOS (peruog) Avdsip vuynss19s (snoeuury) syUsval vuDASDE) Aqiamos SNSABQSUDA] XDUOCT snoeuury SnyDNIYUap xDUuOGE —orpTuosere T[e are satsads Sutmoyoy ayy, (Aqiamos ¥ drrepoig) -passolg = oy tuo Seay ue}ezeyy Yauang vsoprya [. orzeusiid ue9009 snoeuury eyisodmo0gj =— aq tuo Sey ueIpUy DIDSALA YpJaU2}Ia T. oreusiid yoreue’y ayisoduiog = a tuoSe1y oueqsug viumdaapida xvuoq oryeurstid Agyue py optsodurog aytuosery lopenoy Aagsw xvuoq oryeustid (sneeuurq) ayisoduioy = oq uoSeIy — saTfayaAas vypsoyap siydvs pF IohPT rang = ASoyeA0 UIT Aypeoo'y aure N azejaume] 1aXe] JouUT Jewry 104] soUU] oyeyjowmey 1a4P] Jouuy asury yyeauaq 1aAv] JOUUL Ul JUasaId spurq WsUg aze|jawe] 19AP] Jouuy oquin y}eRauAaq vorijsoAu tepad ‘1aAP] Jauur ur spurq wstad uy y, azeyJaue] 1aAv] Jauuy JoAR] JOUUL 9}e][aWIeT] minori}zsoAur iyped yoryy A170, azeJawey] JaXr] Jauuy] aze[jawey] 1aAv] ouUy JaAxy] Jauu spurq wsug azeyjowrey] IAP] tauuy JaAv] JIUUT 9ye]]oUILT JaAP] JOUUT UT gsarnqny ayeyoure] IaAe] JouuyT azE[JawL| 1oXe] JoUUT pepurq ‘auy 190A] 1auUT suolyeArasqQ, O1RUISL = ARUSUg IVRUISUg IVRUIStIg onewsug — aryewsig _ o1yewsiag — o1yeusiig _ o1yeuIsiig _ oneusug — oneusug _— orewmsig = onewsug -- o1eusig — onewsig _— orewsug — o1yeUIsug oWeUsug IeWSUg jojpONppy reurea ————_,—_—__S PovIISOAT snoauasouloyy reyjaurry -pessoro xojdu105 snoauasowoyy Te[jawey] -passoio xajdui07 TeTPPUe] -passoi9 xajduro5 Teyaurey ~passoi9 xajdurosy Ae]jouey -passo19 xajduio5 snoouasouloy{ Ie]jaurry -passoio xajduio9 rryawry] -passo1g Ie]auTey -passoig, IETaMUe] ~passoig Te]jourey -passory re[[aury ~passo1g IP][aWI] -passo1g reyawey snoauasowloyy -passo1g sNoauasoul0yy -passoip, ayyaurey, = Tey_aurey -passoi9 xajdurog = -passo19 qeyawme, = reypaurey -passoi9 xajduiog = -passor9g, IET[AUTE] ~passoio xayduioz = qeyjauey snoauasouloyyY = -passo1g IeT[aUIEL ~passo19 xajdui09 = qe]jaurey snoauasouloyyY -passois IPT[aUrRy ~passoio xajduiog _ reyaurey ~passor9 xayduio9g — reyjourey -passo19 xajdui09—1ejjauTeT /snoauasowoyy -passo1g Ieyaurey = TeTaurEy -passo19 xajduioy © -passoig Ieyaurey -pesso19 xajduroo = re[JauIe] /snoauasowiopy{ = -passoig aeyjewrey = Teyjaurey -passo19 xajduiog = -passoig qeyawe, = reyjourey ~passo19 xajduiog §=-passo1g rah] rahe] qouut = @| PPIN VGOOVNITTAL OL aTav Jopenoy sopenoy sayjayaAag Ayev Jopenosy SeIpuy “MA (snaeuury) DIADUIADI DIPLSLAPS uossiQ ¥ Arqsita 155049 SNJANIAIOS (peruoD) avdsip purnss195 (snaeuurq) syisvaf vunasvy AqiaMos SNSAANSUDA] XDUOCT shoruury snyonayuap xouocy —o1pluosere qe are sarsads Aurmoyjoy ayy ~passo1 ayuoseiy UPzEZEY o1yeusud uPra00 apsodui0>) oqyuosely ueipuy aeustid ayisoduioy = ayuOSeIy = SaIpuy “AA oneusud apisodwoj = aytuosery = pueyuaarsy IET[aUIE] Pipensny apuosery ‘I9ART UeMS Ieyjauey ~passo1g, ayluosery vIpuy oryeustid urauel ayisoduioy —ajisoduuo9 -I9}IpIT oreursiid aysodwog ajtuoseiy Jopenog, onewsud apisoduioy = ay tuoSery Jopenoy oryewsud ayisoduioy) «= aytuoSv1y puysuaan() oneusud ayisodui09 ayimosery sayjquy oreustid ayisoduro> ayuosery lopenoy oryeusuid aytsodurosy ayuosery — sajjayaAag qey[aurey -passo1g ayuosery = sayjayoAag oryeusuid ayisoduro) apuosery saurddyryg Ie[awey -passolg = aytuoseiy ure}ig oryeusiid. ayisodwoy aytuose1y Bouyy “S Iepourey -passoig =: o}tuosvIy Iopenoy reyjourey| -passolg = ayuosery = PLAINS oryewsid aysodmoy = ayruosvry urezyugT orewsiud elperjsny ayisoduioy «= aytuOSeIy ‘IaANT UEMS onewsud ayisoduioy «= aytuosery aueqsug oneusiud ayisoduioy = aytuose1y iopenog oryeustid eysodwoj = =©aytuosery — saqqayoAag IaAP] qayng = ABoyezauTyY AqyRo0'T (Aqaamos 3» duapoig) yauang vaopyya T snovuury pypsaia yypaeyya 7 snavuury DjDIpvA BUNA TL. (urjoury) DIADIIDI DULOID (poo) vypipy4rg vUyjaoUajoS (Aqaamos) soydup nuyjnouszos ayraurerg, snyyy -[18tays snpAnaajos ds aaumas (swepy ‘@ 9) DSONJAo] 9]91UI9S (snaeuurq) vyourgors nusvdoravjnag (snaeuury) syafonsuy vispdo2dnjnag (aa]Suads) vusnw DiADjNIVGOAIS ayepery wunjopod svsvdiupmO (tddyrqa) DIDIPDA DIIOU Wnsq saXeysaq SUIP1IIO DIJOMMEST (snaeuurq) vo1yjj0q DUOID IY (snaeuurq) wingaors vqnaayy (Aquamos) DIVDSNAAOI SYIULIAOD AT (joreure7) DIDIpYA DABS] (3805 ep) sno xpuoqy urjaury vqvf xvuoq yorewey vuudapidea xvuoq Agquueyy aadsy xouoqy (snevuury) vyoaoyap siydvs y owen SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 267 Fic. 11. Radial section of Donax faba. CP = composite prismatic, CL = crossed- lamellar, CCL = complex crossed-lamellar, PM = pallial myostracum. Fic. 12. Radial section of Hecuba scortum. CP = composite prismatic, CL = crossed- lamellar, PM = pallial myostracum, CCL = complex crossed- lamellar. Fic. 13. Detail of a radial section of Hecuba scortum showing alignment of crystallites in the outer composite prismatic layer (CP), and of lamellae in the middle crossed-lamellar layer (CL). Fic. 14. Radial section of Macoma balthica. CL = crossed-lamellar, CCL = complex crossed-lamellar, PM = pallial myostracum. Fic. 15. Radial section of Solenotellina sipho. CL = crossed-lamellar, CCL = complex crossed-lamellar, PM = pallial myostracum. 268 TAYLOR, KENNEDY AND HALL GAIMARDIACEA (Plate 6, figs 5 & 6) Only one species was examined from this small superfamily, it is aragonitic. Gaimardia trapezina (Lamarck) from Tierra del Fuego has a two layered shell. Both layers are made up of homogenous structure; the constituent crystallites are very small about 0-5 p in diameter and have irregular rounded outlines so that the structure appears granular (Plate 6, figs 5 & 6). The two layers are separated by a thin prismatic pallial myostracum. ARTICACEA Four species of this group have been examined structurally and six mineralogically. The shell is entirely aragonitic but there is some variation of structure within the superfamily. In Trapezium and Coralliophaga there is an outer crossed-lamellar layer and an inner complex crossed-lamellar layer bounded by the trace of the pallial line. The primary lamels of the outer layer are very coarse and are arranged concentrically to the shell margin; this layer also forms the hinge. There is a thin prismatic pallial myostracum dividing the layers. In Arctica islandica there are two shell layers, but in contrast to the other members of the family, both layers are built of homogeneous structure. The two layers are separated by an extremely fine prismatic pallial myostracum along which there is a distinct break in growth lines and shell layering. Details of the fine structure of the homogeneous structure were given in Taylor e¢ al (1960). A specimen of Calyptogena ponderosa, a member of the problematical family the Vesicomyidae was examined. Members of this family exhibit certain similarities with both the Veneracea and the Arcticacea, Boss (1968). The shell structure consists of two homogeneous layers and in this respect resembles the Arcticacea more than any other superfamily and is thus placed here for convenience. A similar conclusion was reached by Oberling & Boss (1970). DREISSENACEA (Plate 6, fig. 3; Text-fig. 16) This is a small group of byssate freshwater bivalves of which we have examined one species structurally and three mineralogically. The shell is aragonitic. In Dreissena polymorpha an outer crossed-lamellar layer (Plate 6, fig. 3) and an inner complex crossed-lamellar layer are present ; the former forms most of the hinge. The layers are separated by a thin prismatic pallial myostracum. The primary lamels of the outer layer are arranged concentrically. puelsuq JON aUOse1y “SION, mnouumy wuniprsrg (pnerre9) ayuoseiy DUIAgOSUOI DUAAKD ureyig, (Aqramos) eyuosiy ‘ueloeureds snuaofiauna vyn2194075 ureyig: (stz10]{{) oyuosery ‘ueroeureds DIVPAOI HINILQLOD Ieyjaurey Ieyourey eIpuy (AevIn) vaurmny “> ur se raXv] JOUUT PuYsIpU. -passo1o xejduioy —-passoro Ajouryy ayUOseIy ‘ero, eSuey, saproursgda vgraoja A I9AV] JouUT 84} Ur yueseid are suustid orjeusud Tejaurey Ie][auey] (soXeysaq) ad4}-ye0e.14s04n Jo sjoays Uy, ‘uIyI -passo1o xaqduog -passo19 Ajouyy oyUOsery Topenoy —vjnumoun vposaujog snonurzuoostp oryeurstid TeTaUILy] Ieyourry] St tanoeijsoAum yeryyed oy, ‘UIQ -passo19 xajduro9 -passoio Ajout sy ayuosery —enServor N Aes vjoyfur DUadhy reyjauey IeT[OUIeT ezued Ny aAoge se rae] JoUUT PUYSIPUT -Passoio xefduiog = -passozo AToung ayruoseiy axe] “ds vynaiqaoa IeT[OUIe] Ie][aurey saAeysoq aAoqge se rave] JouUT PUHSIPpUy -passoro-xefduroy —-passoro Ajoury opuoseiy eIpuy suap1990 »jn91q4075 evaoesdoury ou UI Se ‘sUUINJO Ie ma jon AIOA = - = (snoeuury) Aywueurmoig “1aAe] IguUT O1peUISII TnOser urequg DILPUD]SL DIYIAP Ul spueq o1yeUIStId UY, = 4oUTZSIpuT ‘UI snoouasowoyy snoauasoul0 yy yt V oe snoauaSow0yy SnosuesoWOFT = aH UOFVIW —ODTXA JO" wsoaapuod vuasoji4Xjvg eacrel (snoeuury) mall nizaqva. oeuUIsLig -passoio xejduioy = re] auIe]-passo1g opuoseIy Uuedd0 UeIpUy mnsu0]g0 mnizadvA I, i (uyjeun) vsvydoyposoa eaves tuoSer SeOnT[ON ninyqoutp4og — oeuUslig -passo1o xejdui0g IeT[eure[-pessoig = =-_- oo Vv IoyONppy Teed 1K owen n Asojer1oUur HLEIO'T suo1yeazesqo a JoAe] IoUUT IaAP] 19jNO T TAL eoelysoAy VaOVOILOUNV II ATaVL ayuosery ajmoser aymosiy aqymosery Teyjaurey Ielaurey vaurmny “> ut se 1aXhv] JaUUT — qouystpuy -passo1s xajdmog = -passo19 Ajautg ayuosery Jahr] dguut ay} ur quasaid are suistid oneusud Ie[[aurey aepjeuey adAq-[eoe13sohur yo sjaays unT. — ‘ary, -passo19 xajdurog = -passo19 Alou aymuosery snonuruoostp onemsud IPT[AUIET IET[AUIL] SI tnoriysoAu perped ayy - ‘uly. -passo1o xajdurog = -passoro Ajauty aymosery Te]aUey] Ieyjowry aAoge se 1aAP] JoUUT youysipuy -passo19 xajdurog = -passos9 Ajoutg ayuosery TeT[aurey aeyjaurey aAoge se rahe] 1auuy = qouNstpuy -passoro-xajduioy =-passor9 Ayjoutsy aytuosery raoesdoury ay} Ul se ‘suUINjoo rejndax AraA ul pasuerre sqj}eT sMoys IAP] apyewey Teyjaurey Ie|[auUE] passoio-xajdu09 any — youysipuy -passo1-xajdurog =-passoi9 Ajaurg = ayuoSery JoyoNppy reread suoljzeArasqaQ, u—,—~ iaAv] Jauuy Jake] JaynQ = ASoyerauTyy PoRISOAW VaOV TNOIEUOD br atavye 91njon44s oreusud Tey[aurey — ‘uly, -passo1d xajdurog reyjemiry-passoig «= aTUOS ey susud adA}-jvor.4soAur jo spurq ayewsud oeusud TRUM reyjaurey JoAv] JOUUT UI spueq ws ‘ayy, ‘uly, -passor9 xajdu0g snoauasou10fy oyuosery suisud ad4q-jroerysoAur jo spueq qahey oonewsud onewsud WM reypauey dauul ul spueq wsud Aur UI ‘uly. -passo19 xajduiog snoauasowoyy ayuosery a0yoNppy ened suolzeAtasqg —, aoAP] Jouuy rake] 1aynQ = ABoyerauryy PoRmsoAyy VdAOVSSOTS €r alavy ajuoseiy ayuosery apiuoseiy orewsud aP]aue] adury sui0y Iafvy 133nOG — uly. -passoin xajdwiop re[[aure[-passoig = a} tuosery 10};9Nppy remled suorjzeArasqQ —_ aaAvy rouUT yakep iajng = ASoyesauryy eoerysoAqy VaOVNASSIaua ZI ATaVy, aquos1y ayuoseiy pepueq Ayueurmorg “raAe] rout oneusud ur spurq o4eusud uy youNstpuy ‘army snoauasouofy snoauasowoy, = ayuosery = snoauasou0yy snooussowoy, apluosery reyjaurey — ayeusug -passois xejduioy reljauey-passory = — aqruosery aeyjourey _ oneUIsUgd -passoio xajdmog xreyjaueE[-passoig — aytuoSery aoyonppy remed. SuoIZeAIASqO YY saAv] 1ouuy Joke] 193nQ = ABoyeroUNY PRORIYSOAT, VOOVOLLOUV II alavy, uea9Q UeIpUy purysuq TaN “SHON tng wnipisty (preyed) DULAGOSUOD DUIAKD, urequg (Aqramos) ‘avioemieds siusofiauna vjna1q4oD urequag (staz0qy) ‘aerseureds DIVPAOI DINIWQAOD erpuy (fe15) : ‘Q1O[PSuLY saproursgaa 04140194 (safeyseq) aJopenoy vjynumouv vposamcog enservoiny Aes vyoyfur vuaska | ezueANy aqeT «ds mng19q409 saXeysaq, eIpuy suapi1990 MynI1q40D, (qoreurey) uedef vauuungy, v1nI1qQ409 Aqyrpeso"T saroadg (aaaayq) uedef y24muM) wipavzorayr Ayeay (sneeuury) 9uao04sla[q snupuny snssojy (snaeuurq) } UPJ JO IST snununy snssopy Agr e007 saroads meq, ‘gua00q, Uupundg vuassiaaqy uapauag UBA PLUasIN puvriafy DUuassiaAqy (seqred) eiqeiessag § vy daowdjod vuassiaaq purysug (seqrea) “s}ON vydsowdjodg vuassizaq - AqyRo0T sarsads | Urey, ‘“uerqry (Aqiamos) ‘snoaoe}a19 suuaaofipaso? v9 94 ures ‘ueeuey (Aqiamos) ‘Arensay, vung 091947 (snoeuury) Wreqig DIYPUDISS VII ssog ? OoIXaW JO" vsoaapuod vuasojgxqv9 (snavuury) wnsuojqo wnrzadva (uyjeurg) vsvydouzosos seonqjoW vsvydoyyos05 AYIyeIOT owen SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 269 GLOSSACEA (Plate 6, fig. 1) Two species of this very small superfamily have been examined structurally and mineralogically. The shell consists of aragonite. In Glossus humanus there is an outer layer which is largely homogeneous and an inner complex crossed-lamellar layer which is bounded by the trace of the pallial line. In some specimens and in some parts of the shell there is a faint vertical structure similar to crossed-lamellar structure (Plate 6, fig. 1). The inner layer is built of rather fine lamels and in all specimens there are many thin sheets of myo- stracal prisms. In very old individuals where the inner layer is thick these prism sheets become abundant and closely spaced; in some irregularities develop and spherulite patterns appear in some sections. This is probably due to the develop- ment of corrugations upon the accretionary surface. In Meiocardia lamarckii there is an outer crossed-lamellar layer in which the lamels are distinct and arranged concentrically; within this there is an inner complex crossed-lamellar layer bounded by a thin prismatic pallial myostracum. CORBICULACEA (Plate 6, figs 2 & 4) Six species of this superfamily have been examined structurally and ten miner- alogically. The shell is totally aragonite. Again two shell layers are present, an outer crossed-lamellar layer which forms the hinge and teeth and an inner complex crossed-lamellar layer bounded by the trace of the pallial line. In the outer layer the lamels are rather fine and arranged concen- trically (Plate 6, fig. 4). In most forms no prismatic pallial myostracum is visible only a dense granular zone separating the layers. Prisms are however detectable in Polymesoda and Velorita cyprinoides. The inner complex crossed-lamellar layer is always built of rather coarse laths, and in all species of Corbicula examined and in Velorita these are arranged in columns as in the Limopascea (Taylor et al, 1969). Tubules were seen in Pisidium amnicum. Fic. 16. Radial section of Dveissena polymorpha. CL = crossed-lamellar, CCL = com- plex crossed-lamellar, PM = pallial myostracum. 270 TAYLOR, KENNEDY AND HALL VENERACEA (Plate 7, figs 1-5; Plate 8, figs 1-5; Text-figs 17-22) This superfamily includes a large number of extant genera and species, conse- quently we have examined over fifty species structurally and mineralogically. The shell is aragonitic in all species. Both Boggild (1930) and Oberling (1964) stated that the distribution of shell structure types is highly variable, thus in order to ascertain if there is any systematic variation we have listed and discussed the species examined at family and sub-family level. (Table 15). The shell structural variations found are indeed more variable than any other superfamily. The various combinations are shown diagrammatically in Text-fig. 17. The apparently most important structural distinction is that between species having an outer composite prismatic layer and those without. The other variations exhibited between crossed-lamellar, complex crossed-lamellar and homogeneous structures almost always show transitions and all gradations between these structures may be found. There is thus in many species a basically three layered shell consisting of an outer composite prismatic layer, a middle crossed-lamellar/homogeneous layer and an inner complex crossed-lamellar/homogeneous layer. In most other species the shell is basically two layered with an outer crossed-lamellar/homogenous layer and an inner complex crossed-lamellar/homogeneous layer. The composite prismatic layer consists of radially aligned primary units made up of smaller crystallites radiating from a central axis (Plate 7, figs 1-5). Each of these smaller crystallites may be several mm. in length and 40 yp in diameter, but the size varies greatly from species to species. Each of these crystallites is surrounded by a sheath of organic matrix. Closer examination shows that each of the larger crystallites Fic. 17. Diagram showing the main types of shell layering found in the Veneracea. CP = composite prisms, CL = crossed-lamellar, CCL = complex crossed-lamellar, H = homogeneous, PM = pallial myostracum. orzeustid ‘UIgL snosuesomo0y{ — Ie][Pure]-passo1y oymuOseIy Iopenoy ‘ds wnjrg orzeustid . ‘ug snoomasourozy — TeTSUIE[-PsssolQ = ayTMOseIY uve0_ UPIpUyT (aysuryH) seufly sor SpIeMut snosuasouroy Surm0s;8q (Aqiamos) snosuasowmofzy —_— ‘Ie[JoWIe]-psssolg ayuoserIy wpyonbs vysyjoo0Aav Jy spremur orzeustid Ie][SUILT snosussomoy Ssururosaq (snoeuury) uly -passoio xejduro9g — /zeyJowR]-pesso1g = op uoSery purjsusen() SISWAAJSDI pyguorory Te][eure]-passo1o oryeutstid xe,dutoo (Aqiamos) ‘TY L /snoaussoul0 yy Ie][ewle]-pessorg oeustid oysoduiog = oy 10 Serv iopenoq DULIAAIGSD DYIU0IOLT oryeurstid IeT[OWUIeT : (Austq10,p) ug -pessoio xajduio9g — Iv][ewWe]-pesso1g apuosery iopenoy sesuadod nysuosjaumv ty snoouesour0y/rey[aurey (Aqiamos) -passoio xe[durog — Ie][OUIe]-passo1g ayuOseIy Jopensy vsourdsyjnum vysuososagsa Fy SpIeMUrI snosuesou0Yy (snoeuury) Or} eUIsIIg snoauasowofzy{ — Ie[[aUIe]|-psssolg aytuosery SOIPUT S89. auoup pysuosonagsA EI Se - $$ Lr nme mr spreMuUr oreurstid snosuesouloy suru0seq (e3s09 ep) ‘UIT L snosussouloyY == ‘UTY} Ie] [eure]-pesso1g oneustid oyisodwioj «= oq uO SeIy Ayeqy DINIOLAS SnUua A spieMut orpeurstid snosuesowoy Suru0seq (aa9a37) ‘UIT L snosuasouloyy Ie][OUIL]-passo1g oreusid oyisoduog «= oq uo sey Aled DyDID snua A :OeULIOUS A winoe1ysoAu leured JaAe] IOUUT I9AR] O[PPTIN IaAR] IojNOQ ASoyeroUrp, Aqeo0'_T setoeds VAOVUANAA Cr alavy oneusud ‘arg snooussoul0yy oeuisud ‘arg snoamasomoyy snoauasowoyy oneusiud IeTOUIET ‘arg ~passoi9 xajdui0oz Ap[]aure|[-passo1o ogeusud xayduio ‘un /snoauasomoyy oreuisrid Ieyewmey ‘UL -passor9 xajdu09 snoauasouloy/rTeyjauiey ~passo19 xajdui0od ones snoauasowoyy snoauasowloyy Avjjaure|-passo1o onrusid xaduroo “yourjstpuy /snoauasowoyy snoauasouloyy jourjsipur ‘oryeulsug snoauasowoyy orewisiig snoaguasowiofyy o1yeusug snoauasowoyy snoauss0Wwoyy snoauasouloyy swsud yeorsjsoAur QiM reyjaurey o1eusitg, ~passo19 xaydwio05, suisuid yeoejsoAur YF TeTOUeT O1PeUISLLT -passor9 xajdui0d, susid [eoe.1jysoXur FIA reyaurey, ores, -passoio xejdui0> sjaays wstid ‘reyjaurey oryeulsig -passoio xojdw09 onjemsud wy snoauasouro py qouysipuy snoauasoul0 py swistd eoe1jsoAur jo sasue] FIM Ie[jeurey o1eUIsiig -passo19 xajdui0> qouljsipuy snoauasowoyy o,qeurstid ‘ory snoauasowofy orjeusiid “uy snoouasouloyy onrusud ‘Uy snoauasowoyy orpewsid “ary, snoauasowoyy wnoerrjsodw lemred aaAv| rouUy IPTJUIE]-passorg, snoauasowoyy spiemut snoauadomoy Suraros9aq ‘1e]]aUIe]-passo1g spreMut snoauasowoy Surmo.q ‘Ie][ewIe]-Passo1n spreaut snoaussomoy Suru0.8q “ury} Te[[euUe]-passory spreaur shoauesowoy Surm0.eq Av][oure]-passo1g a94R] B1PPIL TETTSUTE]-Passory IETPUTE[-passorg SpIeaur snoausgomoy surm029q ‘IPTJEWE]-passorg Spremur snoausasomoy Suraro.aq /zeyjaure]-passo1g oyeustud azisodaiog AP|[EUIE]-passoin IP|[AUIE]-passo1g, SpIeMUI snoauasomoy Ie[jaure]-passory spremul snoauesowoy rpyjaurey passoro-Ajauny spreaur snoauasouoy Suru1098q aeyjeurey-passor9 ATaULT spreMul snoauasouroy Iey[aUIe]-passo10y oyeusud aptsoduiog spreMul snoauasowoy zeyjawey-passoro Ayjaury spieMul snoauasouloy oyut Suissed /zey]aue]-passoro yeIpexy rejjawel-passors Apauty rey[aurey-passos9 ATaut SpIePMUI snoauasow0Yy /zey[aure]-passorg spIeMul Shoauasowoy /aeyjaurey-passo1g SpIeMuUI snoauasoulOYy /xe|Jaure]-passo1g spieaur snoauesowoy /zeyJaure[-passory spivMur snosuasowoy /xeyjaue|-passor) SpIeMur snosuesomoy /xeyjaue[-passorg onewsid aysodu09 spremut snoauasouroy Suruz09aq apyjeure|-passoro Ajouty arzeuistid ayztsodu09 spreaut shoauasowoy Suruo0s9q ‘e]]aUIL]-passorg oyewusud ajisoduo9 oeursid azisodmo5 A9AP] 1930 VaOVUANAA Cr alavy ayuosery ayruosery opuosery apuosery ayruosery ayuosery ayuosery ayuosery ayuosery opuosery apuosery ayuosery apuosery ayuosery ayuosery ayuosery apuosery ayuosery oyuosery aqyuosery ayuose1y oyuosery ayuosery ayuosery opuoseiy ayuosery ayuosery ayuosery ASoypesouryy Jopensyy ue29Q UeIpuy puejsuaan Jopenoy Jopenoy ropenoy Sorpuy 4S9\\ ueadQ ULIpUy sopenogq uapy aopenogq uva0Q UeIpuy uea0Q UerIpuy epnuueg vyeqsny sayjayoAes wea0Q UeIpUy sotpuy sey ogtoeg-opuy uepy Bag pet “WS SouOL POLY 3S9\\ ureqgy > sade PoLIOWy yeaquag, Ayeay Ayes Agreo07T psourdsyyni vy2uor04aSC Fy a ds wong (ujauip) swelfo avr (Kqzamos) ppypnbs vjsyyo204620 JV (sneeuurq) S1sug4jso2 DYIUOI0NT (&qramos) _ viurdsagsy DYIUOIONT | (Sa3q20,p) sasuaphndg Dyauoayjauioy (Aqzemos) (snoeuury) auorp py2u0I0495h FT wnpuuny (su: upped wnivafoy (uyeurp) — tunyworavarp wnravafoy (enoeuurq) MAGAdas 9IK1D, (eaooy) uInoeIJSOAU Teled oreusiuid JoAR] IoUUT snoauesowoTyy TeT[awry] -passoi9 xefdwo09g Ie][oury] JA] a|PPUY SPpIPMUI snosuasOWOY Suruio9aq ‘Te[[PUIe]-passorg, ‘ary -passoi9 xatdui05 onewsiid Ie][OUIe] ‘aIyL -passor1o xafduiog o1eustid ‘uy snoaussowoyy{ oneusid Ie][owWe] uy -passoio xejduiog oryeurstid Ie]awr] ‘UI L -passo19 xepdurog oryeustid Ie[[owey] ‘aI L -passo19 xejdui09 winoe13soAur lemed JoAR] IOUUT spIeMUI snoauesowo0Yy ‘Te[[PUIL[-passo1n spieMur Snosuasowo0Yy ‘Ie][9UIe[-passo1g spreMut snoauesouloy Surtu0seq Ie][aure|-passo1g JOA] OTPPHAL IAP] 19jNO snoauasowio yy Ie] [aure|-passo1y Ie[[aurey-passo.y Iv]Jaure]-passo1g oeutstid aytsoduiog orneusid ayisoduog oneustid oz1soduiog suustid a}z1soduro9 IAP] 19}NnO panuyuoy SI aATaVy ASoperouTpy ayuosery ayuOseIy ayluoseiy ayUOseIy apUOseIy ayuoseiy apuoseiy ayuoseIy ABSoleIOUTI AyyRooT sotoadg Joyuedie5 elULOFIeD vunydvipqns wjjasadoo5 Saeplyeradoo5 (snrzyayq) UvdURLIA}Ipay, vspydoyjy vjorragag yoreureT uleyug srusofippjoyd vjonaag Aq19MOS ilopenoq DIDINIYUaP VIOINAJIG PeepPlOOIjag (snaeuury) puelsuq DIMDUaIAaU DIADUAIAATT (Aern) iopenoy vsosapuog viuisog tajuedie5 lIopenoyy apuun puso BUT[OIeD “S AyyeooT ‘ds miuisog :@BUIIUISO(CT saroads winorsysoAUL remled aoAv] t9uuy snoauasowoyy ; IE]PWe] -passoi9 xajduio> onvusud IP[[PWe] ‘Un -passo19 xaydwo) oeusid TE[[PW] “UL -passo1o xajdui0> oyrusiud ‘uy. snoauasowoyy onrusud “UL snoauasowopy Teyjaue] oneurstuid -passo19-xodu09 wy, /snoauasouo yy onrwsud Uy, snoauasowopy oneusud “UY snoauasowopy oneuisiid ‘uy snoauasouloyy oeusud ‘uIq snoouasomoyy, oneUsiid snoauasowoy, oneusiud ‘UL snoauasowoyy snoauasowoy/1e][awey o1yeusug -passoro xajdwi05 oneusid ‘uy. snoauasowoyy orjemsud “uly snoauasowoy oeusud ‘UL snoauas0woy IP]PPUe] oneuisug -passo19 xajdwio05 shoauasowopy onewsud apyawey “UU ~passo19 xajdwiod, oyewsud IeTaWey] “uly -passo19 xajduio05 oneusud Iv|pawe] “UNLL -passo19 xajdui09 oneusud IpyawWe] “Un -passoi9 xajdwiog winoriysoAu reured JaAK] JaUUT 49hE] 2|PPHN spreMul snoauasouloy Suruo9aq ‘TE|[PULeT-passo1y SpIeA\UI snosuasoWOY “Te[JOUIE|-passosg SpreMur snosuasOWoy Surwi099q /xejjaue|-passo1g spieavul snoauasowoy Surui099q /aeyjaue|-passo1g spieul snoauasowoy, Surm0s9q ‘re][PUrIe]-passoly snoauasouloy Surm09aq ‘re[[aurey-passo1y Sp1eMUur snoauasouIOY, surw03aq ‘Ie[[eUre[-passo1y spieaul snoauasouloy Surm03aq qe] [aule]-passo1y spieaut snoauasomo0y, ‘Ie|[aWeE]-passory) SpIPAUl snoauasowoy “1e][aWe[-passo1y Spreaut shoouasowmoy Surum0saq Te|jawr]-passo1y JOE] ATPPH raXej 1930 snoauasouloyy Te|]aWE]-passo1y Iv|[PUIR|-passory av[Jaue]-passo1y oyneusud aytsoduro5 oneusud oy1soduro5 spivMur snoauasowoy Surur0saq ‘re[aure[-passo1y SPIEMUI snosuasOUOY Suru0saq AP][PWIE]-passoly oeusud oytsodwo5 oyrusiud aysodmo5 SpreAur snoauasomoy suru09aq /2eyJaurel-passory onrvusud ayisodwo5 onevusud aytsodwo> spieMUI Snoauasowoy /ze][aute]-passo1y oneusud ayisoduio> oneusiud aytsoduioy onvusiud apsodui05 Av][aWL]-passos9 ApOULT spreMul snoauasowoy Surm099q ‘TE][AWIe]-passo1y, SpreMul oly}ue9U09 ‘apis}no uo yerper ‘re|]aUIe[-passosy oneusiud apsoduo5 ogerustud opsodurog stustid a}tsoduio9 A9Av| 19300 panuyuoy) SX aTavy, ASojes9uryy aquosery aquoseiy anuosely ayuoseiy ayuosery aywuosery ayuosery ayuosery opuosriy oyuosery opuo0sery ayuosriy aywuosery oyuosery aymosery aquosery apuosery ayuosery ayuosery ayuosery ayuosery ayuosery ayuosery ASoyperouryy AQye90T TIUIO;TTeD urauLta}payy ureyug: AOpenoy puepsug uedef nag vrapeyy JOpEnogy Jopenog PIUIOSIED SaIpuy 4S, Sarpuy 4S9\\ purleez Many pea, uosyor{ pod uo UeIpUy eury wWs'D dopenog JOpeNogy eurporeg *S Aqiypeo07T seroads Jojuadie> pupydoipqns v11242400), saeptjeradoo5 (snizjayq) vsvydoysy RpoIad suusofipojoyd vjoraag Aquamos Dypjnayuap wjoratad TSePIOOLQeg (snaeuury) DIADUIIAIUL DLADUIIAI TA stsuaopal vavyjorad (Aqranos) vuviandad vaoyqoary (euoatg) vsoffe vaoydourT ' (disaporg, ay Aqtanos) vpins sisqouowm) AqiaMog vsosnaqns auorys Aquamog vyjajppun auory Ds (snoeuury) viydod auory (urjaury) vvynuvsd auoryy (Avag) manqysinys ane (uyjaur5) DUDINLZDAQ DIPADIO[DUOL Fr sepUTUOIy | (oreurey), vyouadd sidnaaua, (snoeumry) vyrsayny sodvr (snovuury) syyxe miydog (uaq30],) pumas vuuay PavUTUItaS) ———Ee SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 271 is made up of further smaller units about 0-5 » in diameter and diverging in a feathery manner from the central axis of the larger crystallites (Plate 7, fig. 4). The outer layer is separated by various degrees of distinctiveness from the under- lying crossed-lamellar layer which has concentrically aligned primary lamels (Plate 7, fig. 1). This middle layer changes to homogeneous structure when traced towards the shell interior. In some species the distinctly crossed-lamellar portion of the layer is almost entirely suppressed (Plate 7, fig. 2). The crossed-lamellar and homo- geneous portions of the shell cannot be designated as separate layers for they vary in extent both between and within a species. In two layered Veneracea the outer part of the outer layer consists of crossed- lamellar structure (Plate 8, figs 2 & 4) which passes transitionally inwards into homogeneous structure (Plate 8, fig. 3). The orientation of the lamels in the outer layer is controlled by the type of shell margin present in each species. In the Veneracea the marginal areas are variable with margins which are reflected, inflected, shelf-like or combinations of these. A further complication to the shape of the shape of the margin may be ribbing and strong concentric sculpture. With a reflected shell margin the first order lamels in the outer region of the outer shell layer le Fic. 18. Diagram showing radial sections of three types of shell margin found in the Veneracea. A. Here the margin is slightly reflected; there will be a gradual change from crossed-lamellar structure on the outside to homogeneous structure inwards. B. In this case the margin is strongly reflected and in the position marked by the dotted line there will be a sharp structural change. C. The margin is even more strongly reflected than in B and similarly there will be a sharp change in structure along the dotted line. B* 272 TAYLOR, KENNEDY AND HALL subparallel to the outer shell surface, although at the time of secretion, they were aligned normal to the secreting surface. With an inflected shell margin (Text-fig. 18) the first order lamels retain more of a concentric alignment. Again, when traced towards the shell interior the crossed-lamellae pass into homogeneous structure. The point at which the change takes place is usually where the reflection or inflection of growth lines changes rapidly. Further inwards from these points the growth increment lines are much more closely bunched suggesting a slower growth rate. The change from crossed-lamellar structure to homogeneous can thus be interpreted as a result of differential growth rates imposed by geometrical constraints caused by the shape of the shell margin. In all cases the inner shell layer consists of complex crossed-lamellar or homo- geneous structures or combinations of the two. Sheets of myostracal prisms are also common. Fic. 19. Radial section of Chione subrugosa. CP = composite prismatic, CL = crossed- lamellar, h = homogeneous, PM = pallial myostracum. GR © H Fic. 20. Radial section of Chamalea striatula. CP = composite prisms, CL = crossed- lamellar, H = homogeneous, PM = pallial myostracum. ’ SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 273 Subfamily VENERINAE Most species in this group possess the three layered shell; composite prisms, crossed-lamellar/homogeneous and with one exception an inner homogeneous layer. Two species have only two layered shells. In Periglypta puerpera the inner layer is largely constructed of myostracal prisms with small areas of complex crossed- lamellar structure. Subfamily CIRCINAE All species in this group have the basic two layered shell. In most species there is a well defined prismatic pallial myostracum. The inner layer is variable; in the species of Gafrarium examined it is largely made up of myostracal prisms with small amounts of complex crossed-lamellar structure. In most other species it is homogeneous. Fic. 21. Radial section of Tivela hians. CP = composite prisms, H = homogeneous, PM = pallial myostracum. Fic. 22. Radial section of Lioconcha castrensis. CL = crossed-lamellar, H = homo- geneous, PM = pallial myostracum, CCL = complex crossed-lamellar. 274 TAYLOR, KENNEDY AND HALL Subfamily SUNNETINAE In Sunetta solanderi the margin is strongly reflected and the structure consists of outer crossed-lamellar structure, with the lamels radially aligned, which pass inwards into homogeneous structure. The inner layer is homogeneous with thin sheets of myostracal prisms. Subfamily MERETRICINAE In Tivela hians there is a three layered shell with the middle layer consisting entirely of homogeneous structure and also a homogeneous inner layer. Tivela ponderosa and Meretrix have a two layered shell. Subfamily PITARINAE Most of this family have a two layered shell but in Lioconcha asperrima there is an outer composite prismatic layer. Subfamily DOSININAE The three species examined in this group, all have a three layered shell, with an outer composite prismatic, a middle crossed-lamellar/homogeneous and an inner complex crossed-lamellar layer. A similar three layered shell has been described from Dosinia japonica by Kobayashi (1966). Subfamily CYCLININAE The one species examined has a two layered shell. Subfamily GEMMINAE Both species examined have a two layered shell. Subfamily TAPETINAE In Tapes litterata and Venerupis there is a three layered shell but only two layers in Paphia textils. Subfamily CHIONINAE Both three and two layered shells are found in this family and the variation may be found within species of one genus. In Mercenarvia mercenaria the crossed- lamellar portion of the middle layer is often indistinct but the structure is revealed by electron microscopy. Most species have an inner homogeneous layer. SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 275 Subfamily PETRICOLIDAE This group has two layered shells; in the outer layer the crossed-lamellar structure does not grade inwards into homogeneous structure, as in most other Veneracea. Subfamily COOPERELLIDAE In the one species examined both layers consisted of homogeneous structure. Order MYOIDA MYACEA (Plate 11, fig. 4; Text-figs 23-25) Six species have been examined structurally and twelve mineralogically. The shell is aragonitic. The superfamily divides naturally into two groups the Corbulidae and the Myidae, these are discussed in turn. In the family Corbulidae the shell is inaequivalve, the left valve being the smaller. The outermost part of the left valve consists solely of periostracum which fits like a flap against the right valve when the shell is closed (Yonge, 1946). In both valves, the periostracum may line the inner margin of the shell for some distance in preserved specimens. Two main shell layers are present, an outer crossed-lamellar layer which forms most of the hinge and an inner complex crossed-lamellar layer which is bounded by the trace of the pallial line. In the outer layer the lamels are arranged concen- trically to the shell margin. In adult specimens where the growth rate is slower, marginal thickening has taken place and the periostracal flaps and extensions are frequently incorporated into the shell proper, by subsequent deposition. The periostracal flap of the left valve may even become incorporated into the shell of the right valve. In most species there is a well developed pallial myostracum and within this there is the inner shell layer of complex crossed-lamellar structure which is fairly coarse. In all species there are commonly sheets of myostracal prisms interbedded with the normal structure. In Corbula crassa and C. tunicata there are well developed myostracal pillars arising from the trace of the pallial line. In addition to the marginal periostracal flaps the animal appears capable of laying periostracum-like material down as a sheet, over all the inner surface of the shell. In some species, this happens several times in the life of the animal (Text-fig. 25). In the family Myidae a rather different arrangement is seen, there being three distinct shell layers. There is an outer homogeneous layer, a middle crossed- lamellar layer and within the pallial line an inner layer which consists of either complex crossed-lamellar or homogeneous structures. The outer layer consists of granular crystals (Plate 11, fig. 4) about 5 yin length and 2-5 u in diameter with no obvious crystal form but with a slight elongation towards the shell margin. Although this layer is called homogeneous it differs from all other homogeneous 76 TAYLOR, KENNEDY AND HALL Fic. 23. Radial section of Platydon cancellata. HH = homogeneous, CL = crossed- lamellar, PM = pallial myostracum, CCL = complex crossed-lamellar. er CCL Fic. 24. Detail of shell layers in radial section of Platydon cancellata. HH homogeneous, CL = crossed-lamellar, PM = pallial myostracum, CCL = complex crossed-lamellar. Fic. 25. Radial section of both valves of Corbula gibba showing the periostracal flaps (PF) and sheets (PS) incorporated into the shell a characteristic of this family. Other letter- ing; CL = crossed-lamellar, CCL = complex crossed-lamellar, PM = pallial myostracum, L = ligament. SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 277 structures we have recognized by appearing grey instead of brown in thin section. It is probable that the outer layer may have been derived by the degeneration of a phylogenetically earlier prismatic layer. The middle layer consists of very fine primary lamels, arranged concentrically. There is a thin, prismatic pallial myo- stracum in Platydon cancellata but only a sharp change in shell banding in Mya arenavia. Distinct sheets of myostracal prisms are often found in the inner layer. GASTROCHAENACEA Three species of this small superfamily of rock borers have been examined struc- turally and mineralogically. The shell is aragonitic. The shell consists of two layers, an outer crossed-lamellar layer and an inner layer which may be complex crossed-lamellar or homogeneous. In all three species the outer layer has concentrically arranged lamels which pass transitionally inwards into homogeneous structure. The inner layer of Gastrochaena gigantea is complex crossed-lamellar but that of G. ovata and G. truncata is homogeneous, the former being distinctly lamellate. A thin prismatic pallial myostracum is present in all three species. HIATELLACEA (Plate 9, figs 1-4; Text-fig. 26) This is a very small superfamily consisting of four extant genera, one of which Panopea, is divided into two subgenera Panopea and Panomya. Five species have been examined structurally and mineralogically, all are aragonitic. In Panopea s.s. the shell consists of three layers, an outer simple prismatic layer which may be very thin, a middle homogeneous layer and an inner layer which may be homogeneous or complex crossed-lamellar. The simple prisms of the outer layer have rather irregular boundaries and orientations (Plate 9, fig. x). They vary in width between 30-50 pu and are made up of smaller platy crystallites between 0-5—- I'5 win width, which radiate from the central prism axis (Plate 9, fig. 3). The rate Fic. 26. Radial section of Panopea zeylandica. P = prisms, H = homogeneous, PM = pallial myostracum. oneusig ojeursig oI}euIsIg o1},eWISsTIg O1VeUsL oyeUsNg IzeUsig uryy ‘o1yeuIsLig oreusi dg oTeuUIsitg to;ONppV Temred (eee yee eoesoAy 278 sreqid pur sjooys JeoeI}SOAUL YIM Ieyourey -pessoio xe[duro9g sreyid pue sjoays yeoes}soAur Y}IM Ie][eUIe] -passoi9 xeyduiog Ie][eWIe] -pessoio xefdurog sieyid pue spueq JeoeI}SOAUL YIM IeTOUIE] -passoio xefduro9g sjeays peoesysohur YPM reyoUrEy -passo1o xajdurog sjeays [eor1}soAur YQIM Iepjeurey-passo1o xeduroo Ajouty sjeays [eoeIjsoAur YIM Ieyjaurey-passo19 xeduroo Ajouty JoAe] Jouuy Ie][eure]-pesso1g Ie]Joure|-passoig ouy * Ie][PUIe]-Pesso1n JAP] OTPPIN Ie][eUIe]-passo1g Te][eure[-passo1g Ie]]eure|-passoig Ie][owWe]-pesso1g snoauesowoy ‘rejnueis ‘Aoi snoauesowoy ‘rejnueis ‘Adar snosuesowoy qepnueis ‘Asin ioe] I9jNO VaOVAW gl alavy, ayuosery ayuosery ayuose1y apyUOseIy opymoseiy oyMOsely apuosery ASoyer9uryy Jopenoq purjsusan() sojdeny uede[ BIUIOFITED ureyig uleqig. Ayyeo0T uoss[O DUuYyscyjamy vjnqroz0hAvD ayepely vordoApay vjnqQao7 (IANO) 29918 nINGQ40g SPUIF] VSsv41I HjNGLOD pero) vyopjaquva UopKyw) gq snoeeuury DIDIUNA, VDATAT sneeuury DIADUAAD DATA setoads SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 279 of divergence from the axis is high, so that the crystallites appear almost horizontal in relation to the axis (Plate 9, fig. 2). The middle homogeneous layer at high magnifications is seen to consist of short crystallites aligned in two directions (Plate 9, fig. 4) suggesting perhaps a transition to crossed-lamellar structure. The outer shell surface of Panopea is ornamented by granules arranged into rows radiating from the umbo. In Hiatella, Cyrtodaria and Panomya there are only two layers. In all cases, the outer layer is homogeneous and excepting Panomya the inner layer is also homo- geneous. In Panomya the inner layer may be homogeneous with thin prismatic sheets or it may consist of complex crossed-lamellar structure. In a specimen of Hiatella arctica from Spitzbergen the inner layer consisted almost entirely of myo- stracal prism sheets. Prismatic adductor myostraca were seen in Panomya norvegica and Hiatella arctica. The presence of the outer simple prismatic layer in Panopea may be of considerable phylogenetic significance and this is discussed further in the conclusions. PHOLADACEA (Plate ro, figs 1-4; Plate 11, figs 1-3; Text-figs 27-29) Two families constitute this superfamily the Pholadidae and the Teredinidae; these are discussed separately. Eleven species of Pholadidae have been examined structurally and mineralogically ; all species examined consisted of aragonite. Two main types of shell structure were found in this family. In three species examined there was a three layered shell consisting of an outer simple prismatic layer, a middle crossed-lamellar layer and an inner complex crossed-lamellar or homogeneous layer. The outer layer consists of prisms (Plate 11, fig. 3) very similar to those found in Panopea (Hiatellacea p. 34); P+CL Fic.27. Radial section of Bayneacandida, as shown in Text-fig. 28 there is an interdigitation of the outer prismatic layer with the middle crossed-lamellar layer making differentiation at this magnification difficult. P = prisms, CL = crossed-lamellar, CCL = complex crossed-lamellar. oryeuIsiig oreUsiig o1ryeUISIIg, oreUIsitg oreusitg wimnovi}SOAUW [eed uryy ‘oryeustag ung} ‘oes ury} ‘o1VeUIsTIg umnorizsoAur [eed 280 snoouasoul0 x] snoauesoulopy opU0seIy snoauss0wo py snoouesoulopyy oyuosery sjeays wistid ZIM snoauesou04 IO Ie][eure[-passo1o xe[duroD snoouss0umloyY «= oy TOS eIW , Ie[[aure] passo1o xo[duioo 10 snosuasoul0fY += snoauesouroyy sutstid aydurts ay MOsIy Ie][ewWe]-pesso1o xe[duioo 10 snoauesouloyy + snosauesowofy suistid afduris ayuOsIy IoAV] IouUy Jah] ITPPUA IeAe[ IajngQ = ASoyerIOUTT VaOV TIaLVIH QI alavy snoauesowoy, snosuesowoy/1e[[eure|-pesso1y opuOseIy Ie}[eure]-passoio xejdui05D snosuesomoy/reyjoure[-pesso1j =» a TuOs eI Ie[jowrey]-pessoio xetdui05) snoauesowoy/re[joure|-pessoig, ayuosery ieAPy IouUyT JoAey{ 19jnQ Asoyes9uTyy VAOVNAVHDOULSVD 41 alavy (snoeuury) uleyug DIYIAD YIIAIVLET PIquinjoD (1apsueds) ysug onbyis viavpopc a (1a[8ueds) var7ams0u eas GHON (vimouvg) vadouvg Aqiemos elperjsny syvasnv vadouvng (preutey » Aond) purylesz MON voipunjiaz vadouvg Ayzyeo0'T saroeds ueyezeyy = AqiaMosg vywIUNnA, DUaDYIOYSY euleueg Aqiamos vyyno yuavyI04svy) eIpuy soAeysaq vajunsis nuanyr04svy Aypeo0T[ satoads SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 281 the prisms are irregular in size, length, and orientation in contrast to the more regular arrangement found, for example, in the Unionacea. The prismatic layer is not deposited continuously, for as seen in Text-fig. 28, the imbricating concentric ornament of this family is formed by a cyclical deposition and non-deposition of the prismatic structure (Plate 11, figs 1-2). A ventral ‘shoot’ of crossed-lamellar structure corresponds with each period of non-deposition of prisms. The crossed- lamellar layer is usually thin and the first order lamels short and coarse. The inner layer within the pallial line may be homogeneous or complex crossed-lamellar and is frequently lamellate and may contain sheets of myostracal prisms. In Zirfaea crispata there is also a three layered shell but the outer layer consists of grey homogeneous structure. The individual crystallites (Plate ro, figs 1 & 2) are approximately 5-10 p» long and 2-4 » wide with a slightly elongate shape. In this species the concentric ornament consists entirely of homogeneous structure. The structure resembles that of the Myidae and is conceivably derived from the simple prisms described above for other pholads. In all other species of Pholadidae examined there is a two layered shell, with the outer ribbing and ornament being formed from crossed-lamellar structure. The PM CGE Fic. 28. Detail of radial section of Baynea candida showing the alternation of prismatic and crossed-lamellar structure in the outer layer. SP = simple aragonite prisms, CL = crossed-lamellar, PM = pallial myostracum, CCL = complex crossed-lamellar. H CL Fic. 29. Detail of a radial section of Zirfaea crispata showing outer homogeneous (H), and middle crossed-lamellar layer (CL). 282 TAYLOR, KENNEDY AND HALL strong umbonal reflections characteristic of the Pholadacea are formed from complex crossed-lamellar structure. Deposits of myostracal prisms occur beneath the adductor, pallial and other muscle attachment sites. One species of Teredinae was examined structurally and mineralogically. The shell and tube are both aragonite. The shell is nearly hemispherical in shape and complicated as in the Pholadidae by apophyses, shelves and condyles associated with the wood-boring habit (Turner, 1966). The shell is basically two layered, with an outer crossed-lamellar (Plate ro, figs 3 & 4) and an inner complex crossed-lamellar layer. The outermost part of the outer layer shows strongly reflected growth lines and the crossed-lamels are con- sequently radially aligned. In addition the lamels are very fine and present an almost homogeneous appearance. The complex crossed-lamellar layer is restricted to the umbonal ridge. Pads of myostracal prisms were seen beneath the large posterior adductor and beneath the anterior adductor which is situated on the umbonal reflection. The ventral condyle has homogeneous structure. The calcareous tube which is secreted by the mantle surrounding the siphon tips consists of layers of irregular granular crystals about 5-10 » in diameter. Sub-Class ANOMALODESMATA Order PHOLADOMYINA PHOLADOMYACEA (Plate 12, figs 1-4) Because of lack of available material of this rare superfamily only a small fragment of Pholadomya candida was studied. It is aragonitic. The shell is basically three layered with an outer very thin simple prismatic layer a middle nacreous layer and within the pallial line an inner nacreous inner (Plate 12, fig. 1). The thin outer layer (Plate 12, fig. 1) also forms the surface granules arranged in radiating rows from the umbo. The middle nacreous layer appears to be ‘Treppen’ structure of Wise (1970) (Plate 12, fig. 4) and the inner nacre to be sheet nacre. The inner part of the inner layer consists of alternations of thin sheets of nacre with layers of myostracal prisms which form the dominant component (Plate 12, figs 2 & 3). Because of the very limited sampling it is not certain how typical the myostracal prism layers are of the whole shell. PANDORACEA (Plate 13, figs 1-4; Text-figs 30 & 31) Sixteen species were examined structurally and mineralogically. The shell is aragonitic throughout. Representatives from all seven families recognized by Moore (1969) were examined and two distinct structural arrangements were found. One of these, with both layers consisting of homogeneous structure, is found in the Thracidae alone; all the (10]8ueds) qourjsIpuy snooussowopy{ — snooussowoyy opuosery Urey DIDAASOA DIADPIGSND) (uoesprd 3 YIFID) snoouesouloTzy — snoauasouloyy oyuoseIy osulog sisuaurys viavpidsna (sres) yOuT|sIpuy snoauasouloyy _ snoaussowoyy oyuoseIy AeMION DIYIAD DIADPLGSND, (dorpusjso,, 2 SAN) Or}yeUIsLIg a1oeu 4904S aINeN snosuasowoyy ayuosery ure}yig DIvINUDAS DAMLOACT TUTYSousT 3 TALS o1jyeustig aroeu 420qS aroeu Ie[noyueT suistid afduits aqyuosery ATeqYI vajuasay 01004299 (zeqoSt.J) oreUulsiig aroeu 4204S aioeu IepNoyUeT suustid ofdurts apyuoOseiy orzuelyVy DUvISAKDYSAP VUPAOIYAI A (yo00[y 2 WOseI\-Poo\) UlY} ‘o1yeUIsIIg arpoeu 4a0qS aroeu IepnoyUeT sustid afdurts ayuosery sueulepuy vauanga voAlmnzy eoe1}SOAU [eed IaAv] IoOUUT IoAP] OPP IoAP] 193NO ASoyer9uryy Aypeoo0'T sotoads VaDVANOUOd Saat ee RRNA Yh a So Aaa Ca a, ee ata (Aq1amos) oneusiig Ie[[eure[-passoio xajdui0D — Ie][PWL][-passolg s}uoCseIy JOpenoy vyoumunov snjoydvivgd ojeUISHIg Ie]jawe[-pesso10 xefduiog =e] [auUe]|-pesso1n orpewusiig ayuosely UlepI_ snoeuury snjAjowp svjoyq oyeuIslig Ie[joure[-passo1o xajdwog = e[[aure[-pesso1p ojeusig ouosery Jopenoy BUT[OJN[ SISUaO]1YI SYIOYT oneWsig sIe[jeurej-pessoio xejdwiog = re[Jeure]-passo1p oeusiig ojuo0seiy uleyg (sneeuury) vpypuva vausvg Iojon eITTe, \ i Lat pe eS, eS IoAe] IOUT IaAP] IPP yoke, iayng = ABorerouTY «= APTTBOOT satoads eoelsoAyL Vaovdav 1OoHd 61 aAlaV (za[8ueds) qouystpuy snoauadowoyy _ snosuesomoy aymuosery ureqig DIDAISOA DIADPLGSND (uoaSptdg -p WUD) snoauasomoyzy — snoauasomoyy ayuosery oauiog stsuaurya viavpigsnd (sres) youyjsipuy snoauasomofy _ snoauesomoyy ayuosery APMION poyrav miapprgsng (doxpuaqsa 9 SAN) oneuisug a10eU 4204S aIDEN snoauasowoyy aqyuosery Urey: DyIDINUDAT DAMOAOT Turysausy Y APS t oryeustig aIOeU 3994S aioeu Ie[noyueT swsud ajduris ayuosery Ayes vaquasav 1jOryI99q (zayosta) onrusug aioeu 4aaqS aiseu IPpnoyUeT suisud ajduits ayiuoseiy onuely puvisadpysap DYpAomyad A | (q200TV 3 WOSETN-PooA\) ] uly} ‘oeUWsug aajoru 4a0qS aroeu IejnoyUe’y suistd ajduns ayuosery surmepuy VIUANGI DOALINT poviysoAur [ered aoAv] Iauuy raAhP] BPP I@AP] 1BINO ASopesrsuryy AQYEI0'T satoads VaAOVANOUOd Iz aTAV, | (Aerator) } snoaussomoyy snoauasouloyy «=: aus eI ureqig DIINSNSOYIUA DIIWAY T. (qorewre7T) ary, snoauasowoyy snoasuasomoy, = ayTuOSeIy ureyig, puyoasnyd DIIDAY T, arr, snoauasowoyy snoauasomoyy = a} UOSeIy ureyuig: POONA VxBAUOI DIIDAY T. o1yeuIsiigy aioeu yaays asoeuseynoquey susud aldwig oymuoSrry PIUIOFTLD “ds vkpsouuayy (Aamqyoqn4s) onewsug onjeuIsug aroeu jooys axeurepnonjuey susud ajdung = aytuosery eiyperysny sapromoun DuDy20a AT yorewre’] orusig o1yeusiig aloeu yaayS aioeurepNoqueyT suisud ajdung —ajtuosery SaTeAA “SN ppiqyy snasvyjop1aD suisud jeovsysofur pooMm onewsug pur aroeu yaays azeuiynoguey = susudajdug = aytuos ery PIUeUUISE pounusn} DAOPVA TT suustid jeorsysofur (preurrey 2p Aon) | o1yeulsug pur o1eu yaayg aroeuaynojuey = swstd aduig —ayuoSvay pueppny vyoiays DAOPoA TT suisud yeorsrjsofur | oryewisug oeWsiig pur sioeu jaoys aioe ayjnoyuey — suustid eydurr aqymuoser AqiaMos s1nadq vAopowy, [Nor [GUS “ Vv 4: IS S1nadq DAOpOA (snaeuurq) aroeu JaoyS asoeurepnoquey suisud adung o}tuosery DUYDUD DINUAIOT (aayostyy 2 ASSOID) areu 3a94S aneN susudaduig aymuosery elyperjsny ispsun vusapof{[o Jayoeunyos onewsud ‘ayy aioeu jyaays aioeurejnoguay stustad ayduirs ayuosery eoreure[ stappambanur viuo)qrdag aioeu yaaysS aioeurEnoyquey swsudoduig aymosery “W'S We}seq Av vjnauyia, vAOPUDT aioeu jaaysS arureynoyjueyT stusud ajduis ayuosely sajden (Furpoyy) vpiqyy vaopuvgy onewmsug oewsud ‘uy aieu JaayS aieurENOQueT swsud aduig aywoseiy sopenoy Aqianosg vjwnz4y véopung 10;9Nppy reured ; H——__.,—_—_——_ raXp] Jouuy IaA¥| Aa[PPUN rahe] tayng = Adoyeszouryy AqyRoo'y satsads poesqsoAy VaOVUOUNVd oz aTaV onPMUsUg WjeWISUg Ie[[aure[-passos9 xajdu10D — AP[JaWe]-passorg —- ayTuOSeay ureyligy (snaeuury) syvavu opedsaT snoauasowoy | oneusug Ij7eUIsIg IeTJawWe[-passo19 xajduiog e[Jaure]-passory ‘e[NURIS ‘AoIg — ayTUOS LIV ureyug (snavuuryq) wjodsz49 vevfarz uopny, oeWIsUg IPe[jaure[-pessos9 xajduroD _ Iej[aure[-passorg — aus eI ureyuig: DUuDiquiogso] vapLpvjOYT oneWSsUg Ie[JawWe]-passor9 xajdur0oy — ieyjaweEEpassory = ayuoSery pepruuy, (shoeuury) ypiys visaAv]y (Aqzamos) onewsug Ie}[auwre[-passo19 xajdui0> _— AP[[aWL]-posso1y apuosery sIOpenoy pypuruna sopoyqvav oneWsUg IepjewE-pesso19 xayduiog = re]Jawr]-passo1g oryeusig ayuoseiy uleyig snaeuury smpAjovp svjoyd NEWSU Ie[jaure[-passor9 xajdwioy 41] Jaure[-passo1y oryeusiig ayuosery ropenoq VULLOJ SISuaO]IYI SVJOYT oOAeUSUg Ie[Jawe|-passo19 xajdwioy re[Jaure|-passo19g oneusug a}uosery ulryug (sheeuury) ppipun? vauavg ; soyonppy reurea uN ,—~ JaXe] Jouu 4akr] a[PPUL aah c y c Ae Ae, y Saas I T AIPPU J 4a3nQ =AsopesoUNyY —- ABIPRIOT saroads : WaOvVdv10Hd 61 alav ee g =] Wr Roe Hr o ds. = TSM ee ee = Rep ond Ga 8 a ee oo SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 283 other families have a three layered shell consisting of an outer simple prismatic layer a middle lenticular nacre layer and an inner sheet nacre layer. The outer simple prism layer is very thin and frequently worn off much of the shell. In most species examined a thin pallial myostracum separated the middle and inner nacreous layers. In Myadora striata most of the inner layer consists of myostracal prisms. Radial rows of granules are present on the outside of the shell in many species. In the three species of Thracia examined, the two layered shell consists of homo- geneous structure in both layers. Two species were examined at high magnifications and the inner surface of the shell appears granular with irregular crystals about 3 p in diameter. In section these crystals are slightly flattened and have a slightly laminar arrangement. On the outside of the shell patterns of granules are seen (Tebble, 1966, fig. 103), when these are examined more closely they are seen to be isolated spherulitic structures (Plate 13, figs 1-4). These spherulites are made up of smaller crystallites (Plate 13, fig. 3) about 3 » in length. The spherules form columnar growths intercalated with layers of periostracum (Plate 13, fig. 2). Eventually as growth proceeds these spherulites merge together and the crystal arrangement passes into a uniform homogeneous structure. Fic. 30. Radial section of the flat right valve of Pandora albida. SP = aragonite simple prisms, N = nacre, PM = pallial myostracum. Fic. 31. Radial section of Cleidothaerus albida. SP = simple prisms, LN = lenticular nacre, PM = pallial myostracum, SN = sheet nacre. 284 : TAYLOR, KENNEDY AND HALL Order POROMYOIDA POROMYACEA (Plate 13, figs 5; Plate 14, figs 1-5) Seven species were examined structurally and mineralogically. The shell is aragonitic. This superfamilyis represented by three families, the Poromyidae, the Cuspidaridae and the Verticordidae. The latter have a three layered shell consisting of an outer, simple prismatic layer, a lenticular nacre a middle layer and a sheet nacre inner layer. The middle and inner layers are separated by a thin sheet of pallial myostracal prisms. The shell structure is generally similar to that of most of the Pandoracea; in Euciroa the prisms are irregular (Plate 13, fig. 5) and resemble those of Panopea (Hiatellacea). In Povomya granulata there is a three layered shell as above but the outer layer consists of granular homogeneous structure probably phylogenetically derived from a structural breakdown of simple prismatic structure (Plate 14, figs 2 & 4). The Cuspidaridae and some Poromyidae both have a two layered shell with homo- geneous structure in both layers. The granules of the homogeneous layers are about 2 p in size (Plate 14, fig. 5) and generally similar in appearance to those of the Thracidae. The pallial myostracum was indistinct in the species examined and shows a discontinuity rather than a distinct structure. CLAVAGELLACEA (Plate 15, figs 1-5; Text-fig. 32) This is a small highly aberrant superfamily which consists of three extant genera Clavagella, Humphreysia and Penicillus which show a progressive fusion of the true shell with the calcareous tube. The valves are however free when young. Three species were examined structurally and mineralogically ; the shell and tube are both aragonite. In Clavagella aperta the valves are fairly large and only one is fused with the siphonal tube. The valves consist of a thin simple prismatic outer layer with a sheet nacre inner layer. Thin sheets of myostracal prisms are secreted beneath the muscle attachment scars. Valves of a juvenile Humphreysia strangei were examined, these had a simple prismatic outer layer and inner sheet nacre layers (s). The outside of the valves is finely pustulate. In Penicillus s.s. the true shell is seen as two valves occupying a saddle shaped area incorporated into the side of the tube (Text-fig. 32). The tube is extended pos- teriorly as a hollow cylinder and anteriorly as a perforated disc (the watering pot). The true valves are covered by a thin periostracum which is inserted from the outside of the shell to line the inside of the tube at the edge of the saddle shaped area. The valves consist of two layers, an outer extremely thin simple prismatic layer with an TABLE 22 satadayL SUVTITId TVOVULSOATL SNOANADONOY, UVTIANVI-GaSsOuD XATAWOD UVTITANVI-GaSsouy daLvitoy AUOVN LAAHS AMOVN UVINOIINAT SWSIud ALISOdNWOD SWSIUd ATMWIS ALIOTVD SWSIud ATMWIS ALINOOVAY ADOTVUANIPL SUPERFAMILY 4 x eo Nuculacea < Nuculanacea satan Ee Sg) Ree SARA CIA tots, as) ca MTR Ca eal BAIR ns ws sco Saget (RLY 3. SUVTIId TVOVLSOATY So ey tele See erp oho SR RpGibd eetteraetbei bn try (oy 1st ee ren meats snoaNaooWoRT SOME FS pi Gag nS HOG ECCS ee a Ye a ‘ AVITANVT-dassoud XATAWOD se A PP RP Pe pe ddd a od YVTTANVT-Gassou) St A Rt KK RMR OM aax1vi10q Buc, WRCHe cae cs cale Men ae Ra ie utes i ceOsamen O: oMiao cet ons AUOVN LAaHS hog eck S. ciMp ONG fog oe BRACE. oy etn oe gear cae orkc. ect etig ce AYOVN UVINOLINET Pai. ace go ee hMia ola ple hen Bo edo auooposemomany. ci ance Goud Be NSIT a : ame Ta) h5 Se oth, tha: Ouee eH TG Cet Ae enna cre Bree Mb o aod v7 Q BS SWSId ATAWIS ALIOTVD SB tm Rh cM eC TS SOican e CSO CDM RAM. “AF: 0 tena oc swSIud AIdNIS ALINOOVAY aA te "ost ole Eee VE SEN ce cio coma Ipetiers. etero oo art 0 eA AA OCVO000N0 > AOOTVHENIN | eg oeopiwpA7y =e, . paspuojday pa2ouInn] 222D10¢d || aa, « oaaphw Da 2DUaDYIOL}SOS) pasDppjoug MYOIDA D9DD||a;DIH paspAwoppjoudg Da 2DIOpU_Y paap||aHDA0|>5 paspAwolog PHOLADO MYOIDA BadeUIIag AUIOS PUL RadvaIsO ayy WIP]-Ppassolo a[ppru savy YIIYM esIepE[OY, aulos ynq snoauasouroy Ayureul are eadeT[azeIF] VY. uMOoUyU) rg ainjonys pazyeyos uaaI5 eu/susud a 1s aWOSe1y = (snonunuos) antg asurio MoroR Aj1umeyradns ay} uo pasodut Injon4ys [Jays ureul oy PIAE pue Aroysty peotdojoay ££ “ong , a3 parpkwosoy 2 = a SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 287 important in the Ordovician where most of the radiation of the major lineages and shell structure combinations probably took place. However because of dissolution, recrystallisation and replacement the original shell fabrics have disappeared or have been altered out of recognition. The relationships of the various bivalve superfamilies are discussed in terms of shell structure variations below. Subclass PALAEOTAXODONTA Two superfamilies belong in this group; the Nuculacea are generally regarded as the most primitive living bivalves (Yonge, 1959) but the other superfamily the Nuculanacea are considered to be as highly specialized as any similar group through- out the bivalves (Yonge, 1959). The origin of the Nuculacea can be traced back to the Upper Cambrian Ctenodonta (Cox, 1959) and the group appears to have remained relatively unchanged morphologically throughout their subsequent history. The nacreous and composite prismatic shell is different from that of any other family. However, we consider that the difference between simple aragonite prisms (possibly the ancestral condition) such as found in the Unionacea and Pholadomyacea and the composite prisms of the Nuculacea is slight and arises from differences in the degree of mantle reflection at the shell margin. The extant Nuculanacea have a homogeneous shell but as shown by Cox (1959) and Taylor, Kennedy & Hall (1969) this has not always been the case. Subclass CRYPTODONTA Solemya has been considered to be a protobranch (Palaeotaxodonta) by Yonge (1939, 1959) but as Newell (1965) has pointed out, it is becoming increasingly apparent that the Solemyacea have been separated from the rest of the protobranchs from at least the Devonian and are not obviously related to the Nuculacea. The shell structure and in particular the character of the outer prismatic layer, is distinc- tive, but nevertheless can be readily derived from simple aragonite prisms. Our observations tend to support Newell’s opinion that the Solemyacea belong to a separate subclass the Cryptodonta. The extinct Palaeozoic order Praecardioida is placed in the Cryptodonta, but there is very little evidence of any relationship to the Solemyacea. We have no shell structure information on this group. Allen & Sanders (1969) have recently described the anatomy and discussed the affinities of the Recent genus (Nucinella classified in the Limopsacea, in the Treatise) which they consider to be a monomyarian ‘solemyid’ and possibly related to the extinct actinodont group (i.e. Cycloconchacea). Although there are several anatomical resemblances of Nucinella to Solemya other characters resemble those of the Nuculacea. Our observations of the shell structure show that it is homo- geneous structure similar to that of Nuculana but unlike Solemya or the Nuculacea. But 1f Nucinella is either a nuculacean or a solemyacean then reference to Text-fig. 33 will show that both these groups were probably derived from a cycloconchacean ancestor. c* 288 TAYLOR, KENNEDY AND HALL Subclass PTERIOMORPHIA The Arcacea are generally thought to be derived from the Cyrtodontacea of the lower Ordovician (Cox, 1959, 1960) but as pointed out by Morris (1967) the connec- tion is not firmly established. Newell (1954) has considered that the ‘cyrtodontids’ are also the ancestors of the Pteriacea, Pectinacea and also possibly the Mytilacea. These latter groups probably became separate from the ‘“‘cyrtodontid stock” rather earlier (Text-fig. 33). The Arcacea have a crossed-lamellar and complex crossed- lamellar shell structure with tubules and myostracal pillars. This structure is very different from that of the rest of the Pteriomorphia, but similar to that of some heterodonts such as the Carditacea. The only character which is common between the Arcacea and the rest of the Pteriomorphia is the filibranch gill and it seems to us that there is no close relationship between the groups and the Arcoida (Arcacea & Limopsacea) should possibly be considered as a separate subclass related to the Heterodonta. This does not of course deny a once common ancestry. The Mytilacea have a very distinctive prismatic, calcitic, outer shell layer, some- times called ‘fibrillar’ (Oberling, 1964). This particular structure is found in no other bivalve group. The work of Osborn (1970) on mammalian teeth has shown that all the different prism-like structures may not be very different from each other. The Mytilacea may have arisen directly from the lower Ordovician-Permian family the Modiomorphidae and have no apparent derivatives. Newell (1965) placed the Pinnacea in the order Mytiloida (implying relationship) but the simple calcite prisms, general shell form and anatomy suggest derivation from the Pteriacea. As mentioned above the Pteriacea and the Pectinacea are both considered to have been derived from a cyrtodontid ancestor (Cox, 1960; Newell, 1938). Although they have different shell structures this does not rule out a common ancestor. The occurrence of an outer prismatic layer in oysters, the early post larval stages of some pectens (Jackson, 1890) and in some species of Propeamussium suggests that the foliated layer in these forms may have originally been derived from aragonite nacreous structure by a change in the calcium carbonate polymorph. The super- family Ambonychiiacea which ranges from middle Ordovician to upper Devonian has been extensively discussed by Pojeta (1966); it includes many ‘Pteria’-like forms. Recently we have examined an Ambonychia from the upper Ordovician (Ashgill) from near Girvan, Scotland which has some shell structure preserved. As might be expected it showed nacreous inner layers, but unfortunately the outer layer was recrystallised, but was probably calcite prisms. Newell & Boyd (1970) have recently described the earliest known members of the Anomiacea, from the Permian. This superfamily is probably derived from the Pectinacea. The same is probably true of the Limacea. The Ostreacea first appeared in the Permian and were probably derived from a Pectinacean ancestor the Pseudomonotidae (Newell, 1961; Newell & Boyd, 1970). The shell structure characters support this suggestion. SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 289 Subclass PALAEOHETERODONTA The Unionacea and Trigonacea have a very similar shell structure of aragonite simple prisms and lenticular and sheet nacreous layers. There has long been debate as to the possible relationship of these two families (Cox, 1960). The anatomical evidence suggests that they may be distinct groups, whereas the palaeontological evidence is ambiguous and unsatisfactory. As well as the morphological and shell structure similarities, they have a character in common which is usually overlooked ; this is the possession of calcareous gill spicules recorded for the Unionacea by Ridewood (1904) and for the Unionacea and Trigonacea (Atkins, 1938). They are the only bivalve superfamilies to possess these spicules. Subclass HETERODONTA The Lucinacea are known from the Silurian to Recent and can be traced through the Babinkacea back to the middle Ordovician (McAlester, 1965, 1966). McAlester has argued that the Lucinacea are a distinct bivalve group and should be considered as a separate subclass. Certainly the Lucinacea have been distinct for a long period of time and only the Leptonacea and Cyamiacea can be related to them. However Boss (1969) considers from anatomical and shell morphological evidence that the Lucinacea are closely connected to other bivalves of the heterodont subclass. The Lucinacea have a three layered shell of an outer composite prismatic layer, a middle crossed-lamellar layer and an inner complex crossed-lamellar layer. This combina- tion is also found in the Tellinacea and some Veneracea. The shell structure evidence thus supports the opinion of Boss (1969) that the Lucinacea belong to the Heterodonta, but reference to Text-figure 33 will show that they have been distinct from the rest of the heterodont stock for a long time. The Tellinacea are known from the Upper Triassic to Recent but their phylogenetic relationships are obscure. As noted above the three layered shell structure is found in the Lucinacea and Veneracea. The Solenacea may have arisen from the Tellinacea in the late Cretaceous or early Cainozoic (Davies, 1935; Morris, 1967). In the process they must have lost the outer composite prismatic layer, as indeed have some of the Tellinacea. The Astartacea, Carditacea, Chamacea, Cardiacea, Tridacnacea, Mactracea, Arcticacea, Veneracea, Corbiculacea, Dreissenacea and the Glossacea all appear to be generally related (Text-fig. 33). The shell structure is generally similar in all these groups with only relatively small variations (Table 22). The most important variation is the three layered shell in some Veneracea. Most of these families arose in the Mesozoic and Cainozoic, and Stanley (1968) has discussed this spectacular radiation. The most striking trend is the appearance and extensive radiation of the infaunal siphonate feeders, which Stanley relates to the development of siphons and the closure of the mantle cavity by mantle fusion. Most of the families involved in this radiation have a two layered shell of crossed-lamellar structure and complex crossed-lamellar structures. In some families one or both layers may consist of homogeneous structure, but in these cases it is obviously derived from the structures mentioned. 290 TAYLOR, KENNEDY AND HALL The Mesozoic Veneroida were probably derived from either the Crassatellacea (Stanley, 1968), which first appeared in the Devonian, or from the Carditacea which also appeared in the Devonian (Morris, 1967). These two families probably have a common origin in the lower Palaeozoic from a cyrtodontacean stock (Text-fig. 33). Yonge, (1969) has recently stressed the similarities between the Crassatellacea and the Carditacea. The Chamacea which first appeared in the upper Cretaceous are thought on the basis of shell structure and anatomical characters to have been derived from the Carditacea (Kennedy, Morris & Taylor, 1970). The Cardiacea first appeared in the Trias, but no obvious ancestor can be cited from older rocks. The Tridacnacea can be readily derived from the Cardiacea in the Eocene or late Cretaceous (Stasek, 1962). The Mactracea appear similar to the Cardiacea in shell structure details but there is no real evidence of any relationship. The Arcticacea, Veneracea and Corbiculacea may have been derived from the Jurassic forms Pseudotrapezium and Pronella (Casey, 1952; Morris, 1967). The Arcticacea and the Veneracea are probably very closely related. Although Arctica shows a homogeneous shell structure traces of crossed-lamellar structure may sometimes be seen. Other members of the Arcticacea show crossed-lamellar and complex crossed-lamellar structure. The Veneracea show two distinct types of shell structure; this may be a result of the loss of the outer composite layer in some forms or a polyphyletic origin for the Veneracea. The Dreissenacea are a group of fresh water byssate anisomyarian bivalves which appeared in the Cainozoic. Because of their mytilid-like shell, their relations have remained obscure, but it has been realized for some time that they are unrelated to the Mytilacea (Yonge & Campbell, 1968). The shell structure shows great similarity in micro-details to that of the Corbiculacea and it is reasonable to suppose that the Dreissenacea arose from the fresh and brackish water Corbiculacea. Morton (1970) has made a study of the morphological changes seen in fossil forms, demonstrating a progression from the Corbiculacea to the Dreissenacea. However, the idea of some relationship to the Mytilacea has not entirely disappeared (Purchon & Brown, 1969). Subclasses MYOIDA and PHOLADOMYOIDA The Myoida and Pholadomyoida although classified in separate subclasses show obvious similarities and we consider that all the superfamilies in these subclasses can be derived from a ‘“‘pholadomyacean”’ stock which has been in existence since the middle Ordovician. Other workers however, consider the resemblances to be the result of morphological convergence (Runnegar, 1966, 1967). Pholadomya s.s. has a shell structure of simple aragonite prisms and middle and inner nacreous layers. Panopea of the Hiatellacea (Myoida) is anatomically and morphologically very similar to Pholadomya (even including surface granules) but has an outer prismatic layer, a middle homogeneous and an inner complex crossed- lamellar later. Other members of the Hiatellacea have shells consisting of homo- geneous structure alone. It seems very probable that the Hiatellacea have been SHELL STRUCTURE AND MINERALOGY OF BIVALVIA 291 derived from the “‘pholadomyoid” stock. Some Pholadacea have a shell structure of simple prisms, crossed-lamellar and complex crossed-lamellar layers. The structure of the outer layer closely resembles that of Panopea. It seems that the Pholadacea may have arisen from the Pholadomyacea in the early Jurassic; the genera Myopholas and Giradotia would seem to be transitional forms (Morris, unpub.). The Myacea consist of two families, the Myidae (Palaeocene-Recent) and the Corbulidae (L. Jurassic-Recent). It does not seem very likely on anatomical and shell morphological grounds that the Corbulidae gave rise to the Myidae. This is supported by the fact that the Corbulidae have a two layered and the Myidae a three layered shell. The origin of the Corbulidae might perhaps be found in the Permian pholadomyoid forms such as Pyramus amd M egadesmus (see figures in Runnegar, 1967). The Myidae would seem to have been independently derived from the ‘pholadomyoid” stock at a much later date. Most Pandoracea have a shell structure of simple prisms and two nacreous layers, this and anatomical characters suggest a derivation from the Pholadomyacea in the Trias or lower Jurassic. The Thracidae (family of Pandoracea) have today a largely homogeneous shell, the outermost part of which retains a vestige of prismatic structure. However in the Cretaceous the Thracidae had a prismato-nacreous shell and apart from shell structure there is little to differentiate the Thracidae, from other Pandoracean families such as the Laternulidae. The origins of the Poromyacea are obscure but certainly the Cuspidariidae can be traced back to the Trias (Cox, 1960; Morris, 1967) and have probably arisen from the Edmondiacean genus Solenomorpha. The Edmondiacea appear to be a hetero- geneous Palaeozoic group closely related to the Pholadomyacea. Some of the Poromyacea have a prismato-nacreous shell and others are entirely homogeneous. Although the superfamily has a septibranch gill there are many anatomical resem- blances to the Pandoracea. The Clavagellacea are a highly aberrant group but anatomical characters, the nacreo-prismatic shell and the surface granules suggest a close affinity with the Pandoracea. Evidence from the Monoplacophora (Erben, et al, 1968), Archaeogastropoda (Wise, 1970; Wise & Hay, 1968), Nautilus (Grégoire, 1962) and some of the oldest bivalve lineages strongly suggests that the ‘‘primitive”’ shell structure of the bivalves is a simple aragonite prism outer layer and middle and inner nacreous layers. Sub- sequent evolutionary radiation of the shell structures has been a result of the increased exploitation of different habitats and different modes of life. Taylor & Layman (1972) have stressed the functional significance of bivalve shell structures and present evidence correlating structure with mode of life. However we need much more information on the course of evolutionary change in shell structures and it is probable that in time sufficient well preserved Palaeozoic material will be discovered in order to document these changes. 292 TAYLOR, KENNEDY AND HALL ACKNOWLEDGEMENTS We are very grateful for the constant interest, assistance and advice of Dr. Noel Morris, particularly in the preparation of the conclusions. We should also like to thank Dr. R. P. S. Jefferies, Mr. R. J. Cleevely and Mr. C. P. Palmer for advice and assistance in various ways. We are especially grateful to the staff of the electron microscope unit at the British Museum (Natural History) under the direction of Mr. B. Martin for patient advice and assistance. REFERENCES ALLEN, J. A., & Sanpers, H. L. 1969. Nucinella servei Lamy (Bivalvia: Protobranchia) a monomyarian solemyid and possible living actinodont. Malacologia, 7 : 381-396. Atkins, D. 1937. On the ciliary mechanisms and interrelationships of Lamellibranchs. Part II: Types of Lamellibranch gill and their food currents. Q. Jl. microsc. Sci., 79 : 375-421. — 1938. On the ciliary mechanisms and interrelationships of lamellibranchs. VII, Latero- frontal cilia of the gill filaments and their phylogenetic value. @Q. Jl. microsc. Sci., 80: 346-436. Boacaitp, O. B. 1930. The shell structure of the mollusks. K. danske Vidensk. Selsk. Skyr., 2 : 232-325. Boss, K. J. 1968. New species of Vesicomyidae from the Gulf of Darien, Caribbean Sea (Bivalvia, Mollusca). Bull. mar. Sci., 18 : 731-748. 1969. Lucinacea and their heterodont affinities (Bivalvia). Nautilus, 82 : 128-131. Boyp, D. W., & NEwett, N. D. 1968. Hinge grades in the evolution of crassatellacean bivalves as revealed by Permian genera. Am. Mus. Novit., 2328 : 52. Casey, R. 1952. Some genera and subgenera, mainly new, of Mesozoic heterodont lamelli- branchs. Pyvoc. malac. Soc. Lond., 29 : 121-176. Cox, L.R. 1959. The geological history of the Protobranchia and the dual origin of taxodont Lamellibranchia. Pyvoc. malac. Soc. Lond., 33 : 200-209. 1960. Thoughts on the classification of the Bivalvia. Pyoc. malac. Soc. Lond., 34 60-88. Davies, A.M. 1935. Tertiary Faunas. Vol.1. The composition of Tertiary faunas. 406 pp. London. ErRBEN, H. K., Frays, G., & Stent, A. 1968. Uber die Schalenstruktur von Monoplacophoren. Akad. Wiss. Lit. Mainz. 1968, no. 1 : 1-24. GHISELIN, M. T., DEGEns, E. T., SPENcER, D. W., & Parker, R. H. 1967. ins ->- os a ON Fic. Fic. Fic. Fic. Fic. WwWwne = Be PLATE 15 All figures are scanning electron-micrographs of Penicillus sp. Inner surface of nacreous layer. _ 2,300. Fractured section of nacreous layer of valves showing sheet nacre. _ 2,400. Fractured section of the tube showing the flat platy Gaal Mes * 2,400. As Fig. 3 but showing the stacks of platy crystallites. < 1,200. Surface granules, covered by periostracum on the paisiae of the true valves. granules are arranged in rows which radiate from the umbo. x 1,300. The Bull. By. Mus. nat. Hist. (Zool.) 22, 9 PLATE 15 " Z “J ! “a ga” = 7 ‘i - . é eae ™ = 7 : r | : , vy P r r A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) Kay, E. Arison. Marine Molluscs in the Cuming Collection British Museum (Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 1965. (Out of Print.) £3.75. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and Valenciennes. Pp. 180; 11 Plates, 15 Text-figures. 1967. £4. TaytLor, J. D., KENNEDY, W. J. & Hatt, A. The Shell Structure and Mineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates, 77 Text-figures. 1969. £4.50. Haynes, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur) 1962-1964. {10.80. aut Printed in England by Staples Printers Limited at their Kettering, Northants, establishment INDEX TO VOLUME 22 The page numbers of the principal references and the new taxonomic names are printed in bold type. Abramis . 4 , 9 48 abyssinica, Genetta 240, 247 abyssinicus, Arvicanthis nilotieus 229-230 Tab. 6 (opp. p. 263), Pl. 3 219, 220, 222, 228-229, 246, 248 Tab. 6 (opp. p. 263) Tab. 4 (opp. p. 260) Tab. 19 (opp. p. 282) Acanthocardia Acomys aculeata, Acanthocardia aculeata, Cardita . acuminata, Parapholas . acuminatus, Rhinolophus 185, 194 acuminatus audax, Rhinolophus_ . 5 mG acuminatus sumatranus, Rhinolophus. . 185 acuta, Dussumieria < ¢ 5 80 adversus, Myotis . d 9 188, 194 adversus carimatae, Myotis . ° - 188 adversus moluccarum, Myotis 5 188 aeruginosus, Lithobius . I14, 117, 133-134 aestuarius, Gilchristella 65, 68, 69 aethiopicus, Paraechinus 4 3 246 aethiopicus, Phacochoerus. . ale, 221, 222, 243 aethiops, Cercopithecus .220, 221, 225 afer, Orycteropus 217, 242 afer, Papyrocranus 3, 5, 7, 18-20, 21 affinis, Carditamera Tab. 4 (opp. p. 260) affinis, Pitar Tab. 15 (opp. p. 270) affinis macrurus, Rhinolophus c 182 africana, Dreissena 268, Tab. 12 africana, Ilisha_ . ; 0 . A ola africanus, Equus . 9 0 a 2K) agilis, Lithobius 113, 129, 143 agilis pannonicus, Lithobius . F 5 113 Agripoma Tab. 15 (opp. p. 270) alata, Venus Tab. 15 (opp. p. 270) albella, Sardinella : : 63 albicauda, Ichneumia Di, 221, 240 albida, Cleidothaerus Tab. 20 (opp. p. 282), 283 albida, Pandora Tab. 20 (opp. p. 282), 283 albipes, Praomys . « 221, 234, 247, 248 albipinnis, Taphozous longimanus . 5 182 albocaudata, Ruane 4 2477) Albula 5 9 “ a 8, 10, 44 Alcelaphus . 216, 244, 245 alexandrinus, Rattus rattus 6 : . 235 Allopolybothrus . 5 : 3 » 108 Alosinae. = 6 4 + 70-72 alosoides, Hiodon Shah ye alpestris, Lithobius dentatus! 108, 109 alpini, Procavia habessinica . 241, 242 alpinus, Lithobius 130, 143 alternatum, Laevicardium Tab. 6 (opp. p. 263), Pl. 3 altitudinis, Cynopterus eae ETO amazonica, Ilisha 5 5 6 Gif Ambonychia a ¢ : 0 zoe amethystina, Caryocorbula_ . e27e Amia . 3 . 5 : 8, 10, 48 Amiantis Tab. 15 (opp. p. 270) Ammodorcas : : 5 246 amnicum, Pisidium amomoides,, Myochama amphibius, Hippopotamus 219, 242-243 amplexicaudatus, Rouseltus . 173, 193 amplexicaudatus amplexicaudatus, Rouseltus 173 Tab. 20 (opp. p. 282) angasi, Offadesma Tab. 20 (opp. p. 282) anguilloides, Mormyrops 3, 6, 7, 38-40 angulatus, Cynopterus . : : = 2o4! Tab. 14 ‘(opp. Pp. 269) Tab. 20 (opp. p. 282) anatina, Laternula angulatus, Cynopterus brachyotis . 6 175 angulatus, Cynopterus sphinx 175-179 annae, Dosinia Tab. 15 (opp. p. 270) annectans, Myotis : 188-190, 193 annectans, Pipistrellus . 188, 189, 194 annectens, Pteropus hypomelanus . ee Anodontostoma . : 6 9 O 81 anomala, Polymesoda Tab. 14 (opp. p. se Pl. 6 [anomalata] Anomalocardia 270, Tab. 15 Anomalodesmata . Fs 5 a en zs2 Anomiacea . Tab. 22 (opp. p. 284), 288 antarcticum, Cyamium . : : ae Z00 antillarum, Crassatella . Tab. 5 (opp. p. 261) anubis, ;, Ale 2 219, 220, 221, 222, 225-226 aperta, Clavagella . 4 284 Arapaima - 3,5, 7, 14-16, 20, 50, 52 Arcacea Tab. 22 (opp. p. 284), 288 Archaeogastrepoda 0 5 c a aia Arcoida . : . 286, Fig. 33 Arctica Tab. 11 (opp. p. 268) arctica, Cuspidaria . 282, Tab. 21 arctica, Hiatella . 279, 280 256, 268, Tab. II (opp. p. 268), Tab. 22 (opp. p. 284), 289 arcuata, Pandora Tab. 20 (opp. p. 282) arenaria, Mya E : 278) argentea, Pecchiola Tab. 21 (opp. p. Bess argyrotaenia, Clupea 4 Arcticacea arkelli, Thryonomys_ . A c eee) os aristotelensis, Parasilurus. i : 53 Artiodactyla 242-244 Arvicanthis 217, 229-232, 246, 247, 248 Asaphis Tab. 10 ° (ope p. 266), Pl. 5 Ascaridida . I51I-170 asiaticus, Bodo . F b 5 5 99 asperatus, Lithobius’ . elas asperrima, Lioconcha Tab. 15 5 (opp. P- 270), 274 Astartacea . : é 289 296 INDEX Astarte Tab. 5 5 (ope. p. 261), 262, Pls 1 and 2 Astartidae ni 261-263 Atactodea Tab. 8 {oep- p- 264) atricauda, Clupea 2 80 audax, Rhinolophus acuminatus 2 b 185 aulacopus, Lithobius. . i b20) auratus, Engraulis aureus, Canis australe, eeguissar haa australis, Cardita australis, Gouldia "50, 71-79, 82, Pl a 217, 221, 239 Tab. 6 (opp. p. 263) Tab. 4 (opp. p. 260) Tab. 15 iPER: p. 270) australis, Panopea 3 . 280 baelama, Thrissina 9 0 0 5 82 Bagrus c : é c 53 baloghi, Lithobins z é : ah SEES balthica, Macoma Tab. 10 (opp. p. 266), 267 bane, Petrocephalus 3316; /7; S9n37, Barnea 279, 281, Tab. 19 (opp. p. 282), Pl. 11 beaulieui, Scotomanes (Parascotomanes) Igt bebe, Hyperopisus 3, 6, 7, 40-44 beisa, Oryx gazella 2 222, 244 bianalis, Spratellomorpha-. 8 e 68 bicirrhosum, Osteoglossum_ . -3, 5,7, 12-13 bicolor, Crocidura 3 5 5 225 biradiata, Solentellina . Tab. 10 (opp. p. 266) Bivalvia 4 253-294, Pls 1-15 blanfordi, Hesperoptenus ci 191, 194 blanfordi, Sphaerias z 181, 194 blicki, Arvicanthis 9 é > As) Bodo . é 87- 102, Pls 1-6 boehmi, Equus burchelli 216, 242 boelama, Engraulis 5 a 82 borealis, Astarte . Tab. 5 (opp. p- 261), 263, Pl. 2 borealis, Cardita . Tab. 4 (opp. p. 260) borealis, Lucina Tab. 1 (opp. p. ae borneensis, Coilia 5 borneensis, Rhinolophus 183, 185-187, ee oe borneensis spadix, Soe o : 6 187 Bothropolys E : a ra ea bottazzii, Gpelacomysis. 207, 208-209 Bovidae 243-244 brachyotis, Cynopterus 175-179, 194 brachyotis altitudinis, Cynopterus . : 179 brachyotis angulatus, Cynopterus . a 175 brachyotis brachyotis, Cynopterus 179-181 brachyotis hoffeti, Cynopterus D é 177 brachyotis insularum, Cynopterus . é 177 brachyotis scherzeri, Cynopterus . cee sf brachyotis titthaecheilus, gsr! 2 bs brachysoma, Clupea. : 5 80 brachysoma, Pellona_. F 3 81 brachysoma, Sardinella a . 63, 80 brandii, Dreissena Tab. 12 (opp. p. 268) braziliana, Anomalocardia Tab. 15 (opp. p. 270) breviceps, Engraulis. 3 5 82 breviceps, Setipinna. : Fi 82 brevis, Myadora . Tab. 20 (opp. p. 282) brockmani, Praomys fumatus 234-235 brockmani internus, Claviglis 3 A 236 broggi, Solecurtus Tab. 10 (opp. p. 266) Brooker, B. E. . 87-102, Pls 1-6 brucei brucei, Heterohyrax 242, 248 brucei hararensis, Heterohyrax 6 nea bucculentus, Lithobius . 7 125-126 buchholzi, Pantodon 5, 7, 16-18, 50 bulgaricus, Lithobius_ . “ 2 en tAG burchelli boehmi, Equus 216, 242 burmanica, Clupea : c 70 burneti, Tellidora Tab. 10 (opp. p. 266) buselaphus, Alcelaphus 216, 245 buselaphus swaynei, Alcelaphus 216, 244 buxtoni, Tragelaphus . 6 c e247) cadornae, Pipistrellus 191, 193 cahirinus, Acomys 228, 229 cahirinus cineraceus, Acomys 4 5 Bh) calcaratus, Lithobius 109-110, 114, 116, 136, 137 calcarea, Macoma Tab. 10 (opp. p. 266) calcarina, Tellina . 5 : 5 Fe 266 calyculata, Cardita Tab. 4 (opp. p. 260), 261 Calyptogena Tab. 11 ere: p. 268) calva, Amia : SS LO camelopardalis, Giraffa 5 . - 245 Campylomormyrus - 3,6, 7, 33-35, 49, 49, 50 cancellata, Platydon 276, 277, 278 candida, Barnea 279, 281, Tab. 19 (opp. p. 282), Pl. 11 candida, ee 282, Pl. 12 Canidae A F 2 2 . 239 Canis . 5 2 “ . 217, 221, 239, 247 cantoris, Coilia . : $ 83 canus, Pteropus hypomelanus “ 3 174 capensis, Lepus 226, 227, 246 capensis starcki, Lepus 3 £ 226 Capra 3 5 5 . 246 Cardiacea 255, Tab. 6 (opp. p. 263), 289 Cardilia Tab. 8 (opp. p. 264) Cardita Eco Tab. 5 (opp. p. 261), Pl. 2 Carditacea . 255, 260—261, Tab. 4 (opp. p. 260), Tab. 22 (opp. p. 284), 289 Tab. 6 (opp. p. 263) Tab. 10 (opp. p. 266) . 137-138, 139, 143 Cardium carinaria, Strigilla carinatus, Lithobius Carnivora : : 2 239-241 Caryocorbula 3 eB = E278 caschive, Mormyrus 3, 6, 23, 23-28 castaneus, Scotophilus . : : 192 castrensis, Lioconcha Tab. 15 (opp! Pp. 270), 273 catharia, Agripoma Tab. 15 (opp. p. 270) catostoma, Petrocephalus 3, 7) 35, 36, 37 caudatus, Bodo, 87-102, pls 1-2, 4-6 Cerastoderma Tab. 6 (opp. p. 263) Pl. 3 Cercopithecus F 220, 221, 225 cerina, Gouldia Tab. 15 (opp. p. 270) chabaudi, Cucullanus . 5 3 154 chacunda, Anodontostoma . F < 81 chacunda, Chatoessus . P c é 81 INDEX 297 Chama 258-259 Chamacea 255, 258-259, Tab. 22 (opp. Pp. 284), 289, 290 Chamalea . : 0 272 chanleri, Recents ‘ : 3 231, 232 chapra, Clupea_ . 6 5 a 81 chapra, Gadusia . 0 . 7 2ox chaseni, Rhinolophus 183, 187, 194 Chatoessus . 59, 73-74, 81 chiloensis, Pholas Tab. 19 (opp. p. 282) Chilopoda 103-150 Tab. 21 (opp. p. 282) Tab. 15 (opp. p. 270) chinensis, Cuspidaria chinensis, Cyclina Chione Tab. 15 (opp. p. 270), 272 Chioninae Tab. 15 (opp. p. 270), 274 Chionopsis . 3 Tab. 15 (opp. p. 270) Chirocentridae . ° : B 6 80 Chirocentrus 9 : o 5 a 80 Chironax 3 ‘ 181, 193 chitala, Notopterus : : : r 18 Chlamydoconchacea 255, 259, Tab. 22 (opp. p. 284) ciliatum, Cardium Tab. 6 (opp. p- 263) cineraceus, Acomys cahirinus 220) cinnamomeus, Lithobius : 135-136 Circa . 2 : - Tab. 15 (opp p. 270) Circinae Tab. 15 (opp. p. 270) Circomphalus Tab. 15 (opp. p. 270) civetta, Viverra 221, 240 clarkei, Ammodorcas. . A = 246 Clavagellacea 256, 284, Tab. 22 (opp. p. 284), 291 Clavagella . 0 0 5 é 6 ys Claviglis . 5 . : , : 236 Cleidothaerus 282, Tab. 20 (opp. p. 282), 283 clisea, Matronella Tab. 8 (opp. p. 264) Clupea . 58, mess 70-72, 80, 81 Clupeidae 61-77, 80 Clupeinae : : 61-63, 80 Clupeoidea . 5 é a . - 57-85 cobbi, Neodavisia fi F : 260 Codakia c Tab. 1 (opp. p. 256), 257, Ply coelophyllus, Rhinolophus . 183, 186 coelophyllus shameli, ea ais 183, 194 Coelops 187, 194 Coilia . C a : . F 5 83 Colobus , . 219, 220, 221, 225 Colomys . 247 columbella, Lucina Tab. I t (opp. p. 256), 257 commersonianus, Engraulis . c 82 commersonii, Stolephorus’ . 5 82 communis, Lithobius “110-211, 118 concamerata, Thecalia . Tab. 4 (opp. p. 260) condorensis, Pteropus hypomelanus ee) Congothrissa é : A 6 68 consobrina, Cyrena Tab. 14 (opp. p. 269) continentis, Myotis hasseltii . 188, 194 consors, Trachycardium Tab. 6 (opp. p. 263), Pl. 3 convexa, Thracia . Tab. 20 (opp. p. 282) Cooperella Tab. 15 (opp. p. 270) Cooperellidae Tab. 15 (opp. p. 270), 275 Coralliophaga 268, Tab. 11 (opp. p. 268) coralliophaga, Coralliophaga Tab. 11 (opp. p. 268) corallina, Mactra . Tab. 8 (opp. p. 264) Corbett, G. B. 211-252 Corbicula 2 Tab. 14 (opp. p. 269) Corbiculacea 256, 269, Tab. 14 (opp. p. 269) Tab. 22 (opp. p. 284), 289 Corbis a E Tab. 1 (opp. p. 256), 257 Corbula 5 275, 276, 278 Corbulidae . 275, 291 cordata, Corbicula cordiformis, Arctica Tab. 14 (opp. p. 269) Tab. 11 (opp. p. 268) coriaceus, Lithobius 124-125 Corica . : c 81 corrugata, Florimetis Tab. 10 (opp. p. 266) corsicus, Lithobius impressus 5 : 108 cortius, Chatoessus = 9 b 0 81 Costia, Ichthyobodo_ . ° ° 98 coucha, Mastomys : a : Z 235 coucha lateralis, Mastomys . a 3235) crassa, Corbula 9 275,278 Crassatella . Tab. 5 (opp. p. 261), Pls 1 and 2 Crassatellacea 255, 201-263, Tab. 5 (opp. p. 261), Tab. 22 (opp. p. 284) Crassatellidae 261-263 Crassinella . Tab. 5 (opp. p. 261) crassipes, Lithobius 114, 115, 131 crassum, Laevicardium Tab. 6 (opp. p. 263) crawshayi, Lepus 5 ez crenata, Venerupis Tab. 15 ‘(opp. p- 270) Cricetidae . a 5 0 227 crispata, Zirfaea . 281, Tab. 19 ‘(opp. Pp. 282) cristata, Hystrix . 219, 239, 246 cristatus, Proteles 240, 248 Crithidia 99, 100 Tab. 15 ‘(opp. Pp- 270) Tab. 7 (opp. p. 264) crocea, Circa crocea, Tridacna . Crocidura 220, 221, 222, 224-225, 246, 247, 248 Crocuta 3 ; -217, 221, 241 crocuta, Crocuta . 5 217, 241 Cryptodonta 5 4 5 3 OT) Ctenodonta : , 0 5 ~ 286 Cucullanus . 4 I51I-170 Cultellus 264, Tab. 9 (opp. p. 264) cuneiformis, Corbicula . Tab. 14 (opp. p. 269) curtipes, Lithobius 113-116, 131 curtipes turkestanicus, Lithobius . = eD5) Cuspidaria . Tab. 21 (ope: Pp. hah Pl. 14 cuspidaria, Cuspidaria . welia4 Cuspidaridae j é : : zon! cuspidata, Cuspidaria . polka Cyamiacea . 255, 260, Tab. 22 (ope. p. 284) Cyamium . 6 Aes Cyathomonas 5 6 98 Cyclina Tab. 15 cpp: i: 270) Cyclininae Tab. 15 (opp. p. 270), 274 298 INDEX cyclotis cyclotis, Murina 193, 194 Cynopterus . 175-180, 194 Cyphomyrus 3, 6, 7, 30-31, 59, 51 cyprinoides, Marcusenius 3, 6, 7, 10, 31, 32 cyprinoides, Velorita . 269, Tab. 14 Cyrena Tab. 14 (opp. p. 269) Cyrtodaria . é 279, 280 dactylus, Pholas . Tab. 19 (opp. p. 282), Pl. 11 Dasymys 5 221, 232 Day, Francis, pollections of . c » 59-61 Dayella 5 . 59-70 decipiens, Crassatella Tab. 5 ene e 26x); Plia defassa, Kobus 216, 222, 243 deflorata, Asaphis Tab. 10 (opp. p. 266), Pl. 5 deignani, Myotis . ; eh 3188 deliciosus, Mormyrops . 5 2 me Yanete) dembeensis, Pelomys . 5 . “| 247 Dendromus 5 - 219, 236, 247 dentatus, Lithobius 108, 128 dentatus var. alpestris, Lithobius . A 108 denticulata, Petricola Tab. 15 (opp. p. 270) deshayesiana, Verticordia Tab. 21 (opp. p. 282) dilatata, Crassatella Tab. 5 (opp. p. 261) dillwyni, Meretrix Tab. 15 (opp. p. 270) dimidiatus, Acomys : 220, 229, 248 dimidiatus mullah, Acomys . . e222 dione, Hysteroconcha Tab. 15 (opp. p. 270), Pl. 8 Tab. 10 (opp. p. 266) 2 227-228 3, 6, 7, 30-31, 51 Tab. 10 (opp. p. 266) 2 Q 81 diphos, Solenotellina (Dipodillus), Gerbillus . discorhynchus, Cyphomyrus . 4 dispar, Scissulina . ditchela, Pellona . c : ditchoa, Pellona . 5 81 divaricatum, Gafrarium Tab. 15 Neca! p- 270) Divaricella . Tab. 1 (opp. p. 256), 257 divergens, Ctena . Tab. 1 (opp. p. 256) docmac, Bagrus : 53 Donax Tab. 10 > (ope. p. 266), 267, Pl. 4 dorab, Chirocentrus é 9 c 80 Dorcatragus és 4 5 5 : 246 doriana, Crocidura 221, 224 Dorosomatinae 73-74, 81 Dosinia Tab. ce (opp. p. 270), 274 Dosininae ; Tab. 15 (0 OPP. P- 270), 274 Dreissena Tab. 12 (opp. p. 268), 269, Pl. 6 Dreissenacea 256, 268, Tab. 12, Tab. 22 (o 28 dubosequi, Lithobius_ . a8 ei : oe = Dussumieria 5 : < . 68, 80 dussumieri, Coilia b 6 > O 83 dussumieri, Engraulis . 5 ci Tite dussumieri, Thryssa 77-79, 82 Eason, E. H. 103-150 eburnea, Divaricella Tab. x (opp. Pp. 256) eburnea, Euciroa Tab. 21 (opp. p. 282), Pls 13 and 14 ecaudatus, Megaerops echinata, Acanthocardia Tab. 6 (opp. p. 263), Pl. 3 edax, Bodo . Z : 5 2 f 99 Edmondiacea z 5 291 edule, Cerastoderma Tab. 6 (opp: ce 263), Pl. 3 effosa, Lirophora . Tab. 15 (opp. p. 270) 181, 193 Egeria Tab. 10 (opp. p. 266) Ehirava ie 65, 67, 68, 69, 81 Elephantulus 223, 246 elephas, Campylomormyrus 6, 7, 33-35 ellipta, Astarte Tab. 5 (opp. p. 261) elongata, Ilisha . : 5 TROL elongatus, Lithobius” . : s a weeros elongata, Oriocrassatella 6 - = 263 elongata, Pellona ; 7 ; 81 elopsoides, Dussumieria F . 5 80 Elops . A 7 6 - . 8, 10, 48, 50 emini, Taterillus . 222, 228, 248 Engraulidae 77-79, 82 Engraulis 59, 77-79, 82 Ensis . c Tab. 9 (opp. p. 264), 265, Pl. 4 ensis, Ensis epidermia, Domax Tab. 9 (opp. p. 264) Tab. 10 (opp. p. 266) Equidae . 5 é nay z4aZ Equus 3 3 é 0 “216, 242, 246 Erinaceus . c 246 erlangeri, Gazella Posaimening? © 6 244 erlangeri, Procavia habessinica 0 . 241 erycina, Amiantis Tab. 15 (opp. p. 270) erythrocephalus, Lithobius . 109, 116, 129, 143 erythropus leucoumbrinus, Xerus . : 227 Escualosa_ . 0 a : 80 etruscus, Suncus . | Baa 246, 248 Euciroa Tab. 21 (opp. p. 282), 284, Pls 13 and 14 Eucrassatella Tab. 5 (opp. p. 261), 262 Eupolybothrus - 108, III, 112, 140 exogyra, Chama . : . “ 3 259 exoleta, Mactronella Tab 8 (opp. p. 264), 265, Pl. 4 faba, Donax Tab. 10 (opP: Pp. 266), 267, Pl. 4 fagani, Lepus whytei 226-227 fasciatus, Eupolybothrus 5 é III, 140 fasciculata, Crithidia . b é 5 99 Felidae 0 c < : F 241 Felis| . c : . 221, 222, 241, 246 ferganensis, Taghobias : a : 5 II5 festivus, Lithobius F : 119 fijiensis, Lucina Tab. 1 (opp. p- 256), 257, Pl. x filigera, Pellona . 5 82 Fimbria Tab. I (opp. p. 256) fimbriata, Clupea_ 2 = 80 fimbriata, Corbis . Tab. 1 (opp. p- 256), 257 fimbriata, Sardinella_ . : - 63, 80 fishes, clupeoid . 5 z - 57-85 Fishes, Osteoglossomorph : ; - 1-55 flavescens, Crocidura 224, 246 INDEX 299 flavopunctatus, Lophuromys 217, 221, 233, 247, 248 flesus, Platichthys s . 153 floridana, Cardita. Tab. 4 ‘(opp. p. 260) Florimetis Tab. 10 (opp. p. 266) florium toxopei, Murina 5 : i 193 flounder : b hs 153 fluminea, Gaapicnla fluviatilis, Ehirava forficatus, Lithobius 107, 120, I2I, 122, 124, 125, 140 forficatus var. villosus, Lithobius . , 120 formosus, Scleropages fossor, Lithobius . fragilis, Gastrana . Tab. 14 ‘(opp. p. 269) 64, 65, 68, 69, 81 . : 7 122-123, 124 Tab. 10 (opp. p. 266) Fragum 9 Tab. 6 (opp. p. 263) frithii, Coelops A - 187, 194 fulvorufula, Redunca . 6 5 5 247 fumatus, Praomys é 222, 246, 248 fumatus allisoni, Praomys_ . C 6 235 fumatus brockmani, Praomys 234-235 furthii, Ilisha f : 5 5 3 717 Gafrarium Tab. 15 (opp. p. 270), 273, Pl. 8 Gaimardia . : ‘5 | 208, P16 Gaimardiacea 256, 268, 284, Tab. 22 gairdneri, Scotophilus kuhlii . 191-192 Galago 225, 246 gambianus multicolor, Heliosciurus re eed Gastrana Tab. 10 (opp. p. 266) Gastrochaena 277, 280 Gastrochaenacea oe 277, 280, Tab. 22 (opp. p. 284) Gazella . 216, 222, 244, 245, 246 gazella, Oryx 6 : 5 gazella beisa, Oryx 222, 244 gelada, Theropithecus 226, 247 geminorum, Pteropus hypomelanus ya gemma, Gemma . Tab. 15 (opp. p. 270) Gemma Tab. 15 (opp. p. 270) Gemminae . Tab. 15 (opp. p. 270), 274 Genetta 221, 240, 247 genetta, Genetta . " 5 4 : 240 Gerbillinae . 5 e . “ i BEF Gerbillus 222, 227-228 gibba, Corbula 276, 278 gibbosa, Eucrassatella “Tab. 5 5 (opp. p. a 262 gibbosa, Sardinella 2 63 Gibson, D. I. ii 170 gigantea, Gastrochaena 277, 280 gigas, Arapaima . 3,5, 7, 14-16, 52 Gilchristella 4 65, 67, 68, 69 Giraffa 4 6 - 245 glaber, Heterocephalus | 0 e240) glabrata, Atactodea Tab. 8 (opp. p. 264) glabratus, Lithobius . 106, 113, 125, 127 Gliridae : : 3 B 5 Een230; Glossacea 256, 269, Tab. 13, Tab. 22 (opp. p. 284), 289 Glossus R 269, Tab. 13, Pl. 6 Gnathonemus 4 =D 7, 32-33, 40, 49, 50 godanahiai, Gudusia_. : : 70 Gonialosa_ . a : é 73-74, 81 goslingi, Colones . 247 Gouldia a Tab. 15 cers p- 270) gracilitarsis, Lithobius piceus 6 é 124 granulatus, Lithobius . , 5 5 131 granti, Gazella 216, 244, 245 granulata, Chione Tab. 15 (opp. p. 270) granulata, Poromya Tab. 21 opp. | p. 282), 284, Pl. 14 Graphiurus . 236, 240 Greenwood, P. EL 5 5 . 1-55 gregorianus, Thryonomys 238-239, 248 grimmia, Sylvicapra. . 243 griseifrons, Dasymys incomtus c 6 232 groenlandicus, Serripes . Tab. 6 (opp. p. 263) grossipes, Eupolybothrus . III, 112, 119 grossipes, Lithobius 105, 106, 111, 112 gryphina, Chama a 3 . 258 guanocostense, Trigonocardia Tab. 6 (opp. p. 263) Gudusia 4 70-72, 81 guentheri, Madoqua : 5 : . 243 guereza, Colobus . 219, 221, 225 Gymnarchidae - 3,4, 44-48, 49, 54 Gymnarchus 3, 6, 7, 8, 10, 23, 44-48, 49, 59, 51, 52, 53 habessinica, Procavia . 220, 241-242, 248 habessinica alpini, Procavia . 241, 242 habessinica erlangeri, Procavia eat habessinica scioana, Procavia : é 242 habessinicus, Lepus”. 5 5 226, 227 hamadryas, Papio 5 F E 222, 226 hamiltonii, Engraulis . 5 Q d 82 hamiltoni, Thryssa F - 82 hararensis, Heterohyrax brucei 4 ~ 242 hararensis, Madoqua phillipsi é . 243 harringtoni, Taterillus . Fi . 0 228 harrisoni, Thryonomys . ; c 238-239 hardwoodi, Gerbillus. 5 zzz harwoodi, Gerbillus (Dipodillus) 227-228 hasselquisti, Mormyrus a 3, 6, 7, 28-29 hasseltii, Dussumieria . 4 A : 80 hasselti, Myotis . & s : 188, 194 hasseltii contintis, Myotis ‘ 2 188, 194 haueri, Chama . é 9 b . 258 heathi, Scotophilus G 192 heathi (?) watkinsi, Scotophilus 192-193 hecki, Tachyoryctes : 237-238 Hecuba Tab. 10 (opp. p. a: 267, Pl. 4 hecuba, Donax . 0 2) 206) Heliosciurus 6 b a 227 Hemicardia Tab. 6 (opp. p- 263) hemicardia, Hemicardia Tab. 6 (opp. p. 263) henryi, Isichthys . -3, 6, 7, 37-38 Herklotsichthys . C fi . 61, 80 300 INDEX Herpestes . . B : d 240) Hesperoptenus : IQI, 193, 194 Heterocephalus . e240) heterochrous, Cucullanus 153) ne 159, 161-167 Heterodonta Fs 256-282, 289 Heterohyrax 242, 248 Heterotinae J g . 14-16 Heterotis = 35 5, oh a 16, 18, 50, 52 hians, Tivela Tab. 15 (opp. p. 270), 273, 274, Pl. 7 Hiatella 279, 280 Hiatellacea 256, 277, 280, Tab. 22 (opp. p. 284), 290 Ell) J. EBs; 9 171-196 Hilsa . : 5 4 3 53, 72, 81 hindei, Geocitinra! . : za) hindei marrensis, Crocidura . = oe Hiodon 3, 5, 7,9, 10, 48, 50, 51 Hiodontidae 3, 9-11 Hippopotomidae . 6 242-243 Hippopotamus. 7 é 2 a 209) Hippopus Tab. 7 (opp. p. 264), Pl. 3 hippopus, Hippopus Tab. 7 (opp. p. gue Pl. 3 hoevenii, Pellona . c < 81 hoffeti, Cynopterus brachyotis 2 c 177 horsfieldii, Myotis : z & 188 hortensis, Lithobius 121 122 humanus, Glossus 269, Tab. 13 (opp. p. 269), Pl. 6 Humphreysia 5 3 a me 284 Hyaenidae . “ 2 240-241 hydropica, Corbula o 5 278 Hyperopisus = 5h a Ts fee 49, 50, 51 hypomelanus, Pteropus 174-175, 193 hypomelanus annectens, Pteropus . 174 hypomelanus canus, Pteropus ¢ 5 174 hypomelanus condorensis, Pteropus a 174 hypomelanus geminorum, Pteropus ci 174 hypomelanus lepidus, Pteropus’. 174 Hyracoidea 2 241— 242 Hysteroconcha Tab. 15 (ope. p. 270), Pl. 8 Hystrix 5 : 219, 239, 246 ibex nubiana, Capra. 5 . - 246 ibex walie, Capra i : “ 246 Ichneumia . o -219, 221, 240 Ichthyobodo ‘ 5 ; : , 98 Ictalurus . 5 E . a 7 53 Ictonyx P “ ¢ , 222, 239 Ilisha . : c 9 - 74-77, 81, 82 ilisha, Clupea 5 = - : 5 81 ilisha, Hilsa : ‘ 6 - - 53, 81 imberbis, Muriculus’ . 5 5 on a 2A, imberbis, Tragelaphus . 5 9 . 244 imbricata, Venericardia Tab. 4 (opp. p. 260), 261 imhausi, Lophiomys . : 3 a BAG immutabilis, Lithobius . 128-129 impressus, Lithobius 106, 108 impressus corsicus, Lithobius - : 108 impressus impressus, Eupolybothrus (Allopolybothrus) . > - 108 inaequivalvis, Periploma Tab. 20 (opp. p. 282) incarnatum, Vasticardium Tab. 6 (opp. p. 263) incomptus, Dasymys 221, 232 incomtus griseifrons, Dasymys : 5 ze incrassata, Astarte Tab. 5 (opp. p. 261), 262, Pl. 1 incrassata, Cardita Tab. 4 (opp. p. 260) indica, Ilisha . 2 3 - 77, 81 indica, Pellona . : 5 C : 81 indicus, Engraulis % i 6 82 indicus, Stolephorus” . - - 3 82 inerbis, Tregelaphus’ . : : = S22) inermis, Lithobius : : “ s 118 infinitesimus, Suncus . 223-224 inflata, Cyrena Tab. 14 (opp. p. 269) Ingle, R. W. 197-210 Insectivora . 223-225 insularum, Cynopterus brachyotis a A W77. intermedia, Circa Tab. 15 (opp. p. 270) internus, Claviglis brockmani : = 236; internus, Graphiurus murinus 6 5 236 jostoma, Chama . 5 e 258 Isichthys 6, Wh 37-38) 49, 50, 51 islandica, Arctica Tab. 11 Gee p. 268) Isomys z 2 5 : 230 jacksoni, Otomys : A A Zaz japonica, Dosinia A - c ee jebelae, Arvicanthis P : 230 jedoensis, Perothaca Tab. 15 (opp. p. 270) Jenkinsia 68 kammalensis, Engraulis c 4 82 kammalensis, Thryssa . : . é 82 kanagurta, Clupea : : 81 kannume, Mormyrus 3, & a 2357, 28 kapirat, Notopterus , : Tee kelee, Hilsa 2 5 , . a 92s Bit Kelia . d , - 6 259 Kennerlya . p : Tab. 20 (opp. p- 282) kijabius, Mus ; : 0 235 klunzei, Clupea_. 5 2 fi 2 80 Kobus : 216, 222, 243 Koch collection (Chilopeda: Lithebiowospee) 103-150 kuhliui, Scotophilus . A 6 192, 194 kuhlii kuhli, Scotophilus 5 S be ee kuhlii gairdneri, Scotophilus . < 191-192 lacernatus, Arvicanthis 230, 232 lacerda, Mormyrus 3, 6, 7, 28 lactuosus, Cultellus lacustris, Sphaerium Laevicardium Tab. 6 (opp. Pp. 263), Pl. : Laeviscutella i 68 Tab. 9 (Opp. p- ae Pl. INDEX 301 Lagomorpha 226-227 lamarckii, Meiocardia Tab. 13 (opp. p. 269) Lamelliconcha Tab. 15 (opp. p. 270) lamellosa, Crassatella . Tab. 5 (opp. p. 261) lamellosa, Fimbria : Tab. 1 (opp. p. 256) laminiferum, Cyamium a : 5 260 lanzae, Lithobius Fs 4 2 . IIo lapidicola, Lithobius . a 2 110 lateralis, Mastomys coucha . E235 Laternula Tab. 20 ‘(opp. p- 282) Leggada . : : oe eye leichardti, Scleropages . = 5,7, 13-14 leiogaster, Clupea 5 a 80 leiogaster, Sardinella . : 0 ° 80 Leishmania é a 0 99 Lemniscomys 220, 230 233, 245, 248 leo, Panthera 3 af é 4 2109, 241 Lepidomysidae P 197-210 Lepidomysis A 5 3 199, 201, 207 Lepidophthalmus 5 . 199, 201 Lepidopidae a a 5 5 : 207 Lepidops : 199, 201 lepidus, Pteropus hypomelanns ° 5 174 Leporidae 226-227 Leptonacea . 255, 250, Tab. 22 “(opp. p- 284) leptopus, ata : TL, ee Lepus . : 226-227, 246 leschenaulti, Pellona 2 “ “ 4 81 leschenaulti, Roussettus i . F 173 leschenaulti leschenaulti, Rousettus 6 173 leucoumbrinus, Xerus erythropus . a 2H! leucozonica, Mactra liana, Lucinisca Tab. 8 (opp. p. 264) Tab. 1 (opp. p. 256) libyca, Felis silvestris . a E a ere lile, Clupea . 6 6 © : B 80 Limacea Tab. 22 (opp. p. 284) Limopsacea . Tab. 22 (opp. p. 284) linguafelis, Scutarcopagia Tab. 10 (opp. p. 266) Lioconcha Tab. 15 (opp. p. 270), 273, 274 Lirophora : Tab. 15 (opp. p. 270) Lithobiomorpha . . 103-150 Lithobius 103-150 lithophaga, Petricola Tab. 15 (opp. p. 270) Litocranius . “ 222,244 litoralis, Eupolybothrus ean ee?) litoralis, Lithobius 105, 140 litterata, Tapes Tab. 15 (opp. p. 270), 274 longibarbis, Gnathonemus -3, 6, 7, 32-33 longiceps, Clupea x 5 80 longiceps, Sardinella. ; 2 z 80 longimanus albipinnis, Taphozous . PLS longimanus longimanus, Taphozous 182, 193 longipes, Spelaeomysis . 208-209 Lophiomys . 2 5 6 ¢ 247 Lophuromys 217, 221, 233, 247, 248 Loripes Tab. 1 (opp. p. 256) Lorisidae . 225 loscombiana, Pholadidea Tab. 19 ‘(opp. p- 282) lovati, Dendromus 5 e247) Loxocardium Tab. Gl (opp. p. 263) lubricus, Lithobius lucifugus, Lithobius Lucina Lucinacea 253) SN Tab. 1, 284, Tab. 22, 289 lucinalis, Loripes . Tab. 1 (opp. p. 256) 136-137 130, 131, 136, 143 Tab. 1 (opp. p. 256), 257, Pl. 1 Lucinisca 6 : Tab. 1 (opp. p. 256) Lucinopsis . o d Tab. 1 (opp. p. 256) lunatula, Crassinella . Tab. 5 (opp. p. 261) lutrella, Crocidura 3 a : . 224 Lycaon 3 7 : , c 6 239 macilentus, Lithobius 129-130 Macoma 266, Tab. 10, 267 Macrocallista Tab. 15 (opp. p. 270) macrocephalus hecki, Tachyoryctes 237-238 macrocephalus macrocephalus, Tachyoryctes 237-238, 247, 248 Macroglossus 6 f 182, 194 macrolepis, Mastomys . : : 235 macrops, Lithobius a 137, 138 Macroscelididae . 5 c 223 macrurus, Rhinolophus affinis ° ek 82 Mactra Tab 7 (opp. p. 264) Pl. 4 Mactracea 256, 264, Tab. 7, 265, 284, Tab. 22, 289 Mactroderma bi é Tab. 8 (opp. p. 264) Mactronella Tab. 8 (opp. p. 264), 265, Pl. 4 maculosum, Trachycardium Tab. 6 (opp. p. 263) madagascariensis, Sauvagella : . 67, 69 Madoqua 243, 246 magna, Serobicularia Tab. 10 loro! p. 266) mahomet, Mus 217, 233-234 malabaricus, Dayella neat 69, 81 malabaricus, Engraulis 5 5 82 malabaricus, Spratelloides eae 85, pl. 16 malabaricus, Thryssa . & 82 malayanus, Rhinolophus ‘183, 187, 194 Mammals (other than bats) 211-252 manmina, Chatoessus . 0 é 5 81 manminna, Gonialosa 73, 74, 81 Marcusenius 3, 9, 7, 31-32, 49, 50 marmorea, Cardita Tab. 4 (opp. p. 260), 261, Pl. marrensis, Crocidura hindei . : 224 Martesia Tab. 19 “(opp. p. 282) massaicus, Lemniscomys striatus . 6 233 (Mastomys), Praomys . 0 a 7235 mearnsi, Arvicanthis . 3 ‘5 230 Megadesmus . 6 " 291 Megaerops 5 é ase 193 Megalops. . 5 ¢ , . 10, 48 megaloptera, Ilisha 2 : c Bahra megaloptera, Pellona . 2 : : 82 megalotis, Dorcatragus 7 e ~ 1246 megalotis, Otocyon a : 0 e | 26h) Megapipistrellus . 7 : G titsle) 302 INDEX melanocephalus, Chironax 181, 193 melanocephalus, Lithobius - 106, 126-127 melanops, Lithobius I13, 125, 126, 127, 128 melanura, Clupea F : é 80 melanura, Sardinella . : : 80 Meletta ; 3 6 2 4 61 Mercenaria Tab. 15 (opp. p. 270), 274, Pls 7 and 8 mercenaria, Mercenaria Tab. 15 (opp. p. 270), 274, Pls 7 and 8 Meretricinae Tab. 15 (opp. p. 270), 274 Meretrix 3 Tab. 15 (opp. p. 270) martini, Cardilia . Tab. 8 (opp. p. 264) Matronella . : . Tab. 8 (opp. p. 264) maxima, Tridacna Tab. 7 (opp. p. 264) Meiocardia . Tab. 13 (opp. p. 269) Microdillus . 5 5 c 246 microps, Lithobius : ; 4 F 133 minimus, Lithobius : 4 : 128 minimus sobrinus, Macroglossus 182, 194 minor, Rhinolophus~—. : 2 183 minutus, Cucullanus minutus, Lithobius modesta, Gonialosa modestus, Chatoessus e535 Msso16c) 165 rir, 118 - 73-74, 81 50, Stes 81, pie 2a moluccensis, Clupea—. < 80 Monoplacophora . : 2 a Zon Monotarsobius “aRG. 132, 133 montanus, Lithobius 112, 119 mordax, Lithobius 119-120 Mormyridae 3, 22-44, 52-53 Mormyrops . X 3, 6, 7, 38-40, 50, 51 Mormyrus - 3,6, 7, 23-29, 44, 49, 50 motius, Pellona . F : 81 Mulina Tab, 8 (opp. p. 264) mullah, Acomys diniidiatus 222, 229 multicolor, Heliosciurus gambianus 9 227 multispinosa, Hysteroconcha Tab. 15 (opp. p. 270) Muriculus . : f me Rey. Muridae 228-236 Murina 193, 194 murinus, Graphianas 2 236, 246 murinus ? internus, Graphiurus = . 236 murinus saturatus, Graphiurus 4 236 Mus . : 217, 219, 233-234, 235, aay, 248 muscorum, Lithobius . . 121, 122 Mustelidae . ¢ , y 239 mutabilis, legneaiis 107, I10, III, 116, 117, 118, 134-135 muticus, Lithobius 107, 109, 110, 116-117, 135, 136 Mya . 5 « 278, ,2k 20 Myacea 256, 275, 278, 284, Tab. 22, 291 Myadora e Tab. 20 (PP. p. 282), 283 Myidae F 5 : a) e200 Myochama . Tab. 20 Wenn p. 282) Myoida 275) a 33 (Opp. p. 286), 290 Myomys . : 5 2 BY Myotis : 188-190, 193, 194 Myoxus 0 0 : ' 236 Mysidacea: Lepidomysidae 197-210 mystacalis, Dendromus 219, 236 mystax, Engraulis é fe £ 5 82 mystax, Thryssa . 2 . 82 Mytilacea Tab. 22 2 (Opp: p. 284), 288 Mytiloida Fig. 33 (opp. p. 286) nasus, Nematalosa F : - 83 natalensis, Praomys 220, 221 natalensis, Praomys (Mastomys) a . 235 Nautilus a : 5 291 navalis, Teredo Tab. 19 (opp. p. ea! Pl. 10 nebulosus, Ictalurus. é 53 necator, Ichthyobodo (Costia) 4 6 98 Nematalosa 0 3 5 3 zt 83 Nematoda: Ascaridida . A : 151-170 Neodavisia . o 3 < : =) 4260) Neolithobius 5 . : c 5 120 nigri, Xenomystus 5,7, 18, 21, 22 nigripalpis, Lithobius 140-142 Nilopegamys : 5 E « S2A7 niloticus, Arvicanthis aby, 230, 231, 232, 246 niloticus abyssinicus, Arvicanthis 229-230 niloticus, Gymnarchus . 3, 6, 7, 8, 23, 44-48 niloticus, Heterotis 3, 5, 7, 14, 50 niobe, Crocidura . 224-225, 248 norvegica, Panomya, Panopea (Panamya) 279, 280 Notopteridae 3, 18-22, 51-52 Notopterus . 7, 18, 22 nubiana, Capra ibex . 3 5 iy 8240 Nucinella . : a : : 287 Nuculacea Tab. 22 (opp. p. 284), 287 Nuculanacea Tab. 22 (PP: p. 284), 287 nudicornis, Lithobius . 108 Nuculoida Fig. 33 (opp. p. 286) obliqua, Astarte . Tab. 5 (opp. p. 261) obliquum, Loxocardium Tab. 6 (opp. p. 263) oblongum, Trapezium Tab. 11 (opp. p. 268) occidens, Corbicula Tab. 14 (opp. p. 269) occidens, Psammotea Tab. 10 (opp. p. 266) occidentalis, Crocidura . 5 5 =. Rez Offadesma . Tab. 20 (opp. p. 282) omalii, Astarte , Tab. 5 (opp. p. 261) oncopelti, Crithidia E < 5 é 100 Opisthopterus. : : : : 82 Oreotragus . : . 219, 222, 243, 248 oreotragus, Oreotragus . . 219, 222, 243, 248 Oriocrassatella . > 263 orobinus, Claviglis é 5 0 2) 236, orobinus, Myoxus : ° 236 Orycteropus : 1217, 2 221, 242 Oryx . . 5 G 5 222, 244 Osteoglossidae 3, 12-16 Osteoglossinae . c : : - 12-14 Osteoglossomorpha : . 1-55 Osteoglossum $b 7h 12-13, 16, 17, 48,50 Ostreacea Tab. 22 (CEP: p. 284) Otocyon f : é 239 Otomys 0 217, 247 ourebi, Ourebia 220, 243 Ourebia : 220, 243 ovata, Gastrochaena . : 277, 280 oweni, Scientilli . 3 5 3 3 259 Palaeoheterodonta 7 6 3 3 289 Palaeotaxodonta . 5 » 287 Tab. 10 (opp. p. 266) Tab. 8 (opp. p. 264) palatam, Quidnipagas . pallida, Mulina Pandora Tab. 20 (opp. p. 282), 283 Pandoracea 256, 282, Tab. 19, Tab. 22 aA: p. i 291 Pangasius . 154 pangasius, Pangasius o : c =) 154 pannonicus, Lithobius agilis. 4 113 Panomya 277; 270, 280 Panopea 277, 279, 280, Pl. 9 Panopea (Panomya) 280, 284, 290 Panthera 219, 241 Pantodon 3, 5,7, 16-18, 48, 50 Pantodontidae 0 3, 16-18 Paphia Tab. 15 (opp. p. 270), 274 paphia, Chione Tab. 15 (opp. p. 270) Papio . a 219, 220, 221, 222, 225-226 Papyridea Tab. 6 (opp. p. 263) Papyrocranus 3, 5» 7, 18-20, 21, 22 Paraechinus é c 246 Parapholas . Tab. 19 (opp. p. 282) Parascotomanes, Scotomanes : c IQI Parasilurus . 5 0 é a : 53 pardus, Panthera 2 c c 5 ue) parisiensis, Lithobius - ¢ 5 apa parkinsoni, Acanthocardia Tab. 6 (opp. p. 263) Parvicardium Tab. 6 AREB: p- 263) pasha, Leggada . ‘i Hy 3234 pasha, Mus . aS: 234, 248 paytensis, Lamelliconcha Tab. 15 (opp. p. 270) Pecchiola 5 . Tab. 21 (opp. p. 282) Pectinacea . . Tab. 22 (opp. p. 284) Pectinator . a : -me240 pectinatum, Cite Tab. 15 (opp. p. 270), Pl. 8 peeli, Microdillus . é 4 F 246 pelidnus, Lithobius 107, TIO, II7, 135, 136 pelliceus, Arvicanthis . 5 a PR) Pellona 6 50, 74-77, 81, 82 Pellonulinae 63-70, 81 pellucida, Chama . 258, 259 Pelomys Q ° 247 Penicillus 284, 285, Pl. 15 peregrinus, Thebes ¢ E ez Periglypta . Tab. 15 (opp. p. 270), 273 Periploma ceebalbs120) (eep- p. 282) Perissodactyla . 2 d 242 Peristedion . “ c 5 48 Perothaca Tab. 15 (gam p- 270) INDEX 393 peruviana, Lirophora Tab. 15 (opp. p. 270) Petricola Tab. 15 (opp. p. 270) Petricolidae Tab. 15 (opp. p. 270), 275 Petrocephalus 3, 6, 7, 35-37, 49, 50 Petromyzon A : “ 154 Phacochoerus le Ae 221, 222, 243 phasa, Setipinna . 3 3 5 5 82 phaseolina, Thracia Tab. 20 (opp. p. 282), Pl. 13 phillipsi, Madoqua 5 - . 246 phillipsi, Tatera robusta 0 d 0 228 phillipsi hararensis, Madoqua : . 243 Pholadacea . 256, 279, Tab. 19 (opp. p. 282), Tab. 22 (opp. p. 284) Pholadidae . 5 ' 279 Pholadidea . Tab. 19 cpp: p. 282) pholadiformis, Petricola | Tab. 15 (opp. p. 270) Pholadomya 282m blare Pholadomyacea 256, 282, Tab. 22 (opp. p. 284) Pholadomyoida Fig. 33 (opp. p. 286), 290 Pholas . Tab. 19 (opp. p. 282), Pl. 11 Pholadomyina. 5 7 282 piceus, Lithobius . wen225 123-1 24, 140 piceus gracilitarsis, Lithobius 5 124 piceus piceus, Lithobius 7 é Sey piceus verhoeffi, Lithobius . 2 = 124 pictus, Lycaon . 239 pila, Lucina 256, Tab. I (opp. p. 256) Pinnacea Tab. 22 (opp. p. 284) Pipistrellus . - 188, 191, 193, 194 Pisidium Tab. 14 (opp. p. 269) Pitar . : < . Tab. 15 (opp. p.270) Pitarinae Tab. 15 (opp. p. 270), 274 plana, Arctica . Tab. 11 (opp. p. 268) planeri, Petromyzon . 0 154 planicostata, Venericor Tab. 4 “(opp. p. 260) Platichthys ¢ é é 153 Platydon 276, 277, 278 plicata, Circomphalus Tab. 15 (opp. p. 270) plicata plicata, Tadarida ° 193, 194 Polymesoda Tab. 14 (opp. p. 269), Pl. 6 polymorpha, Dreissena Tab. 12 (opp. p. 268), 269, Pl. 6 ponderosa, Calyptogena Tab. 11 (opp. p. 268), 274 ponderosa, Dosinia Tab. 15 (opp. p. 270) panderosa, Tivela Tab. 15 (opp. p. 270) porcus, Potamochoerus 6 . 243 Poromya Tab. 21 (opp. p: 282), 284, Pl. 14 Poromyacea 256, Tab. 21, Tab. 22 (opp. p. 282), 284, Tab. 22 (opp. p. 284), 291 Poromyidae 6 0 5 é EZ S| Poromyoidea : 6 5 a : 284 Poropolys . é F A 0 : 142 Potamochoerus . 9 243 Praomys 220, 221, 222, mae 235, 246, Ap 248 Primates é 225-226 primula, Myotis 189, 190, 194 Pristigasterinae 74-77, 81 304 INDEX pristigastroides, Ilisha . = 5 1777, Procavia 220, 222, ass 247, 248 proconodon, Mus . So eey producta, Mactra Tab: 8 (opr. ?. 264), Pl. 4 Proteles 240, 248 Protozoa 5 é 87-102 Prowazekia ‘ : 97 Psammotea : Tab. 10 (opp. p. 266) Pseudochama : 3 258 Pteriacea Tab. 22 ‘(opp. p- 284) Pteriomorphia 288 Pteroida Fig 33 fone p- 286) Pteropus 174, 193 pubescens, Lithobius 139-140 puerpera, Periglypta . 2 273 punctata, Codakia Tab. 1 = (ee Pp- 256), 257 punctatus, Herklotsichthys . 2 . 61,80 purava, Engraulis 3 z é - 82 purava, Thryssa . 2 c : f 82 pustula, Kelia. S A 2 e250) punctulatus, Lithobius 111-112 pusillus, Lithobius 5 pusillifrater, Lithobius pusillus pusillus pusillifrater, Lithobius 109, I10, 115 109g, 110, 118 10g, 110, 118 Pyramus . 7 2 5 ede pyriformis, Tetrahymena a “ 0 99 quadragesimalis, Coilia = E : 83 quadrisulcata, Divaricella Tab. 1 (opp. p. 256), 257 Tab. 5 (opp. p. 261) Tab. 10 (opp. p. 266) quexii, Astarte Quidnipagas quinterensis, Spelaeomysis 208-209 Raconda . 4 6 2 “ 82 radians, Pseudochama : F 0 5 258 radiata, Crassatella radiata, Egeria radiata, Psammotea Tab. 5 (opp. p. 261), Pl. 2 Tab. 10 (opp. p. 266) Tab. 10 (opp. p. 266) radiata, Solenotellina . : é eels radiata, Tellina Tab. 10 (opp. p. 266), Pl. 5 Raeta 5 5 Tab. 8 (opp. p. 264), 265 raffertyi, Arvicanthis . : : 5 230 ramearati, Coilia . . 6 i . 83 Rattus a ; : 221, 235 rattus, Rattus. P 6 221 rattus alexandrinus, Rattus ‘ 5 - 235 Redunca. 6 : 3 a 247, reptans, Arvicanthis 3 231, 232 reticulata, Periglypta Tab. 15 (Opp. p. 270) revoili, Elephantulus. . 5 3 246 reynaldi, Coilia . 5 e : < 83 Rhinolophus Ee 182-187, 194 ringicula, Papyridea Tab. 6 (opp. p. 263) robinsoni, Rhinolophus o : xe HOF, robusta, Tatera 222, 228, 248 Rodentia 5 ¢ 9 227-239 rosea, Scientilla . 5 f p S250) rostrata, Cuspidaria Tab. 21 (opp. p. 282) Rousettus : 173, 194 rubescens, Arvicanthis . Z ' 5 230 tubiginosa, Genetta . o 24a tufescens, Elephantulus 223, 246 russellii, Engraulis : - 0 a 82 tusseliana, Raconda. 5 c - 82 tutilus, Xerus . 5 = 6 Pal Salmo : a 5 : . 2 48 saltans, Bodo 87-102, Pls 1-6 samharensis, Mus é 5 “ 285 sanguineus, Herpestes . : 5 . 240 Sardinella . ; t 3 5 . 61, 80 saturatus, Arvicanthis . 0 oP mrz30) saturatus, Graphiurus murinus - = 1236 saurus, Elops . 2 : 6 >, 8.0) Sauvagella . 6 ‘ . 67, 69 scherzeri, Cynopterus beaebneees c : 177 schraderi, Genetta tigrina . F ~ 240 Scientilli . : 0 5 259 scioana, Procavia Nabe, : 242 Scissulina Tab. 10 opp. p- 266) Sciuridae . : A 227 sclateri, Evinaeeds c ~ 246 Scleropages 3) 5, 7, 13-14, 50 scobinata, Scutarcopagia Tab. 10 (opp. p. 266) scortum, Donax hecuba 266, 267 scortum, Hecuba Tab. 10 ”ePe- p- ssa 267, Pl. 4 Scotomanes 2 IQI Scotophilus : - 191-192, 194 scripta, Circa Tab. 15 (opp. p. 270) scriptus, Tragelaphus 5 219, 221, 244 Scrobicularia Tab. 10 (opp. p. 266) Scutarcopagia c Tab. 10 (opp. p. 266) Semele 206, Tab. Bae} (opp. p-. 266), Pl. 5 senegalensis, Galago senticosum, Trachycardium Tab. 6 (opp. p. 263) 225, 246 sericea, Crocidura 222, 224, 248 sericea sericea, Crocidura - 22M: serratum, Laevicardium Tab. 6 (opp. p. 263) Serripes Tab. 6 (opp. p. 263) Setipinna : 5 0 Fi 82 setirostris, Enpraulis 2 7 a : 82 setirostris, Thryssa : 4 4 = 82 serval, Felis 2 F 221, 241 servatus, Lepidomysis . 5 R ope eae servatus, Lepidophthalmus . z > 9208 servatus, Lepidops c 5 dj =) 200 servatus, Spelaeomysis . - 197-200, 201-210 shameli, Rhinolophus 183-185, 186 shameli, Rhinolophus ea aatlata - To4 shoana, Tatera robusta = 225) Sierrathrissa 5 = z 68 siligorensis thaianus, Myotis 190, 194 Siliqua Tab. 9 opp: p. 264) siliqua, Cyrtodaria . 7, 6280 INDEX 305 Tab. 9 CPE Pp. 264), 265, Pl. 4 siliqua, Ensis +221, 241, 246 silvestris, Felis silvestris libyca, Felis. : : 5 241 simensis, Canis . 2 5 7 247 sindensis, Clupea . 61- as. 80, Pl. 1a sindensis, Sardinella”. 5 3 . 61, 80 sipho, Solenotellina a : J 5 AG sladeni, Ilisha 74-77, 82 sladeni, Pellona 50: Unie 82, Pl. 2b soborna, Corica . : 81 sobrinus, Macroglossus 1 minimus 182, 194 soemmerringi, Gazella . 222,244, 245, 246 soemmerringi erlangeri, Gazella_. > 244 solanderi, Sunetta Tab. 15 (opp. p. 270), 274 Solecurtus . Tab. 10 (opp. p. 266) Solemya . a : ear Solemyacea Tab. 22 (opp: p- 284) Solemyoida . ‘ Fig. 33 (opp. p. 286) Solen . 4 n Tab. 9 (opp. p. 264) Solenacea Bas, Tab. 9 (opp. p. 264), 265, Tab. 22 (opp. p. 284) Solenotellina Tab. 10 (opp. p. 266), 267, Pl. 5 solida, Spisula Tab. 8 (opp. p. 264), 265 somalica, Heterohyrax . ez 42 somalicus, Arvicanthis . 230-231, 232, 246, 248 sordidus, Lithobius 122, 123 Soricidae F 223-225 soror, Tatera valida! a . 228 sowerbyi, Cardita Tab. 4 (opp. p. 260), 261, Pl. 2 spadix, Rhinolophus borneensis’ . TS spekei, Pectinator 9 a 6 bp LO Spelaeomysis 197-210 Sphaerias - 0 : 181, 194 Sphaerium . c 6 6 PING sphinx, Cynopterus 177, 194 sphinx angulatus, Cynopterus 175-179 spinipes, Lithobius . 5 6 119 spinosior, Bothropolys . : 142 Spisula Tab. 8 (opp. Pp. 264), 265 splendens, Tachyoryctes 236, 247 spondyliodes, Chama . 6 250) Spratelloides 4 50, 64, 68, 81, 85 Spratellomorpha . > 5 - 67, 68 squalida, Macrocallista. Tab. 15 ‘\(cpp-p. . 270) sqamosa, Tridacna Tab. 7 (opp. Dp. 204, IE 82 Stolephorus strangei, Humphreysia . 5 a Ay! starcki, Lepus capensis : 2 - 226 stelmioides, Cucullanus 5 5 0 154 Stenocephalemys . 0 4 247 222, 244 Tab. 5 (opp. p. 261) Tab. 19 (opp. p. 282) Tab. 20 (opp. p. 282), 283 strepsiceros, Tregelaphus striata, Astarte striata, Martesia . striata, Myadora . striatula, Chamalea 5 2/72) striatula, Venus Tab. 15 (opp. P. 270), Pl. 7 striatus, Ictonyx . : 222, 239 striatus, Lemniscomys . . 220, 232-233, 245, 248 striatus massaicus, Lemniscomys . Fe 233 striatus wroughtoni, Lemniscomys é 233 Strigilla 2 . Tab. 10 (opp. p. 266) strigillatus, Solecurtus . Tab. 10 (opp. p. 266) stutchburyi, Chione . Tab. 15 (opp. p. 270) subdiaphana, Cooperella Tab. 15 (opp. p. 270) subimbricata, Venus . Tab. 15 (opp. p. 270) suborbicularis, Kelia . a ; 5 259 subrugosa, Chione Tab. 15 (opp. p. 270), 272 sueziense, Parvicardium Tab. 6 GEE p- 263) suhia, Clupea 5 5 7o Suidae : A A 2 : a 243 sulcata, Astarte Tab. 5 (opp. p. 261), Pl. 1 sulcata, Cardita Tab. 4 (opp. p. 260) sulcatus, Lithobius 132-133, 143 sumatranus, Rhinolophis, acuminatus . 185 Suncus = 223, 246, 248 Sunetta Tab. 15 (opp. p. 270), 274 Sunnetinae . Tab. 15 (opp. p. 270), 274 swaynei, Alcelaphus buselaphus 216, 244 swinderianus, Thryonomys . 4 HPL) Sylvicapra . 0 : : : R243) Tachyoryctes » 236-238, 247, 248 Tadarida Q , : 193, 194 Talwar, P. K. é x . 57-85 Tapes B 5 Tab. 15 (opp. p. 270), 274 Tapetinae Tab. 15 (opp. p. 270), 274 Taphozous . A 182, 193, 194 tardoore, Opisthopterus 3 F : 82 tarentolae, Leishmania . 3 : 99 tartoor, Opisthopterus . 82 tasmanica, Myadora Tab. 20 iopps p. 282) Tatera 222, 228, 248 Taterillus : 222, 228, 248 taty, Engraulis . 0 : : 82 taty, Setipinna . : 5 o 5 82 taylori, Tatera robusta 5 5 5 228 telara, Engraulis . 5 82 Tellidora ° Tab. 10 (opp: p- 266) Tellina Tab. 10 (opp. p. 266), Pl. 5 Tellinacea 256, 266, Tab. 10 (opp. p. 266), Tab. 22 (opp. p. 284), 289 temminckii, Scotophilus 192, 194 tenellus, Mus : 210, 234, 248 Teredinidae a 0 a é 9 279 Teredo Tab. 19 (opp. p. 282), Pl. 10 testicularis, Isomys 5 a 0 - 230 Tetrahymena : 2 3 2 : 99 textilis, Paphia Tab. 15 (opp. p. 270), 274 thaianus, Myotis siligerensis . 0 190, 194 theobaldi theobaldi, Says iy 182, 194 Theropithecus 226, 247 Thonglongya, K. . a 171-196 thoracata, Escualosa . 2 80 Thracia Tab. 20 (pp. 1 P. 282), 293, Pl. 13 Thracidae . : c 291 Thrissina . e : 6 0 : 82 Thryonomyidae 2 238-239 Thryonomys . 220, 238-239, 248 INDEX 306 Thryssa 3 77-79, 82 Thyasira Tab. I (opp. p. 256) tickelli, Hespereptenits 5 191, 193 tigerina, Codakia &e Tab. 1 (opp. p. a 257, Pl. 1 tigrina schraderi, Genetta 9 240 tigrina, Genetta : 240 titthaecheilus, Cynopterus eercas D 175 Tivela Tab. 15 (opp. p- ais 27352745 Elz, toli, Clupea 81 toli, Hilsa 3 7 : 81 tortuosa, * snele . Tab. 10 (opp. p. 266), Pl. 5 toxopei, ina florium 193 Trachycardrim Tab. 6 (opp. P. B88) Pes Tragelaphus 219, 221, 222, 244, 247 transmarinu®, Lithobius 119, 120 transversus, Donax trapezia, Ga mardia Tab. 10 (opp. p. 266) : . ~ PING trapezina, Gaimardia : 268 Trapezium . Tab. Il (opp. p. 268) tri, Engraulis : 82 tri, Stolephorus 82 tricuspis, Lithobius 125, 129 Tridacna 264, Tab. 7 fear P. 264), Pl. 3 variegata, Clupea 59, 70-72, 81, Pl. 1c variegata, Gudusia 70-72, 81 variegatus, Lithobius - 107, 134, 135 varius, Lithobius . . 3 ; 117 sticardium Tab. 6 (opp. p. 263) velata, Mactroderma Tab. 8 (opp. p. 264) Velorita : . Tab. 14 (opp. p. 269) velox, Lithobius . a 2 - 125 venator, Lithobius 5 127-128 venenosa, Clupea : ° - 61,80 venenosa, Meletta 61 Veneracea 256, 270, Tabs 15 (opp. p: 270), Tab. 22 (opp. p. 284), 289 Venericardia Tab. 4 (opp. p. 260), 261 Venericor Tab. 4 (opp. p. 260) Venerinae Tab. 15 (opp. p. 270), 273 Veneroida . 256-275, Fig. 33 (opp. p. 286), 289 Venerupis Tab. 15 (opp. p. 270), 274 Venus : . Tab. 15 (opp. p. 270), Pl. 7 verhoeffi Lithobius piceus’”. Be ees verrucosa, Venus . Tab. 15 ‘(opp. p- 270) Verticordia . Tab. 21 AeEp. p- 282) Verticordidae é * lai! vicinus, Lithobius 4 , é 3 115 victoriae, Marcusenius . 3, 6, 7, 32 villiosusulca, Thracia Tab, 20 (opp. p. 282) villosus, Lithobius forficatus . é . 120 violacea, Mactra . Tab. 8 (opp. p. 264) virgata, Tellinella Tab. 10 (opp. p. 266) viriatus, Lithobius E e 140 vittatus, Donax Tab. 10 Soca p- 266) Viverra 5 é 221, 240 Viverridae . 5 = : 2 > 240 vulpes, Albula_. f : é a 8 walie, Capra ibex - 246 walleri, Litorcanius 222, 244 Tridacnacea 255, 264, Tab. 7 (opp. p. 264), Tab. 22 (opp. p. 284), 289 Trigonacea . Tab. 22 (opp. p. 284), 289 trigonata, Crassatella Tab. 5 (opp. p. 261) Trigonocardia Tab. 6 (opp. p. 263) Trigonoida . Fig. 33 (opp. p. 286) trilineatus, Lithobius : 120, 121 triliniata, Pandora Tab. 20 (opp. p. 282) truncata, Cyathomonas 4 0 98 truncata, Gastrochaena 277, 280 truncata, Mya 278 earn truncata, Solen Tab. 9 eee: p- 264) Trypanosomatidae 87-102 Tubulidentata. 5 e242 tumidum, Gafrarium Tab. 15 (opp. p. 270) tunicata, Corbula =“ 275 turkestanicus, Lithobius I15, 116 turkestanicus, Lithobius curtipes . A 115 typus, Otomys 5 6 217, 236 undata, Lucinopsis undatella, Chione undulata, Raeta . unedo, Fragum Tab. 8 (opp. p. 264), 265 Tab. 1 (opp. p. 256) 270 Tab. 6 (opp. p. 263) Unionacea 284, Tab. 22 (opp. p. 284), 289 Unionoida 5 a 33 (opp. p. 286) urinaria, Prowazekia 5 97 valida, Tatera 248 valida soror, Tatera 228 validus, Lithobius 112 variegata, Cardita Tab. 4 (opp. p. 260), 261 watkinsi, ere heathi . 192-193 Whitehead, P. J. P . 57-85 whytei, Lepus - *226 whytei fageni, Lepus 226-227 wroughtoni, Lemniscomys striatus | c 233 wroughtoni, Scotophilus 5 fF : 192 Xenomystus 3,5, 7, 18, 21-22, 51 Xerus " . 5 8 bh BF Yalden, D. W. 211-252 zaphiri, Arvicanthis . s - 230 zeylandica, Panopea a, 280, Pl. 9 Zirfaea 281, Tab. 19 (opp. p.. 282), Pl. 10 zoomastigophora: Protozoa zunasi, Sardinella